Arellano y Halffter (2003) Diversidad Gama

Acta Zoologica Mexicana (n.s.
) 90: 27-76 (2003)
GAMMA DIVERSITY: DERIVED FROM AND A DETERMINANT

OF ALPHA DIVERSITY AND BETA DIVERSITY.
AN ANALYSIS OF THREE TROPICAL LANDSCAPES
Lucrecia ARELLANO y Gonzalo HALFFTER
Instituto de Ecología, A.C. Departamento de Ecología y Comportamiento Animal
Apartado Postal 63, 91000 Xalapa, Veracruz, MÉXICO
E-mail: lucreci@ecologia.edu.mx halffter@ecologia.edu.mx
RESUMEN
Utilizando tres grupos taxonómicos en este trabajo examinamos como las diversidades alfa y beta
influyen en la riqueza de especies de un paisaje (diversidad gamma), así como el fenómeno
recíproco. Es decir, como la riqueza en especies de un paisaje (un fenómeno histórico-biogeográfico)
contribuye a determinar los valores de la diversidad alfa por sitio, por comunidad, la riqueza
acumulada de especies por comunidad y la intensidad del recambio entre comunidades. Los grupos
utilizados son dos subfamilias de Scarabaeoidea: Scarabaeinae y Geotrupinae, y la familia Silphidae.
En todos los análisis los tres grupos taxonómicos son manejados como un grupo indicador: los
escarabajos copronecrófagos. De una manera lateral se incluye información sobre la subfamilia
Aphodiinae (Scarabaeoidea), escarabajos coprófagos no incorporados al manejo del grupo indicador.
Los paisajes estudiados son tres (tropical, de transición y de montaña), situados en un gradiente
altitudinal en la parte central del estado de Veracruz. Partimos de las premisas siguientes. La
diversidad alfa de un grupo indicador refleja el número de especies que utiliza un mismo ambiente
o recurso en un lugar o comunidad. La diversidad beta espacial se relaciona con la respuesta de los
organismos a la heterogeneidad del espacio. La diversidad gamma depende fundamentalmente de
los procesos histórico-geográficos que actúan a nivel de mesoescala y está también condicionada
por las diversidades alfa y beta. Es a nivel de paisaje o mesoescala donde las acciones humanas
como cambio y fragmentación de comunidades, tienen sus efectos más importantes, efectos que en
muchas ocasiones escapan al análisis ecológico puntual. En el conjunto de los tres paisajes se
realizaron muestreos regulares en 67 sitios, más muestreos complementarios en 69 lugares más. Se
estudiaron 26 tipos de comunidades vegetales. Se capturó un total de 16,152 ejemplares de 60
especies, 52 especies de Scarabaeinae, 4 de Geotrupinae y 4 de Silphidae. En el paisaje tropical la
comunidad más rica en especies es la selva baja caducifolia; en el paisaje de transición es el bosque
mesófilo. Ambas, son las comunidades naturales más importantes de sus pisos altitudinales. Por el
contrario, en el paisaje de montaña la mayor riqueza se encuentra en praderas y potreros, un tipo de
comunidad favorecido o incluso creado por la intervención humana. Esto se explica por la expansión
a estos lugares de especies heliófilas del Altiplano mexicano. En el paisaje tropical los potreros
presentan una riqueza en especies próxima a la de las selvas, pero una composición parcialmente
diferente, caracterizada por la dominancia de especies heliófilas y coprófagas, a las que se suman
las especies más ubicuistas compartidas con la selva. En el paisaje de transición se puso en relieve
la importancia para la conservación de la fauna del bosque mesófilo, de los cafetales de sombra
poliespecífica. Estos cafetales, el tipo de comunidad con cubierta arbórea que ocupa la mayor
superficie en este paisaje, permiten a los grupos estudiados la intercomunicación entre los fragmentos
remanentes de bosque mesófilo. Para los escarabajos que constituyen el grupo indicador, a nivel de
paisaje (no puntualmente) la fragmentación de las comunidades naturales no parece haber
ocasionado pérdidas en el número de especies. Aparentemente, la perturbación humana ha sido
superada por razones distintas en cada paisaje. En el tropical porque existe una fauna heliófila
característica de los potreros, fauna que incluso ha aumentado con dos especies invasoras recientes.
En el paisaje de transición por el efecto de los cafetales de sombra poliespecífica que crean una
matriz de intercomunicación. En el de montaña porque la expansión de las praderas ha ampliado las
condiciones favorables para las especies heliófilas. Estos resultados no tienen forzosamente que
repetirse con otros grupos de organismos.
Palabras Clave: diversidad alfa, beta y gamma; paisajes antropizados; Scarabaeinae; Geotrupinae;
Silphidae; Veracruz.
27
Arellano & Halffter: Gamma diversity: an analysis of three tropical landscapes.
ABSTRACT
Using three taxonomic groups of beetles we examine how alpha and beta diversity influence the
species richness of a landscape (gamma diversity), and vice versa. That is, how the species richness
of a landscape – which is a historical and biogeographical phenomenon – contributes to the values
of alpha diversity (1) at a given site, (2) in a community, (3) in terms of cumulative species richness
by community, and also contributes to (4) the intensity of species exchange between communities.
To explore this question, we used two subfamilies of Scarabaeoidea: Scarabaeinae and Geotrupinae,
and the family Silphidae. In all analyses these three taxonomic groups are considered as a single
indicator group: the copronecrophagous beetles. Information is also included on the subfamily
Aphodiinae (Scarabaeoidea), coprophagous beetles not included in the indicator group. Several types
of vegetation located in three landscapes (tropical, transition and mountain) were studied, and these
are located along an altitudinal gradient in the central part of the state of Veracruz, Mexico. We base
this study on the following concepts. The alpha diversity of an indicator group reflects the number of
species that use a given environment or resource in a given place or community. Spacial beta diversity
is related to the response of organisms to spatial heterogeneity. Gamma diversity depends primarily
on the historical and geographic processes that operate on the mesoscale level and is also affected
by alpha and beta diversity. It is on this scale of landscape that human actions, such as the
modification and fragmentation of vegetation, have their most important effects. These are, however,
often beyond the scope of ecological analyses carried out on a local scale. In the three landscapes,
sampling was carried out regularly at 67 sites, with complementary sampling at another 69 sites.
Twenty-six types of vegetation communities were studied. A total of 16,152 specimens representing
60 species were captured (52 species of Scarabaeinae, 4 Geotrupinae and 4 Silphidae). In the tropical
landscape the community richest in species was low deciduous forest. In the transition landscape,
cloud forest was the richest. Each of these communities is the most representative of their respective
altitudinal bands. In contrast, the greatest species richness in the mountain landscape occurred in the
mountain grasslands and pastures; types of community favoured by or even created by human
intervention. This is explained by the expansion of heliophilous species from the Mexican High Plateau
into these areas. In the tropical landscape the species richness of the pastures is similar to that of its
forests, but with a partially different composition which is characterized by the dominance of
heliophilous and coprophagous species; the latter, in addition to the more ubiquitous species that are
shared with the tropical forest. In the transition landscape the cloud forest and the coffee plantations
with polyspecific shade are important in the context of conserving the fauna. This type of community
offers arboreal cover and occupies the majority of this landscape, allowing the groups of insects
studied to move between remnant fragments of cloud forest. On the landscape scale but not locally,
the fragmentation of natural communities does not appear to have reduced the number of species for
the beetles of the indicator group. In each landscape disturbance by human activity appears to have
been overcome for distinct reasons. In the tropical landscape we find the heliophilous beetle fauna
characteristic of pastures, and this has increased by two species of recent invaders. In the transition
landscape, the coffee plantations with polyspecific shade create a communication matrix, while in the
mountain landscape the expansion of the mountain pastures has made conditions more favourable
for heliophilous species. These results are not necessarily expected for other groups of organisms.
Key Words: alpha, beta and gamma diversity; anthropogenic landscape; Scarabaeinae; Geotrupinae;
Silphidae; Veracruz.
INTRODUCTION
The most widely used level of biological organization in the study of biodiversity
is the species or organismal level (the last name sensu Harper & Hawksworth
1994). The most compelling reason is that species, in spite of the different criteria
applied by different schools of systematics, are easily detectable and quantifiable
in nature. The number of species of an indicator group that can be found in a
particular site, community or region is a variable that can be measured without
any notable technical or conceptual challenges.
28
Acta Zool. Mex. (n.s.) 90 (2003)
Spatial scale is very important in the evaluation of species diversity since the
processes that influence biodiversity vary with scale. So, at the local or
community level, ecological processes exert the greatest influence: niche
structure, biological interactions, environmental variables, etc. At the regional
level, evolutionary and biogeographical aspects (dispersal, extinction, speciation,
etc.) are the most important. On the mesoscale or landscape scale both sets of
processes affect the number and quality of species (Ricklefs & Schluter 1993).
To study the dynamic relationships among the different types of diversity we
studied three different landscapes along an altitudinal gradient in the central zone
of the state of Veracruz, Mexico (Fig. 1). We worked with three groups of insects
to form a reliable indicator group1. The usefulness of these taxa has been
demonstrated in previous studies (Halffter & Favila 1993, Favila & Halffter 1997,
Halffter 1998 a, b, Halffter & Arellano 2002). They include two subfamilies of
Coleoptera Scarabaeoidea: Scarabaeinae (family Scarabaeidae) and
Geotrupinae (family Geotrupidae), plus a third group of necrophagous beetles,
family Silphidae. The subfamily Aphodiinae (Scarabaeoidea: Scarabaeidae)
forms an important part of the guild of coprophagous beetles and, in the
mountain landscape, replaces part of the Scarabaeinae group. This group was
only sampled occasionally for several reasons, but most importantly owing to the
fact that when this study began, the identification of the Aphodiinae of Mexico
was very difficult. Currently there is more information available, but further study
of this group is still required (M. Dellacasa, com. pers.). In addition, we did not
expect the Aphodiinae to be as numerically important as they turned out to be.
In order to correct this error as far as possible, the species of Aphodiinae
captured during complementary sampling of the landscapes and reports gleaned
from the literature are included in Appendix 1. In the Discussion we use the
information summarized in this Appendix, however it was not included in tables
or in the data presented in the Results section.
The use of landscape units for the evaluation, monitoring and conservation of
the diversity of species has been considered by a growing number of specialists
(Noss 1983, Franklin 1993, McNaughton 1994, Forman & Collinge 1996, Harris
et al. 1996, Miller 1996, Noss 1996, Halffter 1998a,b) as having notable
theoretical and practical value since it allows for the integral analysis of a
complex problem. In a landscape, the composition and number of species
changes from one community to another. This means that the species diversity
of a landscape (gamma diversity) includes the richness of species in the
individual communities that make up the landscape (alpha diversity) and the
degree of difference between those communities (beta diversity). The alpha
diversity of an indicator group reflects the number of species that use a given
environment or resource. Beta diversity represents the response of the
1) The term “indicator group is used in different contexts in ecology: to designate groups that reveal
the occurrence of disturbance (such as the stage) or for groups that allow us to estimate the species
richness of a site or landscape. As Favila and Halffter (1997) remark, “... the function of the indicator
group is to make possible the approximation of an answer to a complex and labourious problem, that
of measuring and monitoring total biodiversity". It is in this sense that the term indicator group is used
in the present study.
29
organisms to spatial heterogeneity. Gamma diversity depends primarily on

historical and evolutionary processes that operate on the mesoscale level and is
also affected by alpha and beta diversity (Whittaker 1972, 1977). It is on the
landscape or mesoscale level that human activity (contamination, degradation,
change and fragmentation of ecosystems) has effects that often elude ecological
analysis carried out on a local scale (Halffter 1998a, b). Modifications caused by
human activities have resulted in the creation of landscapes that are complex
mosaics of primary vegetation, secondary communities, pastures, annual and
perennial crops. These landscapes represent a challenge for the study of
biodiversity.
Veracruz is one of the three states in Mexico with the greatest biological
diversity, but it is also one of the areas in the country with the highest degree of
anthropogenic disturbance. In Veracruz, the main consequence of human activity
has been the fragmentation of natural communities such that what remains are
patches of the original vegetation with differing degrees of modification and that
retain some of the components of the original forests (for Scarabaeinae see
Halffter et al. 1992, Halffter & Arellano 2002). The central part of Veracruz is a
region where the disturbance occurred a long time ago and so it is difficult to find
anywhere that has not been disturbed. Before the Spanish Conquest this area
was densely populated by indigenous peoples who, among other activities,
practiced slash and burn agriculture. From the 16th century onwards, sheep
ranching (practically abandoned today) and cattle ranching (Barrera-Bassols &
Rodríguez 1993) were added to this form of agriculture and then came the
cultivation of sugar cane (Prieto 1968, Rodríguez 1970, Ponce & Núñez 1992).
In the anthropogenic landscapes of central Veracruz our research team is
carrying out various projects on biodiversity related to the present study. With
dung beetles (Scarabaeinae and Geotrupinae) and Silphidae we have studied the
biogeographical elements of regional biodiversity (Halffter et al. 1995). Using only
Scarabaeinae we have compared the biodiversity of different tropical rainforests
with that of an induced pasture (Halffter et al. 1992). In addition, we have
examined the relative importance of arboreal cover with respect to food
availability and its influence on species diversity and the structure of the
Scarabaeinae guild (Halffter & Arellano 2002). Using bats as an indicator group,
we have also analyzed the relationship between the different levels of diversity
(Moreno & Halffter 2001).
Using Scarabaeinae, Geotrupinae and Silphidae to form an indicator group in
this study we examine how alpha and beta diversity influence species richness
(gamma diversity) in three landscapes of central Veracruz. We also explore the
inverse phenomenon. That is, how the species richness of a landscape – a
historical, biogeographical element affected by anthropogenic transformations,
mainly community fragmentation – contributes to determining the values of alpha
diversity of a given site, of a given community, and the cumulative species
richness of a community, as well as the intensity of species exchange between
communities.
30
MATERIALS AND METHODS
Some definitions
In the interest of clarity, we present the meaning of some of the terms used in
this manuscript, including several terms generally used in ecology.
Alpha diversity is the number of species (belonging to the indicator group we
have defined for our work) found at a site or in a community. Although not
specified in the majority of studies on biodiversity, alpha diversity corresponds to
the number of species that exist over a short period of time during which the
number of species does not change. The rate of change in the number of species
at a given site varies greatly over longer periods of time owing to emigration, the
local extinction of some species and the arrival of others previously not present.
Alpha diversity can be expressed for a site or a community type, and in both
cases the total number of species or mean values can be used. The mean value
is the arithmetic mean of the values recorded for the sites of a given landscape
or at the sampling sites of a certain type of community.
The cumulative species richness of a community is the sum of the number of
species belonging to the indicator group collected in that community over a given
period of time. Beta diversity is the difference in the number of species between
two sites, two types of community or two landscapes. This difference could
originate from available space and changes associated with space when the
diversity of the places being compared is obtained simultaneously. It could also
result from comparing alpha diversities that were obtained at different times.
Gamma diversity is the total number of species recorded for the group of sites
or communities that make up a landscape.
Community refers mainly to different types of vegetation, but in some cases,
also to areas which have been visually characterized for practical purposes.
Pasture refers to deforested areas where grass has been sown (or grown
spontaneously) and used for livestock, almost exclusively cattle, in the region.
A site is the minimum area, in terms of space and time, that has a sample of
a given functional assemblage or community (definition as given by G. Halffter
and C.E. Moreno, pers. com.). For a study such as this, it is the same as “point”
or “place”.
High altitude vegetation includes all the community types found above 3,500
m asl. The use of this term is general since the beetles of the indicator group are
scarce (both in species and individuals) in this type of vegetation and show no
marked affinity towards any given community at these altitudes. The most
extensive community types at these altitudes are: pine forest (Pinus hartwegii) with
Juniperus monticola and Berberis schiedeana in the shrub layer and Muhlenbergia
macroura, Stipa ichu, Trisetum spicatum and Calamagrostis tolucensis in the herbaceous
understorey. This type of vegetation is found between 3,500 and 4,000 m asl.
Mountain grassland, found between 4,000 and 4,200 m asl, is a predominantly
herbaceous community that has the shrub species mentioned above, but with
species different to those found in the pine forest (see Narave 1985).
31
Landscape units, sampling sites and capture methods

A landscape is a unit of space, with geographic limits and a specific area,
characterized by its relief, climate, soil and vegetation (Noss 1990, Castillo-
Campos 1995, Geissert & Castillo-Campos 1997). Each landscape has its own
biogeographical history (Halffter 1998 a,b) which emerges as a result of two
types of processes that are constantly interacting: ecological or current, and
historical-evolutionary; the latter referring as much to the biota as to the physical
components of the landscape. Of the current processes that determine the
characteristics of each species assemblage, some are natural (occurring without
direct human intervention) and some are the direct or indirect, desired or
undesirable, result of human activity.
In agreement with generally accepted criteria for climatological and
physiographical limits (INEGI, SEDUE), for this study we have classified the
landscapes as three successive levels along an altitudinal gradient that ranges
from sea level to 4,250 m asl in the central region of the State of Veracruz (19º
90'-19º 25' lat. N, 96º30'- 97º20' long. W) (Fig. 1). The delineation of the
landscapes was compared with site classification by species composition using
the program Statistica (Statsoft 1991, see Fig. 2), and refined using information
from a study by Halffter et al. (1995), which included a biogeographical analysis
of copronecrophagous beetles from the central region of Veracruz. The
landscapes are:
a)Tropical. 0-1,000 m asl. Coastal plains. Soil: feozems, luvisols and rendzinas
(Zolá 1987). Characterized by limestone, sandstone and rocks of volcanic
origin (INEGI 1988). Climate: Aw'(i)g, warm, subhumid with rain during the
summer (García 1981). Mean annual temperature ranges from 22.3°C to
24.5°C. Total annual precipitation: 1,500 - 2,000 mm. Vegetation communities:
coastal sand dunes, mangrove, halophyte vegetation in the swales of fixed
dunes and around mangroves, deciduous forest on the more elevated terrain,
tropical oak forests, medium semi-deciduous forest in the most humid ravines
(Castillo-Campos 1985, 1991, Acosta 1986, Robles 1986, Cházaro-Basáñez
1992). Land use: agriculture with irrigation, sugar cane and mango cultivation,
extensive cattle ranching.
b)Transition. 1,000 - 2,000 m asl. Terrain with tablelands and hill and valley
formations. Soils: andosols, lithosols and feozems (INEGI 1988), on volcanic
ash, basalt, cinder, lapilli and andesites (Zolá 1987). Climate: on the border of
two climate types, C(fm)w"b(i)g, humid temperate with rain year round, and
(A)C(fm)w"a(i)g, semi-warm humid (Soto & Angulo 1990, Angulo 1991). Mean
annual temperature ranges from 12.26°C to 22.3°C. Total annual precipitation:
1,200 - 2,500 mm. Vegetation communities: medium altitude oak forest, cloud
forest, oak forest, pine-oak forest (Castillo-Campos 1991, Zamora 1992,
Narave 1985). Land use: predominantly coffee plantations, but also corn
cultivation and dairy cattle ranching.
c) Mountain. > 2,000 m asl. Soils: andosols (De Luna 1983). The most common
rock types are volcanic ash, basalt and andesites. Climate: C(fm) humid
temperate with rain year round, C(m) humid temperate with rain during the
32
summer, CW2 subhumid temperate with rain during the summer, CW1" dry and
cool subhumid temperate (Soto & Angulo 1990). Mean annual temperature
ranges from 11.04°C to 12.85°C. Total annual precipitation: 800 - 1,500 mm.
Vegetation communities: pine forest, fir forest and mountain pastures (Narave
1985). Land use: mainly dairy cattle ranching, but also seasonal agriculture
(corn, wheat, potato, oats, etc.) with apple, pear and plum orchards (Narave
1985).
Part of these landscapes is covered by an extensive zone of rocky lava flows

(type Aa) formed by numerous basalt extrusions originating from a string of small
volcanos (such as El Volcancillo, Xocotepec and La Joya) located along the lava
flow (Geissert 1994). We sampled at sites located on: a) the La Joya-Acajete flow
which is estimated to have occurred no more than 36,000 years ago and is
covered by a layer of ash that is continuous and of varying thickness; b) an
extrusion covered by a discontinuous layer of ash of varying thickness that is
5,000 to 10,000 years old, begins near La Joya and extends towards Rafael
Lucio, El Duraznal, Teapan and Jilotepec; c) the chaotic rocky lava flows which
begin at the Volcancillo and end near Actopan, more than 50 km from their origin
(Fig. 3). The latter are recent; less than 5,000 years old according to C14 dating
(Geissert 1994). At most this formation is 2,400 years old (Ortega 1981; for more
information see Negendank et al. 1985).
Based on site classification, for the analysis of the alpha and beta diversity we
decided to separate the communities located on the rocky lava flows from those
found on soils that are not rocky since, in most cases, these communities formed
separate groups (Fig. 2).
In the tropical landscape, samples were taken at 18 sites and complementary
samples at 20 additional sites. Five community types were studied. In the
transition landscape samples were taken regularly at 29 sites and additional
sampling carried out at 30 other sites. Thirteen community types were studied.
The heterogeneity in vegetation of this landscape required a greater sampling
effort. In the mountain landscape samples were taken at 20 sites and
complementary sampling was carried out at 19 additional sites. Eight community
types were studied. Figure 1 shows the locations of the sampling sites.
For regular sampling, we baited traps with human excrement, carrion
(decomposing squid) or fruit (mango, guava, mamey), and specimens were also
collected from small cadavers, fruit, wild animal scat and livestock dung (cattle,
horse and sheep). During regular sampling we placed baited traps at each site
each month for a 24 hour period. The bait was placed on the surface of four traps
(two with cow dung and two with decomposing squid), and at the bottom of six
traps (three with human excrement and three with rotting squid). The designs of
the pitfall traps are described in Halffter and Arellano (2002). Regular sampling
covered a total of 723 capture days with the participation of two people per day:
190 days in the tropical landscape, 375 days in the transition landscape and 158
days in the mountain landscape. Sampling was carried out mainly during the
rainy season (May to October) of 1990, and from 1994 through 1996.
33
Figure 1
Location of the region studied in Veracruz State, Mexico. The circles represent the regular sampling sites and the
triangles represent complementary or occasional sampling sites.
34
Figure 2
Dendrogram of the cluster analysis done of the data from the regularly sampled sites. Similarity was generated
using the Euclidian distance (converted to percentage) and the unweighted pair group method with arithmetic
average linkage rule (UPGMA) was used. The three landscapes separate into three large groups of sites. The
names of the vegetation communities are indicated by their abbreviations in letters. See the reference list.
Dlink/Dmax is a standarization of the Euclidean Distance to 0-100 scale.
For the complementary sampling, beetles were only collected from excrement.
This occasional sampling allowed us to complete the list of species for a given
landscape. As complementary methods in forests and coffee plantations, we
used a NTP 80 trap (Morón & Terrón 1984) that was checked every 30 days. In
pastures, specimens were collected from ten dung piles (mainly cow and horse).
During the occasional sampling (1989-1997) specimens were only collected from
excrement that was encountered, and these were used to complete species lists
for each landscape.
Data analysis
To evaluate sampling efficiency for each site, species accumulation models
and Abundance-base Coverage Estimator (ACE), non-parametric estimators of
species richness, were used (Colwell 1997). The unit of effort used was a day of
sampling, including all methods of capture, excluding the data from occasional
or complementary sampling.
35
Abbreviations used for plant communities in figures, tables & appendices:

Plant Community abbreviations
Tropical Landscape
Tropical deciduous forest tf
Pasture adjacent to tropical deciduous forest pt
Secondary vegetation adjacent to tropical deciduous forest svt
Pasture adjacent to tropical deciduous forest on rocky lava ptr
flow
Tropical deciduous forest on rocky lava flows tfr
Transition Landscape
Cloud forest cf
Coffee plantation with polyspecific shade cp
Pasture adjacent to pine-oak forest on rocky lava flows ppor
Pasture adjacent to cloud forest pc
Oak forest of
Pasture on rocky lava flows, adjacent to pine-oak forest prpo
Coffee plantation with monospecific shade cm
Oak forest on rocky lava flows ofr
Pine-oak forest on rocky lava flows por
Pine-oak forest pof
Gap in cloud forest gc
Secondary vegetation adjacent to oak forest svo
Pasture on rocky lava flow, adjacent to oak forest prof
Mountain Landscape
Grassland adjacent to pine forest gp
Pine forest pf
Pine-alder forest pa
Grassland adjacent to fir forest gf
Fir forest ff
Grassland adjacent to pine-alder forest gpa
Pine forest on rocky lava flows pfr
High altitude vegetation hav
Randomized data (100 times, using EstimateS Colwell 1997) were adjusted to
two asymptotic models: the linear dependence model and Clench’s model (see
Soberón & Llorente 1993, León-Cortés 1994, León-Cortés et al. 1998) using
SIGMASTAT (Jandel 1995). We did not use the results obtained from the Linear
Dependence Model because it underestimated richness values. Only the
asymptote values obtained with Clench’s model were used to to select those of
the 67 regularly sampled sites where capture effort provided close to the
maximum diversity expected. 85% of the total estimated fauna, though arbitrary,
36
is considered an acceptable degree of efficiency. When the data did not meet the
criteria of the accumulation model, the ACE non-parametric estimators proposed
by Colwell & Coddington (1995) was calculated applying the criterion of 85% for
the selection of adequately sampled sites. In the end, 56 sites were selected for
diversity analysis. These analyses were also carried out at the level of community
type for each landscape.
The folloling were calculated: alpha diversity for each site, mean alpha diversity
for each community type, mean alpha diversity for each landscape, and
cumulative species richness for each community.
Figure 3
The relatively exposed (50-70%) rocky lava flow Volcancillo-Actopan, in the region of central Veracruz, Mexico.
Modified from Ortega (1981).
Beta diversity was calculated for all pairs of sites from the same community,
after which mean beta diversity per community was obtained. Beta diversity was
also calculated for all pairs of sites from adjacent communities, these values
indicating the highest exchange rates. Whittaker’s modified index was used
(Harrison et al. 1992):
β = {[(S/a)-1]/(N-1)} x 100
where:
S = the number of species recorded for the landscape (gamma diversity)
a = the mean number of species in a given community type (mean alpha
diversity)
N = the number of study sites
37
This index is similar to that of Whittaker (1972), except that it is expressed as

a percentage. The closer the value is to zero, the more similar communities are.
Higher values indicate greater differences (less similarity) between communities.
Using the program Orden (Ezcurra 1990), an uncentred, unstandardized Principal
Components Analysis (56x58) was conducted to graph the trends in beta
diversity. Sites that have a similar number of species and species composition
are closer together (Ter Braak 1983, Cody 1993). It is important to note that
estimation of real variability of axes (with respect to means) was calculated by
eliminating the first Eigen value, because in uncentered presence/absence
analysis, Axis 1 is affected by the decision not to center data, and data variability
increases artificially (Montaña & Ezcurra, 1991).
The gamma diversity for each of the three landscapes, measured as the
cumulative number of species captured, was calculated using the following index
proposed by Schluter & Ricklefs (1993):
γ = α x isd x sd
where:
α = the mean number of species per site in a landscape unit,
isd= the inverse of the species dimension; that is, 1/the mean number of
communities or locations occupied by a species,
sd = sample dimension or total number of sites sampled.
Gamma diversity can be obtained from the general species list for each
landscape. However, the use of the formula above gives us an idea of which
component of gamma diversity is the most important in each landscape, whether
it is the mean alpha diversity, landscape heterogeneity or the number of
communities occupied by species. This allows us to compare the different
components of gamma between landscapes.
To estimate the degree of dissimilarity in the species composition of the
landscapes, we calculated complementarity between pairs of landscapes (Colwell
& Coddington 1995). The complementarity for landscapes A and B is expressed
as:
CAB= UAB/SAB
where UAB is the sum of the species unique to each of the two landscapes,
calculated as:
UAB = a + b- 2c
with:
a is the number of species of landscape A,
b is the number of species of landscape B,
c is the number of species in common to landscapes A and B
and where SAB is the total species richness of both landscapes combined, as
follows:
SAB = a + b- c
For the data on the cumulative richness and gamma diversity of tropical low
deciduous forest and the pastures adjacent to them, we included species
captured by G. Halffter at Laguna Verde, a site close to the Gulf of Mexico where
our group has carried out studies over the past 20 years and where more than
95% of the existing species have been collected (see Appendix 2).
38
RESULTS
A total of 16,152 specimens belonging to 60 species were captured from the

67 sites that were regularly sampled. Of these, 52 species belong to
Scarabaeinae, 4 to Geotrupinae and 4 to Silphidae. Of the total 15,716
individuals belonging to 60 species were caught in the 56 sites selected for
diversity analysis.
The different estimates of sampling quality by community type were
satisfactory: more than 85% of the estimated species were caught (see tables 1
and 2).
Table 1
The number of copronecrophagous beetle species recorded, as well as the parameters and
predictions of Clench’s asymptotic accumulation model fitted for each plant community from
three landscapes in central Veracruz. a is the slope at the beginning of sampling, b is a
parameter related to the shape of the curve indicating the accumulation of new species
during sampling, a/b is the asymptote, R2 is the coefficient of determination, n is the number
of samples for each community and % is the percentage of the asymptote recorded. See
abbreviations list for the complete names of the plant communities.
Community Clench’s Model
spp. n R2 a b a/b %
obs.
Tropical landscape
tfr 7 14 999 2105 258 8 8750
pt 14 51 1000 3788 253 15 9333
svt 17 19 999 3793 195 19 8947
Transition landscape
cp 16 63 992 2476 145 17 9411
cm 10 23 814 2536 242 10 10000
gc 8 22 999 1552 169 9 8888
of 11 16 997 4030 385 10 10000
svo 7 8 997 1064 1356 8 8750
prpo 11 12 997 294 230 13 8462
pc 12 40 972 1338 95 14 8571
pof 8 13 846 3621 398 9 8888
ppor 13 34 990 2845 210 14 9286
prof 6 18 995 2733 392 7 8571
ofr 9 8 997 3222 318 10 9000
Mountain landscape
pa 5 7 999 3421 687 5 10000
ff 3 7 1000 1272 371 3 10000
gpa 5 7 996 1428 259 6 8333
gf 2 7 967 1819 816 2 10000
pfr 2 7 995 2662 1202 2 10000
gp 16 54 989 1117 68 16 10000
hav 2 17 994 277 120 2 10000
39
Table 2
The number of copronecrophagous beetle species recorded and prediction of nine non-
parametric estimators of species richness (Colwell & Coddington 1995) for some plant
communities of central Veracruz, Mexico.
Tropical Transition Landscape Mountain
Landscape Landscape
Community tf cf por pf
No. of samples 91 71 29 53
No. of species 19 18 8 8
observed
ACE 20 18 8 8
ALPHA DIVERSITY
Alpha diversity by site

Species distribution in each site and community, as well as the values for alpha
diversity are given in Appendix 3.
In the tropical landscape, alpha diversity on sites 1, 2, 3, 4, 5 and 6 was
significatively higher (Kruskal, H = 73.11, d.f. = 15, P < 0.001, Dunnet’s Method,
P < 0.05) than on sites 7 and 8, two tropical forests growing on the fairly exposed
rocky lava flows. In the transition landscape there were significant differences
among the alpha diversity of sites with 5-7 species - sites with pine-oak forest on
rocky lava flows, monospecific coffee plantations, pastures and secondary
vegetation- and the others with higher alpha diversity (Kruskal, H = 118.43, d.f.
= 23, P < 0.001, Dunnet’s Method, P < 0.05). In the mountain landscape
differences were observed (Kruskal, H = 82.03, d.f. = 15, P < 0.001, Dunnet’s
Method, P < 0.05), as at 3,900 m asl and higher, none of the species belonging
to the groups studied were found. (For biogeographical analysis explaining the
presence of these beetles at these altitudes, see Halffter et al. 1995, Peck &
Anderson 1985).
Mean alpha diversity by community

There were significant differences in the mean alpha diversity of communities
among landscapes (ANOVA, F = 17.21, d.f. = 2, P < 0.001). The values for the
communities of the transition and tropical landscapes were similar (Tukey, q =
1.382, P > 0.05), while those of the mountain and tropical landscape (Tukey, q =
7.021, P < 0.05), and those of the transition and mountain landscape (Tukey, q
= 7.289, P < 0.05) were not.
The effect of substrate type and the age of the rocky lava flow sites
In the tropical landscape there were significant differences in number of
species (t = 3.936, d.f. = 6, P = 0.008), and in number of individuals (t = 3.035,
d.f. = 6 , P = 0.007) between the tropical forests growing on the fairly exposed
40
rocky lava flows (< 5,000 years old) and those growing on other types of soil.
Significant differences in the number of species (Mann-Whitney, Tx = 658.0, P =
0.02) and in the number of individuals (Mann-Whitney, Tx = 682.0, P = 0.05) were
also found between the forests on the rocky lava flows and the pastures adjacent
to them. These differences were notable at La Concepción where only 5 species
and 42 individuals were found in the forest, but 10 species (277 individuals) were
caught in the adjacent pasture. There were no significant differences in number
of species (t = 0.804, d.f. = 4, P = 0.466), and in number of individuals (t = 0.887,
d.f. = 4, P = 0.425) between pastures adjacent to tropical forests growing on the
fairly exposed rocky lava flows and the other sites.
For the transition landscape, there were no significant differences in the
number of species caught in forests located on the relatively older (5,000 -
10,000 year old) soils of the rocky lava flows and those on other soils (Mann-
Whitney, Tx = 13, P = 0.381), nor were there differences in the number of
individuals (t = 0.752, d.f. = 4, P = 0.494). There were no significant differences
in the number of species collected in the forests on the rocky lava flows and their
adjacent pastures (Mann-Whitney, Tx = 2117.00, P =0.709), nor in the number
of individuals (Mann-Whitney, Tx = 2474.5, P = 0.730). In spite of this, we did
record marked differences between a pine-oak forest growing on the
predominantly rocky soil of a lava flow (Teapan) and its adjacent pasture, both
in the number of individuals (forest = 139, pasture = 224) and in the number of
species (forest = 6, pasture = 11). In the transition landscape, for sites that occur
on an ancient lava flow (36,000 years old) where the soil is not very rocky, there
were no significant differences between the forest and the adjacent pasture in the
number of species (Mann-Whitney, Tx = 210.0, P = 0.359), or in the number of
individuals captured (Mann-Whitney, Tx = 135.5, P = 0.781).
For the mountain landscape there were no significant differences in the number
of species (t = 2.079, d.f. = 6, P = 0.083) or in the number of individuals captured
(t = 0.609, d.f. = 6, P = 0.565) between forests located on the predominantly
rocky lava flows (5,000+ years old, Geissert 1994) or on other soils.
In the mountain landscape, there were no significant differences in the number
of species (Mann-Whitney, Tx = 240, P = 0.372), or in the number of individuals
(Mann -Whitney, Tx = 244, P = 0.378) between the forest and the pastures that
grow on the rocky lava flows.
Cumulative species richness (cumulative alpha diversity) of the vegetation

communities
Table 3 shows the cumulative richness of the different community types. Low
deciduous tropical forest had the greatest number of species (22), followed by
pastures adjacent to these forests and the lowest values (< 3 species) were
recorded for high altitude vegetation and pine forest on the rocky lava flows.
For the three landscapes there were significant differences among
communities in mean alpha diversity and cumulative species richness (t = -2.883,
P = 0.007). In various communities cumulative species richness was
approximately double the mean alpha diversity (Table 3).
41
Table 3
Mean alpha diversity and cumulative community species richness (CSR) of
copronecrophagous beetles in three landscapes of central Veracruz, Mexico.
Mean Alpha min max
Community Diversity/ no. no. CSR
community spp. spp.
± S.E.
Tropical Landscape
low deciduous forest 11.33 ± 0.97 8 15 22
pasture next to low deciduous forest 11.25 ± 0.84 9 14 21
secondary vegetation next to low deciduous forest 11.50 ± 2.00 9 13 17
pasture next to low deciduous forest on rocky lava flows 9.5 ± 2.50 7 12 14
low deciduous forest on rocky lava flows 4.5 ± 0.50 4 5 7
cloud forest 11.67 ± 0.88 10 13 18
coffee plantation with polyspecific shade 12.50 ± 0.50 12 13 16
pasture next to pine-oak forest on rocky lava flows 8.33 ± 1.76 5 11 13
pasture next to cloud forest 8.5 ± 1.50 7 10 12
oak forest 9.00 ± 1.00 8 10 11
pasture on rocky lava flows, next to pine-oak forest 11.00 ± 0.00 11 11 11
coffee plantation with monospecific shade 6.50 ± 1.50 5 8 10
oak forest on rocky lava flows 9.00 ± 0.00 9 9 9
pine-oak forest on rocky lava flows 5.67 ± 0.33 5 6 8
pine-oak forest 8.00 ± 0.00 8 8 8
gap in cloud forest 7.00 ± 1.00 6 8 8
secondary vegetation next to oak forest 7.00 ± 0.00 7 7 7
pasture on rocky lava flows, next to oak forest 6.00 ± 0.00 6 6 6
Mountain Landscape
grassland next to pine forest 5.75 ± 0.96 3 7 13
pine forest 3.25 ± 0.63 2 5 7
pine-alder forest 5.00 ± 0.00 5 5 5
grassland next to fir forest 2.00 ± 0.00 2 2 2
fir forest 3.00 ± 0.00 3 3 3
grassland next to pine-alder forest 5.00 ± 0.00 5 5 5
pine forest on rocky lava flows 2.00 ± 0.00 2 2 2
high altitude vegetation 0.67 ± 0.33 1 1 2
BETA DIVERSITY
Shared and exclusive species

The number of species shared among the different community types for each
landscape was high (Table 4). In the tropical landscape the forests had the
highest percentage of exclusive species (27.27%), followed by pastures adjacent
to forests (14.29%) and pastures next to forests on the rocky lava flows (7.14%).
Neither the forests on the rocky lava flows, nor the sites with secondary
vegetation next to forests had any exclusive species.
In the transition landscape the greatest percentage of exclusive species was
found in coffee plantations with polyspecific shade (12.5%), followed by oak
forests (9.09%), pastures next to cloud forest (8.33%) and cloud forests (5.55%).
42
None of the other communities had any exclusive species (Table 4).
The communities with the highest number of exclusive species were found in
the mountain landscape. In this landscape, unlike the others, no community
shared 100% of its species with another (Table 4).
Table 4
Total number of species, number of exclusive species and shared species in each
community type for the three landscapes studied. Community designation (first column)
follows the text. In the last four columns on the right, all the community types have been
grouped into four major community types. The dashes indicate the same community type
in the column and the row (no comparison possible) or that no samples were collected from
this community type in the corresponding landscape.
Number of species shared with
total # of # of
other other secondary coffee
Community species exclusive
forests pastures veg + gaps plantations
spp.
.
Tropical Landscape
low deciduous forest 22 6 6 13 13 -
pasture next to low deciduous forest 21 3 13 13 12 -
secondary vegetation next 17 0 13 12 - -
to low deciduous forest
pastures next to low deciduous forest on 14 1 9 13 10 -
rocky lava flows
low deciduous forest on rocky lava flows 7 0 6 6 5 -
cloud forest 18 1 13 10 8 16
oak forest 11 1 8 7 7 6
oak forest on rocky lava flows 9 0 9 6 - 9
pine-oak forest 8 0 8 8 - 3
pine-oak forest on rocky lava flows 8 0 9 8 - 5
coffee plantation with monospecific 10 0 10 8 - 8
shade
coffee plantation with polyspecific shade 16 2 14 11 - 8
pastures next to cloud forest 12 1 10 4 - 10
pastures on rocky lava flows, next to oak 6 0 3 6 3 5
forest
gap in cloud forest 8 0 8 4 - 8
secondary vegetation next to oak forest 7 0 7 2 4 5
pastures next to pine-oak forest on rocky 13 0 11 13 9 5
lava flows
pastures on rocky lava flows, next to 11 0 9 10 - 5
pine-oak forest
Mountain Landscape
pine forest 7 2 - 4 - -
pine-alder forest 5 2 - 3 - -
pine forest on rocky lava flows 2 2 - - - -
fir forest 3 1 - 2 - -
pastures next to pine forest 13 9 - 4 - -
pastures next to pine-alder forest 5 1 - 2 - -
pastures nect to fir forest 2 2 - - - -
high altitude vegetation 2 2 - - - -
43
Mean beta diversity among vegetation communities

The values of mean beta diversity among communities showed different
tendencies depending on the landscape. In the tropical landscape, the greatest
mean beta diversity was found between the tropical deciduous forests that grow
on rocky lava flows (77.5%), followed by the species exchange between different
types of tropical deciduous forests and their adjacent pastures (>68%). Beta
diversity between deciduous tropical forests and their secondary vegetation was
53.25%. This is reflected in the Principal Components Analysis where axis 2
separates deciduous tropical forests from their adjacent secondary vegetation
and from neighbouring pastures (Fig. 4). The lowest value for mean beta diversity
was recorded between different sites with secondary vegetation, between
deciduous tropical forests or between pastures (Table 5).
Figure 4
Uncentred, non-standardized Principle Components Analysis of sites with more than 85% of the estimated fauna.
The names of the vegetation communities are indicated by their abbreviations in letters. See the reference list.
44
Table 5
Mean beta diversity by community for three landscapes in the Cofre de Perote region of central
Veracruz, Mexico. β = Whittaker’s index (modified), S.E. = standard error. A value of “indef” for
S.E. is given when the beta value was obtained by comparing only one pair of sites. Mean beta
diversity for patches with the same type of community are expressed to determine the
heterogeneity of the most typical communities for each landscape. Mean beta diversity is also
given where different community types are adjacent and the type of contiguity specified is the
most common for that landscape (e.g. forests cf. pastures).
Community Beta Diversity

β (%) Std min max
error
Tropical Landscape
low deciduous forests 3733 343 1000 6470
pasture next to low deciduous forest 3902 302 2220 5455
secondary vegetation 3000 indef. 3000 3000
low deciduous forests on rocky lava flows 7750 250 7500 8001
pastures next to low deciduous forest on rocky lava flows 4440 indef 4440 4440
low deciduous forests cf. pastures 6861 1464 4290 10000
low deciduous forests on rocky lava flows cf. pastures 6957 1263 4444 10008
low deciduous forests cf. adjacent secondary vegetation 5326 1464 4290 6361
cloud forests 3737 682 2380 4540
coffee plantation with monospecific shade 5385 indef. 5385 5385
coffee plantation with polyspecific shade 2727 indef. 2727 2727
oak forests 1579 indef. 1579 1579
cloud forests cf. oak forests on rocky lava flows 3547 730 2220 4740
cloud forests cf. oak forests 4764 416 3330 5790
pine-oak forests on rocky lava flows 3485 758 2727 5000
pine-oak forests on rocky lava flows cf. pine-oak forests 3161 1334 770 5380
pastures next to cloud forest 2860 indef. 2860 2860
gaps in cloud forest 1428 indef. 1428 1428
coffee plantation with monospecific shade cf. those with polyspecific 4641 494 3333 5550
shade
pastures next to pine-oak forest 4737 135 4550 5000
pine-oak forests cf. pastures 3826 656 2730 5000
cloud forests cf. pastures 5555 1111 4444 6666
oak forests cf. adjacent secondary vegetation 1761 indef. 1760 1760
pine-oak forest on rocky lava flows cf. pastures 3240 510 2730 3750
cloud forests cf. coffee plantations with monospecific shade 5555 indef. 5555 5555
cloud forests cf. coffee plantations with polyspecific shade 1430 indef. 1430 1430
oak forests cf. adjacent pastures on rocky lava flows 6250 indef. 6250 6250
cloud forests cf. gaps 1580 indef. 1580 1580
Mountain Landscape
pine forests 3059 631 1130 5000
pine forests cf. grasslands 5777 1309 4000 8333
pine forests cf. pine forests on rocky lava flows 3292 715 1500 5000
pine forests cf. pine-alder forests 3571 937 1111 5550
pine forests cf. fir forests 4718 428 4290 6000
grasslands next to pine forest 6392 799 4660 10000
grasslands next to pine forest cf. grasslands next to pine-alder forest 3713 625 2000 5000
grasslands next to pine forest cf. grasslands next to fir forest 7920 829 6000 10000
high altitude vegetation 10000 indef. 10000 1000
45
In the transition landscape the greatest mean beta diversity occurred between
oak forests and neighbouring pastures on the rocky lava flows (62.50%), between
cloud forests and coffee plantations with monospecific shade (55.55%), and
between cloud forests and their adjacent pastures (55.55%). This is seen (Fig.
4) in the PCA where, on axis 3, cloud forests are separated from their pastures
and from a coffee plantation with monospecific shade. The latter was the poorest
in species (5) and had been subjected to treatment with herbicides. This coffee
plantation had 38% fewer species than the other plantation examined (also with
monospecific shade) in this study. The PCA also shows that the beta diversity of
oak forest on rocky lava flows is closer to that of cloud forests and their gaps than
to that of other oak forests. This can be explained by the fact that the oak forest
on the rocky lava flows shares two thirds of its species with cloud forests and
their gaps, but only 44% of its species with other oak forests above 1,600 m asl.
The communities with the greatest similarity in their composition were cloud
forests and coffee plantations with polyspecific shade, cloud forests and their
gaps, and oak forests and their adjacent secondary vegetation (Table 5, Fig. 4).
Cloud forests and their gaps, and the coffee plantations with polyspecific shade
form a small group on axis 3 of the PCA (Fig. 4). Axes 2 and 3 explain 29.86%
of the total variance.
GAMMA DIVERSITY
In the tropical landscape gamma diversity was 31 species, 30 Scarabaeinae

and one Silphidae. Gamma diversity in the transition landscape was 30 species,
26 Scarabaeinae and four Silphidae. In the mountain landscape, gamma diversity
was 17 species, 11 Scarabaeinae, four Geotrupinae and two Silphidae (see
Appendix 2, Appendix 3, and percentages of species exclusive to each
landscape in Table 6).
Table 6
Gamma diversity of copronecrophagous beetles for three landscapes in central Veracruz,
Mexico. Exclusive species, shared species and complementarity (excluding Aphodiinae).
Landscape
Indicators of Gamma Diversity Tropical Transition Mountain
Total # of species 31 30 17
% species exclusive to the
landscape 61.29 40 47.06
% species shared
with the Tropical landscape - 29.03 3.22
with the Transition landscape 29.03 - 26.67
with the Mountain landscape 3.22 26.67 -
% complementarity
with the Tropical landscape - 82.69 98.87
with the Transition landscape 82.69 - 79.49
with the Mountain landscape 98.87 79.49 -
46
There were significant differences in the number of sites where each species
was found in each landscape (Kruskal, H = 7.606, P = 0.022). In the mountain
landscape, species were distributed in many fewer sites than in the transition
(Dunnet’s Method, q’ = 2.65, P < 0.05), or the tropical landscape (Dunnet’s
Methos, q’ = 2.279, P < 0.05).
From the general list of species (Appendix 3), some of the ubiquitous species,
such as Dichotomius colonicus, Onthophagus hoepfneri, Onthophagus chevrolati retusus
and Oxelytrum discicolle, were collected from more than one landscape. Euoniticellus
intermedius, an invader species that has only been in the region for a few years
(first record in 1995, see Montes de Oca & Halffter 1998), was found in the
pastures of the tropical and transition landscapes from sea level up to 1,400 m
asl.
The species composition of the tropical landscape was very different to that of
the other two. This is seen in the Principal Component Analysis (Fig. 4). The
complementarity of the tropical and mountain landscapes was 98.87%, for the
tropical and transition landscapes it was 82.69%, and for the transition and
mountain landscapes it was 79.49%.
The tropical landscape shared 29.03% of its species with the transition
landscape and only 3.22% with the mountain landscape. The tropical landscape
had the highest proportion of exclusive species (61.29%). The transition
landscape shared 26.67% of its species with the mountain landscape, a value
very similar to the percentage of species shared with the tropical landscape. In
spite of its transitional character, 40.00% of the species of the transition
landscape were exclusive to it. 47.06% of the species of the mountain landscape
were exclusive (Table 6).
DISCUSSION AND CONCLUSIONS
We present the discussion of our results in two parts. In the first we discuss the
three levels of biodiversity – alpha, beta and gamma – for each of the three
landscapes studied. In the second part we draw some general conclusions and
make some closing remarks.
ALPHA DIVERSITY
Tropical landscape
The complete dominance of the Scarabaeinae subfamily (97% of the total
species) within the group of copronecrophagous beetles of this landscape is a
biogeographical phenomenon characteristic of Tropical America (see Halffter &
Matthews 1966). In addition to the Scarabaeinae and one species of Silphidae
there were some species of Aphodiinae, almost all of the genus Ataenius (see
Appendix 1).
The origin of the greatest difference found in the number of species per site is
ecological, and this difference depends on the type of soil, its depth, shallowness,
or scarcity as occurs where there are rocky lava flows (see below); and above all
the effect on substrate of arboreal cover.
47
In the low deciduous forests, the alpha diversity of Scarabaeinae ranges from
8 to 15 species (mean: 11.33). These data coincide with those published for other
low deciduous tropical forests in Mexico where there has been disturbance by
humans: 12 species in Jojutla, Morelos State (Deloya et al. 1987), 15 species in
Tepexco, Puebla State (Deloya 1992), and 16 species in El Crucero, Veracruz
State (Pensado-Cadena com. pers.). On the Pacific slope 13 and 19 species
were caught at two sites of tropical deciduous forest in the Manantlán Reserve
in the States of Jalisco and Colima (García-Real 1995). At all of these sites, the
number of species per site is much lower (from a quarter to a half, if not less),
than the number found in Tropical America in sites with rainforest (see Montes
de Oca & Halffter 1998, Halffter & Arellano 2002). The latter is even true when,
for the comparison, we do not consider the number obtained in a given site or the
mean value, but rather the cumulative richness for a given type of community. In
the deciduous tropical forests of central Veracruz, cumulative species richness
is 22. Comparing this with published data that we are certain refer to cumulative
alpha richness, we have very similar values for similar ecological and
biogeographical conditions: 20 species in the Manantlán tropical deciduous forest
(García-Real 1995) and 22 species in patches of tropical forest in Tolima,
Colombia (Escobar 1997). On the other hand, for southern Veracruz (Los
Tuxtlas), in fragments of tropical rainforest, the cumulative richness of
copronecrophagous beetles is 30 species (Estrada et al. 1998). In this region (Los
Tuxtlas), for the entire landscape including fragments of rainforest, cocoa, coffee
and citrus plantations, as well as pastures, the gamma diversity is 36 species,
which emphasizes the greater predominance of the rainforest species (Estrada
et al. 1998) as compared to that found in the tropical landscape we studied.
In the pastures of the tropical landscape species exchange is notable (see beta
diversity), but the number of species remains fairly constant: from 9 to 14 species
per site (mean: 11) with a cumulative richness for the community of 21 species
(see Appendix 3).
In secondary vegetation next to patches of deciduous tropical forest, there
were 9 to 13 species per site (mean: 11), with a cumulative richness of 17
species. These areas have dense vegetation which allows some forest species
to coexist with heliophilous species from the open areas (Appendix 3). These
data indicate an interesting constancy in the number of species, both for sites
and for communities. This, even though there are some very important qualitative
differences between the species of the forest (and, in part, in the surrounding
secondary vegetation) and those of the pastures; species of the latter being more
heliophilous and predominantly coprophagous.
The tropical forests that grow on the rocky lava flows are quite different from
the three community types mentioned above, in that they are much poorer in
species (4 to 5 species, cumulative richness: 7). None of these species is
exclusive to this type of community, but rather all are the most widely spread and
most ecologically tolerant. The volcanic extrusion on which these forests grow is
one of the youngest of the lava flows in the three landscapes studied (less than
5,000 years old). The soil, formed by the disintegration of the lava, accumulates
in fissures and depressions but is only a few centimetres thick. This could
48
adversely affect nesting for many Scarabaeinae species. As such, the number
of individuals of the few species collected is lower than that for the other
communities of the tropical landscape.
The species composition of the communities in the tropical landscape depends
on the degree of arboreal cover (see Halffter & Arellano 2002). The more
continuous the canopy, the greater the number of species from the forest and the
fewer the number of heliophilous species. In the most intact forests (Jalcomulco,
for example) more than 70% of the species are exclusively forest dwelling, while
in sites with greater degrees of disturbance, and consequently reduced arboreal
cover, as much as 70% of the species are heliophilous. The same has been
observed in Colombia (Escobar 1997).
In the most disturbed patches of tropical forest, some species become very
abundant. This is the case with Canthon (Canthon) cyanellus and Deltochilum lobipes.
Favila and Halffter (1997) documented how the rank of C. cyanellus varies in terms
of abundance according the degree of disturbance in the landscape. This species
went from being the 11th of 27 species in the rainforest of Chajul, Chiapas, to
ranking 12th of 24 species in the Los Tuxtlas, Veracruz rainforest, and to being
the first of 18 species in pastures with forests remnants in Laguna Verde,
Veracruz. One possible explanation offered by the authors is that C. cyanellus
prefers the environment at the forest’s edge and fragmentation increases the
extent of this environment or similar conditions.
Kramer (1997) has indicated that the existence of abundant populations of
some species of plants and animals appears to be a general characteristic of
fragmented systems. Some species respond to an increase in the borders of the
original community (the forest) resulting from fragmentation. Debinski & Holt
(2000) and literature cited therein provide a review of the studies on the effects
of fragmentation.
Biogeographically, the three suprageneric taxa (Scarabaeinae, Geotrupinae
and Silphidae) of the indicator group found in this landscape show an overlap
between the species that follow a Neotropical distribution typical of the tropical
landscape, and those that follow the Nearctic and Paleoamerican mountain
pattern characteristic of the mountain landscape. However, there is a notable
number of exclusive species (see Table 4, Appendix 3 and biogeographical
analysis in Halffter et al. 1995). The same happens with Aphodiinae (see
Appendix 1).
With 10 to 13 species (mean: 11.67), the alpha diversity in the cloud forest was
among the highest found in the three landscapes; only bettered by that of the
deciduous forests and their neighbouring pastures in the tropical landscape. In
similar a cloud forest near the Orizaba Volcano, Veracruz, Pensado-Cadena
(pers. com.) captured 8 to 15 species (mean: 11.5) per site.
With respect to the species richness of cloud forest, it is important to consider
that those we have studied, as well as most of the other surviving patches of this
type of forest in the region, have been intensely disturbed and fragmented by
human activity.
49
At sites located in the gaps of cloud forest, i.e. with no arboreal cover, we
captured 8 species. This represents a poor subset of the cloud forest fauna. As
expected, 100% of the species caught are shared with the cloud forest.
The alpha diversity for sites of coffee plantations with polyspecific shade (12 -
13 species) is similar to that of the neighbouring patches of cloud forest. These
plantations have the greatest area with arboreal cover in this landscape and
comprise an important element of the matrix by linking the patches of cloud
forest. Their three layers of vegetation (Jiménez-Avila & Correa 1980, Aguilar-
Ortíz 1982, Cházaro-Basáñez 1982, Nestel 1990, Nestel et al. 1993) make them
similar to the disturbed cloud forests that still exist. In addition, at least
temporarily, there is more excrement available, especially human. This explains
the abundance of some exclusively coprophagous forest species such as
Dichotomius satanas (see Appendix 3).
There is a contrast between the results for coffee plantations with polyspecific
shade, and those with monospecific shade. In the latter, the number of species
is lower (5 to 8 species per site) than the number found in neighbouring cloud
forests. These data are similar to those found in similar coffee plantations (4 to
7 species per site; Morón & López-Méndez 1985, Morón 1987, Nestel et al. 1993).
In the pastures next to cloud forests, site alpha diversity (7-10 species, mean:
8.5) was lower than that of the forests. Species inhabiting the pastures belong
only to the Scarabaeinae family. These pastures serve as corridors that allow the
heliophilous and thermophilous species such as Dichotomius colonicus and
Euoniticellus intermedius from the tropical landscape to move to higher altitudes. In
contrast with the tropical landscape, here there is no group of species that is
specific to deforested areas, although some (such as Onthophagus chevrolati retusus
and Dichotomius colonicus) are more abundant in the pastures and a few, such as
Canthon humectus and Phanaeus amethystinus, are only found in this type of
community. The latter two species are more abundant (and biogeographically
characteristic) of the mountain landscape and, as such, their presence in the
pastures of the transition landscape represent an altitudinal expansion down to
lower levels.
The alpha diversity recorded for the oak forest (8-10 species, mean: 9) was
lower than that obtained by Pensado-Cadena (pers. com.) for the region near the
slope of the Orizaba Volcano in Veracruz (15 species). These notable differences
are explained by differences in the type of oak forest and their altitudes. The
forest of Pensado-Cadena’s study was located at 750 m asl and at 1,600 - 1,800
m asl in the present study. The comparison and the number of species of the
indicator group in the oak forests studied by Pensado-Cadena are similar to the
values we recorded for the forests of the tropical landscape.
Secondary vegetation next to oak forest had 7 species, all of which were also
found in the oak forests, and therefore represents a species-poor subset of the
oak forest fauna.
The mean alpha diversity in the pine-oak forests of the transition landscape (8
species) was less than that found in the cloud and oak forests. As altitude
increases (towards more temperate conditions) the number of Scarabaeinae
species decreases (pine-oak forests are at a higher altitude). Mean alpha
50
diversity in the pine-oak forests we studied was greater than that reported by
García Real (1995) for the Manantlán mountain range or by Pensado-Cadena
(pers. com.) in the region of the Orizaba Volcano, both of whom captured a mean
of 6 species.
In pastures next to pine-oak forest alpha diversity varied from site to site (5 -
11 species), but the mean was similar to the value for pastures next to cloud
forest (8.5), even though the composition was very different. In pastures next to
pine-oak forest we found species from the mountain landscape, such as Canthon
humectus.
We attribute the variation among pastures to differences in the time elapsed
since deforestation. Where less time has passed since the forest was cut, there
has been less opportunity for the process of assemblage of new beetle
communities. Similarly, in communities growing on rocky lava flows (discussed
below), as the assemblage process matures, more niches are occupied and the
number of species increases.
At the transition landscape the rocky lava flows are 5,000 to 10,000 years old,
on a site that is 36,000 years old (Geissert 1994). This means the soils are
deeper than those of the tropical landscape. Consequently, the number of
species of the indicator group is equivalent to that recorded on the other kinds of
soils (Table 3).
The mean alpha diversity for sites in the transition landscape (8.54 species)
was similar to that obtained for the tropical landscape (10.25 species), and
suggests a regularity in the number of species at sites located in the tropical and
subtropical areas of Mexico that share a similar biogeographical history.
As for the tropical landscape, the cumulative species richness in the transition
landscape was almost double the mean alpha diversity and indicates high values
of temporal species exchange. In the cloud forests, characterized by a high
degree of heterogeneity in their vegetation, cumulative richness was 18 species;
a little less than that found in the tropical forests. In pastures next to cloud forest
cumulative richness was only 66.66% of that recorded for the adjacent forests.
We have already mentioned that there is no specific group of species in the
treeless environments of the transition landscape. Also noteworthy is the lack of
Scarabaeinae roller species in the pastures. This absence truly leaves an empty
niche given the high availability of dung which is only partially used by burrower
species. There is ample documentation confirming that the coprophagous beetle
fauna of various tropical and subtropical areas is made up of roller and digger
Scarabaeinae species, and that the differences in their behaviour allow them to
exploit excrement in different ways. In our study region, the roller species are
found in the pastures of the tropical and mountain landscapes. They are not
however found in the pastures of the transition landscape. The presence of
Canthon humectus in some of the higher altitude pastures of the transition
landscape is clearly the result of downward expansion from the mountain
landscape and there is only one necrophagous roller species in the forests. This
phenomenon has also been observed for sites at similar altitudes and with similar
ecological conditions in Colombia (Escobar 1997, Escobar & Chacón de Ulloa
2001).
51
Mountain landscape
This landscape has species with biogeographical affinities that are distinct to
those of the species of the tropical landscape (see Halffter et al. 1995). In terms
of the representation of the indicator group there is a decrease in the relative
importance of the Scarabaeinae and an increase in that of Geotrupinae and
Silphidae (all together 6 species; see Appendix 3). There is also a substitution in
Aphodiinae: Aphodius and relatives are the dominant genera and there is only one
species of Ataenius in a site at a relatively low altitude in the landscape (see
Appendix 1 and Lobo & Halffter 2000). Furthermore, this is the only one of the
three landscapes studied where the richness of the indicator group is higher in
treeless sites (3 to 7 species) than in the forests (2 to 5 species; see Appendix
3 and Table 3). In biogeographical terms, this landscape has more affinities with
the nearctic element of North America. Sites without arboreal cover share
geographic and ecological continuity with the large expanses of the Mexican High
Plateau that have similar conditions. For ecological reasons the Scarabaeinae
have greater affinity for higher light conditions. In this landscape there is a
convergence with what is found in southern Europe: much greater species
richness in the grasslands (see, for example, Kadiri et al. 1997).
In the grasslands of the mountain landscape during the day, insolation levels
result in more temperate conditions. This community type has its own
Scarabaeoidea (Scarabaeinae plus Geotrupinae) fauna. In the conifer forests
(Pinus, Pinus-Alnus and Abies) which are much less heterogeneous than the tropical
and subtropical forests, and colder than the treeless areas, the number of
Scarabaeoidea and Silphidae is low (2 to 5 species). García-Real (1995) found
6 species of beetles in similar forests in the Manantlán mountain range. In
highland vegetation, a harsh environment with marked changes between day and
night time, we only found two species of Silphidae. Close to the Orizaba Volcano,
Pensado-Cadena (pers. com.) did not find a single species belonging to the
indicator group above 3,000 m asl.
In the mountain grasslands Onthophagus hippopotamus was found, along with 4
species of Aphodius, inside pocket gopher burrows (Rodentia Geomyidae:
Cratogeomys merriami perotensis Merriam). These species are found between 3,000
and 3,100 m asl (see Appendix 1; Lobo & Halffter 1994). We captured O.
hippopotamus at 2,450 m asl, the lower limit of its altitudinal range.
Silphidae show a preference for forests, both in this and the transition
landscape. Within the genus Nicrophorus, there is an ecological and altitudinal
substitution. N. olidus is mainly found in the cloud and oak forests of the transition
landscape, while N. mexicanus is found in pine forests of the transition and
mountain landscapes. Other authors (Arellano 1992, Halffter et al. 1995,
Navarrete-Heredia 1995, Martínez-Morales et al. 1997, Arellano 1998, Navarrete-
Heredia & Quiroz-Rocha 2000) confirm this segregation. Tanatophilus graniger,
which has a clear northern affinity, is abundant in the forests of the mountain
landscape. The only elements of the indicator group found in forests growing on
the rocky lava flows were two species of Silphidae (Appendix 3).
The cumulative richness of 13 species in the mountain grasslands is almost
double that of the richest of the neighbouring forests (Table 3). Under the most
52
open conditions, Phanaeus amethystinus and Ontherus mexicanus are found and
these species were also collected in the transition landscape. The rest of the
species form a group characteristic of the mountain landscape, a few of which
are also found on the Mexican High Plateau.
BETA DIVERSITY
Tropical landscape
In previous studies (Halffter et al. 1992, Halffter & Arellano 2002) we have
indicated that there are two different groups of fauna in the tropical landscape:
one of forest Scarabaeinae species and one with heliophilous species,
characteristic of pastures. In spite of this, the number of species shared by
different community types is relatively high (Table 4). The most characteristic
fauna is that of the tropical forests (22 species in total, 6 exclusive). This fauna
shares more than half of its species with the pastures and with secondary
vegetation. Our results coincide with those obtained by Escobar & Medina (1996)
for a region of Colombia with forests and pastures where the primary vegetation
is rainforest. In contrast, in a landscape with rainforest fragments, plantations and
pastures in southern Veracruz (Los Tuxtlas), the proportion of forest species is
markedly greater than that of the pasture species (Estrada et al. 1998).
Analyzing the beta diversity among the sites of a given community (Table 5),
the exchange was low compared to those sites with better conditions for the
beetles, and high among those sites with harsh environments. The highest beta
diversity was found for tropical forests on the rocky lava flows (77.5%). This
clearly indicates the relatively greater number of non-resident individuals
(metapopulations, tourist species) under harsher conditions. A good many of
these individuals could arrive from nearby communities to take advantage of
occasional food, but they do not nest because of the scarcity of soil. Among
these individuals, large species dominate (>70%) as do those with a greater
ecological tolerance.
Species exchange among cloud forests was 37.37% (Table 5). On comparing
beta diversity among sites with forest or with pastures for a given location, the
greatest exchange was recorded between a small but well preserved fragment
of cloud forest and its neighbouring pasture. The latter had species very
characteristic of treeless areas, such as Dichotomius colonicus and Scatimus ovatus,
that do not enter the forest. Taken together (Table 5) the differences between
cloud forests and pastures are great. In contrast, between cloud forests and
coffee plantations with polyspecific shade, the exchange was limited (14.3%, see
Table 5). This similarity (lower beta) is confirmed for the number of shared
species between the most widely distributed natural community (i.e. the cloud
forest) - even though it is currently fragmented and partially altered - and the
anthropogenic community that is most densely treed and covers the greatest
area of this landscape (i.e. the coffee plantation with polyspecific shade).
53
This is very important for the conservation of the species originally limited to
cloud forest. In other groups of organisms, such as for birds in both Chiapas
(Greenberg et al. 1997) and in central Veracruz (Aguilar-Ortiz 1982) a similar
phenomenon occurs. According to Muñoz et al. (2000) for the amphibians and
reptiles of the El Triunfo Reserve in Chiapas, coffee plantations with polyspecific
shade function as a community that preserves the species of both cloud forest
and pine-oak forest, as well as connecting the patches of forest that still exist.
The Scarabaeinae and Silphidae of the transition landscape move through this
mosaic of cloud forest, gaps in the forest and coffee plantations with polyspecific
shade. To a lesser degree they enter the pastures and coffee plantations with
monospecific shade. Thus, the greatest values for mean beta diversity (>55%)
are recorded between cloud forest and the pasture, or between the cloud forest
and coffee plantations with monospecific shade (Table 5).
Mountain landscape
There is a high degree of species exchange between the pasture and fir forests
(>79%) or pine forests (>63%). This exchange is a consequence of the rich
copronecrophagous fauna that is characteristic of treeless places (see Table 5);
of which only a very small proportion enter the forest.
GAMMA DIVERSITY
We have seen how gamma diversity for the three landscapes is determined by
the relationship between alpha and beta diversity. However, gamma diversity
does have its own value, a historical value, on which the number and composition
of species found in a landscape depends.
In the tropical landscape, the richness of the indicator group is primarily a result
of the Scarabaeinae which are characteristic of the tropical forest, although there
is a group of species associated with open conditions. In contrast, for the
mountain landscape, the most important element is comprised of the species of
the indicator group that are associated with open areas. In the transition
landscape the species associated with the cloud forest dominate.
In the tropical and transition landscapes, the dominant fauna corresponds to
the natural community that was most extensive before human intervention. The
fact that there are more species in the grasslands of the mountain landscape,
even though this is not the dominant natural community, is related to the
thermophilous and heliophilous characteristics of the Scarabaeinae. (These
beetles are not abundant in other cool, temperate forests of the rest of the world).
In the three landscapes that we have studied, the fragmentation of the natural
communities does not appear to have caused the loss of species (see Closing
Remarks below). In the tropical landscape the fragmentation of the tropical
forests has favoured the expansion of species from open areas. These
heliophilous and coprophagous species include two recent invaders
(Digitonthophagus gazella and Euoniticellus intermedius) which are very abundant in
some pastures. Fragmentation has also favoured the expansion of ubiquitous
54
species, or those that take advantage of the edges of communities, such as

Canthon (Glaphyrocanthon) leechi and Canthon (C.) cyanellus. In contrast,
fragmentation has resulted in a contraction in the distribution of specialists such
as Canthon (Glaphyrocanthon) femoralis that exploit the excrement of arboreal
mammals, especially that of monkeys. For now, as far as we can determine, no
species has disappeared. This has to do with the topography of the landscape
and the type of deforestation that has occurred. Gorges with steep slopes help
the narrow strips of tropical forest to survive. The majority of the pastures are not
totally denuded of trees since gorges, living fences with trees and trees in the
pastures help to maintain the connectivity of the landscape (Guevara et al. 1998).
In a tropical landscape such as that of central Veracruz the deciduous or semi-
deciduous tropical forest has alternated between forest and open areas over
centuries. Hence, the heliophilous species are more numerous than in the parts
of Tropical America that were originally covered with tropical rainforest. In the
latter, the effects of deforestation are much more drastic (Howden & Nealis 1975,
Klein 1989, Turner 1996, Laurence & Bierregard 1997, Estrada et al. 1998).
Nevertheless, even where rainforest completely dominates, there are some
heliophilous species (those of the gaps or edges) that respond favourably to
forest fragmentation (for related information on the butterflies of the Amazon, see
Tocher et al. 1997).
In the transition landscape there are no species that are characteristic of the
pasture. Phanaeus amethystinus which shows a consistent affinity for open
conditions, is more abundant on the lower levels of the mountain landscape.
Although Onthophagus incensus, Copris incertus and Ontherus mexicanus are found
more often in the pastures, unlike in the tropical landscape, there is not a clear
separation between the fauna of the pastures and that of the forest. Under these
conditions the coffee plantations with polyspecific shade provide an important
connection between the surviving fragments of cloud forest. These plantations
occupy a wide area and, as qualitatively and quantitatively shown in this
research, are an example of the ideas of Víctor M. Toledo and colleagues
(Moguel & Toledo 1999, Toledo et al. 1994) with respect to how diversified agro-
silvicultural exploitation can help conserve biodiversity.
The absence of a group of fauna characteristic of the pasture in the transition
landscape is confirmed by the lack of an ecological group: the roller beetles (see
Alpha Diversity), and also by the good number of cow pats that go unexploited.
The latter is emphasized by the unequal development of the grasses.
It is evident how fragmentation and the change of communities can reduce the
number of species that are found in a landscape. On losing the conditions that
are favourable for a given population, it disappears. But, how can a certain
degree of fragmentation and landscape modification result in an increase in the
original number of species? For this to happen, two conditions must be met. First,
the fragments of the original community that persist must be of a sufficient size
and there must be adequate interconnection so that populations are not lost. The
required size and degree of connectivity varies greatly from one species to the
next. See below for a discussion of species that can adapt to a metapopulation
dynamic.
55
Second, the new conditions created by the change open opportunities. They
allow colonization by species that were previously outside of the landscape, and
not necessarily in contiguous places. This occurs in the three landscapes that we
analyzed. In the tropical landscape, deforestation creates or expands pastures
and this allows heliophilous species and those that prefer cow dung to extend
their distribution. Deforestation in the transition landscape permits heliophilous
species at both higher and lower altitudes to move downwards or upwards. And
finally, in the mountain landscape deforestation creates conditions that are more
favourable for the penetration of some tropical species, but especially the
heliophilous species of the Mexican High Plateau. It is evident that the altitudinal
gradient favours these types of displacement associated with the modification
communities that previously were the most continuous and wooded.
An idea of the relative richness of each landscape is given by the mean alpha
diversity by landscape (that is, the mean of all the alpha diversity values found
in all the communities of a given landscape). These values are 10.25 species for
the tropical landscape, 8.45 species for the transition landscape and 3.37 for the
mountain landscape. These data are notably influenced by the richness of the
main component of the indicator group: the Scarabaeinae, which is much more
abundant in species in the tropical landscape.
Complementarity among landscapes

Species composition is very distinct in the three landscapes and so
complementarity is greater than 79%. The proportion of species exclusive to each
landscape are 70.97% in the tropical, 47% in the mountain, and 43% in the
transition landscape; the latter sharing the most species with the other two (Table
6).
Complementarity between landscapes was calculated with and without
Aphodiinae species. Complementarity between the tropical and transition
landscapes is C12 = 82.69 (with Aphodiinae) and 77.11 (without), and between
the tropical and mountain landscapes it is C13 = 97.87 and 94.81, indicating
totally different fauna (only one species is shared). Between the transition and
mountain landscape complementarity is C23 = 79.49 and 35.00. Here, the
marked change that occurs on including Aphodiinae is the result of this subfamily
having many species that are shared between this last pair of landscapes.
CLOSING REMARKS
1. As far as our data allows us to speculate, the effects of human activity
(deforestation, changes in landscape use to pastures or crops, alteration and/or
fragmentation of the forests) has not affected the total number of species of the
indicator group in any of the three landscapes. We do however have doubts
about the cloud forest which is the foundation of gamma diversity in the transition
landscape. These doubts can only be resolved by a comparative study of a well
preserved and biogeographically equivalent cloud forest with the cloud forest of
central Veracruz (see Appendix 4).
56
It is interesting that disturbance by human activity appears to have been

overcome by the beetles of the indicator group for different reasons in each
landscape. In the tropical landscape it is a result of the heliophilous fauna, for
which the number of species has even increased with the recent arrival of two
invaders. This fauna also complements the tropical forest fauna that lives in the
remnant fragments, and a small proportion of it is adapted to the pastures. This
is why the pasture fauna is not poor in species, as might be expected in a
landscape once dominated by deciduous tropical forests (see Montes de Oca
2001 for an analysis of pasture fauna in Los Tuxtlas, southern Veracruz and its
relationship to the forest and forest remnants). In the transition forest, where we
feel the impact on the forest has been greater, the coffee plantations with
polyspecific shade have mitigated anthropogenic effects by creating a matrix that
connects the fragments of cloud forest. In the mountain landscape the indicator
group is not naturally species rich in the cool temperate forests. The expansion
or creation of grasslands has increased the availability of conditions favourable
for heliophilous species.
It remains to be seen whether these findings can be applied to other groups of
animals and plants. We do not expect all biota to have the same capacity for
homeostasis in the face of anthropogenic changes as the indicator group of
beetles we have studied. From the work done by our research group Julián
Bueno-Villegas (pers. com.) has found 23 species (preliminary data) of Diplopoda
in the cloud forest and only 2 in coffee plantations, with only one species shared
between the two communities. Diplopods are very sensitive to the loss of forest
cover and the introduction of agricultural and animal husbandry practices (Bueno-
Villegas & Rojas 1999).
An interesting sign is that throughout the sampling of our indicator group we
have not detected any extinctions (aside from our doubts about the cloud forest).
This, in spite of the disappearance or big decrease in birds, medium-sized and
large fauna (often as a direct result of hunting). Here, we refer to extinction at the
landscape level since it is evident that at the local level, the number of species
can be greatly affected by anthropogenic modifications. The homeostasis of the
landscapes is an unexpected result of the topographic heterogeneity of central
Veracruz, and of the previously mentioned factors which, to a certain degree,
buffer the effects of anthropogenic changes. It is important to remember that for
the Scarabaeinae, the most abundant of our indicator group, the main change
has not occurred recently, but rather from the time of the 16th century when cattle,
sheep and horses were introduced to Mexico. Additionally, in recent years
evidence has been published (along with models for its analysis) indicating that
some species can survive as metapopulations, even in highly fragmented
landscapes (Wahlberg 1996, and literature cited therein).
It is important to note that in the tropical and transition landscapes
(Scarabaeinae are poor indicators for mountain forests), there is the feeling that
in the type of forest that originally dominated (i.e. the main “container” of species
richness) fragmentation and alteration are very close to or have passed
sustainable limits. This is especially apparent for the cloud forest, where the
composition and richness of the indicator group is very similar to the values found
57
for the coffee plantations with polyspecific shade. This, added to the visual
impression of alteration, makes us aware of the fact that it has not been possible
for us to work with intact cloud forest, but rather only with remnants which are all
in some way simpler than the original community. We used the word “feeling”
above because we recognize that an analysis of the fragments that takes into
consideration its size and shape, as well as the number of species, is required
for the three landscapes. This would allow us to predict the point beyond which
the number of species decreases dramatically.
Klein (1989) has published the only study to date regarding the number of
copronecrophagous Scarabaeoidea (essentially Scarabaeinae) species and
forest fragment size. This study was done in Manaos, Brazil and has been
repeatedly cited to illustrate the catastrophic effects of deforestation and
demonstrate the loss of species from tropical evergreen forest that occurs as
fragment size decreases. However the landscape, even in the Amazon, has other
components. Klein mentions that four species of Canthon (Glaphyrocanthon) appear
in a gap of the forest. They are heliophilous species that are dominant at the
edges and in gaps, just as Canthon (Gl.) leechi is in our study.
As our study shows for the tropical landscape, heliophilous species can even
be found in regions originally dominated by tropical forest. The transformation of
forest to pastures and the new availability of cattle dung allows these species to
become more abundant and evident. (An analysis of the richness of dung beetle
fauna in a highly fragmented landscape of southern Veracruz that has rainforest
remnants, plantations and pastures can be found in Estrada et al. 1998.)
Thus, a question arises: Can the analysis of the species richness of a forest or
of the dominant community allow us to predict what is happening to the diversity
and species composition in the entire landscape? Our answer is no, although we
must remember that we are working in a very heterogeneous region where there
is obvious complementarity between the different communities of the landscape.
Smith et al. (1997) propose a reevaluation of the role of ecotones in the
generation and maintenance of the biodiversity of tropical forests. Davis et al.
(2000) show, for the conditions of Borneo, how forest species that live in more
exposed conditions (edges, riverbanks) are more adaptable than those of the
forest interior to anthropogenic conditions, such as forest plantations.
2. The title of this study reflects our interest in the relationships between the three
expressions of biodiversity: alpha, beta and gamma. Gamma diversity depends
on the biogeographical history of the landscape and is the most stable expression
of biodiversity. It is also that which has greater homeostasis in the face of human
activities. The value of alpha diversity depends on the biogeographical and
historical events, but even more on the assemblage of the community being
studied. In this study we have tried to see whether the characteristics of regional
species richness (gamma) depend mainly on local richness (alpha) or on the
exchange between communities (beta). Ricklefs (2000) also addresses this
question with a different focus in his analysis.
It is important to understand the double sense in which the relationships
between local diversities and landscape diversity manifest. The alternatives are:
how many, and which species depend on the historical biogeographical
58
phenomena that have acted both on the biota and on the landscape. The value
is expressed as gamma, but gamma is “constructed” from different values of
alpha. This is why we say that gamma diversity is a determinant of alpha and
beta diversity, but at the same time is also conditioned by the values of alpha and
beta.
In the tropical landscape, gamma largely depends on the species richness of
the community that was originally dominant (alpha): deciduous (or semi-
deciduous) tropical forest. There is a notable degree of complementarity with the
heliophilous fauna of the pasture that results in gamma (30 species) being higher
than the cumulative richness for the deciduous forest (22 species) or the pastures
(21 species, see Table 3). In the transition landscape, cloud forest had the
highest richness and was most responsible for gamma diversity, though
anthropogenic modification (i.e. deforestation) allows some species to ascend
from the tropical landscape and others to descend from the mountain landscape.
These species add to the gamma richness. In the mountain landscape the total
number of species (17) is close to the cumulative richness of the pastures (13),
but very different from that of even the richest forests (7 species). The creation
or expansion of the grasslands creates the right conditions for the addition of a
rich heliophilous fauna.
So, the relationship between the cumulative richness of the community that
originally was the most important and the gamma diversity is different for each
of the three landscapes, and these differences depend on whether or not there
was an ecologically distinct species assemblage during the original conditions,
and on the response of that assemblage to changes introduced by humans. We
return to the title of our study and our central question: What is the order of
importance of the factors that determine the diversity of a landscape? And our
answer varies with the biogeographical history of the landscape. In the three
landscapes, the species exchange between forested areas and their
corresponding grasslands results in a value for gamma diversity that is superior
to the highest recorded value of alpha diversity. Hence beta diversity is also very
important to the gamma diversity of coprophagous beetles.
Moreno & Halffter (2001) did a study of bats in a valley of the tropical
landscape with the same objectives as those of this study. They found that of the
20 species that comprised the gamma diversity, 18 were found in the richest
community (subdeciduous tropical forest) and the remaining two were found in
the riparian vegetation. In bats, the landscape’s species richness depends on that
of the richest community to a greater extent than it does in beetles.
In a study of frogs in the cloud forests, coffee plantations and pastures of the
transition landscape, the expression of the components of diversity of beetles and
frogs (two taxonomically and biologically distinct groups) is similar for parts of the
landscape with similar characteristics (E. Pineda, pers. com.). Gamma diversity
for the frogs is 21 (22 for beetles), and mean alpha diversity is 8.16 species for
the frogs (9 for beetles). Species exchange between sites for both beetles and
frogs is high and greater than between communities, suggesting that in this
mosaic spatial heterogeneity is linked to the heterogeneity inherent to groups of
species and that differences in the degree of conservation of each site lead to
differences in the composition of the indicator groups.
59
3. Alpha diversity is one of the most frequently used values to characterize

biodiversity. Although it is well defined in spatial terms, its large variation over
time is not often considered. Along with Moreno and Halffter (2001), we
distinguish the alpha diversity of a site or a community (which requires a varying
amount of time to sample) from the cumulative species richness of a community
(referred to as cumulative alpha diversity by Moreno and Halffter), which is the
sum of all the species collected from a community during a given period of time.
Taking into consideration the great variations that can occur between these two
values (Table 3), it is interesting to ask which of these two values is most
meaningful for the characterization of a community and its contribution to the
biodiversity of the landscape (gamma).
The differences in the values of diversity are not constant. They are much
greater in the communities that are the richest and most representative of each
landscape, and in some communities the cumulative richness can be double the
mean alpha diversity. These communities might be remnants of the natural
vegetation or communities resulting from human activity. This is the case for the
tropical forests and their neighbouring pastures in the tropical landscape, the
cloud forests and coffee plantations with polyspecific shade in the transition
landscape, and for the grasslands next to pine forest, as well as the pine forests
themselves in the mountain landscape.
At the other extreme, in the poorest communities (including those on the rocky
lava flows) cumulative richness and mean alpha diversity have approximately the
same value. There is no evidence of species accumulation over time. This means
that for the indicator group, and perhaps for many others, the communities that
are richest in species have the greatest degree of species exchange over time.
This conclusion is quite contrary to one’s first impression. The communities that
are richest in species correspond to the most complex assemblages: those where
there should be fewer empty sites. How is it that under these conditions there are
so many tourist species and even so much species exchange? One possible
explanation is that the massive availability of herbivore and omnivore dung
(cows, horses, pigs, sheep and even humans) is a new phenomenon across the
entire gradient of landscapes, having occurred over the last 400 years for
domestic livestock and several millennia for humans. The assemblage process
has not finished; quite the contrary. Human activities such as deforestation, the
introduction of livestock and the very presence of humans, have created new
niches that have yet to be fully taken advantage of. In the transition landscape,
this is quite clear in the unused dung piles that are abundant in the pastures.
An additional explanation exists. The richest communities are those that have
the greatest number of rare species which, are the most likely to be missed
during sampling. The number of species will be affected by both tourist species
(occasional passersby) and individuals from the source populations. In both
cases these beetles stay in these sites precisely because of the heterogeneity
of the community which allows them to temporarily survive in the ecological
spaces that are available.
Consistent with the importance of species exchange, the number of species
exclusive to communities is low. The exception being the pastures next to pine
forest in the mountain landscape where the highest absolute number of exclusive
species was recorded. The pastures of the mountain landscape are a community
60
that was created or at least expanded by deforestation. The species of this

community – even though they are exclusive in relation to the other communities
of the same landscape (see Table 4) – represent a peculiar assemblage of
species from distinct origins and come together in response to the open
conditions and food availability (cattle dung).
The presence of exclusive species in communities of anthropogenic origin in
the tropical and transition landscapes is the result of two invader species (only
one of which is found in the transition landscape), and of the existence even in
undisturbed conditions of a few species that prefer gaps (see comments in
Closing Remarks 1).
The rocky lava flows have a drastic effect on the indicator group, especially the
Scarabaeinae. In each of the three landscapes, the communities on this kind of
soil are the poorest in species. In the transition and mountain landscapes the
indicator group is only represented by the Silphidae that are less sensitive than
the Scarabaeoidea to soil quality. In the tropical landscape the Scarabaeinae that
are found on the rocky lava flows are the most abundant and ubiquitous of the
landscape, but are always present in limited numbers (marginal specimens of
nearby metapopulations). One exception is the abundance of Canthidium
puncticolle and Canthon (Gl.) leechi, two species that are very small in size that were
captured in a pasture on the rocky lava flows (see Appendix 3).
The comparison of the numeric values of mean alpha diversity by landscape
or of the gamma diversity, disregards the very important replacement of species
of different biogeographic origin that occurs in the indicator group (see Halffter
et al. 1995, Lobo & Halffter 2000). In the tropical landscape, of 32 species, 31 are
Scarabaeinae and one is Silphidae. In the transition landscape, of 30 species, 25
are Scarabaeinae, 1 is Geotrupinae and 4 are Silphidae. In the mountain
landscape, of a total of 17 species, 11 are Scarabaeinae, 4 are Geotrupinae and
2 are Silphidae. Biogeographical replacement also occurs in the Aphodiinae. The
genus Ataenius dominates in the tropical and transition landscapes, while Aphodius
group of genera dominates in the mountain landscape (see Lobo & Halffter
2000).
4. The temporal and spatial dimensions of alpha diversity. Both this study and
that of Moreno & Halffter (2001) demonstrate that alpha diversity has values that
are far from fixed. Alpha diversity, that is the number of species that are found in
a community, is one of the important characteristics and perhaps the most
important in terms of reflecting the history of the community. It also contributes
decisively to conditioning the structure and function of the community. These
relationships between the number of species and structure and function require
an in-depth review since, there does not appear to be confirmation of the linear,
deterministic sequence that most authors have maintained to date.
If the community is an assemblage of species that varies in time and space, we
should not be surprised that alpha diversity also varies in time and space. In this
study we have explored spatial variation. What we call alpha diversity by site is
the number of species found in a site when the sampling effort has resulted in the
capture of at least 85% of the estimated number of species present. Mean alpha
diversity of a community is the mean obtained by sampling the different sites that
correspond to the same community in a landscape. This value does not take into
consideration the spatial and stochastic variation among the samples (sites)
61
taken from an entity (community). The cumulative richness of species for a

community is a cumulative value that ignores spatial and temporal variations
within the sampling period.
We have not analyzed the differences in alpha diversity that the temporal
dimension might cause, such as those resulting from the timing, duration and
frequency of sampling, or how alpha diversity varies over the years and longer
cycles of time.
This study clearly indicates that the expressions of alpha diversity we have
studied, i.e. those associated with space, vary greatly depending on the type of
community. If we consider the differences related to temporal variation, we
believe that these too would depend on the type of community. The richest
communities, such as the tropical forests exhibit a greater variation in alpha
diversity over time owing to the low probability of capturing rare species. In
simpler communities, such as those growing on the rocky lava flows in the
tropical landscape, there is less variation in the number of species captured,
since the species are more widespread.
ACKNOWLEDGEMENTS
We are grateful to Lorrain Giddings and Juan Chávez (Instituto de Ecología, A.C.) for
their help in the preparation of the maps. Daniel Geissert (Instituto de Ecología, A.C.) kindly
provided information on the age of lava flows and assisted in establishing the boundaries
of the landscapes. Both Leonardo Delgado (Instituto de Ecología, A.C.) and Mario Zunino
(Universidad de Urbino, Italia) helped with the identification of Onthophagus species.
Margarita Soto and her research team (Instituto de Ecología, A.C.) generously helped us
obtain environmental parameters for the sites studied. We thank Rafael Sánchez for his
help with the field work. The authors especially wish to express their gratitude to Miguel
Pensado-Cadena, for permission to cite information collected an as yet unpublished study
on coprophagous beetles of the Orizaba Volcano gradient. We are also grateful to the
following colleagues for providing unpublished information: Imelda Martínez (Instituto de
Ecología, A.C.), Marco Dellacasa (Universidad de Pisa, Italia), Julián Bueno (Instituto de
Ecología, A.C., Field Museum of Natural History, Chicago), Eduardo Pineda (Doctoral
candidate in the Graduate Program of the Instituto de Ecología, A.C.) and Antonio Muñoz
(ECOSUR. San Cristóbal de las Casas, Chiapas). Claudia E. Moreno (Universidad
Autónoma del Estado de Hidalgo) kindly read this text and made insightful suggestions.
We thank Bianca Delfosse who not only carefully translated this article from the original
in Spanish, but also provided valuable help by reviewing the text and offering many
suggestions.
This study forms part of the following CONABIO (Comisión Nacional para el
Conocimiento y Uso de la Biodiversidad, México) projects: "Parameters for measuring
biodiversity and its changes: Stage II - Development of Examples” (CONABIO
FB532/K038/97), and "Parameters for measuring biodiversity and its changes: Stage III”
(CONABIO FB733/U030/00). This study is also part of the project for the development of
methods of measuring biodiversity funded by the Regional Office of UNESCO for Latin
America and the Caribbean (883.624.0). The last part of the elaboration of this document
was carried out as part of the project “The Effects of fragmentation on biodiversity: the case
of the Los Tuxtlas rain forest and the dung beetles” (CONACYT, 37514V). These projects
have been instrumental in our efforts to develop a new strategy for measuring biodiversity
on the scale of landscapes. The first author is grateful to the American Museum of Natural
History for support in the form of the Theodore Roosevelt scholarship which helped offset
field expenses.
62
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Recibido: 11 de julio 2002

Aceptado: 20 de marzo 2003
68
Appendix 1
The Aphodiinae species (Coleoptera: Scarabaeidae) of Central Veracruz, Mexico.The
landscape in which the species are found in the Cofre de Perote Region of central
Veracruz, Mexico. 1 = Tropical, 2 = Transition, 3 = Mountain. Sources for data collection
are: A = occasional captures by Arellano and Halffter, B = captures made by Imelda
Martínez and colleagues, C = data from Marco Dellacasa, as well as reports from the
literature. Only one source is cited for cases where several authors report a species for a
given location. Sources of bibliographic information are in Roman numerals: I = Lobo &
Halffter (2000); II = Deloya (2000), III = Deloya & Lobo (1995), and IV = Deloya (1994). The
local name of the site, its altitude and, where possible, the number of individuals captured
or reported (ND = no data) are given. The taxonomic study used for the determination of
Aphodiinae was Dellacasa et al. (2001) and for Eupariini, Deloya (2000).
Species Landscape Source Site Altitude No.

(m asl) ind.
Trichonotuloides glyptus Bates, 1887 3 A San José Aguazuelas 2900 2
3 I Los Pescados 3000 19
3 A Estación Las Lajas 3000 1
3 A km 11 Cofre de Perote 3300 2
3 C El Triunfo 2600 6
3 C Tembladeras 3100 5
Agrilinus prope duplex LeConte, 1878 3 I Los Pescados 3000 15
3 I El Conejo 3300 2
3 A km 11 Cofre de Perote 3300 42
Agrilinus azteca Harold, 1863 2 A Barrio La Ermita 1900 1
3 I Chololoyan 2500 1
3 A Camino a Las Lajas 2900 7
3 C Cruz Blanca 2400 2
Chilotorax pumilio Schmidt, 1907 2 I Ixhuacán 1900 10
2 I Acajete 2000 16
3 I Toxtlacoaya 2300 18
3 I Cruz Blanca 2400 57
Chilotorax multimaculosus Hinton, 1834 3 I Cruz Blanca 2400 4
Agrilinus lansbergei (Harold, 1874) 1 C Palma Sola 4 2
Planolinus prope tenellus Say, 1823 3 I San José Aguazuelas 2950 2
Blackburneus (sensu lato) guatemalensis ab scotinus (Bates, 2 C Las Animas 1400 6
1887)
2 C El Fresno 1800 2
Trichonotulus perotensis Lobo y Deloya, 1995 3 III Los Pescados 2900-3100 90
Platyderides pierai Lobo y Deloya, 1995 3 III Los Pescados 2900-3100 36
Blackburneus (sensu lato) charmionus (Bates, 1887) 1 A Otates 480 1
3 C Oxtlapa 2100 5
3 C 2 km from Oxtlapa 2400 2
3 C San José Paso Nuevo 2300 1
2 C km 14.5 Xico-Oxtlapa 2050 5
Cephalocyclus hogei (Bates, 1887) 3 I San José Aguazuelas 2950 23
3 A Plan del Vaquero 2900 3
3 A Camino a Las Lajas 2920 26
Planolinus vittatus Say, 1825 2 B Alto Lucero 1100 2
2 I Ixhuacán 1900 17
69
Appendix 1 Continuation
(m asl) ind-
Planolinus vittatus Say, 1825 3 B Rancho Los Salazares 2450 21
2 C Xalapa 1400 1
3 C El Triunfo 2600 2
3 C Los Laureles 2500 1
3 C Oxtlapa 2100 8
3 C 2 km from Oxtlapa 2400 14
3 C San José Paso Nuevo 2300 75
3 C San José Aguazuelas 3000 2
3 C Tonalaco 2350 11
Gonaphodiellus opistius Bates, 1887 2 I Acajete 2000 506
2 C El Fresno 1800 92
2 C La Joya 2000 7
3 I San José Aguazuelas 2950 1
3 I Ixhuacán 1900 2915
3 C Rancho Los Salazares 2450 35
3 C A 2 km de Oxtlapa 2400 2
3 C Tetelcingo 2200 50
3 C Km 14.5 Xico Oxtlapa 2050 1
Agrilinus sallei Harold, 1863 2 C Las Animas 1400 2
2 C Alto Lucero 1100 4
2 C Parque Ecológico Clavijero 1360 2
2 C Quiahuistlán 60 88
1 C Palma Sola 5 2
1 C La Estancia, Palma Sola 10 2
1 C El Toche, Chavarrillo 600 3
1 C Los Lirios, Actopan 5 15
1 B La Mancha 0 17
1 B Rancho el Tajo, Actopan 5 23
1 A Cerro El Metate 30 3
1 A Barranca Grande 980 1
1 A Dos Rios 990 30
Nialaphodius nigrita Fabricius, 1801 1 B Quiahuistlán 60 10
1 B Villa Rica 4 4
1 A Cerro El Metate 30 2
1 A Dos Ríos 990 2
1 C Palma Sola 5 5
1 C El Tajo, Actopan 5 30
1 C La Estancia, Palma Sola 10 12
1 C Los Lirios, Actopan 5 68
2 B Alto Lucero 1100 1
2 C Km 14.5 Xico Oxtlapa 2050 1
Labarrus pseudolividus Balthasar, 1941 1 B Quiahuistlán 0 5
1 B Villa Rica 4 6
1 B Rancho Los Lirios, 5 3
Actopan
1 C Palma Sola 5 1
1 C El Tajo, Actopan 5 54
1 C Quiahustlán 60 26
70
(m asl) ind.
Labarrus pseudolividus Balthasar, 1941 1 C La Estancia, Palma Sola 10 4
2 B Alto Lucero 1100 22
2 A El Fresno 1920 36
2 C Club Hípico, Las Animas 1400 5
2 C Las Animas 1400 66
3 C Los Laureles 2500 1
3 C Oxtlapa 2100 1
3 C Tembladeras 3100 1
Ataenius apicalis Hinton, 1837 1 II Villa Rica 10 ND
1 II Actopan 200 ND
1 IV Cotaxtla 50 ND
Ataenius euglyptus Bates, 1887 3 A Las Vigas 2450 1
Ataenius languidus Schmidt, 1910 1 II Puente Nacional 6 2
1 IV Cotaxtla 50 2
Ataenius cibrithorax Bates, 1887 2 IV Xalapa 1400 ND
Ataenius imbricatus Melsheimer, 1844 1 II Cotaxtla 50 ND
2 IV Xalapa 1400 ND
Ataenius holopubescens Hinton, 1938 1 IV Cotaxtla 50 ND
Ataenius abditus (Haldeman, 1848) 2 A Rancho Briones 1360 ND
Ataenius sculptor Harold, 1868 1 IV Cotaxtla 50 1
2 A Ixhuacán 1900 14
Ataenius strigicauda Bates, 1887 1 II Cotaxtla 50 ND
1 IV Veracruz 5 ND
2 IV Xalapa 1400 ND
Ataenius jalapensis Bates, 1887 2 II Xalapa 1400 ND
Ataenius liogaster Bates, 1887 1 II Villa Rica 10 ND
2 IV Xalapa 1400
Ataenius texanus Harold, 1874 1 IV Puente Nacional 60 1
Ataenius glabriventris Schmidt, 1911 1 IV Cotaxtla 50 4
Ataenius capitosus Harold, 1867 3 IV Las Vigas 2400 ND
Ataenius figurator Harold, 1867 1 IV Córdoba 900 ND
Ataenius rickardasi Hinton 1938 1 IV Veracruz 5 ND
1 IV Cotaxtla 50 1
2 IV Huatusco 1250 2
2 IV Fortín de las Flores 1000 1
Literature cited
Dellacasa, G., P. Bordat & M. Dellacasa. 2001. A revisional essay of world genus-group taxa of
Aphodiinae. Mem. Soc. Entomol. Ital., 79: 1 - 482.
Deloya, C. 1994. Distribución del género Ataenius Harold, 1867 en México (Coleoptera: Scarabaeidae:
Aphodiinae, Eupariini). Acta Zool. Mex. (n.s.) 61: 43-56.
Deloya, C. & J. M. Lobo. 1995. Descripción de dos nuevas especies mexicanas de Aphodius de los
subgéneros Platyderides y Trichonotulus (Coleoptera: Scarabaeidae: Aphodiidae) asociadas con
Pappogeomys merriami (Rodentia: Geomyidae). Folia Entomol. Mex., 94: 41-55.
Deloya, C. 2000. Revisión de las especies mexicanas del género Ataenius Harold, 1867 (Coleoptera:
Scarabaeidae: Aphodiinae, Eupariini). Master Degree Thesis. Facultad de Ciencias. Universidad
Nacional Autónoma de México. 214 pp.
Lobo, J. M. & G. Halffter. 2000. Biogeographical and ecological factors affecting the altitudinal variation
of mountainous communities of coprophagous beetles (Coleoptera, Scarabaeoidea): a
comparative study. Ann. Entomol. Soc. Amer., 93 (1):115-126.
71
Appendix 2
A list of the Scarabaeinae species found in Laguna Verde, Veracruz, Mexico.
Community
Species tf pt
Onthophagus batesi Howden y Cartwright, 1963 0 1
Onthophagus schaefferi* Howden y Cartwright, 1963 0 1
Onthophagus hoepfneri* Harold, 1869 0 1
Digitonthophagus gazella (Fabricius, 1787) 0 1
Dichotomius amplicollis (Harold, 1869) 1 0
Dichotomius colonicus (Linneo, 1767) 0 1
Copris incertus (Say, 1835) 1 0
Canthidium puncticolle Harold, 1868 1 0
Copris lugubris Boheman, 1868 0 1
Phanaeus scutifer Bates, 1887 0 1
Phanaeus tridens Castelnau, 1840 0 1
Coprophanaeus pluto (Harold, 1863) 1 0
Sisyphus mexicanus Harold, 1863 0 1
Eurysternus mexicanus Harold, 1869 1 0
Euoniticellus intermedius Reiche, 1849 0 1
Canthon (Glaphyrocanthon) circulatus Harold, 1868 1 0
Canthon (G.) leechi (Martínez Halffter y Halffter,1964) 1 0
Canthon (G.) antoniomartinezi Rivera y Halffter, 1999 1 0
Canthon (C.) cyanellus Le Conte, 1859 1 0
Canthon indigaceus chevrolati, Harold, 1868 0 1
Deltochilum lobipes Bates, 1887 1 0
Deltochilum scabriusculum Bates, 1887 1 0
Alpha Diversity
* Mario Zunino identified these species and recommends a taxonomic revision of this group
of Onthophagus.
72
Appendix 3
The distribution and alpha diversity of Scarabaeinae, Geotrupinae and Silphidae species
in sites with expected capture rates above 85%.
Tropical Landscape
Community tf tfr pt ptr svt
Site 1 2 3 4 5 6 1 2 1 2 3 4 1 2 1 2
Species
Onthophagus batesi Howden y Cartwrigt, 1963 1 1 1 0 0 0 0 0 1 0 1 1 1 0 1 0
Onthophagus schaefferi Howden y Cartwrigt, 1963 0 0 0 0 0 0 1 0 1 1 1 1 1 0 0 0
Onthophagus hoepfneri Harold, 1869 1 1 1 1 1 1 0 0 0 0 0 0 0 0 1 1
Onthophagus igualensis Bates, 1887 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0
Digitonthophagus gazella (Fabricius, 1787) 0 0 0 0 0 0 0 0 1 1 1 1 1 0 1 0
Dichotomius amplicollis (Harold, 1869) 1 1 1 1 1 1 1 1 0 0 0 0 0 0 1 0
Uroxys boneti Pereira y Halffter, 1961 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0
Canthidium puncticolle Harold, 1868 1 1 0 0 1 1 0 0 0 1 0 1 1 1 0 1
Dichotomius colonicus (Linneo, 1767) 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1
Copris incertus (Say), 1835 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 1
Copris lugubris Boheman, 1868 0 0 0 0 0 0 0 0 1 0 1 1 1 1 1 1
Phanaeus endymion Harold, 1863 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0
Phanaeus tridens Castelnau, 1840 1 1 0 0 1 0 0 0 0 0 1 1 1 1 0 0
Coprophanaeus telamon Harold, 1863 1 1 1 1 1 0 0 1 0 1 0 1 0 1 0 0
Coprophanaeus pluto (Harold, 1863) 1 1 0 0 0 1 1 0 1 0 0 0 1 0 0 1
Sisyphus mexicanus Harold, 1863 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0
Eurysternus mexicanus Harold, 1869 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0
Euoniticellus intermedius Reiche, 1849 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0
Canthon (Glaphyrocanthon) femoralis (Chevrolat,1834) 0 1 1 0 0 1 0 0 0 0 0 0 0 0 1 0
Canthon (G.) leechi (Martínez Halffter y Halffter,1964) 1 1 1 1 1 1 1 0 1 1 0 1 1 0 1 1
Canthon (G.) zuninoi Rivera y Halffter, 1999 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0
Canthon (G.) moroni Rivera y Halffter, 1999 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0
Canthon (G.) antoniomartinezi Rivera y Halffter, 1999 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Canthon (Canthon) indigaceus chevrolati Harold, 1868 0 1 1 1 1 0 0 0 0 1 1 0 1 0 1 1
Canthon (C.) cyanellus Le Conte, 1859 1 1 1 1 1 1 0 0 1 1 0 1 0 0 1 1
Deltochilum gibbosum Bates, 1887 0 0 1 1 1 1 1 1 1 1 0 0 0 1 0 1
Deltochilum lobipes Bates, 1887 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Deltochilum scabriusculum Bates, 1887 0 1 1 0 1 0 0 1 0 1 0 1 1 0 1 0
Oxelytrum discicolle (Brullé, 1840) 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0
Alpha Diversity 10 14 15 8 10 11 5 4 11 11 9 14 12 7 13 10
73
Community cf of ofr por pof cm cp pc prof gc svo ppor prpo
Site 1 2 3 1 2 1 1 2 3 1 1 2 1 2 1 2 1 1 2 1 1 2 3 1
Species
Onthophagus schaefferi Howden y
0 0 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0
Cartwright,1963
Onthophagus hoepfneri Harold, 1869 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1
Onthophagus incensus (Say, 1835) 1 1 1 0 0 1 1 1 1 0 1 0 1 1 1 1 1 1 1 0 1 1 1 1
Onthophagus cyanellus Bates, 1887 1 1 1 1 1 1 1 0 1 1 0 0 1 1 0 1 0 1 1 1 1 1 1 1
Onthophagus corrosus Bates, 1887 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1
Onthophagus nasicornis Harold, 1869 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Onthophagus mextexus Howden, 1879 0 1 0 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 1 0 1 1
Onthophagus subcancer Howden, 1973 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Onthophagus aureofuscus Bates, 1887 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Onthophagus chevrolati retusus Harold, 1869 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 1 0 0 0 1 1 1 1
Onthophagus rhinolophus Harold, 1869 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
Dichotomius amplicollis (Harold, 1869) 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0
Dichotomius colonicus (Say, 1835) 0 1 1 0 0 0 0 0 0 0 1 0 1 1 1 1 0 0 0 0 0 0 0 0
Dichotomius satanas Harold, 1867 1 1 1 1 0 0 0 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 0
Ateuchus illaesum Harold, 1868 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
Scatimus ovatus Harold, 1862 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0
Ontherus mexicanus Harold, 1869 1 1 0 1 1 0 0 1 0 1 0 0 0 0 0 0 1 0 0 1 1 1 1 1
Copris incertus (Say, 1835) 1 0 1 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 0 1 0 1 0
Phanaeus endymion Harold, 1863 1 0 1 0 0 1 0 0 0 0 1 1 1 1 0 0 0 0 1 0 0 0 0 0
Phanaeus amethystinus Harold, 1863 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1
Coprophanaeus telamon Harold, 1863 1 0 1 0 0 1 0 0 0 0 1 1 1 1 1 1 0 1 1 0 0 0 0 0
Eurysternus magnus Castelnau, 1840 0 1 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0
Euoniticellus intermedius Reiche, 1849 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
Canthon humectus (Say, 1832) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0
Deltochilum mexicanum Burnmeister, 1848 1 1 1 1 1 1 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0
Onthotrupes nebularum (Howden, 1964) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0
Oxelytrum discicolle (Brullé, 1840) 0 1 1 1 1 1 1 1 1 1 0 1 1 1 0 0 0 1 1 1 0 1 1 1
Nicrophorus olidus Matthews, 1888 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 0 0 0 1 1 0 0 1 0
Nicrophorus mexicanus Matthews, 1888 0 0 0 1 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 1 1
Tanatophilus graniger (Chevrolat, 1833) 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1
Alpha Diversity 10 13 12 10 8 9 6 6 5 8 8 5 13 12 10 7 6 6 8 7 9 5 11 11
74
Montane Landscape
Community pf pfr ff pa gp gf gpa hav
Site 1 2 3 4 1 1 1 1 2 3 4 1 1 1 2 3
Species
Onthophagus mextexus Howden, 1970 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0
Onthophagus aureofuscus Bates, 1887 0 1 1 1 0 0 1 0 0 0 1 1 0 0 0 0
Onthophagus chevrolati retusus Harold, 1869 1 0 0 0 0 1 0 1 1 1 0 1 0 0 0 0
Onthophagus fuscus orientalis Zunino y Halffter, 1988 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0
Onthophagus lecontei Harold, 1871 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0
Onthophagus chevrolati chevrolati Harold, 1869 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Onthophagus hippopotamus Harold, 1869 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
Copris armatus (Harold), 1869 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0
Ontherus mexicanus Harold, 1869 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0
Phanaeus amethystinus Harold, 1863 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
Canthon humectus (Say), 1832 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0
Onthotrupes nebularum (Howden), 1964 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0
Halffterius rufoclavatus (Jekel), 1865 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0
Onthotrupes herbeus (Howden), 1964 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Ceratotrupes bolivari Halffter y Martínez, 1962 1 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0
Nicrophorus mexicanus Matthews, 1888 1 1 1 1 1 0 1 0 1 0 0 0 0 1 0 0
Tanatophilus graniger (Chevrolat), 1833 0 1 1 0 1 1 1 0 0 0 0 0 0 0 1 0
Alpha Diversity 3 5 3 2 2 3 5 6 7 7 3 5 2 1 1 0
75
Appendix 4
List of the cited species for the Cofre de Perote region, but not collected in this article.
In searching for information on the species that might have escaped our collection efforts in
the three landscapes in the Cofre de Perote region, we have carried out a thorough review of
both collections and the literature. The species that have cited in some way for the region, but
not collected by us are listed here.
Nicrophorus marginatus Fabricius, cited by Peck and Anderson (1985) for Xalapa. We have not
collected this species, nor have any of our colleagues who carry out studies in the area of
Xalapa. N. marginatus is widely distributed in southern Canada, the majority of the United States
of America, and enters northern Mexico (Coahuila, Durango) and extends to Mexico City D.F.
and Puebla (in the centre of the Mexican High Plateau). According to Peck and Anderson’s data
(1985) this is the only record of this species for Veracruz State.
Copris laeviceps Harold is cited by Matthews (1965) for Xalapa, but was based on specimens
collected by Hoege (British Museum). We are quite familiar with the distribution and ecological
requirements of this species. It is associated with the rainforest and conditions that are more
tropical than those of Xalapa. It is possible that this identification is the result of the erroneous
labelling that frequently occurred in the 19th century.
Copris rebouchei Harold is a species from the Balsas River Basin. Matthews (1965) includes a
site in Coatepec which would be the most eastern of its distribution area. Coatepec is within our
study region and is not an impossible site for the collection of C. rebouchei. However, if the
specimens on which Matthews bases the record correspond to this site, then it is totally marginal
to its main area of distribution. The other site reported in Veracruz: Presidio, is clearly an error.
Many years ago, one of the authors (G. Halffter) received from a collector who lived in Presidio
material erroneously referenced to this location; and it is from there that the specimens labelled
from Presidio come.
Phanaeus mexicanus Harold is cited by Edmonds (1994) for sites near our transect, but located
either on the Pico de Orizaba volcano slope or in the most tropical and rainy region of Los
Tuxtlas. P. mexicanus appears to have a vicariant distribution with P. scutifer which is found in
the transect.
Phanaeus sallei Harold is cited by Edmonds (1994) for Banderilla, Xalapa and Cosautlán - all
places that are located in our transect. We have not collected this species which tends to have
the rest of its distribution in more tropical areas than the places cited. Even so, its presence as
cited by Edmonds does not seem impossible since it has been collected in cloud forest in
Chiapas State, in the Montebello Lagoons and in rainforest. Perhaps its present is restricted to
the lowest parts of some ravines in our transect.
Literature Cited
Edmonds, W. D. 1994. Revision of Phanaeus MacLeay, a New World Genus of Scarabaeinae dung beetles
(Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science, 443: 1-105. Natural History Museum
of Los Angeles County.
Matthews, E. G. 1961. A revision of the genus Copris Müller of the Western Hemisphere (Coleoptera:
Scarabaeidae). Entomologia Americana, 6 (N.S.): 1-137.
Peck, S. B. & R.S. Anderson. 1985. Taxonomy, phylogeny and biogeography of the carrion beetles of Latin
America (Coleoptera: Silphidae). Quaestiones Entomologicae, 21: 247-317.
76

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Acta Zoologica Mexicana (n.s.

) 90: 27-76 (2003)

GAMMA DIVERSITY: DERIVED FROM AND A DETERMINANT

organisms to spatial heterogeneity. Gamma diversity depends primarily on

MATERIALS AND METHODS

Landscape units, sampling sites and capture methods

Part of these landscapes is covered by an extensive zone of rocky lava flows

Abbreviations used for plant communities in figures, tables & appendices:

This index is similar to that of Whittaker (1972), except that it is expressed as

A total of 16,152 specimens belonging to 60 species were captured from the

Alpha diversity by site

Mean alpha diversity by community

Cumulative species richness (cumulative alpha diversity) of the vegetation

Shared and exclusive species

Mean beta diversity among vegetation communities

Community Beta Diversity

In the tropical landscape gamma diversity was 31 species, 30 Scarabaeinae

DISCUSSION AND CONCLUSIONS

species, or those that take advantage of the edges of communities, such as

Complementarity among landscapes

It is interesting that disturbance by human activity appears to have been

3. Alpha diversity is one of the most frequently used values to characterize

that was created or at least expanded by deforestation. The species of this

taken from an entity (community). The cumulative richness of species for a

Acosta, P. R. 1986. La Vegetación de la Sierra de Manuel Díaz, Veracruz, México. Tesis

Deloya,C., G. Ruíz Lizárraga & M. A. Morón. 1987. Análisis de la entomofauna necrófila en la

Martín-Piera F. & J. M. Lobo. 1993. Altitudinal distribution patterns of copro-necrophage

Recibido: 11 de julio 2002

Species Landscape Source Site Altitude No.

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