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Journal of Biogeography (1998) 25, 10931103

The geographic ranges of mammalian species in South America:


spatial patterns in environmental resistance and anisotropy
Ani :N: Ricci ro
1
, JonN H. L:v1oN
2
and Ti x M. Bi:cxniN
2 1
Centro Regional Universitario Bariloche-UNC,
Unidad Postal Universidad Nacional del Comahue (8400) S.C. de Bariloche, Rio Negro, Argentina,
2
NERC Centre for Population Biology, Imperial College at Silwood Park, Ascot, Berkshire, SL5 7PY, U.K.
Abstract. We analyse the geographical distribution of South occurs at the limit between the Guayano-Brazilian and
Andean-Patagonean subregions. R
50
data do not support the Americanmammalianspecies using twobiogeographic indices:
latitudinal Rapoport eect: the most widespread species locate environmental resistance (R
50
) and anisotropy (A
50
). R
50
in the eastern portions of Brazil, the most restricted ones are quanties the loss of biotic resemblance occurring from any
in association with the Andes. A
50
data support Janzens point in the map to the rest of the continent. A
50
quanties
prediction: the eect of mountains on species distributions is
the extent towhichthe perimeter: area ratioof the geographical
greater in equatorial and central regions of the Andes rather
ranges of all species whose distributions overlap at any
than in the south. R
50
-and A
50
-contour maps reveal that,
particular location depart from the perimeter: area ratio of a
mainly due to the eects of the lie of the land and likely
circle. We test for the latitudinal Rapoport eect that predicts
dierences in the history of the fauna, the continent has a
an increase in the range-sizes of mammalian species, and
biogeographic texture which must have major constraints
hence a decrease in the values of R
50
, towards the south. We
on local ecological patterns and processes. This stresses the
test for the eect of mountains on species ranges, given that
importance of considering the role of biogeographic structure
Janzens argument that mountain passes are higher in the
in the analyses of geographical gradients in species
tropics implicitly predicts greater anisotropy in the tropics.
distributions.
Continental spatial patterns of variation in R
50
and A
50
suggest
a biogeographic division of South America consistent with
most classical zoogeographical classications proposed for the Key words. South America, mammals, zoogeography,
geographical range, Rapoport eect, latitude. continent. Rapid change in mammalian range-sizes and shapes
Resumen. Analizamos la distribucio n geograca de areas geogracas de los mam feros ocurren en el l mite entre
especies de mam feros sudamericanos usando dos ndices las subregiones Guayano-brasilen a y Andino-patago nica.
biogeogracos: resistencia ambiental (R
50
) y anisotrop a R
50
no sustenta el efecto latitudinal de Rapoport: las especies
(A
50
). R
50
cuantica la perdida de la resemblanza bio tica mas ampliamente distribuidas se localizan en el este de
desde cualquier punto del mapa hacia el resto del continente. Brasil, y las de distribucio n geograca mas restringida
A
50
cuantica co mo la relacio n per metro: area de los rangos asociadas a los Andes. Los datos de A
50
convalidan la
geogracos de todas las especies cuyas distribuciones se prediccio n de Janzen: el efecto de las montan as sobre la
superponen en una localidad particular se apartan de la distribucio n geograca de las especies es mayor en regiones
relacio n per metro: area de un c rculo. Probamos el efecto centrales y ecuatoriales de los Andes y menor en el sur. Los
latitudinal de Rapoport, que predice un incremento en el mapas de R
50
y A
50
revelan que, principalmente debido a
taman o de los rangos geogracos de las especies de
las caracter sticas naturales de la supercie continental y
mam feros y, por lo tanto, una disminucio n en los valores
probables diferencias en la historia de la fauna, el continente
de R
50
hacia el sur. Probamos el efecto de las montan as
tiene una textura biogeograca que debe ejercer restricciones
sobre los rangos geogracos de las especies, dado que el
mayores sobre los patrones y procesos ecolo gicos que
argumento de Janzen los pasos montan osos son mas altos
ocurren a escala local. Esto subraya la importancia de
en los tro picos impl citamente predice mayor anisotrop a
considerar el papel de la estructura biogeograca en analisis
en los tro picos. Los patrones continentales de variacio n
de gradientes geogracos en la distribucio n de las especies.
espacial de R
50
y A
50
sugieren una divisio n biogeograca
de America del Sur consistente con las mayor a de las
Palabras claves. America del Sur, mam feros, clasicaciones zoogeogracas clasicas propuestas para el
continente. Cambios rapidos en el taman o y forma de las zoogeograf a, rango geograco, efecto Rapoport, latitud.
Correspondence: Adriana Ruggiero, Centro Regional Universitario Bariloche-UNC, Unidad Postal Universidad Nacional del Comahue, (8400) S.C.de
Bariloche. Rio Negro, Argentina.
1998 Blackwell Science Ltd 1093
1094 A. Ruggiero, J. H. Lawton and T. M. Blackburn
overlap at any particular location depart from the perimeter:
INTRODUCTION
area ratio of a circle. The basic underlying idea is that a
continent can be considered to be isotropic for any location Studies of spatial patterns in range-size at a continental
scale usually focus on the kind of geographical gradients if the geographical ranges of species inhabiting that
particular site approach circularity. In other words, low that occur on single linear dimensions of species distribu-
tions. For instance, the latitudinal and altitudinal extent of values of anisotropy at any given location suggest that
species living at that site face the same amount of species have been used to test for the latitudinal and
altitudinal versions of the Rapoports rule (e.g. Stevens, environmental resistance in all directions from the centre
toward the outside limits of their distribution. The presence 1989, 1992; France, 1992; Rohde et al., 1993; Macpherson
& Duarte, 1994; Roy et al., 1994; Lyons & Willig, 1997). of a barrier, whether physical or biological, that limits range-
expansion along any particular linear dimension distorts the Although the value of analysing spatial patterns of species
distribution using information contained in two (or more) geographical ranges of species inhabiting that location, and
this will be reected in an increase in Rapoports index of linear dimensions of geographical range combined has been
theoretically recognized (Gaston, 1994), in practice anisotropy. The indices of environmental resistance and
anisotropy are therefore independent descriptors of the empirical studies have rarely done this; examples include
Pagel et al. (1991), Letcher & Harvey (1994), Blackburn & geometry of the geographical ranges of species: for example,
a given location can have a low value of anisotropy yet it Gaston (1996a), and Williams (1996). To the extent that
dierent linear measurements of range-size are usually well may have a low or a high value of environmental resistance.
The goal of the present analysis is to elaborate maps correlated (e.g. Stevens, 1992; Quinn et al., 1996), and that
spatial variation in dierent linear dimensions of species that depict continental spatial patterns of variation in
environmental resistance and anisotropy (hereafter ranges along latitude, longitude or elevation can be easily
compared, there seems to be a general belief that uni- abbreviated as R
50
and A
50
), which are used to suggest a
biogeographic division of South America exclusively on the dimensional analyses of spatial patterns in species
geographical ranges are sucient. The present analysis, basis of variation in the sizes and shapes of the geographical
ranges of mammalian species. Geographic gradients in the however, will try to show that indeed there is considerable
added value in the kind of results that can be obtained spatial patterns of R
50
and A
50
also allow us to test two
theoretical predictions about the geographical distributions fromthe study of species distribution by using more detailed
information on the size and shape of species ranges. of species.
In the present study we re-analyse data on the geo-
graphical distributions of South American mammalian 1. The Rapoport eect (Stevens, 1989) predicts an
increase in the size of the geographical ranges of species species (data from Ruggiero, 1994) to examine how the size
and shape of geographical ranges vary as a function of with latitude. A previous analysis of the latitudinal
extent of mammalian species in South America provided latitude and longitude. However, instead of taking into
account the spatial pattern of variation in single linear equivocal evidence, suggesting that the pattern varied
depending upon the taxa considered (Ruggiero, 1994). dimensions of species distributions (for instance, in
Ruggiero (1994) the latitudinal extents of these mammalian In the present analysis, the spatial pattern of variation
in R
50
is used to infer the relative dierences in the size species are used to test for the latitudinal Rapoport eect
(Stevens, 1989)), here we consider spatial variation in the of the geographical ranges of mammals in tropical vs.
temperate regions of the continent. Adecrease in R
50
must shape and size of species ranges along both (latitudinal
and longitudinal) gradients simultaneously, using two occur towards the south if the range-sizes of mammalian
species increase with latitude as the Rapoport eect biogeographic indices introduced by Rapoport (1975, 1979,
1982): environmental resistance and anisotropy. predicts.
2. Mountains act to deformthe shape of the geographical Rapoport (1975, 1979, 1982) used his index of environ-
mental resistance (a formal denition will be given later) to ranges of species (Rapoport, 1975, 1982; Brown &
Maurer, 1989; Lawton et al., 1994; Brown, 1995). The infer the eects of physical and biological barriers on the
size of the geographical distributions of species inhabiting major NS lines of orientation of the Andean mountain
ranges in South America are expected to deform the a particular location. According to Rapoport, the magnitude
of environmental resistance at any given location depends shape of the geographical ranges of mammalian species.
We use the spatial pattern of variation in A
50
to test for on (i) the size of the geographical ranges of all species
recorded at that point and (ii) the loss of biotic resemblance an increase in the asymmetry of the geographical ranges
of species toward the west of the continent. More occurring from that point to the rest of the continental
surface. Clearly (i) and (ii) are interrelated: the smaller the specically, Janzens (1967) argument that mountain
passes are higher in the tropics, because a lowland size of the geographical ranges of species living at a dened
location, the higher the loss of species with distance from tropical species that attempts to cross a mountain pass
will face environmental conditions more extreme than it that point, with this being translated into a higher value of
environmental resistance. normally experiences, whereas this is less likely to be
true for lowland species resident at higher latitudes, Rapoports (Rapoport, 1975, 1979, 1982) index of
anisotropy (a formal denition will be given later) quanties implicitly predicts greater anisotropy (distortions and
restriction of range) in the tropics. We test this prediction the extent to which the perimeter: area ratios of the
geographical ranges of all species whose distributions here.
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
Environmental resistance and anisotropy in South America 1095
METHODS The creation of equiprobabilistic maps
The rst step in obtaining an equiprobabilistic (=isodensity)
Choice of the data
map (sensu Rapoport, 1975, 1979, 1982) is to consider
species richness at an arbitrary central point in this case Distributional information was compiled for 536 species
a cell in the grid-map-as 100%. Then, the proportion of in the following taxa: Marsupialia, Edentata, Chiroptera,
species at this central point present at every one of the Primates, Carnivora, Artiodactyla, Perissodactyla, Lago-
remaining cells in the grid-map is calculated, thus obtaining morpha and Rodentia (only Hystricognathi) (Cabrera, 1957;
a probability of nding a species present at that central Harris, 1968; Hershkovitz, 1977; Franklin, 1982; Honacki
point in any part of the rest of the continent. Equal et al., 1982; Koopman, 1982; Mares & Ojeda, 1982; Mayr
probability cells are linked by contour lines to produce the & Brand, 1982; Streilein, 1982; Wetzel, 1982; Melquist, 1984;
equiprobabilistic map (Fig. 1). In the present study, we Eisenberg, 1987; Emmons, 1990). These are same data
analysed the 481 equiprobabilistic maps obtained by analysed in Ruggiero (1994), which should be consulted
considering each one of the cells in the grid map in turn, for details. To minimize the inuence of taxon-dependent
as the central point. Probability values for each cell in the variation in the spatial patterns of species distributions (see
grid map were then calculated using P.An.D.A. (Programa Ruggiero, 1994), the present study uses data from all species
Para el Analisis de Datos Areogracos developed by E.M. throughout.
Lopasso, see Lopasso & Ruggiero, 1995). Contour lines Most of the sources used to delimit species ranges follows
were obtained by the application of a geostatistical technique the nomenclature in Honacki et al. (1982). However, a
that uses an estimation of the semivariance of the data to number of nomenclatural changes have occurred for Central
perform interpolations (kriging: see Matheron (1963) and and South American mammals since 1982 (Patterson, 1996).
Rossi et al. (1992)). Sixty-two Neotropical taxa considered to be species in 1982
are nowregarded as synonyms (adding to the ranges of other
species), 173 have resurrected from synonymy (subdividing
other species ranges), and sixty additional species have
The estimation of the environmental resistance (R
50
)
been newly discovered and named. These changes aect
and anisotropy (A
50
)
31.5% of Neotropical mammal species, based on a total of
937 in the region (see Patterson, 1996). Applying these
Environmental resistance for any given cell in the grid map
changes to the 536 species used by Ruggiero (1994) results
was dened by selecting the 50% probability level for the
in a revised total of 577 species. To ensure that the patterns
correspondent equiprobabilistic map (see Fig. 1). The choice
we document herein are not simple consequence of the
of 50% is arbitrary (see Rapoport, 1975, 1979, 1982 for a
arbitrary choice of one or other of these taxonomies, we
discussion). The area enclosed by this 50% isodensity line
perform all the analyses using both. For the revised
(a
50
) was measured and the environmental resistance (R
50
)
classication, we used the descriptions in Wilson & Reeder
for that cell calculated as R
50
=1-(a
50
/a
c
), where a
c
is the total
(1993), which document most of the changes that have
continental area. R
50
can take values between 0 (indicating
occurred both nomenclaturally and geographically, to make
maximum expansion of species, and low losses of faunistic
the necessary changes on each species range. Fifteen
resemblance with distance from the central point) and 1
species were excluded from the analyses because they have
(maximum resistance, and quick loss of faunistic similarity).
an uncertain taxonomic status or we lack of enough
Once a value of R
50
is calculated for each one of the 481
information to adequately delimit their geographical ranges.
cells in the grid map, equal R
50
cells are linked to obtain
These species are: Thylamys pallidior, Marmosops handleyi,
an R
50
-contour map, as described in the previous section.
Didelphis aurita (Didelphimorphia); Callithrix kuhlii,
Anisotropy for any given cell in the grid map was
Leontopithecus caissara, Callicebus dubius (Primates);
calculated using the length of the perimeter of the 50%
Proechimys bolivianus, Proechimys steerei, Mesomys
probability level (i
50
) (see Fig. 1). Anisotropy (A
50
) for that
obscurus (Rodentia); Lonchophylla dekeyseri, Anoura
cell is estimated as (A
50
)=(i
50
/8a
50
)3.54, where 8a
50
is the
latidens, Sturnira luisi, Artibeus mbriatus, Artibeus
square root of the area covered by the 50% probability level
obscurus, Tadarida espiritosantensis (Chiroptera).
and 3.54 is roughly the ratio of perimeter: 8area for a
circle. A
50
approaches zero when the geographical ranges
of species inhabiting the central point tend to be circular.
Data analysis
The more the shape of the geographical ranges of species
departs from circularity, the more anisotropy increases.
The geographical ranges of each mammal species were
Values of A
50
for each of the 481 cells in the grid map were
drawn on a standardized grid map (481 quadrats, 200 km
calculated, and used to produce an A
50
-contour map.
on a side) overlaying an azimuthal projection (scale=1: 40,
000,000). The presence-absence (10) of each species in each
cell of the grid map was recorded. Based on these data,
481 equiprobabilistic maps (=isodensity maps) (sensu
RESULTS
Rapoport, 1975, 1979, 1982) were obtained, each cor-
responding to one of the cells in the grid map; these 481 maps Spatial patterns in R
50
and A
50
based on the updated data
base (N=577 species) dier slightly from those obtained were used to estimate values of environmental resistance and
anisotropy for every cell in the grid map, as explained below. from the original data base (N=536 species). Given that
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
1096 A. Ruggiero, J. H. Lawton and T. M. Blackburn
FIG. 1.
Examples of equiprobabilistic maps centred at the northern (a,b,c) and southern (d,e,f) halves of South America. indicates the location
of the central point. The area and perimeter of the 0.5 isoline is used to estimate, respectively, a value of R
50
and A
50
for each of the 481
cells in the grid map (see text for details).
the main trends repeat in both analyses, we only present R
50
values are comparatively low in the northern half of
South America (0.40.6, Fig. 2), and increase progressively here the R
50
-and A
50
-maps corresponding to the analysis of
the updated data base. in a SE-NW direction, from eastern Brazil to the northern
portions of the Andes. As a consequence, within this region,
at least three distinct subregions can be distinguished (1)
Eastern Brazil (R
50
=0.4), extending mostly from the
Trends in environmental resistance (R
50
) and the
Amazon river and its outows, to the south and east of the
division of South America into biogeographic zones
continent. (2) Central Amazonia (R
50
=0.40.5), extending
mostly from the Amazon river to the north and west of The division of South America into two (north and eastern
v. south and western) biogeographic regions is the most South America. (3) North-western South America (R
50
=
0.50.6), generated mainly by the presence of the Andean obvious feature revealed by spatial patterns in R
50
(see
Fig. 2). The division is dened by a clumping of isolines that mountain ranges in the west of the continent. The greatest
values of R
50
(0.6) in the north half of South America highlight regions of rapid change in values of environmental
resistance. Division into northern and southern halves of occur when altitude increases in the northern Andes. In the
southern half of South America, values of R
50
are higher the continent begins at the Gulf of Guayaquil (in Ecuador),
continues close to the Andes to the north of Argentina, and and tend also to increase westward. The Andean region -
with the highest values of environmental resistance (R
50
= then follows a SE direction to reach the Uruguayan coast
(Fig. 2). 0.9)-stands out distinctly in Fig. 2.
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
Environmental resistance and anisotropy in South America 1097
FIG. 2.
Spatial patterns of variation in the values of environmental resistance, R
50
. The 0.5 isolines of each one of the 481 equiprobabilistic maps
(Fig. 1) were used to estimate a value of environmental resistance (R
50
) for each one of the cells in the grid-map. Interpolations among cells
were performed by kriging to produce this R
50
-contour map (see detailed explanation in text).
The relationship between spatial patterns in R
50
and mammalian fauna. Variation in R
50
suggests that range-
sizes are large in the tropical portions of the continent and range-size
decrease toward the south. Moreover, R
50
reveals more
subtle trends than the one predicted by the Rapoport eect. The trends in R
50
clearly suggest that Rapoports eect
(Stevens, 1989) does not hold for the South America The most widespread mammalian species occur in the
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
1098 A. Ruggiero, J. H. Lawton and T. M. Blackburn
eastern regions of Brazil, and those with the most restricted taken by Williams (1996) and Mourelle & Ezcurra (1997),
who used measures of faunal and orist turnover to identify distributions are located in the southern hemisphere, along
the west of South America, clearly in association with transition zones between dierent biogeographic regions.
Despite the simplicity of our approach, it is remarkable the presence of the Andean mountain ranges. Even if
comparisons are restricted to the Andean region, patterns that the biogeographic scheme produced by spatial patterns
of variation in R
50
is consistent with most classical zoo- in R
50
suggest that the northern portion of the Andes
harbours relatively more widespread species than do the geographical classications proposed for the continent; a
similar conclusion was also reached by Williams (1996) for southern portions.
his broader-scale analysis of the global bumble bee fauna.
The most striking similarity in the present case is the
Trends in anisotropy (A
50
)
coincidence between the region of major change in R
50
(at
the limit between the northern and the southern halves of The distinction between the northern and southern half of
South America is also suggested by spatial patterns of South America), and the separation between the Guayano-
Brazilian zoogeographic subregion and the Andean- variation in A
50
(Fig. 3). An increase in the values of
anisotropy occurs toward the south, due mostly to the Patagonian one proposed by Hershkovitz (1969: Fig. 1).
The region also corresponds to the so called subtropical narrowing of the continental landmass. Nonetheless, other
features in the spatial pattern of variation in A
50
seem line proposed by Sclater (1858; quoted in Rapoport, 1968)
and Wallace (1876) (see Rapoport, 1968: Figs 3 and 4; not to be merely the consequence of the geometry of the
continent. The most evident, perhaps, is an heterogeneous Hershkovitz, 1969: Fig. 1). Referring to the biogeography
of all classes of terrestrial animals in the Neotropics, Wallace pattern of peaks in A
50
, following roughly the position and
direction of the major frontier of change in R
50
(compare considered all tropical South America to form a unique
subregion. The R
50
-and A
50
-contour maps quantify what Figs 2 and 3). This region is separated from the other two
most highly anisotropic regions of the continent (i.e. the Wallace (1876) described as the . . . remarkable uniformity
of animal life over all the tropical continental portions of Andes and Patagonia) by a zone with the lowest values of
anisotropy for the continent, which roughly corresponds to South America . . . where . . . even the Andes do not seem
to form such a barrier as has been supposed, with most of the position of the major plains (Pampas and Chaco) in
Argentina. the genera being found on both sides . . . .
The comparatively lower values, and relatively more The northern half of the continent diers from the
southern half in its constancy in values of A
50
. In contrast, uniform patterns, of spatial change in R
50
recorded in
the northern half of South America presumably reect a spatial patterns of variation in A
50
that occur with latitude
along dierent portions of the Andes suggest that mountains relatively low rate of mammalian species turnover between
habitats in the tropics. There is some evidence, for example, do indeed act to deform the shape of the range-size of
mammalian species. However, as Janzen (1967) suggested, suggesting an overlap between the Cerrado mammalian
fauna and neighbouring biomes: fty-ve out of the 100 eects are heterogeneous: mountains seem to have a bigger
eect around the equator with the highest values of A
50
species recorded in the Cerrado also occur in the Amazon
basin, thirty-nine in the Caatinga, fty-three in the Chaco recorded at about this latitude and along most of the
central portions of the Andes, and tend to be less overall and fty-four in the pluvial rainforest of the south-east of
Brazil (Redford & Fonseca, 1986). Mares (1992) pointed in the northern and southern Andes.
out that 68% of mammalian species, and 95.4% of genera,
found in the Amazon lowlands are also distributed in
DISCUSSION
other habitats. However, a dierent picture might emerge
if specic details of habitat use were considered (e.g. the
R
50
-based biogeographic divisions of South America: a
55% of cerrado species proposed to be shared with the
new bottle for old wine?
Amazon are mainly those found in gallery forests that
cross the cerrado rather than cerrado species per se; B.D. The spatial patterns in R
50
(Fig. 2) emerge from an objective,
quantitative analysis of the geographical ranges of 577 Patterson, personal communication). Patterson also pointed
out that, for example, nonvolant mammals in Atlantic species of mammals. It makes no assumptions about, for
instance, the uniqueness of certain species in certain areas, Forest habitats are strikingly dierentiated from those in
other forest types (examples include: Leontopithecus, as has traditionally been the case in biogeography (see, e.g.
Cabrera & Willink (1980), Rapoport (1968) and Morrone Brachyteles (Primates); Chaetomys, Euryzygomatomys,
Kannabateomys, Nelomys (Caviomorpha)) yet this (1996) for a review of the classical biogeographic divisions
of South America, and in particular Cabrera & Yepes distinction is not even faintly suggested in Fig. 2. His
interpretation is that either this is a function of grid scale (1940) and Hershkovitz (1969) for mammalian-based
zoogeographic divisions of the continent). All species are used (e.g. see Anderson & Marcus, 1993), or else the
resistance and anisotropy approaches are more sensitive to used equally in our analysis. The present analysis also diers
from the conceptual approach that cladistic biogeography species richness rather than to species composition, thus
emphasizing shared widespread species more than unique uses to dene hierarchical relationships between dierent
areas of endemism (see e.g. Humphries & Parenti (1986) and restricted ones. Clearly, all these problems deserve more
attention. Nonetheless, patterns in R
50
emerging from the and Morrone & Crisci (1996) for general reviews). The
present analysis is, however, closely related to the approach analysis of the geographical distribution of the South
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
Environmental resistance and anisotropy in South America 1099
FIG. 3.
Spatial patterns of variation in the values of anisotropy, A
50
. The 0.5 isolines of each one of the 481 equiprobabilistic maps (Fig. 1) were
used to estimate a value of anisotropy (A
50
) for each one of the cells in the grid-map. Interpolations among cells were performed by kriging
to produce this A
50
-contour map (see detailed explanation in text).
American mammalian fauna taken as whole helps to birds. It is suggestive that the R
50
-contour map allows
roughly the same three subdivisions for mammals (see understand why Wallace (1876) argued that it was almost
impossible to divide the tropical portions of the continent Fig. 2).
However, the biogeographic subdivision of South into further zoogeographic subdivisions, although he
recalled that Sclater, Salvin and Newton recognized three America revealed by R
50
is not merely a new bottle for old
wine. The R
50
-contour map provides additional information subdivisions (Brazilian, Amazonian and Columbian) for
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
1100 A. Ruggiero, J. H. Lawton and T. M. Blackburn
that helps us understand previous disagreements among in A
50
could reect the eect of some kind of faunistic-
interference eect on species ranges, that could occur when authors working on dierent taxa. For example, as
previously also suggested by Ringuelet (1961), the separation the southern limits of the geographical distribution of species
living in northern half of South America meet the northern between the two subregions is not just as a line, but a region
in itself. Comparison of the spatial patterns in R
50
with limits of distribution of species living in the southern half
(compare Figs 2 and 3). The nal eect may be local previous delimitations of the boundary between northern
and southern faunas (summarized in Rapoport, 1968: 78), penetration zones of one fauna by another squeezing the
ranges of the loosing taxa. makes clear that Wallaces (1876) line roughly coincides
with much of the course of the 0.50.6 R
50
-isolines, whereas Finally, it has been suggested that when the perimeter:
area ratio of species geographical ranges becomes larger the Sclaters (1858) line approximately overlaps the 0.70.8
R
50
-isolines. Thus, at least in the case of the South American demographic and environmental stochasticity become more
important to determine the longevity of populations; mammals, the limit between the Guayano-Brazilian and
the Andean-Patagonian subregions can be idealized as the compressions and distortions of the geographical ranges
tend to be accompanied by a decrease in the abundance of region encompassed within the limits of Wallaces line in
the north and Sclaters line in the south. Note that the populations, that become more vulnerable to the eect of
deleterious disturbances and biotic interactions with the majority of subtropical lines summarized in Rapoport (1968)
for other taxa also lie between the limits of the 0.5 and 0.8 surrounding habitats, most strongly at the edge of species
ranges (see, e.g. Soule & Simberlo, 1986; Graves, 1988). R
50
-isolines on Fig. 2.
Both from this, and the faunal interchange point of view,
further eorts to delimit dierent biogeographic regions in
Spatial patterns in anisotropy: the imprint of
South America must not be regarded simply as an old-
topographic relief and faunistic encounters
fashioned way of doing biogeography. The limit between
the Guayano-Brazilian and Andean-Patagonian subregions Trends in A
50
conrm the widely held perception that the
geographical ranges of species are inuenced by changes in is more than a static line drawn on a map to indicate
changes in faunistic composition; it is potentially a zone of topographic relief. The shape of speciess ranges tend to be
more asymmetric (when compared to the shape of a circle) intense faunistic interactions and large-scale population
dynamics, and deserves more intensive and extensive towards the west of South America. The heterogeneous
pattern of variation in the values of A
50
, with peaks in ecological and biogeographical analysis.
anisotropy occurring at dierent locations along the western
but not along the eastern edge of the continent, is strong
Macroecological predictions in South America
enough to suggest that it arises mainly from the eect of
mountains that tend to deform the geographical ranges of Recent years have seen a resurgence of interest in the
form, determinants and consequences of geographical scale species (see Rapoport, 1975, 1982; Brown & Maurer, 1989;
Lawton et al., 1994; Brown, 1995), rather than merely being patterns in the structure of species assemblages (for major
reviews see Ricklefs & Schluter, 1993; Edwards et al., 1994; the consequence of a coastal eect. High anisotropy in the
west also contrasts with a dramatic decrease in values of Gaston, 1994; Brown, 1995; Rosenzweig, 1995), with the
growing recognition of a distinctive program of research A
50
in the central plain regions of the Pampas and Chaco,
in Argentina (Fig. 3), reinforcing the idea that there is into these issues, termed macroecology (Brown & Maurer,
1989; Brown, 1995). The prime concerns of this discipline indeed an eect of topographic relief on range-size and
shape. Less obviously and, as predicted by Janzen (1967), centre on attempts to understand spatial patterns in the
distribution, abundance and diversity of species at the A
50
-contour map suggests that the eect of mountains
on species distributions is more important in equatorial continental scales (e.g. Brown, 1995). In this regard, the
R
50
-and A
50
-contour maps (Figs 2 and 3) provide means of and central regions of the Andes compared with the southern
Andes (Fig. 3). interpreting spatial patterns in species distributions that
would be exceptionally dicult, if not impossible, to achieve The other most striking feature in the A
50
-contour map
is the region at the border between the northern and southern were only geographical variation in single linear dimensions
of species ranges considered (e.g. Ruggiero, 1994). halves of South America, where there is a heterogeneous
pattern of peaks in A
50
. The direction of this area of The R
50
-contour map suggests that the most striking
dierences between range-sizes of mammalian species in rapid change in mammalian range-sizes and shapes roughly
follows the direction of the arid northwestsoutheast South America do not occur in a NS direction, as the
Rapoport eect predicts (Stevens, 1989) but rather in an diagonal that appears to have existed in South America at
least since late Tertiary (i.e. middle Miocene), after the EW direction, with the most widespread species located in
the eastern portions of Brazil (R
50
=0.4, Fig. 2) and present tropical-temperate climatic zonation was established
in the continent (see, e.g. Pascual, 1984; Pascual & Ortiz the most restricted ones in association with the Andean
mountain ranges (R
50
=0.9, Fig. 2). A tendency for the Jaureguizar, 1990 for a detailed review of the historical
events). Nonetheless, at present, changes in mammalian most geographically restricted mammalian species to occur
in the west of South America was also observed by Ruggiero species distributions over that region could be driven not
just by geography but also by biotic interactions at the (1994; Fig. 5); however, the pattern is clearer using R
50
data.
Trends in R
50
indicate that, at least in South America, boundaries where the two faunas meet. M.A. Aizen
(personal communication) has suggested that these peaks variation in the range-size of species follows a much more
Blackwell Science Ltd 1998, Journal of Biogeography, 25, 10931103
Environmental resistance and anisotropy in South America 1101
complex pattern than that predicted by the Rapoport eect, Blackburn & Gaston, 1996a). It also raises the question:
are the kinds of processes aecting species distributions and hence it turns out to be more obvious why the climatic-
variability hypothesis (Stevens, 1989) is not a universal (and hence also species diversity) in those regions of rapid
change in range-size and range-shape dierent from, or just explanation for range-size variation at continental scale
(see, e.g. Rohde et al., 1993; Roy et al., 1994; Blackburn & more intense than, those in other portions of the continent?
Gaston, 1996a; Rohde & Heap, 1996; Rohde, 1996; Rahbek,
1997). The climatic variability hypothesis does, however,
ACKNOWLEDGMENTS
receive indirect support from the A
50
data; mountain passes
M.A. Aizen suggested the idea that the peaks in A
50
at do indeed appear to be higher in the tropics (Janzen, 1967).
the limit between northern and southern halves of South In other words, on the basis of the present analyses, tropical
America could result from the encounter between two species in the southern hemisphere do show narrower
dierent faunas, thus triggering o one possible environmental tolerances, and hence greater restrictions of
interpretation of the patterns of variation in A
50
presented range from the tropical Andes than from the southern
in this paper. Comments from B.D. Patterson and an Andes (Fig. 3). Why these eects do not translate into
anonymous reviewer were very useful to identify the limits of Stevenss (1989) predictions of larger ranges in more
resolution for this analysis. La Divisio n F sica de Reactores temperate species is unclear. One possibility is that
Avanzados (DiFra) del Centro Ato mico Bariloche (CNEA) dierences in the environmental tolerances of species are
allowed the use of their computer facilities to perform most tempered by variation in land shape with latitude in South
of the analysis of the data. The NERCCentre for Population America.
Biology, Imperial College at Silwood Park (University of The apparent contradiction between spatial patterns in
London) provided the academic environment and logistic R
50
and A
50
in the Andes is intriguing, given that R
50
data
facilities for A.R. to complete the manuscript. A.R. was do not support Janzens prediction. To the extent that A
50
supported by a CONICET fellowship (Resolucio n N 1377). data reects distortions in the shape of species distributions
A fellowship from Fundacio n Antorchas (Project N 13145/ that could occur rather independently of changes in species
10013) supported A.R. during the rst seven months at ranges-sizes (better reected by R
50
data), spatial patterns
the Centre for Population Biology. A.R. would not have of variation in A
50
could be more sensitive to the combined
been able to do this work without the help, support and eects of narrower climatic tolerances of species around the
love of E.M. Lopasso. tropics and a more complex topographic relief at those
latitudes where the highest altitudes in the Andes recorded.
We must, however, give notice that our test of Janzens
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