Ph. Williams et al., Interpreting biogeographical boundaries among Afrotropical birds: spatial patterns in richness gradients and species replacement, J BIOGEOGR, 26(3), 1999, pp. 459-474
Aim We analyse the geographical distribution of 1911 Afrotropical bird spec
ies using indices of three simple biogeographic patterns. The first index,
the frequency of species with range edges (T-e), is formulated to map direc
tly the density of species distribution limits, for comparison with the res
ults of traditional biogeographical classification and ordination procedure
s, in order to show variations in the strength and breadth of transition zo
nes. The other two indices are formulated to seek to distinguish as directl
y as possible between two components within these transition-zone patterns:
contributions from gradients in species richness (T-g); and contributions
from replacements among species (T-r). We test the ability of these indices
to discover the same boundaries among Afrotropical bird faunas as one popu
lar procedure for classifying areas (TWINSPAN) and then use them to look fo
r geographical trends in the different kinds of transition zones.
Location The analysis is restricted to the sub-Saharan or Afrotropical regi
on, excluding the Arabian Peninsula, Madagascar and all offshore islands.
Methods We record the presence of each species in 1961 1 degrees x 1 degree
s grid cells of the map. To apply the three indices, each (core) grid cell
in turn is compared with its neighbouring eight cells in the grid. The rang
e edges index (T-e) counts the number of species with range edges between t
he core cell and the surrounding cells. The richness gradients index (T-g)
counts the largest difference in species richness measured diametrically ac
ross the core cell in any direction when there is a consistent trend in ric
hness along this line of three cells. The species replacements index (T-r)
counts the number of species pairs recorded within a nine-cell neighbourhoo
d that are not corecorded within any of the cells. Values for each of the 1
961 grid cells are calculated and used to produce colour-scale maps of tran
sition zones.
Results Large-scale spatial patterns of variation in density of range edges
(T-e) are consistent with classifications of the same data and with most p
revious biogeographical classifications proposed for the region. Variation
in richness gradients (T-g) and species replacements (T-r) explain differen
t parts of this pattern, with transition zones around humid forests in the
equatorial region being dominated by species replacement, and transition zo
nes around deserts (most extensive in the north and south) bring dominated
by richness gradients.
Main conclusions The three indices distinguish the spatial arrangement and
intensity of different kinds of transition zones, thereby providing a first
step towards a more rigorous mechanistic understanding of the different pr
ocesses by which they may have arisen and are maintained. As an example of
one such pattern shown by our analyses of Afrotropical birds, there is evid
ence for a broad latitudinal trend in the nature of transition zones in fau
nal composition (following the latitudinal distribution of the different ki
nds of habitat transitions), from being dominated by species replacements n
ear the equator to being dominated by richness gradients further from the e
quator.