Ibis (2005), 147, 225–227
© 2005 British Ornithologists’ Union
Blackwell Publishing, Ltd.
Reply
Seeing the woodhoopoe for the trees: a response to
Simmons et al. (2005)
MICHAEL CUNNINGHAM
1
& MICHAEL I. CHERRY
2
*
1
Department of Genetics, University of Pretoria, Pretoria 0002, South Africa
2
Department of Botany and Zoology, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa
We conducted a preliminary mitochondrial DNA
sequence study of woodhoopoes with the aim of
acquiring diagnostic molecular markers to study
interbreeding between two poorly differentiated
forms (Cooper et al. 2001). This study revealed a
surprisingly close relationship between 12 violet-
mantled birds from two localities in northwestern
Namibia and green-mantled birds several thousand
kilometres away in South Africa and Kenya; we
also found negligible differentiation between these
violet-mantled birds and a single green-mantled bird
from a mixed flock in Namibia. These results led us
to question the phenotypic basis on which these
species are recognized, which we found to be inade-
quate for species diagnosis and inconsistent in
application. We are not the first to do so: the Violet
Woodhoopoe Phoeniculus damarensis was first described
by Ogilvie-Grant (1901), but was subsequently syn-
onomized with the Green Woodhoopoe P. purpureus
by Peters (1945). Fry (1978) treats them as separate
taxa, but Clancey (1985, p. 245) states that ‘the cur-
rent policy of according the Violet Woodhoopoe
specific status needs in-depth investigation’, a call
reiterated by du Plessis (1997a).
Observations of phenotypic variation are easily
canalized by systematic interpretation; what one
sees depends on what one expects to see. The central
issue in Namibian woodhoopoes concerns the signi-
ficance of the sole diagnostic trait – adult mantle
coloration (Clancey 1985). Simmons et al. (2005)
apparently believe that variation in mantle colora-
tion reflects a discrete difference between independ-
ently evolved species lineages, with distinct habitat
preferences, distribution and behavioural ecology.
They suggest that individuals with intermediate
mantle coloration arise through hybridization, and
that this process represents a threat to the locally endemic
violet-mantled form. We propose an alternative
interpretation in which the violet-mantled and inter-
mediate forms result from clinal variation across an
aridity gradient, with associated changes in habitat
use and group size probably due to the varying avail-
ability of tree holes for this communally roosting species.
We contend that the ecological gradient hypothesis
is more parsimonious, and more consistent with the
mitochondrial DNA sequence similarity and the
extensive distributional overlap between these inter-
grading forms. We also contend that in the absence of
a consistent phenotypic diagnosis these forms should
be considered conspecific, pending further systematic
research on geographical variation in green, violet
and mixed groups of woodhoopoes in Namibia.
Simmons et al. (2005) have criticized this work on
five grounds: (1) that our sampling is flawed, (2) that
our phylogenetic tree could be interpreted differently,
(3) that violet and green woodhoopoes differ beha-
viourally, (4) that there are biogeographical reasons
suggesting that the Violet Woodhoopoe is a proper
species and (5) that even near-endemic subspecies
should be given special status by conservationists. Here
we deal briefly with each of these points in turn.
(1) Given the degree of distributional overlap, incon-
sistency among authors and observers in the identifi-
cation and distribution of the Namibian Violet
Woodhoopoe P. d. damarensis, and the occurrence of
mixed groups, there is no location that could be un-
equivocally assigned to Violet Woodhoopoes. Six of
our samples came from Hobatere, which is within the
westernmost third of the P. d. damarensis distribu-
tion (du Plessis 1997a), with most records of the
species coming from further east. A comparison with
the distribution map for the Green Woodhoopoe (du
Plessis 1997b) shows virtually complete sympatry
between these two taxa. The remaining six samples
came from Omaruru, an appropriate locality as it is
*Corresponding author.
Email: mic@sun.ac.za