Characterization and Evolution of the Mitochondrial DNA Control Region in
Hornbills (Bucerotiformes)
Wayne Delport,
1,2
J. Willem H. Ferguson,
1
Paulette Bloomer
2
1
Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa
2
Molecular Ecology and Evolution Program, Department of Genetics, University of Pretoria, Pretoria 0002, South Africa
Received: 3 April 2001 / Accepted: 7 December 2001
Abstract. We determined the mitochondrial DNA
control region sequences of six Bucerotiformes. Horn-
bills have the typical avian gene order and their control
region is similar to other avian control regions in that it
is partitioned into three domains: two variable domains
that flank a central conserved domain. Two characteris-
tics of the hornbill control region sequence differ from
that of other birds. First, domain I is AT rich as opposed
to AC rich, and second, the control region is approxi-
mately 500 bp longer than that of other birds. Both these
deviations from typical avian control region sequence are
explainable on the basis of repeat motifs in domain I of
the hornbill control region. The repeat motifs probably
originated from a duplication of CSB-1 as has been de-
termined in chicken, quail, and snowgoose. Furthermore,
the hornbill repeat motifs probably arose before the di-
vergence of hornbills from each other but after the di-
vergence of hornbills from other avian taxa. The mito-
chondrial control region of hornbills is suitable for both
phylogenetic and population studies, with domains I and
II probably more suited to population and phylogenetic
analyses, respectively.
Key words: Mitochondrial DNA — Control region —
Bucerotiformes — Characterization — Repeats — Sec-
ondary structure — Phylogenetics — Population genet-
ics
Introduction
The mitochondrial DNA genome has proven to be useful
in many avian phylogenetic and population studies (re-
viewed by Baker and Marshall 1997; Mindell et al. 1997;
Moore and DeFilippis 1997). Advantages of working
with mtDNA include maternal inheritance (Lansman et
al. 1983), a higher rate of evolution than in single-copy
nuclear DNA (Brown et al. 1979), and variable evolu-
tionary rates of genes within the genome itself (Aquadro
and Greenburg 1983; Cann et al. 1984). Therefore it is
possible to address both phylogenetic and population-
level questions using different genes of the same genome
(Wenink et al. 1994). Traditionally coding genes of the
mitochondrial genome have been reserved for phyloge-
netic studies (Moore and DeFilippis 1997), whereas the
control region has been considered more suitable for
population level studies (Baker and Marshall 1997). The
control region evolves three to five times more rapidly
than the remainder of the mitochondrial genome
(Aquadro and Greenburg 1983; Cann et al. 1984) and
therefore can be difficult to amplify with universal ver-
tebrate primers in a polymerase chain reaction (PCR). In
addition, both nuclear integration (Sorenson and
Fleischer 1996; Zhang and Hewitt 1996), although not
restricted to the control region, and repeat motifs (Des-
jardins and Morais 1990, 1991; Quinn and Wilson 1993;
Ramirez et al. 1993; Randi and Lucchini 1998; Wenink
et al. 1994) could render PCR amplification difficult.
Part of the control region is thought to regulate tran-
scription and replication of the vertebrate mtDNA ge-
nome (Brown et al. 1986; Clayton 1991). These areas
that may have function associated-secondary structure
Correspondence to: Wayne Delport, Molecular Ecology and Evolution
Program; email: wdelport@postino.up.ac.za
J Mol Evol (2002) 54:794–806
DOI: 10.1007/s00239-001-0083-0
© Springer-Verlag New York Inc. 2002