PHYLOGENETIC RELATIONSHIPS OF THE MADAGASCAR PYGMY KINGFISHER (ISPIDINA MADAGASCARIENSIS)

Auk, The, Oct 2005 by Marks, Ben D, Willard, David E

ABSTRACT.-

The avifauna of Madagascar presents a complicated taxonomic and biogeographic problem. Although Madagascar was once connected to Africa, the birds of the island are not all of African origin. The Madagascar Pygmy Kingfisher (Ispidina madagascariensis) is sometimes placed in the African genus Ispidina and sometimes in the Southeast Asian genus Ceyx. We sequenced 755 base pairs of mitochondrial DNA from a fragment of the NADH dehydrogenase subunit II (ND2) and from the complete NADH dehydrogenase subunit III (ND3). We used these data to construct phylogenetic hypotheses for the Alcedinidae. Using these phylogenies, we evaluate previous hypotheses of relationships of I. madagascariensis. Although we cannot reject the hypothesis that I. madagascariensis is indeed a member of Ispidina, tree-building analyses support a relationship not with Ispidina or Ceyx, but instead with members of the African genus Corythornis. These data suggest that certain behaviors and plumage types have evolved several times in the Alcedinidae. Received 22 March 2004, accepted 27 April 2005.

Key words: Alcedinidae, Ceyx, Ispidina, Madagascar, phylogeny.

Relations Phylogénétiques de Ispidina madagascariensis

RÉSUMÉ. -L'avifaune malgache présente un problème taxonomique et biogéographique assez compliqué. Bien qu'il soit reconnu que Madagascar était jadis connecté au reste du continent africain, il est aussi clair que tous les oiseaux de l'île n'ont pas une origine africaine. Ispidina madagascariensis est tantôt placé dans le genre africain Ispidina, tantôt dans le genre asiatique Ceyx. Nous avons réalisé le séquençage de 755 paires de bases d'ADN mitochondrial provenant d'un fragment de la sous-unité II de la déshydrogénase NADH (ND2) et la totalité de la sous-unité III de la déshydrogénase NADH (ND3). Ces données ont été utilisées pour reconstruire les hypothèses phylogénétiques des Alcedinidae. Nous avons évalué les hypothèses précédentes à propos des affinités du martin-pêcheur Ispidina madagascariensis en utilisant ces phylogénies. Bien que nous ne soyons pas en mesure de rejeter l'hypothèse selon laquelle Ispidina madagascariensis est en réalité membre du genre Ispidina, l'analyse des arbres phylogénétiques appuie l'existence d'une affinité non pas avec Ispidina ou Ceyx, mais plutôt avec des membres d'un autre genre africain Corythornis. Ces données suggèrent que certains comportements et types de plumage ont évolué plusieurs fois chez les Alcedinidae.

THE ISLAND OF Madagascar is well known for the high degree of endemism of its flora and fauna. Madagascar split off from Africa ~165 mya and from the Indian subcontinent ~88 mya (Krause 2003). It now sits ~400 km off of the coast of Africa and ~4,000 km from India (Krause 2003). Although the island is close to Africa, the prevailing winds travel from east to west (Donque 1972), a counter against "easy" dispersal of birds from the mainland to Madagascar. Because of its current proximity to Africa and its tectonic history, Madagascar is an especially interesting case study for the biogeographic origins of its biota.

It has long been recognized that the Malagasy avifauna has relationships to the faunas of both Africa and Asia (Keith 1980). Malagasy birds span a spectrum, ranging from members of ancient endemic families (e.g. Cibois et al. 2001, Kirchman et al. 2001) whose closest relatives are yet to be determined, to species that move between Africa and Madagascar for breeding and wintering. Most nonendemic species of Malagasy birds are shared with Africa but not with Asia (e.g. Marsh Owl [Asia capensis], Namaqua Dove [Oena capensis], Madagascar Bee-eater [Merops superciliosus], and Wattled Starling [Creatophora cinerea]). Some endemic species, such as the Madagascar Malachite Kingfisher (Alcedo vintsioides), often treated as a subspecies of the African Malachite Kingfisher (A. cristata), also have clear connections to Africa. However, other species, such as the Madagascar Bulbul (Hypsipetes madagascariensis) and Madagascar Magpie-Robin (Copsychus albospecularis), are included in genera with Asian ranges. A full understanding of the biogeographic relationships of the Malagasy avifauna has been hampered by a lack of knowledge regarding the phylogenetic relationships of these species. For many genera that are shared among Africa, Asia, and Madagascar, no published studies have yet determined the closest relatives of the Malagasy form.

The Madagascar Pygmy Kingfisher (lspidina madagascariensis) is an example of a Madagascar endemic with uncertain affinities, and it has been treated taxonomically in a variety of ways over the past 50 years (Table 1). Delacour (1951) and Fry (1980) included it in the largely Asian genus Ceyx, along with the African Pygmy (I. picta) and Dwarf (Myioceyx lecontei) kingfishers, largely on the basis of bill shape and color. Traylor (1960) considered these characters convergent and placed I. madagascariensis in a broad African genus Corythornis, which included species otherwise treated in Alcedo, lspidina, and Myioceyx. Sibley and Monroe (1990) and Peters (1945) connected it to the small insectivorous African Pygmy Kingfisher (picta) in the genus lspidina, whereas Wolters (1976) placed it in a monotypic genus, Ceycoides.