THE HIPPOBOSCIDAE (INSECTA: DIPTERA) FROM MADAGASCAR, WITH NEW RECORDS FROM THE “PARC NATIONAL DE MIDONGY BEFOTAKA” RAHOLA N.*, GOODMAN S.M.** & ROBERT V.* Summary: Résumé : LES HIPPOBOSCIDAE (INSECTA: DIPTERA) DE MADAGASCAR, AVEC DE NOUVELLES OBSERVATIONS DANS LE PARC NATIONAL DE The Hippoboscidae or “louse-flies” is a family of pupiparous MIDONGY BEFOTAKA Diptera, which in their adult stage are ectoparasites of mammals and birds. This paper presents a comprehensive review of Malagasy Les Hippoboscidae constituent une famille de Diptères pupipares Hippoboscidae. In total, amongst the 213 species of this family dont le stade adulte est ectoparasite de mammifères et d’oiseaux. known worldwide, 14 have been reported in Madagascar, among Nous présentons ici une revue de tous les Hippoboscidae which six are considered as endemic to the Malagasy region. In connus de Madagascar. Parmi les 213 espèces de cette famille addition, data are presented from a collection of 17 Hippoboscidae connues dans le monde, 14 ont été observées au moins une obtained from seven species of forest-dwelling birds in the “Parc fois à Madagascar, parmi lesquelles six sont considérées comme National de Midongy Befotaka”, southeastern Madagascar, in endémiques de la région malgache. Nous présentons aussi la 2003. The flies in this collection belong to three different species: récolte de 17 Hippoboscidae adultes collectés de sept espèces Icosta malagasii (one), Ornithoica podicipis (ten) and Ornithoctona d’oiseaux strictement forestiers en 2003 dans le Parc National de laticornis (six). The two former species were previously only Midongy Befotaka, sud-est de Madagascar. Ces 17 hippobosques known from single specimens in museum collections; the later se répartissent en trois espèces : Icosta malagasii (un), Ornithoica species is distributed across much of the Afrotropical region podicipis (dix) et Ornithoctona laticornis (six). Les deux premières and the records presented herein are the first for Madagascar. de ces espèces étaient précédemment connues par un unique All the seven bird species are new hosts for hippoboscids. We spécimen enregistré dans les collections des muséums ; O. present the first description of the male of Icosta malagasii. An laticornis a une large répartition en zone Afrotropicale, mais illustrated dichotomous determination key of the 14 Malagasy il est trouvé pour la première fois à Madagascar. Chacune species, based on morphological criteria only, is presented. des sept espèces d’oiseaux constitue un nouvel hôte pour les hippobosques collectés. Nous proposons la première description KEY WORDS: Hippoboscidae, birds, review, determination key, host- du mâle d’Icosta malagasii. Nous présentons enfin une clé parasite relationship, taxonomy, Madagascar, Midongy Befotaka. dichotomique illustrée, basée exclusivement sur des critères morphologiques, pour la détermination des 14 espèces malgaches. MOTS-CLÉS : Hippoboscidae, oiseaux, revue, clé de détermination, relations hôte-parasite, taxonomie, Madagascar, Midongy Befotaka. INTRODUCTION larval development takes place within the mother’s abdomen and pupation occurs almost immediately he members of the family Hippoboscidae, after pre-pupae laying (Hutson, 1984). The Hippo- known as “louse flies” or “keds”, are obligate boscidae belongs to the super-family of Hippobos- ectoparasites of mammals and birds. Members coidea, which groups four families of flies that are T bloodsucking at the adult stage for both sexes. The of this Diptera family consist of winged species, some of which have considerable flight capacity, while four families consist of Hippoboscidae, the bat flies other species are flightless and highly apomorphic, Nycteribiidae and Streblidae, and the tsetse flies Glos- having vestigial or no wings. Hippoboscidae belong sinidae. Hippoboscoidea are considered monophyletic, to Pupipara “pupa-bearers”, in reference to the fact but based on current data, the basal relationships that, unlike virtually all other insects, most of the between the four families are not resolved (Dittmar et al., 2006). A recent checklist of Hippoboscidae across the world retains three subfamilies (Ornithomyinae, * MIVEGEC (Maladies Infectieuses et Vecteurs : Écologie, Génétique, Hippoboscinae, and Lipopteninae), 21 genera, and Évolution et Contrôle), IRD 224, CNRS 5290, Université Montpellier 213 species (Dick, 2006). It has been shown that the 1, Université Montpellier 2, Montpellier, France. ** Field Museum of Natural History, Chicago, Illinois, United States two subfamilies Hippoboscinae and Lipopteninae are & Vahatra, Antananarivo, Madagascar. monophyletic groups (Petersen et al., 2007). Correspondence: Vincent Robert, MIVEGEC, Centre IRD, 911, avenue Caution must be taken with the name Hippoboscidae. Agropolis, BP 64501, 34394 Montpellier Cedex 5, France. Tel.: 33 (0)4 67 41 61 27 – Fax: 33 (0)4 67 41 63 30. In old taxonomic treatments, it was used to encom- E-mail: [email protected] pass the Hippoboscidae as defined today, as well as Parasite, 2011, 18, 127-140 Original contribution 127 RAHOLA N., GOODMAN S.M. & ROBERT V. the bat-flies (Nycteribiidae and Streblidae). Irwin et Hippoboscinae, and Lipopteninae – are 11, two, and al. (2003), following the BDWD (BioSystematic Data- zero, respectively. Six species of Ornithomyinae are base of World Diptera), used this old definition and presently considered as endemic to Madagascar. Indi- indicated 760 valid species in the world and 15 in viduals of eight species have been collected from a Madagascar, among which seven are Hippoboscidae single individual host, including three different specific sensu stricto. names proposed for the female flies (Icosta malagasii, Members of the Hippoboscidae are known to act as Ornithomya sorbens, and Ornithoctona idonea), while vectors of many infectious agents: protozoan, bac- the identity of the male remains unknown. teria, helminthes, and perhaps viruses. They certainly The genus Allobosca is composed by only one known transmit mammal Trypanosomatidae of the genus species, A. crassipes. This species, considered endemic Megatrypanum (Baker, 1967; Oyieke & Reid, 2003) to Madagascar, has rudimentary wings and is known and probably transmit avian trypanosomes (Kucera, from several species of lemurs (Ferris & Cole, 1922; 1983). Hippoboscidae are the only known vectors of Maa, 1969c). Proparabosca is a genus of Hippobos- Haemoproteus (an apicomplexan parasite of birds). cidae, with only one known species, P. alata, which is Melophagus ovinus louse flies play a role in the an ectoparasite of the Indriidae Propithecus verreauxi transmission of Bartonella among ruminants (Halos et in Namoroka National Park (Theodor & Oldroyd, al., 2004; Reeves et al., 2006). In Kenya, the fly Hip- 1965). Most of the 171 species of Ornithomyinae are pobosca longipennis is thought to transmit to hyenas bird ectoparasites, with the exceptions of A. crassipes and domestic dogs the larva of the filarial nematoda and P. alata (plus five species that parasite wallabies Acanthocheilonema dracunculoides (Nelson, 1963). in Australia). This filaria has been reported from domestic animals Icosta malagasii is only known by one female. It in Madagascar (Daynes, 1964). The louse fly Icosta was collected as a “parasite sur l’aile d’un Coezach/ americana is currently suspected in the transmission Sahafanjana/N: 115/R. M./ Inst. Sci. Madagascar” (Maa, of West Nile Virus in North America (Farajollahi et 1969b). Sahafanjana is located in close proximity to al., 2005). the “Parc National de Mananara-Nord” and in the zone Although the importance of Hippoboscidae in the between Manambato and Anove (Quentin & Villiers, transmission of mammal and bird parasites in Mada- 1973; Flint et al., 1987). One possibility in the identi- gascar has been suspected and to some extent docu- fication of the “Coezach” is that it is a poor translite- mented (Daynes, 1964; Savage et al., 2009), members ration of the Malagasy vernacular name “Koa”, which of this family remain poorly documented on the island. are members of the endemic genus Coua (subfamily We here present the first comprehensive review of Couinae). In certain Malagasy dialects, the last syl- Malagasy Hippoboscidae (sensu stricto i.e. excluding lable of the vernacular name is accented and harshly Nycteribiidae and Streblidae). In addition, we pro- pronounced. In the Parc Botanique et Zoologique de pose the first illustrated dichotomous determination Tsimbazaza, Antananarivo (previously known as “Ins- key for the genera and species currently known from titut de Recherche Scientifique de Madagascar”) there Madagascar. Recent specimens of hippoboscids have is a male specimen of C. caerulea, collected on 12 been collected from Malagasy vertebrates and remain April 1955 by “R.M.” at Sahafanjana. This information unstudied by fly specialists (e.g., Wright et al., 2009). gives credence to the identity of the “Coezach” as a Herein we present details on a hippoboscid collec- Coua. tion obtained from forest-dwelling birds in the “Parc National de Midongy Befotaka”, previously known as Ornithoctona idonea is only known by one male and Midongy-Sud, southeastern Madagascar. two females with two of these specimens, including the male holotype, being from Madagascar (Falcoz, 1929). Theses two Malagasy specimens formerly in REVIEW OF MALAGASY HIPPOBOSCIDAE the Falcoz collection are now untraceable. Curiously, although the male was designated as the holotype, BASED ON PUBLISHED LITERATURE it remains undescribed. Presently, the only