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Accepted by P. Rasmussen: 24 Jun. 2010; published: 30 Jul. 2010 65
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2010 · Magnolia Press
Zootaxa 2554: 65–68 (2010)
Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy
ALICE CIBOIS1, NORMAND DAVID2, STEVEN M.S. GREGORY3 & ERIC PASQUET4
1Department of Mammalogy and Ornithology, Natural History Museum, CP 64 34 Geneva 6, Switzerland.
210385A, Clark, Montréal, Québec, Canada, H3L 2S3. E-mail: email@example.com
335, Monarch Road, Northampton, Northamptonshire NN2 6EH, U.K.. E-mail: firstname.lastname@example.org
4Département Systématique et Evolution, Muséum National d’Histoire Naturelle, UMR7205-CNRS, CP 51, 57 rue Cuvier, F-75231
Paris Cedex 05, France. E-mail: email@example.com
The island of Madagascar is a renowned hotspot for adaptive radiations. Madagascar has been separated from mainland
Africa since the end of the Jurassic, and from India since the Late Cretaceous. This long isolation, combined with the
island’s large size and relatively few dispersal events has resulted in an avifauna characterized by a low species count and
high endemism: for instance, 80% of the breeding Malagasy songbirds (Passeriformes) are endemic (Hawkins &
Goodman 2003). A first series of papers (Cibois et al. 1999, 2001; Fjeldsa et al. 1999) on the phylogeny of the Malagasy
taxa traditionally classified as Timaliidae, Sylviidae and Pycnonotidae (all families included in the large sylvioid clade)
showed that several of these passerines form an original radiation endemic to the island. Because these results were
based solely on a single kind of molecular marker (mitochondrial DNA sequences), the authors refrained at that time
from giving a name to this clade. More recently, other studies using nuclear markers as well (Beresford et al. 2005;
Johansson et al. 2008a, 2008b) confirm the existence of this Malagasy sylvioid radiation. The species that comprise this
group exhibit a great variety of bill shapes, wing and tail proportions, and tarsus lengths. This diversity in morphology is
linked to varieties of habitat and prey favoured by these insectivorous forest dwellers (Schulenberg 2003). Thus the
endemic Malagasy sylvioid clade rivals other island radiations, including the vangas of Madagascar and the finches of
the Galapagos, in ecological and morphological diversity. Several authors were inclined to consider this group at the
family level, using the name ‘Bernieridae’. To our knowledge the first study using this name was the book “The natural
history of Madagascar”, edited by S. M. Goodman and J. Benstead in 2003, where the name ‘Bernieridae’ appeared in
two chapters (in Tingle et al. (2003: p. 522) and Hawkins & Goodman (2003: p. 1036), although Schulenberg (2003: p.
1131) referred to the Malagasy "warblers" in his chapter on the radiations of passerine birds on Madagascar). An
alternative spelling for the family-group name, ‘Bernieriidae’, can be found in several personal pages on the internet, but
we have not found an occurrence of this in any publication, as defined in the International Code of Zoological
Nomenclature (4th edition, 1999). The name ‘Bernieridae’ was later used in several journal articles (Chouteau &
Fenosoa 2008; Fuchs et al. 2008; Johansson et al. 2008a, 2008b), however, none of these have introduced the family-
group name ‘Bernieridae’ according to the provisions of the International Code of Zoological Nomenclature, i.e. the
nominal taxon was not explicitly indicated as intentionally new (Article 16.1) and the type genus was not cited (Article
16.2). In the present paper, we therefore propose to rectify this situation by correctly introducing the family-group name
for the Malagasy sylvioid radiation.
According to Cibois et al. (2001) and Johansson et al. (2008b), the 10 species that comprise this group are as follows
(another endemic passerine, not yet studied, Randia pseudozosterops Delacour & Berlioz, 1931, could be a possible
addition to this group):
Bernieria madagascariensis (Gmelin, 1789)
Muscicapa madagascariensis Gmelin J.F. 1789, Systema Naturae (ed. 13). Vol. 1, part 2: 940.
Xanthomixis zosterops (Sharpe, 1875)
Bernieria zosterops Sharpe 1875, Proc. zool. Soc. Lond. (1875): 76.
CIBOIS ET AL.66 · Zootaxa 2554 © 2010 Magnolia Press
Xanthomixis cinereiceps (Sharpe, 1881)
Oxylabes cinereiceps Sharpe 1881, Proc. zool. Soc. Lond. (1881): 197.
Xanthomixis apperti (Colston, 1972)
Phyllastrephus apperti Colston 1972, Ibis 114: 89.
Xanthomixis tenebrosa (Streseman, 1925)
Phyllastrephus tenebrosus Streseman 1925, Orn. Monatsb. 33:150.
Oxylabes madagascariensis (Gmelin, 1789)
Motacilla madagascariensis Gmelin J.F. 1789, Systema Naturae (ed. 13). Vol. 1, part 2: 952.
Thamnornis chloropetoides (Grandidier, 1867)
Ellisia chloropetoides Grandidier A. 1867, Rev. Mag. Zool. (2) 19: 256.
Crossleyia xanthophrys (Sharpe, 1875)
Oxylabes xanthophrys Sharpe 1875, Proc. zool. Soc. Lond. (1875): 76.
Hartertula flavoviridis (Hartert, 1924)
Neomixis flavoviridis Hartert 1924, Bull. Brit. Orn. Club 45: 35.
Cryptosylvicola randrianasoloi Goodman, Langrand & Whitney, 1996
Cryptosylvicola randrianasoloi Goodman, Langrand & Whitney 1996 Ibis 138: 154.
The first five species on this list were often placed in the same genus: Phyllastrephus Swainson, 1831. However,
they do not form a monophyletic group, and consequently Cibois et al. (2001) suggested that madagascariensis be
restored to the genus Bernieria Pucheran 1855, and that the remaining species apperti, cinereiceps, zosterops and
tenebrosa (the latter studied by Johansson et al. 2008b), be restored to the genus Xanthomixis Sharpe, 1881. Cibois et al.
(1999) proposed that the genus Hartertula Streseman, 1925 should be used for the species flavoviridis, as this species
belongs to a different clade from the three other Neomixis species; Nguembock et al. (2007) showed that Neomixis is
embedded in the Cisticolidae. The last endemic Malagasy “warbler”, Nesillas typica (Hartlaub, 1860), is not a member of
this endemic radiation and in fact belongs to the Acrocephalidae (Johansson et al. 2008b).
Thus the seven genera that comprise the Malagasy sylvioid radiation are: Bernieria Pucheran, 1855; Oxylabes
Sharpe, 1870; Crossleyia Hartlaub, 1877; Xanthomixis Sharpe, 1881; Thamnornis Milne-Edwards & Grandidier, 1882;
Hartertula Streseman, 1925; Cryptosylvicola Goodman, Langrand & Whitney, 1996. The genus Bernieria would appear
to be the oldest name, but the attribution of the genus is open to interpretation. Indeed the first mention of the name
Bernieria (Bonaparte, 1854) is a nomen nudum: the name is mentioned on page 11: "Bernieria, Bp. (*)", and at the foot
of p. 10: "Deux espèces du Muséum: Bernieria major et Bernieria minor, Bp., Madagascar". However these two species
were not described and Pucheran (1855, page 369) was probably the first to introduce the genus. He provided a
description of the type specimen of Ramphocoenus viridis (= Muscicapa madagascariensis Gmelin, 1789), collected by
Delalande in 1820. He associated this description with the following footnote: “M Charles Bonaparte a fait plus
récemment de cette espèce le type de son genre Bernieria (Comptes rendus, vol. 38, p. 40): c’est pour lui Bernieria
major” (More recently, M Charles Bonaparte made this species, that he named major, the type of his genus Bernieria).
Hartlaub (1860) later presented the description of both Bonaparte’s species, major and minor, based on the same three
specimens held in the Muséum National d’Histoire Naturelle (Paris): the type specimen of Ramphocoenus viridis, which
was first described by Pucheran, and the two syntypes of Bernieria minor, one collected by Sganzin in 1832 (as indicated
on the specimen label, contrary to Hartlaub who mentioned the year 1834), the other by the French surgeon Chevalier
J.A. Bernier in 1834. Sexual dimorphism in size is striking in Bernieria madagascariensis (Fishpool & Tobias 2005): the
two minor specimens correspond to females (as Sharpe (1883) pointed out), whereas the specimen described as major
corresponds to a male.
It could be argued that the mention of Berniera by Pucheran was not adequate: for instance Bonaparte (1854) listed
two names (major, minor) under Bernieria and Pucheran merely cited the first (major). The fact, however, that Bernieria
was previously a nomen nudum does not preclude its later availability for the same or a different concept, and would take
authorship and date from that act of establishment, and not from any earlier publication as a nomen nudum. Therefore
Zootaxa 2554 © 2010 Magnolia Press · 67
BERNIERIDAE (AVES: PASSERIFORMES)
Pucheran’s use of Bernieria, linking the species Ramphocoenus viridis Lesson, 1830, with the former nomen nudum
Bernieria is not contradictory to the Requirements for names published before 1931: he provided a description of the
taxon (Article 12.1) as well as an indication (i.e. the footnote) of the new name (Article 12.2.5). Additionally, Bernieria
Pucheran, 1855 was the accepted citation in Schulze et al. (1926) and Neave (1939) and we choose here to follow this
Although it was not explicitly indicated by Bonaparte or Pucheran, the name Bernieria is clearly derived from
Bernier, the name of one of the collectors. It is not a Greek or Latin word, and under Article 29.3.3, we choose the stem
“Bernier” and keep the most often used spelling for the Malagasy sylvioid clade when treated at the family level:
Bernieridae, fam. nov.
Type genus: Bernieria Pucheran, 1855 [= Bonaparte n.n.]
Type species: Ramphocoenus viridis Lesson, 1830 = Muscicapa madagascariensis Gmelin, 1789 by monotypy.
Description: a group of insectivorous passerine birds (Aves, Passeriformes) endemic to Madagascar, which share a
common ancestor and form an adaptive radiation within the island. The species that comprise this group exhibit a great
variety of bill shapes, wing and tail proportions, and tarsus lengths. Yellow and green are the most common colorations,
with few plumage patterns (e.g. white throat for Oxylabes madagascariensis and Xanthomixis apperti).This group has
been diagnosed as monophyletic based only on genetic results.
We are grateful to Walter Bock, Edward Dickinson, Robert Dowsett, and Roger Safford for productive and helpful
discussion, as well as for much of the bibliographic information presented in this paper.
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Bonaparte, C.-L. (1854) Notes sur les collections rapportées en 1853, par M. A. Delattre, de son voyage en Californie et
dans le Nicaragua. Comptes rendus de l'Académie des Sciences, Paris, 38, 1–11.
Chouteau, P. & Fenosoa, R. (2008) Seasonal effects on foraging behaviour of two sympatric species of couas in the
western dry forest of Madagascar. African Journal of Ecology, 46, 248–257.
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Timaliidae) and warblers (Passeriformes: Sylviidae), based on cytochrome b and 16S rRNA sequences. Molecular
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CIBOIS ET AL.68 · Zootaxa 2554 © 2010 Magnolia Press
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Nguembock, B., Fjeldsa, J., Tillier, A. & Pasquet, E. (2007) A phylogeny for the Cisticolidae (Aves: Passeriformes)
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