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Bernieridae (Aves: Passeriformes): A Family-Group Name For The Malagasy Sylvioid Radiation

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Cibois, Alice, David, Normand, Gregory, Steven M. S., Pasquet, Eric (2010): Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy sylvioid radiation. Zootaxa 2554: 65-68
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: 6568 (2010)
Bernieridae (Aves: Passeriformes): a family-group name for the Malagasy
sylvioid radiation
1Department of Mammalogy and Ornithology, Natural History Museum, CP 64 34 Geneva 6, Switzerland.
210385A, Clark, Montréal, Québec, Canada, H3L 2S3. E-mail:
335, Monarch Road, Northampton, Northamptonshire NN2 6EH, U.K.. E-mail:
4Département Systématique et Evolution, Muséum National d’Histoire Naturelle, UMR7205-CNRS, CP 51, 57 rue Cuvier, F-75231
Paris Cedex 05, France. E-mail:
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
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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molecular systematics of several evolutionary "enigmas". Proceedings of the Royal Society of London B, 272, 849–
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.
Cibois, A., Pasquet, E. & Schulenberg, T.S. (1999) Molecular systematics of the Malagasy babblers (Passeriformes:
Timaliidae) and warblers (Passeriformes: Sylviidae), based on cytochrome b and 16S rRNA sequences. Molecular
Phylogenetics and Evolution, 13, 581–595.
Cibois, A., Slikas, B., Schulenberg, T.S. & Pasquet, E. (2001) An endemic radiation of Malagasy songbirds is revealed
by mitochondrial DNA sequence data. Evolution, 55, 1198–1206.
Fishpool, L. & Tobias, J. (2005) Family Pycnonotidae (Bulbuls). In: del Hoyo, J., Elliot, A. & Christie D. (Eds),
Handbook of the Birds of the World. Volume 10: Cuckoo-Shrikes to Thrushes. Lynx Edicions, Barcelona, pp. 124–
Fjeldsa, J., Goodman, S.M., Schulenberg, T.S. & Slikas, B. (1999) Molecular evidence for relationships of Malagasy
birds. In: Adams, N. J. & Slotow, R. H. (Eds), Proceedings of the 22nd International Ornithological Congress,
Durban. BirdLife South Africa, Johannesburg, pp. 3084–3094.
Fuchs, J., Pons, J.-M., Goodman, S.J., Bretagnolle, V., Melo, M., Bowie, R.C.K., Currie, D., Safford, R., Virani, M.Z.,
Thomsett, S., Hija, A., Cruaud, C. & Pasquet, E. (2008) Tracing the colonization history of the Indian Ocean scops-
owls (Strigiformes: Otus) with further insight into the spatial-temporal origin of the Malagasy avifauna. BMC
Biology, 8, 197.
Hartlaub, G. (1860) Systematische Uebersicht der Vögel Madagascars. Journal für Ornithologie, 44, 6–112.
Hawkins, A.F.A. & Goodman, S.J. (2003) Introduction to the birds. In: Goodman, S. M. & Benstead, J. (Eds), The
Natural History of Madagascar. The University of Chicago Press, Chicago, pp. 267–281.
International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth
edition. International Trust for Zoological Nomenclature c/o The Natural History Museum, London, 306 pp.
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babbler (Mystacornis crossleyi): adding a new niche to the vanga radiation of Madagascar. Biology Letters, 4, 677–
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A review and a new molecular phylogeny based on three nuclear intron markers. Molecular Phylogenetics and
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Neave, S.A. (1939) Nomenclator Zoologicus. A List of the Names of Genera and Subgenera in Zoology from the Tenth
Edition of Linnaeus 1758 to the End of 1935. Zoological Society of London, London, xiv + 957 pp.
Nguembock, B., Fjeldsa, J., Tillier, A. & Pasquet, E. (2007) A phylogeny for the Cisticolidae (Aves: Passeriformes)
based on nuclear and mitochondrial DNA sequence data, and a re-interpretation of an unique nest-building
specialization. Molecular Phylogenetics and Evolution, 42, 272–286.
Pucheran, J. (1855) Mémoires sur les types peu connus de Passereaux dentirostres de la collection du musée de Paris.
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... U tradicionalnoj taksonomiji ova skupina se tretira unutar porodice Pycnonotidae Gray, 1840 (Bernieria i Xanthomixis), Sylviidae (Thamnornis i Cryptosylvicola) i Timaliidae (ostali rodovi), ali prema molekularno-biološkim podacima data skupina spada u zasebnu porodicu (Beresford et al., 2005;Cibois et al., 2010). ...
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U višegodišnjim zoološkim istraživanjima Bosne i Hercegovine autori, a posebno kustos ornitolog Dražen Kotrošan, su sakupili i značajan broj podataka o biodiverzitetu ptica. Spoznate činjenice, uz konstatacije o brojnim taksonomskim i nomenklaturnim promjenama date skupine u nizu publiciranih radova dovele su i do specifične obaveze zvanične aktuelizacije spomenutih podataka, osobito nakon zaključka Vijeća Odsjeka za biologiju Prirodno-matematičkog fakulteta Univerziteta u Sarajevu da prof. dr. Suvad Lelo bude odgovorni nastavnik na nastavnom predmetu (Bio)Sistematika hordata od akademske 2012/2013. godine (na prijedlog dotadašnjeg odgovornog nastavnika prof. dr. Rifat Škrijelj na čemu mu se i ovom prilikom javno zahvaljujemo). Kao zaduženi nastavnik za interpretaciju aktuelne biosistematike ptica dr. Lelo je oformio ekipu, autore ove knjige, koja je priredila rukopis (skriptu predavanja iz ornitologije) da bi obezbijedili studenskoj populaciji adekvatan prijegled podataka o bosanskohercegovačkoj i svjetskoj ornitofauni. Neophodno je istaći da je ovo prva ornitološka knjiga pripremljena u cilju edukacije studenata biologije od osnivanja Prirodno-matematičkog fakulteta Univerziteta u Sarajevu, a koja je proistekla od bosanskohercegovačkih autora. Do sada, skoro pola vijeka, generacije studenata biologije na Prirodno-matematičkom fakultetu u Sarajevu kao osnovnu literaturu iz oblasti sistematike kičmenjaka koristile knjigu prof. dr. Aleksandra Aleksopula „Zoologija kičmenjaka: morfologija, evolucija, ekologija, privredni značaj i sistematika kičmenjaka“ objavljene 1963. godine u Beogradu. Na kraju, ovu knjigu posvećujemo najznačajnijem poznavaocu bosanskohercegovačke ornitofaune, negdašnjem kustosu Zemaljskog muzeja Bosne i Hercegovine u Sarajevu – Otmaru Reizeru (O. Reiser: 1861.-1936.). O. Rajzer je kao kustos ornitolog radio od 1887. do 1914. godine pri čemu je utemeljio bosanskohercegovačku ornitologiju.
... This paper is the first of a series of works on the systematics, diversity and host associations of feather mites associated with Malagasy warblers (Passeriformes: Bernieridae), a group endemic to Madagascar. Malagasy warblers form a recently recognized passerine family (Cibois et al. 2010), established with the realization that many Malagasy species, placed previously in Sylviidae, Pycnonotidae, and Timaliidae, actually form a monophyletic lineage (Cibois et al. 1999(Cibois et al. , 2001. Most phylogenies place Bernieridae close to grassbirds, Locustellidae, a group of sylvioid birds widely distributed in Africa, Asia and Australia, or Donacobiidae with a single species from South America (Alström et al. 2013;Moyle et al. 2016). ...
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A new feather mite genus Bernierinyssus gen. n. (Analgoidea: Pteronyssidae), associated with endemic Malagasy warblers (Passeriformes: Bernieridae), is proposed based on morphological evidence and DNA sequence data. Within this genus, we detected six mite species, including five new species described here: Bernierinyssus angulatus sp. n. from Crossleyia xanthophrys, B. bernieriae sp. n. from Bernieria madagascariensis, B. bifenestratus sp. n. from Hartertula flavoviridis, B. randiae sp. n. from Randia pseudozosterops, B. xanthomixis sp. n. from Xanthomixis zosterops (type host) and X. cinereiceps, and B. oxylabis (Mironov and Wauthy 2005) comb. n. (transferred from Pteronyssoides Hull). Phylogenetic relationships of these mites were nearly perfectly congruent with those of their hosts, indicating that ancestral Bernierinyssus probably co-dispersed to Madagascar on the common ancestor of Malagasy warblers and then cospeciated with their hosts. Species of Bernierinyssus are well-delimited based on several lines of evidence: morphology (clear among-specific differences in discrete characters), host associations (one mite species per one host species, except for B. xanthomixis), genetic distances (large COX1 barcoding gap between among-and within-species K2P distances: 8.22-12.38% vs 0-2.9%, respectively), and molecular phylogenetics (all species are well-supported, monophyletic clades). Our study suggests that species of the genus Bernierinyssus have evolved slower than their avian hosts or co-associated feather lice. Despite the discordance in the mitochondrial DNA evolutionary rates, speciation events in mites largely corresponded to bird species divergences, resulting in a nearly perfect correlation between mite and bird species richness (Eichler's Rule). The mite B. xanthomixis was associated with two avian species, but still formed two distinct shallow lineages (COX1 distance: 1.65%) separated by the host species. The nearly strict host-specificity pattern found in Bernierinyssus contrasts with that of continental feather mites, which tend to be less host-specific and have nearly equal proportions of single-host vs multi-host species.
... This monophyletic assemblage consists of at least 22 species, many of which were originally placed in other families, and is nearly as morphologically diverse as all passerines combined (Jønsson et al., 2012;Reddy et al., 2012;Yamagishi et al., 2001). Furthermore, a group of 11 species previously placed in three different families was found to represent another monophyletic avian assemblage endemic to Madagascar (Cibois et al., 2001), and now comprises the recently described family Bernieridae (Cibois et al., 2010). In light of recent phylogenetic discoveries and the ongoing deforestation of Madagascar (Harper et al., 2007), phylogenetic analysis of Madagascar's unstudied avifauna, particularly forest-dependent Passeriformes, is of critical importance. ...
... Most of the birds formerly known as Malagasy greenbuls, previously placed within the Pycnonotidae, as well as several taxa previously considered amongst the sylviids or timaliids, have been recently recognized as an endemic lineage, the family Bernieridae[27]. The study wasTable). ...
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Background to the work For centuries taxonomy has relied on dead animal specimens, a practice that persists today despite the emergence of innovative biodiversity assessment methods. Taxonomists and conservationists are engaged in vigorous discussions over the necessity of killing animals for specimen sampling, but quantitative data on taxonomic trends and specimen sampling over time, which could inform these debates, are lacking. Methods We interrogated a long-term research database documenting 2,723 land vertebrate and 419 invertebrate taxa from Madagascar, and their associated specimens conserved in the major natural history museums. We further compared specimen collection and species description rates for the birds, mammals and scorpions over the last two centuries, to identify trends and links to taxon descriptions. Results We located 15,364 specimens documenting endemic mammals and 11,666 specimens documenting endemic birds collected between 1820 and 2010. Most specimens were collected at the time of the Mission Zoologique Franco-Anglo-Américaine (MZFAA) in the 1930s and during the last two decades, with major differences according to the groups considered. The small mammal and bat collections date primarily from recent years, and are paralleled by the description of new species. Lemur specimens were collected during the MZFAA but the descriptions of new taxa are recent, with the type series limited to non-killed specimens. Bird specimens, particularly of non-passerines, are mainly from the time of the MZFAA. The passerines have also been intensely collected during the last two decades; the new material has been used to solve the phylogeny of the groups and only two new endemic taxa of passerine birds have been described over the last two decades. Conclusions Our data show that specimen collection has been critical for advancing our understanding of the taxonomy of Madagascar’s biodiversity at the onset of zoological work in Madagascar, but less so in recent decades. It is crucial to look for alternatives to avoid killing animals in the name of documenting life, and encourage all efforts to share the information attached to historical and recent collections held in natural history museums. In times of conservation crisis and the advancement in digital technologies and open source sharing, it seems obsolete to kill animals in well-known taxonomic groups for the sake of enriching natural history collections around the world.
... Molecular-based systematic approaches have revealed that the Sylviidae, as defined in this "traditional" classification, is best broken into several families whose composition remains in a state of flux (e.g. Sibley and Ahlquist 1990, Sefc et al. 2003, Alström et al. 2006, Nguembock et al. 2007, 2008, 2012, Johansson et al. 2008, Cibois et al. 2010, Alström et al. 2011. One such family that has come to be defined with the aid of molecular characters is the Cisticolidae, a large radiation of primarily African warblers (Ryan 2006. ...
The Long-billed Tailorbird (Artisornis moreaui), one of Africa's rarest birds, has a strikingly disjunct distribution, the origin of which has long puzzled biogeographers. One small population (subspecies moreaui) occurs in montane forest in the East Usambara Mountains, a montane sky island near the coast of northern Tanzania, and another (subspecies sousae) on Serra Jeci in northwestern Mozambique, 950 km away. At both sites, the Long-billed Tailorbird co-occurs with its putative sister-species the African Tailorbird (Artisornis metopias). At Serra Jeci, Long-billed Tailorbirds are observed primarily in the mid-canopy and African Tailorbirds in the understorey. This situation contrasts with that in East Usambara Mountains, where both species make extensive use of the understorey (<5 m). The apparent difference in canopy strata occupancy of Long-billed Tailorbird between the East Usambara and Serra Jeci formed the basis for Stuart's (1981) hypothesis that African Tailorbirds have forced Long-billed Tailorbirds out of the understorey at Serra Jeci through competitive exclusion. We sought to 1) determine that the two species of African Tailorbird are indeed sister-species, and 2) test Stuart's (1981) competitive exclusion hypothesis. Phylogenetic analyses of our seven gene dataset (3 mtDNA, 4 introns; 4784 bp) places these two species together in the genus Artisornis. We recover deep genetic divergence with geographic structure among populations of both tailorbird species, a result consistent with long-term co-existence of the two species. Thus, our data are consistent with the two species having achieved equilibrium and with successful resource partitioning having taken place over evolutionary time. From a conservation standpoint, our results suggest that extinction of the Long-billed Tailorbird as a function of competition with African Tailorbird is highly unlikely and should not be viewed as imminent. Finally, our empirical results suggest that mis-specification of the branch-length prior in Bayesian analyses of mitochondrial DNA data can have a profound effect on the overall tree-length (sum of branch-lengths), whereas the topology and support values tend to remain more stable. In contrast, mis-specification of the branch-length prior had a lesser impact on all aspects of the nuclear-only DNA analyses. This problem may be exacerbated when mitochondrial and nuclear DNA analyses are combined in a total evidence approach.
... @BULLET Zootaxa has published several other relatively short descriptions of family group names in Aves, based on an assessment of latest molecular studies (e.g. Cibois et al. 2010, Bowie et al. 2013). At the time of rejection of MS1, I thought little of it, because rejection of papers by journals is normal and happens all the time to everyone. ...
... Therefore, if host sympatric mtDNA structure were due to a scenario of despeciation, distinct parasite lineages also would be present due to codiversification with the previously isolated host lineages. Our preliminary mtDNA sequencing of Xanthomixis zosterops (Sharpe, 1875), a forest-dependent Malagasy passerine belonging to the endemic family Bernieridae (Cibois et al. 2010), showed the presence of multiple sympatric, cryptic, deeply divergent clades that did not sort strongly by geography or recognized subspecies (Salomonsen 1934; Clements et al. 2014), indicating possible despeciation. We test the hypothesis that the X. zosterops clades are merging by examining DNA sequence data from mitochondrial and nuclear markers across much of its known range, as well as sequence data from feather lice (Myrsidea, Phthiraptera). ...
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The merger of formerly isolated lineages is hypothesized to occur in vertebrates under certain conditions. However, despite many demonstrated instances of introgression between taxa in secondary contact, examples of lineage mergers are rare. Preliminary mtDNA sequencing of a Malagasy passerine, Xanthomixis zosterops (Passeriformes: Bernieridae), indicated a possible instance of merging lineages. We tested the hypothesis that X. zosterops lineages are merging by comparing mtDNA sequence and microsatellite data, as well as mtDNA sequence data from host-specific feather lice in the genus Myrsidea (Phthiraptera: Menoponidae). Xanthomixis zosterops comprises four deeply divergent, broadly sympatric, cryptic mtDNA clades that likely began diverging approximately 3.6 million years ago. Despite this level of divergence, the microsatellite data indicate that the X. zosterops mtDNA clades are virtually panmictic. Three major phylogroups of Myrsidea were found, supporting previous allopatry of the X. zosterops clades. In combination, the datasets from X. zosterops and its Myrsidea document a potential merger of previously allopatric lineages that likely date to the Pliocene. This represents the first report of sympatric apparent hybridization among more than two terrestrial vertebrate lineages. Further, the mtDNA phylogeographic pattern of X. zosterops, namely the syntopy of more than two deeply divergent cryptic clades, appears to be a novel scenario among vertebrates. We highlight the value of gathering multiple types of data in phylogeographic studies to contribute to the study of vertebrate speciation.
... The family Bernieridae is endemic to Madagascar and is made up of 11 species (Cibois et al. 2010;Safford and Hawkins 2013). Contrary to the better known Malagasy bird groups such as the family Vangidae (Yamagishi et al. 2001;Eguchi et al. 2002;Rakotomanana et al. 2009), little is known about the natural history of the Bernieridae and detailed information about the breeding biology and the reproductive success is especially scarce. ...
The Spectacled Tetraka Xanthomixis zosterops is a sexually monomorphic species in a family (Bernieridae) endemic to Madagascar and is found in various humid forest types. It is categorised as Least Concern by the IUCN. Despite its commonness, the biology of this species is little known. This study focused on its breeding biology and took place from September to December 2010 at Talatakely, Ranomafana National Park, Madagascar. Thirty nests were monitored and incubating birds were marked. Direct observations were made associated with nest building, incubating and nestling periods. Habitat structure was assessed to detect nesting site choice. The breeding episode, from the nest-building stage to fledging of the last chick, lasted on average 45 d. Only one individual, probably the female, in each presumed pair incubated eggs and brooded hatchlings, whereas both parents fed and cared for the nestlings. The mean clutch size was 2.4 ± 0.2 egg per nest (range 1–3, n = 24), the incubation period averaged 17.7 ± 0.3 d (range 16–19, n = 10), and the nestling period averaged 14.7 ± 0.3 d (range 14–15, n = 3). We estimated the reproductive success to be 13.6% (n = 59 eggs). Nest predation was believed to be the primary cause of nesting failure with 24 eggs (40.7%) and 18 (64.3%) nestlings being predated.
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The bulbuls comprise an ecologically important group of frugivorous, seed-dispersing birds found in Asia and Africa. Although several studies have examined the phylogenetic relationships of subsets of bulbul species, a comprehensive phylogeny of the family Pycnonotidae has hitherto been lacking. We used publicly available sequences generated from previous phylogenetic studies, augmented by new sequences from several unstudied taxa, to create a supermatrix from which to infer the phylogeny of the family. In all, we compared 121 of the 130 bulbul species. Our tree supports the monophyly of the family and comprises an exclusively African and a predominantly Asian clade. Several genera were found not to be monophyletic and we suggest taxonomic changes to provide a more accurate classification based on phylogeny. This article is protected by copyright. All rights reserved.
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Crossley's babbler (Mystacornis crossleyi) is a passerine endemic to Madagascar. Traditionally, it has been classified as a babbler (Timaliidae), although affinities with warblers and vangas have been suggested. We investigated the phylogenetic affinities of Crossley's babbler using sequence data from two nuclear introns (myoglobin intron 2 and beta-fibrinogen intron 5) and one mitochondrial gene (ND2). We present for the first time (to our knowledge) a molecular phylogeny that confidently places this enigmatic species within the vangas (Vangidae). The inclusion of Crossley's babbler within the vangas adds another foraging niche--gleaning small invertebrates from the ground-to this already large adaptive radiation of songbirds.
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The bird fauna of Madagascar includes a high proportion of endemic species, particularly among passerine birds (Aves: Passeriformes). The endemic genera of Malagasy songbirds are not allied obviously with any African or Asiatic taxa, and their affinities have been debated since the birds first were described. We used mitochondrial sequence data to estimate the relationships of 13 species of endemic Malagasy songbirds, 17 additional songbird species, and one species of suboscine passerine. In our optimal trees, nine of the 13 Malagasy species form a clade. although these birds currently are classified in three different families. In all optimal trees, the sister to this endemic clade is a group of Old World warblers including both African and Malagasy birds. The endemic Malagasy songbird clade rivals other island radiations, including the vangas of Madagascar and the finches of the Galapagos, in ecological diversity.
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The deep divergence between the African endemic passerines Picathartidae (rockfowl Picathartes and rockjumpers Chaetops, four species) and the Passerida (ca. 3500 species) suggests an older history of oscines on the African continent than has previously been assumed. In order to determine whether any additional, unexpectedly deep lineages occur in African endemic songbirds, 29 species--including 10 enigmatic focal taxa endemic to southern Africa--were added to a large nuclear sequence dataset gathered from oscine songbirds (Passeri). Phylogenetic analyses of these data resolve many long-standing questions about the affinities of these birds, not all of which were predicted by traditional approaches. The application of a molecular clock indicates that most basal divergences in Passerida occurred in the middle to late Eocene, with divergences between African and Australasian core corvoids occurring somewhat later in the early Miocene. Consistent with inferences for mammals, divergences between Malagasy endemic passerines and their mainland relatives suggests an asynchronous colonization history. This emerging phylogenetic picture reveals that relationships within Old World families are highly informative regarding the early dispersal and radiation of songbirds out of Gondwana. Future analyses will depend on improving resolution of higher-level phylogenetic relationships among these groups, and increasing the density of taxon sampling within them.
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Based on some general similarities in feeding adaptations, a large number of Old World passerine birds were in the past lumped in one broad family, the Sylviidae. Recent molecular studies, starting with the DNA-DNA hybridization work by Sibley et al. [Sibley, C.G., Ahlquist, J.E., 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution, Yale University Press, New Haven, CT], have revealed that this group is in fact a paraphyletic assemblage, mainly in the superfamily Sylvioidea, and within this assemblage a distinct group (the Cisticolidae) can be identified around the genus Cisticola. In this study we try to define natural lineages within it, based on DNA sequence data from 35 ingroup taxa representing 12 putative genera. Both nuclear myoglobin intron II (630 bp in our study) and mitochondrial ND2 (1041 bp) genes were sequenced, and 1671 bp were aligned and subjected to parsimony, maximum likelihood and Bayesian analyses. The results strongly support the monophyly of a cisticolid clade, with the Malagasy warblers Neomixis constituting the deepest branch within the clade. Three major clades receive statistical support, but not all relationships between and within these are well resolved. All species of the genus Bathmocercus belong to the Cisticolidae but in two different clades. The tailorbirds appear also polyphyletic with most species of the genus Orthotomus (but O. cucullatus falling in the outgroup) and the African metopias being in two different clades. Also the genus Apalis is polyphyletic, but all other included genera seem to be confirmed as natural units. Based on these findings we resurrect the genera Scepomycter and Artisornis. Calamonastes is confirmed to be in the Cisticolidae and grouped with Camaroptera. Main basic nest types do not follow the phylogenetic branching, and notably the peculiar "tailorbird" technique of stitching leaves together around the nest is found in different parts of the phylogeny. The basic types of nests seem to be found in particular environments, and the sewing may therefore have evolved in some ancestor of the Cisticolidae and was later lost or modified in some genera or species following the spread of drier habitats from the mid-Miocene.
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The island of Madagascar and surrounding volcanic and coralline islands are considered to form a biodiversity hotspot with large numbers of unique taxa. The origin of this endemic fauna can be explained by two different factors: vicariance or over-water-dispersal. Deciphering which factor explains the current distributional pattern of a given taxonomic group requires robust phylogenies as well as estimates of divergence times. The lineage of Indian Ocean scops-owls (Otus: Strigidae) includes six or seven species that are endemic to Madagascar and portions of the Comoros and Seychelles archipelagos; little is known about the species limits, biogeographic affinities and relationships to each other. In the present study, using DNA sequence data gathered from six loci, we examine the biogeographic history of the Indian Ocean scops-owls. We also compare the pattern and timing of colonization of the Indian Ocean islands by scops-owls with divergence times already proposed for other bird taxa. Our analyses revealed that Indian Ocean islands scops-owls do not form a monophyletic assemblage: the Seychelles Otus insularis is genetically closer to the South-East Asian endemic O. sunia than to species from the Comoros and Madagascar. The Pemba Scops-owls O. pembaensis, often considered closely related to, if not conspecific with O. rutilus of Madagascar, is instead closely related to the African mainland O. senegalensis. Relationships among the Indian Ocean taxa from the Comoros and Madagascar are unresolved, despite the analysis of over 4000 bp, suggesting a diversification burst after the initial colonization event. We also highlight one case of putative back-colonization to the Asian mainland from an island ancestor (O. sunia). Our divergence date estimates, using a Bayesian relaxed clock method, suggest that all these events occurred during the last 3.6 myr; albeit colonization of the Indian Ocean islands were not synchronous, O. pembaensis diverged from O. senegalensis about 1.7 mya while species from Madagascar and the Comoro diverged from their continental sister-group about 3.6 mya. We highlight that our estimates coincide with estimates of diversification from other bird lineages. Our analyses revealed the occurrence of multiple synchronous colonization events of the Indian Ocean islands by scops-owls, at a time when faunistic exchanges involving Madagascar was common as a result of lowered sea-level that would have allowed the formation of stepping-stone islands. Patterns of diversification that emerged from the scops-owls data are: 1) a star-like pattern concerning the order of colonization of the Indian Ocean islands and 2) the high genetic distinctiveness among all Indian Ocean taxa, reinforcing their recognition as distinct species.
Coquerel's Coua (Coua coquereli) and Red-capped Coua (Coua ruficeps) occur in the western dry forest of Ankarafantsika in northwest Madagascar. This kind of forest is characterized by an alternating of a dry and a rainy season. Although they belong to the same genus, the two species differ in appearance, with Red-capped Coua being slender than Coquerel's Coua. We analyse their respective foraging strategies, according to the seasonal variation. The foraging behaviour of both species was different and was also influenced by the seasonal variations. They tended to use the same main substrates but differed in the proportion of foraging techniques according to the season. Seasonal variations probably have an important effect on the prey availability (estimated by the rate of prey capture), the nature of prey and also alternative foraging substrates used, forcing the two species to use different techniques and probably to capture different prey. These different foraging strategies could maintain the coexistence between these two species.
The phylogenetic relationships of the Timaliidae (babblers) and Sylviidae (warblers) have long challenged ornithologists. We focus here on three Malagasy genera currently assigned to the Timaliidae, Mystacornis, Oxylabes, and Neomixis, and on their relationships with other babblers and warblers using the sequences of two mitochondrial genes (cytochrome b and 16S rRNA). Maximum parsimony analyses show that the Malagasy "babblers" are not related to any of the other African and Asian babblers. The genus Mystacornis is neither a babbler nor a warbler. The other Malagasy "babblers" are members of warbler groups (the monophyly of the Sylviidae is not demonstrated). Oxylabes madagascariensis and Hartertula flavoviridis (we recognize Hartertula as a genus for the species flavoviridis, previously Neomixis flavoviridis) constitute, with two presumed sylviine taxa, Thamnornis chloropetoides and Cryptosylvicola randrianasoloi, a warbler radiation endemic to the island of Madagascar. The other Neomixis species (tenella, striatigula, and viridis) belong to another warbler group comprising cisticoline taxa. These results show that the Timaliidae did not disperse to Madagascar. Rather, the island has been colonized, independently, by at least two clades of warblers, probably originating from Africa, where the Sylviidae radiation has been the most extensive.
The avian clade Passerida was first identified based on DNA-DNA hybridization data [C.G. Sibley, J.E. Ahlquist, Phylogeny and Classification of Birds, 1990, Yale University Press, New Haven, CT]. Monophyly of the Passerida, with the exception of a few taxa, has later been corroborated in several studies; however, the basal phylogenetic relationships have remained poorly understood. In this paper, we review the current knowledge of the phylogenetic relationships within Passerida and present a new phylogeny based on three nuclear introns (myoglobin intron 2, ornithine decarboxylase introns 6 and 7, as well as beta-fibrinogen intron 5). Our findings corroborate recent molecular hypotheses, but also identify several hitherto unrecognized relationships.
Notes sur les collections rapportées en 1853, par M. A. Delattre, de son voyage en Californie et dans le Nicaragua
  • C.-L Bonaparte
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.