Download Two-phase extinction of “Southern Hemispheric” birds in the

Palaeobio Palaeoenv (2011) 91:325–333
DOI 10.1007/s12549-011-0062-4
ORIGINAL PAPER
Two-phase extinction of “Southern Hemispheric”
birds in the Cenozoic of Europe and the origin
of the Neotropic avifauna
Gerald Mayr
Received: 27 July 2011 / Revised: 12 September 2011 / Accepted: 21 September 2011 / Published online: 13 October 2011
# Senckenberg Gesellschaft für Naturforschung and Springer 2011
Abstract A considerable number of fossil birds from the
Cenozoic of Europe belong to taxa whose extant representatives are only found in the Southern Hemisphere. This study
presents the first detailed analysis of the stratigraphic
occurrences of these groups. Two well-separated extinction
phases can be distinguished: one in the Paleogene, which
concerned birds with crown group representatives in South
America, Madagascar and Australia, and the second in the
Miocene, which involved taxa that are today found in Africa
or have a pantropic distribution. It is hypothesised that this
unusual pattern is the result of a successive action of biotic and
abiotic factors. South America was least affected by both
extinction phases and served as a refugium for bird groups,
which had a more widespread distribution in the Paleogene.
Keywords Biogeography . Fossil birds . Climatic cooling .
Grande coupure
Introduction
As early as the 19th century it was noted that the Cenozoic
avifauna of Central Europe includes representatives of
avian groups that today inhabit the tropic and subtropic
regions, such as parrots and trogons (e.g. Milne-Edwards
1867–1871). As our knowledge of the Cenozoic avifaunas
of the Northern Hemisphere increased, it has become
evident that many other avian taxa have a complex
biogeographic history.
G. Mayr (*)
Sektion Ornithologie, Senckenberg Forschungsinstitut
und Naturmuseum Frankfurt,
Senckenberganlage 25,
60325 Frankfurt am Main, Germany
e-mail: [email protected]
Europe in particular has an extensive and well-studied
fossil record, which for some geological intervals provides
a detailed picture of the prevalent avifaunas and indicates
that the extant distribution of many Southern Hemispheric
avian “families” is relictual (Mayr 2009a; Mourer-Chauviré
1982; Olson 1989; contra Cracraft 2001). Olson (1989: p.
2024) even concluded that “almost all the major taxa that
are ‘characteristic’ of the southern continents are actually
relicts of once more widespread groups that occurred in the
Northern Hemisphere in the Paleogene or even later”.
In particular, a large number of birds existed in the
Cenozoic of Europe whose closest extant relatives only
occur in South or Central America (Mourer-Chauviré
1999). It has been briefly noted that the extinction of the
“South American” taxa precedes that of other “Southern
Hemispheric” avian groups in the Miocene (Blondel and
Mourer-Chauviré 1998), but the extent, timing and cause of
these extinctions remain elusive. Whereas some authors
assume that ecological competition played a major role
(Mayr 2009a; Olson 1989), others consider global cooling
during the Cenozoic to be the main cause (Hawkins et al.
2006; Lindow and Dyke 2006; Martin 2010).
Here I give the first detailed stratigraphic account on the
occurrence of “Southern Hemispheric” birds in Europe and
show that the extinction dates of these taxa correlate closely
with the distribution area of their extant representatives.
Successive and independent actions of biotic and abiotic
factors best explain the observed two-phase extinction pattern.
Methods
The stratigraphic occurrences of 71 bird species from Cenozoic
Northern Hemispheric localities are analysed whose crown
group representatives either have a pantropical distribution or
326
are only found on the Southern Hemispheric continents, i.e.,
South America, Africa (including Madagascar) and Australia.
The Podargidae (frogmouths), whose crown group representatives live in Australia and southeast Asia, are also considered.
Taxonomic sampling is restricted to well-constrained exemplar
species per stratigraphic unit. With the exception of few
records of Anhingidae (darters) there are no aquatic birds from
Cenozoic Northern Hemispheric fossil sites whose crown
group representatives have a pantropic distribution or are
restricted to the Southern Hemisphere, and all taxa considered
in this analysis represent terrestrial or arboreal birds. The
temporal focus is on Eocene and Miocene fossils because the
poorly known Paleocene avifaunas include no taxa of
relevance for the present study, whereas Pliocene avifaunas
have an essentially modern higher level composition. The data
are based on both own examination of fossils (mainly
Paleogene European taxa) and a literature review.
Results and comments
Figure 1 summarises the known temporal distribution of
avian higher level taxa with a fossil record in Europe and
North America and whose extant representatives have a
pantropic or Southern Hemispheric distribution. The individual species and their stratigraphic occurrences are listed
in the table in Appendix 1.
All taxa which disappeared in the Miocene occur today
in the Ethiopian zoogeographic realm or have a pantropic
extant distribution. Some of these groups, i.e., Coliiformes
(mousebirds), Trogoniformes (trogons) and Psittaciformes
(parrots), have a long evolutionary history in Europe. For
others, there exists an earlier fossil record in Africa, and
these birds may have dispersed into Europe during the
Miocene thermal maximum. Such was probably the case
for Struthionidae (ostriches), which first occur in the early
Miocene of Namibia (Mourer-Chauviré et al. 1996),
Bucorvinae (ground hornbills), which were reported from
the middle Miocene of Morocco (Brunet 1971), and
Musophagiformes (turacos), which have a fossil record in
the early Oligocene of Egypt (Rasmussen et al. 1987).
In contrast, all taxa that became extinct in the Paleogene
have crown group representatives in South or Central
America, Madagascar or southeast Asia and Australia.
Pervasive as the “South American” pattern appears, the list
of involved groups may not even be complete, as only
published species with a substantial fossil record were
considered. Not included, for example, are putative early
Eocene Anhimidae (screamers) from Europe and North
America, which have not yet been formally described (Mayr
2009a), as well as putative Galbulae from the early Eocene
North American Green River Formation (Weidig 2010),
whose tentative identification needs further verification.
Palaeobio Palaeoenv (2011) 91:325–333
Taking into account that North American avifaunas are less
well known than the European ones, the general pattern in
North America is similar to that seen in Europe. This is
especially evident for forest-dwelling taxa, with early Eocene
North American avifaunas showing a high taxonomic
congruence with those from the middle Eocene of Europe.
However, due to the earlier disappearance of paratropical
forests in North America, many of the arboreal taxa disappeared earlier there than in Europe (Mayr 2009a). None of
the Miocene forest dwelling taxa with African affinities have
been recorded in North America, and Balearicinae (crowned
cranes), which inhabit open habitats, represent the only
“Ethiopian” faunal element in the Miocene of North America.
Paleogene and early Cenozoic avifaunas of Asia and the
Southern Hemisphere are poorly known. Anseranatidae
(magpie geese) occur in the late Oligo-Miocene of Australia
(Worthy and Scanlon 2009), and stem group representatives
of the Cariamae (seriemas and allies) and Cathartidae (New
World vultures) have been reported from the Paleogene of
South America (Alvarenga 1985; Mayr 2009a). Some
species from the Paleogene of South America are closely
related to birds from contemporaneous European deposits,
and it is likely, albeit not yet proven, that stem group
representatives of the European taxa with South American
affinities also lived in the early Cenozoic of South America
(Mourer-Chauviré 1999).
Discussion
After the mid-Eocene, global cooling proceeded during the
Cenozoic, and by the end of the Miocene climatic optimum
maximum, there had been a drop of about 11°C in the
minimum cold months temperature (Fig. 1; Böhme 2003).
Emergence of a marked climatic seasonality towards the late
Miocene (Mosbrugger et al. 2005) limited food availability in
the cold northern hemispheric winters. Accordingly, the
extinctions after the onset of the late Miocene cooling
involve taxa whose extant representatives have poor dispersal
capabilities and a predominantly insectivorous or frugivorous
diet, such as Coliiformes, Musophagiformes, Bucorvinae,
Psittaciformes, and Ramphastidae (barbets sensu Dickinson
2003). These groups still occur today in the warm regions of
the Old World, and at least the latter three are species rich
and widely distributed. Their disappearance between the
stratigraphic units MN 6 and MN 11 (Fig. 1) corresponds
well with Central European extinctions of thermophilic
reptiles during the same period (Böhme 2003).
The Paleogene extinctions, however, do not correlate with
major climatic events. Although the mean annual temperatures
declined significantly by about 15°C during the Eocene, they
were still equable during the Oligocene (e.g. Mosbrugger et al.
2005). As noted above, thermophilic reptiles, such as
Palaeobio Palaeoenv (2011) 91:325–333
327
Fig. 1 Stratigraphic occurrence of avian taxa with Southern Hemispheric extant relatives in the Palaeogene and Miocene of Europe
(black lines) and North America (grey lines). The stratigraphic
positions of actual fossils (Appendix 1) are indicated by squares;
question marks denote uncertain identifications (squares) or unknown
exact stratigraphic dates (encircled). The grey ellipses highlight the
two major extinction periods. The curve on the right shows marine
oxygene isotope records as a global temperature proxy reconstruction
(from Zachos et al. 2001). Stratigraphy follows Legendre and Lévêque
(1997) for the Palaeogene (MP) units and Böhme (2003) for the
Neogene (MN) ones. Austral Australian region, Md Madagascar
crocodilians and chameleons, were found in Central Europe
until the middle Miocene (MN 6), with Varanidae persisting
even until the late Miocene (MN 11; Böhme 2003). It is
unlikely that birds were more affected by global cooling than
these ectothermic groups. Climate-driven extinctions are
further not expected to have had such a strong correlation
with the distribution area of the closest extant relatives, which
mainly survived in the Neotropical region but not in the
geographically closer Old World zones with similar climates.
Because most of the taxa that disappeared in the
Paleogene have affinities to South America, which was
isolated from the northern continents until the Pliocene
closure of the Panamanian Isthmus (Smith et al. 1994),
biotic factors are more likely to have caused their
extinction. The extant representatives of these groups have
very different life histories, and previous attempts to explain
their disappearance have failed to find an explanation (Mayr
2009a). However, it went unnoticed that their last occurrences
correspond with the disappearance of flightless birds in the
early Cenozoic of continental Europe.
During most of the Eocene, Europe was geographically
isolated from Asia by the Turgai Strait (Fig. 2), and the
comparatively high number of medium-sized early Paleogene
flightless birds indicates a low predation pressure at that time
(Mayr 2009a). Closure of this seaway at the Eocene/
Oligocene boundary commenced a faunal turnover known
as the Grande Coupure. The immigration of new taxa from
Asia led to the extinction of numerous archaic European
mammals (e.g. Hooker 2010), but the immediate effects of the
Grande Coupure on birds are not well understood (Mayr
2009a). Taking into account that the exact stratigraphic
occurrences of the latest European Nyctibiidae (potoos) and
Leptosomidae (courols) are unknown, only Cariamae have a
fossil record after the early Oligocene. The latest unambiguously established occurrence of a flightless bird in the
Paleogene of continental Europe, a coracoid that was assigned
328
Palaeobio Palaeoenv (2011) 91:325–333
Fig 2 Phylogenetic interrelationships of fossil and extant “caprimulgiform” and apodiform birds (after Mayr 2009a). The dotted lines
point to distribution areas on a palaeomap of the continents in the
middle Eocene (after Smith et al. 1994). The daggers denote extinct
taxa, the asterisks indicate those with a worldwide distribution
to Strigogyps dubius (“Ameghinornis minor”), also stems
from the early Oligocene (stratigraphic unit MP 23) of France
(Mourer-Chauviré 1983: p. 127), and there is no record of
flightless birds until the late Miocene occurrence of ostriches.
In modern insular environments, predation by mammals
and snakes is the main cause of avian extinctions, and apart
from true carnivores, rats and mice pose a particular threat
for eggs and nestlings (e.g. Wanless et al. 2007). The new
mammalian taxa, which arrived in Europe just after the
Grande Coupure, indeed included muroid rodents as well as
carnivorans, such as viverrids and felids (Rose 2006). If a
lower predation pressure in the Eocene allowed more
exposed nesting sites, arboreal birds are expected to have
been seriously affected by the new immigrants (e.g. Sieving
1992; Galetti et al. 2009). Although this hypothesis can not
be verified through the fossil record, it is notable that extant
“caprimulgiform” and apodiform birds, which are among
the groups with the most distinctive Southern Hemispheric
pattern (Fig. 2), exhibit strikingly different nesting behaviours, ranging from ground breeding (Caprimulgidae,
nightjars), cave nesting (Steatornithidae, oilbirds), hole breeding (Aegothelidae, owlet-nightjars) and saliva use (Apodidae,
swifts), to elaborate nest constructions (Trochilidae). The
independent evolution of such divergent strategies is in line
with the hypothesis of strong selective forces acting on the
breeding biology of the Paleogene representatives of the
respective lineages. Psittaciformes and Trogoniformes, by
contrast, which were not affected by the Paleogene extinctions,
excavate nesting cavities in tree trunks (Aegothelidae are
dependent on existing ones). Coliiformes build open-bowl
nests like passeriform birds, whose sophisticated nest building
behaviour is considered a key factor to explain their adaptive
radiation (Olson 2001).
Carnivorous marsupials also occurred in the Cenozoic of
South America (e.g. Rose 2006), but contrary to the
Northern continents there were no pre-Pliocene large-scale
immigrations of new taxa, and birds thus coevolved with
their predators. It is the sudden immigration of new
predators in the Cenozoic of Europe, which is considered
to have had a dramatic impact on avian communities.
In conclusion, the extinctions of “Southern Hemispheric”
taxa in Europe are best explained by a two-phase extinction
model that combines late Miocene extinctions due to the
emergence of Northern Hemispheric winters and earlier
Paleogene extinctions after faunal turnovers. Contrary to
previous hypotheses (Lindow and Dyke 2006), climatic
cooling does not explain Paleogene extinctions of “Southern
Hemispheric” taxa. Likewise, not all Southern Hemispheric
taxa are relics of groups that were once more widespread on
the Northern Hemisphere (contra Olson 1989) and, as noted
above, some of the African taxa seem to have dispersed into
Europe during the Miocene.
It is especially the Neotropic realm and, to a lesser
degree, Madagascar and Australia, which served as refugia
for avian groups that had a more widespread distribution on
the Northern Hemisphere during the early Cenozoic. There
are also some non-avian faunal elements in the Paleogene
of Europe, whose closest extant relatives are found in South
Palaeobio Palaeoenv (2011) 91:325–333
329
America, Madagascar, or Australia, such as Marsupialia
and Boidae (e.g. Morlo et al. 2004). The high number of
“Neotropic” elements in Paleogene European avifaunas is,
however, unparalleled by other vertebrates and can be
explained by the much greater dispersal capabilities of
birds. If these groups did not already have a wide
distribution in the Paleogene, they could certainly better
respond to environmental changes by dispersal than nonvolant vertebrates.
Acknowledgements I thank Ursula Göhlich, Naturhistorisches
Museum Wien, for photos of coliiform birds from Kohfidisch in
Austria. Comments by B. Lindow and the reviewers Z. Bochenski and
C. Mourer-Chauviré improved the manuscript.
Appendix 1
Table 1 Species used as reference points for Fig. 1
Family
Fossil species
Geographic
occurrence
Stratigraphic occurrence
References
Anseranatidae
Anatalavis (Nettapterornis)
oxfordi Olson
Anserpica kiliani Mourer-Chauviré
et al.a
Masillapodargus longipes Mayr
Quercypodargus olsoni
Mourer-Chauviré
Idiornis tuberculata Peters
?Idiornis anthracinus Mayr
Idiornis gallicus (Milne-Edwards)
Idiornis cursor (Milne-Edwards)
Idiornis gracilis (Milne-Edwards)
Eutreptornis uintae Cracraft
England
MP 8
France
MP 29/30
Olson (1999), Mlíkovský
(2002)
Mourer-Chauviré et al. (2004)
Germany
France
MP 11
MP 16
Mayr (2009a)
Mourer-Chauviré (2006)
Germany
Germany
France
France
France
Utah, USA
MP 11
MP 13
MP 16, 17a, 19
MP 22, 23, 28
MP 23, 25, 28
Middle Eocene (Uinta Formation)
Podargiformes
Cariamae
Bathornis veredus Wetmore
Paracrax gigantea Cracraft
Cathartidae
Nyctibiidae
Steatornithidae
Momotidae
Todidae
Trochilidae
Leptosomidae
Bathornis fricki Cracraft
Diatropornis ellioti
(Milne-Edwards)b
Phasmagyps patritus Wetmore
Paraprefica kelleri Mayr
Euronyctibius kurochkini
Mourer-Chauviré
Prefica nivea Olson
Protornis glarniensis von Meyer
gen. et sp. indet.
Palaeotodus escampsiensis
Mourer-Chauviré
Palaeotodus itardiensis
Mourer-Chauviré
Palaeotodus emryi Olson
Parargornis messelensis Mayr
Jungornis geraldmayri
Mourer-Chauviré
Eurotrochilus inexpectatus Mayr
Plesiocathartes kelleri Mayr
Plesiocathartes geiselensis Mayr
Colorado, USA
South Dakota,
USA
Wyoming, USA
France
Mayr (2009a)
Mayr (2002)
Mourer-Chauviré (2006)
Mourer-Chauviré (2006)
Mourer-Chauviré (2006)
Brodkorb (1967), Mayr
(2009a)
Late Eocene (Chadron Formation) Brodkorb (1967), Mayr (2009a)
Early Oligocene (Poleslide
Brodkorb (1971), Mayr (2009a)
Member, Brule Formation)
Late Oligocene (Arikareean)
Brodkorb (1971), Mayr (2009a)
MP 16, 17b
Mourer-Chauviré (2002, 2006)
Colorado, USA
Germany
France
Late Eocene (Chadronian)
MP 11
“MP 16 –28”
Mayr (2009a)
Mayr (2009a)
Mourer-Chauviré (2006)
Wyoming, USA
Early Eocene (Green River
Formation)c
“MP 21–24”
Late Miocene (early Hemphillian)
MP 19
Olson (1987)
Switzerland
Florida, USA
France
Olson (1976), Mlíkovský (2002)
Becker (1986)
Mourer-Chauviré (2006)
France, Germany MP 23
Mourer-Chauviré (2006)
Wyoming, USA
Olson (1976)
Germany
France
Early Oligocene (Orellan, Brule
Formation)
MP 11
MP 17b
Germany
Germany
Germany
MP 23d
MP 11
MP 13
Mayr (2009a)
Mourer-Chauviré and Sigé
(2006)
Mayr and Micklich (2010)
Mayr (2008)
Mayr (2002, 2008)
330
Palaeobio Palaeoenv (2011) 91:325–333
Table 1 (continued)
Family
Struthionidae
Balearicinae
Sagittariidae
Coliiformes
Fossil species
Stratigraphic occurrence
References
Plesiocathartes europaeus Gaillard France
“MP 16–28”
Plesiocathartes wyomingensis
Weidig
Struthio chersonensis (Brandt)
Balearica rummeli (Mlíkovský)
Balearica exigua Feduccia and
Voorhies
Pelargopappus schlosseri
(Milne-Edwards)
Pelargopappus magnus
(Milne-Edwards)
Selmes absurdipes Peters
Celericolius acriala Ksepka and
Clarke
Palaeospiza bella Allen
Germany
Nebraska, USA
Early Eocene (Green River
Formation)c
MN 9–MN 11/12
MN 2-3
Late Miocene (late Clarendonian)
Mourer-Chauviré (2002),
Mayr (2008)
Mayr (2008)
France
MP 23, 28
France
MN 2a
Germany
Wyoming, USA
France
MP 11
Early Eocene (Green River
Formation)c
Latest Eocene (Florissant
Formation, 34 Ma)
MP 16, 17b, 19
Germany
MP 23d
Mayr (2009a)
France
MN 2a
France
MN 6, 7-8
Austria
France
MP 11e
MN 5
France
MN 8
Ballmann (1969a), Mlíkovský
(2002) , Mayr (2010)
Mlíkovský (2002), Mayr
(2010)
Mlíkovský (2002), own obs.
Ballmann 1972), Mlíkovský
(2002)
Ballmann (1969a), Mlíkovský
(2002)
Ballmann (1969b), Mlíkovský
(2002)
Primocolius sigei
Mourer-Chauviré
Oligocolius brevitarsus Mayr
Limnatornis paludicola
(Milne-Edwards)
Necrornis palustris
(Milne-Edwards)
gen. et sp. indet.
Musophagiformesf gen. et sp. indet.
Phoeniculidae
Bucorvinae
Psittaciformes
Trogoniformes
Veflintornis (Apopempsis) meini
(Ballmann)
Phirriculus pinicola Mlíkovský
and Göhlich
Euroceros bulgaricus Boev and
Kovachev
Psittacopes sp.
Cyrilavis (“Primobucco”) olsoni
(Feduccia and Martin)
Psittacopes lepidus Mayr and
Daniels
Quercypsitta sudrei MourerChauviré
Archaeopsittacus verreauxi
(Milne-Edwards)
Xenopsitta fejfari Mlíkovský
Bavaripsitta ballmanni Mayr
and Göhlich
Pararallus dispar
(Milne-Edwards)
Conuropsis fratercula Wetmore
Septentrogon madseni
Kristoffersen
gen. et sp. indet.
Masillatrogon pumilio Mayr
Primotrogon wintersteini Mayr
Paratrogon gallicus
(Milne-Edwards)
Geographic
occurrence
Wyoming, USA
Colorado, USA
France, Germany MN 2a, MN3a, MN4
Mlíkovský (2002)
Mourer-Chauviré (2001)
Feduccia and Voorhies (1992)
Mourer-Chauviré and Cheneval
(1983)
Mourer-Chauviré and Cheneval
(1983)
Mayr (2009a)
Ksepka and Clarke (2010a)
Ksepka and Clarke (2009)
Mourer-Chauviré (2006)
Bulgaria
MN 11-12
Boev and Kovachev (2007)
England
Wyoming, USA
Mayr and Daniels (1998)
Mayr (2009a), Martin (2010)
Germany
MP 8
Early Eocene (Green River
Formation)c
MP 11
France
MP 17a
Mourer-Chauviré (2006)
France
MN 2a
Czech Republic
Germany
MN 3
MN 6
Milne-Edwards (1867-71),
Mlíkovský (2002)
Mlíkovský (2002)
Mayr and Göhlich (2004)
France
MN 6
Nebraska, USA
Denmark
Late Miocene (lower Snake
Creek beds)
MP 7
Mayr (2009a)
England
Germany
MP 8
MP 11
Mayr (2009a)
Mayr (2009b)
France
France
MP 23
MN 2a
Mayr (2009a)
Milne-Edwards (1867–1971),
Mlíkovský (2002)
Mayr and Daniels (1998)
Cheneval (2000), Mlíkovský
(2002)
Brodkorb (1971)
Palaeobio Palaeoenv (2011) 91:325–333
331
Table 1 (continued)
Family
Fossil species
Ramphastidae
Suboscinesh
Geographic
occurrence
Stratigraphic occurrence
References
Rupelramphastoides knopfi Mayrg Germany
Capitonides europeus Ballmann
Germany
MP 23d
MN 3
Capitonides protractus Ballmann
Germany
MN 6
Capitonides sp.
France
MN 7-8
cf. Pogoniulus
gen. et sp. indet.
gen. et sp. indet.
gen. et sp. indet.
gen. et sp. indet.
Eurylaimidae indet.
Austria
Florida, USA
France
France
Germany
Germany
MN 10
Early Miocene (Hemingfordian)
MP 23
MP 26
MP 28
MN 3
Mayr (2009a)
Ballmann (1969b), Mlíkovský
(2002)
Ballmann (1983), Mlíkovský
(2002)
Ballmann (1969b), Mlíkovský
(2002)
Mlíkovský (2002)
Olson (1985)
Mayr and Manegold (2006)
Mourer-Chauviré (2006)
Manegold (2008)
Ballmann (1969b), Mlíkovský
(2002)
For European species, the Mammalian Paleogene (MP) and Mammalian Neogene (MN) stratigraphic units are indicated. Quotation marks indicate
that the exact stratigraphic position within a range is uncertain
a
The affinities of this species, which is only known from the coracoid, are uncertain (see Mayr 2009a)
b
Diatropornis may be the sister taxon of a clade including Cathartidae and the New World Teratornithidae (Mayr 2009a), which would, however,
not affect the conclusions of the present study
c
Following Ksepka and Clarke (2010a, b), the age of the Fossil Butte Member of the Green River Formation, from which most fossil birds stem,
is set at 52 Ma
d
See Mayr and Micklich (2010) for stratigraphic assignment
e
See Daxner-Höck (2004) for the age of the Kohfidisch site
f
Identification of a putative musophagid from the late Oligocene of Germany (Ballmann 1970) can not be upheld (Mayr 2009a)
g
Rupelramphastoides exhibits a barbet-like overall morphology, but the two known skeletons do not allow for an unambiguous phylogenetic
placement of the taxon, which may be outside crown group Pici (Mayr 2005)
h
I agree with Mlíkovský (2002: p. 257) that the identification of alleged Dendrocolaptidae in the middle Miocene (MN 6) of France (Cheneval
2000) is ill-founded
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Becker JJ (1986) A fossil motmot (Aves: Momotidae) from the late
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