REVIEWS

Evolution and history of the western

Palaearctic avifauna
Jacques Blondel and Cécile Mourer-Chauviré

T
here is still much un- Recent data accumulated from fields as Cretaceous forms, such as Ambi-
certainty and controversy varied as avian palaeontology, ortus and Gansus, the toothed birds
about the early history palaeobotany, historical biogeography and of the Late Cretaceous (Hesper-
of birds, which began dur- molecular phylogenetics provide a ornithiformes and Ichthyornithi-
ing the Mesozoic [Late Jurassic, completely renewed picture of the origin, formes), as well as Neornithes. By
145 million years ago (Mya)]. The evolution and distribution of modern birds. the end of the Cretaceous only
debate concerns the phylogenetic Although the origin of birds is still Neornithes remained.
relationships among the many controversial, their Tertiary history is now Regardless of the arguments
Mesozoic lineages, the origin of well known. The reconstruction of concerning the timing of the
modern birds (Neornithes), and palaeoenvironments and the identification origins of modern birds or the
the consequences of the mass ex- of shifts in major vegetation belts and effects of the K–T mass extinc-
tinction of dinosaurs at the Cre- habitats during the Pliocene–Pleistocene tions on avian evolution, it is un-
taceous–Tertiary (K–T) boundary epochs have added to these recent disputed that the fossil record
(65 Mya), a key episode in verte- developments. Together they provide a indicates that all orders of birds
brate history. There are two main new perspective on speciation and living today were present in the
hypotheses. The first is that this extinction rates since the late Pliocene western Palaearctic during the
devastating crisis not only af- and the establishment of modern ten million years that followed
fected dinosaurs, but also caused avifaunas in the western Palaearctic. the end of the Cretaceous (in the
the extinction of most birds (and Paleocene and Early Eocene,
mammals). This resulted in a se- 65–55 Mya) (Ref. 1). The large
vere bottleneck that was followed Jacques Blondel is at the Centre d’Ecologie number of fossils that have been
by a period of intense avian diver- Fonctionnelle et Evolutive/CNRS, UPR 9056, found from this epoch in the
34293 Montpellier cedex 5, France
sification in the Early Tertiary. (blondel@cefe.cnrs-mop.fr); Cécile Mourer-Chauviré is Northern Hemisphere, especially
The bottleneck consisted of a few at the Centre des Sciences de la Terre/UMR 5565, in Europe6–8, reveal that several
surviving lineages (or perhaps Université Claude Bernard Lyon I, 43 Boulevard du 11 diagnostic characteristics of mod-
even only a single lineage) of the novembre 1918, 69622 Villeurbanne cedex, France ern avifauna were already pres-
Late Cretaceous, called ‘transi- (mourer@cismsun.univ-lyon1.fr). ent at that time (Box 1). Among
tional shorebirds’, comparable to the 44 families identified in the
extant Charadriiformes (waders)1. Late Eocene avifauna of Europe,
The alternative hypothesis is 30 are Recent (Holocene) families
that most modern avian (and mammal) orders originated and 22 of them still have representatives in the modern avi-
much earlier, in the Early Cretaceous (140 Mya). The exten- fauna of the western Palaearctic.
sive avian diversification in this epoch was characterized The Paleogene avifauna included diversified nonpass-
by species that acquired specializations for a variety of erine tropical birds, which were widespread in the entire
habitats2. According to this view, the presence of many lin- Northern Hemisphere as a result of the absence of climatic
eages of modern birds at the end of the Cretaceous is and physical barriers. During the Paleogene (65–25 Mya),
indicative of highly selective extinction among vertebrates continents were more or less flat, without ice sheets and
during the K–T mass extinction crisis. Consequently, it large cordilleras. Moreover, the earth’s climate was sub-
casts doubt on the more widespread hypothesis of an tropical, with no marked seasonality, and forests domi-
explosive cladogenesis of modern birds at the beginning of nated by oaks, laurels and palms prevailed as far north as
the Tertiary2. The earlier diversification of avifauna is sup- northern Canada, Greenland and northern Scandinavia12.
ported by recent molecular data, which indicate that most The concurrence between the temperate floras of the west-
modern orders diverged long before the Early Tertiary, ern Palaearctic, eastern North America and eastern Asia
about 100 Mya (Refs 3 and 4). today is another indication of the homogeneous ecological
conditions that existed over these regions13.
The fossil record of modern avifauna before the The numerous fossils provide good evidence that most
Quaternary modern orders and families of birds appeared between
Proponents of the two hypotheses have many argu- the Early Eocene and the Late Oligocene–Early Miocene
ments with which to defend their viewpoints. The debate is (23 Mya). This was not only an epoch of intense diversifi-
still centred around which epochs modern avian orders cation, but also one in which the Recent fauna of the North-
originated from, because the timing depends on whether ern Hemisphere was established7,8. The gradual global cool-
dates are calibrated from molecular data3,4 or from ing, which started in the mid-Oligocene (30 Mya), resulted
palaeontological records1,5. The two schools, however, do in the appearance of seasonal climates at mid-latitudes and
agree that all modern birds derive from a group (the the contraction of the tropics towards the Equator14. Com-
Ornithurae) that differentiated in the Early Cretaceous1,2. bined with the increasing isolation of Eurasia from North
The Ornithurae descended from forms that subsequently America, these climatic changes favoured diversification
gave rise to Archaeopteryx and a group of diversified Creta- and continental specialization of avifauna in new types of
ceous birds, the Enanthiornithes. Ornithurae include Early habitat (temperate forests, grasslands and steppes). From

488 0169-5347/98/$ – see front matter © 1998 Elsevier Science. All rights reserved. PII: S0169-5347(98)01461-X TREE vol. 13, no. 12 December 1998

the Miocene onwards, evolution of birds generally involved
differentiation at the genus and species levels within exist-
ing families.
In contrast to all other orders and families of extant birds
that occurred in the western Palaearctic in the Eocene, the
Columbidae (pigeons), Anatidae (ducks), Podicipediformes
(grebes), Psittacidae (parrots) and Passeriformes did not
appear as important components of the fauna in the North-
ern Hemisphere until the Late Oligocene–Early Miocene8,15,16.
Today, Passeriformes constitute the most abundant and
widely distributed order of living birds (60% of the world
avifauna), occurring on all continents except Antarctica15,
but the niches they currently occupy were presumably
filled in the Eocene and Oligocene by small nonpasserine ar-
boreal birds, such as Sandcoleiformes (extinct order related
to mousebirds), Coliiformes (mousebirds), Caprimulgi-
formes (nightjars and frogmouths), Coraciiformes (rollers
and allies) and Apodiformes (swifts and allies), which are
common in the extensive Early Tertiary fossil deposits of
both North America and Europe17. It has been proposed
that passerines originated in the protocontinent Gond-
wanaland18 (which subsequently split to form the southern
continents, where the diversity of primitive passerines is
the highest), and then spread north at the beginning of the
Miocene. The recent discovery of passerine fossils in the
Early Eocene strata of Australia19 provided the first evi-
dence that this continent could have been the birthplace of
passerines.
All the changes that started in the Late Oligocene
resulted in the withdrawal of tropical birds from the North-
ern Hemisphere. These have been progressively replaced
by temperate bird assemblages. Most of the major taxa
that are now restricted to the tropics are relics of groups
that were widespread in the early Tertiary in what is cur-
rently the temperate zone of the Northern Hemisphere8,15
(Fig. 1). Of all the families presently confined to continental
Africa, only the Hamerkop (Scopidae) and the Shoebill
(Balaenicipitidae) are unknown as fossils outside the Afro-
tropical realm15.
Quaternary development of the western Palaearctic
avifauna
The climatic upheavals of the Late Tertiary reached a
peak in the Late Pliocene and the Pleistocene (i.e. occur-
ring over the past 2.3 million years20,21) and were charac-
terized by a series of about 20 strong short-term wet–dry
and cool–warm fluctuations. Over timescales of 103–105
years, these climatic changes continuously forced major
geographical shifts in species distributions and rearrange-
ments in species assemblages21–23. Using fossil pollen data
to draw ‘isopoll maps’ (maps with lines of equal relative
pollen abundance for a taxon), palaeobotanists have
mapped the range extensions and shifts in distribution of
several important tree species since the last glacial maxi-
mum22,23 (18 000 BP). Although direct evidence is sparse,
birds and other animal communities must also have
‘tracked’ their habitats as these shifted in response to en-
vironmental change22. During glacial episodes, virtually no
arborescent vegetation persisted north of the large Eur-
asian mountain ranges (Pyrenees, Alps, Carpathians and
Himalayas), causing the forest biota of the western Palae-
arctic to be concentrated in the south and south-east of
Europe (e.g. the Mediterranean basin). During interglacial
periods, the forest biota re-expanded to the north22. Many
fossils of extant bird species that currently live in the far
north have been found at sites in southern Europe that date
back to the last glacial episode, indicating the magnitude of
TREE vol. 13, no. 12 December 1998
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Box 1. Fossil birds of the Tertiary
Since the mid-seventies, the description and revision of Tertiary fossil birds have
provided an enormous amount of information on the epochs of differentiation
and past distribution of Recent orders and families. The map shows the Euro-
pean distribution of the 503 Paleogene (65–25 Mya, encircled) and Neogene
(25–1.6 Mya) localities9 that provided 445 fossil species10. European sites have
played a major role in deciphering Paleogene avifaunas, with such localities as
Walton-on-the-Naze (early Eocene, 53 Mya) in the UK, Messel (mid-Eocene,
49 Mya) in Germany, and the famous group of 27 localities spread over an area
of about 300 km2, the so-called ‘Phosphorites du Quercy’ (mid-Eocene to Late
Oligocene, 40 to 26 Mya) in France (square on the map). Localities of similar age
in North America belong to the early Eocene (Green River and Willwood)7, but
these avifaunas are much less diverse and still incompletely studied. Preliminary
identifications in these deposits provided 25 families as compared with 55 families
at the UK site7 and 42 at the French site8. In the Quercy site, which extends over
14 million years, fossils of small mammals provided cues for building biochrono-
logical scales, which are internationally accepted as Paleogene reference levels,
making it possible to date fossil birds accurately. The large number of discrete
fossil deposits at the Quercy site provides a better picture of the diversity of the
avifauna than just a single site. Europe has also been an important source of
Neogene fossils – 270 localities have provided significant avian fossils9 as com-
pared with 138 in North America11, although the number of Neogene species is
very similar in the two regions (229 in Europe9 and 211 in North America11).
these migrations24. Given the alternating full-glacial and
interglacial episodes, two types of north–south shift (and
vice versa) have been hypothesized to occur repeatedly
across continents25.
First, species with very large contemporary distribution
ranges expanded to the north during interglacial intervals
without leaving their southern refugia. They alternated
between reduced and often fragmented populations during
unfavourable glacial periods and large extensive popu-
lations during favourable interglacial periods. The homo-
geneity and low level of regional endemism of the forest
avifauna throughout the western Palaearctic have been
explained as a consequence of these periodic shifts, which
repeatedly mixed populations, thus preventing allopatric
differentiation at the species level25. Although species dif-
ferences are not apparent, it has recently been shown that
differences at the intraspecific level can be detected. From
their ‘genetic signals’ (i.e. their phylogeography, which tests
whether current genetic patterns among species were
shaped by common responses to unique historical events),
Taberlet et al.26 have revealed traces of local population
differentiation for several animal (mammals, amphibians
and insects; using mtDNA) and plant species (using chloro-
plast DNA), which are postulated to have occurred during
489
REVIEWS

Fig. 1. History of bird families in the Tertiary (65–1.6 Mya) reconstructed from European fossils dating from that period. The central column is divided into the
epochs of the Tertiary. The number of nonpasserine families found in European deposits in each epoch is indicated in bold text. The number of families that
became extinct (stippled areas) tends to decrease during the Tertiary. Arrows indicate the regions where families that occurred in Europe (encircled area) at
different epochs of the Tertiary, but which are no longer present, lived († = extinct; numbers of extinct families given in plain text) or still live after they disappeared
from Europe. Note the explosive radiation of families in the Lower (or Early) Eocene (~55 Mya). Families that were (or are still) present in the Neotropics mostly
disappeared from Europe during the Paleogene (Eocene to Upper Oligocene, 55–25 Mya), and those that were (or are still) present in other regions of the world
disappeared from Europe later in the Neogene (25–1.6 Mya). An explanation still has to be proposed for this difference.

glacial times in southern refugia. A strong phylogeographical
structuring, recently demonstrated for both North American
and European bird species27, strongly suggests that similar
processes of intraspecific differentiation also occurred in
birds.
Second, there were large-scale north–south cyclic mi-
grations. For example, species that currently only occur in
the far north, such as the snowy owl Nyctea scandiaca (a
species common in fossil deposits of southern Europe), were
present in the Mediterranean only during glacial periods24.
These large-scale cyclic migrations also occurred in sea-
birds that bred along the coast of southern Europe and in
the Mediterranean during glacial episodes28. Remains of
the great auk Pinguinus impennis (extinct in Iceland by
1844) have been found in several Pleistocene and Holocene
deposits along the coasts of Iberia, France and Italy (as far
east as Calabria)29.
Effects of Pleistocene climatic events on western
Palaearctic avifauna
Compared with that of the two other large forested re-
gions of the Northern Hemisphere (eastern North America
and eastern Asia), the bird fauna (and particularly forest
avifauna) of the western Palaearctic is relatively species
poor. Controlling as much as possible for the size of areas,
Mönkkönen and Viro30 have shown that the western
Palaearctic (including North Africa) has 50% fewer forest-
associated bird species than eastern Asia and 40% fewer
than eastern North America. Europe is similar in size to
China (~9 500 000 km2, including Manchuria), but European
species richness (about 500 bird spp.) is only about half
that reported for China. In the western Palaearctic, less
than half of the terrestrial avifauna is associated with for-
est compared with two-thirds in eastern North America
and eastern Asia30.
Similar patterns of species impoverishment have been
found in freshwater fish between North America and the
western Palaearctic31 and in tree species between all three
regions of the Northern Hemisphere13. Currently, there
are three times as many tree species in mesic forests of
eastern Asia (729 species) than in eastern North America
(253 species) and six times more than in European forests
(124 species) (Ref. 13). The low taxonomic diversity of tree
species in European forests has been attributed to the lim-
ited potential refugia available to them during glacial epi-
sodes32, resulting in high extinction rates during Pleisto-
cene glaciations33.
Large and repeated habitat changes during the Pleisto-
cene have undoubtedly accelerated the extinction rates of
birds of the western Palaearctic24, just as they did for plants
and large mammals in North America34. Comparative analy-
ses using molecular phylogenies of 11 lineages of North
American songbirds also indicate a general decrease in
rates of species diversification during the Pleistocene34.
However, molecular evolutionary trees cannot distinguish
between reduced speciation rates (e.g. as a consequence

490 TREE vol. 13, no. 12 December 1998


of progressive niche filling through evolutionary time35)
and higher extinction rates as the cause of net changes in
diversification rates.
Assuming similar rates of speciation over the whole
Northern Hemisphere during the Pleistocene, the relative
impoverishment of the western Palaearctic bird fauna (com-
pared with the two other regions of the Northern Hemi-
sphere) could be caused by higher extinction rates in the
western Palaearctic, because there was no connection with
the tropical biomes further south21. In contrast to those of
the western Palaearctic, the extant bird faunas of both
northeastern North America and eastern Asia share many
species in common with southern tropical regions. For ex-
ample, 76% of passerine species breeding in North America
have either conspecific populations or congeneric species
that breed in the Neotropics36. In eastern Asia, two orders
and nine families are tropical, whereas Afro-tropical influ-
ence in the fauna of the western Palaearctic is negligible25.
The most likely explanation for these differences is that
the biota of both eastern North America and eastern Asia re-
mained connected to the tropics over the whole Tertiary–
Quaternary, making it possible for a continuous interchange
between tropical and temperate avifaunas13,21. In contrast,
the massive east–west oriented barriers (mountain ranges,
seas and large desert belts) ‘trapped’ biota of the western
Palaearctic in their southern refugia during glacial periods,
preventing them from finding refuge in tropical regions fur-
ther south, and preventing tropical species from colonizing
northern regions. Since the Pliocene, the Sahara desert has
never been sufficiently forested to provide a dispersal corri-
dor for forest birds between the Afro-tropics and Eurasia37.
The origin of modern species
There has been a long-standing controversy about the
spatiotemporal context in which extant bird species evolved.
Wetmore’s38 contention that most modern species date from
the Pliocene has been challenged by Selander39 and others
who have argued that Pleistocene glacial cycles have been
conducive to speciation in most groups of birds, a tenet
strongly supported by many studies concerned with Pleis-
tocene refuge theory40,41. However, recent studies using
molecular systematics (based on mtDNA) provide evidence
that many species are much more ancient34,40,42 (Box 2). In
what are presumed to be the most recently evolved song-
birds, the average percentage divergence of mtDNA is higher
than that expected for a Pleistocene differentiation. For ex-
ample, according to conventional mtDNA clock calibration,
only 11 of the 35 pairs (31%) of North American sister-species
date to a separation during the Quaternary40 (1.6 Mya), the
remaining having separated previously. Similar divergence
times to those reported for North American birds have
been estimated from mtDNA differentiation for birds of the
western Palaearctic27. The somewhat controversial DNA–
DNA hybridization technique gives divergence times of a
similar order of magnitude35. For example, it was recently
shown that the Sylvia complex, a group of 17 closely related
species of Mediterranean warblers, started to differentiate
as far back as 6.3–6.8 Mya (Ref. 46).
Does this mean that Pleistocene environmental changes
had little impact on extant avian diversity? It seems a para-
dox that speciation and extinction rates did not increase
significantly in the face of the extreme environmental in-
stability of the Pleistocene. Diversification rates are re-
ported to have actually decreased through the Pleistocene
compared with the Pliocene34, and Klicka and Zink40 re-
cently concluded that the contribution of recent ice ages in
the differentiation of bird species is a ‘failed paradigm’.
TREE vol. 13, no. 12 December 1998
REVIEWS
Box 2. Dating species from mitochondrial DNA
Mitochondrial DNA (mtDNA), especially the encoded cytochrome b gene, provides
useful information to reconstruct phylogenies and date divergence events in
many animals, including birds. Nucleotide evolution of mtDNA has been esti-
mated as 2% per million years in geese (Anatidae)43, a figure similar to that pro-
posed for divergences in a complex of Hawaiian honeycreepers (Drepanididae),
based on the geological history of the islands44. A large number of studies in
many groups of birds gave values of a similar order of magnitude, although they
depend on generation times, being lower in species with long generation times
(e.g. cranes, Gruidae). For most congeneric species of birds that have been stud-
ied, mtDNA divergence far exceeds the 2% that would be expected if divergence
took place during the Pleistocene (~1 Mya). For 35 pairs of North American song-
birds postulated to have differentiated during the Pleistocene (in the framework
of the refuge theory), mean values of estimates of mtDNA sequence divergence
are 5.1% with only six pairs having values lower than 2% (Ref. 40). In partridges
of the genus Alectoris, a group of closely related and largely allopatric species
that have for long been thought to be of recent Pleistocene origin, calibration
rates of nucleotide evolution (using mtDNA of cytochrome b) indicate divergence
values falling between 1.7% and 8.6% (Ref. 45). These data indicate that only a
small proportion of extant species differentiated during the Pleistocene.
Large-scale climatic events producing evolutionary changes
in flora and fauna date back to the Early Pliocene20. Events
conducive to evolutionary differentiation in populations
(such as vicariance events or bottlenecks) have occurred
since that time and continued during the Pleistocene26.
Avise and Walker27 have emphasized the importance
of the Pleistocene in differentiation processes that may
have started in the Pliocene. In 63 species from both North
America and Europe they demonstrated that Pleistocene
events promoted substantial microevolutionary genetic
diversification – 37 of these species displayed significant
geographically-oriented phylogroups (i.e. genetically dis-
tinct populations) within matrilineal gene trees. Separ-
ation dates of these phylogroups fell within the Pleistocene
for 28 pairs with the most recent split dating back to about
200 000 BP. Differentiation in glacial refugia during the Pleis-
tocene and subsequent range expansions of two or more
phylogeographical units have been proposed for explaining
these patterns in birds27 (and in other groups of animals
and plants26).
If Pliocene climatic events were as effective as those that
operated during the Pleistocene in producing phylogeo-
graphical differentiation20, then many species that entered
the Quaternary were presumably already separated into dis-
tinctive phylogeographical units, and were themselves likely
candidates for further evolutionary divergence and speci-
ation27. Thus, species-level divergences initiated in the Plio-
cene as a result of climate-induced environmental change
might have been completed during the Pleistocene. If avian
speciation can be considered as a gradual process, by which
species originate from suites of diverging phylogeographi-
cal units over long timespans, rather than a point event in
history, then the role of Pleistocene events in accelerating
speciation processes that had been initiated earlier becomes
more important27. This new insight from molecular studies
could help reconcile the formerly diverging views on the
tempo and mode of speciation events in modern birds.
Prospects
There are still large gaps in our understanding of the
early history of birds, but current progress in the fields of
palaeontology and molecular techniques will help overcome
these. A promising avenue will be to combine the use of
molecular tools (that are likely to gain even higher powers
of resolution) with an understanding of the environmental
events that have driven the geographical structure of
491
REVIEWS
genetic variation across different temporal scales. This will
allow us to correlate phylogenetic patterns with effects such
as distance, geographical barriers and putative refugia at
various epochs of the past.
The interpretation of phylogeographical structure,
already found in many species in North America47 and
Europe27, indicates the timescale of speciation processes.
Molecular analyses of population differentiation are a power-
ful tool for investigating whether consistent geographical
correlates of structured haplotypes are found among spe-
cies in other regions of the Northern Hemisphere. Compara-
tive phylogeography will help to reconstruct the geography
of genetic variation in bird faunas and to date it in relation
to the separate histories and geographical configuration of
the western Palaearctic, North America and Eastern Asia.
Acknowledgements
We are most grateful to Marcel Lambrechts, Doyle McKey,
Michel Raymond, and three referees for their most helpful
comments and suggestions on this article. René Ferris
drew the figures.
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TREE vol. 13, no. 12 December 1998