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Th e A uk 12L (4) :11 5Tl1 60, 2004
MIOCENE SONGBIRDS AND THE COMPOSITION
OF THE EUROPEAN PASSERIFORM AVIFAUNA
ArsnscHr MaNscoro,l GrnAtD MayR,2'a aN» CÉcIrs Mounsn-CHAUvIRÉ3
tlnstitut für Biologie und Zoologie, Freie Unioersitrit Berlin, Kônigin-Luise-Strasse 1-3, D-1-4195 Berlin, Germany;
zSektion Ornithalogie, Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60j25 Frankfurt am Main, Germany; and
3llMR 5125 Paléoenoironnements et Paléobiosphère, Centre des Sciences de la Terre, Unittersité Claude Bernard-Lyon 1
2743 Bouleaard du 11 Nooembre, F-69622 Vlleurbanne Cedex, France
AssrRAcr. -Songbirds (Passeriformes) occur in the fossil record of the Northem Hemisphere
around the early Oligocene. It has recently been suggested that the maior passeriform lineages
diverged in Gondwana in the mid- to late Cretaceous and that the oscines, which include all
extant European songbirds, originated on the Australian continental plate. Suboscines are as-
sumed to have originated in western Condwana. Although there is an abundant fossil record
of songbirds in Europe, few attempts have been made to set those remains in a phylogenetic
context. Our examination of fossil songbirds from three middle Miocene localities in Germany
and France shows that many lack the derived morphology of the hypotarsus that characterizes
extant Eupasseres (a taxon that comprises oscines and suboscines). We assume that these fossil
taxa are outside the crown-group of Eupasseres, which indicates the presence of an ancient
songbird avifauna in the Miocene of Europe, in addition to the few fossil Eupasseres already
described in the literature. Receiaed 31 October 2003, accepted 24 lune 2004.
RpsuurN.-Las aves canoras (Passeriformes) aparecen en el registro fôsil del Hemisferio
Norte alrededor del Oligoceno temprano. Recientemente se ha sugerido que los linajes
principales de Passeriformes se separaron en Gondwana durante el Cretâcico medio a tardio
y que los oscines, que incluyen todas las aves canoras vivientes europeas, se originaron en
la plataforma continental de Australia. Se supone que los suboscines se originaron en el
oeste de Gondwana. Aunque para Europa existe un registro f6sil abundante, se han hecho
pocos intentos de poner estos restos f6siles en un contexto filogenético. Nuestra evaluaci6n
de las aves canoras fôsiles de tres localidades del Mioceno medio ubicadas en Alemania y
Francia muestra que muchas de estas aves no presentan la morfologia derivada dei hipotarso
que caracteriza a los Eupasseres vivientes (un taxôn que incluye los oscines y suboscines).
Suponemos que estos taxa fôsiles estân fuera del grupo terminal Eupasseres, 1o que indica la
existencia de una avifauna de Passeriformes antigua en el Mioceno de Europa, ademâs de los
pocos f6siles de Eupasseres ya descritos en la literatura.
SoNcsrnos oR pASsERINEs (Passeriformes) are
the most species-rich group of birds, compris-
ing more than half of all extant bird species
(Sibley and Monroe 1990). The phylogenetic
relationships between these birds have long been
controversial (see Sibley and Ahlquist 1990 for a
review of the history of classification), but recent
molecular anâlyses have greatly improved our
understanding of passeriform phylogeny. Those
studies have shown that New Zealand wrens
(Acanthisittidae) are the sister group of all other
extant passerines (the Eupasseres, which are fur-
ther divided into suboscines and oscines; Fig. 1;
Barker et al. 2002; Ericson et al. 2002, 2003).
aAddress correspondence to this author. E-mail
gerald.mayr@senckenberg.de
All extant European songbirds belong to the
oscines, which are assumed to have arisen on the
Australian continental plate (Barker et al. 2002).
With the exception of the sister taxa broadbills
(Eurylaimidae) and pittas (Pittidae) (e.9. Ericson
et al. 2003), members of the suboscines are today
restricted to North, Central, and South America.
That pattem of distribution has led to the
assumptions that (1) Passeriformes originated
in the Southern Hemisphere and (2) the sepa-
ration of New Zealand from Gondwana in the
mid-Cretaceous (some 82-85 mya) resulted in
the split into the Acanthisittidae and Eupasseres
lineages (Ericson et al. 2003). The split between
the oscine and suboscine lineages is assumed
to have occurred in the late Cretaceous, when
the South American and Indian tectonic plates
became isolated (Ericson et al. 2003).
1155
11,s6 MaNscoro, Mavn, aup Mounsn-CHauvlnÉ [Auk, Vol. 121
Passeriform birds exhibit a highly derived
and very uniform osteology that facilitates their
distinction from nonpasseriform birds but aggra-
vates recognition of subgroups within the taxon.
For that reasory few attempts have been made
to set fossil songbirds in a phylogenetic context.
However, we discovered that the hypotarsus-a
bony strucfure at the proximal tarsometatarsus
that conducts the tendons of the flexor muscles of
the toes -exhibits a phylogenetically informative
variation within passeriform birds.
We examined hypotarsi of representatives of
all but 5 (for which no skeletons were available)
of the 46 extant "families" of Passeriformes
recognized by Sibley and Monroe (1990)
and of fossil songbirds from three middle-
Miocene localities (all from the mammalian
biostratigraphic level MN 6, about 14-15 mya)
in southern Germany (Petersbuch 39 and
Nôrdlinger Ries) and France (Sansan), which
yielded abundant remains of small passeriform
birds. Surprisingly, many of the well-preserved
fossil hypotarsi we studied differ distinctly from
the pattem typical of crown-group Eupasseres.
Mrrnnral aNo Mrruops
We examined hypotarsi of the following extant
taxa in the collections of the Institut für Biologie
und Zoologie, Freie Universitât Berlin; the Museum
of New Zealand Te Papa Tongarewa, Wellington,
New Zealand; the Natural History Museum London;
Forschungsinstitut Senckenberg, Frankfurt am
Main, Germany; the National Museum of Natural
History, Washington D.C.; and the Museum für
Naturkunde, Berlin (species are listed following the
higher-level passerine sequence of Sibley and Monroe
1990; "Corvidae" were shown to be a paraphyletic
group by Barker et aL.2002 and Ericson et al. 2002):
Acanthisittidae : Acanthisitta chloris, Xenicus gilaizten-
iris. Suboscines: Pittidae: Pitta guajana, P. sordida, P.
granatina. Eurylaimidae (inciuding Phiiepittidae;
see Prum 1993): Smithornis capensis, Corydon
sumatranus, Cymbirhynchus macrorhynchus, P sarisomus
dalhousiae, Philepitta castanea. Tyrannidae: Xolmis
irrupero, Machetornis rixosus, Tyrannus melancholicus,
Megarhynchus pitangua, Pitangus sulphuratus, Phy-
totoma rara, Lipaugus subalaris, Xipholena punicea,
Cephalopterus ornatus, Procnias nudicollis, Rupicola
peruztiana, Pipra erythrocephala, Chiroxiphia caudata,
Manacus manacus. Thamnophiiid ae: Thamnophilus cae-
r ulescens, T. ruficapillus. Furnariidae: F urnarius rufus,
P s eudo col ap t es b oiss onne au t ii, Sy nda cty I a r ufo sup er cili at a,
Dendrocincla cf. fuliginosa, Lepidocolaptes fuscus, Sit-
tasomus griseicapillus. Formicariidae: Formicarius sp.
Rhinocryptidae:. Pteroptochos megapodius. Oscines:
Passeriformes
-1
;- Eupasseres -1
Acanthisittidae Suboscinês Oscines
.-ô,,\ --*<^
(.'-l /(\4
\ >-^ ^- \ - -->J
L)L lL,-fdl v/ ) \
çrnffiÿ - Dfhr{Y}\fdr
Frc. 1. Left hypotarsi of extant (A, B) and fossil (C,
D) passeriform birds in comparison. (A) South Island
Wren (Xenicus gilaioentris, Acanthisittidae). (B) Gamet
Pitta (Pitta granatina, suboscines, Eupasseres); this
pattern corresponds to the derived six-canal pattern
of Eupasseres. (C) Fossil species from the middle
Miocene (Mlikovskÿ 1996) of Petersbuch 39, Germany
(specimen SMF Av 496). (D) Fossil species from the
middle Miocene (Cheneval 2000) of Sansan, France
(specimen MNHN SA 1263c). Images are not to scale.
Abbreviations of tendinal canals: fhl = musculus Jlexor
hallucis longus, ldl = musculus flexor digitorum longus,
tp = musculus flexor perforatus, fpp = musculus flexor
perforans et perforatus; numerals indicate number of
the toe suppiied by the tendon. Phylogeny is based on
recent molecular studies (Barker et al. 2002; Ericson et
al.2002, 2003).
The earliest fossil record of songbirds is
from the Eocene of Australia, though the frag-
mentary fossils do not allow an assignment to
any specific taxon within Passeriformes (Boles
1995a). From the early Miocene of Australia,
crown-group oscines have been reported and
even assigned to distinct extant genera (Boles
'1995b, L997), thus corroborating molecular phy-
logenies that indicate an origin of oscines on the
Australian continental plate. There is no record
of early-Tertiary Passeriformes from Africa, and
only a single fragmentary passeriform humerus
is known from the early Miocene of South
America (Noriega and Chiappe 1993).
The earliest passeriform fossils from Europe
are from the early Oligocene of Germany (Mayr
and Manegold 2004) and France (Roux 2002).
Songbirds have not been identified among the
numerous avian remains from Eocene fossil sites
in Europe and North America (Mourer-Chauviré
L995,Mayr 2000), and it is assumed that they were
still restricted to the Southem Hemisphere by that
time (Olson 1988, Maclean 1990, Cracraft 2001).
October 20041
Climacteridae: Climacteris leucophaea. Menuridae: Men-
u r a n o a a eh o I I an di ae. P tilonorhy nchi d ae : P t il o n o rhy n chu s
oiolace us. Maluridae: Malurus splendens. Meliphagidae:
Myzomela rubrata, Acanthochaera inauris. Pardalotidae:.
P ardalotus substriatus. Petroicidae: Eopsaltria australis.
Irenidae: lrena puella. Orthonychidae: Orthonyx spal'
dingi. Pomatostomidae: Pomatostomus superciliosus.
Laniidae: Lanius collurio, L. excubitor. Vireonidae:
Vireo olittaceus. "Corvidae": Corcorax melanorhamphos,
S truthidea cinere a, P achy cephala rut'iaentris, P. lanioides,
Turnagra capensis, Pica pica, Corztus monedula, Seleucidis
melanoleuca, Paradisaea rubra, Artamus pelewensis, Ori-
olus oriolus, Dicrurus adsimilis, Grallina cyanoleuca,
Aegithina tiphia, F alculea p alliata. Picathartidae: Pica-
thartes oreas. Bombycillidae: Bombycilla garrulus.
Cinclidae: Cinclus cinclus. Muscicapidae: Turdus mer-
ula, T. philomelos, Zoothera citrina, Ficedula hypoleuca,
Erithacus rubecula, Luscinia megarhynchus. Sturnidae:
Cosmopsarus regius, Sturnus aulgaris, Mino anais,
Gr acula religiosa, Buphagus ery throrhynchus, D umetella
carolinensis, Mimus poly glollos. Sittidae: Sitta europae a.
Certhiidae: Certhia sp., Troglodytes troglodytes. Paridae:
Remiz p endulinus, P arus caeruleus, P. maj or. Aegithalidae:
Aegithalos caudatus, Ae. concinnus. Hirundinidae:
Hirundo rustica, Delichon urbica. Reguiidae: Regulus
regulus. Pycnonotidae: Pycnonotus jocosus, Chlorocichla
falkensteini, Hypsipetes madagas cariensis, N e oLest es tor'
quatus. Cisticolidae: Cisticola galactotes. Zosteropidae:
Zosterops erythropleurus, Z. palpebrosa. Sylviidae: Acro-
cephalus schoenobaenus, Phylloscopus trochilus, Garrulax
leucolophus, Stachyris polio cephala, Tur doides s quamiceps,
Leiothrix argentauris, Sylztia atricapilla, Hippolais ic-
terina. Alaudidae: Galerida cristata, Alauda arttensis,
Eremophila alpestris. Nectariniidae: Dicaeum hirun-
dinaceum, Anthreptes singalensis, Nectarinia asiatica.
Passeridae: Passer domesticus, Motacilla alba, Anthus
pratensis, Prunella, Quelea quelea. Fringillidae: Fringilla
coelebs, F. montifringilla, Carduelis carduelis, C. spinus, C.
chloris, Coccothraus t es coccothr austes, Emberiza citrinella,
Seiurus aurocapillus, Tangara nigroztiridis, T. uassorii,
Diglossa lafresnayii, lcterus dominicensis, Agelaius hu'
mer alis, Quiscalus niger.
The fossil specimens are deposited in the
following institutions: Muséum National d'Histoire
Naturelle, Paris, France (MNHN); Forschungsinstitut
Senckenberg, Frankfurt, Germany (SMF); and
Staatliches Museum für Naturkunde, Stuttgart,
Germany (SMNS). Anatomical terminology follows
Baumel and Witmer (1993).
We focused on the hypotarsus because we found
that element to be especially informative in regard to
characterization of some maior ciades of passeriform
birds (see below). Some of the examined specimens
consist of complete bones with the highly characteris-
tic distal end of the passeriform tarsometatarsus (e.9.
trochleae of the second, third, and fourth toe arranged
in a line in distal aspect; and trochleae of the second
and fourth toe very narrow). A derived feature on the
1157
proximal end that confirms passeriform affinities of
the fossils is the presence of an ossified retinaculum
extensorium; the shaft of the tarsometatarsus further
shows a marked crista plantaris lateralis, which occurs
only in songbirds and cuckoos (Cuculidae).
The pattern of hypotarsal canals in the fossils
reflects the original condition of the bone and is not
the result of the breakage of bony ridges (Fig. 2). The
fossils are certainly from adult birds, because the
tarsal cap is completely fused with the metatarsals
and the proximal end of the bone does not show the
blurred surface characteristic of juvenile birds. Other
skeletal elements of passeriform birds in the material
(carpometacarpi, humeri) cannot reliably be assigned
to a given type of tarsometatarsus and thus are not
considered here.
REsurrs
The hypotarsus of the Acanthisittidae, sister
taxon of the Eupasseres, bears only two ossified
canals for the deep flexor tendons of the toes
(fhl and fdl in Fig. 1B); whereas the superficial
flexor tendons (fp24 and fpp2-3 in Fig. 18) run
through bony grooves on its plantar surface
Frc. 2. Scanning electron microscope photographs
of the proximal end of a left tarsometatarsus from
the middle Miocene of Sansan, France (specimen
MNHH SA 1263c). (A) Proximal view on hypotarsus.
(B) Dorsal view. (C) Plantar view. (D) Medial view.
Abbreviations: cpl = crista plantaris lateralis, fhl = ossi-
fied canai for tendon of musculus lTexor hallucis longus,
re = ossified retinaculum extensorium (broken). Arrows
indicate the bony ridges that border the canal for the
tendon of musculus flexor digitorum lozgrzs (note that
these do not show any sign of breakage).
Miocene Songbirds
1158 MaNrcom, Mavn, ewo Mounnx-CuauvrnÉ [Auk, Vol. 121
Miocoraus larteti from the middte Miocene of
Sansan (specimens MNHN SA 1267, MNHN
SA 1289, MNHN SA 1491). Surprisingly, the
canals for the superficial tendons fp3-4 and
fpp3 are plantarly open in all of the 13 well-
preserved fossil hypotarsi from the German
localities Petersbuch (specimens SMF Av 487-
496) and Nôrdlinger Ries (specimens SMNS
86822, SMNS 86825, SMNS 86826) (Fig. 1C).
The canals for those tendons are also plantarly
open in 8 of the 26 well-preserved specimens
from Sansan (specimens MNHN 5A1259-1262,
MNHN SA 1263a, b, c, d) (Figs. 1D and 2), in
which only the lateral canal for the deep flexor
tendons (fhl) is completely ossified, a condi-
tion we found in the South American Lesser
Woodcreeper (Lepidocolap t es fus cus, Furnariidae,
Suboscines; Fig. 3L) but not in the closely
related Dendrocincla cf. fuliginosa and Siftasomis
griseicapillus (see Iresteàt et al. 2002).
DrscussroN
Outgroup comparisons with any extant non-
passeriform avian taxon show the unique six-
canal pattern to be derived within passeriform
birds. A six-canal pattem is present in almost
all members of suboscines and oscines, includ-
ing basal taxa (e.9. Barker et al.2002), and thus
almost certainly was also present in the last
common ancestor of crown-group Eupasseres. It
is the first morphological apomorphy of crown-
group Eupasseres identified so far and supports
the molecular results conceming the sister-
group relationship between Acanthisittidae and
Eupasseres.
]udging from current phylogenies, the pres-
ence of plantarly open canals for the super-
ficial tendons fp3-4 and fpp3 in a few extant
Eupasseres is most likely a reversal to the primi-
tive condition in those unrelated taxa nested
deeply within crown-group Eupasseres (e.g.
Barker et al. 2002). Apart from the suboscines
Xipholena punicea and Lepidocolaptes fuscus, all
the birds in question are ground-dwelling non-
perching birds. Although it thus appears as if
there was a functional correlation between that
way of living and ossification of the hypotarsus,
other ground-dwelling songbirds, such as pittas
(Pittidae; Fig. 18) and lyrebirds (Menuridae),
exhibit the Eupasseres' typical six-canal pattem.
The considerable variation in the hypotarsi
of the fossil taxa (Fig. 1C, D) indicates a high
(Fig. 1A). In crown-group Eupasseres, in con-
trast, the flexor tendons typically run through
six ossified canals arranged in a unique complex
pattern found in no other living or fossil avian
taxon (Figs. 1B and 3). In most Eupasseres, the
canals for the superficial tendons are completely
ossified (some canals may be fused, as in some
Turdidae and Zosteropidae; see Figs. 3I, ]); very
rarely, the canal for fp3-4 is plantarly open (e.g.
in all Eurylaimidae investigated; Fig. 3K).
Among the numerous passeriform taxa
we examined, only in the South American
Pteroptochos megapodius (suboscines, Rhino-
cryptidae) and Xipholena punicea (suboscines,
Tyrannidae), the African Picathartes oreas
(oscines, Picathartidae), and the Palaearctic
Cinclus cinclus (oscines, Cinclidae) are the
canals plantarly open for both the superficial
tendons fp3-4 and fpp3 (Figs. 18 and 3M-O).
That condition was also mentioned by Feduccia
and Olson (1982) for the rhinocryptid genera
Scelorchilus and Myornis and illustrated by Rich
et al. (1985) for the two species of Australian
Atrichor nis (oscines, Menuridae).
We found the derived "six-canal pattern"
of crown-group Eupasseres to be present in
,a.î
n(9 «-{l
.\"9,,
(1 )
Y^ -'
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BY9
(() \
Yn(ÿ-
FQ9
(û,
I:00à
),râ/
K*
ra-î
rer-
G.rn
\U§
L-
Frc. 3. Selected hypotarsi of extant suboscines (A-D,
K-M) and oscines (E-I, N-O). Note that the excep-
tions from the Eupasseres type are over-represented
in the figure. (A) Tyrannus melancholicus (Tyrannidae).
(B) Phytotoma rara (Tyrannid,ae). (C) Thamnophilus
caerulescens (Thamnophilidae). (D) Furnarius rufus
(Furnariidae). (E) Dicrurus adsimilis (Dicruridae). (F)
Paradisaea rubra (Paradisaeidae). (C) Falculea palliata
(Vangidae). (H) Hirundo rustica (IJir:undinidae). (!
Turdus merula (Turdidae). (!) Zosterops erythropleurus
(Zosieropidae). (K) Cymbirhynchus macrorhynchus
(Eurylaimidae). (L) Lepidocolap tes fuscus (Furnariidae).
(M) Pteroptochos megapodius (Rhinocryptidae). (N)
Picathartes oreas (Picathartidae). (O) Cinclus cinclus
(Cinciidae). Images are not to scale.
K1)\
"\efl
rG1 )
\ooZ
]X a/
lUo/
H\J ar) C-r)
Rf, ,WBT-
r;) a.:t) {)
W \ ry' lo;[
M N \l o tL
October 20041
taxonomic diversity and a radiation of at least
one extinct passeriform taxon (there is no evi-
dence that the fossil birds in question are mem-
bers of a monophyletic group). If the absence
of the derived Eupasseres pattern in the fossil
specimens was also a secondary reversal to the
primitive conditiory it raises the question of
why taxa without the six-canal pattem are the
exception among extant passerines, whereas
they appear to be abundant and diverse in the
middle Miocene. Because of the geological age of
the specimens, we consider it likely that absence
of the six-canal pattern in the fossil hypotarsi is
indeed primitive and indicates that the taxa are
outside the crown-group Eupasseres.
The as-yet-unnamed early-Oligocene song-
bird from Germany described by Mayr and
Manegold (2004) is too poorly preserved for
detailed comparisons. However, a plesiomor-
phic morphology of the coracoid also suggests
that it is outside the crown-group Eupasseres.
The hypotarsus morphology of that specimen is
unknown. The early-Oligocene songbird from
France mentioned by Roux (2002) has not yet
been described.
Few late Oligocene or early to middle Miocene
songbirds have been described from Europe
so far, and all were considered to be within
the Eupasseres-representatives either of the
suboscines (Ballmann 1969, Mourer-Chauviré
1995, Cheneval 2000) or of the oscines (Mourer-
Chauviré et al.-1.989, Cheneval 2000). Although
the hypotarsus of most fossil specimens is either
unknown or too poorly preserved to allow rec-
ognition of the canal pattern, some identifica-
tions of Eupasseres in the Miocene of Europe
are certainly correct-for example, Miocorous
larteti (see above) or the as-yet-unnamed taxon
of Eurylaimidae reported by Ballmann (1969)
from the early Miocene of Germany. We thus
do not question the existence of Eupasseres
in the Miocene of Europe (see also Steadman
[1981] concerning the presence of Eupasseres
in the Miocene of North America). However,
our study indicates for the first time that fossil
songbirds outside the crown-group Eupasseres
(i.e. taxa other than suboscines and oscines)
may have been present in the Miocene of
Europe. Our study focuses on fossils from one
period, the middle Miocene (MN 6). Further
studies of older (Oligocene or early-Miocene)
songbird remains are needed for learning more
1159
about the temporal distribution of these taxa.
With accumulation of more fossil material, it
may also be possible to assign other skeletal
elements to the tarsometatarsi, which may
provide additional clues on the phylogenetic
position of these fossil songbirds.
AcrNowrEocurNrs
We thank R. Bôttcher (SMNS) for the ioan of fos-
sil specimens and J. Cooper (The Natural History
Museum, London) for information on the hypotar-
sus of some extant songbirds. We further thank D.
Steadman and an anonymous reviewer for comments
on the manuscript. A.M. is supported by Deutsche
Forschungsgemeinschaft (GK 503).
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