GeolCarp_Vol67_No5_463

GEOLOGICA CARPATHICA

, OCTOBER 2016, 67, 5, 463 – 469

doi: 10.1515/geoca-2016-0029

www.geologicacarpathica.com

A tiny short-legged bird from the early Oligocene of Poland

ZBIGNIEW M. BOCHENSKI

, TERESA TOMEK

and EWA SWIDNICKA

Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, 31-016 Kraków, Poland;

bochenski@isez.pan.krakow.pl, tomek@isez.pan.krakow.pl

Department of Palaeozoology, Chair of Evolutionary Biology and Ecology, University of Wrocław, Sienkiewicza 21,

50-335 Wrocław, Poland; gama@biol.uni.wroc.pl

(Manuscript received February 3, 2016; accepted in revised form September 22, 2016)

Abstract: We describe an articulated partial leg of an Oligocene bird. It is one of the smallest avian fossils ever

recorded. Its slender and exceptionally short tarsometatarsus, hallux as long as the tarsometatarsus and stout

moderately curved claws agree with stem-group Apodidae (swifts), stem-group Trochilidae (hummingbirds), and

stem-group Upupidae/Phoeniculidae (hoopoes/woodhoopoes). Unfortunately, due to the poor preservation of

the specimen its more precise affinities remain unresolved. The specimen differs in many details from all other tiny

Palaeogene birds and therefore most probably it represents a new taxon but it is too fragmentary to describe it. It is just

the twelfth avian fossil from the Oligocene marine deposits of the Outer Carpathians and Central Palaeogene Basin

— a huge area that covers south-eastern Poland, north-eastern Czech Republic and northern Slovakia — and therefore

it adds to our very limited knowledge on the avifauna of that region. The remains of land birds from Jamna Dolna and

other sites of the region can be attributed to the general sea level fall at that time, which led to limitation of

the connection with the open ocean and resulted in many shallow shoals, temporary islands and exposed dry land areas

along the coast.

Keywords: Fossil birds, Menilite shales, Carpathian flysch, Palaeogene.

Introduction

The Menilite beds of the Carpathian flysch zone that are

found in north-eastern Czech Republic and south-eastern

Poland, and the Central Palaeogene basin of northern Slovakia

are extremely rich in Oligocene fish fossils. Only in

south-eastern Poland, many thousand of fish fossils were

recovered from more than 200 outcrops between the mid

1970s and mid 1990s (Kotlarczyk et al. 2006). However,

animal fossils other than fish are extremely rare. So far only

eleven avian specimens have been described from the marine

deposits of the Outer Carpathians and Central Palaeogene

Basin: two procellariiforms (Gregorová 2006; Elzanowski et

al. 2012), one galliform (Tomek et al. 2014), one humming-

bird (Bochenski & Bochenski 2008), one putative upupiform

(Kundrát et al. 2015), one piciform (Mayr & Gregorová

2012), four passerines (Bochenski et al. 2011, 2013b,

2014a, b) and Aves indet. (Bochenski et al. 2010). Here, we

describe an extremely small, incomplete avian leg that

resembles those of stem-group apodid, trochilid or upupid

birds.

The apodiform birds are nowadays classified within the

order Caprimulgiformes (del Hoyo & Collar 2014); they

include three living families: the globally distributed true

swifts (Apodidae), the Southeast Asian tree swifts

(Hemiprocnidae), and the hummingbirds (Trochilidae),

which nowadays occur exclusively in the New World (del

Hoyo et al. 1999). The Palaeogene fossil record of swifts is

relatively rich but the systematic position of many fossil taxa

is often unclear and widely disputed. According to Mayr

(2009) there are two extinct families: Eocypselidae (Eocy

pselus) – a stem group taxon of swifts from the early Eocene

of the UK and Denmark (Harrison 1984; Dyke et al. 2004),

and Aegialornithidae (Aegialornis and Primapus) with

several species from the late Eocene to early Oligocene of

France and the UK (Harrison & Walker 1975; Mourer-

Chauviré 1988). The earliest stem group members of the

Apodidae belong to the genus Scaniacypselus from the early

Eocene of Europe; other representatives include Procypseloides

from the late Oligocene of France and Collocalia from the

late Oligocene/early Miocene of Australia (Mayr 2009).

The late Eocene Cypseloides mourerchauvireae from France

described by Mlíkovsky (1989) in Apodidae is believed to be

a junior synonym of Aegialornis gallicus (Aegialornithidae)

(Mayr 2003, 2009). The earliest Trochilidae include several

genera: Jungornis (described by Karhu (1988) in Jungorni-

thidae), Ar

gornis, Parargornis, Cypselavus (all three classi-

fied by Mourer-Chauviré (2006) into Cypselavidae) and

Eurotrochilus Mayr, 2004. Hemiprocnidae have no Palaeo-

gene fossil record (Mayr 2009). In North America, the Palaeo-

gene fossil swifts include Eocypselus rowei described from

the Eocene Green River Formation (Ksepka et al. 2013).

The upupiform birds are nowadays classified within the order

Bucerotiformes which includes, among others, the African

and Eurasian hoopoes (Upupidae) and the African

woodhoopoes (Phoeniculidae) (del Hoyo & Collar 2014).

The stem-group upupids are known from the Eocene deposits

of the United Kingdom (London Clay) and Germany (Messel

and the Geisel Valley) (Mayr 1998, 2000, 2006). Three

species of the extinct family Messelirrisoridae are among the

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, 2016, 67, 5, 463 – 469

most abundant small to tiny birds in Messel; they are

distinguished by their long beak, short tarsometatarsus and

very long hallux (Mayr 2009).

Material and methods

The osteological terminology used here follows that of

Baumel & Witmer (1993). Dimensions are given in milli-

metres and refer to the greatest length along the longitudinal

axis of the bone. The very small size of the specimen excludes

most avian orders. The fossil was compared with skeletal

specimens of extant nightjars and allies, hummingbirds and

swifts, hoopoes, coliiforms and passerines from the osteo lo-

gical collection of the Institute of Systematics and Evolution

of Animals, Kraków, Poland, and Palaeogene species of

the afore-mentioned taxa described in the literature (Peters

1985; Karhu 1988; Mourer-Chauviré 1988; Mayr 2003a,

2004, 2007, 2009, 2010, 2015; Mourer-Chauviré & Sigé

2006; Louchart et al. 2008

;

Mayr & Micklich 2010; Ksepka

et al. 2013).

The fossil was found at the former village of Jamna Dolna

(geographical coordinates of Jamna Dolna: 49

37’45.0’’ N,

34’08.0’’ E), situated about 8 km south-east of the village

of Bircza, Podkarpackie Voivodeship, SE Poland (Fig. 1).

It is a natural exposure of the Menilite strata with more

than 15-m thick deposits in the high escarpment of the Jam-

ninka stream, a right tributary of the Wiar River. In the

geologi cal literature (Jerzmańska 1967a, b, 1968; Jerzmańska

& Kotlarczyk 1968; Kotlarczyk et al. 2006), the exposure is

known as Jamna Dolna 1, to distinguish it from other smaller

outcrops located in the same area. The specimen consists of

one slab (Figs. 2, 3) with imprints of a partial left avian leg.

It was found by Albin Jamróz who passed it to the Institute of

Systematics and Evolution of Animals PAS, Kraków, Poland

where it is housed (ISEA AF/JAM1). ISEA AF/JAM1 is pre-

served on the surface of soft, light brown siliceous marly

shale, collected from the horizon (Unit C, probably layer

C-4) of the Kotów Chert Member, the lower part of the

Menilite Formation of the Skole Unit in the Outer Carpa-

thians. The specimen was found within the ichthyofaunal

assemblage of the IPM-1C Subzone (according to Kotlarczyk

et al. 2006, fig. 20) that includes such fossil fish as Aeoliscus

heinrichi (Heckel, 1850), Anenchelum glarisianum Blainville,

1818, Capros rhenanus (Weiler, 1920), Oligophus moravicus

(Paucă 1931), Zenopsis clarus Daniltshenko, 1960, and other

taxa, which are characteristic of the IPM1 Zone (Kotlarczyk

et al. 2006, p. 65)

The description of fish taxa can be found

in Jerzmańska (1968) and Świdnicki (1986). The fossili-

ferous horizon is dated to the Early Oligocene (Rupelian,

approximately 32.5 m.y.a.) and correlated with the calcareous

nannoplankton of the NP22 biozone by Martini (1971) (see

Berggren et al. 1995; Kotlarczyk et al. 2006).

Description and comparison

As it is often the case in other bird fossils from the Oligo-

cene of Poland (Bochenski & Bochenski 2008; Bochenski et

al. 2010, 2011, 2013a, b, 2014a, b), particular elements in

ISEA AF/JAM1 are broken longitudinally and preserved as

imprints partly lined with remnants of bone. As a result,

a mixture of an imprint and the inner side of a bone rather

than its external surface is visible, which hinders compa-

risons with fossil and extant specimens. Better preserved

specimens are found very rarely in Poland (Elzanowski et al.

2012; Tomek et al. 2014).

Measurements (maximum length in mm). Tibiotarsus as

preserved, 13.8; tarsometatarsus, 6.3; os metatarsale I, 1.4;

hallux: proximal phalanx, 4.1; hallux: claw, 2.1; first phalanx

of digit II, ~1.6; second phalanx of digit II, ~2.9; claw of

digit II, 2.1; first phalanx of digit III, ~1.9; second phalanx of

digit III, ~2.1; third phalanx of digit III, 2.4; claw of digit III,

2.1; first phalanx of digit IV, ~1.5; second phalanx of digit IV,

1.4; third phalanx of digit IV, 1.2 (fourth phalanx of digit IV

and claw of digit IV could not be measured because they are

partly under the hallux claw).

Tibiotarsus. The tibiotarsus is visible in lateral view. Its

proximal part is missing, a fragment of the condylus lateralis

is visible but it is too poorly preserved to allow meaningful

comparisons. A short fragment of the fibula is also

preserved.

Tarsometatarsus. The tarsometatarsus is visible in lateral

view but the surface of the bone is missing and only its inner

Fig. 1. The location of the village of Jamna Dolna (asterisk)

in south-eastern Poland, where the specimen ISEA AF/JAM1 was

found.

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part can be seen. The bone is relatively slender and about half

the length of the tibiotarsus. With the length of just 6.3 mm it

is among the shortest fossil tarsometatarsi ever recorded; it is

by far shorter than that in any extant and extinct Passeri-

formes, Coliiformes or Caprimulgidae. In the Palaeogene,

there were three groups of birds with representatives that had

equally small tarsometatarsi: swifts, hummingbirds and hoo-

poes. ISEA AF/JAM1 is most similar in length and slender-

ness to the early Oligocene Eurotrochilus inexpectatus

(Trochilidae) (6.4–6.7 mm, see Mayr 2004, 2007; Louchart

et al. 2008) and Eocene Scaniacypselus szarskii (Apodidae)

(5.5–5.9 mm, see Peters 1985; Mayr 2015), whereas among

Palaeogene stem lineage upupiforms its size resembles most

small specimens of Messelirrisor parvus (≥7.2 mm, Mayr

1998) and SNMZ 27188 (7.1 mm, Kundrát et al. 2015).

No details of hypotarsal canals and/or furrows nor the exact

arrangement of the trochlea metatarsi II, III and IV are

visible.

Regarding the tibiotarsus / tarsometatarsus proportion,

ISEA AF/JAM1 is most similar to the Palaeogene Scania

cypselus szarskii (Apodidae), Eurotrochilus inexpectatus

(Trochilidae) and species of the genus Messelirrisor (Messe-

lirrisoridae) (Table 1). Also many extant hummingbirds show

similar proportion, contrary to extant swifts that have rela-

tively shorter tarsometatarsus, and extant hoopoes whose

tarsometatarsus is relatively longer. Passerines — even such

short-legged species as swallows — have proportionally

longer tarsometatarsus. Similar results based on different set

of taxa are obtained when the measurements of ISEA AF/

JAM1 are plotted on a log diagram with other groups of

birds: ISEA AF/JAM1 is between the clusters of Messelirrisor

and extant hummingbirds, well apart from other extinct and

extant taxa (Kundrát et al. 2015, fig. 9).

Toes. The foot has anisodactyl arrangement of toes, with

digits II, III, and IV directed forward and digit I (hallux)

directed backward. The phalangeal formula is 2–3–4–5. As

in Eurotrochilus inexpectatus, (Trochilidae), Scaniacypselus

szarskii (Apodidae) and stem-group upupiforms including

Messelirrisor spp. and SNMZ 27188, all digits are relatively

long although this is especially evident in the hallux which in

ISEA AF/JAM1 is similar in length to the tarsometatarsus

and in the afore-mentioned taxa just a little shorter (Peters

1985; Mayr 1998; Kundrát et al. 2015); in extant Apodidae

and Hemiprocnidae as well as all Passeriformes, Coliiformes

and Caprimulgidae the hallux is less than half the length of

the tarsometatarsus. As in Eurotrochilus inexpectatus, Scania

cypselus szarskii and also the putative upupiform SNMZ

27188 from Slovakia, the proximal phalanx of the hallux is

clearly more than half the length of the tarsometatarsus

(Peters 1985; Mayr 2004, fig. 2; Kundrát et al. 2015), whereas

in Palaeogene representatives of other Apodidae (Eocypselus

Fig. 2. Specimen ISEA AF/JAM1 of an avian foot from south-

eastern Poland, Jamna Dolna 1, early Oligocene, ca. 32.5 Mya.

a — Slab; b — Interpretative drawing of the slab.

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vincenti, Eocypselus rowei) and Messelirrisoridae (Messe

lirrisor spp.) it is definitely shorter (Mayr 2010; Ksepka et al.

2013, fig. 1). As in extant hummingbirds (Mayr2004, p. 863),

the long hallux attaches to the tarsometatarsus approximately

at the beginning of its distal third, whereas in Scaniacypselus

szarskii (Apodidae) it attaches to the tarsometatarsus at its

mid-length (Peters 1985, p. 153). As in Eurotrochilus

inexpec tatus and the putative upupiform SNMZ 27188 from

Slovakia but contrary to the modern common swift Apus

apus, the os metacarpale I exhibits an elongated shaft (Mayr

2004, fig. 2; Kundrát et al. 2015); other details are too poorly

preserved for a meaningful comparison. On digits II and III

the penultimate phalanx is the longest (its length in digit IV

is unknown). Similar to Eurotrochilus inexpectatus (Trochi-

lidae), Messelirrisor spp. and SNMZ 27188 (both upupi-

forms), the proximal phalanges of digits II, III and IV are

relatively long; in many extant Apodidae (e.g., Apus, Aero

nantes, Collocalia, Cypsiurus, Panyptila) they are extremely

short but in the Eocene Eocypselus vincenti and

Scaniacypselus szarskii they were not so much abbreviated

(Peters 1985; Mayr 2010, 2015). The claws are robust, mode-

rately curved and their tubercula flexoria are rounded and

rather well-developed; in extant swifts tubercula flexoria are

elongated proximo-distally.

Discussion

It seems that there are just two avian orders — Caprimulgi-

formes and Bucerotiformes — with some representatives

that show a combination of characters observed on ISEA AF/

JAM1. The extremely small size of the specimen, very short

and slender tarsometatarsus, hallux as long as the tarso-

metatarsus, proximal phalanges of all digits elongated

(extremely so in the hallux), and stout, rather massive, mode-

rately curved claws with well-developed tubercula flexoria

make the specimen look similar to a stem taxon of either

swifts, hummingbirds or hoopoes/woodhoopoes. Other avian

Fig. 3. Enlarged fragment of Figure 2a. Abbreviations: d I, pp — digit I, proximal phalanx; d I, up — digit I, ungual phalanx;

d II, p1 — digit II, phalanx 1; d II, p2 — digit II, phalanx 2; d II, up — digit II, ungual phalanx; d III, p1 — digit III, phalanx 1;

d III, p2 — digit III, phalanx 2; d III, p3 — digit III, phalanx 3; d III, up — digit III, ungual phalanx; d IV, p1 — digit IV, phalanx 1;

d IV, p2 — digit IV, phalanx 2; d IV, p3 — digit IV, phalanx 3; d IV, p4 — digit IV, phalanx 4; d IV, up — digit IV, ungual phalanx.

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higher-level taxa including Coliiformes and Passeriformes

can be excluded. Unfortunately, due to the poor preservation

of the specimen and its incompleteness its more precise affi-

nities remain unresolved. The specimen differs in many

details from all other tiny Palaeogene swifts, hummingbirds

and hoopoes and therefore most probably it represents a new

taxon but in our opinion it is too fragmentary to describe it.

Nevertheless, it is a valuable addition to our knowledge of

Palaeogene birds from the Carpathians — a large region that

has yielded only a handful of avian specimens so far.

Most Palaeogene swifts, hummingbirds and also hoopoes

are represented by the wing and/or pectoral girdle bones.

Consequently, the taxonomies of the groups are largely based

on those elements. The leg bones are only seldom preserved

and even rarer are the pedal digits described above. This

study, based on a specimen with known lengths of particular

phalanges, is a small step forward to fill the gap in our

knowledge.

Although there is no way to know what the rest of our

specimen looked like, the relative proportions of leg and foot

Table 1: Total lengths and ratios of two leg bones of chosen extant and extinct taxa with short tarsometatarsus. Measurements are given in

millimetres. A dagger (†) indicates extinct taxa, and an asterisk (*) arithmetic mean of left and right bones. Measurements after:

Ksepka et

al. (2013);

Mayr (2010);

Peters (1985);

Mayr (2015);

Mayr (2004);

Mayr (2007);

Louchart et al. (2008);

Mayr (1998);

Kundrát et al.

2015;

Mayr (2000); other specimens measured by the authors in the skeletal collection of ISEA PAS.

Taxon

Number

TBT

TMT

TBT/TMT

Aves indet. (present study)

†ISEA AF/JAM1

~13.8

6.3

2.19

†Eocypselidae
†Eocypselus rowei

WDC-CGR-109

20.1*

10.0*

2.01

†Eocypselus vincenti

MGUH 26729

20.2

10.9

1.85

Apodidae
†Scaniacypselus szarskii

LNK-Me 301, holotype

12.0

5.5

2.18

†Scaniacypselus szarskii

SMF-ME 3409A+B

~11.4

~5.9

1.93

Apus apus

A/4117/84

24.7

10.5

2.35

Apus apus

A/5858/01

25.6

10.1

2.53

Hirundapus caudacutus

A/5001/91

36.3

15.7

2.31

Trochilidae
†Eurotrochilus inexpectatus

SMNS 80739/4, holotype

14.9

6.5

2.29

†Eurotrochilus inexpectatus

SMNK-PAL 5591, 2

slab of holotype

~15.0

6.7

2.24

†Eurotrochilus sp.

NT-LBR-040

15.1

6.8

2.22

Eupeptomena macroura

A/5441/95

14.3

6.2

2.31

Ramphodon naevius

A/5516/96

13.2

6.5

2.03

Clytolaema rubricauda

A/5539/96

13.5

6.3

2.14

Calypta costae

A/4174/85

10.7

4.2

2.55

Phaethornis bourcieri

A/4074/84

11.2

4.3

2.60

†Messelirrisoridae
†Messelirrisor halcyrostris

SMF-ME 1883, holotype

16.9

9.6

1.76

† Messelirrisor halcyrostris

SMF-ME 11117a+b

15.5*

8.8*

1.76

† Messelirrisor halcyrostris

SMF-ME 10987a+b

15.3

9.2

1.66

†Messelirrisor grandis

SMF-ME 600

~19.0

~10.4

1.83

† Messelirrisor grandis

SMF-ME 108.33

18.2*

9.6*

1.90

†Messelirrisor parvus

SMF-ME 2793, holotype

15.8

8.4

1.88

† Messelirrisor parvus

SMF-ME 2466

14.0*

7.2*

1.94

† Messelirrisor parvus

SMF-ME 1180

14.8

7.9

1.87

† Messelirrisor parvus

SMNK-Me 300

15.8*

8.0*

1.98

† Messelirrisor parvus

SMNK-Me 776

15.0

8.1

1.85

Stem-group upupiform

† SNM-Z 27188

SNM-Z 27188

~19.0

7.1

2.68

Upupidae

Upupa epops

A/4006/84

36.7

21.8

1.68

Upupa epops

A/216/61

38.5

24.5

1.57

Passeriformes

Hirundo rustica

A/6106/02

21.2

11.5

1.84

Delichon urbica

A/5334/94

20.9

11.6

1.80

Riparia riparia

A/3133/76

19.4

10.2

1.90

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bones and the general shape of claws indicate a bird that

perched regularly. Short legs with long toes preclude a ground

dwelling bird and the claws are too robust and show too little

curvature for a bird that would cling to vertical tree trunks.

Thus, the specimen must have been a flying non-ground

dwelling bird that perched regularly. A similar but not iden-

tical, unspecialized foot was observed in the putative upupi-

form SNMZ 27188 (Kundrát et al. 2015) and stem-group

apodid Scaniacypselus (see Mayr 2015). It is noteworthy that

with the exception of two procellariiforms, all other speci-

mens — including ISEA AF/JAM1 — recorded from the

Outer Carpathians and Central Palaeogene Basin are land

birds that must have lived in the forests or shrubs near the

shoreline. The predominance of terrestrial birds in marine

deposits is nothing unusual; it was also observed in the

Eocene Fur Formation of Jutland in Denmark (Kristoffersen

2002), London Clay Formation in southern England

(Mlíkovský 2002), or the Oligocene Wiesloch-Frauenweiler

in southern Germany (Mayr 2009).

Although remains of Oligocene birds are extremely rare in

Poland (Bochenski et al. 2013a), the exposure at Jamna

Dolna 1 has already yielded several such fossils. The present

find is the oldest within the site (Unit C, ca. 32.5 Mya),

closely followed by a nearly complete passerine bird Jamna

szybiaki (Unit E, ca. 31.5 Mya) (Bochenski et al. 2011).

Moreover, three isolated feathers of unknown birds were

reported (Units G–H, ca. 31.0 Mya) (Bieńkowska-Wasiluk

2010, text-fig. 41A– C). The remains of land birds from

Jamna Dolna and other sites in the region can be attributed to

the general sea level fall at that time, which led to limitation

of the connection with the open ocean and resulted in many

shallow shoals, temporary islands and exposed dry land areas

along the coast (e.g., Rögl 1999; Kotlarczyk et al. 2006).

Acknowledgements: We thank Krzysztof Wertz (Institute

of Systematics and Evolution of Animals, Polish Academy

of Sciences, Kraków, Poland) for his help in preparing the

illustrations and comments on the early version of the manu-

script, and Andrea Pereswiet-Soltan for taking the photo-

graphs in Figure 3.

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