Historical Biology

An International Journal of Paleobiology

ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20

An illustrated summary of the lower Miocene

carnivores (Mammalia, Carnivora) of Tuchoice,
Czech Republic
Oldich Fejfar & Elmar Heizmann
To cite this article: Oldich Fejfar & Elmar Heizmann (2016) An illustrated summary of the
lower Miocene carnivores (Mammalia, Carnivora) of Tuchoice, Czech Republic, Historical
Biology, 28:1-2, 316-329, DOI: 10.1080/08912963.2015.1029923
To link to this article: http://dx.doi.org/10.1080/08912963.2015.1029923

Published online: 01 Oct 2015.

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Date: 02 October 2015, At: 00:18

Historical Biology, 2016

Vol. 28, Nos. 1 2, 316329, http://dx.doi.org/10.1080/08912963.2015.1029923

An illustrated summary of the lower Miocene carnivores (Mammalia, Carnivora) of Tuchorice,

Czech Republic
Oldrich Fejfara* and Elmar Heizmannb
a

Department of Paleontology, Faculty of Science, Charles University, Praha, Czech Republic; bStaatliches Museum fur Naturkunde,
Stuttgart, Germany
(Received 11 March 2015; accepted 12 March 2015)

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Fossils were discovered at the historical sites of Tuchorice, Middle Bohemia, Czech Republic in the mid-nineteenth
Century and first studied by Eduard Suess. After rediscovery in 1972, the localities were systematically collected.
At least 17 species and 14 genera of 8 families of Carnivora are recorded in the Early Miocene of Tuchorice and are
illustrated here.
Keywords: Early Miocene; Carnivora; natural trap; travertine cascade; thermal spring

Introduction

The freshwater limestones Facies of Tuchorice

Early Miocene mammalian remains from freshwater

limestones near Tuchorice have been known since the
nineteenth century (Reuss & v. Meyer 1851). The first
study of mammalian fossils from there was of
fragmentary rhinoceros teeth reported by Suess (1861),
which were given to him by Reuss (Figure 1).
Fragmentary teeth from the freshwater limestone were
sold in the 1880s by a local teacher, Karl Ihl, to the
Naturhistorisches Museum in Vienna. Much richer finds
from the travertines near Tuchorice were collected many
years later by O. Fejfar. The present synopsis provides
historical photographs and reports and illustrates the
carnivores from these historically important localities.
Most of the fossils illustrated here were collected since
1974 and are curated in the National Museum in Prague.
The Tuchorice fossil localities are now by law Protected
Paleontological Sites.
The freshwater limestones of Tuchorice were
believed to be the Lower Miocene equivalent of the
underlying marls of the main brown coal seam located in
the North Bohemian Tertiary basin (Figure 2). New
collections and comparisons of similar mammalian taxa
have shown the more advanced character of some
species, i.e., indicating a somewhat higher level in the
Miocene mammalian zone MN 3 (MN 3b). Thus, these
limestones are believed either contemporaneous with the
main brown coal seam or may be even somewhat
younger. The unusually rich accumulation of predators
resulted from their suffocation by CO2 released from the
thermal springs, which deposited travertines along an
active tectonic line.

Close to the southern margin of the Chomutov-Most

brown coal basin, at the sites near Tuchorice and Lipno
(Figure 3), several dislocated remnants of freshwater
limestones have been preserved. Genetically, they belong
to the sedimentary sequence of the basin but were isolated
by postdepositional tectonic movements and subsequent
erosion and are exposed now as relics. The freshwater
limestones of Tuchorice and Lipno were produced in three
different facies by several strong thermal mineral springs
along the southern margin of the major tectonic North
Bohemian Rift. The bones and teeth from there are often
fragmented through the mineralisation of gypsum.
Facies 1 is referred to as a cascade; it represents the
immediate surroundings of the hot springs and is exposed
in the SE portion of the village Tuchorice. This is the
original locality discovered around 1860 by A.E. Reuss
and described together with Suess (1861). The site was
rediscovered by Pavel Ctyroky in 1972 and since 1974
until 2000 has been systematically collected by O. Fejfar,
E. Heizmann and K. Heissig (for mammals) and C. Bu;
zek, Z. Kvacek and M. Konzalova (for palaeobotany and
palynology).
The outcrop (Figure 4; the Reuss Suess section) of the
mammal-bearing fossiliferous strata is on the southeast
edge of the village Tuchorice. The bright sequence of
travertines, dark-toned layers, grey marls and solid
calcareous banks produce rich Early Miocene floral and
faunal records including leaves, fruits and seeds, encrusted
insects and especially vertebrate remains. The vertebrates
are often in dark-tone lenses and comprise disarticulated
fragmentary bones and teeth of small and large mammals,

*Corresponding author. Email: fejfar@natur.cuni.cz

q 2015 Taylor & Francis

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Figure 1. Four renowned scientists who participated in the exploration of the Tertiary Period in Bohemia: August Emanuel Reuss,
Eduard Suess, Max Schlosser and Gustav Laube.

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Figure 2. Geological sketch of the Middle Brown Coal Basin, region of Chomutov Most. The N S profile between points A0 A
(shown in Figure 3) crosses the Ohre-Eger Rift in the central basin between Chomutov and Tuchorice. Brown coal seams are worked
today mostly in open pits. 1, Early Miocene near Merkur Nord, brown coal seam and strong humus clays at the calcareous base produces
an MN 3 mammal fauna and freshwater molluscs); 2, tectonic relics of Lower Miocene freshwater limestones (warm spring cascades) at
Tuchorice and Lipno, stratigraphically equivalent to the base of the brown coal seam in Merkur-Nord, with flora, molluscs and MN 3
mammal fauna; 3, Late Miocene noncalcareous clays and mudstones; 4, Late Cretaceous sandstones and spongilites; 5, Upper Paleozoic;
6, Erzgebirge phyllite and mica-schist in the region of Saxothuringia (separated by the Eger rift from tectonic system of the Krusne hory
Mts.).

amphibians, reptiles and rare endocranial parts and bones

of birds. In contrast to other facies of the limestones of
Tuchorice, this facies lacks mollusc remains, probably an
effect of the warm springs.
Facies 2 consists of solid slab-like limestone exposed
in an old quarry on the north-eastern margin of the village

Tuchorice (Figure 5) and in the abandoned quarries at

Lipno. It represents irregular, thinly bedded sediments
deposited in smaller ponds in close proximity to the hot
springs. These localities are rich in freshwater and
terrestrial molluscs (Wenz 1917) as well as the scattered
remains of mammals (e.g., Aureliochoerus, Palaeomeryx)

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Figure 3. (A) N S profile of the lignite basin along points A0 A of Figure 2 (1, brown coal seams, dark carbonaceous clays and
mudstones; 2, Lower Miocene freshwater limestones; 3, tertiary sands and clays; 4, Upper Cretaceous; 5, Permo-Carboniferous; 6,
crystalline core of the Krusne hory Mts.). (B) Detail of the southern part of the transect through freshwater limestones of Tuchorice and
Lipno (1, Ohre River Quaternary deposits; 2, Lower Miocene freshwater limestones; 3,4, tertiary sands, clays; 5, lower Turonian Planer;
6, Cenomanian sandstones and mudstones.

Figure 4. Profile through travertine facies showing 1974 excavations (Tuchorice 2 in Figure 3; in the Reuss Suess section). 1, Holocene
clay; 2, 3, sterile, irregularly layered fresh lime; 4, grey marl; 5, grey marl of the cavernous tufa (site 7389); 6, 7, soft tufa with
unidentifiable bone fragments, curled leaves, seeds of Phoenix, Toddalia, Celtis, natural casts of insects including the diving beetle,
Ditiscidae and two endocranial casts of birds; 8, 9, dark grey to black calcareous clay with disarticulated and fragmented bones and teeth
(sites 7382, 7389, 7391, 7392); 10, dark sideritic layer with gypsum veins (site 7390).

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O. Fejfar and E. Heizmann

Figure 5. Stratified freshwater limestones at Tuchorice 1 (Figure 3). Tuchorice 1 and Lipno are the classic sites of freshwater land snail
faunas (Reuss 1861; Klika 1891; Wenz 1917) yielding large mammals (Amphicyon, Hemicyon Aureliochoerus, Palaeomeryx, and tapirid)
and leaves of Livistona Bruder (1890) and seeds and fruits of Carya and Celtis, collected since the 1850s and housed in the museums in
Vienna and Prague.

Table 1. Early Miocene carnivores from the freshwater limestones of Tuchorice, Czech Republic.
Amphicyonidae
Amphicyon bohemicus (Schlosser 1899)
Amphicyon schlosseri Dehm 1950
? [Amphicyon helbingi Dehm 1950]
Megamphicyon major giganteus
Hemicyonidae
Hemicyon stehlini Hurzeler 1944 small form
? [Sp Phoberocyon ] great species
Thaumastocyonidae or Tomocyonidae
Tomocyon (? nov. gen.), nov. sp.
Ursidae
Ballusia elmensis (Stehlin 1917)
Ursavus isorei (Ginsburg & Morales 1998)
Procyonidae
Broiliana nobilis Dehm 1950
Stromeriellida
Stromeriella franconica Dehm 1950
Franconictis humilidens (Dehm 1950)
Viverridae
Semigenetta elegans Dehm 1950
Mustelidae
Martes laevidens Dehm 1950
Palaeogale hyaenoiodes Dehm 1950
Laphyctis vorax Dehm 1950
Amphictis antiquus Pomel 1853

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Figure 6. Left: lithographic plate of carnivore teeth identified as Amphicyon intermedius Meyer from Suess (1861). Right: photographs
of the same specimens and others from the collections of Charles University, Department of Geology-Palaeontology (GSKU). Figure
numbers from Suess, where applicable, follow photograph numbers in this caption. On the right, 1 (5a, on the left) and 2 (5b) are here
identified as Tomocyon (? nov. gen., nov. sp.), right C1 midsection in lingual and buccal views. Amphicyon bohemicus (Schlosser 1899), 3
(12a,b), left M3 in lingual view; 4, M2 occlusal view; 5, M3 occlusal view (11 a,b); 6 (lingual), 7 (occlusal). Hemicyon stehlini Hurzeler
1944, left P4 (6a c). 8: Amphicyon sp., left I3, medial and labial views (2a,b. 9 12, Amphicyon bohemicus (Schlosser 1899), left P4,
lingual, occlusal and buccal views (paratype; 7a d); 13, right M1, occlusal view (GSKU 4329; 10). 14, Amphicyon bohemicus (Schlosser
1899), M1 (9a,b). 15 17: Amphicyon bohemicus (Schlosser 1899), left M1, holotype, in lingual view (15; 8b), in buccal view (16; 8a) and
in occlusal view (17; 8c).

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Figure 7. Isolated amphicyonid teeth from Truchorice 1 in the Vienna Natural History Museum, Paleontological Department collected
by a local teacher Karl Ihl in 1890. Amphicyon bohemicus (Schlosser 1899), (1) left P4 in occlusal and buccal views, (3) right P4 in
occlusal and buccal views; (4) left M1; (5) juvenile left M2; (6) M2 isolated protocone; 2: Megamphicyon major-giganteus, left M3; 7,
Hemicyon stehlini Hurzeler 1944, right M1 without the mesial part.

Figure 8. Amphicyon sp. (cf. schlosseri Dehm 1950), from Tuchorice; 1 3, left mandible with C M2 in lateral, medial and occlusal
views; 4, M2 occlusal view; 5, M2 occlusal view.

Figure 9. Amphicyon bohemicus (Schlosser 1899) from Tuchorice (site 7391); lower dentition: (1, occlusal view; 2 lingual, view) M1
M3; (3, occlusal and lingual views) P3 M3; (4, occlusal and lingual views) M1 3; (5, occlusal and lingual view) P2 M2; (6, occlusal and
lingual views) P4 M2.

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Figure 10. Amphicyon bohemicus (Schlosser 1899) from Tuchorice (site 7391); lower dentition: (1 3, lingual, occlusal and buccal views)
M1 2; (4 5, lingual and occlusal views) M1 3; (6 7, buccal and occlusal views) M2, M3; (8, occlusal view) M2; (9, occlusal view) M2 3.

Figure 11. 1 5, 11 17: Amphicyon bohemicus (Schlosser 1899) from Tuchorice (site 7391). 10, 18: Amphicyon helbingi Dehm 1950
from Wintershof-West (WW); upper dentition. 110: P4 (left: buccal, middle: occlusal, right: lingual; 7 10: occlusal); 11, 12: M2 occlusal;
13 16: M3 occlusal. GKU4329 originally illustrated by Suess (1861).

Figure 12. Megamphicyon major giganteus from Tuchorice (site 7389). 1, M1 M3, occlusal view; 2 3, M1 buccal and lingual
views; 4, P4 M3 and P3 M1 and 5, M3, occlusal view.

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Figure 13. Hemicyon stehlini Hurzeler, 1944, from Tuchorice (site 7391); lower dentition. 1 3, P3 M3; 4 6, P2 M3. Hemicyon cf.
stehlini: 7 9, P3 M3.

Figure 14.

Hemicyon stehlini Hurzeler 1944 from Tuchorice (site 7391); lower dentition. 1 6: P4 M3, occlusal, buccal and lingual views.

Figure 15. Hemicyon stehlini Hurzeler, 1944 from Tuchorice (sites 7391, 7392); upper dentition. 1, 2, P3 M1 occlusal and buccal views;
3a, P3 M2 occlusal view; 3b, P2 3occlusal and buccal; 4, P3 4, buccal and lingual; 5, M2 occlusal and distal; 6, M2 occlusal; 7, M1
occlusal and distal; 8, M1 occlusal and distal; 9, M2 occlusal and distal; 10, P4 occlusal and lingual (Suess 1861, Taf.II, 6a c; numbers 5
6 in Figure 6 herein); 11: P4 occlusal, buccal and lingual.

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Figure 16. Tomocyon (? nov. gen.), nov. sp. from Tuchorice (site 7391). 1 3: left mandible with P4 M2 and the alveoli for P3 and M3; 4:
composite P4 M2 occlusal; 5: M2 occlusal; 6: M1 2 occlusal; 7: P4 buccal and lingual; 8: M1 distal and mesial; 9: M2 distal and mesial.

Figure 17. Tomocyon (? nov. gen.), nov. sp., from Tuchorice (site 7391); lower dentition. 1: P4 M1; 2: M1 2; 3: P4 M2, lingual, occlusal
and buccal views.

Figure 18. Tomocyon (? nov. gen.), nov. sp., from Tuchorice (site 7391); lower dentition. 1 and 2, P4 M3 buccal, occlusal and lingual
views; 3, M1, buccal, occlusal and lingual views; 4 8, M2 occlusal views.

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Figure 19. Ursavus (Ballusia) elmensis (Stehlin 1917) from Tuchorice (sites 7391, 7392, 7389) and from Wintershof-West (WW;
original illustrations in Dehm 1950). 1 6: M2 occlusal and buccal; 7: P4, buccal, occlusal and lingual views; 8: P4, occlusal and lingual
views; 9: P4, occlusal and lingual views.

Figure 20. Ursavus (Ballusia) elmensis (Stehlin 1917) from Tuchorice (sites 7391, 7389, and Wintershof-West, WW, originally
illustrated by Dehm 1950). 1, M2 occlusal and mesial views; 2, M2 fragment, occlusal and mesial views; 3, M1, occlusal view; 4, M1 distal
and occlusal views; 5, M2 occlusal and mesial views; 6, M2, occlusal; 7, P4 occlusal and lingual views.

Figure 21. Ursavus isorei (Ginsburg & Morales 1998) from Tuchorice (site 7391). 1, M1 occlusal and mesial views; 2, M1 occlusal and
distal; 3, M1? lingual fragment occlusal; 4, occlusal; 5, M3 M2 occlusal; 6: M2 occlusal and lingual; 7, M1 occlusal and buccal.

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Figure 22. Franconictis, Stromeriella and Palaeogale from Tuchorice (sites 7382, 7389, 7390, 7391). Franconictis (Plesictis)
humilidens (Dehm 1950): 1, M1 buccal, occlusal and lingual views; 2, occlusal; 3, M2 occlusal and buccal. Stromeriella franconica Dehm
1950: 4 7, M2 occlusal and buccal; 8, M2 occlusal. Palaeogale hyaenoiodes Dehm 1950: 9 11: P4, P3, P4 left lingual, right buccal; 12
15: p4 , p4 7, p3, p1 2 occlusal, 15: right buccal; 16: P4 upper lingual, down occlusal; 17, P4 buccal and lingual.

Figure 23. Palaeogale hyaenoiodes Dehm 1950 from Tuchorice (sites 7389, 7390, 7391): 1, left lower jaw with P4 and M1 buccal, 2,
occlusal and detail of P4 and M1, and 3, buccal detail; 4, right jaw fragment with M1 occlusal and buccal; 5, M2 mesial, lingual and
occlusal; 6, M1 buccal, occlusal and lingual; 7 9: three P3s buccal and occlusal; 10 12: three P4s buccal and occlusal.

Figure 24. Stromeriella and Broiliana from Tuchorice. Stromeriella franconica Dehm 1950 (sites 7382, 7389, 7390, 7391): 1, 2, M1
buccal, occlusal and lingual; 3, M1 occlusal; 4, M1 occlusal, mesial and distal; 5, M1 mesial and occlusal; 8, M2 mesial, occlusal and distal; 9,
M3 occlusal and distal. Broiliana nobilis Dehm 1950 (sites 7391, 7389): 6, M1 (lingual part) occlusal; 7, M1 mesial, occlusal and distal.

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Figure 25. 1 4: Semigenetta, Franconictis, Laphyctis, and Amphictis from Tuchorice (sites 7389, 7390, 7391). Semigenetta elegans
Dehm 1950: 1, P3 occlusal, lingual and buccal; 2, P3 occlusal, lingual and buccal; 3, P4 occlusal, lingual and buccal; 4, M1 (without
talonid) lingual, occlusal and buccal; 5, M1 (talonid) buccal; (HM 1950). Franconictis (Plesictis) humilidens: 6, M1 lingual, occlusal and
buccal. Laphyctis comitans Dehm 1950: 7, M1 left lingual, occlusal and buccal. Amphictis antiquus Pomel 1853; 8, P3 occlusal, buccal
and lingual; 9, P3 occlusal, buccaland lingual.

Figure 26. Laphyctis vorax seu Laphyctis comitans Dehm 1950 from Tuchorice (sites 7389, 7391): 1, M1 lingual, occlusaland buccal; 2,
M1 (without talonid) lingual, occlusal and buccal; 3, 4, two P4s lingual, occlusal and buccal; 5, M1 occlusal, mesial and distal; 6, M1
occlusal.

Figure 27. Laphyctis vorax seu Laphyctis comitans Dehm 1950 from Tuchorice (sites 7389, 7391): 1, M1 without paraconid (completed
with M1 from Wintershof-West, WW 13282) occlusal, buccal and distal; 2, M1 buccal, occlusal, lingual; 3, P3 occlusal, buccal and
lingual; 4, P3 occlusal, buccal and lingual; 5, P3 occlusal, buccal and lingual; 6, P3 occlusal, buccal and lingual; 7, P2 occlusal, buccal and
lingual; 8, P1 occlusal, buccal and lingual; 9 11, three M2s occlusal and lingual.

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and plants (Livistona, Carya fruit casts, Celtis fruit

stones).
Facies 3 consists of dark carbonaceous clays and grey
marls containing abundant Celtis fruit stones either
concentrated in lenses in the travertine cascades of the
Reuss Suess section in the southeast of Tuchorice
(Figures 6 and 7) or underlying the solid limestones of
the Tuchorice and Lipno quarries (Figure 5).
A list of the Early Miocene Carnivora from Tuchorice
is given in Table 1, and all taxa are illustrated in
Figures 6 27. A more detailed study is in preparation by
Jorge Morales, Juan Abella (Madrid), Elmar Heizmann
(Stuttgart), Oldrich Fejfar, Jan Wagner (Prague) and
Martin Sabol (Bratislava).

329

Disclosure statement
No potential conflict of interest was reported by the authors.

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Mammalia) et les formes apparentees du Miocene inferieur et moyen
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ber die Baren und barenahnlichen Formen des
Schlosser M. 1899. U
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ber die groen Raubtiere der osterreichischen
Suess E. 1861. U
Tertiarablagerungen [On the large Carnivores in the Austrian
tertiary deposits]. Sitzb Akad Wiss Math-nat Kl. 43:217232.
Wenz W. 1917. Zur Altersfrage der bohmischen Susswasserkalke [On
the age of the freshwater limestones in Bohemia]. Jahrb Nassau Ver
Naturk Wiesbaden. 70:183.