EURASIAN MIDDLE AND LATE MIOCENE HOMINOID ......African apes. Together with analysis of environmental data, Eurasian mammals support the hypothesis that the descendant of a Eurasian
This document is posted to help you gain knowledge. Please leave a comment to let me know what you think about it! Share it to your friends and learn new things together.
Transcript
EURASIAN MIDDLE AND LATE MIOCENE HOMINOID PALEOBIOGEOGRAPHY
AND THE GEOGRAPHIC ORIGINS OF THE HOMININAE
by
Mariam C. Nargolwalla
A thesis submitted in conformity with the requirements
Chapter 2 – Data Quality .................................................................................................... 14 Introduction......................................................................................................................... 14
Loss of Biological Information in the Fossil Record...................................................... 14 Materials and Methods........................................................................................................ 16
Spatio-temporal Distribution of Faunas.......................................................................... 16 Method of Analysis......................................................................................................... 17
Results................................................................................................................................. 19 Relationship between duration of temporal intervals & number of localities/number of
localities & CI................................................................................................................. 19 Large Mammal Completeness ........................................................................................ 24 Small Mammal Completeness ........................................................................................ 29 Identification of “Marker Taxa” ..................................................................................... 33
Discussion ........................................................................................................................... 38 Overall Completeness and Data Quality......................................................................... 38 Ecology of “Marker Taxa” ............................................................................................. 40 Distribution of sample localities ..................................................................................... 40
Chapter Summary & Conclusions ...................................................................................... 44 Chapter 3 – Provinciality, paleoenvironments, in situ evolution and dispersal ................. 45
Introduction......................................................................................................................... 45 Zoogeographic Provinces and Purpose of Study ............................................................ 45 Barriers to dispersal & paleoenvironmental events in the Miocene of Eurasia.............. 46 Previous method and study of provinciality ................................................................... 53
Discussion ......................................................................................................................... 148 Resolution of analysis ................................................................................................... 148 Faunal provinces in comparison to previous studies .................................................... 151 Faunal provinces: In situ evolution and dispersals ....................................................... 157 Paleoenvironmental Influence on Faunal Provinces..................................................... 179 Inter- and Intracontinental Faunal Exchange and Dispersal Pathways......................... 180
Introduction....................................................................................................................... 186 First appearances and relations among Eurasian Miocene apes ................................... 186 Griphopithecus and cf. Griphopithecus........................................................................ 187 Dryopithecus/Pierolapithecus ...................................................................................... 203
Table 1.1: Morphological characters shared with African apes and Dryopithecus .................. 3 Table 1.2: Temporal association of the Eurasian MN Zones.................................................... 6 Table 2.1: Temporal ranges of Eurasian large mammal genera ............................................. 25 Table 2.2: Temporal ranges of Eurasian small mammal genera............................................. 30 Table 2.3: Temporal range and paleoecology of well-sampled Eurasian small mammals..... 34 Table 2.4: Temporal range and paleoecology of well-sampled Eurasian large mammals ..... 36 Table 3.1: Localities included in analysis............................................................................... 59 Table 3.2: Summary of provincial composition (species-level) ........................................... 140 Table 4.1: Eurasian – Afro-Arabian shared taxa (MN5 and earlier) .................................... 192 Table 4.2: FAs and centres of origin of German and Turkish MN5 taxa ............................. 193 Table 4.3: European-Turkish shared Astaracian taxa ........................................................... 200 Table 4.4: Spanish large and small mammal FAs, MN6-MN9 ........................................... 206 Table 4.5: MN7/8 Spanish – Central European shared taxa................................................. 210 Table 4.6: Shared Spanish taxa............................................................................................. 213
vii
List of Figures
Figure 2.1: Distribution of Sample Localities......................................................................... 16 Figure 2.2: (a-g) ...................................................................................................................... 20 Figure 2.3: Relative completeness in Eurasian large mammal taxa ...................................... 24 Figure 2.4: Relative completeness in Eurasian small mammal taxa...................................... 29 Figure 2.5: Regional or bioprovincial zones previously assessed for CI................................ 41 Figure 2.6: Regional and total large mammal completeness ................................................. 43 Figure 2.7: Regional and total small mammal completeness ................................................. 43 Figure 3.1: Physiographic features in study area .................................................................... 53 Figure 3.2: Distribution of localities....................................................................................... 69 Figure 3.3a: MN5 large mammal species dendrogram........................................................... 76 Figure 3.3b: MN5 large mammal genera dendogram............................................................. 77 Figure 3.3c: MN5 small mammal species dendrogram .......................................................... 78 Figure 3.3d: MN5 small mammal genera dendogram ............................................................ 79 Figure 3.4a: MN6 large mammal species dendrogram........................................................... 84 Figure 3.4b: MN6 large mammal genera dendogram............................................................. 85 Figure 3.4c: MN6 small mammal species dendrogram .......................................................... 86 Figure 3.4d: MN6 small mammal genera dendogram ............................................................ 87 Figure 3.5a: MN7/8 large mammal species dendrogram........................................................ 93 Figure 3.5b: MN7/8 large mammal genera dendogram.......................................................... 94 Figure 3.5c: MN7/8 small mammal species dendrogram ....................................................... 95 Figure 3.5d: MN7/8 small mammal genera dendogram ......................................................... 96 Figure 3.6a: MN9 large mammal species dendrogram......................................................... 102 Figure 3.6b: MN9 large mammal genera dendogram........................................................... 103 Figure 3.6c: MN9 small mammal species dendrogram ........................................................ 104 Figure 3.6d: MN9 small mammal genera dendogram .......................................................... 105 Figure 3.7a: MN10 large mammal species dendrogram....................................................... 110 Figure 3.7b: MN10 large mammal genera dendogram......................................................... 111 Figure 3.7c: MN10 small mammal species dendrogram ...................................................... 112 Figure 3.7d: MN10 small mammal genera dendogram ........................................................ 113 Figure 3.8a: MN11 large mammal species dendrogram...................................................... 118 Figure 3.8b: MN11 large mammal genera dendogram......................................................... 119 Figure 3.8c: MN11 small mammal species dendrogram ...................................................... 120 Figure 3.8d: MN11 small mammal genera dendogram ........................................................ 121 Figure 3.9a: MN12 large mammal species dendrogram....................................................... 127 Figure 3.9b: MN12 large mammal genera dendogram......................................................... 128 Figure 3.9c: MN12 small mammal species dendrogram ...................................................... 129 Figure 3.9d: MN12 small mammal genera dendogram ........................................................ 130 Figure 3.10a: MN13 large mammal species dendrogram..................................................... 136 Figure 3.10b: MN13 large mammal genera dendogram....................................................... 137 Figure 3.10c: MN13 small mammal species dendrogram .................................................... 138 Figure 3.10d: MN13 small mammal genera dendogram ...................................................... 139 Figure 3.11: Geographic distribution of faunal provinces .................................................... 155 Figure 3.12a: Dispersal and in situ evolution in Eurasian faunas (MN5)............................. 171 Figure 3.12b: Dispersal and in situ evolution in Eurasian faunas (MN6) ............................ 171
viii
List of Figures (Continued) Figure 3.12c: Dispersal and in situ evolution in Eurasian faunas (MN7/8).......................... 171 Figure 3.12d: Dispersal and in situ evolution in Eurasian faunas (MN9) ............................ 171 Figure 3.12e: Dispersal and in situ evolution in Eurasian faunas (MN10)........................... 171 Figure 3.12f: Dispersal and in situ evolution in Eurasian faunas (MN11) ........................... 171 Figure 3.12g: Dispersal and in situ evolution in Eurasian faunas (MN12) .......................... 171 Figure 3.12h: Dispersal and in situ evolution in Eurasian faunas (MN13) .......................... 171 Figure 3.13: West Asian-Eastern European dispersal routes................................................ 181 Figure 3.14: Dispersal routes into and trans-Pannonian Basin ............................................. 183 Figure 4.1: Distribution of Griphopithecus localities ........................................................... 188 Figure 4.2a: MN3/5 dispersal pathways into Europe (modified from Popov et al. 2004) .. 197 Figure 4.2b: MN4 dispersal pathways into Europe via Turkey (modified from Rögl 1999. 199 Figure 4.3: Distribution of Dryopithecus localities (including Pierolapithecus) ................ 204 Figure 4.4: Late Astaracian Eurasian immigrations ............................................................. 206 Figure 4.5: Distribution of Ouranopithecus localities .......................................................... 218
1
Chapter 1 - Introduction
On the Birthplace and Antiquity of Man – We are naturally led to enquire,
where was the birthplace of man at that stage of descent when our
progenitors diverged from the Catarhine stock? The fact that they belonged
to this stock clearly shews that they inhabited the Old World; but not
Australia nor any oceanic island, as we may infer from the laws of
geographical distribution. In each great region of the world the living
mammals are closely related to the extinct species of the same region. It is
therefore probable that Africa was formerly inhabited by extinct apes closely
allied to the gorilla and chimpanzee; and as these two species are now
man’s nearest allies, it is somewhat more probable that our early
progenitors lived on the African continent than elsewhere…
C. Darwin (1871, p182) Since the publication of Darwin’s The Descent of Man, and Selection in Relation to
Sex in 1871, this passage has been well cited, illuminating Africa as the geographic place of
origin and diversification of the African ape and human (hominine) lineage. An African
origin hypothesis is logical on many counts. As Darwin (1871) pointed out, because our
closest living relatives, the gorilla and chimpanzee, are restricted to Africa, the last common
ancestor of African apes and humans most likely evolved there as well. Furthermore, the
earliest fossil hominins (species more closely related to Homo than to Pan) are similarly
restricted to Africa. African paleoenvironments are thought to have provided the ecological
requirements necessary for ancestral hominines to evolve there in the late Miocene (Bernor
1978; Cote 2004; Pilbeam 1997; Pilbeam & Young 2004). However, until recently, very few
fossil apes were known from the late Miocene of Africa. Cote (2004) has suggested that this
is a consequence of a poor fossil record, riddled with preservational and sampling biases, and
2
that of the late Miocene fossil localities that are known, very few sample the type of
environment in which fossil apes could thrive.
Darwin’s (1871) speculation of Africa as the birthplace of the hominine lineage was
immediately followed in the next sentence by an interesting observation; curiously enough,
one that is largely overlooked:
…. But it is useless to speculate on this subject; for two or three
anthropomorphous apes, one the Dryopithecus of Lartet, nearly as large as
a man, and closely allied to Hylobates, existed in Europe during the
Miocene age; and since so remote a period the earth has undergone many
great revolutions, and there has been ample time for migrations on the
largest scale.
C. Darwin (1871, p182-183)
A Eurasian origin for the Homininae is considered a minority view (Cote 2004; Pilbeam &
Young 2004; Stewart & Disotell 1998). Advocates of this hypothesis suggest that after an
initial intercontinental dispersal from Africa to Eurasia at ~17Ma and subsequent radiation
from Spain to China, fossil apes disperse into Africa at least once in the late Miocene. It is
therefore probable that African hominines are derived from a Eurasian ancestor (Begun 2007,
2005, 2001; Begun et al. 1997; Heizmann & Begun 2001). Support for this hypothesis is
derived from cladistic analyses, identifying synapomorphies supporting a sister relationship
between late Miocene Eurasian fossils and the African ape and human clade (Table 1.1).
These analyses specifically identify late Miocene Eurasian genera, such as Dryopithecus and
Ouranopithecus, as stem hominines, while the few contemporaneous specimens in Africa are
considered to lack synapomorphies linking them to the African ape and human clade (Begun
2007, 2005, 2001, 1994; Begun et al. 1997). A Eurasian origin of the Homininae is also
supported by the biogeographic and phylogenetic patterns in other land mammal groups
3
(Begun 2007, 2005; Folinsbee & Brooks 2007; Made 1999; Stewart & Disotell 1998) and is
consistent with the molecular divergence dates of extant taxa (Stewart & Disotell 1998),
while the African fossil record fails to preserve diagnostic hominines until the latest
Miocene.
Table 1.1: Morphological characters shared with African apes and Dryopithecus
(from Begun 2005, p56)
The Eurasian origin hypothesis has been challenged on several grounds. First, there
lacks consensus regarding the phylogenetic relations among Eurasian Miocene apes and
between these taxa and African hominins, as a consequence of temporal, geographic and
morphological unevenness in fossil representation, as well as differences in interpretation of
the phylogenetic significance of preserved anatomy (Andrews 1992a; Andrews & Bernor
1999; Andrews et al. 1996; Andrews & Martin 1987; Begun 1995, 1994, 1992a; Begun &
4
Kordos 1997; Begun et al. 1997; Harrison & Rook 1997; Martin & Andrews 1993; Moyà-
Solà et al. 2004; Moyà-Solà & Köhler 1993; Pilbeam 1997, 1996; Pilbeam & Young 2004;
Ward et al. 1997). Second, the ancestral hominine morphotype has been reconstructed using
what some consider crown ape synapomorphies, as a suspensory, frugivorous, tropical forest-
dweller, most similar to Pan (Cote 2004; Pilbeam 1997, 1989; Pilbeam & Young 2004).
According to supporters of the African origin hypothesis, no Eurasian ape fits this
morphotype and instead, the last common ancestor of African apes and humans has yet to be
discovered in equatorial Africa. Lastly, from an ecological perspective, late Miocene
Eurasian paleoenvironments would not support intercontinental dispersals of Eurasian apes to
Africa, due to the lack of densely forested conditions and the unavailability of ripe fruit along
dispersal corridors (Andrews 2007; Cote 2004; Pilbeam & Young 2004). As a result,
Eurasian apes underwent extinction during or shortly after the [mid-] Vallesian Crisis at
~9.6Ma, when most forest-dwelling mammals succumbed to the cooler, drier, and more
seasonal climates, and thus bear no relation to the African ape and human lineage
Within each mammalian order there occurred genera with overall complete fossil
records and long temporal durations and would thus serve as good “marker” taxa to clarify
patterns of primate dispersal. Specifically, many of these genera are considered to have
similar niche requirements and/or are associated with non-cercopithecoid primates at fossil
localities. These taxa are listed in the following tables (Tables 2.3 & 2.4). Note that for
many genera, their last appearance actually occurs in the Pliocene, however they are listed
with a LAD of MN13 since Pliocene taxa are not being considered here.
34
Table 2.3: Temporal range and paleoecology of well-sampled Eurasian small mammals * = co-occurs with a non-cercopithecoid primate (E) = environmental tolerance of extant form/relative
Taxon Ecology Temporal Range Reference Lagomorpha Ochotonidae: Eurolagus*
Table 2.4: Temporal range and paleoecology of well-sampled Eurasian large mammals * = co-occurs with a non-cercopithecoid primate (E) = environmental tolerance of extant form/relative
Taxon Ecology Temporal Range Reference Artiodactyla Bovidae: Eotragus*
lightly wooded (lrg body size), closed
MN4-7/8
Gentry et al. (1999)
Miotragocerus* MN5(6?) -12 Gentry et al. (1999) Palaeoreas* grazer MN9-13 Gentry et al. (1999) Prostrepsiceros* grazer MN9-12 Gentry et al. (1999) Tethytragus* closed MN5-7/8 Gentry et al. (1999) Tragoportax* MN9-13 Gentry et al. (1999) Cervidae: Dicrocerus*
forest (E), wooded, proximity to water
MN5-9
Gentry et al., (1999)
Euprox* MN5-10 Gentry et al. (1999) Lagomeryx* forest, wooded, +
underbrush MN4-7/8 Gentry et al. (1999)
Procapreolus* MN9-13 Gentry et al. (1999) Stehlinoceros* wooded MN5-7/8 Gentry et al. (1999) Giraffidae: Bohlinia*
MN10-13
Gentry et al. (1999)
Giraffokeryx* MN5-7/8 Gentry et al. (1999) Helladotherium* associated with open
fauna MN10-13
Gentry et al. (1999)
Paleotragus* associated with forest/ steppe/mixed/open fauna
MN7/8-13 Gentry et al. (1999)
Samotherium* mixed feeder/grazer, associated with mixed/ open fauna
MN9-13 Gentry et al. (1999)
Palaeochoeridae: Taucanamo
forest
MN4-9
Fortelius, Made & Bernor (1996)
Suidae: Aureliachoerus*
MN4-6
Hünermann (1999)
Bunolistriodon* heterogeneous environment, open
MN4-7/8
Fortelius, Made & Bernor (1996), Hünermann (1999)
Conohyus* marshy forest, not open MN5-9 Fortelius, Made & Bernor (1996), Hünermann (1999), Thenius (1952)
Hippopotamodon* woodland/grassland mosaic, open
MN7/8-10 Fortelius, Made & Bernor (1996), Hünermann (1999)
Hyotherium* not open MN4-6 Fortelius, Made & Bernor (1996), Hünermann (1999)
Listriodon* heterogeneous environment, open
MN4-9 Fortelius, Made & Bernor (1996), Hünermann (1999)
Microstonyx* woodland/grassland mosaic
MN9-13 Fortelius, Made & Bernor (1996), Hünermann (1999)
Paraleuasto- choerus*
not open MN6-11 Fortelius, Made & Bernor (1996), Hünermann (1999)
Propotamo- heterogeneous MN7/8-13 Fortelius, Made & Bernor
37
Taxon Ecology Temporal Range Reference choerus* environment, open (1996), Hünermann (1999) Tragulidae: Dorcatherium*
forest, + undergrowth, proximity to water, semi-aquatic
consider all pair-wise dissimilarities among data points (Quinn & Keough 2002).
The current study differs from those previously mentioned in several significant
ways. In contrast to both Bernor (1983 & 1978) and Fortelius et al. (1996a), this analysis
was conducted using raw presence/absence data, in addition to similarity indices. When
using a similarity index, taxonomic identity is lost, due to the numerical representation of
taxa. Therefore, analysis conducted on the raw presence/absence data facilitates the
recognition of key taxa influencing the formation, composition and maintenance of
57
bioprovinces. Over successive time slices, these same taxa can be tracked to better
understand if, when and where they disperse. In contrast to Fortelius et al. (1996a), who
divided their study area into blocks and regions a priori and subsequently assessed similarity,
this study uses the fauna themselves to define faunal provinces. This allows for the
recognition of bioprovinces in relation to landforms and regional topography. In addition, no
study to date has specifically accounted for large scale faunal distribution in relation to the
combined effect of specific climatic, tectonic and eustatic factors. This study uses the most
recently updated and extensive sample of fossil mammals and integrates new data not
previously used. Lastly, the innovative use of GIS to graphically represent environmental
effects together with faunal distribution facilitates the recognition and definition of
bioprovinces, their barriers and fluctuations therein over time.
Materials & Methods
Materials
Taxon occurrence data from the previous chapter were used here to assess
provinciality. Localities were temporally ordered according to their MN designation into a
data matrix and their component taxa were scored as present or absent. Localities falling
within a single MN unit were considered here, although a number of localities correlated
with land mammal ages (i.e., Vallesian, Turolian, etc.) were also included if these localities
had geographic, temporal or taxonomic relevance. Furthermore, localities with more than 10
species were selected for analysis as to minimize taxonomic “noise,” as per Alroy (2004).
However, localities located in poorly represented regions, as well as species-poor primate
localities were also considered. The sample of localities extends across the same geographic
expanse as in the previous chapter, but also includes late Miocene localities in the Middle
58
East and a smaller series of middle and late Miocene African localities to demonstrate the
timing and taxonomic composition of intercontinental faunal exchange (Figure 3.2, Table
3.1). Temporally, the sample of localities spans from 17Ma (base of MN5) to 5.3Ma (base of
MN 14) and includes small mammals (rodents, insectivores, lagomorphs) from 270 localities
and large mammals from 328 localities.
Analysis
Hierarchical cluster analysis was used to explore and determine groupings at the
genus and species level for large and small mammals within individual MN units. I used
single linkage, complete linkage, unweighted pair-group average and Ward’s minimum
variance to determine which linkage method would result in the highest co-phenetic
correlation, as well to observe whether topologies would change when using the different
algorithms. Single linkage determines the distance between clusters using nearest
neighbours, or the minimum distance between objects in different clusters, while complete
linkage performs the opposite way; distances between clusters are determined using the
greatest distance between any two objects in the different clusters. The unweighted pair-
group average method assesses the mean distance between all pairs of objects in different
clusters. Ward’s minimum variance evaluates distance between clusters using an analysis of
variance to minimize the sum of squares of each cluster (Hammer et al. 2007; Legendre &
Legendre 1998). I also conducted identical cluster analyses using the Jaccard, Dice and
Raup-Crick similarity indices, again to observe any differences in clustering and dendrogram
topology. The Jaccard and Dice index both use pair-wise presence/absence data to produce a
coefficient of faunal similarity, however, the latter weights observed joint occurrences more
heavily than absences. The Raup-Crick index uses Monte Carlo randomization to compare
59
the observed number of co-occurrences in a pair of localities with the distribution of co-
occurrences in 200 random iterations (Hammer et al. 2007). Following the similarity
analyses, I verified the clustering patterns with the raw data to ensure that the groups formed
were supported by the data. Not wanting to force the data into a specified number of groups
a priori, I chose to avoid any partition methods of clustering (i.e., K-means, non-hierarchical
linkage). The spatial distribution of fauna and the geographic extent of mountain chains, the
Paratethys and the Tethys, were plotted as individual layers on a series of base maps, using
ArcGIS 9.2.
Table 3.1: Localities included in analysis
MN 4-MN5 (17-15.2Ma)
Locality Country Age Ad Dabtiyah Saudi Arabia MN3-5 Rusinga (-) Kenya MN4 Rusinga (Gumba) Kenya MN4 Rusinga (Hiwegi west) Kenya MN4 Rusinga (Hiwegi) Kenya MN4 Rusinga (Kathwanga) Kenya MN4 Rusinga (Kiahera Hill) Kenya MN4 Rusinga (Kiyune) Kenya MN4 Rusinga (Nyamsingula) Kenya MN4 Rusinga (R 113) Kenya MN4 Rusinga (R 114) Kenya MN4 Rusinga (R105) Kenya MN4 Rusinga (R106) Kenya MN4 Rusinga (R107) Kenya MN4 Rusinga (R3) Kenya MN4 Rusinga (Wayondo) Kenya MN4 Kalodirr Kenya Burdigalian Napak Uganda Burdigalian Napak (Iriri member) Uganda Burdigalian Napak Member Uganda Burdigalian Napak Rhino Site Uganda Burdigalian Göriach Austria MN5 Grund Austria MN5 Muhlbach am Manhartsberg Austria MN5 Obergänserndorf 1 & 2 Austria MN5 Teiritzberg 1 (T1) Austria MN5 Franzensbad Czech Republic MN5
60
Baigneaux-en Beauce France MN5 Esvres - Marine Faluns France MN5 Faluns of Touraine & Anjou France MN5 La Condoue France MN5 Manthelan France MN5 Pontlevoy France MN5 Poudenas-Peyrecrechen France MN5 Rimbez - Lapeyrie base France MN5 Savigné-sur-Lathan France MN5 Contres MN 5 France MN5 Belometchetskaja Georgia MN5 Affalterbach Germany MN5 Engelswies Germany MN5 Gisseltshausen Germany MN5 Griesbeckerzell Germany MN5 Häder Germany MN5 Hambach 6C Germany MN5 Heggbach Germany MN5 Laimering 3 GE 5 Germany MN5 Massendorf Germany MN5 Puttenhausen Germany MN5 Rothenstein 1 Germany MN5 Sandelzhausen Germany MN5 Schellenfeld 2-4 Germany MN5 Walda 2 Germany MN5 Wannenwaldtobel-2 Germany MN5 Ziemetshausen 1b Germany MN5 Antonios (ANT) Greece MN5 Chios Greece MN5 Belchatow B Poland MN5 La Hidroelectrica Madrid Spain MN5 Montejo de la Vega Spain MN5 Puente de Vallecas Spain MN5 Sant Mamet Spain MN5 Somosaguas-Sur Spain MN5 Chatzloch Switzerland MN5 Frohberg Switzerland MN5 Hüllistein Switzerland MN5 Martinsbrünneli Switzerland MN5 Tobel-Hombrechtikon Switzerland MN5 Vermes 1 Switzerland MN5 Vermes 2 Switzerland MN5 Ardiç-Mordoğan Turkey MN5 Çandır (Loc. 3) Turkey MN5 Çandır 2 Turkey MN5 Karaağaç 1 Turkey MN5 Koçgazi Turkey MN5 Paşalar Turkey MN5 Arrisdrift Namibia MN5 Al-Sarrar Saudi Arabia MN5
61
Sinda Congo Congo/Zaire Langhian Kipsaramon 1 Kenya Langhian Kipsaramon 2 Kenya Langhian Maboko Kenya Langhian Majiwa Kenya Langhian Ombo Kenya Langhian
MN6 (15.2-12.5)
Locality Country Age Sinda Congo Congo/Zaire Langhian Kipsaramon 1 Kenya Langhian Kipsaramon 2 Kenya Langhian Maboko Kenya Langhian Majiwa Kenya Langhian Ombo Kenya Langhian Klein Hadersdorf Austria MN6 Trimmelkam Austria MN6 Castelnau-d'Arbieu France MN6 Four France MN6 Four (general) France MN6 Liet France MN6 Sansan France MN6 Simorre France MN6 Diessen am Ammersee Germany MN6 Stätzling Germany MN6 Steinberg Germany MN6 Thannhausen Germany MN6 Unterneul Germany MN6 Matraszolos1-2 Hungary MN6 Samsonhaza Hungary MN6 Subpiatrã 2/1R Romania MN6 Devínská Nová Ves - Bonanza Slovakia MN6 Devínská Nová Ves - Fissures Slovakia MN6 Devínská Nová Ves - Sandhill Slovakia MN6 Armantes 7 Spain MN6 Arroyo del Val Spain MN6 Elgg Switzerland MN6 Kreutzlingen Switzerland MN6 Rümikon Switzerland MN6 Schwamendingen Switzerland MN6 Stein am Rhein Switzerland MN6 Wiesholz Switzerland MN6 Zeglingen Switzerland MN6 Bagici Turkey MN6 Catakbagyaka Turkey MN6 Inönü I (Sinap 24A) Turkey MN6 Al Jadidah Saudi Arabia MN6 Sevastopol (Sebastopol) Ukraine Sarmatian
62
Fort Ternan Kenya Serravallian Fort Ternan 2 (Serek) Kenya Serravallian Ngorora Kenya Serravallian Nyakach 10 (Kaimogool North) Kenya Serravallian Nyakach 10 (Kaimogool North) Kenya Serravallian Nyakach 11 (Kaimogool South) Kenya Serravallian Nyakach 11(Kaimogool South) Kenya Serravallian Nyakach 8 (Kadianga West) Kenya Serravallian Nyakach 8 (Kadianga West) Kenya Serravallian Ngorora Kenya Serravallian-Tortonian
MN7/8 (12.5-11.2Ma)
Locality Country Age St. Stephan im Lavanttal Austria MN7/8 La Grive St. Alban France MN7/8 Poudenas-Cayron France MN7/8 St. Gaudens France MN7/8 Massenhausen Germany MN7/8 Steinheim Germany MN7/8 Felsotárkány 1 Hungary MN7/8 Felsotárkány 3/2 Hungary MN7/8 Felsotárkány 3/2 (Güdör-kert) Hungary MN7/8 Felsotárkány 3/8 Hungary MN7/8 Felsotárkány-Felnémet Hungary MN7/8 Opole 2 Poland MN7/8 Przeworno 2 Poland MN7/8 Comanesti-1 Romania MN7/8 Subpiatrã 2/2 Romania MN7/8 Can Feliu Spain MN7/8 Can Mata 1 Spain MN7/8 Castell de Barberà Spain MN7/8 Escobosa Spain MN7/8 Hostalets de Pierola Inferior Spain MN7/8 Nombrevilla-2 Spain MN7/8 Sant Quirze Spain MN7/8 Anwil Switzerland MN7/8 Bois de Raube 3 Switzerland MN7/8 Bayraktepe 1 Turkey MN7/8 Pismanköy (Yenidibekkoyu) Turkey MN7/8 Sariçay Turkey MN7/8 Sofca Turkey MN7/8 Yeni Eskihisar 1 Turkey MN7/8 Yenieskihisar Turkey MN7/8 Bled Douarah Tunisia 12.5-9.5Ma Sevastopol (Sebastopol) Ukraine Sarmatian
63
Fort Ternan Kenya Serravallian Fort Ternan 2 (Serek) Kenya Serravallian Ngorora Kenya Serravallian Nyakach 10 (Kaimogool North) Kenya Serravallian Nyakach 10 (Kaimogool North) Kenya Serravallian Nyakach 11 (Kaimogool South) Kenya Serravallian Nyakach 11(Kaimogool South) Kenya Serravallian Nyakach 8 (Kadianga West) Kenya Serravallian Nyakach 8 (Kadianga West) Kenya Serravallian Ngorora Kenya Serravallian-Tortonian
MN9 (11.2-9.5Ma)
Locality Country Age Götzendorf Austria MN9 Mariathal Austria MN9 Vösendorf Austria MN9 Suchomasty Czech Republic MN9 Doué-la-Fontaine France MN9 Jujurieux France MN9 Priay II France MN9 Udabno I Georgia MN9 Eppelsheim Germany MN9 Esselborn Germany MN9 Hammerschmiede Germany MN9 Höwenegg Germany MN9 Melchingen Germany MN9 Wartenberg Germany MN9 Wissberg Germany MN9 Rudabánya Hungary MN9 Sümeg Hungary MN9 Atavaska Moldova MN9 Buzhor 1 Moldova MN9 Kalfa Moldova MN9 Varnitsa Moldova MN9 Belchatow A Poland MN9 Ballestar Spain MN9 Can Llobateres I Spain MN9 Can Ponsic Spain MN9 Can Ponsic I Spain MN9 El Firal Spain MN9 Hostalets de Pierola Superior Spain MN9 Los Valles de Fuentidueña Spain MN9 Nombrevilla Spain MN9 Pedregueras 2A Spain MN9 Pedregueras 2C Spain MN9 Peralejos 5 Spain MN9 Sant Miquel de Taudell Spain MN9 Santiga Spain MN9
Locality Country Age Eichkogel-upper Austria MN10 Kohfidisch Austria MN10 Ambérieu 2A France MN10 Ambérieu 2C France MN10 Douvre France MN10 Lo Fournas 1993 France MN10 Montredon France MN10 Soblay France MN10 Salmendingen Germany MN10 Nikiti 1 (NKT) Greece MN10 Nikiti 2 (NIK) Greece MN10 Pentalophos 1 (PNT) Greece MN10 Pyrgos Vassilissis Greece MN10 Ravin de la Pluie (RPL) Greece MN10 Xirochori1(XIR) Greece MN10 Poksheshty Moldova MN10 Can Purull Spain MN10 La Cantera Spain MN10 La Roma 2 Spain MN10
65
La Tarumba I Spain MN10 Los Aguanaces 5A & B Spain MN10 Masia de la Roma 11 Spain MN10 Masia de la Roma 3-9 Spain MN10 Masía del Barbo Spain MN10 Masía del Barbo 2 Spain MN10 Masía del Barbo2B Spain MN10 Puente Minero 2 & 8 Spain MN10 Terrassa Spain MN10 Gülpinar Turkey MN10 Yulafli (CY) Turkey MN10 Sinap 49 Turkey C4AR.1N Sinap 12 Turkey C4AR.2N Sinap 84 Turkey C4AR.2R Eldari I Georgia 10.1-9Ma Udabno II Georgia MN9-MN10 Awash 1 Ethiopia MN9-MN10 Nakali Kenya MN9-MN10 Sig 1 Algeria MN9-MN10 Feid El Atteuch R1 Algeria MN10 Tafna Algeria MN10
MN11 (9-8.2Ma)
Locality Country Age Ambérieu 1 France MN11 Ambérieu 3 France MN11 Bernardière France MN11 Dionay France MN11 Lobrieu France MN11 Mollon France MN11 DornDürkheim1 Germany MN11 Halmyropotamos (HAL) Greece MN11 Halmyropotamos (HAL)+A90 Greece MN11 Pikermi Greece MN11 Prochoma Greece MN11 Ravin des Zouaves 5 Greece MN11 Samos Greece MN11 Samos (A-1) Greece MN11 Vathylakkos 3 (VAT) Greece MN11 Csakvar Hungary MN11 Baccinello V1 Italy MN11 MonteBamboli Italy MN11 Alfambra Spain MN11 Crevillente 2 Spain MN11 La Gloria 10 Spain MN11 Los Aguanaces Spain MN11 Los Aguanaces 3 Spain MN11 Masada Ruea 2 Spain MN11
biradiculus) and sciurid rodents (Heteroxerus grivensis, H. rubricati), in addition to an
erinaceid insectivore (Mioechinus butleri). The final province recognized in the small
mammal species is comprised of faunas from African localities. This province lacks any
shared species with Eurasian localities.
At the genus level, there is considerably less resolution in the definition of small
mammal faunal provinces due to the preponderance of cosmopolitan genera (Figure 3.3d).
For example, there are many small mammal genera that are endemic to Turkey, including
murid, glirid, pteromyid, sciurid, zapodid (jumping mice, birch mice and jerboas), and
ctenodactylid rodents. Although these endemics serve to group Paşalar and Çandır, other
Turkish localities are not as tightly clustered (i.e., Koçgazi and Karaağaç 1). A similar
condition is evident in the Spanish localities, which are dispersed throughout the dendrogram
75
because of a larger proportion of cosmopolitan taxa and only two endemic genera
(Fahlbuschia and Armantomys). The more central European province, which in other
analyses shared common taxa with Germany, France, Austria, Switzerland, the Czech
Republic and Poland, is maintained by the MN5 small mammal genera and is well-supported
by several castorid (beaver), pteromyid, glirids, murid and sciurid rodents, as well as soricid,
erinaceid and dimylid insectivores. Similarly, the African province is well maintained by a
large number of endemic genera, however, the cosmopolitan erinaceid insectivore, Galerix,
which is widespread in Europe, is also known from two of the Rusinga Island localities in
Kenya. The erinaceid insectivore, Amphechinus, is common to both localities in Eurasia
(Georgia and Germany), Kenya (several of the Rusinga localities) and Namibia. The
ctenodactylid rodent, Sayimys, is also present at localities in Turkey and Saudi Arabia.
76
Figure 3.3a: MN5 large mammal species dendrogram
Ad Dabtiyah SA Napak (IR) KE Arrisdrift NA Kipsaramon KE Kipsaramon KE Maboko KE Majiwa KE Ombo KE Kalodirr KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Sinda Congo CO/ZA Napak UG La Condoue FR Chios GR Antonios GR Belometchetskaja RG Çandır 3 TU Paşalar TU Ardiç-Mordoğan TU Walda 2 GE Griesbeckerzell GE Hambach 6C GE Ziemetshausen 1b GE Savigné-sur-Lathan FR Contres MN5 FR Wannenwaldtobel-2 GE Baigneaux-en Beauce FR Pontlevoy FR Esvres - Marine Faluns FR Göriach AU Heggbach GE Rothenstein 1 GE Poudenas FR Rimbez FR Sant Mamet SP Engelswies GE Sandelzhausen GE Häder GE Puente de Vallecas SP La Hidroelectrica SP Montejo de la Vega SP Grund AU Faluns of T & A FR Manthelan FR Laimering 3 GE Gisseltshausen GE Napak KE
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 similarity
8
16
24
32
40
48
56
cc=0.91
77
Figure 3.3b: MN5 large mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Grund AU Laimering 3 GE Walda 2 GE Griesbeckerzell GE Wannenwaldtobel-2 GE Heggbach GE Contres MN5 FR Sant Mamet SP Savigné-sur-Lathan FR Poudenas FR Rimbez FR Häder GE Puente de Vallecas SP Engelswies GE La Hidroelectrica SP Rothenstein 1 GE Göriach AU Hambach 6C GE Baigneaux-en Beauce FR Esvres - Marine Faluns FR Pontlevoy FR Sandelzhausen GE Faluns of T & A FR Montejo de la Vega SP Antonios GR Al Sarrar SA Ad Dabtiyah SA Çandır 3 TU Paşalar TU Belometchetskaja RG Ardiç-Mordoğan TU Karaağaç 1TU Gisseltshausen GE Chios GR Kipsaramon KE Maboko KE Majiwa KE Ombo KE Napak KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Kalodirr KE Rusinga KE Kipsaramon KE Arrisdrift NA Napak UG Ziemetshausen 1b GE La Condoue FR Manthelan FR Napak KE Sinda Congo CO/ZA
similarity
8
16
24
32
40
48
56
cc=0.81
64
78
Figure 3.3c: MN5 small mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
8
16
24
32
40
48
56
Arrisdrift NA Kipsaramon KE Kipsaramon KE Kalodirr KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Napak UG Rusinga KE Rusinga KE Rusinga KE Napak UG Napak UG Al Sarrar SA Sant Mamet SP Montejo de la Vega SP Esvres - Marine Faluns FR Contres MN5 FR Vermes 2 SW Çandır 3 TU Paşalar TU Somosaguas-Sur SP Teiritzberg 1 AU Obergänserndorf 1 & 2 AU Schellenfeld 2-4 GE Engelswies GE Vermes 1 SW Massendorf GE Rothenstein 1 GE Franzenbad CZ Belchatow PO Sandelzhausen GE Affalterbach GE Gisseltshausen GE Puttenhausen GE Laimering 3 GE Hambach 6C GE Tobel-Hombrechtikon SW Martinsbrünneli SW Chatzloch SW Hüllistein SW Belometchetskaja RG Ziemetshausen 1b GE Wannenwaldtobel-2 GE Frohberg SW Muhlbach AU Grund AU Antonios GR Göriach AU Häder GE Heggbach GE Manthelan FR Savigné-sur-Lathan FR Pontlevoy FR Rimbez FR
similarity
cc=0.90
79
Figure 3.3d: MN5 small mammal genera dendrogram
8
16
24
32
40
48
56
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Al Sarrar SA Rusinga KE Rusinga KE Rusinga KE Napak UG Napak UG Rusinga KE Kalodirr KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Napak UG Rusinga KE Rusinga KE Rusinga KE Rusinga KE Rusinga KE Arrisdrift NA Kipsaramon KE Kipsaramon KE Heggbach GE Manthelan FR Savigné-sur-Lathan FR Çandır 3 TU Paşalar TU Karaağaç 1 TU Muhlbach AU Grund AU Antonios GR Belometchetskaja RG Montejo de la Vega SP Ziemetshausen 1b GE Wannenwaldtobel-2 GE Sandelzhausen GE Laimering 3 GE Teiritzberg 1 AU Obergänserndorf 1 & 2 AU Massendorf GE Tobel-Hombrechtikon SW Puttenhausen GE Vermes 1 SW Engelswies GE Gisseltshausen GE Martinsbrünneli SW Schellenfeld 2-4 GE Franzenbad CZ Hambach 6C GE Belchatow PO Chatzloch SW Esvres - Marine Faluns FR Affalterbach GE Sant Mamet SP Vermes 2 SW Frohberg SW Rimbez FR Contres MN5 FR Rothenstein 1 GE Hüllistein SW Somosaguas-Sur SP Pontlevoy FR Göriach AU Häder GE Koçgazi TU
similarity
64 cc=0.91
80
MN6
In contrast to MN5, the MN6 large mammal species do not form a distinct Turkish
cluster (Figure 3.4a). Although there are a number of species endemic to the region
(including several bovid species, a suid and palaeochoerid, a rhino and a mammutid), the
lack of association in the Turkish localities is due to the combined influence of low species
richness and a lack of shared species among localities, coupled with the presence of
cosmopolitan taxa. The MN6 Spanish faunas are similar in this respect. Although several
carnivore species from this region are shared with France and Slovakia, the low richness of
these faunas and presence of cosmopolitan taxa complicates the placement of the Spanish
locality in the large mammal species dendrogram. This locality is grouped with the single
Turkish locality based on the presence of one shared rhino, Alicornops simorrensis. A
Greek cluster is not recognized for MN6 due to a lack of large mammal genera known from
this region during the MN6 interval. The Central European province of MN5 continues to be
supported during MN6, although by a smaller number of species (cervids and suids).
However, smaller clusters of localities are evident. These include clusters formed by
Slovakian faunas (supported by species of suid, Aureliachoerus aurelianensis; deinotheriid,
pliopithecid, Epipliopithecus vindobonensis; mustelid, Lartetictis dubia and Trocharion
albanense; and ursid, Ursavus brevirhinus), French-German-Swiss faunas (supported by
species of bovid, Eotragus clavatus; cervid, Euprox furcatus; moschid, Micromeryx
flourensianus; rhino, Lartetotherium sansaniensis; pliopithecid, Pliopithecus antiquus; and
deinotheriid, Deinotherium giganteum) and Austrian faunas (supported by species of bovid,
Tethytragus langai; and pliopithecid, Plesiopliopithecus lockeri). The large mammal species
81
from the Saudi Arabian group more closely with the Eurasian faunas, due to the shared
gomphotheriid, Gomphotherium angustidens, which is known from France and Turkey.
There are no common taxa between this region and Africa, nor are there any shared large
mammal species between Africa and Eurasia during this temporal interval.
The Turkish large mammal genera in MN6 form weak clusters with the Central
European group, because of very few endemic genera (the bovid, Turcocerus, and rhino,
Beliajevina) (Figure 3.4b). The Turkish localities are in large part comprised of more widely-
ranging taxa, which are shared with faunas in Central Europe (particularly France), Kenya
and Zaire. The Raup-Crick similarity analysis weakly groups the Zaire and Turkish faunas,
while the Jaccard and Dice analyses do not recognize the single shared widely-ranging rhino,
Brachypotherium, and group the Zaire locality with other African localities. The two
Turkish localities analysed for MN6 do not cluster together because although they share
common genera, these genera are widespread, and no endemic taxa are shared between these
localities. Large mammal genera from Greece are poorly sampled for this temporal interval
and are thus excluded in the MN6 analysis. The Spanish fauna weakly groups with French
and Turkish localities based on the co-occurrence of suid (Listriodon) and rhino (Alicornops)
taxa, in addition to several carnivore genera (including the felid, Pseudaelurus, also known
from Slovakia; hyaenid, Protictitherium; nimravid, Sansanosmilus, also known from
Slovakia; and ursid, Plithocyon). The Central European province is maintained and well-
supported by a diversity of large mammals, however again includes smaller sub-groupings
supported by their constituent taxa. These include a French-German-Swiss-Austrian
grouping and a Slovakian grouping. The former is supported by several rhino genera
(Lartetotherium, Plesiaceratherium, Prosantorhinus), a bovoid (Amphimoschus), cervid
82
(Stehlinoceros), cainotheriid (Cainotherium) and pliopithecid (Pliopithecus). The latter is
supported by mustelid genera (Trocharion, Lartetictis), an ursid (Ursavus), rhino
(Hoploaceratherium), equid (Hippotherium) and pliopithecid (Epipliopithecus). The African
localities share the tragulid, Dorcatherium; gomphotheriid, Gomphotherium; suid,
Albanohyus; and rhinos, Aceratherium and Dicerorhinus, with European faunas, as well as
the rhino, Brachypotherium, and the hominoid, Kenyapithecus, with Eurasian faunas. These
localities share the gomphotheriid, Choerolophodon, with Turkish faunas. The African
localities also share several giraffid and bovid genera exclusively with faunas from Saudi
Arabia.
The MN6 small mammal species create clusters that are very similar to the large
mammals (Figure 3.4c). Due to the lack of small mammal faunas at this time period, the
Turkish province is again poorly differentiated and groups with Swiss faunas based on the
shared castorid, Chalicomys jaegeri (also known from Germany). Similarly, the Spanish
small mammal species are represented in MN6 at a single locality. While comprised of
numerous endemic species, including murid, sciurid and glirid rodents, soricid insectivores,
an ochotonid lagomorph and a castorid, this locality clustered weakly with the Central
European group, based on a single shared soricid insectivore, Miosorex grivensis, which is
also found in France. The lack of small mammals from Greece excludes this region from
analysis. The small mammal species during MN6 form a broad Central European cluster,
which subdivide into well-supported smaller groupings. These include a group formed by
Slovakian species, which despite having many endemics (glirid, murid and pteromyid
rodents; erinaceid, soricid and talpid insectivores), also shares taxa with Hungary (murid,
Anomalomys gaudryi, Eumyarion latior; and glirid rodents, Muscardinus sansaniensis; and
83
dimylid insectivore, Plesiodimylus chantrei), Romania (glirid, Muscardinus sansaniensis),
Germany (murid, Eumyarion weinfurteri; and eomyid rodent, Keramidomys carpathicus),
France (murid, Cricetodon sansaniensis, Democricetodon gaillardi, Eumyarion latior,
Megacricetodon schaubi, Neocometes brunonis; and glirid rodents, Bransatoglis astaracensis
and Muscardinus sansaniensis; dimylid insectivore, Plesiodimylus chantrei; erinaceid,
Lanthanotherium sansaniensis; and talpid, Talpa minuta) and Switzerland (murid rodent,
Democricetodon gaillardi; and dimylid insectivore, Plesiodimylus chantrei). Similarly, the
smaller French cluster, supported by glirid, murid and pteromyid rodents, as well as
erinaceid, heterosoricid and soricid insectivores, shares species with Slovakia, Hungary,
Romania, Switzerland and Germany. The Swiss cluster is formed by fewer endemic species,
including murid, eomyid and sciurid rodents and a soricid insectivore, however the raw data
indicates that the majority of fauna from this region are also shared with France, Germany,
Slovakia, Hungary and Romania. Similar to the large mammal species, the African small
mammals have no species in common with Eurasia or Saudi Arabia.
The MN6 small mammal genera broadly support the species-level observations
(Figure 3.4d). The Turkish localities do not cluster together because although they each
share common taxa with regions to the west, they lack any common genera with each other.
Greek small mammal genera are not known for MN6. The Central European province is still
recognizable and the fauna still support a Slovakian, French and Swiss cluster, however few
endemic genera support these groupings and more commonly, localities in these regions are
comprised mostly of genera known to the Central European region. The African faunas
share no common taxa with either Eurasia or Saudi Arabia, although the latter shares a
sciurid rodent, Atlantoxerus, with Turkish faunas.
84
Figure 3.4a: MN6 large mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
4
8
12
16
20
24
28
Diessen am Ammersee GE Kreutzlingen SW Stein am Rhein SW Four FR Trimmelkam AU Steinberg GE Sansan FR Simorre FR
Castelnau-d'Arbieu FR Inönü I TU Arroyo del Val SP Catakbagyaka TU Liet FR
Rümikon SW Stätzling GE Thannhausen GE Klein Hadersdorf AU
Devínská Nová Ves SL Devínská Nová Ves SL Devínská Nová Ves SL Elgg SW Al Jadidah SA Fort Ternan KE Fort Ternan KE Ngorora KE Kipsaramon KE Kipsaramon KE Maboko KE Majiwa KE
Ombo KE Nyakach KE Nyakach KE Nyakach KE Sinda Congo ZA/CO
similarity
32
cc=0.89
85
Figure 3.4b: MN6 large mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Al Jadidah SA Ngorora KE Kipsaramon KE
Maboko KE Majiwa KE Ombo KE Nyakach KE Nyakach KE Nyakach KE Fort Ternan KE Fort Ternan KE Kipsaramon KE Sinda Congo ZA/CO Devínská Nová Ves SL Elgg SW Trimmelkam AU
Sansan FR Devínská Nová Ves SL
Devínská Nová Ves SL Liet FR Rümikon SW
Stätzling GE Thannhausen GE Klein Hadersdorf AU Castelnau-d'Arbieu FR Simorre FR Arroyo del Val SP Inönü I TU Catakbagyaka TU
Four FR Wissholz SW
Steinberg GE Diessen am Ammersee GE Kreutzlingen SW Stein am Rhein SW Armantes 7 SP
4
8
12
16
20
24
28
32
36
cc=0.87
similarity
86
Figure 3.4c: MN6 small mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Devínská Nová Ves SL Devínská Nová Ves SL
Four FR Four FR Matraszolos 1-2 HU Schwamendingen SW Rümikon SW Zeglingen SW Samsonhaza HU Wissholz SW Steinberg GE Unterneul GE
Sansan FR Armantes 7 SP Liet FR Subpiatrã 2/1R RO
Stätzling GE Catakbagyaka TU Elgg SW Al Jadidah SA Kipsaramon KE Kipsaramon KE
similarity
2.5
5
7.5
10
12.5
15
17.5
20
22.5 cc=0.92
87
Figure 3.4d: MN6 small mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Sansan FR Devínská Nová Ves SL Four FR Four FR Rümikon SW Wissholz SW Zeglingen SW Samsonhaza HU
Unterneul GE Steinberg GE Liet FR
Devínská Nová Ves SL
Matraszolos 1-2 HU Schwamendingen SW
Subpiatrã 2/1R RO
Bagici TU Stätzling GE
Elgg SW
Catakbagyaka TU Al Jadidah SA Kipsaramon KE Kipsaramon KE
similarity
2.5
5
7.5
10
12.5
15
17.5
20
22.5 cc=0.95
88
MN7/8
During this temporal interval, the large mammal species support a weak cluster of
some, but not all Turkish localities (Figure 3.5a). The Yeni Eskihisar and Yeni Eskihisar 1
localities cluster on the basis of few shared species, but interestingly other Turkish localities
group with the Tunisian locality of Bled Douarah on the basis of the shared gomphotheriid,
Gomphotherium angustidens, and the bovid, Protoryx solignaci. Although these localities
preserve a number of endemic species, including mustelids, felids, hyaenids, rhinos, and
bovids, they also contain a number of more cosmopolitan taxa. The other Turkish localities
such as Sofca cluster with those in Germany, while Sariçay clusters with Spain, due to shared
species, including the rhino, Brachypotherium brachypus, in the former and the hyaenid,
Protictitherium crassum, in the latter. Both localities share the chalicotheriid,
Chalicotherium grande, with Germany, France and Spain. Sofca and Sariçay lack any
common large mammal species. Spanish large mammal species in MN7/8 are represented at
more localities than in the preceding intervals. These faunas support clusters of Spanish
localities, in addition to clustering with Turkey (mentioned above), weakly with Switzerland,
based on the moschid, Micromeryx flourensianus, and Austria, based on the cervid, Euprox
furcatus (also known from Poland and the Ukraine) and suid species, Listriodon splendens
(also known from Turkey). The Spanish clusters are supported by bovid (Miotragocerus
chalicotheriid (Chalicotherium), felid (Machairodus) and hyaenid (Percrocuta) genera,
which are all to varying degrees widespread.
The MN7/8 small mammal species from Turkey support a distinct clustering of
localities (Figure 3.5c). This cluster is supported by several murid rodent species (Byzantinia
bayraktepensis, B. eskihisarensis, Myocricetodon eskihisarensis), talpid (Desmanella
cingulata, Desmanodon major) and erinaceid insectivores (Mioechinus tobieni, Schizogalerix
anatolica). Moreover, this cluster shares very few taxa in common with European faunas,
with only two murid (Megacricetodon similes, Neocometes brunonis) and one pteromyid
rodent species (Albanensia albanensis) in common with Switzerland. One exclusion to the
Turkish cluster is Bayraktepe 1, which has only one species in common with the Turkish
cluster of localities, the murid, Byzantinia bayraktepensis. Bayraktepe 1 groups with Spain
and Poland (Jaccard and Dice) and Poland (Raup-Crick), based on the common occurrence of
the castorids, Chalicomys jaegeri (also known from Germany), and Trogontherium minutum
(also known from Switzerland and Hungary). The Spanish small mammal species in MN7/8
91
form a much more distinct cluster than in the previous interval. This cluster is extremely
well supported by murid (Cricetodon lavocati, Democricetodon crusafonti, D. nemoralis,
Eumyarion leemanni, Hispanomys dispectus, Megacricetodon debruijni, M. ibericus) and
glirid rodents (Myomimus dehmi, Muscardinus hispanicus, M. vallesiensis, Tempestia
hartenbergeri), in addition to several soricid (Alloscapanus lehmani, Crusafontina excultus),
erinaceid (Amphechinus golpeae), talpid (Domninoides santafei, Talpa minuta, T.
vallesensis) and heterosoricid insectivore species (Dinosorex sansaniensis). Greek small
mammals are not sampled for this interval. The larger more central European province of
French-Swiss-German-Hungarian-Romanian faunas support smaller divisions of Hungarian-
Romanian localities and French-Swiss-German-Hungarian localities. The former group is
supported by murid (Eumyarion medius) and glirid (Glirulus lissiensis) rodents, in addition
to a larger number of more cosmopolitan species. The latter group is supported by eomyid
(Eomyops oppligeri), glirid (Microdyromys complicatus, Miodyromys aegercii, M.
hamadryas) and murid (Democricetodon brevis) rodents (as well as by murids and
pteromyids also found in Romania), however, still shares many species in common with the
larger Central European cluster. No African small mammals were included in the MN7/8
species-level analysis due to lack of adequate samples for this time period.
The MN7/8 small mammal genera support clear clusters of both Turkish and Spanish
localities (Figure 3.5d). The Turkish cluster is supported by several murids and sciurids, a
glirid rodent, in addition to an erinaceid and talpid insectivore. Similarly, the Spanish
localities are supported by murid, sciurid and glirid rodent, together with soricid and talpid
insectivore genera. No Greek small mammal genera were available from this temporal
interval. The previously clustered Hungary-Romania group also includes Swiss localities,
92
supported by pteromyid genera, although Germany and Hungary also share a number of
common small mammal genera. Collectively, this more central region of Europe shares a
larger number of wide-ranging genera in common with the surrounding regions (ie. Spain,
Turkey), rather than having its own exclusively endemic forms. There are no MN7/8 small
mammal genera from Africa included for this interval.
93
Figure 3.5a: MN7/8 large mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Fort Ternan KE Nyakach KE
Nyakach KE Nyakach KE Fort Ternan KE Ngorora KE
Opole 2 PO Przeworno PO Sevastopol UK Can Feliu SP Nombrevilla 2 SP
St. Stephan AU Sofca TU Massenhausen GE Poudendas-Cayron FR
Sariçay TU Can Mata 1 SP Sant Quirze SP Hostalets de Pierola In SP Castell de Barberà SP La Grive St. Alban FR
Steinheim GE Anwil SW Escobosa SP St. Gaudens FR Yeni Eskihisar TU Yeni Eskihisar TU
Bled Douarah TUN
similarity
3
6
9
12
15
18
21
24
27
cc=0.86
94
Figure 3.5b: MN7/8 large mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Felsotárkány HU Can Feliu SP St. Stephan AU Can Mata 1 SP St. Gaudens FR Nombrevilla 2 SP Opole 2 PO Anwil SW Sariçay TU Massenhausen GE Przeworno PO La Grive St. Alban FR Steinheim GE Sant Quirze SP Castell de Barberà SP Hostalets de Pierola In SP Poudendas-Cayron FR Sofca TU Fort Ternan KE Fort Ternan KE Nyakach KE Nyakach KE Nyakach KE Ngorora KE Bled Douarah TUN Escobosa SP Yeni Eskihisar TU Yeni Eskihisar TU Sevastopol UK
similarity
4
8
12
16
20
24
28
cc=0.86
95
Figure 3.5c: MN7/8 small mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Comanesti-1 RO Felsotárkány HU Bois de Raube 3 SW Felsotárkány HU
Steinheim GE La Grive St. Alban FR
Anwil SW Sant Quirze SP Hostalets de Pierola In SP Castell de Barberà SP Escobosa SP Nombrevilla 2 SP Can Feliu SP Bayraktepe 1 TU
Opole 2 PO Felsotárkány HU Felsotárkány HU Subpiatrã 2/2 RO Yeni Eskihisar TU
Yeni Eskihisar TU Pismanköy TU
Sofca TU
Sariçay TU
similarity
2.5
5
7.5
10
12.5
15
17.5
20
22.5
cc=0.86
96
Figure 3.5d: MN7/8 small mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 similarity
Felsotárkány HU Felsotárkány HU Subpiatrã 2/2 RO Sant Quirze SP Castell de Barberà SP
Anwil SW Felsotárkány HU
Sariçay TU
Opole 2 PO Hostalets de Pierola In SP
Escobosa SP
Nombrevilla 2 SP Can Feliu SP Comanesti-1 RO Bois de Raube 3 SW Steinheim GE Felsotárkány HU
La Grive St. Alban FR Bayraktepe 1 TU Yeni Eskihisar TU
Yeni Eskihisar TU
Pismanköy TU
Sofca TU
2.5
5
7.5
10
12.5
15
17.5
20
22.5
cc=0.76
97
MN9
The MN9 Turkish localities form a distinct cluster, supported by equid
(Cormohipparion sinapensis), giraffid (Palaeotragus coelophrys, also known from Georgia,
and P. roueni), rhino (Ceratotherium neumayri), and hyaenid species (Ictitherium
intuberculatum), as well as the bovid, Protoryx solignaci, which is also known from Tunisia
(Figure 3.6a). Large mammal faunas also support a Moldovan cluster of localities on the
basis of shared equid (Hipparion sarmaticum), felid (Machairodus laskarevi) and percrocutid
species (Dinocrocuta robusta). However, the Moldovan locality of Varnitsa clusters with the
Georgian localities of Udabno 1 and Eldari 1, based exclusively on the shared presence of the
bovid, Tragoportax leskewitschi, and all localities within this cluster are notably species
poor. No Greek faunas are sampled for this time period. The Spanish large mammal species
for this temporal interval support well-defined clusters, based on deinotheriid (Deinotherium
hyaenid (Plioviverrops guerini) and mustelid (Baranogale adroveri) species. The cluster of
Central European localities, restricted to Germany and Hungary, is supported by cervid
(Cervavitulus mimus), hyaenid (Allohyaena kadici), and mustelid species (Eomellivora
wimani), the latter of which is also known from the Ukraine. The Ukrainian localities cluster
on the basis of a small number of fauna, including cervid (Cervavitus variabilis), bovid
(Gazella schlosseri) and mustelid species (Promephitis maeotica), and although this region
also shares species in common with Turkey and Greece due to its easterly location, it also
shares taxa to the west, as far as Spain. The cluster of Italian localities is not surprisingly
supported by species that are completely endemic to this region, including bovid (Etruria
viallii, Maremmia hauptii, M. lorenzi, Tyrrhenotragus gracillimus), suid (Eumaiochoerus
etruscus), mustelid (Mustela majori, Paludolutra campanii, P. maremmana, Tyrrhenolutra
helbingi) and ursid species (Indarctos anthracitis), as well as the enigmatic ape,
Oreopithecus bambolii. The African localities for this interval are notably species-poor, and
although they share no common species between them, they share the giraffid, Palaeotragus
germaini, with Turkey.
Although the genus-level analysis groups localities from Turkey, there are very few
genera shared exclusively by localities in this region (Figure 3.8b). These include the bovids,
Helicotragus and Plesiaddax, in addition to the giraffid, Giraffa. In fact, the faunas from
Turkey share more genera in common with Greece, and to a lesser extent with adjacent
regions, such as Iran. Although the Greek localities contain genera endemic to the region,
there are similarly very few taxa that actually support the clusters. These taxa include the
116
bovid, Pseudotragus; the suid, Propotamochoerus; and the mustelid, Plesiogulo. Like
Turkey, the Greek faunas share more taxa in common with surrounding regions. The same is
true of the Spanish large mammal genera. Although the Spanish localities cluster together,
this is on the basis of two shared genera, the cervid, Lucentia, and the bovid, Birgerbohlinia,
together with endemic genera known from single localities and cosmopolitan taxa. Like the
species-level analysis, the Italian faunas support a cluster of localities based on the endemic
taxa, however this region shares the mustelid, Mustela, with Turkey and the more wide-
spread ursid, Indarctos, with Eurasia. Localities in the Ukraine cluster based on the shared
cervid, Cervavitus, in addition to large mammal genera shared with other Eurasian localities.
The MN11 small mammal species support clusters of Turkish and Greek localities
(Figure 3.8c). These groupings are supported by murid (Byzantinia pikermiensis, Karnimata
provocator) and hystricid species (Hystrix primigenia), in addition to endemic taxa known
from individual localities. The small mammal species from Spain support a large cluster of
localities, based on the murids, Hispanomys freudenthali and Kowalskia occidentalis. Like
the large mammal analysis, fauna from this region tend to be dominated by more widespread
species. In the Central European region, there is a clear cluster of French and French-
German localities. The French cluster is well supported by eomyid (Eomyops catalaunicus,
Graphiurops austriacus), glirid (Muscardinus austriacus), murid (Kowalskia skofleki,
Prospalax petteri, Rotundomys bressanus), and pteromyid rodents (Blackia miocaenica,
Pliopetaurista bressana), as well as erinaceid (Lanthanotherium sanmigueli), soricid
(Crusafontina kormosi, Petenyia dubia), and talpid insectivores (Archaeodesmana vinea,
Talpa gilothi).
117
The genus level analysis of small mammal faunas again support the cluster of Turkish
and Greek localities, based on the gerbillid, Pseudomeriones; the glirid, Myomimus; and the
murid, Byzantinia; in addition to genera with more widespread distribution patterns (Figure
3.8d). The small mammal genera continue to support a large cluster of Spanish localities.
These localities group on the basis of the glirid rodent, Eliomys, in addition to genera
endemic to the region known only from individual localities, and more widespread taxa. The
Central European small mammal genera support a cluster of French localities. These
localities cluster on the basis of glirid (Glirulus, Graphiurops), murid (Rotundomys) and
sciurid rodents (Tamias), and a soricid insectivore (Petenyia). Despite the clustering of these
localities, the French small mammals have more genera in common with Germany, than
within the sample of French localities. Shared genera include the castorid, Trogontherium;
eomyid, Eomyops; murid, Cricetulodon and Prospalax; and pteromyid rodents, Blackia and
Pliopetaurista; and dimylid, Plesiodimylus; erinaceid, Lanthanotherium; and soricid
insectivores, Crusafontina.
118
Figure 3.8a: MN11 large mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Samburu KE Baccinello IT
Monte Bamboli Halmyropotamos GR Samos GR Samos GR Pikermi GR Ravin des Zouaves GR Prochoma GR Vathylakkos 3 GR Maragheh IR Halmyropotamos GR Kalimanci BU Küçükçekmece TU
BalaYaylaköy TU Karacahasan TU Garkin TU
Mahmutgazi TU Çorakyerler TU Kemiklitepe TU
Kemiklitepe TU Piera SP
Crevillente 2 SP Puente Minero SP Grebeniki UK
Novo-Elizavetkovka UK Vivero de Pinos SP Dorn Dürkheim1 GE
Csakvar HU Kayadibi TU Injana IR Halmyropotamos GR
similarity
4
8
12
16
20
24
28
32
cc = 0.85
119
Figure 3.8b: MN11 large mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Baccinello IT Monte Bamboli Injana IR Kayadibi TU Küçükçekmece TU BalaYaylaköy TU Karacahasan TU Mahmutgazi TU Çorakyerler TU Ravin des Zouaves GR Prochoma GR Vathylakkos 3 GR Garkin TU Pikermi GR Halmyropotamos GR Samos GR Samos GR Kemiklitepe TU Kemiklitepe TU Maragheh IR Grebeniki UK Novo-Elizavetkovka UK Vivero de Pinos SP Piera SP Crevillente 2 SP Puente Minero SP Dorn Dürkheim1 GE cc = 0.88
3
6
9
12
15
18
21
24
27
similarity
120
Figure 3.8c: MN10 small mammal species dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Afoud 6 MOR Kemiklitepe TU Samos GR Pikermi GR Çorakyerler TU Crevillente SP Vivero de Pinos SP Los Aguanaces SP
Puente Minero SP Alfambra SP La Gloria 10 SP Los Aguanaces SP Masada Ruea 2 SP Puente Minero SP Peralejos D SP Dorn Dürkheim1 GE Dionay FR Mollon FR Ambérieu FR Ambérieu FR Bernardière FR Lobrieu FR Csakvar HU Baccinello IT
cc = 0.97
2.5
5
7.5
10
12.5
15
17.5
20
22.5
similarity
121
Figure 3.8d: MN11 small mammal genera dendrogram
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Mollon FR Lobrieu FR Crevillente SP Los Aguanaces SP Alfambra SP La Gloria 10 SP Los Aguanaces SP Puente Minero SP Masada Ruea 2 SP Puente Minero SP
Vivero de Pinos SP Peralejos D SP Baccinello IT Pikermi GR Samos GR
Kemiklitepe TU Çorakyerler TU Dionay FR Ambérieu FR Dorn Dürkheim1 GE
2.5
5
7.5
10
12.5
15
17.5
20
cc = 0.94
similarity
122
MN12
The MN12 large mammals species support a cluster of Turkish localities, based on
the shared occurrence of bovid (Gazella gaudryi and Sinotragus occidentalis), equid
(Hipparion matthewi and H. mediterraneum) and hyaenid (Ictitherium hipparionum and I.
robustum) species in this region (Figure 3.9a). A small cluster of Turkish and Iranian
localities also occurs and is supported by species of bovid (Pachytragus laticeps), equid
(Cremohipparion matthewi) and felid (Machairodus giganteus). This region also clusters
broadly with areas of Eastern Europe, based on shared taxa with Greece (Hyaenotherium
Eastern Europe/West Asia Central Europe Spain MN7/8
TU Byzantinia bayraktepensis B. eskihisarensis Myocricetodon eskihisarensis Desmanella cingulata Desmanodon major Mioechinus tobieni Schizogalerix anatolica
TU-RG Palaeotragus coelophrys Byzantinia bayraktepensis B. dardanellensis B. nikosi B. ozansoyi Heramys anatolicus Myocricetodon eskihisarensis Progonomys minus Schizogalerix anatolica S. intermedia S. sinapensis Bellatonoides eroli
Although previous studies include analysis at both the genus and species level, results
are reported for the genus level analysis only. Bernor (1978, 1983) found that clustering at
the genus level was more strongly supported and produced logical results, while clusters at
the species level tended to be weakly supported and often produced different dendrogram
topologies than the genus-level analysis. Bernor (1978) reasoned that analyses at the genus
level were more useful for studies of faunal correlation and zoogeography at the inter-
provincial scale, since paleospecies were subject to personal interpretation and had shorter
temporal and geographic ranges. Interestingly, the results of this study demonstrate the
opposite; in almost all cases, the species level analyses provided similar, if not identical
topologies to the genus-level analyses (differences were usually negligible). Furthermore,
faunal provinces were more clearly defined at the species level. Although few species were
found to support clusters during several intervals, genus-level analysis also often failed to
provide adequate support for clusters of localities. At the species level, the sample of taxa
available for analysis was decreased, due to the omission of indeterminate or cf. species
designations (i.e., Dorcatherium indet., Dorcatherium cf. D. naui). However, species level
analyses provided a precise definition of provincial boundaries, in addition to measure of
species diversity. In this analysis, clustering at the genus level was more inclusive and often
supported by taxa that were geographically widespread continentally and intercontinentally.
Although the results of this analysis at both taxonomic levels were broadly in agreement with
149
each other (and with previous studies, to varying degrees), the provincial signal from
particular taxa can be very different depending on the level of analysis. For instance, while
Dryopithecus laietanus is endemic to Spain, the genus, Dryopithecus, is considered
cosmopolitan, ranging from Spain to Georgia. Overall, the results here suggest that the
choice of genus or species level analysis depends on the specific goals of the study.
Inclusion of small mammal taxa
Previous studies of provinciality either excluded small mammals altogether (Bernor
1978, 1983) or emphasized them to a lesser degree due to potential sampling bias, in
comparison to their large mammal counterparts (Fortelius et al. 1996a). In this study, the
small mammals supported clusters of localities better than the large mammals, particularly in
MN7/8 and MN9, but also in MN10 and MN11. These results suggest that when small
mammals are adequately sampled, they may in fact provide a better indication of
provinciality than their large mammal counterparts. The small mammals generally revealed
similar, but more distinct distribution patterns than the large mammals, perhaps due to an
increased sensitivity to paleoenvironmental barriers. Sensitivity to climatic variables was
discussed in the previous chapter, but it is logical that physiogeographic barriers may also
pose a more stringent restriction on the dispersal capabilities of most small mammal taxa.
The more rapid generation and turnover rates of small mammals may also provide clues as to
why they appear to be more provincially informative. A quicker response to
paleoenvironmental change may be more visible in these taxa than in large mammals.
However, Liow et al. (2008) recently documented higher origination and extinction rates
(and thus shorter temporal ranges) in large mammal genera and species and argue that the
150
“sleep-or-hide” behaviours common in small mammals (including hibernation, torpor,
burrowing) serve as an environmental buffer contributing to higher mean survivorship.
Non-uniform geographic representation
Jernvall and Fortelius (2004) noted that the distribution of Eurasian Miocene fossil
vertebrate localities was spatially and temporally non-uniform, affecting the reliability of
geographic range estimates of fossil mammals. In the study described here, areas of Eastern
Europe are poorly represented in comparison to other regions of Europe and Western Asia.
For example, although a single Greek locality is considered during MN5, localities from this
region are not well represented again until MN10 and onwards. While only further sampling
will assist in better representation of faunas from these regions, this non-uniformity
complicates provincial divisions, particularly between Central Europe and more eastern
regions.
Terrestrial faunas within the Pannonian Basin are one example of this. Surrounded
by the Alps, Dinarides and Carpathians, the Pannonian Basin of Central Europe is a relatively
closed system and corridors for faunal interchange during the middle and late Miocene were
limited by topography and fluctuating paleoenvironments. In a previous study, Nargolwalla
(2006) found that although the Pannonian Basin represents a fairly restricted geographic area,
its constituent faunas are not more similar to each other than to faunas outside of the basin
system, contrary to expectations. Overall, the Pannonian Basin faunas did not cluster as
expected, with few localities demonstrating any similarity (i.e., in MN7/8). The bioprovinces
recognized in MN6, MN9 and MN12 in this study were consistent with previous research,
with the Austrian, Slovakian and Hungarian localities more similar to the Central and
151
Western Europe province. MN7/8 Romanian localities were more similar to Central and
Western Europe than the South Eastern province of Fortelius et al. (1996a). However,
during this interval, the Pannonian lake had regressed slightly from the Vienna Basin,
potentially introducing a corridor for faunal interchange. Similarly, faunas from the MN11
Hungarian locality of Csakvar were most similar to those from the Ukraine, perhaps due to
the presence of an interchange corridor in the region of the Eastern Carpathians, which had
not yet attained their full elevation and were still experiencing lowland conditions (Popov et
al. 2004). Overall, this may explain the similarity between Pannonian Basin localities and
those in Central and Western Europe and South Eastern province in MN7/8 and MN11,
respectively, and demonstrates that although the Pannonian Basin faunas are broadly
consistent with the bioprovincial concept outlined by Bernor (1983) and Fortelius et al.
(1996a), there are several exceptions that further sampling of taxa could clarify. It is
important to note that these conclusions are influenced by the nature of the faunal data in the
Pannonian Basin, where uneven distribution of terrestrial vertebrate localities in this region is
related to a number of factors. The intensity of research has varied over the region, and
suitable outcrops are also unevenly distributed. Further sampling could support the previous
findings, or indicate stronger provincial affinities with either regions to the west or east.
Faunal provinces in comparison to previous studies
Bernor (1983) recognized a Western and Southern European Province, comprised of
late Miocene large mammal faunas from Spain, France and Italy. Fortelius et al. (1996a)
also recognized a similar province, termed “Western Europe,” which also included Portugal.
At the species level, this analysis suggests that Spanish large and small mammal faunas are
152
distinct from those to the east from MN5 to MN9 (Figure 3.11a-c). In MN10, the species
level analysis of large mammals lacks resolution due to the considerable number of taxa
endemic to the region, but known only from single localities (i.e., not shared). During this
interval though, the small mammal species demonstrate a distinct provincial association
within Spain. In MN11, Spanish large mammals again fail to cluster with localities from the
region, but in this case it is due to the presence of cosmopolitan taxa. In fact, only the MN12
large mammal genera and the MN12 and MN13 small mammal species demonstrate distinct
similarities to France and Italy specifically. At the genus level, the similarities that the
Spanish faunas have to other regions are on the basis of more widespread taxa, rather than to
those found exclusively in France or Italy.
Next, Bernor (1983) recognized an Eastern and Central European province, composed
of late Miocene large mammals from Switzerland, Germany, Austria, Hungary, the Czech
Republic, Slovakia and Poland. In contrast, Fortelius et al. (1996a) divide west Central
Europe (Germany and Switzerland), from Austria and the Black Sea region (Hungary,
Romania, Moldova and the Ukraine). Overall, the results of this study tend to more strongly
corroborate those of Bernor (1983), however faunas from Romania, albeit poorly sampled,
are included within the Central European group (Figure 3.11a-c). Although the regions
comprising the larger Central European province cluster into small groupings, these regions
tend to share more taxa in common with the Central European province than they do with
their smaller groupings. In addition, the French faunas, which both Bernor (1983) and
Fortelius et al. (1996a) group with those from Spain and Italy (and Portugal – Fortelius et al.
1996a), share more large and small mammals in common with the rest of Central Europe
than they do with Spain for most of the time period of interest, with the exception of MN12
153
and 13. The Central European province (and smaller groupings) remains distinct from MN5
to MN7/8, after which point, the influx of more cosmopolitan taxa coupled with a low
incidence of shared endemic fauna in this region hinders provincial distinction. After MN9,
sampling becomes an issue since Central European faunas are represented at few, patchily
distributed localities. However after this point, localities from the region tend to have more
similarities to localities from Eastern Europe, likely related to drying of the fore-Carpathian
basin, which was flooded until at least the top of MN9 (Popov et al. 2004).
Mammal faunas are known from Italy from MN11 onwards. Although Bernor (1983)
and Fortelius et al. (1996a) group the Italian faunas with those from Spain and France, this
study suggests that these groupings are based on very few shared genera in MN11 and MN12
(Figure 3.11a-c). During these intervals, the Italian faunas are completely endemic at the
species level and share more taxa amongst themselves than they do with either Spain or
France at the genus level. Furthermore, the geographic and geological evidence of insular
environments in Italy (Moyà-Solà et al. 1999; Harrison & Rook 1997; Rook et al. 1999;
Rook et al. 1996) further support the recognition of these faunas belonging to their own
province. In MN13, however, this previously isolated region is invaded by more
cosmopolitan taxa. Italy shares faunas with Spain, Hungary and Greece at the species level,
but also shares even more fauna intercontinentally with Africa at the genus level.
Bernor (1983) proposed a Romanian and Western Russian province, composed of
localities in Romania, Moldova, the Ukraine and Georgia, as well as a Sub-Paratethyan
province of localities in Turkey, Greece and Iran. Fortelius et al. (1996a) described similar
Balkan (Slovenia, Croatia, Bosnia, Serbia, Macedonia, Albania, Greece and Bulgaria) and
Anatolian provinces (Anatolia, Samos and Georgia). The results of my analysis generally
154
support those of Bernor (1983) (Figure 3.11a-c). Despite geographic proximity, Turkish and
Georgian faunas rarely cluster and only weakly when they do. Turkish faunas tended to be
more similar to themselves, however often displayed low speciosity, few endemics during
certain intervals and a higher proportion of more widespread taxa (i.e., MN6, MN7/8 large
mammals). However, during MN5, MN7/8 (small mammals) and MN9, the Turkish faunas
support a provincial distinction, strongly supported at the species level. At the genus level,
particularly in MN9, this region shares cosmopolitan taxa continentally with the rest of
Europe, as well as sharing taxa with Algeria and Morocco. Similar to Bernor (1983), the
Georgian and Moldovan localities cluster in MN9, but only weakly (based on species-poor
localities). From MN10 to MN12, the Turkish large mammals share more similarities to
Greece, which is finally becoming well sampled during this and successive intervals.
Although these regions share large mammals with those in Bernor’s (1983) Sub-Paratethyan
province, MN11 localities in Moldova (Bernor’s Romanian and Western Russian province)
also share common taxa with Turkey and Greece, as well as regions to the west. However,
MN11 small mammals support an exclusive Greek and Turkish cluster. In MN12, Turkish
and Greek faunas support both a Sub-Paratethyan province, as well as sharing taxa with the
Romanian and Western Russian province (Moldova and Bulgaria). Interestingly, the MN12
small mammal species from Turkey share no taxa in common with Greece or Moldova,
although at the genus level, these faunas group very weakly with those in Greece. Turkish
and Greek faunas cluster only weakly in MN13 at the species level of large mammals, and at
the genus level of large mammals, the lack of endemics, coupled with the preponderance of
cosmopolitan taxa, link this region continentally with the rest of Europe and
155
intercontinentally with localities in Africa. At the species level, the small mammals weakly
group the Turkish and Greek localities.
Lastly, while Bernor (1983) recognized a North African province, Fortelius et al.
(1996a) limited their analysis to Eurasian localities. For the purposes of this analysis,
localities from the expanse of the African continent were lumped into a single province since
they were not the prime focus of this study (Figure 3.11a-c).
Figure 3.11: Geographic distribution of faunal provinces
A. Modified from Bernor (1983)
1 Western and Southern European Province 2 Eastern and Central European Province 3 Romanian and Western USSR Province 4 Sub-Paratethyan Province 5 North African Province
1
23
4
5
156
B. Bioprovinces recognized by Fortelius et al. (1996)
1 Western Europe 4 the Black Sea 2 west Central Europe 5 the Balkans 3 Austria 6 Anatolia
C. Provinces recognized in this study
1 Spanish province 3 Italian province 2 Central European province 4 East European/West Asian Province
1
2 3 4
5
6
1
2
3
4
157
Faunal provinces: In situ evolution and dispersals
This study, together with previous research (i.e., Agustí et al. 1999a; Benammi et al.
1996; Bernor 1983; Made 1999; Steininger et al. 1985; Thomas 1985; Thomas et al. 1982),
identifies the taxa involved and temporal occurrences of intercontinental dispersals between
Eurasia and Africa at various intervals during the late early, middle and late Miocene.
Similarly, patterns of in situ evolution and dispersal have been recognized for specific
mammalian groups and for specific geographic regions in Europe and Western Asia (i.e.,
Agustí et al. 1996b; Bernor et al. 1996c; Fortelius et al. 2003b; Lindsay 1989; Rössner &
Heissig 1999). However, a large-scale, comprehensive account of bioprovincial evolutionary
events is currently lacking. The faunal provinces, as defined in this analysis, provide the
opportunity for evaluation of their constituent faunas and changes in these faunas over time,
allowing for a characterization of continental trends in faunal evolution. These trends are
described below and episodes of dispersal and in situ evolution are summarized in figures
3.12a-h. These figures do not include taxa previously known to a region. Location of origin,
FA or distribution in previous intervals is included in brackets (see map legends).
1. The Spanish Province:
MN5 Spanish proboscideans, although African in origin, are known in Western
Europe prior to this interval (FA MN3/4) (Göhlich 1999). The palaeomerycids are both
known from earlier intervals in Eurasia, as are the cervids and bovids, both of which have
Asiatic origins (MN3 and MN4, respectively) (Gentry et al. 1999). Although the small
mammals known from this region originate in Asia or Eurasia, all taxa are known from
previous intervals in Spain (Daams 1999; de Bruijn 1999; Kälin 1999; Ziegler 1999).
158
In MN6, Agustí (1999) notes several important events in Spain coincident with this
interval, including the replacement of the suid, Bunolistriodon, by the Asian, Listriodon, in
addition to the first appearances of taxa arriving from the east, including the hyaenid,
Protictitherium, bovids, Tethytragus and Hispanomeryx, and the cervid, Euprox.
During MN7/8, several taxa comprising the Spanish province are already known to
the region from previous intervals. These include the previously mentioned, Protictitherium,
and the deinotheriid, Deinotherium. The bovid, Miotragocerus, may also be included in this
group, however while Agustí (1999) recognizes this taxon from MN5 in Spanish deposits,
Gentry et al. (1999) proposes MN7/8 for the entry of this taxon into the region. MN7/8 also
marks the immigration of the bovid Protragocerus (Gentry et al. 1999), the nimravid,
Sansanosmilus jourdani (Ginsburg 1999) and mustelids, Palaeomeles and Trocharion.
Interestingly, this interval documents the arrival in Spain of taxa known from previous
intervals in Europe. These include the suid, Albanohyus (Hünermann 1999); the rhinos,
Hoploaceratherium and Lartetotherium (Heissig 1999a), and hominid and pliopithecid
primates. This pattern of arrival suggests that the Pyrenees were acting as a filter-type
barrier that restricted these taxa from entering prior to MN7/8, particularly since some of
these taxa were known adjacent to the mountains from MN5. Of the small mammals known
for this interval, all of the cricetid rodents (Cricetodon, Megacricetodon, Democricetodon,
Eumyrarion) are previously known in Spain, with the one exception of Hispanomys dispectus
having entered during this interval (Agustí 1999; Kälin 1999). The glirid rodents (i.e.,
Myomimus, Muscardinus) exclusive to this province also are known previously, and although
Tempestia hartenbergeri first appears in this interval in Spain, it is thought to succeed the
occurrence of T. ovilis in MN5. The erinaceid insectivore, Amphechinus golpae, is known
159
from MN7/8 but the genus is known from the region since MN4 (Ziegler 1999). Similarly,
Talpa minuta is first known from MN7/8 in Spain, but T. vallesiensis and Dinosorex
sansaniensis are both known from earlier intervals in Western Europe (Ziegler 1999).
The MN9 large mammal faunas comprising the Spanish province are almost entirely
known from previous intervals in the region (Agustí 1999). Immigrants to the region,
however, include the North American Hipparion catalaunicum, entering from the east; and
the carnivores Limnonyx sinerizi, Indarctos vireti and Thaumastocyon dirus all making their
first appearances during MN9 (although Thaumastocyon is known elsewhere in Europe from
MN5) (Ginsburg 1999). Like the large mammals, the MN9 small mammals from Spain are
mostly known from previous intervals. However, Progonomys hispanicus and potentially
Cricetulodon sabadellensis both have first appearances during this interval in Spain. It is
disagreed upon, however, whether the latter taxon originated from Democricetodon, or
immigrated from the East (Agustí 1999; Agustí et al. 1997; Kälin 1999). According to
Agustí (1999), the late Aragonian (MN7/8) and earliest Vallesian (MN9) deposits in the
Vallès-Penedès Basin share such a similarity in large and small mammal faunas that they are
biostratigraphically indistinguishable, in the absence of Hippotherium.
MN10 marks the mid-Vallesian Crisis, involving the extinction of forest-dwelling
large and small mammals and the replacement of these forms by more arid/open adapted
taxa. In Spain, small suids (i.e., Conohyus), cervids (Amphiprox), bovids (Miotragocerus
and Protragocerus), rhinos (Lartetotherium sansaniense), larger nimravid and amphicyonid
carnivores, as well as smaller carnivores (i.e., Protictitherium) undergo extinction. These
extinctions also affected most cricetids and glirid rodents that previously were abundant in
Spain. Some taxa, however, persist to the end of this interval, including hominids, larger
160
suids, Listriodon and Paraleuastochoerus, and the rodent, Cricetulodon (Agustí 1999). As a
result of these immediate and prolonged extinctions, the Spanish large mammals exclusive to
the region are known from individual localities (rather than being shared), while the small
mammals remain almost identical to previous successions.
The MN11 large mammals from Spain contain a significant number of cosmopolitan
taxa, however they also include the first appearances of the giraffid, Bigerbohlinia schaubi,
the cervid, Lucentia pierensis (Agustí 1999), and the mustelid, Baranogale adroveri.
Localities continue to preserve Hippotherium primigenium from previous intervals, as well as
the hyaenid, Plioviverrops guerini, which is considered to have evolved from the MN7/8 P.
gaudryi (Ginsburg 1999). The small mammals for MN11 are likewise mainly composed of
more cosmopolitan taxa. The two taxa shared among the Spanish localities are common in
earlier intervals (Hispanomys freudenthali and Kowalskia occidentalis) and one of these taxa
(Kowalskia), most likely evolved from a Democricetodon species (Kälin 1999). Agustí
(1999) notes that both of these taxa represent the remaining members of a previously diverse
cricetid radiation. Agustí (1999) also includes Occitanomys sondaari as a taxon common to
this province, having evolved from the late Vallesian O. hispanicus.
MN12 marks the first appearance of the bovid, Hispanodorcas torrubiae (Made
1999), the cervid, Turiacemas concudensis (Gentry et al. 1999), as well as the North
American canid, Canis cipio (Ginsburg 1999). Agustí (1999) also notes the appearances of
Palaeoryx and Gazella during this interval. These taxa combine with the large mammals that
continue their occurrences from previous intervals, including giraffid, equid, mustelid and
hyaenid species. The Spanish small mammals remain similar to the preceding interval,
however include the first appearance of the North American castorid, Dipoides, which
161
although known from Central European localities since MN11, becomes common only in
Spain in MN12 and MN13 (Hugueney 1999).
MN13 marks the entry of the cervid, Pliocervus turolensis, and bovid, Parabos
(Agustí 1999); the hippo, Hexaprotodon crusafonti (Made 1999); and the North American
canid, Nyctereutes (Ginsburg 1999). The remainder of the large mammals exclusive to the
Spanish province during this interval are known previously from the region. Small mammal
first appearances include Blancomys sanzi (Fejfar 1999) and Castillomys crusafonti. In
addition to the latter taxon, Agustí (1999) also notes the first appearances of lineages that
persist into the Pliocene, including Apodemus, Stephanomys, and ‘Cricetus.’ Interestingly,
Agustí (1999) also notes that although most of these taxa can be related to previous genera,
these taxa are considered as new immigrants, rather than evolving in situ from previous
forms.
2. The Central European Province:
All of the large mammals exclusive to the Central European province first appear
from Africa or Asia, either at the genus or species level, prior to MN5 in this region (Gentry
et al. 1999; Ginsburg 1999; Fortelius et al. 1996b; Made 1999), with a few notable
exceptions. These include the cervid, Dicrocerus elegans, and moschid, Micromeryx
flourensianus, both of which possibly descended from earlier forms in the region (Gentry et
al. 1999); the mustelid, Trocharion albanense, whose origins are uncertain since it is known
from Europe, Asia and North America at this time; and the primates, Pliopithecus antiquus
and cf. Griphopithecus, having either an Asian origin (in the former) or African origin (both),
but both arriving from the east. The Central European small mammal faunas are similar in
162
that a number of taxa exclusive to the region are known from previous intervals (Bolliger
1999; Daams 1999; Fejfar 1999). Although the insectivores, Plesiosorex germanicus and
Proscapanus sansaniensis first appear in MN5, both might be descended from species known
earlier in the region (Ziegler 1999). Both Keramidomys carpathicus and Muscardinus
sansaniensis are known to have first appearances in MN5 as immigrants arriving from the
east. Bernor et al. (1996b) also note the number of small and large mammal lineages
“holding over” from previous intervals, and particularly recognize the prevalence of relictual
small mammals in Central and Western Europe.
The MN6 large and small mammals are similarly comprised of a number of taxa
known from previous intervals. Interestingly, two of the small mammals possibly evolved
from earlier forms. Anomalomys gaudryi either migrated more westward or alternatively
evolved from A. minor in the southern Alpine region (Bolliger 1999) and Neocometes
brunonsis evolved from N. similis, known from the previous interval (Fejfar 1999). The
MN6 faunas also document a number of first appearances, including the mustelid, Lartetictis
dubia of uncertain origin (Ginsburg 1999); the cervid, Euprox furcatus, of European origin
(Gentry et al. 1999); the bovid, Tethytragus from Anatolia; and the pliopithecids,
Epipliopithecus and Plesiopliopithecus. Bernor (1983) and Made (1999) note a significant
immigration of Sub-Paratethyan and African faunas into this region at the base of MN6,
which included primates, proboscideans (Deinotherium and Platybelodon), bovids
(Protragocerus) and suids (Kubanochoerus and Listriodon).
The MN7/8 large and small mammal faunas document fewer first appearances than
MN6 or MN9. Only the mustelid, Trochotherium cyamoides, of uncertain origin and the
hominid, Dryopithecus fontani, first appear during this interval. Made (1999) also
163
recognizes the arrival of the suid, Propotamochoerus, from Asia during this interval. Among
the small mammals, Eomyops oppligeri, and Miodyromys hamadryas, have first appearances
in MN7/8 although both genera are previously known (Daams 1999; Engesser 1999).
The MN9 Central European large and small mammal faunas lack provincial
distinction, however, Bernor (1983) notes that this interval begins with a large scale
immigration of large mammals from the Sub-Paratethyan region and Asia. According to
Steininger et al. (1985), the first appearance of the equid, Hipparion, in MN9 coincided with
the immigration of Asian small mammals into Europe, including soricids, (Blarinella,
Anourosorex, Petenyiella), cricetids (Microtocricetus and Kowalskia), as well as the bovid,
Tragocerus. These authors also recognize the influence of African immigrants, including the
rhino, Diceros; hyaena, Ictitherium; and saber-toothed felid, Machairodus.
The MN10 Mid-Vallesian Crisis in Central Europe records the extinction of many of
the same forest-dwelling small and large mammal taxa as in Spain. This faunal turnover
event perhaps contributes to the lack of provincial definition in the MN10 large mammal
faunas, also exacerbated by the small number of localities sampled for this interval. Bernor
et al. (1996b) also note a significant drop in faunal diversity that likely had an effect on
provinciality in MN10. Bernor et al. (1996b) have observed a decrease in the number of
entries into this region, recognizing only the occurrence of the hyaenid, Adcrocuta eximia,
and large suid, Microstonyx, during this interval. Made (1999) additionally recognizes the
first occurrence of the African Pliohyrax in MN10. The MN10 small mammals retain a
number of taxa known from earlier intervals, however, Archaeodesmana vinea and
Graphiurops, both of uncertain origin, and Prospalax, possibly of Asian origin (Bolliger
1999), all have their first appearances during this interval. Bernor et al. (1996b) noticed that
164
the MN11 faunas from Central and Western Europe are relictual, due to the retention of more
forest adapted taxa. This is demonstrated through the first appearance of only two large
mammals, Cervavitulus mimus (Gentry et al. 1999) and Eomellivora wimani, although the
latter genus is known previously in Spain (Ginsburg 1999). The small mammal faunas from
this region are known from the previous interval and none of the taxa that are exclusive to
this region have a first appearance during MN11.
Although the Central European province lacks any endemic forms during MN12 and
MN13, Bernor et al. (1996b) indicate that the trend observed in MN11 continues into the
following intervals, however also recognize a lack of data in this region. In MN12, these
authors recognize the first appearance of the murid, Stephanomys, and in MN13, they note
the expansion of more eastern “Pikermian” faunas into Central and Western Europe (Bernor
et al. 1996b&d).
3. The Italian Province:
Large mammal taxa known from Italy document the presence of three distinct
bioprovinces, two of which (Abruzzi-Apulia and Tusco-Sardinian) preserve endemic
mammals. The third province, Calabria-Sicily, preserves land mammals also known from
North Africa (Rook et al. 2006). Although taxa from the Abruzzi-Apulia province do not
appear to have relations to other known forms, taxa from the Tusco-Sardinian province have
affinities predominantly with European mammals (Moyà-Solà et al. 1999; Rook et al. 2006,
1999). According to Rook et al. (2006), these faunas are indicative of land connections with
Europe at the beginning of the late Miocene, occurring in the region of the south-western
Alps and the northern Apennines. The non-endemic Baccinello V3 fauna also indicate
migration pathways from the northern Apennines into southern Tuscany, which are supported
165
by the distribution of localities along this route (Fine Valley, Casino, Velona and Baccinello
V3) (Rook et al. 2006).
4. The Eastern European-South Asian Province
The Eastern European/South Asian faunas from MN5 are dominated by Turkish large
and small mammals during this interval. As noted by previous researchers, the large
mammal faunas from Turkey are derived from Europe and possibly Asia, while many are
endemic to Turkey. Of the taxa endemic to the Turkish localities in MN5, the bovids,
Hypsodontus and Tethytragus are both known from Europe (MN5-MN6 in the former; MN6-
MN7/8 in the latter, Gentry et al. 1999), with the former having an East Asian origin (Gentry
& Heizmann 1996). Among the suids, although both Listriodon and Bunolistriodon are
present, Made (1999) notes that the former taxon evolved from the latter in the Indian
subcontinent. This would suggest that the replacement of Listriodon by Bunolistriodon
within Europe was not the result of in situ evolution, but rather of an immigration event.
Fortelius et al. (1996b), however, suggests that an evolutionary transition between the two
genera is unlikely, but that Listriodon also has a South Asian origin in MN5. Made (2003)
recognizes the suid, Schizochoerus, from the Paratethys area in MN5. The mustelid,
Ischyrictis, and the bovid, Turcocerus, have their first appearances in Turkey in MN5 (Begun
et al. 2003a-b; Nagel 2003). The giraffid, Giraffokeryx, originating in Africa (Gentry &
Heizmann 1996) is known from MN6 in Europe and is considered to be dentally advanced
from Georgiomeryx, occurring contemporaneously in Greece (Gentry et al. 1999). The
hyaenid, Protictitherium, is known previously in Western Europe (MN4) (Ginsburg 1999;
Made 1999), as is the cervid, Heteroprox (MN5-MN7/8) (Gentry et al. 1999). According to
Bernor et al. (1996b) the former taxon appears in Turkey after extending its initial range
166
eastward. Although Gomphotherium is known previously from Europe (MN4 in Germany,
Spain), this taxon is also known from Turkey during this interval. Begun et al. (2003a & b)
additionally observe that Griphopithecus and Orycteropus are represented in MN5 in Europe
after an initial African migration. These authors also recognize a close relationship between
Aceratherium and Plesiaceratherium from MN4-MN5, and Beliajevina and Hispanotherium,
which disappears in Europe in late MN5-early MN6. Heizmann & Begun (2001) observe a
larger number of taxa that first occur in Europe (MN4/5) likely before their occurrences at
Paşalar, including Amphicyon major, Plithocyon, felids, Pseudaelurus quadridentatus and P.
larteti, suid, Conohyus, and tayassuid, Taucanamo. Fortelius et al. (1996b) also recognize
Taucanamo from MN4 and note that the Turkish species (T. inonuensis) is derived relative to
the MN5 T. sansaniensis known from Europe. Begun et al. (2003a & b) make the same
observation and further suggest the European Bunolistriodon lockharti to be also be more
primitive than those occurring in Turkey. The Turkish small mammals are derived from a
number of regions. For example, Cricetodon, Megacricetodon, Glirulus and Schizogalerix
are all known previously in the region (de Bruijn et al. 2006; Mein 2003). Pliospalax,
Albanensia and Forsythia all have their first appearances during MN5, while the European
Oligocene genus Peridyromys (Daams 1999) and European MN3 genus, Desmanodon
(Ziegler 1999), represent the rare occurrence of taxa that disperse from west to east into
Turkey (de Bruijn et al. 2003; de Bruijn & Ünay 1996; Engesser & Ziegler 1996).
The MN6 Turkish faunas lack the provincial distinction evident in MN5. Despite
this, the bovid species remain unchanged from the previous interval, as does the
palaeochoerid, Taucanamo inonuensis. According to Made (1999), the first appearance of
Protoryx in the Turkish deposits in this interval is the result of evolution from Tethytragus
167
(which is also still present from the previous interval). Bunolistriodon splendens known
from the previous interval is replaced by B. latidens, however no phylogenetic relationship
has been suggested between these taxa. The mammutid, Zygolophodon, is known from MN4
in Turkey (Madden 1980). Similarly, the castorid, Chalicomys jaegeri is known from the
previous interval in the region, however this taxon is cosmopolitan and is known as early as
MN4 in Central Europe (Hugueney 1999).
Most of the MN7/8 large mammals shared among the Turkish localities are
previously known in the region. However, the hyaenid, Protictitherium cingulatum, first
appears in this interval (Werdelin & Solounias 1991), as does the machairodont cat,
Miomachairodus pseudailuroides, the latter originating in Africa (Hoek Ostende et al. 2006).
An indeterminate species of the palaeomerycid, Triceromeryx, also has a first appearance in
MN7/8 in Turkey, however this taxon is known from much earlier deposits in Spain (MN4)
(Janis & Lister 1985). Among the small mammals from MN7/8, two species of Byzantinia
have first appearances during this interval after possibly originating in Asia (Bernor et al.
1996b; de Bruijn & Ünay 1996; Rummel 1999). According to Wessels (1999) and Wessels
et al. (2003), the origin and migration patterns of Myocricetodon eskihisarensis are not clear,
but this taxon may have originated on the Arabian Peninsula. Although the genus,
Desmanella, is known from the late Oligocene of Europe, this taxon makes a first appearance
in Turkey in MN7/8 and likely evolved from a Central European congener and dispersed into
Turkey, together with Desmanodon (Engesser & Ziegler 1996). According to Engesser &
Ziegler (1996), Schizogalerix anatolica known from this interval forms a lineage with the
previously known S. pasalarensis from Paşalar.
168
The early Vallesian marks a number of immigrations into Turkey from Asia, Africa
and North America, in addition to genera previously known from the region. Bernor et al.
(2003) recognize the first appearance of the North American equid, Cormohipparion, at
10.692Ma, which they consider to be distinct from Hippotherium primigenium from Central
and Western Europe. The rhino, Ceratotherium neumayri, also has a first appearance in
Turkey in MN9 after immigrating from East Africa (Bernor et al. 1996b; Heissig 1999a).
The hyaenid, Ictitherium intertuberculatum, first appears in MN9, although the origins of this
taxon are unclear (Bernor et al. 1996b; Werdelin & Solounias 1996). The bovid, Protoryx, is
previously known from the region and the giraffid species, Palaeotragus coelophrys and P.
roueni are thought to succeed Giraffokeryx. Gentry & Heizmann (1996) also note that the
former giraffid is the likely ancestor of the latter taxon. Among the small mammals, a
number of taxa are previously known in Turkey. Byzantinia bayraktepensis and B. ozansoyi
are previously known, while B. dardanellensis and B. nokosi have their first appearances in
MN9 (de Bruijn & Ünay 1996; Ünay et al. 2003). Similarly, Schizogalerix anatolica is
previously known, while S. intermedia and S. sinapensis have their first appearances.
Selänne (2003) notes a gradual evolutionary change among these taxa. Among the
immigrant taxa from Asia are Progonomys minus and Bellatonoides eroli (Bernor et al.
1996b; Made 1999; Sen 2003). The latter taxon is considered to occur later than the
Hipparion datum (base of MN9) and is contemporaneous with the appearance of
Progonomys (Sen 2003).
Most of the MN10 large mammal faunas from the regions of Turkey and Greece are
known previously. However, first appearances in the region include Chilotherium kiliasi,
which originated in Asia and occurs towards the top of MN9 (Bernor et al. 1996b; Heissig
169
1996). The giraffids Bohlinia attica and Helladotherium duvernoyi are considered to have an
MN10 entry into the region (Gentry & Heizmann 1996), however Gentry (2003) suggests
that the ancestors of the former taxon are perhaps known from the Eurasian middle Miocene.
The Asian chalicotheriid, Ancylotherium, also makes a first appearance during this interval,
however entered Europe no later than the middle Miocene (Made 1999). Although the
aardvark, Orycteropus, is previously known from the region (O. seni), Heissig (1999b)
recognizes a second immigration during the Vallesian of O. gaudryi. Most of the small
mammals are previously known to the region, however the gerbillid, Pseudomeriones, is
considered the first of its kind to appear in this region after an Asian origin (Agustí &
Casanovas-Vilar 2003; Ünay et al. 2003).
Among the MN11 large mammals known from this region, most are previously
known. Giraffa has a first appearance during this interval, however Gentry et al. (1996)
suggest that this taxon is probably related to the earlier Bohlinia. Similarly, although the
genus is known from the previous interval, Chilotherium samium makes its first appearance
in MN11. All of the small mammals known for this interval in Eastern Europe and Turkey
are previously known to the region.
The MN12 large mammals are dominated by ungulates, specifically bovids, which
continue their trend of expansion and diversification from the early Turolian (Gentry &
Heizmann 1996). The large mammals are all known previously either from the region or
from more central areas of Europe. An exception is the first appearance of the bovid,
Sinotragus, which is known previously from China (Geraads et al. 2002). The felid,
Machairodus giganteus, is also known from the previous interval, but is thought to have
descended from the Spanish MN9 species, M. alberdiae (Ginsburg 1999). The hyaenid,
170
Hyaenotherium wongi, is known previously from MN9 in Germany (Bernor et al. 1996b).
The MN12 small mammals known to this region are also all previously known, however
Rhagapodemus and Micromys are both considered to have descended from Apodemus, also
known to the region (de Bruijn et al. 1996).
The MN13 large mammal faunas from the region are previously known, mostly from
the same region, but some from more central regions of Europe. The small mammals are
similar in this respect, however the insectivore, Deinsdorfia, makes a first appearance in
MN13 from an unknown origin (Ziegler 1999). Allocricetus also makes a first appearance,
however according to Kälin (1999) this taxon is not recognized until MN14.
171
Figure 3.12a: Dispersal and in situ evolution in Eurasian faunas (MN5)
(Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets)
* All faunas in MN5 Spanish province known from previous intervals EU Europe EA Eurasia AS Asia AF Africa
Keramidomys carpathicus (EU) Muscardinus sansaniensis (EU) Trocharion albanense (EU?) Dicrocerus elegans (EU?) Pliopithecus antiquus (AS? AF?) cf. Griphopithecus (AF) Micromeryx flourensianus (in situ) Plesiosorex germanicus (in situ?) Proscapanus sansaniensis (in situ?)
Figure 3.12b: Dispersal and in situ evolution in Eurasian faunas (MN6) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) EU Europe EA Eurasia AS Asia AF Africa
Chalicomys jaegeri (EU?) Protoryx (in situ) Bunolistriodon latidens (in situ?)
173
Figure 3.12c: Dispersal and in situ evolution in Eurasian faunas (MN7/8) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) EU Europe EA Eurasia AS Asia AF Africa Ar Saudi Arabia
Propotamochoerus (AS) Eomyops oppligeri (in situ?) Miodyromys hamadryas (in situ?)Trochotherium cyamoides (?) Dryopithecus fontani (?)
Protictitherium cingulatum (EU) Triceromeryx (EU) Desmanella (EU) Byzantinia x 2 (AS) Miomachairodus pseudailuroides (AF)Myocricetodon eskihisarensis (Ar) Schizogalerix anatolica (in situ)
174
Figure 3.12d: Dispersal and in situ evolution in Eurasian faunas (MN9) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) EU Europe EA Eurasia AS Asia AF Africa NA North America
Ictitherium intertuberculatum (EU?) Progonomys minus (AS) Bellatonoides eroli (AS) Ceratotherium neumayri (AF) Cormohipparion (NA) Palaeotragus coelophrys & P. roueni (in situ) Schizogalerix intermedia & S. sinapensis (in situ) Byzantinia dardanellensis & B. nokosi (in situ?)
175
Figure 3.12e: Dispersal and in situ evolution in Eurasian faunas (MN10) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) * mainly extinctions in Spain in MN10 EU Europe EA Eurasia AS Asia AF Africa
Figure 3.12f: Dispersal and in situ evolution in Eurasian faunas (MN11) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) EU Europe EA Eurasia A Eurasia AS Asia AF Africa
Lucentia pierensis (EU?) Baranogale adroveri (EU?) Pliovivverops guerini (in situ) Kowalskia occidentalis (in situ) Occitanomys sondaari (in situ) Bigerbohlinia schaubi
Figure 3.12g: Dispersal and in situ evolution in Eurasian faunas (MN12) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) * in situ from EU M. alberdiae EU Europe EA Eurasia AS Asia AF Africa
Figure 3.12h: Dispersal and in situ evolution in Eurasian faunas (MN13) (Taxa previously known to a region are not included. Location of origin, FA or distribution in previous interval is included in brackets) * genus known from AF & EU, P. meini from EU only EU Europe EA Eurasia AS Asia AF Africa
Canthumeryx X X X X Bunolistriodon X X Germany X X X Kubanochoerus Belometchetskaja X Dorcatherium X X X X X X X Eotragus France, Austria X Aceratherium X X? X X X? X? X Brachypotherium X X Germany, France X X X Lartetotherium X X Chalicotherium X X France X X Amphicyon X X Germany, France X X Ysengrinia France X Hyainailouros France X Pseudaelurus X X Germany, France X X X Orycteropus X Deinotherium X X France (1) X X Platybelodon Belometchetskaja X Gomphotherium X X X X
Galerix cosmopolitan
Europe X Amphechinus Germany, Georgia X X Sayimys X X
193
As previously mentioned, the Eurasian Griphopithecus localities date to MN5 and
MN6, however Engelswies is considered to be slightly younger and thus marks the first
shared species 2 (7%) 1 (14%) 3 (8%) shared genera 8 (31%) 3 (43%) 11 (33%)
Engelswies
shared species 2 (7%) 0 (0%) 2 (8%) shared genera 2 (8%) 0 (0%) 2 (6%)
Çandır/Paşalar + Engelswies
shared species 1 (3%) 0 (0%) 1 (3%) shared genera 5 (19%) 2 (29%) 7 (21%)
Unshared
shared species 24 (83%) 6 (86%) 30 (83%) shared genera 11 (42%) 2 (29%) 13 (39%)
Total
species 29 7 36 genera 26 7 33
This pattern is likely influenced by the low species richness at some of the localities,
particularly Klein Hadersdorf, and when considering the centres of origin of the taxa
represented at the MN6 Griphopithecus localities, this pattern ceases to exist almost entirely.
Of the MN6 large and small mammals at Devínská Nová Ves and Klein Hadersdorf, only
two to four taxa first appear in Turkey, while 10-13 first appear in Europe. 15 additional taxa
known from these localities (including Griphopithecus) are indeterminate, either due to
simultaneous first occurrences in Turkey and Europe or endemism, and thus determination of
their centres of origin would certainly clarify this issue. From the current faunal evidence, a
biogeographic association between cf. Griphopithecus from Engleswies and G. darwini from
the Pannonian Basin is more plausible than a close relation of the latter taxon and
Griphopithecus from Turkey.
201
A further consideration is the relationship between Griphopithecus and
Kenyapithecus, the latter recognized now as the second (less common) species (K. kizili) at
Paşalar and slightly later at Fort Ternan (K. wickeri) (Andrews & Kelley 2007; Begun 2001;
Humphrey & Andrews 2008; Kelley 2008; Kelley et al. 2008). Although the presence of a
second hominoid at Paşalar is accepted based on morphological and metric variation in the
dental sample from this locality (i.e., Andrews & Kelley 2007; Humphrey & Andrews 2007;
Kelley 2008; Kelley et al. 2008; Kelley & Alpagut 1999; Martin & Andrews 1993; Waddle
et al. 1995; Ward et al. 1999), this represents the only occurrence of sympatric large bodied
apes at a Eurasian Miocene locality. This is curious, considering that the norm is occurrence
of single ape taxa and rarely, an ape with a pliopithecoid. Previously, Begun (2002, 2001),
Begun et al. (2006a, 2003) and Heizmann & Begun (2001) suggested that K. wickeri from
Fort Ternan bridged the temporal gap between previously occurring Griphopithecus and
subsequent European taxa and that this taxon represented an early Serravallian return of
hominoids to Africa during a cycle of sea level lowering, contemporaneous with the
appearance of Griphopithecus in the Pannonian Basin. Dispersal back to Africa has been
previously shown to be supported faunally by the appearance of rodent (Tong & Jaeger
1993), ruminant (Gentry & Heizmann 1996), suid (Bunolistriodon – although this taxon is
previously known in Africa) and carnivore taxa (Thomas 1985) at Fort Ternan that are
considered to be Eurasian in origin. Interestingly, Begun (2002) and Begun et al. (2003) also
suggest that Limnopithecus legetet from Fort Ternan may also have affinities to Eurasian
pliopithecoids. From the study of bioprovinciality in chapter 3, the pattern of faunal
association between Turkey and Kenya is supported by Brachypotherium, Aceratherium and
202
Dicerorhinus. Fort Ternan and Devínská Nová Ves also share Albanohyus pygmaeus,
although this taxon is cf. at Fort Ternan.
More recently, Andrews & Kelley (2007) observed that, with the exception of minor
morphological details of the anterior dentition, the degree of similarity between the species of
Kenyapithecus from Turkey and Kenya makes distinguishing between the two taxa difficult.
Kelley et al. (2008) and Andrews & Kelley (2007) recognize a suite of synapormorphies
which, according to them, are clear evidence of a phylogenetic and paleobiogeographic link
between the earlier occurrence at Paşalar and Fort Ternan at ~14Ma. These authors suggest
that after an initial dispersal from Africa of the more primitive Griphopithecus, the more
derived Kenyapithecus dispersed back to Africa.
In sum, the results of this analysis together with previous research suggest that a
descendant of Afropithecus extended its range into Eurasia ~17Ma. The current state of the
fossil record supports a first appearance of cf. Griphopithecus at Engelswies in MN5, where
it most likely extended its range eastward thereafter. Slightly later in MN5, Griphopithecus
is recognized at Çandır and Paşalar, together with Kenyapithecus kizili at the latter locality.
Since the fossil assemblage at Paşalar is considered to represent a restricted temporal period
and approximates a natural assemblage (Andrews 1995), the occurrence of two large bodied
hominoids at this locality were likely coincident and thus represents the only occurrence of
sympatric apes in Eurasia. There is some support for the hypothesis that Griphopithecus
from Turkey represents the first appearance of this taxon in Eurasia (rather than Engelswies),
however the eastern range extension hypothesis is more supported faunally. Similarly, the
majority of fauna support a biogeographic relation link between the Central European cf.
203
Griphopithecus and G. darwini from the Pannonian Basin. However, a small number of taxa
also support an association between the latter and G. alpani from Turkey.
Dryopithecus/Pierolapithecus
Several hypotheses have been put forth with the purpose of explaining the phyletic
relations of Griphopithecus to later European hominids, specifically Dryopithecus. The last
occurrence of Griphopithecus (Klein Hadersdorf, top of MN6) immediately precedes the first
appearance of Dryopithecus (D. fontani) in the Pannonian Basin (St. Stephan, MN7/8) and
despite morphological differences in these taxa, an ancestor-descendant relationship is
possible (Agustí et al. 1999; Andrews & Bernor 1999; Andrews et al. 1996; Begun et al.
2006a). Alternatively, the appearance of Dryopithecus in Europe could represent a separate
immigration event from Africa (Agustí et al. 1999; Andrews & Bernor 1999; Made 1999).
Agustí et al. (1999) suggest an additional hypothesis, proposing that Griphopithecus and
Dryopithecus immigrated to Europe together and perhaps due to taphonomic processes,
Dryopithecus does not appear in the fossil record until much later on. The recent discovery
of Pierolapithecus catalaunicus from MN7/8 deposits in Spain must also be considered with
these hypotheses for the appearance of middle and late Miocene apes. According to Begun
(2006) and Begun et al. (2008, 2006b), this taxon is conspecific with D. fontani in France
and Austria and thus the nomen, Pierolapithecus, is a junior subjective synonym of
Dryopithecus. Begun et al. (2008) suggested that this taxon diversified into at least two
genera, one which is more endemic and one which is more cosmopolitan, following the
pattern of non-primate land mammals in their respective regions. A biogeographic
association between the latest occurring Griphopithecus and the first appearance of
204
Dryopithecus and/or Pierolapithecus can be evaluated through faunal continuity between
localities (Figure 4.3). Lastly, Begun suggested that a relative of a western Eurasian
hominid, such as Dryopithecus or Ouranopithecus, dispersed back to Africa in the late
Miocene and represents the sister group to the African ape and human lineage (Begun 2005,
2002, 2001; Begun et al. 1997). Review of the taxa involved in intercontinental dispersals
during the temporal range of Dryopithecus can potentially clarify this issue.
Figure 4.3: Distribution of Dryopithecus localities (including Pierolapithecus)
* lack of concensus Late Astaracian (MN7/8) SS St. Stephan (Austria) Dryopithecus fontani LG La Grive St. Alban (France) Dryopithecus fontani SG St. Gaudens (France) Dryopithecus fontani H de P Hostalets de Pierola Inferior (Spain) Pierolapithecus catalaunicus CV Can Vila Dryopithecus laietanus ? SQ Sant Quirze Dryopithecus laietanus ? C de B Castell de Barberà Dryopithecus laietanus ? Vallesian (MN9 - MN10) R Rudabánya Dryopithecus brancoi M Mariathal Dryopithecus brancoi? Me Melchingen Dryopithecus brancoi? T Trochtelfingen Dryopithecus brancoi? W Wissberg Dryopithecus brancoi? EF El Firal Dryopithecus fontani/crusafonti? CL Can Llobateres Dryopithecus laietanus CP Can Ponsic Dryopithecus crusafonti G Götzendorf ? S Salmendingen Dryopithecus brancoi? LT La Tarumba Dryopithecus laietanus P Polinýa Dryopithecus laietanus U Udabno Dryopithecus garedziensis
H de
SG
LSS
RMG
U
W S, *Me,
E
CL, CP
*CV, *SQ,*C de P, LT
H de P
SG
LG SS
RMG
U
W *S, *Me, *T
*E
CL, CP
*CV, *SQ,*C de B P, LT
205
In Central Europe, Made (1999) noted considerably fewer immigrations in MN7/8
than in the preceding and subsequent intervals, which would support the argument that
Dryopithecus is derived from locally occurring Griphopithecus, rather than the result of a
separate immigration event. Although relatively few taxa are shared between the MN6
Griphopithecus and MN7/8 Dryopithecus/Pierolapithecus localities in comparison to the
total number represented at these localities (13 of 109 genera, eight of 128 species), this is
almost certainly a reflection of the overwhelming taxonomic richness of the MN7/8 reference
locality, La Grive, in comparison to earlier Griphopithecus localities or Hostalets.
Nevertheless, these localities do in fact share taxa. The occurrence of Albanohyus in Spain in
MN7/8 is the result of a westward immigration into the region, which also likely included
Dryopithecus/Pierolapithecus. Despite the high degree of endemism in Spain from MN5 to
MN9, the MN7/8 Spanish faunas include approximately double the immigrants in either the
preceding or subsequent intervals (Table 4.4), and therefore, if Dryopithecus/
Pierolapithecus immigrated prior to MN7/8, it is not preserved in the MN6 deposits. Central
Europe, in contrast, shows the opposite pattern; fewer immigrations in MN7/8 (only three)
than MN6 and MN9 and instead the MN7/8 Central European province is formed largely of
taxa already known to the region (Figure 4.4). This latter observation also supports the
hypothesis that the MN7/8 occurrence of Dryopithecus in Central Europe is most likely the
result of in situ evolution from a taxon previously known to the region.
206
Table 4.4: Spanish large and small mammal FAs, MN6-MN9
Pliopithecus Sansanosmilus jourdani Palaeomeles Trocharion Hispanomys dispectus * uncertain FA – Genry et al. (1999) FA = MN7/8, Agustí (1999) FA = MN5 ** no concensus on whether this taxon represents in situ evolution from Democricetodon or whether it is an immigrant from the east (Agustí 1999, Agustí et al. 1997, Kälin 1999) Figure 4.4: Late Astaracian Eurasian immigrations
The second hypothesis of a dispersal from Africa for Dryopithecus is also supported,
albeit weakly, by the faunal data. Made (1999) identified two dispersal events during
MN7/8. Although this temporal interval was not coincident with a regression, sea levels
FA Central Europe Trochotherium cyamoides (?) Propotamochoerus (As) Dryopithecus fontani Eomyops oppligeri Miodyromys hamadryas
FA E Europe/W Asia Miomachairodus pseudailuroides (Af) Protictitherium cingulatum (Eu) Triceromeryx (Eu) Byzantinia (As) Desmanella (Eu) Desmanodon (Eu)
207
were already very low and likely did not restrict intercontinental exchange pathways. The
first, the “Tethytragus event,” at 12.5Ma involved Albanohyus and possibly Dryopithecus
entering Europe, while the hyaenid, Percrocuta, appears in Africa. Towards the end of this
interval, Made (1999) noted a “Propotamochoerus event,” in which Protoryx disperses to
Africa. In the previous chapter, Protoryx solignaci was found to be shared between Turkish
localities and the Tunisian locality of Bled Douarah, however the Turkish occurrence of this
taxon predates (MN6-MN7/8) the Tunisian occurrence (MN7/8-MN9). The proboscidean,
Tetralophodon longirostris, also possibly demonstrates the same pattern as Protoryx,
although the temporal range of this taxon is not terribly well constrained. The earliest
occurrence of T. longirostris in Europe is at Nombrevilla 9 (Spain) during the Aragonian (18-
11.2Ma), but is later known at Sant Quirze (Spain) with a more definite age of MN7/8. In
Africa, Tetralophodon cf. T. longirostris is known from Djebel Krechem el Artsouma
(Tunisia), however this locality, like Nombrevilla 9, is dated from the Vallesian to the
Turolian (11.2Ma-5.3Ma). Tetralophodon sp. is similarly known from the Chorora
Formation in Ethiopia from the Serravallian to the Tortonian (13.65-7.25Ma). Therefore, it
is possible that this taxon, together with Protoryx, dispersed to Africa towards the top of
MN7/8, however the temporal resolution of the associated localities must be refined. Despite
the significant number of taxa shared between Africa and Eurasia during MN7/8, the vast
majority are previously known in both regions and are thus uninformative in clarifying the
appearance of Dryopithecus. In addition, Central Europe during MN7/8 documents very few
immigrations (only three), supporting the hypothesis that Dryopithecus is derived from a
taxon already known to the region. It is still possible that the occurrence of Dryopithecus in
208
Europe is the result of an immigration from Africa, although very few taxa support this
hypothesis.
The third hypothesis, of an earlier immigration of Dryopithecus along with
Griphopithecus but lack of retrieval for taphonomic reasons, is particularly interesting, but
difficult to test. However, the results of the completeness analysis in Chapter 2 lend some
interesting insights. The appearance of Griphopithecus in MN5 corresponds to an interval
that is extremely well sampled with CIs between 92.5 and 98 for the large mammals and
between 90.7 and 96.6 for the small mammals, both well above the cutoff of 70. These
results would suggest that the likelihood of Dryopithecus being unsampled during this
interval following an immigration with Griphopithecus is unlikely, particularly since the
small mammals, who are more prone to taphonomic bias, are extremely complete. However,
if Dryopithecus immigrated from Africa at the base of MN6, during the extensive
interchange identified by Bernor (in Steininger et al. 1996), there is slightly more of a
possibility that it is not being sampled during this interval. Although most of the CIs remain
above the cutoff of 70, they have noticeably decreased from both the preceding and
subsequent interval (76.3-86.2 for the large mammals and 68.1-77.5 for the small mammals).
The results of the completeness analysis therefore suggest that it is unlikely that
Dryopithecus immigrated with Griphopithecus in MN5 and is not being sampled. Although
there is a slight possibility of this taxon going undetected in MN6, all measures of
completeness suggests that although there are more sampling gaps, this interval still remains
relatively well sampled, with the exception of the strict index for the small mammals.
Furthermore, in Spain, few entries (except in MN7/8) and the endemic nature of Spanish
faunas would also suggest that Dryopithecus and/or Pierolapithecus were not present in this
209
region until the influx of immigrations in MN7/8. In sum, of the three hypotheses put forth
to explain the appearance of Dryopithecus in Central Europe and Spain in MN7/8, the most
well supported scenario is that this taxon evolved from a species previously known to the
region.
In 2004, a large bodied ape was described from the upper MN7/8 locality of Els
Hostalets de Pierola and a new nomen, Pierolapithecus catalaunicus, was erected to
distinguish this taxon from other early, middle and late African and Eurasian Miocene
hominoids (Moyà-Solà et al. 2004). Based on their analysis of the preserved morphology of
this taxon, Moyà-Solà et al. (2004) proposed that Pierolapithecus represents a stem hominid.
Begun et al. (2006b) and Begun and Ward (2005), however, suggested that Pierolapithecus
is instead most likely a stem hominine and also that this taxon is conspecific with D. fontani
from Austria and France based on dental similarities, temporal and geographic proximity,
and the paleobiogeographic trends in other terrestrial mammals (Begun 2006; Begun et al.
2008, 2006b). With respect to the latter, these authors found that despite the endemism of
the Spanish faunas, 59% of the total Hostalets mammals are also shared with La Grive,
including an 82% overlap in carnivore taxa (Table 4.5). Furthermore, in the previous
chapter, Hostalets, together with Sant Quirze, was found to cluster broadly with La Grive and
Steinheim. Within Spain, Agustí (1999) identified MN8 in the Vallès-Penedès Basin through
the entry of Hispanomys, Palaeotragus, Protragocerus and Tetralophodon. Interestingly, of
these immigrant taxa, Hostalets shares Palaeotragus with other MN7/8 primate-bearing
localities, such as Castell de Barberà, and Protragocerus with Castell de Barberà and Sant
Quirze. Therefore, it seems unlikely that while having these immigrants in common,
Hostalets would receive a different genus of ape. These data, together with the influx of
210
first appearances in MN7/8 in Spain, clearly indicate faunal continuity between the Spanish
and Central European bioprovinces and therefore support the hypothesis that Pierolapithecus
is at least congeneric with Dryopithecus.
Table 4.5: MN7/8 Spanish – Central European shared taxa
Locality Hostalets (Spain)
La Grive (France)
St. Gaudens (France)
St. Stephan (Austria)
Taxon Chalicotherium grande X X X Listriodon splendens X X X Euprox furcatus X X X Albanohyus pygmaeus X X Protragocerus chantrei X X Plithocyon armagnacensis X X Pseudaelurus quadridentatus X X Pseudaelurus lorteti X X Hemicyon goriachensis X X Sansanosmilus jourdani X X Semigenetta sansaniensis X X Protictitherium crassum X X Thalassictis X X Alicornops simorrensis X X Lartetotherium sansaniensis X X Deinotherium giganteum X X The results of the previous chapter support the endemic nature of the Spanish lineage from
MN7/8 to MN9, with very few immigrations and the virtually unchanged nature of the large
and small mammal faunas. Agustí (1999) also noted that the MN7/8 to MN9 Spanish faunas
are biostratigraphically indistinguishable in the absence of Hippotherium. Across the MN9
- MN10 transition, the Spanish faunas continue to support a single endemic ape lineage in
Spain. The results presented in the previous chapter indicate that the MN10 Spanish
localities (including La Tarumba) continue to cluster together. Furthermore, comparison of
the MN10 Spanish Dryopithecus locality, La Tarumba, (Polinýa II has only a single taxon),
reveals that this locality shares 67% of its constituent species and 73% of its genera with
MN9 Spanish localities.
211
Following a re-analysis of the Spanish apes, Begun (1992b) described a new species
of Dryopithecus, D. crusafonti, from Can Ponsic in the Vallès-Penedès Basin, northeastern
Spain. Similar in age to Can Llobateres (MN9), Can Ponsic is considered to be slightly older
based on its rodent faunas. This locality is also similar to Can Llobateres in terms of its
faunal composition, although slightly less abundant, but with similar depositional and
paleoecological settings (Begun 1992b and references therein). Begun (1992b, 1991)
recognized this new species based on dental differences with other Spanish and European
Dryopithecus, including distinctive molar occlusal morphology, large and broad upper
molars and very high crowned upper central incisors with well-developed lingual pillars.
Begun (1992b) also noted similarities in two lower molars from Can Ponsic (IPS 1813 and
1816) to the El Firal mandible, which he attributed to D. cf. D. crusafonti. Begun (1992b)
observed that D. crusafonti is most similar in dental characters to D. laeitanus and dissimilar
from D. fontani and the Central European D. brancoi.
In contrast, Ribot et al. (1996) reviewed the morphology of Dryopithecus from
Vallès-Penedès, as well as El Firal, and using the range of variation observed in extant apes,
concluded that sufficient morphological or metrical distinction was currently lacking to
justify the recognition of two separate species in the samples from Can Ponsic and other sites
in Vallès-Penedès. These authors attributed all material from Vallès-Penedès to D. laietanus
and recognized D. crusafonti as a junior subjective synonym of D. laietanus. Specifically,
Ribot et al. (1996) found that lower molar cusp morphology of El Firal distinguished this
specimen from the entire sample from Vallès-Penedès and were unable to find any
morphological features that El Firal shared uniquely with Can Ponsic, to the exclusion of
other specimens from Vallès-Penedès. However, like Begun (1992b), Ribot et al. (1996)
212
also observed differences between El Firal and St. Gaudens, but noted a number of
distinctive features that the El Firal mandible shares with D. fontani that are not seen in other
Dryopithecus specimens in Spain. In conclusion, these authors concur with the majority of
prior research (i.e., Andrews et al. 1996) in recognizing the El Firal mandible as belonging to
D. fontani.
As previously mentioned, there is little change and very few immigrations from
MN7/8 to MN9 in Spain as a whole. The results from chapter 3 indicate that for large and
small mammal species, Can Ponsic groups with Can Llobateres, and for large and small
mammal genera, Can Ponsic groups overall with Spanish localities. Although the degree of
similarity between Can Ponsic and Can Llobateres is not tremendous, this is likely due to the
size of the Can Llobateres faunal list (79 versus 52 taxa). Upon closer examination, the
similarity between these localities is much more evident. Of the fauna present at Can Ponsic,
69% of species and 83% of the genera are shared with Can Llobateres (Table 4.6). El Firal,
shares 62% of its species and 75% of its genera with Can Llobateres. Although the latter
finding is almost certainly being influenced by the considerably smaller faunal list at El Firal
(16 taxa), there are still a number of taxa at El Firal that are not known at either Can Ponsic
or Can Llobateres, including Dicrocerus elegans, Amphicyon pyrenaicus and
Gomphotherium angustidens. From chapter 3, the large mammals from El Firal clustered
broadly with Spain at the species level, but with Wissberg and the Central European cluster
at the genus level. Dicrocerus elegans is shared between El Firal and Wissberg. Amphicyon
pyrenaicus is unique to El Firal and the only other occurrence of Gomphotherium
angustidens in Spain during MN9 is at Santiga. These findings indicate that the faunas from
Can Ponsic and to a lesser extent, El Firal, are similar enough to Can Llobateres to suggest
213
that, coupled with the very low immigration in MN9, it is perhaps unlikely that an additional
species of Dryopithecus (i.e., D. crusafonti) dispersed into the region. However, the
replacement of four other mammalian species perhaps due to in situ evolution (Semigenetta
sansaniensis or S. grandis to S. ripola, Dorcatherium crassum to D. jourdani or D. naui,
Hispanomys dispectus to H. thaleri and Ischyrictis mustelinus to I. petteri) suggests that
perhaps D. crusafonti evolved from Dryopithecus previously known. The presence of
several taxa that are shared with other localities in Central Europe also indicates that perhaps
El Firal is somewhat distinct from the Vallès-Penedès faunas.
Table 4.6: Shared Spanish taxa
Locality Can Ponsic I El Firal Can Llobateres Taxon
Listriodon splendens X X Parachleuastochoerus steinheimensis X X Parachleuastochoerus huenermanni X Propotamochoerus palaeochoerus X X X Dorcatherium jourdani X Micromeryx flourensianus X X Euprox dicranoceros X X Miotragocerus pannoniae X X Palaeotragus indet. X X Sansanosmilus jourdani X X Amphicyon cf. major X X Ursavus primaevus X X Protictitherium gaillardi X X Machairodus aphanistus X X Promeles indet X Indarctos vireti X X Mesomephitis medius X X Limnonyx sinerizi X Martes cf. andersoni X X Thalassictis montadai X Plesiodimylus chantrei X X Galerix socialis X X Lanthanotherium sanmigueli X X Dinosorex sansaniensis X X Talpa minuta X X Talpa vallesensis X Postpalerinaceus vireti X X Prolagus crusafonti X X Hoploaceratherium tetradactylum X
214
Locality Can Ponsic I El Firal Can Llobateres Taxon
Alicornops simorrensis X X Aceratherium incisivum X X X Dicerorhinus steinheimensis X Lartetotherium sansaniensis X X cf. Hippotherium catalaunicum X X X Chalicotherium grande X X X Tapirus priscus X X X Deinotherium giganteum X X Cricetulodon hartenbergeri X Miodyromys hamadryas X Myoglis meini X X Muscardinus hispanicus X X Albanensia cf. grimmi X X Heteroxerus cf. grivensis X Heteroxerus rubricati X Spermophilinus bredai X X Hispanomys thaleri X X Trogontherium minutum X X Miopetaurista crusafonti X X Keramidomys carpathicus X Chalicomys jaegeri X X X Dicrocerus elegans X Amphicyon pyrenaicus X Dihoplus schleiermacheri X X Gomphotherium angustidens X Tetralophodon longirostris X X Paraleuastochoerus crusafonti X
Dryopithecus brancoi is known from the type locality of Salmendingen, Germany,
and also from the large sample from Rudabánya, Hungary. According to Begun (2002)
isolated molars that may belong to this species are also known from Melchingen, Ebingen,
Trochtelfingen and Wissberg (Germany), Mariathal (Austria) and Udabno (Georgia).
Unfortunately no faunal list for Ebingen could be located and the list for Trochtelfingen
includes only the primate occurrence. In chapter 3, these localities lacked any clear
association with each other and upon further comparison of their component faunas to
Rudabánya (due to its taxonomic diversity), each had very few taxa in common. This is
perhaps due in part to the lack of species richness at these localities (i.e., only Wissberg has
215
an appreciable number of taxa [18] in comparison to the 72 taxa at Rudabánya). Therefore,
the lack of faunal associations between these localities prevents any conclusions regarding
the taxonomic identity of the specimens from Germany, Austria and particularly Georgia.
However, with regard to the latter locality, the late Astaracian marks the dispersal of several
European taxa into Eastern Europe/Western Asia, including Protictitherium cingulatum,
Triceromeryx, Desmanella and Desmanodon (MN7/8). It is therefore possible that the later
primate occurrence at Udabno 2 in MN11 (Gabunia et al. 2001) is the result of this earlier
dispersal.
The relations of Dryopithecus to living African apes have been suggested by some
researchers (Begun 2005, 2002, 2001; Begun et al. 1997; Stewart & Disotell 1998).
Interestingly, two intercontinental dispersal events occur during the temporal range of
Dryopithecus, either or both of which certainly could have involved this taxon. First, the
Astaracian (MN6-MN7/8) marks the appearance of Protictitherium, Hypsodontus,
Protragocerus, Protoryx solignaci, Percrocuta and possibly Machairodus in Africa.
Although the first three of these taxa have Asian origins, they are known from Eurasia before
they appear in Africa. Similarly, Percrocuta also has Asian origins but follows the same
pattern as the previous three taxa. The origins of Machairodus is uncertain, however this
taxon is known from Europe (MN7/8-MN9) slightly before it is known in Africa (MN9).
The early Vallesian marks a second significant and diverse dispersal to Africa (Made 1999),
which included a founding population of hipparionine horses derived from Eurasian
Hippotherium primigenium (Bernor & Harris 2003), tetraconodonine suids such as
Conohyus (represented in Africa as Nyanzachoerus) (Harris & Leakey 2003), and possibly
Sivachoerus, Progonomys, Myocricetodon and Atlantoxerus. According to Bernor & Harris
216
(2003), the first occurrence of hipparionines in Africa is at Chorora between 10.5 and 9.3Ma.
The eastern range extension of Dryopithecus into Georgia, together with two dispersal
events, both including numerous other mammalian groups, indicates that Dryopithecus or a
close relative most certainly had the opportunity to disperse along with these other groups to
Africa.
In sum, the patterns in faunal dynamics during the middle and late Miocene more
fully support the hypothesis that Dryopithecus evolved from a taxon previously known in
Central Europe, rather than arising from a dispersal into the region. Pierolapithecus is most
likely a junior subjective synonym of Dryopithecus, although this analysis was unable to
determine whether this taxon is conspecific with D. fontani. Trends in the Spanish faunas do
not support a dispersal of an additional species, D. crusafonti, into the region, however the
possible occurrence of in situ evolution in other mammalian groups does not preclude the
evolution of this species from previously occurring Dryopithecus. This analysis was unable
to provide further insight into the species allocation of isolated molars from Melchingen,
Ebingen, Trochtelfingen and Wissberg (Germany), Mariathal (Austria) and Udabno
(Georgia), due to a lack of correlation between faunas at these localities. However, it is
possible that the latter specimen is the result of an eastward migration of Central European
Dryopithecus in the late Astaracian, together with other large and small mammals. It is also
possible that Dryopithecus or a close relative dispersed to Africa during a significant
migration event in the late Astaracian and/or earliest Vallesian.
Ouranopithecus
217
Ouranopithecus macedoniensis was originally recognized from the late Vallesian of
Greece. More recently, a new species was identified in the Turolian of Turkey, O. turkae
(Güleç et al. 2007). An isolated P4 from the Chirpan District of Bulgaria at ~7Ma is also
considered to resemble Ouranopithecus (Spassov & Geraads 2008) (Figure 4.5). According
to Bernor et al. (1996b), the first appearance of Ouranopithecus is likely to be the result of an
African immigration or alternatively a vicariant lineage of MN8-MN9 Dryopithecus. The
mandible attributed to Graecopithecus freybergi has also been considered to be synonomous
with Ouranopithecus by some (Andrews et al. 1996; Made 1999; Martin & Andrews 1984;
Pilbeam 1996; Szalay & Delson 1979), while others consider features of the corpus and
molar dentition to signify a separate taxon (i.e., Begun 2002; Koufos & de Bonis 2004,
2005). The evolutionary relations of this taxon are contentious. de Bonis & Koufos (2004,
p257) consider that “if Ouranopithecus occurred in Africa, it would even be a plausible
ancestor for Australopithecus,” due to similarities with the latter, which they consider to be
synapomorphies (de Bonis & Koufos 1997). Others, however, consider these similarities to
be the result of parallel evolution in response to environmental conditions favouring hard
object consumption. Recently, two large bodied hominoids were described from the late
Miocene of Africa, Nakalipithecus nakayamai and Chororapithecus abyssinicus, of which
the morphology of the former has been likened to Ouranopithecus (Kunimatsu et al. 2007).
Through the previous analysis of the non-primate mammalian faunas from these localities,
the following questions will be addressed: is Ouranopithecus the result of an African
immigration or a vicariant lineage of earlier Dryopithecus; do faunal differences exist
between Pyrgos Vassilissis and the other Greek localities to suggest that Graecopithecus and
218
Ouranopithecus are distinct genera; and lastly, what, if any, relationship does
Ouranopithecus have with late Miocene African species?
Figure 4.5: Distribution of Ouranopithecus localities
RP Ravin de la Pluie N Nikiti 1 X Xirochori 1 P Pyrgos Vassilissis
B Chirpan, Bulgaria C Çorakyerler The Greek and Turkish Ouranopithecus localities share 25% of their large mammal
faunas and also share taxa in common with Iran, Iraq, and to a lesser extent, Bulgaria.
Establishing a biogeographic link between these faunas and those occurring earlier in Europe
is complicated by the Vallesian Crisis, such that almost all of the forest-dwelling taxa that
characterize the primate localities in MN9 have gone extinct by the late Vallesian (Agustí et
al. 2003). However, the giraffid, Decennatherium?, from Ravin de la Pluie and
Tragoportax from Nikiti 1 have close affinities with the Spanish Vallesian species (de Bonis
C B
P
RP N
X
219
& Koufos 1999). In addition, the MN9 primate localities of Can Llobateres and Rudabánya
share a number of large mammal genera and species in common with later MN10 and MN11
Greek and Turkish localities, including Aceratherium incisivum, Chalicotherium goldfussi,
2006). Furthermore, although hominines, and specifically Pan, demonstrate a clear
preference for soft fruit consumption, reliance on fallback foods in times of food scarcity is
documented in many populations (Hladik 1977; Stanford et al. 1994; Tutin et al. 1997;
Yamakoshi 1998). Therefore, dismissing the dispersal of Eurasian apes into Africa in the late
Miocene on the basis of a lack of required habitats is clearly unwise.
Nargolwalla & Begun (2005) noted an interesting trend in a significant number of
large and small terrestrial mammals that persist beyond the Vallesian crisis. Many small and
large mammal taxa were characterized by an increase in body size and/or dental complexity,
in response to the shift in ecological conditions. Therefore, perhaps thick enamel and
megadonia were key adaptations that allowed a descendent of a Eurasian ape to disperse back
to Africa in the late Miocene, in much the same way that these adaptations perhaps allowed
232
Griphopithecus to disperse and radiate in Eurasia in the early middle Miocene (Begun 2002).
Although the circumstances surrounding the origins of the Homininae remain unclear, the
late Miocene fossil record in Africa and Eurasia continues to develop. Survey and
excavation in the Black Sea region, Eastern Mediterranean, Saudi Arabia and North Africa
will undoubtedly unearth new fossils and further opportunity for more spatially and
temporally continuous paleoenvironmental reconstruction along intercontinental dispersal
crossroads. These, combined with advances in analytical methods promise fascinating
insights into questions of African ape and human origins.
233
References Cited
Aguilar J-P, WA Berggren, M-P Aubry, DV Kent, G Clauzon, M Benammi & J Michaux.
2004. Mid-Neogene Mediterranean marine-continental correlations: an alternative interpretation. Palaeogeogr Palaeoclimatol Palaeoecol 24:165-186.
Agustí J. 1999. A critical re-evaluation of the Miocene mammal units in Western Europe:
dispersal events and problems of correlation. In: Agustí J, L Rook & P Andrews, (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p84-112.
Agustí J, M Garcés & W Krijgsman. 2006. Evidence for African-Iberian exchanges during
the Messinian in the Spanish mammalian record. Palaeogeogr Palaeoclimatol Palaeoecol 238:5-14.
Agustí J, I Casanovas-Vilar & M Furió. 2005. Rodents, insectivores and chiropterans
(Mammalia) from the late Aragonian of Can Missert (Middle Miocene, Vallès-Penedès, Spain). Geobios 38: 575-583.
Agustí J, A Sanz de Siria & M Garcés. 2003. Explaining the end of the hominoid experiment in Europe. J Hum Evol 45:145-153. Agustí J & I Casanovas-Vilar. 2003. Neogene gerbils from Europe. In: Reumer JWF & W
Wessels (Eds) Distribution and Migration of Tertiary Mammals in Eurasia. A Volume in Honour of Hans DeBruijn. Deinsea 10:13-22.
Agustí J & M Antón. 2002. Mammoths, sabertooths, and hominids: 65 million years of mammalian evolution in Europe. New York: Columbia University Press. Agustí J, L Cabrera, M Garcés, W Krijgsman, O Oms & JM Parés. 2001a. A calibrated mammal scale for the Neogene of Western Europe. State of the art. Earth-Sci Rev 52:247-260. Agustí J, L Cabrera & M Garcés. 2001b. Chronology and zoogeography of the Miocene hominoid record in Europe. In: Bonis L de, GD Koufos & P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 2: Phylogeny of the Neogene Hominoid Primates of Eurasia. Cambridge: Cambridge University Press, p2-18. Agustí J, L Cabrera, M Garcés & M Llenas. 1999a. Mammal turnover and global climate
change in the late Miocene terrestrial record of the Vallès-Penedès basin. In: Agustí J, L Rook & P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p397-412.
234
Agustí J, L Cabrera, M Garcés & JM Parés, 1997. The Vallesian mammal succession in the Vallès-Penedès basin (northeast Spain): Paleomagnetic calibration and correlation with global events. Palaeogeogr Palaeoclimatol Palaeoecol 133:148-180.
Allen KMS. 1990. Interpreting Space. In: Allen KMS, SW Green & EBW Zubrow (Eds) Interpreting Space: GIS and Archaeology. New York: Taylor and Francis, p383-386. Alroy J. 1994. Appearance event ordination: a new biochronological method. Paleobiology 20:191-207. Alroy J, RL Bernor, M Fortelius & L Werdelin. 1998. The MN System: Regional or Continental? Mitt Bayer Staatsslg Paläont hist Geol 38: 243-258. Andrews P. 2007. The biogeography of hominid evolution. J Biogeogr 34:381-382. Andrews P. 1996. Palaeoecology and Hominoid Palaeoenvironments. Biological Reviews 71:257-300. Andrews P. 1995. Time resolution in the Miocene fauna from Paşalar. J Hum Evol 28:343- 358. Andrews P. 1992a. Evolution and environment in the Hominoidea. Nature 360:641-646. Andrews P. 1992b. An ape from the south. Nature 356:106. Andrews P. 1990. Owls, Caves and Fossils: Predation, Preservation, and Accumulation of
Small Mammal Bones in Caves, with an Analysis of the Pleistocene Cave Faunas from Westbury-sub-Mendip, Somerset, UK. London: British Museum of Natural History.
Andrews P & J Kelley. 2007. Middle Miocene dispersals of apes. Folia Primatol 78:328- 343. Andrews P & T Harrison. 2005. The Last Common Ancestor of Apes and Humans. In: Lieberman DE, RJ Smith & J Kelley (Eds) Interpreting the Past: Essays on Human, Primate, and Mammal Evolution. Boston: Brill Academic Publishers, Inc, p103-121. Andrews P & RL Bernor. 1999. Vicariance Biogeography and Paleoecology of Eurasian Miocene Hominoid Primates. In: Agustí J, L Rook & P Andrews (Eds)
Hominoid Evolution and Climatic Change in Europe, vol 1: Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p454-487.
Andrews P, DR Begun & M Zylstra. 1997. Paleoecology of Miocene hominoids. In: Begun
DR, CV Ward & MD Rose (Eds) Function, Phylogeny and Fossils: Miocene Hominoid Origins and Adaptations. New York: Plenum Press, p29-58.
235
Andrews P, T Harrison, E Delson, RL Bernor & L Martin. 1996. Distribution and Biochronology of European and Southwest Asian Miocene Catarrhines. In: Bernor
RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p168-207.
Andrews P & LB Martin. 1987. Cladistic relationships of extant and fossil hominoids. J Hum Evol 16:101-118. Andrews P, WR Hamilton & PJ Whybrow. 1978. Dryopithecines from the Miocene of Saudi Arabia. Nature 274:249-250. Antunes MT. 1979. Hispanotherium fauna in Iberian Middle Miocene, its importance and paleogeographical meaning. Ann Géol Pays Hellén 1:19-26. Avery DM. 2003. Early and middle Pleistocene environments and hominid biogeography: micromammalian evidence from Kabwe, Twin Rivers and Mumbwa Caves in central Zambia. Palaeogeogr Palaeoclimatol Palaeoecol 189:55-69. Badgley C. 1990. A statistical assessment of last appearances in the Eocene record of mammals. In: Bown TM & KD Rose (Eds) Dawn of the Age of Mammals in the northern part of the Rocky Mountain Interior. Geological Society of America Special Papers 243:153-167. Badgley C & PD Gingerich. 1988. Sampling and faunal turnover in early Eocene mammals. Palaeogeogr Palaeoclimatol Palaeoecol 63:141-157. Becker A. 2002. The Jura Mountains—an active foreland fold-and-thrust belt? Tectonophysics 321:381– 406. Begun DR. 2007. Fossil record of Miocene hominoids. In: Henke W & I Tattersall (Eds) Handbook of Paleoanthropology, vol II: Primate Evolution and Human Origins. New
York: Springer-Verlag, p921-977. Begun DR. 2006. Revision of the Dryopithecini. J Vert Paleontol 26(S3):40A. Begun DR. 2005. Sivapithecus is east and Dryopithecus is west, and never the twain shall meet. Anthrop Sci 113:53-64. Begun DR. 2002. European Hominoids. In: Hartwig WC (Ed) The Primate Fossil Record. Cambridge: Cambridge University Press, p339-368. Begun DR. 2001. African and Eurasian Miocene hominoids and the origins of the Hominidae. In: de Bonis L, GD Koufos & P Andrews (Eds) Hominoid Evolution and Climate Change in Europe, vol 2: Phylogeny of the Neogene Hominoid Primates of Eurasia. Cambridge: Cambridge University Press, p231-253.
236
Begun DR. 2002. European Hominoids. In: Hartwig WC (Ed) The Primate Fossil Record. Cambridge: Cambridge University Press, p339-368. Begun DR. 2001. African and Eurasian Miocene hominoids and the origins of the Hominidae. In: de Bonis L, GD Koufos & P Andrews (Eds) Hominoid Evolution and Climate Change in Europe, vol 2: Phylogeny of the Neogene Hominoid Primates of Eurasia. Cambridge: Cambridge University Press, p231-253. Begun DR. 1995. Late Miocene European orang-utans, gorillas, humans, or none of the above? J Hum Evol 29:169-180. Begun DR. 1994. Relations Among the Great Apes and Humans: New Interpretations Based on the Fossil Great Ape Dryopithecus. Yearb phys Anthrop 37:11-63. Begun DR. 1992a. Phyletic Diversity and Locomotion in Primitive European Hominids. Am J Phys Anthrop 87:311-340. Begun DR. 1992b. Dryopithecus crusafonti sp. nov., a new Miocene hominoid species from Can Ponsic (Northeastern Spain). Am J phys Anthrop 87:291-307. Begun DR. 1991. European Miocene catarrhine diversity. J Hum Evol 20:521-526. Begun DR, MC Nargolwalla & L Kordos. 2008. Revision of the Dryopithicinae:
Phylogenetic and paleobiogeographic implications. Am J Phys Anthrop 135(S46):66. Begun DR, MC Nargolwalla & MP Hutchison. 2006a. Primate diversity in the Pannonian Basin: In situ evolution, dispersals, or both. Beiträge zur Paläontologie 30:43-56. Begun DR, CV Ward, AS Deane, TL Kivell, MC Nargolwalla & ND Taylor. 2006b. Stem hominine or hominid? The phylogeny and functional anatomy of Pierolapithecus catalaunicus. Am J Phys Anthrop 129(S42). Begun DR & CV Ward. 2005. Comment on “Pierolapithecus catalaunicus,” a New Middle Miocene Great Ape from Spain.” Science 308:203c. Begun DR, E Güleç, D Geraads. 2003a. Dispersal patterns of Eurasian hominoids: implications from Turkey. In: Reumer JWF & W Wessels (Eds) Distribution and Migration of Tertiary Mammals in Eurasia. A Volume in Honour of Hans DeBruijn. Deinsea 10:23-39. Begun DR, D Geraads & E Güleç. 2003b. The Çandır hominoid locality: Implications for
the timing and pattern of hominoid dispersal events. In: Güleç E, DR Begun & D Geraads (Eds) Geology and Vertebrate Paleontology of the Middle Miocene Hominoid Locality Çandır. Cour Forsch –Inst Senckenberg 240:251-265.
Begun DR, E Güleç, D Geraads. 2001. Dispersal pattern of Eurasian hominoids:
237
implications from Turkey. Distribution and Migration of Tertiary Mammals in Eurasia. The University of Utrecht, 17-19 May, 2001. p8-9.
Begun DR & L Kordos. 1997. Phyletic Affinities and Functional Convergence in
Dryopithecus and Other Miocene and Living Hominids. In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and
Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p389-416.
Behrensmeyer AK. 1991. Terrestrial Vertebrate Accumulations. In: Allison PA & DEG Briggs (Eds) Taphonomy: Releasing the Data Locked in the Fossil Record. New York: Plenum Press, p291-335. Behrensmeyer AK, D Western & D Dechant-Boaz. 1979. New Perspectives in Vertebrate
Paleoecology from a Recent Bone Assemblage. Paleobiology 5 (1):12-21. Bellier E, P Monestiez, J-P Durbec & J-N Candau. 2007. Identifying spatial relationships at multiple scales. Ecography 30:385-399. Benammmi M, M Calvo, M Prévot & J-J Jaeger. 1996. Magnetostratigraphy and
paleontology of Aït Kandoula Basin (High Atlas, Morocco) and the Africa-European late Miocene terrestrial fauna exchanges. Earth Planet Sci Lett 145:15-29.
Bernor RL. 1983. Geochronology and Zoogeographic Relationships of Miocene
Hominoidea. In: Ciochon RL & RS Corruccini (Eds) New Interpretations of Ape and Human Ancestry. New York: Plenum Press, p21-64.
Bernor RL. 1978. The Mammalian Systematics, Biostratigraphy and Biochronology of
Maragheh and its Importance for Understanding Late Miocene Hominoid Zoogeography and Evolution (PhD thesis). University of California, Los Angeles.
Bernor RL & JM Harris. 2003. Systematics and Evolutionary Biology of the Late Miocene
and Early Pliocene Hipparionine Equids from Lothagam, Kenya. In: Leakey MG & JM Harris (Eds) Lothagam: The Dawn of Humanity in Eastern Africa. New York:
Columbia University Press, p388-438. Bernor RL, RS Scott, M Fortelius, J Kappelman & S Sen. 2003. Equidae (Perissodactyla).
In: Fortelius M, J Kappelman, S Sen & RL Bernor (Eds) Geology and Paleontology of the Miocene Sinap Formation, Turkey. New York: Columbia University Press, p220-281.
Bernor RL & M Armour-Chelu. 1999. Family Equidae. In: Rössner GE & K Heissig
238
(Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p193-202.
Bernor RL, V Fahlbusch, H-W Mittmann & S Rietschel. 1996a. The Evolution of Western Eurasian Neogene Mammal Faunas: The 1992 Schloss Reisensburg Workshop
Concept. In: Bernor RL, V Fahlbusch, H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press. p1-4.
Bernor RL, V Fahlbusch, P Andrews, D de Bruijn, M Fortelius, F Rögl, FF Steininger & L Werdelin. 1996b. The Evolution of Western Eurasian Neogene Mammal Faunas: A
Chronologic, Systematic, Biogeographic, and Paleoenvironmental Synthesis. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p449-469.
Bernor RL, GD Koufos, MO Woodburne & M Fortelius. 1996c. The Evolutionary History and Biochronology of European and Southwest Asian Late Miocene and Pliocene Hipparionine Horses. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p307-338. Bernor RL, N Solounias, CC Swisher III & JA Van Couvering. 1996d. The Correlation of
Three Classical “Pikermian” Mammal Faunas – Maragheh, Samos, and Pikermi – with the European MN Unit System. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p137-156.
Bernor RL, V Fahlbusch & S Rietschel. 1993. Evolution of Neogene continental biotopes in
Central Europe and the eastern Mediterranean (15.5-5Ma). Immendingen and Schloss Reisensburn, 5-11th July, 1992. J Hum Evol. 24: 169-171.
Bernor RL & H Tobien. 1990. The mammalian geochronology and biogeography of Paşalar (Middle Miocene, Turkey). J Hum Evol 19:551-568. Bicchi E, E Ferrero, M Gonera. 2003. Palaeoclimatic interpretation based on Middle
Miocene planktonic foraminifera: the Silesia Basin (Paratethys) and Monferrato (Tethys) records. Palaeogeogr Palaeoclimatol Palaeoecol 196:265-303.
Bogart SL & JD Pruetz. 2008. Ecological context of savanna chimpanzees (Pan troglodytes verus) termite fishing at Fongoli, Senegal. Am J Primatol 70:605-612. Böhme M. 2003a. The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe. Palaeogeogr Palaeoclimatol Palaeoecol 195:389-401. Böhme M. 2003b. Migration history of air-breathing fishes reveal Neogene atmospheric
239
circulation pattern. EEDEN – Environments and Ecosystem Dynamics of the Eurasian Neogene, Birth of the New World. Stará Lesná, Slovakia. November 12-16th, 2003. p14.
Boillot G & R Capdevila. 1977. The Pyrenees: Subduction and collision. Earth Planet Sci Lett 35:151-160.
Bojar A-V, H Hiden, A Fenninger, F Neubauer. 2004. Middle Miocene seasonal temperature changes in the Styrian basin, Austria, as recorded by the isotopic composition of pectinid and brachiopod shells. Palaeogeogr Palaeoclimatol Palaeoecol 203:95-105.
Bolliger T. 1999. Family Anomalomyidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p411-420. Bolliger T. 1996. A Current Understanding about the Anomalomyidae (Rodentia): Reflections on Stratigraphy, Paleobiogeography, and Evolution. In: RL Bernor, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p235-245. Boon-Kristkoiz E & AR Kristkoiz. 1999. Order Lagomorpha. In: Rössner GE & K Heissig
(Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p259-262.
Boschetto HB, FH Brown & I McDougall. 1992. Stratigraphy of the Lothidok Range, northern Kenya, and K/Ar ages of its Miocene primates. J Hum Evol 22:47-71. Bradley RS. 1999. Paleoclimatology (2nd Edition). Toronto: Academic Press, Ltd. Brooks DR & DA McLennan. 2002. The Nature of Diversity: An Evolutionary Voyage of Discovery. Chicago: University of Chicago Press. Bruijn H de. 1999. Superfamily Sciuroidea. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p271-280. Bruch AA, D Uhl & V Mosbrugger. 2007. Miocene climate in Europe – Patterns and evolution – A first synthesis of NECLIME. Palaeogeogr Palaeoclimatol Palaeoecol 253:1-7. Bruch AA, T Utescher, V Mosbrugger, I Gabrielyan & DA Ivanov. 2006. Late Miocene climate in the circum-Alpine realm – a quantitative analysis of terrestrial palaeofloras. Palaeogeogr Palaeoclimatol Palaeoecol 238:270-280. Brusch AA. T Utescher, CA Olivares, N Dolakova, D Ivanov & V Mosbrugger. 2004. Middle and late Miocene spatial temperature patterns and gradients in Europe –
preliminary results based on palaeobotanical climate reconstructions. Cour Forsch –Inst Senckenberg 249:15-27.
240
Clauzon G, J-P Suc, F Gautier, A Berger & M-F Loutre. 1996. Alternate interpretation of the Messinian Salinity Crisis: Controversy resolved. Geology 24:363-366. Costeur L, S Montuire, S Legendre & O Maridet. 2007. The Messinian event: What
happened to the peri-Mediterranean mammalian communities and local climate? Geobios 40:423-431.
Cote SM. 2004. Origins of the African hominoids: an assessment of the paleobiogeographical evidence. CR Palevol 3:321–338. Csontos L. 1992. Tertiary tectonic evolution of the Intra-Carpathian area: a review. Acta Vulcanologica 7: 1-13. Daams R. 1999. Family Gliridae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p301-318. Daams R & AJ Van Der Meulen. 1984. Paleoenviromental and paleoclimatic interpretations of micromammal faunal successions in the Upper Oligocene and Miocene of North Central Spain. Paléobiologie continentale 14(2):241-257. Darwin C. 1871. The Descent of Man, and Selection in Relation to Sex. London: John Murray. Daxner-Höck G. 1999. Family Zapodidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p337-342. Deane AS. 2007. Inferring Dietary Behaviour for Miocene Hominoids: A High-Resolution
Morphometric Approach to Incisal Crown Curvature (PhD Thesis), University of Toronto.
Deane AS, EP Kremer & DR Begun. 2008. Broken fingers: retesting locomotor hypotheses
for fossil hominoids using fragmentary proximal phalanges and high-resolution polynomial curve fitting (HR-PCF). J Hum Evol.
de Bonis L & GD Koufos. 1997. The Phylogenetic and Functional Implications of
Ouranopithecus macedoniensis. In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p317-326.
de Bonis L & GD Koufos. 2004. Ouranopithecus et la date de separation des hominoids modernes. CR Palevol 3:255-262. de Bonis L & GD Koufos. 1999. The Miocene large mammal succession in Greece. In: Agustí J, L Rook & P Andrews (Eds) Hominoid Evolution and Climatic Change in
Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p205-237.
241
de Bruijn H, S Mayda, L vd Hoek Ostende, T Kaya & G Saraç. 2006. Small mammals from the Early Miocene of Sabuncubeli (Manisa, SW Anatolia, Turkey). Beiträge zur Paläontologie 30:57-87. de Bruijn H, L vd Hoek Ostende, E Kristkoiz-Boon, M Rummel, C Theocharopoulos & E
Ünay. 2003. Rodents, lagomorphs and insectivores, from the middle Miocene hominoid locality of Çandır. In: Güleç E, DR Begun & D Geraads (Eds) Geology and Vertebrate Paleontology of the Middle Miocene Hominoid Locality Çandır. Cour Forsch –Inst Senckenberg 240:51-88.
de Bruijn H, E Ünay & L vd Hoek Ostende. 1996. The Composition and Diversity of Small
Mammal Associations from Anatolia Through the Miocene. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p266-270.
de Bruijn H & E Ünay. 1996. On the Evolutionary History of the Cricetodontini from Europe and Asia Minor and Its Bearing on the Reconstruction of Migrations and the
Continental Biotope during the Neogene. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p227-234.
Dèzes P, SM Schmid & PA Ziegler. 2004. Evolution of the European Cenozoic Rift
System: interaction of the Alpine and Pyrenean orogens with their foreland lithosphere. Tectonophysics 389:1-33.
Dreyer SK. 1984. The Theory and Use of Methods for the Study of Mammalian Paleoecology (Ph.D. thesis). University College, London. Durden CJ. 1974. Biomerization: An ecologic theory of provincial differentiation. In: CA Ross (Ed) Paleogeographic Provinces and Provinciality. Society of Economic Paleontologists and Mineralogists, Special publication No. 21:18-53. Efremov JA. 1940. Taphonomy: A New Branch of Paleontology. Pan-America Geologist. 74 (2):81-93. Engesser B. 1999. Family Eomyidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p319-336. Engesser B & R Ziegler. 1996. Didelphids, Insectivores, and Chiropterans from the Later
Miocene of France, Central Europe, Greece, and Turkey. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p157-167.
Eronen J & L Rook. 2004. The Mio-Pliocene European primate fossil record: dynamics and habitat tracking. J Hum Evol 47:323-341.
242
Eronen J, L Rook & M Fortelius. 2003. Mammal community structure and primate dynamics during late Neogene. EEDEN – Environments and Ecosystem Dynamics of the Eurasian Neogene, Birth of the New World. Stará Lesná, Slovakia. November 12-16th, 2003. p17.
Esteban M. 1996. An overview of Miocene reefs from Mediterranean areas: general trends
and facies models. In: Franseen EK, M Esteban, WC Ward & J-M Rouchy (Eds) Models for Carbonate Stratigraphy from Miocene Reef Complexes of Mediterranean Areas. Tulsa: Society for Sedimentary Geology, Concepts in Sedimentology and Paleontology 5:3-53.
Esteban M, JC Braga, J Martín, C De Santisteban. 1996. Western Mediterranean Reef
Complexes. In: Franseen EK, M Esteban, WC Ward, J-M Rouchy (Eds) Models for Carbonate Stratigraphy from Miocene Reef Complexes of Mediterranean Areas. Tulsa: Society for Sedimentary Geology, Concepts in Sedimentology and Paleontology Volume 5, p55-72.
Estrada Belli F. 1999. The Archaeology of Complex Societies in South Eastern Pacific Coastal Guatemala: A Regional GIS Approach. England: British Archaeological Reports. Esu D. 1999. Contribution to the knowledge of Neogene climatic changes in western and
central Europe by means of non-marine mollusks Agustí J, L Rook & P Andrews (Eds) The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge:
Cambridge University Press, p328-354. Fahlbusch V & P Mein. 1989. New Perspectives: The Past, the Present, and the Future. In:
Lindsay E, V Fahlbusch & P Mein (Eds) European Neogene Mammal Chronology. NATO ASI Series A 180:625-628.
Fejfar O. 1999. Microtoid Cricetids. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p365-372. Fejfar O, W-D Heinrich. 1989. Muroid Rodent Biochronology of the Neogene and Quaternary in Europe. In: Lindsay E, V Fahlbusch & P Mein (Eds) European
Neogene Mammal Chronology. NATO ASI Series A 180:91-117. Fernández-Jalvo Y. 1995. Small Animal Taphonomy at La Trinchera de Atapuerca (Burgos, Spain): A Remarkable Example of Taphonomic Criteria Used for Stratigraphic
Correlations and Palaeoenvironment Interpretations. Palaeogeogr Palaeoclimatol Palaeoecol 114 (2-4):167-195.
Fernández-Jalvo Y, C Denys, P Andrews, T Williams, Y Dauphin & L Humphrey. 1998.
Taphonomy and palaeoecology of Olduvai Bed-I (Pleistocene, Tanzania). J Hum Evol 34:137-172.
243
Folinsbee KE & DR Brooks. 2007. Miocene hominoid biogeography: pulses of dispersal and differentiation. J Biogeogr 34:383-397. Fortelius M. 1985. Ungulate cheek teeth: developmental, functional and evolutionary interrelations. Acta Zoologica Fennica 180:1-76. Fortelius M, J Eronen, LP Liu, D Pushkina, A Tesakov, I Vislobokova & ZQ Zhang. 2003a. Continental-scale hysodonty patterns, climatic paleobiogeography and dispersal of Eurasian Neogene land mammal herbivores. In: Reumer JWF & W Wessels (Eds) Distribution and Migration of Tertiary Mammals in Eurasia. A Volume in Honour of Hans de Bruijn. Deinsea 10:1-11. Fortelius M, J Kappelman, S Sen & RL Bernor. 2003b. Geology and Paleontology of the Miocene Sinap Formation, Turkey. New York: Columbia University Press, Fortelius M, J van der Made & RL Bernor. 1996. Middle and Late Miocene Suoidea of Central Europe and the Eastern Mediterranean: Evolution, Biogeography, and
Paleoecology. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p348-377.
Fortelius M, L Werdelin, P Andrews, RL Bernor, A Gentry, L Humphrey, H-W Mittmann &
S Viranta. 1996a. Provinciality, Diversity, Turnover, and Paleoecology in Land Mammal Faunas of the Later Miocene of Western Eurasia. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p414-448.
Fortelius M, J Van der Made & RL Bernor. 1996b. Middle and Late Miocene Suoidea of
Central Europe and the Eastern Mediterranean: Evolution, Biogeography, and Paleoecology. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press. p348-377.
Fortin M-J & M Dale. 2005. Spatial Analysis: A Guide for Ecologists. Cambridge: Cambridge University Press. Frakes LA, JE Francis, JI Syktus. 1992. Climate Models of the Phanerozoic. Cambridge: Cambridge University Press. Franzen JL. 1997. Die Säugetiere aus dem Turolium von Dorn-Dürkheim (Rheinhessen,
Deutschland), Teil 1: Carnivora, Proboscidea (Tetralophodontidae), Perissodactyla (Rhinocerotidae, Equidae), Artiodactyla (Suidae). Cour Forsch –Inst Senckenberg, Band 197, Senckenbergische Naturforschende Gesellschaft, Frankfurt am Main.
Franzen JL & G Storch. 1999. Late Miocene mammals from Central Europe. In: Agustí J, L
244
Rook, P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p165-190.
Freudenthal M & E Martin Suárez. 1999. Family Muridae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p401-410. Gabunia L, E Gabashvili, A Vekua & D Lordkipanidze. 2001. The late Miocene hominoid
from Georgia. In: Bonis L de, GD Koufos & P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 2: Phylogeny of the Neogene Hominoid Primates of Eurasia. Cambridge: Cambridge University Press, p316-325.
Gebhardt H. 1999. Middle to Upper Miocene benthonic foraminiferal palaeoecology of the Tap Marls (Alicante Province, SE Spain) and its palaeoceanographic implications. Palaeogeogr Palaeoclimatol Palaeoecol 145:141-156. Gentry AW, GE Rössner & EPJ Heizmann. 1999. Suborder Ruminantia. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p225-258. Gentry A & EPJ Heizmann. 1996. Miocene Ruminants of the Central and Eastern
Paratethys. 1996. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p378-394.
Geraads D, T Kaya & S Mayda. 2005. Late Miocene large mammals from Yulaflı, Thrace region, Turkey, and their biogeographic implications. Acta Palaeontological Polonica 50:523-544.
Geraads D, E Güleç & T Kaya. 2002. Sinotragus (Bovidae, Mammalia) from Turkey, and the Late Miocene Middle Asiatic Province. Neues Jahrbuch für Geologie und Paläontologie Monatschefte 8:477-489. Giese P, K-J Reutter, V Jacobshagen & R Nicolich. 1982. Explosion seismic crustal studies in the Alpine Mediterranean region and their implications to tectonic processes. In: Berckhemer H & K Hsü (Eds), Alpine-Mediterranean Geodynamics. Colorado: American Geophysical Union 7:39-74. Ginsburg L. 1999. Order Carnivora. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p109-148. Göhlich U. 1999. Order Proboscidea. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p157-168. Golonka J. 2004. Plate tectonic evolution of the southern margin of Eurasia in the Mesozoic and Cenozoic. Tectonophysics 381:225-273.
245
Görür N, M Sakinç, A Barka, R Akkök. 1995. Miocene to Pliocene palaeogeographic evolution of Turkey and its surroundings. J Hum Evol 28:309-324. Green SW. 1990. Approaching Archaeological Space. In: Allen KMS, SW Green, EBW Zubrow (Eds) Interpreting Space: GIS and Archaeology. New York: Taylor and Francis, p3-8. Guérin C. 1989. Biozones or Mammal Units? Methods and Limits in Biochronology. In:
Lindsay E, V Fahlbusch & P Mein (Eds) European Neogene Mammal Chronology. NATO ASI Series A 180:119-130.
Güleç E, A Sevim, C Pehlevan & F Kaya. 2007. A new great ape from the late Miocene of Turkey. Anthrop Sci 115:153-158. Güleç E & DR Begun. 2003. Functional morphology and the affinities of the hominoid mandible from Çandır. Geology and Vertebrate Paleontology of the Middle Miocene
Hallam A. 1974. Changing patterns of provinciality and diversity of fossil animals in relation to plate tectonics. J Biogeog 1:213-225. Hamilton WR, PJ Whybrow & HA McClure. 1978. Fauna of fossil mammals from the Miocene of Saudi Arabia. Nature 274:248-249. Hammer Ø, DAT Harper & PD Ryan. 2007. PAST – Palaeotological Statistics, ver. 1.72. http://folk.uio.no/ohammer/past/ Harris JM & MG Leakey. 2003. Lothagam Suidae. In: Leakey MG & JM Harris (Eds),
Lothagam: The Dawn of Humanity in Eastern Africa. New York: Columbia University Press, p485-519.
Harrison T. 2004. The zoogeographic and phylogenetic relationships of early catarrhine primates in Asia. Anthrop Sci 113:43-51. Harrison T. 2002. Late Oligocene to middle Miocene catarrhines from Afro-Arabia. In:
Hartwig WC (Ed) The Primate Fossil Record. Cambridge: Cambridge University Press, p311-338.
Harrison T. 1991. Some observations on the Miocene hominoids from Spain. J Hum Evol 20:515-520. Harrison T & Y Gu. 1999. Taxonomy and phylogenetic relationships of early Miocene catarrhines from Sihong, China. J Hum Evol 37:225-277. Harrison T & L Rook. 1997. Enigmatic anthropoid or misunderstood ape? The phylogenetic
246
status of Oreopithecus bambolii reconsidered. In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations. New York: Plenum, p327-362.
Harzhauser M, C Latal & WE Piller. 2007. The stable isotope archive of Lake Pannon as a
mirror of Late Miocene climate change. Palaeogeogr Palaeoclimatol Palaeoecol 249:335-350.
Heikinheimo H, M Fortelius, J Eronen & H Mannila. 2007. Biogeography of European land mammals show environmentally distinct and spatially coherent clusters. J Biogeogr 34:1053-1064. Heissig K. 1999a. Family Rhinocerotidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p175-188. Heissig K. 1999b. Family Chalicotheriidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p189-192. Heissig K. 1999c. Family Tubulidentata. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p155-156. Heissig K. 1996. The Stratigraphical Range of Fossil Rhinoceroses in the Late Neogene of
Europe and the Eastern Mediterranean. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p339-347.
Heizmann EPJ & DR Begun. 2001. The oldest Eurasian hominoid. J Hum Evol 41:463- 481. Hladik CM. 1977. Chimpanzees of Gabon and chimpanzees of Gombe: Some comparative data in the diet. In: Clutton-Brock TH (Ed) Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, Monkeys and Apes. London: Academic Press, p481- 501. Hoek Ostende L vd, M Morlo & D Nagel. 2006. Magestic Killers: the sabre-toothed cats. Geology Today 22:150-157. Hsü KJ, L Montadert, D Bernoulli, MB Cita, A Erickson, RE Garrison, RB Kidd, F Mèliéres, C Müller & R Wright. 1977. History of the Mediterranean salinity crisis. Nature 267:399-404. Hughes CP. 1973. Analysis of past faunal distributions. In: Tarling DH & SK Runcorn, (Eds) Implications of Continental Drift to the Earth Sciences, vol. 1. New York: Academic Press, p219-230. Hugueney M. 1999. Family Castoridae. In: Rössner GE & K Heissig (Eds) The Miocene
247
Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p281-300. Huismans RS, Bertotti G, D Ciulavu, CAE Sanders, S Cloetingh & C Dinu. 1997. Structural
evolution of the Transylvanian Basin (Romania): A sedimentary basin in the bend zone of the Carpathians. Tectonophysics 272:249-268.
Humphrey L & P Andrews. 2008. Metric variation in the postcanine teeth from Paşalar, Turkey. J Hum Evol 54:503-517. Hünermann KA. 1999. Superfamily Suoidea. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p209-216. Ivanov D, AR Ashraf, V Mosbrugger & E Palamarev. 2003. Climate change about 10Ma in Paratethyan realm of Balkan Peninsula. EEDEN – Environments and Ecosystem Dynamics of the Eurasian Neogene, Birth of the New World. Stará Lesná, Slovakia. November 12-16th, 2003. p 48. Ivanov D, AR Ashraf, V Mosbrugger & E Palamarev. 2002. Palynological evidence for Miocene climate change in the Forecarpathian Basin (Central Paratethys, NW Bulgaria). Palaeogeogr Palaeoclimatol Palaeoecol 178:19-37. Janis C. 1988. An estimation of tooth volume and hypsodonty indices in ungulate mammals, and the correlation of these factors with dietary preferences. Mémoirs du Muséum national d’Histoire naturelle, Paris 53:367-387. Janis CM & A Lister. 1985. The morphology of the lower fourth premolar as a taxonomic character in the Ruminantia (Mammalia; Artiodactyla), and the systematic position of Triceromeryx. Journal of Paleontology 59:405-410. Jernvall J & M Fortelius. 2004. Maintenance of trophic structure in fossil mammal
communities: site occupancy and taxon resilience. The American Naturalist 164(5):614-624.
Johnson JG. 1971. A quantitative approach to faunal province analysis. Am J Sci 270:257- 280. Jones RW. 1999. Marine invertebrate evidence for the palaeogeography of the Oligocene- Miocene of western Eurasia, and consequences for terrestrial vertebrate migration. In
Agustí J, L Rook & P Andrews (Eds) The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p274-308.
Kälin D. 1999. Tribe Cricetini. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p373-388. Kappelman J, S Sen, M Fortelius, A Duncan, B Alpagut, J Crabaugh, A Gentry, JP Lunkka,
248
F McDowell, N Solounias, S Viranta & L Werdelin. 1996. Chronology and Biostratigraphy of the Miocene Sinap Formation in Central Turkey. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p78-95.
Kay RF & PS Ungar. 1997. Dental evidence for diets in some Miocene catarrhines with comments on the effects of phylogeny on the interpretation of adaptation. In: Begun
DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p131-151.
Kelley J. 2008. Identification of a single birth cohort in Kenyapithecus kizili and the nature
of sympatry between K. kizili and Griphopithecus alpani at Paşalar. J Hum Evol 54:530-537.
Kelley J, P Andrews, B Alpagut. 2008. A new hominoid species from the middle Miocene site of Paşalar, Turkey. J Hum Evol 54:455-479. Kelley J & B Alpagut. 1999. Canine sexing and species number in the Paşalar large hominoid sample. J Hum Evol 36:335-341. Kingston JD & A Hill. 1999. Late Miocene palaeoenviornments in Arabia: A synthesis. In:
Whybrow PJ & A Hill (Eds) Fossil Vertebrates of Arabia. New Haven: Yale University Press, p389-409.
Kidwell SM & AK Behrensmeyer. 1993. Taphonomic Approaches to Time Resolution
in Fossil Assemblages. 16th Annual Short Course of the Paleontological Society, Boston, Massachusetts.
Kordos L & DR Begun. 2002. Rudabánya: A late Miocene subtropical swamp desposit with evidence of the origin of the African apes and humans. Evol Anthropol 11:45-57. Koufos GD & de Bonis L. 2005. The Late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomy. Annls Paléont 91:227-240. Koufos GD & L de Bonis. 2004. The late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomy. 5th International Symposium on Eastern Mediterranean Geology, Thessaloniki 1:322- 325.
Koussoulakou A & E Stylianidis. 1999. The Use of GIS for the Visual Exploration of Archaeological Spatio-Temporal Data. Cart Geog Info Sci 26:152-160. Kopp ML. 2007. The late alpine structure of the Greater Caucasus as an element of the Peri-
Arabian collisional area. European Geosciences Union Geophysical Research Abstracts, Volume 9.
249
Krause DW & MC Maas. 1990. The biogeographic origins of late Paleocene-early Eocene
mammalian immigrants to the Western Interior of North America. In: Bown TM & KD Rose (Eds) Dawn of the Age of Mammals in the northern part of the Rocky Mountain Interior. Geological Society of American Special Papers 243:71-105.
Krijgsman W. 2002. The Mediterranean: Mare Nostrum of Earth Sciences. Earth Planet Sci Lett 205:1-12. Krijgsman W, CG Langereis, R Daams & AJ van der Meulen. 1994. Magnetostratigraphic dating of the middle Miocene climate change in the continental deposits of the
Aragonian type area in the Calatayud-Teruel basin (Central Spain). Earth Planet Sci Lett 128:513-529.
Kunimatsu Y, M Nakatsukasa, Y Sawada, T Sakai, M Hyodo, H Hyodo, T Itaya, H Nakaya, H Saegusa, A Mazurier, M Saneyoshi, H Tsujikawa, A Yamamoto & E Mbua. 2007. A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans. Proc natl Acad Sci 104:19220-19225.
Lake MW, PE Woodman & SJ Mithen. 1998. Tailoring GIS Software for Archaeological Applications. J Arch Sci 25:27-38. Leakey MG & JM Harris. 2003. Lothagam: Its Significance and Contributions. In: Leakey
MG & JM Harris (Eds) Lothagam: The Dawn of Humanity in Eastern Africa. New York: Columbia University Press, p 625-670.
Leakey MG & A Walker. 1997. Afropithecus: Function and Phylogeny. In: Begun DR, CV
Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p225-240.
Leakey MG, CS Feibel, RL Bernor, JM Harris, TE Cerling, KM Stewart, GW Storrs, A
Walker, L Werdelin & A Winkler. 1996. Lothagam: A record of faunal change in the late Miocene of East Africa. J Vert Paleontol 16:556-570.
Leakey RE, MG Leakey & AC Walker. 1988. Morphology of Afropithecus turkanensis from Kenya. Am J phys anthrop 76:289-307. Leakey REF & A Walker. 1985. New higher primates from the early Miocene of Buluk, Kenya. Nature 318:173-175. Lindsay EH. 1989. European Neogene Mammal Chronology. NATO ASI Series A 180. Lyman RL. 1994. Vertebrate Taphonomy. Cambridge: Cambridge University Press. Maas MC, MRL Anthony, PD Gingerich, GF Gunnell & DW Krause. 1995. Mammalian
250
generic diversity and turnover in the Late Paleocene and Early Eocene of the Bighorn and Crazy Mountains Basins, Wyoming and Montana (USA). Palaeogeogr Palaeoclimatol Palaeoecol 115:181-207.
Madden CT. 1980. Zygolophodon from Subsaharan Africa, with Observations on the Systematics of Palaeomastodontid Proboscideans. Journal of Paleontology 54:57-64. Made J van der. 2003. Suoidea (pigs) from the Miocene hominoid locality of Çandır in Turkey. In: Güleç E, DR Begun & D Geraads (Eds) Geology and Vertebrate Paleontology of the Middle Miocene Hominoid Locality Çandır. Cour Forsch –Inst
Senckenberg 240:149-180. Made J van der. 1999. Intercontinental relationship Europe - African and the Indian Subcontinent. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p457-472. Made J van der, J Agustí & J Morales. 2003. The Vallesian Crisis in the Iberian and
European mammal records. EEDEN – Environments and Ecosystem Dynamics of the Eurasian Neogene, Birth of the New World. Stará Lesná, Slovakia. November 12-16th, 2003. p61-62.
Magyar I, DH Geary & P Müller. 1999. Paleogeographic evolution of the Late Miocene
Lake Pannon in Central Europe. Palaeogeogr Palaeoclimatol Palaeoecol 147:151-167.
Manly BJF. 2005. Multivariate Statistical Methods: A Primer. Florida: Chapman & Hall/CRC. Maridet O, G Escarguel, L Costeur, P Mein, M Hugueney & S Legendre. 2007. Small
mammal (rodent and lagomorphs) European biogeography from the Late Oligocene to the Pliocene. Global Ecol. Biogeogr 16:529-544.
Martin L & P Andrews. 1993. Renaissance of Europe’s ape. Nature 365:494. Martin L & P Andrews. 1984. The phyletic position of Graecopithecus freybergi von Koenigswald. Cour Forsch –Inst Senckenberg 69:25-40. Martin L & P Andrews. 1983. Species recognition in middle Miocene hominoids. In:
Kimbel WH & L Martin (Eds) Species, Species Concepts and Primate Evolution. p393-427.
Martín JM, JC Braga, I Sánchez-Almazo. 1999. The Messinian record of the outcropping
marginal Alboran Basin deposits: significance and implications. In: Zahn R, MC Comas & A Klaus (Eds) Proceedings of the Ocean Drilling Program, Scientific Results, Volume 161 (Sites 974-979 Mediterranean II). Texas: Ocean Drilling Program. p543-551.
251
McCrossin ML & BR Benefit. 1997. On the relationships and adaptations of Kenyapithecus,
a large-bodied hominoid from the middle Miocene of eastern Africa. In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations. New York: Plenum Press, p241-267.
Mein P. 2003. On Neogene rodents in Eurasia: distribution and migrations. In: Reumer
JWF & W Wessels (Eds) Distribution and Migration of Tertiary Mammals in Eurasia. A Volume in Honour of Hans DeBruijn. Deinsea 10:407-418.
Mein P. 1989. Updating of MN Zones. In: Lindsay E, V Fahlbusch & P Mein (Eds)
European Neogene Mammal Chronology. NATO ASI Series A 180:73-90. Mein P. 1979. European Mammal Correlations. In: Lindsay EH, V Fahlbusch & P Mein
(Eds) European Mammal Geochronology. New York: Plenum Press, p73-90. Mein P. 1975. Résultats du Groupe de Travail des Vertébrés. In: Senes J (Ed) Report on
Activity of RCMNS Working Groups 6. Congress of the Regional Committee of Mediterranean Neogene Stratigraphy, Proceedings 1:78-81.
Merceron G, E Schulz, L Kordos & TM Kaiser. 2007. Palaeoenvironment of
Dryopithecus brancoi at Rudabánya, Hungary: evidence from dental meso and microwear analyses of large herbivorous mammals. J Hum Evol 53:331-349.
Merceron G, L de Bonis, L Viriot & C Blondel. 2005a. Dental microwear of fossil bovids
from northern Greece: paleoenvironmental conditions in the eastern Mediterranean during the Messinian. Palaeogeogr Palaeoclimatol Palaeoecol 217:173-185.
Merceron G, L de Bonis, L Viriot & C. Blondel. 2005b. Dental microwear of the late Miocene bovids of northern Greece: Vallesian/Turolian environmental changes and disappearance of Ouranopithecus macedoniensis? Bull Soc geol. Fr 176:475-484. Middlemiss FA & PF Rawson. 1969. Faunal provinces in Space and Time – some general
considerations. In: Middlemiss FA, PF Rawson & G Newall (Eds) Faunal Provinces in Space and Time. Liverpool: Seel House Press, p199-210.
Moyà-Solà S, M Köhler, DM Alba, I Casanovas-Vilar & J Galindo. 2004. Pierolapithecus catalaunicus, a New Middle Miocene Great Ape from Spain. Nature 306:1339-1344. Moyà-Solà S, J Quintana, J Antonio Alcover & M Köhler. 1999. Endemic Island Faunas of the Mediterranean Miocene. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p435-442. Moyà-Solà S & M Köhler. 1993. Recent discoveries of Dryopithecus shed new light on
evolution of great apes. Nature 365:543-545.
252
Nagel D. 2003. Carnivora from the middle Miocene Hominoid locality of Çandır. In: Güleç E, DR Begun & D Geraads (Eds) Geology and Vertebrate Paleontology of the Middle Miocene Hominoid Locality Çandır. Cour Forsch –Inst Senckenberg 240:113-132.
Nargolwalla MC. 2007. Middle and late Miocene terrestrial mammals from the Pannonian Basin, Central Europe. J Vert Paleontol 26(3):104a. Nargolwalla MC, MP Hutchison & DR Begun. 2006. Middle and late Miocene terrestrial vertebrate localities and paleoenvironments in the Pannonian Basin. Beiträge zur Paläontologie. 30:347-360. Nargolwalla MC & DR Begun. 2005. Late Miocene hominid biogeography and extinction
patterns. Am J Phys Anthrop 126(S40):155. Ni X & Z Qiu. 2002. The micromammalian fauna from the Leilao, Yuanmou hominoid locality: Implications for biochronology and paleoecology. J Hum Evol 42:535-546. Ohta S, K Kaiho & T Takei. 2003. Relationship between surface-water temperature and ice-
sheet expansion during the middle Miocene. Palaeogeogr Palaeoclimatol Palaeoecol 201:307-320.
Olson EC. 1980. Taphonomy: Its History and Role in Community Evolution. In
Behrensmeyer AK & A Hill (Eds) Fossils in the Making: Vertebrate Taphonomy and Paleoecology. Chicago: University of Chicago Press, p5-19.
Pagani M, KH Freeman, MA Arthur. 1999. Concentrations and the Expansion of C4 Grasses. Science 285:876-880. Peresson H & K Decker. 1997. The Tertiary dynamics of the northern Eastern Alps
(Austria): changing palaeostresses in a collisional plate boundary. Tectonophysics 272(2-4):125-157.
Pickford M. 1989. Dynamics of Old World Biogeographic Realms During the Neogene:
Implications for Biostratigraphy. In: In: Lindsay E, V Fahlbusch & P Mein (Eds) European Neogene Mammal Chronology. NATO ASI Series A 180:413-442.
Pielou EC. 1979. Biogeography. New York: Wiley Pilbeam D. 1997. Research on Miocene Hominoids and Hominid Origins: The last three
decades. In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p13-28.
Pilbeam D. 1996. Genetic and morphological records of the Hominoidea and hominid origins: A synthesis. Mol Phylogenet Evol 5(1):155-168.
253
Pilbeam D. 1989. Human fossil history and evolutionary paradigms. In: Hecht MK (Ed)
Evolutionary Biology at the Crossroads. New York: Queens College Press, p117- 138.
Pilbeam D & N Young. 2004. Hominoid evolution: synthesizing disparate data. CR Palevol
3:303-319. Pilbeam D, M Morgan, JC Barry & L Flynn. 1996. European MN Units and the Siwalik
Faunal Sequence of Pakistan. In: Bernor RL, V Fahlbusch, H-W Mittmann (Eds) TheEvolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p96-105.
Popov SV, F Rögl, AY Rozanov, FF Steininger, IG Shcherba M & Kovac. 2004.
Lithological-Paleogeographic maps of Paratethys. Cour Forsch –Inst Senckenberg 250: 1-46.
Potts R. 2004a. Paleoenvironmental Basis of Cognitive Evolution in Great Apes. Am J
Primatol 62:209-228. Potts R. 2004b. Paleoenvironments and the Evolution of Adaptability in Great Apes. In:
Russon AE & DR Begun (Eds) The Origins of Thought: The Evolutionary Origins of Great Ape Intelligence. Cambridge: Cambridge University Press, p78-127.
Potts R, T Jorstad & D Cole. 1996. The Role of GIS in Interdisciplinary Investigations at Olorgesailie, Kenya, at a Pleistocene Archaeological Locality. In: Aldenderfer M & HDG Maschner (Eds) Anthropology, Space, and Geographic Information Systems. New York: Oxford University Press, p202-213.
Potts R & AK Behrensmeyer. 1992. Late Cenozoic Terrestrial Ecosystems. In:
Behrensmeyer AK, JD Damuth, WA DiMichele, R Potts, H-D Sues & SL Wing (Eds) Terrestrial Ecosystems through Time: Evolutionary Paleoecology of Terrestrial Plants and Animals. Chicago: University of Chicago Press, p419-541.
Pruetz JD. 2007. Evidence of cave use by savanna chimpanzees (Pan troglodytes verus) at
Fongoli, Senegal: Implications for thermoregulatory behaviour. Primates 48:316- 319.
Quinn GP & MJ Keough. 2002. Experimental Design and Data Analysis for Biologists. Cambridge: Cambridge University Press. Rantitsch G. 1997. Thermal history of the Carnic Alps (Southern Alps, Austria) and its palaeogeographic implications. Tectonophysics 272:213-232. Ribot F, J Gibert & T Harrison. 1996. A reinterpretation of the taxonomy of Dryopithecus from Vallès-Penedès. J Hum Evol 31:129-141.
254
Rögl F. 1999a. Mediterranean and Paratethys palaeogeography during the Oligocene and
Miocene. In: Agustí J, L Rook & P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p8-23.
Rögl F. 1999b. Circum-Mediterranean Miocene Paleogeography. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. Germany: Verlag Dr. Friedrich Pfeil, p39-48. Rögl F. 1999c. Oligocene and Miocene Palaeogeography and Stratigraphy of the Circum- Mediterranean Region. In: Whybrow PJ & A Hill (Eds) Fossil Vertebrates of Arabia. New Haven: Yale University Press, p485-500. Rögl F & FF Steininger. 1984. Neogene Paratethys, Mediterranean and Indo-pacific
Seaways. In: Brenchley P (Ed) Fossils and Climate. New York: John Wiley and Sons, p171-200.
Rook L, G Gallai & D Torre. 2006. Lands and endemic mammals in the Late Miocene of Italy: Constraints from paleogeographic outlines of Tyrrhenian area. Palaeogeogr Palaeoclimatol Palaeoecol 238:263-269. Rook L, P Renne, M Benvenuti & M Papini. 2000. Geochronology of Oreopithecus-bearing successions at Baccinello (Italy) and the extinction pattern of European Miocene hominoids. J Hum Evol 39:577-582. Rook L, Abbazzi L & Engesser B. 1999. An overview of the Italian Miocene land mammal
faunas. In: Agustí J, L Rook, P Andrews (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p191-204.
Ross CA. 1974. Paleogeography and Provinciality. In: Ross CA (Ed) Paleogeographic Provinces and Provinciality. Society of Economic Paleontologists and Mineralogists, Special publication No. 21:1-17. Royden LH. 1988. Late Cenozoic Tectonics of the Pannonian Basin System. In: Royden LH
& F Horváth (Eds) The Pannonian Basin – A Study in Basin Evolution, AAPG Memoir 45, 27-48.
Ruddiman WF & WL Prell. 1997. Introduction to the Uplift-Climate Connection. In:
Ruddiman WF (Ed) Tectonic Uplift and Climate Change. New York: Plenum, pp3-15.
Rummel M. 1999. Tribe Cricetodontini. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p359-364.
255
Russak SM & WC McGrew. 2008. Chimpanzees as fauna: Comparisons of sympatric large mammals across long-tem study sites. Am J Primatol 70:402-409.
Sánchez-Gómez M, JC Balanyá, V García-Dueñas & JM Azañón. 2002. Intracrustal tectonic evolution of large lithosphere mantle slabs in the western end of the Mediterranean orogen (Gibraltar arc). Journal of the Virtual Explorer 8. Sanders WJ & ER Miller. 2002. New proboscideans from the early Miocene of Wadi Moghara, Egypt. J Vert Paleontol 22:388-404. Sarmiento E. 1987. The phylogenetic position of Oreopithecus and its significance in the
origin of the Hominoidea. Am Mus Novitates 2881:1-44. Schellart W. 2002. Alpine deformation at the western termination of the Axial Zone, Southern Pyrenees. Journal of the Virtual Explorer 8. Schlunegger F & G Simpson. 2002. Possible erosional control on lateral growth of the European Central Alps. Geology 30(10):907-910. Schmidt-Kittler N. 1989. A Biochronologic Subdivision of the European Paleogene Based
on Mammals. In: Lindsay E, V Fahlbusch & P Mein (Eds) European Neogene Mammal Chronology. NATO ASI Series A 180:-50.
Schwartz T. 1997. Lateritic bauxite in central Germany and implications for Miocene palaeoclimate. Palaeogeogr Palaeoclimatol Palaeoecol 129:37-50. Selänne L. 2003. Genus Schizogalerix (Insectivora). In: Fortelius M, J Kappelman, S Sen
& RL Bernor (Eds) Geology and Paleontology of the Miocene Sinap Formation, Turkey. New York: Columbia University Press, p69-89.
Sen S. 2003. Muridae and Gerbillidae (Rodentia). In: Fortelius M, J Kappelman, S Sen &
RL Bernor (Eds) Geology and Paleontology of the Miocene Sinap Formation, Turkey. New York: Columbia University Press, p125-140.
Sen S. 1997. Magnetostratigraphic calibration of the European Neogene mammal
chronology. Palaeogeogr Palaeoclimatol Palaeoecol 133:181-204. Sen S. 1996. Present State of Magnetostratigraphic Studies in the Continental Neogene of Europe and Anatolia. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p56-63. Sibuet J-C, SP Srivastava, W Spakman. 2004. Pyrenean orogeny and plate kinematics. Journal of Geophysical Research 109:1-18. Simpson GG. 1960. Notes on the measurement of faunal resemblance. Am J Sci 258a:300-
256
311. Simpson GG. 1940. Mammals and Land Bridges. J Wash Acad Sci 30:137-163. Sokal RR & FJ Rohlf. 1995. Biometry. New York: WH Freeman and Company. Sokal RR & PHA Sneath. 1963. The Principles of Numerical Taxonomy. San Francisco: Freeman & Co. Solounias N, JM Plavcan, J Quade & L Witmer. 1999. The paleoecology of the Pikermian
Biome and the savanna myth. In: Agustí J, L Rook & P Andrews, (Eds) Hominoid Evolution and Climatic Change in Europe, vol 1: The Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge: Cambridge University Press, p436- 453.
Spassov N & D Geraads. 2008. The latest prehumans of Europe: discovery of a late Miocene hominoid in Bulgaria of about 7Ma. Bulgarian Academy of Sciences News 2(54):2-4. Spezzaferri S, Cita MB & McKenzie JA. 1998. The Miocene/Pliocene boundary in the
Eastern Mediterranean: results from sites 967 and 969. In: Robertson AHF, K-C Emeis, C Richter & A Camerlenghi (Eds) Proceedings of the Ocean Drilling Program, Scientific Results, vol 160 (Sites 963-973 Mediterranean I). Texas (Ocean Drilling Program). p9-28.
Sponheimer M, JE Loudon, D Codron, ME Howells, JD Pruetz, J Codron, DJ de Ruiter & JA Lee-Thorp. 2006. Do “savanna” chimpanzees consume C4 resources? J Hum Evol 51:128-133. Stampfli GM, GD Borel, R Marchant & J Mosar. 2002. Western Alps geological constraints on western Tethyan reconstructions. Journal of the Virtual Explorer 7:75-104. Stanford C, J Wallis, H Matama & J Goodall. 1994. Patterns of predation by chimpanzees on red colobus monkeys in Gombe National Park, Tanzania, 1982-1991. Am J Phys
Anthrop 94:213-228. Steininger FF, RL Bernor & V Fahlbusch. 1989. European Neogene Marine/Continental
Chronologic Correlations. In: Lindsay E, V Fahlbusch & P Mein (Eds) European Neogene Mammal Chronology. NATO ASI Series A 180:15-46.
Steininger FF, G Rabeder & F Rögl. 1985. Land Mammal Distribution in the Mediterranean Neogene: A Consequence of Geokinematic and Climatic Events. In: Stanley DJ & F-
C Wezel (Eds) Geological Evolution of the Mediterranean Basin: New York: Springer-Verlag, p560-571.
Stewart C-B & TR Disotell. 1998. Primate evolution – in and out of Africa. Current
257
Biology 8:R582-R588. Susman RL. 1985. Functional morphology of the Oreopithecus hand. Am J Phys
Anthrop 66:235. Swisher III CC. 1996. New 40Ar/39Ar Dates and Their Contribution Toward a Revised
Chronology for the Late Miocene of Europe and West Asia. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p64-77.
Suwa G, RT Kono, S Katoh, B Asfaw & Y Beyene. 2007. A new species of great ape from the late Miocene epoch in Ethiopia. Nature 448:921-924. Szalay FS & E Delson. 1979. Evolutionary History of the Primates. New York: Academic Press. Tarling DH. 1978. The geological-geophysical framework of ices ages. In: Gribbin J (Ed) Climatic Change. Cambridge: Cambridge University Press, p3-24. Thenius E. 1952. Die Säugetierfauna aus dem Torton von Neudorf an der March (CSR). Neues Jahrbuch für Geologie und Paläontologie 96:27-136. Thomas H. 1985. The Early and Middle Miocene Land Connections of the Afro-Arabian
Plate and Asia: A Major Event for Hominoid Dispersal? In: Delson E (Ed) Ancestors, the Hard Evidence. New York: Liss, p42-50.
Thomas H, RL Bernor & J-J Jaeger. 1982. Origines du peuplement mammalien en Afrique due nord durant le Miocène terminal. Geobios 15 :283-297. Tong H & J-J Jaeger. 1993. Muroid rodents from middle Miocene Fort Ternan locality
(Kenya) and their contribution to the phylogeny of muriods. Palaeontographica 229:51-73.
Tutin CEG, R Ham, LJT White & MJS Harrison. 1997. The primate community of Lope’
reserve in Gabon: Diets, responses to fruit scarcity, and effects on biomass. Am J Primatol 42:1-24.
Ünay E, H de Bruijn & G Saraç. 2003. A preliminary zonation of the continental Neogene of Anatolia based on rodents. In: Reumer JWF & W Wessels (Eds) Distribution and Migration of Tertiary Mammals in Eurasia. A Volume in Honour of Hans DeBruijn. Deinsea 10:539-548. Ungar PS. 2005. Dental evidence for the diets of fossil primates from Rudabánya,
northeastern Hungary with comments on extant primate analogs and “noncompetitive” sympatry. Palaeontographica Italica 90:97-111.
258
Ungar PS. 1998. Dental allometry, morphology and wear as evidence for diet in fossil primates. Evol Anthropol 6:205-217. Ungar PS. 1996. Dental microwear of European Miocene catarrhines: Evidence for diets
and tooth use. J Hum Evol 31:335-366. Ungar PS & RF Kay. 1995. The dietary adaptations of European Miocene catarrhines. Proc
Nat Acad Sci USA 92:5479-5481. van Dam JA, L Alcalá, A Alonso Zarza, JP Calvo, M Garcés & W Krijgsman. 2001. The upper Miocene mammal record from the Teruel-Alfambra region (Spain). The MN system and continental stage/age concepts discussed. J Vert Paleontol 21(2):367-385. van Dam JA & GJ Weltje. 1999. Reconstruction of the Late Miocene climate of Spain using
rodent palaeocommunity successions: an application of end-member modelling. Palaeogeogr Palaeoclimatol Palaeoecol 151:267-305.
Vergé J, M Fernàndez & A Martínez. 2002. The Pyrenean orogen: pre-, syn-, and post- collisional evolution. Journal of the Virtual Explorer 8. Vestal AG. 1914. Internal relations of terrestrial associations. Am Naturalist 48:413-445. Waddle DM, LB Martin, DA Stock. 1995. Sexing isolated hominoid canines with special
reference to the Middle Miocene specimens from Paşalar, Turkey. J Hum Evol 28:385-403.
Ward CV, DR Begun & MD Rose. 1997. Function and Phylogeny in Miocene Hominoids.
In: Begun DR, CV Ward & MD Rose (Eds) Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptation. New York: Plenum Press, p1-12.
Ward SC, B Brown, A Hill, J Kelley & W Downs. 1999. Equatorius: a new hominoid genus from the middle Miocene of Kenya. Science 285:1382-1386. Weerd A van de & R Daams. 1978. Quantitative composition of rodent faunas in the Spanish Neogene and paleoecological implications. Proceedings Koninklijke Nederlandse Akademie van Wetenschappen 81(4):448-473. Werdelin L. 2003. Mio-Pliocene Carnivora from Lothagam, Kenya. In: Leakey MG & JM Harris (Eds), Lothagam: The Dawn of Humanity in Eastern Africa. New York: Columbia University Press, p261-328. Werdelin L. 1996. Carnivores, exclusive of the Hyaenidae, from the later Miocene of
Europe and Western Asia. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia
University Press, p271-289.
259
Werdelin L & N Solounias. 1996. The Evolutionary History of Hyaenas in Europe and Western Asia During the Miocene. In: Bernor RL, V Falhbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p290-306.
Werdelin L & N Solounias. 1991. The Hyaenidae: taxonomy, systematics and evolution. Fossils and Strata 30:1-104. Wesselman HB. 1995. Of mice and almost men. In: Vrba ES, GH Denton, TC Partridge
& LH Burckle (Eds) Paleoclimate and evolution with emphasis on human origins. New Haven: Yale University Press, p356–368.
Wessels W, O Fejfar, P Peláez-Campomanes, A van der Meulen & H de Bruijn. 2003.
Miocene small mammals from Jebel Zelten, Libya. Coloquois de Paleontologia 1:699-715.
Wessels W. 1999. Family Gerbillidae. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Pfeil, p395-400. Whybrow PJ & D Clements. 1999. Arabian Tertiary fauna, flora, and localities. In:
Whybrow PJ & A Hill (Eds), Fossil Vertebrates of Arabia. New Haven: Yale University Press, p460-475.
Woodburne MO, RL Bernor, CC Swisher III. 1996. An appraisal of the Stratigraphic and Phylogenetic Bases for the “Hipparion” Datum in the Old World. In: Bernor RL, V Fahlbusch & H-W Mittmann (Eds) The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press, p124-136. Yamakoshi G. 1998. Dietary responses to fruit scarcity of wild chimpanzees at Boussou,
Guinea: Possible implications for ecological importance of tooth use. Am J Phys Anthrop 106:283-295.
Zeck HP. 1996. Betic-Rif orogeny: subduction of Mesozoic Tethys lithosphere under
eastward drifting Iberia, slab detachment shortly before 22 Ma, and subsequent uplift and extensional tectonics. Tectonophysics 254:1-16.
Zhang Z & T Harrison. 2008. A new middle Miocene pliopithecid from Inner Mongolia, China. J Hum Evol. 54:444-447. Ziegler R. 1999. Order Insectivora. In: Rössner GE & K Heissig (Eds) The Miocene Land Mammals of Europe. München: Verlag Dr. Friedrich Pfeil, p53-74.