Introduction   9
Weisenau and Mösskirch Suidae
and the problematic species
Hyotherium medium Von Meyer, 1841a   10
Material and Methods   11
Organisation of Systematic Descriptions   13
Designation of tooth position   13
Measurements   13
Geological Time Scale and Biostratigraphy   13
Oligo-Miocene Biostratigraphy in Europe   13
Occurrences of Hyotherium in Europe
(whether verii, dubii or spurri)   15
Biostratigraphy of Mösskirch   17
Historically early samples of Hyotherium   17
Archives   19
Literature   20
Context of suid studies over time   21
Geochronology   21
Socio-politics   21
Suprageneric classification
of suid-like mammals   22
Historical review the genus Hyotherium
with the focus on Mösskirch Suidae   24
Phase 1: 18181867   27
Phase 2: 18671900   32
Phase 3: 19011971   36
Phase 4: 19712015   45
Systematic Palaeontology   52
Family Suidae Gray, 1821   52
Subfamily Listriodontinae Gervais, 1859   52
Genus Listriodon Von Meyer, 1846   52
Species Listriodon latidens
(Biedermann, 1873)   52
Subfamily Hyotheriinae Cope 1888   53
Genus Hyotherium Von Meyer, 1834   53
Species Hyotherium soemmeringi
(Von Meyer, 1834)   53
Derching, Germany  62
Stätzling, Germany  62
Quinta da Farinheira, Portugal  62
Petersbuch 39, Germany  63
Petersbuch 81, Germany  70
Steyregg bei Wies, Austria  73
Species Hyotherium medium
Von Meyer, 1841a   73
Description of Mösskirch specimens of
Hyotherium medium   74
Comment on the taxonomic status of
Sus (Palaeohyus) wylensis
(Von Meyer, 1859)   79
Comment on the status of
Hyotherium soemmeringi bavaricum
Dehm, 1980   81
Sandelzhausen, Germany  81
Anmühle bei Weidorf, Germany  91
Pfettrachmühle, Germany  91
La Romieu, France  93
Käpfnach, Switzerland  94
Feisternitz bei Eibiswald, Austria  94
Rothenstein 1-13, Germany  98
Engelswies, Germany  106
Baigneaux, France  113
Chilleurs-aux-Bois, France  118
Schönegg bei Wiess and Kirchberg,
Austria   118
Riedern, Switzerland  118
Fohnsdorf, Austria  120
Burg-Balhausen, Germany  120
Faluns d’Anjou, France  121
Münzenberg, Austria  121
Seegraben, Austria  122
Zangtal, Austria  122
Species Hyotherium major (Pomel, 1847)   122
Comments on the status of
Hyotherium major (Pomel, 1847)   122
Wiesbaden (Spelzmühle), Germany  123
Saint-Gérand-le-Puy, France  123
Montaigu-Le-Blin, France  127
Quercy, France  127
Rothenstein 10-14e, Germany  127
Ulm-Westtangente, Germany  131
Öhlinger Tal bei Ulm, Germany  135
Laugnac, France  135
Corronsac, France  144
Gibel, France  144
Hüllistein and Höhronen, Switzerland  144
Hausen bei Pfullendorf  144
Hausen bei Sigmaringen, Germany  144
Haslach bei Ulm, Germany  147
Chinese occurrences of Hyotherium   147
General discussion and conclusions
on Hyotherium   149
Genus Aureliachoerus Ginsburg, 1974   150
Species Aureliachoerus aurelianensis
(Stehlin, 18991900)   150
Artenay, France  150
Grépiac, France  150
Hessler, Germany  156
Büdenheim, Germany  165
Andelsbachtal, Germany  165
Brüttelen, Switzerland  165
Wildensbuch, Switzerland  166
Species Aureliachoerus minus
(Golpe-Possé, 1972)   168
Agreda, Spain  168
Tuchorice, Czech Republic  168
Wintershof-West, Germany  168
Benken, Switzerland 173
Chilleurs-aux-Bois, France  175
Chitenay, France  176
St. Lys, France  177
Oberdorf, Austria  177
Semsettin, Turkey  177
Genus Nguruwe Pickford, 1986   181
Species Nguruwe kijivium
(Wilkinson, 1976)   181
Napak, Uganda  181
Species Nguruwe namibensis
(Pickford, 1986)   189
Genus Xenohyus Ginsburg, 1980b   192
Species Xenohyus venitor
Ginsburg, 1980b   192
Genus indet. meisneri (Von Meyer, 1829)   192
Note concerning the validity of the name
Genus indet. meisneri   192
Genus and Species indet. possibly meisneri
(Von Meyer, 1829)   194
Family Doliochoeridae Simpson, 1945   194
Genus Doliochoerus Filhol, 1882a   194
Species Doliochoerus quercyi
Filhol, 1882a   194
Note on a species name which could
have priority over quercyi   194
Paulhiac, France  196
Aarwangen, Switzerland  201
Massif Central, France  202
Species Doliochoerus pusillus
(Ginsburg, 1974)   203
Tomerdingen, Germany  203
Genus Lorancahyus Pickford & Morales,
1998   209
Species Lorancahyus hypsorhizus
Pickford & Morles, 1998   209
Montaigu-le-Blin, France 209
Genus Propalaeochoerus Stehlin,
18991900   212
Note concerning the type species of
Propalaeochoerus Stehlin, 18991900   212
Species Propalaeochoerus gergovianus
(Croizet, 1846)   213
Cadibona, Italy  214
Quercy, France  215
Hochheim, Germany  218
Haslach bei Ulm, Germany  218
Species Propalaeochoerus suillus
(Pomel, 1853)   218
Species Propalaeochoerus paronae
Dal Piaz, 1930   223
Genus Bransatochoerus nov.   223
Species Bransatochoerus elaverensis
(Viret, 1929)   223
Genus Dubiotherium Hellmund, 1992   247
Species Dubiotherium waterhousi
(Pomel, 1853)   247
La Sauvetat (Pérignat), France  248
Status of Palaeochoerus massiliensis
Viret, 1929 and Doliochoerus aquensis
Repelin, 1930   248
St.-André, France  250
St.-Henri, France  252
Les Milles, France  252
Marseille, France  254
Quercy, France  254
General Discussion on Oligo-Miocene
Suoidea    256
Conclusions   257
Acknowledgements   260
References   261
Annex 1   270

This monograph treats bunodont suoids mainly from Europe, some from Africa and in passing mentions some Chinese material. It is an in-depth revision of the Hyotheriinae (Suidae) and Doliochoeridae (previously called the Palaeochoeridae) from Oligocene and Early to basal Middle Miocene levels, for which a summary of European Oligo-Miocene biostratigraphy is provided.

Much of the early work on Oligo-Miocene suids of Europe was carried out prior to the publication of the “Origin of Species” by Darwin (1859) a period during which ranges of morphological and metric variation were poorly understood or underestimated. In order to understand the historical background of the naming of genera and species of hyotheres and doliochoeres prior to Peters, 1869, the first paper which demonstrated the presence of sexual dimorphism in fossil suids, a detailed review of the old literature is included. This review attempts to explain the intellectual milieu, with its preconceptions, contradictions and belief values, within which the nascent taxonomy was done. It is clear that many of the early researchers were aware of the vastness of geological time, even if some of them underestimated its depth, but few if any of them resolved the succession of events – what we would now term biostratigraphy. The palaeontologists were thus severely handicapped when it came to interpreting their fossil samples. It is also clear that until Darwin’s (1859) publication, few researchers interpreted their fossils within a framework of natural selection. Until 1869, most workers on fossil suids did not take into account sexual dimorphism, some of them putting males and females into different genera. None of the earlier studies undertook statistical analyses of the fossils, but over the past Century some localities have yielded important samples, by which intraspecific variability could be documented. The list of European localities which have yielded rich fossil assemblages has increased to the point where it is now possible to be reasonably confident of the ranges of metric and morphological variation in many of the taxa. Nevertheless, there are several genera and species which are still relatively poorly represented in the fossil record.

It is concluded that the holotype of Choeropotamus meisneri from Rappenfluh, Switzerland, is so poorly preserved that the taxon is a nomen vanum. It could represent a suid, but appurtenance to Choeropotamidae cannot be excluded, although, on the basis of its radicular morphology, the specimen is unlikely to represent a doliochoerid. For over a Century, this species was included in Hyotherium on the basis of fossils from sites other than the type locality. Fossils from these sites are here attributed to Hyotherium major which, despite its name, is smaller than Hyotherium medium. Hyotherium medium is described in detail and a lectotype nominated from the sample from Mösskirch, Germany, that was studied by Von Meyer (1834). Much new material of Hyotherium major and Hyotherium soemmeringi is described. Good samples of Aureliachoerus aurelianensis and Aureliachoerus minus are described, which throw a great deal of light on morphometric variability in these species. The genus Aureliachoerus is closely related to the African genus Nguruwe.

The genus Palaeochoerus is likely to represent a suid rather than a peccary-like mammal, which makes continued use of the family name Palaeochoeridae Stehlin, 1909, caduque. Doliochoeridae Simpson, 1945, is here utilised for this family of suoids. Among the Doliochoeridae, there has been a great deal of confusion generated by mis-attribution of fossils between Doliochoerus and Propalaeochoerus. For example, a most recent diagnosis of Doliochoerus includes characters gleaned from specimens of Propalaeochoerus. A new genus of doliochoerid, Bransatochoerus, is erected for the abundant material from Coderet, France, which has unique features of the post-cranial skeleton suggesting that this species dug its own burrows, in which it lived and sometimes died. Additional material of Doliochoerus pusillus is illustrated, as are some interesting samples of Lorancahyus hypsorhizus. Much new material is added to Dubiotherium waterhousi, previously known only by two mandibles.

The monograph ends with comments on the phylogeny of doliochoerids and suids, but it is stressed that a great deal of further study is required to resolve the phylogenetic issues. These studies must take into account the post-cranial skeleton, as well as the dento-gnathic record upon which most previous attempts at phylogeny reconstruction were based. For instance, the species Doliochoerus pusillus possesses metapodials that are suid-like, rather than doliochoerid-like, which makes for a contradiction, as was pointed out by Hellmund & Heizmann (2015).