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European land mammal age

The European Land Mammal Mega Zones (abbreviation: ELMMZ, more commonly known as European land mammal ages or ELMA) are zones in rock layers that have a specific assemblage of fossils (biozones) based on occurrences of fossil assemblages of European land mammals. These biozones cover most of the Cenozoic, with particular focus having been paid to the Neogene and Paleogene systems (i.e. rock layers which are 65.5 to 2.588 million years old), the Quaternary has several competing systems. In cases when fossils of mammals are abundant, stratigraphers and paleontologists can use these biozones as a more practical regional alternative to the stages of the official ICS geologic timescale. European Land Mammal Mega Zones are often also confusingly referred to as ages, stages, or intervals.[1]

Biostratigraphic methods

Mammal zones were, like all biozones, established using geographic place names where fossil materials were obtained. The basic unit of measure is the first/last boundary statement. This shows that the first appearance event of one taxon is known to predate the last appearance event of another. If two taxa are found in the same fossil quarry or at the same stratigraphic horizon, then their age-range zones overlap.

The terrestrial stratigraphy of the Cenozoic is more difficult than that of marine deposits. The geologic timescale of the ICS is therefore based on marine fossils, that don't occur in terrestrial sediments. This makes the correlation of terrestrial deposits with the ICS timescale often difficult. Correlation is possible when marine deposits interfinger with terrestrial deposits (resulting from a series of transgressions and regressions of the sea during deposition), but this isn't the case everywhere. A fine stratigraphic division of the terrestrial record can in most places only be made using fossils of land species. Small mammals are often the best choice as they are quite abundant in the terrestrial record, especially their teeth. Teeth have an even better chance of preservation than bones.

The European mammalian biozones were established for the Paleogene (66-23.03 Mya, 8 zones) and Neogene (23.03-2.58 Mya, 7 zones) separately. Some of these, especially for the Neogene, were already established in the 19th century. The Villafranchian was, for example, introduced by Lorenzo Pareto in 1865. A finer subdivision was established by Pierre Mein in 1975, who divided the Neogene in 17 zones, known as the MN zonation, indicated by the letters MN (Mammal Neogene) and a number.

Similarly, a more detailed subdivision for the Paleogene period was established. There are 30 such Mammal Paleogene zones (MP1 to MP30, numbered from old to young).[2]

Paleogene European mammal zones

Epoch ICS age ELMMZ Age (Ma)
Miocene Aquitanian Agenian younger
Oligocene Chattian Arvernian 23.03–29.2
Suevian 29.2–33.8
Headonian 33.8–37.2
Eocene Priabonian
Bartonian Robiacian 37.2–42.7
Geiseltalian 42.7–48.5
Grauvian 48.5–50.8
Neustrian 50.8–55.0
Paleocene Thanetian
Cernaysian 55.0–55.9
Subdivision of the Paleogene period into European Land Mammal Mega Zones (ELMMZ).

There are 30 Mammal Paleogene zones covering the Paleogene (66-23.03 Mya).

Neogene European mammal zones

European Land Mammal Mega Zones most often have their bases at first appearances (FAD, First Appearance Date) of a certain species or genus. The numbers are higher for younger zones. Due to a redefinition of the boundary between the Neogene and Quaternary periods, MN 17 is now in fact considered a Quaternary biozone.

Biozone Small mammals Large mammals
MN 17 Kislangia gusi, Mimomys tornensis, Mimomys pliocaenicus, Mimomys reidi Eucladoceros
MN 16 Kislangia ischus, Mimomys polonicus, Kislangia cappettai, Mimomys hajnackensis Equus (horse), Mammuthus (mammoth), Homotherium, Megantereon, Ursus etruscus,
Pliohyaena perrieri, Gazellospira torticornis, Arvernoceros ardei, Hesperidoceras merlai, Cervus perrieri
MN 15 Mimomys occitanus, Oryctolagus, Mimomys vandemeuleni, Mimomys davakosi Chasmaporthetes lunensis
MN 14 Promimomys, Trilophomys, Celadensia, Castor (beaver) Sus arvernensis, Croizetoceros, Acinonyx, Lynx issiodorensis
MN 13 Paraethomys, Rhagapodemnus, Stephanomys, Apodemus, Apocricetus Parabos, Paracamelus, Agriotherium, Apocricetus, Nyctereutes, Hexaprotodon
MN 12 Parapodemus barbarae, Huerzelerimys turoliensis Pliocervus, Hispanodorcas, Palaeoryx, Occitanomys adroveri, Procapreolus
MN 11 Parapodemus lugdunensis, Huerzelerimys vireti, Occitanomys sondaari Birgerbohlinia, Lucentia
MN 10 Rotundomys, Pliopetaurista, Schreuderia, Progonomys cathalai Hyaenictis almerai, Adcrocuta eximia, Microstonyx major, Tragoportax gaufryi
MN 9 Cricetulodon Hippotherium, Decennatherium, Machairodus
MN 7/8 Megacricetodon ibericus, Megacricetodon gregarius Parachleuastochoerus, Propotamochoerus, Palaeotragus, Protragocerus, Tetralophodon
MN 6 Megacricetodon crusafonti, Megacricetodon gersi Tethytragus, Hispanomeryx, Euprox, Listriodon
MN 5 Megacricetodon collongensis Miotragocerus, Micromeryx, Heteroprox, Hispanotherium
MN 4 Megacricetodon primitivus Bunolistriodon, Dorcatherium, Chalicotherium, Eotragus, Prodeinotherium
MN 3 Gomphotherium, Procervulus, Lagomeryx, Actoocemas, Palaeomeryx, Brachyodus, Anchitherium, Aureliachoerus, Hemicyon
MN 2 Ligerimys, Prolagus, Lagopsis, Ritterneria manca Teruelia, Lorancameryx, Oriomeryx, Pseudaelurus, Xenohyus, Andegameryx, Amphitragulus
MN 1 Rhodanomys schlosseri, Vasseuromys Hyotherium

Quaternary European mammal zones

The first zonation for the Quaternary of Europe was proposed by Azzaroli in 1967.[3] This was then expanded by Gliozzi et al. in 1997 to make a system of 3 'ages' subdivided into 13 'faunal units'.[4] The scheme does not define boundaries but instead is accompanied by a range chart, where the entry and exit dates for the taxa are indicated. Each zone is named after a reference locality. Most of the reference locations are in Italy but the scheme is used in other European regions. The mammal ages and Faunal Units (FU) after Gliozzi et al. are:

Mammal Ages after Gliozzi et al., 1997[4][5][6][7][8][9]
Mammal Age Faunal Unit sub-age Large Mammals
(Middle Pliocene to Early Pleistocene)
Triversa Early Villafranchian Pliorhinus megarhinus, Sus minor, Alephis liryx, Felsinotherium gervaisi
Montopoli Early Villafranchian Mammuthus gromovi, Equus lioenzwensi, Stephanorhinus etruscus
Costa S. Giacomo Middle Villafranchian Canis, Sus strozzi, Leptobos furtivus, Hystrix refossa, Gazellospira torticomis, Anancus arvernensis
Olivola Late Villafranchian Pachycrocuta brevirostris, Panthera gombaszoegensis, Procamptoceras brivatense, Eucladoceros dicranios, Eucladoceros nestii, Pseudodama, Canis etruscus
Tasso Late Villafranchian Hippopotamus antiquus, Praevibos, Leptobos vallisarni, Equus stehlini, Canis arnensis, Canis falconeri
Farnetta Late Villafranchian Leptobos vallisarni, Eucladoceros dicranios, Eucladoceros nestii, Praemegaceros obscurus, Microtus
Piro Nord Late Villafranchian Bison, Praemegaceros solilhacus
(Middle Pleistocene)
Colle Curti Early Galerian Equus altidens, Equus bressanus, Equus sussenbornensis, Praemegaceros verticornis
Silvia Middle Galerian Cervus elaphus acoronatus
Isernia Middle Galerian Panthera leo, Palaeoloxodon antiquus, Mammuthus trogontherii, Stephanorhinus hundsheimensis, Megaloceros savini, Bos, Bison schoetensacki, Equus caballus, Pseudodama
Fontana Ranuccio Late Galerian Homotherium latidens, Cervus elaphus eostephanoceros, Dama clactoniana, Equus altidens, Equus sussenbornensis
(late Middle Pleistocene to Late Pleistocene)
Torre in Pietra Early Aurelian Canis lupus, Ursus spelaeus, Megaloceros giganteus, Cervus elaphus rianensis
Vitinia Middle Aurelian Dama dama tiberina, Equus hydruntinus
none designated Late Aurelian Dama dama dama, Capra ibex, Coelodonta antiquitatis, Mammuthus primigenius

In 1982, Guérin[10] proposed an alternative scheme, which extended the MN zonation scheme for the Neogene with additional units to cover the Quaternary. There have been further updates since.[11] The MNQ (Mammal Neogene Quaternary) scheme added an additional 12 units in total, MNQ 16–27.

For small mammals there is a third scheme, the MmQ, published by Agustí, Moyà‐Solà, and Pons‐Moyà in 1987.[12] The scheme includes some large mammals for reference and thus has a wider application.[13]

Other continental mammalian biozones


  1. ^ According to Steininger (1999), it is better to just use ELMMZ's in a biostratigraphic sense
  2. ^ Mammal Paleogene zones Archived 2012-10-12 at the Wayback Machine, The Paleobiology Database
  3. ^ Azzaroli, Augusto. 1967. “Villafranchian Correlations Based on Large Mammals.” Giornale di Geologia 35 (1): 21
  4. ^ a b Gliozzi, Elsa, Laura Abbazzi, Patrizia Argenti, Augusto Azzaroli, Lucia Caloi, Lucia Capasso Barbato, Giuseppe Di Stefano, et al. 1997. “Biochronology of Selected Mammals, Molluscs and Ostracods from the Middle Pliocene to the Late Pleistocene in Italy: The State of the Art.” Rivista Italiana di Paleontologia e Stratigrafia 103 (3): 369–88.
  5. ^ Kurten, Bjorn (1968). Pleistocene Mammals of Europe. London: Weidenfeld and Nicolson.
  6. ^ Pandolfi, Luca; Pierre-Olivier, Antoine; Bukhsianidze, Maia; Lordkipanidze, David; Rook, Lorenzo (2021-08-03). "Northern Eurasian rhinocerotines (Mammalia, Perissodactyla) by the Pliocene–Pleistocene transition: phylogeny and historical biogeography". Journal of Systematic Palaeontology. 19 (15): 1031–1057. doi:10.1080/14772019.2021.1995907. ISSN 1477-2019.
  7. ^ Frantz, Laurent; Meijaard, Erik; Gongora, Jaime; Haile, James; Groenen, Martien A.M.; Larson, Greger (2016-02-15). "The Evolution of Suidae". Annual Review of Animal Biosciences. 4 (1): 61–85. doi:10.1146/annurev-animal-021815-111155. ISSN 2165-8102.
  8. ^ Lister, Adrian M.; Sher, Andrei V.; van Essen, Hans; Wei, Guangbiao (2005-01-01). "The pattern and process of mammoth evolution in Eurasia". Quaternary International. Studying Proboscideans: knowledge, Problems and Perspectives. Selected papers from "The world of Elephants" Congress, Rome. 126–128: 49–64. doi:10.1016/j.quaint.2004.04.014. ISSN 1040-6182.
  9. ^ Croitor, Roman (2018). Plio-Pleistocene Deer of Western Palearctic: Taxonomy, Systematics, Phylogeny. Institute of Zoology of the Academy of Sciences of Moldova. ISBN 978-9975-66-609-1. OCLC 1057238213.
  10. ^ Guérin, Claude. 1982. “Première Biozonation du Pléistocène Européen, Principal Résultat Biostratigraphique de l'Étude des Rhinocerotidae (Mammalia, Perissodactyla) du Miocène Terminal au Pléistocène Supérieur d'Europe Occidentale.” Geobios 15 (4): 593–98. DOI:10.1016/s0016-6995(82)80076-4
  11. ^ Faure, Martine, and Claude Guérin. 1992. “La Grande Faune d'Europe Occidentale au Pléistocène Moyen et Supérieur et ses Potentialités d'Information en Préhistoire.” Mémoires de la Société Géologíque de France 160: 77–84.
  12. ^ Agustí, Jordi, Salvador Moyà‐Solà, and Joan Pons‐Moyà. 1987. “La Sucesión de Mamíferos en el Pleistoceno Inferior de Europa: Proposición de Una Nueva Escala Bioestratigráfica.” Paleontología i Eovlució, mem. esp. 1: 287–95.
  13. ^ van der, Made, J. (2018). Quaternary Large‐Mammal Zones. In The Encyclopedia of Archaeological Sciences, S.L. López Varela (Ed.). doi:10.1002/9781119188230.saseas0486



  • Koufos, G.D.; Kostopoulos, D.S. & Vlachou, T.D. (2005). '"Neogene/Quaternary mammalian migrations in Eastern Mediterranean", Belgian Journal of Zoology 135(2): pp. 181–190.
  • Lindsay, Everett (October 1997). "Eurasian mammal biochronology: an overview". Palaeogeography, Palaeoclimatology, Palaeoecology. 133 (3–4): 117–128. Bibcode:1997PPP...133..117L. doi:10.1016/S0031-0182(97)00083-7.
  • Mein, P. (1975). Report on activity RCMNS-Working groups, 1971–1975, pp. 78–81, Bratislava.
  • Rose, Kenneth D. (2006). The beginning of the age of mammals. Baltimore, Md.: Johns Hopkins Univ. Press. ISBN 978-0801884726.
  • Steininger, F.F. (1999). "Chronostratigraphy, Geochronology and Biochronology of the Miocene "European Land Mammal Mega-Zones" (ELMMZ) and the Miocene "Mammal Zones (MN-Zones)"". In Rössner, G. E.; Heissig, K. (eds.). The Miocene land mammals of Europe. München: Pfeil. pp. 9–24. ISBN 3-931516-50-4.
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European land mammal age
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