Maramena (Miocene of Greece)

Where: Central Macedonia, Greece (41.2° N, 23.5° E: paleocoordinates 40.9° N, 23.2° E)

• coordinate estimated from map

• small collection-level geographic resolution

When: Lefkon Formation, Turolian (8.7 - 5.3 Ma)

• may be the same level as Ano Metochi 2-3

Environment/lithology: alluvial fan; conglomerate and sandy marl

• A polymict poorly sorted conglomerate containing pebbles of schist, gneiss and granite. This coarse-grained series grades upwards into arkosic sands and finally into clays with occasional lignites. Within the top part of the formation, two "landslide" breccias occur. The lower of these consists of metamorphic rocks, the upper one of granite.

Size class: mesofossils

Reposited in the AMPG

Primary reference: H. de Bruijn. 1989. Smaller mammals from the Upper Miocene and Lower Pliocene of the Strimon basin, Greece. Part 1. Rodentia and Lagomorpha. Bollettino della Societa Paleontologica Italiana 28(2-3):189-195 [J. Alroy/E. Leckey/E. Leckey]more details

Purpose of describing collection: taxonomic analysis

PaleoDB collection 32189: authorized by John Alroy, entered by Erin Leckey on 09.06.2003, edited by Lars van den Hoek Ostende, Torsten Liebrecht and Evangelos Vlachos

Creative Commons license: CC BY (attribution)

Taxonomic list

unclassified
  -
Milax sp.3
or Tandonia sp.
Norbertia hellenica n. gen. n. sp.3
Ommatotriton sp.2
M1: a vertebra (UU MAA 7489) and two femurs (UU MAA 7248 and UU MAA 7509).
Bivalvia
 Cardiida - Sphaeriidae
Pisidium casertanum3 Poli 1791 pea clam
Gastropoda
 Architaenioglossa - Aciculidae
"cf. Platyla sp." = Acme (Platyla)3
"cf. Platyla sp." = Acme (Platyla)3 Moquin-Tandon 1855 snail
 Rissooidea - Bithyniidae
Bithynia sp.3 Leach 1818 snail
 Pulmonata - Ellobiidae
 Pulmonata - Clausiliidae
Clausiliidae indet.3 Gray 1855 slug
 Pulmonata - Cochlicopidae
Cochlicopa sp.3 Ferussac 1821 slug
 Pulmonata - Parmacellidae
Parmacella sp.3 Cuvier 1804 slug
nov
 Helicoidea -
Helicoidea indet.3 Rafinesque 1815 slug
sp. nov.
 Limacoidea - Limacidae
cf. Limax sp.3 Linnaeus 1758 slug
 Basommatophora - Planorbidae
Planorbarius sp.3 Dumeril 1806 pulmonate
cf. conru-mantelli
Gyraulus sp.3 Charpentier 1837 pulmonate
Planorbis sp.3 Muller 1774 pulmonate
Pisces
  -
"Pisces indet." = Osteichthyes3
"Pisces indet." = Osteichthyes3 Huxley 1880 bony fish
Osteichthyes
 Salientia -
Latonia sp.2 von Meyer 1843 frog
M1: two fragmentary frontoparietals belonging to a single individual (UU MAA 7445 and UU MAA 7462), three premaxillae (UU MAA 7130), 66 maxillae (UU MAA 7120, UU MAA 7128, UU MAA 7131, UU MAA 7156, UU MAA 7249, UU MAA 7250, UU MAA 7458, and UU MAA 7464-UU MAA 7472), seven angulars (UU MAA 7154 and UU MAA 7529), an atlas (UU MAA 7459), 17 presacral vertebrae (UU MAA 7125, UU MAA 7158a, UU MAA 7442-UU MAA 7444, UU MAA 7452, UU MAA 7454, UU MAA 7457, UU MAA 7460, and UU MAA 7461), three sacral vertebrae (UU MAA 7455 and UU MAA 7522), four urostyles (UU MAA 7446, UU MAA 7450, and UU MAA 7521), a scapula (UU MAA 7520), four costae (UU MAA 7523), 14 humeri (UU MAA 7155, UU MAA 7158b, UU MAA 7453, and UU MAA 7463), and nine ilia (UU MAA 7447-UU MAA 7449, UU MAA 7456, and UU MAA 7528); M2: a fragmentary frontoparietal (UU MAA 7192); M3: seven maxillae (UU MAA 7223, UU MAA 7224, UU MAA 7227, UU MAA 7238, UU MAA 7239, UU MAA 7240, and UU MAA 7241), a presacral vertebra (UU MAA 7228), a humerus (UU MAA 7225), and an ilium (UU MAA 7229).
 Salientia - Ranidae
Ranidae indet.2 Rafinesque 1814 frog
M2: three ilia (UU MAA 7195); M3: three humeri (UU MAA 7231).
Pelophylax sp.2 Fitzinger 1843 frog
M2: an angular (UU MAA 7194); M3: a coracoid (UU MAA 7233) and three ilia (UU MAA 7232).
 Salientia - Hylidae
Hyla sp.2 Laurenti 1768 tree frog
M1: seven humeri (UU MAA 7119, UU MAA 7244, UU MAA 7251, and UU MAA 7474-UU MAA 7476) and an ilium (UU MAA 7118); M2: three humeri (UU MAA 7193).
 Caudata - Urodela
Cryptobranchidae indet.2 Cope 1889 hellbender
M1: an atlas (UU MAA 7441), probably also a second atlas (UU MAA 7530)
Salamandridae indet.2 Gray 1825 newt
M1: an atlas (UU MAA 7490), four vertebrae (UU MAA 7153, UU MAA 7485, UU MAA 7488, and UU MAA 7243), and two ribs (UU MAA 7508 and UU MAA 7519).
Salamandrina sp.2 Fitzinger 1826 spectacled salamander
M1: two vertebrae (UU MAA 7483 and UU MAA 7484)
Lissotriton sp.2 Bell 1839 newt
(Lissotriton vulgaris group) M1: two vertebrae (UU MAA 7517 and UU MAA 7518).
Salamandra sp.2 Laurenti 1768 salamander
M1: 12 trunk vertebrae (UU MAA 7494- UU MAA 7505) and six caudal vertebrae (UU MAA 7117, UU MAA 7491-UU MAA 7493, UU MAA 7506, and UU MAA 7507).
Reptilia
 Testudines -
Testudines indet.2 Batsch 1788 turtle
M1: an incomplete unidentified neural (UU MAA 7136), three unidentified peripherals (UU MAA 7137, UU MAA 7140, and UU MAA 7142), and several shell fragments (UU MAA 7138, UU MAA 7139, UU MAA 7147 [five fragments]); M3: an unidentified peripheral (UU MAA 7254), a shell fragment (UU MAA 7252), and several fragments of different elements (one caudal vertebra, one pelvic fragment, three neurals, four costals, one peripheral, and 24 shell fragments) (UU MAA 7255).
 Testudines - Pantestudinidae
? Testudinidae indet.2 Batsch 1788 turtle
M1: an isolated ?limb osteoderm (UU MAA 7135).
Testudo sp.5 Linnaeus 1758 turtle
 Testudines - Pangeoemydidae
Mauremys aristotelica2 Vlachos et al. 2019 pond turtle
M1: three incomplete unidentified neurals (UU MAA 7143, UU MAA 7144, and UU MAA 7146); M3: a nuchal (UU MAA 7283), a fragmentary unidentified neural (UU MAA 7253), a left costal VI (UU MAA 7284), unidentified costals (UU MAA 7286 and UU MAA 7288), a left peripheral III (UU MAA 7289), a right peripheral III (UU MAA 7290), unidentified peripherals (UU MAA 7285, UU MAA 7287, UU MAA 7291, UU MAA 7292, UU MAA 7293, UU MAA 7295, and UU MAA 7297), and a right hyoplastron (UU MAA 7296).
 Squamata -
Squamata indet.2 Oppel 1811 squamates
M1: a premaxilla (UU MAA 7093), eight maxillae (UU MAA 7342, UU MAA 7355, UU MAA 7356, UU MAA 7360, and UU MAA 7428), six dentaries (UU MAA 7103, UU MAA 7105, and UU MAA 7427), 14 fragments of tooth bearing bones (UU MAA 7102, UU MAA 7311, UU MAA 7354, UU MAA 7393, UU MAA 7405, and UU MAA 7406), three axes (UU MAA 7346 and UU MAA 7378), three post-axial cervical vertebrae (UU MAA 7322, UU MAA 7348, and UU MAA 7789), 13 trunk vertebrae (UU MAA 7110, UU MAA 7345, UU MAA 7382-UU MAA 7384, UU MAA 7421), two cloacal vertebrae (UU MAA 7380 and UU MAA 7381), eight caudal vertebrae (UU MAA 7347, UU MAA 7385-UU MAA 7387, and UU MAA 7390), a fragment of caudal vertebra (UU MAA 7310), 11 ribs (UU MAA 7111 and UU MAA 7419), seven humeri (UU MAA 7309 and UU MAA 7352), two pelvic girdles (UU MAA 7349 and UU MAA 7376), 14 femurs (UU MAA 7109, UU MAA 7308, UU MAA 7351, UU MAA 7377, and UU MAA 7420), and a tibia (UU MAA 7350); M2: a fragment of a presacral vertebra (UU MAA 7299) and a caudal vertebra (UU MAA 7300); M3: a fragment of tooth bearing bone (UU MAA 7339) and a caudal vertebra (UU MAA 7221).
 Squamata - Lacertidae
Lacertidae indet.2 Bonaparte 1831 squamates
M1: three maxillae (UU MAA 7031, UU MAA 7032, and UU MAA 7438), a quadrate (UU MAA 7096), three pterygoids (UU MAA 7097-UU MAA 7099), 16 dentaries (UU MAA 7048-UU MAA 7061, UU MAA 7095, and UU MAA 7424), 10 fragments of tooth bearing bones (UU MAA 7087-UU MAA 7092, UU MAA 7094, UU MAA 7391, and UU MAA 7802), and an isolated tooth (UU MAA 7392); M3: a dentary (UU MAA 7220).
 Squamata -
Laterata indet.2 Vidal and Hedges 2005 squamates
M1: 29 dentaries (UU MAA 7026, UU MAA 7106, UU MAA 7357, UU MAA 7361, UU MAA 7379, UU MAA 7402-UU MAA 7404, UU MAA 7435, and UU MAA 7436); M3: a dentary (UU MAA 7206).
Anguimorpha indet.2 Fürbringer 1900 squamates
M1: a dentary (UU MAA 7353).
 Squamata - Varanidae
Varanus sp.2 Merrem 1820 monitor lizard
M1: six teeth (UU MAA 7029, UU MAA 7030, and UU MAA 7148-UU MAA 7151) and 12 caudal vertebrae (UU MAA 7028, UU MAA 7278, UU MAA 7388, UU MAA 7389, UU MAA 7429-UU MAA 7434, UU MAA 7805, and UU MAA 7806); M3: a caudal vertebra (UU MAA 7198).
 Squamata - Anguidae
Anguinae indet.2 Gray 1825 squamates
M1: 188 osteoderms (UU MAA 7160-UU MAA 7170, UU MAA 7262, UU MAA 7265, UU MAA 7268, UU MAA 7271, UU MAA 7275, UU MAA 7306, and UU MAA 7314-UU MAA 7318); M2: eight osteoderms (UU MAA 7187, UU MAA 7188, and UU MAA 7298).
Pseudopus sp.2 Merrem 1820 legless lizard
M1: a maxilla (UU MAA 7171), a dentary (UU MAA 7407), a fragment of tooth bearing bone (UU MAA 7307), two trunk vertebrae (UU MAA 7172 and UU MAA 7328), and a cloacal vertebra (UU MAA 7329); M3: three trunk vertebrae (UU MAA 7212-UU MAA 7214).
Anguis sp.2 Linnaeus 1758 slow worm
M1: 37 trunk vertebrae (UU MAA 7181, UU MAA 7263, UU MAA 7274, UU MAA 7280, UU MAA 7332-UU MAA 7334, UU MAA 7411, UU MAA 7418, and UU MAA 7804), a cloacal vertebra (UU MAA 7269); M3: two trunk vertebrae (UU MAA 7217 and UU MAA 7218).
Ophisaurus sp.2 Daudin 1803 glass lizard
M1: a maxilla (UU MAA 7324) and two dentaries (UU MAA 7176 and UU MAA 7177).
 Squamata - Agamidae
Agaminae indet.2 squamates
M1: a premaxilla (UU MAA 7076), three maxillae (UU MAA 7043-UU MAA 7045); five dentaries (UU MAA 7041, UU MAA 7042, UU MAA 7075, UU MAA 7276, and UU MAA 7305); 15 fragments of tooth bearing bones (UU MAA 7063, UU MAA 7065-UU MAA 7074, UU MAA 7264, UU MAA 7303, UU MAA 7304, and UU MAA 7394); five caudal vertebrae (UU MAA 7062, UU MAA 7064, and UU MAA 7302); M2: a frontal (UU MAA 7191); M3: a premaxilla (UU MAA 7341), a maxilla (UU MAA 7208), and a left dentary (UU MAA 7207).
 Squamata -
Serpentes indet.4 Linnaeus 1758 snake
6 precaudal and caudal vertebrae (AMPG MA 190 to 195), "may have belonged either to Naja or Vipera"
 Squamata - Elapidae
Naja sp.4 Laurenti 1768 cobra
25 fragmentary precaudal vertebrae (AMPG MA 163 to 187)
 Squamata - Colubridae
Periergophis micros n. gen. n. sp.2 Georgalis et al. 2019 colubrid snake
Holotype. Specimen UU MAA 7615 (a posterior trunk vertebra) (Figures 22 and 24.1-2). The holotype originates from M1 site. Paratypes. Specimen UU MAA 7551 (a cervical vertebra) (Figure 23.1-2); UU MAA 7614 (a midtrunk vertebra) (Figure 23.3-8); UU MAA 7618 (a cloacal vertebra) (Figure 23.9-12); UU MAA 7636 (a caudal vertebra) (Figures 23.13-16, 24.3). All paratypes originate from M1 site.
Colubrinae indet.4 Oppel 1811 colubrid snake
36 fragmentary precaudal vertebrae (AMPG MA 101 to 136)
Paraxenophis spanios n. gen. n. sp.2 Georgalis et al. 2019 colubrid snake
Holotype. Specimen UU MAA 7645 (a mid-trunk vertebra) (Figure 27). The holotype originates from M1 site. Paratypes. Specimen UU MAA 7536 (a posterior trunk vertebra) (Figure 29.1-5); UU MAA 7533 (a posteriormost trunk vertebra) (Figure 29.6-8). Both paratypes originate from M1 site.
Natricinae indet.4 Bonaparte 1838 colubrid snake
26 fragmentary precaudal vertebrae (AMPG MA 137 to 162)
Natrix aff. rudabanyaensis2 water snake
M1: 34 precloacal vertebrae (UU MAA 7656-UU MAA 7689) and seven caudal vertebrae (UU MAA 7690-UU MAA 7694, UU MAA 7893, and UU MAA 7904); M3: four precloacal vertebrae (UU MAA 7741-UU MAA 7744).
Natrix sp.2 Laurenti 1768 water snake
M1: six precloacal vertebrae (UU MAA 7695-UU MAA 7700).
 Squamata - Crotalidae
Vipera sp.4 Laurenti 1768 pit viper
"oriental viper", 2 fragmentary precaudal vertebrae (AMPG MA 188, 189)
 Squamata -
Scolecophidia indet.2 snake
M1: three trunk vertebrae (UU MAA 7373-UU MAA 7375).
 Squamata - Scincidae
? Scincidae indet.2 Gray 1825 skink
M1: two dentaries (UU MAA 7027 and UU MAA 7107).
 Squamata - Cordylidae
aff. Palaeocordylus sp.2 Roček 1984 spinytail lizard
M1: 26 maxillae (UU MAA 7046, UU MAA 7047, UU MAA 7077-UU MAA 7086, UU MAA 7101, UU MAA 7273, UU MAA 7358, UU MAA 7362-UU MAA 7365, UU MAA 7370-UU MAA 7372, UU MAA 7395, UU MAA 7396, and UU MAA 7437), 57 dentaries (UU MAA 7001-UU MAA 7025, UU MAA 7034-UU MAA 7036, UU MAA 7039, UU MAA 7040, UU MAA 7104, UU MAA 7108, UU MAA 7266, UU MAA 7267, UU MAA 7270, UU MAA 7279, UU MAA 7359, UU MAA 7366-UU MAA 7369, UU MAA 7397-UU MAA 7401, UU MAA 7425, and UU MAA 7426), and six fragments of tooth bearing bones (UU MAA 7037 and UU MAA 7038); M3: a maxilla (UU MAA 7340), eight dentaries (UU MAA 7199-UU MAA 7205, and UU MAA 7209), and a fragment of tooth bearing bone (UU MAA 7210).
Mammalia
 Primates - Cercopithecidae
Mesopithecus pentelicus3 Wagner 1839 monkey
 Rodentia - Muridae
Apodemus dominans Eurasian field mouse
A. gudrunae+A.dominans=approx. 390 specimens
Apodemus gudrunae van de Weerd 1976 Eurasian field mouse
A. gudrunae+A.dominans=approx. 390 specimens
Micromys "sp. A" Dehne 1841 harvest mouse
approx.
Occitanomys cf. neutrum de Bruijn 1976 mouse
approx.
 Rodentia - Cricetidae
"Kowalskia fahlbuschi" = Neocricetodon fahlbuschi, "Cricetus" "sp. A"
"Kowalskia fahlbuschi" = Neocricetodon fahlbuschi Bachmayer and Wilson 1970 mouse
Kowalskia + Cricetus= approx. 83 specimens
"Cricetus" "sp. A" Leske 1779 mouse
Kowalskia + Cricetus= approx. 83 specimens
 Rodentia - Sciuridae
Tamias "sp. A" Illinger 1811 chipmunk
Tamias + Protospermophilus=approx. 13 specimens
Miopetaurista thaleri3 Mein 1970 squirrel
Hylopetes macedoniensis squirrel
Pliopetaurista + Hylopetes=approx. 50 specimens
? Protospermophilus sp. Gazin 1930 squirrel
Tamias + Protospermophilus=approx. 13 specimens
Pliopetaurista dehneli squirrel
Pliopetaurista + Hylopetes=approx. 50 specimens
 Rodentia - Gliridae
Muscardinus pliocaenicus3 Kowalski 1963 dormouse
Glis cf. minor3 Kowalski 1963 dormouse
Myomimus dehmi3 de Bruijn 1966 dormouse
Myomimus maritsensis de Bruijn et al. 1970 dormouse
approx.
 Rodentia - Eomyidae
Keramidomys aff. carpathicus Schaub and Zapfe 1953 rodent
probably less than 3 specimens
 Rodentia - Castoridae
Castoridae indet. Gray 1821 beaver
probably less than 3 specimens
 Lagomorpha - Leporidae
Alilepus turolensis rabbit
Alilepus + Prolagus = 41 specimens
 Lagomorpha - Ochotonidae
Prolagus michauxi pika
Alilepus + Prolagus = 41 specimens
 Lipotyphla - Erinaceidae
Schizogalerix macedonica n. sp.1 Doukas et al. 1995 hedgehog
 Lipotyphla - Talpidae
"Ruemkelia dekkersi" = Archaeodesmana1 mole
Desmanella dubia1 Rümke 1976 mole
Urotrichus sp.1 Temminck 1841 greater Japanese shrew-mole
Talpa fossilis1 Petenyi 1864 mole
 Perissodactyla - Rhinocerotidae
Rhinocerotidae indet.3 Gray 1821 rhinoceros
 Perissodactyla - Equidae
Hipparion sp.3 de Christol 1832 hipparionine horse
2 species
 Tribosphenida - Soricidae
Sorex sp.1 Linnaeus 1758 long-tailed shrew
Deinsdorfia kerkhoffi n. sp.1 Doukas et al. 1995 red-toothed shrew
Petenyia hungarica1 Kormos 1934 red-toothed shrew
Amblycoptus jessiae n. sp.1 Doukas et al. 1995 red-toothed shrew
 Chiroptera -
Chiroptera indet.3 Blumenbach 1779 bat
 Carnivora - Mustelidae
Lutra affinis3 Gervais 1859 otter
Martes lefkonensis n. sp.3 marten
Promephitis sp.3 Gaudry 1861 mustelid carnivore
 Carnivora - Viverridae
Viverridae indet.3 Gray 1821 civet
 Carnivora - Hyaenidae
Chasmaporthetes sp.3 Hay 1921 hyaena
 Proboscidea - Gomphotheriidae
 Ungulata - Suidae
 Ungulata - Giraffidae
Samotherium cf. boissieri3 Forsyth Major 1888 giraffe
 Ungulata - Bovidae
Boselaphini indet.3 Knottnerus-Meyer 1997 antelope
Tragoportax gaudryi3 Kretzoi 1941 antelope
Tragoportax cf. amalthea3 Roth and Wagner 1854 antelope
Gazella sp.3 Blaineville 1816 gazelle
 Ungulata - Cervidae
Pliocervus graecus n. sp.3 deer
 Theriiformes -
Erinaceus samsonowiczi1 Sulimski 1959 mammal