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N number of complete specimens Table 1 The minimum, mean and maximum values (in mm) of the maximum length and maximum width and the length/width ratio of th Palaeoleuciscus sp. and Barbus s.1. lapilli, following Schulz-Mirbach and Reichenbacher (2006)

Table 1 N number of complete specimens Table 1 The minimum, mean and maximum values (in mm) of the maximum length and maximum width and the length/width ratio of th Palaeoleuciscus sp. and Barbus s.1. lapilli, following Schulz-Mirbach and Reichenbacher (2006)

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Fig. 1 Lesvos Island in the NE Aegean Sea. GAV fossiliferous locality of Gavathas, LRN fossiliferous locality of Lapsarna, NHMLPF Natural History Museum of the Lesvos Petrified Forest; the dark grey color shows the protected area of the Lesvos Petrified Forest One of the most renowned geological monuments in the Aegean area, the Lesvos Petrified Forest, which covers a large part of western Lesvos Island (also known as Lesbos) in Greece (Fig. 1), has been providing information on the Early Miocene subtropical flora of the Aegean—Anatolian area for many decades. The petrified tree trunks, branches,
Fig. 1 Lesvos Island in the NE Aegean Sea. GAV fossiliferous locality of Gavathas, LRN fossiliferous locality of Lapsarna, NHMLPF Natural History Museum of the Lesvos Petrified Forest; the dark grey color shows the protected area of the Lesvos Petrified Forest One of the most renowned geological monuments in the Aegean area, the Lesvos Petrified Forest, which covers a large part of western Lesvos Island (also known as Lesbos) in Greece (Fig. 1), has been providing information on the Early Miocene subtropical flora of the Aegean—Anatolian area for many decades. The petrified tree trunks, branches,
Fig. 2 Synthetic stratigraphical column with the rock types in western Lesvos (after Mourouzidou 2001 and Koufos et al. 2003, modified). 7 Neopaleozoic—Triassic basement rocks (mainly schists, phyllites and carbonate rocks); 2 Tectonic nappe of the ophiolitic sequence overthrusting the previous unit; 3 Neogene lacustrine deposits (alternating strongly silicified marly limestones and marls with thin lignitic intercalations); 4 lower ignimbrite (small lava conglomerates and volcanic ashes); 5 pyroclastic deposits (mud flows with a debris flow in the upper part); 6 upper ignimbrite (black glassy lavas); 7 Lava Unit (dacites and dakito-andesites); 8 recent alluvial deposits. The white bone shows the vertical position of the Lapsarna fossiliferous layer in the upper part of the Neogene lacustrine deposits in the Gavathas-Lapsarna Basin. Layers 4-6 correspond to the Acid Volcanic Unit of Pe-Piper (1980). Details on the geological setting and characteristics of the LRN locality, as well as maps, photographs of the sampled section and a synthetic stratigraphical column of western Lesvos can be found in Vasileiadou and Zouros (2012) Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
Fig. 2 Synthetic stratigraphical column with the rock types in western Lesvos (after Mourouzidou 2001 and Koufos et al. 2003, modified). 7 Neopaleozoic—Triassic basement rocks (mainly schists, phyllites and carbonate rocks); 2 Tectonic nappe of the ophiolitic sequence overthrusting the previous unit; 3 Neogene lacustrine deposits (alternating strongly silicified marly limestones and marls with thin lignitic intercalations); 4 lower ignimbrite (small lava conglomerates and volcanic ashes); 5 pyroclastic deposits (mud flows with a debris flow in the upper part); 6 upper ignimbrite (black glassy lavas); 7 Lava Unit (dacites and dakito-andesites); 8 recent alluvial deposits. The white bone shows the vertical position of the Lapsarna fossiliferous layer in the upper part of the Neogene lacustrine deposits in the Gavathas-Lapsarna Basin. Layers 4-6 correspond to the Acid Volcanic Unit of Pe-Piper (1980). Details on the geological setting and characteristics of the LRN locality, as well as maps, photographs of the sampled section and a synthetic stratigraphical column of western Lesvos can be found in Vasileiadou and Zouros (2012) Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest Fig. 3. Freshwater truncatelloid Gastropoda from Lapsarna, Lesvos Island, Greece. Bithynia sp. opercula: a LRNg-22; b LRNg-25; c LRNg-21; Emmericia sp.: d LRNg-10; e LRNg-13; f LRNg-4; g LRNg-3. Hydrobiidae indet.: h LRNg-31; i LRNg-32; note that the specimen of i shows, quite misleadingly, a deeper suture than that of h, which is a result of deformation rather than a morphological difference; ? Bania sp.: j LRNg-55; k LRNg-57; 1 LRNg-65; m LRNg- 43; n LRNg-45; o LRNg-43, protoconch of m; scale bars equal 1 mm (a-g), 0.5 mm (A-n) and 0.1 mm (0)
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest Fig. 3. Freshwater truncatelloid Gastropoda from Lapsarna, Lesvos Island, Greece. Bithynia sp. opercula: a LRNg-22; b LRNg-25; c LRNg-21; Emmericia sp.: d LRNg-10; e LRNg-13; f LRNg-4; g LRNg-3. Hydrobiidae indet.: h LRNg-31; i LRNg-32; note that the specimen of i shows, quite misleadingly, a deeper suture than that of h, which is a result of deformation rather than a morphological difference; ? Bania sp.: j LRNg-55; k LRNg-57; 1 LRNg-65; m LRNg- 43; n LRNg-45; o LRNg-43, protoconch of m; scale bars equal 1 mm (a-g), 0.5 mm (A-n) and 0.1 mm (0)
Fig. 4 Freshwater pulmonate Gastropoda from Lapsarna, Lesvos Island, Greece. ? Radix sp.: a LRNg-93; b LRNg-94; c LRNg-95, apical view; Lymnaeidae indet.: d LRNg-71; e LRNg-81; f LRNg-81, apical view; Gyraulus sp.: g LRNg-110; h LRNg-30; i LRNg-127; j LRNg-101, protoconch; k LRNg-110, protoconch of g; Planorbarius sp.: 1 LRNg-141; m LRNg-142; n LRNg-141, protoconch of n; o LRNg-142, protoconch of m. Scale bars equal 1 mm (a-f, [-m), 0.5 mm (g-i, n, 0) and 0.1 mm (, k) Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
Fig. 4 Freshwater pulmonate Gastropoda from Lapsarna, Lesvos Island, Greece. ? Radix sp.: a LRNg-93; b LRNg-94; c LRNg-95, apical view; Lymnaeidae indet.: d LRNg-71; e LRNg-81; f LRNg-81, apical view; Gyraulus sp.: g LRNg-110; h LRNg-30; i LRNg-127; j LRNg-101, protoconch; k LRNg-110, protoconch of g; Planorbarius sp.: 1 LRNg-141; m LRNg-142; n LRNg-141, protoconch of n; o LRNg-142, protoconch of m. Scale bars equal 1 mm (a-f, [-m), 0.5 mm (g-i, n, 0) and 0.1 mm (, k) Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest Fig. 5 Terrestrial gastropods from Lapsarna, Lesvos Island, Greece. Helicidae indet.: a LRNg-151; b LRNg-152; note that a was restored from two separate SEM images, resulting in some graphical issues: Limax sp.: c LRNg-183; d LRNg-181; Milax sp.: e LRNg-172; f LRNg-171. Scale bars equal 1 mm
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest Fig. 5 Terrestrial gastropods from Lapsarna, Lesvos Island, Greece. Helicidae indet.: a LRNg-151; b LRNg-152; note that a was restored from two separate SEM images, resulting in some graphical issues: Limax sp.: c LRNg-183; d LRNg-181; Milax sp.: e LRNg-172; f LRNg-171. Scale bars equal 1 mm
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest 2003 XVII 47-50, figured in Schulz-Mirbach anc Reichenbacher 2006: Fig. 26) and of Palaeoleuciscus sp. | from Mauensee (BSPG 2003 XVIII 84-89, figured in Jos et al. 2007: Fig. 5). The LRN lapilli are morphologically similar to the recent lapilli from Germany. The lapilli o: Palaeoleuciscus sp. from Le Locle display a much thicker mineralised area and dorsal lobe and a thin posterior part, whereas the LRN lapilli of a similar size show a flatter dorsal side and almost equally thick anterior and posterior lobes. Furthermore, the Le Locle specimens are less
Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest 2003 XVII 47-50, figured in Schulz-Mirbach anc Reichenbacher 2006: Fig. 26) and of Palaeoleuciscus sp. | from Mauensee (BSPG 2003 XVIII 84-89, figured in Jos et al. 2007: Fig. 5). The LRN lapilli are morphologically similar to the recent lapilli from Germany. The lapilli o: Palaeoleuciscus sp. from Le Locle display a much thicker mineralised area and dorsal lobe and a thin posterior part, whereas the LRN lapilli of a similar size show a flatter dorsal side and almost equally thick anterior and posterior lobes. Furthermore, the Le Locle specimens are less
Fig. 7 Freshwater fish remains from Lapsarna, Lesvos Island, Greece; pharyngeal teeth in occlusal view unless otherwise specified. Cyprinion sp.: a LRNt-55, left pharyngeal bone with Al, A2, B1 and B2; b LRNt-279, left Al; c LRNt-347, right A2; d LRNt-57, left B1; e LRNt-67, right B. Scale bar equals 1 mm Comments. Cyprinion Heckel, 1843 is a western Asian genus of small-sized barbin fishes that occupy rivers and tributaries (Alkan Uckun and Goékce 2015). According to Banarescu and Herzig-Straschil (1995), the distribution of the eight recent species comprises the northern (Orontes, Qweiq and Tigris—Euphrates drainage basins; C. macros- tomus species group) and the southern (southern Arabian Peninsula, southern and central Iran, Afghanistan, and Pakistan west of the Indus river; C. watsoni-microphthal- mus species group) parts of western Asia. Although it is the first time that fossil records of Cyprinion have been described, isolated pharyngeal teeth resembling the LRN material are known as aff. Cyprinion from early Middle Miocene sediments of western Anatolia (Dumlupinar, Calikli localities: reported as Varicorhinus/Capoeta by Bohme and Ilg 2003). According to BOhme et al. (2003), aff. Cyprinion characterizes period B (Late Burdigalian— Middle Serravallian, MN 4-MN 6, ~18—13.5 Ma) of the Anatolian freshwater fish evolution. The early Miocene Cyprinion from Lesvos might represent a new species, which, however, should only be named based on articu- lated skeletons. The genus Cyprinion is also the only barbel group in western Eurasia that has seven branched anal fin rays, instead of the five typical of nearly all barbins (Banarescu and Herzig-Straschil 1995). The Early Miocene (MN 3) species “Barbus” bohemicus Obrhelova, 1967 is described as having seven branched anal fin rays (Obrh- elova 1967), which makes a generic attribution within Cyprinion most plausible. Also, based on its short dorsal fin ray (7-9 branched fin rays), it strongly resembles the southwestern Asian species group CC. watsoni- microphthalmus.
Fig. 7 Freshwater fish remains from Lapsarna, Lesvos Island, Greece; pharyngeal teeth in occlusal view unless otherwise specified. Cyprinion sp.: a LRNt-55, left pharyngeal bone with Al, A2, B1 and B2; b LRNt-279, left Al; c LRNt-347, right A2; d LRNt-57, left B1; e LRNt-67, right B. Scale bar equals 1 mm Comments. Cyprinion Heckel, 1843 is a western Asian genus of small-sized barbin fishes that occupy rivers and tributaries (Alkan Uckun and Goékce 2015). According to Banarescu and Herzig-Straschil (1995), the distribution of the eight recent species comprises the northern (Orontes, Qweiq and Tigris—Euphrates drainage basins; C. macros- tomus species group) and the southern (southern Arabian Peninsula, southern and central Iran, Afghanistan, and Pakistan west of the Indus river; C. watsoni-microphthal- mus species group) parts of western Asia. Although it is the first time that fossil records of Cyprinion have been described, isolated pharyngeal teeth resembling the LRN material are known as aff. Cyprinion from early Middle Miocene sediments of western Anatolia (Dumlupinar, Calikli localities: reported as Varicorhinus/Capoeta by Bohme and Ilg 2003). According to BOhme et al. (2003), aff. Cyprinion characterizes period B (Late Burdigalian— Middle Serravallian, MN 4-MN 6, ~18—13.5 Ma) of the Anatolian freshwater fish evolution. The early Miocene Cyprinion from Lesvos might represent a new species, which, however, should only be named based on articu- lated skeletons. The genus Cyprinion is also the only barbel group in western Eurasia that has seven branched anal fin rays, instead of the five typical of nearly all barbins (Banarescu and Herzig-Straschil 1995). The Early Miocene (MN 3) species “Barbus” bohemicus Obrhelova, 1967 is described as having seven branched anal fin rays (Obrh- elova 1967), which makes a generic attribution within Cyprinion most plausible. Also, based on its short dorsal fin ray (7-9 branched fin rays), it strongly resembles the southwestern Asian species group CC. watsoni- microphthalmus.
Fig. 8 Amphibian remains from Lapsarna, Lesvos Island, Greece. ? Mioproteus sp.: a-b LRNa-2249, trunk vertebra (a ventral view; b dorsal view); ¢ LRNa-2248, trunk vertebra (lateral view); Anura indet.: d LRNa-2253, maxillary fragment (lingual view). Scale bar equals 1 mm Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
Fig. 8 Amphibian remains from Lapsarna, Lesvos Island, Greece. ? Mioproteus sp.: a-b LRNa-2249, trunk vertebra (a ventral view; b dorsal view); ¢ LRNa-2248, trunk vertebra (lateral view); Anura indet.: d LRNa-2253, maxillary fragment (lingual view). Scale bar equals 1 mm Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest
2275, trunk vertebra (ventro-lateral view); Serpentes indet.: LRNr- 2276, trunk vertebra (h ventral view; i anterior view); Squamata indet.: 7 LRNr-2281, osteoderm (dorsal view). Scale bar equals 1 mm Fig. 9 Reptilian remains from Lapsarna, Lesvos Island, Greece. Crocodylia indet.: a-d isolated teeth: a LRNr-2071 (lingual view); b LRNr-2057 (lingual view); c LRNr-2058 (buccal view); d LRNr- 2059 (occlusal view); Lacertidae indet.: LRNr-2255, dentary frag- ment (e lingual view; f dorsal view); ? Natricinae indet.: g LRNr-
2275, trunk vertebra (ventro-lateral view); Serpentes indet.: LRNr- 2276, trunk vertebra (h ventral view; i anterior view); Squamata indet.: 7 LRNr-2281, osteoderm (dorsal view). Scale bar equals 1 mm Fig. 9 Reptilian remains from Lapsarna, Lesvos Island, Greece. Crocodylia indet.: a-d isolated teeth: a LRNr-2071 (lingual view); b LRNr-2057 (lingual view); c LRNr-2058 (buccal view); d LRNr- 2059 (occlusal view); Lacertidae indet.: LRNr-2255, dentary frag- ment (e lingual view; f dorsal view); ? Natricinae indet.: g LRNr-
Table 2 Faunal list of the Early Miocene locality of Lapsarna, Lesvos (micromammalian taxa follow Vasileiadou and Zouros 2012, modified following the discovery of additional glirid material)
Table 2 Faunal list of the Early Miocene locality of Lapsarna, Lesvos (micromammalian taxa follow Vasileiadou and Zouros 2012, modified following the discovery of additional glirid material)
Related topics:
Systematics (Taxonomy)PaleontologyVertebrate PaleontologyInvertebrate PaleontologyFreshwater fish taxonomyPetrified Forest of Lesvos
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