Quercus mediterranea Unger
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publication ID |
https://doi.org/ 10.5281/zenodo.5374994 |
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persistent identifier |
https://treatment.plazi.org/id/F3021B7D-B158-3E1A-7742-36E91DF1FB91 |
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treatment provided by |
Marcus |
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scientific name |
Quercus mediterranea Unger |
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Quercus mediterranea Unger View in CoL ( Figs 2 View FIG H-K; 5J)
MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.6; 32.1.2.17; 32.1.2.79; 32.1.2.155; 32.1.3.2; fragmentary leaves nos. 32.1.2.24; 32.1.2.43; 32.1.2.147; 32.1.2.229; 32.1.2.233; 32.1.2.248; 32.1.3.4.
MATERIAL EXAMINED. — Complete leaf no. 32.1.3.1; DESCRIPTION
fragmentary leaves nos. 32.1.2.72; 32.1.2.158.
Five complete leaves very variable in shape, 3.3 to
6.9 cm long and 1.8 to 4.7 cm wide. Shape broadly
DESCRIPTION oval to ovate or obovate. Base acute or rounded, One almost complete and two incomplete leaves; rarely cordate and apex acute, obtuse or slightly broadly oval to elongate, slightly asymmetric in emarginate. Leaf margin simple serrate at least in shape; at least 10 cm long and 4.5 cm wide. Mar- the upper half of the lamina, occasionally entire or gin entire, base widely cuneate to rounded, apex undulate. Teeth widely spaced, rather small, more not preserved. Petiole c. 0.5 cm long. Primary vein or less sharply pointed. Venation craspedodromous, stout, slightly bent. Venation brochidodromous, midvein stout, in the upper part more slender and 8-10 pairs of curved secondary veins arising at 35- in some cases slightly sinuous. Secondary veins 60° from the midvein, regularly and widely spaced, distinct, in 5 to 9 pairs, arising at an angle of 30 joined together in a series of arches along the mar- to 80° from the primary vein, irregularly spaced, gin. Tertiary veins percurrent, straight or irregularly rarely forked, running straight or bent towards the branched. Leaf texture possibly leathery. leaf margin entering the teeth.Tertiary veins poorly visible, irregular, simple or branched. Leaf texture
REMARKS rather chartaceous or coriaceous.Also, 7 fragmentary Size, shape and venation pattern of these specimens leaves are assigned to this fossil species.
seems to fit the description of the fossil species Juglans acuminata . This taxon is present in many European REMARKS
Neogene deposits. Leaflets of Juglans sect. Juglans Quercus mediterranea is a characteristic common show entire margins (e.g., J. regia L.). Therefore, element in Neogene deposits especially in southern Juglans acuminata may be related to modern Jug- Europe. In Greece, this species is represented in lans regia that grows in mixed mesophytic forests numerous Miocene and Pliocene plant assemblages in southeast Europe and North Asia. However, of mainland and islands (Knobloch & Velitzelos Knobloch & Kvaček (1976) had studied the epi- 1987; Velitzelos & Gregor 1990). Leaf remains of dermal anatomy, and considered Juglans acuminata Quercus mediterranea from the Vrysses outcrops to be related to the North American species Juglans correspond much better to the records from Vegora rupestris Engelm. (late Miocene, N Greece) and Kymi (early Miocene Since several species of Pterocarya Kunth also Evia Island, central Greece) on the basis of grosshave leaves with entire margins, it can not be com- morphology (Velitzelos et al. 2002; Kvaček et al. pletely ruled out that the Vrysses leaves belong to 2002; Kvaček & Walter 1989) while leaves assigned this taxon. In the Makrilia plant assemblages from to Q. mediterranea from Makrilia (late Miocene,
Miocene leaf assemblage from Vrysses, western Crete eastern Crete, S Greece) seem to differ slightly. At dulate or simply serrate with small widely spaced the latter locality oaks are represented by a few teeth, serration starting in the lower third of the leaves, which are rather small. They hardly reach a lamina. Base cordate, apex acute maybe acuminate. length of 2.5 cm ( Sachse 1997). Venation craspedodromous. Midvein prominent Based on studies of leaf epidermis tissue from more or less curved, slightly zig-zag at the upper Lava (North Greece) that indicated an extremely part. Secondary veins more than 10 pairs, regularly thin cuticle, it was suggested that Q. mediterranea disposed, quite dense, diverging at angles of 40 to was a mesophytic deciduous oak (Kvaček & Walter 80° and running straight, parallel to each other, 1989). However newly obtained epidermal charac- across lamina, entering marginal teeth. Higher teristics show that in spite of the thin cuticle, the order venation poorly preserved. epidermis of either leaf side is sclerenchymatous underlain by a hypodermis (Kvaček et al. 2002). REMARKS Thus Q. mediterranea probably was a sclerophyllous Such leaves may be assigned to the genus Quercus , element able to survive short dry periods. representing probably an evergreen oak quite dif- Quercus mediterranea is perhaps ancestral to the ferent from Q. mediterranea especially in leaf size modern evergreen species Quercus ilex L. and Quer- and shape and secondary veins pattern. Leaf remains cus coccifera L., although both do not correspond from early Miocene plant assemblage of Kymi figured in all details of leaf anatomy and morphology to by Unger (1867: taf. VI, figs 23-27) and identified the fossils. Today these oaks are present through- as Quercus drymeja Unger by Velitzelos et al. 2002 out the Mediterranean basin and on coastal slopes (revised list of Kymi material), are very close in mortowards the Atlantic (South France) and Black Sea, phology to Vrysses specimens. However more and on relatively dry soils. better preserved material is necessary to elucidate According to Barbero et al. (1992), Q. ilex thrives the systematic relationships of these specimens. in a wide climatic range (from warm-temperate to subtropical and from medium humid to semi-arid) due to numerous morphological adaptations. In Family ULMACEAE the northern Mediterranean realm, Q. ilex forms woodland with several laurophyllus elements such Ulmaceae gen. et sp. indet. as Arbutus L., Hedera L., Ilex L. and Viburnum L., ( Fig. 3B View FIG ) while in south it thrives together with more xerothermic taxa such as Juniperus L., Phillyrea Tourn. MATERIAL EXAMINED. — Complete leaf no. 32.1.2.70. ex Adans., Genista , Ephedra L. and Buxus . In the eastern Mediteranean basin Q. ilex is replaced by DESCRIPTION Q. coccifera . One small, broadly elliptic leaf, about 1.8 cm long and 1 cm wide. Apex incomplete, probably acute to acuminate, base rounded, strongly asymmetrical.
Quercus sp. Leaf margin serrate at the upper half part of lamina
( Figs 3A View FIG ; 5Q View FIG ) with several wide, simple teeth. Venation craspedodromous. Midvein prominent, quite thick and
MATERIAL EXAMINED. — Fragmentary leaves nos. strongly curved. From the primary vein arise four 32.1.2.19; 32.1.2.95; 32.1.2.142.
pairs of secondaries at angles of 35 to 55°, rather strong, more or less curved, running into the tooth
DESCRIPTION apices. No other details of the venation preserved. Three leaf fragments, long petiolate (up to 1.5 cm). Complete leaf length estimated at 7 to 9 cm or REMARKS perhaps more, width about 2.5 to 3.2 cm. Shape Such a leaf shape (leaf margin, asymmetrical base, of lamina ovate to broadly elliptic, leaf margin un- laminar shape) suggests a relationship with the
Zidianakis G. et al.
family Ulmaceae . However this specimen lacks margin dentate with obtuse teeth. Petiole not prefurther diagnostic features for a determination to served. Venation actinodromous with three prigenus level. The Ulmaceae is a family of tropical mary veins. Midvein slightly bent, laterals curved and mainly temperate trees and shrubs, with 16 and forked. Four or five pairs of curved secondary genera (Heywood et al. 1993). Due to very small veins arising from the midvein at 30-55°, looping size (nanophyll) and simple dentate margin, the with the laterals and the adjacent secondaries near Vrysses specimen may represent a slightly xero- the margin.Tertiary veins oblique forming meshes morphic element. between secondaries.
REMARKS Family SALICACEAE Shape , teeth at the leaf margin and venation of Genus Salix L. these specimens are similar to lamina of the modern species Populus tremula . Leaf remains of this cf. Salix sp. poplar have been found in several European Neo- ( Fig. 3C View FIG ) gene deposits. Today Populus tremula trees grow in open mesophilous forests up to 1000 m of altitude
MATERIAL EXAMINED. — Fragmentary leaf no. in Europe, North Africa, Asia Minor and Siberia
32.1.2.39. ( Leroy & Roiron 1996).
DESCRIPTION
Basal half of a probably narrow elliptic, slightly Family BUXACEAE asymmetrical leaf (3.9 cm long and 1.7 cm wide); Genus Buxus L.
entire leaf possibly twice that long. Leaf base acute,
margin finely serrate with simple acuminate teeth, Buxus pliocenica Saporta & Marion regularly spaced. Midvein medium, thick, well ( Figs 3E View FIG ; 5M View FIG )
developed and curved. Higher order venation not visible. M ATERIAL EXAMINED. — Complete leaf no.
32.1.2.140. REMARKS
Shape and margin of this leaf show resemblance to DESCRIPTION those of extant Salix L. However , because the upper One complete leaf, narrowly elliptic, with short part of the leaf is missing, as well as higher order petiole, 3.6 cm long and 1.2 cm wide. Leaf margins venation, a definite identification is not possible. entire, apex acute, base cuneate. Primary vein thin and almost straight. Secondary veins fine, arising at an angle of 30 to 70°, closely spaced, parallel
Genus Populus L. to each other, branched once to many times in acute angles without higher order veins rejoining
Populus tremula L., foss. secondaries.
( Fig. 3D View FIG )
REMARKS MATERIAL EXAMINED. — Complete leaf no. 32.1.3.7; Buxus pliocenica fossils have been known in Mifragmentary leaf no. 32.1.2.160.
ocene and Pliocene deposits of Eurasia, mostly in the form of leaf impression and compression, rarely DESCRIPTION as pollen or fossil fruits. Saporta & Marion (1876) One complete and one fragmentary leaf. Lamina treat Buxus pliocenica as an ancestor of Buxus sembroadly ovate, 4.5 to 4.8 cm long and about 5.5 to pervirens L. However, Kvaček et al. (1982) consider 5.7 cm wide. Apex probably obtuse, base rounded, Buxus pliocenica closely allied to the B. sempervirens -
Miocene leaf assemblage from Vrysses, western Crete
Zidianakis G. et al.
B. colchica - B. hyrcana group. According to these DESCRIPTION authors this group replaced the large-leaved Buxus One complete, very well preserved leaflet; 3.3 cm lineage which became extinct because of the climatic long and 1.5 cm wide, with oval outline. Margin of deterioration during the late Miocene. Extant Buxus lamina entire, rounded at the apex and quite asymsempervirens is a typical element of sclerophyllous metrical at the base. Texture chartaceous, petiolule vegetation in Mediterranean basin while Buxus missing. Venation brochidodromous. Primary vein hyrcana Pojark. and Buxus colchica Pojark. occur in thin, running irregularly (zig-zag) to the apex of pure stands or in underwood in lowland deciduous leaflet. Five pairs of secondaries arising at an angle broad-leaved mesophilous forests in the Hyrcanian of 30 to 80° from the midvein, running more or region (between the southern shores of the Caspian less sinuous, joining each other by loops near the Sea and the Elburz Mountains) and in the Colchis margin. The lateral secondary at one leaflet side is (Western Georgia). prominent, extending to the upper third of the middle of the leaflet. Intersecondary veins two or three,
weakly developed. Higher order venation branching
Family ROSACEAE out and joining the secondaries to form a network of rather large, irregularly shaped areoles.
Rosaceae gen. et sp. indet.
( Figs 3F View FIG ; 5G, S View FIG ) REMARKS
This type of leaflets is related to the subfamily Cae- MATERIAL EXAMINED. — Complete leaf no. 32.1.2.41; salpinioideae and especially to the artificial genera fragmentary leaves nos. 32.1.2.8; 32.1.2.55. Cassiophyllum and Caesalpinites by the asymmetrical form, the size and the venation pattern. These DESCRIPTION genera occur in European deposits from the Oli- One complete and two fragmentary leaves with short gocene to Miocene. The determination of modern petiole. Leaf length 2.7 to 3.5 cm (may be more) equivalents is difficult because the leaflets of many and width 1.3 to 2 cm. Shape of lamina ovate or species have a similar morphology, as for instance oval, rounded at the base acute at the apex. Margin Scohtia humboltioides Oliv., Wagetia spicata Daez finely serrate with small, simple and sharp teeth. and some Cassia species (Shakryl 1992) . Primary vein well developed and straight. Higher order veins not visible.
Fabaceae type 2 REMARKS ( Fig. 3H View FIG ) The features observed in these fossils and especially leaf margin suggest an affinity to the family MATERIAL EXAMINED. — Almost complete leaflet no.
32.1.2.78. Rosaceae . Today this large family comprises about 3300 species. It is cosmopolitan with maximum diversity in the northern temperate area (Heywood DESCRIPTION et al. 1993). One leaflet 3.8 cm long (complete about 4 cm) and
1.5 cm wide, narrow elliptic to lanceolate, slightly asymmetrical,entire-margined.Apex of lamina acute,
Family FABACEAE base obtuse or rounded, asymmetrical (part of the base is missing). Primary vein strong, S-like curved.
Fabaceae type 1 Higher order venation fine, hardly visible.
( Fig. 3G View FIG )
REMARKS MATERIAL EXAMINED. — Complete leaflet no. Entire margins and asymmetry of this specimen 32.1.2.161. are features that suggest a relationship with the
Miocene leaf assemblage from Vrysses, western Crete
Fabaceae . Unfortunately, poor preservation of the Weber. The name Acer pseudomonspessulanum is leaflet does not allow a more precise determination. used as the Neogene ancestors of the modern spe- Fabaceae is a very large plant family (approximately cies of the series Monspessulana Pojark., especially 17000 species) with great variety of habitat and a A. monspessulanum and A. sempervirens and their cosmopolitan distribution in tropical, subtropical intermediate forms ( Leroy & Roiron 1996). Leaf and temperate zones (Heywood et al. 1993). The and fruit remains of Acer pseudomonspessulanum are small-sized leaflet of Vrysses may belong to a more known from several Neogene deposits in Europe. or less xeric element. Today Acer monspessulanum extends from central,
south Europe and northwest Africa to north Iran
(south shores of the Caspian Sea), while A. sem-
Family ACERACEAE pervirens occurs at the east Mediterranean basin.
Genus Acer L. Acer monspessulanum is a deciduous element while
A. sempervirens is considered as deciduous or semi -
Acer pseudomonspessulanum Unger evergreen element. Both of these maples are trees
( Figs 3 View FIG I-L; 5L, P, R) or shrubs and common elements of more or less xerothermic mixed evergreen and deciduous forests. MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.46; Today Acer sempervirens grows on Crete at moun- 32.1.2.61; 32.1.2.67; 32.1.3.8; fragmentary leaves nos. tain slopes between 300 and 1700 m (Turland et 32.1.2.28; 32.1.2.29; 32.1.2.31; 32.1.2.37; 32.1.2.40; 32.1.2.45; 32.1.2.58; 32.1.2.62; 32.1.2.91; 32.1.2.244; al. 1993). 32.1.3.9; 32.1.3.10.
DESCRIPTION Acer sp. Four complete and 12 incomplete leaves, are assigned ( Figs 3M View FIG ; 5K View FIG ) to this fossil species. Lamina deeply trilobate, 2.1 to 4 cm (or somewhat more) long and 3 to 6.3 cm MATERIAL EXAMINED. — Almost complete leaves nos.
32.1.2.12; 32.1.2.56;
fragmentary leaves nos. 32.1.2.14;
wide, with short petiole. Leaf texture coriaceous. 32.1.2.49; 32.1.2.75.
Base of lamina rounded or cordate, margin entire. The median lobe more or less broader than the lat- DESCRIPTION eral lobes. Apices of lobes acute to obtuse. Venation Two almost complete leaves and 3 fragments,5-lobed, actinodromous with three primary veins, straight 3.1 cm to 3.9 cm (possibly more) long and 4.4 to to slightly bent. Secondary veins fine, dense, aris- 5.8 cm wide. Margins entire with acuminate apex. ing at an angle of 60 to 80° from the primaries, Leaf base cordate, venation actinodromous with branching near the margin. Higher order veins five primary veins. Lateral primaries arising under poorly preserved. angles of 40 to 55° from the midvein, while basal primaries arise under angles of 80 to 110°. Five to
REMARKS seven pairs of hardly visible secondary veins, arise Most of these fossil leaves (the larger specimens) are at 30 to 55° from primary veins and run in curves most likely related to Acer monspessulanum L. but towards the leaf margin. some of them, with sharper sinuses and less broad lobes with acuminate apices which stretch upwards, REMARKS are related to Acer sempervirens L. In fact, remains of Some of these specimens are comparable to Acer inte- Acer pseudomonspessulanum from Vrysses do not differ gerrinum, a common element of European Neogene in gross-morphology from the records in Makrilia, deposits. In Greece, A. integerrinum is represented from where this leaf type was described as Acer series in the late Miocene fossiliferous deposits of Licudi, Monspessulana Pojark. Material from Vegora that central Greece (Velitzelos & Gregor 1990). is also similar in its variability has been determined Acer laetum C.A.Mey. and Acer pictum Thunb. as A. pseudomonspessulanum and A. integrilobum are considered to be the possible analogue among
Zidianakis G. et al.
extant maples on the basis of their gross-morphology Dicotylophyllum sp. 1 ( Leroy & Roiron 1996). Acer laetum thrives from ( Fig. 4A View FIG ) the Caucasus to the Himalayas and the mountains of central China while A. pictum grows on the slopes MATERIAL EXAMINED. — Complete leaf no. 32.1.2.1. of northern China and Japan.
DESCRIPTION
One entire leaf, reniform, 2.4 cm long and about
Family RHAMNACEAE 3.2 cm wide, stalked, entire-margined. Leaf texture
Genus Ziziphus Mill. evidently fine.Venation actinodromous with seven primary veins. Midvein straight, stronger than the
cf. Ziziphus ziziphoides (Unger) Weyland lateral primaries. All primary veins forked before
( Fig. 3 View FIG N-P) reaching leaf margin. Secondary veins on the me-
dial primary vein widely and irregularly spaced
MATERIAL EXAMINED. — Almost complete leaves nos. in four pairs, also forked. Higher order venation 32.1.2.3; 32.1.2.5; 32.1.2.22; 32.1.2.180; fragmen- not visible. tary leaves nos. 32.1.2.10; 32.1.2.32; 32.1.2.106; 32.1.2.209.
REMARKS
DESCRIPTION A very interesting leaf form. In morphology (ve- Four almost complete leaves and four fragments. nation, margin, shape), it is comparable with the Lamina broad oval to ovate 3.2-5.8 cm long (maybe foliage of modern Cercis silliquastrum L. ( Fabaceae ), more) and 1.9-4.1 cm wide, simply dentate at the differing mainly by having smaller leaf size than upper two thirds of the leaf margin. Apex acute to living element. Cercis silliquastrum is a Mediterobtuse, base partly asymmetric. Venation acrodro- ranean, deciduous tree. mous with three stout primary veins running in convergent arches toward the leaf apex.Higher order veins very fine, hardly visible, vertical to midvein. Dicotylophyllum sp. 2
( Figs 4B View FIG ; 5H View FIG )
REMARKS The specimens described above probably belong to MATERIAL EXAMINED. — Almost complete leaf no. the species Ziziphus ziziphoides , a common element 32.1.2.92. in the European Paleogene and Neogene. The genus Ziziphus consists of about 100 species of decidu- DESCRIPTION ous or evergreen trees and shrubs distributed in One almost complete (only a small part of its apex the tropical and subtropical regions of the world is missing) leaf, narrow oblanceolate 6.2 cm long (Johnston 1963). This leaf form may be related to (complete leaf somewhat more) and 2.2 cm wide. a non-sclerophyllous species of Ziziphus not iden- Lamina with entire margin, gradually narrowing tical with recent species of the Mediterranean area towards the base. Primary vein straight, very strong and the Near East. The nearest extant relatives of and thick. Leaf texture coriaceous. Secondary veins Ziziphus ziziphoides considered to be the decidu- fine, hardly visible, arising at 30 to 50° from the ous Z. sinensis and Z. glabrata both thrive in Asia midvein. (Givulescu 1962). However, this leaf type may also represent a completely different family (see Sachse REMARKS 1997; Dicotylophyllum type 3). This leaf with poorly preserved venation cannot be reliably assigned to any specific genus. The general shape of this specimen suggests that it
Family incertae sedis represented a broad-leaved humid or mesic ele-
Genus Dicotylophyllum Saporta ment.
Miocene leaf assemblage from Vrysses, western Crete
Zidianakis G. et al.
Dicotylophyllum sp. 3 broadly lanceolate to obovate, apex acute to obtuse
( Fig. 4C View FIG ) and base obtuse to slightly cordate. Margin slightly wavy or dentate with small simple teeth. Venation MATERIAL EXAMINED. — Complete leaf no. 32.1.2.16. semicraspedodromous. Primary vein prominent,
somewhat straight. Secondary veins thin, densely DESCRIPTION spaced at angles of 45 to 80°, running almost straight, One complete leaf (only petiole missing), narrowly parallel to each other across the lamina, dichotomised elliptic, slightly asymmetrical leaf; 5 cm long and near the margin before entering teeth.Tertiary veins 1.2 cm wide. Apex rounded, base obtuse, somewhat percurrent or simply forked forming irregular meshes asymmetrical, leaf margin entire.Primary vein stout, with quaternary veins. Also two fragmentary leaves S-like curved. Secondary veins fine, rather dense, are assigned to this form.
hardly visible.
REMARKS
REMARKS This fossil leaf taxon is similar in gross-morphol- It has not been possible to determine the affinity ogy to numerous genera in different families such of this leaf due to poor preservation. as Rhamnaceae and Fagaceae .
Dicotylophyllum sp. 4 ( Figs 4D View FIG ; 5O View FIG ) Dicotylophyllum sp. 6 ( Figs 4F View FIG ; 5I View FIG )
MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.27; MATERIAL EXAMINED. — Fragmentary leaves nos. 32.1.2.9; 32.1.2.33; 32.1.2.60; 32.1.2.144. 32.1.2.107.
DESCRIPTION DESCRIPTION Four complete leaves, small, long petiolate, oblan- Two fragmentary leaves with broad obovate outceolate to obovate leaves, 4 to 5.5 cm long and line, 5 to 5.7 cm long (perhaps more) and 2.2 to 1 to 1.5 cm wide. Leaf apex obtuse or rounded, 3.2 cm wide. Petiole rather slender, up to 1.2 cm base attenuate and leaf margin entire. Texture co- long. Base of lamina heart-shaped, apex absent, riaceous. Primary vein strong, more or less curved. with undulate or finely, simply dentate margin. No other details of the venation visible. Midvein fine, hardly visible, straight or slightly bend. Higher order venation invisible. Leaf texture REMARKS coriaceous. The affinity of these specimens is unknown and much better preserved material is necessary for a REMARKS reliable identification. The poor preservation makes it difficult to clarify the systematic relationship of these specimens.
( Figs 4E View FIG ; 5B, N View FIG ) Dicotylophyllum sp. 7
( Fig. 5T View FIG )
MATERIAL EXAMINED. — Complete leaves no. 32.1.3.4;
32.1.3.6; fragmentary leaves nos. 32.1.2.34; 32.1.2.35. MATERIAL EXAMINED. — Complete leaf no. 32.1.2.44.
DESCRIPTION DESCRIPTION
Two almost complete petiolate leaves. Length of the One complete, petiolate, slightly asymmetrical, lamina 2.5 to 3.9 cm and width 1.7 to 1.9 cm. Leaf broadly oval leaf with entire margin. Leaf length
Miocene leaf assemblage from Vrysses, western Crete
5.9 cm, width 2.2 cm. Leaf base obtuse, apex round- veins at least seven pairs, hardly visible, very deli- ed. Venation probably semicraspedodromous. Pri- cate, rather widely spaced, arising at angles of 55 mary vein quite strong, S-like curved. Secondary to 80° from the midvein, running nearly straight
Zidianakis G. et al.
to somewhat bent towards the margin. Probably 3.6 long and 3 cm wide). Leaf apex not preserved, secondaries forking near the margin. Further vena- base obtuse, margin hardly visible, undulate. Leaf tion pattern not preserved. texture coriaceous. Venation craspedodromous, primary vein stout, S-like curved. Secondaries at REMARKS least seven pairs, strong, parallel, regularly spaced, The systematic affinity of this leaf taxon remains more or less straight, directed towards the margin. unclear. Higher order veins not visible.
REMARKS Dicotylophyllum sp. 8 Such leaves may be related to the genus Fagus L., ( Fig. 4G View FIG ) but such affinity cannot be proved. More and much better preserved material is necessary to elucidate MATERIAL EXAMINED. — Almost complete leaf no. the position of this specimen. 32.1.2.223.
DESCRIPTION Dicotylophyllum sp. 10 One leaf almost complete (only a small part of apex ( Figs 4I View FIG ; 5C View FIG ) missing), broadly ovate, quite asymmetrical, 5.7 cm long (complete leaf about 6 cm) and 4.3 cm wide. MATERIAL EXAMINED. — Complete leaf no. 32.1.2.93; fragmentary leaves nos. 32.1.2.4; 32.1.2.94. Apex of lamina acute or acuminate and base heartshaped. Margin simply dentate with moderately large and acute teeth, irregularly spaced. Venation DESCRIPTION actinodromous with five primary veins.Primary vein One complete and two fragmentary leaves, petirather stout, S-like. Basal primaries quite weak, aris- olate, with elliptic outline. Their length is 5 to ing from the midvein at angles of 100 to 120° while 6.2 cm (maybe somewhat more) and their width lateral primaries stronger, arising at angles of 60 to 1.7 to 1.9 cm. Leaf margin entire, acute at the apex 80°. Four pairs of curved secondary veins arising from and cuneate at the base. Primary vein prominent the midvein, widely spaced, dichotomised near the and straight. Secondary veins fine, hardly visible. margin.Tertiary veins usually simple forked forming Higher order venation not preserved. polygonal meshes with quaternary veins. REMARKS REMARKS The systematic affinity of these three specimens Crucial characters are missing and thus it is im- is unclear. possible to determine the systematic affinity of this specimen. Dicotylophyllum sp. 11 ( Figs 4J View FIG ; 5E View FIG ) Dicotylophyllum sp. 9 ( Fig. 4H View FIG ) MATERIAL EXAMINED. — Complete leaf no. 32.1.2.159; fragmentary leaves nos. 32.1.2.98; 32.1.2.131.
MATERIAL EXAMINED. — Fragmentary leaf no. DESCRIPTION
32.1.2.30. One complete leaf, the largest in the Vrysses assemblage, 17 cm long and 4 cm wide (size of lamina
DESCRIPTION more than 4.500 mm 2). Also two fragmentary leaves,
One fragmentary leaf, broadly elliptic or obovate, smaller, up to 12 cm long and 2 to 2.5 cm wide.
2.6 cm long and 2.5 cm wide (entire leaf about Leaf apex obtuse or acute, base cuneate, entire-
Miocene leaf assemblage from Vrysses, western Crete margined, with broad elliptic outline. Primary Cyperaceae . The Poaceae and Cyperaceae are two vein characteristically thick and strong, straight large families of monocotyledons, cosmopolitan. or slightly bent. Higher order venation not visible Only few ecological plant formations lack Poaceae , probably duo to thick coriaceous texture. while many formations are dominated by them (Heywood et al. 1993).
REMARKS
Such leaf specimens are usually considered to belong to magnoliid families, such as Magnoliaceae Poaceae / Cyperaceae type 2
or Lauraceae . However a relationship with other ( Fig. 4L View FIG )
families cannot be excluded.
MATERIAL EXAMINED. — Fragmentary leaves nos. 32.1.2.54; 32.1.2.198.
( Fig. 5A View FIG ) DESCRIPTION
Two fragments of broad linear leaves, wide, more MATERIAL EXAMINED. — Complete leaf no. 32.1.2.42. than 2 cm wide, with parallel venation.
DESCRIPTION REMARKS A small complete leaf, short petiolate, 3.2 cm long These specimens probably also represent leaves of and 1 cm wide. Lamina narrow obovate, with apex Poaceae or Cyperaceae . rounded, base acute and entire margin. Primary vein quite strong and thick, S-like curved. Secondary veins poorly preserved, arise from the midvein at FLORISTIC COMPOSITION angles of 30 to 50°. Leaf texture chartaceous.
The Vrysses plant assemblage is documented by REMARKS foliage. More than 260 specimens of leaf mega- This leaf lacks diagnostic features. fossils, in many cases very fragmentary, were examined. A total of 31 taxa of vascular plants have been described based on their macro morpho-
Class MONOCOTYLEDONAE logical features ( Table 1). Nineteen of them were
Families POACEAE / CYPERACEAE relatively well preserved and identified at least to family level.Two genera of conifers and 29 taxa of
Poaceae / Cyperaceae type 1 angiosperms, shrubs, trees and, rarely herbs have
( Fig. 4K View FIG ) been documented.
Conifers are represented in the assemblage by MATERIAL EXAMINED. — Fragmentary leaves nos. Tetraclinis and Pinus shoots and leaves. Fertile 32.1.2.57; 32.1.2.63; 32.1.2.151; 32.1.2.169; 32.1.2.186; parts, such as cones, cone scales or seeds were not 32.1.2.189; 32.1.2.190; 32.1.2.191; 32.1.2.192; 32.1.2.199; 32.1.2.203; 32.1.2.208; 32.1.2.224; found. 32.1.2.228; 32.1.2.236; 32.1.2.245. Angiosperm taxa make up approximately 95%
of the total flora. Among the angiosperms, Acer- DESCRIPTION aceae and Fagaceae are most common. Fagaceae is Sixteen fragments of narrow strap-like leaves about represented by the genus Quercus , mainly by the 1 to 1.5 cm wide, that bear parallel venation. probably evergreen oak Quercus mediterranea , which predominates and Quercus sp. considered also to REMARKS belong to evergreen oaks. Two leaf-types belong to These fossil leaves are similar in gross-morphol- Aceraceae ; a small 3-lobed leaf identified as Acer ogy to numerous genera mainly in Poaceae and pseudomonspessulanum , considered to be more or
Zidianakis G. et al.
Family Genus Species/ Leaves/ Leaf mar- Maximum leaf D, L, S a Dispersal type series needles shoots gin entire size class mechanism
GYMNOSPERMAE
Pinaceae Pinus sp. 17 needles - - - wind
Cupressaceae Tetraclinis sp. 4 twig fragments - - - wind
ANGIOSPERMAE
Dicotyledonae
Lauraceae Daphnogene polymorpha 2 yes microphyll L endozoochory
Hamamel-?“ Parrotia ” pristina 1 no microphyll D autochory idaceae
Juglandaceae ? Juglans acuminata 3 leaflets yes notophyll D dyschory
Fagaceae Quercus mediterranea 12 no microphyll S dyschory
Quercus sp. 3 no microphyll S dyschory
Ulmaceae gen. indet. - 1 no nanophyll D/S wind
Salicaceae ? Salix sp. 1 no microphyll D wind
Populus tremula , foss. 2 no microphyll D wind
Buxaceae Buxus pliocenica 1 yes microphyll L/S autochory/ myrmecochory
Rosaceae gen. indet. - 3 no microphyll D wind/ exozoo-/endozoochory
Fabaceae gen. indet. type 1 1 leaflet yes microphyll D/S -
gen. indet. type 2 1 leaflet yes microphyll D/S -
Aceraceae Acer pseudomons- 16 no microphyll D/S wind pessulanum
Acer sp. 5 no microphyll D wind
Rhamnaceae ? Ziziphus ziziphoides 8 no microphyll D endozoochory
Family incertae Dicotylophyllum sp. 1 1 yes microphyll D/L/S -
sedis Dicotylophyllum sp. 2 1 yes microphyll L -
Dicotylophyllum sp. 3 1 yes microphyll S -
Dicotylophyllum sp. 4 4 yes microphyll L/S -
Dicotylophyllum sp. 5 4 no microphyll D/S -
Dicotylophyllum sp. 6 2 no microphyll D -
Dicotylophyllum sp. 7 1 yes microphyll L/S -
Dicotylophyllum sp. 8 1 no microphyll D -
Dicotylophyllum sp. 9 1 no microphyll D -
Dicotylophyllum sp. 10 3 yes microphyll L/S -
Dicotylophyllum sp. 11 3 yes notophyll L -
Dicotylophyllum sp. 12 1 yes nanophyll S -
Monocotyledonae
Poaceae / gen. indet. type 1 16 - - - wind/
Cyperaceae nautochory/ exozoo-/ endozoochory gen. indet. type 2 2 - - - wind/ nautochory/ exozoo-/ endozoochory less xeromorphic and a 5-lobed leaf represents most naceae may be represented by irregularly dentate, likely a mesophytic deciduous species. 3-veined leaves that are close in appearance to the Fabaceae are represented by a few leaflets that have Neogene species Ziziphus ziziphoides . Leaves of been subdivided into two morphotypes ( Fabaceae Lauraceae seem to have been present in the assemtype 1 and type 2). Juglandaceae seem to be also blage as well by the narrow leaf type of Daphnogene present. The leaflets of the Vrysses plant assemblage polymorpha . Imprints of Populus tremula , foss., Salix may belong to Juglans acuminata . The family Rham- sp. and possibly “ Parrotia ” pristina occur rarely.One
Miocene leaf assemblage from Vrysses, western Crete leaf of Buxus sp. has been identified. Rosaceae and from different habitats and the high percentage of Ulmaceae may have been present as well. Mono- small-sized leaves (Ferguson 1985; Spicer 1991). cots are represented by numerous specimens of the More than 85% of the taxa identified in Vrysses families Poaceae and Cyperaceae . However many outcrops are classified as microphyll (225-2.025 leaf impressions of the Vrysses plant assemblage mm2), while 7.4% as nanophyll (25-225 mm 2) have doubtful affinities. They are considered to ( Table 1). The allochthonous nature of the assembelong to 12 taxa of angiosperms, and classified as blage is also reflected by the absence of different Dicotylophyllum . organs, such as seeds and fruits of the same taxon Considering the number of specimens, four woody in the assemblage and by the high percentage of taxa are most common in the assemblage: Pinus sp. , heavily fragmented leaves that reach 64% of the Acer pseudomonspessulanum , Quercus mediterranea examined specimens. and Ziziphus ziziphoides ( Table 1). Tetraclinis sp. Plant debris have probably been transported by and Acer 5-lobed are found also more frequently. river floods into their offshore burial site. Alterna- More than one species may have existed among tively, plant remains may have drifted on the sea Aceraceae , Fagaceae Salicaceae and Fabaceae (all surface and reached the locality of sedimentation two taxa). after having been blown by strong winds from high The majority of the genera identified at Vrysses coastal cliffs. are related to elements now growing in the North- According to its composition, the Vrysses plant ern Hemisphere. The modern living relatives of assemblage is a heterogeneous mixture of higher Buxus pliocenica , Populus tremula , foss., Acer pseudo- plants with different ecology. Considering the monspessulanum and Juglans acuminata thrives in specific features of the modern associations (Wolfe Eurasia while Daphnogene polymorpha has its clos- 1979; Klötzli 1988; Song 1983, 1988) that can est counterparts in Asia and America and Ziziphus be related to the Vrysses fossil leaves, the outline ziziphoides in Asia. A few taxa have affinities to of a regional vegetation reconstruction has been living Mediterranean species such as Tetraclinis and developed, that is characterized by a floristic dif- Quercus mediterranea while extant Parrotia is a relic ferentiation based on water efficiency and altitude found in Iran (Zohary 1947; Wang 1961; Zohary (Gastaldo & Ferguson 1998). & Orshan 1966; Meusel et al. 1965, 1978; Strahler – a) Riparian paleocoenoses composed of representa- & Strahler 1992; Achhal et al. 1980; Quezel 1988; tives of the wood elements Salix sp. and perhaps Barbero et al. 1992; Turland et al. 1993; Arabatzis Populus tremula and Juglandaceae , as well as Poaceae 1998 , 2001). The core of the nearest living relatives and Cyperaceae herbs. These elements were forming of the fossil taxa grows today in the temperate to a belt of a riparian forest along the banks of inflowsubtropical zone, whereas only a few taxa extend ing streams. Subtropical, partly evergreen elements, to the tropical zone ( Lauraceae ) (Walter & Lieth such as Daphnogene polymorpha may have played 1967; Browics 1982; Klötzli 1988). also a certain role in these lowland forests. – b) Sclerophyll forest paleocoenoses consisted in general of microphyllous, broad-leaved taxa TAPHONOMY AND VEGETATION (such as Quercus mediterranea , Quercus sp. , possi- ANALYSIS bly Buxus pliocenica , Acer pseudomonspessulanum , Ulmaceae and several Fabaceae ) and conifers (Pi- The Vrysses Formation was most likely deposited in a nus, possibly Tetraclinis ) forming woodland at dry shallow, semi-enclosed marine embayment, bordered lowland areas. by islands and shoals ( Meulenkamp et al. 1979). – c) Mixed-Mesophytic forest coenoses. Sclerophyl- The fossil assemblage contains autochthonous and a lous evergreen forests of lowlands were gradually large proportion of allochthonous elements. This is replaced by mixed mesophytic forests of higher elevaconfirmed mainly by the composition of the taphoc- tions along an altitude gradient. The canopy of these oenosis that consists of a mixture of plant remains forests was dominated by deciduous representatives
Zidianakis G. et al.
)
C
°
(30 of sclerophyllous elements (about 15-52%) reduces temperature 20
the – modern b accuracy) Co-existence taxon of analog this approach correlation of the species. Table. recognized 2 shows the in annual 10 temperature the Vrysses plant requirements assemblage. A and range their of mean 15-16 annual °C for Mean 0 mean Several annual of the temperature Vrysses taxa is also seem estimated to have. their 0 20 40 60 80 natural habitat today in the Euxino-Hyrcanian Prov-
Percentage of entire-margined taxa ince. These taxa comprise Parrotia , Buxus , Juglans
FIG. 6. — Correlation of percentage of species with entire-mar-
and Acer . The average annual temperature of the gined leaves in the humid and mesic broad-leaved forests of Hyrcanian region varies from 15°C in the west to eastern Asia with mean annual temperature, after Wolfe (1979). 17.5°C in the east. The temperature of the warmest ---, percentage of entire-margined taxa in the Vrysses (western month ranges between 28-35°C while that of the
Crete) plant assemblage.
coldest month ranges from 1.5 to 4°C.
Rainfall varies substantially from 530 mm in of Acer 5-lobed, Populus tremula , Ziziphus ziziphoides , the east to 1350 mm in the west. In most areas Juglandaceae and probably Acer pseudomonspessula- of this region precipitation is all year round with num and Ulmaceae . In the stratum of lower trees maxima in spring, late fall and winter, the eastern and shrubs a mixture of evergreen ( Buxus pliocenica parts experience three months of summer drought and Daphnogene polymorpha ), as well as deciduous (see www.rifr-ac.ir/index_eng.htm). If we take this elements (“ Parrotia ” pristina, Rosaceae ) and conifers region as model for the Neogene climate, a range of ( Tetraclinis sp. ) seem to have existed. 15-17.5°C for mean annual temperature (MAT), It is also interesting to consider dispersal mecha- and more than 800 mm for mean annual precipitanisms of the recorded taxa. About 23% of the tion (MAP) is estimated.
identified plant taxa were dispersed either endo- or The average small leaf size of the assemblage exozoochorously by birds or mammals (see Table 1) (microphyllous flora) may be not only caused by that implies also a diverse paleofauna. taphonomic processes but also by climatic factors,
such as periodic drought. The relatively high proportion of sclerophyllous xeromorphic elements PALEOCLIMATE RECONSTRUCTION in the plant assemblage may confirm the latter hypothesis.
Analyses of Neogene plant assemblages are in gen- On the basis of physiognomic features, leaf reeral a very useful tool to extract proxy data for mains of Vrysses are divided in three main catpaleoclimatic reconstructions. We tried to apply egories: 1) a broad-leaved deciduous component two different methods in order to estimate climatic which predominates with approximately 37-59%; parameters of the broader Vrysses area during the 2) broad-leaved evergreens (laurophyllous taxa) reach late Miocene: the leaf margin analysis and the co- about 11-30%; and 3) the sclerophyllous compoexistence approach. However because of the lim- nent with the relatively high proportion of 15-52% ited number of taxa and the unresolved taxonomic ( Table 1). The broad-leaved deciduous elements problems due to poor preservation of the material are characteristic for humid, temperate climatic only rough estimations were possible. conditions, while the broad-leaved evergreen taxa – a) Leaf margin analysis. The proportion of taxa for humid, subtropical and sclerophyllous taxa for with entire-margined species in the Vrysses plant sub-humid, subtropical conditions (Berger 1954; assemblage reaches 48% ( Table 1), suggesting a Kovar-Eder et al. 2006).
mean annual temperature of 15-16°C (Zidianakis Thus, the Vrysses plant assemblage may indicate et al. 2004) (Fig. 6). The relatively high percentage a warm-temperate, humid climate for the late
Miocene leaf assemblage from Vrysses, western Crete
Taxa NLR MAT of NLR (°C)
Tetraclinis sp. Tetraclinis articulata 15.60-19.90
“ Parrotia ” pristina Parrotia persica 13.60-15.80
Juglans acuminata Juglans regia 10.10-17.30
Quercus mediterranea Quercus ilex , Quercus coccifera 11.70-19.50
Populus tremula , foss. Populus tremula 1.60-19.00
Buxus pliocenica Buxus sempervirens 9.70-17.30
Acer pseudomonspessulamum Acer pseudomonspessulanum , 9.60-20.50 Acer sempervirens
Ziziphus ziziphoides Ziziphus sinensis 7.30-27.60
Miocene. The existence of a dry period during the flora was determined using nannoplankton and year is probable, although this seasonality was clearly dinoflagellate cysts as late Miocene (7.7-8.6 Ma; not yet as pronounced as today. Using the climatic Sachse & Mohr 1996) and is thus slightly older than classification of Köppen (1931), the Vrysses paleo- the Vrysses flora. As it was expected, the Makrilia climate is most likely ranging between Cfa (warm assemblage shares many arboreal taxa with the temperate with warm, humid summers) and Csa Vrysses flora. More than 70% of the taxa identified (warm temperate with warm, arid summers) cli- in Vrysses exist also in Makrilia. Subtropical lauromates. The climatic change from pre-late Miocene phyllous elements, such as Daphnogene polymorpha , summer wet climates to summer dry climates was deciduous elements, such as Acer 5-lobed, Populus , possibly caused by the successive westward retreat of Salix and Ziziphus , conifers, such as Tetraclinis , Pinus the Paratethys between the Oligocene and the mid- and xeromorphic taxa, such as Quercus mediterranea , dle Miocene (Popov et al. 2004) and the changing Acer pseudomonspessulanum and Buxus pliocenica are topography of eastern Asia ( Sachse 1997). common in both floras. However, the Makrilia assemblage is more diverse than the one from Vrysses.
Considering the composition of the Makrilia flora
DISCUSSION it clearly contains relatively more deciduous taxa than the Vrysses plant assemblage. However, it is
The discovery of a new Miocene leaf assemblage in not clear whether this is a real signal or the result of the eastern Mediterranean realm offers the opportu- taphonomic processes or in the nearby hinterland nity for a better understanding of the floristic and of the Vrysses area floras of high elevations might climatic development of this region. Neogene floras have simply not existed.
occur in central and northern Greece, as well as in The late Miocene floras of continental Greece several Aegean islands including Crete (Knobloch & (Vegora, Likudi, Prosilion, Lava) and other islands Velitzelos 1987; Velitzelos & Gregor 1990; Sachse (Chios, Kerkyra) consist mainly of conifers and & Mohr 1996, Velitzelos et al. 2001). If the Vrysses deciduous broad-leaved elements, such as Ginkgo , plant assemblage is compared with the known Greek Taxodiaceae , Pinaceae , Sassafras , Alnus , Fagus , deplant assemblages on the basis of physiognomy and ciduous Quercus , Populus , Zelkova , Ulmus and floristic composition, the closest match seems to Acer (Velitzelos et al. 2001) . Obviously, not only be with the late Miocene assemblage of Makrilia, on Crete but in entire Greece, the exotic elements eastern Crete ( Sachse 1997, 2004). of the early Miocene were gradually replaced by The plant macro-remains of Makrilia occur in deciduous broad-leaved trees and shrubs, forming semi-pelagic silty layers and represent a plant assem- forests that resembled modern mixed mesophytic blage of which 94% of the leaf taxa are microphyll woods of Eastern Asia, the Colchis and the Caspian or nanophyll ( Sachse 1997). The age of the Makrilia area. This change most likely indicates a decrease
Zidianakis G. et al.
of mean annual temperature during the Miocene pliocenica and Daphnogene polymorpha ), deciduous in Greece, as it seems to have happened in all the (“ Parrotia ” pristina, Rosaceae ) and conifers (Tetraother Tethys and Paratethys areas (Gregor 1990; clinis sp.) occurred in the lower strata.
Kovar-Eder 1987). The floristic composition and physiognomy of the Furthermore, the relative increase of xeromorphic assemblage, as well as a comparison with published taxa, such as evergreen oaks ( Quercus drymeja Unger, Miocene Greek assemblages indicate a warm-tem- Q. mediterranea and Q. sosnowskyi Kolakovskii ), as perate, humid climate for this period, possibly with well as Acer pseudomonspessulanum and Chamaerops not very pronounced dry periods for the end of the humilis (Velitzelos & Schneider 1978; Velitzelos Tortonian in the eastern Mediterranean area.
et al. 2001; Kvaček et al. 2002) may indicate the beginning of weak drought periods during the last
Miocene in Greece and the onset of the Csa climate Acknowledgements type sensu Köppen (1931) (warm temperate with We would like to thank Prof. M. Mylonas, director arid summers). The existence of summer drought of the Natural History Museum of Crete, for his may be more obvious in the plant assemblage of helpful suggestions, and Dr. A. Nikolaidis, direc- Vrysses indicating already the beginning Messinian tor of the Mediterranean Agronomic Institute of salinity crisis (Hsu et al. 1973). Chania, for providing samples from this institution. For the free access to the Berlin Herbarium of the Botanic Garden for comparative studies we thank CONCLUSIONS Dr. R. Vogt. This work would not have been possible without the help of the staff of the Natural The upper Tortonian (6-7.5 Ma) Vrysses plant as- History Museum of Crete, A. Kagiambaki, M. Trisemblage consists of a mixture of leaf taxa that had kali, G. Iliopoulos, E. Nikolakakis and P. Lyberakis. most likely originally grown in riparian to possibly Also, comments of Dr. D. Franceschii, Dr. S. Sen mid altitude forests. and of an anonymous reviewer greatly improved The Vrysses assemblage represents a microphyll this paper.
flora consisting of 31 taxa. Remains of Acer pseudomonspessulanum , Pinus sp. , Quercus mediterranea and Ziziphus ziziphoides predominate. The assem- REFERENCES blage contains deciduous mesophytic elements
(approximately 37-59%), thermophilous evergreens ACHHAL A., AKABLI O., BARBERO M., BENABID A., M’HIRIT (appr. 11-30%) and xerophytic elements (appr. A., PEYRE C., QUEZEL P. & RIVAS- MARTINEZ S. 15-52%). 1980. — À propos de la valeur bioclimatique et A comparison of the Vrysses plant remains with dynamique de quelques essences forestières au Maroc. Ecologia Mediterranea 4: 211-248.
recent taxa and their ecological needs leads to the ARABATZIS T. I. 1998. — Shrubs and Trees of Greece, following reconstruction: riparian forests with vari- vol. 1. Technological Education Foundation of Kaous taxa of Salix sp. and possibly Populus tremula , vala, Kavala, 292 p.
Juglandaceae existed, including possibly several ARABATZIS T. I. 2001. — Shrubs and Trees of Greece, thermophilous evergreens, such as Daphnogene vol. 2. Technological Education Foundation of Kavala, Kavala, 435 p.
polymorpha . Sclerophyllous vegetation comprising BAILEY I. W. & SINNOTT E. W. 1915. — A botanical possibly Quercus mediterranea , Quercus sp. , maybe index of Cretaceous and Tertiary climates. Science Buxus pliocenica , Acer pseudomonspessulanum , Ul- 41: 831-834.
maceae and several legumes stood most likely on dry BAILEY I. W. & SINNOTT E. W. 1916. — The climatic lowland soils. Mixed mesophytic deciduous trees, distribution of certain types of angiosperm leaves. American Journal of Botany 3: 24-39.
such as Acer sp. , Populus tremula , Juglans acuminata, BARBERO M., LOISEL R. & QUEZEL P. 1992 . — Biogeo- Ziziphus ziziphoides and possibly Acer pseudomons- graphy, ecology and history of Mediterranean Quercus pessulanum and Ulmaceae , with evergreen ( Buxus ilex ecosystems. Vegetatio 99/100: 19-34.
Miocene leaf assemblage from Vrysses, western Crete
BELZ G. & MOSBRUGGER V. 1994. — Systematisch- green broad-leaved (non-ombrophilous) forest in the paläoökologische und paläoklimatische Analyse von subtropical and temperate zones. Veröffentlichungen Blattfloren im Mio-Pliozän der Niederrheinischen des Geobotanischen Institutes der ETH, Stiftung Rübel Bucht (NW-Deutschland). Palaeontographica Abt. 98: 169-195.
B 223: 19-156. KNOBLOCH E. & KVAČEK Z. 1976. — Miozäne Blätter- BENDA L., HILTERMANN H., KUSS S. E. & SYMEONI- floren vom Westrand der Böhmischen Masse. Zedice DIS N. K. 1968. — Der erste Mastodon-Fund der Rozpravy Ustredniho Ustava Geologickeho 42: 1-131. Insel Kreta. Annales géologiques des Pays helléniques KNOBLOCH E. & VELITZELOS E. 1987. — New leaf floras 21: 167-177. in the Neogene of Greece. Bulletin of the Geological BERGER V. W. 1954. — Flora und Klima im Jungtertiär Survey, Praqua 62 (3): 157-164.
des Wiener Beckens. Zeitschrift der Deutschen Geolo- KÖPPEN W. 1931. — Grundriss der Klimakunde. Walter gischen Gesellschaft 105 (2): 228-233. de Gruyter and Co., Berlin, 230 p.
BROWICS K. 1982. — Chorology of Trees and Shrubs in KOVAR- EDER J. 1987. — Pannonian (Upper Miocene) Southwest Asia and Adjacent Regions vol. 1. Polish vegetation character and climatic inferences in the Scientific Publishers, Warzawa, 172 p. central Paratethys Area. Annalen des Naturhistorischen BRUCH A. A., UTESCHER T.,OLIVARES C. A., DOLAKOVA Museums in Wien 88: 117-129.
N., IVANOV D. & MOSBRUGGER V. 2004. — Mid- KOVAR- EDER J. 1988. — Obermiozäne (Pannone) Floren dle and late Miocene spatial temperature patterns aus der Molassezone Österreichs. Beiträge zur Palä- and gradients in Europe. Preliminary results based ontologie Österreich 14: 19-121.
on palaeobotanical climate reconstructions. Courier KOVAR- EDER J. & KVAČEK Z. 1995. — Der Nachweis Forschungsinstitut Senckenberg 249: 15-27. eines fertilen Zweiges von Tetraclinis brachyodon DERMITZAKIS M. D. & PAPANIKOLAOU D. 1981 . — (Brongniart) Mai et Walter from Rodoboy, Croatia Paleogeography and geodynamics of the Aegean (Middle Miocene). Flora 190: 261-264.
region during the Neogene. Annales géologiques des KOVAR- EDER J., KVAČEK Z., ZASTAWNIAK E., GIVULESCU Pays helléniques, hors serie 3: 246-290. R., HABLY L., MIHAJLOVIC D., TESLENKO J. & WAL- FASSOULAS C. 2001. — Field Guide of Crete Geology. TER H. 1996. — Floristic trends in the vegetation Natural History Museum of Crete, Irakleio, 103 p. of the Paratethys surrounding areas during Neogene FERGUSON D. K. 1985. — The origin of leaf-assemblages, time, in BERNOR R. L., FAHLBUSH V., MITTMANN new light on an old problem. Review of Palaeobotany H.-W. (eds), The Evolution of Western Eurasian Neo- and Palynology 46: 117-188. gene Mammal Faunas III. Columbia University Press, FRYDAS D. & KEUPP H. 1996. — Biostratigraphical results New York: 395-413.
in Late Neogene deposits of NW Crete, Greece, based KOVAR- EDER J., KVAČEK Z., MARTINETTO E. & ROIRON on calcareous nannofossils. Berliner Geowissenschaftliche P. 2006. — Late Miocene to Early Pliocene vegetation Abhandlungen, Reihe E 18: 169-189. of southern Europe (7-4 Ma) as reflected in the mega- GASTALDO R. A. & FERGUSON D. K. 1998. — Recon- fossil plant record. Palaeogeography Palaeoclimatology structing Tertiary communities: introductory remarks. Palaeoecology 238: 321-339.
Review of Palaeobotany and Palynology 101: 3-6. KUSS S. E. 1980. — Führer zur Kreta – Exkursion. Geo- GIVULESCU R. 1962. — Die fossile Flora von Valea logisch-Paläontologisches Institut der Universität Neagra. Palaeontographica 110: 128-187. Freiburg, Freiburg i.Br., 51 p.
GREGOR H. J. 1990. — Contributions to the Late KVAČEK Z. 1989. — Fossil Tetraclinis Mast. (Cupres- Neogene and Early Quaternary floral history of the saceae). Journal of the National Museum, ser. B, Praqua Mediterranean. Review of Palaeobotany and Palyno- 155 (1-2): 45-56.
logy 62: 309-338. KVAČEK Z. & WALTER H. 1974. — Bemerkenswerte und HEYWOOD V. H., MOORE D. M., RICHARDSON I. B. K. seltene cinnamomoide Blätter aus dem Grenzbereich & STEARN W. T. 1993. — Flowering Plants of the des Oligo-Miozäns Mitteleuropas. Abhandlungen des World. B.T. Butsford Ltd, London, 335 p. Staatliche Museums für Mineralogie und Geologie zu HICKEY L. J. 1973. — Classification of the architecture Dresden 21: 197-221.
of dicotyledonous leaves. American Journal of Botany KVAČEK Z. & WALTER H. 1989. — Paleobotanical 60 (1): 17-33. studies in Fagaceae of the European Tertiary. Plant HSU K. J., RYAN W. B. F. & CITA M. B. 1973. — Late Systematics and Evolution 162: 213-229.
Miocene desiccation of the Mediterranean. Nature KVAČEK Z., BŮŽEK Č. & HOLÝ F. 1982. — Review of 242: 240-244. Buxus fossils and a new large-leaved species from the JOHNSTON M. C. 1963. — The species of Ziziphus Miocene of central Europe. Review of Palaeobotany indigenous to United States and Mexico. American and Palynology 37: 361-394.
Journal of Botany 50: 1020-1027. KVAČEK Z., VELITZELOS D. & VELITZELOS E. 2002. — KLÖTZLI F. 1988. — On the global position of the ever- Late Miocene Flora of Vegora Macedonia N Greece.
Zidianakis G. et al.
University of Athens, Faculty of Geology, Depart- SACHSE M. 2004. — Die neogene Mega- und Miment of Historical Geology and Paleontology, Korali kroflora von Makrilia auf Kreta und ihre aussagen Publications, Athens, 175 p. zur Klima- und Vegetationgeschichte des östlichen
LEROY S. A. G. & ROIRON P. 1996. — Latest Pliocene Mittelmeergebietes, in GREGOR H. J. (ed.), Flora pollen and leaf floras from Bernasso palaeolake (Es- Tertiaria Mediterranea. Die tertiären Floren des candorgue Massif, Hérault, France). Review of Pal- Mittelmeergebietes. Vegetationsgeschichte, Phyaeobotany and Palynology 94: 295-328. tostratigraphie, Paläoökologie, Paläoklimatologie,
MEULENKAMP J. E., DERMITZAKIS M., GEORGIADOU- Paläogeographie. Documenta Naturae, München DIKEOULIA E., JONKERS H. A. & BEÖGER H. 1979. — 6 (12): 1-323.
Field Guide to the Neogene of Crete. Department of SACHSE M. & MOHR B. A. R. 1996. — Eine obermiozäne Geology and Paleontology, University of Athens, Makro- and Mikroflora aus Südkreta (Griechenland), Athens, Series A, 32 p. und deren paläoklimatische Interpretation-Vorläufige
MEUSEL H., JÄGER E., RAUSCHERT S. & WEINERT T. Betrachtungen. Neues Jahrbuch für Geologie und Pa- 1965. — Vergleichende Chorologie der zentraleuropä- läontologie Abhandlungen 200: 149-182.
ischen Flora, vol. 1. VEB Fisher, Jena, 583 p. SACHSE M., MOHR B. & SUC J.-P. 1999. — The Makrilia-
MEUSEL H., JÄGER E., RAUSCHERT S. & WEINERT T. flora. A contribution to the Neogene History of the 1978. — Vergleichende Chorologie der zentraleuropä- climate and vegetation of the Eastern Mediterranean, ischen Flora, vol. 2. VEB Fisher, Jena, 418 p. Acta Palaeobotanica Suppl. 2: 365-372.
MOHR B., KÖHLER J. & ADAMEK- JACOBS E. 1991. — A SAPORTA G. & MARION A. F. 1876. — Recherches sur les late Miocene leaf flora from southern Crete (Greece). végétaux fossiles de Meximieux. Archives du Muséum Paneuropean Palaeobotanical Conference Vienna: 26. d’Histoire naturelle de Lyon 1: 131-335.
MOSBRUGGER V. & UTESCHER T. 1997. — The coexistence SHAKRYL A. K. 1992. — Leguminosae species from the approach – a method for quantitative reconstructions Tertiary of Abkhazia, in HERENDEEN P. S. & DILCHER of Tertiary terrestrial palaeoclimate data using plant D. L. (eds), Advances in Legume Systematics. Part 4. fossils. Palaeogeography Palaeoclimatology Palaeoecol- The Fossil Record. The Royal Botanic Gardens, Kew: ogy 134: 61-86. 189-206.
NEGRI A. & VILLA G. 2000. — Calcareous nannofossil SHATILOVA I. I. & STUCHLIK L. 2001. — On the history biostratigraphy, biochronology and paleoecology at of the development of the family Hamamelidaceae in the Tortonian/Messinian boundary of the Faneromeni Eurasia. Palaeontographica 252: 235-244.
Section (Crete). Palaeogeography Palaeoclimatology SONG Y. 1983. — Die räumliche Ordnung der Vegeta- Palaeoecology 156: 195-209. tion Chinas. Tuexenia, N.S. 3: 131-157.
PALAMAREV E. 1989. — Palaeobotanical evidences of SONG Y. 1988. — Broad-leaved evergreen forests in the Tertiary history and origin of the Mediterranean Central Japan in comparison with Eastern China. sclerophyll dendroflora. Plant Systematics and Evolu- Veröffentlichungen Geobotanisches Institut ETH, Stiftion 162: 93-107. tung Rübel 98: 197-224.
PETERS J. M. 1985. — Neogene and Quaternary vertical SPICER R. A. 1991. — Plant taphonomic processes, in tectonics in the south Hellenic arc and their effect ALLISON P. A. & BRIGGS D. E. G. (eds), Taphonomy. on concurrent sedimentation processes. GUA, S1, Releasing the Data Locked in the Fossil Record. Plenum 23: 1-247. Press, New York: 71-113.
POPOV S. V., RÖGL F., ROZANOV A. Y., STEININGER F.F., STRAHLER A. H. & STRAHLER A. N. 1992. — Mod- SHECHERBA I. G. & KOVAC M. (eds) 2004. — Litho- ern Physical Geography. Wiley & Sons, New York, logical-Paleogeographic maps of Paratethys. Courier 638 p.
Forschungsinstitut Senckenberg 250: 1-46. TEN VEEN J. H. 1998. — Neogene outer-arc evolution
QUEZEL P. 1988. — Esquisse phytogéographique de la in the Cretan segment of the Hellenic arc: tectonic, végétation climatique potentielle des grandes îles médi- sedimentary and Geodynamic reconstructions. Geoterranéennes. Bulletin d’Écologie 19 (2-3): 121-127. logica Utraiectina 160: 1-184.
ROTH J. L. & DILCHER D. L. 1978. — Some consid- TURLAND N. J., CHILTON L. & PRESS J. R. 1993. — Flora erations in leaf size and leaf margin analysis of fossil of the Cretan Area. The Natural History Museum, leaves. Courier Forschungsinstitut Senckenberg 30: London, 439 p.
165-171. UNGER F. 1867. — Die fossile Flora von Kumi auf
SACHSE M. 1997. — Die Makrilia-flora (Kreta, Grie- der Insel Eubpoea. Kaiserlichen Akademie der Wischenland) – Ein Beitrag zur Neogenen Klima- und senschaften, Mathematisch. Naturwissenschaftlichen Vegetationsgeschichte des östlichen Mittelmeergebietes. Classe 27: 27-87.
Unpubl. PhD Thesis No. 12 250 ETH, Eidgenössi- UTSESCHER T. & MOSBRUGGER V 1999. — Palaeoflora schen Technischen Hochschule Zürich, Switzerland, Database, Palaeoclimate Reconstructions Using Tertiary 312 p. Plant Taxa. Bonn University, accessible via inter-
Miocene leaf assemblage from Vrysses, western Crete net: http://www.geologie.uni-bonn.de/palaeoflora/ University Press, Harvard, publ. no. 5, 313 p.
palaeoflora_home.htm WALTER H. & LIETH H. 1967. — Klimadiagramm- VELITZELOS E. & SCHNEIDER H. E. 1978. — Jungter- Weltatlas. VEB Gustav Fischer, Jena, 253 p.
tiäre Pflanzenfunde aus dem Beckenvon Vegora in WOLFE J. A. 1979. — Temperature parameters of hu- West-Mazedonien. 3. Mitteilung: Eine Fächerpalme mid to mesic forests of Eastern Asia and relation to ( Chamaerops humilis L.). Annales géologiques des Pays forests of other regions of the Northern Hemisphere helléniques 29 (2): 796-799. and Australasia. Geological Survey Professional Paper VELITZELOS E. & GREGOR H. J. 1990. — Some aspects 1106: 1-37.
of the Neogene floral history in Greece. Review of WOLFE J. A. 1985. — Distribution of major vegetational Palaeobotany and Palynology 62: 291-307. types during the Tertiary. Geophysical Monograph 32: VELITZELOS E., KVAČEK Z. & VELITZELOS D. 2001. — 357-375.
The cenophytic floras of Greece, in ZOUROS N. C. ZIDIANAKIS G. 2002. — The Late Miocene Flora of Vrysses (ed.), International Symposium for Protected Areas. (Western Crete-Greece). A Contribution to the Climate Lesvos Natural History Museum of Petrified Forest, and Vegetation History of Crete. Master of Science Thesis, Lesvos, Greece, 639 p. University of Crete, Irakleio, Greece, 118 p.
VELITZELOS E., KVAČEK Z., DENK T.&VELITZELOS D. ZIDIANAKIS G., MOHR B. & FASSOULAS C. 2004. — The 2002. — Field Trip Guidebook Evia Island Neogene late-Miocene flora of Vrysses, western Crete. A con- Sedimentary Basin of Kymi/Aliveri. University of Ath- tribution to the climate and vegetation history, in ens, Faculty of Geology, Department of Histori- CHATZIPETROS A. A. & PAVILDES S. B. (eds), 5th cal Geology and Paleontology, Korali Publications, International Symposium on Eastern Mediterranean Athens, 61 p. Geology, Proceddings Vol. 1. Thessaloniki, Greece: VIDAKIS M., TRIANTAPHYLLIS M., & MYLONAKIS I. 515-518.
1983. — SheetVrysses. Geological Maps of Greece (Scale ZOHARY M. 1947. — A vegetation map of Western 1:50000). IGME, Athens. Palestine. Journal of Ecology 13: 1-19.
WANG C. W. 1961. — The Forests of China with a Sur- ZOHARY M. & ORSHAN G. 1966. — An outline of the vey of Grassland and Desert Vegetation. Maria Moors geobotany of Crete. Journal of Botany 14 Suppl.: Cabot Foundation for Botanical Research, Harvard 1-49.
Submitted on 23 December 2005; accepted on 17 April 2007.
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