Quercus mediterranea Unger

Zidianakis, Giannis, Mohr, Barbara A. R. & Fassoulas, Charalampos, 2007, A late Miocene leaf assemblage from Vrysses, western Crete, Greece, and its paleoenvironmental and paleoclimatic interpretation, Geodiversitas 29 (3), pp. 351-377: 358-377

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http://doi.org/10.5281/zenodo.5374994

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scientific name

Quercus mediterranea Unger
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Quercus mediterranea Unger   ( 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.

Dicotylophyllum sp. 5  

( 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.

Dicotylophyllum sp.   12

( 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.

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Submitted on 23 December 2005; accepted on 17 April 2007.

Kingdom

Plantae

Phylum

Tracheophyta

Class

Magnoliopsida

Order

Fagales

Family

Fagaceae

Genus

Quercus