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351GEODIVERSITAS • 2007 • 29 (3) © Publications Scienti ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com

A late Miocene leaf assemblage

from Vrysses, western Crete, Greece,and its paleoenvironmentaland paleoclimatic interpretation

Giannis ZIDIANAKISDepartment of Biology, University of Crete,Voutes, GR-71305 Irakleio, Crete (Greece)

[email protected]

Barbara A. R. MOHRInstitute of Palaeontology, Museum of Natural History,

Invalidenstr. 43, D-10115 Berlin (Germany)barbara. [email protected]

Charalampos FASSOULASNatural History Museum, University of Crete,

GR-71409 Irakleio, Crete (Greece)[email protected]

Zidianakis G., Mohr B. A. R. & Fassoulas C. 2007. — A late Miocene leaf assemblage fromVrysses, western Crete, Greece, and its paleoenvironmental and paleoclimatic interpretation.Geodiversitas 29 (3) : 351-377.

ABSTRACT A new leaf impression ora is described from late Miocene marly sediments(6-7.5 Ma) near the village of Vrysses in northwestern Crete, Greece. Te stud-ied plant remains represent more than 30 taxa of conifers and angiosperms.Deciduous broad-leaved woody plants such as Acer pseudomonspessulanum, 5-lobed Acer , Ziziphus ziziphoides , “Parrotia ” pristina , Populus tremula , Salixsp. and Juglandaceae, prevail in this leaf assemblage. Tese plants are associatedwith sclerophyllous, evergreen taxa (Quercus mediterranea , Quercussp., Buxus pliocenica and Pinussp.) and a few subtropical Miocene elements (Daphnogene polymorpha , etraclinis sp.). A regional vegetation reconstruction is developedbased on the ecological affi nities of the nearest living relatives. Te climateconditions are deduced from the physiognomy and composition of the Vryssesplant assemblage. Tey indicate a warm-humid climate possibly with a weaklydeveloped dry period during the summer months.

KEY WORDSLate Miocene,

Crete,Greece,leaves,

paleovegetation,paleoclimate.

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352 GEODIVERSITAS • 2007 • 29 (3)

Zidianakis G.et al.

IN RODUC ION

During the last decades progress had been made inunderstanding Late Neogene eastern Mediterraneanvegetation and climate. For many years the resultsof paleoclimatic studies were highly controversialand often hampered by the lack of precise dating ofplant bearing strata. Terefore paleoclimatic recon-structions for Miocene climates differed, rangingfrom humid climates with maximum precipitationduring summer (e.g., Gregor 1990) to stronglyseasonal climates with minimum precipitationduring summer (e.g., Palamarev 1989). Recentclimate reconstructions for the European erti-ary using the Coexistence Approach revealed thatspatial temperature gradients became more stronglydeveloped during the early late Miocene and musthave become even more differentiated towards theend of the Miocene (Bruchet al. 2004). Studies ona well dated late ortonian plant assemblage fromeastern Crete near the village Makrilia conrmedthat besides laurophyllous and deciduous taxa,the sclerophyllous Mediterranean component wasmore pronounced than in central European oras

of similar age (Sachseet al. 1999) indicating pos-sibly a seasonal climate with a (weakly developed)summer drought.

In western Crete plant megafossils have neverbeen collected systematically so far. Meulenkampet al. (1979) were the rst who recognized plantmacro-fossils in the laminated marls of the Vryssesbasin. During the last decades more fossils cameto light in small quarries where Neogene limestoneis worked for building purposes. In order to studyNeogene vegetation and climate of this part of theisland, a survey on Neogene sedimentary rocks allover western Crete was undertaken (Zidianakis2002) for new outcrops containing plant macro-remains.

Although stratigraphy and fauna of Cretan Neo-gene strata are well studied (Meulenkampet al. 1979;Kuss 1980; Dermitzakis & Papanikolaou 1981; Pe-ters 1985; Frydas & Keupp 1996; ten Veen 1998),vegetation studies are rare and mainly focused oneastern Crete (Mohret al. 1991; Sachse & Mohr1996; Sachse 1997; Sachseet al. 1999).

Te discovery and study of the Vrysses plant as-semblage offers the opportunity for a comparison

RÉSUMÉUne paléoore du Miocène supérieur de Vrysses, Crète occidentale (Grèce) et soninterprétation paléoenvironnementale et paléoclimatique.Une nouvelle ore constituée d’empreintes foliaires a été découverte dans dessédiments du Miocène supérieur (6-7,5 Ma) près du village de Vrysses au nordde la Crète (Grèce). L’étude des restes de plantes concerne plus de 30 taxa deconifères et d’angiospermes. Des arbres feuillus décidus, tels qu’ Acer pseudomonspessulanum, Acer à cinq lobes, Ziziphus ziziphoides , “Parrotia ” pristina , Populustremula , Salix sp. et Juglandaceae, dominent cette paléoore. Ces plantes sontassociées à des taxa sclérophylles (Quercus mediterranea , Quercus sp.,Buxus plioce-nica etPinus sp.) et à quelques éléments subtropicaux du Miocène (Daphno gene polymorpha , etraclinis sp.). Un modèle de végétation régionale est développésur la base des affi nités écologiques des plus proches parents vivants. Les condtions climatiques sont déduites de la physiognomie et de la composition de lapaléoore de Vrysses et elles indiquent la présence d’un climat chaud et humideavec peut-être une légère période sèche pendant les mois d’été.

MOTS CLÉSMiocène supérieur,

Crète,Grèce,

feuilles,paléovégétation,

paléoclimat.



354 GEODIVERSITAS • 2007 • 29 (3)

Zidianakis G.et al.

Lefka Ori Mountains, the largest mountain area ofthe island with many summits over 2000 m. Tebasin develops in a graben structure inclined towardsthe sea that fragments the same Mesozoic carbonaterocks that build the Lefka Ori Mountains (Vidakiset al. 1983).

Te Vrysses Formation, that rests unconformablyon Mesozoic crystalline limestone and dolomitesof the Plattenkalk and ripali units (Vidakiset al. 1983), covers most of the area around Vrysses. Ac-cording to Meulenkampet al. (1979), the VryssesFormation is mainly composed of bioclastic andreefal, light-colored limestones and alternations oflaminated shallow water, white to grayish marls. Arich fauna represented by molluscs (mainlyOstrea )occurs in the limestone beds, whereas molluscs,echinoderms and remnants of sh and plants arefound in the marls. Based on planktonic foraminiferaMeulenkampet al. (1979) correlated the formationto theGloborotalia conomiozea biozone which tracesthe ortonian/Messinian boundary in the Medi-terranean basin (Negri & Villa 2000). Accordingto the mammal bio-stratigraphy the formation isplaced between the MN12 and MN13 biozones.Te age of the formation is thus regarded to be lateortonian-Messinian (6-7.5 ma). In lignite layers

of the Vrysses basin of about the same age, Bendaet al. (1968) found fragments of a mastodon tooth.No younger sediments, except of Holocene riverbeddeposits, are cropping out in the area.

Te Vrysses beds are almost horizontal or slightlyinclined to the northeast. In the study area limestoneand marls can be approached in naturally erodedoutcrops or in active quarries. Te typical sectionin the study area can be lithologically subdividedinto ve distinct horizons. Te lowermost (M1) isformed by massive white limestone, normally verycompetent, that shows irregular and angular fracture.Its thickness is at least 1 m and it is poor in fossilswhich are well preserved. Te next horizon (M2) isconstituted of well laminated yellowish to grayishmarls and marly limestone of an overall thicknessof 50 cm. Most of the plant remains are found inthe marly limestone. In these strata animal remainsare limited to a few insect wings and sh scales. Tethird horizon (M3) is made of a thin bed of yellow,well stratied marl with few fossils and high organic

component. Te M4 horizon comprises a whitishsandy to marly limestone of an average thicknessup to 1 m. Tis horizon is full of mollusks, mainlygastropods and ostracodes and has a moderatecompetency and hardness. Finally, the uppermosthorizon (M5) is formed by a stratied white sandylimestone that is partly covered by soil. A few leaffossils have been found in this horizon too.

Paleogeographic studies of Crete (Meulenkampetal. 1979; Dermitzakis & Papanikolaou 1981) suggestthat during the Miocene, Crete was fragmented intosmaller and larger islands, some of them located atthe present mountain areas. Te Lefka Ori area wasat that times a hilly area, surrounded by shallowmarine, lagoonal and delta environments. Duringthe late Miocene tectonic activity formed an elon-gated marine realm at the northern anks of theLefka Ori Mountains that was extending eastwardsto the Rethimnon area and was characterized byshallow bays, reefs and small islands. During thisperiod and prior to the Messinian salinity crisis thelight-colored limestone and marls of the VryssesFormation were deposited.

MA ERIAL AND ME HODS

More than 250 specimens of plant macro-remainswere excavated from the two quarries mentionedabove. In addition, 12 specimens were collectedfrom stone walls in the Mediterranean AgronomicInstitute of Chania that originated from the Vryssesquarries. All studied plant fossils are preserved asleaf impressions. Te specimens were studied af-ter careful cleaning and in a few cases mechanicalpreparation was applied. Unfortunately, the leaveswere in most cases fragmented, very rarely complete.Te lack of organic material on nearly all of theleaf specimens did not allow any cuticular analysis.However, the ne-grained marls maintained manyleaf characteristics for a large number of specimens. A Leica MZ8 stereoscope was used to observe detailsof the leaf morphology and a Canon digital camera(Powershot G4) for the photographs. ransparenttracing paper was used for drawings of the leaves.Te majority of the material is stored in the Pal-aeontological Department of the Natural History



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Miocene leaf assemblage from Vrysses, western Crete

GEODIVERSITAS • 2007 • 29 (3)

A

B

C

D E F

G

H I

JK

FIG. 2. — Photographs of fossil leaves from the Vrysses outcrops (western Crete): A , B , Pinus sp., needles nos. 32.1.2.168 and 32.1.2.48;C , D , Tetraclinis sp., twig fragments nos. 32.1.2.126 and 32.1.2.141; E , Daphnogene polymorpha (A.Braun) Ettingsh., no. 32.1.2.20;F , cf. “ Parrotia ” pristina (Ettingsh.) Stur, no. 32.1.3.5; G , cf. Juglans acuminata Braun ex Unger, no. 32.1.3.1; H -K , Quercus mediter-

ranea Unger, nos. 32.1.2.24, 32.1.2.155, 32.1.3.2 and 32.1.2.79. Scale bars: 1 cm.



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Museum of Crete in Irakleio while the 12 specimensfrom stone walls are housed in the Mediterranean Agronomic Institute of Chania.

Te determination and systematic affi nities of theVrysses specimens is based on morphological char-acteristics of the leaves (shape and size of lamina,margin type, leaf venation, etc.) through studiesof herbarium specimens of the Natural HistoryMuseum of Crete as well as published descriptionsand illustrations of Neogene plant assemblages ofEurope and modern regional oras.

In order to estimate climatic parameters of theVrysses area during the Late Neogene, we appliedtwo different methods that t Vrysses materialcharacteristics and preservation conditions:– 1) leaf margin analysis (after Bailey & Sinnott1915, 1916; Roth & Dilcher 1978; Wolfe 1979,1985). Tis method correlates leaf margin shapewith mean annual temperatures (MA ). In tropicforests most canopy species are entire-marginedin order to conserve water. Moving to the temper-ate zone, the amount of transpiration is generallyless critical and the number of woody species withtoothed leaves tends to increase. So the percent-age of entire-margined species can be used as anapproach to reconstruct the temperature regime.Studies on humid to mesic broad-leaved forestsof eastern Asia have shown that the percentageof species with entire-margined leaves is in directproportion to the mean annual temperature of theparticular forest (Wolfe 1979);– 2) co-existence approach (Mosbrugger & Utescher1997; Gastaldo & Ferguson 1998). Tis methoduses nearest living relatives of the fossil taxa in or-der to estimate several of the main parameters ofpaleoclimate. It is based on the assumption thatNeogene plant taxa had similar climatic require-ments as their modern nearest relatives in order todetermine the climatic interval in which all relativesof the assemblage can co-exist.

SYS EMA ICS

All leaf taxa found in the Vrysses outcrops are brieydescribed below, along with some notes on possiblesystematic relationships as well as geographic and

stratigraphic occurrences. Tey comprise remainsof gymnosperms, such as Pinaceae (Fig. 2A, B)and Cupressaceae (Fig. 2C, D), dicotyledonous(Figs 2E-K; 3; 4A-J; 5) and monocotyledonousangiosperms (Fig. 4K-L). Te material is listed withthe collection numbers of Natural History Museumof Crete. Te terminology of leaf architecture isbased on Hickey (1973).

Division GYMNOSPERMAEFamilyPINACEAEGenusPinusL.

Pinus sp.(Fig. 2A, B)

M ATERIAL EXAMINED. — Specimens nos. 32.1.2.23;32.1.2.48; 32.1.2.71; 32.1.2.100; 32.1.2.129; 32.1.2.154;32.1.2.156; 32.1.2.162; 32.1.2.163; 32.1.2.168;32.1.2.170; 32.1.2.183; 32.1.2.185; 32.1.2.201;32.1.2.212; 32.1.2.213; 32.1.2.239.

DESCRIPTIONSeventeen fascicles of two leaves or isolated leaffragments have been recovered, linear in outline,quite thin. Some of these pine needles are morethan 12 cm long.

R EMARKSTe taxon described above is possibly related tothe modern Mediterranean long leaf speciesPinushalepensis Mill. orPinus pinaster Aiton because ofthe unusual leaf length.

FamilyCUPRESSACEAEGenus etraclinisMast.

Tetraclinis sp.(Fig. 2C, D)

M ATERIAL EXAMINED. — wigs nos. 32.1.2.73; 32.1.2.125;32.1.2.126; 32.1.2.141.

DESCRIPTIONFour twig segments of the genusetraclinis composedof small, closely attached leaves fused in veticles of four.Lengths of specimens varying from 0.5 to 3 cm.



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R EMARKSBecause of poor preservation these samples are notidentiable to the species level.etraclinis is a com-mon element in Oligocene and Miocene strata ofEurope. During the Miocene existed possibly twospecies (Kvaček 1989; Kovar-Eder & Kvaček 1995),

etraclinis salicornioides (Unger) Kvaček andetra-clinis brachyodon (Brongn.) Mai & Walter. Both dif-fered mainly in epidermal structure of their leaves.

etraclinis salicornioides is considered to have beena thermophilous element of the undergrowth ofmixed evergreen and deciduous forests (Kovar-Ederet al. 1996), though. brachyodonwas most likely amore xeromorphic element growing mainly in low-land forest along sea coasts (Kvaček 1989). oday,a single species exists,etraclinis articulata Mast.,which thrives in the Atlas Mountains (Morocco)and is well adapted to the Mediterranean climate(Achhalet al.1980; Kovar-Ederet al. 1996).

Division ANGIOSPERMAEClass DICO YLEDONAE

FamilyL AURACEAEGenusDaphnogeneUnger

Daphnogene polymorpha (A.Braun) Ettingsh.(Figs 2E; 5D)

M ATERIAL EXAMINED. — Complete leaf no. 32.1.2.20;fragmentary leaf no. 32.1.2.26.

DESCRIPTIONOne complete leaf and one fragmentary specimen.Leaves petiolate, about 4 to 4.5 cm long and 1 to1.2 cm wide with entire margin. Shape of laminanarrowly elliptic or oblanceolate, obtuse at theapex, cuneate at the base. Venation acrodromous,triveined. Midvein thin, straight, lateral primaryveins sub-basal, slender, running in convergentarches toward the leaf apex. No additional detailsof the venation visible.

R EMARKSTese narrow triveined leaves are comparable tofoliage of fossilDaphnogene polymorpha . Such leaveswere formerly described under the generic name

CinnamomumSchaeff. However neither morphologynor anatomy allowed to obtain an exact correlationof this leaf type to the modern taxa of Lauraceaefamily (Kvaček & Walter 1974). Tus the name ofthe fossil genusDaphnogeneis preferred.Daphno- gene polymorpha is a common species in EuropeanNeogene deposits, with remarkable variability inleaf forms. Te plant is considered to represent apaleotropical element of riparian forests along riversand mixed mesophytic forests (Belz & Mosbrugger1994) where it survived in this southerly area tillLate Neogene.

FamilyH AMAMELIDACEAEGenusParrotiaC.A.Mey.

cf. “Parrotia ” pristina (Ettingsh.) Stur(Fig. 2F)

M ATERIAL EXAMINED. — Complete leaf no. 32.1.3.5.

DESCRIPTIONOne long petiolate, broadly obovate complete leaf(c . 5.5 cm long, 3.5 cm wide). Base rounded, slightlyasymmetrical, apex broadly rounded. Margin den-tate, with a few wide, obtuse teeth mainly at theupper part of lamina. Venation craspedodromous,primary vein slightly curved. Five pairs of second-ary veins, straight or slightly curved, branchingfrom the midvein at an angle of 30 to 40°. Tebasal pair of secondaries runs to the upper thirdof the lamina.

R EMARKSLeaf shape and venation are similar to the fos-sil species “Parrotia ” pristina. However, leaves ofthis fossil species usually show an acute or evenacuminate apex. In the specimen of Vrysses, theapex is broadly rounded thus the identicationremains somewhat uncertain. “Parrotia ” pristinaleaves have been found in numerous Miocene andPliocene outcrops throughout Europe (Shatilova &Stuchlik 2001). Due to the considerable similar-ity of these leaves in morphology and anatomicalfeatures to the recent generaFothergilla L. andParrotia , Kovar-Eder (1988) proposed to use the



358 GEODIVERSITAS • 2007 • 29 (3)

Zidianakis G.et al.

genus nameParrotia with quotation marks. ExtantParrotia is monotypic. Te only species,Parrotia

persica(DC.) C.A.Mey. is a deciduous mesic woodyplant, restricted in its distribution. It occurs in thecoastal area south of the Caspian Sea and to theCaucasus up to 1500 m altitude.

Family JUGLANDACEAEGenus JuglansL.

cf. Juglans acuminata Braun ex Unger(Figs 2G; 5F)

M ATERIAL EXAMINED. — Complete leaf no. 32.1.3.1;fragmentary leaves nos. 32.1.2.72; 32.1.2.158.

DESCRIPTIONOne almost complete and two incomplete leaves;broadly oval to elongate, slightly asymmetric inshape; at least 10 cm long and 4.5 cm wide. Mar-gin entire, base widely cuneate to rounded, apexnot preserved. Petiolec . 0.5 cm long. Primary veinstout, slightly bent. Venation brochidodromous,8-10 pairs of curved secondary veins arising at 35-60° from the midvein, regularly and widely spaced, joined together in a series of arches along the mar-gin. ertiary veins percurrent, straight or irregularlybranched. Leaf texture possibly leathery.

R EMARKSSize, shape and venation pattern of these specimensseems to t the description of the fossil species Juglansacuminata.Tis taxon is present in many EuropeanNeogene deposits. Leaets of Juglans sect. Juglans show entire margins (e.g., J. regiaL.). Terefore, Juglans acuminata may be related to modern Jug-lans regia that grows in mixed mesophytic forestsin southeast Europe and North Asia. However,Knobloch & Kvaček (1976) had studied the epi-dermal anatomy, and considered Juglans acuminata to be related to the North American species JuglansrupestrisEngelm.

Since several species ofPterocarya Kunth alsohave leaves with entire margins, it can not be com-pletely ruled out that the Vrysses leaves belong tothis taxon. In the Makrilia plant assemblages from

eastern Crete pollen ofPterocarya was a commonelement (Sachse & Mohr 1996).

FamilyF AGACEAEGenusQuercusL.

Quercus mediterranea Unger(Figs 2H-K; 5J)

M ATERIAL EXAMINED. — Complete leaves nos. 32.1.2.6;32.1.2.17; 32.1.2.79; 32.1.2.155; 32.1.3.2; fragmentaryleaves 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.

DESCRIPTIONFive complete leaves very variable in shape, 3.3 to6.9 cm long and 1.8 to 4.7 cm wide. Shape broadlyoval to ovate or obovate. Base acute or rounded,rarely cordate and apex acute, obtuse or slightlyemarginate. Leaf margin simple serrate at least inthe upper half of the lamina, occasionally entire orundulate. eeth widely spaced, rather small, moreor less sharply pointed. Venation craspedodromous,midvein stout, in the upper part more slender andin some cases slightly sinuous. Secondary veinsdistinct, in 5 to 9 pairs, arising at an angle of 30to 80° from the primary vein, irregularly spaced,rarely forked, running straight or bent towards theleaf margin entering the teeth. ertiary veins poorlyvisible, irregular, simple or branched. Leaf texturerather chartaceous or coriaceous. Also, 7 fragmentaryleaves are assigned to this fossil species.

R EMARKSQuercus mediterranea is a characteristic commonelement in Neogene deposits especially in southernEurope. In Greece, this species is represented innumerous Miocene and Pliocene plant assemblagesof mainland and islands (Knobloch & Velitzelos1987; Velitzelos & Gregor 1990). Leaf remains ofQuercus mediterranea from the Vrysses outcropscorrespond much better to the records from Vegora(late Miocene, N Greece) and Kymi (early MioceneEvia Island, central Greece) on the basis of gross-morphology (Velitzeloset al. 2002; Kvačeket al. 2002; Kvaček & Walter 1989) while leaves assignedto Q. mediterranea from Makrilia (late Miocene,



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eastern Crete, S Greece) seem to differ slightly. Atthe latter locality oaks are represented by a fewleaves, which are rather small. Tey hardly reach alength of 2.5 cm (Sachse 1997).

Based on studies of leaf epidermis tissue fromLava (North Greece) that indicated an extremelythin cuticle, it was suggested thatQ. mediterranea was a mesophytic deciduous oak (Kvaček & Walter1989). However newly obtained epidermal charac-teristics show that in spite of the thin cuticle, theepidermis of either leaf side is sclerenchymatousunderlain by a hypodermis (Kvačeket al. 2002).Tus Q. mediterranea probably was a sclerophyllouselement able to survive short dry periods.

Quercus mediterranea is perhaps ancestral to themodern evergreen speciesQuercus ilex L. andQuer-cus coccifera L., although both do not correspondin all details of leaf anatomy and morphology tothe fossils. oday these oaks are present through-out the Mediterranean basin and on coastal slopestowards the Atlantic (South France) and Black Sea,on relatively dry soils.

According to Barberoet al . (1992),Q. ilex thrivesin a wide climatic range (from warm-temperate tosubtropical and from medium humid to semi-arid)due to numerous morphological adaptations. Inthe northern Mediterranean realm,Q. ilex formswoodland with several laurophyllus elements suchas ArbutusL., HederaL., Ilex L. andViburnumL.,while in south it thrives together with more xero-thermic taxa such as JuniperusL., Phillyreaourn.ex Adans., Genista , EphedraL. and Buxus . In theeastern Mediteranean basinQ. ilex is replaced byQ. coccifera .

Quercus sp.(Figs 3A; 5Q)

M ATERIAL EXAMINED. — Fragmentary leaves nos.32.1.2.19; 32.1.2.95; 32.1.2.142.

DESCRIPTIONTree leaf fragments, long petiolate (up to 1.5 cm).Complete leaf length estimated at 7 to 9 cm orperhaps more, width about 2.5 to 3.2 cm. Shapeof lamina ovate to broadly elliptic, leaf margin un-

dulate or simply serrate with small widely spacedteeth, serration starting in the lower third of thelamina. Base cordate, apex acute maybe acuminate.Venation craspedodromous. Midvein prominentmore or less curved, slightly zig-zag at the upperpart. Secondary veins more than 10 pairs, regularlydisposed, quite dense, diverging at angles of 40 to80° and running straight, parallel to each other,across lamina, entering marginal teeth. Higherorder venation poorly preserved.

R EMARKSSuch leaves may be assigned to the genusQuercus ,representing probably an evergreen oak quite dif-ferent fromQ. mediterranea especially in leaf sizeand shape and secondary veins pattern. Leaf remainsfrom early Miocene plant assemblage of Kymi guredby Unger (1867: taf. VI, gs 23-27) and identiedasQuercus drymejaUnger by Velitzeloset al. 2002(revised list of Kymi material), are very close in mor-phology to Vrysses specimens. However more andbetter preserved material is necessary to elucidatethe systematic relationships of these specimens.

FamilyULMACEAE

Ulmaceae gen. et sp. indet.(Fig. 3B)


DESCRIPTIONOne small, broadly elliptic leaf, about 1.8 cm longand 1 cm wide. Apex incomplete, probably acuteto acuminate, base rounded, strongly asymmetrical.Leaf margin serrate at the upper half part of laminawith several wide, simple teeth. Venation craspe-dodromous. Midvein prominent, quite thick andstrongly curved. From the primary vein arise fourpairs of secondaries at angles of 35 to 55°, ratherstrong, more or less curved, running into the toothapices. No other details of the venation preserved.

R EMARKSSuch a leaf shape (leaf margin, asymmetrical base,laminar shape) suggests a relationship with the



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family Ulmaceae. However this specimen lacksfurther diagnostic features for a determination togenus level. Te Ulmaceae is a family of tropicaland mainly temperate trees and shrubs, with 16genera (Heywoodet al. 1993). Due to very smallsize (nanophyll) and simple dentate margin, theVrysses specimen may represent a slightly xero-morphic element.

FamilyS ALICACEAEGenusSalixL.

cf. Salix sp.(Fig. 3C)

M ATE RIAL EXAMINED. — Fragmentary leaf no.32.1.2.39.

DESCRIPTIONBasal half of a probably narrow elliptic, slightlyasymmetrical leaf (3.9 cm long and 1.7 cm wide);entire leaf possibly twice that long. Leaf base acute,margin nely serrate with simple acuminate teeth,regularly spaced. Midvein medium, thick, welldeveloped and curved. Higher order venation notvisible.

R EMARKSShape and margin of this leaf show resemblance tothose of extantSalix L. However, because the upperpart of the leaf is missing, as well as higher ordervenation, a denite identication is not possible.

GenusPopulusL.

Populus tremula L., foss.(Fig. 3D)

M ATERIAL EXAMINED. — Complete leaf no. 32.1.3.7;fragmentary leaf no. 32.1.2.160.

DESCRIPTIONOne complete and one fragmentary leaf. Laminabroadly ovate, 4.5 to 4.8 cm long and about 5.5 to5.7 cm wide. Apex probably obtuse, base rounded,

margin dentate with obtuse teeth. Petiole not pre-served. Venation actinodromous with three pri-mary veins. Midvein slightly bent, laterals curvedand forked. Four or ve pairs of curved secondaryveins arising from the midvein at 30-55°, loopingwith the laterals and the adjacent secondaries nearthe margin. ertiary veins oblique forming meshesbetween secondaries.

R EMARKSShape, teeth at the leaf margin and venation ofthese specimens are similar to lamina of the mod-ern speciesPopulus tremula . Leaf remains of thispoplar have been found in several European Neo-gene deposits. odayPopulus tremula trees grow inopen mesophilous forests up to 1000 m of altitudein Europe, North Africa, Asia Minor and Siberia(Leroy & Roiron 1996).

FamilyBUXACEAEGenusBuxusL.

Buxus pliocenica Saporta & Marion(Figs 3E; 5M)

M AT ERIAL EXAMINED. — Complete leaf no.32.1.2.140.

DESCRIPTIONOne complete leaf, narrowly elliptic, with shortpetiole, 3.6 cm long and 1.2 cm wide. Leaf marginsentire, apex acute, base cuneate. Primary vein thinand almost straight. Secondary veins ne, arisingat an angle of 30 to 70°, closely spaced, parallelto each other, branched once to many times inacute angles without higher order veins rejoiningsecondaries.

R EMARKSBuxus pliocenica fossils have been known in Mi-ocene and Pliocene deposits of Eurasia, mostly inthe form of leaf impression and compression, rarelyas pollen or fossil fruits. Saporta & Marion (1876)treatBuxus pliocenicaas an ancestor of Buxus sem-

pervirensL. However, Kvačeket al. (1982) considerBuxus pliocenica closely allied to theB. sempervirens -



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A B C D

E

F

GH I

J K

L

M N O

P

FIG. 3. — Photographs of fossil leaves from the Vrysses outcrops (western Crete): A , Quercus sp., no. 32.1.2.142; B , Ulmaceae gen.et sp. indet., no. 32.1.2.70; C , cf. Salix sp., no. 32.1.2.39; D , Populus tremula L., foss., no. 32.1.3.7; E , Buxus pliocenica Saporta &Marion, no. 32.1.2.140; F , Rosaceae gen. et sp. indet., no. 32.1.2.55; G , Fabaceae type 1, no. 32.1.2.161; H, Fabaceae type 2, no.32.1.2.78; I-L , Acer pseudomonspessulanum Unger, nos. 32.1.2.61, 32.1.3.8, 32.1.2.46 and 32.1.2.67; M , Acer sp., no. 32.1.2.12;N-P , cf. Ziziphus ziziphoides (Unger) Weyland, nos. 32.1.2.3, 32.1.2.180 and 32.1.2.5. Scale bars: 1 cm.



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Zidianakis G.et al.

B. colchica -B. hyrcana group. According to theseauthors this group replaced the large-leavedBuxus lineage which became extinct because of the climaticdeterioration during the late Miocene. ExtantBuxussempervirens is a typical element of sclerophyllousvegetation in Mediterranean basin whileBuxushyrcanaPojark. and Buxus colchicaPojark. occur inpure stands or in underwood in lowland deciduousbroad-leaved mesophilous forests in the Hyrcanianregion (between the southern shores of the CaspianSea and the Elburz Mountains) and in the Colchis(Western Georgia).

FamilyR OSACEAE

Rosaceae gen. et sp. indet.(Figs 3F; 5G, S)


DESCRIPTIONOne complete and two fragmentary leaves with shortpetiole. Leaf length 2.7 to 3.5 cm (may be more)and width 1.3 to 2 cm. Shape of lamina ovate oroval, rounded at the base acute at the apex. Marginnely serrate with small, simple and sharp teeth.Primary vein well developed and straight. Higherorder veins not visible.

R EMARKSTe features observed in these fossils and espe-cially leaf margin suggest an affi nity to the familyRosaceae. oday this large family comprises about3300 species. It is cosmopolitan with maximumdiversity in the northern temperate area (Heywoodet al. 1993).

FamilyF ABACEAE

Fabaceae type 1(Fig. 3G)

M ATE RI AL EXAMINED. — Complete leaflet no.32.1.2.161.

DESCRIPTIONOne complete, very well preserved leaet; 3.3 cmlong and 1.5 cm wide, with oval outline. Margin oflamina entire, rounded at the apex and quite asym-metrical at the base. exture chartaceous, petiolulemissing. Venation brochidodromous. Primary veinthin, running irregularly (zig-zag) to the apex ofleaet. Five pairs of secondaries arising at an angleof 30 to 80° from the midvein, running more orless sinuous, joining each other by loops near themargin. Te lateral secondary at one leaet side isprominent, extending to the upper third of the mid-dle of the leaet. Intersecondary veins two or three,weakly developed. Higher order venation branchingout and joining the secondaries to form a networkof rather large, irregularly shaped areoles.

R EMARKSTis type of leaets is related to the subfamily Cae-salpinioideae and especially to the articial generaCassiophyllum andCaesalpinites by the asymmetri-cal form, the size and the venation pattern. Tesegenera occur in European deposits from the Oli-gocene to Miocene. Te determination of modernequivalents is diffi cult because the leaets of manyspecies have a similar morphology, as for instanceScohtia humboltioides Oliv.,Wagetia spicata Daezand someCassia species (Shakryl 1992).

Fabaceae type 2(Fig. 3H)

M ATERIAL EXAMINED. — Almost complete leaet no.32.1.2.78.

DESCRIPTIONOne leaet 3.8 cm long (complete about 4 cm) and1.5 cm wide, narrow elliptic to lanceolate, slightlyasymmetrical, entire-margined. Apex of lamina acute,base obtuse or rounded, asymmetrical (part of thebase is missing). Primary vein strong, S-like curved.Higher order venation ne, hardly visible.

R EMARKSEntire margins and asymmetry of this specimenare features that suggest a relationship with the



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Fabaceae. Unfortunately, poor preservation of theleaet does not allow a more precise determination.Fabaceae is a very large plant family (approximately17000 species) with great variety of habitat and acosmopolitan distribution in tropical, subtropicaland temperate zones (Heywoodet al. 1993). Tesmall-sized leaet of Vrysses may belong to a moreor less xeric element.

Family A CERACEAEGenus AcerL.

Acer pseudomonspessulanum Unger(Figs 3I-L; 5L, P, R)

M ATERIAL EXAMINED. — Complete leaves nos. 32.1.2.46;32.1.2.61; 32.1.2.67; 32.1.3.8; fragmentary leaves nos.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;32.1.3.9; 32.1.3.10.

DESCRIPTIONFour complete and 12 incomplete leaves, are assignedto this fossil species. Lamina deeply trilobate, 2.1to 4 cm (or somewhat more) long and 3 to 6.3 cmwide, with short petiole. Leaf texture coriaceous.Base of lamina rounded or cordate, margin entire.Te median lobe more or less broader than the lat-eral lobes. Apices of lobes acute to obtuse. Venationactinodromous with three primary veins, straightto slightly bent. Secondary veins ne, dense, aris-ing at an angle of 60 to 80° from the primaries,branching near the margin. Higher order veinspoorly preserved.

R EMARKS

Most of these fossil leaves (the larger specimens) aremost likely related to Acer monspessulanum L. butsome of them, with sharper sinuses and less broadlobes with acuminate apices which stretch upwards,are related to Acer sempervirens L. In fact, remains of Acer pseudomonspessulanum from Vrysses do not differin gross-morphology from the records in Makrilia,from where this leaf type was described as Acer seriesMonspessulana Pojark. Material from Vegora thatis also similar in its variability has been determinedas A. pseudomonspessulanum and A. integrilobum

Weber. Te name Acer pseudomonspessulanumis used as the Neogene ancestors of the modern spe-cies of the series Monspessulana Pojark., especially

A. monspessulanum and A. sempervirens and theirintermediate forms (Leroy & Roiron 1996). Leafand fruit remains of Acer pseudomonspessulanumareknown from several Neogene deposits in Europe.

oday Acer monspessulanum extends from central,south Europe and northwest Africa to north Iran(south shores of the Caspian Sea), while A. sem-

pervirens occurs at the east Mediterranean basin. Acer monspessulanum is a deciduous element while A. sempervirensis considered as deciduous or semi-evergreen element. Both of these maples are treesor shrubs and common elements of more or lessxerothermic mixed evergreen and deciduous forests.

oday Acer sempervirens grows on Crete at moun-tain slopes between 300 and 1700 m ( urlandetal . 1993).

Acer sp.(Figs 3M; 5K)

M ATERIAL EXAMINED. — Almost complete leaves nos.32.1.2.12; 32.1.2.56; fragmentary leaves nos. 32.1.2.14;32.1.2.49; 32.1.2.75.

DESCRIPTIONwo almost complete leaves and 3 fragments, 5-lobed,

3.1 cm to 3.9 cm (possibly more) long and 4.4 to5.8 cm wide. Margins entire with acuminate apex.Leaf base cordate, venation actinodromous withve primary veins. Lateral primaries arising underangles of 40 to 55° from the midvein, while basalprimaries arise under angles of 80 to 110°. Five toseven pairs of hardly visible secondary veins, ariseat 30 to 55° from primary veins and run in curvestowards the leaf margin.

R EMARKSSome of these specimens are comparable to Acer integerrinum, a common element of European Neogenedeposits. In Greece, A. integerrinum is representedin the late Miocene fossiliferous deposits of Licudi,central Greece (Velitzelos & Gregor 1990).

Acer laetum C.A.Mey. and Acer pictum Tunb.are considered to be the possible analogue among



364 GEODIVERSITAS • 2007 • 29 (3)

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extant maples on the basis of their gross-morphology(Leroy & Roiron 1996). Acer laetum thrives fromthe Caucasus to the Himalayas and the mountainsof central China while A. pictum grows on the slopesof northern China and Japan.

FamilyR HAMNACEAEGenus ZiziphusMill.

cf. Ziziphus ziziphoides (Unger) Weyland(Fig. 3N-P)

M ATERIAL EXAMINED. — Almost complete leaves nos.32.1.2.3; 32.1.2.5; 32.1.2.22; 32.1.2.180; fragmen-tary leaves nos. 32.1.2.10; 32.1.2.32; 32.1.2.106;32.1.2.209.

DESCRIPTIONFour almost complete leaves and four fragments.Lamina broad oval to ovate 3.2-5.8 cm long (maybemore) and 1.9-4.1 cm wide, simply dentate at theupper two thirds of the leaf margin. Apex acute toobtuse, base partly asymmetric. Venation acrodro-mous with three stout primary veins running inconvergent arches toward the leaf apex. Higher orderveins very ne, hardly visible, vertical to midvein.

R EMARKSTe specimens described above probably belong tothe species Ziziphus ziziphoides , a common elementin the European Paleogene and Neogene. Te genus Ziziphus consists of about 100 species of decidu-ous or evergreen trees and shrubs distributed inthe tropical and subtropical regions of the world(Johnston 1963). Tis leaf form may be related toa non-sclerophyllous species of Ziziphus not iden-tical with recent species of the Mediterranean areaand the Near East. Te nearest extant relatives of Ziziphus ziziphoidesconsidered to be the decidu-ous Z. sinensisand Z. glabrataboth thrive in Asia(Givulescu 1962). However, this leaf type may alsorepresent a completely different family (see Sachse1997;Dicotylophyllum type 3).

Family incertae sedis Genus Dicotylophyllum Saporta

Dicotylophyllum sp. 1(Fig. 4A)


DESCRIPTIONOne entire leaf, reniform, 2.4 cm long and about3.2 cm wide, stalked, entire-margined. Leaf textureevidently ne. Venation actinodromous with sevenprimary veins. Midvein straight, stronger than thelateral primaries. All primary veins forked beforereaching leaf margin. Secondary veins on the me-dial primary vein widely and irregularly spacedin four pairs, also forked. Higher order venationnot visible.

R EMARKS A very interesting leaf form. In morphology (ve-nation, margin, shape), it is comparable with thefoliage of modernCercis silliquastrumL. (Fabaceae),differing mainly by having smaller leaf size thanliving element.Cercis silliquastrum is a Mediter-ranean, deciduous tree.

Dicotylophyllum sp. 2(Figs 4B; 5H)

M ATERIAL EXAMINED. — Almost complete leaf no.32.1.2.92.

DESCRIPTIONOne almost complete (only a small part of its apexis missing) leaf, narrow oblanceolate 6.2 cm long(complete leaf somewhat more) and 2.2 cm wide.Lamina with entire margin, gradually narrowingtowards the base. Primary vein straight, very strongand thick. Leaf texture coriaceous. Secondary veinsne, hardly visible, arising at 30 to 50° from themidvein.

R EMARKSTis leaf with poorly preserved venation cannotbe reliably assigned to any specic genus. Tegeneral shape of this specimen suggests that itrepresented a broad-leaved humid or mesic ele-ment.



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A B C D

E

F

G

H

I L

J

K

FIG. 4. — Photographs of fossil leaves identi ed in Vrysses outcrops (western Crete): A , Dicotylophyllum sp. 1, no. 32.1.2.1; B , Dicotylophyllum sp. 2, no. 32.1.2.92; C , Dicotylophyllum sp. 3, no. 32.1.2.16; D , Dicotylophyllum sp. 4, no. 32.1.2.33; E , Dicotylophyllum sp. 5, no. 32.1.3.6;F , Dicotylophyllum sp. 6, no. 32.1.2.9; G , Dicotylophyllum sp. 8, no. 32.1.2.223; H, Dicotylophyllum sp. 9, no. 32.1.2.30; I, Dicotylo-

phyllum sp. 10, no. 32.1.2.93; J , Dicotylophyllum sp. 11, no. 32.1.2.159; K , Poaceae/Cyperaceae type 1, no. 32.1.2.191; L, Poaceae/ Cyperaceae type 2, no. 32.1.2.54. Scale bars: 1 cm.



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Dicotylophyllum sp. 3(Fig. 4C)


DESCRIPTIONOne complete leaf (only petiole missing), narrowlyelliptic, slightly asymmetrical leaf; 5 cm long and1.2 cm wide. Apex rounded, base obtuse, somewhatasymmetrical, leaf margin entire. Primary vein stout,S-like curved. Secondary veins ne, rather dense,hardly visible.

R EMARKS

It has not been possible to determine the affi nityof this leaf due to poor preservation.

Dicotylophyllum sp. 4(Figs 4D; 5O)

M ATERIAL EXAMINED. — Complete leaves nos. 32.1.2.27;32.1.2.33; 32.1.2.60; 32.1.2.144.

DESCRIPTION

Four complete leaves, small, long petiolate, oblan-ceolate to obovate leaves, 4 to 5.5 cm long and1 to 1.5 cm wide. Leaf apex obtuse or rounded,base attenuate and leaf margin entire. exture co-riaceous. Primary vein strong, more or less curved.No other details of the venation visible.

R EMARKSTe affi nity of these specimens is unknown andmuch better preserved material is necessary for areliable identication.

Dicotylophyllum sp. 5(Figs 4E; 5B, N)

M ATERIAL EXAMINED. — Complete leaves no. 32.1.3.4;32.1.3.6; fragmentary leaves nos. 32.1.2.34; 32.1.2.35.

DESCRIPTIONwo almost complete petiolate leaves. Length of the

lamina 2.5 to 3.9 cm and width 1.7 to 1.9 cm. Leaf

broadly lanceolate to obovate, apex acute to obtuseand base obtuse to slightly cordate. Margin slightlywavy or dentate with small simple teeth. Venationsemicraspedodromous. Primary vein prominent,somewhat straight. Secondary veins thin, denselyspaced at angles of 45 to 80°, running almost straight,parallel to each other across the lamina, dichotomisednear the margin before entering teeth. ertiary veinspercurrent or simply forked forming irregular mesheswith quaternary veins. Also two fragmentary leavesare assigned to this form.

R EMARKSTis fossil leaf taxon is similar in gross-morphol-ogy to numerous genera in different families suchas Rhamnaceae and Fagaceae.

Dicotylophyllum sp. 6(Figs 4F; 5I)

M ATERIAL EXAMINED. — Fragmentary leaves nos. 32.1.2.9;32.1.2.107.

DESCRIPTION

wo fragmentary leaves with broad obovate out-line, 5 to 5.7 cm long (perhaps more) and 2.2 to3.2 cm wide. Petiole rather slender, up to 1.2 cmlong. Base of lamina heart-shaped, apex absent,with undulate or nely, simply dentate margin.Midvein ne, hardly visible, straight or slightlybend. Higher order venation invisible. Leaf texturecoriaceous.

R EMARKSTe poor preservation makes it diffi cult to clarifythe systematic relationship of these specimens.

Dicotylophyllum sp. 7(Fig. 5 )


DESCRIPTIONOne complete, petiolate, slightly asymmetrical,broadly oval leaf with entire margin. Leaf length



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A B C D E F

G

H

I J

K

L

N

O

M

P

Q

R S

T

FIG. 5. — Drawings of selected fossil leaves from Vrysses (western Crete): A , Dicotylophyllum sp. 12, no. 32.1.2.42; B , N, Dicotylo- phyllum sp. 5, nos. 32.1.3.4 and 32.1.2.34; C , Dicotylophyllum sp. 10, no. 32.1.2.4; D , Daphnogene polymorpha (A.Braun) Ettingsh.,no. 32.1.2.26; E , Dicotylophyllum sp. 11, no. 32.1.2.98; F, cf. Juglans acuminata Braun ex Unger, no. 32.1.3.1; G , S , Rosaceae gen.et sp. indet., nos. 32.1.2.41 and 32.1.2.8; H , Dicotylophyllum sp. 2, no. 32.1.2.92; I, Dicotylophyllum sp. 6, no. 32.1.2.107; J , Quercus

mediterranea Unger, no. 32.1.2.147; K , Acer sp., no. 32.1.2.56; L, P , R , Acer pseudomonspessulanum Unger, nos. 32.1.3.9, 32.1.2.37and 32.1.2.31; M , Buxus pliocenica Saporta & Marion, no. 32.1.2.140; O , Dicotylophyllum sp. 4, no. 32.1.2.144; Q , Quercus sp., no.32.1.2.95; T, Dicotylophyllum sp. 7, no. 32.1.2.44. Scale bar: 1 cm.

5.9 cm, width 2.2 cm. Leaf base obtuse, apex round-ed. Venation probably semicraspedodromous. Pri-mary vein quite strong, S-like curved. Secondary

veins at least seven pairs, hardly visible, very deli-cate, rather widely spaced, arising at angles of 55to 80° from the midvein, running nearly straight



368 GEODIVERSITAS • 2007 • 29 (3)

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to somewhat bent towards the margin. Probablysecondaries forking near the margin. Further vena-tion pattern not preserved.

R EMARKSTe systematic affi nity of this leaf taxon remainsunclear.

Dicotylophyllum sp. 8(Fig. 4G)

M ATERIAL EXAMINED. — Almost complete leaf no.32.1.2.223.

DESCRIPTIONOne leaf almost complete (only a small part of apexmissing), broadly ovate, quite asymmetrical, 5.7 cmlong (complete leaf about 6 cm) and 4.3 cm wide. Apex of lamina acute or acuminate and base heart-shaped. Margin simply dentate with moderatelylarge and acute teeth, irregularly spaced. Venationactinodromous with ve primary veins. Primary veinrather stout, S-like. Basal primaries quite weak, aris-ing from the midvein at angles of 100 to 120° whilelateral primaries stronger, arising at angles of 60 to80°. Four pairs of curved secondary veins arising fromthe midvein, widely spaced, dichotomised near themargin. ertiary veins usually simple forked formingpolygonal meshes with quaternary veins.

R EMARKSCrucial characters are missing and thus it is im-possible to determine the systematic affi nity ofthis specimen.

Dicotylophyllum sp. 9(Fig. 4H)

M ATE RIAL EXAMINED. — Fragmentary leaf no.32.1.2.30.

DESCRIPTIONOne fragmentary leaf, broadly elliptic or obovate,2.6 cm long and 2.5 cm wide (entire leaf about

3.6 long and 3 cm wide). Leaf apex not preserved,base obtuse, margin hardly visible, undulate. Leaftexture coriaceous. Venation craspedodromous,primary vein stout, S-like curved. Secondaries atleast seven pairs, strong, parallel, regularly spaced,more or less straight, directed towards the margin.Higher order veins not visible.

R EMARKSSuch leaves may be related to the genusFagusL., but such affi nity cannot be proved. More and muchbetter preserved material is necessary to elucidatethe position of this specimen.

Dicotylophyllum sp. 10(Figs 4I; 5C)


DESCRIPTIONOne complete and two fragmentary leaves, peti-olate, with elliptic outline. Teir length is 5 to6.2 cm (maybe somewhat more) and their width1.7 to 1.9 cm. Leaf margin entire, acute at the apexand cuneate at the base. Primary vein prominentand straight. Secondary veins ne, hardly visible.Higher order venation not preserved.

R EMARKSTe systematic affi nity of these three specimensis unclear.

Dicotylophyllum sp. 11(Figs 4J; 5E)


DESCRIPTIONOne complete leaf, the largest in the Vrysses assem-blage, 17 cm long and 4 cm wide (size of laminamore than 4.500 mm2). Also two fragmentary leaves,smaller, up to 12 cm long and 2 to 2.5 cm wide.Leaf apex obtuse or acute, base cuneate, entire-



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margined, with broad elliptic outline. Primaryvein characteristically thick and strong, straightor slightly bent. Higher order venation not visibleprobably duo to thick coriaceous texture.

R EMARKSSuch leaf specimens are usually considered to be-long to magnoliid families, such as Magnoliaceaeor Lauraceae. However a relationship with otherfamilies cannot be excluded.

Dicotylophyllum sp. 12(Fig. 5A)


DESCRIPTION A small complete leaf, short petiolate, 3.2 cm longand 1 cm wide. Lamina narrow obovate, with apexrounded, base acute and entire margin. Primary veinquite strong and thick, S-like curved. Secondaryveins poorly preserved, arise from the midvein atangles of 30 to 50°. Leaf texture chartaceous.

R EMARKSTis leaf lacks diagnostic features.

Class MONOCO YLEDONAEFamiliesPOACEAE/C YPERACEAE

Poaceae/Cyperaceae type 1(Fig. 4K)

M ATERIAL EXAMINED. — Fragmentary leaves nos.32.1.2.57; 32.1.2.63; 32.1.2.151; 32.1.2.169; 32.1.2.186;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;32.1.2.228; 32.1.2.236; 32.1.2.245.

DESCRIPTIONSixteen fragments of narrow strap-like leaves about1 to 1.5 cm wide, that bear parallel venation.

R EMARKSTese fossil leaves are similar in gross-morphol-ogy to numerous genera mainly in Poaceae and

Cyperaceae. Te Poaceae and Cyperaceae are twolarge families of monocotyledons, cosmopolitan.Only few ecological plant formations lack Poaceae,while many formations are dominated by them(Heywoodet al. 1993).

Poaceae/Cyperaceae type 2(Fig. 4L)

M ATERIAL EXAMINED. — Fragmentary leaves nos.32.1.2.54; 32.1.2.198.

DESCRIPTION

wo fragments of broad linear leaves, wide, morethan 2 cm wide, with parallel venation.

R EMARKSTese specimens probably also represent leaves ofPoaceae or Cyperaceae.

FLORIS IC COMPOSI ION

Te Vrysses plant assemblage is documented byfoliage. More than 260 specimens of leaf mega-fossils, in many cases very fragmentary, were ex-amined. A total of 31 taxa of vascular plants havebeen described based on their macro morpho-logical features ( able 1). Nineteen of them wererelatively well preserved and identied at least tofamily level. wo genera of conifers and 29 taxa ofangiosperms, shrubs, trees and, rarely herbs havebeen documented.

Conifers are represented in the assemblage byTetraclinis andPinus shoots and leaves. Fertileparts, such as cones, cone scales or seeds were notfound. Angiosperm taxa make up approximately 95%

of the total ora. Among the angiosperms, Acer-aceae and Fagaceae are most common. Fagaceae isrepresented by the genusQuercus , mainly by theprobably evergreen oakQuercus mediterranea , whichpredominates andQuercus sp. considered also tobelong to evergreen oaks. wo leaf-types belong to Aceraceae; a small 3-lobed leaf identied as Acer pseudomonspessulanum, considered to be more or



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Family Genus Species/ type series

Leaves/ needles shoots

Leaf mar-gin entire

Maximum leafsize class

D, L, S a Dispersalmechanism

GYMNOSPERMAEPinaceae Pinus sp. 17 needles - - - windCupressaceae Tetraclinis sp. 4 twig fragments - - - wind

ANGIOSPERMAEDicotyledonae

Lauraceae Daphnogene polymorpha 2 yes microphyll L endozoochoryHamamel-idaceae

?“ Parrotia ” pristina 1 no microphyll D autochory

Juglandaceae ? Juglans acuminata 3 lea ets yes notophyll D dyschoryFagaceae Quercus mediterranea 12 no microphyll S dyschory

Quercus sp. 3 no microphyll S dyschoryUlmaceae gen. indet. - 1 no nanophyll D/S windSalicaceae ? Salix sp. 1 no microphyll D wind

Populus tremula , foss. 2 no microphyll D windBuxaceae Buxus pliocenica 1 yes microphyll L/S autochory/

myrmecochoryRosaceae gen. indet. - 3 no microphyll D wind/

exozoo-/endo-zoochory

Fabaceae gen. indet. type 1 1 lea et yes microphyll D/S -gen. indet. type 2 1 lea et yes microphyll D/S -

Aceraceae Acer pseudomonspessulanum

16 no microphyll D/S wind

Acer sp. 5 no microphyll D windRhamnaceae ? Ziziphus ziziphoides 8 no microphyll D endozoochoryFamily incertae

sedisDicotylophyllum sp. 1 1 yes microphyll D/L/S -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 -

MonocotyledonaePoaceae/ Cyperaceae

gen. indet. type 1 16 - - - wind/ nautochory/ exozoo-/ endozoochory

gen. indet. type 2 2 - - - wind/ nautochory/ exozoo-/ endozoochory

T ABLE 1. — Summary of the taxa of the Vrysses ora (western Crete), their dispersal mechanisms and physiognomic signatures. Note: a ,

taxa which cannot be restricted to one single category, are included in all assigned categories. Percentage of entire-margined taxa: 48.1%(13 taxa). Percentages of notophyll, microphill and nanophyll taxa: notophyll 7.4% (2 taxa), microphyll 85.2% (23 taxa) and nanophyll 7.4%(2 taxa). Percentages related to deciduous, laurophyllous and sclerophyllous taxa: deciduous 37-59% (10-16 taxa), laurophyllous11-30% (3-8 taxa) and sclerophyllous 15-52% (4-14 taxa). Abbreviations: D, deciduous; L, laurophyllous; S , sclerophyllous.

less xeromorphic and a 5-lobed leaf represents mostlikely a mesophytic deciduous species.

Fabaceae are represented by a few leaets that havebeen subdivided into two morphotypes (Fabaceaetype 1 and type 2). Juglandaceae seem to be alsopresent. Te leaets of the Vrysses plant assemblagemay belong to Juglans acuminata . Te family Rham-

naceae may be represented by irregularly dentate,3-veined leaves that are close in appearance to theNeogene species Ziziphus ziziphoides . Leaves ofLauraceae seem to have been present in the assem-blage as well by the narrow leaf type ofDaphnogene polymorpha . Imprints of Populus tremula , foss.,Salix sp. and possibly “Parrotia ” pristina occur rarely. One



371



leaf ofBuxus sp. has been identied. Rosaceae andUlmaceae may have been present as well. Mono-cots are represented by numerous specimens of thefamilies Poaceae and Cyperaceae. However manyleaf impressions of the Vrysses plant assemblagehave doubtful affi nities. Tey are considered tobelong to 12 taxa of angiosperms, and classied asDicotylophyllum.

Considering the number of specimens, four woodytaxa are most common in the assemblage:Pinussp., Acer pseudomonspessulanum, Quercus mediterranea and Ziziphus ziziphoides ( able 1).Tetraclinissp.and Acer 5-lobed are found also more frequently.More than one species may have existed among Aceraceae, Fagaceae Salicaceae and Fabaceae (alltwo taxa).

Te majority of the genera identied at Vryssesare related to elements now growing in the North-ern Hemisphere. Te modern living relatives ofBuxus pliocenica , Populus tremula , foss., Acer pseudo-monspessulanum and Juglans acuminata thrives inEurasia whileDaphnogene polymorpha has its clos-est counterparts in Asia and America and Ziziphusziziphoides in Asia. A few taxa have affi nities toliving Mediterranean species such asetraclinis and Quercus mediterranea while extantParrotia is a relicfound in Iran (Zohary 1947; Wang 1961; Zohary& Orshan 1966; Meuselet al. 1965, 1978; Strahler& Strahler 1992; Achhalet al. 1980; Quezel 1988;Barberoet al.1992; urlandet al. 1993; Arabatzis1998, 2001). Te core of the nearest living relativesof the fossil taxa grows today in the temperate tosubtropical zone, whereas only a few taxa extendto the tropical zone (Lauraceae) (Walter & Lieth1967; Browics 1982; Klötzli 1988).

APHONOMY AND VEGE A ION ANALYSIS

Te Vrysses Formation was most likely deposited in ashallow, semi-enclosed marine embayment, borderedby islands and shoals (Meulenkampet al. 1979).Te fossil assemblage contains autochthonous and alarge proportion of allochthonous elements. Tis isconrmed mainly by the composition of the taphoc-oenosis that consists of a mixture of plant remains

from different habitats and the high percentage ofsmall-sized leaves (Ferguson 1985; Spicer 1991).More than 85% of the taxa identied in Vryssesoutcrops are classied as microphyll (225-2.025mm2), while 7.4% as nanophyll (25-225 mm2)( able 1). Te allochthonous nature of the assem-blage is also reected by the absence of differentorgans, such as seeds and fruits of the same taxonin the assemblage and by the high percentage ofheavily fragmented leaves that reach 64% of theexamined specimens.

Plant debris have probably been transported byriver oods into their offshore burial site. Alterna-tively, plant remains may have drifted on the seasurface and reached the locality of sedimentationafter having been blown by strong winds from highcoastal cliffs. According to its composition, the Vrysses plant

assemblage is a heterogeneous mixture of higherplants with different ecology. Considering thespecic features of the modern associations (Wolfe1979; Klötzli 1988; Song 1983, 1988) that canbe related to the Vrysses fossil leaves, the outlineof a regional vegetation reconstruction has beendeveloped, that is characterized by a oristic dif-ferentiation based on water effi ciency and altitude(Gastaldo & Ferguson 1998).– a) Riparian paleocoenoses composed of representa-tives of the wood elementsSalix sp. and perhapsPopulus tremula and Juglandaceae, as well as Poaceaeand Cyperaceae herbs. Tese elements were forminga belt of a riparian forest along the banks of inow-ing streams. Subtropical, partly evergreen elements,such asDaphnogene polymorphamay have playedalso a certain role in these lowland forests.– b) Sclerophyll forest paleocoenoses consistedin general of microphyllous, broad-leaved taxa(such asQuercus mediterranea , Quercussp., possi-blyBuxus pliocenica , Acer pseudomonspessulanum, Ulmaceae and several Fabaceae) and conifers (Pi-nus , possibly etraclinis ) forming woodland at drylowland areas.– c) Mixed-Mesophytic forest coenoses. Sclerophyl-lous evergreen forests of lowlands were graduallyreplaced by mixed mesophytic forests of higher eleva-tions along an altitude gradient. Te canopy of theseforests was dominated by deciduous representatives



372 GEODIVERSITAS • 2007 • 29 (3)

Zidianakis G. et al.

M e a n a n n u a

l t e m p e r a

t u r e

( ° C )

Percentage of entire-margined taxa

30

20

20 40 60 80

10

00

FIG. 6. — Correlation of percentage of species with entire-mar-gined leaves in the humid and mesic broad-leaved forests ofeastern Asia with mean annual temperature, after Wolfe (1979).--- , percentage of entire-margined taxa in the Vrysses (western

Crete) plant assemblage.

of Acer5-lobed, Populus tremula , Ziziphus ziziphoides , Juglandaceae and probably Acer pseudomonspessula-num and Ulmaceae. In the stratum of lower treesand shrubs a mixture of evergreen (Buxus pliocenicaand Daphnogene polymorpha ), as well as deciduouselements (“Parrotia ” pristina , Rosaceae) and conifers(Tetraclinis sp.) seem to have existed.

It is also interesting to consider dispersal mecha-nisms of the recorded taxa. About 23% of theidentied plant taxa were dispersed either endo- orexozoochorously by birds or mammals (see able 1)that implies also a diverse paleofauna.

PALEOCLIMA E RECONS RUC ION

Analyses of Neogene plant assemblages are in gen-eral a very useful tool to extract proxy data forpaleoclimatic reconstructions. We tried to applytwo different methods in order to estimate climaticparameters of the broader Vrysses area during thelate Miocene: the leaf margin analysis and the co-existence approach. However because of the lim-ited number of taxa and the unresolved taxonomicproblems due to poor preservation of the materialonly rough estimations were possible.– a) Leaf margin analysis. Te proportion of taxa

with entire-margined species in the Vrysses plantassemblage reaches 48% ( able 1), suggesting amean annual temperature of 15-16°C (Zidianakiset al. 2004) (Fig. 6). Te relatively high percentage

of sclerophyllous elements (about 15-52%) reducesthe accuracy of this correlation.– b) Co-existence approach. able 2 shows themodern taxon analog of the species recognized inthe Vrysses plant assemblage and their mean annualtemperature requirements. A range of 15-16°C formean annual temperature is also estimated.

Several of the Vrysses taxa seem to have theirnatural habitat today in the Euxino-Hyrcanian Prov-ince. Tese taxa comprise Parrotia , Buxus , Juglansand Acer . Te average annual temperature of theHyrcanian region varies from 15°C in the west to17.5°C in the east. Te temperature of the warmestmonth ranges between 28-35°C while that of thecoldest month ranges from 1.5 to 4°C.

Rainfall varies substantially from 530 mm inthe east to 1350 mm in the west. In most areasof this region precipitation is all year round withmaxima in spring, late fall and winter, the easternparts experience three months of summer drought(see www.rifr-ac.ir/index_eng.htm). If we take thisregion as model for the Neogene climate, a range of15-17.5°C for mean annual temperature (MA ),and more than 800 mm for mean annual precipita-tion (MAP) is estimated.

Te average small leaf size of the assemblage(microphyllous ora) may be not only caused bytaphonomic processes but also by climatic factors,such as periodic drought. Te relatively high pro-portion of sclerophyllous xeromorphic elementsin the plant assemblage may conrm the latterhypothesis.

On the basis of physiognomic features, leaf re-mains of Vrysses are divided in three main cat-egories: 1) a broad-leaved deciduous component

which predominates with approximately 37-59%;2) broad-leaved evergreens (laurophyllous taxa) reachabout 11-30%; and 3) the sclerophyllous compo-nent with the relatively high proportion of 15-52%( able 1). Te broad-leaved deciduous elementsare characteristic for humid, temperate climaticconditions, while the broad-leaved evergreen taxafor humid, subtropical and sclerophyllous taxa forsub-humid, subtropical conditions (Berger 1954;Kovar-Ederet al. 2006).

Tus, the Vrysses plant assemblage may indicatea warm-temperate, humid climate for the late

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http://www.rifr-ac.ir/index_eng.htm



373



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 10.10-17.30Quercus mediterranea Quercus ilex , Quercus coccifera 11.70-19.50Populus tremula , foss. Populus tremula 1.60-19.00Buxus pliocenica Buxus sempervirens 9.70-17.30

Acer pseudomonspessulamum Acer pseudomonspessulanum , Acer sempervirens

9.60-20.50

Ziziphus ziziphoides Ziziphus sinensis 7.30-27.60

T ABLE 2. — Mean annual temperature (MAT) requirements for nearest living relatives (NLR) of taxa of the Vrysses (western Crete) as-semblage. Data are taken from Palaeo ora Database (Utescher & Mosbrugger 1999).

Miocene. Te existence of a dry period during theyear is probable, although this seasonality was clearlynot yet as pronounced as today. Using the climaticclassication of Köppen (1931), the Vrysses paleo-climate is most likely ranging between Cfa (warmtemperate with warm, humid summers) and Csa(warm temperate with warm, arid summers) cli-mates. Te climatic change from pre-late Miocenesummer wet climates to summer dry climates waspossibly caused by the successive westward retreat ofthe Paratethys between the Oligocene and the mid-dle Miocene (Popovet al. 2004) and the changingtopography of eastern Asia (Sachse 1997).

DISCUSSION

Te discovery of a new Miocene leaf assemblage inthe eastern Mediterranean realm offers the opportu-nity for a better understanding of the oristic andclimatic development of this region. Neogene orasoccur in central and northern Greece, as well as inseveral Aegean islands including Crete (Knobloch &Velitzelos 1987; Velitzelos & Gregor 1990; Sachse& Mohr 1996, Velitzeloset al. 2001). If the Vryssesplant assemblage is compared with the known Greekplant assemblages on the basis of physiognomy andoristic composition, the closest match seems tobe with the late Miocene assemblage of Makrilia,eastern Crete (Sachse 1997, 2004).

Te plant macro-remains of Makrilia occur insemi-pelagic silty layers and represent a plant assem-blage of which 94% of the leaf taxa are microphyllor nanophyll (Sachse 1997). Te age of the Makrilia

ora was determined using nannoplankton anddinoagellate cysts as late Miocene (7.7-8.6 Ma;Sachse & Mohr 1996) and is thus slightly older thanthe Vrysses ora. As it was expected, the Makriliaassemblage shares many arboreal taxa with theVrysses ora. More than 70% of the taxa identiedin Vrysses exist also in Makrilia. Subtropical lauro-phyllous elements, such asDaphnogene polymorpha ,deciduous elements, such as Acer 5-lobed,Populus , Salix and Ziziphus , conifers, such asetraclinis , Pinus and xeromorphic taxa, such asQuercus mediterranea ,

Acer pseudomonspessulanum and Buxus pliocenica arecommon in both oras. However, the Makrilia as-semblage is more diverse than the one from Vrysses.Considering the composition of the Makrilia orait clearly contains relatively more deciduous taxathan the Vrysses plant assemblage. However, it isnot clear whether this is a real signal or the result oftaphonomic processes or in the nearby hinterlandof the Vrysses area oras of high elevations mighthave simply not existed.

Te late Miocene oras of continental Greece(Vegora, Likudi, Prosilion, Lava) and other islands(Chios, Kerkyra) consist mainly of conifers anddeciduous broad-leaved elements, such asGinkgo,

axodiaceae, Pinaceae,Sassafras , Alnus , Fagus , de-ciduousQuercus , Populus , Zelkova , Ulmus and

Acer(Velitzelos et al. 2001). Obviously, not onlyon Crete but in entire Greece, the exotic elementsof the early Miocene were gradually replaced bydeciduous broad-leaved trees and shrubs, formingforests that resembled modern mixed mesophyticwoods of Eastern Asia, the Colchis and the Caspianarea. Tis change most likely indicates a decrease



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

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