Revision of Tertiary Fagus cupules from Russia, Transcaucasia and western Siberia

Feddes Repertorium 113 (2002) 3–4 , 193–210 Berlin, August 2002

© WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim, 2002 0014-8962/02/3-408-0193 $ 17.50+.50/0

Swedish Museum of Natural History, Department of Palaeobotany, Stockholm

TH. DENK

Revision of Tertiary Fagus cupules from Russia, Transcaucasiaand western Siberia

With one Figure, 3 Tables and 7 Plates

Summary

Tertiary cupule and nut remains of Fagus fromTranscaucasia, Russia and western Siberia whichhave been ascribed to various modern species, arerevised and compared to European Tertiary cupulesand nuts. Specimens, previously ascribed to Faguscf. orientalis and F. cf. grandifolia from the Plio-cene and Late Miocene of western Transcaucasia,belong to the same morphotype. They cannot bedistinguished from Late Miocene and Pliocenecupules from Europe referred to as Fagus deucali-onis UNGER emend. DENK & MELLER. Middle Mio-cene to Late Oligocene cupules from western Sibe-ria ascribed to F. cf. grandifolia, F. cf. japonica andF. cf. longipetiolata belong to a second morphotype,and also clearly fall within the morphological vari-ability encountered in cupule and nut remains ofF. deucalionis of the same age from Europe. Thefossils ascribed to F. cf. lucida remain enigmaticalthough they differ from extant F. lucida by havingwinged nuts and spine-like appendages. In general,diagnostic characteristics of cupules and nuts inFagus are easily lost during transport and fossiliza-tion, which makes fossile cupules and nuts difficultto assign to distinct taxa and to compare to singlemodern species.

Zusammenfassung

Revision tertiärer Fagus-Kupulen aus Russ-land, Transkaukasien und Westsibirien

Früchte und Kupulen von Fagus aus dem TertiärTranskaukasiens, Russlands und Westsibiriens, diezu verschiedenen modernen Buchenarten gestelltwerden, sind Gegenstand einer Revision und werdenmit Früchten und Kupulen aus dem Tertiär Europasverglichen. Fossilien aus pliozänen und obermiozä-nen Ablagerungen West-Transkaukasiens, ursprüng-lich zu Fagus cf. orientalis und F. cf. grandifoliagestellt, gehören zu einem Morphotyp (MT1). Siesind identisch mit spätmiozänen und pliozänenKupulen und Früchten Europas (F. deucalionisUNGER emend. DENK & MELLER). Mittelmiozäneund spätpliozäne Kupulen aus Westsibirien, welchezu F. cf. grandifolia, F. cf. japonica und F. cf. lon-gipetiolata gestellt wurden, entsprechen einemzweiten Morphotyp (MT2), der ebenso in die mor-phologische Variabilität von F. deucalionis fällt.Kupulen und Früchte, welche als F. cf. lucida be-stimmt wurden, weichen etwas von MT1 ab, unter-scheiden sich aber von der modernen Art F. lucidadurch geflügelte Früchte und borstenartige Kupula-Anhängsel. Im Allgemeinen sind diagnostischeMerkmale von Kupulen und Früchten bei Fagussehr unvollständig erhalten, da sie bei Transport undEinbettung im Sediment leicht verloren gehen. Ausdiesem Grund ist es problematisch, fossile Kupulenund Früchte von Fagus verschiedenen Arten zuzu-schreiben, bzw. sie mit rezenten Arten zu verglei-chen.

Introduction

The genus Fagus (beech, Fagaceae) is a rela-tively small genus comprising 8–13 tree spe-cies displaying a northern hemispheric dis-

junct distribution in North America and Mex-ico, Europe and south-western Asia, and EastAsia (SHEN 1992). Beech trees are economi-cally important as timber (PETERS 1997). ForTertiary palaeontology Fagus is a key taxon,

194 Feddes Repert., Berlin 113 (2002) 3–4

indicating fossil plant assemblages to have beenlinked to various temperate climates (Cfa cli-mate sensu KÖPPEN; CAO 1995). Fagus madeits first appearance in the Early Tertiary in thenorthern Pacific region (FOTJANOVA 1982).Earliest leaves probably belonging to the genuswere reported from the Eocene of Kamchatka(FOTJANOVA 1982; BUDANTSEV 1997), where-as earliest cupules and nuts unambiguouslyassignable to Fagus are from the Early Oligo-cene of western North America (MEYER &MANCHESTER 1997). From the northern PacificBasin Fagus appears to have spread westwardsalong the Parathetyan coast to Central Asia andEurope, as well as via western North Americato Mexico and eastern North America.

Cupules and nuts from the Tertiary havebeen referred to several species (UNGER 1847;REID & REID 1915; MÄDLER 1939; DOROFEEV1982; UEMURA 1982). In his treatment of Ter-tiary cupules in “Magnoliophyta FossiliaUSSR” DOROFEEV (1982) assigned cupulesand nuts originating from Oligocene to Plio-cene deposits in Russia and Asia to five mod-ern species. Specimens from the Late Oligo-cene of western Siberia were ascribed to F. cf.longipetiolata (at present in China), and speci-mens from the Miocene to F. cf. japonica (Ja-

pan) and F. cf. lucida (China). Middle and LateMiocene specimens from western Siberia,Caucasia, and the European part of Russiawere ascribed to F. cf. grandifolia (at presentconfined to Mexico and eastern North Amer-ica). Finally, F. cf. orientalis was recorded forthe Pliocene of Transcaucasia, leading to mod-ern F. orientalis (F. sylvatica subsp. orientalis)native of this region (DOROFEEV 1982). In arecent paper DENK & MELLER (2001) havedemonstrated that various characteristics pre-viously used to dinstinguish between fossilFagus species based on cupules and nuts oftenonly represent different stages of decay ratherthan biologically significant characters thatcould be used for species delimitation. Giventhe few characteristics preserved, EuropeanTertiary Fagus species based on cupules andnuts combine features of several modern spe-cies, and therefore cannot be compared to par-ticular modern species (DENK & MELLER2001).

Here I revised cupules and nuts described in“Magnoliophyta Fossilia USSR” and estab-lished, whether comparisons to particular mod-ern species are possible or not. In addition, thephylogenetic significance of the studies cu-pules and nuts is discussed.

Table 1Origin and revised taxonomy of cupules examined for this study, and Fagus species based on leaves that arepossibly linked to cupule/nut species

Revised nomenclaturethis article

TaxonDOROFEEV 1982

Occurrence

F. deucalionis emend.aMT1b (Pl. II)

Fagus cf. orientalis K531 Duab, AbkhasiaEarly Pliocene

F. deucalionis emend.MT1 (Pl. III)

Fagus cf. grandifolia K530 Apsheronsk, Caucasiaearly Late Miocene

F. deucalionis emend.MT1 (Pl. III)

Fagus cf. grandifolia K412 Wolnaya Vershina, Russia (Europe)Middle to Late Miocene

F. deucalionis emend.MT2 (Pl. V)

Fagus cf. grandifolia K502 Romanovo, Western SiberiaMiddle Miocene

F. deucalionis emend.(Pl. III, IV)

Fagus cf. lucida K517 Kireevskoye, Western SiberiaMiocene

F. deucalionis emend.MT2 (Pl. VI)

Fagus cf. japonica K467 Polevskoy, Western SiberiaMiocene

F. deucalionis emend.MT2 (Pl. VII)

Fagus cf. longipetiolata K509 Kosyulina, Western SiberiaLate Oligocene

a DENK & MELLER (2001), b MT = Morphotype

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Materials and methods

All specimens examined for the present studyare housed in the palaeobotanical collectionsof the I. V. Komarov Institute of the RussianAcademy of Sciences in St. Petersburg (BIN).Collection numbers, origin and age of the cu-pules and nuts are given in Table 1. Map refer-ences in Table 1 refer to TAKHTAJAN (1982).For comparision material of all modern speciesof Fagus as well as cupules and nuts from theTertiary of Europe and North America havebeen studied (DENK & MELLER 2001). Formodern Fagus species the taxonomic conceptof SHEN (1992) is followed (Table 2).

Results

Degradation of modern cupules throughtransportation: a modern analoguefor Tertiary cupules

Observations of water-transported cupules ofthe modern species Fagus sylvatica along theeastern Black Sea coast showed several typesof morphological decay of outer and innervalve surfaces (for details see DENK & MELLER2001). A series of types of degradation of the

outer cupule surface is presented in Plate I.Because very similar patterns of alteration canbe observed in fossil cupules, the systematicsignificance of these patterns, i.e.“decurrent”versus “non-decurrent” bases of cupule ap-pendages, is suggested to be low.

Systematics

Fagus cf. orientalis (K531)

(revised Fagus deucalionis UNGER emend.DENK & MELLER, Morphotype 1)

Of 42 specimens, 14 are more or less completecupules, 24 are single valves or heavily de-cayed cupules, and four are nutlets. Valves are7–11 mm long and 3–7 mm wide. The pedun-cle is thickened at the point of insertion into thecupule; it either merges into the cupule, orabruptly inserts into it. Its diameter is up to3 mm at the point of insertion, and 1–2 mm atits most proximal preserved part. Sometimesthe peduncle is quite flattened due to preserva-tion; it is always broken. The number of ap-pendages per cupule valve is not countable dueto abrasion of parts of the outer cupule surface.The appendages are always spine-like and

Table 1 (continued)

Map reference TAKHTAJAN 1982 Species based on leaves possibly linked to cupule/nut remainsTAKHTAJAN 1982; Locality, Age, Map, Reference

no. 79 (map 2) F. herthae (UNGER) ILJINSK.no. 15 (map 2) Pizunda, Pont, 227/2; Barmush, Sarmatian 23/2; Armavir, Late

Sarmatian, 17/2; Nakhichevanj, Late Sarmatian, 201/2

no. 53 (map 1) F. herthae (UNGER) ILJINSK.Gryaznushka, 66/1; Gurovo 68/1, Middle to Late MioceneF. juliae JAKUB., Kammenij Brod, Middle to Late Miocene, 104/1

no. 246 (map 3) –

no. 128 (map 3) –

no. 228 (map 3) F. antipofii HEERBerekmak, Oligocene, 30/3; Kinzebulatovo, Oligocene, 125/3

no. 135 (map 3) F. antipofii HEERRhezenka, Oligocene, 240/3

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Table 2Recent classification of Fagus by SHEN (1992)

Fagus engleriana SEEMEN(incl. F. multinervis NAKAI)

China, South Korea

Fagus japonica MAXIM. JapanFagus okamotoi SHEN Japan

Fagus longipetiolata SEEMEN(incl. F. brevipetiolata HU)

China, Vietnam

Fagus bijiensis C.F. WEI & Y.T. CHANG* Western ChinaFagus tientaiensis a T.N. LIOU* Eastern ChinaFagus lucida REHDER & WILSON ChinaFagus chienii CHENG* Western ChinaFagus hayatae PALIBIN China mainland, TaiwanFagus crenata BLUME JapanFagus sylvativa L.(incl. F. orientalis LIPSKY, F. moesiaca (MALY) CZECZ.)

Europe, south-western Asia

Fagus grandifolia EHRH.(incl. F. mexicana MARTINEZ)

Eastern North America, Mexico

a Fagus subgenus Engleriana, b Fagus subgenus Fagus, c Fagus longipetiolata species aggregate,* based on one collection of a single tree

display different stages of preservation [bestcomparable to steps 4, 5, (8), 9, and 10 inPlate I]. They are broken and rather fragile,about 0.2 mm or less in cross section, oftenflattened and then forming triangle-shapedsharp ridges on the outer cupule surface(Plate II, Figs. 6, 7, 9–12). In rare cases theymay be stout, >0.2 mm in cross section. De-pending on the type of preservation, the outercupule surface may be ridged or not. Heavilydecayed cupules display smooth outer sur-faces (Plate II, Figs.1–3). The length of nuts is7–10 mm. Nutletes may be winged in theirupper parts (Plate II, Fig. 16) or unwinged(Plate II, Fig. 15).

Remarks : Based on the few systematicallyuseful characteristics preserved, this taxon combinesfeatures of serveral modern species, i.e., F. crenata,F. grandifolia, and F. hayatae, but is not assignableto any particular extant species. Clearly, it is notcomparable to the modern F. orientalis (= F. sylva-tica subsp. orientalis), which displays cupule sizesof (10–) 15–30 (–37) mm (DENK & MELLER 2001;Table 3). The name Fagus deucalionis UNGERemend. DENK & MELLER should be used for thesespecimens, referring to an organ species that isconspicuously homogeneous through the Tertiary ofEurasia, but may have belonged to more than onegenetically different species.

Fagus cf. grandifolia (K530)

(revised Fagus deucalionis, Morphotype 1)

Of 18 species one is a complete cupule, five arecupules with more or less fragmentary valves,11 are two connected valves or single valves,whereas one is a fragmented nut. Valves are6–11 mm long and 3–5 mm wide. In onespecimen (Plate III, Fig. 1) the preserved partof the peduncle is 4.5 mm long. The insertionof the peduncle into the cupule is heart-shapedin most cases, the peduncle abruptly insertinginto the cupule. The peduncle is thickened atthe point of insertion (Plate III, Figs. 1, 2). Ingeneral, the number of appendages is not count-able, but exceeds 25 per valve (Plate III, Fig. 3).The appendages are spine-like, broken. Theydisplay different kinds of preservation [bestcomparable to steps 4, 5, (8), 9, and 10 in PlateI], and may be fragile or stout (0.2–0.4 mm indiameter). The outer cupule surface may as-sume a strongly ridged appearance, often dis-playing sharp ridges (Plate III. Figs. 3–6), butmay also be almost smooth (Plate III, Fig. 2).Accordingly, the bases of appendages mayappear decurrent as well as non-decurrent. Asingle broken nut is more than 7 mm long, andappears to have been winged in its upper parts.

a

b

c

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Remarks : The specimens show similarities tomodern F. crenata, F. grandifolia and F. hayatae(Table 3). They cannot be ascribed to a particularmodern species, and should be referred to as F. deu-calionis.



Of three specimens, two are cupules and one isa single valve. The valves are 8–11 mm longand 4–5.5 mm wide. No peduncle is preserved,but the insertion of the peduncle into the cu-pule is heart-shaped (Plate III, Fig. 7). Thenumber of appendages per cupule valve ismore than 30. The appendages are alwaysbroken and display various kinds of preserva-tion [best comparable to steps (5), 7 and 9 inPlate I]. They are spine-like, fragile or stout.The outer cupule surface is slightly ridged orwith wart-like protuberances. Bases of appen-dages are not decurrent.

Remarks : The specimens are comparable toseveral modern species, i.e., F. crenata, F. grandi-folia, and F. hayatae. They fall well within thevariability of Fagus deucalionis.

Fagus cf. lucida (K517)

(revised Fagus deucalionis, aff. Morphotype 1)

Of eight specimens, three are more less com-plete cupules, four are valves, and one is afragmented nut. Valves are up to 9 mm longand to 5.5 mm wide. The peduncle is thickenedwhen merging into the cupule (Plate III,Figs. 8, 9). Its diameter is up to 2.5 mm at thepoint of insertion, and 1.0 to >1.0 mm at itsmost proximal preserved part. All pedunclesare broken. The number of appendages percupule valve is >35 when countable (Plate IV,Figs. 8, 9). Appendages are spine-like. Stagesof decay correspond to steps (3, 4), 5, 7, 9, and10 (Plate I). Depending on the degree of decay,appendages are slightly bending outwards(Plate III, Fig. 10) or appressed to the cupulesurface, they are always broken. Outer cupulesurfaces display slightly decurrent bases ofappendages (Plate III, Fig. 10; Plate IV,Figs. 8, 9). The two nuts preserved are around10 mm long, and thus slightly longer than thevalves. One appears to be winged in its upperparts, whereas the other one is wingless. Onecupule contains nuts (Plate IV, Figs. 1–7). The

angles of the nuts are badly preserved (Plate IV,Fig. 5) but they appear to have been winged intheir upper parts (Plate IV, Figs. 3, 7). The nutsare slightly longer than the cupules valves.

Remarks : DOROFEEV (1982) considered thebadly preserved outer cupule surface of these speci-mens most similar to Fagus lucida. The valves ofthe few available specimens, however, displayspine-like appendages, and not scale-like ones, asfound in F. lucida. Moreover, the nuts are (partly)winged in the fossil specimens, whereas F. lucidanever has winged nuts. In principal, these specimensare comparable to several modern species, namely toF. hayatae and F. grandifolia. They should be in-cluded within F. deucalionis.



Of four specimens, two are cupules and two aresingle valves. The valves are 10–14 mm longand 4.5–6.5 mm wide. No peduncle is pre-served, but the point of insertion of the pedun-cle into the cupule is heart-shaped (Plate V,Figs. 5–8) The number of appendages percupule valve is more than 30. Appendages arespine-like and display different stages of pres-ervation (best comparable to steps 5 and 7in Plate I). Appendages are fragile (0.15–0.25 mm in diameter) and sometimes bend-ing outwards (Plate V, Fig. 9) or stout(>0.3 mm in diameter) and appressed (Plate V,Figs. 1–4, 7). Bases of appendages form ridgeson the outer cupule surface, which makes them“decurrent”. A single specimen contains nuts,which are as long as the cupule, i.e., 11 mmlong (Plate V, Figs. 7–9). They are winged intheir upper parts (Plate V, Fig 9).

Remarks : These specimens are larger thanthose of the preceeding samples. They are bestcomparable to F. tientaiensis (F. longipetiolataspecies aggregate), F. crenata, and F. grandifolia(Table 3). They should be included within Fagusdeucalionis.

Fagus cf. japonica (K467)


Of 21 specimens, four are complete cupules,five are fragmented cupules, seven are singlevalves, and five are nuts. Valves are 9–14 mmlong and 4–7 mm wide. The peduncle isslightly thickened when inserting into the cu-


pule, but clearly distinct from the cupule itself(not merging into it; Plate VI, Figs. 6–8). Itsdiameter is up to 3 mm at the point of insertion,and 1.0–1.5 mm at its most proximal preservedpart. Sometimes the peduncle is quite flatteneddue to preservation. It is almost broken. Thenumber of appendages per cupule valve is>25–>50. They are spine-like and displaydifferent stages of preservation (steps 2, 4–9 inPlate I). They are bending outwards and bro-ken. Their diameter may be roundish, about0.2–0.25 mm in cross section, to stronglyflattened. Depending on the type of preserva-tion the outer cupule surface may be ridged ornot. In addition, some valves display wart-likeprotuberances. In one specimen the innercupule surface is conspicuously ribbed. Thelength of the nuts is 9.0–12.5 m (Plate VI,Figs. 9–12). Their shape resembles nuts ofmodern Fagus crenata and F. sylvatica. Wingsare developed along the upper angles of thenuts (Plate VI, Figs. 10, 11).

Remarks : The name F. cf. japonica is proba-bly due to confusion with modern (herbarium) mate-rial. It might refer to a modern specimen of F. cre-nata, figured in ILJINSKAYA (1982, pl. 136, fig. 12)and erroneously ascribed to F. japonica. The fossilshave nothing in common with the modern F. japo-nica but resemble F. tientaiensis (F. longipetiolataspecies aggregate), F. crenata, and F. grandifolia.They should be included within Fagus deucalionis.

Fagus cf. longipetiolata (K509)


Of eight specimens one is a complete cupule,another a cupule with broken apex, and six aresingle valves. Valves are 9–17 mm long and5.0–6.5 mm wide. The peduncle is thickenedat the point of insertion, and either merginginto the cupule or inserting abruptly. Its di-ameter is up to 4 mm at the point of insertionand 1–2.5 mm at its most proximal parts(Plate VII, Figs. 1, 2, 4). Sometimes it isstrongly flattened. It is always broken. Thenumber of appendages per cupule valve ismore than 35 in one specimen (Plate VII,Fig. 8). Appendages are always spine likeand display different stages of preservation(steps 1, 5, 9 in Plate I). In general they arecomparatively well preserved and not flattened.They are broken and not appressed to the

valve. The appendages are stout, and more than0.25 mm in cross section. Appendages thatappear to be conspicuously stout may be thebasal part of a fused bundle of appendagesbefore they separate and diverge. The cupulesurface is not ridged and thus bases of appen-dages do not appear “decurrent”. The innersurface of the cupule valves may be smooth ormarkedly ribbed.

Remarks : This species resembles at least threemodern species, namely F. sylvatica, F. crenata, andF. longipetiolata. Due to the absence of nuts itshould be placed into Fagus deucalionis.

Relationships among the studied taxa

Based on the length and width of the cupulevalves examined, two groups can be distin-guished (Fig. 1). Specimens ascribed to F. cf.grandifolia (K412), F. cf. orientalis and F. cf.grandifolia (K530), all of which are fromsouth-western Asia and the European partof Russia from Middle Miocene to LowerPliocene deposits, form one group character-ized by “small” cupules/valves (Morphotype1 = MT1), whereas F. cf. grandifolia (K502),F. cf. longipetiolata and F. cf. japonica, allfrom Oligocene to Miocene deposits of westernSiberia, form a second group having “large”cupules (MT2). Fagus cf. lucida is somewhatisolated from these groups (Fig. 1) but wouldrather belong to MT1. It is not possible to un-ambiguously decide whether these morpho-types represent two distinct species or mor-photypes of a single species. The few availablenuts are either 7–10 mm (MT1), or 9.0–12.5 mm long (MT2). Except for F. cf. lucidaand F. cf. orientalis, where they are eitherwinged or unwinged, nuts display wingededges in the upper parts.

Discussion

The results of the present study do not confirmthe previous assignment of several cupule/nutspecimens from the Tertiary of Russia, Cauca-sia and Central Asia to particular modernFagus species. Whereas some specimens donot at all resemble modern species they wereascribed to (F. cf. orientalis, F. cf. lucida,

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Fig. 1Scatter diagram. Length and width values for complete valves are plotted. Two groups are recognized: Faguscf. longipetiolata, F. cf. japonica and F. cf. grandifolia K502, (upper part of diagram: Morphotype 2 = MT2),and the remaining taxa (lower part; MT1). Lines are trend lines

F. cf. japonica), others show similarities tomore than one modern species (F. cf. grandi-folia, F. cf. longipetiolata). In addition, it isdifficult to establish whether fossils from dif-ferent localities represent distinct species ornot. This is mainly because a number of cupulecharacteristics previously considered diagnos-tic of different fossil Fagus species (e.g.,decurrent versus non-decurrent appendages,MAI 1997) may merely represent differentstages of cupule decay (compare Plate I; DENK& MELLER 2001).

Similarities between the Middle to LateMiocene and Pliocene F. cf. grandifolia (K530and K412) and Fagus cf. orientalis from Rus-sia and Transcaucasia (Table 1; Fig. 1) aremainly due to their relatively small cupules.Both, F. cf. orientalis and F. cf. grandifoliaK530 exhibit two kinds of cupule valves,namely broad ovate-elliptic ones (Plate II,Figs. 1, 3; Plate III, Fig. 1) and conspicuouslynarrow ones (Plate II, Figs. 9, 10; Plate III,Fig. 5). Outer cupule surfaces often displaysharp ridges (Plate II, Figs. 7, 11, 12; Plate III,Fig. 4). Nuts are clearly winged and as long as

the cupule valves. Due to the absence of distin-guishing characters these specimens are sug-gested to belong to one morphotype (MT1) of asingle biological species, rather than to threespecies of two subgenera of Fagus (DOROFEEV1982).

A second group of taxa suggested to bemore closely related comprises the westernSiberian Oligocene to Miocene F. cf. lon-gipetiolata, F. cf. grandifolia (K502), andF. cf. japonica. Again, this is mainly due totheir relatively larger size as compared to theremaining taxa examined. Moreover, theydisplay similar ovate-elliptic shapes of cupulevalves (compare Plate V, Figs. 3, 5–7, F. cf.grandifolia K502; Plate VI, Fig. 1, F. cf. ja-ponica; and Plate VII, Figs. 1, 2, F. cf. lon-gipetiolata). Fagus cf. grandifolia K502 andF. cf. japonica also display cupules with broadovate-roundish valves. Both have clearlywinged nuts, whereas no nuts are preserved inF. cf. longipetiolata. Patterns of decay of theouter cupule are similar to step 1 (in Plate I) inF. cf. longipetiolata, to steps 5 and 7 in F. cf.grandifolia K502, and to steps 5 to 9 in F. cf.


Table 3Cupule/nut characteristics of modern species of Fagus that are comparable to fossil species

Cupule valve length(mm)

Nut morphology

F. engleriana (7–)15–20(–25) Winged apical edgesF. japonica 5–8 UnwingedF. okamotoi 5–7 Prominently wingedF. longipetiolata 15–25 Unwinged, or only slightly winged

[Prominently winged in some populations]F. bijiensisa 12–15 UnwingedF. tientaiensisa 12–15 WingedF. lucida 7–15 UnwingedF. chieniia 10–16 Slightly wingedF. hayatae 7–10 Prominently wingedF. crenata 9–20 Prominently to not wingedF. sylvaticab (10–)15–30 Prominently wingedF. grandifolia 7–18 Prominently to not winged

a Based on one collection of a single tree; b including F. moesiaca and F. orientalis

japonica. In general, these specimens are notdistinguishable from the European fossil spe-cies Fagus deucalionis. They appear to belongto a second morphotype (MT2). The only fea-ture distinguishing them from the specimens ofthe first group of species is their larger size.

In light of a conspicuous amplitude of thesize of cupules in modern Fagus species (SHEN1992; DENK 1999a, b; Table 3), and the Euro-pean fossil species F. deucalionis (DENK &MELLER 2001), the two groups recognizedhere, and separated geographically and (partly)stratigraphically, are considered as two mor-photypes of the same species F. deucalionis.Fagus cf. lucida does not clearly fall within thevariability of the two morphotypes recognizedabove. This species is based on a few badlypreserved specimens. In principle, it couldrepresent a population slightly deviating fromthe specimens assigned to MT1.

Alternatively, MT1 and MT2 could havebelonged to two different species. Modernanalogues for such a scenario are found, forexample, in the cupule/nut complex of themodern species Fagus crenata and F. hayatae(Table 3). These species are very similar butdiffer considerably in size.

Cupules and valves ascribed to MT2 mayhave been linked to leaves of Fagus antipofii(Table 1) from approximately the same regionand stratigraphic position. In such a case, both

leaves and cupules/nuts would have been con-siderably homogeneous suggesting them tobelong to one species. At least for the Oligo-cene and Early Miocene it seems possible thatmost Fagus remains (pollen, leaves and cu-pules/nuts) of Eurasia belong to few if not asingle one biological species (DENK 2000). Theabsence of major ecological radiations duringan early phase of expansion of the genus couldexplain the uniformity of Fagus fossils duringthe Oligocene and Early Miocene. It wouldalso be in agreement with the high amount ofmorphological parallelisms in modern Fagusspecies.

In case of MT1 (Europe, western Transcau-casia) leaves that were possibly associated withthe fossil cupules and nuts are less homogenous(F. herthae, F. juliae; Table 1). These leavesdisplay morphological affinities to severalEast Asiatic and North American species andto a lesser degree to European/south-westernAsiatic ones. Together with their relativelysmall cupules they do not resemble modernFagus sylvatica from Europe and south-western Asia. This may indicate a late differ-entiation of modern European/west Asiatic andEast Asiatic species, and would be in agree-ment with observations in European Fagusremains from the Late Miocene and Pliocene.The few cupules found so far in Late Miocenedeposits of Macedonia (northern Greece) are

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relatively large, pointing to extant Fagus syl-vatica (VELITZELOS et al. 2002, in press). Incontrast, leaves are extremely variable dis-playing morphotypes that are clinally con-nected and either very similar to modernChinese species Fagus longipetiolata, or tothe modern Fagus sylvatica subsp. orientalis(DENK, unpubl. data). In Pliocene deposits ofFrankfurt cupules are relatively smaller,pointing to several East Asiatic and NorthAmerican species (MÄDLER 1939). They co-occur with leaves that are conspicuously vari-able showing close similarities to the modernChinese species Fagus hayatae subsp. pa-shanica and the Japanese Fagus crenata(DENK, unpubl. data).

Acknowledgements

Alexander Ruhr i thanked for translating the chap-ter about Fagaceae in “Magnoliophyta FossiliaUSSR”. Special thanks go to Irina I l j inskaya ,Lev Budan tsev , Lena Golevneva and OlgaArbuzova for providing help in the palaeobotani-cal collections in the I. V. Komarov Botanical In-stitute of the Russian Academy of Sciences in St.Petersburg.

References

BUDANTSEV, L. Y. 1997: Late Eocene Flora ofwestern Kamchatka. – St. Petersburg.

CAO, K.-F. 1995: Fagus dominance in Chinesemontane forests. – Landbouwuniversiteit, Wage-ningen, Ph. D. Diss.

DENK, TH. 1999a: The taxonomy of Fagus in west-ern Eurasia, 1: Fagus sylvatica subsp. orien-talis (= Fagus orientalis). – Feddes Repert. 110(1–2): 177–200.

DENK, TH. 1999b: The taxonomy of Fagus in west-ern Eurasia, 2: Fagus sylvatica subsp. sylvatica.– Feddes Repert. 110 (5–6): 381–412.

DENK, TH. 2000: The northern hemispheric genusFagus in the Oligocene: one or many species? –IOPC 6, Quinhuangdao, China.

DENK, TH. & MELLER, B. 2001: Systematic signifi-cance of the cupule/nut complex in livingand fossil Fagus. – Intern. J. Plant Sci. 162 (2):869–897.

DOROFEEV, P. I. 1982: Fagus L. II: 72–73. Speciesidentified after cupules. – In: A. TAKHTAJAN

(ed.), Magnoliophyta fossilia URSS, Vol. 2.Ulmaceae – Betulaceae. – Leningrad (St. Peters-burg).

FOTJANOVA, L. I. 1982: Fagus napanensis FOTJAN.sp. n.: 64. – In: A. TAKHTAJAN (ed.), Magnolio-phyta fossilia URSS, Vol. 2. Ulmaceae – Betu-laceae. – Leningrad (St. Petersburg).

ILJINSKAYA, I. A. 1982: Fagus L.: 60–73. – In: A.TAKHTAJAN (ed.), Magnoliophyta fossilia URSS,Vol. 2. Ulmaceae – Betulaceae. – Leningrad(St. Petersburg).

KVA�EK, Z.; VELITZELOS, E. & VELITZELOS, D.2001: A monograph of the Late Miocene flora ofVegora, Northern Greece. – in press.

KVA�EK, Z. & WALTHER, H. 1991: Revision dermitteleuropäischen Fagaceen nach blattepider-malen Charakteristiken. IV. Teil Fagus. –Feddes Repert. 102 (7–8): 471–534.

MÄDLER, K. 1939: Die Pliozäne Flora von Frankfurtam Main. – Abh. Senckenberg. Naturforsch.Ges. 446: 1–202.

MAI, D. H. 1997: Die oberoligozänen Floren amNordrand der Sächsischen Lausitz. – Palaeon-tographica, B, 244: 1–124.

MEYER, H. W. & MANCHESTER, S. R. 1997: TheOligocene Bridge Creek Flora of the John DayFormation Oregon. – Univ. California, Publ.141: 1–195.

PETERS, R. 1997: Beech Forests. – Geobotany 24:1–169. – Dordrecht.

REID, C. & REID, E. M. 1915: The Pliocene floras ofthe Dutch-Prussian border. – Medel. Rijksops.Delftst. 6: 1–178.

SHEN, C.-F. 1992: A monograph of the genus FagusTOURN. ex L. (Fagaceae). – The City Univ. ofN.Y., Ph. D. Diss.

TAKHTAJAN, A. (ed.) 1982: Magnoliophyta FossiliaURSS, Vol. 2. Ulmaceae – Betulaceae. – Lenin-grad (St. Peterburg).

UEMURA, K. 1982: Fagus remains from the Pleisto-cene beds in the Atsuni Peninsula, Central Japan.– Mem. Nat. Sci. Mus. (Tokyo) 13: 35–43.

UNGER, F. 1847: Chloris protogaea. – Leipzig.

Address of the author:

Dr. Thomas Denk, Swedish Museum of NaturalHistory, Department of Palaeobotany, Box 50007,104 05 Stockholm, Sweden.e-mail: [email protected]

Manuscript received: January 2nd, 2002.


Explanations to Plates I to VII

PLATE I

Stages of decay in water transported modern cupulesof Fagus sylvatica

Fig. 1: Slightly decayed valve with spine-like bro-ken and complete, twisted appendages, S116046. —Fig. 2: Apical part of valve, all appendages broken,their bases non-decurrent, triangle-shaped, S115973.— Fig. 3: Apical part of valve, all appendages bro-ken, their bases decurrent, S115979. Fig. 4: — Morestrongly decayed valve displaying decurrent bases ofappendages that appear corrugated, S115962. —Fig. 5: Cupule displaying vertically and denselyarranged decurrent bases of appendages, which areabraded in the basal parts, S115966. — Fig. 6: Cu-pule with broken, spine-like appendages displayingwart-like non-decurrent bases, S115972. — Fig. 7:Heavily decayed cupule with broken appendages.Bases are decurrent, S115976. — Fig. 8: Heavilydecayed cupule with broken appendages; their basesforming sharp ridges. — Fig. 9: Heavily decayedcupule. Bases of broken appendages appear wart-like in the apical part, and smooth due to abrasion inbasal parts. — Fig. 10: Heavily decayed cupule withcompletely smooth outer surface due to abrasion,S115968

Scale bar = 10 mm

PLATE II

Fagus orientalis (K531)

Figs. 1, 3: Cupule with ovate-elliptic valves, apicalparts broken. Outer cupule surface heavily decayed,displaying vertically running decurrent bases ofappendages. Peduncle merging into cupule. —Fig. 2: Cupule with oblong valves, peduncleabruptly inserting into cupule. — Fig. 4: Narrowelliptic cupule valve with lop-sided insertion ofpeduncle. — Fig. 5: Close-up of Fig. 4: Apical partof valve with ridged outer cupule surface. — Fig. 6:Cupule valve with peduncle. — Fig. 7: Close-up ofFig. 6: outer cupule surface displaying verticallyrunning sharp ridges. — Fig. 8: Cupule valve withridged outer surface and few broken appendageswith non-decurrent bases (left part of valve). —Fig. 9: Narrow elliptic valve merging into brokenpeduncle. — Fig. 10: Close up of Fig. 9 showingvertical ridges and broken appendages on erodedouter cupule surface. — Fig. 11: Narrow-triangularshaped valve. — Fig. 12: Close-up of Fig. 11showing sharp vertical ridges on outer cupule sur-face. — Figs. 13–16: Nuts: 13: – relatively large nutwith broken wings, 14 – nut with broken wings, 15 –

relatively small unwinged nut, 16 – close-up ofapical part of a nut with fragmentary wing

Scale bar = 10 mm in Figs. 1, 2, 4, 9; 5 mm in Figs. 3, 6, 8,10, 11; 3 mm in Figs. 5, 7, 12

PLATE III

Figs. 1–6: Fagus cf. grandifolia (K530)

1 – cupule with part of peduncle preserved; cupulevalves broad-ovate, peduncle abruptly inserting intocupule; 2 – cupule with narrow-elliptic valves,peduncle merging into cupule; 3 – valve displayingvertically arranged appendages with slightly decur-rent bases; 4 – close up of 3; 5 – lanceolate-ellipticvalve; 6 – close-up of Fig. 5 showing decurrentbases of appendagesFig. 7: Fagus cf. grandifolia K412; cupule withbroad-ovate valves and heart-shaped insertion ofpeduncleFigs. 8–11: Fagus cf. lucida (K517)8 – cupule displaying narrow ovate-elliptic valvesthat are slightly contorted; the broken pedunclemerging into cupule; 9 – same cupule from back;10 – valve displaying eroded outer surface withbroken spine-like appendages, 11 – close-up of 10

Scale bar = 10 mm in Figs. 5 – 7, 11 – 13; 5 mm in Figs. 1,3, 9, 14; 3 mm in Figs. 2, 4, 8, 10, 15

PLATE IV

Fagus cf. lucida (K517)

Fig. 1: Cupule with narrow elliptic valves displayingeroded outer surfaces, the two nuts are longer thanthe valves. — Fig. 2: Same cupule in different view,one valve is broken. — Fig. 3: Close-up of upperpart of Fig. 2 showing nut with fragmented wing. —Fig. 4: Same cupule in different view; nut appears tobe unwinged. The gap between the base of the nutand the point where it is attached to the cupule isindicated by broken lines. — Fig. 5: Close-up ofFig. 4 showing apical margin of nut, which appearsto be unwinged due to smoothly broken wings. —Fig. 6: Same cupule in different view. — Fig. 7:Close-up of Fig. 6 showing fragmented wing of oneof the two nuts, indicated by an asterisk. — Fig. 8:Broad valve displaying vertically arranged slightlydecurrent spine-like appendages. — Fig. 9: Close-upof Fig. 8, compare to Plate V, Figs. 2, 4

Scale bars = 5 mm in Figs. 1, 2, 4, 6, 8; 2 mm in Figs. 3, 5,7, 9

PLATE V


Fig. 1: Valve with vertically arranged decurrentappendages. — Fig. 2: Close-up of Fig. 1 showing

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ridges of bases of appendages. — Fig. 3: Cupulewith smooth surface in its lower parts and brokenappendages in upper parts; broken peduncle merginginto cupule. — Fig. 4: Close-up of Fig. 3 showingvertically arranged spine-like appendages. — Fig. 5:Cupule with ovate-elliptic valves, no pedunclepreserved. — Fig. 6: Same cupule in different view.— Fig. 7: Cupule with decurrent appendages andtwo nuts that are as long as the valves. — Fig. 8:Same cupule in different view. — Fig. 9: Close-upof Fig. 8 showing winged nuts indicated by an as-terisk

Scale bar = 10 mm in Figs. 1, 3, 4, 5, 6, 7, 8; 3 mm inFigs. 2, 4, 9

PLATE VI

Fagus cf. japonica (K467)

Fig. 1: Cupule with ovate elliptic valves, outer sur-face heavily decayed, displaying wart-like protru-sions. — Fig. 2: Cupule with broad-ovate valves,displaying spine-like broken, recurved appendages.— Fig. 3: Same cupule in different view — Figs. 4,5: Close-ups of Fig. 3 under different light. Brokenand partly recurved appendages. — Figs. 6, 7: Cu-pule under different light. Outer surface with sharp

ridges, decurrent; nuts as long as the valves. Brokenpeduncle abruptly inserting into the cupule. —Fig. 8: Cupule with broad ovate valves; containingnuts; the broken peduncle abruptly inserting intocupule. — Figs. 9–12: Nuts. 9, 10 – nut with brokenwing, 11 – detail of nut angle showing part of wing,12 – basal part of nut showing the triangle shapednut-scar

Scale bar = 10 mm in Figs. 1, 2, 3, 6, 7, 8, 9; 3 mm inFigs. 4, 5, 11, 12

PLATE VII

Fagus cf. longipetiolata (K509)

Figs. 1, 2: Different views of a complete cupuleshowing the thickened peduncle at the point ofinsertion into the cupule. — Fig. 3: Close-up ofFig. 1 showing broken, spine-like non-decurrentappendages. — Fig. 4: Valve with broken peduncleand few appendages. — Fig. 5: Close-up of Fig. 4.— Fig. 6: Narrow ovate cupule valve. — Fig. 7:Close-up of Fig. 6 showing broken spine-like ap-pendages. Ovate-elliptic cupule valve. — Fig. 9:Close-up of Fig. 8

Scale bar: = 10 mm in Figs. 1, 2, 4, 6, 8; 3 mm in Figs. 3,5, 7, 9


Plate I

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Plate II


Plate III

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Plate IV


Plate V

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Plate VI


Plate VII

Documents

Revision of Tertiary Fagus cupules from Russia, Transcaucasia and western Siberia