The Aptian–Cenomanian Flora of Primor’e, Part 1: Floral Assemblages

613

Stratigraphy and Geological Correlation, Vol. 13, No. 6, 2005, pp. 613–631. Translated from Stratigrafiya. Geologicheskaya Korrelyatsiya, Vol. 13, No. 6, 2005, pp. 58–76.Original Russian Text Copyright © 2005 by Volynets.English Translation Copyright © 2005 by

åÄIä “Nauka

/Interperiodica” (Russia).

INTRODUCTION

The Early Cretaceous flora of Primor’e was underexamination during almost a hundred years. It attractedattention of many well-known experts in paleobotany:A.N. Krishtofovich, V.D. Prinada, B.M. Shtempel’,V.A. Vakhrameev, V.A. Samylina, and V.A. Krassilov.Krishtofovich (1921) who studied Mesozoic plantremains in southern Primor’e termed the oldest andyoungest ones as the Mongugai and Nikan floras. Hewas first to discover here the remains of early floweringplants:

Pandanophyllum ahnertii

in the Razdol’nayadepression and

Aralia lucifera

Krysht. and

Cissites pro-dromus

Krysht. in Partizanskaya depression (Krishto-fovich, 1928, 1929; Krishtofovich and Pavlov, 1928).Prinada described the Mesozoic flora of Primor’e in amonograph that has not been published yet. Manuscriptof the book is deposited in the “Primorgeologiya”funds. Some of plant remains studied by Prinada aredescribed in books entitled “Data on Paleontology”(1956) and “Fundamentals of Paleontology” (1963a,1963b). Shtempel’ (1959a, 1959b, 1960, 1962) investi-gated the Early Cretaceous flora from the Partizanskayadepression and published some new data on paleobot-any of other regions. Vakhrameev (1959) described acollection of fossil plants from western coastal out-crops of the Lake Khanka. Samylina (1961) studied anddescribed representatives of cycadophytes and conifersfrom the Razdol’naya depression and fruits of theangiosperm genus

Nyssidium.

Krassilov (1967) sug-gested a rather detailed stratigraphic chart for theLower Cretaceous of southern Primor’e and distin-

guished the Ussuriisk (Barremian), Lipovtsy (Aptian)and Galenki (Albian) formations in the Razdol’nayadepression, and the Staryi Suchan (Barremian), Sever-nyi Suchan (Aptian) and Frentsevo (Albian) formationsin the Partizanskaya depression. The Early Cretaceousflora of southern Primor’e, which is represented by180 taxa, 58 new species included, has been studied tosubstantiate the chart. Krassilov and Shorokhova (1989)identified plant remains from five localities of theAlchan Formation in synonymous depression and themiddle Albian–Cenomanian age of this subdivision. In1985, Nevolina (1990) first established the late Cenom-anian age of fossil flora from the Stolbovaya sequenceof the Alchan depression. Data on the Aptian–Cenoma-nian floras from the Alchan, Partizanskaya andRazdol’naya depressions, which are presented in thiswork, specify stratigraphic ranges of certain subdivi-sions and elucidate further the taxonomic compositionof their floras.

The Aptian–Cenomanian time span characterizes animportant period of transition from Mesophytic toCenophytic vegetation and associated appearance offlowering plants representing one of the high-rank taxa.In Primor’e, the Aptian–Cenomanian floras are studiedbest of all in the Alchan, Razdol’naya and Partizan-skaya depressions (Fig. 1). In the Alchan and Partizan-skaya depressions, the flora-bearing beds are interca-lated with strata containing fossil fauna. Localities offossil floras are confined to the Assikaevka (Aptian–ini-tial middle Albian) and Alchan (middle–upper Albian)formations and to the Stolbovaya sequence (Cenoma-

The Aptian–Cenomanian Flora of Primor’e, Part 1: Floral Assemblages

E. B. Volynets

Institute of Biology and Soil Science, Far East Division, Russian Academy of Sciences, Vladivostok

Received February 25, 2004; in final form, December 7, 2004

Abstract

—Abundant and diverse Aptian–Cenomanian floras from deposits of the Alchan, Razdol’naya, andPartizanskaya depressions of Primor’e are investigated. Stratigraphic ranges of 370 identified taxa are estab-lished. Studied floras are divided into floral assemblages (FA) and subassemblages. Fossil plants of the Alchandepression are divided into five floral assemblages: the early and late Assikaevka FAs, the latter with two sub-assemblages, the early Alchan FA consisting of early, middle and late subassemblages, the middle and lateAlchan FAs, the last one again divisible into three subassemblages, and the Stolbovaya FA. The early and lateLipovtsy, early to late Galenki, and early Korkino FAs represent six floras of the Razdol’naya depression. Dis-criminated in the Partizanskaya depression are the late Staryi Suchan, early and late Severnyi Suchan, Frentsevo(with two subassemblages), Kangauz, Romanovka, Brovnichi, and Dadanshan FAs. The Aptian–Cenomanianflora of the Alchan depression and flora of the terminal late Albian from basal beds of the undivided KorkinoGroup, the Razdol’naya depression, are comprehensively characterized for the first time.

Key words

: Aptian, Albian, Cenomanian, depression, flora, assemblage, subassemblage, Primor’e.

614

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2005

VOLYNETS

nian) of the Alchan depression. In the Razdol’nayadepression, they are known in the Lipovtsy (Aptian andpresumably initial lower Albian) and Galenki (lowerand first half of the upper Albian) formations and in theundivided Korkino Group (uppermost Albian). In thePartizanskaya depression, plant remains occur in theupper Staryi Suchan Subformation (Aptian–lowermostAlbian) and in the Severnyi Suchan (lower–initial mid-dle Albian), Frentsevo (middle–initial upper Albian),Kangauz (middle upper Albian), Romanovka (upper-most Albian), Brovnichi (uppermost Albian–lowermostCenomanian) and Dadanshan (upper Cenomanian) for-mations.

MATERIALS

Abundant plant remains studied in this work (fewhundred thousand specimens) have been collected fromAptian–Cenomanian deposits of Primor’e in collabora-tion with my colleagues S.I. Nevolina and S.A. Sho-rokhova and by many field geologists: G.L. Amel’-chenko, V.F. Lushnikov, V.L. Mudrovoi, A,N. Naid-enko, V.N. Okovityi, S.V. Kovalenko, A.V. Oleinikov,V.V. Golozubov, A.T. Kondaurov, L.A. Gusakov, andothers. The collected plant fossils are stored in Vladi-vostok at the institute of Biology and Soil Science, Rus-

sian Academy of Sciences, and the Primorsk Expedi-tion of Geological Survey.

Plant remains characterize more than 200 burials offloras. They are represented mostly by impressions ofleaves and leafed shoots, by mineralized stems andwood, and by fruits. Being in satisfactory preservationstate, the remains exhibit fine details of plant organs,but phytoleims are usually rare or considerably altered,unsuitable for getting preparations of cuticular tissueappropriate for examination.

CHARACTERIZATIONOF APTIAN–CENOMANIAN FLORAL

ASSEMBLAGES FROM PRIMOR’E

According to their taxonomic composition, theAptian–Cenomanian floras of Primor’e (Table 1) aredivided into 19 floral assemblages (FA)

Northwestern Primor’e, the Alchan Depression

The Aptian–Cenomanian terrigenous and volcano-genic deposits widespread in the Alchan depression aredivided into the Assikaevka (Aptian–initial middleAlbian) and Alchan (middle–upper Albian) formationsand Stolbovaya (Cenomanian) sequence (Amel’chenko

et al.

, 2001; Nazarenko and Bazhanov, 1989; Markevich

0 60 km

SEA

OF

JAPA

N

N

S

China

Bikin R.

B. Ussurka R.

Ussuri R.

Lake Khanka

Ussuriisk

Partizansk

Vladivostok

1 2 3

Fig. 1.

Cretaceous depressions of Primor’e: (1) Razdol’naya; (2) Partizanskaya; (3) Alchan.

STRATIGRAPHY AND GEOLOGICAL CORRELATION

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2005

THE APTIAN–CENOMANIAN FLORA OF PRIMOR’E, PART 1: FLORAL ASSEMBLAGES 615

et al.

, 2000;

Regional…

, 2001). These deposits containabundant and diverse remains of fossil plants, whichhave been collected during geological survey and bios-tratigraphic research (Amel’chenko

et al.

, 2001; Voly-nets, 1997, 2000; Krassilov and Shorokhova, 1989).Fossil plants are divisible into five FAs, which succes-sively replace each other upward in the section.

The early Assikaevka floral assemblage (FA)

is dis-tinguished at the base of the lower Assikaevka Subfor-mation in lower courses of the Bikin and Assikaevkarivers (Volynets, 2001). The assemblage is not verydiverse (18 taxa). Dominant among floral remains arecycadophytes (6 taxa) of genera

Pterophyllum

and

Nils-sonia

, the most abundant species

Pterophyllum sut-schanense

and

Nilssonia

ex gr.

brongniartii

included.Subdominant ferns (5 taxa) include genera

Birisia,Onychiopsis, Polypodites, Coniopteris (Dicksonia)

,and

Cladophlebis.

Two species

Polypodites ver-estchaginii

and

Onychiopsis psilotoides

are especiallyabundant. Remains of ginkgoales are rare, representedby

Ginkgoites

cf.

coriacea

(Table 1, Fig. 2). Consider-able percentages of

Elatides asiatica, Athrotaxites ber-ryi, Pityophyllum

sp. and

Podozamites

sp. are alsocharacteristic.

Lower beds of the lower Assikaevka Subformationyield shells of bivalve mollusks

Nuculopsis

sp.,

Ento-lium

sp. indet,

E.

cf.

utukokense

Imlay,

Eumorphotis

sp.,

Pleuromya

sp. indet., while bivalves

Nucula

sp.indet.,

Isognomon

sp.,

Trapezium (?)

sp.,

Gresslia (?)

sp.,

Thracia

sp. indet. and ammonite

Hulenites

sp. havebeen found in the subformation upper part (Table 2). Inopinion of V.P. Konovalov, sediments containing fossilsare of the Aptian–Albian age (Markevich

et al.

, 2000).

The late Assikaevka FA

is characteristic of lowerand upper beds of the upper Assikaevka Subformation,which are exposed in lower and middle courses of theBikin River (Volynets, 2001). This FA is more diverse(54 taxa), containing abundant fern species

Onychiop-sis psilotoides, Anemia dicksoniana, Gleichenites por-sildii, Birisia onychioides, B. alata

, and representativesof genera

Ruffordia, Dicksonia, Osmunda, Adiantopt-eris, Alsophilites, Coniopteris, Arctopteris, Lobifolia

and

Cladophlebis.

Species of frequently occurringconifers and cycadophytes are

Elatides asiatica,Podozamites

ex gr.

lanceolatus, Athrotaxites berryi,Sequoia reichenbachii

, and

Nilssonia

ex gr.

brongn-iartii.

Representatives of genera

Zamiophyllum, Ptero-phyllum, Ptilophyllum, Taeniopteris, Athrotaxopsis,Sphenolepis, Torreyites, Taxites, Ginkgo, Baiera

, and

Desmiophyllum

occur as single specimens. In addition,leaves of angiosperm genera

Sapindopsis, Laurophyl-lum, Dicotylophyllum, Quercophyllum

, and fruits

Nys-sidium

and

Onoana

appear for the first time in theassemblage. Czekanowskiales, caytoniales, and horse-tails are of insignificant abundance (one taxon per each,Table 1, Fig. 2). The considerable proportion of“young” genera

Anemia, Osmunda, Birisia, Arctopt-eris, Sequoia, Sphenolepis

and

Taxites (“Cephalotax-

opsis”)

is a distinctive feature of the assemblage. Cze-kanowskiales and early angiosperms are encountered atthis level for the first time. The late Assikaevka FA isdivided in two subassemblages.

The early subassemblage

from the lower part of theupper Assikaevka Subformation is dominated by fernsof genera

Onychiopsis, Birisia, Ruffordia, Dicksonia,Gleichenites, Lobifolia

and

Cladophlebis.

The mostfrequent fern species are

Onychiopsis psilotoides, Gle-ichenites porsildii

and

Cladophlebis frigida.

Subdomi-nant cycadophytes are represented by genera

Zamio-phyllum, Pterophyllum, Ptilophyllum, Nilssonia

, in par-ticular by abundant species

Nilssonia

ex gr.

brongniartii.

Abundant among conifers are

Elatidesasiatica

and

Podozamites

ex gr.

lanceolatus

, which areaccompanied by appearing

Torreyites

cf.

dicksonioides

(Table 1, Fig. 2).

The late subassemblage

from the upper AssikaevkaSubformation is also dominated by ferns (18 taxa) ofgenera

Osmunda, Anemia, Ruffordia, Adiantopteris,Onychiopsis, Gleichenites, Alsophilites, Dicksonia,Birisia, Coniopteris, Arctopteris, Lobifolia

and

Cla-dophlebis.

Species most frequently occurring in thisgroup are

Anemia dicksoniana, Gleichenites porsildii,Birisia onychioides

and B. alata. Subdominant conifers(16 taxa) include first appearing representatives of gen-era

Athrotaxopsis, Sequoia, Sphenolepis

and

Taxites

inaddition to

Elatides asiatica, Podozamites

ex gr.

lan-ceolatus

and

Athrotaxites berryi.

The associated cyca-dophytes (5 taxa) belong to genera

Zamiophyllum,Pterophyllum, Ptilophyllum, Nilssonia

and

Taeniopt-eris.

Among single specimens of ginkgoales genera

Ginkgo

and

Baiera

, the frequently occurring species is

Ginkgo

ex gr.

adiantoides

, whereas

Baiera

cf.

polymor-pha

is extremely rare. The first appearing angiospermgenera are

Sapindopsis, Laurophyllum, Dicotylophyl-lum, Quercophyllum, Nyssidium

and

Onoana.

Remainsof czekanowskiales, caytoniales and horsetails are rare(Table 1, Fig. 2).

Plant remains from lower and upper parts of the sub-formation are found in association with ammonites

Phyllopachyceras

ex gr.

theresae

(Anders.),

Anagaud-ryceras aurarium

(Anders.), and bivalve mollusks(Table 2). The latter are represented by

Thracia

ex gr.

sanatae-crucis

Picter et Camp. and

Aucelina

cf. cauca-sica (Buch) in the lower part, by Entolium utucocensisImlay, Macromya sp., Plyuromia cf. kelleri Imlay andAucelina caucasica (Buch) in the middle one. Bivalvespecies found in the upper interval are Leda sp., Ento-lium utucocensis Imlay, Aucelina caucasica (Buch),Inoceramus ex gr. altifluminis McLearn, Thracia cf.stelcki McLearn, and Gresslya sp. In opinion of Konov-alov (see in Markevich et al., 2000), this molluscanfauna corresponds in age to the middle Albian (mostprobably to the first half of the substage).

The early Alchan FA established in the lowerAlchan Subformation (Volynets, 1997, 2001) is repre-sented by 40 taxa. Dominant plant remains (15 taxa)

616

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 13 No. 6 2005

VOLYNETS

Table 1. Taxonomic composition of Aptian-Cenomanian floral assemblages from Primor’e (taxa present in assemblages aremarked by asterisks)

Taxa

Assemblages

Alchan depression Razdol’nayadepression Partizanskaya depression

Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an

Subassemblages (S/a) S/a

earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

BryophytesMarchantites yabei Krysht. et Pryn. * * * *M. ex gr. yabei Krysht. et Pryn. *M. aff. jimboi (Krysht.) Krysht. *Thallites sp. * * * * *Thallites sp. 1 * *Thallites sp. 2 *

LycopodsLycopodites prynadae Krassil. * * * * * * *L. nicanicus Krassil. * *L. ovatus Deng *L. cf. prynadae Krassil. * *L. cf. macrostomus Krassil. *L. aff. multifurcatus Lee *Lycopodites sp. * * * * * * * * * *Isoetites sp. * * * * *Synlycostrobus ex gr. tyrmensis Krassil. *

HorsetailsEquisetum ramosus Samyl. * *E. yokoyamai Sew. *E. burejensis (Heer) Krysht. * * *E. ex gr. burejensis (Heer) Krysht. * * *E. cf. ramosus Samyl. * * *Equisetum sp. * * * * * * * * * * * * * * * * * *Equisetostrobus sp. A *Equisetostrobus sp. *

FernsOsmunda denticulata Samyl. * * * * * * * * * * * * *O. cf. denticulata Samyl. *Osmunda sp. A * *Osmunda sp. * * * *Ruffordia goeppertii (Dunk.) Sew. * * * * * * * * *Ruffordia ex gr. goeppertii (Dunk.) Sew. * * * * * *Ruffordia sp. *Pelletieria ussuriensis (Pryn.) Krassil. * * *Stachypteris turkestanica Tur.-Ket. *S. ketovae Krassil. *Anemia dicksoniana (Heer) Krassil. * * * * * * * * * * * * * * * * * * * *A. asiatica Vachr. * * *A. sphenopteroides J. Skog * *A. sutschanica Volynets Sp. nov. * * * *Anemia sp. * *Gleichenites porsildii Sew. * * * * * * * * * * * * *G. aff. porsildii Sew. *G. zippei (Corda) Sew. * * * *G. gieseckianus (Heer) Sew. * * * * * *Gleichenites sp. * * * * * * * * * * * * *Nathorstia pectinata (Goepp.) Krassil. * *Matonidium sp. * *Alsophilites nipponensis (Oishi) Krassil. * * * * * *Cyathea sp. sujfunensis Volyn. *C. aff. tyrmensis (Sew.) Krassil. * *Cyathea sp. 1 * *Gleicheniopsis sujfunensis Krassil. * *



Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

Dicksonia concinna Heer * * * * * * * * * * * * * * *Birisia onychioides (Vassilevsk. et K.-M.) Samyl. * * * * * * * * * * * * * * * *B. (Cladophlebis) jelisejevii (Krysht.) Philipp. *B. (Acanthopteris) alata (Pryn.) Samyl. * * * * * * *B. oerstedtii (Heer) E. Lebed. * * * *B. aff. oerstedtii (Heer) E. Lebed. *B. samylinae E. Lebed. * * *B. aff. samylinae E. Lebed. *B. ochotica E. Lebed. * * * *Birisia (Acanthopteris) sp. A *Birisia sp. * * *Acanthopteris (Birisia) gothanii Sze *Acanthopteris sp. 1 *Acanthopteris sp. 2 *Coniopteris brevifolia (Font.) Bell *C. asplenioides Kiritchk. * * * * *C. setacea (Pryn.) Vachr. * * *C. chankaensis Krassil. *C. (Dicksonia) burejensis (Zal.) Sew. * * *C. aff. silapensis Samyl. *C. ex gr. arctica (Pryn.) Samyl. * * * * * * *C. yukonensis Bell * *Coniopteris sp. * * * * * * *Eboracia lobifolia (Phill.) Thomas * * *E. ex gr. lobifolia (Phill.) Thomas *E. aff. kataevensis Vachr. *Onychiopsis psilotoides (St. et W.) Ward * * * * * * * * * * * * * * * * * * * *Adiantopteris yuasensis (Oishi) Krassil. * * * *A. sewardii (Yabe) Vassil. * * * *A. grandis Vachr. emend. Volynets *A. minimus Vachr. *A. aff. gracilis (Vassilevsk.) Vassilvesk. *Arctopteris rarinervis Samyl. *A. aff. heteropinnula Kiritchk. * * *A. kolymensis Samyl. * *A. aff. kolymensis Samyl. * *A. cf. kolymensis Samyl. * * * *A. obtusipinnatus Samyl. *Arctopteris sp. 1 *Asplenium rigidum Vassilevsk. * *A. lenaensis Kititchk. *Vargalopteris rossica Pryn. * * *Eogymnocarpium aff. sinense Li et Yeh * *Eogymnocarpium sp. A *Eogymnocarpium sp. B * *Teilhardia tenella (Pryn.) Krassil. * * * * * * * * * * *Hausmannia sp. 1 *“Polypodites” polysorus Pryn. * * *P. verestchaginii Krassil. * *Weichelia reticulata St. et Weeb * *Acrostichopteris pluripartita (Font.) Berry * *Acrostichopteris sp. *Temskya sp. *Lobifolia novopokrovskii (Pryn.) Rasskaz. et Lebed. * * * * * *Cladophlebis frigida (Heer) Sew. * * * * * * * * * * * * * * * * *

618


VOLYNETS

Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

C. denticulata (Brongn.) Font. * * * *C. distans Oishi *C. opposita Pryn. * * * * * * * *C. virginiensis Font. * * * * * *Cladophlebis sp. * * * * * * * * * * * * *Sphenopteris elegants Oishi * *S. stricta (Newb.) Bell *Sphenopteris sp. * * * *“Salvinia” sp. 1 *“Salvinia” sp. 2 *

CaytonialesCaytonia orientalis Krassil. * * *Sagenopteris variabilis (Velen.) Velen. * * *S. mantellii (Dunk.) Schenk. *S. aff. mantellii (Dunk.) Schenk. *Sagenopteris sp. * * * * * * * *

CycadophytesAnomozamites sp. A *Anomozamites sp. * *Dictyozamites grossinervis Yok. *D. cordatus (Krysht.) Pryn. * *D. imamurae Oishi *Dictyozamites sp. A *Dictyozamites sp. * *Williamsonia pacifica Krysht. *Zamiophyllum ivanovii (Krysht. et Pryn.) Krassil. * * *Z. buchianum (Ett.) Nath. *Zamiophyllum sp. * *Cycadeoidea bikinensis Krassil. *Cycadeoidea sp . A *Otozamites aff. schenkii (Heer) Tanai *O. cf. schenkii (Heer) Tanai *Otozamites sp. 1 *Otozamites sp. 2 * *Otozamites sp. * * * * * * * *Zamites borealis Heer * *Z. aff. borealis Heer *Zamites sp. A *Subzamites sp. *Neozamites verchojanensis Vachr. * *N. denticulatus (Krysht. et Pryn.) Vachr. *Pterophyllum sutschanense Pryn. * * * * * * * * * * * * *P. burejense Pryn. * * * * * * *P. manchurense (Oishi) Krassil. *Pterophyllum (Tyrmia pterophylloides Pryn.) sp. *Pterophyllum sp. * * * * * *Leptopterophyllum pterophylloides (Pryn.) Krassil. * *Ptilophyllum sp. * * *Cycadites sulcatus Krysht. et Pryn. * * *Cycadites sp. * * * *Cycadolepis sp. *Pseudocycas sp. *Paracycas aff. brevipinnatus Delle *Nilssoniopteris rhitidorachis (Krysht.) Krassil. * *N. robusta (Krysht. et Pryn.) Krassil. *



Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

N. aff. prynadae Samyl. *Nilssoniopteris sp. * * *Encephalartopsis vachrameevii Volynets sp. nov. * *Encephalartopsis sp. A *Neodoratophyllum sulcatum (Pryn.) Krassil. *Ctenis yokoyamae Krysht. * *C. kaneharai Yok. *C. cf. borealis (Daws.) Bell *Ctenis sp. * * * *Pseudoctenis eathiensis (Rich.) Sew. *P. aff. eathiensis (Rich.) Sew. *Pseudoctenis sp. *Nilssonia densinervis (Font.) Berry * * * * * * * * * *N. prynadii Vachr. *N. ex gr. orientalis Heer * * * * * * * * * * *N. nicanica Pryn. * * * * * *N. canadensis Bell * * * * *N. mediana (Leck. ex Bean MS) Fox-Strang. * * * *N. lobatidentata Vassil. *N. yukonensis Hollick *N. ex gr. brongniartii (Mant.) Dunk. * * * * * * * * * * * * * *Nilssonia sp. * * * * * * * * * *Nilssonia sp. 1 *Nilssonia sp. 2 *Taeniopteris bikinensis Schoroch. * *T. cf. bikinensis Schoroch. *Taeniopteris sp. *“Nikania” sp. *

GinkgoalesGinkgo concinna Heer * *G. pluripartita (Schimp.) Heer * * * * *G. huttonii (Sternb.) Heer *G. ex gr. adiantoides (Ung.) Heer * * * * * * * * * *G. cf. coreacea Florin *Ginkgo sp. * * * * * * * *Baiera manchurica Yabe et Oishi *B. cf. polymorpha Samyl. *Baiera sp. *Sphenobaiera cf. uninervis Samyl. *Sphenobaiera sp. * * * * * *Pseudotorellia ex gr. angustifolia Dolud. * *Pseudotorellia sp. * * * * *

CzekanowskialesPhoenicopsis ex gr. angustifolia Heer * *Phoenicopsis sp. *Czekanowskia aff. vachrameevii Kiritchk. et Samyl. *C. cf. ninae Kiritchk. et Samyl. *C. ex gr. rigida Heer * * * * *Leptostrobus mollis Pryn. *Leptostrobus sp. *

ConifersPodozamites ex gr. lanceolatus (L. et H.) Schimp. * * * * * * * * * * * * * *P. tenuinervis Heer * * * * * * * *Podozamites sp. * * * *Araucariodendron heterophyllum Krassil. *

620


VOLYNETS

Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

A. oblongifolium Krassil. *Araucariodendron sp. * *Ussuriocladus racemosus (Halle) Krysht. et Pryn. *Pagiophyllum orientale Krysht. et Pryn. * *Pagiophyllum sp. * * * *Podocarpus harrsii Krassil. * *P. sujfunensis Krassil. *P. cf. nicanicus Krassil. *Podocarpus sp. *Nageiopsis ussuriensis Krysht. et Pryn. *Nageiopsis ex gr. heterophylla Font. * *Pseudolarix dorofeevii Samyl. * *P. (Pityocladus) kolymensis Samyl. * *P. aff. dorofeevii Samyl. * * *Pseudolarix sp. * *Pityocladus pseudolarixoides Samyl. *Pityocladus sp. A * *Pityophyllum ex gr. nordenskioldii Heer * * * * * * *Pityophyllum sp. * * * * * * * * * * * *Pityospermum aff. prynadae Krassil. *P. prynadae Krassil. *Pityospermum sp. * * *Pityospermum sp. 1 * *Pityospermum sp. 2 *Pityostrobus piceoides Samyl. *Pityostrobus sp. * * * * * *Pityostrobus sp. A * *Pityolepis sp. 1 * * * * *“Picea” sp. 1 *Torreya nicanica Krassil. *Torreyites dicksonioides (Daws.) Bell *T. cf. dicksonioides (Daws.) Bell * * * * * *Taxites brevifolius (Font.) Samyl. * * * * * * * * * *T. acuminatus (Krysht. et Pryn.) Volynets comb.nov. * * * *T. heterophyllus (Holl.) Samyl. *Taxites sp. A * * *Taxites sp. * *Elatides asiatica (Yok.) Krassil. * * * * * * * * * * * * * * * * * * * * *E. splendida Bell * *Elatides sp. A *Sequoia fastigiata (Sternb.) Heer * * *S. reichenbachii (Gein.) Heer * * * * * * * * * * * * * * * * *S. ex gr. reichenbachii (Gein.) Heer *S. ambigua Heer * * * * * *Sequoia sp. 1 * *Sequoia sp. 2 *Sequoia sp. *Cyparissidium gracile Heer * * *Cyparissidium sp. *Sphenolepis sternbergiana (Dunk.) Schenk * *S. ex gr. sternbergiana (Dunk.) Schenk * *S. kurriana (Dunk.) Schenk *Sphenolepis sp. * *Athrotaxites berryi Bell * * * * * * * * * * * * *A. sutschanicus Krassil. *



Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

Athrotaxopsis expansa Font. emend. Berry * * * * * * * * * * * * * *Athrotaxopsis sp. *“Cupressus” sp. * * *Tsuja cretacea (Heer) Newb. *Brachyphyllum douglassi Bell *B. aff. mamillare Brongn. *B. japonicum (Yok.) Oishi *B. ex gr. obesum Heer * * *Brachyphyllum sp. * * * *Elatocladus obtusifolius Oishi * * * *E. gracillimus (Holl.) Sveschn. *E. smittiana (Heer) Sew. *Elatocladus sp. * * * * * * *“Abietites” angusticarpus Font. *Schizolepis neimengensis Deng *S. cretacea Samyl. *Schizolepis sp. ASchizolepis sp. 1 *Schizolepis sp. A *Stachyotaxus sp. A *Coniferites sp. * * *Conites sp. 1 *Conites sp. 2 *Conites sp. A (Metasequoia sp.?) *Conites sp. B *Conites sp. * * *

AngiospermsMagnoliaephylum sp. *Magnoliaephyllum sp. 1 *Magnoliaephyllum sp. 2 * *Sapindopsis variabilis (Font.) Berry * * * * * *S. aff variabilis (Font.) Berry * *S. brevifolia Font. * * *Sapindopsis sp. * * * * * * * *Cinnamomoides ex gr. ievlevii Samyl. *Vitiphyllum (Cissites) sp. *V. (Cissites) parvifolium Font. *Cissites aff. uralensis Krysht. *Cissites sp. * * * *Sassafras ussuriensis Krassil. *S. aff. ussuriensis Krassil. * * *S. cretaceous Font. *Sassafras sp. A * * *Sassafras? sp. * *Quercophyllum ex gr. grossedentatum Font. * *Q. ex gr. tenuinerve Font. *Quercophyllum sp. * *Celastrophyllum ex gr. oppositifolius Samyl. * *C. cf. serrulatus Samyl. *Celastrophyllum sp. * * *Lindera jarmoljukii E. Lebed. *Araliaephyllum obtusilobum Font. *A. rotundilobata (Newb.) Fritel *A. samargense Nevol. *A. cf. parvidensis (Holl.) Philipp. *

622


VOLYNETS

Table 1. (Contd.)

Taxa

Assemblages


Ear

ly A

ssik

aevk

a

Lat

e A

s-si

kaev

ka

Ear

lyA

lcha

n

Lat

eA

lcha

n

Stol

bova

yaE

arly

Lip

ovts

yL

ate

Lip

ovts

yE

arly

Gal

enki

Mid

dle

Gal

enki

Lat

e G

alen

kiE

arly

Kor

kino

Lat

e St

aryi

Suc

han

Ear

ly S

ever

nyi S

ucha

nL

ate

Seve

rnyi

Suc

han

Fren

t-se

vo

Kan

gauz

Rom

anov

kaB

rovn

ichi

Dad

ansh

an


earl

yla

te

earl

ym

iddl

e la

te

earl

ym

iddl

e la

te

earl

yla

te

Araliaephyllum sp. 1 * * * *Araliaephyllum sp. 2 *Araliaephyllum sp. * * * * * *Aralia lucifera Krysht. *Cercidiphyllum aff. sachalinensis Krysht. *Cercidiphyllum sp. * * *Tetracenton aff. potomacence (Ward) Iljinsk. *Artocarpidium sp. *Laurophyllum sp. 1 * * * * *Laurophyllum sp. 2 * *Laurophyllum sp. * * * * * * *Diospiros sp. A *Ficus? glascoeana Lesq. *Andromeda spatulata Bell *Menispermites sp. * * * *Platanus newberryana Heer *P. cuneifolia (Bronn.) Vachr. *P. embicola Vachr. *P. oleinicovii Nevol. *P. cf. primaeva Lesk. *Platanophyllum sp. 1 *Platanophyllum sp. 2 *Platanophyllum sp. A *Platanophyllum sp. * * * *Protophyllum aff. sterbergii Lesq. *Protophyllum sp. *Pseudoprotophyllum sp. *Paraprotophyllum sp. *Credneria sp. *“Scheffleraephyllum” sp. 1 *“Scheffleraephyllum” sp. 2 *Dicotylophyllum sp. 1 * *Dicotylophyllum sp. 2 (Platonoid leaf fragments) *Dicotylophyllum sp. A *Dicotylophyllum sp. * * * * * * *Dalembia sp. A *Quereuxia angulata (Newb.) Krysht. *Pandanophyllum ahneritii Krysht. *Potamogeton sp. A *Nyssidium orientale Samyl. *Nyssidium sp. * *Onoana nicanica Krassil. *Onoana sp. *Kenella harrisiana Samyl. *K. filatovii Samyl. *Kenella sp. *

Plants of unclear taxonomic affinityDesmiophyllum sp. * * * * * * * * * * *Machairostrobus sutschanicus Krassil. * *Zamiopsis sujfunensis (Krysht. et Pryn.) Krassil. *Sujfunophyllum dichotomum Krassil. *Chankanella vachrameevii Krassil. *Carpolithes cinctus Nath. *Carpolithes sp. * * * * * * * * * * *



correspond to ferns of genera Osmunda, Anemia, Dick-sonia, Birisia, Arctopteris, and much less frequent Ruf-fordia, Gleichenites, Alsophilites, Coniopteris, Onychi-opsis, and Acrostichopteris species. Among conifers,species Podozamites ex gr. lanceolatus, Elatides asiat-ica, Torreyites cf. dicksonioides, Sequoia reichen-bachii, Athrotaxites berryi, and Athrotaxopsis expansaare common, whereas Pseudolarix sp. and Sequoiaambigua are rare. Permanent components of cycado-phytes are Pterophyllum sutschanense, Taeniopterisbikinensis, Nilssonia ex gr. brongniartii, and N. densin-

ervis associated with rarer Ctenis and Pseudoctenisremains. The most characteristic species of ginkgoalesis Ginkgo ex gr. adiantoides, while Sphenobaiera isextremely rare (Table 1, Fig. 2). Distinctive features ofthe assemblage are prevalence of ferns (37.5%) andconifers (27.5%) but lower percentage of cycadophytesand absence of flowering plant remains. The assem-blage is divided into three subassemblages.

The early subassemblage is established in basal andoverlying beds of the lower Alchan Subformationexposed in lower courses of the Bikin River. Ferns

10

10

20

30

40

50

60

70

80

90

100

2 3 4 5 6 7 8 9 10 11 12

1 2 3 4 5 6 7 8 9 10 11

%

Fig. 2. Percentages of different plant groups in Cretaceous flora of the Alchan depression: (1) bryophytes; (2) lycopods; (3) horse-tails; (4) ferns; (5) caytoniales; (6) cycadophytes; (7) ginkgoales; (8) czekanowskiales; (9) conifers; (10) angiosperms; (11) plantsof unclear taxonomic affinity. Floral assemblages by names: (1) early Assikaevka; (2) late Assikaevka (early subassemblage); (3)late Assikaevka (late subassemblage); (4) late Assikaevka in general; (5) early Alchan (early subassemblage); (6) early Alchan (mid-dle subassemblage); (7) early Alchan (late subassemblage); (8) early Alchan in general; (9) late Alchan (early subassemblage); (10)late Alchan (middle subassemblage); (11) late Alchan (late subassemblage); (12) Stolbovaya.

624


VOLYNETS

abundant in subassemblage (14 taxa) belong to generaOsmunda, Onychiopsis, Ruffordia, Dicksonia, Birisia,Coniopteris and Cladophlebis. Species Gleichenitesporsildii and Anemia dicksoniana most abundantamong fern remains occur in association with frequentOsmunda denticulata, Onychiopsis psilotoides, Cla-dophlebis opposita, and rare Gleichenites zippei,Acrostichopteris sp., Alsophilites nipponensis, Coniop-teris setacea, Arctopteris aff. kolymensis and Cla-dophlebis frigida. Cycadophytes of genera Pterophyl-lum, Nilssonia and Taeniopteris are not abundant, andTaeniopteris bikinensis is more frequent than other cyc-adophyte species. Species prevailing among conifersare Sequoia reichenbachii and Elatides asiatica(Table 1). Prevalence of ferns (53.8%) and consider-able abundance rate of Sequoia reichenbachii andElatides asiatica (Fig. 2) represent distinctive featuresof this subassemblage.

The middle subassemblage from respective intervalof the Lower Alchan Subformation is established in theSol’da, Malaya Sakhalinka and Krutoberezhnaya riverbasins. Conifers of this subassemblage include abun-dant Athrotaxites berryi and Elatides asiatica, less fre-quent Sequoia reichenbachii, and appeared Sequoiaambigua. Most frequent species of subdominant fernsare Anemia dicksoniana, Dicksonia concinna, Birisiaonychioides and Cladophlebis frigida. Remains of cyc-adophytes (Ctenis sp. and Pseudoctenis sp.) and gink-goales (Ginkgo ex gr. adiantoides and Sphenobaierasp.) are very rare. Typical of the subassemblage aredominance of conifers (37.1%) with a considerable

proportion of Sequoia reichenbachii and high percent-ages of fern remains representing genera Anemia andOsmunda (Fig. 2).

The late subassemblage is distinguishable in upperpart of the lower Alchan Subformation exposed at theMt. Tochil’naya site on the Bikin River right side. It isdominated by ferns, including Anemia dicksonianaoccurring most frequently among them. Subdominantcycadophytes include numerous remains of Pterophyl-lum sutschanense and Nilssonia densinervis. Percent-age of conifers, especially of Elatides asiatica and Ath-rotaxopsis expansa, is high like before. Remains ofLycopodites, Equisetum, Ginkgo, Nageiopsis, and Tax-ites occur as single specimens. Dominant ferns andconifers (up to 31.3%) and considerable abundance(18.8%) of cycadophytes (Fig. 2) are the distinctive fea-tures.

The late Alchan FA from the upper Alchan Subfor-mation is described from the Alchan–Bikin–Marevkainterfluve (Volynets, 1997, 2001). This assemblage isvery diverse in taxonomic composition (over 150 taxa).It consists of dominant ferns and conifers (41 taxa pereach group) and of subdominant angiosperms and cyc-adophytes (26 taxa per each group). The associatedplant remains represent ginkgoales, czekanowskiales,and rare caytoniales, mosses, lycopods and horsetails(Table 1, Fig. 2). The FA is divided into three subas-semblages.

The early subassemblage from the lower part of theupper Alchan Subformation is established in middlecourses of the Bikin River. It is dominated by ferns,

Table 2. Stratigraphic chart and floral assemblages of the Alchan depression

Biostratigraphic subdivisions Lithostratigraphicsubdivisions

Stage Substage Fauna-bearing beds(Markevich et al., 2000)

Floralassemblages

Subassem-blages

Formation(sequence)

Subformation(member)

Cenoma-nian

Upper StolbovayaStolbovaya

Upper

Lower Lower

Albian

UpperEogunarites sp. Late Alchan

Late

Alchan

UpperMiddle

Early

Subarcthoplites talkeetnanus, Paragas-troplites ex gr. spiekeri Early Alchan

Late

Lower

Middle

Middle

Early

Phyllopachyceras ex gr. theresae, Ana-gaudryceras aurarium, Inoceramus ex gr. fitifluminis, Aucellina caucasica

Late AssikaevkaLate

Assikaevka

Upper

LowerEarly

Inoceramus anglicus, Aucellina caucasi-ca, A. anadyrensis, A. ucturiensis et al. Middle

Aptian Hulenites sp., Nucula sp., Isognomon sp. et al. Early Assikaevka Lower



especially by most abundant representatives of generaOsmunda, Gleichenites and Arctopteris. Subdominantconifers consist of genera and species identical to thoseof the early Alchan FA. Nevertheless, representatives ofgenera Podozamites (P. tenuinervus), Brachyphyllum,Pseudolarix, and Pityocladus occur more frequentlyamong them. Cycadophytes (Neozamites), caytoniales(Caytonia, Sagenopteris) and czekanowskiales (Phoe-nicopsis and Czekanowskia) are rare. The subassem-blage is distinct because of presence of dicot generaSapindopsis, Sassafras, Quercophyllum, Araliaephyl-lum, Laurophyllum and Dicotylophyllum. The high tax-onomic diversity of subassemblage is determined byagain appearing angiosperms, caytoniales, and somerepresentatives of conifer genera Brachyphyllum andPityocladus.

The middle subassemblage distinguished in themiddle part of the upper Alchan Subformation in lowerand middle courses of the Alchan, Bikin and Marevkarivers is also very diverse (more than 140 taxa). Domi-nant ferns (40 taxa) include the most representativegenera Gleichenites (G. porsildii, G. aff. porsildii,G. zippei, G. gieseckianus and other species), Birisia(B. onychioides, B. jelisejevii, B. alata, B. oerstedtii,B. samylinae and other forms), Coniopteris (C. asple-nioides, C. setacea, C. ex gr. arctica, C. sp.), Arctopt-eris (A. kolymensis, A. aff. kolymensis, A. aff. het-eropinnula, A. sp. 1) and Eogymnocarpium (E. aff. sin-ense, E. sp. A, E. sp. B). The group of subdominantconifers (37 taxa) consists of abundant Pinaceae(Pseudolarix, Pityophyllum, Pityocladus, Pityosper-mum, Pityostrobus, Pityolepis, “Picea”) and Taxodi-aceae (Elatides, Athrotaxites, Athrotaxopsis, Sequoia).Angiosperms are represented by 29 taxa. Besides thecommon genera Sapindopsis, Sassafras, Quercophyl-lum, Araliaephyllum, Laurophyllum and Dicotylophyl-lum, this group includes new representatives from generaCinnamomoides, Vitiphyllum, Cissites, Celastrophyllum,Lindera, Menispermites, Magnoliaephyllum, Dalembiaand those with platanoid leaves belonging to genusDicotylophyllum. Cycadophytes are also diverse (22taxa), dominated by subtropical Cycadeoidea bikinen-sis and Zamiophyllum ivanovii. Ginkgoales (3 taxa) andczekanowskiales (2 taxa) are less diverse. Rare plantremains represent caytoniales, horsetails, lycopods,bryophytes, and fossil fruits of dicot Kenella. Thegrown species diversity of ferns, angiosperms and flow-ering plants is main distinctive feature of the subassem-blage. Beds with plant remains contain larva sheaths ofTrichoptera (Zherikhin and Sukacheva, 1990), wings ofinsects, fins and spines of fishes, and shells of gastro-pods.

The late subassemblage from upper beds of theAlchan subformation, the Bikin River exposures, isdominated by angiosperms (6 species) of genera Sassa-fras, Sapindopsis, Quercophyllum, Araliaephyllum,and Laurophyllum, the abundant species Araliaephyl-lum sp. included. Subdominant groups are conifers(Elatides asiatica, Athrotaxopsis expansa, Sequoia

reichenbachii, S. sp. 1), cycadophytes (Otozamites sp.,O. sp. 1, O. sp.2, and Nilssoniopteris aff. prynadae) andczekanowskiales. Ferns (Gleichenites sp., Anemia dick-soniana, Onychiopsis psilotoides) are less abundantand diverse than in underlying sediments. Horsetailsand fruits are found as single specimens. Prevalence offlowering plant remains and decreased diversity inother groups of higher plants are the characteristic fea-tures of this subassemblage (Fig. 2).

The Stolbovaya FA is known from the upper part ofsynonymous sequence exposed in middle reaches of theAlchan River (Nevolina, 1990; Volynets, 2001; Voly-nets and Nevolina, 2003). It includes 43 taxa, dominantamong which are angiosperms frequently representedby large platanoid leaves of genera Platanophyllum,Pseudoprotophyllum and Paraprotophyllum. Commoncomponents are leaves of Araliaephyllum, “Schef-fleraephyllum” and Menispermites. The dicot genusQuereuxia is a newcomer. Representatives of generaPodozamites, Athrotaxopsis, Pityophyllum, andSequoia are abundant in subdominant group of coni-fers, which includes also less frequent species Taxites(“Cephalotaxopsis”) heterophyllus?, Sequoia reichen-bachii, Brachyphyllum douglasii, Elatocladus smitti-ana, and first representatives of the genus Thuja. Theassociated fern genera are Coniopteris, Gleichenites,Anemia, Sphenopteris and Cladophlebis. Leaves andgenerative organs of aquatic fern “Salvinia” have beenfound as well. Cycadophytes (Nilssonia sp.) and gink-goales (Ginkgo ex gr. adiantoides) are rare. Remains ofaquatic monocot “Potamogeton” sp. A are found as sin-gle specimens (Table 1, Fig. 2). Phyllopod shells aredetected in flora-bearing beds. Dominant angiosperms(48.8%) represent characteristic feature of the assem-blage.

Southern Primor’e, the Razdol’naya Depression

The Aptian–Cenomanian succession of the depres-sion is composed of coaliferous and volcaniclastic-sed-imentary deposits attributed to the Lipovtsy (Aptian)and Galenki (lower–initial upper Albian) formationsand to the undivided Korkino Group of the upperAlbian-Cenomanian (Nazarenko and Bazhanov, 1989;Resolutions…, 1994; Regional…, 2001). An abundantand diverse fossil flora collected from these depositsduring the last 20 years is well studied in taxonomicaspect and divided into six floral assemblages (Fig. 3).

The early Lipovtsy FA from the lower part of synon-ymous formation is known from outcrops of theRazdol’naya River and Murav’ev-Amurskii Peninsula(Volynets and Nevolina, 2003). Characteristic of theassemblage is a considerable taxonomic diversity(64 taxa). Dominants are ferns (25 species), abundantremains of Ruffordia goeppertii, Nathorstia pectinata,Alsophilites nipponensis, and “Polypodites” polysorusincluded, and cycadophytes. Most frequent among thelatter are Pterophyllum burejense, Nilssoniopteris rhit-idorachis, Zamiophyllum ivanovii, Cycadites sulcatus,

626


VOLYNETS

Ctenis yokoyamae, Nilssonia nicanica, and N. ex gr.brongniartii. Species Araucariodendron heterophyl-lum, Elatides asiatica, Podocarpus harrisii, Athrotax-ites berryi and Brachyphyllum japonicum prevail in thegroup of conifers. Remains of caytoniales, ginkgoales,bryophytes, lycopods and horsetails are rare (Table 1).Predominance of ferns (40.5%) and cycadophytes

(32.8%) associated with a considerable proportion ofconifers (Fig. 3) is diagnostic feature of the assem-blage.

The late Lipovtsy FA is typical of the upper part ofthe Lipovtsy Formations in the Razdol’naya Riverbasin and Murav’ev-Amurskii Peninsula (Volynets andNevolina, 2003). It is more diverse than the previousone (88 taxa). Ferns dominant in the assemblage(32 species) are most frequently represented by speciesof genera Gleichenites, Onychiopsis, Dicksonia,Coniopteris, Nathorstia, Eboracia, and Lobifolia.Newcomers among ferns are “young” generaOsmunda, Birisia, Anemia and Arctopteris. Subdomi-nant plant remains belong to cycadophytes (21 species)and conifers (18 species). Species Nilssoniopteris rhit-idorachis, Cycadites sulcatus, Zamites borealis, Zami-ophyllum ivanovii, Dictyozamites grossinervis, D. cor-datus, Pterophyllum sutschanense, and Pseudocteniseathiensis are abundant among cycadophytes. Espe-cially abundant are Nilssonia densinervis, N. canaden-sis, N. ex gr. orientalis, N. nicanica, N. prynadii,N. mediana, and N. ex gr. brongniartii. Representativesof genera Athrotaxopsis, Athrotaxites, Elatides,Podocarpus, Brachyphyllum, and Podozamites fre-quently occur in the group of conifers. Remains of cay-toniales (Sagenopteris), ginkgoales (Ginkgo, Baiera,Pseudotorellia), czekanowskiales (Phoenicopsis, Cze-kanowskia), lycopods (Lycopodites), horsetails (Equi-setum) and Zamiopsis of unclear taxonomic affinity arerare (Table 1, Fig. 3). The increased taxonomic diver-sity distinguishing the FA is determined by presence of“young” genera Osmunda, Birisia, Anemia, Sequoiaand by appearance of new czekanowskiales.

The early Galenki FA from the lower part of synon-ymous formation is established in the Razdol’nayaRiver basin and Murav’ev-Amurskii Peninsula (Voly-nets and Nevolina, 2003). It is not very diverse(31 taxa), dominated by cycadophytes and ferns. Cyca-dophytes are represented by genera Dictyozamites,Pterophyllum, Leptopterophyllum, and Nilssonia. Mostabundant species of this group are Dictyozamites cor-datus, Leptopterophyllum pterophylloides, Nilssonia.densinervis, and N. ex gr. orientalis. Abundant amongferns are Onychiopsis psilotoides, Teilhardia tenellaand Anemia dicksoniana. The subdominant group ofginkgoales includes abundant Ginkgo ex gr. adian-toides, but specimens of G. concinna are rare. SpeciesLycopodites prynadae prevails among lycopods,Podozamites tenuinervis among conifers, and Sagenopt-eris aff. mantellii among remains of caytoniales (Table 1,Fig. 3). Distinctive features of the FA are predominanceof Early Cretaceous cycadophytes (30 %) and higherproportion of “young” fern genera.

The middle Galenki FA is established in the middleof the same formation in exposures of the Razdol’nayaand Barabashevka rivers and Murav’ev-Amurskii Pen-insula (Volynets, Nevolina, 2003). This FA is quitediverse (69 taxa) and dominated by ferns (27 taxa). The

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Fig. 3. Percentages of different plant groups in Cretaceousflora of the Razdol’naya depression (symbols as in Fig. 2).Floral assemblages by names: (1) early Lipovtsy; (2) lateLipovtsy; (3) early Galenki; (4) middle Galenki; (5) lateGalenki; (6) early Korkino.



abundant fern species are Birisia alata, Anemia dickso-niana, Dicksonia concinna, Lobifolia novopokrovskii,and representatives of the genus Adiantopteris (4 spe-cies). Species most frequently occurring in the group ofsubdominant conifers (14 taxa) are Podozamites tenu-inervis, Taxites brevifolius and Sequoia reichenbachii.The group of cycadophytes (10 taxa) persistentlyincludes abundant remains of Neozamites denticulatus,Pterophyllum sutschanense, Nilssonia canadensis andN. ex gr. orientalis. Genera Sapindopsis, Dicotylophyl-lum and Cercidiphyllum are common among dicots,whereas Laurophyllum is rare. Single remains representPseudotorellia sp., Sujfunophyllum dichotomum,monocot Pandanophyllum ahnertii, and fruits Onoanaand Nyssidium (Table 1). Dominant ferns (39.2%) andconifers (20.4%), a considerable part of which is repre-sented by genera Birisia, Anemia, Dicksonia, Adian-topteris, Taxites, and Sequoia, and appearance of newangiosperm taxa (Fig. 3) characterize specific composi-tion of the assemblage.

The late Galenki FA is established in theRazdol’naya and Amba river basins on the Murav’ev-Amurskii Peninsula and near the Lake Khanka, whereit is confined to the upper part of synonymous forma-tion (Volynets, Nevolina, 2003). Being of diverse com-position (86 taxa), the assemblage is dominated byferns (27 taxa) and cycadophytes (25 taxa). The mostdiverse fern genera are Coniopteris, Anemia andOsmunda. Fern species occurring frequently areOsmunda denticulata, Anemia dicksoniana, Gleicheni-tes porsildii, Birisia oerstedtii, Cyathea aff. tyrmensis,Eogymnocarpium sinense and Lobifolia novopok-rovskii. Representatives of the genus Temskya are rare.Species Pterophyllum burejense, P. sutschanense, Lep-topterophyllum pterophylloides, Encephalartopsisvachrameevii, Nilssonia canadensis, N. ex gr. brongn-iartii, and appeared representatives of the genus Ctenis(4 species) are most abundant in the group of cycado-phytes. Among conifers (13 taxa), there are frequentPodozamites tenuinervis, Sequoia reichenbachii, Tax-ites acuminatus, T. brevifolius and Athrotaxopsisexpansa. Taxa listed above occur in association withextremely rare remains of caytoniales, ginkgoales andflowering plants (Table 1). Taxonomic diversity charac-teristic of the assemblage is determined by diversifiedferns (31.2%) and cycadophytes (29.1%), whereasabundance of conifers is lower (15.1%) andangiosperms represent a minor part (3.5%) of all plantremains (Fig. 3).

The early Korkino FA from the lower part of undi-vided synonymous group is established in the AmbaRiver basin. Being not diverse (20 taxa), it is dominatedby ferns (9 taxa), the most frequent species Gleicheni-tes porsildii included. Sequoia reichenbachii is mostabundant species in the group of subdominant conifers(4 taxa). Remains of lycopods, horsetails, cycado-phytes, ginkgoales and angiosperms are rare (Table 1).The FA exemplifies a sharp diversity decrease in all

the groups of higher plants and rarity of angiosperms(Fig. 3).

Partizanskaya Depression

The Aptian–Cenomanian continental coaliferous,marine and volcanogenic-terrigenous deposits of thedepression correspond to succession of the StaryiSuchan (Aptian), Severnyi Suchan (lower–initial mid-dle Albian), Frentsevo (middle Albian), Kangauz(upper Albian), Romanovka (upper Albian), Brovnichi(upper Albian–lower Cenomanian) and Dadanshan(upper Cenomanian) formations (Volynets, 1998; Voly-nets et al., 2001; Regional…, 2001). The depositsyielded abundant and diverse plant remains, which aredivided into eight floral assemblages after thoroughtaxonomic investigation.

The late Staryi Suchan FA confined to the upper partof synonymous formation is known from upper reachesof the Partizanskaya River (Volynets, 1998; Volynets,Nevolina, 2003). Characteristic of the FA is a moderatetaxonomic diversity (36 taxa) and predominance offerns (13 taxa). Species frequently occurring among thelatter are Gleichenites gieseckianus, Onychiopsis psilo-toides, Dicksonia concinna and Anemia dicksoniana.Subdominant conifers (8 taxa) are represented mostlyby Elatides asiatica, Athrotaxopsis expansa andSequoia reichenbachii. Species Nilssonia ex gr.brongniartii is abundant among cycadophytes (5 taxa).Horsetails (especially Equisetum cf. ramosus), cayto-niales and lycopsids are rare. The FA includes singlespecimens of czekanowskiales (Leptostrobus mollis)and ginkgoales (Pseudotorellia ex gr. angustifolia).Dominant ferns (36%), subdominant conifers (22.2%),and appearance of “young” species Anemia dicksoni-ana and Sequoia reichenbachii are the distinctive fea-tures of the FA (Table 1, Fig. 4).

The early Severnyi Suchan FA from the lower partof synonymous formation has been studied in middlereaches of the Partizanskaya River (Volynets and Nev-olina, 2003). It includes 33 taxa, 14 of them represent-ing dominant ferns. The most abundant fern species areOsmunda denticulata, Gleichenites porsildii, Anemiadicksoniana and Birisia onychioides, whereas Birisiaalata, Coniopteris asplenioides, Teilhardia tenella andCladophlebis virginiensis are rare. Subdominant coni-fers (8 taxa) are represented primarily by abundant spe-cies Athrotaxopsis expansa and Elatides asiatica, whileTaxites brevifolius and Araucariodendron oblongifo-lium occur in contrast as single specimens. Dominantamong cycadophytes (6 taxa) are Nilssonia canadensis.Equisetum ex gr. burejensis is frequently occurring spe-cies of horsetails. Among 2 taxa of ginkgoales, themore frequent taxon is Ginkgo ex gr. adiantoides(Table 1). Specifics of the FA consists in dominance offerns (42.4%) and conifers (24.2%) and in perceptibleproportion of remains representing genera Osmunda,Anemia and Athrotaxopsis, whereas Taxites species arerare (Fig. 4).

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The late Severnyi Suchan FA known from the Parti-zanskaya River basin is confined to the upper part of thesame formation (Volynets and Nevolina, 2003). Thismoderately diverse assemblage of 46 taxa is again dom-inated by ferns (18 taxa), the abundant species of whichare represented by Osmunda denticulata, Dicksoniaconcinna, Birisia onychioides, Gleichenites zippei,Anemia dicksoniana and Cladophlebis frigida. Newthough rare fern species, which augment diversity ofthis group, are Coniopteris asplenioides, C. setacea,Gleichenites zippei, Birisia ochotica, Anemia sutschan-ica, Arctopteris obtusipinnatus, Lobifolia novopok-

rovskii and Pelletieria ussuriensis. Species abundant inthe group of subdominant conifers (14 taxa) arePodozamites ex gr. lanceolatus, Elatides asiatica, Ath-rotaxites berryi, Pseudolarix dorofeevii and Schizolepisneimengensis, whereas Podozamites tenuinervis, Tor-reyites dicksonioides, Sequoia reichenbachii, Pityocla-dus pseudolarixoides, Taxites brevifolius, and Schizole-pis cretacea occur less frequently. Among cycado-phytes (6 taxa), frequent species are Pterophyllumsutschanense and Nilssonia ex gr. brongniartii, whileNilssonia densinervis and plant remains from generaAnomozamites and Otozamites are rare. Similarly rare

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Fig. 4. Percentages of different plant groups in Cretaceous flora of the Partizanskaya depression (symbols as in Fig. 2). Floral assem-blages by names: (1) late Staryi Suchan; (2) early Severnyi Suchan; (3) late Severnyi Suchan; (4) Frentsevo (early subassemblage);(5) Frentsevo (late subassemblage); (6) Kangauz; (7) Romanovka; (8) Brovnichi; (9) Dadanshan.



are ginkgoales Ginkgo ex gr. adiantoides and Sphe-nobaiera sp. The other persistent components ofassemblage are horsetails (3 taxa), single Leptostrobusand Dicotylophyllum sp. representing the early flower-ing plants (Table 1, Fig. 4). Osmunda, Anemia, Birisia,Arctopteris, Torreyites, and Sequoia, the evolutionaryadvanced fern and conifer genera, characterize, alongwith appearance of flowering Dicotylophyllum sp., spe-cific character of the assemblage.

The Frentsevo FA from synonymous formation isknown from northern and central areas of the Partizan-skaya River basin, eastern coast of the Bay of Ussuri-isk, and from exposures of the Sukhodol River (Voly-nets and Nevolina, 2003). This rather diverse assem-blage of 55 taxa is dominated by ferns (18 taxa),including their abundant species Osmunda denticulata,Anemia dicksoniana, Birisia onychioides, Vargolopt-eris rossica, Teilhardia tenella and Lobifolia novopok-rovskii. The less abundant fern species are Anemia sut-schanica, Birisia alata and B. ochotica. The group ofsubdominant conifers (13 taxa) includes Athrotaxopsisexpansa, Athrotaxites berryi, Sequoia ambigua and S.reichenbachii occurring frequently and associated withless frequent Podozamites tenuinervis, Elatocladusobtusifolius and Sphenolepis kurriana. Abundant Nils-sonia densinervis is main species among 5 taxa of cyc-adophytes. Remains of ginkgoales are extremely rare,mainly represented by Ginkgo pluripartita. Lycopsidspecies Lycopodites prynadae and L. ovatus are persis-tent components of the assemblage. Single representa-tives of bryophytes and horsetails are present as well.Rare plant remains represent flowering taxa “Magnoli-aephyllum” sp., Dicotylophyllum sp. A, Sapindopsisvariabilis, Sassafras ussuriensis, Artocarpidium sp.,Laurophyllum sp. and fruits Nyssidium, Kenella andCarpolithes (Table 1, Fig. 4). Thus, the FA is dominatedby ferns (35.1%) and conifers (22.8%), which are asso-ciated with less abundant cycadophytes (8.8%) andearly angiosperms (14%). The FA can be divided intothe early and late subassemblages. The early one con-fined to the lower part of the Frentsevo Formation con-sists of fern and gymnosperm typical of the assemblagein general, but it is lacking angiosperms. The late sub-assemblage from the formation upper part includes thesame taxa of ferns and gymnosperms, which are asso-ciated, however, with species of early flowering plantslisted above (Table 1). Konovalov who studied mollus-can shells (Table 3) from the “Nadvelikanovskii Hori-zon” of the formation identified species Quadratotrigo-nia (Transitrigonia) fudsinensis Mirol., Pterotrigoniahokkaidoana (Yeh.), Ussuritrigonia subpyriformisKonov., and “Callista” pseudoplana Yabe et Nagao,which are of the Albian age (Markevich et al., 2000).

The Kangauz FA of synonymous formation has beenestablished in exposures of the Partizanskaya River andeastern coast of the Bay of Ussuriisk (Volynets andNevolina, 2003). This moderately diverse assemblageof 48 taxa includes fens and conifers in equal propor-tions (13% each). The dominant fern species are Birisia

alata, B. ochotica and Anemia dicksoniana. Prevailingspecies of conifers are Athrotaxites berryi and Sphenol-epis sternbergiana. The associated remains of flower-ing plants (10 taxa) are represented by leaves of Sapin-dopsis variabilis, Aralia lucifera and Vitiphyllum (Cis-sites) parvifolium, and fruits Kenella sp. Lycopsids ofthe assemblage (5 taxa) belong to genera Lycopodites,Isoetites, and Synlycostrobus. Species Dictyozamitessp. and Nilssonia sp. of cycadophytes and generaMarchantites and Thallites of bryophytes are rare(Table 1, Fig. 4). Thus, a considerable proportion ofearly diverse flowering taxa (20.8%) is a characteristicfeature of the assemblage. It was formerly suggested(Krishtofovich, 1929; Krassilov, 1967) that the well-known flora containing Aralia lucifera is confined tothe Suchan Group of the Tret’ya Kamenka River basin.However, Konovalov who reexamined the successionof flora-bearing beds established that this flora is actu-ally from the Kangauz Formation of the Korkino Group(Konovalov 1964; Markevich et al., 2000). On the leftside of the Tigrovaya River, lower beds of this forma-tion contain shells of Aucellina caucasica (Buch.). Inaddition, A.A. Yakushina identified Albian bivalves andgastropods in the same formation (Markevich et al.,2000).

The Romanovka FA from synonymous formationhas been discovered in the Partizanskaya riverhead area(Volynets and Nevolina, 2003; Oleinikov et al., 1989).Being of a low diversity (14 taxa), this assemblage isdominated by conifers of genera Elatides, Elatocladus,and Pityophyllum, the most frequent species Elatidesasiatica and Pityophyllum ex gr. nordenskioldiiincluded. Ferns (Gleichenites, Anemia, Onychiopsis)and cycadophytes, predominantly Otozamites sp., areless abundant. The associated remains of caytoniales(Sagenopteris) occur as single specimens. Representa-tives of “young” genera Menispermites and Tetracen-ton appear in the group of flowering plants (Table 1). Ingeneral, ferns and gymnosperms are of a sharplyreduced taxonomic diversity in this FA demonstratingappearance of evolutionary advanced taxa in the groupof flowering plants (Fig. 4).

The Brovnichi FA from outcrops of the Partizan-skaya River and Serebryanka Creek is confined to theeponymous formation (Volynets et al., 2001; Volynetsand Nevolina, 2003). Dominant among 31 taxa of theassemblage are angiosperms (13 taxa) frequently repre-sented by species of genera Cercidiphyllum, Araliae-phyllum, Sapindopsis, and occasionally by Platano-phyllum remains. Subdominant conifers (9 taxa),include abundant Athrotaxopsis expansa and Brachy-phyllum sp. Associated ferns (6 taxa) belong to generaRuffordia, Gleichenites and Birisia, the last of which ismore diverse than others (three species). SpeciesOtozamites aff. schenkii is most frequent among cyca-dophytes (Table 1). Specific aspect of the FA consists ina high proportion (41.9%) of diverse flowering plants(Fig. 4).

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The Dadanshan FA is characteristic of the epony-mous formation in middle reaches of the PartizanskayaRiver (Volynets and Nevolina, 2003). It consists of35 taxa, which represent dominant angiosperms(19 taxa), subdominant conifers (9 taxa), and associ-ated ferns and ginkgoales. Most frequent fern speciesare Anemia dicksoniana and Birisia oerstedtii, whereasGleichenites, Birisia cf. samylinae and Cladophlebisare rare. In addition to abundant Sequoia reichenbachii,the group of conifers includes single representatives ofgenera Podozamites, Brachyphyllum, Elatocladus, Tax-ites, and Torreyites. Platanaceae of genera “Platanus,”Platanophyllum, Protophyllum, and Credneria, theabundant Platanus newberryana, Protophyllum sp. andAraliaephyllum sp. 1 included prevail over otherangiosperm taxa. Among the latter, there are small-leaved Celastrophyllum sp., Dicotylophyllum sp. 1 andDicotylophyllum sp. Species Araliaephyllum samar-gense, Menispermites sp., Cissites aff. uralensis, andCissites sp. are of a low abundance (Table 1). Theassemblage is distinctly dominated by angiosperms(54.3%), mainly by Platanaceae (Fig. 4). Concurrentfauna is represented by fossil fish fins and fresh-watermollusks, which are of the Albian-Cenomanian ageaccording to determinations of Yakushina (1964).

All the floral assemblages considered above eluci-date general trends in evolution of the Albian floras.Ferns dominated in the floras almost throughout theAlbian, while gymnosperms were of subordinate sig-nificance during this period and proportions ofangiosperms grew with time.

In conclusion, it is necessary to note some peculiar-ities, which discriminate the Middle Cretaceous flora ofthe Alchan depression from other concurrent floras.

Typical of the flora under consideration are mixing ofnorthern and southern plant taxa, diversity of cycado-phytes, and abundance of Taxodiaceae among conifers.

ACKNOWLEDGMENTS

The work was supported by programs of Presid-ium of the Russian Academy of Sciences, projectnos. 04-1-P12-009 and 04-1-P25-053.

Reviewers A.B. Herman, V.S. Markevich,and M.A. Akhmet’ev

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Table 3. Stratigraphic chart and floral assemblages of the Partizanskaya depression

Biostratigraphic subdivisions Lithostratigraphic subdivisions

Stage Substage Fauna-bearing beds (Markevich et al., 2000) Floral assemblages Subassem-

blages Group Formation

Cenoma-nian

Upper Dadanshan Dadanshan

LowerBrovnichi

Korkino

Brovnichi

Albian

UpperRomanovka Romanovka

Aucellina caucasica Kangauz Kangauz

Quadratotrigonia, Pterotrigo-nia, Ussuritrigonia, “Callista,” Isognomon, Lima et al.

FrentsevoLate

Suchan

Frentsevo

MiddleEarly

Late Severnyi SuchanSevernyi Suchan

Lower Early Severnyi Suchan

Late Staryi Suchan Staryi Suchan(upper)Aptian Upper



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Documents

The Aptian–Cenomanian Flora of Primor’e, Part 1: Floral Assemblages