J . L i n n . 8 0 ~(Bot.),61,384,pp.
.
207-218
With 1 plate and 3 tezt-jigures
Printed i n Great Britain
July, 1968
Early Cretaceous angiosperms of the Soviet Union
based on leaf and fruit remains
BY V. A. SAMYLINA
Palaeobotanical Laboratory, Komarov Botanical
Institute of the Academy of Sciences, U.S.S.R.,
Leningrad
Five newly collected Early Cretaceous angiosperms from the Kolyma River in Siberia are
described. There are two new species of leaf, Cinnamomoides ievlevii and Celastrophyllum
aerrulatus and anew genus of fruit, Kenella harrisiana together with two leaves, Nelumbites aff.
minimus Vachrameev and Celastrophyllum aff. hunteri Ward. Early Cretaceous angiosperm
records for the Soviet Union are given in a table and their signifieance is discussed. It is
argucd that by the end of the Early Cretaceous whcn the flora underwent a substantial change
angiosperms were already distinguished by their high state of morphological development
judging by leaf form and systematic composition. They were widely spread occurring in
regions of both temperate and warm humid climates. They appear to have been plants of
mountains, including trees and herbs, terrestrial and aquatic. With the advent of physical and
climatic changes these angiosperms were ready for rapid expansion. Their domination of the
vegetation of the Late Cretaceous was not as inexplicable as previously thought.
INTRODUCTION
All discoveries of new angiosperms from floras which are older than those dominated by
angiosperms of the Late Cretaceous are interesting and valuable as we know too little of
their early history. I n this paper some new forms of angiosperm leaf and fruits are described
from the Albian deposits from the east bank of the Kolyma River in Eastern Siberia. This
material was collected by the geologists S. I. Filatov, L. V. Ievlev together with the
author. These specimens were studied and are kept in the Komarov Botanical Institute in
Leningrad.
SYSTEMATIC DESCRIPTION
Genus Cinnamomoides Seward, 1925
Cinnamomoides ievlevii sp. nov.
(Pl. 1, figs 1 4 ; Text-fig. 1A-C)
Holotype. No. 511 : 1, Palaeobotanical Laboratory, Komarov Botanical Institute,
Leningrad.
Paratypes.Nos511:2-8,517:
1,2.
Locus typicus.,Omsukchan,Sugoi River, Kolyma basin, East Siberia.
Stratus typicus. Toptanskaja suite :Late Albian.
Name. This speciesis named after L. V. Ievlev.
Diagnosis. Leaves petiolate, ovate, entire with smooth margins; rounded t o tapering a t
the base, tapered acuminate a t the apex. Blade size 2 to 7 cm long by 0.7 to 4.5 cm wide,
petiole about one-third the length of the blade. Venation, trinerved; midrib, straight
reaching well into the apical region. Lateral primaries, opposite, inserted a t c . 40" to
208
V. A. ~AMYLINA
midrib, 1 to 2 mm above the base, curving upwards and running two-thirds to threequarters the length of the leaf. Secondaries, a t least two to three alternate pairs, inserted
30 to 35" to the midrib in the upper half of the leaf, curving upwards. I n small leaves,
primaries nearly parallel to the margin. I n larger leaves, lateral primaries give off a series of
camptodromous veins on their outer sides, these form fairly large loops. Finer veins not
preserved.
Description. These leaves vary in size from the smallest, which is 19 mm long by 7 mm
wide; its greatest width is in the basal third of the leaf. Among fairly large leaves there is
one about 6 cm long by 3 cm wide. The maximum width of the largest leaf of which only the
lower half is preserved, amounts to 4.5 cm. Petioles appear to be long: in one of the largest
leaves the petiole fragment preserved attains 18 mm. The smaller leaves had proportionally shorter petioles.
Comparison. The genus Cinnamomoides was established by Seward for leaves whose
venation resembled that of Cinnamomum leaves, in particular C. camphora. But the present
leaves cannot be attributed to the family Lauraceae with any certainty because of the
absence of finer featyres of agreement. One may note that in leaves of most living species of
Cinnamomum lateral primaries are given off higher than in those of Cinnamomoides. This
feature brings them nearer to certain species of Lindera (for instance, L. fruticosa Hemsl.
and L. strichnifolia Vill.). The most characteristic features of the new leaves described here
are : the ovate outline, comparatively small size of the blade and, abaxial looping branches
of the primary laterals. C. ievlewii is closely related t o C. owalis (Dawson) Bell from nearly
contcmporaneous sediments in Western Canada. The leaves of C . owalis are distinguished
by their elongate-elliptical outline and primary laterals running parallel to the margin.
Genus Nelumbites Berry, 1911
Nelumbites aff. minimus Vachrameev
(Pl. 1, figs 14, 15; Text-fig. 1D-E)
Material. 511 : 10 A, B.
Horizon and Locality. Toptanskaja suite : Late Albian ; Omsukchan, Sugoi River,
Kolyma basin, East Siberia.
Description. There are impressions (counterparts) of two leafy shoots and some incomplete detached leaves. They are all preserved on a single piece of the black aleurolite.
The shoots are thin, about 1 to 1.2 mm in diameter. The leaves are peltate, small, oval to
almost orbicular, with smooth or slightly undulate margins. The largest completely
preserved leaf (Text-fig.1 E) is 16 mm in length by 13 mm in width. The petiole is attached
to the lower surface of the leaf, not far from the lower margin; it is thin (less than 0.5 nini
wide), up to 12 mm long. The venation is palmate, camptodromous but the veins are
difficult to distinguish, thin, generally seven to nine primaries, radiating from the top of
petiole to the margin. The median primary is straight, non-dichotomising and gives off
two or three pairs of secondaries. Two or three primaries also do not dichotomise, running
downwards. The remaining veins divide dichotomously one to three times close to margin.
Near the impressions of leafy shoots but not organically connected with them, there is an
impression of the female structure, preserved as if in longitudinal section and resembling
an overturned triangle with a concave base, rounded angles and a tapered apex, that
continues into the petiole. Judging by the coaly lustre of the impression and sculpture, this
structure was fleshy and contained elongated bodies, arranged vertically. The structure
described may be interpreted as an impression of a n inversally conical, accrescent receptacle with fruits inside, like those of Nelumbo.
Comparison. I n size, outline and venation the leaves described much resemblc those of
Nelumbites minimus from the Middle Albian deposits of Western Kazachstan (Vakhrameev, 1952).A single distinguishing character of Kolymean leaves is a greater number of
Early Cretaceous angiosperms o j U.S.S.R.
209
primaries and, accordingly, a lesser angle between them. This character makcs them more
like N . tenuinervis (Font.) Berry from Patapsco formation of the Atlantic coast, U.S.A.,
but leaves of t,helatter arc larger.
B
b
Text-fig. 1. A-C. Cimnamomdes ievlevii sp. nov. A, Loaves of avoragesize, holotypo, no. 51 1 :
1 ; x 1. B, C, small loaves, partttypesnos 5 1 1: 4-5. B x 2, C x 1 .
D-E. Nelurtahites aff. minimus Vachrameov. D, Leafy shoots, tlctached leaves and R fruit, no.
511: 10,x 1.E,TholcafintheccntroofD.x 2.
Genus Celastrophyllum Goeppert, 1854
Celastophyllum aff. hunteri Ward
(Pl. 1 , figs 8-10; Text-fig. 2A-D)
Materinl. 517 : 11-22.
Horizon and lomlity. Toptanskaja suite : Late Albian; Omsukchan, Sugoi River, Kolyma
basin, East Siberia.
Description. Entire, elongate, tapered leaves, acuminate a t the apex and decurrent,
cuneate a t the base. The margins are large-serrate, but entire in the basal region. The size of
leaves is variable. Average size 5 to 6 cm by 1 to 1.5 cm. There are small leaves : 2.5 to 3 cm
14
210
V. A. SAMYLINA
long by 7 to 8 mm wide (PI.1, fig. 8) ; the largest leaf is about 8 cm long by about 2 cm wide
(Text-fig.2 C).Venation, pinnate. Secondaries and finer veins, thin, hardly distinguisl~ablc.
Secondaries, attached a t 30 to 45" to the midrib, curving upwards and dividing two or
three times.
Comparison. Among the known species of Celastrophyllum, the impressions described
resemble most of all C . hunteri Ward from the Patapsco formation of the Atlantic coast,
U.S.A. Detailed comparison of Kolymean and American specimens is difficult as the
preservation of both is poor. In correlation between length and width Kolymean leaves are
comparable with C .albaeformis Ward which is also from the Patapsco formation, but leaves
of the latter have a rounded apex and base. I n outline of blade and marginal features there
is a resemblance with leaves of the same poor preservation, described by W. A. Bell (1056)
as C . acutidens Font. from the Albian deposits of Western Canada. The differcncc from C .
serrulatus from the Kolymean sediments will be given further in the description of the
latter.
Celastrophyllum serrulatue sp. nov.
(PI. 1, figs 5-7; Text-fig. 2E-F)
Holotype. No. 51 1 : 11, Palaeobotanical Laboratory, Komarov Botanical Institute,
Leningrad.
Paratypes. Nos 511 : 12-22.
Locus typicus. Omsukchan, Sugoi River, Kolyma Basin, East Siberia.
Stratus typicus. Toptanskaja suite : Late Albian.
Diagnosis. Leaves entire, oval, short petiolate, with nearly acuminate apex and a
decurrent cuneate base; margin serrulate but entire a t extreme base; bladc up to 0 cm in
length by 4 cm in width. Venation pinnate, camptodronious; secondaries once or twico
divided, 10 to 12 pairs in number, attached a t about 45" to midrib.
Description. The leaves are medium size to small. The largest leaf (incomplete)was about
6 cm long by about 4 cm wide. More often the leaves are 3.5 to 4 cm long by 1.8 to 2.3 cm
wide. Three leaves are preserved attached to a shoot o f 4 mm diameter by a short petiole
about 5 mm long.
Near to the margin the secondaries curve upward somewhat and dichotomise once or
twice to meet secondaries next above and to form loops. Tertiaries are very thin, and
indistinct, leaving secondaries a t a fairly open angle (70 to 80").
Comparison. The most characteristic features in which C . serrulatus differs from other
species of Celastrophyllum, are the serrulate margin and crowded secondaries. These
features are present in all the leaves, independently of their size. I n size and outline the
Omsukchanean leaves most resemble C. newberrianum Hollick (in Newberry, 1805) from
the Upper Cretaceous deposits of U.S.A. (New Jersey). C. aff. hunteri described here from
the same deposits differs in its narrower leaves, larger marginal serrations nnd lesser
attachment angle of secondaries.
PLATE
1
Figs 1-4. Cinnummoides ievleuii sp. nov. Leaves. Fig. 1. Holotype no. 511 : 1. Figs 2-4. Paratypes
nos511 : 7,4,2. All x 1.
Figs 5-7. Celastrophyllumserrulatussp. nov. Leaves. Fig. 6. Holotype no. 511: 11. Figs. 5, 7. Paratypes
nos 12,13. All x 1.
Figs 8-10. Cehstrophyllumaff. hunteri Ward. Leaves, nos 517: 9,7,5.All x 1.
Figs 11-13. Kenella havisiann gen. et sp. nov. Fruits. Fig. 13. Holotype no. 510: 1. Figs 1 1 , 12.
Paratypesnos 510: 3. Figs 11.13, x 1 ; Fig. 12, x 2.
Fig8 14,15. Nelumbites aff. minimus Vachrameev.Fig. 14. Leafy shoot and detached leaves, no. 51 1 : 10.
Fig. 16.TheleafmarkedwiththearrowinFig. l4.Fig. 14, x 1;fig. 15,x2.
Plate 1
Early Cretaceoua angiosperms of U.S.S.R.
E
~
(.\
Ii
21 1
F
!
D
Toxt-fig. 2. A-D. Celostro/,lLyllumaff. kunteri Ward. Incomplete leaves, nos517: 5, 3, 6, 10. x 1.
E-F. Celastrophylluinserrulutus sp. nov. F, Holotype leaf, no. 51 1 : 11 ;x 1 . E, Paratype no.
511: 12; x 1. G.KeneZlohclrrisianagen.etsp.nov.Holotypefruits,no.510:
1 ; x 2.
Angiosperm of uncertain aginities
Genus Kenella gen. nov.
Diagnosis. Fruit ; small, elongate with a tapered apex and elongate poorly developed
surface ribs.
Type species. Kenella harrisiana sp. nov.
Discussion. This reproductive structurc, established as new genus, is considered to be an
angiosperm fruit, belonging presumably to the Dicotyledons. It must be mentioned that
angiosperm leaves were not collected from the outcrops where the fruits occurred, although
their presence is probable since a leaf impression of Dalbergites sp. was found in another
outcrop of the same deposit and a great number of angiosperm leaves were found in nearly
contemporaneous deposits of the Buor-Kemiusskaja suite on the left bank of the Kolyina
River (Zyrianka-Syliap coal-fields). The fruits described have characteristic features and
may be easily distinguished. Among the fruits of living plants closely related ones to these
are unknown to me. Of the fossil genera one must point out certain resemblance with
fruits of Nyssidium from Cretaceous deposits. The latter, however, have more clearly
pronounced and prominent ribs and a rounded or nearly rounded apex and their surface is
never covered with bristles. There is a certain resemblance in general fruit outline and in
the presence of elongated ribs with fruit impressions described by Kryshtofovich (1958)as
Quereuxia aculeata. But these are smaller than Kenella and have a t least two spreading
spines at the apex.
14*
212
V. A. SAMYLINA
Kenella h r r i e i a n a sp. nov.
(Pl. 1, figs 11-13; Text-fig. 2 G)
Holotype. No. 510 : 1, Palaeobotanical Laboratory, Komarov Botanical Institute,
Leningrad.
Paratypes. Nos 510: 2 4 .
Locus typicus. Omsukchan, Ken River, Kolyma River, East Siberia.
Stratus typicus. Omsukchanskaja suite :Albian.
Name. This speciesis named after Professor T. M. Harris.
Diagnosis. Fruits, elongate-ovate about 16 mm in length by 5 mm in maximum width,
with three or four elongate ribs on the surface of the impression, covered by thin crowded
bristles.
Description. Fruits small, elongate-ovate, about 16 mm in length by about 5 mm in
greatest width which is in the basal third of the fruit, gradually becoming narrow a t the
base and apex. The extreme top of the apex and the stalk of the fruit are never preserved.
The fruit surfaces have poorly developed elongate ribs (three or four on a n impression)
which come nearer to each other a t the apex. Most probably, these ribs are strongly
developed vascular bundles. I n some specimens one may observe that neighbouring ribs
become connected by very thin transverse veins. The surface of fruits (probably the ribs
only) bear numerous thin bristles about 1 mm long. Judging by the fact that in two cases
the fruit remains are not flat but bulky rtnd are actually moulds, it is likely that fruits were
also bulky and polygonal (with six t o eight angles) in cross-section when alive.
Comparison. The structures from the same deposits similar in appearance and differing
from K . hurrisiana chiefly in the absence of bristles on the surface are attributed by me to
the same genus, but probably represent a different species.
BRIEF SURVEY OF THE EARLY CRETACEOUS ANGIOSPERMS OF THE
SOVIET UNION AND DISCUSSION OF THE MATERIAL
The apparently sudden appearance and rapid distribution of angiosperms during the
Late Cretaceous, may be chiefly attributed to the fact that we know too little of the Early
Cretaceous angiosperms. In the whole world, there are so far known but few localities of
fossil floras of the end of the Early Cretaceous; this gives rise to a somewhat exaggerated
notion of limited distribution of angiosperms a t that time.
In the Soviet Union before the 1950’sonly a few forms of Early Cretaceous angiosperms
were known, i.e. Pandanophyllum ahnertii Krysht., Cissites prodromus Krysht. and Aralia
Zuciferera Krysht. from the South Primorye and Aralia kolymensis Krysht. from the Kolyma
basin. There was the leaf of Cissites from western Kasachstan. Stratigraphical investigations during the last 15 years have revealed a number of new sites where Early Cretaceous
angiosperms occurred in the Soviet Union. I n addition more complete collections were
made from the previously known localities (see the map). Outside of the U.S.S.R. in
Eurasia only in Portugal are there Lower Cretaceous deposits with angiosperms (Saporta,
1894 ; Teixeira, 1948). Accordingly, the area of distribution of the Lower Cretaceous
deposits from which angiosperm remains are known occur both from the Siberian and
Indo-European paleofloristic regions (see Vakhrameev, 1964); but within the latter they
occur only in areas outside the arid belt.
At present about 80 species of Early Cretaceous angiosperms are known from the Soviet
Union. In Table 1 a complete list of them is given with localities and ages, as well as the
reference t o the paper where the description may be found. The Table does not include the
names of problematical angiosperms or species of indisputable angiosperms from deposits
of insecurely esta.blishedage, possibly even Late Cretaceous.
As it is evident from the complete list given most Early Cretaceous angiosperms were
Early Cretaceous angiosperms of U.S.S.R.
213
identified as organ-genera or form-genera. This is normal since most living genera apparently did not exist a t that time. The names of certain living genera in the list (Sassafras,
Cercidiphyllunt, Aralia) to a considerable extent are attributed to paleobotanical tradition
of associating certain types of fossil leaves with the generic names of living plants, and bear
evidence not so much of the identity of the genus but of possible affinity. Most of the
angiospermous remains were collected in deposits of Albian age. The area of the earliest
occurrence of angiosperm remains is the Far East (South Primorye), whence some species
from pre-Albian deposits are known, i.e. two forms of Nissidium from Barrcmian and
Aralia lucifera, Onoana nicaiaica, Paitda,nophyllum ahnertii from Aptian. Among them
Nyssidiunt and Onoana are in the form of fruit impressions and Aralia and Pandanophyllum
as leaf impressions. The flora of Cercal in Portugal appears to be somewhat younger than
the remaining Albian floras with early angiosperms. Both the areas, the South Primorye
Test-fig.3. DistributionmapshowingtheregionsinEurasiawheredepoaitsofLower
Cretaceous
age with leaves and fruits of early Angiosperms occur.
.,The boundary between the Siberian and Indo-European paleofloristic regions after
Vakhrameev (1964). 0 , Areas with Lower Cretaceous deposits bearing angiosperm remains.
. ...
and Cercal, are included in the Indo-European paleofloristic region and located a t latitude
40 to 50" N. The future will show whether this fact reflects a regular phenomenon of the
earlier origin of angiospcrms a t southern latitudes (Axelrod, 1959) or is accidental. A t
present one may state that the known pre-Albian angiospermous remains do not have any
obvious morphologically primitive features, in which they would differ from Albian
angiosperms. As soon as we find fossil remains of obvious angiosperms we immediately
come across a great systematic diversity. I n addition, one should point out that the generic
composition of angiospcrms from different localities of Eurasia, even those from diverse
paleofloristic regions, differs but little between localities. There is every reason to think
that the fossil genera which have been found so far only in one of the paleofloristic regions
of Eurasia may occur in the other one. As regards the species composition of the Early
Cretaceous angiosperms almost every locality is specific in spite of latitudinal location.
Thus, in three known areas of distribution of the Lower Cretaceous deposits with angiosperms in the Indo-European region (they all are situated a t latitude 40 to 50" N) there
are no species of angiosperms in common. The absence of a large number of identical or
?Fieussp.
Prototrochodendroidesjncuticu Budantsev e t
Kiritchkova
Trochodeiulroides ( ? ) sp.
Family Platanaceae?
? Platanus sp.
? Protophyllum sp.
Family Moraceae
Artocarpidium sp.
Family hloraceaa?
Morophyllum denticulatum Budantsev et
Kiritchkova
?Ficustschuschlzakulensis Vachrameev
C . sujfuneme Krassilov
Cercidiphyllum sp.
Family Ranunculaceae?
Raiiunculaecarpus quinquecurpellutusSamylina
aif. Ranunculaecarpusap.
Family Cercidiphyllaceae
Cercidiphyllum potmnacense (Ward)
Vachrameev
N . aff.minimus Vachrameev
N . tenuivervis (Fontaine)Berry
Dicotyledones
Family Lauraceae
Sassafras kolymnsis (Kryshtofovich)
Baikovskaja
S. ussuriensis Krassilov
Cinnamamoides ievlevii sp. nov.
Laurophyllum sp.
Family Lauraceae?
Laurophyllum? sp.
Family Xelumbonaceae
Nelumbites minimus Vakhrameev
Sames of plants
Krassilov, 1967
Here
Krassilov. 1967
Only the name in : Ifiitchkova & Slastenov, 1966
Vakhrameev, 1952
Suchan, South Primorye, Albian
Omsukchan, Kolyma basin, Late Albian
Sujfun, South Primorye, Albian
Lepiske River, Lena basin, Late Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Omsukchan, Kolyma basin, Late Albian
Chushkakul Mountains, Western Kazachstan,
&fiddleAlbian
Vakhrameev, 1952; Samylina, 1960
Kiritchkova & Slastenov, 1966
Kiritchkova& Budantsev, 1967
Only the name, here
Samylina, 1960
Krassilov, 1967
Kiritchkova & Budantsev, 1967
Vakhrameev, 1952
Chushkakul Mountains, Western Kazachstan,
Middle Albian; Zyrianka River, Kolyma
basin, Albian
Sujfun, South Primorye, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Lepiske River, Lena basin, Albian
Sitte River, Lena basin, Albian
Omsukchan, Kolyma basin, Late Albian
Zyrianka River, Kolyma basin, Albian
Suchan, South Primorye, Albian
Sitte River, Lenabasin, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Lepiske River, Lena basin, Late Albian
Only the name in: Kiritchkova & Slastenov, 1966
Krassilov, 1967
Vakhrameev, 1952
Samylina, 1960
Only the name in: Kiritchkova& Slastenov, 1966.
Zyrianka River, Kolyma basin, Albian
Lepiske River, Lena basin, Late Albian
Here
Vakhrameev, 1952
Kryshtofovich, 1938; Sainylina 1960
References
Zyrianka River, Kolyma basin, Albian
Localities, ages
Table 1. The complete list of the early angiosperm from the Soviet Union
P
E:
z
E*
?
c
zizyphoides sp.
Family Rhamnaceae
C . aff. ovale Vachrameev
6.serrdatussp. n.
Celastrophyllum sp.
Celastrophyllum sp.
C . kolymensis Samyline
C . cf. kolymeiwis Samylina
C . ovale Vachrameev
Family Celastraceae?
Celastrophyllum aff. hunteri Ward
C . kazachstanense Vachrameev
O n o a m nicanica Krassilov
Family Icacinaceae
Family Sapindaceae?
Supindopsis cf. angustn (Heer)Seward et Conway
Family Kyssaceae
Nyssidium orientale Samylina
Xyssidium sp.
Family Araliaccae
Araliaecarpum kolymensis Samylina
Aralia lucifera Kryshtofovich
Arulia sp.
Dalbergites sp.
Dulbergttes sp. 1
Dalbergites sp. 2
Dulbergitessp. 3
Family Leguminosites?
? Dulbergitessp.
Family Rosaceae
Crutaegitesborealis Samylina
C . cf. borealis Samylina
C.borealisf.sinuosa Samylina
Family Leyminosae
Leguminosites karatscheensisVachrameeb
Names of plants
Samylina, 1960
Kryshtofovich, 1929; Krassilov, 1967
Vakhrameev, 1952
Zyrianka River, Kolyma basin, Alhian
Suchan, South Primorye, Aptian, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Zyrianka River, Kolyma basin, Albian
Omsukchan, Kolyma basin, Late Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Zyrianka River, Kolyma basin, Albian
Lepiske River, Lena basin, Late Albian
Chushkakul Mountains, Western Kazachstan,
Mid& Albian; Lepiske River, Lena basin,
Late Slbian
Zyrianka River, Kolyma basin, Albian
Onisukchan, Kolyma basin, Late Albian
Zyrianka RiL-er, Kolyma basin, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Samylina, 1960
Samylina, 1960
Here
Samylina, 1960
Vakhrameev, 1952
Samylina, 1960
Kiritchkovah Slastenov, 1966
Vakhrameev, 1952; Kiritchkova Pi Slastenov, 1966
Here
Vakhrameev, 1932
Krassilov. 1967
Samylina, 1961
Samylina, 1961
Suchan, South Primorye, Barremian
Suchan, South Primorye, Barremian
Sujfun, South Primorye, Aptian
Krassilov, 1967
Vakhrameev, 1952
Suchan, South Primorye, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Vakhrameev, 1952
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Zyrianka River, Kolyma basin, Albian
Lepiske River, Lena basin, Late Albian
Lepiskc River, Lena basin, Late Albian
Lepiske Kirer, Lena basin, Late Albian
Samylina, 1960
Only the name in: Kiritchkova & Slastenov, 1966
Only the name in: Kiritchkova & Slastenov, 1966
Only the name in: Kiritchkova 8r Slastenov, 1966
Samylina, 1960
Kiritchkova 6 Slastenov, 1966
Samylina, 1960
References
Zyrianka River, Lena basin, Albian
Lepiske River, Lena basin, Late Albian
Zyrianka Rix-er,Kolyma basin, -4lbian
Localities, ages
Table 1. (contd.) The complete list ofthe early an,giosperms from the Soviet Union
D. obliquum Samylina
D.zyyrjankense Samylina
DicotylophyUum sp.
Dicotylophyllumsp.A
Dicotylophyllum sp. B
Dicotylophyllumsp. C
DicotylophyUumsp.D
Dicotylophyllum sp. E
DicotylophyUumsp.A
DicotylophyUumsp.B
Dicotylophyllum sp. 1
DicotylophyUum sp. 2
DicotylophyElumsp. 3
Kenella harrkiana gen. et sp. n.
Kenella sp.
Rogersia? denticulata Samylina
Dieotylophyllumbilobatum Vachrameev
C. prodromus Kryshtofovich (nomennudum)
cissites sp.
cissitessp. 1
cissitessp. 2
cissites sp. 3
Monocotyledones
Family Cyperaceae
Carecopsiscomrpacta S a m y h a
c.l a Samylina
Family Pandanaceae?
PandanophyUum ahnertii Kryshtofovich
Angiosperms of uncertain affinities
Carpolithes karatscheelzsisVachrameev
Family Vitaceae
Cissites cf. paruifolius Berry
Names of plants
Kryshtofovich, 1945,1950
Krassilov, 1967
Kiritchkova & Slastenov, 1966
Only thename in: Kiritchkova & Slastenov, 1066
Only the name in: Kiritchkova & Slastenov, 1966
Samylina, 1960
Samylina, 1960
Kryshtofovich, 1929
Vakhmmeev, 1952
~
Albian
Zyrianka River, K o l y m basin,
Zyrianka River, Kolyma basin, Albian
Sujfun, South Primorye, Aptian
C h u s h k W Mountains, Western Kazachstan,
Middle Albian
Omsukchan, Kolyma basin, Albian
Omsukchan,Kolyma basin, Albian
Zyrianka River, Kolyma basin, Albian
Chushkakul Mountains, Western Kazachstan,
Middle Albian
Zyrianka River, Kolyma basin, Albian
Zyrianka River, Kolyma basin, Albian
Sujfun, South Primorye, Albian
Zyrianka River, Kolyma basin, Albian
Zyrianka River, Kolyma basin, Albian
Zyrianka River, Kolyma basin, Albian
ZyriankaRiver, Kolyma basin, Albian
Zyrianka River, Kolymabasin, Albian
Omsukchan, Kolyma basin, Late Albian
Omsukchan,Kolymabasin, Late Albian
Lepiske River, Lena basin, Late Albian
Lepiske River, Lena basin, Late Albian
Lepiske River, Lena basin, Late Albian
Samylina, 1960
Samylina, 1960
Krassilov, 1967
Samylina, 1960
Samylina, 1960
Samylina, 1960
Samylina, 1960
Samylina, 1960
Only the name in: Filatov & Samylina, 1966
Only the name, here
Kiritchkovak Slastenov, 1966
Only the name in: Kiritchkova & Slastenov, 1966
Only the name in: Kiritchkova & Slastenov, 1966
Here
Only the name, here
Samylina, 1960
Vakhrameev, 1952
Vakhrameev, 1952
Middle Albian
Suchan, South Primorye, Albian
Suchan, South Primorye, Albian
Lepiske River, Lena basin, Late Albian
Lepiske River, Lena besin, Late Albian
Lepiske River, Lena basin, Late Albian
References
chushkakul Mountains, Western Kazechstan,
Localities, ages
Table 1. (contd.) The complete list of the early angiosperm from the Soviet Union
Early Cretaceous angiosperms of U.S.S.R.
217
similar angiosperm species in approximately contemporaneous floras of similar climatic
conditions may be attributed, I think, to the fact that although Albian angiosperms were a
strongly differentiated plant group, their systematic composition is still being revealed and
each new locality adds information in this respect.
If the assumption of the Early Cretaceous angiosperms being mountainous plants is
correct (Arnold, 1947; Takhtajan, 1948,1954; Vakhramecv, 1947,1952) and if in addition
their habitats were a t fairly high altitude, they could not have been thermophilic plants,
even within the Indo-European region. Mountainous conditions of early angiospermous
habitats could be the reason for comparatively rare findings of their remains in pre-Upper
Cretaceous deposits and their great systematic diversity as the abundance of various t.ypes
of habitats in mountains must have inevitably promoted intensive species formation,
I n the literature the small-leaved nature of most Early Cretaceous angiosperms is
repeatedly emphasized (Kryshtofovich, 1946,1950; Vakhrameev, 1952 ;Takhtajan, 1954 ;
Samylina, 1959). So far there is no accepted explanation for this fact. Possibly, it is
likewiseassociated with the mountainous habitats which had the inhibiting effect on plants
and prevented the development of large-leaved forms. The leaves of Early Cretaceous
angiosperms were morphologically fairly diverse both in the form of the leaf blade and the
type of its margin and in venation. One may say that towards the end of the Early Cretaceous the basic morphological types of dicotyledonous leaves already existed but on
the microphyllous background. Judging by the frequency of occurrence, entire elongate
leaves were most widespread among dicotyledons (Celastrophyllum, Cinnummoides,
Laurophyllum, etc.). I n the Siberian paleofloristic region they comprise more than half of
the dicotyledonous species and about one-third of the dicotyledons of the Indo-European
region. Fairly often, especially in localities of the Indo-European region, impressions of
dissected leaves, trilobate or pinnate, occur (Sassafras,Aralia, Crataegites,Artocarpidium).
As for the character of the leaf blade margin apart from the smooth-edged leaves, leaves
with a denticulate margin are as frequent among the Early Cretaceous angiosperms.
Moreover one may point out in dicotyledons of the Indo-European region a tendency to
predominance of smooth-edged leaf species (12 species out of 22, i.e. 54 yo)and respectively
in dicotyledons of the Siberian region a tendency to predominance of denticulate leaf
species (24 species out of 42, i.e. 57 yo).It is of interest that in the living floras of temperate
climate plants with denticulate leaves also predominate whereas in tropical floras the
smooth edged forms are predominant.
Most of the remains of early angiosperms appear to have belonged to small trees or
shrubs, But by the end of the Cretaceous the angiosperms had already begun to conquer
aquatic habitats and to cover the soil as the findings of the aquat,icplants, Nelumbites and
Hydrocotylophyllum and grassy ones, Carecopsistestify.
REFERENCES
ARNOLD, C. A,, 1947. An introductiontopaleobotany.New York & London.
AXELROD,
D., 1959. Poleward migration ofearlyangioperm floras.Science, 130,3369: 203-207.
BELL,W. A., 1956. Lower Cretaceousflorasof Western Canada. Ottawa.
FILATOV,
S. I. & SAMYLINA,
V. A., 1966. Tho stratigraphy and flora of Lower Cretaceous deposits of the
Balyghychan-Sugoydownwarping. Dokl. Akad. NaukSSSR, 1661 (1): 18f5-189. (Russian text.)
KIRITCHEOVA,
A. I. & BUDANTSEV,
L. U., 1967. A new find of the Lower Cretaceous flora, including
angiosperms, in Yakutia. Bot.J., 52 ( 7 ) : 937-943. (Russian text with English summary.)
KIRITCHKOVA,
A. I. & SLASTENOV,
J. L., 1966. Tho stratigraphy and flora of Lower Crctaceous deposits
of Lepiske River (Western Priverhojanio). Trans. A12-Union geol. prospect. Inst., 249: 147-169.
(Russian text.)
KnAssIr.ov, V. A , , 1907. The l h l y Cretaceous flora of the Southern Primorye and its st'gni'cance for
strciti~/rtrphy.
(Russiantext.)Moscow.
KrtYsHTomvIcH, A. N., 1929. Discovery of the oldest dicotyledons of Asia in the equivalents of the
Potomac Group in Suchan, Ussuriland, Siberia. Bull. du Comiti Geol., 48 (9): 1357-1390. (Russian toxt
with English Summary.)
218
V. A. SAMYLINA
KRYSHTOFOVICH,
A. N., 1938. Upper Cretaceousplants from the Kolyma River. Contr. to the knowledge of
the Kolyma-Indigirkaland, ser. 2,15: 1-16. (Russian text with English summary.)
KRYSHTOFOVICH,
A. N., 1945. Paleobotany. In Achievements of Biological Sciences during 25 year8
(1917-1942): 199-217. (Russiantext.)Moscow.
KRYSHTOFOVICH,
A. N., 1950. Evolution based onpaleobotanical data. I n Botanical problems, 1 : 5-37.
(Russian text.) Moscow & Leningrad.
KRYSHTOFOVICH,
A. N., 1958. The Cretaceous flora of the Anadyrland. Inat. Bot. Acnd. Sci. TISSR,
Acta, ser. V I I I ,Paleobotanica, 3 : 7-71. (Russian text with English summary.)
NEWBERRY,
J . S., 1655. Flora of the Amboy Clays. Momgr. U.S. geoLSurv., 26.
S A M Y L I N AA.,
, ~ 1959.
.
New occurrences of angiosperms from the Lower Cretaceous of the Kolyma basin.
Bot.J.,44 ( 4 ) :483-491. (Russiantext with English summary.)
SAMYLINA,
V. A., 1960. Angiosperms from the Lower Cretaceous of the Kolyma basin. Bot. J., 45 (3):
335-352. (Russian text with English summary.)
SAMYLINA,
V. A., 1961. hrew data on the Lower Cretaceous flora of the southern part of the Maritime
Territory of the R.S.F.S.R. Bot.J.,46 (5): 634-645. (Russian text with English summary.)
SAPORTA,
G., 1694. Florafosaile duPortuga1. Lisbon.
TAKHTAJAN,
A, L., 1946. The morphological evolution of Angiosperm. (Russian text.) Moscow.
TAKHTAJAN,
A. L., 1954. Originsofangios~mrtousplants.(Russiantest.) Moscow.
TEIXEIRA,
C., 1948. Flora MesozoicaPortuguesa, I. Lisbon.
VAKHRAMEEV,
V. A., 1947. The role of geographical conditions in development and distribution of
angiospermous floras in Cretaceous time. BuU. Mosc.Soc. Nut. geol. Div., 6 : 3-17. (Russiantext.)
VAKHRAMEEV,
V. A,, 1952. The stratigraphy and the fossil flora of the Cretaceous deposits of wcstcrn
Kazakhstan. Regional stratigraphy ojthe USSR, 1, (Russian text.) Moscow.
VAKHRAMEEV,
V. A., 1964. Jurassic and early Cretaceous floras of Eurasia and the paleofloristic
provinces of this period. Trans. geol. Inst. Acad.Sci. USSI1,102. (Russian text.)