Upper Cretaceous pollen flora from the Vilui Basin, Siberia: Circumpolar and endemic Aquilapollenites , Manicorpus , and Azonia species

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Upper Cretaceous pollen flora from the Vilui Basin,Siberia: Circumpolar and endemic Aquilapollenites,Manicorpus, and Azonia speciesChrista‐Charlotte Hofmann a & Reinhard Zetter a

a Department of Palaeontology , Faculty of Earth Sciences , Geography and Astronomy ,University of Vienna , Vienna, AustriaPublished online: 17 Dec 2007.

To cite this article: Christa‐Charlotte Hofmann & Reinhard Zetter (2007) Upper Cretaceous pollen flora from the ViluiBasin, Siberia: Circumpolar and endemic Aquilapollenites, Manicorpus, and Azonia species, Grana, 46:4, 227-249, DOI:10.1080/00173130701763142

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Upper Cretaceous pollen flora from the Vilui Basin, Siberia:Circumpolar and endemic Aquilapollenites, Manicorpus, and Azonia

species

CHRISTA-CHARLOTTE HOFMANN & REINHARD ZETTER

Department of Palaeontology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, Vienna, Austria

AbstractA detailed LM and SEM examination of the Upper Cretaceous Timerdyakh Formation microflora from the Vilui Basinyielded 13 Aquilapollenites, two Manicorpus, and three Azonia species. Comparisons with existing taxa from the RussianFederation and North America are impeded by the fact that many previously described forms were investigated using LMonly, while SEM was very rarely used. Consequently, some of the pollen types reported in this study could not be ascribedto any previously documented taxa and have been described as nine new species. Further one previously described varietyhas been renamed. Except for two Azonia species and four Aquilapollenites species that also occur in Alaska, Canada and theRocky Mountains in USA, the remaining taxa described here are restricted to the Late Cretaceous Khatanga-Lena-Subprovince. The several species unique to the Vilui Basin might indicate separate evolutionary paths for the generaAquilapollenites and Manicorpus in North America and both northern and southern Asia during the Upper Maastrichtian.

Keywords: Upper Cretaceous, Aquilapollenites, Manicorpus, Azonia, Vilui Basin, Siberia

The three form genera Aquilapollenites (Rouse)

emend. Srivastava, Manicorpus Mchedlishvili emend.

Srivastava, and Azonia Samoilovich are widely

distributed in Upper Cretaceous sedimentary rocks

of Canada, the USA, northern Europe (Mull,

Scotland), Siberia, Sakhalin, and Japan and conse-

quently they are valuable stratigraphical markers in

the Northern Hemisphere (Bolchovitina, 1959;

Bratseva, 1965, 1969; Chlonova, 1961; Dawson et

al., 1994; Farabee, 1990; Farabee & Canright, 1986;

Funkhouser, 1961; Jarzen, 1977; Jarzen & Norris,

1975; Krutzsch, 1970; Martin, 1968; Mchedlishvili,

1961; Nichols & Sweet, 1993; Norris et al., 1975;

Samoilovich, 1965, 1967; Stanley, 1961; Srivastava,

1966, 1972, 1981,1994a; Srivastava & Rouse, 1970;

Takahashi, 1994; Tschudy, 1969; Tschudy &

Leopold, 1971; Wiggins, 1976). A number of

botanical affiliations have been proposed for the form

genus Aquilapollenites, such as with the Santalaceae/

Santalales (Funkhouser, 1961; Jarzen, 1977; Norton

& Hall, 1969), Loranthaceae (Erdtman, 1971; Jarzen

& Norris, 1975) or that they represent several

distinctly related families (Farabee, 1990; Srivastava

& Rouse, 1970). Chlonova (1967) equated the

Oculata types, such as the Azonia species, with the

Balsaminaceae family; whilst Samoilovich &

Mchedlishvili (1961) and Wiggins (1976) suggested

that the group is extinct and only occurred in the

Cretaceous.

The three form genera occur relatively frequently,

and sometimes abundantly, in Campanian to

Maastrichtian sedimentary rocks of the Vilui Basin,

mainly in the Timerdyakh Formation, which gener-

ally comprises fluvial and associated deposits (Spicer

et al., submitted). The shapes of Aquilapollenites are

very unusual in comparison to modern pollen types:

Polar and equatorial areas can be extremely exagger-

ated, producing polar and equatorial projections. If

isopolar, these triprojectate pollen have been assigned

to several different genera including Triprojectus

Mchedlishvili or Parviprojectus Mchedlishvili or

Integricorpus Mchedlishvili (all in Samoilovich &

Correspondence: Christa-Charlotte Hofmann, Department of Palaeontology, Geozentrum, University of Vienna, Althanstrasse 14, A–1090 Vienna, Austria.

E-mail: [email protected]

(Received 27 June 2007; accepted 25 October 2007)

Grana, 2007; 46: 227–249

ISSN 0017-3134 print/ISSN 1651-2049 online # 2007 Taylor & Francis

DOI: 10.1080/00173130701763142

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Mchedlishvili, 1961), whilst very heteropolar forms

were assigned to Manicorpus. All these form genera,

except Manicorpus, were later included in the form

genus Aquilapollenites by Stanley (1979) and Tschudy

& Leopold (1971). Manicorpus was kept as a separate

genus for species with an x/y ratio less than 0.5 (length

from the reduced pole to equator divided by the

length of the elongated pole to the equator; see

Srivastava, 1968 and Srivastava & Rouse, 1970).

Many former studies were undertaken using only

light microscopy, but there were exceptions, where

SEM was partly used (Farabee, 1990; Farabee &

Canright, 1986; Srivastava, 1972, 1975; Takahashi,

1994; Tschudy, 1969). This has in part impeded the

comparison of taxa from Canada/USA with those

from Siberia, as the light microscopy images often

are not sufficiently detailed to differentiate between

the proposed species.

The unusual shape of Oculata (Azonia) pollen types

has also been a subject of discussion (Samoilovich,

1961; Stanley, 1961; Chlonova, 1967; Wiggins, 1976)

and it is still unclear where the poles and equators of

these grains lie because they have never been found in a

tetrad configuration. Chlonova (1967) proposed an

affinity with the Balsaminaceae, in particular to the

genera Impatiens and Jollydora, and compared the

position of the Oculata apertures (pores or colpi)

with the equatorial apertures of the genera mentioned

above.

In this study, we present detailed descriptions of all

the Aquilapollenites, Manicorpus, and Azonia forms

found in the Upper Cretaceous Timerdyakh

Formation of the Vilui Basin, Siberia, using both

light microscopy and detailed SEM images.

Comparisons with examples documented from else-

where demonstrate that a few of these Siberian taxa

are unique, but some forms belong to a circumpolar

flora, which includes Alaska and Northern Canada.

The concept of a Khatanga-Lena-Subprovince

(Samoilovich, 1967; Chlonova, 1981) thus may be

useful, describing a northernmost palynomorph

assemblage of the Russian Federation, which prob-

ably extended into Alaska and Canada. However, the

subprovince was originally defined by the absence of

tropical and subtropical families, a circumscription

that cannot be confirmed for the rest of the microflora

(Spicer et al., submitted). Taxa described from Mull,

Scotland (Martin, 1968; Srivastava, 1975) were

linked to Siberian taxa in the southerly located

Yenisei-Amur-Subprovince (Samoilovich, 1967).

These included both Aquilapollenites pachypolus

Martin (previously Parviprojectus striatus Mche-

dlishvili in Samoilovich & Mchedlishvili, 1961: one

LM image and two drawings) and Aquilapollenites

subtilis Mchedlishvili (Samoilovich & Mchedlishvili,

1961: two LM images and two drawings).

Material and methods

Seventeen samples of Campanian/Maastrichtian age

from the Timerdyakh Formation (Albian to

Maastrichtian in age) were analysed from logged

sedimentological profiles of riverbank outcrops

along the Tyung River in the centre of the Vilui

basin, adjacent to Locality 4215 of Vachrameev and

Pushcharovski (1954). The fieldwork was done by

Anders Ahlberg, Alexei Herman, Maria Moiseeva

and Robert Spicer. Only samples T4 (youngest),

T5b, T7, T9, T10, T11, T13, T14 and T15 (oldest)

yielded Aquilapollenites, Manicorpus and Azonia

pollen (summarized in Table I), with a decrease in

diversity of all three genera towards the youngest

strata. The investigated material comprises very fine-

grained to fine-grained floodplain, palaeosol, peat,

lake and mud clast/drape sediments. The succession

is characterized by an increased reworking of over-

bank fines by river channel migration and channel

cannibalism, thus witnessing a very dynamic hydro-

logical and sedimentary environment (Spicer et al.

submitted).

Sample preparation was done following standard

procedures: The samples were crushed with a mortar

and pestle, and the resulting rock powder treated

with standard wet chemical processes using HCl and

HF. The organic extract, which was not sieved in

order to retain palynomorphs smaller than 10 mm,

was acetolyzed and mixed with glycerine and stored

in small glass bottles. For examination under LM a

drop of well-mixed extract was evenly distributed on

a glass slide. For LM photography (Nikon Coolpix)

the investigated pollen grains were transferred by a

micromanipulator (a hair on a preparation needle)

into a clean drop of glycerine on a new slide. For

further examination under the SEM (JEOL 6400),

the pollen grains were again moved with a micro-

manipulator to a SEM stub and carefully washed

with 100% alcohol to remove the glycerine.

Overview LM and SEM images were used to

measure the dimensions of the pollen grains. The

defining measurements used for the genus

Aquilapollenites followed Srivastava (1968, Figure 3)

and Srivastava and Rouse (1970, Figure 1). The

whole packet with described specimens on the SEM

stubs, the electronic photomicrographs (CD), and

prints of the plates are stored in the type collection

of the Department of Palaeontology at the University

of Vienna (Austria) under the inventory number:

3830.

Results

Detailed SEM imaging of pollen grains (18 taxa)

yielded 13 Aquilapollenites species, with one taxon

displaying three variants, two Manicorpus species,

228 C.-C. Hofmann and R. Zetter

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and three Azonia species. The results give no

additional information on probable botanical affilia-

tions of these genera. We agree with Erdtman

(1971): ‘……These morphological characteristics

constitute a morphological complex which, as far

as I know, does not appear in other plants…..’

This paper describes eight new species of

Aquilapollenites (A. hermanii, A. samoilovichiae, A.

striatolongus A. fergusonii, A. spiceri, A. heteropolaris,

A. srivastavae, A. ovatus), three variants of

Aquilapollenites turbidus Tschudy & Leopold, most

of which can only be distinguished with certainty by

SEM investigation and one new Manicorpus species

(M. truncatus). We also suggest a new name (nom.

nov.): Azonia lindensis (formerly Azonia calvata var.

lindensis Samoilovich). With the exception of

Aquilapollenites chlonovae, A. hermanii, and A. mche-

dlishvilii (all three reticulate forms), A. turbidus, and

Azonia recta and A. calvata, which occur also on the

Alaskan peninsula and northern America, the

remaining species described here are thought to be

endemic to the Khatanga-Lena-Subprovince of the

Russian Federation.

Description of taxa

A list of the measurements of all the described taxa is

given in Table II. Where not otherwise stated all

images referred to in the synonymy lists are

LM images. The taxa are ordered according to

the sculpture commencing with the reticulate,

striatoreticulate and striate Aquilapollenites species,

then the spinulose species, followed by two

Manicorpus species and at last three Azonia species.

Genus Aquilapollenites Rouse emend. Srivastava 1968

Aquilapollenites chlonovae (Chlonova) Srivastava

(Figure 1 A–F & I)

1959 Aquilapollenites trialatus Rouse, in

Bolchovitina: Plate 8 Figure 113c

1961 Aquilapollenites reticulatus Chlonova, in

Chlonova: Plate 14 Figures 107 & 107a (drawing)

1961 Integricorpus bellum Mchedlishvili, in

Samoilovich & Mchedlishvili: Plate 70 Figures 1 a–

d & 2 a, b

non 1961 Parviprojectus reticulatus Mchedlishvili,

in Samoilovich & Mchedlishvili: Plate 73 Figures 2,

3

1965 Integricorpus clarireticulatus Samoilovich, in

Samoilovich: Plate 1 Figure 2

1967 Integricorpus sp.1, in Samoilovich: Plate 3

Figure 16

1967 Integricorpus sp.2, in Samoilovich: Plate 3

Figure 19

1967 Integricorpus cf. bellum Mchedlishvili, in


1967 Integricorpus clarireticulatus Samoilovich, in


cf. 1969 Aquilapollenites clarireticulatus Samoilovich,

in Tschudy: Plate 2 Figures 2, 3; Plate 3 Figure 6

Table I. Occurrences of Aquilapollenites, Manicorpus, and Azonia taxa in the samples of the Timerdyakh Formation, with short descriptions

of the sedimentary rock facies type.

Sample Lithology Aqu

ilapol

lenites

chlo

nov

ae

A.

her

manii

A.

sam

oilo

vic

hia

e

A.

mch

edlish

vilii

A.

stri

ato

longu

s

A.

turb

idus

A.

rom

bicu

s

A.

ferg

uso

nii

A.

spic

eri

A.

pro

ceru

s

A.

het

erop

olari

s

A.

sriv

ast

avae

A.

ovatu

s

Manic

orpus

tenue

M.

trunca

tus

Azon

iaca

lvata

A.

rect

a

A.

linden

sis

T4 upper mudball x x

T5b mudfilled

channel

x x x

T7 Base of

lacustrine

unit

x x x x x x x x

T9 peatball x x x x x x x x x

T10 lacustrine

siltstone

x x

T11 laminated

coarse

siltstone

x x x x x x x

T13 lacustrine

shale

x x x x x

T14 mudball x x x x x x x x x x

T15 floodplain

siltstone

x x x x x x x x x x x x

U. Cretaceous pollen flora from Vilui Basin, Siberia 229

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Description Tricolpate pollen grains with long colpi,

subisopolar with slightly flattened poles, thus giving

a more-or-less angular outline. Pollen grains are very

variable in their overall size and width of brochi of

the reticulum (polar axes ranging from 25–56 mm,

equatorial axes ranging from 22–63 mm). The polar

projections display a coarse reticulum), in which

brochi decrease in size and shape towards the

equatorial area (Figure 1 A–F). The brochi in the

polar areas ranges from 1 to 5 mm min diameter and

display rudimentary freestanding columellae

(Figure 1 I). The equatorial area is characterized

by more-or-less parallel long striae running from the

colpi to the equator (ring-like thinning; Figure 1 D).

The striae tend to be fused near the colpus margins

(Figure 1 F). The equatorial ring-like thinning

separates the pollen grains into two halves

(Figure 1 B & E). The equatorial projections, if

not broken off, are thin and elongate (lengths range

from 5 to 20mm), faintly striate in the proximal

region and smooth to slightly perforate in the distal

region. The most important LM features are the

equatorial ring-like thinning and the very coarse

reticulations.

Remarks. The pollen grains of this taxon are very

variable in size and when using light-microscopy

alone it is difficult to recognize that all sizes occur in

one species, only. Both small and large pollen grains

of this taxon may co-occur in the same sample; there

is no correlation between pollen size and

stratigraphic level. The common preservation as

half pollen grains led to the assumption that both

endexine and ectexine are thinned in the equator.

The original name Aquilapollenites reticulatus

Chlonova (1961) was changed to Aquilapollenites

chlonovae nom. nov. by Srivastava (1968) using a

description translated from Russian, because A.

reticulatus Chlonova 1961 is a later homonym of A.

reticulatus Stanley (1961). This species was then

discovered in Alaska by Tschudy (1969), who

renamed it Aquilapollenites clarireticulatus (former

name Integricorpus clarireticulatus Samoilovich,

1965). Tschudy’s (1969) circumscription of this

species was rather wide containing a heterogeneous

assemblage of forms also including pollen grains that

in our study are recognised as a separate species

described as Aquilapollenites hermanii (see below).

The figures of Aquilapollenites clarireticulatus on

plates two and three of Tschudy (1969) are a

mixture of LM photographs of Aquilapollenites

chlonovae and SEM and LM photographs of grains

that we here include in the new species A. hermanii.

This demonstrates that using exactly the same pollen

grain for the LM and SEM photography has

undeniable merits.

Illustrated specimens. 3830VB07-9/2/1 and 3830VB07-

14/10/1

Occurrence. Not abundant but frequent in samples

T7, T9, T11, T14 and T15.

Aquilapolleniteshermaniisp.nov.(Figure 1G,H&J,K)

1969 Aquilapollenites clarireticulatus Samoilovich, in

Tschudy:Plate 2Figures 4&8,Plate 3Figures 1,5&7

1997 Aquilapollenites catenireticulatus Srivastava, in

Takahashi: Figure 61 (SEM image)

Derivatio nominis. The species name honours the

Russian palaeobotanist Alexei Herman, whose work

covers the Cretaceous macroflora of the northern

hemisphere.

Specific diagnosis. As for the genus, but with the

following additions: the new species differs from the

holotype of A. catenireticulatus (LM images,

Srivastava, 1968, Plate 1, Figures 7–9) and from

the SEM images of A. catenireticulatus in Farabee &

Canright, (1986, Plate 12, Figures 6–10), as well as

from LM images of Parviprojectus reticulatus

Mchedlishvili in Samoilovich & Mchedlishvili

(1961), Parviprojectus dolium (Samoilovich, 1965),

and Integricorpus reticulatus (Mchedlishvili) Stanley.

These all lack the equatorial ring-like thinning (LM)

and the parallel striations perpendicular the

equatorial area (SEM), features that characterise

A. hermanii. In the new species, the striae are

arranged more densely, the equatorial ring-like

thinning is more rugulate and the distal equatorial

projections are broader than in Aquilapollenites

samoilovichiae described below.

Holotype. 3830VB07-9/2/2, Figure 1 G, H & J, K

Paratypes. 3830VB07-7/20/1, 3830VB07-14/26/1

r

Figure 1. A–F & I. Aquilapollenites chlonovae. G, H & J, K. A. hermanii. A, D & G. LM overview of pollen grains (ca6900). B, C & E, F,

H–K. SEM micrographs. B, E & H. Equatorial pollen view. C, F, I–K. Details of exine sculpture: (C) reticulum with free-standing

columellae, (F) colpus margin with parallel fused striae, (I) high reticulum at polar area, (J) equatorial area and distal end of equatorial

projection, (K) striate micro-sculpture. Scale bars 21 mm (C, F, I–K); 10 mm (B, E, H).


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Table II. Short description of microstructures and measurements of the different Aquilapollenites, Manicorpus and Azonia taxa found in the Timerdyakh Formation. The method of measuring the

polar axes, polar projections, equatorial axes, equatorial projections, and equatorial areas follows Srivastava (1968, Figure 3) and Srivastava & Rouse (1970, Figure 1).

Taxa No. Polarity

Polar axis

length

(mm)

Polar

projections

length

(mm)

Polar

sculpture

Polar projection

sculpture

Equatorial

axis

length

(mm)

Equatorial

projection

length (mm)

Equatorial

projection

shape

Equatorial

area length

(mm)

Equatorial

sculpture

Aquilapollenites

chlonovae

18 subisopolar 25–57 9–29, 11–22 reticulate reticulate 22–63 5, 11 thin elongate 6–14 long striae fineretic

A. hermanii w30 subisopolar 24–34 6–11, 7–15 striate parallel striate 25–33 6, 9 round wedge 7–11 pa.striae.,rugulate

A. samoilovichiae 3 subisopolar 27–28 9–12, 11–13 striate striato-reticulate 29–31 8, 9 thin elongate 7–8 pa.striae.,rugulate

A. mchedlishvilii 2 subisopolar 26–30 7–12, 9–13 reticulate-striate striato-reticulate 24–34 4, 8 thin elongate 6–10 striato-reticulate

A. striatolongus 1 subisopolar 20 4 & 4 striate perforate striate, perforate 21 8 round wedge 12 striate

A. turbidus var.1 4 subisopolar 21–30 3–5, 5–6 foveate perforate smooth perforate 29–33 29–30 round wedge 12–18 perforate, angular spines

A. turbidus var. 2 2 subisopolar 28 6 & 7 bald perforate smooth perforate 36 12 round wedge 18 perforate, few spines

A. turbidus var. 3 3 subisopolar 24 4 & 5 bald bit perfor. smooth 28 10 round wedge 15 numerous spines

A. rombicus 2 subisopolar 28 3 & 4 perforate scaly perforate scaly 33.5 11 round wedge 21 spiny-scale perf.

A. fergusonii 2 subisopolar 33 7.5 & 11 bald perforate microretic scaly 38 17 elong. round. 14 scaly perforate

A. spiceri 8 het. & subis. 35–45 11–13, 12–17 bald slight perf. microretic scaly 51–56 21–23 elong. round. 11–13 flat-scaly microballs

A. procerus 13 subisopolar 29–50 16–17, 18–19 bald perforate reticulate spiny 38 13–17 elong. round. 12–15 flat-scaly

A. heteropolaris 4 heteropolar 28–35 7–10, 10–14 bald perforate perf. spiny scales 36–40 13–15 elong. round. 11–15 perforate scaly (few)

A. srivastavae 2 subisopolar 40 11 & 13 bald perforate verrucate perforate 36 14 round wedge 17 verrucate

A. ovatus 3 subisopolar 32 9 & 11 bald perforate perf. microspheres 18 3 small wedge 12 perf. microsphres

Manicorpus tenue 5 heteropolar 23 2 & 9 microreticulate microretic. spiny/blunt 32 11 round wedge 15 flat spiny, perforate

M. truncatus 4 heteropolar 27 8 & 2 microreticulate microretic. spiny/blunt 32 11 big angular 15 foveate spiny

Azonia lindensis 3 isopolar 21 reticulate 30 perforate rugulate

A. calvata 5 isopolar 26–35 reticulate 35–42 perforate rugulate

A. recta 6 het. & iso 14–23 reticulate 24–40 verrucate

232

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Figure 2. A–D. Aquilapollenites samoilovichiae. E–H. A. mchedlishvilii. I–L. A. striatolongus. A, E & I. LM overview of pollen grains (ca6900).

B–D, F–H & J–L. SEM micrographs. B, F & J. Equatorial pollen view. C, D, G, H, K & L. Details of exine sculpture: (C) conical equatorial

projection, (D) striatoreticulate polar area, (G) striate colpus margin and tongue-like equatorial projection, (H) striatoreticulate

mesocolpium, (K) parallel striate mesocolpium, (L) striate perforate polar area. Scale bars 21 mm (C, D, G, H, K, L); 10 mm (B, F, J).


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Figure 3. A–P. Aquilapollenites turbidus: A–H. A. turbidus var. turbidus; I–L. A. turbidus var. paucispinus; M–P. A. turbidus var. compactus. A,

E, I, M, N. LM overview of pollen grains (ca6900). B–D, F–H, J–L & O, P. SEM micrographs; B, J, O. Equatorial pollen view; (F)

polar view. C, D, G, H, K, L & P. Details of exine sculpture: (C) angular spinae, (D, G) foveolate polar area, (H) colpus margin with

‘dragoncomb’, (K) mesocolpium with few spines, (L) bald polar area and ‘dragoncomb’, (P) angular spinae of the ‘dragoncomb’ and

perforations. Scale bars 21 mm (C, D, G, H, K, L & P); 10 mm (B, F, J, O).


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Type locality. Timerdyakh Formation, outcrop

Tyung River sample T9, centre of the Vilui Basin,

Eastern Siberia.

Type stratum and age. Timerdyakh Formation

(Campanian/Maastrichtian), peat ball (brown clay

matrix with plant hash) preserved within fluvial

channel facies.

Description. Tricolpate, compact, subisopolar pollen

grains (polar axes: 24–34 mm, equatorial axes: 25–

33 mm) with three small rounded, wedge-shape

projections (length: 6–9 mm) and striate, micro-

reticulate to perforate tectum. The LM images

display a characteristic ring-like thinning in the

equator (Figure 1 G), where the nexine does not

reach into the equatorial projection. It appears that

the nexine is thinner than the sexine. Under the

SEM, the striae are long (up to 10 mm) and are

more-or-less parallel, densely arranged, running

perpendicular to the colpi and the equatorial area

(Figure 1 H & K). On the polar projection, the striae

produce a striato-micro-reticulum. The striae

generally end at the equatorial ring-like thinning or

get crinkled, to produce a more rugulate pattern

(Figure 1 J). In the distal region of the equatorial

projections the striae are fused to a perforate tectum

(Figure 1 K). The only occasionally visible colpus

membrane is microverrucate.

Remarks. This is one of the most abundant and

relatively frequent pollen species occurring in the

Timerdyakh Formation.

Occurrence. Abundant and frequently occurring in

samples T7, T9, T11, T13, T14, and T5.

Aquilapollenites samoilovichiae sp. nov. (Figure 2 A–

D)

cf. 1967 Parviprojectus sp. in Samoilovich: Plate 3

Figure 24

Derivatio nominis. This species name is in honour of

S. R Samoilovich, a famous palynologist from the

Russian Federation.


following additions: A. samoilovichiae differs from

Aquilapollenites pachypolus Martin (Scotland: Martin

1968, Plate 106 Figures 9–14) and Parviprojectus

striatus Mchedlishvili (Samoilovich & Mchedlishvili,

1961) in having a ring-like thinning at the equator

and very narrow cone-shape equatorial projections.

It differs from A. hermanii in its microreticulate

sculpture at the equator and the very narrow cone-

shaped equatorial projections.

Holotype. 3830VB07-15/18/1, Figure 2 A–D

Paratype. 3830VB07-9/2/3



Eastern Siberia.


(Campanian/Maastrichtian), floodplain siltstone

(brownish grey, non-stratified).

Description. Tricolpate, less compact, subisopolar

pollen grains (polar axes: 27–28 mm, equatorial

axes: 29–31 mm) with three narrow, cone-shaped

equatorial projections (7–8 mm) and a striato-

reticulate tectum (Figure 2 B). Under the SEM,

the striae are relatively long, running (arranged

parallel) more-or-less perpendicular towards the

colpi and towards the equatorial area (Figure 2 C,

D). At the equatorial ring-like thinning (visible

under LM as a lighter line; Figure 2 A), the striae

again produce a micro-reticulum. On the equatorial

projection, the striae run parallel to the equatorial

axis and look like a bundle of fused striae in the distal

pointed end (Figure 2 C). Under LM the nexine

looks more-or-less as thick as the sexine and reaches

to the middle of the equatorial projection.

Remarks. A. samoilovichiae is less abundant and less

frequent than A. hermanii, with which it is easily

confused under the LM.

Occurrence. Not abundant, occurring only in

samples T9 and T15.

Aquilapollenites mchedlishvilii Srivastava (Figure 2

E–H)

1961 Parviprojectus reticulatus Mchedlishvili,

in Samoilovich & Mchedlishvili: Plate 73

Figures 2a & 3

cf. 1965 Parviprojectus reticulatus Mchedlishvili, in

Bratseva: Plate 4 Figures 5 & 7

cf. 1967 Parviprojectus dolium Samoilovich, in

Samoilovich: Plate 3 Figures 1a & 2a

1971 Aquilapollenites reticulatus Mchedlishvili in

Tschudy & Leopold: Plate 1 Figures 12 a, b

1990 Integricorpus reticulatus Mchedlishvili, in

Farabee: Plate 2 Figures 14 & 17 (SEM image)

1993 Aquilapollenites reticulatus Mchedlishvili in

Nichols & Sweet: Plate 2 Figure 30, LM

Description. Tricolpate, subisopolar pollen grains

(polar axes: 26–30 mm, equatorial axes: 24–34 mm)

with an oval outline and three narrow, tongue-shaped


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equatorial projections (4–7 mm length) and a striato-

reticulate tectum (Figure 2 E, F). On the polar

projections, the distal end of the colpus is bordered

by a slightly protruding bulge (Figure 2 F). The striae

tend to be arranged perpendicular and denser around

the colpi, but the equatorial area is continuously

striato-reticulate. The equatorial projections are

striate, perforate and thin.

Remarks. Under LM and SEM, this form does not

display the equatorial ring-like thinning typical of A.

hermanii, A. samoilovichiae and A. chlonovae and

therefore can be distinguished from these forms. It

somewhat resembles the SEM image of A.

catenireticulatus Srivastava Figured in Farabee &

Canright (1986: Plate 12 Figures 9 and 10), but

differs from this specimen in its thin, tongue-shaped

striate perforate equatorial projections. The LM

image is comparable with the LM image of

Integricorpus reticulatus Mchedlishvili and the SEM

image of Integricorpus reticulatus (Mchedlishvili)

Stanley Figured in Farabee (1990, Figure 14 &

17). However, the species name ‘reticulatus’

has been already used by Stanley (1961) and

Chlonova (1961). Consequently, Srivastava (1968)

gave this species a new name, Aquilapollenites

mchedlishvilii, stating that Aquilapollenites reticulatus

Stanley is different from Parviprojectus reticulatus

Mchedlishvili. Later, Tschudy and Leopold (1971)

regarded these as a single species. This problem can

be only resolved by further investigations and

comparisons of SEM images.

Illustrated specimen. 3830VB07-10/26/1

Occurrence. Not abundant, occurring in samples

T11 and T14.

Aquilapollenites striatolongus sp. nov. (Figure 2 I–L)

Derivatio nominis. The species name striatolongus

stands for the characteristic long striae, which

more-or-less extend from pole to pole.


following additions: A. striatolongus is the only

species with parallel striae extending from pole to

pole and can be differentiated from Bratsevaea sp.

Takahashi Figured in Farabee (1990): Plate 2

Figure 18 by the parallel striae on the polar

projections, which in Bratsevaea are not parallel

arranged and produce a more irregular striato-

reticulate pattern.

Holotype. 3830VB07-15/26/1, Figure 2 I–L



Eastern Siberia.



(brownish grey, non-stratified).

Description. Tricolpate, subisopolar (despite the

same lengths, one polar projection is broader than

the other) pollen grains (polar axis: 20 mm,

equatorial axis: 22 mm) with narrow polar

projections (length: 4 mm each), and three broad,

rounded wedge-like equatorial projections (Figure 2

I, J). Under SEM the entire pollen grain is parallel-

striate from pole to pole (Figure 1 J); these striae can

be rarely, if at all, seen under LM (I). The polar

projections have few perforations between truncating

striae (Figure 2 L). In general, striae result from

narrow folding of the tectum, whilst the striae on the

equatorial projections are more flattened and fused,

particularly at the distal ends (Figure 2 K).

Remarks and occurrence. This species is rare in our

samples and occurs only in sample T15.

Aquilapollenites turbidus Tschudy & Leopold (Figure 3

A–P)

1961 Aquilapollenites quadrilobus Rouse, in

Samoilovich & Mchedlishvili: Plate 67 Figures 4 & 3


Bratseva: Plate 2 Figures 1–11



1969 Aquilapollenites rombicus Samoilovich, in

Bratseva: 27 Figures 5 & 7

1997 Aquilapollenites quadrilobus Rouse emend.

Srivastava & Rouse, in Takahashi: Figures 55–58

(SEM)

r

Figure 4. A–D. Aquilapollenites rombicus. E–H. A. fergusonii. I–N. A. spiceri. A, E & I. LM overview of pollen grains (ca6900). B–D, F–H,

J–N. SEM micrographs. B, F, J. Pollen in equatorial view. C, D, G, H, K–N. Details of exine sculpture: (C) detail of the scale-like spinae,

(D) colpus margin with ‘dragoncomb’, (G) densely spaced broad spinae, (H) tectate-perforate polar area, (K) round scale-like spinae on

the distal equatorial projection, microspheres and perforation, (L) irregular spinae of the ‘dragoncomb’ colpus margo; (M) equatorial

projection with scale-like spinae, (N) tectate cap of the polar area. Scale bars 21 mm (C, D, G, H, K–N); 10 mm (B, F, J).


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Figure 5. A–H. Aquilapollenites procerus. I–M. A. heteropolaris. A, E, I. LM overview of pollen grains (ca6900). B–D, F–H & J–M. SEM

micrographs. B, F, J. Pollen in equatorial view. C, D, G, H, K–M. Details of exine sculpture: (C) equatorial projection with numerous flattened

spinae and regularly composed ‘dragoncomb’, (D, G) triangular shaped, perforate tectate cap of the polar areas, (H) slim polar projection with

microreticulum and few, rudimentary spinae, (K) polar projection with tectate polar area and perforate spiny sculpture, (L, M) equatorial

projections with perforations, flattened spinae and ‘dragoncomb’ colpus margins. Scale bars 21 mm (C, D, G, H, K–M); 10 mm (B, F, J).


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Figure 6. A–D. Aquilapollenites srivastavae. E–G. A. ovatus. H–K. Manicorpus tenue. L–O. M. truncatus. A, E, H, L. LM overview of pollen

grains (ca6900). B–D, F, G, I–K & M–O. SEM micrographs. B, F, I & M. Pollen grains in equatorial view. C, D, G, J, K, N & O.

Details of exine sculpture: (C) verrucate sculpture of the central body and flattened spinae of the distal equatorial projection, (D) tectate,

perforate polar area and verrucate sculpture of the polar projection, (G) perforate tectum with rounded spinae, (J) flattened rounded spinae

of the equatorial area, (K) perforated polar area and ‘dragoncomb’ on the equatorial projection, (N) perforate equatorial area with angular

spinae; (O) perforate to foveolate polar area. Scale bars 21 mm (C, D, G, J, K, N & O); 10 mm (B, F, I, M).


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1971 Aquilapollenites turbidus Tschudy & Leopold

in Tschudy & Leopold: Plate 2 Figures 7a, b & 9

1977 Aquilapollenites quadrilobus Rouse, in Jarzen:

Figure 5.1

1994 Aquilapollenites turbidus Tschudy & Leopold,

in Dawson et al.: Plate 1 Figure 8

1997 Aquilapollenites quadrilobus Rouse emend.


(SEM)

Remarks. This variable taxon (three variants are

described below) has been continuously confused with

the heteropolar (rarely sub-isopolar) A. quadrilobus

Rouse emend Srivastava 1968 (Srivastava, 1968;

Srivastava & Rouse, 1970) by Russian and Japanese

workers, despite the fact that spines have been

described on the polar projection and poles and that

these are clearly visible under the light microscope.

Aquilapollenites turbidus Tschudy & Leopold var.

turbidus (Figure 3 A–H)

cf. 1965 Aquilapollenites quadrilobus Rouse, in

Bratseva: Plate 2 Figures 7, 8

cf. 1969 Aquilapollenites rombicus Samoilovich, in

Bratseva: 27 Figure 7

1997 Aquilapollenites quadrilobus Rouse emend


(SEM)

Description. Tricolpate (long colpi), compact,

subisopolar pollen grains (polar axes: 21–30 mm,

equatorial axes: 29–33 mm), with three big, rounded

wedge-like equatorial projections (Figure 3 A, B).

The short polar projections (3–6 mm) are angular-

foveolate, perforate (some angular fovae have

perforations) and spineless (bald polar area;

Figure 3 B, D & G), whilst the equatorial area and

projections are regularly perforated and covered with

widely spaced supratectal, crystal-like spines (length:

1–2.5 mm; Figure 3 C & H) pointing towards the

mesocolpium and, sometimes, unevenly distributed

granules. Each side of the colpus has a margo that is

composed of a distinct row of widely spaced, fused

and flattened spines (‘dragoncomb’; Figure 3 B, F &

H) with quite deep perforations between the teeth

(Figure 3 H). These two rows are generally fused

and look like a ribbon with a zigzag pattern on both

sides.

Remarks. All the Siberian A. turbidus specimens are

spineless on the polar projections (‘bald’), subispolar

and, on average, considerably smaller than the

specimen measured by Tschudy & Leopold (1971;

polar axes: 25–35 mm, equatorial axes: 31–45 mm).

The A. turbidus var. turbidus documented here only

superficially resembles an SEM image of A. aptus

Srivastava from the Lance Formation, Wyoming

(Farabee & Canright, 1986); the Siberian form

has much more rounded, angular-foveolate poles

and less wedge-shaped equatorial projections.

Differentiating Aquilapollenites turbidus var. turbidus

from A. turbidus var. compactus using only the LM is

very difficult, if at all possible.

Illustrated specimen. 3830VB07-9/1/1 and 3830VB07-

9/18/1

Aquilapollenites turbidus var. paucispinus var. nov.

(Figure 3 I–L)


Samoilovich & Mchedlishvili: Plate 67 Figure 3

cf. 1969 Aquilapollenites rombicus Samoilovich, in

Bratseva: 27 Figure 5

Description. Tricolpate (long colpi), compact

subisopolar pollen grain (polar axis: 24 mm,

equatorial axis: 28 mm), with three big, rounded

wedge-like equatorial projections (10 mm, Figure 3

I, J). The polar projections are short (4–5 mm) and

smooth with few, faint perforations (Figure 3 L).

The equatorial projections are covered with very few

flattened, supratectal spines pointing towards the

mesocolpium (Figure 3 K). Each side of the colpus

margin has a margo that is characterized by a distinct

row of irregularly spaced, flattened and fused spines

that becomes more diffused towards the poles. Both

rows are generally fused and produce a zigzag ribbon

(Figure 3 L).

Remarks. Under the LM, this variant can be

recognized, if at all, only by the fewer spines and

more elongated equatorial projections.

r

Figure 7. A–D. Azonia calvata. E–J. A. recta. K–N. A. lindensis. A, E, H & K. LM overview of pollen grains (ca 6900). B–D, F, G, I, J &

L–N. SEM micrographs. B, C, F, I & L. Pollen grains in equatorial view: (F) grain with symmetrical outline and slit-like aperture; (I) grain

with asymmetrical outline and pore-like aperture; (L) grain with symmetrical outline and pore-like aperture. D, G, J, M & N. Details of

exine sculpture: (D) high columellae and muri of the polar flange and sponge-like aperture area, (G) verrucate aperture area; (J) polar

flange reticulum; (M) reticulum of polar flange, with high columellae; (N) pore-like aperture and rugulate to verrucate aperture area. Scale

bars – 1 mm (D, G, J, M & N); 10 mm (B, C, F, I & L).


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Illustrated specimen. 3830VB07-5b/25/1

Aquilapollenites turbidus var. compactus var. nov.

(Figure 3 M–P)



Description. Tricolpate (long colpi), compact

subisopolar pollen grain (polar axis: 28 mm,

equatorial axis: 36 mm), with three big, rounded,

wedge-like equatorial projections (12 mm, Figure 3

M & O). The polar projections are short (6–7 mm)

and smooth without any perforations (Figure 3 O),

whilst the equatorial projections are perforated

(Figure 3 P) and covered with several angular,

crystal-like supratectal spines that all point towards

the mesocolpium (Figure 3 O). Each side of the colpus

has a margo that is characterized by a distinct row of

fused, more-or-less regularly spaced, angular spines

(‘dragoncomb’). Both rows are generally fused to

produce a zigzag ribbon (Figure 3 O).

Remarks. Under LM, this variant is easily mixed up

with A. turbidus var. turbidus and might be

differentiated, if at all, only by its compact outline.


Occurrence. All variants are relatively common,

occurring in samples T4mu, T5b, T7, T9, T11,

T13, T14, and T15.

Aquilapollenites rombicus Samoilovich (Figure 4 A–D)


Samoilovich: Figure 3a, b (drawing), plate 1

Figure 3a–d



cf. 1969 Duplosporis borealis (Chlonova) Bonda-

renko, in Bratseva: Plate 18 Figures 2 & 4

non 1993 Aquilapollenites rombicus Samoilovich, in

Nichols & Sweet: Plate 1 Figures 14 & 33

cf. 1997 Aquilapollenites proteus Simpson, in

Takahashi: figs. 59–60 (SEM image)

Description. Tricolpate (long colpi), subisopolar

pollen grain (polar axis: 28 mm, equatorial axis:

33 mm) with three big, rounded wedge-like

equatorial projections (Figure 4 A, B) and a

perforated tectum, densely covered with scale-like

spines, which all point towards the mesocolpium

(Figure 4 B, C). The short polar projections are

perforate and the polar tectum looks foveolate. The

colpus margin displays a margo that is characterized

by a distinct row of densely spaced, flattened and

fused, blunt spines (‘dragoncomb’; Figure 4 D).

Locally, assembled masses of granules are visible

(Figure 4 C, D).

Remarks. The LM image can be misidentified as A.

aptus Srivastava (Farabee & Canright, 1986), but the

SEM image of A. rombicus displays a dense cover of

scaly spines, whereas spines in A. aptus are widely

spaced and the poles are more pointed. The sur-

face shown in SEM slightly resembles the sur-

face of Aquilapollenites scabridus Tschudy (SEM

image, 1969), which has more pronounced polar

projections. The LM images of A. rombicus

Samoilovich in Nichols and Sweet (1993; Plate 1

Figures 14 & 33) display two reticulated forms and

are assumed to be another species.

Illustrated specimen. 3830VB07-15b/19/1

Occurrence. Relatively rare taxon, occurring in

samples T13 and T15.

Aquilapollenites fergusonii sp. nov. (Figure 4 E–H)

cf. 1961 Aquilapollenites granulatus Mchedlishvili, in

Samoilovich & Mchedlishvili: Plate 68 Figure 1a

(LM); non 1b and 1c (drawings)

Derivatio nominis. The species name is in honour of

the palaeobotanist David K. Ferguson, who works

mostly in Neogene strata and is known for his plant

taphonomy work.


following additions: A. fergusonii differs from A.

granulatus Mchedlishvili by not having any big

spines. This is well illustrated by the more detailed

drawings of A. granulatus by Mchedlishvili

(Samoilovich & Mchedlishvili, Plate 68 Figure 1b,

c). A. fergusonii differs from A. rhombicus by the

elongated polar projections, with a prominent solid

tectate cap and broader scale-like spines and from

other taxa by the unique uniform cover of densely

spaced, broad, flattened scale-like spines (reptile-like).

Holotype. 3830 VB07-7a/15/1, Figure 4 E–H




Eastern Siberia.


(Campanian/Maastrichtian), base of lacustrine unit

(laminated clay and siltstone).


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Description Tricolpate (long colpi), subisopolar


38 mm), with three big, distally rounded equatorial

projections (length: 17 mm; Figure 4 E, F). The

equatorial area and projections are perforate and

densely covered with large, blunt scale-like spines

(1–3 mm min diameter; Figure 4 G), which point

towards the mesocolpium (reptile-like), whilst the

polar projections are perforate to fissurate and tend

to be fused towards the polar regions, to produce a

solid tectum (bald polar area) (Figure 4 H). The

colpus margin is characterized by a margo that is

composed of a distinct row of fused and flattened

scale-like spines (‘dragoncomb’).

Remarks and occurrence. A rare taxon occurring only

in sample T7.

Aquilapollenites spiceri sp. nov. (Figure 4 I–N)

cf .1965 Aquilapollenites procerus Samoilovich, in

Samoilovich: Figure 4a, b (drawings), Plate 2

Figure 1 a–d

Derivatio nominis. The species name refers to the

palaeobotanist Robert A. Spicer whose work in

palaeobotany covers a wide range, including

Cenophytic macrofossils, plant taphonomy and

climate modelling.


following additions: Aquilapollenites spiceri generally

has longer polar axes than equatorial axes, which is

the opposite of A. procerus. It differs from A. procerus

by its hemispheric, more-or-less tectate cap at the

polar areas and the margo, which in A. spiceri is

composed of fused, irregularly sized and spaced

spines. Aquilapollenites spiceri can be also

differentiated from A. procerus Samoilovich by the

perforated polar projections and their blunt scale-

like spines.

Holotype. 3830VB07-15/18/2, Figures 4 I–N

Paratype. 3830VB07-7/24/1, 3830VB07-9/24/2



Eastern Siberia.



(brownish-grey, non-stratified).

Description. Tricolpate (long colpi), subisopolar to

heteropolar pollen grain (polar axes: 35–45 mm,

equatorial axes: 51–56 mm) with broad, distally

rounded elongated projections (length: 21–23 mm;

Figure 4 I, J). Under LM and SEM the polar areas

display a clearly visible, small tectate hemispherical

cap with a few perforations that transform into a

distinct narrow band with more-or-less regular

brochi (Figure 4 I & N). The slim polar

projections (11–13 mm; 12–17 mm) are perforate to

micro-reticulate and densely covered with relatively

blunt, scale-like supratectal spines (Figure 4 J & N).

The equatorial area is perforate and densely covered

with angular scale-like spines (Figure 4 M), which

change into less densely packed, flattened, rounded,

scale-like spines towards the distal ends of the

equatorial projections (Figure 4 L). In between

these rounded scales, there are perforations and

small accumulations of granules. The distal ends of

the equatorial projections and the area adjacent to

the colpus margins are more-or-less free of scale-like

spines, conspicuously perforate and covered with

granules (Figure 4 K). Each side of the colpus

margin is characterized by a margo consisting of a

row of a few, irregularly spaced and sized, fused and

flattened spines (‘dragoncomb’; Figure 4 L). In

between the spines are large gaps with freestanding

columellae (Figure 4 M).

Remarks. This taxon is very variable in overall size

and shape and length of its polar and equatorial

projections. In most specimens, the polar axes are

definitely shorter than the equatorial axes. This is in

contrast to the LM images of A. procerus

(Samoilovich, 1965), which show longer polar

axes. On the other hand, a hand-drawn sketch

of one specimen of Aquilapollenites procerus

(Samoilovich, 1965: text Figure 4a, b, and Plate 2

Figure 1 a, b) displays a tectate hemispherical cap at

the polar area. On the other specimen (text

Figure 5a & b and Plate 2 Figure 2 a–d) the

hemispherical nature of the tectate cap is not

clearly visible.

Occurrence. This quite abundant pollen taxon occurs

in samples T9, T11, T14 and T15.

Aquilapollenites procerus Samoilovich (Figure 5 A–H)

1965 Aquilapollenites procerus Samoilovich, in

Samoilovich: text Figure 5 a, b, Plate 2 Figure 2a–d

Description. Variably sized, tricolpate (long colpi),

subisopolar pollen grains (polar axes: 29–50 mm,

equatorial axes: 27–41 mm), with three broad,

distally rounded equatorial projections (lengths:

11–17 mm; Figure 5 A, B & E, F) and extremely

elongated, club-shaped polar projections (lengths:

7–1 mm; 8–19 mm) with their greatest widths at the

poles. These tectate caps of the polar projections are


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smooth, perforate to foveolate and have a more-or-

less triangular outline (Figure 5 D & G), with

corners pointing towards the mesocolpium areas

(Figure 5 B & F). The other parts of the polar

projections are reticulate or microreticulate and are

partly covered with angular, relatively blunt spines

(Figure 5 G, H). The variably shaped equatorial

projections are perforate at the distal areas and

covered everywhere with numerous small scale-like

spines (flattened spines) that point in the direction of

the mesocolpium (Figure 5 C). Accumulations of

granules are mostly visible in the equatorial area.

Each colpus margin is characterized by a margo of a

distinct row of small, regularly spaced, fused and

flattened spines (Figure 5 C). If the two rows are

fused, they produce a zigzag ribbon.

Remarks. A. spiceri and A. procerus initially look very

similar under the LM but the latter generally has

longer and microreticulate to reticulate polar

projections, and longer polar axes than equatorial

axes. There are other differences, including the

tectate triangular shape at the polar areas, the club-

like shape of the polar projections, the distribution

and form of the scale-like spines on the equatorial

projections, and the regularly spaced flattened spines

at the margo. It is difficult to establish whether or

not one of the two grains of A. procerus described by

Samoilovich (1965) is actually A. spiceri.

Illustratedspecimen. 3830VB07-13-1/11/1and3830VB07-

9/26/1

Occurrence. A relatively frequent and common

species that occurs in samples T7, T9 T11, T13,

T14 and T15.

Aquilapollenites heteropolaris sp. nov. (Figure 5 I–

M)

Derivatio nominis. The species name refers to the

heteropolarity of the pollen grain.


following additions: Aquilapollenites heteropolaris can

be distinguished from A. procerus by the different

long polar projections (heteropolarity), which are

clearly perforate and not reticulate as in A. procerus,

and the fewer, larger and widely spaced flattened

spines on the equatorial projections. The taxon

differs from A. spiceri by the sculpture of the

equatorial projections: A. heteropolaris has few,

widely spaced and flattened spines up to the distal

end whilst A. spiceri has more densely spaced

rounded scale-like spines, but the distal ends are

only perforated and covered with distinct granules.

The margo of A. heteropolaris is composed of

regularly spaced spines, whereas A. spiceri has a

margo of irregularly sized and spaced, fused spines.

Holotype. 3830VB07-13-1/11/2, Figure 5 I–M




Eastern Siberia.


(Campanian/Maastrichtian), lacustrine shale (finely

laminated claystone, medium grey with brownish

tinge).

Description. Tricolpate (long colpi), heteropolar,

relatively compact pollen grains (polar axes: 28–

35 mm, equatorial axes: 36–40 mm), with three

broad, distally rounded equatorial projections

(lengths: 13–15 mm) and differently elongated

polar projections (7–13.5 mm; Figure 5 I, J). The

polar-regions are smooth and compact (slightly

triangular) with very few perforations (Figure 5 K).

The polar projections are perforate and covered with

more-or-less rounded, blunt spines that point

towards the poles (Figure 5 K). The equatorial

projections are slightly perforated near the colpi

and display irregularly scattered, flattened spines

(1–1.5 mmm length) that point towards the

mesocolpium area (Figure 5 L). Each margin of

the colpus has a margo of a distinct row of

fused, flattened spines (‘dragoncomb’; Figure 5

M); the two rows may be fused, producing a zigzag

ribbon.

Remarks and occurrence. A relatively rare taxon

occurring in samples T7 and T13

Aquilapollenites srivastavae sp. nov. (Figure 6 A–D)

Derivatio nominis. The species name is in honour to

the palynologist Satish K. Srivastava who worked on

Cretaceous palynology, particularly on the genus

Aquilapollenites.


following additions: Aquilapollenites srivastavae is

characterized by its unique, mainly verrucate

sculpture, which transforms at only the distal ends

of the equatorial projections into scale-like spines. It

differs from A. procerus by having a hemispheric

tectate cap and broader, more-or-less tectate polar

projections with distinct verrucate sculpture. A.

srivastavae can be differentiated from A. spiceri, by


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its shorter equatorial projections, which have densely

spaced, flattened spines at their distal ends and a

very narrow margo of small fused spines. A.

srivastavae has numerous and smaller sculpture

elements (verrucae) and thus differs from A.

heteropolaris, which is characterized by fewer and

bigger spines on the equatorial projection.

Holotype. 3830VB07-7/25/2, Figure 6 A–D

Paratype. 3830VB07-7/3



Eastern Siberia.


(Campanian/Maastrichtian), base of lacustrine unit

(laminated clay and siltstone).



36 mm) with three broad, distally rounded

equatorial projections (length: 14 mm) and

elongated but differently shaped (one wide and

one slim) polar projections (11 and 13 mm; Figure 6

A, B). The polar areas are compact, hemispherical

tectate caps with few perforations (Figure 6 D)

and the remaining polar projections are slightly

perforate and covered densely with verrucae. The

distal ends of the equatorial projections are

perforate and covered densely with small (0.5 mm)

flat spines, which transform towards the

mesocolpium area into more-or-less rounded

verrucae (Figure 6 C). Each colpus margin is

covered by a row of small, flattened, fused spines

representing the margo.

Remarks and occurrence. A rare taxon, occurring only

in sample T7.

Aquilapollenites ovatus sp. nov. (Figure 6 E–G)

Derivatio nominis. The species name ‘ovatus’ refers

to the more-or-less oval shape.


following additions: Aquilapollenites ovatus differs

from all the other genera by its unique sculpture of

scattered, minute rounded granule-like spines and

short wedge-shaped equatorial projections.

Holotype. 3830VB07-9/1/2, Figure 6 E–G




Eastern Siberia.


(Campanian/Maastrichtian), peatball (brown clay

matrix with plant hash) preserved within fluvial

channel facies.



18 mm), with three short, wedge-shaped equatorial

projections (ca. 3 mm; Figure 6 E, F). The polar

areas are smooth and faintly perforate, whilst the

areas of the polar and equatorial projections are

equally perforate and display more-or-less regularly

but widely spaced tiny rounded spines (0.2–0.3 mm

min diameter (Figure 6 G).

Remarks. The rounded granule-like spines are a

unique sculpture and are assumed to represent

strongly reduced spines that are known from many

of the other Aquilapollenites taxa described above.

Occurrence. Rare, occurs only in sample T9.

Genus Manicorpus Mchedlishvili emend. Srivastava

1968

Manicorpus tenue (Figure 6 H–K)

cf. 1961 Manicorpus tenue Mchedlishvili, in

Samoilovich & Mchedlishvili: Plate 71 Figure 4a–c

cf. 1969 Manicorpus tenue Mchedlishvili, in

Bratseva: Plate 18 Figures 1, 2 (LM images)

Description. Tricolpate, heteropolar (x/y504),

pollen grain (polar axis: 23 mm; equatorial axes:

32 mm) with three broad, perpendicularly oriented,

elongated equatorial projections and one reduced

pole (Figure 6 H, I). The polar areas are smooth and

perforate (Figure 6 K), whilst the remaining polar

projection is foveolate to micro-reticulate and

covered with very blunt supratectal spines

(Figure 6 J). The equatorial projections are

perforate and regularly covered with small,

flattened, blunt spines that point towards the

mesocolpium areas. Each colpus margin is

characterized by a margo consisting of small,

flattened, fused spines (Figure 6 K).

Remarks. According to Srivastava (1968) and

Srivastava & Rouse (1970), this shape with an x/y

ratio (length from the reduced pole to the equatorial

diameter divided by the length of the elongated pole

to equatorial diameter) of less than 0.5 should be

accounted to the genus Manicorpus. The taxon


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described here has bald polar areas and numerous

more flattened spines, in contrast to the North

American taxa, such as the strongly heteropolar

Aquilapollenites polaris, A. cosmos and A. firmus

(Srivastava, 1994: Plate 2 Figures 5–7) and A.

delicatus (Farabee,1990: Figure 5).

Illustrated specimen. 3830VB07-15/26/2.

Occurrence. This taxon occurs occasionally in

samples T5b, T7, T14, and T15.

Manicorpus truncatus sp. nov. (Figure 6 L–O)

Derivatio nominis. The species name ‘truncatus’

refers to the truncated equatorial projections.

Specific diagnosis As for the genus, but with the

following additions: Manicorpus truncatus differs

from M. tenue by its perforate to foveolate tectum,

foveolate polar areas, and the truncated shape of the

equatorial projections. It can be differentiated from

A. cosmos (Srivastava, 1994a, Plate 2 Figure 6),

which also has truncated equatorial projections, by

having bald (spineless) polar areas.

Holotype. 3830VB07-15-1/26/2, Figures 6 L–O

Paratype. 3830VB07-15/2673



Eastern Siberia.



(brownish-grey, non-stratified).

Description. Tricolpate, heteropolar (x/y50.4),

pollen grain (polar axis: 27 mm; equatorial axes:

32 mm), with three broad, elongated and distally

truncated equatorial projections (Figure 6 L, M).

The polar areas are smooth and perforate to

foveolate, whilst the remaining polar projection is

micro-reticulate and covered with very blunt spines

(Figure 6 O). Under the micro-reticulum a layer

with perforations is visible. The equatorial

projections are regularly perforate to foveolate and

evenly covered with small, flattened, scale-like spines

that point towards the mesocolpium areas (Figure 6

N). Each colpus margin has a margo consisting of a

row of small, flattened, fused spines.

Remarks. According to Srivastava (1968) and

Srivastava & Rouse (1970), this shape with an x/y

ratio (length from the reduced pole to the equatorial

diameter divided by the length of the elongated pole

to equatorial diameter) of less than 0.5 should be

accounted to the genus Manicorpus.

Occurrence. This is a relatively rare taxon occurring

in samples T7, T14 and T15.

Genus Azonia Samoilovich 1961

Azonia calvata (Samoilovich) Wiggins (Figure 7 A–D)

1961 Kryshtofoviana calvata Samoilovich, in

Samoilovich & Mchedlishvili: Plate 78 Figure 4a–b

(LM images)

non 1961 Wodehouseia calvata Samoilovich var.

lindensis, in Samoilovich & Mchedlishvili: Plate 3

Figure 4a–d (LM images)

1961 Deplexipollis calvatus Chlonova, in Chlonova:

Plate 13 Figures 97/97a (drawing)

cf. 1976 Azonia calvata (Samoilovich) Wiggins, in

Wiggins: Plate 3 Figures 5, 6

1981 Wodehouseia calvata Chlonova, in Chlonova:

Plate 2 Figure 10

non 1994 Azonia calvata (Samoilovich) Wiggins,

in Nichols & Sweet: Plate 1 Figure 23

Description. Aperturate (4 pore-like apertures,

‘binigeminate’ after Wiggins, 1974) pollen grains

with an elliptical to rounded rectangular outline in

equatorial view (long equatorial axis: 35 mm;

definition after Samoilovich, 1961; Wiggins, 1976).

The equatorial flanges are protruding and tectate

(Figure 7A–C), whereas the polar areas (axis 26 mm;

definition after Samoilovich, 1961; Wiggins, 1976) are

characterised by a reticulated flange composed of very

high columellae (1.52v2 mm) and muri (ca. 1.5–

2 mm; Figure 7 C, D), whilst the equatorial flanges are

tectate to slightly perforate. The inner parts of the

flanges are smooth and tectate and surround the

aperture area, which is perforate and displays a

sponge-like and microverrucate texture at high

magnification (69500 under SEM; Figure 7 D).

Remarks. This taxon is easily recognizable under the

LM and apparently was also found on the Alaska

Peninsula (Wiggins, 1976).

Illustrated specimen. 3830VB07-14/21/1 and 3830VB-

07-15/26/4

Occurrence. Not so common taxon in samples T14

and T15.

Azonia recta Bolchovitina (Figure 7 E–J)

1959 Pollenites rectus Bolchovitina, in Bolcho-

vitina: Plate 8 Figure 112a, b (drawing)


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1961 Azonia recta (Bolchovitina) Samoilovich, in




1967 Azonia sp. in Samoilovich: Plate 3 Figure 29


Chlonova: Plate 2 Figure 14

cf.? 1976 Azonia recta (Bolchovitina) Samoilovich,

in Wiggins: Plate1 Figures 14–16

cf.?1994 Azonia recta Bolchovitina) Samoilovich,

in Dawson et al.: Plate 1 Figure 11

Description. A flattened aperturate (4 pore- and/or

short colpi-like apertures, depending on the state of

preservation), slightly heteropolar or isopolar pollen

grain. Well preserved forms display colpi (Figure 7

F), badly preserved forms tend to have more

destroyed aperture areas (Figure 7 I). Pollen grains

have an elongate elliptical, slightly lobate at the poles

(creating two small protrusions at the pole), outline

in equatorial view (Figure 7 E, F & H, I). The long

equatorial axis varies between 24–36 mmm and the

polar axis between 14–23 mm. In some examples,

one polar area may be more concave than the other

and one may be more-or-less straight (Figure 7 I).

The elliptical field around the apertures is micro-

verrucate to slightly micro-rugulate, perforate

(Figure 7 G) and the rest (polar and ?equatorial

flanges) is reticulate to foveolate, the brochi are

irregularly shaped (Figure 7 J).

Remarks. Either a single taxon with considerable

variation in size, polarity and aperture shape or a

taxon with many variants. This taxon also occurs in

the southern part of the Khatanga-Lena-

Subprovince and on the Alaska Peninsula

(Bolchovitina, 1959; Wiggins, 1976). However,

Chlonova (1981) stated, that the pollen types

included into Azonia recta by Wiggins (1976) is not

identical with the holotype, which has four polar

protrusions with highly reflective (light areas) that

look like pores.

Illustrated specimen. 3830VB07-7/13/1 and 3830VB07-

15/18/1

Occurrence. A frequently and relatively common

taxon occurring in samples T7, T9, T10, T11,

T14, T15.

Azonia lindensis nom. nov. (Figure 7 K–N)

1965 Wodehouseia calvata Samoilovich var.

lindensis, in Samoilovich: Plate 3 Figure 4a, b

1967 Wodehouseia calvata Samoilovich var. linden-

sis, in Samoilovich: Plate 3 Figure 26

Description. Aperturate (4 pore-like apertures;

Figure 7 N), isopolar pollen grain with a more-or-

less elliptical outline in equatorial view (Figure 7 K,

L). The distal ends of the long equatorial axis

(30 mm) are perforate to foveate and the more

expanded polar flanges are reticulate at the distal

poles and more foveate to perforate towards the

apertures. The height of the columellae is more than

2 mm, the height of the muri around 1 mmm

(Figure 7 M). The field around the apertures is

perforate and micro-rugulate (Figure 7 N).

Remarks. The aperture region (rugulate) and the

equatorial flange (not so prominent and generally

microreticulate) differ considerably from Azonia

calvata, described above, with its prominent,

tectate equatorial flanges and microverrucate

aperture region.


Occurrence. Relatively rare in samples T9 and T10.

Discussion

The overall north hemispheric distribution of

Aquilapollenites, Manicorpus, and Azonia led to the

assumption that some of the taxa had a circumpolar

distribution during the Upper Cretaceous. This is

probably true for two of the Azonia species as well as

for three reticulate and one spinulose Aquilapollenites

species that occur both in the Vilui Basin and in

Alaska, Canada and the Rocky Mountain area of the

USA. Unfortunately, there are some uncertainties

about the Azonia species: Azonia recta and A.

calvata, both of which were also described from the

Alaskan Peninsula (in Wiggins, 1976) and Canada

(north-western Alberta: Dawson et al., 1994;

Northwest Territories & Yukon Territory: Nichols

& Sweet, 1994) may not be the same species as the

Siberian ones. Chlonova (1981) claimed that the A.

recta pollen described by Wiggins (1976) is not

identical with the type material (see systematic part)

and that the pollen ascribed to Azonia calvata

(Wiggins, 1976; Nichols & Sweet, 1993) does not

really look identical to the Russian material. Further

SEM investigations of the Alaskan material would

solve this problem by establishing whether they are

variants of the two species, or entirely different

species.

Of the four reticulate and striato-reticulate spe-

cies, only the rare A. samoilovichiae is endemic to the

Khatanga-Lena-Subprovince, whilst A. chlonovae

and A. hermanii were also discovered in Alaska by

Tschudy (1969). This leads to the speculation as to

whether or not the Alaskan Peninsula belonged to


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the Khatanga-Lena-Subprovince during the Late

Cretaceous (Maastrichtian).

Further, there are two Vilui Aquilapollenites species

with an even wider distribution: Aquilapollenites

mchedlishvilii (without equatorial ring-like thinning)

and the spinulose bald Aquilapollenites turbidus, have

both been found in the Lower Campanian to upper

Maastrichtian strata of Canada (Dawson et al.,

1994; Nichols & Sweet, 1993) and the Rocky

Mountain area (Montana, Dakota, Wyoming,

Colorado - Farabee & Canright, 1986; Tschudy &

Leopold, 1971).

However, despite some doubts about the Azonia

affiliations, the species discussed above indicate a

strong floral connection between Siberia, Alaska and

the Rocky Mountain (western Cordillera) area at

around Middle (Senonian) to Late Cretaceous time,

a hypothesis, which has been already hinted at by

Norris et al. (1975) and Stanley (1970).

Aquilapollenites turbidus might be the oldest (since

Lower Campanian) and also the most southerly

occurring (Colorado) of the ‘‘cosmopolitan’’

Aquilapollenites species. It has been regularly con-

fused with Aquilapollenites quadrilobus by Russian and

Japanese workers. However, Aquilapollenites quad-

rilobus from North America is characterized by

spines on the polar areas, whilst Aquilapollenites

turbidus forms from Siberia, Sakhalin, Canada and

USA have none. The variability in shape and

sculpture of the Siberian Aquilapollenites turbidus

forms (SEM) can be anticipated on the type material

Figured by Tschudy and Leopold (1971, Plate 2

Figures 7–10), who described the species first

from the Rocky Mountain area, occurring in Lower

Campanian to upper Maastrichtian strata (Tschudy

& Leopold, 1971: Montana to Colorado). One

might speculate that this species evolved further

and developed several variants since the Lower

Campanian.

All the remaining spinulose Aquilapollenites and

Manicorpus species from Siberia and Sakhalin of

Upper Campanian to Maastrichtian age are bald,

being spineless on the poles. This is an important

characteristic that separates them from all the other

spinulose forms (except A. turbidus) documented

from the southerly lying Yenisei-Amur-Subprovince,

Scotland and North America. Furthermore,

the spinulose Aquilapollenites and Manicorpus all

display the very distinct ‘dragoncomb’-like margo

(two distinct rows of fused spines) that can also

be partly observed (often not fused spines) on a

few North American and Scottish Maastrichtian

forms, such as A. delicatus, A. quadrilobus, A. aptus,

and A. polaris (see SEM images in: Farabee,

1990; Farabee & Canright 1986; Srivastava, 1975,

1994a, b).

Together with Aquilapollenites samoilovichiae,

nearly all (except Aquilapollenites turbidus) of the

described spinulose Aquilapollenites and Manicorpus

species are endemic to the Khatanga-Lena-

Subprovince of northern Asia. It is suggested, that

during the uppermost Cretaceous the evolution of

the spinulose north Russian taxa led to the loss of

spines on the polar projections resulting in bald

poles and probably to the further development of a

very distinct ‘dragoncomb’- like margo consisting of

two rows of fused flattened spines.

Conclusions

The Timerdyakh Formation of the Vilui Basin

yielded nine samples that were, amongst others

species, rich in Aquilapollenites (13 taxa), Manicorpus

(two taxa), and Azonia (three taxa) species. Nine

new species (eight Aquilapollenites and one

Manicorpus species) have been described and

distinguished by LM and SEM analysis. All

spinulose forms, including Manicorpus are charac-

terized by spineless polar areas and margos (colpus

margins) that are reminiscent of a ‘dragoncomb’.

Three of the four reticulate-striatoreticulate forms

are distinguished by a ring-like thinning in the

equator. Two of the three Azonia species and

Aquilapollenites chlonovae, and A. hermanii were

recorded from the Alaskan peninsula, whilst A.

turbidus and A. mchedlishvilii were recorded even

further south from the Rocky mountain area. Of the

18 taxa identified, 12 are suggested to be endemic

to the Khatanga-Lena-Subprovince and represent

the unique evolution of these form genera during

the Upper Cretaceous.

Acknowledgements

We thank A. Ahlberg, A. Herman, M. Moiseeva,

and R. A. Spicer for collecting the samples in the

Vilui Basin, N. Kravcik for chemical sample

preparation and digitizing the photographic plates,

D. Ferguson for helping with the nomenclature and

discussions, Hugh Rice for correcting the English,

and A. Ostrovsky and B. I. Sharenkoff for help with

Russian names. The comments of the reviewers D. J.

Nichols and A. R. Sweet are greatly acknowledged.

The study was funded by RFBS/INTAS grant

950949.

References

Bolchovitina, N. A. (1959). Sporen und Pollenkomplexe der

mesozoischen Ablagerungen des Viluibeckens und ihre stratigra-

phische Bedeutung. Moskva: USSR Acad. Sci. Publ. H. Tr.

Geol. Inst. USSR Acad. Sci. 24 (in Russian).


Dow

nloa

ded

by [

Uni

vers

ity o

f O

klah

oma

Lib

rari

es]

at 0

0:48

28

Aug

ust 2

014

Bratseva, G. M. (1965). Pollen and spores in Maastrichtian deposits

of the Far East. Moskva: USSR Acad. Sci. Publ. H. Tr. Geol.

Inst. USSR Acad. Sci. 129 (in Russian).

Bratseva, G. M. (1969). Palynological studies of upper Cretaceous

and Palaeogene of the Far East. Moskva: USSR Acad. Sci. Publ.

H. Tr. Geol. Inst. USSR Acad. Sci. 207 (in Russian).

Chlonova, A. F. (1961). Pollen und Sporen der hoheren

Oberkreide im Ostteil der westsibirischen Tiefebene.

Novosibirsk: Siberia Div. USSR Acad. Sci. Tr. Geol. Inst.

USSR Acad. Sci. 7 (in Russian).

Chlonova, A. F. (1967). Possible botanical relationships of the

pollen of the morphological type ‘‘Oculata’’. Rev. Palaeobot.

Palynol., 5, 217–226.

Chlonova, A. F. (1981). Senonian (Late Cretaceous) palynofloral

provinces in circumpolar areas of the northern hemisphere.

Rev. Palaeobot. Palynol., 35, 315–324.

Dawson, F. M., Kalkreuth, W. D. & Sweet, A. R. (1994).

Stratigraphy and coal resource potential of the Upper

Cretaceous to Tertiary strata of northwestern Alberta. Geol.

Surv. Can. Bull., 466, 1–64.

Erdtman, G. (1971). On the affinities of Aquilapollenites Rouse.

Grana, 11, 15–17.

Farabee, M. J. & Canright, J. E. (1986). Stratigraphic palynology

of the lower part of the Lance Formation (Maastrichtian) of

Wyoming. Palaeontographica B, 199, 1–89.

Farabee, M. J. (1990). Triprojectate fossil pollen genera. Rev.

Palaeobot. Palynol., 65, 341–347.

Funkhouser, J. W. (1961). Pollen of the genus Aquilapollenites.

Micropalaeontology, 7, 193–198.

Jarzen, D. M. & Norris, G. (1975). Evolutionary significance and

botanical relationship of Cretaceous angiosperm pollen in the

western Canadian interior. Geosci. Man, 11, 47–60.

Jarzen, D. M. (1977). Aquilapollenites and some Santalalean

genera. Grana, 16, 29–39.

Krutzsch, W. (1970). Taxonomie syncolp(or)ater und morpho-

logisch benachbarter Pollengattungen und-Arten (Sporae

dispersae) aus der Oberkreide und dem Tertiar. Teil II:

Aquilapollenites (5Triprojectacites). Pollen Spores, 12, 103–122.

Martin, A. R. H. (1968). Aquilapollenites in the British Isles.

Palaeontology, 11, 549–553.

Nichols, D. J. & Sweet, A. R. (1993). Biostratigraphy of Upper

Cretaceous non-marine palynofloras in a north-south transect

of the Western Interior basin. In W. G. E. Caldwell & E. G.

Kauffmann (Eds). Evolution of the Western Interior Basin

(pp. 539–584). Calgary AL: GAC Sp. Pap. 39.

Norris, G., Jarzen, D. M. & Awai-Thorne, B. V. (1975).

Evolution of the Cretaceous terrestrial palynoflora in western

Canada. Geol. Assoc. Can. Spec. Publ., N13, 333–364.

Norton, N. J. & Hall, J. W. (1969). Palynology of the Upper

Cretaceous and Lower Tertiary in the type locality of the Hell

Creek Formation, Montana, U.S.A. Palaeontographica B, 125,

1–64.

Samoilovich, S. R. (1965). The description of new pollen species

of the Upper Cretaceous angiospermic flora. Tr. VNIGRI, 239,

121–141 (in Russian).

Samoilovich, S. R. (1967). Tentative botanico-geographical

subdivision of northern Asia in Late Cretaceous time. Rev.

Palaeobot. Palynol., 2, 127–139.

Samoilovich, S. R. & Mchedlishvili, N. D. (1961). Pollen and

spores of Western Siberia. Jurassic to Palaeocene. Leningrad

(St. Petersburg): State Sci.-Techn. Publ. House

(Gostoptechizdat). Tr. VNIGRI, 177 (in Russian).

Srivastava, S. K. (1966). Upper Cretaceous microflora

(Maastrichtian) from Scollard, Alberta, Canada. Pollen

Spores, 8, 497–552.

Srivastava, S. K. (1968). Reticulate species of Aquilapollenites and

emendation of genus Manicorpus Mchedlishvili. Pollen Spores,

10, 665–699, 7 pls.

Srivastava, S. K. (1972). Palaeoecology of pollen genera

Aquilapollenites and Manicorpus in Maastrichtian deposits of

North America. In J. E. Gill (Ed.), Proc. 24th Int. Geol. Congr.,

Montreal 1972, Sect. 7 Palaeontol (pp. 111–120). Gardenvale,

Que.: Harpell’s Press Coop.

Srivastava, S. K. (1975). Maastrichtian microspore assemblages

from the interbasaltic lignites of Mull, Scotland.

Palaeontographica B, 150, 125–156.

Srivastava, S. K. (1981). Evolution of Upper Cretaceous

phytogeoprovinces and their pollen flora (La Habra, Calif.,

U.S.A.). Rev. Palaeobot. Palynol., 35, 155–173.

Srivastava, S. K. (1994a). Evolution of Cretaceous phytogeopro-

vinces, continents and climates. Rev. Palaeobot. Palynol., 82,

197–224.

Srivastava, S. K. (1994b). Palynology of the Cretaceous-Tertiary

boundary in the Scollard Formation Alberta of Canada, and

global KTB events. Rev. Palaeobot. Palynol., 83, 137–158.

Srivastava, S. K. & Rouse, G. E. (1970). Systematic revision of

Aquilapollenites Rouse 1957. Can. J. Bot., 48, 1591–1601.

Stanley, E. A. (1961). The fossil pollen genus Aquilapollenites.

Pollen Spores, 3, 329–352.

Stanley, E. A. (1970). The stratigraphical, biogeographical,

paleoautoecological and evolutionary significance of the fossil

pollen group Triprojectacites. Bull. Georg. Acad. Sci., 28,

1–44.

Takahashi, M. (1997). Fossil spores and pollen grains of

Cretaceous (Upper Campanian) from Sakhalin, Russia. J.

Pl.Res., 110, 283–298.

Tschudy, B. D. (1969). Species of Aquilapollenites and Fibulapollis

from two Upper Cretaceous localities in Alaska. U.S. Geol.

Surv. Prof. Pap, 643-A, 1–48.

Tschudy, B. D. & Leopold, E. B. (1971). Aquilapollenites (Rouse)

Funkhouser–selected Rocky Mountain taxa and their strati-

graphic ranges. In R. M. Kosanke & A. T. Cross (Eds),

Symposium on Palynology of the Late Cretaceous and Early

Tertiary. Geol. Soc. Am. Sp. Pap., 127, 113–167.

Vachrameev, V. A. & Pushcharovski, Y. M. (1954). On the

geological history of the Vilui Depression and the adjacent part

of the Priverkhoyansk Marginal Trough. Geol. Quest. Asia 1,

588–628.

Wiggins, V. D. (1976). Fossil Oculata pollen from Alaska. Geosci.

Man, 15, 51–76.


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Documents

Upper Cretaceous pollen flora from the Vilui Basin, Siberia: Circumpolar and endemic Aquilapollenites , Manicorpus , and Azonia species