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Mauldinia hirsuta sp. nov., a New Member of the Extinct Genus Mauldinia (Lauraceae) from the Late Cretaceous (Cenomanian‐Turonian) of Kazakhstan


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A new species of the extinct genus Mauldinia (Mauldinia hirsuta) is based on three-dimensionally preserved inflorescence fragments and flowers from the Late Cretaceous (Cenomanian-early Turonian) of the Sarbay Quarry, northwestern Kazakhstan, Central Asia. It is distinguished from previously described species of Mauldinia by its smaller size and the broad, obovate, sometimes angular outline of the lateral inflorescence units, narrow floral tube, and the presence of a dense indumentum of stiff long hairs occurring on all surfaces of the inflorescence axis and lateral inflorescences units, basal part of inner tepals, as well as stamen filaments and carpels. The new finding considerably extends the geographic range of Mauldinia and provides additional evidence for a floristic link between eastern North America, Europe, and Central Asia during the mid- and Late Cretaceous. In addition to the distinct Mauldinia fossils, the Sarbay flora have also yielded a few isolated lauraceous floral organs that are probably part of the Mauldinia plant.
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Mauldinia hirsuta sp. nov., a New Member of the Extinct Genus Mauldinia (Lauraceae) from the
Late Cretaceous (Cenomanian‐Turonian) of Kazakhstan
Author(s): Suembikya Frumin, Helena Eklund, and Else Marie Friis
International Journal of Plant Sciences,
Vol. 165, No. 5 (September 2004), pp. 883-895
Published by: The University of Chicago Press
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Suembikya Frumin,* Helena Eklund,yand Else Marie Friis1
*Department of Evolution, Systematics, and Ecology, Hebrew University, Givat-Ram, 91904 Jerusalem, Israel; and
yDepartment of Palaeobotany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden
A new species of the extinct genus Mauldinia (Mauldinia hirsuta) is based on three-dimensionally preserved
inflorescence fragments and flowers from the Late Cretaceous (Cenomanian–early Turonian) of the Sarbay
Quarry, northwestern Kazakhstan, Central Asia. It is distinguished from previously described species of
Mauldinia by its smaller size and the broad, obovate, sometimes angular outline of the lateral inflorescence
units, narrow floral tube, and the presence of a dense indumentum of stiff long hairs occurring on all surfaces
of the inflorescence axis and lateral inflorescences units, basal part of inner tepals, as well as stamen filaments
and carpels. The new finding considerably extends the geographic range of Mauldinia and provides additional
evidence for a floristic link between eastern North America, Europe, and Central Asia during the mid- and Late
Cretaceous. In addition to the distinct Mauldinia fossils, the Sarbay flora have also yielded a few isolated
lauraceous floral organs that are probably part of the Mauldinia plant.
Keywords: angiosperms, Cenomanian-Turonian, fossil flowers, fossil inflorescences, Late Cretaceous,
Kazakhstan, Lauraceae, Mauldinia hirsuta.
The Lauraceae are a large eumagnoliid family (sensu Soltis
et al. 2000) with 2500–3500 species of mainly pantropical
distribution (Rohwer 1993). The fossil record documents
that the family was diverse and widespread by the early Late
Cretaceous. The Cretaceous record is particularly rich in dis-
persed leaves (Upchurch 1984; Crabtree 1987; Kvac
ˇek 1992),
but there is also an increasing number of fossil reproductive
structures such as inflorescences and flowers (Drinnan et al.
1990; Crane et al. 1994; Herendeen et al. 1994; Mickle
1996; Eklund and Kvac
ˇek 1998; Eklund 1999, 2000; Taka-
hashi et al. 1999, 2001). Fossil wood is less common (Heren-
deen 1991; Poole et al. 2000), and pollen grains are rare
because of the weakly developed pollen wall (Herendeen
et al. 1994).
Assessing the systematic and biogeographic diversification
in early Lauraceae on the basis of isolated fossil organs is not
a straightforward task. Flowers of extant Lauraceae generally
exhibit the same basic structure with variation mostly ex-
pressed in the androecium, and while they are usually easily
recognized as Lauraceae, they are difficult to classify pre-
cisely in the family on the basis of the flowers alone (Rohwer
1993). Leaves of extant Lauraceae also show rather uniform
morphology, and it is difficult to classify fossil material on
the basis of leaf morphology and epidermal features alone.
Therefore, many fossil Lauraceae are grouped in unnatural
organ genera such as Laurophyllum Go
¨ppert (1854) for
leaves and Laurocarpum Reid & Chandler (1933) for fruits.
The extinct genus Mauldinia Drinnan et al. (1990) is the
earliest Lauraceae for which inflorescence and floral details
are known. Although exhibiting a rather general floral plan,
fossils assigned to Mauldinia clearly constitute a natural spe-
cies complex defined by their unique and distinct compound
inflorescences with flattened bilobed lateral inflorescence
units bearing sessile trimerous flowers. Mauldinia was first
described from the Cenomanian of North America on the
basis of fragments of inflorescences and isolated flowers
assigned to Mauldinia mirabilis (Drinnan et al. 1990).
Mauldinia was subsequently discovered also from the Ceno-
manian of the Czech Republic (Mauldinia bohemica; Eklund
and Kvac
ˇek 1998) and from the Santonian of North America
(Mauldinia sp.; Herendeen et al. 1999). In this work, we de-
scribe a new species (Mauldinia hirsuta) from the Cenoma-
nian or Turonian Sarbay flora (northwestern Kazakhstan).
This discovery considerably extends the geographical range
of Mauldinia and provides additional evidence of a floristic
link between North America, Europe, and Central Asia dur-
ing the mid-Cretaceous. A similar distribution was shown
previously for two other mesofossils from the Sarbay flora,
the Late Cretaceous Liriodendroidea Knobloch & Mai
(1984) (Magnoliaceae) known from seeds and fruits (Knob-
loch and Mai 1986; Frumin and Friis 1996; Frumin and
Friis 1999) and Platanaceae known from fossil flowers (Friis
et al. 1988; Krassilov and Shilin 1995). A floristic link was
also indicated earlier on the basis of palynological studies
that placed northwestern Kazakhstan in the easternmost part
of the Aptian–Cenomanian/Turonian Boreal–European paly-
nofloral subprovince on the border to the Boreal-Arctic
1Author for correspondence; e-mail
Manuscript received October 2003; revised manuscript received February
Int. J. Plant Sci. 165(5):883–895. 2004.
Ó2004 by The University of Chicago. All rights reserved.
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subprovince (Herngreen and Chlonova 1981; Herngreen et al.
Material and Methods
The fossils described here are from the Sarbay mesofossil
flora extracted from sediment samples provided by S. G. Zhi-
lin (St. Petersburg), who collected them in 1969 from the Sar-
bay Iron Quarry (52°589000N, 63°079000E) near the town of
Rudnyy, Kustanay Region, northwestern Kazakhstan (Frumin
and Friis 1996, 1999). The fossil-bearing sediments occur in
a small depression in the southern part of the western wall of
the quarry at level VI from the base (S. G. Zhilin, personal
communication). The terrestrial Cretaceous sediments in the
Sarbay area are referred to as two sedimentary suits, which
both contain plant fossils. The Novokozyrevskaya suite is in-
terpreted as lagoonal deposits and consists mainly of kaolin-
itic clays rich in bauxite and iron. The overlaying Shetirgiz
suite, from which the Sarbay mesofossil flora was recovered,
was probably deposited in a lacustrine to alluvial environ-
ment. The plant bed is ca. 1 m thick and consists of gray to
dark gray sandy clay with subordinate lenses of sand. It over-
lays a ca. 6-m-thick horizon of bauxitic clay and a 5–15-m-
thick weathered sequence and is superimposed by marine
sands of the Santonian-Campanian Ayat suite containing the
extinct molluscs Inoceramus cardissoides Goldfuss and Ino-
ceramus pachti Arck. and leaf imprints of Geinitzia formosa
Heer typical for the Senonian (Levina et al. 1990). The Creta-
ceous sequence is discordantly overlain by Palaeogene and Qua-
ternary sediments (S. G. Zhilin, personal communication).
Palynological and geological data indicate a Cenomanian–
early Turonian age of these two suites; the basal part of the
Novokozyrevskaya suite may extend into the late Albian
(Levina et al. 1990).
The mesofossil material was extracted and prepared for
further investigation using standard methods of sieving and
chemical treatment (Friis et al. 1988). Specimens selected for
SEM were mounted on stubs using nail polish and coated
with ca. 100 A
˚of gold. They were then investigated using
a Phillips 515 scanning electron microscope at 15 kV and/or
a Hitachi S-4300 field emission scanning electron microscope
at 5 kV. Specimens for cuticle preparations were treated in
Schulze’s solution (NHO
and KClO
) and washed in dis-
tilled water. After removal of the oxidation products by
KOH (<5%), the specimens were washed again in distilled
water and mounted on stubs for SEM investigations or embed-
ded in glycerine or Histo-mount for light microscope studying
using a Dialux 20 (Leitz, Wetzlar, Germany) equipped with
Wild Photoautomat MPS 55 and a Zeiss Axioskop 2 Plus
equipped with an Axiocam and an Axiovison 3.1 software
(Zeiss). Measurements were made from SEM and Axioskop
micrographs and in light microscope. Specimens described
in this work will be stored in the collections of the Komarov
Botanical Institute, St. Petersburg, Russia.
The Sarbay Mesofossil Flora
In total, 56 taxa have been described from the Shetirgiz
suite. The study of the mesofossil flora is only in its initial
phase, and most taxa remain to be described. The mesofossil
flora is very rich and consists of three-dimensional, generally
well-preserved lignite fossils. It includes a variety of mega-
spores of Isoetaceae, leaf fragments and sporangia of ferns,
as well as twigs, needles, cones, cone scales, and seeds of
conifers, including one species of the extinct taxodiaceous ge-
nus Alapaja (Frumina et al. 1995) and abundant angiosperm
fossils. In addition to the species of Mauldinia described in
this work, six angiosperm taxa have been described previ-
ously from the mesofossil flora. These include four species of
the extinct magnoliaceous genus Liriodendroidea (Frumin
and Friis 1996, 1999) and two species assigned to extinct
taxa of Illiciaceae, Illiciospermum pusillum and aff. Illicio-
spermum sp. (Frumin and Friis 1999). Shilin (1986) also de-
scribed a leaf macroflora from the Sarbay Quarry. This flora
is dominated by impressions of platanaceous leaves. Platana-
ceous staminate heads, Sarbaya radiata Krassilov & Shilin
(1995), and fruits assigned to an extinct genus and species,
Donica zhilinii Hvalj (2001) of uncertain affinity, are further
angiosperm meso- and macrofossils described from the Sar-
bay quarry. Rich palynological assemblages from the same
site (Levina et al. 1990) are characterized by abundant ferns,
while the occurrence of conifers and angiosperms varied con-
siderably between the assemblages ranging from 0% to 50%.
Among the conifers, Pinaceae may constitute up to 20%.
Systematic Description
Genus—Mauldinia Drinnan, Crane, Friis & Pedersen
Species—Mauldinia hirsuta Frumin, Eklund &
Friis sp. nov. (Figs. 1–6)
Derivation of specific epithet. Hirsute (Latin): covered
with fairly coarse and stiff long erect or ascending straight
hairs (Stearn 1998).
Diagnosis. Inflorescence axis with two different types of
scars. Lateral inflorescence units broadly obovate in outline,
sometimes with angular margins. Lobes of lateral units more
or less obtriangular in outline. Scales of lateral units membra-
nous without perforations. Each lateral unit with five sessile
flowers. Outer tepals short, broadly triangular; inner tepals
longer, narrowly obovate to spathulate with an indistinct keel.
Receptacle concave, forming a narrow floral tube. Staminal
appendages flat and differentiated into stalk and triangular
head. Staminodes short, narrow, cuspidate, and dorsiventrally
flattened. Resin bodies globular, occurring in tissue of lateral
inflorescence units and carpels. Indumentum dense, occurring
on all surfaces except staminodes and staminal appendages.
Subsidiary cells of stomata often divided several times.
Holotype. 1700a-51 (plates 1E,3A,4A–4C).
Other specimens. 1700a-44; 1700a-52–1700a-57; 1700b-
56; 1700b-66–1700b-67; 1700b-70–1700b-76; 1700b-101–
1700b-111; 1700b-122; 1700b-148–1700b-149; 1700b-184–
1700b-186; 1700b-188–1700b-196; 1700b-198–1700b-249.
Ca. 1500 specimens.
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Fig. 1 Mauldinia hirsuta sp. nov. A, Lateral units attached to inflorescence axis; note the bract supporting the lateral unit (sb) (1700b-203). B,
C, Inflorescence axes showing big scars with sinuous apex and decurrent keel-shaped base (dr), bracts supporting the lateral units (sb), and smaller
rhomboidal scars (rs) (1700b-205, 1700b-204). D, Adaxial view of deeply bilobed lateral unit showing rounded attachment scar (as) and scalelike
structures (1700b-70). E, Abaxial view of lateral unit with several attached flowers (holotype: 1700a-51). F, Abaxial view of lateral unit showing
a thick cuticle (1700b-149). G, Detail of Bshowing decurrent keel-shaped base (dr) at the base of scar after detached lateral unit, bract supporting
the lateral unit (sb), and thick cuticle of axis with hair bases. All SEM. Scale ¼1 mm, except for G.
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Fig. 2 Mauldinia hirsuta sp. nov. A, Lateral unit showing the apical rim, three flowers () still attached, and sterile scales (sc) (1700b-148). B,
Adaxial view of bilobed lateral unit showing five scars from detached flowers (fs) (1700b-107). C, Lateral unit with carpel (c) at the base and two
flowers () sitting on lobes (1700b-211). D, Isolated lobe of lateral unit with flower bud (b) (1700b-240). E, Base of lateral unit (adaxial view)
showing basal part of two flowers () still attached, flower tube (ft) formed by receptacle, and floral bract (fb) at the base of flower (1700a-54). F,
Detail of Dshowing floral bud with incurved inner tepals. All SEM. Scale ¼1 mm, except for F.
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Fig. 3 Mauldinia hirsuta sp. nov. A, Lateral unit with flowers showing attachment scar, bracts supporting the lobes (lb), and several flowers
with short outer (ot) and long inner tepals (it) (holotype: 1700a-51). B, Detail of cuticle of lateral unit showing hair bases (outer surface) (1700b-
149). C, Adaxial view of lateral unit showing sterile scales (sc) (1700a-54). DF, Detached flowers showing short flower tube (ft) and outer (ot)
and inner tepals (it) (1700b-101; 1700b-103; 1700b-104). G, Fragmented flower showing the inner tepal (it) and outer whorl of stamens (st)
(1700b-102). H, Cellular details of lamina of inner tepal (outer surface) (1700b-185). I, Cellular detail of adaxial side of lateral unit showing
indumentum (1700b-211). All SEM.
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Type locality. Sarbay Quarry, near the town Rudnyy,
Northern Kazakhstan (52°589000N, 63°079000E).
Age. Late Cretaceous (Cenomanian-Turonian).
Dimensions. Maximal length of axis fragments: 5.5 mm;
breadth of axis: 0.8–(1.47)–2.0 mm. Larger scars on axis:
0.7–(1.17)–2.0 mm 30:35–(0.56)–0.8 mm; smaller scars:
0.15–0.23 mm in diameter. Bracts supporting lateral inflores-
cence units: 0.4 mm long; 0.05 mm broad. Lateral inflores-
cence units: 1.7–(2.6)–3.5 mm long; 1.5–(2.45)–3.2 mm
broad. Attachment scar on lateral units: 0.67–0.86 mm in
diameter. Sterile scales: 0.15–0.40 mm in diameter to
0.3–0.5 mm 30:16–0.25 mm. Prophylls supporting lobe of
lateral units: 0.27 mm long; 0.82 mm broad. Prophylls sup-
porting flowers: 0.32–0.43 mm long; 0.23–0.41 mm broad.
Scars after detached flowers: 0.3–0.4 mm in diameter. Flower:
1.3–1.7 mm long; 0.55–0.90 mm broad. Outer tepals:
0.6–(0.8)–1.4 mm long; 0.3–(0.5)–0.6 mm broad. Inner
tepals: 1.4–(1.6)–1.9 mm long; 0.3–(0.6)–0.9 mm broad.
Stamen: ca. 1.2 mm long; breadth of filament: 0.15 mm.
Staminode-like appendages: 0.57–0.73 mm long; 0.16 mm
broad. Staminodes: 1.0 mm long; 0.15–0.20 mm broad.
Receptacle tube: 0.25 mm long; 0.25 mm broad.
Fig. 4 Mauldinia hirsuta sp. nov. A, Fragmented flower from holotype showing short cuspidate staminodes (st), hairy fruit wall (fw) with two
globular appendages (ga), and filament with broken anther (f) (1700a-51). B, Detail of Abefore opening showing cellular detail of lamina of inner
tepals (it) and filament of stamen (f) with staminal appendage (sa). C, Fragmented flower from same specimen as Bshowing filament (f) of anther
with two staminal appendages (sa). D,E, Immature fruits showing remnants of flower perianth (fp) at the base, abscission zone (az) at the base of
style, and remnants of style (s) (1700b-196, 1700b-189). All SEM. Scale ¼0:5 mm, except for B.
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Description and Remarks on the Species
Inflorescences. The material includes a few fragments of
inflorescence axes sometimes with lateral inflorescence units
attached, but the most common type of inflorescence frag-
ments are detached lateral units. These are often without
flowers, but some have flowers still attached.
The inflorescence is compound, consisting of an elongate
axis, 0.8–(1.5)–2.0 mm in diameter (fig. 1A–1C), bearing
many lateral inflorescence units in a spiral arrangement at
a spacing of ca. 0.8–1.0 mm. The lateral units are supported
by a broadly obtriangular and persistent bract, 0.4 mm long
and 0.05 mm broad, with a short decurrent keel-shaped base
(fig. 1B,1C,1G).
There are two types of scars along the inflorescence axis.
Larger scars, 0.7–(1.2)–2.0 mm long and 0.4–(0.6)–0.8 mm
broad, are ovate to narrowly ovate (lanceolate) in outline
and sometimes with a sinuous apex (fig. 1B,1C). These are
probably scars from fully developed lateral inflorescence
units. Smaller scars occur interspaced among the larger scars
(fig. 1B,1C). They are more or less rhomboidal in outline,
0.1–0.2 mm in diameter, and are here interpreted as scars
from undeveloped lateral units. The longest axis fragment
observed has six larger scars from detached, mature lateral
The lateral inflorescence units are 1.7–(2.6)–3.5 mm long
and 1.5–(2.5)–3.2 mm broad, flattened or concave, and
deeply bilobed and broadly obovate in outline, with an irreg-
ular, sometimes distinctly angular margin. They have a thick
and leathery appearance and an irregularly wrinkled surface
(fig. 1D–1F). Typically the two lobes overlap. The material
also includes many individual lobes that have separated from
the lateral units. The individual lobes are more or less ob-
triangular in outline, ca. 2.6 mm long and 1.9 mm broad,
with irregular lobed margins. The distal margins of the lobes
are often strongly incurved toward the axis, and lateral mar-
gins are also sometimes incurved (fig. 2A,2D). Each lobe is
Fig. 5 Mauldinia hirsuta sp. nov. A, Cuticle of inflorescence axis, inner side; note longitudinally elongated rows of polygonal cells (1700b-
242). B, Detail of Ashowing strongly cutinized hair bases. C, Cuticle of abaxial surface of lateral unit, inner surface; note thickness of cuticle,
rounded hair bases (hb) on outer surface, and imprints of thick-walled epidermal cells on inner surface (1700b-226). D, Detail of Cshowing thin-
walled cells surrounding paracytic elongated stoma with divided subsidiary cells (sc) elongated parallel to the pore and two polar cells (pc)
elongated parallel to the stoma axis; note numerous strongly cutinized hair bases (hb). All SEM.
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supported by a broadly triangular prophyll, 0.3 mm long and
0.8 mm broad (fig. 3A). At the base of detached lateral inflo-
rescence units, there is a distinct and rounded to ovate, some-
times bilobed attachment scar, 0.7–0.9 mm in diameter (fig.
1D; fig. 2B,2C,2E; fig. 3A).
Flowers are borne on the adaxial surface of the lateral in-
florescence units (fig. 1A,1E; fig. 2A–2D; fig. 3A). The num-
ber of flowers per lateral unit is five (fig. 2B), with one
flower in a median position, proximal to the bifurcation of
the unit, and two flowers on each of the lobes borne at
slightly different heights. The lowermost flowers apparently
develop before the uppermost (fig. 2C). Each flower is sub-
tended by a thin ovate prophyll, ca. 0.4 mm long and 0.2 mm
wide (fig. 2E). Scars from detached flowers, ca. 0.3–0.4 mm in
diameter, have been observed in some empty lateral units, but
most of the lateral units are without flowers or scars from
flowers. Whether these units were originally without flowers
or the flower scars were obscured is unclear.
Small membranous, glabrous, and shiny scales are scat-
tered on both adaxial and abaxial surfaces of the lobes.
Those of the abaxial surface and in a distal position on the
adaxial surface are typically 0.2–0.4 mm in diameter and
with an irregular rhomboidal to circular outline (figs. 1D,
3C). Those of the adaxial surface occurring in the position of
the flowers (fig. 2A) are narrowly triangular in outline, 0.3–
0.5 mm long and 0.2–0.3 mm broad, and resemble tepals.
Flowers. The flowers are usually strongly compressed
and difficult to study, but a few three-dimensionally pre-
served specimens with more details intact have also been
found. The perianth is often complete at the base, but other
floral organs are typically fragmentarily preserved or strongly
Flowers are actinomorphic, ca. 1.3–1.7 mm long and 0.6–
0.9 mm wide, bisexual, and sessile (fig. 2A; fig. 3A,3D,3E).
The receptacle is concave, forming a short and narrow cup,
ca. 0.3 mm long, bearing the floral organs on the rim of the
cup (figs. 2E,3D–3F). The flowers have a trimerous perianth
and androecium. The perianth consists of two whorls of
straight tepals (fig. 3A,3D–3F). The outer tepals are short
and ovate, 0.6-(0.8)-1.4 mm long and 0.3-(0.5)-0.6 mm
broad. The inner tepals are about one-third to twice as long
as the outer tepals, obovate to spathulate, sometimes folded
along an indistinct median keel, 1.4-(1.6)-1.9 mm long, and
0.3-(0.6)-0.8 mm broad. A single immature flower was ob-
served attached to an inflorescence lateral unit (fig. 2D,2F).
It is obovate with the inner tepals folded forming a rounded
Because of the compressed nature of the flowers, the struc-
ture of the androecium is poorly known. However, our ob-
servations indicate that it is similar to other species of
Mauldinia, with three whorls of fertile stamens followed by
a fourth, inner whorl of staminodes. Remnants of filaments
indicate that stamens extended above the inner tepals at an-
thesis. Also, in better-preserved specimens, the distinction of
stamens from staminodes is problematic because the anther
is only slightly differentiated from the filament. However, the
presence of stamens, ca. 1.2 mm long, with paired staminal
appendages has been documented (fig. 4B,4C). The append-
ages are flat, ca. 0.6–0.7 mm long and 0.2 mm broad, adnate
to the filament, and differentiated into a stalk bearing an an-
gular head. The innermost whorl of the androecium consists
of short and narrow staminodes, ca. 0.3 mm long and 0.2
mm broad, cuspidate in outline and dorsiventrally flattened
(fig. 4A).
The gynoecium is ovate (fig. 2C) and tapers apically into
a short and slender style. Ovary and style are separated by
a distinct joint.
Fruits. The material includes two immature fruits,
0:830:5 mm and 1:030:7 mm (fig. 4D,4E). The lower half
of the fruit is surrounded by a cupule composed of the
Fig. 6 Mauldinia hirsuta sp. nov. A,B, Cuticle of lateral bilobed
structure (abaxial side) showing irregularly arranged triangular to
rectangular and polygonal cells, numerous actinocytic hair bases (hb),
elongated paracytic stomata (ps) with divided subsidiary cells, and pa-
pillae (pl) (1700b-246). C, Cuticle of lateral bilobed structure (adaxial
side) showing numerous elongated actinocytic hair bases (hb) among
longitudinally elongated triangular to rectangular and polygonal cells
(1700b-246). All light microscope micrographs. Scale ¼10 mm.
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enlarged receptacle and remnants of the perianth. Apically,
the fruit tapers into the style. There is a distinct abscission
zone between the fruit and the style. All surfaces of fruit and
perianth are covered by stiff trichomes and trichome bases.
Epidermis features and cuticle. The cuticle of inflores-
cence axes and lateral inflorescence units is thick, ca. 0.01
mm (fig. 5C), while that of floral organs is thin.
The epidermal cells of all inflorescence and floral parts
studied are irregularly rectangular to triangular or polygonal
in shape with straight anticlinal walls. Those of the inflores-
cence axes are longitudinally arranged, 10–20 mm long and
4–12 mm wide (fig. 5A,5B).
The abaxial and adaxial epidermis of the lateral inflores-
cence units is distinct. Epidermal cells of the adaxial side are
10–(17)–24 36–(8)–10 mm and arranged in longitudinal rows
in the middle part while irregularly arranged at the apex.
Epidermal cells of the abaxial side are smaller than those on
the adaxial side and irregularly arranged (fig. 5C,5D). At
the base, they are triangular to rectangular in shape, 4–(12)–
18 32–(7.5)–12 mm; in the middle, they are more elongated,
8–(16)–22 34–(11)–13 mm, and arranged in indistinct longi-
tudinal rows, while at the apex cells are intermediate in size,
9–(12)–14 34–(8)–10 mm.
Epidermal cells of the tepals are rectangular to polygonal
in outline, longitudinally elongated measuring 28–(37)–
40 312–(18)–27 mm (outer tepals) and 14–(21)–31 38–(10)–
15 mm (inner tepals; fig. 3H). Epidermal cells over the me-
dian keel of the inner tepals are narrower, less than 10 mm
wide, and arranged in longitudinal rows.
Stomata were observed scattered on the abaxial surface of
the lateral inflorescence units (fig. 5D; fig. 6A,6B). They are
elliptical in shape and randomly orientated, thin-walled, and
paracytic. Subsidiary cells are often transversely and/or longi-
tudinally divided, and there are two polar cells orientated
with their long axis parallel to the long axis of the stoma.
Guard cells are characterized by lamellar thickening. Length
of stomatal complex and size of surrounding cells vary
together with size of epidermis cells. In the basal and
middle part, stomata are ca. 24 mm long, and at the apex,
they are ca. 18 mm long. The subsidiary cells measure 11 33–
(18 310)–36 310 mm. The width of guard cells is ca. 5–8 mm.
Elongate papillae occur on the epidermis of the lateral
units (fig. 6A). On the abaxial surface, they are present near
the distal margin, ca. 18 mm long and 7 mm broad. On the
adaxial surface, they are distributed near the base and distal
margins and are ca. 7 mm long.
A dense indumentum of ascending and unicellular simple
trichomes, up to 0.3 mm long and with actinocytic bases, oc-
curs on most surfaces of inflorescences (fig. 3I) and flowers.
The density of the indumentum varies considerably from
specimen to specimen, and on the axes and on inner tepals, it
may range from densely hairy to almost glabrous. Frequently
trichomes are abraded, but their strongly cutinized bases are
typically well preserved (figs. 1G,3B; fig. 5B–5D; fig. 6A
6C). Trichome bases are circular to elliptical or triangular in
shape. At the inflorescence axes, they vary from triangular,
ca. 5 mm in diameter, to elliptical, up to 13 mm long (fig.
1G). Trichome bases of the lateral inflorescence units are
generally circular and irregularly distributed (fig. 3B). Basally
on the abaxial side, trichomes are numerous with very small
trichome bases, ca. 2–3 mm in diameter. In the middle and
apical part, they are less numerous and bigger, ca. 6–7 mmin
diameter (fig. 5C,5D). On the adaxial side, trichomes are
more evenly distributed, with trichome bases circular, ca. 7–
11 mm in diameter, in the basal and apical part, while in the
middle part they are elliptical and longitudinally elongated
(8–16 mm long) (fig. 6C).
Both outer and inner tepals have irregularly scattered tri-
chomes (up to 0.2 mm long) and circular trichome bases (5–
13 mm in diameter) at the base of the abaxial surface (fig.
3D). On the inner tepals, trichomes also occur on the median
keel and the adaxial surface with trichome bases ca. 6–13
mm in diameter (fig. 3E,3F), while the adaxial surface of the
outer tepals are almost smooth. Trichomes on stamens are
ca. 0.1 mm long and occur scattered on the filament (fig. 4B,
4C), while the staminal appendages and staminodes are gla-
brous (fig. 4A,4C). Trichomes are up to 0.3 mm long and
densely spaced over the surface of the ovary and fruits (fig.
Other Reproductive Structures of
Possible Lauraceous Affinity
In addition to the many reproductive fragments that can
unambiguously be assigned to Mauldinia, the Sarbay fossils
also include a few dispersed reproductive organs (a single sta-
men adnate to a tepal, a staminode, a flower fragment with
gynoecium preserved, and a fruit) that probably also belong
to the Mauldinia plant but cannot be unambiguously as-
signed to the genus because of lack of diagnostic features.
However, because they provide additional information on
lauraceous reproductive organs from the Sarbay assemblage,
a short description of these organs is given in the separate
sections below.
Tepal with adnate stamen (1700b-188; fig. 7A). This
specimen is a fragment of a tepal, ca. 1.6 mm long, with
slightly incurved margins and with a stamen adnate to the
base of tepal. The stamen is ca. 1.5 mm long and differenti-
ated into a slender filament, 1 mm long and 0.07 mm broad,
and an ovate anther, 0.5 mm long and 0.2 mm broad. Anther
dehiscence is introrse by two narrow, elliptical valves. The
tepal and filament are glabrous, while the anther is densely
hairy. The stamen is similar to those of the third androecial
whorls in Mauldinia flowers from other areas (borne oppo-
site to the inner tepals and with introrse dehiscence by two
narrow valves). It is most likely that the tepal and stamen are
detached from a Mauldinia flower, but because of incomplete
knowledge of the stamens in Mauldinia hirsuta, we are reluc-
tant to assign this specimen to this species.
Staminode (N1700b-190, fig. 7B). The fossil is ca. 1 mm
long with a distinctly flattened stalk, ca. 0.3 mm wide, and
with a triangular expanded apex, ca. 0.4 mm wide, very simi-
lar to staminodes of Mauldinia described from other areas.
The supposed staminode is adnate to a fragmented organ, per-
haps a fruit wall, with a strongly cutinized and hairy surface.
Flower fragment with gynoecium (1700b-122; fig.
7C). The specimen is a fragment of a charcoalified flower
with a concave receptacle forming a short and narrow floral
tube bearing remnants of a tepal, filament, and gynoecium.
The tepal is straight, ca. 0.9 mm long, with incurved margins
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and a hairy lamina. The filament is broken apically, but the
length indicates that the stamens extended above the tepals
at anthesis. The gynoecium is ovate, ca. 0.3 mm long, and
tapers apically into a short and slender style, ca. 0.2 mm
long, with a sessile stigma, ca. 0.05 mm in diameter. Ovary
and style are separated by a distinct joint. Small globular ap-
pendages, 0.04 mm in diameter, occur at the base of the
style. All flower organs, except for the style, are covered by
a dense indumentum.
The size of the tepal, presence of a distinct joint between
style and ovary, globular appendages at the base of the ovary,
and a dense indumentum indicate that the specimen belongs to
M. hirsuta. However, information about the androecium struc-
ture is too incomplete to make more detailed comparison.
Fig. 7 Other reproductive structures of Lauraceous aff. A, Fragment of flower showing tepal (t) and bisporangiate stamen (st) with valvate
dehiscence (1700b-188). B, Staminode attached to fragment of fruit wall (fw) (1700b-190). C, Fragment of hairy flower showing tepal, filament
and hairy carpel with globular appendage (ga), and a glabrous style (1700b-122). D, Circular fruit with thick cuticle, remnants of flower perianth
(fp) at the base, and elongated attachment scar (1700b-197). E, Detail of Dshowing thick cuticle with scattered hair bases (hb). All SEM.
Scale ¼1 mm, except for E.
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Fruit (N1700b-197; fig. 7D–E). The fruit is circular in
outline and flattened, ca. 2.2 mm in diameter, and with rem-
nants of a perianth at the base. The attachment scar is broad
and elongated. The cuticle is thick, strongly wrinkled, and
with numerous trichome bases. Although the fruit differs in
its larger size from the known immature fruits of other Maul-
dinia species, the general shape, the presence of floral rem-
nants at the base, and the cuticle features indicate that this
fossil may also belong to Mauldinia.
Although not all floral features are known for the Sarbay
fossils, the flattened bilobed lateral inflorescence units bear-
ing densely spaced, trimerous flowers clearly place the fossils
in the extinct genus Mauldinia. This genus was based on
a single species, Mauldinia mirabilis, described from the Cen-
omanian Mauldin Mountain flora of Maryland, United States
(Drinnan et al. 1990). Subsequently another species, Mauldi-
nia bohemica, was described from the Cenomanian Peruc-
Korycany Formation of the Czech Republic (Eklund and
ˇek 1998), and an additional unnamed species was re-
corded from the Late Santonian Allon Flora of central Geor-
gia, United States (Mauldinia sp.; Herendeen et al. 1999).
The floral features are well preserved in many of the North
American and Bohemian fossils and unambiguously place the
fossils in the family Lauraceae (Drinnan et al. 1990; Eklund
and Kvac
ˇek 1998). In addition to the shared general features,
the Sarbay Mauldinia exhibits several unique features that
distinguish it from previously described species (table 1), and
the material has therefore been assigned to a new species,
Mauldinia hirsuta.
The unique broad and flattened lobes of the lateral inflo-
rescence units of Mauldinia have been interpreted as derived
from a simple racemose inflorescences by triplication of flow-
ers followed by condensation, fusion, and planation of all
branches of the lateral inflorescence units (Eklund and Kvac
1998). In M. hirsuta, the lobes have weakly developed inden-
tations and provide no further information on the derivation
of the lobes.
The main diagnostic features for species separation within
Mauldinia are the shape and size of lateral units and the type
of sterile scales occurring on the units (table 1). The lateral
units of M. hirsuta have very little reminiscence of the
branching system. The individual lobes are very short and
broad with a rounded or angular margin and with only
weakly pronounced scales or bracts/prophylls along the mar-
gin of the lobes. In this respect, the Sarbay fossils are most
closely similar to the unnamed Mauldinia sp. from the Allon
flora. Mauldinia mirabilis has somewhat longer lobes, and
typically the apical part of the lobes is distinctly divided, in-
dicating a suppressed branching system. Mauldinia bohemica
has even longer lobes and very distinct bracts/prophylls and
scales arranged distichously along the margins of the lobes.
Mauldinia hirsuta is also most comparable to the North
American M. mirabilis in having five flowers per lateral inflo-
rescence unit in contrast to M. bohemica, which typically has
seven flowers per unit. The flowers of Mauldinia are small,
sessile, bisexual, actinomorphic, and trimerous. The perianth
consists of two whorls of three tepals. In all species, tepals of
the outer whorl are about half the length of those of the in-
ner whorl. The shape of the tepals, however, varies from spe-
cies to species, with the ratio width of the outer to inner
tepal being ca. 1 : 1 in the Sarbay flowers and M. mirabilis
while almost 1 : 2 in M. bohemica. Tepals and details of
flower structure were not described for the Allon Mauldinia.
The androecium consists of four alternating whorls. The
outer three whorls consist of fertile stamens, while the inner
whorl consists of staminodes. The stamen is differentiated in-
to a filament and an anther, which dehisces by two apically
hinged valves. Associated with the anthers of the third whorl
are paired, clavato-saggitate appendages. Details of stamen
structure are unknown for M. hirsuta, but our observations
indicate that the general androecium organization is similar
to other species of Mauldinia. The glabrous staminal append-
ages and staminodes of M. hirsuta are similar to those of
M. bohemica but differ from the hairy androecium of M.
At the cellular level, M. hirsuta is comparable to both M.
mirabilis and M. bohemica in shape and orientation of epi-
dermal cells, and all species have paracytic stomata. Mauldi-
nia bohemica differs from M. hirsuta and M. mirabilis by the
distinct perforation of the scales of the lateral units, probably
indicating the presence of densely spaced ethereal oil cells in
the Bohemian material.
All Mauldinia species currently known have an indumen-
tum of unicellular trichomes. Mauldinia hirsuta is, however,
distinguished from both M. mirabilis and M. bohemica in
having a much denser indumentum. Only Mauldinia sp. from
the Allon flora apparently has a similarly dense indumentum.
The density of the indumentum varies considerably from
specimen to specimen in the Sarbay material; this was also
noted for other species of Mauldinia (Drinnan et al. 1990;
Eklund and Kvac
ˇek 1998).
Another compound inflorescence with two-parted lateral in-
florescence units thought to be related to the Lauraceae was
described by Eklund (2000) from the Santonian/Campanian
Neuse River locality of North Carolina, United States, as
taxon A. It resembles Mauldinia in having sessile flowers and
bisporangiate anthers but differs in the lack of staminal ap-
pendages and staminodes and in its umbellate inflorescence
with two large free bracts/prophylls supporting the flowers.
Other lauraceous reproductive organs described from the
Late Cretaceous are isolated flowers similar to those of Maul-
dinia in having shorter outer and longer inner tepals. Among
these are three taxa with well-preserved androecial features:
Neusenia tetrasporangiata Eklund (2000), described from the
Santonian-Campanian Neuse River locality, North Carolina,
United States, and ‘‘hypogynous flower type 2,’ and Lauran-
thus futabensis, described by Takahashi et al. (1999, 2001)
from Lower Coniacian sediments of the Kamikitaba Assem-
blage, northeastern Japan. These are all distinguished from
Mauldinia in having tetrasporangiate anthers. Two other
lauraceous flowers, ‘‘taxon B’ (Eklund 2000) from the
Santonian-Campanian Neuse River locality, North Carolina,
United States, and Perseanthus crossmanensis Herendeen
et al. (1994) from the Turonian of New Jersey, United States,
have staminal appendages and perianth arrangement similar
to Mauldinia, but incomplete preservation and the lack of in-
florescence characters impede a more detailed comparison.
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The lauraceous fossils from Kazakhstan clearly belong to
the extinct genus Mauldinia, which is characterized by the
unique bilobed lateral inflorescence units. With this new rec-
ord, the geographic range of the genus is expanded from
North America over Europe to Asia. The genus was appar-
ently a conspicuous element of the Northern Hemisphere
mid-Palaeolatitude floras during the early to middle part of
the Late Cretaceous. It is interesting to note that the new spe-
cies, Mauldinia hirsuta, is most similar to the younger spe-
cies, Mauldinia sp. from the Late Santonian Allon flora of
Georgia, United States, and most distinct from the Bohemian
species Mauldinia bohemica, which is of approximately the
same age as the Sarbay fossils and closer geographically.
We thank Sergej G. Zhilin, St. Petersburg, for providing
the sediment material and information on the locality;
Table 1
Descriptive Characters of Mauldinia Species
Species Mauldinia hirsuta Mauldinia sp. Mauldinia bohemica Mauldinia mirabilis
Lateral unit Wide-obovate lateral
unit with wide-
obovate, angulate
lobes; indistinct
branching system
of lateral unit
Wide-obovate lateral
unit with wide-
obovate, angulate
lobes; indistinct
branching system
of lateral unit
Ovate lateral unit with
lobes; distinct
branching system
of lateral unit
Ovate lateral unit with
lobes; distinct
branching system of
lateral unit
Scales on lateral unit Nonperforated No scales Perforated No scales
Axis width (mm) 0.8–2.0 Not known 1.3–(1.7)–2.4 0.8
Scar after partial
inflorescence Encircled ovate with
sinuous apex
Not known Encircled elliptical Encircled elliptical
Additional scars Rhomboidal, regular Not known Not known Elliptical, rare
Lateral unit size (mm) 1.7–3.5 length, 1.5–3.2
5–6 length, 6–7 breadth 1.9–(5.5)–8.0 length,
1.9–(6.2)–8.8 breadth
2.0–3.5 length, 2.0–
(1.8)–3.5 breadth
Lobe size (mm) 1.7–3.5 length, 0.7–1.6
5–6 length, 3–4 breadth 5.5 length, 3.1 breadth 2.0–3.5 length, 1.8
No. flowers 5 Not known 3–5–(7)–9 5
Flower size (mm) 1.3–1.7 length, 0.6–0.9
2.3 length 1.5–2.6 length, 0.6–1.4
1.5–3.6 length, 1.0–2.0
Floral tube length (mm) 0.20–0.25, abruptly
Not known 0.09–(0.19)–0.29, not
0.50, not separated
Tepal shape Outer: obovate; inner:
indistinctly keeled,
often folded
Outer: triangular; inner:
indistinctly keeled,
often folded
Outer: triangular; inner:
distinctly keeled and
Outer: triangular; inner:
distinctly keeled and
Stamen whorls Not known Not known 3 3
Staminal appendages Stalk with flat angular
head, 0.6–0.7 mm
Not known Stalk with glandular
head, 0.8–1.2 mm
Stalk with flat clavate-
sagittate head, 0.7–
1.5 mm long
Staminodes whorls Cuspidate, 0.3 mm long Not known Narrow, tongue shaped,
1.4 mm long
Narrow with
rhomboidal head, up
to 1.0 mm long
Stigma Not known Not known Not known Expanded, flattened
Indumentum Dense on axis, lateral
unit, inner tepals,
filaments, carpel;
absent on staminodes
and staminal
Dense on adaxial
surface of lateral
unit (?)
Scattered on lateral unit,
carpel, pistil,
occasionally present
on filaments and
Dense on filaments and
staminal appendages,
present on inner
surface of tepals,
occasionally scattered
on staminodes; absent
on lateral unit and
ovary surface
Stomata on lateral unit Paracytic with
transversely or
longitudinally divided
subsidiary cells; rare,
restricted to the distal
area on the abaxial
surface of lateral unit
Not known Paracytic/paracytic with
one transversely
divided subsidiary
cell; restricted to the
distal area on the
abaxial surface
Paracytic/paracytic with
one longitudinally
divided subsidiary
cell; on both surfaces
of lateral unit
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thanks are also due to David Cantrill and Thomas Denk,
Stockholm, for the help with cuticle preparation, assistance
with digital images, and valuable discussions. Yvonne Ar-
remo, Stockholm, is thanked for help with SEM and Alexej
Hvalj for help with material. This work was supported by
the HighLat program (Swedish Museum of Natural History,
project HPRI-CT-2001-00125) to S. Frumin, a grant from
the Swedish Natural Science Research Foundation to E. M.
Friis, and support from Helge Ax:son Johnsons Stiftelse to
H. Eklund.
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... In Europe Eklund and Kvaèek (1998) described inflorescence fossils of Lauraceae from the Cenomanian of Bohemia, Czech Republic while Viehofen et al. (2008), reported the occurrence of the species Mauldinia angustiloba from the Cretaceous of Germany. Such fossils have also been collected from the Cretaceous of East Asia (Ferumin et al. 2004). Ferumin et al. (2004) also described Mauldinia hirsuta from the Late Cretaceous of Kazakhstan, while Takahashi et al. (1999Takahashi et al. ( , 2001 reported inflorescence fossils of Lauraceae from similar-aged rocks in Japan. ...
... Such fossils have also been collected from the Cretaceous of East Asia (Ferumin et al. 2004). Ferumin et al. (2004) also described Mauldinia hirsuta from the Late Cretaceous of Kazakhstan, while Takahashi et al. (1999Takahashi et al. ( , 2001 reported inflorescence fossils of Lauraceae from similar-aged rocks in Japan. On the basis of above fossil record it is clear that Lauraceae had widely distributed in Laurasia, Gondwana, and in the tropical zone around the Tethys ocean. ...
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In the present investigation, a morpho-taxonomic study has been carried out on recently recovered plant macrofossils from the Siwalik Group (Middle Miocene) sediments of Sarkaghat, Mandi District, Himachal Pradesh, India. This has revealed the occurrence of two new fossil leaves belonging to the extant taxa, Artocarpus heterophyllus Lam. and Phoebe opaca Blume. of the tropical dicotyledonous families, Artocarpaceae and Lauraceae respectively. As the nearest living relatives of the macrofossils reported here, are restricted to the southern part of India and southeastern Asia (Sri Lanka, Malaya, Indonesia, and Philippines), it is plausible that tropical forests under moist conditions were prevalent during the Upper Miocene times in this region. The recovered data also indicates prevalence of tropical climatic conditions during the Miocene in the Sarkaghat and nearby area.
... Currently, Cupressaceae still has a worldwide distribution and can be found in a wide variety of habitats, including the West Indies (Judd et al., 1999). Similarly, Lauraceae had a high diversity and wide distribution range by the Late Cretaceous (Poole et al., 2000;Frumin et al., 2004) comprising Gondwana (Poole et al., 2000), North America (Little et al., 2009), and Asia (Frumin et al., 2004). In the present, the family is predominantly found in tropical-subtropical latitudes of southeast Asia and northern South America (Judd et al., 1999;Chanderbali et al., 2001). ...
... Currently, Cupressaceae still has a worldwide distribution and can be found in a wide variety of habitats, including the West Indies (Judd et al., 1999). Similarly, Lauraceae had a high diversity and wide distribution range by the Late Cretaceous (Poole et al., 2000;Frumin et al., 2004) comprising Gondwana (Poole et al., 2000), North America (Little et al., 2009), and Asia (Frumin et al., 2004). In the present, the family is predominantly found in tropical-subtropical latitudes of southeast Asia and northern South America (Judd et al., 1999;Chanderbali et al., 2001). ...
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The Caribbean islands are one of the most important hotspots of endemism and biodiversity globally, and the scenario of unique examples of biological radiations. Although our knowledge of the current and recently extinct diversity in the area is strong, the origin and evolution of most groups in the region remain obscure because of the absence of fossils from deep time periods. The existence of temporal islands on the Caribbean plate can be traced back to the late Mesozoic, but little evidence of the paleo-communities that once inhabited the archipelago and their relationship with the older lineages in the region has been discovered. Simultaneously, the relationship of the early Caribbean archipelago with the Late Cretaceous–early Paleogene biotic interchange between the Americas has remained unsolved. Here we describe the first evidence of a Late Cretaceous terrestrial community in the region based on several remains recovered at three upper Campanian–lower Maastrichtian localities in Central Cuba. The fossil assemblage includes four specimens referable to a midsize pterosaur on the base of morphological and paleohistological characters, as well as seeds and casts of leafy shoots of plants of the families Cupressaceae and Lauraceae. Fossils fruits of a new taxon closely related to Chlorocardium are of particular interest because they correspond to the first direct evidence of the role played by the Caribbean seaway and islands in the First American Biotic Interchange.
... Republic (Eklund & Kvaček, 1998), M. hirsuta from the Cenomanian-Turonian of Kazakhstan (Frumin et al., 2004), and M. angustiloba from the middle Cretaceous of Germany (Viehofen et al., 2008). A further, unassigned Mauldinia was reported from the Santonian of Georgia, USA (Herendeen et al., 1999). ...
... Similarly to Mauldinia sp. from Gard, some Mauldinia species were also reported from marginal-littoral areas. The paleoenvironments of M. bohemica and M. hirsuta were interpreted as floodplains with large, tidally-influenced rivers and lacustrine to fluvial with lagoon inputs, respectively (Eklund & Kvaček, 1998;Frumin et al., 2004). Mauldinia-like specimens from the Albian-Cenomanian of the Dakota Formation, Kansas (Retallack & Dilcher, 1981;Kovach & Dilcher, 1988) have been recovered from tidal to lagoonal environments. ...
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La transition Crétacé inférieur-Crétacé supérieur (environ 100 millions d’années) marque une période cruciale pour l’évolution de certaines plantes à graines. Elle correspond à la rapide radiation des plantes à fleurs et à la réorganisation écologique des Angiospermes et des conifères. Cependant, notre connaissance des plantes à graines durant le Crétacé moyen et en Laurasie reste encore très partielle. Pendant la dernière décennie, la microtomographie synchrotron s’est avérée une technique d’imagerie très performante pour l’étude des structures internes de divers fossiles. Jusqu’à maintenant, les applications de cette technique n’ont été que très limitées en paléobotanique.C’est dans ce contexte que cette thèse s’est intéressée : (1) au développement de nouvelles approches d’imagerie par tomographie, utiles en paléobotanique, et sur différents types de préservation ; (2) à l’utilisation de ces nouvelles approches pour compléter notre connaissance de la morpho-anatomie, la systématique, la diversité, et l’écologie des plantes à graines laurasiatiques du Crétacé moyen. Ainsi, un peu plus d’une centaine de spécimens (e.g. inflorescences, fleurs, fruits, et grains de pollen d’Angiospermes ; cônes and axes feuillés de conifères) provenant de gisements paléontologiques français ont été étudiés par tomographie synchrotron.Pour la première fois, de nombreuses fleurs très diversifiées sont décrites dans les dépôts cénomaniens du Gard (Sud-Est de la France). Elles montrent une remarquable préservation en trois dimensions. Dans certains cas, toutes les unités florales du périanthe, de l’androcée, et du gynécée sont préservées. Un nouveau protocole combinant microtomographie par contraste de phase de propagation et nano-holotomographie a été développé pour étudier des petits spécimens isolés tels que des fleurs préservées sous forme de fusain ou de lignite (taille de voxel proche de 50 nm). Les données tomographiques permettent de décrire les fleurs de la morphologie générale à la paroi des grains de pollen, in situ, dans les étamines. Certaines de ces fleurs montrent des structures florales inédites. Elles sont majoritairement attribuées à des Lauraceae.De plus, des nodules siliceux, opaques, denses, et contenant des inclusions végétales sont ici signalés dans les dépôts cénomaniens de Charente-Maritime (Ouest de la France). Ils contiennent d’abondants restes de conifères (e.g. Brachyphyllum, Frenelopsis, Geinitzia et Glenrosa). Ils sont préservés sous forme de perminéralisation siliceuse et en trois dimensions. L’essentiel des tissus est préservé. Ce travail propose un protocole tomographique multi-échelles, haute résolution et haute énergie, utile à l’étude d’inclusions végétales contenues à l’intérieur de nodules rocheux de grande dimension. Le conifère Glenrosa est décrit pour la première fois, de la morphologie générale des structures végétatives et reproductives à l’histologie. Nous discutons la systématique et la paléoécophysiologie de ce genre. Les caractéristiques xéromorphes de Glenrosa suggèrent que ce conifère était probablement adapté pour tolérer des conditions difficiles induites par les milieux côtiers. Ce travail soutient que pendant le Cénomanien, les environnements les plus ouverts sur la mer étaient dominés par les conifères alors que les milieux littoraux plus internes et protégés montraient quant à eux des flores à dominante angiospermienne.Pour conclure, cette recherche doctorale contribue considérablement à notre connaissance des Angiospermes et des conifères de la Laurasie, pendant le Crétacé moyen. De plus, les nouvelles approches tomographiques proposées dans ce travail pourront être reproductibles pour l’étude d’autres flores, ou d’autres spécimens paléontologiques, quelle que soit leur origine géographique et stratigraphique.
... Lauraceae/?Mauldinia(Fruit) (Eklund and KvaČek (1998) (2007) like Olmosoxylon and Laurinoxylon (Ulminium), but they cannot be included in a particular extant taxon (e.g., Stern, 1954;Süss, 1958;Page, 1967;Estrada-Ruiz et al., 2010b; Table 1). Reviews on the fossil record of the family can be found in Kostermans (1957), Chandler (1964Chandler ( , 1978, Mai (1971Mai ( , 1999Mai ( , 2001, Drinnan et al. (1990), Herendeen (1991), Kvaček (1992), Rohwer (1993), Herendeen et al. (1999), Crane et al. (1994), Manchester (1994), Pingen et al. (1994), Mickle (1996), Eklund and KvaČek (1998), Frumin et al. (2004). The described plants in northern Mexico suggest this family was an important component of the paleovegetation in transitional or near coast areas (Wheeler and Lehman, 2000;Estrada-Ruiz et al., 2008), and that the family reached important morphological/anatomical diversity early in their lineage history, but it is necessary to understand how this diversity defines taxa through whole plant reconstructions (Table 1). ...
A new wood type for the Baja California Cretaceous adds to the plant diversity so far known for the area where gymnosperms seem to be dominant. It was collected near El Rosario, Baja California, from rocks of the Rosario Formation, in a sedimentary sequence that comprises ca. 1200 m of non-marine to deep marine sediments from Upper Campanian to Lower Danian age. The wood is characterized by having semiring porous growth rings, predominantly radial multiples of 2–7 with occasional clusters and some solitary vessels, simple perforation plates, alternate intervascular pits, oval to large elliptical vessel element-ray pits with reduced borders, septate thin-walled fibers, 1–4 seriate heterocellular rays, scares paratracheal, vasicentric and marginal parenchyma and oil cells associated with ray parenchyma. All these characters are found in Lauraceae, however, none of the extant taxa of the family have all these characters and even among fossil woods the characters in the Baja California material are better described only among the diverse Laurinoxylon, but vessel grouping, growth ring type, absence of marginal parenchyma, and slightly thicker rays suggest the presence of a new taxon, Rosarioxylon bajacaliforniensis Cevallos-Ferriz, Catharina & Kneller. By the end of the Cretaceous the family formed part of the plant community that represents a western extension of vegetation types more completely described from areas in the margins of the southern limits of the Western Interior Sea. The new taxon is proposed to highlight anatomical differences and geographic isolation from similar taxa and further suggests a large distribution of Lauraceae in what appears to be conifer dominated communities.
... The bulk of the angiosperm mesofossil record has been recovered from Cretaceous sediments collected in eastern North America, central Portugal, and southern Sweden (Atkinson et al., 2018;Friis and Pedersen, 2012;Friis et al., 2013aFriis et al., , b, 2015Friis et al., , 2017aFriis et al., , b, 2018aFriis et al., , b, c, d, e, 2019Herendeen et al., 2017;Martínez et al., 2016;Mendes and Friis, 2018;Mendes et al., 2014;Schönenberger et al., 2012). In contrast, the mesofossil record from Asia is much less extensive, and so far angiosperm mesofossils have been recovered only from Kazakhstan and Japan Friis, 1996, 1999;Frumin et al., 2004;Takahashi et al., 1999aTakahashi et al., , b, 2001aTakahashi et al., , b, 2002Takahashi et al., , 2007Takahashi et al., , 2008aTakahashi et al., , b, 2014Takahashi et al., , 2017. ...
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A preliminary description is provided of a new assemblage of small, three-dimensional and charcoalified mesofossils from the Tamagawa Formation (late Turonian–middle Santonian; Upper Cretaceous) of the Kuji Group in northeastern Japan. The new mesofossils yield excellent structural details and include wellpreserved circinate shoots of ferns together with conifer leafy-shoots, seeds and probable pollen cones, and variety of angiosperm fruits and seeds, including fruits of Cornales and seeds of Nymphaeales. The new mesofossil assemblage is complementary to the previously published macrofossil flora from the Kuji Group.
... We are not aware of any extant Lauraceae with pronounced monochasial parts in the cymes of the thyrsoids. Additional Mid-to Late Cretaceous Mauldinia species with apparently cymose lateral inflorescence units were described from other localities in North America (Herendeen et al. 1999;Friis et al. 2006), Europe (Viehofen et al. 2008), and Kazakhstan (Frumin et al. 2004). Pragocladus (Cenomanian) also has compound inflorescences with an elongate main axis and several lateral axes with two to four flowers of unknown ramification pattern (Kvacek and Eklund 2003). ...
Premise of research. This is the first comparative study of inflorescence morphology through all seven families of the order Laurales (Atherospermataceae, Calycanthaceae, Gomortegaceae, Hernandiaceae, Lauraceae, Monimiaceae, and Siparunaceae) and the larger subclades of these families. Methodology. We studied 89 species of 39 genera from herbarium specimens and partly from liquid-fixed material, focusing on the branching patterns in the reproductive region. In addition, we used the information from the literature. Pivotal results. There are recurrent branching patterns. Botryoids, thyrsoids, and compound botryoids and thyrsoids are the most common forms. Panicles, racemes, and thyrses are rare. Panicles and racemes occur in some highly nested Lauraceae and thyrses in Hernandiaceae. Thus, the presence of thyrso-paniculate inflorescences is not characteristic for Laurales, in contrast to the statement by Weberling. Conclusions. An evolutionary interpretation is still difficult because the existing molecular phylogenetic analyses are not fine grained enough and also because the previous phylogenetic results are not robust enough to make firm conclusions within the order. However, the present structural results show that there are trends of occurrence of certain patterns in families or subclades within families, and these may be useful in a morphological matrix of magnoliids (see work by Doyle and Endress for basal angiosperms).
... Another early lauralean reproductive structure Mauldinia, based on dispersed flowers and fragments of inflorescences, was widespread in the Cenomanian floras of the Northern Hemisphere (Drinnan et al., 1990;Eklund and Kva cek, 1998). Mauldinia hirsuta from the CenomanianeTuronian of Kazakhstan (Frumin et al., 2004) is the oldest unequivocal record of Laurales in Asia and comes from the same deposits as "Lindera" jarmolenkoi leaves. ...
Diversity of palmately lobed leaves of angiosperms of the early–middle Albian floras of the Kolyma River Basin, the Omsukchan Coal Basin, Khabarovsk and Primorye Regions was studied. Leaf fossils, previously compared with those of genera Aralia, Sassafras and Lindera, now reassigned to the fossil genus Araliaephyllum. Four new combinations and one new species are published: A. kolymense (Kryshtofovich) Golovneva, comb. nov., A. luciferum (Kryshtofovich) Golovneva, comb. nov., A. ussuriense (Krassilov) Golovneva, comb. nov., A. ievlevii (Samylina) Golovneva, comb. nov., and A. popovii Golovneva, sp. nov. The type material has been restudied in detail, and lectotypes have been selected to all newly typified species. These species share many lauralean morphological and venation features. They represent the most likely early members of this group. This relationship is based on detailed study of the leaf architecture and comparison with other fossils with studied epidermal features. This study expands our knowledge of radiation and biogeography patterns of early angiosperms in northeastern Asia.
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Premise: Significant paleobotanical discoveries in recent decades have considerably improved our understanding of the early evolution of angiosperms and their flowers. However, our ability to test the systematic placement of fossil flowers on the basis of phylogenetic analyses has remained limited, mainly due to the lack of an adequate, angiosperm-wide morphological data set for extant taxa. Earlier attempts to place fossil flowers phylogenetically were, therefore, forced to make prior qualitative assessments of the potential systematic position of fossils and to restrict phylogenetic analyses to selected angiosperm subgroups. Methods: We conduct angiosperm-wide molecular backbone analyses of 10 fossil flower taxa selected from the Cretaceous record. Our analyses make use of a floral trait data set built within the framework of the eFLOWER initiative. We provide an updated version of this data set containing data for 28 floral and two pollen traits for 792 extant species representing 372 angiosperm families. Results: We find that some fossils are placed congruently with earlier hypotheses while others are found in positions that had not been suggested previously. A few take up equivocal positions, including the stem branches of large clades. Conclusions: Our study provides an objective approach to test for the phylogenetic position of fossil flowers across angiosperms. Such analyses may provide a complementary tool for paleobotanical studies, allowing for a more comprehensive understanding of fossil phylogenetic relationships in angiosperms. Ongoing work focused on extending the sampling of extant taxa and the number of floral traits will further improve the applicability and accuracy of our approach.
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The Winton Formation is increasingly recognised as an important source of information about the Cretaceous of Australia, and, more broadly, the palaeobiogeographic history of eastern Gondwana. With more precise dating and stratigraphic controls starting to provide temporal context to the geological and palaeontological understanding of this formation, it is timely to reassess the palaeoenvironment in which it was deposited. This new understanding helps to further differentiate the upper, most-studied portion of the formation (Cenomanian–Turonian) from the lower portions (Albian–Cenomanian), allowing a coherent picture of the ecosystem to emerge. Temperatures during the deposition of the Upper Cretaceous portion of the Winton Formation were warm, with high, seasonal rainfall, but not as extreme as the modern monsoon. The landscape was heterogeneous, a freshwater alluvial plain bestrode by low energy, meandering rivers, minor lakes and mires. Infrequent, scouring flood events were part of a multi-year cycle of drier and wetter years. The heavily vegetated flood plains supported abundant large herbivores. This was the final infilling of the great Eromanga Basin.
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The Cenomanian-Turonian Shet-Irgiz Formation of clay and sand is present in the iron quarries (Sokolovka and Sarbay). Among numerous plant remains are undoubted taxodiaceous seeds, Alapaja Dorof., and, possibly, Taxodiastrum Dorof. and Quasisequoia Friis. -Authors
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Patterns and trends in angiosperm evolution can only be inferred using a well-resolved and well-supported phylogenetic tree, and many inferences of character evolution across the angiosperms rely on reconstructions at or near the base of the tree. For this reason, inferences of phylogeny and evolution in basal angiosperms have implications for understanding evolution in the angiosperms as a whole. Recent studies have concluded that the first three branches of extant angiosperms are Amborella, Nymphaeales, and a clade of Austrobaileya, Trimenia, and Illiciales; relationships among basal angiosperms following these early diverging branches are less clear. In this article, we (1) review the evidence for these basal relationships inferred from sequence data for three genes, (2) address the relationships among the clades of noneudicot angiosperms, including the monocots, based on both this large analysis of the angiosperms and new, more focused analyses of basal angiosperms alone, and (3) consider the implications of these phylogenetic relationships for the evolution of floral traits, particularly of the perianth. Using the best tree currently available and data on floral characters taken from the literature, we hypothesize that the early angiosperm flower had spiral phyllotaxis, with a small but indeterminate number of undifferentiated perianth organs. From this simple flower, a whorled floral arrangement, with a fixed number of sepals and petals, evolved quite early in angiosperm evolution. Reversions to spiral phyllotaxis, changes in merosity, and losses of floral parts occurred several times in the early evolutionary history of the angiosperms.
A preliminary conspectus of the fossil flora from the Allon locality, in Crawford County central Georgia, documents the presence of at least 63 distinct plant organs, based on macrofossils and mesofossils, but excluding dispersed pollen and spores. The fossils are preserved in a clay lens within the Buffalo Creek Member of the Gaillard Formation, and are of late Santonian (Late Cretaceous) age. In general, macrofossils are sparse at this site, but abundant mesofossils have been isolated from bulk samples by sieving. The flora includes sporophytes and gametophytes of fossil mosses (e.g., Eopolytrichum antiquum, Campylopodium allonense), ferns (e.g., cf. Boodlepteris), and conifers, but is dominated by the diverse and abundant remains of angiosperms. Angiosperms in the flora include cf. Detrusandra (Magnoliales), Mauldinia sp. (Lauraceae), Allonia decandra (Hamamelidaceae), and Parasaurauia allonensis (Actinidiaceae). Especially abundant are flowers and cupules of two species of Fagaceae sensu late (Protofagacea allonensis, Antiquacupula sulcata). The flora also includes fossil flowers of Caryanthus sp. (Juglandales/Myricales) and Bedellia pusilla (cf. Betulaceae), which comprise the first record of Normapolles-producing flowers from North America. The structural and systematic diversity of angiosperms in the Allon flora is comparable: to that at other Turonian-Campanian sites in eastern North America and Europe. Together, these fossil floras indicate that angiosperms, and especially eudicots, were already diverse at this relatively early stage in angiosperm evolution. The source vegetation represented by the Allon fossil assemblage was dominated by angiosperms, probably with taxodiaceous conifers also common. preservation of most of the mesofossils as charcoal indicates that fire may have been an important factor contributing to frequent disturbance of the source plant community.
Synchronous first occurrences of pollen and leaves indicate that angiosperms entered the Northern Rocky Mountain (NRM) region during the Middle Albian, approximately 8 Ma later than such shifts in floras from southern Laurasia. The earliest angiosperm pollen and leaf flora corresponds to the Potomac sub-Zone IIB in that it exhibits a pretricolporate palynoflora and a comparable grade of evolution based on leaf rank. The Albian megaflora is characterized by abundant leaves of the Platanophyll, Protophyll, and Pentalobaphyll morphotypes. Sapindophylls are common and diverse. The flora contains early North American occurrences of the widespread Upper Cretaceous leaf form genera Trochodendroides and Cinnamomoides. Pentalobaphylls (Araliaephylls) appear early and assume a more important position in the lowland vegetation of the NRM region than in other regions. Magnoliidae, Hamamelididae, and Rosidae are represented in the Albian flora. The Albian megaflora is a regional variation of a largely cosmopolitan Laurasian flora. Latest Albian and Cenomanian megafloras suggest the development of north-south provincialism within the region. Post-Cenomanian to Campanian floras show gradual diversification with various Platanaceae and Hamamelidaceae as numerical dominants. Pinnate palms are present by the early Campanian. Higher-level taxa present in the region in the Early Campanian include Magnoliales, Laurales, Chloranthales, Nymphaeales, Menispermaceae, Trochodendrales, Platanaceae, Hamamelidaceae, Cercidiphyllales, Fagales, Rosidae, and palmate Dilleniidae.