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

Abstract

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.

Full text

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
Source:
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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MAULDINIA HIRSUTA SP. NOV., A NEW MEMBER OF THE EXTINCT GENUS
MAULDINIA (LAURACEAE) FROM THE LATE CRETACEOUS
(CENOMANIAN-TURONIAN) OF KAZAKHSTAN
Suembikya Frumin,* Helena Eklund,yand Else Marie Friis1
,
y
*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.
Introduction
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 else.marie.friis@nrm.se.
Manuscript received October 2003; revised manuscript received February
2004.
883
Int. J. Plant Sci. 165(5):883–895. 2004.
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subprovince (Herngreen and Chlonova 1981; Herngreen et al.
1996).
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
3
and KClO
3
) 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%.
Observations
Systematic Description
Order—Laurales
Family—Lauraceae
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 (fl) 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 (fl) 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 (fl) 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.
886
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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). D–F, 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.
887
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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
units.
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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FRUMIN ET AL.—MAULDINIA HIRSUTA SP. NOV. (LAURACEAE) FROM KAZAKHSTAN
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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
compressed.
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
apex.
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.
4A,4D,4E).
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
891
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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.
Discussion
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
Kvac
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
ˇek
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.
mirabilis.
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.
893
FRUMIN ET AL.—MAULDINIA HIRSUTA SP. NOV. (LAURACEAE) FROM KAZAKHSTAN
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Conclusion
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.
Acknowledgments
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
anguste-obovate
lobes; distinct
branching system
of lateral unit
Ovate lateral unit with
anguste-obovate
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
breadth
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
breadth
5–6 length, 3–4 breadth 5.5 length, 3.1 breadth 2.0–3.5 length, 1.8
breadth
No. flowers 5 Not known 3–5–(7)–9 5
Flower size (mm) 1.3–1.7 length, 0.6–0.9
breadth
2.3 length 1.5–2.6 length, 0.6–1.4
breadth
1.5–3.6 length, 1.0–2.0
breadth
Floral tube length (mm) 0.20–0.25, abruptly
separated
Not known 0.09–(0.19)–0.29, not
separated
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
folded
Outer: triangular; inner:
distinctly keeled and
folded
Stamen whorls Not known Not known 3 3
Staminal appendages Stalk with flat angular
head, 0.6–0.7 mm
long
Not known Stalk with glandular
head, 0.8–1.2 mm
long
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
appendages
Dense on adaxial
surface of lateral
unit (?)
Scattered on lateral unit,
carpel, pistil,
occasionally present
on filaments and
staminodes
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
894 INTERNATIONAL JOURNAL OF PLANT SCIENCES
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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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