AN EARLY CRETACEOUS ANGIOSPERM FOSSIL OF POSSIBLE SIGNIFICANCE IN ROSID FLORAL DIVERSIFICATION

2008·7.4 MB·English

by  George O Poinar

AN
EARLY CRETACEOUS ANGIOSPERM
OF
FOSSIL
POSSIBLE SIGNIFICANCE ROSID FLORAL
IN DIVERSIFICATION
George
0. Poinar, Kenton L Chambers
Jr.
Department ofBotany and Plant Pathology
Oregon
State University
Oregon
Corvallis, 97331,
U.S.A.
Ron
Buckley
INTRODUCTION
Burmese amber AD
is first mentioned in works dating from 100, when an amber trade route was established
AD
between that country and China. From 100 until 1936, the Burmese amber mines amber
supplied
for
jewelry and
fossils. Social upheavals closed the mines for 63 years, but in 1999, extraction procedures
be-
gan anew. The fossil was derived from a mine first excavated in 2001, in the Hukawng southwest
Valley, of
Maingkhwan
in the state of Kachin (26°20'N, 96°36'E) in Burma (Myanmar). This new amber known
site,
Bum
Summit
as the Noije 2001 Site, was assigned to the Upper Albian of the Early Cretaceous on
the basis
&
of paleontological evidence (Cruickshank Ko
2003), placing the age at 97 to 110 mva. Nuclear magnetic
Summit
2001
an
Site indicate araucarian (possibly Agathis) tree source for the amber (Poinar 2007).
et
al.
Within
the past few years, amber from this area has yielded a wealth of arthropod and plant remains
(Poinar et al. 2005). While insects dominate, some very interesting angiosperm have been
fossils recovered
&
up
now,
to including flowers with possible affinities to the families Monimiaceae (Poinar Chambers 2005)
and
Cornaceae
(Poinar et al. 2007), as well as two early bambusoid grasses (Poinar 2004). In the present
we
paper, describe an additional flower from the same location, with similarities to and extant members
fossil
of eurosid orders Oxalidales, Cucurbitales, Sapindales, and eudicot order
Saxifragales.
MATERIALS AND METHODS
An
entire, well-preserved female flower with staminodes # embedded
(accession ab-294) round
is in a piece
mm
made
mm. Examination and photographs were with
amber with diameter of 40 and a thickness of 5
of a
SMZ-10 R 80x and Nikon Optiphot microscope at 800x.
Nikon microscope a
stereoscopic at
a
DESCRIPTION
&
CO.
Lachnociona G.O.Poinar, K.L. Chambers R. Buckley, gen. nov. Type Species: Lachnociona teniae Poinar, K.L.
&
Chambers R. Buckley, sp. nov.
5-merous,
specimen
unisexual, the type functionally pistillate,
Diagnosis.— Flowers small, pedicellate,
recurved, lightly
ovate-lanceolate, ciliate,
actinomorphic, apparently apetalous; sepals regular, separate,
5,
conspicuous,
hirsutulous on both surfaces, venation obscure; staminodes 10, in 1 or 2 whorls, filaments
anthers
and hooked in some, the fully preserved ones all non-functional,
slender tapering, the tip retrorsely
by on 3 staminodes; gynoecium 5-carpellate, densely lanate-hirsute,
absent or represented thin vestigal tissue
by pubescence, con-
perhaps and the disc present) concealed the styles
ovary superior or half-inferior, (if
it
recurved-divergent in the upper 1/4-1/3, the stigma probably
sharply acute,
nivent or partly united, their tips
unknown.
and
decurrent ventrahy; staminate flower fruit
&
myanmar
Lachnociona teniae G.O.Poinar, K.L. Chambers R. Buckley, sp. nov. (Figs. 1-4). Type: (Burma):
SW Myanmar, 2001 unknown
northern
amber mine Hukawng of Maingkhwan in the state of Kachin (26°20'N, 96°36'E),
in the Valley;
,
specimen can be made by contacting Ron Buckley at [email protected].
Arrangements study the type
to
mm;
mm,
mm; mm; width length of
Description.—Flower length 5.6 flower width 5 sepal length 3.60 1.8
mm; most broken chewed length of remaining
on 0.25-0.36 staminodes exserted, or off;
trichomes sepals
mm;
mm;
mm; width anther 0.20 length
0.11-0.88 of largest
anthers
1.80-1.90 length of vestigial
filaments
mm;
gynoecial column 3.30 mm; length of divergent upper lobes of style 1.20 length of subtending bract
of
mm.
on mm; distance from base of bract to summit of hirsutulous pedicel 2.90
pedicel 1.10
—
Genus name from the Greek "lachnos," woolly hair, and "kion," column, with reference to the
Etymology.
named honor Mrs. Terri Millinoff.
gynoecium. Species in of the collector,
portion of the
projecting stylar
staminodes,
apparent apetaly, 10
The salient features of the fossil, as described above, are its unisexuality,
The
gynoecium, column, and acute, spreading stigmatic lobes.
densely pubescent, 5-carpellate stout stylar
expanded
from
judging the distal
superior
most reasonable interpretation of the ovary is that it is either or,
The anther tissue on 3 of the filaments is noteworthy, suggesting
region of the pedicel, half-inferior. vestigial
by hence derivation from hermaphrodite ancestors. In framing a hypothesis for the
unisexuality abortion,
which above group
we best the
have searched present-day families
systematic position of Lachnociona, for fit
assumptions made about features not clearly visible in the fossil. Because of the
reasonable are
of
traits, if
could not resolve the choice
pigmented microscopic study of this structure
thick covering of darkly hairs,
As recurved perianth parts with dark coloration at their base
between fused or connivent styles. well, the
we
beneath them, and thus interpret the perianth
prevented observation of any outer perianth parts
tiny,
deciduous.
with the corolla either lacking or
as a calyx,
and Cunoniaceae
Brunelliaceae
Possible candidates have been identified as the closely associated families
mainly
both groups occurring in the
(APG 2001 onwards; 2005),
2003; Stevens Soltis et
(Oxalidales) al.
II
We
Asian
Australian-Southeast region.
Hemisphere with in the case, in the
Southern representatives, latter
interpret the stout, densely hairy, darkly colored (in the fossil) columnar structure to be, all or in part, 5
Schonenberger
(Schonenberger 2001;
connivent-appressed The Late Cretaceous fossil Platydiscus et al.
styles.
&
with
von Balthazar 2006; Friis et 2006) was included in our analysis, because of its initial association
al.
and Aphanopetalum (Aphanopeta-
order Cucurbitales,
(Anisophylleaceae), of
Cunoniaceae. Also, Anisophyllea
was
came under consideration. Eurycoma (Simaroubaceae, order Sapindales)
of order Saxifragales,
laceae),
added comparison involving a different interpretation of the perianth of the fossil.
for a
members
Brunelliaceae
Table summarizes a comparison of floral traits of Lachnociona with four living of
1
and
Aphanopetalum and Eurycoma. Brunelliaceae
with Anisophyllea,
together Platydiscus,
,
Poinar et Lachnociona gen. and n
al., terriae sp.
number
Sepal
3.
-
4. Sepals spreading - +, upright
-
absent
5. Petals - +, present
-
Ovary
8. superior - +, ±, inferior
-
Ovary apocarpous syncarpous
9. - +,
number
Carpel
10.
-0
Styles united -+, separate
-
acute
2. Style tips - +, capitate
1
-
3. Stigma decurrent - +, terminal
1
Gynoecium -
14. hairy - glabrous
+,
5. Free styles connivent - +, spreading -
1
+(large)
Cunoniaceae, sometimes united into one family, have always been considered as closely
relat
theories of angiosperm phylogeny (e.g., Engler 1930a, b; Thorne 1992; Cronquist 1981). The fossil is similar
to Brunellia in being unisexual, apetalous, 10-stamened, and having a 5-carpelled, pubescent gynoecium
with acute style tips whose stigmatic surface is probably ventral and decurrent. Brunellia is apocarpous, the
carpels at most being immersed in the disc (Orozco Pardo 2002; Kubitzki 2004). separate may be
Its styles
vertical in early flower (Cuatrecasas 1970; Kubitzki 2004), with recurved In Cunoniaceae, the carpel
tips.
number
and
is usually 2 the stigmas, except for Vesselowskya, are terminal and often capitate (Bradford
et
However, numbers known,
al. 2004). higher carpel are in Pullea, Aistopetalum, Spiraeanthemum, and
e.g.,
Journal of the Botanical Research Institute of Texas 2(2)
Acsmithia, the latter two genera comprising a possible morphological link with Brunelliaceae (Orozco Pardo
&
DNA
2002; but see Bradford Barnes, 2001, for evidence supporting their inclusion in Cunoniaceae). Genera
and and
of Cunoniaceae with unisexual flowers include Spiraeanthemum, Pancheria, Vesselowskya, the latter
genus unique in the family in showing a ventrally decurrent stigma like that of Brunellia (Dickison 1989,
is
&
Hufford Dickison 1992) and putatively like that of Lachnociona. That the ovary of the fossil might
4.3;
fig.
be half or more inferior (judging from the expanded apex of the floral pedicel) was mentioned above; in
&
would resemble Schizomeria (Matthews Endress 2002) and Pullea (Dickison 1975) and differ from
this,
it
may
and The be and
Spiraeanthemum Vesselowskya. styles erect partially c
=
Figs. 2-3. Flower of Lachnociona tei in Burmese amber. 2. Long trichomes on surface of sepals. Scale bar
least in early flower), as illustrated for Pullea and Acsmithia by Dickison (1975, 15h, and 1989, 4.2B,
fig. fig.
respectively) and Hoogland (1979, fig. 1). Of the above-mentioned genera, petals are absent in Brunellia, Spi-
raeanthemum, and None modern
Acsmithia, Pullea, (in pistillate flowers) Vesselowskya (Dickison 1989). of the
genera listed have styles pubescent nearly to the tip like Lachnociona.
Aphanopetalum (Aphanopetalaceae) is included in Table has a 4-carpelled gynoecium with half-
1; it
and column
inferior ovary, a stout stylar with spreading, terminally-stigmatic tips (Dickison 1975,
fig. la).
Dickison shows median
(op. cit.) a longitudinal section of a flower nearing anthesis; its thick style of 4
much
fused carpels with spreading, bluntly pointed tips very resembles the stout (but more acutely tipped)
gynoecium
of Lachnociona. Significant differences are evident, however, in the androecium of 8 large,
fertile,
nearly sessile anthers of Aphanopetalum, the 4-merous calyx, and the presence of 4 poorly developed
pet-
The
als. flowers of Aphanopetalum are glabrous, as well. This genus has an interesting taxonomic history,
&
having been placed originally in Cunoniaceae (Engler 1930b; Dickison 1975; Hufford Dickison
1992),
but it was found by Bradford and Barnes (2001) to be better placed in or near Saxifragaceae, according to
their analysis of the TrnL-F intron of the chloroplast tRNA-Leu gene. Based on the morphological work of
Journal of the Botanical Research Institute of Texas 2(2)
=
mm.
Fig. 4. Drawing of flower of Lachnociona teniae in Burmese amber. Scale bar 1
Dickison et al. (1994) and the molecular phylogenetic studies by Fishbein et al. (2001) and Hermsen et al.
now
(2006), Aphanopetalum placed as a monotypic family of the order Saxifragales (Stevens 2001 onwards;
is
2005).
Soltis et
al.
Platydiscus a well-studied charcoalified Late Cretaceous fossil flower from sand and clay sediments
is
of the Kristianstad Basin in Scandia, Sweden (Schonenberger et 2001, see drawing of a reconstruction
al.
in Friis et 2006). The perianth is 4-merous, with small, densely glandular petals and spreading sepals,
al.
stamens are 8 and and the gynoecium of 4 carpels has a half-inferior ovary and connivent, erect styles.
fertile,
The spreading stylar tips are reconstructed as acute with a ventral slit; the stigmatic region is not described
may
but well have followed the in Brunellia and Vesselowskya (our suggestion, not the authors').
slit as, e.g.,
The connivent styles, with broken off tips, are well illustrated by Schonenberger et al. (2001, figs 34 and
40). Features of the gynoecium of this fossil, as illustrated and reconstructed, are thus quite reminiscent of
Lachnociona as we visualize Differences between our fossil and Platydiscus are that the latter is 4-merous
it.
throughout, bisexual, and petaliferous. Its age, given as Late Santonian-Early Campanian, is approximately
&
le latter family in the order Cucurbitales (see also Tobe Raven 1988). However, this ordinal
&
Zhang
well borne out by recent molecular studies (Schwarzbach Ricklefs 2000; et 2006).
s al.
1189
The many ways Cunoniaceae and
flowers of Anisophyllea are similar in to those of Platydiscus, as cited in detail
by Matthews et (2001). Like Lachnociona, Anisophyllea unisexual, with two whorls of stamens, separate
al. is
&
acute styles, and stigmas decurrent ventrally (Matthews et 2001; Matthews Endress 2004). differs
al. It
in being petaliferous and usually 4-merous throughout, and ovary fully inferior (Table These and
its is 1).
numerous other traits of Anisophylleaceae are discussed by Matthews and Endress (2004), giving support
to its placement in Cucurbitales despite its divergence in floral form from other families of the order.
An
alternative interpretation of the perianth of Lachnociona, namely that the conspicuous organs are
petals and that an inconspicuous calyx is out of sight beneath them, leads to comparison with a different
We
rosid group, the family Simaroubaceae of order Sapindales. have selected the Southeast Asian genus
Eurycoma (Engler 1931, 175; Nooteboom 1962, 206) for such a comparison (Table In Eurycoma,
fig. p. 1).
the conspicuously hairy flowers are unisexual, the 5-lobed superior ovary of 5 free carpels bearing 5 erect,
connate or connivent styles which are acute, recurved at the tip, and stigmatic ventrally. In fruit, the 5 carpels
and The androecium
separate spread far apart, as occurs also in Brunellia. of pistillate flowers consists of 5
much
normal-sized stamens with abortive anthers, alternating with 5 scale-like, modified staminodes. The
5 hairy petals are spreading-erect and have 5 small sepals at their base. Other than stamen number, there-
fore, and reflexed rather than erect petals, Lachnocii hypothesize that a
small calyx is hidden below the "petals." The fossil ambiguous relationship to Simaroubaceae,
t l
we
which cannot resolve with certainty. In phyloge Simaroubaceae, sensu stricto, are placed
;s,
in the rosid clade (Fernando 1996; Gadek e PG 2003; Stevens 2001 onwards;
II et al. II Soltis et
&
2005; Schonenberger von Balthazar 2006).
al.
summary,
In the case for postulating a relation: ociona to the clade containing Brunelliaceae
and Cunoniaceae rests on the similarity of the folk floral features: unisexuality, 5-merous ac-
1
tinomorphic perianth, apetaly, abundant pubescen of 10 staminodes in the pistillate flower, 5
may
carpels, a stout gynoecial column (which be) c styles, and sharply acute stylar
t
A
tips implying a ventrally decurrent stigmatic zone. present, would be consistent
if
,
with these families, but if the ovary is interpreted as would exist to Pullea and a few
&
other Cunoniaceae (Bauera, Dickison, 1975; Cerati Codia, Hufford Dickison,
f. i; f.
A
1992) as well as to Aphanopetalum and the complete decurrent "sutural" stigma
i.
&
characteristic of Brunellia (Kubitzki 2004) and (Dickison 1989; Hufford
is Vi
:
Dickison
1992).
Our
suggestion of a connection with Brunellia mnot be proven, given the
dif-
we
ficulty in observing certain critical features of the However, believe that the
g
..
placement Fernando
totality of floral features favors of the fo eudicots (e.g., et al.
&
1995; Soltis Soltis 2004; Soltis et 2005; Scheme 2006). Except for Saxifragales,
al.
large group of angiosperms.
is
grains in Late Barremian-
Dllen
Early Aptian strata (about 120 Myr) shows that eudicot angi. established in geographically
ges of crown and stem groups
of basal eudicots were presented by Anderson et (2005) us :nces and multiple fossil refer-
al.
ence points. Their inferred stem and crown group ages for rosic and 108 mybp, respectively,
>p
mybp mybp
and for Saxifragales, the figures are 108 and 102 analysis, Crepet et (2004)
al.
:
minimum
94+ mybp.
dated the age of the "rosid clade as
I" B<
Wilkstrom which stem group
et al. (2001), dates the of the ro:
(2006, rapid diversification of eudicots, with
al. p. 31), "(a) thi
end been
before the of the Albian, has.... inferred, using mole
Burm
eudicot flowers that have recently been described from
&
Poinar Chambers 2005; Poinar et 2007), together with t present report, may help open a window
al.
;
to particular features of floral evolution in this period of eudio
.
may
The flower described here have been insect-pollinated, as proposed earlier by Santiago-Blay et
al.
(2005), and could have been visited by Melittosphex burmensis, a small bee recently described from these
it
&
deposits (Poinar Danforth 2006). The flower also shows evidence of herbivory, as there obvious damage
is
and by chewing
the filaments anthers insects.
to
ACKNOWLEDGMENTS
We
The authors thank Roberta Poinar for reading earlier drafts. are grateful for technical suggestions by 2
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