LASIAMBIX DOMINICENSIS GEN. AND SP. NOV., A EUDICOT FLOWER IN DOMINICAN AMBER SHOWING AFFINITIES WITH FABACEAE SUBFAMILY CAESALPINIOIDEAE

2008·13.2 MB·English

by  George O Poinar

AND A FLOWER
EUDICOT
LASIAMBIXDOMINICENSIS GEN. SR NOV., IN
DOMINICAN AMBER SHOWING WITH
AFFINITIES
FABACEAE SUBFAMILY CAESALPINIOIDEAE
L Chambers
George Kenton
0. Poinar,
Jr.
Department ofZoology
Oregon
State University
USA.
Oregon
97331,
Corvallis,
INTRODUCTION
Amber known
well preservative qualities of both animal and plant remains (Poinar 1992, Poinar
for
is its
&
Amber
Poinar 1999). flowers, maintained in three-dimensional form, are often preserved with such
they can be assigned extant genera (Spahr Such can be useful in paleontological
detail that to 1993). fossils
&
Poinar
reconstructions (Poinar 1999).
The amber from however legume
majority of described flowers originate Baltic deposits (Spahr 1993);
(Dilcher 1992; Poinar 1991) and palm (Poinar 2002a, 2002b) flowers have been described from Do-
et
al.
&
Burmese amber
minican amber, and there are also descriptions of Early Cretaceous flowers in (Poinar
Chambers
2005; Poinar 2007).
et
al.
During an ongoing investigation of remains in Dominican amber, 6 flowers in 5 separate pieces of
floral
amber were discovered, which could be linked through the possession of an identical cup-shaped, puberulent
One
hypanthium with deltate-ovate, spreading-recurved sepals. flower, with petals present, is probably at
show
and
a pre-anthesis stage; 3 flowers, with petals absent, are at or near anthesis; 2 flowers early fruit
pod hypanthium. The
development, with the stipe of the held in the exceptional preservation of the
still
and amber and documentation unique
flowers clarity of the allow description photographic of their features.
new
These flowers, which are described below as a genus, possess a combination of characters that favors
an assignment Fabaceae subfamily
Caesalpinioideae.
to
Journal of the Botanical Research Institute of Texas 2(1)
464
AND METHODS
MATERIALS
mountain
amber containing the flowers originated from mines in the northern range
All 5 pieces of fossil
and Dating
Dominican between Puerto Santiago.
Republic, the of Plata
Septentrional) of the cities
(Cordillera
20-15 mya on
Dominican amber with youngest proposed age of based foraminifera
the
controversial,
of
is
&
MacPhee and 45-30 mya based on coccoliths (Cepek in Schlee
1996) the oldest as
(Iturralde-Vincent
Lower
Upper Eocene
Most amber secondarily deposited in turbiditic sandstones of the to
1999). of the is
Mamey
Group
Miocene (Draper
1994).
et
al.
The morphology cup (hypanthium) and are assumed to have
6 flowers are similar in the of their floral
number
from same plant The amber piece containing the flower assigned to catalogue
originated the species.
mm
mm
and
thickness
Sd-9-154A (Holotype) circular in outline, measures 13 in diameter, 9 in greatest
is
measures
weighs 6 grams. The amber piece containing specimen Sd-9-154B is roughly triangular in shape,
mm
mm mm
The amber
3 thick and weighs 3 grams. third piece (Sd-9-
12 along the 7 the base,
sides, at is
mm
mm mm
The
and weighs grams.
154C) rectangular, measures 8 by 7 along the sides, 2 in thickness 2
is
fourth amber piece (Sd-9-154D), which contains the specimen with the petals, is semicircular in outline,
mm
mm
mm
and weighs gram.
thickness
measures 17 in greatest length, 9 in greatest width, 7 in greatest 1
The amber which contains the specimen with the trapezoidal in outline,
piece (Sd-9-154E), fruits, is
fifth
mm
mm
mm
and weighs
thickness
measures 17 in greatest length, 12 in greatest width, 8 in greatest 1
Oregon
amber maintained University.
gram. All specimens are deposited in the Poinar collection at State
&
Chambers
;>inar,
hypanthium;
with cup-shaped calyx isosepalous,
Flowers symmetrical, perfect (bisexual), a
small, bilaterally
stamens
deciduous;
imbricate or valvate; corolla isopetalous, petals free, early- 3,
sepals deltate-ovate, free, 5,
5,
incurved, anthers
hypanthium strongly
from rim opposite 3 adjacent sepals, filaments short, linear,
arising of
exceeding
hypanthial barely
bilocular, dorsifixed, introrse; ovary superior, sessile at base of cup; style single,
stamens and between the two sepals that lack stamens; stigma
stamens, positioned excentrically opposite the
deciduous.
simple, not enlarged; (immature) obovoid, tapered to a short, thick stipe, style
fruit
mm
hypanthium cup-shaped, densely tomentulose
none;
Flowers 3.6-3.7 in diameter, bilateral, pedicel
mm
mm
rim
opening 1.4-2.2 wide, bearing a distinct to
pubescent within, 1.7-2.1 long, the
externally,
on
both
tomentulose
densely surfaces,
which stamens and petals are attached; disc none; sepals deltate-ovate,
mm mm
deciduous,
spreading with reflexed 0.9-1.6 long, 1.0-1.6 at widest point; petals erect, early-
tip,
mm mm
margins
rounded
1.3-1.9 long, 0.8-1.1 wide, oblanceolate, bluntly at tip, thick-textured, ciliate;
stamens incurved, exserted from hypanthium, the filaments attached to hypanthial rim, linear,
short, slightly
mouth
hypanthium,
thickened point of insertion, glabrous, the anthers positioned horizontally over the of
at
mm
dehiscence
blunt-tipped, cordate base, 0.4-0.7 long, glabrous,
bilocular, slightly dorsifixed, ovate, at
mm
pubescent, 0.6-1.0
by lengthwise ovary in hypanthial cup, lightly style linear,
introrse sessile
slits;
(immature) obovoid, probably
pubescent proximally, stigma terminal, undivided, not enlarged; fruit
long,
mm mm
hypanthium,
mostly enclosed in the
1-few-seeded, 4.8-4.9 long, 3.7-3.9 wide, the short, thick stipe
and not
surface thick-textured rugose (as preserved), style persistent.
—
Genus name from the Greek woolly, and "ambix," cup, referring to the tomentulose
Etymology. "lasios,"
name Dominican country
from Republic, the
hypanthium specimens. Species derived the
that unifies the 6
where were
the found.
fossils
new
Poinar et Lasiambix dominicensis, a genus and species of a Tertiary eudicot flower
al.,
=
Figs. 1-3. Lasiambix dominicensis gen. and sp. nov. 1. Top view of holotype (accession # Sd-9-154A); arrow points to style. Scale bar 0.82 mm. 2.
=
mm.
Oblique view of holotype (accession # Sd-9-154A); arrow points to style. Scale bar 0.82 3. Top view of specimen Sd-9-154C; arrow points to
Research of Texas
Journal of the Botanical Institute 2(1)
=
mm.
4-7. Lasiambixdominicensis gen. and sp. nov. 4. Side view of holotype (accession # Sd-9-1 54A). Scale bar 0.84 5. D<
Figs.
=
mm.
=
men Sd-9-154C; arrow points to style. Scale bar 0.28 mm. 6. Side view of specimen Sd-9-154D; arrow points to loose petal. Scale bar 0.77 7.
=
mm.
Side view of specimen Sd-9-154E showing developing fruit; note stamens at base (arrows). Scale bar 0.57
The we believe unite these 6 specimens into a single taxon are evident in the photographic
features that
floral
Although development, the flowers have a remarkably similar
different stages of
illustrations (Figs. 1-7). at
hypanthium, with respect shape, calyx lobes, and pubescence. All have a truncate base where they broke
to
One specimen with an immature retains pair of stamens attached
a
off of their inflorescences. fruit (Fig. 7)
new
Poinar et al., Lasiambix dominicensis, a genus and species of a Tertiary eudkot flower
»0M&M
%
r
=
mm.
#
>v. (accession Sd-9-154A). 8. Top view. Scale bar 0.57 9. Oblique view. Scale b
Journal of the Botanical Research Institute of Texas 2(1)
468
and one
has
opposite the calyx lobes. The specimen in early flower (Sd-9-154D, Fig. 6) all 5 petals visible,
we The seen
what term an early-deciduous petals are clearly
corolla.
detached the base, illustrative of
at
is
The unsuitable
when specimen viewed from the top as well as the sides. center of this flower is for
the
is
The rim on
and sharp hypanthial the
remains
study due an accumulation there of apparent insect frass.
to
with
shows where presumably were attached, alternate the sepals.
near anthesis the petals It
3 flowers or
at
ambiguous whether were imbricate or valvate in the bud, perhaps because of differ-
remains the sepal lobes
on
and samples hand. Because no or pores are evident the
ent degrees of spreading separation in the at slits
by
we which supported observations of
an dehiscence,
back, or of the anthers, suggest introrse is
sides, tips
from
The mature uncertain, although
one stamen. nature of the fruit
the adaxial side of strongly inflexed is
non-compressed, few-seeded
shape and thick-textured exocarp of the 2 specimens here (Fig. a 1- or
the 7),
Although members of the Fabaceae, several extant genera of the
drupaceous suggested. atypical for
fruit
is
si by
small consisting
drupaceous Dialium characterized fruits
Cassieae have similar indehiscent, fruits. is
one seed surrounded by but somewhat dry, edible aril (Gentry 1993). Few-seeded, ovoid, fleshy-
of a soft,
Lewis This
2005).
or thick-walled fruits are not unusual in Fabaceae subfamily Caesalpinioideae (e.g. et al.
reduced
together with well- developed hypanthium, a superior simple pistil, regular corolla,
fruit-type, a
stamen number, and a zygomorphy of androecium and pistil, are what lead us to suggest assigning the fossil
and
family subfamily.
to this
numbers
stamens,
including of
The legumes extremely diverse in characteristics,
caesalpinioid are floral
and
form
presence or absence of a hypanthium, numbers and form of petals, length of filaments, of anthers
compendium
The
and symmetry, modification unisexuality,
stigmas, shape structure of fruits, floral to etc.
&
(Cowan
by and Raven with taxonomic treatments of the relevant tribes Polhill 1981;
Polhill (1981), its
& &
modern
with
Irwin Barneby 1981 Polhill Vidal 1981), was consulted in a search for genera the particular
;
were examined Taubert
and genera in
descriptions of
combination of features seen in the Illustrations
fossils.
No
examples have been found correspond in
Hutchinson and Lewis that all
(2005).
(1964), et
(1894), al.
numerous
caesalpinioid genera.
with but described are present separately in
respects Lasiambix, traits
its
sym-
A
hypanthium occurs frequently in the subfamily. Bilateral
puberulent
cup-shaped, externally
stamens such genera
common, and accompanies reduction three in as
metry (zygomorphy) also this a to
is
As by Tucker
discussed (1998,
Dialium,
Tamarindus, Macrolobium, Gilbertiodendron, Elizabetha, Apuleia, etc.
number
stamen groups genera
in of related
2002) there are parallel evolutionary trends towards reduction of
From number
and the basic
examples were Detarieae-Amherstieae Dialiinae). 10,
in separate tribes (her
some stamen
may and during development, of the
stamens be reduced few (some Dialium
to as as 1 spp.),
may and become rudimentary staminodial
growth or Gilbertio-
form (e.g.
primordia that early cease their
mentioned
and 3-stamen genera
Tucker Except Dialium Apuleia, the fall
dendron, Macrolobium; 2002). for
defined by analysis of the chloroplast trnL intron
Amherstieae (including Macrolobieae) clade, as
into the
and
by Doyle
(Bruneau 2001). This group also recognizable in the rbcL analyses et al. (1997) Kajita
et is
al.
one
but
genera leading of petal
However, the corolla evolution in these particular to loss all
(2001).
et
al.
Cowan 2-many-seeded, mostly dehiscent pods, plus the usual
form an adaxial vexillum 1953) the
to (e.g.
,
discussed
absence hypanthium, argue against a relationship to Lasiambix. Cynometra, also in this clade, is
of
a
below.
zygomorphic,
show some resemblance Lasiambix in their
and to
Dialium Apuleia, of subtribe Dialiinae,
and Barneby
few-stamened and few-seeded, indehiscent According to Irwin (1981,
flowers their fruits. p.
2-seeded
Dialium "can be defined accurately and rigorously by pod, a sort of indehiscent, 1- or nut,
101) its
and
exocarp
crustaceous
sometimes sometimes compressed consisting of a a
scarcely laterally,
strongly,
termed
when key the "drupa-
mealy endocarp enveloping the seeds." In their 100) fruit is
pulpy, ripe
(p.
and
hypogynous and have only or 2 petals usually
ceous." The flowers of Dialium, a pantropical genus, are 1
hypanthium and stamens, but
2 stamens. The related Apuleia, of South America, possesses a 3 its pistil is
and pod compressed and coriaceous, not ovoid. Despite
3-merous,
not perianth
exserted, sessile, its is its is
new 469
Poinar et Lasiambix dominicensis, a genus and species of a Tertiary eudicot flower
a!.,
their few-seeded, indehiscent pods, neither of these few-stamen genera of Dialiinae suggests a close affinity
and more numerous
Other possible relationships of the are with genera having regular flowers
fossils
One
stamens but with similar type of few-seeded fleshy such example Vouacapoua of tribe Caesal-
a fruit. is
hypanthium,
pinieae (Lewis 2005). This genus possesses regular flowers with 10 short stamens, but
et its
al.
much
sepals, and short petals resemble Lasiambix, and fruits are round, thick-walled, 1-seeded, and
its
&
The presumed
by genus
dehisce a single furrow (Lewis et 2005, 161; Polhill Vidal 1981, 89). is to
al. p. p.
Haston
have affinity to the Peltophorum group of tribe Caesalpinieae (Bruneau et al. 2001; et al. 2003), but
DNA
support connection weak.
as yet the for this
is
A
similar type of fruit occurs in the large genus Cynometra of tribe Detarieae (Lewis et al. 2005, p.
pod and
described as thick, turgid, rugose to verrucose (rarely smooth), 1-seeded, regularly in-
89). Its is
Dwyer
dehiscent with spongy pericarp (Hutchinson 1964, 235; 1958); lacks a floral disc, ovary
a its is
p. it
may
and be hypanthium (Dwyer However, has 10
usually eccentric in the 1958, 322).
hairy, style
its p. it
(Dwyer
long-exserted stamens, a shorter hypanthium, and a usually showier perianth than in Lasiambix
1958). Other differences, besides these, are that sepals are glabrous adaxially (except C. cubensis Rich.)
its
Modern-day
and has
filaments are united very briefly the base. Arcoa, of Hispaniola, turgid, 1-seeded,
at
its
indehiscent fruits (Lewis et al. 2005, p. 132), but its flowers differ considerably from Lasiambix in being
unisexual, bearing 12 stamens, and having an exserted pistil with capitate stigma (Herendeen et
ca. al.
2003). Another Caribbean genus, Stahlia (tribe Caesalpinieae), has fleshy, 2-seeded, indehiscent fruits, regular
5-merous and puberulent (Hutchinson
perianth, imbricate sepals, well-developed, dorsally petals 1964,
p.
235; Lewis 2005, 145). differs from Lasiambix principally in having 10 erect, connivent stamens
et al. p. It
and
with filaments densely woolly in the lower half. Placement of Stahlia near the large genus Caesalpinia its
Simpson
segregates was established by the molecular and morphological studies of et (2003).
al.
The examples mentioned, falling into several distinct clades within the subfamily, illustrate the paral-
make
and morphology placement
lelisms in floral fruit that difficult to suggest a specific, definitive of
it
A
Lasiambix. large amount of recent phylogenetic research, both morphological and molecular, was reviewed
and
by Lavin estimate the time frame of early diversification of this other subfamilies of
et (2005), to
al.
and
Leguminosae. Earlier estimates of a Late Cretaceous origin of the family a "basal" position for Caesal-
Herendeen no wisdom."
longer accepted "conventional
pinioideae (e.g. Polhill et al. 1981; et al. 1992), are as
my
some 60
Rather, the record supports an origination in the Paleocene, allowing for diversification
fossil
with provided by
within the family (Lavin 2005). In molecular phylogenetic studies, best resolution
et
al.
was
matK with
the chloroplast gene, divergence of 6 early-formed lineages of caesalpinioids found, along
the similarly old crown nodes of many papilionoid and mimosoid clades. Caesalpinioids are viewed as a
paraphyletic grade from which stem the mimosoids and papilionoids, but because of the very early, rapid
modern
branching of the ancestors of clades, the caesalpinioid grade "harbor(s) neither the oldest diversi-
nor some legume (Lavin 2005).
fication other quality of antiquity" et
al.
example
our suggested placement of Lasiambix correct, offers a well-preserved of several special-
If is
it
ized floral characteristics of Tertiary-age caesalpinioid legumes. These include the loss of all but 3 stamens
and accompanying bilateral symmetry of the flowers, an absence of petals at anthesis, and presence of few-
What syndrome by with
seeded, probably indehiscent pollination represented flowers short,
fleshy, fruits. is
incurved stamens and no apparent nectar Perhaps nectar was produced the base
disc, difficult to say. at
is
of the ovary, which was not visible to us. In this case small bees might have been able to enter the flowers
unknown,
and remove pollen from the arched, introrse anthers. The ecology of the plants likewise other
is
than that they grew in proximity Hymenaea in moist tropical forests such as those described by Poinar
to
many
modern-day
and Poinar (1999). Such habitats are typical for genera of Caesalpinioideae.
we
unknown,
have
Note: In the because the orientation of the flowers not tried to suggest
illustrations, is
(Cowan
a placement of the stamens as either abaxial, as in Macrolobium et al. 1953) or adaxial as in Dialium
(Tucker
1998).
et
al.
Journal of the Botanical Research Institute of Texas 2(1)
Oregon Uni-
Aaron and Botany Department,
The authors thank Richard Halse, Liston, Gerald Carr, State
John Herbarium, and Margriet Weatherwax, Jepson Herbarium, University
University
versity; L. Strother,
Ana Lawrence Smithsonian
Rancho Garden; Skog,
Robert Thorne, Santa Botanic
of California, Berkeley;
and Schlumpberger,
Systematic Botany, University of Zurich; Boris O.
Institution; Peter Endress, Institute for
Munchen, concerning
discussions the place-
Ludwig-Maximilians-Universitat,
Systematic Botanical Unit, for
We anonymous
and two
ment Steven Manchester, Roberta Poinar
of these flowers. are also grateful to
fossil
comments on manuscript.
reviewers versions of the
for earlier
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