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PUBLISHED: 15 FEBRUARY 2016 | ARTICLE NUMBER: 16005 | DOI: 10.1038/NPLANTS.2016.5

An asterid ower from neotropical

mid-Tertiary amber
George O. Poinar Jr1* and Lena Struwe2*
Fossils preserved in amber may provide signicant palaeoevolutionary and biogeographical data regarding the evolution of
life on Earth1. Although amber is particularly noted for its
detailed preservation of arthropods, the same degree of preservation can be found for vascular plant remains2. Mid-Tertiary
Dominican amber is a rich source for such fossils, and representatives of several angiosperm families have been described.
However, no fossilized examples of the large asterid plant
clade have yet been reported. Here we describe the rst
fossil neotropical owers found in amber from a representative
of the asterids. The asterids are one of the largest lineages of
owering plants, containing groups such as the sunower,
potato, coffee and mint families, totalling over 80,000
species3. The new fossils are only known as owers, more precisely corollas with stamens and styles. We here describe them
as a new species, Strychnos electri sp. nov, in the plant family
Loganiaceae (Gentianales).
The order Gentianales contains over 16,000 species in ve
families, with the majority in Rubiaceae. The new fossil is placed
in the Loganiaceae, a relatively small family comprising 1315
genera of mostly tropical and subtropical herbs, shrubs, trees and
lianas4,5. The genus Strychnos is pantropical and has at least 200
species, making it the largest genus of Loganiaceae worldwide and
in the Neotropics5. Species of Strychnos occur as woody lianas,
shrubs or trees in both seasonally dry and wet areas of tropical
and some subtropical areas6,7. In the New World, Strychnos is distributed from Mexico to Bolivia with one species in the West
Indies. In the Old World, Strychnos occurs throughout tropical
Africa and Madagascar, India, Sri Lanka, Southeast Asia and northern
portions of Australia.
Dating of Dominican amber is imprecise, with the youngest proposed age being 1520 million years (Myr) ago based on foraminifera8
and the oldest at 3045 Myr ago based on coccolith evidence9.
The asterid crown lineage has been estimated to have a median
origin age of 110128 Myr ago in the Cretaceous10,11, using minimum
fossil dates and relaxed molecular-clock-based dating analyses. The
total fossil record for asterids is particularly poor compared with
other angiosperm groups. The monophyletic, closely related asterid
orders Gentianales, Solanales and Lamiales later diverged within the
Lamiid clade of the asterids at a median time of 8698 Myr ago10.
The age of the Loganiaceae (Gentianales) is estimated at a median
time of 60 Myr ago (Late CretaceousEarly Palaeogene).
Strychnos Linnaeus, 1753
Strychnos electri sp. nov.
Etymology. Elektron (Greek). Amber. The species epithet indicates the geologic source of this species.
Holotype. California Academy of Sciences Geology Type collection
cat. no. 73176, G. Poinar no. Sd-9-47A, 1988 (Figs 1ac and 2b,c).

Additional material. California Academy of Sciences Geology

Type collection cat. no. 73631 (paratype), G. Poinar no. Sd-9-47B,
1988 (Fig. 2a).
Locality. Dominican Republic, Province of Santiago;
Municipality of Santiago de los Caballeros between Puerto Plata
and Santiago, La Bcara amber mine, 19.34 N, 71.23 W.
Horizon. Mid-tertiary (4515 Myr ago), El Mamey formation,
Dominican Republic.
Diagnosis. Flowers bisexual, actinomorphic, 5-merous. Corolla
sympetalous, trumpet-shaped, 8.79.0 mm long, abaxially densely
tomentose with non-branched, wool-like trichomes, adaxially glabrous in mouth; tube 4.8 mm 2.63.3 mm (upper width), gradually
narrowing towards the base; lobes spreading to reexed, triangularovate, 1.31.6 1.11.3 mm, valvate, reexed, densely tomentose
on outside, without pubescence in corolla mouth or on inside of
corolla lobes, apex acute. Stamens equal in number to and alternate
with corolla lobes, inserted in corolla lobe sinuses, partially exserted
from corolla mouth; anthers sessile, bilocular, longitudinally introrse,
obovate, 0.60.8 mm long, sagittate, apex rounded to bluntly acute,
bases acute; pollen spherical, 1218 m in diameter. Ovary not
a

Figure 1 | Strychnos electri sp. nov. holotype Sd-9-47A in Dominican

amber. a, Lateral view of ower. Scale bar, 1.1 mm. b, Top view of ower.
Arrow shows stigma. Note cleft anther bases. Scale bar, 0.5 mm. c, Lateral
view of top of corolla tube with tips of anthers. Scale bar, 0.3 mm.

Department of Integrative Biology, Oregon State University, Corvallis, Oregon 97331, USA. 2 Department of Ecology, Evolution and Natural Resources and
Department of Plant Biology and Pathology, Rutgers University, New Brunswick, New Jersey 08901, USA. * e-mail: poinarg@science.oregonstate.edu;
struwe@aesop.rutgers.edu
1

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Figure 2 | Strychnos electri sp. nov. in Dominican amber. a, Top view of

paratype Sd-9-47B. Scale bar, 1.7 mm. b, Trichomes on surface of oral tube
of holotype Sd-9-47A. Scale bar, 50 m. c, Pollen grains adjacent to anther
of holotype Sd-9-47A. Scale bar, 37 m. d, Lateral view of ower of the
extant Strychnos pseudoquina. Scale bar, 1.4 mm.

seen; style one, simple, lamentous, exserted from corolla (at least
810 mm long past corolla lobes); stigma capitate (Figs 1 and 2).
Notes. Both specimens retain the corolla, staminal unit and style
that have separated from the ower base after anthesis.
Consequently, the calyx and ovary are missing. We have compared
the amber fossils with owers from Strychnos taxa with external

DOI: 10.1038/NPLANTS.2016.5

corolla hairs using herbarium specimens and literature6,7. The new

fossil species can be distinguished from extant neotropical species
of Strychnos by having a combination of these morphological traits:
short corollas (under 10 mm long), prominently exserted styles in
proportion to the corolla tube (810 mm long part exserted outside
corolla mouth), corolla tube surface densely tomentose on the
outside, lack of dense pubescence inside the corolla mouth and apparently sessile anthers (Table 1). The two fossilized owers differ
slightly in size and colour, but share the same morphological characters and are considered the same species. The only representative of
Strychnos that occurs in the West Indies today (including
Hispaniola and Cuba) is S. grayi Griseb.6. However, the corolla of
S. grayi has a glabrous outer surface, is sparsely bearded in the
corolla mouth and ranges from 15 to 20 mm in length12.
The general bauplan of these fossil owers clearly place them
among the sympetalous asterid lineage of owering plants. Most
families of this clade have pentamerous, actinomorphic owers
with petals basally fused into a corolla tube, and ve stamens alternating with the corolla lobes and inserted in the corolla tube. Major
families with this oral bauplan are Apocynaceae, Asteraceae,
Boraginaceae, Convolvulaceae, Loganiaceae, Rubiaceae and
Solanaceae3, members of the Lamiid clade of the asterids. The placement of the fossils in the order Gentianales and in the family
Loganiaceae is supported by the following characters: corolla
trumpet-shaped and bearing ve valvate corolla lobes in bud
(non-overlapping, edge-to-edge lobes in bud), ve symmetric
stamens alternating with the corolla lobes and a single long style.
Convolvulaceae and Solanaceae also often have these characters,
but their petals are usually thinner towards the edge of their
lobes, and this is not discernible in these fossils. Rubiaceae has an
inferior ovary and interpetiolar stipules, characters that are unfortunately not known for this fossil. Of the genera in Solanaceae,
Cestrum appears to be the most similar, but the fossils strongly
extended style and thick petal edges are more similar to Strychnos
than Cestrum. The Apocynaceae has mostly contorted bud aestivation (twisted buds) and stamens included in the corolla tube, and
Asteraceae has fused anthers and a bilobed style. An earlier tentative
placement of this fossil2 was as a member of Thymelaeaceae, which
has owers that supercially resemble these, but with stamens
inserted opposite the calyx lobes (which are petaloid and true
petals are absent or scale-like), and not alternate with fully
formed corolla lobes, as in this fossil.
Placement in tribe Strychneae of Loganiaceae and specically in
the worldwide tropical genus Strychnos of these fossils is based on
the presence of these characters that all are in agreement with
extant Strychnos species: valvate corolla lobes of even thickness,

Table 1 | Floral morphological comparison of morphologically similar extant Strychnos species with Strychnos electri.
Strychnos electri
Distribution

Total corolla length

Corolla tube length
Corolla lobe length
Pubescence on outside of
corolla tube

Mid-Tertiary amber
from Dominican
Republic
1011 mm
8.08.7 mm
1.11.3 mm
Densely tomentose

Strychnos panamensis
var. hirtiora
Belize, Costa Rica,
Guatemala, Honduras,
Mexico, Panama
1723 mm
1520 mm
23 mm
Abundant or sparse,
short to long, straight
hairs
Tufts of pubescence

Corolla mouth pubescence

between anthers

Absent from corolla

mouth

Anther position

Sessile

On distinct laments

Anther length

0.60.8 mm

About 1 mm

Strychnos pseudoquina
Brazil, Paraguay

78 mm
57 mm
23 mm
Sparse long hairs

Dense tufts of
woolly
pubescence
Sessile or on very
short laments
About 1 mm

Strychnos tomentosa

Strychnos toxifera

Brazil, French Guiana,

Guyana, Suriname,
Venezuela
1520 mm
1316 mm
24 mm
Densely woolly
tomentose, hairs dense
and straight
Dense tufts of
pubescence

Brazil, Colombia,
Ecuador, Guyana,
Panama, Venezuela
1719 mm
About 15 mm long
24 mm
Abundance of long, thin
silky hairs and papillae

On short laments
(1 mm long)
About 1 mm

On short laments
(23 mm long)
About 1 mm

Dense tufts of woolly

pubescence

Data from Krukoff and new observations.

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DOI: 10.1038/NPLANTS.2016.5

the comparatively long corolla tube (versus shorter lobes) and a

corolla tube narrow at the base and becoming wider upwards, introrse and basixed anthers, long and liform style and a capitate
stigma. Of these characters, the ones that vary in living species are
corolla shape and size and the extent of hairiness of the corolla.
While the owers of these fossils are not complete, and vegetative
as well as some oral parts are missing, we feel condent that
their placement in Strychnos is the best-supported taxonomic placement of this fossil based on current data. We also see no need to
erect a new genus for this fossil because its morphology ts well
into the current circumscription of Strychnos.
Amber from mines in the Dominican Republic originates from
resin-producing trees of the species Hymenaea protera 13. The
species composition of the Dominican amber forest (of midTertiary age) has been characterized on the basis of both animal
and plant fossils as a tropical moist forest1. Plant fossils previously
recovered from Dominican amber include monocots (Araceae,
Arecaceae and Poaceae), basal angiosperms (Lauraceae and
Myristicaceae) and rosids (Anacardiaceae, Burseraceae, Celastraceae,
Chrysobalanaceae, Fabaceae, Meliaceae, Rhamnaceae and
Ticodendraceae)2. The fossils show that the original forest contained
a distinct canopy layer composed of legumes such as algarroba
(Hymenaea protera), cativo (Prioria spp.) and nazareno (Peltogyne
spp.), with emergent trees like caoba (Swietenia; Meliaceae) extending
through the canopy. The subcanopy and understory were represented
by royal palms (Roystonea) and gs (Ficus; Moraceae). The shrub
layer included other types of palms as well as acacias. Grasses like
pega-lega (Pharus) and bambusoids (Alarista) colonized the forest
oor. Orchids, bromeliads, ferns and vines covered the trees, and
various lianas were also part of this tropical forest.
In total, the asterids contain ten orders and 98 families and
approximately 80,000 species, representing about one-third of
angiosperm diversity (assuming around 250,000 species of
angiosperms14). The asterid clade has two major subclades, the
Lamiids and the Campanuliids, and a few more basally placed
orders and families. Recent studies have shown that several
Lamiid family stem lineages (for example, Bignoniaceae,
Gentianaceae, Solanaceae and Verbenaceae) are likely to have originated in South America in the Late Cretaceous, with subsequent dispersal to North America and the rest of the world to reach their
current distributions11,15,16. This implies that many related families
in the orders of Gentianales, Lamiales and Solanales, such as
Loganiaceae, had also formed by Early Cretaceous and that the
Neotropics are likely to be part of their ancestral areas.
The Loganiaceae has a problematic and poor fossil record, and
S. electri represents the rst conrmed Palaeogene/Neogene fossil of
this family. Frasier5 reinvestigated two reported fossils of this family,
a leaf impression of Strychnos sp.17 and seeds of Geniostoma sp.18
and concluded that due to damage and incomplete preservation,
neither could be conrmed as belonging to the Loganiaceae.
In conclusion, S. electri provides evidence that highly derived
asterid groups were present in mixed neotropical forests by the
mid-Tertiary. This suggests that many other present-day neotropical asterid genera and families had also evolved by this time, but
their remains have not yet been discovered. This fossil adds an
important piece to the evolutionary jigsaw puzzle of neotropical
Caribbean forest elements that existed in the mid-Tertiary, long
before North and South America were connected by the Panama
land bridge.

Methods

The two fossil owers are in separate pieces of Dominican amber. They originated from
a mine in the northern mountain range (Cordillera Septentrional) of the Dominican

Republic between Puerto Plata and Santiago on the Caribbean island of Hispaniola.
The amber occurred in the El Mamey formation, which is a Tertiary shalesandstone
deposit interspersed with a mixture of pebbles. Most of the amber was localized in a
ne-grained micaceous carbonaceous laminated greywacke within this deposit1.
Observations and photographs were made with a Nikon SMZ-10 R stereoscopic
microscope and Nikon Optiphot compound microscope with magnications up to 600.
Helicon Focus Pro X64 was used to stack photos for better clarity and depth of eld.
Comparative studies of herbarium materials of extant neotropical Strychnos
species were made by the second author at the herbaria of the Academy of Natural
Sciences of Drexel University in Philadelphia, Pennsylvania, and the New York
Botanical Garden, Bronx, New York.

Received 24 June 2015; accepted 12 January 2016;

published 15 February 2016

References
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Acknowledgements
The authors graciously thank K. Gandhi of Harvard University for help with Latin
nomenclature and D. Bhattacharya of Rutgers University for comments on an earlier
version of this article.

Author contributions
G.O.P. provided the data and analysis of the amber fossils (measurements and
morphological data), all photographs except the extant Strychnos species and was the
primary author for the palaeontological parts of the manuscript. L.S. provided data, photo
and text regarding extant taxa in the Gentianales and other asterids, evaluation of asterid
dating times and was the primary author of the new species description.

Additional information
Reprints and permissions information is available online at www.nature.com/reprints.
Correspondence and requests for materials should be addressed to G.O.P. and L.S.

Competing interests
The authors declare no competing nancial interests.

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