ArticlePDF Available

Abstract and Figures

We describe fossil fruits collected from outcrops of the Salamanca Fm. (Paleocene, Danian, 63.3–61.9 Ma) at the Estancia Las Violetas locality, Chubut, Argentina that show affinities with members of the Subtribe Attaleinae, Tribe Cocoseae, Subfamily Arecoideae within the Arecaceae. The fossils are preserved as permineraliza-tions, and were examined by longitudinal, tangential and cross-sections, as well as by the application of Computed Tomography Scanning technology (CT Scan). The fruits are ovoid drupes with 3 longitudinal grooves delimiting three valves; displaying apical stigmatic remains and a single apical germination pore. The exocarp and mesocarp are fused and anatomically indistinguishable one from another; they contain longitudinal fibrous bands with brachysclereids. A centrally placed seed occupies the single locule entirely; the seeds are deltoid with a basal hilum and acuminate tip while the endosperm is ruminate. The taxonomic position of the fossils was explored using phylogenetic analyses of molecular sequences combined with morphological data. Along with the suite of morphological characters that points to an affinity with the Subtribe Attaleinae, Tribe Cocoseae, subfamily Arecoideae, the results of the combined phylogenetic analyses confirm the taxonomic placement. This report constitutes the first confirmed record for the Subtribe Attaleinae worldwide and the first record of fossil palm fruits from Argentina.
Content may be subject to copyright.
Arecaceae Fossil Fruits from the Paleocene
of Patagonia, Argentina
Mary K. Futey
1
& Maria A. Gandolfo
1,4
&
Maria C. Zamaloa
2
& Rubén Cúneo
3
&
Gerardo Cladera
3
1
L. H. Bailey Hortorium, Department of Plant Biology, Cornell University, Ithaca, NY 14850, USA
2
Departmento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad
de Buenos Aires, Intendente Güiraldes 2620, C1428EHA Buenos Aires, Argentina
3
Museo Paleontológico Egidio Feruglio, Avenida Fontana 140, CP 9100 Trelew, Chubut, Argentina
4
Author for Correspondence; e-mail: mag4@cornell.edu
#
The New York Botanical Garden 2012
Abstract We describe fossil fruits collected from outcrops of the Salamanca Fm.
(Paleocene, Danian, 63.361.9 Ma) at the Estancia Las Violetas locality, Chubut,
Argentina that show affinities with members of the Subtribe Attaleinae, T ribe Cocoseae,
Subfamily Arecoideae within the Arecaceae. The fossils are preserved as permineraliza-
tions, and were examined by longitudinal, tangential and cross-sections, as well as by the
application of Computed Tomography Scanning technology (CT Scan). The fruits are
ovoid drupes with 3 longitudinal grooves delimiting three valves; displaying apical
stigmatic remains and a single apical germination pore. The exocarp and mesocarp are
fused and anatomically indistinguishable one from another; they contain longitudinal
fibrous bands with brachysclereids. A centrally placed seed occupies the single locule
entirely; the seeds ar e deltoid with a basal hilum and acuminat e tip while the endosperm is
ruminate. The taxonomic position of the fossilswasexploredusingphylogeneticanalyses
of molecular sequences combined with morphological data. Along with the suite of
morphological characters that points to an affinity with the Subtribe Attalein ae, Tribe
Cocoseae, subfamily Arecoideae, the results of the combined phylogenetic analyses
confirm the taxonomic placement. This reportconstitutesthefirstconfirmedrecordfor
the Subtribe Attaleinae worldwide and the first record of fossil palm fruits from Argentina.
Keywords Arecaceae
.
Fossil fruits
.
Paleocene
.
Salamanca Formation
.
Patagonia
.
Argentina
Introduction
Arecaceae is a large and economically important family of monocots with ca. 2300
species in 183 genera; with a worldwide distribution restricted to the sub-tropics to
tropics (Dransfield et al., 2008). Of all monocots, Arecaceae has one of the richest
fossil records both temporal ly and spatially (Daghlian, 1981); fossils assigned to it are
found on every continent except Antarctica and interestingly often display a
Bot. Rev.
DOI 10.1007/s12229-012-9100-9
previously higher level of diversity and distribution range than observed today (Pan et
al., 2006). Harley (2006) provides a review of the fossil record of palms that includes
several organs, alth ough the most abundant are stems, fruits, seeds, leaves and pollen
grains. Because of the distinctive diagnostic features of Arecaceae, taxonomic place-
ment of fossils within the family is almost always guaranteed; nevertheless, due to
character vari ability within the family the majority of the fossils assigned to
Arecaceae cannot be placed with confidence within any modern taxa below familiar
or subfamiliar level (Harley, 2006; Dransfield et al., 2008). The earliest reliable
macrofossils are stems placed within the morphogenus Palmoxylon Schenk dated to
the Turonian of France (Crié, 1892) and Coniacian-Santonian of New Jersey (Berry,
1916) and palmate leaves appearing in the Coniacian-Sant onian of South Carolina
(Berry, 1914). Currently Hyphaeneocarpum aegyptiacum (Vaudois-Miéja & Lejal-
Nicol, 1987) from the Aptian of Egypt is the oldest record of fossil palm fruits.
Although in South America modern genera of the family are highly represented,
only 12 extant species within 8 genera are found naturally in Argentina (Zuloaga &
Morrone, 1996)specificallyinthenorth,northeastandinanareaknownas
Mesopotamia (area that comprises the Misiones, Corrientes and Entre Ríos provinces;
Fig. 1). Remarkably, the oldest fossil palm records from Argentina come from the
Upper Cretaceous of Patagonia (Ancibor, 1995; Ottone, 2007) while the most recent
ones are from the Pliocene of Mesopotamia (Lutz, 1980, 1984). Patagonian
Arecaceae fossils are represented by stems, leaves and pollen grains. Based on the
pollen record, the family is represented by several species of the genera Arecipites,
Confertisulcites, Liliacidites, Longapertites, Monosulcites, Psilamonocolpites,
Sabalpollenites, and Spinozonocolpites (Gandolfo et al., 2010). Argentinean fossil
stems are referred to the morphogenus Palmoxylon Schenk 1882 and consist of 9
species that date from the Upper Cretaceous to the Pliocene (Ancibor, 1995; Argujio,
1979, 1981; Gandolfo et al., 2010; Lutz, 1980, 1984; Ottone, 2007; Petriella, 1972;
Romero, 1 968). The family is also represented by phytoliths (Zucol et al., 2007) and
as of yet, undescribed pinnate and bifid palm leaves and fruits (pers. obs). Herein, we
describe fossil fruits from Patagonia, Argentina with a clear affinity to the members of
the subtribe Attaleinae, tribe Cocoseae within the subfamily Are coideae.
Material and Methods
Geologic Background and Age
The fossil remains were collected from outcrops of the Salamanca Formation, which
are exposed at the Estancia Las Violetas locality, Chubut province, Argentina (Fig. 1;
Somoza et al., 1995). Here, a 10 m thick section composed of heterolithic facie s
(conglomerates and fine to medium grained sandstones) was deposited under littoral
shallow marine conditions, including severa l tidal channel deposits represented by
glauco
nitic
bioturbated sandstones that bear the local fossil flora. The associated
fossil logs have intense bioturbation of the Terodolites ichnofacies.
Based on paleomagnetic data, the section at Ea. Las Violetas has been assigned to
Chron 27r of the Geomagnetic Polarity Time scale dated between 63.3 and 61.9 my
(late Danian).
M.K. Futey et al.
Specimen Preparation and Examination
The examined specimens consi st of 25 permineralized fossil fruits. Selected fossils
were cleaned using traditional paleobotanical techniques, using a small pneumatic
hammer, and degagement. Ultra thin and double polished longitudinal, tangential and
transversal sections were performed on two selected specimens at Texas Petrographic
Services, Inc. with the goal of observing anatomical and histological characters.
Sections were examined using an Olympus BX60 microscope and photographed
with a Nikon D70 a Sony DXC-9000. A section set is housed at the Paleobotanical
Collection, Cornell University, USA (CUPC 1521 ab) and the remainder set is
housed at the Paleobotanic al Collection of th e Museo Paleontológic o Eg idio
Feruglio, Trelew, Argentina under the numbers MPEF-Pb- 3767 ac.
A fossil fruit and a fossil seed are housed at the Paleobotanical Collection, Cornell
Univers ity, USA under the numbers CUPC 1522 and CUPC 1523 respectively;
remainder fossils are housed at the Museo Paleontológico Egidio Feruglio under
Fig. 1 Map of Ar gentina show ing
the extant distribution of Arecaceae
(hash-mark) and locality where
fossils were collected (circle)
Arecaceae fossil fruits from Argentina
the number MPEF-Pb-3766 (holotype) and numbers MPEF-Pb 3768, 3769, 3826
3830 (3829 and 3830 are two sets of several fossils each). Some of these fossils were
subjected to X-ray tomography imaging that was conducted at the Cornell Imaging
Center using a GE CT 120 eXplore, with 2 Gy to 8 Gy of radiation and 120,000 eVof
voltage. This technique was used for observing rocks that contain fossils and isolated
fossil fruits. They were placed within the CT Scan machine without any additional
preparation. The images produced from the CT scan were viewed as series of still
images or as movies. Within the rocks, the scan uncovered the presence of 2 fruits
that could not be observed otherwise. Additionally, the scans revealed anatomical
details such as the position of the germination pore, the thickness and relation to each
other of the fruit layers and endosperm features that were not observed in the ultra
thin sections. The use of CT scan technology was able to reveal and clarify several
characters that were instrumental in describing and classifying these fossils while
leaving the specimens fully intact.
Combined phylogenetic analyses were conducted using sequences downloaded
from Genbank (Table 1) and morphological data that were compiled from literature
and personal observation of herbarium material relevant to this study (Tables 2 & 3,
Appendix 1). Nineteen taxa were selected from all five Arecaceae subfamilies with
emphasis on New World species. Three members of Dasypogonaceae were selected
as outgroups and coded from molecular data only. All taxa were scored at generic
level, and some terminals are re presented by more than a species, for example
molecular data for the genus Bactris comes from a combination of the species B.
gasipaes (rbcL)andB. maraga (PRK). The morphological matrix consisted of
fourteen characters and was modified from Baker et al. (2009)withadditional
characters, of longitudinal grooves, mesocarp furrows, fibers in mesocarp, endocarp
grooves, and number of seeds, coded by the examination of herbarium material and
fossil specimens (Table 3). Two genes were selected to provide adequate resolution
and taxa coverage: low-copy nuclear gene phosphoribulokinase (PRK) and chloro-
plast gene ribulose-bisphosphate carboxylase (rbc L). Both sequences were aligned
using MUSCLE v.3.831 (Edgar, 2004). For the fossil taxon all molecular data was
scored as absent. The combined matrix (Appendix 1)containedatotalof202
informative characters. Equal weighted parsimony analyses were conducted using
NONA (Goloboff, 1998 ) as implemented in WinClada (Nixon, 1999). 300 heuristic
search replicates (mult*300) with random starting trees were conducted; a maximum
of 10 trees were held in each replicate (hold/10) and TBR branch swapping (max*)
was conducted on all held trees. A strict consensus tree of the resulting most
parsimonious trees was constructed using TNT (Goloboff et al., 2008), of 500
replicates with 10 replicates each and TBR branch swapping. The bootstrap values
were placed on the strict consensus tree.
Results
Systematics
Family: Arecaceae Schultz 1832
Subfamily: Arecoideae
M.K. Futey et al.
Table 1 List of extant species used to compile the combined molecular and morphological data. All
species used for coding morphological characters are housed at the L.H. Bailey Hortorium Herbarium,
Cornell University (BH). Molecular sequences (rbcL and PRK) were downloaded from Genbank
Species BH Accession numbers Genbank PRK
Genbank rbcL
Outgroup
Calectasia intermedia AF206743
Dasypogon bromeliifolius AM110246
Kingia australis AM110245
Ingroup
Acrocomia aculeata 37261 AM110212 AY601224
Attalea allenii 37276 AJ404829
Attalea phalerata AY601240
Bactris gasipaes 37280 AM110214
Bactris maraga AY601214
Barcella odora 37275 AY044630
Butia yatay 37262
Ceroxylon quindiuense 37282 AJ404781
Ceroxylon amazonicum EF128386
Chamaedorea microspadix 37281 AJ404787
Chamaedorea metallica EF491095
Cocos nucifera 37283 AM110211 AY601232
Copernicia macroglossa 37278
Elaeis guineensis 37263 AJ404830 AY601221
Geonoma congesta 37284 AM110219
Geonoma edulis HM140599
Lytocaryum hoehnei 37273; 37279
Lytocaryum insignis 37264
Lytocaryum weddellianum 37274; 37265 AY044633 AY601249
Mauritia flexuosa 37285 AJ404777
Nypa fructicans 37286 AJ404778 AJ831357
Pseudophoenix vinifera 37277 AJ404780
Pseudophoenix sargentii EF128364
Sabal bermudana 37287 AJ404766
Sabal mexicana EF667947
Socratea exorrhiza 37288 AM110205 AF453378
Syagrus flexulosa 37266
Syagrus oleracea 37269
Syagrus romanzoffiana 37267; 37268
Syagrus smithii 37270 AJ404827 AY601260
Syagrus sp. 37272
Synechanthus warscewiczianus 37289 AJ404786
Synechanthus fibrosus EF491103
Trithrinax campestris 37271 AJ40474
Arecaceae fossil fruits from Argentina
Tribe: Cocoseae Mart.
Genus: Tripylocarpa gen. nov. Gandolfo and Futey
Type species: Tripylocarpa aestuaria sp. nov. Gandolfo and Futey
Generic Diagnosis. One seeded ovoid drupe, exocarp and mesocarp fused and
indistinguishable one from another; three external longitudinal grooves delimiting
three valves, endocarp thin, not sculptured; apical stigmatic remains; one locule, seed
deltoid; endosperm homogeneous and ruminate ; apical germination pore and basal
hilum.
Specific Diagnosis. As for the genus Tripylocarpa.
Holotype: MPEF-Pb-3766; Museo Paleontologico Egidio Feruglio, Trelew,
Chubut Province, Argentina.
Paratypes: MPEF-Pb-3767 a c; 3768, 3769, 38263830 and CUPC 1521 ab and
15221523.
Type locality: Las Violetas, Salamanca Formation, Chubut Province, Patagonia,
Argentina.
Table 2 Morphologic al matrix compiled from herbarium observation s. References to all previously
published matrices serving as sources for the following defining and/or scoring the characters as well as
primary literature consulted in scoring are listed following each character. All characters are nonadditive.
Character numbering corresponds to WinClada numbering; numbering for NEXUS files will start with 1
rather than 0
0123456789101112 13
Mauritia flexulosa 01000000 00 1 0 0
Trithrinax campestris 00000000 00 1 0 0
Sabal bermudana 10 000 0 0 0 1 0 [01]
Nypa fruticans 10 001 0 00 1 0 1
Ceroxylon quindiuense 10 000 0 0 0 1 0 [01]
Pseudophoenix vinifera 10 000 0 00 1 0 1
Geonoma congesta 10 001 0 00 1 0 0
Socratea exorrhiza 00 001 0 00 1 0 0
Chamaedorea microspadix 10 000 0 00 1 ? 0
Synechanthus warscewiczianus 10 001 0 0 0 1 [01] 0
Barcella odora 00 001 1200 ? 0 0
Elaeis guineensis 00 001 1200 1 0 0
Acrocomia aculeata 00 001 1100 0 0 0
Bactris gasipaes 00 001 1100 1 0 0
Cocos nucifera 00 001 1000
0
00
Attalea allenii 00 011 1000 1 0 0
Syagrus smithii 00 011 1010 0 [01] 0
Lytocaryum weddellianum 00 101 1011 0 [01] 0
Tripylocarpa aestuaria 00 111 0 00 0 1 0
M.K. Futey et al.
Age and stratigraphy: Danian, Early Paleocene, Salamanca Formation.
Etymology: Tripylocarpa refers to the three valves characteristic of the fruits (from
the Greek trion+pylo+karpus); the specific epithet aestuaria refers to environment
where the fossils were deposited.
Description: The fossils are petrified ovoid, drupaceous fruits, 17.123.2 mm long
and 14.318.4 mm wide (Fig. 2a). The fruits have 3 main longitudinal grooves that
delimit equal areas of exo/mesocarp, and extend from the apical pore to the most
basal portion of the fruit (Fig. 2b). The three equal areas are interpreted as the fruit
valves. A germination pore is clearly positioned on the apical area of the fruits and it
is formed where the three valves reach each other (Fig. 3b & c). The fruits probably
had a basal peduncle based on the presence of a constricted area opposite to the
germinal pore that is interpreted here as part of the peduncle (Fig. 2e). The exocarp
and mesocarp are fused and anatomically indistinguishable one from another
(Fig. 3a). The exo/mesocarp is 1.53.0 mm wide with no distinct layers (Fig. 2f);
while the endocarp is thin and unsculptured (Fig. 2h). Sclerenchyma bands with
associated brachysclerids run longitudinally through the mesocarp and a layer of
tanniferous cells internal to the endocar p (Fig. 3a). The stigmatic remains are apical
(Fig. 2g). Each fruit carried a single seed; each seed is 1016 mm long and 1014 mm
wide (Fig. 2h); they are deltoid in shape with an acute to acuminate apex, and basally
truncate and they have a hilum that is basal (Fig. 2i). These correspond externally to
the germination pore and basal peduncle respectively. The endosperm is ruminate
(Fig. 3d).
Comments: Cocoseae fruits are more commonly preserved as endocarps and can
often be easily identified due to the 3 pores and occasional ridges in the endocarp.
While these fossils lack certain Cocoseae endocarp features, the exquisite preserva-
tion allows the observation of several key characters linking these fossils with
Table 3 Morphological character list and characters coded
Fruit surface, Exo-/mesocarp
0. Stigmatic remains: 0) apical to sub-apical; 1) lateral to basal [6, 15].
1. Pericarp scales: 0) absent; 1) present [6].
2. Pericarp warts: 0) absent; 1) present [6].
3. Longitudinal grooves: 0) absent; 1) present [6, 20, 32].
4. Mesocarp furrows: 0) absent; 1) present [6, 20].
5. Fibers in mesocarp: 0) absent; 1) present [6, 17].
Endocarp
6. Endocarp sculpturing : 0) smooth; 1) sculptured [6, 20].
7. Endocarp pores: 0) absent; 1) present [6].
8. Endocarp pore position: 0) basal; 1) lateral; 2) apical [6, 15].
9. Endocarp grooves: 0) absent; 1) present [15].
Seed
10. Hilum position: 0) basal; 1) lateral [6].
11. Raphe: 0) absent; 1) present [6, 15].
12. Endosperm: 0) homogenous; 1) ruminate [6, 20, 32].
13. Number of seeds: 0) one; 1) more than one [6].
Arecaceae fossil fruits from Argentina
Cocoseae. The fruits externally display several unique characters within Arecaceae,
such as the furrowed surface and the three striations trisecting the fruit into equal
Fig. 2 ab, ei: Tripylocarpa aestuaria Gandolfo and Futey. cd. Lytocaryum weddellianum (H. Wendl.)
Toledo. ab. Tripylocarpa aestuaria. a. MPEF-Pb 3768- Side view of the fossil fruit showing a suture
(arrow). b. MPEF- Pb- 3766- Top view of the holotype. Note the three longitudinal sutures (arrows)
delimiting the three valves. The three sutures culminate at the germination pore. cd. Lyt ocaryum
weddellianum- BH #37247. Side view showing one suture (arrow). d. Top view showing 3 longitudinal
sutures where the epicarp and mesocarp will split (arrows) and the germination pore. ei. Tripylocarpa
aestuaria. e. MPEF-Pb - 3769. Bas al view sho wing peduncle remains (arrow ). f. MPEF-Pb- 3826-
Longitudinal section showing pericarp and seed cavity (locule). Note that there are no differentiation
between the epicarp and mesocarp. g. MPEF-Pb- 3768- Top view showing the stigmatic remains (arrow)
and germination pore. h. MPEF- Pb- 3827- Single seed surrounded by remains of pericarp, note the basal
hillum. i. MPEF- Pb 3828- Deltoid and basally truncate seed, note the acute apex. All scale bars0 1cm
M.K. Futey et al.
areas. In longitudinal and transverse secti ons, it is possible to observe the central
position of the unique seed and some anatomical characters of the exo, meso and
endocarp. It is clear that there is no distinction between the exo and mesocarp,
therefore we are interpreting them as fused (Fig. 3a). The sections also reveal the
presence of parallel bands of longitudinal fibers in the mesocarp, along with the
apical position of the germination pore (Fig. 3b &c)
Berry (1926) describes palm endocarps from the Middle Eocene of Peru that he
compares with two modern genera of the tribe Cocoseae (Astrocaryum and Attalea)
based on the size, the presence of longitudinal fiberous bands and overall appearance.
Gómez-Navarro et al. (2009) and Tripathi et al. (1999) both describe Cocos fruits
from the Paleocene of Colombia and Tertiary of Madhya Pradesh, India. While in
both cases the endocarp pores are not preserved, the large size and longitudinal ridges
on the fruit surface clearly indicate affinity with modern Cocos. The fruits described
herein are clearly distinguishable from previous Cocoseae fossils based on the much
smaller size and distinguishing characters found in these fossil fruits.
Phylogenetic Analysis
The heuristic search produced 21 most parsimonious trees with 689 steps with a
consistency index (CI) of 76 and retention index (RI) of 64. The strict consensus
resulted in the collapse of 11 nodes and 749 steps with a CI of 69 and a RI of 51. In all
Fig. 3 ad: Tripylocarpa aestuaria Gandolfo and Futey. a- MPEF-Pb 3767a- Transversal section showing
the fused exocarp and mesocarp layers and fiberous bundles (arrows). b. MPEF-Pb 3767b- Longitudinal
section through the germination pore, note the channel left at the pericarp. cd. MPEF-Pb- 3768- c. Still
image from CT scan showing the germination pore (arrow), the pericarp, the single locule and remains of
the seed. The white area is interpreted as the thin endocarp (arrow). d. Still image from CT scan showing
endosperm ruminations (arrows). Scale bar figs. a and b0 1 mm.; c and d0 2.3 mm
Arecaceae fossil fruits from Argentina
trees Tripylocarpa aestuaria is placed in the Attaleinae subtribe with 65 % support
(Fig. 4)
Discussion
The most recent taxonomic classification of palms based on molecular data, primarily
from the chloroplast genome, resulted in the recognition of five subfamilies:
Calamoideae, Coryphoideae, Nypoideae, Ceroxyloideae and Arecoideae, with the
previously recognized subfamily P hytelephantoideae included in Ceroxyloideae
(Asmussen et al., 2006). Inclusion of Tripylocarpa within Calamoideae and
Nypoideae can easily be ruled out based on gross morphological characters; fruits
of the Calamoideae have distinctive imbricate scales while the monotypic Nypoideae
fruits are laterally compressed and irregularly angled (Dransfield et al., 2008),
characters never observed in Tripylocarpa. Fruits of the remaining 3 subfamilies,
Coryphoideae, Ceroxyloideae and Arecoideae, are typically one-seeded and not as
strikingly distinct as those of the Calamoideae and Nypoideae. Longitudinal fibrous
bands without bundle sheaths in the mesocarp are found throughout the Arecoideae
and occasionally in Coryphoideae, but are lacking in fruits of the Ceroxyloideae
(Essig, 1999). Additionally, all seeds of Ceroxyloideae have a homogenous endo-
sperm (Dransfield et al., 2008). Fruits of Coryphoideae tend to be globose and often
have a corky periderm, unlike the ovoid, smooth skinned Tripylocarpa.
The results of our phylogenetic analysis clearly place Tripylocarpa within the
subfamily Arecoideae, tribe Cocoseae, subtribe Attaleinae. Although Tripylocarpa
lacks the characteristic 3 or more well developed endocarp pores typical of tribe
Cocoseae, the suite of anatomical and morphological characters point to an affinity
with Cocoseae. The fossils are morphologically similar to fruits of Cocoseae, sharing
Fig. 4 Strict consensus tree of 21
most parsimonious trees 689
steps long from combined
morphological and molecular
dataset as analyzed using NONA.
CI0 69, RI0 51. Numbers on
branches indicate bootstrap values
M.K. Futey et al.
characters such as ovoid to obovoid shape, apical stigmatic remains and, excluding
Cocos, a relatively small size. The fibrous bands running parallel and longitudinally
through the mesocarp is also a typical character of Cocoseae fruits. While many
arecoid fruits have a smooth epicarp surface, some members of the Cocoseae subtribe
Attaleinae, such as Attalea, Syagrus and Lytocaryum, have furrows or striations in the
epicarp similar to those found in Tripylocarpa.
Specimens of Tripylocarpa displ ay a ch aracter found only in th e genus
Lytocaryum, a member of the subtribe Attalinae. Lytocaryum consists of 4 species
(L. itap ebiensis, L. hohnei, L. insigne and L. weddellianum) restricted to southeast
Brazil (Noblick & Lorenzi, 2010) and has at times been placed in the larger, closely
related genus Syagrus (Glassman, 1987). All 4 species have the 3 distinctive longi-
tudinal markings running from apex to base on the epicarp which are strikingly
similar to those found on these fossils (Fig. 2c & d). In L. hohnei, L. insigne and L.
weddellianum, the exo/mesocarp dehisces at maturity along the grooves exposing the
endocarp and releasing the seed. The shared presence of this unique character within
Arecaceae is strong evidence for the affinity of these fru its t o Lytocaryum.
Additionally, Tripylocarpa has ruminate endosperm, which is also produced in L.
hohnei (Noblick & Lorenzi, 2010).
While the Patagonian palm fossil record is extensive, the types of fossil remains
found lack diagnosti c characters that allow for precise taxonomic placement. In the
case of both pollen and stems designation below the subfamily level is difficult.
Seven speci es of Palmoxylon that have been described from Patagonia have been
suggested to have an affinity to Sabaloid, Coryphoid, Phoenicoid, Bactroideae and
Cocoseae palms based primarily on anatomical characters (Romero, 1968; Arguijo,
1979, 1981; Ancibor, 1995; Ottone, 2007), of those only three species come from
Chubut (Gandolfo et al., 20 10). Romero (1968)describedfossilstemremains
collected at the Cerro Abigarra do locality, also from the Salamanca Fm. (Chubut),
and erected the species Palmoxylon patagonicum. This species shows intermediate
characters with the tribes Sabaleae and Cocoseae, nevertheless the characters pre-
served do not allow a more precise taxonomical placement. Two other Palmoxylon
species of Danian age are described for the Cerro Bororó Fm that outcrops in Chubut,
P. bororoense and P. vaterum (Arguijo, 1979, 1981). Both species show affinities to
the tribes Sabaleae and Cocoseae, but as with P. patagonicum, they cannot be placed
with confidence in any one particular tribe. Although, at this time it is impossible to
discuss potential relationships among the Palmoxylon species and Tripylocarpa, it is
remarkable that the three Patagonian Palmoxylon species show potential affinities
with Cocoseae as does Tripylocarpa. Therefore, the finding of Tripylocarpa provides
unequivocal evidence of the presence of this tribe in Patagonia.
Although palm pollen is relat ively variable when compared to other monocots, its
taxono
mic
utility is limited when applied to the fossil record due to the tendency of
morphologically distinct pollen types to be limited in phylogenetic range (Harley &
Baker, 2001). While several species of palm pollen have been de scribed from
Patagonia, only in the case of Spinozonocolpites has any suggested affinities below
the subfamily level affinities beenascribed,inthiscasetomodernNypa
(Archangelsky, 1973).
Whether Tripylocarpa has any relationship with previously described palm macro-
and microfossils cannot be made or ruled out due to the lack of taxonomic resolution.
Arecaceae fossil fruits from Argentina
Outside of Patagonia records of Cocoseae fruits, and more commonly endo-
carps, are reported from several localities. Coconut-like endocarps from both
the North and South islands of New Zealand were found in deposits dated to
mid-Eocene, Miocene and Pliocene (Ballance et al., 1981; Berry, 1926; Campbell
et al., 2000). Fliche (1896) describes a coconut-like endocarp and fruit from the
Cenomanian of northern France; which he compares to Astrocaryum and Syagrus
based on size and shape of the fossil. Hollick (1928) and Maury (1930) described
endocarps a ssigned to Palmocarpon from the Oligocene of Puerto Rico and the
Maastrichtian of Brazil that they both compared with members of Cocoseae.
However the poor preservation of all these fossils makes any assignment, even to
Arecaceae, dubious. Comparisons among these species and Tripylocarpa are impos-
sible to establish at this time due to the poor preservation of the previously described
fossils and the lack of their reliable taxonomic placement; consequently, Tripylocarpa
are the oldest and unequi vocal record for fruits belonging to the subtribe Attaleinae
and tribe Cocoseae.
In contrast to the current drier conditions found in Chubut today, several lines of
evidence from the Salamanca Fm point to a moister, sub-tropical environment during
the Paleocene. Many taxa found in the paleoflora, such as Akania, Malvaceae, and
Lauraceae leaves and Araucariaceae cone scales (Berry, 1937; Iglesias, 2007) are
associated with thermophillic families. Indeed the presence of palms alone is highly
indicative of higher moisture and temperature levels as modern palm distribution is
heavily influenced by water availability in addition to warm temperatures and
therefore palms are rarely found outside of 34° latitude (Bjorholm et al., 2005 ).
These suggested paleoclimatic condition s are also supported by leaf-margin and
leaf-area analyses on the fossils from two localities in the Salamanca Fm, that
predicted a mean annual temperature of 14.1 °C±2.6 °C and a mean annual precip-
itation of at least 115 cm (+50/35 cm) (Iglesias et al., 2007).
Conclusions
There is no doubt that the fossil fruits herein described belong to the Subtribe
Attaleinae, Tribe Cocoseae, Subfamily Arecoideae. Tripylocarpa aestuaria taxonom-
ic placement is based on the combination of morphological characters preserved as
well as the results of the inclusion of thefossilwithinaphylogeneticcontext.
Remarkably, to our knowledge, Tripylocarpa aestuaria is the first palm fossil taxon
to be included in an analysis that includes molecular and morphological data. The
detailed preservation of these fossils, which allows the observation of descriptive
characters and its phylogenetic placement, makes it a prime candidate for use as a
calibration point fossil in any future molecular clock analys es involving Arecaceae,
the tribe Cocoseae or the subtribe Attaleinae.
This addition to the fossil record of Patagonia expands our know ledge of the
Salamanca paleoflora and is further evidence of a previous subtropical environment
in the Paleocene of Patagonia. Further work on any undescribed palm leaf, stem and
endocarp fossils from Patagonia would only help to resolve the taxonomic affinities
M.K. Futey et al.
of previously described fossil materials, especially stems, and will illuminate our
understanding of the evolutionary history of the palms.
Acknowledgments The authors are grateful to Drs. S. Archanglesky and R. Somoza for making available
the fossil remains studied herein. We thank Ing. M. Riccio for the CT Scan images, Dr. J. Reveal for helping
with Greek for naming of the fossil, J. Svitko and P. Balcells for figure preparation. This research was
supported by NSF grants DEB 0830020, DEB- 0918932 and DEB 0919071, and the Fulbright Foundation
(MAG).
Literature Cited
Ancibor, E. 1995. Palmeras fósiles del Cretácico Tardío de la Patagonia, Argentina (Bajo de Santa Rosa,
Río Negro). Ameghiniana 32: 287299.
Archangelsky, S. 1973. Palinología del Paleoceno de Chubut. I. Descripciones sistemáticas. Ameghiniana
10: 339399.
Argujio, M. H. 1979. Palmoxylon bororoense n. sp. de la Formación Cerro Bororó (Paleoceno), provincia
de Chubut, República Argentina. Physis C. 38: 8796.
——— 1981. Palmoxylon vaterum n. sp. del Paleoceno (Daniano) de la provincia del Chubut, Argentina.
Physis Sección C. 39: 4959.
Asmussen, C. B., J. Dransfield, V. Deickmann, A. S. Barfod, J. Pintaud & W. J. Baker. 2006. A new
subfamily classification of the palm family (Arecaceae): evidence from plastid DNA phylogeny.
Botanical Journal of the Linnean Society 151: 1538.
Baker, W. J., V. Savolainen, C. B. Asmussen-Lange, M. W. Chase, J. Dransfield, F. Forest, M. M.
Harley, N. W. Uhl & M. Wilkinson. 2009. Complete generic- level phylogenetic analyses of
palms (Arecaceae ) with comparisons of supertree and supermatrix approa ches. Syste matic
Biology 58: 240256.
Ballance, P. F., M. R. Gregory & G. W. Gibson. 1981. Coconuts in Miocene turbidites in New Zealand:
possible evidence for tsunami origin of some turbidity currents. Geology 9: 592595.
Berry, E. W. 1914. The Upper Cretaceous and Eocene floras of South Carolina, Georgia. U. S.
Geological Survey, Professional Paper. 84: 1200.
——— 1916. A petrified palm from the Cretaceous of New Jersey. American Journal of Science 41: 193197.
——— 1926. Cocos and Phymatocaryon in the Pilocene of New Zealand. American Journal of Science
212: 181184.
——— 1937. A Paleocene flora from Patagonia. Johns Hopkins University Studies in Geology 12:
3350.
Bjorholm, S., J. Svenning, F. Skov & H. Balslev. 2005. E nvironment al and spacial cont rols of palm
(Arecaceae) species richness across the Americas. Global Ecology and Biogeography 14:
423429.
Campbell, J. D., R. E. Fordyce, A. Grebneff & P. A. Maxwell. 2000. Fossil coconuts from mid-Cenozoic
shallow-marine sediments in southern New Zealand. Otago University, Geology Department, Dunedin
[Poster abstract].
Crié, L. 1892. Recherches sur les Palmiers silicifiés des terains Crétacés de lAnjou. Bulletin de la Socíeté
dÉtudes Scientifiques dAngers 21: 97103.
Daghlian, C. P. 1981. A review of the fossil record of monocotyledons. The Botanical Review 47: 517
555.
Dransfield, J., N. W. Uhl, C. B. Asmussen, W. J. Baker, M. M. Harley & C. E. Lewis. 2008. Genera
Palmarum. Kew Publishing, Kew.
Edgar, R. C. 2004. MUSCLE: a multiple sequence alignment method with reduced time and space
complexity. BMC Bioinformatics 5: 113.
Essig, F. B. 1999. Trends of specialization in the palm pericarp. In: A. Henderson & F. Borchsenius (eds).
Evolution, variation and classification of palms. The New York Botanical Garden Press, Bronx.
Arecaceae fossil fruits from Argentina
Fliche, P. 1896. Etudes sur la flore fossile de lArgonne Albian-Cenomanien. Bulletin de la Societe des
Sciences de Nancy. 14: 114306.
Gandolfo, M.A., M.C. Zamaloa & N. Caviglia. 2010. Revisiting the Argentinean monocotyledon
fossil record. VI Southern Connection Congress, San Carlos de Bariloche, Argentina. [Symposium
abstract].
Glassman, S. F. 1987. Revisions of the palm genus Syagrus Mart. and other selected genera in the Cocos
alliance. Illinois Biological Monographs 56: 1230.
Goloboff, P. A. 1998. NONA. Computer program and software. Published by the author, Tucuman.
———, J. A. Farris & K. C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24:
774786.
Gomez-Navarro, C., C. Jaramillo, F. Herrera, S. L. Wing & R. Callejas. 2009. Palms (Arecaceae )
from a Paleocene rainforest of northern Colombia . American Jou rnal of Botany 96: 1300
1312.
Harley, M. M. 2006. A summary of fossil record for Arecaceae. Botanical Journal of the Linnean Society
151: 3967.
——— & W. J. Baker. 2001. Pollen aperture morphology in Arecaceae: application within phylogenetic
analyses, and a summary of the fossil record of palm-like pollen. Grana 40: 4577.
Hollick, A. 1928. Paleobotany of Porto Rico. Scientific survey of Porto Rico and the virgin islands. New
York Academy of Sciences 7: 177393.
Iglesias, A., P. Wilf, K. R. Johnson, A. B. Zamuner, N. R. Cúneo, S. D. Matheos & B. S. Singer. 2007.
A Paleocene lowland macroflora from Patagonia reveals significantly greater richness than North
American analogs. Geology 35: 947950.
Lutz, A. I. 1980. Palmoxylon concordiense n. sp. del Plioceno de la provincia de Entre Ríos, República
Argentina. Actas del II Congreso Argentino de Paleontología y Biostratigrafía y I Congreso Latin-
oamericano de Paleontología 3: 129140.
——— 1984. Palmoxylon yuqueriense n. sp. del Plioceno de la provincia de Entre Ríos, República
Argentina. Actas del II Congreso Argentino de Paleontología y Biostratigrafía: 197207.
Maury, C. J. 1930. O Cretáceo da Parahyba do Norte. Serviço Geológico e Mineralógico do Brasil, Rio de
Janeiro. Monographias 8: 1305.
Nixon, K. C. 1999. WinClada version 1.87. Published by the author. Ithaca, NY. [Available at http://
www.winclada.com].
Noblick, L. R. & H. Lorenzi. 2010. Lytocaryum, including a new species from Bahia, Brazil. Journal of
the International Palm Society 54: 517.
Ottone, E. 2007. A new palm trunk from the Upper Cre taceous of Argentina. Ameghiniana 44:
719725.
Pan, A. D., B. F. Jacobs, J. Dransfield & W. J. Baker. 2006. The fossil history of palms (Arecaceae) in
Africa and new records from the Late Oligocene (28-27 Mya) of north-western Ethiopia. Botanical
Journal of the Linnaean Society 151: 6981.
Petriella, B. 1972. Estudio de maderas petrificadas del Terciario Inferior del area Central de Chubut (Cerro
Bororo). Revista del Museo de La Plata 6: 159262.
Romero, E. J. 1968. Palmoxylon patagonicum n.
sp.
del Terciario inferior de la Provincia de Chubut,
Argentina. Ameghiniana 10: 417432.
Schenk, A. 1882. Die von den Gebr üdern Schlage nwelt in Indein gesammelten fossilen Hölzer.
Botanischer Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 3: 353358.
Somoza, R., G. Cladera & S. Archangelsky. 1995. Una nueva tafoflora paleocena de Chubut, Patagonia.
Su edad y ambiente de depositación. VI Congreso Argentino de Paleontología y Biostratigrafía, Actas:
265269.
Tripathi, R. P., S. N. Mishra & B. D. Sharma. 1999. Cocos nucifera like petrified fruit from the Tertiary
of Amarkantak, MP, India. Paleobotanist 48: 251255.
Vaudois-Miéja, N. & A. Lejal-Nicol. 1987. Paléocarpologie africaine: apparition dès lAptien en Égypte
dun Palmier (Hyphaeneocarpon aegyptiacum n.sp.). Comptes Rendus de lacadémie des Sciences
304: 233238.
Zucol, A. F., M. Brea, E. Bellosi, A. A. Carlini & G. Vucetich. 2007. Preliminary phytolith analysis of
Sarmiento Formation in the Gran Barranca (central Patagonia, Argentina). Pp 189195. In: M. Madella
& D. Zurro (eds). Plants, people and places: recent studies in phytolith analysis. Oxbow Books,
Oxford.
Zuloaga, F. O. & O. Morrone (eds.) 1996. Catálogo de las plantas vasculares de la República Argentina. I.
Pteridophyta, Gymnospermae y Angiospermae (Monocotyledoneae). Monographs in Systematic Botany
60: 1323.
M.K. Futey et al.
Calectasia ----------------------------------------- --
---- ----- --- --------------------------------------
----------------------------------------------------
----------------------------------------------------
----------------------------------------------------
----------------------------------------------------
----------------------------------------------
-------------------------0023233220333000213223233000203301000332
0133033030111132013012300110002030132030313322102103311202300
13113100111220233112113200200210220 2102 122302 132 1120031331301322301
0322010013232322013203220133011023133203123301000220120321301101031
2022132332332222022000031003030332133032302133031133302011333332022
00223313233011001032333013311033232223003230333223 33100021113012021
3130123132202203132120033111113313303311000013331100221112111103221
03110023320012 0203 0023 320010023 0322 31231111303322203230 1303 30001100
0013222033031121000200330322302021223330320032313012322322013320333
3011002203203200001232001310100110333032123322020201123331330333323
2112002100333030002101022112000122232000310002201033013320032130112
102230103232002000320300000222110303332110202003302202 3311303123003
2103201301330013223220331013212003013021332213333303321121201003221
1301331331010331031212100 321032102330 33203020102 00000310322 3032103
3331232301302130002103301 230323132232 20203103033 10121322301 0230230
2230001322002222021202020 320133322233 33233203330 33012320320 3333033
20000020112002312122303333133301310020332223 1313 032111223233332111 2
32213312222223033103 2333 22103 03211321113201120003133322020320331123
0130102333221220220013330220101113322223003210113223210210213003123
2302133302002123232301002 131230032022 20123203133 21312320022 3003200
0330311212002130110003220231132021302112132133232002303220022120310
00331200331200110230203002130203- - - ------------------------------
-------------------------------------???????????????????????????
Kingia --------------------- --------------------- --------------
------- --------------------- --------------------- ------------
--------- --------------------- --------------------- ----------
----------- --------------------- --------------------- --------
------------- --------------------- --------------------- ------
--------------- --------------------- --------------------- ----
----------------- --------------------- --------------------- --
------------------- ---------------------0033201330330301311
3201301200011000203013203031332210210331120230013113100111220
2331121132022002102222102122302132112003133130132230103220100132323
220132032201330110231332031233010002201203213011010312 0001123323122
2202200003100303033213303230213303113330201133333200200223313233013
0010323330133110332322230032303332233310002111301202131301231322022
Appendix 1. Combined Molecular and Morphological Matrix
Arecaceae fossil fruits from Argentina
0313212003311101331330 33 110000133311002231121131032210311002 33200 02
020300233200100230322312311313033222032301303300011000 0332220330311
2100020013012230202122333032003231301212232201332033330110022032032
000012320013101001103330321233220202011233313303333232 1120021003330
3000212102213200012223200031000220103301332003213011210223010323200
2000320300000222112303332110202003322202131130312300321032013013300
1322222033101321000301302133231300330332112120300322113013313310103
3103121210032103210233033 203020102000 00310322303 21033331232 3013021
3000210330121032313223220 203103033101 21322301023 02302230001 3220022
2200123202032013330223333233203330330123203203333033200000201120223
1212230333333310131002033222313130321132232330321112322133102222230
331032333221030321132111320112000 3133 322020320 331123013010233322122
0220013330220101113322220003210113223210210210003121232213330200212
3232301002131230032022201232031332131232002230032000330311232002130
23000322021 1 132021- - - ------------------------------------------------
---------------------------------------???????????????????????????
Dasypogon -----------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 3010003320133033030
13113201301200011000203013203031332210210331120230013113100111220233
11211320220021022221021223021321120031331301322301032201001323232201
32032201330110231332031233010002201203213011010312000112332312202022
00003100303033213303230213303113330201133333200200223313233013001032
33301331103323222300323033322333100021113012021313012313220220313212
00331110133133033110000133311002211121131032230311002332000202030023
32001002303223123113130332220323013033000110000332220330311210002001
30122302021223330320032313012122322013320333301100220320320000123200
13101001103330321233220202011233313303333232112002101333330002121022
11200012223200031000220103301332003213011210223010323200200032030000
02221103033321102020033022023311303123003210320130133001322322033101
32100030130233322131033033211212010032211301331331010331031232100321
03210233033203020102000003103223032103333123230130213000210330123032
31322322020310303310121122301023023022300013220022220012320203201333
22233332332033303321232032033330332000002011202231212230333313310131
00203322231313032113223233132111232213310222223033103233322103032113
21113201120003133322020320331123013010233322122022001333022010111332
22200032101132232102102100031212322133302002123232301002131230032022
201232031332131232002230032000330311232002130210003220211132021- - - - - -
-------------------------------------------------------------------
-------------------------------------------------------------------
- - - - - - - - - - - - - - - - - ???????????????????????????
M.K. Futey et al.
Mauritia 001000-0001000- ------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
- - - - - - - - - - - - - - - - - - - - -300021002323322033300021322323300020330100
033201330330301311320130120001100020301320303133221021 0331120230013
1131001112202331121132022002102222102102302132112003133130132230103
2201001323232201320322013301102313320312330100022012032130110103120
0011233231222202200003100303033213303230213303113330201133333200200
2233132330130010323330133110332322230032303332233310002111300202131
301231322022 03 132 12 003 31111133133033110000133311002231 0211310322103
1100233200020203002332001002303223123113130332220323013033002110000
3322203303112100020013012230202122333032003231301212232201332033330
1100220320320000123200131010011033303212332202020112333133033332321
1200212133303000212102211200012223200031000220103301332003213013210
223010323200200032031000022211230333211020200332220233 1130312300321
0320130133001322222033101321000301302133221310330332112120300122113
0133133101031103121210032103210233033203020102000003103223032103333
1232301302130002103301230 323132232202 03103033101 22022301023 0232223
0001322002222001232020320 133322233332 33203330330 12320320333 3033200
00020112002312122303133333301310020332223131 3032 110223233031111232 2
13310222223033103233 3221 03032 11321113201120003133322220320331123013
0102333221220220011330220101113322220003210111223210230213003122232
2133302002123232301002131 230032022201 23203133213 12320022300 3200033
03112320021302100032 2021113 2001302 132112133232002303220002020310003
31202333200110230203002132203- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
--------------------------------------???????????????????????????
Trithrinax 000000-0001000- -----------------------------------------
--------------------------------------------------------------
------- ------------------------------------------------------
----- --------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
-----------------100010202013000210023213220333000213223233000203
301000332003303303013113201301200011000203013203031332 2102103311202
30013113100111220233112113202200210 2222 102122 302 132 1120031331301322
3010322010013232322013203220133011023133203123301000220120321301101
03120001123323122220220000310030303321330323021 33031133302011333332
0020022331323301300103233301331103323222300323033322333100021113012
021313012313220 220 313 212 0033111013313303311000013331100223112113103
2210311002332000202030023320010023032231231131303322203230130330001
1000033222033031121000200130122302021223330320032313012122322013320
Arecaceae fossil fruits from Argentina
3333011002203203200001232001310100110333032123322020201123331330333
3232112002101333030002121022112000122232000310002201033013320032130
1321222301032320020003203200002221323032321102020033222023311303123
0032103201301330013222220331013210003013021332213103303321121203001
2211301331331010311031212100321032102330332030201020000031032230321
0333312323013021300021033 012303231322 32202031030 33101210223 0102302
3222300013220022220012320 203201333222 33332332033 30330123203 2033330
33200000201120023121223031333333013100203322231 31303211022323313211
12322133102222230331032333221030321132111320112000313332222032033112
301301023332212202200133302201011133222200032101112232102302130031222
32213330200212323230100213123003202220123203133213123200223003200033
03112320021302100032202111320013021321121332320023032200210203100033
1200331200110230203002132203- - - - - - - - - ?????????????????????????
Sabal 0100-0001(01)00- - - - -123222301023131200021031000213021031200213
120002113201333203213303230223032310113321101- - - - - - - - - - - - - - - - - - - - -
0020033221020210130033323 322303003030 20210330310 33022232033 3213030
1030230031301331303031302100032133103000- - - - - - - - - - - - - - - - -
3200130011300100010002103112021023002310003003210 10000103- - - - -
01322013100003- - - - - -030131330133103132100- - - -33333233203011-
3333333300200002000202221202222220222322211210113203330332000310320
20100330203032132023131032331331020303102003031103-113331331021013-
13322033013201021332 3020 33213 3111 3322 1020111210200210030321 32032112
3003120023333011201010033003311322320100320002000223213202023312003
2232032000200222232002301331002010233301131331201200221311013233310
32203311132 32220 2200213 0021321 31030 - - - - - ??? ????? ??????? - - - - - - -
1101000102000130002100232132203330002132232330002033010003320133033
030131132013012000110002030 1320 303 133 221021 033112023001311310011122
0233112113202200210222210212230213211200313313013223010322010013232
322013203220133011023133203123301000220120321301101031 2000112332312
2220220000310030303321330323021330311333020113333320020022331323301
3001032333013311033232223003230333223331000211130120213130123132202
2031321200 33 1110133133 033110000133311002211121131032210311002332000
2020300233200100230322312311313033222032301303300011000033222033031
1210002001301223020212233303200323130121223220133203333011002203203
200001232001310100110333032123322020201123331330333323 2112002101333
3300021210221120001222320003100022010330133200321301321222301032320
0200032032000022211030333211020200332220213113031230032103201301330
0132222203310132100030130213322131033033211212030012211301331331010
3110312121003210321023303320302010200000310322303210333312323013021
3000210330123032313223220 203103033101 21022301023 02322230001 3220022
2200123202032013332223333233203330330123203203333033200000201120023
1212230313333330131002033222313130321102232331321112322133102222230
331032333221030321132111320112000 3133 322220320 331123013010233322122
022001333022010111332222 000321 0111223 2102302 1300312223221 3330200 212
3232301002131230032022201232031332131232002230032000330311232002130
210003220211132001302132 1121332320023032200221203100033120033120011
M.K. Futey et al.
0230203002132203- ----------------------???????????????????????????
Nypa 0100-0001110- - - - -------23332200021031000213021031200213120002
11320133320321330323322303231- - - - - - - - - - - - - - - - - - - - - - - - - - - - -
03010230031301330303031322100032133103000- - - - - - - -------------320
0130011- - - - - - - - - - - - - 03330-320 0030 0320201 0033020 30- - - - - - -
023131032331331020302102003231103-313331333021013-
133220230132010213323020332132 111332210201110102002100303 2132032 132
3003120223333011201010033003311322120100320032200223213202023312003
22320320002002222320023013311031102333011313332032002213110132333- - -
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -???????????????- - - - -
1101000102000130002100232132203330002132232330002033010003320133033
030131132013012000110002030 1320 303 133 221021 033112023001311310011122
0233112113202200210222210212230213211200313313013223010322010013232
322013203222133011023133203123301000220120321301101031 2000112332312
2220220000310030303321330323021330311333020113333320020022331323301
3001032333013311033232223003230333223331000211130120213130123132202
2031321200311111133133033110000133311002211101131032210311002332000
2020300233200100230322312311313033222032301303300011000033222033031
1210002001301223022212233303200323130121223220133203333011002203203
200001232001310100110333032123322020201123331330333323 0112002100333
0300021210201120001222320003100022010330133200321301121222301032320
0200032032000022211030333211020200332220233113031030032103201301330
0132222203310132100030130213322131033033211212030012211301331331010
3110312121003210321023303320302010200000310322303210333312323013021
3000210330123032313223220 203103033101 22022301023 02322230001 3220022
2200123202032013332223333233203330330123203203333033200000201120023
1212230313333330131002033222313130321102232330301112322133102222230
331032333221030321132111320112000 3133 322220320 331123033010233322122
02200133302201011133222200032101112232102302130031222322133302002123
23230100213123003202220123203133213123200223003200033031123200213021
00032202111320013021321121332320023032200220203100033120133120011023
0203002132203- - - - - - - - - - - - - - - - - - - - - - -???????????????????????????
Ceroxylon 0100-0001(01)00- - - - - - - - - - - - - - - - - - - - - - - - - - - - -200021302
01333223213303233223032310133321101021211302-332330230
1332002003122102 021 0130 033 3233223 0300 303 0202 100323 1033 023 2331 03321-
32003030230030301331203031302100032133103000- - - - - - - -
000013001130013001300210313203120300231000300321010000103------
11322013000003- - - - - - -030030330133103232100- - ----0333333223011-
33333310000200010- - --------222030032020100330
2030323320231 31 032 331 33102 030 310 20 03031103313313330210 13 -13322 033 0
132010203323020332132111 332210 2011101 0200210 030321 32032132300312 002
3333011201012033003311320220100320002000223213202023312233223203200
0200222232102101331203110233- - - - - - - - - - - - - - ???????????????- - - -231
011010001020001300021002323322033300021322323300020330100033201
330330301311320130120001100020301320303133221021033112023001311310
Arecaceae fossil fruits from Argentina
011122023311211320220021022221021223021321120031331301322301032201
0013232322013 203220 133011023133203123301 000220 120321 30110103120001
1233231222202 200003 100303 03321 330323 021330311333020113333320020022
33132330130 0103 23330 13311033232223003 23033 3223 33100 02111 3012 02131 3
012313220220313212003311111331330331100001333110022111211310322103
1100233200020203002332001002 3032231231131303322203230130330001100
00332220330311210002001301223020212233303200323130121223220133203
333011002203203200001 2320013 10100 11033303212332202 020112333133033 3
32321120021003330300021210 2211200012223200031000 22010 330133200 3213
01321222301 0321 20020 00320 3100 002221123033321102020033222023 3113031
23003210320130133 00132222 20331013 2100030 13021332 21310330332 1121203
001221130133133101031103121210032103210233 033203 02010 200000310 3223
0321033331232301302130002 103301230323 13223220203 10303310121 2223010
2302322230001322002222001 232020320133 32223333233 20333033012 3203203
3330332000002 0112002312122303133333 30131 002033 222313 13032110223233
030111232213310222223033103233322103032113211132011200031333222203
20331123013010233322122022001333022010111132222000321011122321023
0213003122232213330200212 323230100213 12300320022 01232031332 1312320
022300320003303112320021302100032202111320013021321121332320023332
2002202031000331200331200110230203002132203- - - - - - - - - - - - - - - - -
------------------------------------------------
--???????????????????????????
Pseudophoenix 0100-0001100- - - - --------------------------20002131
201333203213303233223032310- - - - - - - - - - - - - - - - - - - - - - - - - - -----------
- - - 0332232332031213200103023003020133300303133210003210310320000000
00000003201030110001000310313203100- - - - ------------33332333223011-
33333130020200020- --------------3322022021110113 2033003320203
003202013223020303233202313103233133102030310203303110- - -31333133
(03)021013-13322033013201021332302133223211113221020113010200
2102303213203211230031200233330113010100330233113203201103200022002
232132020233120332232032000200222232002301331003110233- - ----------
- - - - - - - - - - ???????????????- - - - - -1101000102000130002100232332203330002
1322123300020330100033201330330301311320130120001100020301320303133
2210210331120230013113100111220 23 3112111202200210222210 2122 30 21 32 11
2003133130132230103220100132323220132032201330110231332031233010002
2012032130110103120111123323122220220000310030303321330 3230 21330 311
3330201133333200200223313233013001032333013311033232223003230333223
331000211130 120 213130 123 132 2022 031321 200 331110133133033110000133311
002211121131032210311002332000202 0300 233200100 2303 22312 311313033222
0323013033000110000332220330311210002001301223020212233303200323130
1212232201332033330110022032032000012320013101001103330321233220202
0112333133033332321120021003330300021210221120001222320003100022010
3301332003213013212223010323200200032031000022211230333211020200332
2202331130312300321032013013300132222203310132100030130213322131033
0332112120300122113013313310103110312121003210321023303320302010200
0003103223032103333123230 130213000210 33012303231 32232202031 0303310
M.K. Futey et al.
1212223010230232223000132 200222200123 20203201333 22233332332 0333033
0123203203333033200000201120023121223031333333013100203322231313032
1102232330301112322133102222230331032333221030321132111320112000313
3322220320331123013010233322122022001333 022010111332222000321011122
3210230213003122232213330 200212323230 10021312300 32002201232 0313321
3123200223003200033031123200213021000322021113200130213211213323200
230322002200031000331203331203110230203002132203- - - - - - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - ???????????????????????????
Geonoma 0100-0001010- -----------3131202021031000213021031200213120
002133201333203213303233223032310133321101021011302-
3323302321312002 003 3221 020 2131300 3332 312 2303 003030 2021 003 2310 33023-
13320332132-03030230030301331003031301100032133103000- - ------
000013001130013- - - - - - - -0313203 1003002 310003 00321 0100001 03- - - -
01322013201003- - - - - - - 030030330133103032100- - - - - - - - 3333333223011-3
333331003020- ---------------203310332020300320201003302030321120
231310323313- - - - - - - -00303102003- - - - - - - - 1101- - - - - - - - -3231333021013-
1332203321320102133030103321301113322102011321020021003032132032332
30031200(02)33330112010100330033113203201003200022000232112
0202331213322320320002002222320003013310031102333011313312012002213
11013233310322030111- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -??????
?????????----------------------------30100033201330330301311320
1301200011000203013203031 332 21 021 0331120230013113100111220233112113
2022002102222102122302132112003133130132230103220100132323220132032
2013301102313320312330100022012032130110103120001123323122220220000
3100303033213303230213303113330201133333200200223313233013001032333
0133110332322230032303332233310002111301202131301231322022031321200
3311111331330331100001333110022111211310322303110023320002020300233
200100230322312311313033222032301303300011000033222033 0311210002001
3012230202122333032003231301212232201332033330110022032032000012320
0131010011033303212332202020112333133033332321120021213330300021210
2211200012223200031000220103301332003213013212223010323200200032031
0000222112303332110202003322202131130312300321032013013300132222203
310132100030130213322131033033211212030012211301331331 0103110312121
0032103210233033203020102 000003103223 03210333312 32301302130 0021033
0123032313223220203103033 101222223010 23023222300 01322002222 0012320
2032013332223333233203330330123203203333033200000201120023121223031
3333330131002033222313130321102232330301112322133102222230331032333
221030321132111320112000313332222 0320 331123013010233322122022001333
02201011133222200032101112232102302130031222322133 3020021 2323230100
2131230032002201232031332131232002230032000330311232002130210003220
211132001- -------------------------------- --------------------
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - ???????????????????????????
Socratea 000- -0-0001010- - - - - - - - - - -2313120202123
1020213021031200213120002133201333203013303233223032310133321101021
011312-332332230131200200332210202101300333233223032030302021003
Arecaceae fossil fruits from Argentina
211033023133203321320030302300303013310030313021000321331000000- - - -
- - - 0000130011300130010002103-3203100300231000300321030000103- - - - - - -
01322013001003- - -0300303- -013103222100- ----------------------
3331333223011033333310001200020- - - - - - - -2220202111101132033
1033202030032020100330203032332023131032331331020303102003 1101-
303331333021013-1332203321320102133000- ----11133221020
1110102002100303213203233230033200233330112010100130033113203201003
2000220022321320202331213322320320002002222320003013310231102333011
3133320320022131101323331032203011132- - ---------------- ----------
- - - -???????????????- - ------------------------------3010003320133033
03013113201301200011000203013203031332210210331120230013113100111220
23311211320220021022221021223021321120031331301322301032201001323232
20132032201330110231332031233010002201203213011010312000112332312202
02200003100303033213303230213303113330201133333200200223313233013001
03233301331103323222300323033322333100021113012021313012313220220313
21200331110133133033110000133311002211121131032230311002332000202030
02332001002303223123113130332220323013033000110000332220330311210002
00130122302021223330320032313012122322013320333301100220320320000123
20013101001103330321233220202011233313303333232112002101333(03)
30002121022112000122232000310002201033013320032130132122230103232002
00032031000022211230333211020200332220233113031230032103201301330013
222220331013210003013021332213103303321121203001221130133133101031103
121210032103210233033203020102000003103223032103333123230130213000210
330123032313223220203103033101212223010230232223000132200222200123202
032013332223333233203330330123203203333033200000201120023121223031333
333013100203322231313032110223233031111232213310222223033103233322103
032113211132011200031333222203203311230130102333221220220013330220101
113322220003210111223210230213003122232213330200212323230100213123003
20022012320313321312320022300320003 3031123200213021000322021 1132001- - -
--------------------------------------------------------------------
---------------???????????????????????????
Chamaedorea ?100-0001000- - - - - - - - - - - 2313320232103100021102103322021
2120002131201333203213303233223032333133321101023211302-
330332230133200200332210222103310333233223030030302021003331033023333
20332110003030230130301331003031302100032133103300- - - - - - - -
000002000131013 0010002 3031320311030023100030032101 0000103-----------
01322013000003- - - - -030 0- - -310131 1032020 00- - - - - - - - - - 30133 332203011-
333331320000000 20- - - - - - - - - -00202220202100011 1100110 0331033 2020300320 20
100333303232322023131032331331020303102003013323133333133 3021013-
1332203321322102133030203321301113322102011321020021003032132032312
3203120023333011201010033003311320320100320002200223213202023312033
22320320022002222320023033210031002333011313302032002213110112333- -
------------------???????????????- - - - - - - - - - - 100010200013000210
0023233220331000212223233000203301000332013303303
013113201301200011000203013 2030 313 322 102103 311202300131131001112202
0331121132022002102222102122302132112003133130132230103220100132323
M.K. Futey et al.
022013203220133011023133203123301000220120321301101031 2000112332312
0222022000031003330332133032322133031133302011333332002002233132330
0130010323330133110332322230032303332233310002111301202131301231322
0022031321 20 03 3111013313303 3110000133311002211121131032230311002332
0000202030023320010023032231231131303322203230130330001100003322203
3031121000200130122302021223330320032313012122322013320333301100220
320320000123200131010011033303212332202020112333133033 3323211200212
1333030002121022132000122 232000310002 20103301332 00321301321 2223010
3232002000320320000222112303232110202003322202331130312300321032013
013300132222203310132100030130113322131033033211212030 0122113013313
3101031103121210032103210233033203020102000003103223032103333123230
1302130002103301230323132 232202031030 33101212223 01023023222 3000132
2002222001232020320133322233332332033303301232032033330332000002011
2002312122303133333301310020332223131303211022323303011123221331022
22230331032333221030 32113211132011200031333222203203311230130102333
221220220013330220101113 322220 0032101112232102302130031222322133302
0021232323010021312300320022012320313321312320022300320003303112320
0213021000322 021113200130213211213323200030322002212031000331200331-
203110230203002132203- - - - - - - - - - - - - - - - - - - - - ???????????????????????????
Synechanthus (01)100-0001010- ----------23133202021031000213021033220
213120002131201331203213303233223032310133321101223211322-
3321302301332002003322102 021033103322 33223030030 30202100333 1033023
33320332112003030230030301331003031302100032133103320- - - ---------
3210312203110300231000300321012000103- - - - - - - - -21322013222003- - - - - - -
030030310131123202000- - - - -30131332203013-33333313200000020- - ------
2020222020210001211011120331033202030032020120333303232332223131232
3313310203031020030103231331331303021013-133220332132010213 303020
3321301113322102011321020021003032132032312320312002333301120121001-
3003311320120120320002200223213102023312033223203200220022223200230
33210131102333011313312012002213110112333- ------------------------
-------???????????????- -------------200013000210023233220
3330002132232330002033010003320133033030131132013012000110002030132
030313322102103 31120230013113100111220233112113202200210 222210 2122 3
0213211200313313013223010322010013232322013203220133011023133203123
3010002201203213011010312001112332312222022000031003030332133032302
1330311333020113333320020022331323301300103233301331103323222300323
0333223331000211130120213130123332202 20313 2120 033111013313303311000
01333110022111311310322303110023320 0020 203002 332 001 002 3032231 231131
3033222032301303300011000033222033031121000200130122302021223330320
0323130121223220133203333011002203203200001232001310100110333032123
3220202011233313303333232112002100333030002121022112000122232000310
0022010330133200321301321222301032320020003203100002221123032321102
0200332220233113031230032103201301330013222220331013210003013021332
2131033033211212030012211301331331010311031212100321032102330332030
2010200000310322303210333 312323013021 30002103301 23032313223 2202031
0303310121222301023023222 300013220022 22001232020 32013332223 3332332
Arecaceae fossil fruits from Argentina
0333033012320320333303320000020112002312122303133333301310020332223
13130321102232330301112 32 21331 022 22 230 33103 23 332 21 03032 113211132011
2000313332222032033112301301023332212202200130302201011133222200032
1011122321023021300312223221333020021232323010021312300320022012320
31332131232002230032000330311232002130210003220211132001302132112133
23200230322002212031000331200331203110230203003132203- - - - - - - - - - - ---
----------------------------------???????????????????????????
Barcella 00012000?010- - - - - - - - - - - -3133200021031000213 021031200213120
00213320133320321330323322303231313332110002101130- - - -312330230
2332002003322102021003001 332032030320 30302021003 23103302323 3203321
3200303021003030133102303130210003213310300000020010000130011300133
21000200313203100300212000300321010000113- - - -01322
013000003030030030330033103232100- - - - 33333332223011-33333-
30000200020- - - - - - - - - - -22202022222210021111011320
3310332020300320301003302033321320231310323313310203031020131101-
303331333021013-13322033213201021130301033213011133221020
1110102002100303213203233232031200233330110010100330033113203201003
2000220022321320202331203322320320002002222320003013310031102333011
313312012002213130132333- ---------------------???????????????
0323101101000102000130002100232332203330002132232330002033010003320
1330330301311320130120001100020301320303133221021033112023001311310
0111220233112113202200210222210212230213211200313313
0132230103220100132323220132032201300110231332031233010002201203213
0110103120001123323122220220000310030303321330333021330311333020113
3333200200223313233013001032333013311033232223003230333223331000211
1301202131 30 12 31 32 2022 03 132 12 00 33 1111133133033110000133311002211121
1310322303110031320002020300233200100230322312311313033222032301303
3000110000332220330311210002001301223020212233303200323130121
223220133203333011002203203200001232001310100110333032 1233220202011
2333133033332321120021213330300021210221120001222320003100022010330
1332003213013212223010323200200032031000022211230333211020200332220
2331130312300321032013013300132222203310132100030130213322131033033
2112120300122113013313310103110 3121210032103210233033203020102 00000
3103223032103333123230130 213000210330 12303231322 32202031030 3310121
222301023023222300013220022220012320203201333
2223333233203330330123203203333033200000201120023121223031333333013
1002033222313130321102232330301112322133102222230331032333221030321
13211132011200031333222203203 311230130102333221220 220 013330 220 10111
3322220003210111223210230213003122232213330200212323230100213123003
200220123203133213123200223003200033031123200213021000322021113
2001302132112133232002303220022020310003312003312001102302030021322
0302032002020300003002123210300331023003311323
3323313313003320332100330???????????????????????????
Elaeis 000- -120001010 --------------------121302103120021
312000213320133320321330323322303231313332110002101130- -3103
M.K. Futey et al.
3023023320020033221020210 030033323320 30320303020 21003231033 0232332
033213200303021003030 13310230313021000321331030000- - - - - -0000
130012300133210002003 132031003002120003003210 10000113- - - - -0132
2013000003- - - -030030310133103232100- - -33333331223011-333
33310000202022-------20202222221002111101132033103320
2030032030100330203332132023131032331331020303102013- -1101-
303331333021013-1332203321320102113030103321301113322
102011101020021003032132032332320312002333301100101003300331
1320320100320002200223213 202023312033 22320320002 00222232000 3013310
03110233301131331201200221313013133310- --------------???????????????-
3231011010001020001300021002323322033300021322323300020330100033201
3303303013113201301200011000203011203031332210210331120230013113100
1112202331121132022002102222102122302132112003133130132230103220100
132323220132032201330110231332031233010002201203213011 0103120001123
3231222202200003100303033213303230213303113330201133333200200223313
2330130010323330133110332322230032303332233310002111301202131301231
3220220313212003311111331330331100001333110022111211310322303110031
3200020203002332001002303223123113130332220323013033000110000332220
3303112100020013012230202122333032003231301212232201332033330110022
032032000012320013101001103330321233220202011233313303 3332321120021
0033303000212102211200012223200031000220103301332003213013212223010
3232002000320310000222112303332110202003322202131130312300321032013
0133001322222033101321000301302133221330330332112120300122113013313
3101031103121210032103210233033203020102000003103223032103333123230
1302130002103301230323132 232202031030 33101212223 01023023222 3000132
2002222001232020320133322233332332033303301232032033330332000002011
2002312122303133333301310020332223131303211022323303011123221331022
22230331032333221030 32113211132011200031333222203203311230130102333
221220220013330220101113 322220 0032101112232102302130031222322133302
0021232323010021312300320022012320313321312320022300320003303112320
02130210003220211132001302132112133232002303220022020310003312003312
00110230203002132203- - - - - - - - - - - - - - - - - - - -???????????????????????????
Acrocomia 000- -110000010----------------------130210312002131200021
33201333203223303233223032310133321100021211302-310330
2301312002003322102021003 003332332030 30030302021 00323103302 3213203
32132003030230010301331023031302100032133102100- - - - - -00001
3001130013021000200313203100300231000300321010000113- - - - - -
01322013000003- - -030030330133103232100- - -33333332223011-
33333130000200000-------22220222221201111101132033103320
20330- - - - - 0200032132023131032331331020303102013- -1101-
323331333021013-13322033213201021130321033211011131221020
1110102002100303213203213230031200233330110110100330033113201201001
2000220022321320202331213322320320002002222320023013330031102333011
31331201200221311033233310-22030111322200220- - - - - - - - - - - - - - - - - - - - - -
----???????????????----------------------------
301000332013303303013113201301200011000203013203031332 2102103311202
Arecaceae fossil fruits from Argentina
30013113100111220233112113202200210 2222 102122 302 132 1120031331301322
301032201001323232201320322013301102313320312330100022012032130
1101031200011233231222202200003 10030303321330323 0213303 1133302011
3333320020022331323301300103233301331103323222300323033322333100021
113012021313012313220220313212003311101331330331100001333110022211
2113103223031100233200020203002332001002303223123113130332220323013
0330001100003322203303112100020013012230202122333032003231301
2122322013320333301100220320320000123200131010011033303212332202020
1123331230333323211200212133303000212102211200012223200031000220103
3013320032130 132122 230103 23200 200032 031000022 211230333211020200332
2202131130312300321032013013300132222203310132100030130233322131
033033211212030012211301331331010311031212100321032102
33033203020102000003103223032103333123230130213
0002103301230323132232202 031030331012 12223010230 23222300013 2200222
2001232020320133322233332332033303301232032033330332000002011200231
2122303133333301310020332223131303211022123303011123221331022222303
310323332210303211321113201120003 1333 222203203 311230130102333221220
220013330220101113322220 003210 1112232 1023021 300312 22322133302002 123
2323010021312300320022012320313321312320022300320003303112320021302
10003220211132001- ----------------------------------------------
- - - - - - - - - - - - - - - - - - - - - -??????????????????????? ????
Bactris 000--110001010- - - - - - - - - - - - -3110021031000213021031200 21312000
2133201333203223303233223032310133321100021211102-
3123302301332002 003 3231 020 2100300 3332 332 0303 003030 2021 003 2310 33023-
23320332132003030230010301331023031302100032133102100- -------0000
13001130013021000200313203100300231000300321010000113- - - - - -
01322213000003- - -03003033013310323210003333333333223011-
33333130200202020-----222020222121120011110113203310332020300
32020100330203032132023131032331331020303102013--1101-
323331333021013-13322033213201021330321033213011133221020
1110102002100303213203213230031200233330110010100330033113203201001
2000220022321320202331213322320320002002222320023013330031102333311
31331201200221311013233310-22030111- - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - -???????????????- - - - ----------------------------30100033201330
33030131132013012000110002030132030313322102103 31120230013113100111
2202331121132022002102222102122302132112003133130132230103220100132
323220132032201330110231332031233010002201203213011010 3120001123323
1222202200003100303033213303230213303113330201133333200200223313233
0130010323330133110332322230032303332233310002111301202131301231322
0220313212 00 33 1110133133033 1100001333110022111211310322303110023320
0020203002332001002303223123113130332220323013033000110000332220330
3112100020013012230202122333032003231301212232201332033330110022032
032000012320013101001103330321233220202011233312303333 2321120021013
3303000212102211200012223200031000220103301332003213013212223010323
2002000320320000222112303332110202003322202331130312300321032013013
3001322222033101321000301302133221310330332112120300122113013313310
M.K. Futey et al.
1031103121210032103210233033203020102000003103223032103333123230130
2130002103301230323132232 202031030331 01212223010 23023222300 0132200
2222001232020320133322233332332033303301232032033330332000002011200
2312122303133333301310020332223131303211022323303011123221331022222
303310323332210303211321113201120 0031 333222203 2033 11230130102333221
220220013330220101113322 220003 2101112 2321023 021300 31222322133302 002
123232301002131230032002201232031332131232002230032000
330311232002130210003220211132001- - ------------------------------
--------------------------------------------------------------
- - - - - - - - - - - - - - - - - - - - - - - - -???????????????????????????
Cocos 000- -100000010232030322002001232221010231332000210310002130210
31220211122002131201333203213303233 22303231011332110002301130233323-
3223013320020033 221 0202 130 3003332 3322 303 0030 302021 0230 310 3302 32332-
0312112023030230030301331023032302100032133103000- - - - - 000011001130
2130210002003132031203002310003003210100201130132201322013000003- - -
03003233013310 32321000--103333332230 11-33 333130000200020---02022
2320222223022120011110111223310332020300320201003302030
3210202313303- - - - - -010303102003031103-310333313021012-
3332203303320322133030203321201113-2210201110102002100
3032132032312300332002333301120101003300331132032010032000220022321-
3202023312033223203200020022223200230133100311223330113133120120022-
1311013233330322030 11132122302200013002- -------------------------
---------???????????????------------------------
301000332013303303013113201301200011000203013203031332 2102103311202
30013113100111220233112113202200210 2222 102122 302 132 1120031331301322
3010322010013232322013203220133011023133203123301000220120321301101
0312000112332312222022000031003030332133032302113031133302011333332
0020022331323301300103233301331103323222300323033322333100021113012
021313012313 22 022 03 13212 003 311101331330 33110000133311002211121131 03
2230311002332000202030023320010023032231231131303322203230130330001
1000033222033031121000200130122302021223330320032313012122322013320
3333011002203203200001232001310100110333032123322020201123331321333
3232112002100333030002121022112000122232000310002201033013320032130
132122230103232002000320310000222112303332110202003322 2021311303123
0032103201301330013222220331013210003013021332213103303321121203001
2211301331331010311031212100321032102330332030201020000031032230321
0333312323013021300021033 012303231322 32202031030 33101212223 0102302
3222300013220022220012320 203201333222 33332332033 30330123203 2033330
33200000201120023121223031333333013100203322231 31303211022323303011
123221331022222303310323332210 3032113211132011200031333222203203313
230130102333221220220013330220 101113322220003210111223210 2302130031
2223221333020021232323010021312300320022012320313321112320022300320
00330311232002130210003220211132001- - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - -???????????????????????????
Attalea 000 100001011232030322002001232221010231312000
210310002130210312002131200021332013302032133032332230323
Arecaceae fossil fruits from Argentina
101333211000220113021332330230133200200332210202100
3003332332230300303020210232310330232331033213200
3032230030301331023031302100032133103000- - - - - - -000313001
130213021000200313203120300231000300321010000113- - - - -013220
13000033- - -030030330133103232100- - -02333333223011-33333132000
200020- - - - - 02022222302212001111211120331031201030032020100330203-
32132023133032331331020303102003031103-
31333133302101123332203303320122133030203321301113-
221020111010200210030321 320323 12300332002333 30112010100330033113203
2010032000220022321320202331203322320320002002222320023013312031122
33301131331201200221 31101323331032203011132322302200013002132131030
111 3 2 20 0 3 ? ? ? ?? ? ? ? ? ?? ? ? ? ? - - - -------33000210023233220333000
2132232330002033010003320133033030131132013012000110002030132030313
3221021033112023001311310011122023311211320 2200210 222 210 212 2302132 1
1200313313013223010322010013232322013203220133011023133203123301000
2201203213011010312000112332312222022000031003030332133032302113031
1333020113333320020022331323301300103233301331103323222300323033322
3331000211130120 2131 301231322 0220 3132 12003 3111 0133 13303 311000013331
100221112113103223031100233200020 203002332 0010 0230 32231 231131303322
2032301303300011000033222033031121000200130122302021223330320032313
0121223220133203333011002203203200001232001310100110333032123322020
2011233313213333232112002101333030002121022112000122232000310002201
0330133200321301321222301032320020003203100002221123033321102020033
2220213113031230032103201301330013222220331013210003013021332213103
3033211212030012211301331331010311031212100321032102330332030201020
0000310322303210333312323 013021300021 03301230323 13223220203 1030331
0121222301023023222300013 220022220012 32020320133 32223333233 2033303
3012320320333303320000020112002312122303133333301310020332223131303
21102232 33 030111232213312 222 22 303 31032 33 322 10 3032113211132011200031
333222203203313230130102333221 2202200 1333022 010111 3322220 0032101112
2321023021300312223221333 020021232323 01002131230 03200220123 2031332
1112320022300320003303112320021302100032202111320013021321121332320
0030322002212031000331200331200112230203002132203- - - - - - - - - - - - - - - -
???????????????????????????
Syagrus (01)00--10001001123203032200200123222101023131200021031000
3130210312002131200021332013332032233032332230323101133211000212113
0233323302301332002003322 102021003003 33233223030 03030202102 3231033
02323320330132003030230030301331023031302100030133103300----
0000130011300130210002003132031003 0023100330032101 0000113- - - - -
01322013000003- - - -230030330133103232100- - - -03333333223011-
33333130000200020- - - -022222221022120010110111203310332020
30032020100330203032132023131032331331020303102003031103-
313333313021012-3332203303320122133030203321301113-
221020111010200210030321 320323 12300332002333 30112010100330033113203
2010032000220022321320202331203322320320002002222320023013310031122
33301131331201200221311013233310322030111323- - - -00013
M.K. Futey et al.
002132131030111322003???????????????-323101101000102000130002
1002323322033300021322323300020330100033201330330301311320130120001
100020301320303 133 221 021 03311202300131131001112202331121132 022 00210
2222102122302132112003133130132230103220100132323220132032201330110
2313320312330100022012032130110103120001123323122220220000310030303
321330323021130311333020 1133333200200223313233013001032333013311033
232223003230333223331000211130120213130123132202203132120 0331110133
13303311000013331100221112113103223 031100233200020203002 332 0010023 0
3223123113130332220323013033000110000332220330311210002001301223020
2122333032003231301212232201332033330110022032032000012320013101001
103330321233220202011233313213333232112002121333030002 1210221120001
2223200031000220103301332003213013212223010321200200032031000022211
2303332110202003322202331130312300321032013013300132222203310132100
0301302133221310330332112120300122113013313310103110312121003210321
0233033203020102000003103 223032103333 12323013021 30002103301 2303231
3223220203103033101212223 010230232223 00013220022 22001232020 3201333
2223333233203330330123203203333033200000201120023121223031333333013
1002033222313130321102232330301112322133102222230331032333221030321
1321113201120003133322220320331 3230 13010 23332212 20220 0133 3022010111
3322220003210111223210230213003122232213330200212323230100213123003
2002201232031332111232002230032000330 311232002130210003220211132001
3021321121332320003032200221203100033120033120011023020
3002132203- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - -???????????????????????????
Lytocaryum (01)00- - - -10111001023203032200200123222101023
1312000210310003130210312 002131200021 33201333203 21330323322 3032310
133321100021011302133033023013320020033221021210030033323
3223030030302021023231033 223233203321 32003032230 03030133102 3031302
100032133103000----000313001130213021000200313203
100000231000300321010000113- - - -01322033000003- - -
030030330133103232100- - -03333333203011-33333130000200020- -----0222
223330223202111101112033103 32010300 32020 1003 3020 303213202 3131 03233 1
33102030310200301110--311330333021013-333220330
3320122133030203321301113-2210201110102002100303213201231230033200
2333301120101003300331132032010032000220022321320202331203322320300
0020022203200230133100311223330113133120120022231101323331032203011
132222302200003002132131030111322003???????????????
0323101101000102000130002100232332203330002132232330002033010003320
1330330301311320130120001100020301320303133221021033112023001311310
011122023311211320220021 022221021 2230 21321120031331301322301032201 0
0132323220132032201330110231332031233010002201203213011010312000112
3323122220220000310030303321330323021130311333020113333320020022331
3233013001032333013311033232223003230333223331000211130120213130123
1322022031 32 12 00 3311101331330 33 1100001333110022111211 31 03 2230 311002
3320002020300233200100230322312311313033222032301303300011000033222
0330311210002001301223020212233303200323130121223220133203333011002
Arecaceae fossil fruits from Argentina
203203200001232001310100110333032123322020201123331321 3333232112002
1213330300021210221120001222320003100022010330133200321301321222301
0323200200032031000022211230333211020200332220233113031230032103201
3013300132222203310132100030130213322131033033211212030012211301331
3310103110312121003210321023303320302010200000310322303210333312323
0130213000210330123032313 223220203103 03310121222 30102302322 2300013
2200222200123202032013332 223333233203 33033012320 32033330332 0000020
1120023121223031333333013100203322231313032110223233030111232213310
2222230331032333221030321132111320112000313332222032033132301301023
33221220220013330220101113322220003210111223210230 2130031 2223221333
020021232323010021312300320022012320313321112320022300 3200033031123
2002130210003220211132001302132112133232002303220022120310003312003
31200110230203002132203030- - - - - - - - - -00000030021232103003310230033
113233323313313003320332100330???????????????????????????
Tripylocarp a100--0-0100011- ----------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------------------------------------------------------
--------------
M.K. Futey et al.
... It includes microfossils (Camacho, 1967;Méndez, 1966;Chebli and Serraiotto, 1974), bivalves and brachiopods (Chebli and Serraiotto, 1974), echinoids (Parma and Casadío, 2005), turtles and alligators (Staesche, 1929;Simpson, 1937;Bona, 2005Bona, , 2007Bona and De La Fuente, 2005;Bona et al., 2018), and mammals (Bonaparte et al., 1993;Bonaparte and Morales, 1997;Pascual et al., 1992Pascual et al., , 2002Forasiepi and Martinelli, 2003;Gelfo and Pascual, 2001;Gelfo, 2007;Gelfo et al., 2007Gelfo et al., , 2009. Plant remains are also abundant, with leaves (Martínez, 1992;Iglesias, 2007;Iglesias et al., 2007;Jud et al., 2017), flowers (Jud et al., 2017(Jud et al., , 2018a(Jud et al., , 2018b, palm stipes (Romero, 1968), palm fruits (Futey et al., 2012), and palynomorphs (Archangelsky, 1973(Archangelsky, , 1976Romero, 1973;Archangelsky and Romero, 1974 following Figari et al. (1999), and the position of the study area. B-Geological map of the Estancia Las Violetas area, showing the position of the Las Violetas forest (modified from Raigemborn et al., 2010 andRuiz et al., 2017). ...
... Paleoclimatic inferences from all these fossils arrive at the same conclusion inferred by other proxies that suggest that the Paleocene flora from the Salamanca Formation developed under humid and warm-temperate conditions, and without limiting factors. The presence of thermophilic groups such as palms (Romero, 1968;Futey et al., 2012;Ruiz et al., 2017), alligators (Bona, 2007;Bona et al., 2018), and podocarps that grow in nearly exclusively high rainfall environments suggest a warm and humid climate during the deposition of the Salamanca Formation. Mean annual temperature of 14.1 ± 2.6°C and mean annual precipitation of at least 115 cm (+50/-35 cm) were estimated based on standard techniques of leaf-margin analyses . ...
... At this locality, four fossiliferous levels (L1, L2, L3, and L4; Fig. 2) were defined throughout the outcrops of the unit. These levels include conifer and dicot permineralized trunks, branches and twigs, palm stipes, palm fruits and seeds, and leaf impressions (Somoza et al., 1995;Futey et al., 2012;Ruiz et al., 2017), and are evidence of forested coastal settings that developed during the early Paleocene at middle-high paleolatitudes of South America. This fossil forest provides an excellent opportunity to (1) expand the knowledge of Paleocene dicot wood anatomy by describing and taxonomically assigning new dicot fossil woods recovered in the fossiliferous level L3 from the Salamanca Formation, and (2) increase paleoclimate information from South America by incorporating climatic inferences based on fossil wood anatomy of the assemblage cropping out in fossiliferous levels L2 and L3 from the Las Violetas Fossil Forest. ...
Article
Las Violetas Fossil Forest of the Salamanca Formation was a mixed forest of Paleocene age located in the Golfo San Jorge Basin, central Patagonia of Argentina, which includes conifers (previously studied), dicots and palms. In the present work, the dicot fossil woods outcropping in one of the fossiliferous levels (L3) of the Salamanca Formation are described, taxonomically assigned, and their affinities are discussed. They are three new species, including a new genus related to the Myrtaceae and Laurales. The complete fossiliferous assemblage is used to infer the climate and environment of the fossil forest. This assemblage represents a parautochtonous fossil plant association that lived either on the margins of channels or exposed bars in well-drained soils, probably tropical red-soils, close to their depositional setting (tidal channels and bars of an estuary). Warm and humid conditions and porous host-rocks favored the silicification of woods and pigmentation with Fe, resulting in yellow-orange and green fossil woods. We apply growth ring analysis to the conifer woods. For the dicots, wood anatomical characters influenced by the environment were analyzed, and the Vulnerability and Mesomorphy indices were used. These methods, comparisons with extant forests and the sedimentology, suggest that the Las Violetas Fossil Forest was an evergreen forest that developed under uniform growing seasons that ended abruptly, with an abundant water supply and high mean annual temperatures.
... Fossils of Tripylocarpa aestuaria Gandolfo & Futey originate from the Danian Salamanca Formation (63.3-61.9 Ma) of Argentina (Futey et al., 2012), and were placed in subtribe Attaleinae of tribe Cocoseae based on morphological comparisons and phylogenetic analysis. It has since been used as a calibration fossil for Attaleinae in a variety of studies (e.g., Meerow et al., 2015;Freitas et al., 2016;Barrett et al., 2019). ...
... It has since been used as a calibration fossil for Attaleinae in a variety of studies (e.g., Meerow et al., 2015;Freitas et al., 2016;Barrett et al., 2019). However, we think affinities with Cocoseae are equivocal because T. aestuaria has a single apical germination pore (Futey et al., 2012) rather than three pores, as is characteristic of tribe Cocoseae. Germination pores in Attaleinae are either lateral or subbasal but never apical; subapical pores are found in Bactridinae. ...
Article
Full-text available
Premise: Fossils are essential for understanding evolutionary history because they provide direct evidence of past diversity and geographic distributions. However, resolving systematic relationships between fossils and extant taxa, an essential step for many macroevolutionary studies, requires extensive comparative work on morphology and anatomy. While palms (Arecaceae) have an excellent fossil record that includes numerous fossil fruits, many are difficult to identify due in part to limited comparative data on modern fruit structure. Methods: We studied fruits of 207 palm species, representing nearly every modern genus, using X-ray microcomputed tomography. We then developed a morphological data set to test whether the fossil record of fruits can improve our understanding of palm diversity in the deep past. To evaluate the accuracy with which this data set recovers systematic relationships, we performed phylogenetic pseudofossilization analyses. We then used the data set to investigate the phylogenetic relationships of five previously published fossil palm fruits. Results: Phylogenetic analyses of fossils and pseudofossilization of extant taxa show that fossils can be placed accurately to the tribe and subtribe level with this data set, but node support must be considered. The phylogenetic relationships of the fossils suggest origins of many modern lineages in the Cretaceous and early Paleogene. Three of these fossils are suitable as new node calibrations for palms. Conclusions: This work improves our knowledge of fruit structure in palms, lays a foundation for applying fossil fruits to macroevolutionary studies, and provides new insights into the evolutionary history and early diversification of Arecaceae.
... The presence of Paleocene macrofloras in the Salamanca Formation of southern Chubut Province, Argentina, has long been known (Berry, 1937a), but for many decades they were the focus of few studies, and their precise ages remained uncertain (see Clyde et al., 2014). Most paleobotanical work from the Salamanca Formation had focused on the abundantly preserved, permineralized fossil wood and occasional fruits that so far represent relatively few, though significant, taxa of conifers and angiosperms (Petriella, 1972;Petriella and Archangelsky, 1975;Romero, 1968;Ragonese, 1980;Brea et al., 2005Brea et al., , 2007Brea et al., , 2008Brea et al., , 2011Futey et al., 2012;Ruiz et al., 2017Ruiz et al., , 2020. More recently, several macrofloral compression assemblages from the Salamanca and overlying Peñas Coloradas formations were intensively collected in the vicinity of Sarmiento, southern Chubut, and found to be outstandingly preserved and diverse (Brea and Zucol, 2006;Iglesias, 2007;Iglesias et al., 2007;Raigemborn et al., 2009). ...
... Palmate leaves without a costa are placed in the coryphoide-lepidocaryoid clade (Moore and Uhl, 1982;Uhl et al., 1987). None of the several estipites (Romero, 1968;Arguijo, 1979Arguijo, , 1981 and fruits (Futey et al., 2012) known from the Salamanca Formation corresponds to palmate leaved taxa. ...
Article
Full-text available
Early Paleocene macrofloras from the Southern Hemisphere are little known, despite their significance for understanding plant evolution, biogeography, and global variation in recovery after the end-Cretaceous extinction. As a foundation for systematic and paleoecological work, we describe 51 angiosperm leaf morphotypes from three distinct, precisely dated early to late Danian time intervals, using collections from the Salamanca and Peñas Coloradas formations in the San Jorge Basin, Chubut Province, Patagonia, Argentina. These rich floras were previously analyzed but with minimal descriptions. The assemblages comprise the first stratigraphically controlled and quantitatively collected floras for the early Paleocene of the Southern Hemisphere. Botanical affinities of the angiosperm morphotypes are not formally assigned here, but we informally associate some of them with families including Arecaceae, Fabaceae, Cunoniaceae, Lauraceae, Nothofagaceae, Rhamnaceae, and Rosaceae; in addition, leaves of Menispermaceae and other Rhamnaceae were formally described in previous work. Other families potentially present in these assemblages include Akaniaceae, Anacardiaceae, Apiaceae, Araceae, Bixaceae, Juglandaceae, Malvaceae, Sapindaceae, and Urticaceae. Remarkably, there is little floral turnover or change in dominance through the Danian floral sequence spanned by the studied localities, even among estuarine vs. continental depositional environments. This finding indicates a homogeneous, generalist, long-lived floral association following the K-Pg extinction, similar in these respects to many North American Danian floras. However, the richness of the Danian Patagonian floras, from paleolatitudes >50 degrees South, along with other lines of evidence from the region, suggests differences in the response of terrestrial ecosystems in southern South America to the terminal Cretaceous event from those of the Northern Hemisphere. The flora appears to be largely paleo-endemic in nature and shows several compositional links to the Eocene floras of Patagonia, emphasizing the importance of diversification within Patagonia after the end-Cretaceous event as a factor leading to the hyperdiverse Eocene regional floras. Plain language and multilingual abstracts at: https://palaeo-electronica.org/content/2021/663-1124/3258-patagonia-danian-forests-abstracts
... The association between palm bruchines and palms could have arisen as early as the Late Cretaceous, as the fossil record of both palms (Matsunaga and Smith, 2021) and palm bruchines (Poinar, 2005; see Section 4.3 below) extend back to this time. Early records of crown-group Cocoseae include permineralized fruits of subtribe Attaleinae recovered from Danian deposits in Patagonia, Argentina (Futey et al., 2012), indicating that the Cocoseae most likely diverged during the Late Cretaceous, as is also suggested by divergence-age estimations (Meerow et al., 2015). ...
Article
Modern Neotropical rainforests are characterized by the high intensity and host specificity with which insects feed on plants. Previous studies have shown that, during the middle–late Paleocene, the leaves of the early evolving Neotropical rainforests of tropical South America were heavily herbivorized by insects. Yet, less attention has been given to insect damage found on fossil fruits and seeds, despite the host specialization of many disseminule predators in modern forests. Here, we present and describe borings found on a fruit compression fossil of cf. Cocos (coconut) from the middle–late Paleocene Cerrejón Formation (58–60 Ma) of Colombia. We interpret the borings as constructed by palm seed beetles (Chrysomelidae: Bruchinae: Pachymerina) based on size, number, position, plant reaction tissue, and plant host selection. This occurrence provides the earliest record of an ecological interaction between seed beetles and palms, suggesting that this host-specific interaction has been consistently maintained for several tens of millions of years.
... Aside from these, no other records are known of this morphotype in Patagonia, although it reappeared in the Pliocene of the Entre Rios Province (~35 • S; Lutz 1980Lutz , 1984. Futey et al. (2012) described Tripylocarpa aestuaria, the only known Argentinean fossil palm fruits, which belong to the Subtribe Attalainae, Tribe Cocoseae, Subfamily Arecoideae. These were collected at the Salamanca Formation (Danian) at Estancia Las Violetas Locality, Patagonia. ...
Article
Clearly, past and present plant diversity is due to the many evolutionary processes that are the manifestation of the organisms’ responses to changes in the environment. Most current evolutionary studies (such as population genetics, epigenetic mechanisms, phylogeography, and phylogenetics) provide information on a few generation-times and are focused on microevolutionary events. Some of these studies have as major goals the estimation of divergence times, construction of phylogenies, and explanations of biogeographical distributions. Nevertheless, the majority of these studies use fossils only as calibration points and mostly ignore critical information that only fossils can provide about the course of evolution in general. Herein, we reevaluate macroevolutionary changes that are critical for understanding past and present diversity and provide concrete examples in deep time (Cretaceous-Paleogene) from several sites in Patagonia, Argentina. We discuss Extinction, Retraction, Expansion, Diversification, and Coevolution (Codiversification) processes that have significantly shaped modern Patagonian vegetation.
... During the last 15 years, persistent fieldwork in Argentinean Patagonia has yielded several superbly preserved fossil flowers and fruits (Zamaloa et al., 2006;Gandolfo et al., 2009Gandolfo et al., , 2011Futey et al., 2012;Hermsen et al., 2012;Hermsen and Gandolfo, 2016;Gandolfo and Hermsen, 2017;Jud et al., 2017Jud et al., , 2018aWilf et al., 2018Wilf et al., , 2019, which in many cases represent the oldest records globally for the taxa and the clades they belong. Among these, Gandolfo et al. (2011) and Hermsen et al. (2012) 1). ...
Article
Full-text available
PREMISE: Fossils provide fundamental evidence of the evolutionary processes that crafted today’s biodiversity and consequently for understanding life on Earth. We report the finding of Myrtaceidites eucalyptoides pollen grains preserved within the anthers of a 52-million-year-old Eucalyptus flower collected at Laguna del Hunco locality of Argentinean Patagonia and discuss its implications in understanding the evolutionary history of the iconic Australian genus Eucalyptus. METHODS: Pollen grains were extracted from the flower’s anthers and were then observed under light microscopy and scanning electron microscopy. The phylogenetic position of the fossil was investigated by adding pollen data to a previously published total-evidence matrix and analyzing it using parsimony. RESULTS: We erect the species Eucalyptus xoshemium for the fossil flower. Pollen extracted from E. xoshemium belongs to the species Myrtaceidites eucalyptoides, which, until now, was only known as dispersed pollen. The numerous pollen grains recovered from the single flower allowed estimation of M. eucalyptoides’ variability. Results of the phylogenetic analysis reinforce the position of this fossil within crown group Eucalyptus. CONCLUSIONS: The discovery of these pollen grains within a Patagonian Eucalyptus fossil flower confirms the hypothesis that Myrtaceidites eucalyptoides represents fossil pollen in the Eucalyptus lineage, extends the geographic and stratigraphic fossil pollen record, and supports an earlier age for crown-group eucalypts.
... Fossil occurrences from the Salamanca and overlying Peñas Coloradas Formations are consistent with humid, mesothermal, frost-free climates (Bonaparte et al. 1993;Brea et al. 2005;Iglesias et al. 2007;Palazzesi and Barreda 2007;Raigemborn et al. 2009;Futey et al. 2012;Clyde et al. 2014;Ruiz et al. 2017Ruiz et al. , 2020. Palynofloras of the Salamanca Formation and related strata over larger areas reflect a range of environments, from mangrove swamps to lowland and upland forests (Petriella and Archangelsky 1975;Zamaloa and Andreis 1995;Volkheimer et al. 2007). ...
Article
Full-text available
The Cretaceous-Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extir-pation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macro-floras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study.
... (2) Tripylocarpa aestuaria from the late Danian of Argentina (Salamanca Formation). This fossilized fruit shows mesocarp with furrows or striations and three distinctive longitudinal markings running from apex to base of the epicarp, similar to Attaleinae (Futey et al., 2012). From the tribe Iriarteeae we used (3) Socratea brownii from the Late Oligocene of Mexico (Quinta Formation). ...
Article
Background and aims: Cunoniaceae are woody plants with a distribution that suggests a complex history of Gondwanan vicariance, long-distance dispersal, diversification, and extinction. Only four out of c. 27 genera in Cunoniaceae are native to South America today, but the discovery of extinct species from Argentine Patagonia is providing new information about the history of this family in South America. Methods: We describe fossil flowers collected from early Danian (early Palaeocene, ~64 Mya) deposits of the Salamanca Formation. We compare them with similar flowers from extant and extinct species using published literature and herbarium specimens. We used simultaneous analysis of morphology and available chloroplast DNA sequences (trnL-F, rbcL, matK, trnH-psbA) to determine the probable relationship of these fossils to living Cunoniaceae and the co-occurring fossil species Lacinipetalum spectabilum. Key results: Cunoniantha bicarpellata gen. et sp. nov. is the second species of Cunoniaceae to be recognized among the flowers preserved in the Salamanca Formation. Cunoniantha flowers are pentamerous, complete, the anthers contain in-situ pollen, and the gynoecium is bicarpellate and syncarpous with two free styles. Phylogenetic analysis indicates that Cunoniantha belongs to crown-group Cunoniaceae among the core-Cunoniaceae clade; although it does not have obvious affinity with any tribe. Lacinipetalum spectabilum, also from the Salamanca Formation, belongs to the Cunoniaceae crown-group as well, but close to tribe Schizomerieae. Conclusions: Our findings highlight the importance of west Gondwana in the evolution of Cunoniaceae during the early Palaeogene. The co-occurrence of Cunoniantha bicarpellata and Lacinipetalum spectabilum, belonging to different clades within Cunoniaceae, indicates that the diversification of crown-group Cunoniaceae was underway by 64 million years ago.
Article
Slow rates of molecular evolution at low taxonomic levels hamper studies of relationships among species and subsequent biogeographic and evolutionary analyses. An example is the genus Brahea, which is among the most poorly understood lineages of American palms and is characterized by a wide variety of growth forms and intermediate morphological features. We generated c. 400 kb of genome-scale data from all three genomes for the 11 currently described Brahea spp. to infer phylogenetic relationships, reconstruct ancestral growth form, estimate ancestral geographical ranges and test for niche differences among closely related species with geographical overlap. Relationships receive strong support and conform to previous subgeneric assignments, except for placement of the dwarf species B. moorei in Brahea subgenus Erythea. Our robust phylogenetic hypothesis reveals trends in growth form, including an overall increase in height in the B. armata clade and independent evolution of dwarf forms from taller ancestors in the B. pimo and B. dulcis clades. Ancestral range estimation reveals roles of dispersal (e.g. B. edulis on Guadalupe Island) and putative divergence within geographical regions in some cases (e.g. in the B. armata clade) but is equivocal in others (e.g. in the B. pimo clade). We find evidence of niche non-equivalency among species in the B. armata clade in northwestern Mexico and some evidence of niche non-equivalency between B. berlandieri and B. dulcis, the former of which is synonymized under B. dulcis. Our findings have implications for the complex biogeographic history in Central America and Mexico, suggesting that divergence within regions and dispersal are the predominant processes of species diversification. Future studies should include population-level sampling across the genus, along with morphological and ecological information, to assess distinctness among species and, particularly, levels of gene flow, in an integrative fashion.
Article
These fossil palm fruits (19 species) are near to the living Hyphaene thebaica Mart. This fossil is preserved as an subovoid, hard endocarp with basal scar of vascular bundles and apial scar of the germ-pore. It has been found in Aptian rocks near Assouan Lake. Therefore, from this time up to now, in Egypt, this palm was probably existing by hot and dry climate.-after English summary
Article
The structure of silicified axes of palm was studied. The age of plantiferous beds, after Andreis et al. (1991) is Late Cretaceous. The fossil structures were compared with actual palm axes. On the basis of the characteristics of vascular bundles; interfasicular parenchyma; fibres and phloem, the materials were assigned to three Subfamilies: Sabaloideae, Coccoideae, and Phoenicoideae. The endosperm structure of fossil fruits was also studied, showing great similarity with the fruits of actual Phoenix canriensis. -English summary
Article
Description is given of a petrified coconut like fruit measuring 13 x 10 x 6 cm collected from the Tertiary sediments of Amarkantak (M.P.), India. The fruit wall is thick and differentiated into a narrow epicarp, a wide fibrous mesocarp and a sclerosed thick endocarp. The seed coat is 3-4 cells thick, adhered to, or free from the endocarp. The pulp or endosperm of the seed is made of loosely arranged thin walled parenchyma. This is the first report of a C. nucifera like petrified fruit from India.
Article
A new morphospecies of palm trunk, Palmoxylon pichaihuensis sp. nov., is proposed on the basis of several specimens recovered from a succession of tuffs and breccias at Pichaihue, Neuquén Province, Argentina. The specimens comprise atactosteles that display densely crowded fibrovascular bundles in the peripheral zone, and less congested fibrovascular bundles, randomly scattered or arranged in irregular circles in the central zone; sclerenchyma reniform; xylem with 1-4 large metaxylem vessels and 6-12 smaller protoxylem elements in the central zone, and an f/v ratio of 1.3-2:1. The Pichaihue fossil assemblage is similar to, and probably coeval with the flora of Bajo de Santa Rosa, Río Negro Province, reflecting a warm and relatively humid climate, at ca. 40- 45° S, in southern South America during the Late Cretaceous.