“at the end of the world,” reflecting provenance
from the last stages of the Gondwanan supercontinent and from Patagonia, popularly known
as “the end of the world.” Type material: repository Museo Paleontológico Egidio Feruglio,
Trelew, Argentina (MPEF-Pb); holotypus hic
designatus MPEF-Pb 6434a,b (Fig. 1, A to C)
(lowercase letters indicate part and counterpart),
from Laguna del Hunco, Huitrera Formation, early
Eocene, Chubut Province, Argentina, quarry LH13
( 16), collected 7 December 2002 (fig. S18); paratype MPEF-Pb 6435a,b (Fig. 1, G and H), Laguna
del Hunco quarry LH04 ( 16), 21 November 2006.
Description: based on two dispersed, apparently
mature, compressed fruiting calyces; only the holotype preserves berry and pedicel remains. Pedicel
preserved length 18. 8 mm, width 1 mm at inflection point of insertion. Calyx basally invaginated,
highly inflated to completely surround the berry,
five-lobed, angled, partly open at apex, length by
width 25.2 mm by 13.2 mm (1.9:1) on holotype,
21. 6 mm by 17. 6 mm (ratio 1.2:1) on paratype.
Calyx lobes equally sized, sinuses angular and
incised one-quarter to half the total calyx length,
tips acute triangular. Venation with one robust
primary meridional vein per lobe, terminating
at lobe apex and alternating with secondary veins.
Secondary veins arising near the base, visually
distinct from the primaries and other vein orders,
and bifurcating close to the lobe sinuses (Fig.
1C). Intersecondary veins arising near the base,
visually distinct from secondaries and tertiaries,
dichotomizing into the random, irregular reticulum of tertiary through at least quinter-nary veins that fill most of the vein field. Some
basal veins compressed into partial detachment
from the calyx body, the remnants visible against
the matrix. Berry round, flattened and coalified
from fossilization, filling the full width of the
calyx and appearing by surface relief to extend
apically almost to the lobe sinuses; seeds not
The fossils preserve several features that, together, are diagnostic of the Physalinae clade
of Solanaceae ( 20, 21, 23, 25). These include
the highly inflated, lantern-shaped, pedicellate,
five-parted, angled, regular calyces covering a
large, fleshy berry, along with the combination
of (i) invaginated base, (ii) robust primary veins
that terminate in the lobe tips (not the lobe
sinuses), and (iii) secondary veins that fork before the sinus. Nonphysaline species of Solanaceae with ICS lack some or all of these three
features (e.g., those in Deprea, Nicandra, Solanum,
Withania, and the Juanulloeae clade) (figs. S5, S11,
and S13) ( 23, 24). Within Physalinae, there are
several problematic species of Physalis s.l. and
other genera that diverge below the core Physalis
clade (Fig. 2) (equivalent to P. subgenus Rydbergis)
of relatively homogeneous species ( 13, 23–25).
Among these taxa, Alkekengi officinarum calyces are the most similar to the fossils, but they
have a much larger size, consistently rounded
shape, and nearly closed apex. Otherwise, each
of these basal physaline species structurally differs from the fossils and core Physalis ( 20, 24, 25)
(figs. S2 to S17). All these observations highlight
the close relationship of the fossils with core
Both the MP and ML results (Fig. 2 and fig. S1)
support the fossils’ affinity with Physalinae and
Physalis. The newly identified species is placed
in Physalinae with strong support in both MP
and ML analyses and at a basal polytomy of (MP,
weak support) or within (ML, strong support) the
crown of core Physalis. Both the MP and ML
topologies are generally consistent with and
reproduced most major clades from ( 10), with
robust support of the critical Physalinae clade.
However, each tree recovered the closely related ( 10) Iochrominae and Deprea as collapsed
into a single clade that also included Nicandra,
which has Deprea-like calyx morphology but is
not closely related to that genus. Nevertheless,
the overall agreement of the MP and ML tree
topologies with ( 10) is notable in light of the
limitations of the analysis. Notably, no species
from outside Physalinae were misplaced among
Physalinae, and no Physalinae species were misplaced into other clades. Considering the unknown additional organs of this extinct species,
the incomplete knowledge of calyx morphology in extant Solanaceae, and problematic resolution among extant basal species of Physalinae
( 13, 25), we suggest that the newly identified
species can be used, conservatively, to constrain
the divergence of Physalinae to a minimum of
Physalis infinemundi sp. nov. represents a
derived lineage of Solanaceae in Gondwanan
South America at 52.2 Ma, pushing back considerably the evolutionary timing of the family.
Compared with recent molecular divergence estimates ( 10), these fossils are considerably older
than the 30 Ma crown for the entire Solanaceae family as well as the 11 Ma divergence of
Physalinae. A second recent study similarly placed
the Physalinae divergence at only 9 Ma ( 13).
Our results reinforce the emerging pattern
wherein numerous fossil plant taxa from Gond-
wanan Patagonia and Antarctica are substan-
tially older than their corresponding molecular
dates ( 26, 27), demonstrating Gondwanan history
for groups conjectured to have post-Gondwanan
origins under entirely different paleogeographic
and paleoclimatic scenarios. Likewise, the de-
rived position of the newly identified fossil spe-
cies shows that the origins and diversification
of Solanaceae must have taken place at a much
earlier time than previously thought, consid-
erably before final Gondwanan breakup. Other
regions of Gondwana are also likely to have
played prominent roles in Solanaceae evolu-
tion, especially Antarctica, which has produced
other important asterid fossils ( 27). Moreover,
the newly identified fossils directly help to
resolve temporal inconsistencies between the
evolutionary timing of Solanaceae and its her-
bivores and mutualists ( 28). The large fossil
berry strongly implicates trophic associations
with animals, as seen in extant Physalis ( 29).
Today, Physalis inhabits South, Central, and
North America, and Mexico is its center of di-
versity (2). Thus, the fossils establish a rare link
to extant New World floras from late-Gondwanan
Patagonian assemblages, whose living relatives
are mostly concentrated in the Old World tropics
The discovery of 52.2-Ma fossil inflated calyces
demonstrates an ancient history for ICS and
implies that the Eocene world was already populated with derived solanaceous reproductive
structures. The potential adaptive functions of
the inflated calyx are scarcely discussed in the
literature ( 30), but the lakeside rainforest paleoenvironment led us to consider flotation dispersal and protective drying for the berry. Simple
kitchen-sink and stream experiments on several
Physalis species confirmed that intact calyces
hold stable air pockets around the berry, enabling
flotation for several days in water and preventing
wetting of the berry during rains. These insights
suggest potential origins of ICS in ancient humid,
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This work was supported by NSF grants DEB-1556666 and
DEB-1556136, National Geographic Society grant 7337-02, and the
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