Sagaria cilentana gen. et sp. nov.-A New Angiosperm Fructification ...

Sagaria cilentana gen. et sp. nov.-A New Angiosperm Fructification from the
Middle Albian of Southern Italy
Sergio Bravi a , Maria Rosaria Barone Lumaga b , James E. Mickle c, *
a
Dipartimento di Scienze della Terra, Università di Napoli ‘‘Federico II’’, Largo San Marcellino 10, 80138 Napoli, Italy
b
Orto Botanico, Università di Napoli ‘‘Federico II,’’ via Foria 223, 80139 Napoli, Italy
c
Department of Plant Biology, Box 7612, North Carolina State University, Raleigh, N.C. 27695-7612 U.S.A
article info
Article history:
Received 29 June 2007
Received in revised form
24 November 2009
Accepted in revised form 9 December 2009
Available online 11 January 2010
Keywords:
Basal eudicot
Early Cretaceous
Fossil angiosperm
Italy
Plattenkalk
Ranunculaceae
1. Introduction
abstract
Few Cretaceous age angiosperms have been reported from Italy.
Late Cretaceous angiosperm leaves and seeds of uncertain affinity
have been described from the Venetian Alps in northeastern Italy
(Gomez et al., 2002) and a leaf ascribed to Sapindopsis has been
described from the Middle Albian of the southern Apennines (Bravi
et al., 2004). The present study describes a newly discovered
angiosperm fructification from the Albian of southern Italy. A single
specimen has been recovered from fine-grained limestones of the
‘‘plattenkalk’’ of the Middle Albian at Monte Alburni near Petina,
province of Salerno, region of Campania, Italy. This is the first
discovery of a fossil angiosperm fructification from this region and
represents the oldest angiosperm megafossil described from the
Italian peninsula. This fossil also provides new information about
basal eudicot angiosperms.
2. Geological setting and methods
The specimen comes from the ‘‘plattenkalk’’ locality of Monti
Alburni that is located approximately 1.5 km SSW of the town of
* Correspondence author.
E-mail address: james_mickle@ncsu.edu (J.E. Mickle).
0195-6671/$ – see front matter Ó 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cretres.2009.12.001
Cretaceous Research 31 (2010) 285–290
Contents lists available at ScienceDirect
Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes
A single specimen of a new fossil angiosperm, Sagaria cilentana gen. et sp. nov., has been recovered from
limestones of the ‘‘plattenkalk’’ of the Middle Albian at Monti Alburni, near Petina, Campania, Italy.
Associated fossils include Pagiophyllum sp., Podozamites sp. and Zamites sp. The angiosperm has a main
axis approximately 8 cm long with three attached leaves and a distal cyme bearing three fructifications.
The fructifications display divided distal tips indicating three - four lobes. Four cm from the base,
underneath the main axis, there is a lobed structure, presumably a leaf. An unattached leaf near the
specimen is in a position that suggests that it comes from the same plant. The leaf has an elongate
petiole, obtuse base and on the right side of the lamina are three lobes. Affinities are uncertain but show
similarities in overall morphology to basal eudicots, especially Ranunculaceae.
Ó 2010 Elsevier Ltd. All rights reserved.
Petina [Fig. 1; circa 40 31’ 18’’ N 15 22 05’’ E; I. G. M. sheet 198,
I SE–Auletta; 1:25.000], in the Provence of Salerno in Campania,
Italy (Bravi and Garassino, 1998). Plant fossils are found within
thinly laminated limestones and dolomitic limestones termed
‘‘plattenkalk’’ (Fig. 2). In addition to abundant invertebrate and
vertebrate remains (mainly decapod crustaceans and fishes),
frequent plant remains are also encountered, including Pagiophyllum
sp. and Brachyphyllum sp. (Barone Lumaga et al., 2005;
Fig. 3A), and Podozamites sp. and Zamites sp. (Bravi and Garassino,
1998). The depositional environment of the strata at this locality is
interpreted as a strongly restricted, shallow lagoonal area not far
from land, with upper subtidal-intertidal conditions; it evolves
upward to more subtidal marine conditions into a limited carbonate
platform lagoonal environment (Bravi and Garassino, 1998).
The specimen was studied by light microscopy and photographed
under visible and ultraviolet (l ¼ 312 nm) spectra. Ultraviolet
photography was done using a Transilluminator Lourmat UV
TFX-20 M.
3. Systematic paleobotany
Order: aff. Ranunculales Lindley 1833
Family: aff. Ranunculaceae Jussieu 1789
Genus: Sagaria gen. nov.

286
Derivation of Name: Sagaria is named in honor of the collector of
the specimen; Giovanni Sagaria.
Type species: Sagaria cilentana sp. nov.
Generic Diagnosis. Angiosperm, axis with leaves and fruits.
Leaves lobate, petiolate. Distal tips of foliar lobes mucronate.
Inflorescence cymose. Fruits borne by an elongate peduncle, on an
expanded cup-shaped recepticle, up to 1.1 cm long, and 0.6 cm
wide and composed of at least three follicles with a maximum
width of 3 mm and partially fused in their basal three-quarters.
Distal tips of the follicles mucronate.
Derivation of specific epithet. From the area, Cilento, in which the
specimen was found.
Holotype. Specimen 301a housed at Museum of Paleobotany and
Ethnobotany, at Orto Botanico University of Naples. (Fig. 3 A) [part]
and specimen 301b housed at the Paleontology Museum of
Magliano Vetere (Cilento National Park), Provence of Salerno
(Fig. 3B) [counterpart].
Type locality. Monti Alburni locality located approximately
1.5 km SSW of the town of Petina [Fig. 1; circa 40 31’ 18’’ N 15 22
05’’ E; I. G. M. sheet 198, I SE–Auletta; 1:25.000], in the Provence of
Salerno in Campania, Italy.
Type Stratum. ‘‘Plattenkalk’’ at Monte Alburni (Bravi and Garassino,
1998).
Age. Middle Albian.
Material. The species is represented by a single specimen
preserved as part and counterpart of a coalified compression.
Sagaria cilentana sp. nov.
Figs. 3–5
Specific diagnosis. Characteristics as for the genus.
Description of specimen
The specimen consists of a main axis with four attached leaves
and a distal cyme bearing three fructifications. The specimen is
S. Bravi et al. / Cretaceous Research 31 (2010) 285–290
Fig. 1. Map showing the location of the plattenkalk of Monte Alburni in Campania.
approximately 8 cm long, with the main axis about 1.75 mm wide
and remaining at a relatively uniform width. In addition, there is
a detached leaf that is clearly derived from the specimen. The
specimen displays five nodes (Fig. 3A,B). The disposition of the
nodes indicates an alternate or helical arrangement of appendices.
The two most proximal nodes show axes that are broken off but at
their bases are approximately the same diameter as the main axis of
the specimen (Fig. 3A).
About 4 cm from the base of the specimen and lying underneath
the main axis there is a lobed structure, presumably a leaf (Fig. 3A,B,
4G). It is quite folded and details of the morphology are difficult to
discern.
Two small leaves are found in an axillary position approximately
0.5 cm from the insertion of a lateral appendage (Figs. 3A,B, 4D,H).
These leaves are probably in early stages of development. Both
leaves are lobate and about 2 mm long. One leaf, more clearly
visible, displays three distinct lobes (Fig. 4H); at the apex of each
lobe is a small, pointed tip (Fig. 4H arrow). Based on their position,
these leaves could represent stipules or immature leaves of a small
branch, however, the lack of such structures at other nodes indicates
that these are leaves on a branch just beginning to develop.
In proximity to the specimen, part of a lobed leaf is visible on the
matrix (Fig. 3A,B, 4D,E). The petiole of this leaf is not in direct
connection with the main axis (Figs. fig3A, 4D,H); however, from
the position of the petiole (Fig. 4H), and morphology of the leaf
(Fig. 4D), it is highly probable that it comes from the same plant as
the axis. The leaf has a petiole that is 4 mm long and about 0.5 mm
wide. The broken base of the petiole matches the broken end of the
adjacent appendage from the axis of the Sagaria specimen
(Fig. 4D,H). The leaf shows an obtuse base, and on the right side of
the lamina three lobes with deep sinuses are visible (Fig. 4E). Lobes
are about 2 mm deep and 1.5 mm wide (Fig. 4D,E). At the apex of
each lobe is a small pointed tip (Fig. 4D,E arrow). Primary venation
is faint but medial (Fig. 4D). It is not possible to discern secondary
venation, but the position of the lobes and the mucronate tip
suggest a median secondary vein at a perpendicular angle to the

primary veins. The form of the leaf and broken petiole base strongly
indicate that this leaf is from the Sagaria specimen.
At the distal end of the axis a dichasium subtended by a 5 mm
long axillary bract bears three fructifications (Figs. 3A,B). One
fructification is borne singly on what appears to be a continuation
of the main axis (Fig. 3A), the other two are borne on a branching
lateral axis, giving a cymose appearance to the specimen (Fig. 3A,B).
The lateral axis is subtended by a leaf (Fig. 4I arrow) whose
attachment was visible most clearly under ultraviolet light. The leaf
is folded proximally and the lamina is partially obscured by the
lamina of the detached leaf (Figs. 3A, 4I).
The first fructification is 1.1 cm long, and 0.6 cm wide (Figs. 3A,B,
4A). This fructification is borne on a pedicel 1 cm long and 1 mm
wide that supports one enlarged, cup-shaped receptacle (Fig. 4A).
Fine striations are visible on the pedicel proximal to the receptacle
(Fig. 5A, lower arrow). Three follicles are visible (Fig. 4A, 5A,B). The
follicles have a maximum width of 3 mm and are fused at their
basal three-quarters. The tips of the follicles are mucronate (Fig. 4A,
5A,B). On the first fructification there is a single longitudinal line
visible (Fig. 5A, upper arrow) on one of the lobes, especially on the
counterpart (Fig. 5B, arrow). A similar line is also visible on the part,
on another lobe (Fig. 5A, arrow). These lines appear to represent
dehiscence slits. Because these putative dehiscence slits are seen
only on part or counterpart suggests that they occur only on one
S. Bravi et al. / Cretaceous Research 31 (2010) 285–290 287
Fig. 2. Geological section of the plattenkalk of Monte Alburni in Campania.
face of the fruit wall. This further suggests that the fructification is
composed of fused or partially fused follicles. At a distance of
approximately 1.5 mm from the receptacle two linear structures
appear in the illustrations (Fig. 4A) approximately 2.5 mm long that
appear to be bract-like leaves. However, these structures are
actually artifacts of the matrix fracture pattern from collection
(Fig. 5A) and are not bracts.
The other two fructifications are of similar size and show only
the outer surfaces (Figs. 3A,B, 4B,C). The distal tips are not as
distinct in the matrix as the first fructification, but clearly display
three, possibly four lobes (Fig. 4B,C). They are borne on a branching
axis (Fig. 3A,B) with the first of the two fructifications borne nearly
sessile; the other is on a pedicel about 0.5 cm long. The fructifications
are inserted on an expanded receptacle that is 2.0 mm long
and expands to 2.3 mm in diameter (Fig. 4F). One of the fructifications
has basal markings that are suggestive of scars of other
floral parts (Fig. 4F). There is no other evidence of a calyx or corolla
present on the fossil.
4. Discussion
Because it is incompletely preserved, taxonomic placement of
this fossil is somewhat problematic. One possibility is that it is
a gnetophyte. However, the reproductive structures do not appear

288
Fig. 3. Sagaria cilentana part and counterpart. A, entire specimen; arrows indicate
positions of nodes. Note Pagiophyllum sp. on the right; scale bar ¼ 1 cm. B, counterpart
to A; scale bar ¼ 1 cm.
to be paired, as occurs in Gnetales (Foster and Gifford, 1989; Krassilov
and Bugdaeva, 2000) and the leaves are unlike any known
gnetophyte, living or fossil, in phyllotaxy or morphology (Foster
and Gifford, 1989; Taylor et al., 2009). The fossil does not closely
resemble any other group with the exception of the dicotyledonous
angiosperms.
The overall form of the fruit, the fruit’s position with respect
to the receptacle, and possible scars of subtending organs indicate
that Sagaria possessed a superior ovary. It is impossible to
ascertain if the flower was perfect or imperfect, or if sepals,
petals or either were present in the living plant. The degree of
fusion of the fruit of Sagaria is also difficult to interpret, especially
in light of the incomplete preservation of two of the three
specimens preserved. The best preserved specimen shows
a degree of fusion reflecting about G of the distance from the
base to distal tip. This may represent the actual developmental
degree of fusion or may represent a stage in which all of the
follicles were fused and as dehiscence occurred, the follicles
separated from each other as occurs in a number of species (e. g.,
Anderberg, 2001: delpsta1.htm).
In the absence of cuticle, sepals, petals, stamens, clear scars of
these floral parts or associated pollen, affinities of this fossil are
problematic to ascertain. The general features of Sagaria, including
a capsular fruit composed of at least three follicles and fruits borne
in a cyme are most consistent with those of ‘‘basal’’ eudicots as
outlined by APG (1998, 2003). Among the basal eudicots, specific
affinities of Sagaria are not entirely clear, but shape, disposition,
fusion and apparent mode of dehiscence of the fruit, apparent
herbaceous habit, and lobed leaves are most similar to those seen in
Ranunculales, especially Ranunculaceae (Cronquist, 1981) such as
Delphinium (Moraldo et al., 1981; Anderberg, 2001). Certainly,
S. Bravi et al. / Cretaceous Research 31 (2010) 285–290
fused fruits occur in other families such as the Trochodendraceae
(Cronquist, 1981) but the overall form of the fructification is most
similar to Ranunculaceae (Cronquist, 1981).
A number of fossil Ranunuculaceae are known (see Pigg and
DeVore, 2005). Among these are forms known from the Early
Cretaceous, including Teixeiraea lusitanica (von Balthazar et al.,
2005), a fossil similar to Thalictrum (Friis et al., 1994), Hyrcantha
karatscheensis (Krassilov et al., 1983), H. decussatus (Dilcher et al.,
2007), Achaenocarpites capitellatus (Krassilov and Volynets, 2008)
and Ternicarpites floribundus (Krassilov and Volynets, 2008).
Teixeiraea lusitanica consists of a charcoalified fossil flower bud
from the late Aptian or early Albian of Portugal that has been
attributed to Ranunculales, possibly Ranunculaceae (von Balthazar
et al., 2005). Teixeiraea has no vegetative features preserved, thus
few features of this species are preserved in common for comparison
with Sagaria. Teixeiraea is unisexually staminate, while Sagaria
is bisexual or unisexually carpellate. Additional specimens of each
species may permit more direct comparison.
Early Cretaceous fruits from the Famalicão and Vale di Agua
localities in Portugal similar to Thalictrum (Ranunculaceae) are
reported by Friis et al. (1994). These fruits clearly differ from
Sagaria. The fruits reported by Friis et al. (1994) are about one-third
the size of Sagaria and have marked longitudinal ribs in contrast to
the lack of ribs in Sagaria.
Hyrcantha karatscheensis is known from the Middle Albian of
Kazakhstan (Krassilov et al., 1983). The gynoecium of H. karatscheensis
possesses three or five free carpels and a persistent
calyx is present. Leaves are pinnately compound (Krassilov et al.,
1983). Sagaria differs from H. karatscheensis in that Sagaria has
fused carpels, lacks a persistent calyx, and displays simple, lobate
leaves.
Hyrcantha decussatus is known from the Barremian or Aptian of
the Yixian Formation of China (Leng and Friis, 2003; Dilcher et al.,
2007) and shows 3–4 carpels that are basally fused for about onehalf
of their length, but as in Sagaria, it is difficult to ascertain if the
degree of fusion is due to development or dehiscence. The size of
the carpels are similar, 1.25 cm long in H. decussatus and 1.1 cm in
Sagaria, but the arrangement of fruits is decussate in H. decussatus
(Leng and Friis, 2003; Dilcher et al., 2007) whereas they do not
appear to be so in Sagaria. Further, an ochrea is reported at the
nodes and base of the fruits in H. decussatus (Dilcher et al., 2007),
a characteristic not present in Sagaria.
Achaenocarpites capitellatus and Ternicarpites floribundus were
described from the Middle Albian of the Primorye Region of Russian
Far East by Krassilov and Volynets (2008). Both are referred to
Ranunculaceae, and are both known from whole or nearly whole
plants that show a rhizomatous habit. Achaenocarpites is characterized
by fruiting heads composed of 16 radially spreading
achenes while Sagaria differs in fruit type and number of fruits per
fruiting structure. A small leaf associated with Achaenocarpites is
trilobed (Krassilov and Volynets, 2008) and similar in appearance
to the small leaf attached to the axis of Sagaria. Ternicarpites is
known from several branching axes with terminal fruits. The fruits
are ternate follicles 6–8 mm long, showing a longitudinal keel of
two parallel ridges and free to the base (Krassilov and Volynets,
2008). Fruits of Sagaria are larger, about 11 mm long, partially
fused, and do not show a keel. Leaves associated with Ternicarpites
are more deeply lobed than those of Sagaria, with the leaves of
Ternicarpites appearing nearly pinnate (Krassilov and Volynets,
2008) while the leaves of Sagaria are more laminar.
In addition to leaves discussed above, a few angiosperm leaves
from the Early Cretaceous also display a deeply lobed morphology
as seen in Sagaria. One of these is Vitiphyllum Fontaine. (1889; Doyle
and Hickey, 1976) from the Lower Cretaceous of eastern North
America. Both the leaves associated with Sagaria and Vitiphyllum

are simple, lobed leaves but Vitiphyllum shows a greater degree and
complexity of lobing than seen in those associated with Sagaria;
there is probably little affinity between these genera. Another is the
genus Araripia Mohr and Ecklund (2003). Sagaria differs from
Araripia in that Sagaria leaves are more highly lobed and the
venation in Sagaria is inferred to be at a more perpendicular angle.
The base of the leaf attributed to Sagaria is similar to that seen in
S. Bravi et al. / Cretaceous Research 31 (2010) 285–290 289
Fig. 4. Fruits and leaves of Sagaria cilentana. A, single fruit; scale bar ¼ 1 mm. B, first fruit of a pair of fruits on branch; scale bar ¼ 1 mm. C, second fruit of paired fruits on branch;
scale bar ¼ 1 mm. D, detached lobed leaf showing probable point of attachment and axillary branch with two small attached leaves; scale bar ¼ 1 mm. E, lobes of detached leaf in D.
Arrow indicates the small mucronate tip of leaf lobe; scale bar ¼ 1 mm. F, base of fruit in B, showing possible scars of floral parts; scale bar ¼ 1 mm. G, stem with lamina of leaf
compressed underneath; scale bar ¼ 1 mm. H, base of petiole with axillary branch bearing small lobed leaves. Arrow indicates small mucronate tip of leaf lobe; scale bar ¼ 1 mm. I,
base of cyme with subtending petiole (arrow); scale bar ¼ 1 mm.
Ranunculus. Araripia is attributed to Lauraceae (Mohr and Ecklund,
2003).
Sagaria adds to the growing number of basal eudicots known from
the Early Cretaceous flora. While the exact morphology and affinities
of Sagaria are uncertain, its characteristics place it most closely with
the Ranunculales. The great diversity displayed by Sagaria, Teixeiraea
(von Balthazar et al., 2005), Thalictrum-like fossil (Friis et al., 1994),

290
Fig. 5. Fruit of Sagaria cilentana. A, fruit in Fig. 4a showing the presumed line of dehiscence (upper arrow) and fine longitudinal striations on pedicel (lower arrow); scale
bar ¼ 1 mm. B, counterpart of A, showing the presumed line of dehiscence (arrow); scale bar ¼ 1 mm.
Hyrcantha (Krassilov et al., 1983; Leng and Friis, 2003; Dilcher et al.,
2007), Achaenocarpites, and Ternicarpites (Krassilov and Volynets,
2008) suggests that early radiation of ranunculoid eudicots was well
underway during Early Cretaceous time.
Acknowledgement
The authors thank Valerie Knowlton for her assistance in
preparing the figures. We also thank Drs. Peter Endress, James
Doyle, David Dilcher and the reviewers for their constructive
comments.
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