www.elsevier.com/locate/revpalbo

Review of Palaeobotany and Paly

Fossil Nelumbonaceae from the La Colonia Formation

(Campanian–Maastrichtian, Upper Cretaceous),

Chubut, Patagonia, Argentina

Maria A. Gandolfoa,*, Ruben N. Cuneob,1

aL.H. Bailey Hortorium, Department of Plant Biology, Cornell University, Ithaca, NY 14853-5908, USAbMEF-Museo Paleontologico Egidio Feruglio, Trelew, Chubut, Argentina

Received 29 April 2004; received in revised form 5 August 2004; accepted 23 September 2004

Abstract

We describe here fossil leaves and fruits assignable to the family Nelumbonaceae Dumortier 1828. Fossils were collected at

the Canadon del Irupe locality, La Colonia Formation (Campanian–Maastrichtian, Upper Cretaceous), Chubut Province,

Patagonia, Argentina. The fossil leaves are simple, with symmetrical and peltate lamina, orbicular in shape, and an entire

margin. The venation is truly actinodromous with at least 15 primary veins, the secondary veins are poorly developed and

intercalated with the primary veins, the third category vein is opposite, and the areoles are well-developed and 4- to 5-sided.

These features are characteristic of the extant genus Nelumbo Adamson 1763, and the presence of these characters in the fossils

allows their placement within this genus. Reproductive structures, similar to the fruit-receptacle of Nelumbo with fruits in situ,

were found associated but not in organic connection with the leaves. These fossils constitute the only and oldest record for the

family in the Southern Hemisphere. This record indicates that Nelumbo was more widespread in the past than today.

D 2004 Elsevier B.V. All rights reserved.

Keywords: Nelumbonaceae; fossils; Upper Cretaceous; La Colonia Formation; Patagonia; Argentina

1. Introduction

What little is known about Maastrichtian floras of

Southern South America is based primarily on five

floras, four from Chile (Rıo de las Chinas, Quiriquina,

0034-6667/$ - see front matter D 2004 Elsevier B.V. All rights reserved.

doi:10.1016/j.revpalbo.2004.09.007

* Corresponding author. Fax: +1 607 255 5407.

E-mail addresses: mag4@cornell.edu (M.A. Gandolfo)8

rcuneo@mef.org.ar (R.N. Cuneo).1 Fax: +54 2965 432100.

Pichasca, and Cerro Guido) and one from Argentina

(Estancia La Sara), which have been characterized

mostly by palynological data (Troncoso and Romero,

1998). Although special attention has recently been

given to the Upper Cretaceous La Colonia Formation

due to its diverse and rich vertebrate fauna found in its

continental facies (including fishes, turtles, snakes,

plesiosaurs, birds, mammals, and dinosaurs; Albino,

2000; Bonaparte, 1985; Gasparini and De La Fuente,

2000; Pascual et al., 2000; Rougier et al., 2001), its

nology 133 (2005) 169–178

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178170

paleoflora is scarcely known. Nevertheless, new

paleobotanical collections have revealed the presence

of an unexpectedly diverse plant assemblage.

Previous to this report, only two records of fossil

plants from the La Colonia Formation were known;

both records are of aquatic ferns. Archangelsky et al.

(1999) described megaspores and microspore massulae

of the fossil species Paleoazolla patagonica, while

Cuneo and Archangelsky (2000) reported megafossils

and microfossils with affinities to the extant genus

Regnellidium within the family Marsileaceae. Associ-

ated with these aquatic ferns, there are other pterido-

phytes, gymnosperms, and several angiosperms. This

contribution is focused on the taxonomy and descrip-

tion of the aquatic angiosperm family Nelumbonaceae

from the La Colonia Formation paleoflora.

2. Geology and age

The La Colonia Formation outcrops in an area

known as the Somuncura Massif; it is exposed between

Telsen and Sierra Rosada, Chubut province, Patagonia,

Argentina (Fig. 1). This formation was recognized by

Pesce (1979) as a homogeneous series of upward fining

Fig. 1. Locality map of outcrops of the La Colonia Formation

sediments, and it is divided into three portions (lower,

middle, and upper). Argillaceous sediments are char-

acteristic of the lower and middle portions of the

sequence. They are considered continental and thought

to probably represent lacustrine or lagoonal deposits.

The upper portion of the sequence changes to marginal

and shallow marine environment (Ardolino and Del-

pino, 1987; Ardolino and Franchi, 1996).

Ardolino and Franchi (1996) assigned a Senonian

(pre-Campanian) age to the lower and middle

sections. Later, Page et al. (1999) considered that an

unconformity in the base of the unit is inter-Senonian,

and, consequently, they assumed that the lower and

middle sections are at least Campanian in age. The

age of the upper section, based on ostracod and

foraminifera data collected in several localities is

Campanian–Maastrichtian, Upper Cretaceous (Ardo-

lino and Franchi, 1996).

3. Materials and methods

The fossil specimens studied here are fragments

and complete leaves and fruit impressions. They were

recovered from clays deposited in a littoral lagoon

(Maastrichtian, Upper Cretaceous) at Chubut province.

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178 171

environment. Although they seem to be quite fragile,

they are well preserved, suggesting low degree of

transportation. These fossil remains were found in

Fig. 2. Stratigraphic column of the Canadon d

association with other angiosperm leaves (at least four

morphotypes), another aquatic angiosperm (Nym-

phaeaceae seeds, Gandolfo et al., 2002), aquatic ferns

el Irupe locality, La Colonia Formation.

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178172

(Paleoazolla and Regnellidium), other pteridophytes,

and gymnosperms.

Fossil remains were collected at the Canadon del

Irupe locality (Fig. 2), La Colonia Formation, Chubut

province, Argentina and are housed at the Paleobotan-

ical collection of the Museo Paleontologico Egidio

Feruglio under the initials MPEF-Pb-Mz. Leaves,

receptacles, and fruits of extant species of Nelumbo

used for comparisons were observed from the herba-

rium collection of the L.H. Bailey Hortorium (BH).

Fossil leaves were described following the manual

of leaf architecture by the Leaf Architecture Working

Group (1999), and leaves and fruits were studied using

a stereomicroscope Zeiss Stemi SV-11. Photographs

were obtained with a Nikon 995 digital camera.

4. Systematic descriptions

4.1. Leaves

Genus Nelumbo Adamson 1763.

Nelumbo puertae Gandolfo and Cuneo, sp. nov.

(Plate IA–F).

Diagnosis: Simple, centrally peltate microphyll,

lamina orbiculate (L/W: 1), margin entire; first vein

category actinodromous, at least 12 primary veins

diverging regularly from the center of the lamina,

bifurcating at least once, and forming festooned

brochidodromous archs; poorly developed secondary

veins intercalated with the primary veins; third and

fourth category veins opposite percurrent; areoles

well-developed, 4- to 5-sided; marginal ultimate

venation looped, freely ending ultimate veins

unbranched to 1-branched, highest vein order fourth.

Holotype: MPEF-Pb-Mz 864-A and B; Plate IA.

Paratypes: MPEF-Pb-Mz 865–901; Plate IB, C and F.

Type locality: Canadon del Irupe; Mirasol Chico

Creek, Los Altares (4369-IV), Chubut province,

Patagonia, Argentina.

Horizon: uppermost part of the La Colonia Formation

at Canadon del Irupe locality.

Studied material: MPEF-Pb-Mz 864–901.

Derivatio nominis: puertae, in honor of Pablo Puerta,

Argentinean paleontological explorer.

Remarks: The leaves are simple microphylls charac-

terized by their symmetrical peltate lamina. The lamina

is rounded to orbicular with an entire margin, is at least

4 to 11 cm in diameter, and probably of membrana-

ceous texture (Plate IA–C). The petiole is inserted

centrally at the lamina, and, from its point of insertion,

the primary veins emerge radially (Plate IA,B,D,F ).

The first category venation is truly actinodromous, with

12–18 primary veins, emerging from the center of the

leaf (insertion point of the petiole) at acute angles (25–

308) in relation to other primary veins (Plate IA,B,D).

The course is straight, and the veins bifurcate once

maintaining the course; they then abruptly curve to join

another primary vein or a bifurcation of a primary vein,

forming brochidodromous arches (Plate IG). There are

veins of smaller caliber intercalated with the primary

veins that are interpreted as poorly developed secon-

dary veins. These veins have straight to slightly sinuous

course, and they are fused to third order veins forming a

bherring boneQ pattern. The third category vein is

opposite percurrent, with the veins emerging uniformly

from the primary veins at right to obtuse angles (90–

1008), their course is straight. The fourth category veinis the highest order and has an opposite percurrent

model (Plate ID,E). The areoles are well developed,

and they are 4- to 5- sided. Inside the areoles, the freely

ending ultimate veins that are unbranched to 1-

branched are noticeable. In addition to the festooned

brochidodromous arches formed by the primary veins,

there are other arches formed by veins of less caliber;

they originate a looped ultimate marginal venation

(Plate IC–E).

4.2. Reproductive structures

Associated Nelumbo-like structures.

(Plate IIA–C).

Structures inferred as reproductive organs were

found associated with the leaves. The remains consist

of numerous impressions similar to those described by

Samylina (1968) as ban inversely conical, accrescent

receptacle with fruits inside, like those of NelumboQand by Upchurch et al. (1994) as probably ba floral

axis.QThe Patagonian fossils are probably conical fruit-

like receptacles, at least 2.5 cm long and 1.1–1.5 cm in

diameter (Plate IIA–C). From the distal flat portion of

the receptacle, there are between 5–14 protuberances

that we have interpreted as fruits. They are surrounded

Plate I.

A–F Leaves of Nelumbo puertae Gandolfo and Cuneo, sp. nov.

A. General aspect of a leaf showing a round and symmetrical peltate lamina. MPEF-Pb-Mz 864 A.

B. Another leaf showing the general aspect of an orbicular and symmetrical peltate lamina. MPEF-Pb-Mz 901.

C. Detail of the entire margin and primary venation. MPEF-Pb-Mz 899.

D. Detail of the point of insertion of the petiole. Note the central origin of the actinodromous first category venation. MPEF-Pb-Mz. 864 A.

E. Detail of the primary veins forming brochidodromous arches and of the high category venation. MPEF-Pb-Mz 877.

F. Folded lamina exposing the central position of the petiole. MPEF-Pb-Mz 875.

G–H. Nelumbo lutea Willdenow.

G. Detail of the central position of the petiole and pint of origin of the first category venation.

H. Detail of the entire margin and the primary veins forming brochidodromous arches. Also see detail of the high category venation.

Bar=1 cm for a–h.

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178 173

Plate II.

A–C Nelumbo-like fossil receptacle and fruits.

A. Receptacle laterally compressed. MPEF-Pb-Mz 914.

B. Top view of a receptacle with fruits in situ, note the wall of the receptacle. MPEF-Pb-Mz 903.

C. Detail of the fruits, note the apertures (respiratory pores). MPEF-Pb-Mz 920.

D–E. Nelumbo lutea Willdenow.

D. Top view of a receptacle with fruits in situ.

E. Detail of the fruits showing the respiratory pores. Bar=1 cm for figs. A–B and D–E, bar=0.5 cm for C.

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178174

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178 175

by the thick wall of the receptacle. The wall of the

receptacle produces a rim surrounding the fruits. If our

interpretation is correct, each fruit is entrenched into a

single cavity and therefore separated from each other.

The fruits are rounded to oblong in shape and have a

small aperture at the top (Plate IIC). We assume that

these apertures are representing small respiratory pores.

Type locality: Canadon del Irupe; Mirasol Chico

Creek, Los Altares (4369-IV), Chubut province,

Argentina.

Horizon: uppermost part of the La Colonia Formation

at Canadon del Irupe locality.

Studied material: MPEF-Pb-Mz 902–920.

5. Discussion

From the morphological data gathered from the

Patagonian fossil leaves, it is clear that they belong to

the monotypic family Nelumbonaceae, allowing the

assignment of these leaves to the living genus

Nelumbo Adanson 1763. Nelumbo puertae leaves

are almost indistinguishable from extant ones.

Nelumbo, within the monotypic family Nelumbona-

ceae Dumortier, comprises two extant species,

Nelumbo nucifera Gaertner and N. lutea Willdenow.

Both species are aquatic perennial, have peltate

orbicular leaves, and produce nut-like indehiscent

fruits that are free within an accrescent receptacle

(Cronquist, 1981; Williamson and Schneider, 1993;

Takhtajan, 1996; Wiersema, 1997). Hickey and Wolfe

(1975) characterized the leaves of the family Nelum-

bonaceae as simple, truly peltate with entire margin,

and truly actinodromous with numerous primaries.

Upchurch et al. (1994) presented a list of leaf features

for modern Nelumbo. These features are the orbicular

lamina, the entire margin, the central petiole, the

medial primary vein giving rise to secondary veins

immediately beneath the leaf apex, the intercostal

primary loops uniform in size and shape, the tertiary

veins regular and transversely oriented, and, finally,

hexagonal areoles (Plate IG–H). The new fossil

species, N. puertae, shares five of the seven characters

enunciated by Upchurch et al. (1994). Nelumbo

puertae differs from extant species of Nelumbo in

the presence of poorly developed secondary category

venation, and the shape of the areoles. In addition, the

lamina of the fossil species is smaller than the extant

species in which sizes vary between 10–100 cm in

diameter (Williamson and Schneider, 1993; Wier-

sema, 1997). Furthermore, the fossil species has fewer

primary veins than the extant ones (19–23 veins,

Knowlton, 1930; personal observation, Plate IH).

Although the fossil record for the Nelumbonaceae is

extensive, there is no fossil record from the Southern

Hemisphere; all fossil species described as belonging

to the family come from the Northern Hemisphere.

Several fossil species of Nelumbo were described from

the Northern Hemisphere for sediments of Cretaceous

to Miocene in age. Kuprianova and Tarasevich (1983)

provided a summary of the fossil record for Nelumbo

pollen grains. The oldest leaves described as belonging

toNelumbo come from the Albian of Portugal (Saporta,

1894), later redescribed by Teixeira (1945). All the

Portuguese fossil species range from 20–30 cm in

diameter, have more than 20 primary veins, and the

tertiary vein category seems to be random reticulate

(see Teixeira, 1945, plate 1–3). However, their correct

taxonomic position is doubtful. These fossils are only

portion of leaves. They were exhumed from the same

sediments and locality where other leaves are attributed

as members of the family Nymphaeaceae, hence it is

possible that they are indeed members of the latter and

not Nelumbonaceae. A new reinvestigation of these

fossils is necessary to really assess their taxonomic

placement, which is beyond the scope of this paper.

Moreover, Collinson et al. (1993) and Drinnan et al.

(1994) considered as valid for Europe only the reports

of Nelumbo of Tertiary age; also Drinnan et al. (1994)

considered that possible leaves and reproductive

structures of Nelumbonaceae are present at the late

Albian of eastern North America.

In North America, there are at least 30 fossil

species of Upper Cretaceous and Tertiary age that

have been assigned to Nelumbo, among them N.

crossii Knowlton 1930, N. dawsoni (Hollick) Berry

1935, N. tenuifolia (Knowlton) Knowlton 1922 and

N. weymouthi R. W. Brown 1933. All these fossil

species were erected during the first half of the last

century and they are based mostly on single speci-

mens (Knowlton, 1930). They were described as

having entire margins and actinodromous venation,

and primary veins varying from 11 to 23; however,

many of the descriptions and illustrations lack enough

information to allow detailed comparisons with the

new species N. puertae.

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178176

Matsuo (1954) described the species Nelumbo

orientalis on the basis of leaf impressions from the

Upper Cretaceous of Japan, and he considered this

record as the oldest for the family known from East

Asia. Nonetheless, later on Matsuo (1962) transferred

this species to the species Nelumbium orientalis, a

fossil species also based on leaves remains, without

any explanation.

The possibility that the fossils described here could

belong to the fossil genus Nelumbites Berry 1911a

was also considered. The fossil genus Nelumbites was

erected by Berry (1911) to place fossil leaves with

characters similar to those of living Nelumbo.

Upchurch et al. (1994) emended its diagnosis. Based

on the revised diagnosis, we disregard the placement

of the Patagonian fossils within the genus Nelumbites

because they differ in lamina shape, position of the

petiole, tertiary category pattern, the ultimate venat-

ion, and lack of a midvein.

The fossil genus Protonelumbo Knowlton was

rejected as well as suitable for the fossils described

here. Knowlton (1930), who erected this fossil genus,

expressed that Protonelumbo leaves are similar to

those of Nelumbo but they differed in their margins.

Protonelumbo shows obtuse teeth or lobes, one for

each rib, with each lobe separated by deep rounded

sinuses. N. puertae has entire margin.

As mentioned earlier the family comprises two

modern species, N. lutea, a native of North and

Central America, and N. nucifera, which is wide-

spread in warmer parts of Asia and Australia. Both

species grow in ponds, lakes, slow streams and in

waters up to 2 m deep (Williamson and Schneider,

1993). The orbicular shape and the centered petiole of

N. puertae suggest that it had aquatic leaves as well.

Consequently, it is possible that N. puertae lived in a

very similar environment. This interpretation is also

supported by cooccurrence of the two aquatic ferns

(Regnellidium and Paleoazolla) and the sedimento-

logical data that defined the sediments as lacustrine/

lagoonal (Ardolino and Franchi, 1996).

With respect of the fruit-like receptacles, the

characters preserved are consistent to those of living

Nelumbo and with those described by Upchurch et al.

(1994) of the Quantico locality of the Lower Creta-

ceous Potomac Group of Virginia. Nelumbo produces

conical receptacles where the fruits (or disseminules

for Collinson, 1980) are free from each other, and

each of the fruits has a small respiratory pore

(Cronquist, 1981; Takhtajan, 1996; Plate IID,E).

Unfortunately, they are not in organic connection

with the leaves, and it would be incorrect to include

the receptacles within N. puertae. Until their relation-

ships with N. puertae can be determined, we prefer to

place them within the genus Nelumbo without a

species assignation.

Nelumbonaceae is considered as a bbasalQ eudicotfamily. Although it is not our intention to discuss at

length its taxonomic position, it is fair to mention

that the placement of Nelumbonaceae within the

order Nymphaeales was originally disputed by

Takhtajan (1970) who studied the morphology of

its pollen grains. Unexpectedly, based on molecular

data, the family Nelumbonaceae appears to be a

member of the Proteales clade formed by Protea-

ceae, Platanaceae and Nelumbonaceae, three fami-

lies considered in the past to be only distantly

related (Soltis et al., 2000, Qiu et al., 2000, Nandi

et al., 1998). Within the Proteales clade, Nelumbo-

naceae is the sister group of the pair Proteaceae+-

Platanaceae.

The fossils described here are of particular

significance. The discovery of leaves and fruits of

Nelumbo in Patagonia provide a new dimension to

our knowledge of the fossil record for the family

Nelumbonaceae and its present and past distribution.

As mentioned previously, today the family is

restricted to tropical and subtropical areas of North

and Central America and Asia and Australia.

However, this report supports the idea that, in the

past the family was more widespread. The fossils

described in this contribution constitute the only

fossil record for the family Nelumbonaceae in the

Southern Hemisphere.

Acknowledgements

This work was supported by the Franklin Research

Award of the American Philosophical Society awarded

to M. A. Gandolfo. In particular, we are in debt with

Ana Archangelsky and Gerardo Cladera that provided

information and the stratigraphic column of the local-

ity, and Pablo Puerta, Cynthia Gonzalez, Jose Carbal-

lido, Mariano Caffa, Maximo Dellocca of the Museo

Paleontologico E. Feruglio for assistance during

M.A. Gandolfo, R.N. Cuneo / Review of Palaeobotany and Palynology 133 (2005) 169–178 177

fieldwork. Harufumi Nishida helped with the Japanese

literature consulted. We thank Kathleen B. Pigg and an

anonymous reviewer for their useful comments on the

original manuscript. We also want to express our

gratitude to the Curator of the L. H. Bailey Hortorium

Herbarium (BH), Cornell University for allowing the

use of the collection. Donna and Tim Schender

provided leaves and fruits of extant Nelumbo.

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