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Palaeontologia Electronica
http://palaeo-electronica.org
PE Article Number: 11.3.13A
Copyright: Palaeontological Association October 2008
Submission: 29 December 2007. Acceptance: 29 April 2008
Axsmith, Brian J., Escapa, Ignacio H., and Huber, Phillip. 2008. An Araucarian Conifer Bract-scale Complex from the Lower Jurassic
of Massachusetts: Implications for Estimating Phylogenetic and Stratigraphic Congruence in the Araucariaceae. Palaeontologia
Electronica Vol. 11, Issue 3; 13A:9p; http://palaeo-electronica.org/2008_3/153/index.html
AN ARAUCARIAN CONIFER BRACT-SCALE COMPLEX FROM THE
LOWER JURASSIC OF MASSACHUSETTS: IMPLICATIONS FOR
ESTIMATING PHYLOGENETIC AND STRATIGRAPHIC CONGRUENCE
IN THE ARAUCARIACEAE
Brian J. Axsmith, Ignacio H. Escapa, and Phillip Huber
ABSTRACT
The conifer family Araucariaceae has an extensive Mesozoic fossil record, but no
unambiguous megafossils of this group have been described from the Newark Super-
group of eastern North America. A bract-scale complex attributable to Araucaria is
described from the Lower Jurassic Portland Formation of Massachusetts. Although
known from a single specimen, this discovery is significant as the first bona fide
megafossil of the Araucariaceae from the Newark Supergroup and one of the few early
Mesozoic examples from all of North America. Furthermore, this bract-scale complex is
proposed as the earliest known occurrence of Araucaria section Eutacta based on its
wedge-like shape, the centrally placed ovule that was retained at maturity, and lateral
wings. An analysis of the relationship between the most current phylogenetic hypothe-
ses for the sections of Araucaria and the temporal information from the rich fossil
record of the genus indicates low levels of congruence. Clearly, more paleobotantical
and phylogenetic research is needed to provide a robust estimate of this important
conifer family’s evolutionary history.
Brian J. Axsmith. Department of Biology, University of South Alabama, LSCB 124, Mobile, AL 36688, USA,
baxsmith@jaguar1.usouthal.edu
Ignacio H. Escapa. CONICET, Museo Paleontologico Egidio Feruglio, Fontana 140, Trelew, Chubut,
Argentina, iescapa@mef.org.ar
Phillip Huber. GeoScience Books, P.O. Box 1036, Faribault, MN 55021 raregeologybooks@earthlink.net
KEY WORDS: Eutacta; phylogenetics; Mesozoic; Newark Supergroup; Portland Formation
INTRODUCTION
The conifer family Araucariaceae occurs
exclusively in the Southern Hemisphere today, but
it was widely distributed in both hemispheres dur-
ing the Mesozoic (Stockey 1982, 1994; Stockey
and Ko 1986; Hill 1995; Del Fueyo and Archangel-
sky 2002; Kunzman 2007a, 2007b). The mid-
Jurassic through Cretaceous record is particularly
rich in many regions, but early Mesozoic occur-
rences are uncommon and often ambiguous, espe-
cially in North America. Rare araucarian ovulate
cones and cone scales have recently been
described from the Late Triassic Chinle Formation
of Arizona and New Mexico (Axsmith and Ash
AXSMITH ET AL.: JURASSIC ARAUCARIAN
2
2006) and the Lower Jurassic Moenave Formation
of Utah (Tidwell and Ash 2006). Putative araucar-
ian megafossils have also been reported from the
Newark Supergroup of eastern North American
(e.g., Wanner and Fontaine 1900; Bock 1954), but
these are not generally accepted as convincing
evidence for the family (Cornet 1986; Stockey
1994; Axsmith and Ash 2006). It is in this context
that an unambiguous araucarian bract-scale com-
plex from the Lower Jurassic Holyoke Dam locality
of Massachusetts is described. Although known
from a single specimen, this discovery is significant
as the first certain megafossil of the Araucariaceae
from the Newark Supergroup and one of the few
early Mesozoic examples from all of North Amer-
ica. Furthermore, this bract-scale is similar to those
of Araucaria section Eutacta and may represent
the earliest known representative of this clade.
The extant species of Araucaria are com-
monly placed taxonomically among four sections;
Eutacta, Intermedia, Araucaria (= Columbea) and
Bunya (Endlicher 1847; Wilde and Eames 1952).
Several characters, including aspects of the bract-
scale morphology, are often used to distinguish
among the sections. These classic delimitations
are generally concordant with more recent molecu-
lar phylogenies (Gilmore and Hill 1997; Setoguchi
et al. 1998; Kunzmann 2007b); however, relation-
ships within the genus (i.e., between the sections)
remain unclear. Fossil species of Araucaria (as
well as Araucarites – see below) are represented
by impression/compression remains of vegetative
and reproductive organs from the Jurassic of the
Northern and Southern Hemispheres, and most of
these have been assigned to one of the sections of
Araucaria (see Stockey 1982; Del Fueyo and Arch-
angelsky 2002). These early records, along with
the fossil described here, allow for estimates of
phylogenetic/stratigraphic congruence, such as
that presented below.
MATERIALS AND METHODS
Nomenclatural Considerations
Historically, vegetative shoots ascribed to the
Araucariaceae have sometimes been described
using the generic name Araucarites; however, Zijl-
stra and Konijnenburg-van Cittert (2000) proposed
that this name be restricted to megasporangiate
cones and isolated bract-scale complexes. This
practice is acceptable when applied to early repre-
sentatives of the Araucariaceae with uncertain sec-
tional affinities or to more poorly preserved
material. However, the name Araucarites should
not be applied to material assigned to a particular
section, as such a determination must indicate
affinity with the genus Araucaria.
The generic name Araucaria is used here with
reference to the new Holyoke specimen, but it is
not formally named as a new species as it is repre-
sented by a single, isolated specimen that cannot
be definitively diagnosed from several Araucaria
section Eutacta bract-scale complexes. Some
other comparative taxa are referred to here as
Araucarites based on the original descriptions until
a thorough revision of the fossil record of the family
is available.
Locality and Geological Setting
The new Araucaria bract-scale complex was
collected from the South Hadley Falls Member of
the Portland Formation (Olsen et al. 2003, 2005),
from strata that crop out just below the dam in the
middle of the Connecticut River at Holyoke, Mas-
sachusetts (Figure 1). The Portland Formation con-
tains the youngest rocks of the Hartford Group,
Newark Supergroup and has been dated by
palynostratigraphy (Cornet and Waanders 2006)
and vertebrate biochronology (Lucas and Huber
2002) to span the Hettangian, Sinemurian, and
possibly Pliensbachian stages of the Early Juras-
sic. The South Hadley Falls Member is of Hettan-
gian age, and consists of gray and red lacustrine
shale and siltstone beds arranged in ~20 m thick
cycles, each of which has been interpreted as the
depositional product of the 20 Ky precessional
cycle (Olsen 1986; Olsen et al. 2003, 2005). At
Holyoke Dam and nearby exposures, the South
Hadley Falls Member contains an abundant,
allochthonous, low-diversity flora strongly domi-
nated by conifer branches with common equi-
setalian stem fragments and some cycadeoid
leaves. Other fossils from the South Hadley Falls
Member include fossil insect larvae (Huber et al.
2003), fragmentary fishes, and occasional thero-
pod dinosaur and crocodilian footprints (Olsen et
al. 2003, 2005). Although this locality has been
known for some time (see discussion in McDonald
1992) no detailed systematic work on the plant
material has been published despite the use of the
conifer shoots in paleoecological studies (e.g., Cor-
net and Waanders 2006). Based on the abundance
of Pagiophyllum and Brachyphyllum morphotype
leafy shoots, ovulate cone scales putatively similar
to Hirmeriella, and the dominance of dispersed
Classopollis pollen (Cornet et al 1973) at this and
coeval localities in the Hartford and Deerfield
Basins, most of this material has been assumed to
PALAEO-ELECTRONICA.ORG
3
Massachusetts
HD
HD :
Massachusetts
Vermont
Connecticut
Conglomerate
Fluvial Strata
Basalt
Lacustrine strata
Basament inlier
Holyoke Dam
15 km
400 km
References
N
Figure 1. Map of Newark Supergroup basins (left) and inset detail of Hartford Basin (right). Location of Holyoke Dam locality
(HD) indicated by arrow. Figure modified from Olsen, Whiteside, and Huber (2003).
AXSMITH ET AL.: JURASSIC ARAUCARIAN
4
represent the important Mesozoic conifer family
Cheirolepidiaceae. However, it has been previ-
ously suggested that some of the shoots may be
araucarian (Cornet and Waanders 2006). In fact,
the cuticular preservation of the leafy shoots is
poor (contra Cornet and Waanders 2006) and the
familial affinities of any of the specific leafy shoots
remain uncertain. An ongoing restudy of ovulate
cone scales and pollen cones from Holyoke Dam
does suggest dominance of the Cheirolepidiaceae;
however, the bract-scale complex described here
provides unequivocal macrofossil evidence of the
Araucariaceae in this flora.
Fossil Preparation and Study Methods
No special preparation of the fossil was per-
formed; however, it was immersed in ethanol to
increase contrast for photography (Figure 2.1). The
specimen exhibits considerable relief, which
makes it possible to delimit several important char-
acters under incident light, such as the position of
the ovule and the presence and orientation of the
wings. A thin layer of carbonaceous material is
present, but the application of the transfer tech-
nique was not attempted because it is the only
specimen, and attempts at transferring other fossils
from this locality were unsuccessful. The fossil
bract-scale complex specimen (# J 1430) will be
deposited in the collections of the Paleobotany
Division of the Natural History Museum and Biodi-
versity Research Center at the University of Kan-
sas. Comparative material of extant Araurcaria
species was examined and photographed in the
L.H. Bailey Hortorium of Cornell University.
Phylogenetic and Statistical Methods
Although the value of fossils in phylogenetic
reconstruction is controversial, the time of first
appearance of groups in the geological record is
widely used in calibrating phylogenies based on
molecular evidence. Furthermore, fossils are
sometimes used to describe the level of agreement
between the temporal sequence of taxa in the fos-
sil record and the order of branching on phylogenic
trees. Such congruence studies normally employ
one of two approaches; 1) methods that evaluate
the number of inconsistencies between the phylog-
eny and temporal data (Norell and Novaceck 1992;
Huelsenbeck 1994), and 2) methods that measure
calibrated “ghost lineages” or phylogenetically
implied gaps (Sidall 1998; Wills 1999; Pol and
Norell 2001) by which the absolute temporal dis-
parity is evaluated (Brochu and Norell 2000). In this
paper, the second approach is used to evaluate the
congruence between the current phylogenetic
hypotheses for the sections of Araucaria and the
temporal information from the fossil record of the
genus. Specifically, we used the analysis of Set-
Figure 2. Fossil and extant Araucaria bract-scale complexes. (1) Fossil Araucaria bract-scale complex from the Lower
Portland Formation of Massachusetts. Note the central seed-bearing region and lateral wings. # J 1430. (2) Bract-
scale complex of extant Araucaria heterophylla. (From specimen # BH 2732.) Note similarity to fossil in Figure 1.1.
Scale bars = 1.0 cm.
PALAEO-ELECTRONICA.ORG
5
oguchi et al. (1998), which produced one hypothe-
sis of relationships between the sections of
Araucaria and that of Gilmore and Hill (1997),
which produced two topologies–one perfectly con-
gruent with that of Setoguchi et al. (1998) in terms
of the sectional relationships and one different. The
Gilmore and Hill (1997) study used fewer Araucaria
species, but it included at least one species of
each monophyletic section of the genus (sensu
Setoguchi et al. 1998).
The first appearance datum (FAD) used for
each taxon is based on the literature for sections
Araucaria, Bunya, and Intermedia (e.g., Stockey
1994; Setoguchi et al. 1998). The FAD for section
Eutacta is based on the Holyoke bract-scale com-
plex described here. The Manhattan Stratigraphic
Measure (MSM) method originally proposed by
Sidall (1998) and later modified by Pol and Norell
(2001) designated as MSM* was utilized. In addi-
tion, the age uncertainty was considered with the
randomization approach for age ranges (Pol and
Norell 2006) using the MSM* to calculate the strati-
graphic fit for each of 1000 random replicates. This
analysis was performed using the phylogenetic
analysis software program TNT (Goloboff et al. in
press).
DESCRIPTION AND COMPARISONS
The Holyoke Araucaria bract-scale complex is
about 1.67 cm long, with only the most distal por-
tion missing (Figure 2.1). It is 0.8 cm wide at the
base and expands distally to 1.37 cm wide near the
apex. A single obovate seed, which is represented
by a distinct thickening in the middle of the com-
plex surface, is enclosed by the complex tissues.
On each side of the central seed-bearing zone is a
0.2 cm wide wing with longitudinal striations. The
seed-bearing zone is well defined due to the pres-
ence of two deep, longitudinal grooves that delin-
eate it from the wings. The distal part of the
complex shows a distinct thickening that is con-
cave toward the base. No ligule (free portion of the
ovuliferous scale) is visible, but this is probably due
to preservational factors. The basal part of the
complex shows three lobes; the central lobe is pro-
duced by the base of the seed-bearing region, and
the smaller lateral lobes are formed from the bases
of the wings.
The new Araucaria bract-scale complex is
most similar in shape, wing morphology, and ovule
disposition to those of section Eutacta of Araucaria.
Among extant species, it is particularly similar to
those of Araucaria heterophylla (Figure 2.2). In
comparison, bract-scale complexes of section
Araucaria (= Columbea) have a nut-like shape and
entirely lack wings, and those of section Bunya
have thicker woody wings, and the seed is shed
from the complex. Finally, the bract-scale wings of
section Intermedia are broader and thinner than
those of Eutacta.
Araucaria section Eutacta has a rich fossil his-
tory, and during the Mesozoic it was present in
both hemispheres (Stockey 1982; Hill and Brodribb
1999); however, most records are based on vege-
tative remains (Hill and Brodribb 1999). One
exception is Araucarites stockeyi from the Lower
Jurassic of Utah, which is suggested as a repre-
sentative of section Eutacta by Tidwell and Ash
(2006) based on the wedge-shaped bract-scale
complex with a short apical point, and the centrally
placed ovule that was apparently retained at matu-
rity. Although this may indeed be the earliest record
of section Eutacta, the wings of the Holyoke bract-
scale complex described here make it an even
more convincing representative. Araucarites phil-
lipsii from the Jurassic of Yorkshire (Kendall 1949;
Harris 1979; Van Konijnenburg-van Cittert and
Morgans 1999) has been referred to section
Eutacta based on characters of the ovuliferous
cones and seedlings (Stockey 1982). Bract-scale
complexes of Araucarites phillipsii are similar in
shape and size to the Holyoke Araucaria; however,
they do not show the clear delimitation of the seed-
bearing zone, and no distal thickening has been
observed. Araucarites baqueroensis and Araucar-
ites minimus from the Cretaceous of Argentina
have also been included in section Eutacta (Del
Fueyo and Archangelsky 2002). The bract-scale
complexes of these species are similar in general
morphology to the Holyoke specimen, but Araucar-
ites minimus is much smaller (nearly half of the
size) while Araucarite baqueroensis is consider-
ably larger. Another well-known species is Araucar-
ites brodiei, from middle Jurassic of Oxfordshire
(Cleal and Rees 2003). However, these fossils are
notably larger than the Holyoke Araucaria bract-
scale complex, the seed occupies the lower part of
the complex rather than the middle, and no wings
are present. In addition, the seed in Araucarites
brodiei is born in a depression on the bract-scale
but is not embedded in its tissues (Cleal and Rees
2003). In fact, this character suggests that this spe-
cies may not be close to Araucaria at all, as the
seeds of this genus are embedded in the tissues of
the bract-scale complex at least during some onto-
genetic stage. Based on these comparisons, it is
proposed that the Holyoke Araucaria bract-scale
complex provides enough features to be confi-
AXSMITH ET AL.: JURASSIC ARAUCARIAN
6
Figure 3. Stratigraphic fit of phylogenetic hypotheses for Araucaria sections. FADs are considered with uncertainty
intervals (entire lines); the dashed lines represent ghost lineages. (1) Stratigraphic adjustment for the classical
hypothesis (based on Gilmore and Hill 1997, Figures 3a, 4a, 4c and 4d; Setoguchi et al. 1998, Figure 1). (2) Strati-
graphic adjustment for alternative hypothesis (from Gilmore and Hill 1997, Figures 3b, 4b). (3) Frequency histogram
of the difference in MSM*values of both hypotheses obtained in each replicate of the randomization procedure. The
MSM* difference between the classical and alternative hypothesis is positive (or zero) in all replicates indicating that
the alternative hypothesis score is equal to or lower than the classical one in stratigraphic fit (represented by
MSM*value).
PALAEO-ELECTRONICA.ORG
7
dently assigned to Araucaria section Eutacta and
represents the earliest record of this section.
DISCUSSION
Phylogenetic/stratigraphic Congruence
Measures of phylogenetic/stratigraphic con-
gruence for the Araucariaceae utilizing the statisti-
cal tests, fossil taxa, and phylogenies described
above were surprisingly low.
The range of MSM* values obtained for the
sectional relationships within Araucaria is 0.51-
0.84 for the classical phylogenetic hypothesis
(Gilmore and Hill 1997, Figures 3a, 4a, 4c and 4d;
Setoguchi et al. 1998, Figure 1) (Figures 3.1 and
3.3), and 0.5-0.67 for the alternative one (Gilmore
and Hill 1997, Figures 3b, 4b) (Figures 3.2 and
3.3). These relatively low values are probably
related to the incongruence between the FAD
ranges of the sections and their position on both
tree topologies (Figure 3.1-3.2), generating a long
temporal ghost for some lineages. Alternatively, the
broad MSM* range obtained with the FAD dates
used in this study may reflect the high degree of
uncertainty in determining the precise ages of most
of the sections. For example, the Lower Creta-
ceous FAD range for Section Intermedia covers
about 43 million years. Another potential problem
is the lack of fossil taxa as terminals in the phyloge-
netic analyses utilized.
Conclusion
Ideally, a clade with a rich fossil record and a
well-resolved phylogenetic hypothesis should show
high levels of congruence among these data sets.
The Araucariaceae would seem to satisfy the
requirements for high phylogenetic/stratigraphic
congruence, as the family is considered to have a
particularly long and rich fossil record, and sub-
stantial living diversity to provide abundant data for
phylogenetic studies. Nevertheless, the analysis
presented here indicates that much more research
is needed, probably in both areas, to provide a
more consistent estimate of this important conifer
family’s evolutionary history. This will most likely
entail the continued discovery and description of
new fossils as well as a critical re-evaluation of
known fossil taxa. In addition, it is likely that phylo-
genetic studies based only on extant taxa underes-
timate the true complexity of araucarian phylogeny,
as the fossil record indicates high levels of extinct
diversity, including completely extinct sections
(e.g., Yezonia) with unique character combinations
(Ohsawa et al. 1995). The inclusion of well-pre-
served and reconstructed fossil taxa in a combined
analysis of the Araucariaceae should provide new
data regarding those parts of the phylogeny cur-
rently represented by ghost lineages leading to a
more robust phylogenetic hypothesis and improved
phylogenetic/stratigraphic congruence.
Regardless of the causes of the current phylo-
genetic/stratigraphic incongruence for the Araucar-
iaceae, the bract/scale complex described here is
significant as the first bona fide megafossil of the
Araucariaceae from the Newark Supergroup and
one of the few early Mesozoic examples from all of
North America. This fossil is also significant as the
earliest record of Araucaria section Eutacta.
Although plant fossils have been known from the
Newark Supergroup for many years, there is little
question that it remains an underutilized source of
information regarding early Mesozoic plant evolu-
tion.
ACKNOWLEDGEMENTS
The authors are grateful to the staff of the L.H.
Bailey Hortorium at Cornell University for access to
herbarium specimens. This research was sup-
ported by a National Science Foundation Grant
(EAR-0105476) to B.J.A.TNT is freely available,
thanks to a subsidy from the Willi Hennig Society.
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