CSIRO PUBLISHING
www.publish.csiro.au/journals/asb
Australian Systematic Botany, 23, 131–140
Calamoid fossil palm leaves and fruits (Arecaceae: Calamoideae)
from Late Eocene Southland, New Zealand
Samuel J. Hartwich A, John G. Conran A,D, Jennifer M. Bannister B, Jon K. Lindqvist C
and Daphne E. Lee C
A
Australian Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences,
Benham Bldg DP312, The University of Adelaide, SA 5005, Australia.
B
Department of Botany, University of Otago, PO Box 56, Dunedin, New Zealand.
C
Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand.
D
Corresponding author. Email: [email protected]
Abstract. Late Eocene prickly-leaved and scaly-fruited palm macrofossils are described from Pikopiko, Southland,
New Zealand, and compared with extant Arecaceae: Calamoideae. Lamina prickles and scaly fruits support affinities to the
subfamily and tribe Calameae and possible association with the extant genus Calamus. Because isolated calamoid leaf
fragments and fruit are difficult to determine precisely, the fossils are placed into a new form genus (Calamoides) for the
leaves and the existing form genus Lepidocaryopsis for the fruits. These represent the first calamoid-like palm
macrofossils from New Zealand and suggest a subtropical to tropical palaeoclimate at far southern latitudes in the Late
Eocene and an early, widespread vicariant Gondwanan distribution for the subfamily.
Introduction
The palms (Arecaceae) consist of ~190 genera and ~2360 species
(Dransfield et al. 2008; Mabberley 2008), primarily from the
tropics and subtropics, with outliers in warm- to cool-temperate
areas, and are distinctive for their often stout, generally
unbranched, woody stems with a terminal crown of large,
evergreen palmate, pinnate or bipinnate leaves (Dransfield and
Uhl 1998).
Australasia has a relatively poor monocot fossil record,
with most fossils described only recently, providing some
information on biogeography, but less on phylogeny
(Greenwood and Conran 2000; Pole 2007b; Conran et al.
2009). Australia and New Zealand both experienced climate
change during the Cenozoic, with warmer conditions in the
Eocene and Early–Middle Miocene, with possible brief
cooling in the Early Oligocene and major cooling in New
Zealand to cool temperate from the Late Miocene (Lee et al.
2001). Accordingly, the fossil history of Australasia’s monocots
is useful for understanding the past biogeography and climate
of New Zealand because many monocot groups show strong
climatic responses, in particular the palms.
The in situ remains of a Late Eocene fossil forest near
Pikopiko, Southland, New Zealand, are exposed within alluvial
sediments of the Beaumont Formation (c. 35 million years ago;
Lee et al. 2009). Fossil trees from the forest are rooted in
mudstone and coal, spaced 2–5 m apart and distributed over a
30 120-m area in calcite-cemented concretions up to 60 cm in
diameter and 80 cm high. The in situ trees are a part of a
sedimentary succession of a 100+-m thick floodplain swamp
and river-channel assemblage. The sediments formed from the
CSIRO
31 May 2010
fill of a south-flowing river channel that eventually smothered
the growing forest, uprooting and toppling smaller trees.
Macrofossils at the site included monocot leaves with
attached spines or prickles and scale-covered fruits, both with
affinities to Arecaceae subfamily Calamoideae (rattans).
Molecular studies have confirmed Arecaceae as
monophyletic; however, relationships within the family remain
ambiguous (Asmussen et al. 2006). There are five subfamilies
presently recognised, namely the Arecoideae, Ceroxyloideae,
Coryphoideae, Nypoideae and Calamoideae (Dransfield et al.
2005, 2008; Asmussen et al. 2006). Calamoideae are the secondlargest subfamily with 21 genera and ~650 species with a global
distribution, although with the greatest diversity in the wet
tropical forests of South-east Asia (Mathew and Bhat 1997;
Baker et al. 2000a, 2000b; Baker and Dransfield 2008). The
subfamily sits as sister to the remainder of the Arecaceae in
molecular analyses (Baker and Dransfield 2000; Baker et al.
2000b, 2000c; Asmussen and Chase 2001; Hahn 2002; Asmussen
et al. 2006) and anatomical evidence also supports this (Uhl
and Dransfield 1987).
The largest member of the Calamoideae is Calamus, a genus
of spiny climbing palms (rattans) with ~375 species, primarily
in Asia, and extending to Africa, Malesia, Micronesia and
Australia (Mathew and Bhat 1997; Mabberley 2008), and
represented in eastern Australia by eight species, of which five
are endemic (Dowe 1995; Cooper and Cooper 2004). The genus
has relatively small (for palms), pinnate leaves with regularly or
irregularly inserted, often fasciculate, reduplicate, lanceolate or
sigmoid leaflets, usually with a prominent adaxial midrib and
often with subsidiary abaxial ribs and sometimes prominent
10.1071/SB09027
1030-1887/10/020131
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Australian Systematic Botany
spines on the ribs and/or margins (Tomlinson 1961). Some
species also bear spines or prickles along the laminal veins
(Cooper and Cooper 2004), although laminar spines are not
unique to the calamoid palms (Dransfield et al. 2008).
The present study compares the Pikopiko palm leaves and
fossilised fruit with extant Calamus and other Calamoideae, to
determine whether the fossils agree with placement into this
subfamily, as well as to broaden the knowledge of New
Zealand’s Arecaceae diversity during the Late Eocene and the
implications for past climates.
Materials and methods
The fossils were collected from alluvial sediments of the
Beaumont Formation on the eastern bank of the Waiau River
near Pikopiko, 6 km north of Tuatapere, Southland, South
Island, New Zealand (46060 S, 167410 E; NZGS Map
1 : 250 000 Sheet 24 (Invercargill), 1st edition, S22–8037
(Wood 1966)). Most of the palm material came from a loose
40-cm-diameter mudstone block that included fruits and leaves.
Other blocks on a bedding plane were split and extracted fossils
wrapped in cling-film to prevent drying and flaking. Surface
debris was removed and leaf edges freed with wet, fine
paintbrushes and needles. Leaves were photographed with a
Nikon D80 SLR digital camera (Nikon, Tokyo). Specimens
are at the Geology Museum (OU), University of Otago,
Dunedin, New Zealand.
The fossil carbonised palm-leaf material was in fragments,
with some being very small and difficult to clear. When the
fragments were removed from the fine sediment with fine forceps
or paintbrushes, indentations could be seen where laminal
prickles had pressed into the fresh sediment. The fragments of
carbonised material were placed in a watch glass and soaked for
10 min in 50% nitric acid, then washed in tap water. The
fragments were placed in a cell strainer (70-mm mesh) and
soaked in 10% KOH until small pieces of cuticle were
released from the debris; if these pieces were too dark, dilute
bleach was used to lighten them to pale brown. Once cleared, the
cuticle fragments were rinsed in reverse osmosis (RO) water,
mounted on microscope slides in warmed phenol–glycerin jelly
and photographed with a Leica DMR microscope and Leica DC
digital camera (Leica Microsystems, Wetzlar, Germany) under
transmitted and Nomarski DIC lighting.
Leaf material of Calamus aruensis Becc., C. australis
Mart., C. caryotoides Mart., C. moti. F.M.Bailey, C. muelleri
H.Wendl., C. radicalis H.Wendl. & Drude, C. vitiensis Warb. ex
Becc. and C. warburgii K.Schum was obtained for comparison
from the Australian Tropical Herbarium, Atherton, Queensland
(QRS), and to obtain an idea of the degree of inter-species
cuticular variability. Adaxial and abaxial cuticles for each
extant species were mounted on scanning electron microscope
(SEM) stubs, sputter-coated and photographed with an XL20 at
SEM at 1500 magnification. In addition, cuticles were
prepared in a 1 : 1 mixture of 100% ethanol and 25% hydrogen
peroxide at 98.2C until cleared and then cleaned with finegauge needles under an Olympus SZ11 dissection microscope
(Olympus Corporation, Tokyo) to remove debris. Cleaned
cuticles were agitated gently in RO water for 1–2 min to
remove further debris, stained for 60 s in 0.5% aqueous crystal
S. J. Hartwich et al.
violet, re-agitated in RO water until dye leaching ceased,
mounted in phenol–glycerine jelly and photographed with a
Kodak DC4800 3.1 megapixel digital camera (Eastman Kodak
Company, Rochester, NY).
Comparisons between the macrofossils and Calamus
cuticles were based on both the stained cuticles and SEM
micrographs. In addition, the fossils were compared with
published descriptions of leaves and fossil Calamoideae fruits
in Stur (1873), Meschinelli and Squinabol (1892), Berry (1929),
Chandler (1957), Tomlinson (1961), Weyland et al. (1966) and
Schaarschmidt and Wilde (1986). Because the Pikopiko leaf
fragments and fruits are separate, despite being in the same
block, they are treated here as separate form genera; however,
they are discussed together in the light of their combined
similarities to calamoid palms.
Fossil palm-leaf fragments that are pinnate, or of uncertain
attachment, are generally placed in the form genera Phoenicites
Broign. or Amesoneuron Göppert, respectively (Read and
Hickey 1972). Some Cenozoic palm fossils have been placed
previously into Calamus (e.g. Meschinelli and Squinabol 1892;
Jablonsky 1914–1915; Chandler 1957), virtually all on gross
morphology alone. However, even though there are some
features that can allow placement of organically preserved
palm fossils into Calamoideae, generic recognition beyond this
is, at best, dubious.
The Pikopiko leaf fossil is distinguished by prominent
punctate prickle(?) scars along the leaf veins and, in
combination with its epidermal and stomatal characteristics,
this suggests affinities to subfamily Calamoideae, in particular
tribe Calameae. The cuticles are also very close to those of fossil
palms from Germany assigned to Calamoideae (Weyland et al.
1966; Schaarschmidt and Wilde 1986). However, generic
differentiation in Calameae by using cuticles alone is not
possible and the fossil is different enough from extant and all
named fossil palm genera to warrant status as a new form genus.
Unfortunately, the pre-existing generic name Calamopsis
Heer probably refers to a cycad (Read and Hickey 1972), so
cannot be used for our fossil palm.
Similarly, the fossil fruits are placed in the form genus
Lepidocaryopsis Stur, owing to their highly characteristic,
scale-covered fruits. This is a feature unique to subfamily
Calamoideae (Baker et al. 2000a; Dransfield et al. 2008),
although one that does not generally allow further generic
differentiation (Berry 1929). In addition, the association of the
scaly fruits with prickly leaves strengthens the case for
Calamoideae.
Systematic palaeontology
Order: Arecales Bromhead (1840)
Family Arecaceae Schultz Sch. (1832), nom. cons.
et nom. alt.
Subfamily aff. Calamoideae Beilschm. (1833)
Genus Calamoides S.Hartwich, Conran, Bannister, Lindqvist
& D.E.Lee, gen. nov.
Type: C. pikopiko S.Hartwich, Conran, Bannister, Lindqvist &
Lee
New Zealand fossil calamoid palms
Australian Systematic Botany
Calamoides pikopiko S.Hartwich, Conran, Bannister,
Lindqvist & D.E.Lee, sp. nov.
Type locality: East bank of the Waiau River near Pikopiko,
6 km north of Tuatapere, Southland, South Island, New Zealand.
Type horizon: Beaumont Formation, Late Eocene.
(Figs 1, 2A–D)
Holotype: OU31767 (leaves; Fig. 1A, B); Geology Museum (OU),
University of Otago, Dunedin, NZ. Additional specimens: cuticle
microscope slide (OU31767a, Fig. 2A–D); Department of Botany
(University of Otago).
Generic and species diagnosis
Palm leaves with prominent prickle scars along the leaf veins;
adaxial lamina very uniform, with rectangular and longitudinally
(A)
(C)
133
(B)
(D)
(E )
Fig. 1. Fossil Calamoid palms. (A) Fossil leaves of Calamoides pikopiko (white arrows) and associated Lepidocaryopsis zeylanicus fruits
(black arrows) in situ. (B) Leaf detail of C. pikopiko (OU31767), showing punctiform prickle base scars along veins (arrows). (C–E) L. zeylanicus
scaly fruits: (C) OU31767b, (D) acuminate scale detail, and (E) OU31767c with less well preserved scales. Arrows in D, E indicate scales.
Scale bars = 10 cm (A), 10 mm (B), 5 mm (C, E), and 2 mm (D).
134
Australian Systematic Botany
S. J. Hartwich et al.
(A)
(B)
(C)
(D)
Fig. 2. (A–D) Calamoides pikopiko cuticles (OU31767a). (A) Adaxial surface. (B) Abaxial surface. (C) Prickle base (arrow).
(D) Stomata. Scale bars = 25 mm (A, C, D) and 50 mm (B).
extended epidermal cells, which are similar in shape, but smaller
abaxially; anticlinal epidermal cell walls slightly sinuous;
periclinal epidermal cell walls thin, unornamented; stomata
abaxial, tetracytic; subsidiary cells similar in width to
epidermal cells, terminal pair distinctly smaller than lateral.
Description
Fossils incomplete, unattached, apparently linear–lanceolate
pinna fragments at least 20 cm long and 20–25 mm wide
(Fig. 1A, B). Lamina dorsiventral, apex, base and sheath
unknown (Fig. 1A); margins apparently entire; venation
parallel, midrib apparent, no obvious transverse veins;
punctiform depressions 0.5 mm diam. (prickle base scars),
common along veins of at least the adaxial leaf surface (Fig. 1B).
Adaxial epidermal cells rectangular, longitudinally extended
along leaf axis, 35–60 mm long, 15–25 mm wide (Fig. 2A);
anticlinal walls slightly curved, periclinal walls thin,
unornamented. Abaxial cells rectangular, elongated along leaf
axis; intercostal cells ~35–50 mm long, 10–15 mm wide
(Fig. 2B–D); anticlinal walls straight to slightly curved,
periclinal cells thin, unornamented; prickle bases present
(Fig. 2C). Intercostal bands are wider than costal zones
(greater than field of view; Fig. 2D). Costal cells
longitudinally extended, anticlinal walls thicker than in
intercostal cells; costal zones ~50 mm wide (Fig. 2D).
Stomata abaxial (Fig. 2B–D), ~15–20 mm long and 15–20 mm
wide (Fig. 2D), intercostal, arranged sparsely in longitudinal
rows; lateral subsidiary cells ~30–40 mm long and 8–10 mm
wide; terminal subsidiary cells ~20 mm wide and 10 mm wide
(Fig. 2D); anticlinal subsidiary-cell wall thickness similar to that
of epidermal cells.
Etymology
The generic name is derived from the resemblance of the leaves to
the extant genus Calamus L., and the specific epithet refers to the
fossil site at Pikopiko.
Genus Lepidocaryopsis Stur (1873)
Type: L. westphaleni Stur (1873)
Lepidocaryopsis Berry (1929) nom. illeg. auct. non Stur
Lepidocaryopsis zeylanicus S.Hartwich, Conran, Bannister,
Lindqvist & D.E.Lee, sp. nov.
(Fig. 1C, D)
Holotype: OU31767b (fruit; Fig. 1C, D); Geology Museum (OU),
University of Otago, Dunedin, NZ; paratype: OU31767c (fruit; Fig. 1E);
Geology Museum (OU), University of Otago, Dunedin, NZ.
Type locality: eastern bank of the Waiau River near Pikopiko,
6 km north of Tuatapere, Southland, South Island, New Zealand.
Type horizon: Beaumont Formation, Late Eocene.
Species diagnosis
Fruit ovoid to globose, with an epicarp of elongate, rhomboidal,
apically acuminate, imbricate scales; fruits and scales smaller
than for other Lepidocaryopsis species.
New Zealand fossil calamoid palms
Description
Fruit spheroid to ovoid, 17–19 13–17 mm (Fig. 1C–E),
smooth, scale-covered, longitudinal lines prominent, transverse
lines less obvious; scales elongate, rhomboidal, 3.5–5 1.5–2 mm, imbricate, bluntly acuminate (Fig. 1D).
Etymology
The specific name is derived from the origin of the fossil in
New Zealand.
Discussion
Fossil comparisons
The fossil cuticles are monocotyledonous, based on having rows
of longitudinally oriented stomatal complexes and epidermal
cells; a result of the typically parallel-veined leaves (Fig. 1A).
They also have the monocot features of distinct, paired polar and
lateral subsidiary cells (Pole 2007b) and stomata generally of
more-or-less equal size within the leaf (Dunn et al. 1965;
Conover 1991).
The leaf fragments are typical for palms (Tomlinson 1961;
Dransfield et al. 2008), possessing a dorsiventral lamina and
a more-or-less prominent midrib (Fig. 1). The adaxial epidermis
is uniform (Fig. 2A), with slightly smaller abaxial cells
(Figs 2B–D). There are prickle bases along the abaxial veins
(Figs 1B, 2C). Stomata are apparently absent adaxially, restricted
to intercostal regions abaxially, and the terminal subsidiary cells
are short (Fig. 2D).
When compared with extant Calamus species, laminar
prickles (or their bases) occur not only in the fossil (Figs 1B,
2C), but are also seen in many extant species, although
laminar prickles also occur in some species of Phoenicae and
Cocoseae (Dransfield et al. 2008). Similar prickle scars have
also been seen on the lamina of fragments of Spinopalmoxylon
rhenanum Weyland et al. (1966) that was also associated with
Calamoideae.
The abaxial epidermal cells of the fossil are slightly larger
than those of the examined extant Calamus species, but show
similar wall thickness; the fossil’s anticlinal epidermal cell walls
are less curved or undulate than in modern counterparts, whereas
the periclinal walls are similar in form. The adaxial epidermal
cells appear to be somewhat larger in the fossil than those of
the living Calamus, and are similarly less rectangular than the
abaxial cells of the extant species. Stomata are present adaxially
in some extant Calamus spp., but not in others, and the stomata
of the fossil are similar to those of the extant species, albeit
slightly smaller longitudinally and less frequent. The subsidiary
cells in the fossil are slightly larger, and have thinner anticlinal
walls than those in the extant species examined.
In contrast, these non-undulate cuticles are very close to
those of the Eocene calamoid palm remains described (but not
named) from Messel, Germany by Schaarschmidt and Wilde
(1986), as well as to a lesser degree, some of the cuticular
descriptions and illustrations of epidermal tissues for
Spinopalmoxylon rhenanum in Weyland et al. (1966).
When compared with other extant palm groups, differences
in leaf form and/or cuticular morphology largely rule out all
but the Calamoideae (Tomlinson 1961; Dransfield et al. 2008).
Australian Systematic Botany
135
Arecoideae, Corphoideae: Caryoteae and Ceroxylodeae possess
obliquely extended, rhombohedral, or hexagonal adaxial cell
arrangements. Members of Corphoideae: Sabaloideae have
short, wide terminal subsidiary cells and lateral subsidiary
cells that are wider than the other epidermal cells.
Although members of Nypoideae can possess scattered spinelike laminal ramentae (Dransfield et al. 2008), they also display
transversely extended adaxial cells, characteristic guard cells
and very deep lateral subsidiary cells (Tomlinson 1961), which
the fossil does not. Similarly, spiny leaflets do occur in some
Coryphoideae: Phoeniceae and Arecoideae: Cocoseae
(Dransfield et al. 2008), although here the former possess
isolateral laminae, whereas in the latter the adaxial cells are
rhombohedral, transversely or obliquely extended and there
are prominent abaxial costal bands (Tomlinson 1961).
Within Calamoideae, many genera can also be ruled out on
cuticular morphology. Eremospatha (G.Mann & H.Wendl.)
H.Wendl., Laccosperma (G.Mann & H.Wendl.) Drude,
Lepidocaryum Mart., Mauritia L.f., Plectocomiopsis Becc. and
Raphia P.de Beauv. lack hairs or lamina prickles, whereas
Ceratolobus Blume, Eugeissona Griff., Korthalsia Blume and
Myrialepis Becc. possess hairs on both leaf surfaces. Metroxylon
Rottb. lacks hairs, but has small, sunken, thin-walled strongly
sinuous epidermal cells, whereas Daemonorops Blume,
Plectocomia Mart. ex Blume and Salacca Reinw. lack well
defined terminal subsidiary cells (Tomlinson 1961). Calamus,
although being the closest extant match, nevertheless has
undulating to sinuous cell walls, suggesting that the fossil is
not necessarily in this genus either, making assignment to a
modern genus on cuticle alone impossible.
Although there are many fossil Calamoideae reported (see
reviews in Harley (2006) and Dransfield et al. (2008)), fossil leaf
cuticle for the subfamily is rare. However, the few reported
cases from Eocene and Miocene Europe have also noted
absence or weakness of wall sinuosity, a lack of obvious
modern equivalents and difficulty in separating those modern
genera that they tend to resemble on cuticle alone (Weyland
et al. 1966; Schaarschmidt and Wilde 1986), supporting the
placement of our fossil leaves into a new form genus
Calamoides.
One of the most distinguishing features of the Calamoideae
is the imbricately scale-covered fruit (Baker et al. 2000a) and
the scaly fruits at Pikopiko are a close match. The overlapping
fruit scales are in the right orientation, overlap at the right
places, are the right size and shape (Fig. 3B), and agree with
Calamoideae fruits (Dransfield and Uhl 1998; Dransfield et al.
2008), although they are somewhat different because of lack
of clear transverse lines (Fig. 1C, D). They fall within the fruitand scale-size range of genera such as Calamus (Baker et al.
2000a) and Metroxylon (McClatchey et al. 2006) and the
absence of obvious hairs, prickles or warts is consistent with
Calameae fruits (Fig. 3B), although the elongate scales differ
from all extant Calamoideae.
Stur (1873) created the form genus Lepidocaryopsis for
calamoid fruits from Miocene Germany, defining it on the
possession of rhomboidal, bluntly pointed imbricate scales.
Berry (1929) created the same generic name for a fossil
calamoid fruit from Miocene Colombia (as a later homonym),
noting that the species he described was probably close to the
136
Australian Systematic Botany
S. J. Hartwich et al.
(B)
(A)
(C)
Fig. 3. Calamus features and distribution pattern. (A) C. moti leaves showing prickles along the spiny leaflet veins
(arrow). (B) C. australis fruits showing imbricate scales. (C) Distribution of extant Calamoideae. Scale bars = 20 mm.
Photos J. Dowe (A, B).
South American genera Lepidocaryum, Mauritia and the South
American and African genus Raphia, with which it shared much
larger (~1 cm) scales than are seen in our fossil. The precise
relationships of Lepidocaryopsis are uncertain (Moore 1973),
although the scales are similar to those seen in many Old World
calamoids such as Calamus (Cooper and Cooper 2004) and
Metroxylon (McClatchey et al. 2006). In contrast, other scaly
fossil palm fruits from Europe have also been placed into the
‘mixed’ form taxon Calamus daemonorops (Unger) Chandler
(1957, 1963) which, as currently interpreted, includes a wide
range of vegetative, floral and fruit organs (see Dransfield et al.
2008).
Leaf anatomy is considered to be conservative in palms
(Tomlinson 1961), providing a reliable basis for classification
(Mahabale 1965), although caution is required when placing
fossil palms into modern genera on the basis of leaf
morphology alone (Read and Hickey 1972). Hairs and stomata
are of greater significance for palm taxonomy than is gross leaf
shape alone, as they show less variation, and fruits appear to be
even more conservative (Essig 1999), making these features
helpful in the classification of fossil palms (Mahabale 1965).
Fossil history
Read and Hickey (1972) identified a set of characteristics to
recognise fossil palm leaves. These included plicate leaf blades
and segments, pinnately veined and simple or compound form,
or pinnately or palmately veined palmatifid form, and leaf
segments with a strong midvein bounded on each side by two
orders of parallel veins. However, numerous similarities between
different palm leaves makes further identification on leaf form
within the family difficult, indicating the need for caution when
placing fossil palms in modern genera on the basis of leaf
morphology alone (Read and Hickey 1972); for most fossil
palms, this lack of distinctive gross morphological features
limits categorisation below family (Harley 2006).
New Zealand fossil calamoid palms
Arecaceae have a projected Cretaceous origin (Daghlian
1981; Janssen and Bremer 2004; Bremer and Janssen 2006)
and sit near the base of the higher Commelinoids on the
lineage leading to the grasses (Dransfield and Uhl 1998).
Palms display the richest fossil record of all monocots (Harley
2006; Pan et al. 2006), with fossil fruits dating to the Aptian
(Vaudois-Miéja and Lejal-Nicol 1987) and pollen possibly as
early as the Barremian (Dransfield et al. 2008). In Australasia,
leaves showing affinities to Areciflorae were reported by Pole
(1999) from the Albian, and Late Cretaceous palm stems, leaves,
flowers, fruits and pollen are also known (see reviews in
Greenwood and Conran (2000) and Harley (2006)). However,
although all major palm-fossil categories were present by the
Late Cretaceous (Harley 2006; Dransfield et al. 2008), major
palm radiation appears to have occurred mainly in the
Early Cenozoic, with a gradual increase in diversity through
time (Daghlian 1981; Kvacek and Herman 2004). This
diversification was apparently in response to warming during
the early Cenozoic (Morley 1998; Paul et al. 2007; Harrington
2008), with palm diversity and range contracting again following
cooling in the Oligocene and Late Miocene (Daghlian 1981;
Harley 2006).
Modern calamoid palms show a distribution pattern that
almost precisely fits the tropics (Fig. 3C). The Old World
Calamoideae are centred in South-east Asia, and only Calamus
extends to Australia (Dowe 1995). Fossil Calamoideae pollen is
both widespread and ancient (Harley and Baker 2001), being
reported from Cretaceous Africa (Van Hoeken-Klinkenberg
1964; Salard-Cheboldaeff 1981; Schrank 1994), lower Upper
Cretaceous USA (Pierce 1961), the Cretaceous–Cenozoic
boundary Borneo (Muller 1970) and Palaeocene China (Song
et al. 2004). Calamoid macrofossils with affinities to Calamus or
Daemonorops Blume and from a range of organs have been
described from Cenozoic Europe under a variety of names,
mostly as species of Calamus (see reviews in Harley 2006;
Pan et al. 2006; Dransfield et al. 2008) and calamoid fruits are
also known from Miocene Columbia (Berry 1929).
Calamus and/or Metroxylon pollen has been recorded in
Australia and New Guinea (as Dicolpopollis Pfanz) from the
Middle Eocene to the Miocene (Truswell et al. 1987; Macphail
et al. 1994, 1999) and although Oligocene and later Calamus
records might be explained by dispersal owing to close
proximity with South-east Asia (Morley 1998), earlier fossil
records of Calamus favour a Gondwanan origin (Baker and
Dransfield 2000). The presence of Calamus-like pollen in
Australia from at least the Middle Eocene supports a
vicariance explanation, as does this Late Eocene New Zealand
Calamoides and Lepidocaryopsis record, as New Zealand was
isolated from Asia at this time (Lee et al. 2001), making dispersal
less likely. However, Calamus in Australia represents three of
the major lines of evolution seen within the genus (Baker et al.
2000b), suggesting that Calamus either entered Australia
following several radiation events, or that members of at least
one of the lines may be relictual, representing Gondwanan
persistence.
Fossil palms are known in New Zealand from the Late
Cretaceous through to the Pliocene, with Cretaceous pollen of
Trichotomosulcites Couper (1953) and a possible palm fruit
Australian Systematic Botany
137
(Mildenhall 1968), Miocene Rhopalostylis H.Wendl. & Drude
(Arecoideae) pollen (Couper 1952) and Pliocene leaves (Oliver
1928). Rhopalostylis is now represented in New Zealand by
the only extant palm there, R. sapida H.Wendl. & Drude,
which is also the most southerly distributed palm (Wardle
1991). Other New Zealand palm fossils include pollen of Nypa
(= Spinozonocolpites prominatus (McIntyre) Stover and Evans
1973) from the Eocene to Late Oligocene (Pocknall 1989, 1990;
Raine et al. 2008), Miocene and Pliocene endocarps of Cocos
zeylandica Berry (Berry 1926; Ballance et al. 1981), as well as
fronds, fruits and flowers of Phoenicites zeelandica (Ettingsh.)
Pole (1993).
Dicolpopollis cf. D. metroxylonoides Khan pollen attributed
to either Metroxylon (Khan 1976) or Calamus (Truswell et al.
1985) is also reported from Early Miocene New Zealand
(Mildenhall and Pocknall 1989), meaning either that there was
rapid introduction and spread of the Calamoideae into Australia/
Zealandia from South-east Asia (Morley 1998), or that the group
has a Gondwanan origin (Muller 1970; Truswell et al. 1987;
Baker and Dransfield 2000).
Because the fossilised scaly fruits at Pikopiko were within a
few centimetres of the prickly palm-leaf fossils in the same
sediment block (Fig. 1A), it is a reasonable assumption that
this type of fruit, characteristic of Calamoideae palms, belong
to the palm-leaf fossils. Even though a reasonable correlation
exists between the fossil palm and Calamus, the stomatal
morphology has some differences and stomatal undulation and
frequency were higher in all examined extant Calamus species
than in Calamoides, suggesting placement beyond Calamoideae
is not possible for the fossil.
Palaeoclimatic implications
Previous studies in New Zealand and Australia have found
evidence for increased Eocene temperatures at high latitudes
(Greenwood and Christophel 2004), including mesothermal
rainforest fossils from Tasmania (Pole 2007a) deposited
during or close to the warmest known interval of the Cenozoic
(Zachos et al. 2001). These temperatures were due to the absence
of Southern Ocean circumpolar currents and fronts, resulting in
subtropical water flowing to high latitudes (Nelson and Cooke
2001). For example, the mangrove palm Nypa once occurred
in New Zealand and southern Australia, but is represented
today by a single, tropical Indo-Pacific species N. fruticans
Wurmb. confined to within ~18 of the equator; a good
indication of much warmer, high-latitude palaeoclimates (Pole
2007a). Similarly, Carpenter et al. (2007), Pole (2007a) and
McLoughlin et al. (2008) found other warm-growing taxa in
Tasmania, including Lauraceae. This family reaches its present
limits near the southern margin of mainland eastern Australia
and at similar latitudes in New Zealand, with low diversity in
both locations. The world’s southern-most extant palm
(Rhopalostylis sapida) is found as far south as 44S in New
Zealand (Parsons 2007) and this is accepted as the southern-most
modern limit for palms (Endt 1998), although there is evidence
of Rhopalostylis-like palms from Early Oligocene Antarctica
(Thorn 2001).
138
Australian Systematic Botany
The Pikopiko fossil forest grew at a palaeolatitude of ~50S
in an oceanic setting and provides valuable information on
southern hemisphere terrestrial climates in the latest Eocene
(Lee et al. 2009). The Pikopiko forest included a great
diversity and abundance of ferns (six fern macrofossils,
including Cyclosorus Link, Todea Willd. ex Bernh., and 20
microspore fossils), implying that ferns dominated the
understorey, much as they do in modern New Zealand
rainforests (Wardle 1964, 1991). The pollen record at
Pikopiko suggests a warm climate, indicated by the presence
of Cupanieidites Cookson & Pike (cf. Sapindaceae: Cupania L.),
Malvacipollis Harris (cf. Euphorbiaceae: Austrobuxus Miq.)
and a diverse range of epiphyllous fungi including Asterina
Lév., Entopeltacites Selkirk, Callimothallus Dilcher ex Janson.
& Hills, Meliolinites Selkirk ex Janson. & Hills, Quilonia K.P.
Jain & R.C.Gupta and Trichopeltinites Cookson. Various other
microthyraceous shields, fungal germlings, and an assortment
of spores, setae and hyphae also indicate that the prevailing
climate was humid, and at least warm temperate (Bannister
et al. 2003).
The Eocene was a period of increasing temperature
worldwide, with fossil floras and marine paleoclimate
interpretations indicating tropical areas at high northern
hemisphere paleolatitudes (Berggren and Hollister 1974;
Wolfe 1978; Kvacek et al. 2004). Changes in these floras also
indicate abrupt decreases in paleotemperatures near the
Eocene–Oligocene boundary, supported by oxygen isotope
ratios, foraminiferal assemblages and other data (Daghlian
1981), although there are conflicting isotopic and biotic marine
data around New Zealand that suggest waters there were warmer
(Adams et al. 1990; Hornibrook 1992). There were also
changes associated with continental movements throughout the
Cenozoic, leading to the initiation of circumpolar oceanic
circulation patterns (Berggren and Hollister 1974; Daghlian
1981). This then caused steepening latitudinal temperature
gradients from the equator to the poles (Wolfe 1978), leading
to cooling and a major climate change in Oligocene Australasia
(Hornibrook 1992; Crisp et al. 2004).
The presence of Calamus-like palms in New Zealand during
the Late Eocene indicates that the temperature was warm
enough to allow tropical or subtropical plants to grow at
latitudes of ~50S, because modern calamoid palms grow in
high-rainfall tropical or subtropical areas with average annual
temperatures mostly >15C (Xu et al. 2000; Zeng et al. 2000;
McClatchey et al. 2006). The Pikopiko flora supports the
hypothesis of subtropical to tropical conditions being reached
at high latitudes by the Late Eocene; however, as New Zealand
cooled, tropical species either dispersed to warmer biomes or
disappeared (Lee et al. 2001), and Calamus-like palms became
extinct there.
Acknowledgements
The Australian Tropical Herbarium (QRS), Atherton, Queensland provided
material of extant Australian Calamus species, and Adelaide Microscopy
provided valuable assistance with epidermal SEM. The Departments of
Geology and Botany, University of Otago, Dunedin and the School of
Earth and Environmental Sciences, The University of Adelaide, are
thanked for resources to undertake this research. Funds for this study were
provided by the Division of Sciences, University of Otago. John Dowe (James
S. J. Hartwich et al.
Cook University) and Bob Hill (University of Adelaide) are thanked for
comments on the manuscript.
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