Arecaceae: Calamoideae - CSIRO Publishing

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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

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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

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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

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


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)

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(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).

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(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.


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).


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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

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(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).


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


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(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).

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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

Cook University) and Bob Hill (University of Adelaide) are thanked for comments on the manuscript.

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Manuscript received 3 June 2009, accepted 21 January 2010

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