The record of Araucariaceae macrofossils in New ...

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The record of Araucariaceae macrofossils in New Zealand 5

MIKE POLE POLE, M., December, 2008. The record of Araucariaceae macrofossils in New Zealand. Alcheringa 32, 409–430. ISSN 0311-5518.

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The Araucariaceae have a long record in New Zealand, extending back to the Jurassic at least, and Araucaria extends back to at least the Late Cretaceous. This paper reviews the macrofossil record of the family and presents new information based largely on the leaf cuticle record. Agathis, which is the only genus of the family currently growing in New Zealand, has no record before the Cenozoic. All specimens previously identified from pre-Cenozoic strata clearly belong to other taxa or do not show characteristic features of the genus. Araucariaceae macrofossils are virtually ubiquitous in the Cretaceous assemblages of New Zealand but are conspicuous by their absence or rarity in Palaeocene deposits. Their demise may be an expression of events at the Cretaceous–Palaeogene boundary.

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Mike Pole [[email protected]], Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Rd, Toowong QLD 4066, Australia. Received 14.1.2008; revised 20.3.2008.

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Key words: New Zealand, Araucariaceae, Agathis, Araucaria, Araucarioides, Wollemia, Cretaceous, Palaeocene, Miocene, Quaternary, cuticle.

THE ARAUCARIACEAE has a long and widespread history (Stockey 1982, 1994, Veblen et al. 1995). From a virtually global distribution in the Mesozoic, its range is now fragmented to Malesia, Australia, New Zealand, New Caledonia and two areas in South America (southern Brazil-Paraguay and the Argentinian–Chilean Andes). The fossil record of the family has more recently been reviewed by Kershaw & Wagstaff (2001). These authors approached the subject from an ecological point of view. However, while this paper focused on the late Cenozoic of Australia, it did not refer to conifers documented from the mid-Cretaceous Eromanga Basin, eastern Australia (Pole 2000), or the Cretaceous of New Zealand (Bose 1975, Pole 1995, Pole & Douglas 1999). Pole (2000) documented the fossils in 194 fossiliferous mid-Cretaceous samples from seven boreholes in central ISSN 0311-5518 (print)/ISSN 1752-0754 (online) Ó 2008 Association of Australasian Palaeontologists DOI: 10.1080/03115510802417935

Queensland, the largest such study in Australia. Araucariaceae macrofossils were present in at least 25 of these samples and were referable to at least five taxa. Australia has very little exposure of Upper Cretaceous continental strata, hence the macrofossil record is very poorly known. New Zealand began to raft away from Australia about 82 Ma and was not far distant during the Late Cretaceous (Laird 1994, Laird & Bradshaw 2004). The vegetation seems to have been so similar that Mildenhall (1980, p. 203) remarked that for biostratigraphic correlation ‘the same palynological zones and datum planes can be used’. Thus, the New Zealand Late Cretaceous record probably provides a reasonable indication of what was also happening in eastern Australia at the time, and significantly Araucariaceae macrofossils are known from every studied assemblage of this age in New Zealand (see details below). These omissions presumably led to the comment by Kershaw and Wagstaff (2001, p. 404) that ‘By contrast there are few araucarian macrofossils

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recorded from the Middle to Late Cretaceous . . .’. They went on to speculate (p. 409) that the Araucariaceae ‘declined in abundance and range from the Early Cretaceous’ and that this may have been linked to the ‘Middle Cretaceous peak in temperatures and humidity’. Kershaw & Wagstaff (2001, p. 402) also wrote that ‘Only in Australia is there a substantial record of both macrofossils and pollen’. The record elsewhere is not as insubstantial as they may have thought. This paper aims to critically review the published record of Araucariaceae macrofossils in New Zealand, and document several new records.

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Fossil-bearing sediment was broken down in warm water and hydrogen peroxide, sieved, and the cuticle often cleared with further soaking in warm, 40% hydrogen peroxide. In many cases, staining was not necessary. Fossil cuticle was compared against a large reference collection of cuticle from extant conifers. Cuticle preparation of extant leaf material involved soaking in warm 10% aqueous chromium trioxide until clear (usually 2–3 days), then washing, and staining with saffranin. Cuticle fragments were then mounted on microscope slides in thymol glycerine jelly for transmitted light microscopy (TLM) or on aluminium stubs and coated with platinum for scanning electron microscopy (SEM).

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Extant cuticle samples prefixed with ‘AQ’ come from the Queensland Herbarium, and those with ‘NSW’ are from the National Herbarium of New South Wales. Sample numbers prefixed with ‘S-’ are electron microscope stubs, and those with ‘SL’ are fossil hand specimens or slide mounts. ‘OPH’ prefixes extant herbarium material collected by the author. Unless otherwise stated, all material is housed in the Queensland Herbarium.

Extant Araucariaceae

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Three extant genera of Araucariaceae are known: Agathis, Araucaria, and Wollemia. The epidermal morphology of extant Agathis leaves has been documented by several authors. Kausik (1976) described the stomates of Agathis dammara (Lamb.) Richards & Richards, 1826, pointing out that ‘plate-like’ extensions of the subsidiary cells form a deep, epistomatal chamber. If it were not for these, the guard cells would be close to the leaf surface. Page (1980) reviewed the external morphology of Agathis epidermis and noted that the stomates all showed (p. 72) ‘crater-rim like structures around the openings of pores leading to sunken stomata with the rims themselves set in shallow surface depressions’ and ‘varying degrees of surface undulations between the rims’. The surface of each epidermal cell is domed, although this is subdued in some species. Stockey & Atkinson (1993) provided an excellent

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Fig. 1. Extant Araucariaceae and fossil Araucarioides. A, Agathis labillardierei Warburg (1900), TLM view. The outer margin of the outer ring of subsidiary cells is indicated by arrows (AQ423023, scale bar ¼ 50 mm); B, Agathis borneensis Warburg (1900), TLM view. The outer margin of the outer ring of subsidiary cells is indicated by arrows (OPH2113, cultivated specimen, Forestry Research Institute of Malaysia, Kuala Lumpur, scale bar ¼ 50 mm); C, Araucaria araucana (Molina) Koch (1873), section Araucaria, TLM view (OPH5555, Christchurch Botanical Gardens cultivated specimen, scale bar ¼ 50 mm); D, Araucaria hunsteinii Schumann (1889), section Intermedia, TLM view (AQ119970, scale bar ¼ 50 mm); E, Araucaria cunninghamii Aiton ex D. Don (1837), section Eutacta, TLM view (OPH7214, specimen in Mt Mee State Forest, scale bar ¼ 50 mm); F, Araucaria bidwillii Hooker (1843), section Bunya, TLM view (OPH7213, Brisbane Botanical Gardens, cultivated specimen, scale bar ¼ 50 mm); G, Wollemia nobilis Jones et al. (1995), TLM view (NSW329981, scale bar ¼ 50 mm); H, Araucarioides sp., TLM view. Note massive overall thickness of cuticle. Regatta Point, Tasmania, early Eocene (SL4909, scale bar ¼ 50 mm).

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description of the cuticle of extant Agathis. The key characters, which they noted may identify fossil Agathis, are the bilobed polar

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extensions on the guard cells, evident on the internal cuticle surface. Importantly, what appears to be unique to Agathis, although 240

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not ubiquitous within the genus, is that these are genuine lobes, that is, they are somewhat divergent. It is also apparent that when viewed with TLM, Agathis stomatal complexes typically have a very distinctive structure (Fig. 1A, B). Partly this is because massively thickened cuticle of the stomatal rim is contrasted against relatively thin cuticle surrounding it and results in the impression that the rim is ‘suspended’ by the ‘spokes’ of the subsidiary cell walls. Despite the relatively thin cuticle, these ‘spokes’ effectively obscure any clear Florin Ring. The stomatal complexes also have a structure whereby there are commonly two clear rings of subsidiary cells. These are flanked by a further row of cells with generally well-defined straight outer margins. These are the edges of the three rows of cells that gave rise to the stomatal complexes, common throughout the gymnosperms (see numerous drawings in Florin 1931). The outer edge of the inner ring in Agathis is generally very smoothly sub-circular. The edge of the outer ring can be reasonably smooth and clearly defined, or can be much less coherent. The most atypical form is probably the extant New Zealand Agathis australis (D. Don) Lindley, 1829 (see discussion below under ‘Quaternary Araucariaceae’), which lends support to the conclusion of Stockler et al. (2002; see below) that this species is the most basal of the extant Agathis species. Araucaria is grouped into four sections, Bunya, Columbea, Intemedia and Eutacta (Wilde & Eames 1952). The first three of these sections are reasonably similar in that they have stomatal complexes that are predominantly orientated parallel to the long axis of the leaf (Fig. 1C–E) but section Eutacta has stomatal complexes that are randomly orientated, or at least oblique or at right angles (Fig. 1E; and see Bigwood & Hill 1985). Under TLM, Araucaria stomatal complexes typically have a more polygonal or elliptical outline than Agathis (Fig. 1C–F).

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They do not have the clearly visible double ring of subsidiary cells, partly because of crowding and overlapping by cell walls. The structure of Wollemia cuticle has been documented by Chambers et al. (1998). While much more similar to typical Araucaria than Agathis, under TLM the stomatal complexes tend to have an isodiametric and polygonal outline (Fig. 1G). The cuticle is clearly distinct from Araucaria based on the very isodiametric shape of the epidermal cells, whereas elongate epidermal cells are common throughout Araucaria species (Pole 2000). In these features, Wollemia and Agathis australis show some similarities. Wollemia has been compared with the extinct Araucarioides (Chambers et al. 1998; and see below). The stomatal structure is certainly similar (Fig. 1H), but the shoots of Araucarioides are either unknown, or in potential Araucarioides (but without cuticle; McLoughlin et al. 1995) in opposite pairs, not opposite decussate as in adult Wollemia (Jones et al. 1995, Chambers et al. 1998).

Mesozoic Araucariaceae The earliest record of Araucariaceae in New Zealand probably includes the foliage morphotaxon Podozamites and the ovulate conescale Araucarites cutchensis Feistmantel (1876) that have been reported from several Jurassic localities (Arber 1917, Bartrum 1921, Edwards 1934). Podozamites is not comparable with precision to other genera, as the name purely describes gross morphology. When Harris (1969) attempted to make sense of the system of conifer morphotaxa, his concept of Podozamites was that it includes leaves of extant Agathis, and it would also encompass Wollemia and the extinct Araucarioides. However, in the absence of any epidermal information, little is known about the precise affinities of these potential, Jurassic araucarians. The fossil wood genera Araucarioxylon Kraus 1870 (in Schimper), Agathoxylon Hartig, 1848 and

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Dadoxylon Endlicher, 1847 which cover the morphologies produced by the Araucariaceae (but perhaps also other families), have also been described from the New Zealand Jurassic (Crie´ 1889, Edwards 1934, Thorn 2001, 2005). Macrofossils indicate that by the Cretaceous, Araucariaceae were common components of the New Zealand vegetation. Ettingshausen (1887, 1891) described araucarian leaves and wood from Shag Point, and from Pakawau, northwest South Island

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(Fig. 2). Stopes (1914) described Araucarioxylon from Amuri Bluff, northeast South Island. Edwards (1926) described leaves as Araucarites marshalli from the Late Cretaceous of Bulls Point, Kaipara, which he argued were affiliated to Araucaria, together with a fragment of another araucarian leaf, and wood, Dadoxylon kaiparense. Mildenhall & Johnston (1971) described an araucarian ovuliferous cone (Araucarites) from the upper Albian of Mangapurupuru Stream, and reviewed other fossil araucarians

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Fig. 2. Map of New Zealand localities mentioned in the text.

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known in New Zealand at the time (see also Mildenhall’s 1970 checklist of plant macrofossils from New Zealand). This cone has since been re-examined in detail by Cantrill & Raine (2006) as the basis of a new araucarian genus, Wairarapaia. Bose (1975) described Araucaria leaves with cuticle from the Late Cretaceous of Shag Point, and Raine (1990) and Wiffen (1996) have noted araucarian remains preserved with Late Cretaceous dinosaur bones from Mangahouanga Stream. Pole (1995) described araucarian remains from the Cenomanian of Horse Range and Pukeiwitahi, the Campanian of Clutha Mouth and Shag Point, and the Maastrichtian of Fairfield Sand Pit and Kaitangata of the South Island (Fig. 3A–F). Specimens from Shag Point were identified as Araucarioides, a genus first described by Bigwood & Hill (1985) for use as a separate genus of Araucariaceae or to be used when preservation was too poor to allow assignment to an extant genus. Pole (1995) emended the description of Araucarioides to clarify its status as a separate genus, having a distinct flattened, falcate, and strap-like multiveined leaf. Finally, Parrish et al. (1998) listed Agathis (see below) and three species of Araucaria from the Cenomanian of the Clarence River Valley. It is clear that Araucariaceae macrofossils are consistently present in all New Zealand Cretaceous assemblages studied so far. They were an important component of the vegetation, and there were clearly several species, and at least two genera: Araucaria and Araucarioides.

Agathis in the Cretaceous? The most significant mid-Cretaceous palaeobotanical research in New Zealand is the PhD thesis of I.L. Daniel (1989) on the Clarence River Valley. In this unpublished work, two species of Agathis, ‘A. seymouricum’ and ‘A. clarencianum’, were described.

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This identification of Agathis later featured prominently in a reconstruction of the climate of the locality (Parrish et al. 1998), and more recently, Stockler et al. (2002) published a paper entitled ‘New Zealand Kauri (Agathis australis) . . . survives Oligocene drowning’. Their paper documents an investigation into the genetics of Agathis, which concluded that the extant New Zealand species Agathis australis is the most phylogenetically basal of the extant species of Agathis. This led to the claim that since Agathis was present in New Zealand in the mid-Cretaceous (based on Daniel’s work), the genus has an uninterrupted history in the country since then. The claim of Stockler et al. (2002) that there was a ‘drowning’ to survive is based on their misreading of Pole (1994a) where they imagined I had written that New Zealand’s flora has dispersed there since the country was inundated by the sea in the Oligocene. I had not, but I did conclude that it was likely that there had been a complete turnover of vegetation in New Zealand since it was physically isolated by rifting in the Late Cretaceous, with new lineages arriving by long distance dispersal. As this unpublished claim of ancient Agathis has now achieved some prominence, it needs to be reviewed. In fact, Knapp et al. (2007) using DNA evidence suggested the New Zealand Agathis radiated from closest living relatives in the Eocene. In the context of the present paper, it is important to note that Daniel (1989) compared his fossils neither with extant Wollemia (which had not been described at that time; Jones et al. 1995) nor with the earlier described extinct genus Araucarioides (Bigwood & Hill 1985, Hill & Bigwood 1987), both in Australia. Daniel’s ‘Agathis seymouricum’ was described as a flattened, multiveined species with a leaf 20–57 mm long and 5–15 mm wide, with a length:width ratio of 5.5 or less, and unequally amphistomatic. The stomatal

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Fig. 3. Mid-Late Cretaceous Araucariaceae. A, Araucaria taeriensis Pole 1995, TLM view, note distinctly elliptical outline to complexes, Kaitangata (SB0834, scale bar ¼ 100 mm); B, Araucarioides falcata Pole 1995, TLM view, note polygonal outline to complexes, Shag Point (SB0810, scale bar ¼ 100 mm); C, Araucaria desmondii Pole 1995, TLM view, note isodiametric epidermal cells, Horse Range (SB0813, scale bar ¼ 100 mm); D, Araucariaceae, TLM view, note elongate epidermal cells, drill core d.3057, Kaitangata (SL2558, scale bar ¼ 100 mm); E, Araucariaceae, TLM view, note isodiametric epidermal cells, Cave Stream (SL4433, scale bar ¼ 100 mm); F, Araucariaceae, TLM view, note short epidermal cells, Cave Stream (SL4684, scale bar ¼ 100 mm).

complexes in his descriptions and illustrations are dramatically different from all extant Agathis. The cuticular flange, instead of being relatively narrow, is a massively

extended disk that encompasses virtually all of the subsidiary cells. Daniel (1989 p. 82) describes this as: ‘The flanges between the guard cells and subsidiary cells are large,

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circular, coming to a point towards the poles, and nearly covering the subsidiary cells, except at the poles’. Deep cuticular pits are present at the polar ends of the stomata, reflecting guard cells that are reflexed sharply upwards (towards the leaf surface), rather than extending outwards. This gives the stomatal ledge a distinctive appearance. I know of no extant conifers having this form of ledge, although I have described one mid-Cretaceous Australian specimen as Podocarpaceae (Pole 2000, fig. 13), but Daniel’s specimen certainly does not belong in that family. Although no taxonomic comparison is intended, this construction is more typical of cycads (see Hill 1980, fig. 20) and I have described it from Palaeocene leaves that may belong in the Taxodiaceae or Taxaceae (Pole 1998a, fig. 13F, G). As obtaining Daniel’s thesis via the Interlibrary Loan system was not straightforward, for the reader’s convenience I include here two scanning electron microscope (SEM) illustrations of the stomata of another Clarence River taxon (Fig. 4A, B), probably one of Daniel’s (1989) ‘Ginkgo cuneformis’, which shows an identical stomatal ledge form to Daniel’s figures of A. seymouricum (although the subsidiary cell arrangement is quite distinct, and the two are clearly different taxa). This epidermal character makes ‘A. seymouricum’ immediately distinct from extant Agathis species, which have narrow stomatal ledges that diverge near the poles and ‘lie’ in shallow cups at about the same level as the ledge. There is also no indication of the distinctive surface morphology of extant Agathis where individual epidermal cells are domed, and the stomatal pore is surrounded by a rim. No transmitted light microscope photographs were given, which would illustrate the stomatal construction more clearly, and would indicate whether they had the distinctive circle of thin cuticle found in extant species.

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The second Clarence River species, ‘Agathis clarencianum’ also has a flattened and multiveined leaf, 45–130 mm long, 4–18 mm wide, and is hypostomatic, but much more elongate than A. seymouricum, having a length:width ratio of 8.0 or more. The cuticle in Daniel’s (1989) illustrations appears to be more poorly preserved, and the details are unclear, but the stomatal complexes do not have the broad ledges as in ‘A. seymouricum’, as Daniel makes clear in his description. Permission was given to sample cuticle from Daniel’s collection by the Canterbury Museum, where it is now held, in the hope that better SEM specimens might be made, and to prepare specimens for TLM viewing. No cuticle was recovered from ‘Agathis seymouricum’ but small and poorly preserved fragments were recovered from ‘Agathis clarencianum’ leaves. The leaves are hypostomatic, with stomatal complexes and epidermal cells in well-defined rows, and they clearly belong in the Araucariaceae (Fig. 4C–H). However, under TLM, the cuticle does not show the ‘suspended’ ring structure or basic stomatal complex typical of Agathis. SEM observation shows moderately broad stomatal ledges with sharply upturned, non-divergent poles, and very subdued, barely noticeable outer surface topography. There is no indication that these leaves belong in Agathis, and in my opinion these leaves are a species of Araucarioides or Wollemia. The evidence for Agathis in the Clarence River material is thus poor. If Agathis was in New Zealand in the mid-Cretaceous, it might be expected to appear in some of the more diverse conifer-dominated Australian fossil assemblages of a similar or earlier age. Jurassic leaf impressions described as Agathis by White (1981) do not have cuticle and were not regarded as Agathis by Stockey (1982). In addition, they are associated with ovuliferous cone-scales that are quite unlike Agathis. An Agathis reported

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Fig. 4. Mid-Cretaceous Clarence River material. A, ‘Ginkgo cuneformis’ inner SEM view of stomatal complex. Note very broad scales and depressions at the polar ends that represent spaces where the guard cell poles have reflexed abruptly outwards (SL4876, S-1600, scale bar ¼ 10 mm); B, ‘Ginkgo cuneformis’ inner SEM view of stomatal complex (SL4876, S-1600, scale bar ¼ 10 mm); C, Three rock fragments with specimens of ‘Agathis clarencianum’ (scale bar ¼ 10 mm); D, ‘Agathis clarencianum’, TLM view, note lack of ‘suspended rings’ (SL4915, scale bar ¼ 100 mm); E, ‘Agathis clarencianum’, TLM view, note lack of ‘suspended rings’ (SL4915, scale bar ¼ 100 mm); F, ‘Agathis clarencianum’, outer SEM view showing two barely discernible stomatal pores (S-1606, scale bar ¼ 50 mm); G, ‘Agathis clarencianum’, inner SEM view of stomatal complex, showing polar cups that project outwards (with respect to the leaf) and are not divergent. (S-1606, scale bar ¼ 10 mm); H, ‘Agathis clarencianum’, inner SEM view of stomatal complex, showing polar cups that project outwards (with respect to the leaf) and are not divergent (S-1606, scale bar ¼ 10 mm).

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by Cantrill (1992) from the mid-Cretaceous of Victoria has no cuticle, and comparison with Araucarioides was not considered. Chambers et al. (1998) compared the gross morphology with Wollemia. The mid-Cretaceous assemblages of central Queensland, with 26 species of conifer, include no Agathis (Pole 2000). If Agathis were in mid-Cretaceous New Zealand, it might also be expected to be in Late Cretaceous New Zealand assemblages. A dammar (Agathis) reported by Ettingshausen (1887, 1891) from the Late Cretaceous of New Zealand is an impression only and has been shown to be a misinterpreted Araucaria-like scale leaf (the sharp apex was drawn as the false-petiole of an Agathis; Pole 1995). The Kaitangata assemblages have yielded 10 species of conifer with cuticular information (Pole & Douglas 1999), but no Agathis is known. There is no compelling evidence that Agathis existed in the Cretaceous in New Zealand. In fact, in their review of southern conifers, Hill & Brodribb (1999, p. 649) considered that Agathis ‘does not have a convincing pre-Cenozoic record’. Australian records of Agathis also need to be carefully re-examined. Agathis parwanensis (Cookson & Duigan 1951) from the early Miocene appears to be appropriately classified, whereas I consider that Agathis yallournensis Cookson & Duigan, 1951, also from the early Miocene is potentially an Araucarioides or Wollemia based on leaf shape and stomatal outline. Agathis tasmanica from the early Oligocene to early Miocene of Tasmania (Hill & Bigwood 1987) has the external morphology and basic stomatal construction of Agathis, although the bilobed polar extensions are not visible in the illustrations. Agathis sp. from the Eocene of Western Australia (Carpenter & Pole 1995) does not look like an Agathis to this author. The basic stomatal construction and external morphology looks like Agathis, but the

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well-preserved polar extensions are not divergent. It is more likely that this is an Araucarioides. More recently Carpenter et al. (2004, p. 688) published early Eocene specimens from Victoria that have the ‘the distinctive Agathis polar cuticular knobs’ suggesting that their fossils could be ‘assigned to Agathis with confidence’. Their illustration does not show bilobed (diverging) polar knobs, and there is no indication in their illustration that the stomatal complexes were arranged in rows as found in extant Agathis (the construction looks rather similar to the mid-Cretaceous Eutactoides Pole 2000, a taxon that may not be araucarian). The leaves and cuticular characters of extant species of Agathis are very similar and difficult to distinguish (Whitmore 1980). I would thus be surprised if the small differences between Agathis australis and other extant Agathis species originated in the mid-Cretaceous. Agathis, rather than dating from the Cretaceous, or earlier, and being a sure sign of ancient land connections (e.g. Michaux 2001), probably arose in the Palaeogene and crossed ocean-gaps to reach New Zealand.

Cretaceous–Palaeogene boundary decline While Araucariaceae macrofossils have been found in every published Late Cretaceous plant fossil assemblage, the situation in the Palaeogene seems remarkably different. For instance, the mid-late Palaeocene Mount Somers assemblage contains 13 species of conifers, but just one small fragment of probable araucarian cuticle was found among thousands of fragments of other (mostly Podocarpaceae) conifers (Pole 1998a). The only other published case is Lindqvist’s (1986) report of a teredo-bored araucarian log from Palaeocene shoreface sediments, close to

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the Cretaceous–Palaeogene boundary, at Wangaloa, south of Dunedin. No Araucariaceae macrofossils have been found in any sample from the late Palaeocene–early Eocene of Kakahu (Pole 1997). In a terrestrial section that actually crosses the Cretaceous–Palaeogene boundary in the Pakawau region of northwest South Island (Kennedy et al. 2002), Araucariaceae are listed from the latest Cretaceous, but not the earliest Palaeocene. In another Cretaceous– Palaeogene boundary section at Moody Creek Coal Mine, West Coast (Vajda et al. 2001), Araucariaceae fossils, including Araucarioides and associated ovulate conescales are common in the Late Cretaceous, but absent in the Palaeocene (Fig. 5A–D and Pole unpublished data). Thus, a dramatic decline in Araucariaceae around the Cretaceous–Palaeogene boundary is apparent, perhaps involving a reorganization of vegetation structure and basic composition together with extinctions. This change is difficult to interpret, as Araucariaceae pollen is still present in reasonable amounts in the Palaeocene. Vajda & Raine (2003) did note that it was less abundant above the Cretaceous– Palaeogene boundary at Waipara River than below, and Meon (1990) noted that Araucariaceae pollen disappears at the boundary in Tunisia. Based on these observations it is possible that the Cretaceous–Palaeogene boundary is represented at Cave Stream, central Canterbury. Mildenhall (1972) interpreted this section’s age as Late Cretaceous based on the palynology of matrix surrounding the syntype of Dryandra comptoniaefolia Ettingshausen, 1887. However, based on Raine’s (1984) palynological zonation, the taxa seem more indicative of the Palaeocene (see Pole 1998b for a discussion). Recent work at this site has revealed abundant Araucarioides in the lower part of the section, but it is absent in the upppermost samples. In Palaeocene strata bearing

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Fig. 5. Araucarioides and associated ovulate cone scales from the latest Cretaceous of Moody Creek Coal Mine. All scale bars ¼ 10 mm. A, Cone scale, SL4926; B, Leaf, SL4929; C, Cone scale, SL4927; D, Leaf, SL4928.

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Dryandra comptoniaefolia, the only Araucariaceae are uncommon fragments of cuticle that might represent a form of Agathis, the oldest potential Agathis I recognize in New Zealand (Fig. 6A–H).

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Eocene Agathis

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Eocene macrofossils in New Zealand have received little attention, and the publications have not described any Araucariaceae (e.g. Penseler 1930, Pole 1994b). In my

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Fig. 6. Possible Paleocene Agathis, from Cave Stream. A, TLM view of stomatal complexes, note prominent ‘suspended rims’ (SL4588, scale bar ¼ 50 mm); B, TLM view of stomatal complexes, note prominent ‘suspended rims’ (SL4588, scale bar ¼ 50 mm); C, Inner SEM view of stomatal complex, polar cups not apparent (S-1433, scale bar ¼ 10 mm); D, Inner SEM view of stomatal complex, polar cups not apparent (S-1433, scale bar ¼ 10 mm); E, Inner SEM view of stomatal complex, polar cups not apparent (S-1433, scale bar ¼ 10 mm); F, Outer SEM view of stomatal complex showing distinct rim around aperture (S-1433, scale bar ¼ 10 mm); G, TLM view of unusual specimen with distinct papillae in centre of epidermal cells (SL4825, scale bar ¼ 50 mm); H, TLM view of possible Agathis with unusual subsidiary cell morphology (SL4586, scale bar ¼ 50 mm).

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experience of these, and unpublished collections, New Zealand had an angiospermdominated flora at the time, and conifer macrofossils of any kind are not common. However, research in progress on the late

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Eocene of the Waikato Coal Measures of the North Island (Edbrooke et al. 1994) has found small fragments of obviously araucarian cuticle. These specimens from the Huntly Coal Pit have stomates in rows,

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Fig. 7. Possible Eocene Agathis from the late Eocene, Huntly. A, TLM view of stomatal complexes, with distinct ‘suspended rims’ but unusual subsidiary cell morphology. The outer margin of the outer ring of subsidiary cells is indicated by arrows (SL4196, scale bar ¼ 50 mm); B, Close up TLM view of stomatal complexes (SL4196 scale bar ¼ 20 mm); C, Inner SEM view of stomatal complex, note non-diverging polar cups (S-1421, scale bar ¼ 10 mm); D, Inner SEM view of stomatal complex, note non-diverging polar cups (S-1421, scale bar ¼ 10 mm); E, Inner SEM view of stomatal complex, note non-diverging polar cups (S-1421, scale bar ¼ 10 mm); F, Outer SEM view of stomatal complex showing low but distinct rim around aperture (S-1421, scale bar ¼ 10 mm).

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orientated mostly at right angles to the rows, have the typical ‘suspended ring’ appearance of Agathis, have a slight but clearly raised rim around the aperture, and

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have a distinct second (outer) ring of subsidiary cells (Fig. 7A–F). They do not have the bilobed polar knobs that characterize extant Agathis (Stockey & Atkinson

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1993), but like the Palaeocene Cave Stream material, they are consistent with what could be an early form of the genus. 1320

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Araucariaceae remains have been reported from the early Miocene of the Manuherikia Group in the form of both wood and resin by Evans (1937). The wood is informative only at the family level, no characters were provided that allow it to be diagnosed as Agathis, and, to my knowledge, none exist. Evans (1937) recorded agathic acid in resins from coal at Harliwichs Pit and, on this basis, concluded it was produced by Agathis. More recently, such resins around the world have been the focus of research attempting to fingerprint and identify them using nuclear magnetic resonance spectroscopy and cross-polarization and magnetic angle spinning. Lambert et al. (1993, 1999) concluded that resins of Araucaria and Agathis can be distinguished from each other, but that of Agathis and Wollemia (and presumably extinct genera such as Araucarioides) cannot. Lambert & Poinar (2002) found that Agathis-like resins formed a group and had a worldwide distribution. Furthermore, they found that Baltic Amber was distinct, but was probably related to the

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Agathis-like group by chemical maturation. These results are clear evidence that this group of resins is not only formed by Agathis, which has no other evidence of a worldwide distribution. In fact, in the Baltic mid-Cenozoic, where there has been no morphological evidence of Araucariaceae, let alone Agathis, the resin is generally thought to have been produced by the Pinaceae, or a close relative. It has been suggested that the resin may have been produced by an extinct gymnosperm group combining the features of both Araucariaceae and Pinaceae (Larson 1978, Selden & Nudds 2005). Interestingly, agathic acids have also been recorded from Copaifera, a leguminous angiosperm (Lima et al. 2003). Thus, while the resins in the Manuherikia Group and in other localities throughout New Zealand may well have been produced by Araucariaceae, the data do not allow discrimination between Agathis and other genera, which certainly existed. Given these issues, added to the presence of material in the Miocene that is clearly related to Agathis, but still distinct (Pole 2007a), using the resins as a ‘proxy’ for Agathis, as proposed by Lee et al. (2007), is not appropriate. Further south in the South Island, Lindqvist (1990) illustrated araucarian (almost certainly Araucaria) from silicified fluvial sediments of the Gore Lignite

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Fig. 8. Miocene to Quaternary Araucariaceae. A, Araucaria sp., TLM view of stomatal complexes, note short epidermal cells, early Miocene, Bannockburn (SL3270, scale bar ¼ 100 mm); B, Araucaria sp., TLM view of stomatal complexes, note short epidermal cells, early Miocene, Bannockburn (SL3270, scale bar ¼ 100 mm); C, Araucaria nemorosa Laubenfels (1969), TLM view showing typical elongate epidermal cells (OPH4176, scale bar ¼ 100 mm); D, Araucaria schmidii Laubenfels (1969), TLM view showing typical elongate epidermal cells (OPH4179, scale bar ¼ 100 mm); E, Agathis sp., leaf impression, note distinct false petiole, Oruawharo (Medlands Stream), (Auckland University, Geology Department specimen P218, scale bar ¼ 10 mm); F, Agathis sp., ovulate cone scale, Oruawharo (Medlands Stream), (Auckland University, Geology Department specimen P219, scale bar ¼ 10 mm); G, Agathis australis, TLM view of stomatal complexes, Quaternary, Baylys Beach (SL4401, scale bar ¼ 100 mm); H, Agathis australis, close up TLM view of stomatal complex, Quaternary, Baylys Beach. The inner two arrows mark the outer edge of the inner and usually only ring of subsidiary cells. The outer two arrows indicate the outer edge of the original three rows of cells that gave rise to the stomates, i.e. there is only a single ring of subsidiary cells (SL4401, scale bar ¼ 50 mm).

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Measures at Landslip Hill. They appear to be reasonably common at this locality. Pole (1992) recorded relatively common Araucaria shoots, ovuliferous cone scales, and also a microsporangiate cone from a single horizon in the early Miocene Manuherikia Group, at Bannockburn. The Araucaria remains are restricted to within an approximately 1 m thick stratigraphic interval exposed over a 100 m lateral extent. They have not been recorded from any other impression assemblages from the Bannockburn area, assemblages which are, in most cases, overwhelmingly angiosperm-dominated. The fossils were placed in Araucaria section Eutacta based on the morphology of the shoots and the ovuliferous scales with thin, papery wings. Continued work at Bannockburn has revealed a specimen with some attached cuticle (this is essentially an impression-assemblage, but this toughleaved conifer specimen may be unique). The cuticle preservation is less than ideal but is consistent with Araucaria section Eutacta in all but one aspect; the specimen does not show the long, narrow epidermal cells characteristic of extant members of the section (Fig. 8A–D). One possibility is that the fossil may be basal in the Eutacta clade, and that this morphology has become extinct, leaving the more derived taxa with elongate epidermal cells. Since my work on the largely impression-only assemblages of the Manuherkia Group, I have focussed on older strata of the Group, the cuticle-bearing St Bathans Member, and also the Gore Lignite Measures, of the East Southland Group. Araucariaceae cuticle is present, but very uncommon, and most of those specimens that have been located are very fragmentary and somewhat weathered. Presumably, they come from vegetation growing at some distance from the immediate site of burial. They appear to include the same Araucaria taxon as above, together with possible Agathis or Agathis-like species (Pole

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2007a). A recent exception is the Agathis described from the East Southland Group by Lee et al. (2007). These authors illustrate what is clearly an Agathis, although they use the presence of what they called Florin Rings and bilobed polar extensions as important diagnostic characters for the genus. Their use of Florin Rings clearly applies to the markedly raised subsidiary cells, rather than to ‘a special furrow in the cuticular thickening forming a translucent ring [under TLM view]’ (Buchholz & Gray 1948, p. 52). Unfortunately, Lee et al. (2007) did not provide TLM illustrations that would indicate whether Florin Rings or the typical Agathis suspended rim structure is present. Furthermore, there is little about the polar extensions of their material that looks conclusively Agathis (i.e. clearly lobed or divergent) and any different from, for example, those in Wollemia (Chambers et al. 1998). A previously unpublished record is that of a broad ovuliferous cone-scale and the base of what appears to be a conifer leaf narrowing to a false petiole (with no cuticle) in an Auckland University Geology Department collection from the late Miocene locality of Oruawharo (Medlands Stream) on Great Barrier Island (Fig. 8E, F). No cuticle is present, but on simple morphology and their young age, these are likely to be Agathis. A small imbricate conifer shoot from the late Pliocene, or possibly earliest Pleistocene, of the Grey River has been identified as an Araucaria (Pole 2007b). I am not aware of Araucariaceae remains in any other Miocene or Pliocene localities in New Zealand.

Quaternary Araucariaceae Pocknall (1981 p. 272) described Agathis australis pollen, but remarked that ‘specific identification of A. australis pollen in

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Tertiary sediments is made difficult by the presence of morphologically similar pollen produced by other members of the Araucariaceae’. In Australia, a distinction is sometimes made between Araucaria and Agathis pollen; however, Araucariaceae pollen in New Zealand’s Quaternary is generally accepted as being Agathis. There is no positive evidence of Araucaria surviving into the New Zealand mid-Pleistocene, but Agathis wood, leaves, and cone material is common in some North Island peats. For instance, Mildenhall (1985) and Mildenhall et al. (1992) described the palynology of the Baylys Beach lignite, near Dargaville, which is dominated by the compressed remains of Agathis australis foliage and cone scales (pers. obs.). Although its age is not clear, it contains extinct pollen and fruit types, and probably dates from one of the interglacial periods. Agathis australis has the ‘suspended rim’ appearance of typical Agathis, but the epidermal cells are mostly isodiametric and it does not have the clearly visible double ring of subsidiary cells characteristic of other extant Agathis (Fig. 8G, H). I have surveyed several similar midQuaternary peat/lignite deposits in the North Island and find it remarkable that so far there is a general absence of Agathis from other Pleistocene peats in New Zealand and no evidence for Agathis extending beyond its present range in any interglacial period. Ogden et al. (1992) summarized the late Quaternary history of Agathis in New Zealand concluding that prior to about 25 000 years ago, and at least as far back as 40 000 years ago, Agathis was essentially restricted to an area north of about 358S, i.e. the northernmost tip of the North Island. In the mid-late Holocene, there was a general expansion, southwards to its present limit at about 388S. They made the point that this need not have been a physical migration, but more a sequential expansion of local populations. Modelling of the ‘climatic envelope’ of extant A. australis

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(Mitchell 1991) indicates that there is little difference between its actual and predicted distribution, and that climate controls its southern boundary (see also Ecroyd 1982). More recent ‘whole vegetation’ modelling (Leathwick 2001) has reached the same conclusions.

Conclusions Araucariaceae was a common component of the New Zealand vegetation by at least the Cretaceous, although reports of Agathis from that time are not substantiated. The family underwent some sort of decline close to, if not coinciding with, the Cretaceous– Palaeogene boundary; until that time, it seems to have remained generally abundant. Although its macrofossil record seems to disappear around this time, the family clearly remained in New Zealand based on palynological evidence. The decline may have involved selective extinction of those species that tended to inhabit sedimentary basins or the replacement of wetland communities by some other vegetation. Agathis, or perhaps some ‘ancestral’ form, may have arrived in New Zealand soon after the Cretaceous–Palaeogene event. By the midCenozoic, conifers in general had declined, but Araucariaceae, perhaps Agathis, was present in New Zealand. In the early Miocene Araucaria section, Eutacta and Agathis were present, but it is not until the Quaternary that Agathis australis is clearly present. An annotated summary of New Zealand fossils attributed to the Araucariaceae is presented in Appendix 1.

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Acknowledgements My thanks to Dr Norton Hiller of the Canterbury Museum, for permission to sample the Daniel collection, to Dr Neville Hudson, Geology Department, University of Auckland, for permission to photograph the Great Barrier Island material, to

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Dr William Loh, for slide scanning, and to the Christchurch Botanical Gardens, the Brisbane Botanical Gardens, the Forestry Research Institute of Malaysia, Kuala Lumpur, and Queensland Herbarium, for access to their collections. The comments of three anonymous reviewers and editorial assistance were much appreciated.

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POLE, M.S., 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13, 153-197. POLE, M.S., 2007a. Conifer and cycad distribution in the Miocene of southern New Zealand. Australian Journal of Botany 55, 143-164. POLE, M.S., 2007a. Plant macrofossil assemblages during Pliocene uplift, South Island, New Zealand. Australian Journal of Botany 55, 118-142. POLE, M.S. & DOUGLAS, B.J., 1999. Plant macrofossils of the Upper Cretaceous Kaitangata Coalfield, New Zealand. Australian Systematic Botany 12, 331-364. RAINE, J.I., 1984. Outline of a palynological zonation of Cretaceous to Paleogene terrestrial sediments in west coast region, South Island, New Zealand. New Zealand Geological Survey Report 109, 1-82. RAINE, J.I., 1990. Late Cretaceous plant macrofossils from Mangahouanga Stream. Geological Society of New Zealand Annual Conference, Napier 1990. Programme and Abstracts, Geological Society of New Zealand, Lower Hutt, 111. RICHARD, L.C. & RICHARD, A., 1826. Commentatio botanica de Conifereis et. Cycadeis, Cottae, Stuttgart, 4v, xv þ 212 pp., 30 pl. SCHUMANN, K., 1889. Die Flora von Kaiser Wilhelmsland, Asher, Berlin, v þ 137 pp. SELDEN, P.A. & NUDDS, J., 2005. Evolution of Fossil Ecosystems, Manson Publishing and The University of Chicago Press, Chicago, 160 pp. STOCKEY, R.A., 1982. The Araucariaceae: an evolutionary perspective. Review of Palaeobotany and Palynology 37, 133-154. STOCKEY, R.A., 1994. Mesozoic Araucariaceae: morphology and systematic relationships. Journal of Plant Research 107, 493-502. STOCKEY, R.A. & ATKINSON, I.J., 1993. Cuticle micromorphology of Agathis Salisbury. International Journal of Plant Sciences 154, 187-225. STOCKLER, K., DANIEL, I.L. & LOCKHART, P.J., 2002. New Zealand Kauri (Agathis australis (D.Don) Lindl., Araucariaceae) survives Oligocene drowning. Systematic Biology 51, 827-832. STOPES, M.C., 1914. A new Araucarioxylon from New Zealand. Annals of Botany 28, 341-350. THORN, V., 2001. Vegetation communities of a high paleolatitude Middle Jurassic forest in New Zealand. Palaeogeography, Palaeoclimatology, Palaeoecology 168, 273-289. THORN, V., 2005. A Middle Jurassic fossil forest from New Zealand. Palaeontology 48, 1021-1039. UNGER, F., 1864. Fossil Planzenreste aus Neu-seeland. Novara Expedition. Geologisches Teil. 1, 1-13. VAJDA, V. & RAINE, J.I., 2003. Pollen and spores in marine Cretaceous/Tertiary boundary sediments at mid-Waipara River, North Canterbury, New Zealand. New Zealand Journal of Geology and Geophysics 46, 255-273.

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VAJDA, V., RAINE, J.I. & HOLLIS, C.J., 2001. Indication of global deforestation at the Cretaceous–Tertiary boundary by New Zealand fern spike. Science 294, 1700-1702. VEBLEN, T.T., BURNS, B.R., KITZBERGER, T., LARA, A. & VILLALBA, R., 1995. The ecology of the conifers of southern South America. In Ecology of the southern conifers, N.J. ENRIGHT & R.S. HILL, eds, Melbourne University Press, Melbourne, 120-155. WALKOM, A.B., 1921. Mesozoic floras of New South Wales. Part 1- Fossil plants from Cockabutta Mountain and Talbragar. Geological Survey of New South Wales, Palaeontology, Memoir 12, 1-21. WARBURG, O., 1900. Beitra¨ge zur Kenntniss der Vegetation des sud- und ostasiatichen Monsungebietes. Monsunia 1, 42-53.

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WHITE, M.E., 1981. Revision of the Talbragar Fish Bed Flora (Jurassic) of New South Wales. Records of the Australian Museum 33, 695-721. WHITMORE, T.C., 1980. A monograph of Agathis. Plant Systematics and Evolution 135, 41-69. WIFFEN, J., 1996. Dinosaurian palaeobiology: a New Zealand perspective. Memoirs of the Queensland Museum 39, 725-731. WILDE, M.H. & EAMES, A.J., 1952. The ovule and seed of Araucaria bidwillii with a discussion of the taxonomy of the genus. 2. Taxonomy. Annals of Botany n.s. 16, 27-47.

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2015

2010 Jurassic, Curio Bay Late Albian–Cenomanian, Coverham

Appendix 1. New Zealand Araucariaceae macrofossil species.

Podozamites gracilis Arber (1917) Wairarapaia mildenhallii Cantrill & Raine (2006)

Jurassic, Waikawa Age and locality uncertain Late Cretaceous, Kaipara Late Cretaceous, Kaipara Late Cretaceous, Pakawau Age uncertain, Richmond Late Cretaceous, Shag Point Late Cretaceous, Shag Point Late Cretaceous, Shag Point

Jurassic, Mokoia

Late Cretaceous, Shag Point

Late Cretaceous, Shag Point

Late Cretaceous, Malvern Hills Cretaceous, Amuri Bluff

Late Cretaceous, Kaitangata Late Cretaceous, Shag Point Jurassic, Toi toi-Mataura

Cretaceous, Shag Point

Early Miocene, Roxburgh Late Cretaceous, Malvern Hills Late Cretaceous, Kaitangata Late Cretaceous, Shag Point and Malvern Hills

MIKE POLE

Araucarites grandis Walkom, 1921 Araucarites kakanuia Hector (1880) Araucarites marshalli Edwards (1926) Dadoxylon kaiparensis Edwards (1926) Dammara mantelli Ettingshausen (1887) Dammara fossilis Unger (1864) Dammara uninervis Ettingshausen (1887) Dammarites carinata Hector (1880) Dammarites lanceolata Hector (1880)

Araucarites cutchensis Feistmantel (1876)

Araucarites carinaria Hector (1880)

Araucarioxylon ettingshauseni Stopes (1914) Araucarioxylon novae-zeelandiae Stopes (1914) Araucarioxylon zeelandicum Crie´ (1889) Araucarites buchanani Hector (1880)

Araucaria taeriensis (Pole 1995) Araucarioides falcata (Pole 1995) Araucarioxylon australe Crie´ (1889)

Araucaria oweni (Ett.) Pole (1995)

Agathoxylon australe Evans (1937) Araucaria danai Ettingshausen (1887) Araucaria desmondii Pole (1995) Araucaria haastii Ettingshausen (1887)

2000

Early Miocene, Roxburgh Cenomanian, Clarence River

1990

Described in an as-yet unpublished PhD thesis; in my opinion it is not Agathis but more likely a species of Araucarioides or Wollemia Regarded as invalid by Mildenhall (1970) Described in an as-yet unpublished PhD thesis Does not appear to be araucarian and is possibly not a conifer, Regarded as invalid by Mildenhall (1970) Araucarian status not clear; needs further study Based on leaf cuticle The foliage was confirmed as Araucaria based on cuticle by Bose (1975); Ettingshausen (1887), in what would be regarded as unacceptable practise today, also placed fossil wood into the same species; Stopes (1914) transferred the wood to Araucarioxylon ettingshauseni, but was doubtful whether it was araucarian Ettingshausen (1887) described this species as Dammara oweni, apparently interpreting the sharp leaf apex as a false petiole Based on leaf cuticle Based on leaf cuticle Probably araucarian wood, but see Philippe (1993) and Bamford & Philippe (2001) for a discussion on the use of the genus As above and see comments on Araucaria haastii As above Regarded as invalid by Mildenhall (1970) Regarded as valid by Oliver (1950) but listed and illustrated as Araucaria bucanani; regarded as invalid by Mildenhall (1970) Regarded as valid by Oliver (1950) but listed and illustrated as Araucaria carinaria; regarded as invalid by Mildenhall (1970) Specimens placed into this widespread species by Arber (1917) are possibly araucarian ovulate scales, but require detailed study Edwards (1934) placed material into this species, but no illustrations were provided Regarded as invalid by Mildenhall (1970) Probably Araucaria but cuticle unknown Probably araucarian wood Needs further research to determine if this is an Agathis or Araucaria Regarded as invalid by Mildenhall (1970) Probably an Araucaria illustrated upside down Regarded as invalid by Mildenhall (1970) Regarded as valid by Oliver (1950) but listed and illustrated as Agathis lanceolata; regarded as invalid by Mildenhall (1970) Possibly araucarian foliage but cuticle is needed to be certain A well-described ovulate cone scale previously described under Araucarites by Mildenhall & Johnston (1971)

Comments

1985

Cenomanian, Clarence River

1980

Agathis praeaustralis Evans (1937) Agathis seymouricum Daniel (1989)

2005 Age and locality

1975

Agathis clarencianum Daniel (1989)

1995

Taxon

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