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Late Cretaceous (Cenomanian–Turonian) macroflora from the ChathamIslands, New Zealand: Bryophytes, lycophytes and pteridophytes

Chris Mays, Anne-Marie P. Tosolini, David J. Cantrill, Jeffrey D. Stil-well

PII: S1342-937X(14)00108-7DOI: doi: 10.1016/j.gr.2014.03.017Reference: GR 1250

To appear in: Gondwana Research

Received date: 30 October 2013Revised date: 18 February 2014Accepted date: 14 March 2014

Please cite this article as: Mays, Chris, Tosolini, Anne-Marie P., Cantrill, David J., Stil-well, Jeffrey D., Late Cretaceous (Cenomanian–Turonian) macroflora from the ChathamIslands, New Zealand: Bryophytes, lycophytes and pteridophytes, Gondwana Research(2014), doi: 10.1016/j.gr.2014.03.017

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Late Cretaceous (Cenomanian–Turonian) macroflora from the Chatham

Islands, New Zealand: bryophytes, lycophytes and pteridophytes

Chris Maysa,*, Anne-Marie P. Tosolini

b, David J. Cantrill

c,d and Jeffrey D. Stilwell

a

a School of Geosciences, Monash University, Victoria, 3800, Australia

b School of Earth Sciences, The University of Melbourne, Victoria, 3010, Australia

c National Herbarium of Victoria, Royal Botanic Gardens Melbourne, Private Bag 2000, South Yarra, Victoria,

3141, Australia

d School of Botany, The University of Melbourne, Victoria, 3010, Australia

* Corresponding author; School of Geosciences, Monash University, Wellington Road, Victoria, 3800, Australia;

+61 3 9905 5786; Fax +61 3 9905 4903; email: chris.mays@monash.edu

Mays, C., Tosolini, A.-M.P., Cantrill, D.J. and Stilwell, J., 2013. Late Cretaceous (Cenomanian–Turonian) macroflora from the

Chatham Islands, New Zealand: bryophytes, lycophytes and pteridophytes. Gondwana Research XX, XXX–XXX. ISSN

XXXXX-XXXXX.

Abstract. Late Cretaceous fossils from the Chatham Islands, New Zealand, represent an

important high palaeolatitude (~ 70–80°S) flora. Located between eastern and western

Gondwana, and prior to Late Cretaceous continental break-up, these plants grew during a period

of global greenhouse climates. Macrofloral remains of the Tupuangi Formation, Pitt Island,

accumulated in a deltaic floodplain setting with plant material occurring on well-developed,

hydromorphic soil horizons or entrained in sediments overlying the soils. The macroflora

includes a rich angiosperm-conifer-Ginkgo flora with subsidiary ferns, lycophytes and

bryophytes. The components of the assemblage described herein include those of probable

cryptogam affinity, and comprise one thalloid liverwort, one leafy moss (Muscites gracilis sp.

nov.), one lycopod shoot, and three taxa of pteridophytic affinity, including a fern of probable

osmundalean affinity (Cladophlebis auriculipilosus sp. nov.). The floral assemblage shares only

one element with coeval localities across the Southern Hemisphere (Sphenopteris sp. cf. S.

warragulensis), and is characterised by relatively low fern diversity. The unique assemblage is

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attributed, at least in part, to ecological bias associated with local deltaic depositional settings.

However, the relatively low osmundalean fern component follows global floristic trends for the

mid-Cretaceous, which saw a decline in diversity and abundance of ferns during the early phases

of angiosperm diversification and rise to ecological dominance.

Keywords: Cretaceous, fern, bryophyte, lycopod, Chatham Islands.

1. Introduction

The early Late Cretaceous (Cenomanian–Turonian) has long been recognised as an interval of

global greenhouse conditions (Barron, 1983; Arthur et al., 1985; Miller et al., 2005). Marine

fossil evidence suggests that this warming was acutely expressed at polar latitudes, with a

temperature maximum during the Turonian (Huber et al., 1995; Huber, 1998; Bice et al., 2006).

This Cretaceous warming trend has been supported by fossil floral palaeotemperature proxies

from Antarctica (Hayes, 1999; Francis and Poole, 2002), and a Cenomanian floral assemblage

from New Zealand (mean annual temperature: ~ 10°C; Parrish et al., 1998). Atmospheric carbon

dioxide levels were higher than modern levels by a factor of between four and seven (Berner and

Kothavala, 2001; Bice et al., 2006; Wang et al., in press); it has been proposed that such high

CO2 concentrations in ancient atmospheres would lead to a state of ‘carbon fertilisation’,

enhancing polar forest productivity (e.g. Beerling and Osborne, 2002). These global conditions

coincide with major changes to terrestrial ecosystems.

The mid-Cretaceous is characterised as an interval of angiosperm diversification and rise to

ecological dominance (e.g. Wolfe et al., 1975; Crane et al., 1995; Lupia et al., 1999).

Angiosperm diversification was diachronous, occurring at equatorial palaeolatitudes prior to the

mid- and high latitudes. This pattern of diversification is well-established in the Northern

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Hemisphere (Lidgard and Crane, 1988, 1990), with preliminary evidence supporting a

comparable trend for the Southern Hemisphere (Drinnan and Crane, 1990; Cantrill and Poole,

2002; Nagalingum et al., 2002). Over the same interval, relative diversity trends of free-sporing

plants (ferns, lycophytes and non-vascular plants) consistently show an overall decrease

(Northern Hemisphere, Lupia et al., 1999; Antarctica, Cantrill and Poole, 2002; Australia,

Nagalingum et al., 2002), raising the possibility of ecologic replacement (Lupia et al., 1999).

However, the decline of fern diversity during the Cretaceous was not a uniform trend across all

fern groups. Molecular data suggest that some fern groups, most notably the polypods, which are

the most diverse extant clade, show an overall increase in diversity (Pryer et al., 2004; Schneider

et al., 2004). Of the free-sporing plant groups that exhibit a decline in diversity and abundance

during the mid- to Late Cretaceous, osmundaceous ferns appear to have been particularly

affected (e.g. Nagalingum et al., 2002).

Most gondwanan Cretaceous free-sporing plant fossils have been recovered from Lower

Cretaceous strata as a consequence, at least in part, of this mid- to Late Cretaceous floral

overturn. Early Cretaceous cryptogam leaf floras of Gondwana have been primarily reported

from the Antarctic Peninsula (Jefferson, 1981; Cantrill, 1995, 1996, 1997, 1998; Nagalingum,

2003; Cantrill and Nagalingum, 2005; Nagalingum and Cantrill, 2006; Cantrill and Poole, 2012),

and Australia (Seward, 1904; Douglas, 1973; Drinnan and Chambers, 1986; McLoughlin, 1996;

McLoughlin et al., 1995, 2002). These studies consistently show a highly diverse fern

component, with rare lycopods and horsetails, and locally dominant, non-vascular plants. Indeed,

the upper Lower Cretaceous strata of Alexander Island, Antarctic Peninsula, have yielded a

macrofloral assemblage where ferns comprise the most diverse group (Falcon-Lang et al., 2001;

Nagalingum, 2003). This high diversity of free-sporing plants is reflected in the Early

Cretaceous palynological records of both southeast Australia (e.g. Dettmann, 1986, 1994;

Nagalingum et al., 2002; Wagstaff et al., 2013) and the Antarctic Peninsula (Dettmann and

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Thomson, 1987; Cantrill and Poole, 2002), with both regions typically exhibiting a greater

diversity of free-sporing plants than of seed-plants.

There are only a few reports of mid-Cretaceous macrofloral assemblages from South Island,

New Zealand, by Ettingshausen (1887), McQueen (1956) and Daniel (1989). Daniel (1989)

illustrated several cryptogams (pls. 10–11) from the mid-Cretaceous (Albian/Cenomanian) strata

of the Clarence Valley, South Island, and included a comparison of various informally described

fossil pinnules of the extinct osmundaceous Phyllopteroides; several of these were regarded as

species by Parrish et al. (1998), but without formal taxonomic descriptions. Pole and Philippe

(2010) reported dispersed gymnosperm and angiosperm leaf cuticle taxa from the Cenomanian–

Turonian Tupuangi Formation of the Chatham Islands. Recent fieldwork on the Chatham Islands

has yielded an extensive macroflora, which serves as the basis for the present study — the first

of a series of contributions to document this floral assemblage. This is the first systematic study

of probable cryptogamic leaf fossils from eastern Zealandia. This early Late Cretaceous flora

will provide an important data-point for future studies of floristic change across the southern

high latitudes.

2. Geological background

The Chatham Islands are located ~ 850 km east of Christchurch, New Zealand (~ latitude:

44°S; longitude: 176°W), and are the only emergent part of an elongated plateau of continental

crust known as the Chatham Rise (Fig. 1A). The Chatham Rise extends eastward from South

Island, New Zealand, and is a segment of the coherent and mostly submerged subcontinent

Zealandia (including New Zealand, New Caledonia, the Lord Howe Rise and the Campbell

Plateau). Prior to Late Cretaceous continental breakup, Zealandia was conjoined with the

Australian and Antarctic sectors of eastern Gondwana (Fig. 2). The lower Upper Cretaceous

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fossiliferous terrestrial sequence of the Chatham Islands (the Tupuangi Formation) was

deposited on the margins of the Bounty Trough, an extensional basin between the Chatham Rise

and the Campbell Plateau (Fig. 1A). This basin formed prior to, or concurrent with, Zealandia-

Gondwana rifting (Bradshaw et al., 1996; Eagles et al., 2004). During the early Late Cretaceous,

the Chatham Rise was primarily emergent (Wood et al., 1989; Campbell et al., 1993) and located

within the south polar circle (palaeolatitude: ~ 70–80°S; Markwick et al., 2000; Mukasa and

Dalziel, 2000; Stilwell et al., 2006).

<INSERT FIG. 1>

The Tupuangi Formation is the oldest sedimentary stratigraphic unit exposed on the Chatham

Islands (Campbell et al., 1993). On the basis of spore-pollen biostratigraphy, the formation was

originally correlated to the Motuan to Mangaotanean New Zealand chronostratigraphic stages

(upper Albian to Turonian; R.A. Couper in Hay et al., 1970), but Mildenhall (1994) proposed a

correlation to the Teratan New Zealand stage (Coniacian) for the uppermost strata. The

biostratigraphic range was later refined to Ngaterian to Mangaotanean (Cenomanian to

Turonian; Mays and Stilwell, 2013), as no autochthonous Motuan or Teratan spore-pollen

assemblages could be verified. The Tupuangi Formation is overlain by the Kahuitara Tuff. A

probable coeval unit of the Kahuitara Tuff, the Southern Volcanics, has been radiometrically

dated with a maximum age of 85.45 ± 0.6 Ma (Piripauan; Santonian; Panter et al., 2006). This

suggests an unconformity between the Tupuangi Formation and the Kahuitara Tuff (Mays and

Stilwell, 2013).

<INSERT FIG. 2>

The Tupuangi Formation has an estimated exposed thickness of ~ 400 m (Campbell et al.,

1993; Mays and Stilwell, 2013), and has only been reported from stratigraphic sections on Pitt

Island (Māori: Rangiauria; Moriori: Rangiaotea). Pitt Island is the second largest island of the

Chatham Islands archipelago, and is located 22 km southeast of Chatham Island (Fig. 1B). The

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Tupuangi Formation consists primarily of interbedded, poorly-consolidated, quartzofeldspathic

fine sandstone and carbonaceous siltstone facies (Campbell et al., 1993). Lignitic layers (≤ 1.2 m

thick) and large in situ tree stumps with associated root systems are present at discrete horizons

throughout the interbedded silt-/sandstone succession. The Tupuangi Formation has been

interpreted as primarily fluvial to deltaic (Campbell et al., 1993), with palynological evidence of

restricted marine influence from the uppermost (Turonian) strata (Mays and Stilwell, 2013).

3. Materials and methods

<INSERT FIG. 3>

Fossil material from Pitt Island was collected by C. Mays, D.J. Cantrill, J.D. Stilwell and

volunteer field assistants from Monash University, Clayton, Australia, during two field seasons:

January–February, 2009, and February, 2012. Fossils were obtained from outcrops at Waihere

Bay, western Pitt Island (Fig. 3). All specimens were collected from the Tupuangi Formation.

Individual hand samples are provided with unique sample registration numbers, denoted by the

prefix “PL”, and are registered with GNS Science, Lower Hutt, New Zealand. An individual

fossil specimen may comprise multiple samples, e.g., part and counterpart pairs. Localities are

given Fossil Record File numbers, and provided with prefix “CH/f”. To provide stratigraphic

context, stratigraphic logs were made of all reported fossil localities, except for CH/f0774, which

was a non-coastal outcrop and could not be reliably correlated to the coastal Waihere Bay

succession (Fig. 4). Locality and sample details, including figure numbers and part/counterpart

relationships, are provided herein as supplementary data (Table 1).

<INSERT FIG. 4>

Macrophotographs were taken using a Canon Powershot SX120 IS; photomicroscopy was

undertaken using a Leica M205-C, with a Leica DFC290 camera. Line drawings were performed

with a Wacom Bamboo pen tablet. Bold lines of line drawings represent the preserved margins

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of the fossil specimens, but due to a range of taphonomic factors, these may not necessarily be

the margin of the specimen prior to burial; dashed lines represent inferred margins or veins

(unless otherwise stated).

Terms employed in the systematic descriptions conform to the definitions of Ellis et al.

(2009) where relevant; all other taxonomic descriptive terms follow Harris and Woolf-Harris

(2001). Where referred to, extant subclasses, orders, families and genera follow the classification

of Christenhusz et al. (2011). Stratigraphic and sedimentological terms are after Tucker (2011);

palaeosol terms are after Retallack (2001), except where otherwise stated.

4. Results

4.1 Stratigraphic context of fossil localities

The fossil plants are found consistently within massive to weakly laminated to bedded, poorly

consolidated, grey to light brown (chroma: 1–2, as per Munsell, 1975), carbonaceous clay- to

siltstone facies. Laminae are typically defined by beds of leaf remains (generally comminuted);

siltstone fossiliferous lamina surfaces are slightly undulate. Locally, fossiliferous layers are bed-

scale size (1–15 cm) and composed of compressed, carbonised plant remains, but these

fossiliferous beds are not suitable for taxonomy due to the density of fossil material and

distortion of individual specimens. The fossiliferous fine-grained sedimentary packages

represent the upper sections of fining-upward successions 30 to 250 cm thick; these are overlain,

and commonly truncated, by beds of massive to cross-bedded fine- to medium-grained sandstone

(Fig. 4A, B). Fossil roots/rootlets extending below the fossil beds were identified at three

localities (CH/f0775, 0776 and 0778). The sediments at three localities appeared to be partly to

highly weathered as indicated by the light brown colouration (chroma: 2; CH/f0776–0778). The

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fossils from these localities were typically preserved as oxidised impressions. Oxidation appears

to be most intense on outcrop surfaces and adjacent to faults/joints; as such, this feature was

most likely due to late-stage diagenetic factors such as surficial weathering and ground-water

penetration via rock fractures. Localities CH/f0776 and 0778 are rich in sulphide content,

typically pyrite and marcasite.

4.2 Systematic palaeontology

Marchantiophyta Crandall-Stotler and Stotler, 2000

Class incertae sedis

Hepaticites Walton, 1925

Type species

Hepaticites langii Walton, 1925 (lectotype chosen by Schuster, 1966)

Hepaticites sp. (Fig. 5A–C)

Material examined

One specimen as follows: PL1014 (Fig. 5A–C).

Description

Plant thalloid, spreading, prostrate, preserved portion 24 mm in diameter. Individual thallus

branches are 1.2–1.8 mm wide, ≤ 15.5 mm long, dichotomising up to five times, at irregular

intervals (1–3.8 mm); final branch dichotomy diverges at 26–44°, other dichotomies diverge at

39–86°. Thallus apices rounded, margins entire except notched at apices (Fig. 5C). Costa 0.1–0.5

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mm wide where discernible, thinning towards terminal branches; thalli are darkest at costa and

gradationally lighten towards margins. Sporophytic remains not present.

Comparison and remarks

The present taxon is assigned to the fossil-genus Hepaticites, which includes all fossil taxa

having an apparent relationship to the “Hepaticae”, following the system employed by

Oostendorp (1987). Recent phylogenetic reviews of extant liverworts have disbanded Hepaticae,

and recognised three distinct classes under Marchantiophyta: Jungermanniopsida,

Haplomitriopsida and Marchantiopsida (Crandall-Stotler and Stotler, 2000; Crandall-Stotler et

al., 2009). The thalloid organisation of the present taxon is most similar to extant members of

Marchantiopsida or Jungermanniopsida, but the features necessary (e.g. sporophytic remains,

cellular structure, rhizoids) to satisfactorily attribute it to either of these extant classes are not

preserved. The gradual lightening of colour from the midribs suggests that the original tissue

was thinner away from the costal regions and in the thallus apices; this feature is similar to

Hepaticites undulatus Drinnan and Chambers, 1986 but the thalli of the present taxon are much

smaller. Hepaticites minutus Cantrill, 1997 and H. arcuatus (Lindley and Hutton, 1837) Harris,

1942 both feature similar branching patterns, but the thalli of the former are consistently smaller,

whilst those of the latter are larger, and both lack the apical notches that characterise the

presently described specimen. The anthocerophyte Dendroceros victoriensis Drinnan and

Chambers, 1986 is similar to the present specimen by having notched thallus apices and a

branching, prostrate, spreading habit, but the present specimen is smaller and lacks the

preservation quality to express the diagnostic arcuate ribs of D. victoriensis.

<INSERT FIG. 5>

Bryophyta Schimp.

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Class incertae sedis

Muscites Brongniart, 1828

Type species

Muscites tournali Brongniart, 1828

Muscites gracilis Mays and Cantrill sp. nov. (Fig. 5D–G)

Holotype

Holotype specimen comprises two hand samples, part (PL1027; Fig. 5D–F) and counterpart

(PL1028). Holotype is from locality CH/f0777, Waihere Bay, Pitt Island, Chatham Islands, New

Zealand, Tupuangi Formation, Cenomanian.

Other material

Two specimens as follows: PL1033, PL1034. Both from locality CH/f0778.

Etymology

Latin gracilis, meaning slender or narrow, describing the axis and the diagnostic leaf shape.

Diagnosis

Axes furcate into three–five branches; leaves decurrent, linear to lanceolate with involute

margins and prominent costae.

Description

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Branched axis habit, preserved axis length ≤ 24 mm, 0.3–0.8 mm wide; axis furcates into

three–five separate branches at 25–50° at discrete nodes spaced 7–15 mm apart. Axes appear to

bear leaves along their entire length, but insertion relationship is not clear; leaf lengths increase

towards the distal ends of the axes. Leaves are sessile, decurrent and reflexed (deviating from

stem asymptotically, but reflexing typically ≤ 65°, but as much as 180° in rare cases; see Fig. 5E,

F), linear to lanceolate (1.8–3 mm length, 0.2–0.5 mm basal width) and apices are acute; leaf

margins are entire and are involute. Costae are prominent, extending to apices and may be

excurrent. Lamina colour lightens from costa to margins (see Fig. 5G). Sporophytes are not

present.

Comparison and remarks

The present taxon has a branched pattern of strand-like stems and sessile leaves with distinct

midribs, as per the original diagnosis of the fossil-genus Muscites (Brongniart, 1828). These

specimens are similar to various extant taxa of Polytrichopsida (e.g. Polytrichastrum G.L. Sm.).

The gross morphology of the present species resembles the branching fossil pleurocarp

Palaeodichelyma sinitzae Ignatov and Shcherbakov, 2007 which has a probable affinity to

Bryopsida. However, the cellular details are not discernible due to the resolution constraints

imposed by the enclosing coarse sediments. This factor, combined with an absence of preserved

sporophytic details, precludes assignment to class level.

The leaves of this taxon have been flattened against the enclosing sediments in dorsiventral,

lateral and intermediate orientations. The leaves have been described as involute because of the

basiscopic position of the costa on the laterally compressed leaves diverging from the medial

portions of the stem (see Fig. 5G). However, when the leaves are compressed dorsiventrally, as

at the distal end of the stem (see Fig. 5E, F), it is evident that the costae are situated in the

middle of the lamina.

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The linear to lanceolate leaves and abaxial midribs of Muscites gracilis sp. nov. are similar to

M. antarcticus Cantrill, 2000 recovered from Aptian strata of Snow Island, Antarctica; PL1034

from the Tupuangi flora (Fig. 5G) appears particularly similar, this specimen having slightly

broader leaves than the holotype. However, the present taxon consistently has more slender axes,

and longer, thinner leaves. The taxon denoted “gametophyte type 2” described by Drinnan and

Chambers (1986) from Lower Cretaceous strata of southeastern Australia features linear leaves

and a branching habit, but the leaves appear to be excurrent rather than consistently decurrent,

and the axes are too robust to be conspecific with the presently described taxon. This is the first

Cretaceous moss macrofossil to be described from New Zealand.

Lycopodiophyta Cronquist et al., 1966

Lycopodiopsida Bartling, 1830 (as "Lycopodineae")

Order incertae sedis

Lycopodites Lindley and Hutton, 1833 non Brongniart, 1822

Type species

Lycopodites falcatus Lindley and Hutton, 1833 emend. Harris, 1961

Lycopodites sp. (Fig. 5H)

Material examined

One specimen (PL1002).

Description

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Dichotomous secondary axis branches from primary axis at 55°, primary axis width 0.19–

0.24 mm, secondary axis width 0.16–0.2 mm. Homophyllous; leaves sessile, erect, laterally

complanate, alternately arranged; leaves crowded but not overlapping, spaced 0.8–1.2 mm (as

measured from basal margins). Leaf elliptical, apex straight to acuminate (possibly mucronate),

proximal margin of lamina approximately at right angles to stem. Primary axis leaf length 1.4–

1.6 mm, width 0.8–1.0 mm; secondary axis leaf length 1.1–1.2 mm, width 0.5–0.6 mm. Lamina

thickenings evident on proximal (width: 80–150 µm) and distal (width: 60–90 µm) margins.

Fertile organs not present and cellular detail not preserved.

Comparison and remarks

A probable affinity to Lycopodiales has been proposed for various species of Lycopodites,

e.g. L. falcatus Lindley and Hutton, 1833 emend. Harris, 1961 (Harris, 1961); L. victoriae

Seward, 1904 (Douglas, 1973). The same conclusion cannot be made for the present taxon, due

to the absence of preserved homosporous or heterosporous strobili, which would aid in

distinguishing between an affinity with Lycopodiales or Selaginellales, respectively. There

appears to be a distinct size difference between the leaves of the two preserved stems of this

specimen, suggesting a growth habit with subsidiary stems branching from a primary stem. Such

a feature is expressed in various extant species of Selaginella (e.g. S. hordeiformis Baker; S.

novae‑hollandiae (Sw.) Spring). Conversely, the homophyllous habit is suggestive of a

lycopodiaceous affinity (see discussion by Harris, 1961). Furthermore, complanate leaves are

common amongst extant lycpodiaceous species (e.g. Diphasiastrum complanatum (L.) Holub, D.

digitatum (Dillenius ex A. Braun) Holub), whilst this character is rare amongst Selaginellales.

Lycopodites gracilis Oldham and Morris, 1863 and the type species L. falcatus Lindley and

Hutton, 1833 emend. Harris, 1961 are both heterophyllous, but the present taxon is

homophyllous with no evidence of decussate phyllotaxis. Lycopodites victoriae Seward, 1904 is

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homophyllous, but the leaves are typically larger, falcate and have a prominent midrib. The

leaves of L. arberi Edwards, 1934 are erect, similar to the presently described specimen, but are

larger, tend to overlap and have mucronate apices; the present taxon may have this latter feature,

but the apices are not adequately preserved to be described as such. Walkom (1928) described

two lycopodiaceous taxa from Early Cretaceous (Burger, 1980) strata of northeast Australia:

?Lycopodites sp. A and B. The former has a prominent midrib and sub-opposite leaves, whereas

the presently described specimen has alternately arranged leaves with no visible midrib; the

latter has long, lanceolate leaves, rather than the short, elliptical leaves of the Tupuangi taxon.

Lycopodites sp. reported by Rigby (1977) from Middle Jurassic strata of Queensland, Australia

(redrawn and ascribed to L. gracilis (Morris) Seward and Sahni, 1920 by Mcloughlin et al., in

press), has falcate, rather than elliptical, leaves. McQueen (1956) recorded a specimen of

‘Lycopdium cf. volubile Forster f. 1786’ from the Seymour River coal measures (Warder

Formation), middle Clarence Valley, New Zealand, which have since been estimated as Albian–

Cenomanian in age (Douglas and Williams, 1982; Browne and Reay, 1993). The specimen

illustrated is much larger than the present specimen, and exhibits linear, falcate leaves. Upon re-

examination of the specimen recorded by McQueen, Daniel (1989) suggested that it has a

probable podocarpaceous affinity; indeed, the specimen appears to have close resemblance to the

conifer Pagiophyllum, a common element recorded from Albian strata of Alexander Island,

Antarctic Peninsula (as Pagiophyllum sp.; Cantrill and Falcon-Lang, 2001).

An interesting feature of the presently described specimen is the presence of thickened tissue

on both the proximal and distal lamina margins, giving the appearance of a bicostate leaf

architecture; however, no recorded extant member of Lycopodium has such venation (e.g. Chu,

1974). Further, Lycopodiaceae is defined by a single vein (Mirbel in Lamarck and Mirbel, 1802,

in Øllgaard, 1990), whereas a bicostate habit would imply an affinity outside this group. To

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satisfactorily conclude that the present taxon has two marginate costae will require cellular

details that cannot be ascertained from the present specimen.

<INSERT FIG. 6>

Polypodiophyta Cronquist et al., 1966

Polypodiopsida Cronquist et al., 1966

?Osmundales Link, 1833 (as “Osmundaceae”)

Cladophlebis Brongniart, 1849

Type species

Cladophlebis albertsii (Dunker, 1846) Brongniart, 1849

Cladophlebis auriculipilosus Mays and Tosolini, sp. nov. (Fig. 6)

Holotype

Comprises two hand samples, part (PL1036; Fig. 6A, B) and counterpart (PL1035; Fig. 6C–

E). Holotype is from locality CH/f0778, Waihere Bay, Pitt Island, Chatham Islands, New

Zealand, Tupuangi Formation, Cenomanian.

Other material

One specimen from locality CH/f0778 comprising two hand samples: PL1031 (part), PL1032

(counterpart).

Etymology

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Latin auriculi-, from auricula, meaning ear in reference to the diagnostic pinnule auricles;

Latin -pilosus, meaning covered in hair referring to the distinctive hair-like trichomes of the

rachises and rachillae.

Diagnosis

Rachises and rachillae have trichomes with longitudinally elongated bases. Alternating,

sessile pinnae. Pinnules adnate, ovate. Minor auricle on acroscopic pinnule margin, basiscopic

margin decurrent to rachilla, apices convex to rounded. Pecopteroid venation, primary vein

decurrent, distally reflexed.

Description

Frond at least bipinnate (complete frond not preserved), preserved frond 58 mm long (total

length unknown). Rachis ≤ 1.8 mm wide with no lamina wing. Rachis and rachillae feature

trichomes with longitudinally elongated decurrent bases (≤ 1.75 mm long, ≤ 0.2 mm wide; see

Fig. 6C). Preserved pinnae ≤ 45 mm long (total length not known) by ≤ 13 mm wide, distichous

and alternate arrangement, rachillae ≤ 1 mm wide, sessile and inserted laterally or slightly

dorsally on rachis at 65–70°. Pinna planes slightly tilted (5–10°) relative to the plane of the

rachis. Pinnules ≤ 7.5 mm long by ≤ 4.9 mm wide, distichous and sub-opposite arrangement,

inserted laterally on rachilla with adnate attachment; pinnules oblong, apex shape convex to

rounded. Pinnules are typically curved perpendicular to plane of the lamina (?revolute). Pinnules

along proximal rachilla typically discrete, medial pinnules conjoined at bases, complete fusion of

pinnule margins at apices. Isolated pinnules are crowded to partially superimposed, density of

isolated pinnules is approximately 2.3–3.6 per 1 cm along rachillae, increasing apically along

pinna. Pinnule margins entire to sinuous, acroscopic margins are straight to convex but widening

with small auricle and constricted basally, basiscopic margin is straight to convex and basally

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decurrent to the rachilla; basiscopic margins of discrete pinnules typically exhibit overlap with

part of the acroscopic auricle of the subtending pinnule. Venation pattern is pecopteroid; primary

vein, ≤ 0.2 mm wide, diverges from rachilla decurrently, reflexing distally into pinnule, straight

to sympodial in the medial portion of pinnule, angle between medial primary vein and rachilla

65–90°. Primary vein persists for at least two thirds of pinnule length, but evanesces into two

apical secondary veins. From primary vein, ≤ 10 distichously arranged secondary veins diverge

at ≤ 55°, dichotomising once, or less commonly twice, before reaching pinnule margin. Vein

density is approximately 11–15 per 10 mm along the pinnule margin. Initial secondary vein

typically anadromic. Acroscopic secondary vein diverges from primary vein at an angle

approximately parallel to rachilla, dichotomising veinlets terminate at both auricle apex and

sinus. No fertile material was preserved.

Comparison and remarks

The diagnosis of Cladophlebis Brongniart, 1849 describes sterile fronds with pecopteroid

venation, whereas equivalent fertile fronds are referred to several other genera, most notably

Osmundopsis Harris, 1931 emend. Harris, 1961 and Todites Seward, 1900 emend. Harris, 1961

(see review by Balme, 1995). This association of Osmunda-like fertile fronds with their sterile

counterparts provided strong support for a phylogenetic relationship for various members of

Cladophlebis with Osmundales (Phipps et al., 1998). The rachises/rachillae of Osmundaceae

feature grooves and ridges, as per the original familial description by Berchtold and Presl (1820,

fides Kramer, 1990). As such, the presence of diagnostic trichomes on the rachises/rachillae

precludes a definitive association of the presently described species to Osmundaceae.

Furthermore, there have been reported specimens of Cladophlebis that are not likely to be allied

to Osmundales. Cladophlebis cyathifolia Villar de Seoane, 1996 has been found with in situ

Cyathidites-type spores, suggesting an affinity to Cyatheales (Villar de Seoane, 1996, 1999); this

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taxon was later referred to Cyathea cyathifolia (Villar de Seoane, 1996) Villar de Seoane, 1999.

The spores associated with Cladophlebis tripinnata Archangelsky, 1963 emend. Villar de

Seoane, 1996 exhibit both monolete and trilete forms, which Villar de Seoane (1996) interpreted

as indicating an affinity with Dennstaedtiaceae (Polypodiales). However, this is not exclusively a

feature of Dennstaedtiaceae; many species of disparate extant fern groups have been known to

produce both monolete and trilete spores (Selling, 1944; Brown, 1960). A range of fossil species

also exhibit this feature, including members of Marattiales (e.g. Marattia anglica (Thomas)

Harris, 1961; see review by Murdock, 2008) and Dipteridaceae (Polypodiales; e.g. Polyphacelus

stormensis, Yao et al., 1991). This suggests that a mix of spore aperture types from an individual

fertile frond is not a reliable diagnostic feature for any specific fern group; however, no cases of

fertile fossil fronds of unequivocal osmundalean affinity have yet been found with a combination

of monolete and trilete spores. These examples emphasise the probability that Cladophlebis is

not limited to Osmundales. Because Cladophlebis is most commonly associated with

Osmundales, this study tentatively assigns an osmundalean affinity to C. auriculipilosus;

however, without the preservation of fertile material, or the observation of anomocytic stomata

on the lamina surface, uncertainty remains over the assignment of this species to order (or

family), following the recommendation of Seward (1904).

The sterile fronds of Todites williamsonii (Brongniart, 1828) Seward, 1900 emend. Harris,

1961 have sessile, cladophleboid, typically isolated pinnules, similar to the presently described

specimens, but have acute apices that curve acroscopically. The pinnules of Cladophlebis

octonerva Holmes, 2003 from the Triassic of New South Wales, Australia, lack both auricles on

the acroscopic margin and trichomes along rachis/rachilla (Holmes, 2003). Douglas (1973)

reported a similar taxon (Cladophlebis sp. “a”) from the Early Cretaceous (?Berriasian–

Hauterivian) strata of southeastern Australia; however, the pinnules were much more elongate,

were not decurrent on the basiscopic margin, and lacked auricles. There have been several

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reported specimens of Cladophlebis from Cretaceous strata of New Zealand, yet none of these

taxa has equivalent pinnule shape (see Table 2). The species described herein is similar in many

respects to the specimen denoted Cladophlebis sp. by Drinnan and Chambers (1986) from the

Lower Cretaceous (Aptian–Albian; Dettmann, 1986) of Victoria, Australia. Shared features

include: pinnule sizes with entire margins; recurved and flexuose midveins; and variable

presence of an auricle on the acroscopic margins of the laminae. However, the alternate pinnule

arrangement, hastate to falcate pinnule shape, and the consistently catadromic initial secondary

veins of Cladophlebis sp., are not consistent with C. auriculipilosus. Furthermore, upon re-

examination of the original specimen of Cladophlebis sp., the specimen has a distinctly glabrous

rachilla, thus lacking the diagnostic trichomes on the rachillae of C. auriculipilosus.

<INSERT TABLE 2>

Sterile pinnules of extant members of Osmundales typically have better expression of the

venation on the abaxial, rather than adaxial, lamina surfaces (e.g. Osmunda claytoniana L.); if

we assume the same feature for the present specimens, then the prominent grooves on the

impression specimens of C. auriculipilosus (Fig. 6D) represent a mould of the abaxial surfaces

of the pinnae. Therefore, the consistent curvature of the pinnule surfaces would suggest revolute,

rather than involute, lamina margins. Furthermore, members of Osmundales with adnate and

discrete or basally fused pinnules, tend to exhibit sub-planar to revolute pinnule margins (e.g.

Osmundastrum cinnamomeum (L.) C. Presl). There are precedents of revolute pinnule margins

being preserved in various fossil ferns, and these have been attributed to original leaf

physiognomy rather than taphonomy (e.g. Rothwell, 1978; Delevoryas et al., 1992). The

inference of revolute lamina margins provides a method of orienting the frond relative to the

whole plant whilst it was in its living position, despite a lack of preserved cellular details. Where

the pinnule margins are superimposed, the adaxial surfaces of the pinnule auricles most often

underlie the decurrent acropetal pinnule margins (auricle underlap to overlap ratio = 72:29; data

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pooled from both recovered impression samples). A reconstruction is presented (Fig. 7), which

highlights this tendency of the basiscopic margins of the pinnules to overlap the auricles of the

subtending pinnules.

<INSERT FIG. 7>

The pinna planes of the holotype are tilted 5–10° relative to the rachis axis; this angle was

likely greater prior to compaction under sedimentary overburden. All of the preserved pinnae

attached to a rachis have a consistent orientation, with the adaxial surfaces rotated towards the

frond apex. This observation is not a particularly useful diagnostic feature by itself, but this

consistent orientation may represent evidence of a heliotropic adaptation, as expressed by many

extant erect herbaceous ferns; e.g. Dryopteris erythrosora (D.C. Eaton) Kuntze has pinnae that

approach 90° from the plane of the rachis (Iino, 2001). Alternatively, such pinna orientation

could be the result of sedimentary processes, whereby the plant remains have become oriented

with respect to the flow direction during deposition (Bateman, 1999), as is common with oblate

pebble clasts in aqueous depositional environments with a unidirectional flow (SenGupta, 1966).

The fossiliferous beds from which C. auriculipilosus was recovered lacked any discernible

sedimentary structures with which to estimate palaeocurrent directions. Thus, it is not possible to

determine whether the preserved orientation of the pinnae whilst in outcrop was consistent with

the depositional current direction. If further specimens yield a consistent pinna orientation

relative to the rachis, regardless of their orientations relative to the strata, this would point

towards an inherent anatomical feature, consistent with an erect growth habit, rather than being

caused by taphonomic effects.

<INSERT FIG. 8>

Order incertae sedis

Sphenopteris (Brongniart, 1822) Sternberg, 1825

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Type species

Sphenopteris elegans (Brongniart, 1822) Sternberg, 1825

Sphenopteris sp. cf. S. warragulensis McCoy in Stirling, 1892 (Fig. 8)

Synonymy

1995 Sphenopteris sp. cf. S. warragulensis McCoy in Stirling, 1892; McLoughlin et al., p.

283, fig. 4A.

Material examined

Fifteen samples comprising eleven individual specimens with four counterparts (PL1008–13,

PL1015–21, PL1023 and PL1024).

Description

Frond at least pinnate, base and apex not preserved. Preserved rachis length ≤ 62 mm, width ≤

2 mm, prominent longitudinal grooves, no lamina wing. Pinna attachment petiolulate,

distichously and alternately arranged at 10–18 mm intervals along rachis, insertion on rachis at ≤

35°. Pinnae are lanceolate with acute bases and alternate to sub-opposite, compound, pinnatifid

lobes. Lobes are ovate to lanceolate, have acute, lobulate to serrate distal margins (preserved

dimensions: ≤ 17 mm long by ≤ 5.5 mm wide); lobes are interconnected by narrow, interlobate

lamina (≤ 1.2 mm total width) parallel to primary vein (≤ 0.3 mm wide). Venation is

sphenopteroid; primary vein straight to sinuous, preserved length ≤ 39 mm. Single secondary

veins enter each lobe, diverging from the primary vein at ≤ 30°, spaced at ≤ 14 mm intervals,

and persisting to the distal margin. Secondary veins straight to sinuous, ≤ 0.2 mm wide. Tertiary

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veins dichotomise from secondary veins at ≤ 30°, dichotomies spaced at ≤ 2.3 mm intervals.

Tertiary veins dichotomise ≤ six times into lateral veinlets, each terminating at the acute tips of

the distal serrate/lobulate margins; lateral veinlets do not dichotomise. The initial tertiary veins

of each lobe are catadromic. No fertile material was preserved.

Comparison and remarks

The presently described specimens have lamina lobes of widely variable length, width, degree

of vein dichotomy and margin dissection; however, the values of these variables appear to fall

upon a continuous spectrum, suggesting natural intraspecific variation. Thus, there are no

adequate grounds to assign the present specimens to more than one taxon. This conservative

approach is supported by the large, well-preserved specimens of Sphenopteris warragulensis

McCoy, 1892 described by Drinnan and Chambers (1986) from Aptian–Albian strata of

southeastern Australia, which have remarkable intraspecimen variability. Specifically, the

proximal pinnae and associated pinnules were much more likely to be discrete than those closer

to the distal ends of the fronds, the latter becoming “pinnatifid rather than pinnate” (Drinnan and

Chambers, 1986, p. 28). The synonymous Sphenopteris sp. cf. S. warragulensis of McLoughlin

et al. (1995) has a pinna habit described as pinnulate, but under the presently employed

taxonomic scheme (Ellis et al., 2009), these pinnae would be described as lobate rather than

pinnulate, due to the distinctive interconnecting laminar tissue between alternating secondary

veins. The degree of pinnatification for the present taxon is variable across specimens, with most

showing distinct interlobate laminae (e.g. Fig. 8C, D), whilst some specimens have lobes that

approach a pinnulate form (e.g. Fig. 8E, F). Larger preserved specimens will likely reveal that

the variants of the present taxon merely represent different sterile portions of fronds of the same

species.

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The Aptian–Albian southeast Australian specimens of Sphenopteris warragulensis described

by Drinnan and Chambers (1986) featured tertiary veins (‘secondary veins’ therein), which

dichotomise only once or twice, whereas those from the Chatham Islands exhibit up to six

dichotomies; all other features of the presently described specimens fall within the prescribed

variations for S. warragulensis. The similarity of this taxon to fertile fronds of the

leptosporangiate S. warragulensis from southeastern Australia facilitates the assignment of this

taxon to Polypodiopsida, but no order can be definitively ascribed at present. This taxon differs

from S. travisii Stirling, 1900 ex Drinnan and Chambers, 1986 by having longer and more

closely-spaced lobes (pinnules). Unlike the present taxon, the tertiary-order veins in

Sphenopteris göepperti Dunker, 1846 do not dichotomise. The pinnae of S. nordenskjoeldii

Halle, 1913 emend. Rees and Cleal, 2004 exhibit a “pseudo-dichotomous dissection” (Halle,

1913, p. 26), a description that emphasises the lobate (rather than pinnate) arrangement of the

pinnae as seen in the present specimens. However, the veins of S. nordenskjoeldii consistently

terminate in lobules (‘secondary lobes’ of Rees and Cleal, 2004), whereas the present taxon

commonly has serrate lobe margins. The Tupuangi specimens exhibit slightly larger lobes than

Sphenopteris sp. cf. S. warragulensis from Cenomanian strata of Queensland, as described by

McLoughlin et al. (1995), but all other features are comparable.

<INSERT FIG. 9>

Adiantites Göeppert, 1836 emend. Kidston, 1923

Type species

Adiantites oblongifolius Göeppert, 1836 (lectotype chosen by Andrews, 1970)

Adiantites sp. (Fig. 9)

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Synonymy

cf. 2007 Adiantopteris tripinnata Cladera et al., pp. 53, 54, fig. 4B, C.

Material examined

Twenty-nine specimens (no counterparts) as follows: PL1000, PL1001, PL1005–7, PL1022,

PL1025, PL1026, PL1030, PL1039–58.

Description

Rachis ≤ 95 mm long, ≤ 2.2 mm wide, sympodially branching, with shallow longitudinal

grooves and no lamina wing. Pinnae have a distichous and alternative to sub-opposite

arrangement on rachis (at intervals of ≤ 12 mm), and are dimorphic. Petiolules attach at lamina

margin, and typically diverge from rachis decurrently but can be reflexed up to 100° relative to

rachis (Fig. 9E, F). Pinnae have cuneate bases and are alternately pinnatilobate, each lobe has

serrate distal margins. A single primary vein (≤ 0.4 mm wide) enters each pinna, is persistent for

less than a quarter of total pinna length, dichotomising evenly four–ten times at 10–20° each.

Each veinlet terminates at the tip of a lobe distal margin serration, ≤ 27 per lobe, one–two veins

per millimetre along distal margin. Elongated interlobate laminae extend to sinuses, and are

bordered by long, sub-parallel veinlets (Fig. 9B). Most pinnae are arranged alternately,

petiolulate to sub-petiolulate, elliptical to oblong, ≤ 31 mm long (28 mm excluding petiolule)

and ≤ 17.5 mm wide), with four–five distinct lobes; the two (or three) lobes situated on the

acroscopic side of the leaf are consistently longer than the two (or three) basiscopic lobes

(basiscopic to acroscopic lobe length average ratio approximately 1:1.7, as measured from the

initial vein dichotomy to lobe apex; Fig. 9E, F). Second type of pinna (?terminal) is larger (≤ 40

mm long by ≤ 28 mm wide) than lateral pinnae, basal shape is concave to acuminate, five–seven

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alternating distal lobes; lobes are mostly fused along their adjacent margins, basiscopic lobes

approach a pinnatifid habit. Lobes of ?terminal pinnae consistently increase in length distally

(with maximum pinna length equal to the axis of alternation; Fig. 9D), cf. lateral pinnae which

feature acroscopic lobes that are consistently longer than the basiscopic lobes. No fertile material

was preserved.

Comparison and remarks

At present, there are two common generic designations employed to categorise sterile leaf

taxa with foliage similar to the extant Adiantum L.: 1) Adiantites Göeppert, 1836 emend.

Kidston, 1923; and 2) Adiantopteris Vassilevskaja, 1963. Adiantites was initially erected to

comprise all stipitate, fan-shaped fossil leaves with the distinctive anadromous venation pattern

typical of Adiantum. As suggested by his choice of name, Göeppert (1836) suspected a fern

affinity for the genus. He based the diagnosis on various leaf taxa from upper Palaeozoic strata;

however, many of these taxa have since been shown to have probable gymnosperm affinities.

Support for this association include the preservation of Adiantites-type foliage in association

with the seed taxon Wardia White, 1904 (see discussion by White, 1936), and complex axis

vascularity (typical of Medullosales; Jennings, 1985). In terms of stratigraphic distribution, there

appear to be two distinct populations of extinct Adiantum-like taxa in the fossil record: a

Palaeozoic assemblage and a Mesozoic to late Cenozoic assemblage. Adiantopteris was erected

to reflect this, and was intended to apply only to the latter group (Vassilevskaja et al., 1963). In

contrast to the Palaeozoic specimens, the fertile counterparts of the Mesozoic–Cenozoic

Adiantum-like population have consistently featured marginal sori, reflecting a pteridophytic

affinity (e.g. Adiantites lindsayoides Sewell, 1904 emend. Drinnan and Chambers, 1986).

However, the diagnosis of Adiantopteris is not significantly differentiated morphologically from

Adiantites. Furthermore, as noted by Moseychik and Ryabinkina (2012), stratigraphic

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discrepancy does not provide adequate grounds for the reclassification of Adiantites taxa because

it does not account for the possible co-occurrence of gymnosperm and fern groups with

comparable sterile foliage. As such, the presently described material is assigned to Adiantites

rather than Adiantopteris, the former genus having priority.

A frond featuring the in situ attachment of the two described pinna types is not available

amongst any of the preserved specimens. Until the probable terminal pinnae are found in

association with the rachises, the interpretation of a single leaf species with a dimorphic habit

must be regarded as tentative. Based on the pinnate (typically bipinnate) leaf habit of similar

taxa (e.g. Adiantites dispersus Douglas, 1973, Adiantopteris tripinnata Cladera et al., 2007), it is

probable that the observed pattern of increasing sinus depth towards the base of the larger leaves

would continue basiscopically until leaves are pinnatisect, and ultimately petiolate further

proximally along the rachis; this would result in dimorphic leaves with an imparipinnate

arrangement on each axis. The present material is too degraded to gain a satisfactory description

of the pinna arrangement, and precludes assignment to any definite species. Despite this, the

preserved features allow for the exclusion of the present material from some species of

Adiantites. Adiantites lindsayoides from Lower Cretaceous strata of southeastern Australia

(Seward, 1904; Drinnan and Chambers, 1986) has two veins emerging from the base of each

pinna (cf. one for the presently described taxon). The pinnae of A. dispersus are typically 10 mm

long (up to 20 mm), are ‘wedge-shaped’ to weakly lobate and have veins that dichotomise up to

twice (Douglas, 1973); in contrast, the lateral pinnae of Adiantites sp. are much larger, have a

consistent, well-developed pinnatilobate shape, and veins that dichotomise several times. The

present specimens resemble Adiantites sewardi Yabe, 1905 [= Adiantopteris sewardii (Yabe,

1905) Vassilevskaja, 1963] from Lower Cretaceous strata of Korea (Yabe, 1905), eastern Russia

(Krassilov, 1967) and Japan (Kimura et al., 1979), but feature more prominent interlobate

sinuses. Adiantopteris tripinnata, as described by Cladera et al. (2007), matches the presently

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described taxon in all features except the prescribed size range; the specimens herein are all

consistently larger; however, this is not deemed a feature diagnostic enough to exclude the

synonymy outright, as the size differences could be due to a range of uncontrolled environmental

variables. Assignment of the present material to A. tripinnata is not possible until an individual

leaf with both axial and terminal pinnae is found. Specimens of Adiantites have been noted (as

cf. Adiantites and Adiantites sp.) from Cretaceous strata of the Antarctic Peninsula (Cantrill,

1997, and Leppe et al., 2007, respectively), but no formal descriptions were provided therein. To

date, there have been no previously reported specimens of Adiantites (or Adiantopteris) foliage

from New Zealand strata of any age.

Due to the palaeogeographic and geochronological similarity of the present specimens to

Cretaceous Gondwanan foliage of definite leptosporangiate fern affinity (e.g. A. lindsayoides),

and the absence of recorded Mesozoic Adiantites-type pteridosperm material, an assignment to

Polypodiopsida affinity is tentatively proposed for the present taxon until fertile material can

provide a firmer taxonomic placement. Furthermore, this discussion highlights the ambiguity

surrounding the genera presently employed for the assignment of fossil Adiantum-like foliage,

and a taxonomic revision of Adiantites and Adiantopteris is sorely needed.

5. Discussion

5.1 Implications for mid-Cretaceous floral ecology of Gondwana

The Cenomanian Tupuangi Formation flora is one of a handful of high-latitude floras from

the early Late Cretaceous of the Southern Hemisphere and, as such, gives important insights into

ecological changes from this part of the world. Cenomanian tropical sea-surface mean annual

temperatures were ~ 31–37°C, with a Cretaceous maximum during the Turonian (Bice et al.,

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2006); the warming trend was accompanied by a low thermal gradient, whereby the polar

regions experienced the greatest degree of warming over this interval (Huber et al., 2002). This

pattern of warming was mirrored by the rise to dominance of angiosperms in the vegetation and

concomitant ecological changes. Early Cretaceous macrofloras at high southern latitudes are rich

in liverworts (Douglas, 1973; Drinnan and Chambers, 1986; Cantrill, 1997, 2000; McLoughlin

et al., 2002), bryophytes (Drinnan and Chambers, 1986; Cantrill, 2000), and a particularly high

abundance and diversity of ferns (Ettingshausen, 1887; Seward, 1904; Halle, 1913; Arber, 1917;

Edwards, 1934; Medwell, 1954; McQueen, 1955; Douglas, 1973; Drinnan and Chambers, 1986;

Cantrill and Webb, 1987; Cantrill, 1995, 1996, 1998, 2000; Nagalingum, 2003; Cantrill and

Nagalingum, 2005; Nagalingum and Cantrill, 2006; Cantrill and Poole, 2012). These groups

appear to have been impacted by the rise to dominance of angiosperms across the southern high

latitudes (Cantrill and Poole, 2002; Nagalingum et al., 2002) but the severity, timing and the

degree of specific group interactions are not fully understood due to the scarcity of macrofloral

assemblages. For example, Antarctic studies have compared late Albian floras with Coniacian

floras (Cantrill and Poole, 2002; Poole and Cantrill, 2006), but the Tupuangi Formation flora

represents a critical intervening interval and demonstrates that, although these groups were still

present, their diversities and abundances were low. Approximately coeval assemblages have

been recorded from the Albian–Cenomanian (the “Unnamed Formation” at Bajo de los Corrales;

Passalia et al., 2001; Kachaike Formation; Passalia, 2007a, 2007b) and Cenomanian–Coniacian

(Mata Amarilla Formation; Iglesias et al., 2007) of Patagonia, South America (see review by

Archangelsky et al., 2009). Other important coeval floras include those of the Winton

Formation, Queensland, Australia (late Albian–Cenomanian; see review by McLoughlin et al.,

2010), and the Warder Formation, South Island, New Zealand (Ngaterian; ~ late Albian to mid-

Cenomanian; McQueen, 1956; Daniel, 1989; Parrish et al., 1998). With the exception of the

Cenomanian–Coniacian Mata Amarilla flora, all of these assemblages exhibit a diverse and

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abundant fern component, in conjunction with abundant conifers and subsidiary dicotyledonous

angiosperms. In contrast to these assemblages, the Tupuangi Formation macrofloral assemblage

appears to exhibit a depauperate fern component (this study) and a high diversity of

gymnosperms (Pole and Philippe, 2010); however, this conclusion is preliminary until

quantitative intergroup macrofloral data can be obtained. Compared to the Winton assemblage,

the approximately coeval palaeofloras of Patagonia show similarities in angiosperm morphology

(Mata Amarilla flora; Iglesias et al., 2007), and overall floristic components (Kachaike flora;

Passalia, 2007a). It has been noted that these locations represent similar latitudes during the mid-

Cretaceous (Iglesias et al., 2007), and both of these regions were at substantially lower latitudes

than the Chatham Islands during the Cenomanian (Fig. 2). The climatic differences associated

with this latitudinal discrepancy would likely contribute to the unique floral ecology of the

Tupuangi Formation.

The floras of the Warder Formation, middle Clarence Valley, New Zealand, share a similar

palaeolatitude and stratigraphic range with the Tupuangi flora. Despite this, there appears to be

markedly lower fern abundance and diversity in the Tupuangi flora, the most notable absence is

the osmundaceous Phyllopteroides Medwell, 1954 emend. Cantrill and Webb, 1987. From the

Warder flora, Daniel (1989) identified thirty specimens of this genus, and classified them into

four distinct morphological groups. Furthermore, there have been three taxa of Cladophlebis

reported from the Ngaterian Clarence Valley strata (upper Albian to mid-Cenomanian;

McQueen, 1956; Parrish et al., 1998), whereas there were no recorded Cladophlebis from the

Ngaterian strata of the Tupuangi Formation, and only one species (C. auriculipilosus sp. nov.)

from Arowhanan strata (upper Cenomanian). This discrepancy in macrofloral assemblages is

likely due to local palaeoenvironmental influences, as explored in section 5.2 (below).

Because of the uncertainty over their affinities, the Tupuangi fern taxa have limited value for

interpreting the palaeobiogeography of, and ecological interrelationships between, fern groups.

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The fern taxon with arguably the greatest taxonomic certainty is Cladophlebis auriculipilosus,

because of various associations of members of this genus with Osmundaceae. Under the broad

diagnosis presented by Brongniart (1849), Cladophlebis has been a used for many extinct

species of sterile fronds with pecopteroid venation (see Dijkstra and Van Amerom, 1981, for a

review). Various species of Cladophlebis [particularly C. australis (Morris, 1845) Walkom,

1917 and C. denticulata (Brongniart, 1828) Fontaine, 1889 emend. Harris, 1961] were

particularly conservative and successful (Phipps et al., 1998). These have long been recognised

throughout Mesozoic strata across Eastern Gondwana, which includes Australia (Seward, 1904;

Herbst, 1978), Antarctica (Halle, 1913; Gee, 1989) and New Zealand (Arber, 1917; McQueen,

1956; Parrish et al., 1998). An affinity with Osmundaceae has been consistently proposed for

Cladophlebis, based on associations with osmundaceous stems (e.g. Kidston and Gwynne-

Vaughan, 1907; Medwell, 1954) and in situ spores (e.g. Litwin, 1985). In Australia, the relative

abundance of osmundaceous spores shows a progressive decrease during the Early Cretaceous

until the Cenomanian/Turonian, before their abundance stabilises for the most part throughout

the Late Cretaceous (Dettmann et al., 1992; Nagalingum et al., 2002). This relative decrease is

concurrent with an accelerating angiosperm diversification trend at southern high latitudes

(Drinnan and Crane, 1990; Cantrill and Poole, 2002). These data support the notion that the

Osmundales were significantly and adversely affected by Early to early Late Cretaceous

angiosperm diversification in the Southern Hemisphere (Nagalingum et al., 2002). Preliminary

macrofloral data appear to reflect this trend: Cladophlebis is typically the most abundant

component of high southern palaeolatitude macrofloral assemblages during the Early Cretaceous

(e.g. Herbst, 1971; Douglas, 1973; Gee, 1989), followed by a relative paucity of these fossils

during the mid- to Late Cretaceous. The presently recorded occurrence of only two specimens of

a single species of Cladophlebis and total absence of other osmundaceous ferns (e.g.

Phyllopteroides) from the Cenomanian strata of the Chatham Islands, amidst numerous

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specimens of Sphenopteris and Adiantites, lends support for the aforementioned floristic pattern.

However, this support should be considered provisional until a more robust floral

abundance/diversity data-set throughout the stratigraphic succession can be obtained.

Liverwort-rich floras have been recovered from various Cretaceous strata of Gondwana,

including Antarctica (Cantrill, 1997), Australia (Douglas, 1973), India (Banerji, 1989) and South

Africa (Anderson and Anderson, 1985). These commonly form mats of spreading thalloid forms

on regularly disturbed overbank beds of high-energy rivers (e.g. Falcon-Lang et al., 2001). In

contrast, the presently described non-vascular plants were found as solitary elements amongst

leaf litter beds, overlying intermittent deltaic flood deposits (Fig. 4A, B; see section 5.2). From

Alexander Island, Antarctica, specimens of Adiantites (Adiantites sp.) were recorded in

association with the liverwort Marchantites arcuatus Cantrill, 1997 in leaf litter beds

comparable to the Tupuangi flood deposits. Within this context, Cantrill (1997) interpreted

Adiantites as a herbaceous understorey component, and this interpretation is inferred for the

Adiantites taxon of the Tupuangi flora, due to the consistently abundant conifer component (C.

Mays, pers. obs., 2012), which likely formed the overstorey.

5.2 Palaeoenvironmental and taphonomic considerations

The Tupuangi Formation was deposited as part of a vast fluviodeltaic depositional system

along the southern margin of the Chatham Rise (Wood et al., 1989; Campbell et al., 1993)

during the Cenomanian–Turonian (Mildenhall, 1994; Mays and Stilwell, 2013). A wide range of

macrofloral remains, including conifers, ginkgoes and angiosperms (C. Mays and D.J. Cantrill,

pers. obs., 2012), in addition to the free-sporing plants recorded herein, were deposited in leaf

litter horizons. The fining -upward stacking patterns of fine-grained sediments at these localities

are typical of low-flow rate river/delta floodplains (Bridge, 2003). Preserved root systems

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indicate the formation of soils; the soil horizons are characterised by laminated, diverse leaf

beds, or thicker compacted carbonised plant beds. These suggest vegetation re-establishment on

the exposed floodplain deposits, with substantial peats forming in some cases. The massive to

cross-bedded sandstone facies immediately overlying these fossiliferous intervals typically lack

the concave-up geometry typical of meandering river channels, and are interpreted as being

deposited during intermittent fluviodeltaic sheet flow events (Bridge, 2003). Apart from sporadic

root traces, the palaeosols appear to have no discernible pedogenic structures, and sedimentary

depositional structures (e.g. planar bedding, ripples) are typically preserved within 10 cm below

the fossiliferous palaeosol surface. Due to their low chroma and high organic carbon and

sulphide content, these fossil soils are considered gleysols or histosols, which are typical of

regions with a consistently high water table (Mack et al., 1993; Retallack, 2001). The

palaeoenvironmental regime that this suggests for the examined fossil sites is: 1) a fluvial setting

with a sustained high precipitation rate; or 2) a coastal delta with a high groundwater base level.

The intermittent influxes of microplankton in the microfloral record (Mays and Stilwell, 2013),

and a lack of channel structures associated with the deposits, support the latter environmental

interpretation.

The Warder Formation, middle Clarence Valley, and the Tupuangi Formation, the Chatham

Islands, have similar ages and palaeolatitudes (see section 5.1). Furthermore, the Chatham Rise

has been interpreted as primarily emergent throughout the Cenomanian (Wood et al., 1989;

Laird and Bradshaw, 2004), thus no major geographic barriers are inferred between the

Tupuangi and Warder localities during the interval of preservation. Presumably, the

aforementioned factors would promote floral homogeneity between these localities; however,

the macrofloral assemblages appear to be starkly different. The discrepancy between the floras

of the Warder and Tupuangi formations may be explained by biases associated with the different

environments of deposition. The alluvial deposits of the Warder Formation have been interpreted

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as being deposited in a meandering river system with localised lacustrine and swampy settings

(Browne and Reay, 1993; Parrish et al., 1998), whereas the Tupuangi flora localities examined

herein likely represent paralic, deltaic settings (Campbell et al., 1993; Mays and Stilwell, 2013).

Leaf litter beds, like those typical of the Tupuangi and Warder floras, reveal only the floral

ecology proximal to the site of deposition (Burnham, 1989, 1994). As such, differences in

palaeoenvironment may introduce ecological biases by promoting preservation of riparian taxa

within the Warder flora, and flora adapted to marginal marine settings within the Tupuangi

Formation. Local ecological or taphonomic bias is also suggested by the palynological record.

Studies of spore/pollen dispersal in modern deltaic and fluvial systems (e.g. Müller, 1959;

Darrell, 1973; Chmura, 1994) have revealed that levee and overbank deposits can accumulate

large numbers of distally-sourced palynomorphs, and can thus be misleading when interpreting

the local floral ecology; however, deltaic spore/pollen diversity data from such sediments

provides a reliable approximation of the regional floral ecology (e.g. Moss et al., 2005). The

diverse spore assemblage of the Tupuangi Formation (Mildenhall, 1994; Mays and Stilwell,

2013; Mays, in review) belies the relative scarcity of free-sporing plants in the macrofloral

assemblage. However, most of the spore elements of the Tupuangi Formation are common to

coeval strata of mainland New Zealand, including those from middle Clarence Valley (Raine,

1984). This suggests there was a significant degree of ecological homogeneity across Zealandia

during the Cenomanian. As such, non-vascular plants, lycopods and ferns were much more

diverse in the Chatham Islands region than indicated by the locally-derived macrofloral record

represented by the presently reported fossil beds. The differences in fern diversity and

abundance between the Warder and Tupuangi assemblages is likely explained by biases caused

by local depositional settings, rather than climatic, evolutionary or geographic discrepancies.

6. Conclusions

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The Chatham Islands region during the mid-Cretaceous was geographically situated between

the Australian and the Antarctic/South American sectors of Gondwana on the Panthalassan

margin, and is arguably the highest southern latitude (~ 75–80ºS) floral assemblage of the

Cenomanian–Turonian, an interval of intense global greenhouse conditions. Within this context,

the Tupuangi flora represents a crucial biogeographic data-point between the well-described

coeval floras of Australia and South America/Antarctica. Two non-vascular plants, one lycopod

and three taxa of probable fern affinity are described, including two newly described species

(Cladophlebis auriculipilosus sp. nov. and Muscites gracilis sp. nov.), representing the first

macrofloral fossil record of these groups from the Chatham Islands, New Zealand.

The preserved sedimentary record of the fossiliferous beds reflects a deltaic floodplain

environment with a consistently high water table. Plant fossils were typically preserved as leaf

litter mats or thin coal beds immediately overlying gleyed palaeosols, which formed from

immature, waterlogged soils.

The Tupuangi flora fern diversity and abundance is markedly lower compared to various

coeval lower palaeolatitude localities of eastern and western Gondwana. Of particular note is the

relative scarcity of osmundalean ferns compared to Early Cretaceous localities of the high

southern latitudes. This provides provisional support for the decline of this group during the

mid-Cretaceous in the wake of the diversification of angiosperms across high southern latitudes.

Despite an apparent lack of geographic barriers during the Cenomanian, the Tupuangi flora has

yielded a starkly different assemblage, and is relatively depauperate in fern taxa, compared to the

coeval flora of the Warder Formation, mainland New Zealand. This is inferred to be due, at least

in part, to local ecological and/or taphonomic biases associated with differences in depositional

environments between the two localities.

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Acknowledgements

The authors would like to acknowledge the help of volunteer laboratory and field assistants:

Elyse Butterfield, Andrew Giles, Cameron McKenzie, David Pickering and Jesse Vitacca.

Special thanks go to chief preparator, Chava Rodriguez, who directed the efforts to ‘bring life

back’ to these fossils. We would also like to thank Assoc. Prof. Andrew Drinnan of the School

of Botany, University of Melbourne, for his assistance with laboratory facilities. CM would like

to thank the residents of Pitt Island, especially Mr John Preece, Mr Bill Gregory-Hunt and Mrs

Dianne Gregory-Hunt for allowing access to the outcrops on their land, as well as Mr Ken

Lanauze and Mrs Judy Lanauze for their assistance on Pitt Island. Terry and Donna Tuanui

kindly provided support on Chatham Island before and after fieldwork on Pitt Island. Fieldwork

and research supported by a Monash University Faculty of Science Bridging Research Grant and

an Australian Research Council Linkage Grant awarded to JDS. Further financial support was

provided by an Australian Postgraduate Award scholarship awarded to CM.

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Figure captions:

Figure 1: Geography of eastern Zealandia and the Chatham Islands. A) Map of modern eastern

Zealandia. Bold black outline = modern coastline; fine grey outline = 2000 m isobath. B) Map of

Chatham Islands. A detailed map of the studied fossil localities is provided in Figure 3. Adapted

from Campbell et al. (1993) with permission.

Figure 2: Palaeogeographic reconstruction of the Cenomanian (~ 95 Ma), south polar

perspective. Polar circle latitude is based on modern value (66° 33′ 44″S). Ant. = Antarctica;

Aus. = Australia; M. = Madagascar; S.A. = South America; Zeal. = Zealandia. Compiled from

Mukasa and Dalziel (2000), Schellart et al. (2006) and Veevers (2006). Emergent land areas are

modified from Markwick et al. (2000).

Figure 3: Geological map of Waihere Bay. This figure is the complementary inset map for

Figure 1. This map includes the fossil localities reported in this study.

Figure 4: Composite stratigraphic column of the Tupuangi Formation succession at Waihere

Bay, Pitt Island, with expanded stratigraphic logs of the coastal fossil localities reported in this

study. The stratigraphic column has been divided into lower (Fig. 4A) and upper (Fig. 4B) sub-

sections. Lithofacies are after the unit names of Hay et al. (1970), as discussed by Campbell et

al. (1993). New Zealand chronostratigraphic stages are defined by Crampton et al. (2004).

Stratigraphic columns and chronostratigraphic correlations are adapted from Mays and Stilwell

(2013). Ngat. = Ngaterian; Man. = Mangaotanean; ?Piri. = ?Piripauan; NZCS = New Zealand

Chronostratigraphic Stage; SST = sandstone.

Figure 5: Images of marchantiophyte, bryophyte and lycophyte specimens. A–C) Hepaticites

sp., PL1014. A) Microphotograph of specimen taken under ethanol, scale = 5 mm. B) Line

drawing of preserved thalli, dashed lines represent inferred margin, bold lines show where costae

are observed, shaded areas represent thalli that do not appear to be attached to the outlined

specimen, scale = 5 mm. C) Microphotograph taken under ethanol exhibiting notched apices

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(arrows), scale = 0.5 mm. D–G) Muscites gracilis Mays and Cantrill sp. nov. D–F, holotype,

PL1027, scales = 2 mm. D) Composite micrograph of medial portion of stem exhibiting complex

branching habit. E) Distal portion of stem. F) Line drawing of distal portion of stem (as

illustrated in Fig. 5E), featuring examples of both dorsiventrally (D) and laterally (L) flattened

leaves, white areas represent leaves including costae where observed, dashed lines represent

inferred margins/costae of missing or obscured parts, shaded area represents the stem. G) Distal

portion showing costae along basiscopic margin of laterally flattened leaves, PL1034, scale = 1

mm. H) Lycopodites sp., preserved medial portion, PL1002, scale = 1 mm.

Figure 6: Cladophlebis auriculipilosus Mays and Tosolini sp. nov. holotype. A, B)

Compression, PL1036, scales = 10 mm. A) Possible basal portion of frond exhibiting alternating

organisation of pinnae. B) Venation detail under ethanol. C–E) Counterpart impression of

specimen illustrated in Fig. 6A, B, PL1035, scales = 2.5 mm. C) Microphotograph featuring the

elongated trichome bases on rachis and rachilla. D) Medial portion of pinna impression with

pinnule details, photograph taken with illumination at a low angle of incidence (from top of

image) to emphasise venation and curved lamina surface, arrows indicate probable revolute

sinuses on the apical sides of the auricles. Note that this impression represents the abaxial

surface of the pinna. E) Line drawing of pinna illustrated in Fig. 6D featuring pinnule basal

relationships including a tendency for the auricles to underlap the decurrent basiscopic margins

of the acropetal pinnules, dashed lines represent complementary details from compression,

shaded area represents the rachilla.

Figure 7: Artist’s reconstruction of the adaxial frond surface of Cladophlebis auriculipilosus

Mays and Tosolini sp. nov. Scale = 20 mm. Artist: Elyse Butterworth.

Figure 8: Sphenopteris sp. cf. S. warragulensis. A, B) Impression of degraded partial frond,

illustrating pinna arrangement, PL1015, scales = 1 mm. A) Photograph taken with low angle

light to accentuate venation details. B) Line drawing, shaded lobe is not attached to unshaded

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frond. C, D) Partial pinna, PL1024, scales = 5 mm. C) Composite microphotograph, specimen

has been unevenly oxidised with only partial compression preserved. D) Line drawing showing

venation pattern and variably lobulate to serrate distal margins, shaded lobe may not have been

attached to non-shaded pinna. E, F) Impression of partial lobe, PL1019, scales = 2.5 mm. E)

Composite microphotograph. F. Line drawing.

Figure 9: Adiantites sp. A, B) Isolated (?terminal) pinna impression, PL1007, scales = 5 mm. A)

Microphotograph of specimen under ethanol. B) Line drawing, shaded areas indicate interlobate

lamina areas extending to sinuses, partitioning seven separate lobes. C, D) Isolated (?terminal)

pinna with preserved serrate margin details, PL1025, scales = 5 mm. C) Photograph. D) Line

drawing, AA = axis of alternation. E, F) Impression fossil of two leaves attached to rachis,

PL1005, scales = 5 mm. E) Photograph. F) Line drawing. G) Medial portion of rachis showing

arrangement of pinnae, PL1000, scale = 10 mm.

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Figure 1

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Figure 2

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Figure 3

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Figure 4 lower

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Figure 4 upper

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Figure 5

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Figure 6

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Figure 7

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Figure 8

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Figure 9

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Table 2. List of Cladophlebis species recorded from probable Cretaceous strata of New Zealand. Locations for each species are listed in chronostratigraphic order; * = descriptive terms that have been substituted (as per Harris and Woolf-Harris, 2001, and Ellis et al., 2009) to allow for a standardised comparison; † = details not provided in original description, but inferred from illustrated specimen; 1 = Ettingshausen, 1890; 2

= Arber, 1917; 3 = McQueen, 1955; 4 = McQueen, 1956; 5 = Raine, 1981; 6 = Browne and Reay, 1993; 7 = Parrish et al., 1998; 8 = Raine in Kennedy, 2003; 9 = Mays and Stilwell, 2013.

Species Location Age Pinnule

arrangement

Pinnule

length

(mm)

Pinnule

width

(mm)

Pinnule

shape

Pinnule

apex

Pinnule

margin

Pinnule

attachment Comments

Cladophlebis sp. cf.

C. albertsi (Dunker

1846) Seward 1894

Waikato Heads

(2); Clarence

Valley (7)

"Neocomian" (4);

Cenomanian (6) sub-opposite ≤ 7 ≤ 4 falcate acuminate entire †

sessile,

adnate *

Cladophlebis

australis (Morris

1845) Walkom 1917

Waikato Heads

(2); Clarence

Valley (7);

Pakawau (2)

"Neocomian" (4);

Cenomanian (6);

Campanian/

Maastrichtian (3, 8)

not specified 7-33 3.5-8 lanceolate

to falcate

acute to

obtuse entire

sessile,

adnate *

fertile material

preserved

Cladophlebis

obscura

(Ettingshausen 1887)

McQueen 1956

Pakawau (1, 4) Campanian/

Maastrichtian (8)

alternate to

sub-opposite 25-30 5-6

lanceolate

* obtuse entire

sessile,

adnate *

Cladophlebis prisca

(Ettingshausen 1887)

McQueen 1956

Clarence Valley

(7); Pakawau

(1, 4)

Cenomanian (6);

Campanian/

Maastrichtian (3, 8)

alternate to

sub-opposite 25 6-7

lanceolate

*

acute to

obtuse entire

sessile,

adnate *;

acute insertion

Cladophlebis

wellmanii McQueen

1956

Paparoa Coal

Measures (4)

(?)Campanian/

Maastrichtian (5) N/A N/A N/A N/A N/A N/A N/A not pinnulate

Cladophlebis sp. Waikato Heads

(2) "Neocomian" (4)

?alternate to

sub-opposite † ≤ 9.5 † ≤ 6 †

lanceolate

†

?rounded

to obtuse †

?entire

†

sessile,

adnate †

single poorly

preserved

specimen

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Cladophlebis

auriculipilosus Mays

and Tosolini sp. nov.

Pitt Island Cenomanian (9) sub-opposite ≤ 7.5 ≤ 4 ovate rounded entire sessile, partly

adnate

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Graphical abstract

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Late Cretaceous (Cenomanian–Turonian) macroflora from the

Chatham Islands, New Zealand: bryophytes, lycophytes and

pteridophytes

Chris Mays, Anne-Marie P. Tosolini, David J. Cantrill and Jeffrey D. Stilwell

Research highlights

The first record of Cretaceous non-seed macroflora from the Chatham Islands

Two new species described: Cladophlebis auriculipilosus and Muscites gracilis

Fossil fern content provisionally supports regional floral evolutionary trends