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Botanical Journal of the Linnean Society, 2015, 178, 421–440. With 4 figures
The monocot fossil pollen record of New Zealand and
its implications for palaeoclimates and environments
JOHN G. CONRAN1*, DALLAS C. MILDENHALL2, J. IAN RAINE2,
ELIZABETH M. KENNEDY2 and DAPHNE E. LEE3
1
Australian Centre for Evolutionary Biology & Biodiversity and Sprigg Geobiology Centre, School of
Biological Sciences, The University of Adelaide, Benham Building DX 650 312, Adelaide, SA 5005,
Australia
2
GNS Science, PO Box 30-368, Lower Hutt, New Zealand
3
Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand
Received 23 March 2014; revised 15 January 2015; accepted for publication 23 March 2015
The record of fossil monocotyledonous pollen in New Zealand is summarized and discussed in terms of age and
palaeoecological implications. The fossil monocot pollen record covers reports of taxa variously ranging from
the Palaeogene to the present, with diverse groups represented, including Alstroemeriaceae: Luzuriagoideae
(Luzuriaga), Amaryllidaceae (aff. Hymenocallis?), Arecaceae (including Nypa and aff. Metroxylon), Asparagaceae:
Lomandroideae (Arthropodium, Cordyline), Asteliaceae (Astelia and Collospermum), Cyperaceae, Anarthriaceae
(including Hopkinsiaceae and Lyginiaceae) or Flagellariaceae?, Orchidaceae, Pandanaceae, Poaceae, Restionaceae
(aff. Empodisma), Ripogonaceae (Ripogonum), Typhaceae (Typha and Sparganium) and Asphodelaceae (= Xanthorrhoeaceae): Hemerocallidoideae (Dianella and Phormium). There are also monosulcate and reticulate lilioid
and echinate and gemmate arecoid palynomorphs known from the Late Cretaceous. The possible biogeographical
and palaeoecological implications of these records, especially for tropical or subtropical taxa occurring at mid to
high southern latitudes, are also discussed. © 2015 The Linnean Society of London, Botanical Journal of the
Linnean Society, 2015, 178, 421–440.
Additional Keywords: Cretaceous – Eocene – Miocene – palaeoecology – palaeoenvironment – palynology.
INTRODUCTION
Fossil plants were recognized from New Zealand well
over 200 years ago when kauri gum was found in peat
lands in northern North Island, but spores and pollen
were not investigated until the early part of the 20th
century. Early investigations on Pleistocene peats
showed the presence of grasslands, sedge swamps and
other palaeoenvironments dominated by monocot
pollen (e.g. Erdtman, 1924; Cranwell & von Post,
1936). A decade later, the first detailed papers
appeared on fossilized extinct spores and pollen (e.g. Te
Punga, 1949), followed shortly by the seminal papers
and bulletins of Couper (1953, 1954, 1960). These
articles described the first set of extinct monocot pollen
*Corresponding author. E-mail: [email protected]
morphotypes from New Zealand and set the scene for
the numerous studies that followed.
By 2000, some 33 monocot pollen taxa had been
described or identified, and many more have been
identified since (Mildenhall & Raine, 2012). The preQuaternary fossil monocot pollen record in New
Zealand is one of the most complete in the world, and
covers reports of taxa variously ranging from the Late
Cretaceous and Palaeogene to the present (Raine,
Mildenhall & Kennedy, 2011). Diverse groups are
represented, including taxa that are now placed in the
orders Arecales, Asparagales, Liliales, Pandanales and
Poales (sensu APG III, 2009, Chase & Reveal, 2009).
Many of the extinct monocot pollen taxa described
have no precise botanical affinity and so, at this stage,
are not definitive enough for the assessment of evolution and palaeoenvironment; however, they are useful
for discussing diversity, particularly in comparison
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J. CONRAN ET AL.
with the present day. However, care should be taken
with dispersed pollen, as an individual pollen taxon
often represents more than one plant species: a range
of slightly differing morphological types is often
assigned the same form-specific name. Conversely, the
perceived range of morphological variation may be
such that more than one fossil pollen species is instituted for pollen from one plant species, both through
time and at any one fossil locality. In general, however,
diversity is underestimated.
Until the Late Cretaceous (c. 83 Mya), the continent of Zealandia (the area currently defined by the
continental shelf, including New Zealand and New
Caledonia) was a region of the eastern Gondwana
supercontinent, called Tasmantis, which shared a
fossil flora and fauna with Australia (Gibbs, 2006).
Following gradual opening of the Tasman Sea, which
began in the south at about 83 Mya, Zealandia drifted
slowly east and north, reaching its current latitudinal
position at about 40 Mya, although direct land connections between Australia and Zealandia were
severed completely by 65 Mya (Sutherland & King,
2008). The current isolated position of New Zealand,
2000 km east of Australia, has been maintained since
30 Mya, although, at times, there have been intermittent connections to the north-west via New Caledonia
and various volcanic island chains (Lee, Lee &
Mortimer, 2001; Ladiges & Cantrill, 2007).
After initial rifting, the Zealandia continent began
to subside gradually as a result of crustal thinning
and, at c. 40 Mya, was a low-lying landmass, about
three times the size of present-day New Zealand. By
the late Oligocene (c. 25 Mya), the land area was
reduced to a series of large low-lying islands collectively occupying an area of between 50 000 and
20 000 km2 (similar to the current land areas of Tasmania and New Caledonia, respectively). At around
25 Mya, uplift and a concomitant increase in land
area began as a result of the development of a new
plate boundary between the Australian and Pacific
plates: land area increased to about its present size
by the early Miocene. Today, the three large and
hundreds of smaller islands that make up modern
New Zealand comprise only about 10% of the original
land area of Zealandia (Graham, 2008). During
50–10 Mya, the climate of New Zealand was at least
marginally subtropical, compared with the warm to
cool temperate climates in the region today.
Apparent New Zealand monocot diversity increased
with time, with about 17 pollen taxa described from the
Cretaceous, through to almost 100 taxa described from
the Quaternary. For comparison, the modern New
Zealand flora comprises 125 monocot genera in 24
families: these include 36 orchid and 33 grass genera
which have limited fossil records in New Zealand and
elsewhere. The increase in diversity is undoubtedly a
result of several factors, including the development
and expansion of new ecological niches through tectonic activity, which commenced in the early Miocene
and intensified in the late Miocene, and a bias in
research effort towards detailed investigations of Quaternary sediments. The fragmentation of habitats
through orogenic island building led to the expansion
of new ecological palaeoenvironments from the
Miocene to the Pleistocene. This resulted in increased
allopatry, leading to rapid evolution and a high percentage of endemism in some families. Examples are
Cyperaceae and Poaceae, the pollen of which is rare or
absent in sediments older than the latest Miocene and
can, in most cases, only be ascribed to family level.
Diversity and problems besetting the identification
of the taxonomic affinity of fossil pollen have been
discussed briefly by Mildenhall & Raine (2012). They
showed, with several provisos, that pollen from taxon
endemism in the New Zealand flora increased
through the Late Cretaceous and Cenozoic to the
present day. For the Pleistocene, some 50% of pollen
was from endemic plants (but many with close relatives in Australia and South America), whereas, for
the Neogene, it was 45%, the Palaeogene about 30%
and the Late Cretaceous about 12%.
Here, we concentrate on the more important fossil
pollen taxa and describe their general affinities, age
range and palaeoenvironmental significance as an
illustration of the range of fossil monocot pollen
present in New Zealand Cenozoic sediments (Appendix). Data on the numbers of sites in which palynomorphs have been reported are also included. Further
information, including additional references not cited
below, can be obtained from Raine et al. (2011) and
FRED: The New Zealand Fossil Record Electronic
Database (GNS Science, 2014).
MATERIAL AND METHODS
Data were collated from the thousands of monocot
records in Raine et al. (2011) and FRED (GNS Science,
2014), covering the whole of New Zealand, for fossil
sites and mining drill cores (including near offshore)
and samples extracted from the Foulden Maar,
Newvale Mine and Pikopiko fossil deposits discussed
in Conran et al. (2015, this issue). These data were
then collated taxonomically by pollen morphotype and
temporally in order to determine apparent palynomorph diversity and survivorship through time.
RESULTS
At least 130 monocot palynomorphs from more than
20 families have been recognized (Table 1) from New
Zealand, ranging from the Cretaceous to the present
(Appendix). Although many of these represent Qua-
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NEW ZEALAND FOSSIL MONOCOT POLLEN
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Table 1. Summary distribution of New Zealand fossil monocot palynomorphs assignable to modern taxa through
geological time. Classification follows APG III (2009) and Briggs et al. (2014). Details of pollen species-level affinity, age
range and literature are listed in the Appendix
Family
Alstroemeriaceae: Luzuriagoideae
Amaryllidaceae?
Araceae
Arecaceae
Asparagales/Liliales
Asparagaceae: Lomandroideae
Asteliaceae
Cyperaceae
Flagellariaceae/Anarthriaceae
Iridaceae
Juncaceae
Juncaginaceae
Orchidaceae
Pandanaceae
Poaceae
Potamogetonaceae
Restionaceae: Centrolepidoideae
Restionaceae: Leptocaryoideae
Ripogonaceae
Ruppiaceae
Typhaceae
Xanthorrhoeaceae (now Asphodelaceae)
Late
Cretaceous
X
X
ternary records of modern genera or species, there are
still numerous pre-Quaternary taxa reported, indicating that monocots have been a prominent component
of the New Zealand biota since the beginning of the
separation of Zealandia from Tasmantis at 100–
80 Mya (Gibbs, 2006; Sutherland & King, 2008).
When these data are summarized by pollen morphology (Fig. 1A) and time period (Fig. 1B, C), they
show that nearly half are palynomorphs with monosulcate pollen grains (e.g. Fig. 2H), but aletes
(Fig. 2D), trichotomosulcates (Fig. 2Q), disulcatesdicolpates (Fig. 2G), monoporates (Fig. 2I) and polyporates (Fig. 2F) are also present (Fig. 1A). Although
the majority of taxa first appear in the Quaternary
(75 taxa), there were nevertheless 17 monocot palynomorphs present for the first time in the Late Cretaceous, 22 in the Palaeogene and a further 19 in the
Neogene (Fig. 1B).
Comparisons of pollen taxa through time (Fig. 1C)
also show that nearly all of the Cretaceous forms
seem to disappear at the K/Pg boundary, whereas
over half of the Palaeogene taxa persist into the
Neogene (as do two Cretaceous palynomorphs). Nevertheless, the majority of all of these older palynomorphs became at least locally extinct by the
Quaternary, where the pollen signal is dominated by
Palaeogene
Neogene
Quaternary
Holocene
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
modern taxa that are now prominent components of
the modern New Zealand flora (Appendix).
SUMMARY
REVIEW OF SELECTED PRE-QUATERNARY
FOSSIL MONOCOT POLLEN TAXA FROM
(ORDINAL
NEW ZEALAND
AND FAMILY CLASSIFICATION AND TAXON
APG III, 2009 AND BRIGGS,
MARCHANT & PERKINS, 2014)
ORDER FOLLOW
PANDANALES
Lateropora glabra Pocknall & Mildenhall (1984)
Botanical affinity is with kiekie, Freycinetia Gaud.
New Zealand age range is late Palaeogene and
younger.
The monoporate pollen L. glabra (Fig. 2I, J) is
clearly derived from the genus Freycinetia which,
today, grows under warm to cool temperate conditions
in rainforest and coastal scrub throughout New
Zealand (Moore & Edgar, 1970).
Dryptopollenites semilunatus Stover in Stover &
Partridge (1973)
Botanical affinity is with Pandanaceae.
New Zealand age range is Palaeogene.
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J. CONRAN ET AL.
LILIALES
ALSTROEMERIACEAE: LUZURIAGOIDEAE
Liliacidites contortus Mildenhall & Bannister in
Conran et al. (2014a)
Botanical affinity is with Luzuriaga Ruiz & Pav.
New Zealand age range is Neogene and younger.
Monosulcate, reticulate pollen grains with very thin
exines (Fig. 2K) from early Miocene fossil anthers of
Luzuriaga have recently been described (Conran
et al., (2014a). Liliacidites contortus has only been
identified at this one locality, but a few previous
records of Luzuriaga-like pollen from Neogene sediments have yet to be re-examined. Pollen of Luzuriaga parviflora Kunth has been recorded from a
number of Quaternary localities, but the very thin
exine and contorted nature of dispersed Luzuriaga
pollen has undoubtedly limited its fossil record.
Modern Luzuriaga grows in mossy forest floors from
sea level in the south to mountain ranges in the north
(Moore & Edgar, 1970) and, as such, is more indicative of cool temperate than warmer environments.
RIPOGONACEAE (NEOGENE
ONWARDS BUT OLDER
MACROFOSSILS ARE KNOWN)
Figure 1. Summary statistics for New Zealand fossil
monocot pollen diversity. A, Pie chart showing palynomorph diversity by morphotype. B, Pie chart showing
palynomorph diversity by geological epoch. C, Stacked bar
graphs showing monocot palynomorph diversity and relative persistence through time (epoch colours as in B).
This pollen type is a dyad with monosulcate
monads (Fig. 2H) and has been related to Pandanaceae, but not to any modern genus; its palaeoenvironmental significance is not known.
Ripogonum scandens J.R.Forst. & G.Forst. (1776)
New Zealand age range is Neogene and younger.
Microreticulate, monosulcate pollen grains, attributed to the modern taxon R. scandens (supplejack),
have been identified rarely from late Miocene and
younger sediments, but identifications older than late
Miocene need to be treated with caution. Ripogonum
scandens grows in lowland podocarp–broad-leaved
forest throughout New Zealand (Moore & Edgar,
1970) under cool to warm temperate conditions. Nevertheless, there is a range of macrofossil taxa of
Ripogonum J.R.Forst. & G.Forst. known from Eocene
and particularly late Oligocene and early Miocene
sites in New Zealand (Pole, 1993; Conran, Bannister
& Lee, 2009b), and Eocene sites in Australia (Conran,
Carpenter & Jordan, 2009) and South America (Carpenter et al., 2014), showing that the genus has had a
long history in the region.
OTHER LILIACIDITES
SPP.
Traditionally, Liliacidites has been the palynomorph
genus used for ‘Liliaceae’-like fossil pollen grains (e.g.
Raine et al., 2011) (Fig. 2K–O). However, Liliaceae s.l.
(e.g. sensu Krause, 1930; Cronquist, 1981) has been
shown through numerous molecular and other studies
to represent multiple phylogenetic lineages and families now placed in the orders Asparagales and Liliales
(APG III, 2009, Chase & Reveal, 2009; Chase, Reveal
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NEW ZEALAND FOSSIL MONOCOT POLLEN
Figure 2. See caption on next page.
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Figure 2. Examples of monocot palynomorphs described from New Zealand (see Appendix). A, Arecipites cf. Rhopalostylis. B, Arecipites otagoensis. C, Arecipites subverrucatus. D, Assamiapollenites inanis. E, Assamiapollenites incognitus.
F, Cyperaceaepollis neogenicus. G, Dicolpopollis metroxylonoides. H, Dryptopollenites semilunatus. I, Lateropora glabra.
J, Lateropora glabra. K, Liliacidites contortus. L, Liliacidites intermedius. M, Liliacidites kaitangataensis. N, Liliacidites
perforatus. O, Liliacidites variegatus. P, Luminidites phormoides. Q, Luminidites reticulatus. R, Milfordia homeopunctata.
S, Milfordia hypolaenoides. T, Monogemmites gemmatus. U–W, Nupharipollis mortonensis. X, Palmidites maximus. Y,
Sparganiaceaepollenites barungensis. Z–AA, Sparganiaceaepollenites sphericus. AB, Spinizonocolpites prominatus. Photographs, GNS Science; drawings A, L, X from plate 7, Couper (1953).
& Fay, 2009); therefore, assignment to a broad ‘Liliaceae’ should be replaced instead with Asparagales/
Liliales.
other possible Asparagales which this taxon may represent (A. Meerow, USDA, Miami, FL, pers. comm.)
Liliacidites variegatus Couper (1953) (Fig. 2O)
Botanical affinity is with Asparagales/Liliales or, possibly, Atherospermataceae, cf. Laurelia novaezelandiae A.Cunn.
New Zealand age range is Late Cretaceous to
Neogene.
Liliacidites variegatus is one of a number of fossil
reticulate, monosulcate pollen grains placed in the
form genus Liliacidites Couper. Although similar to
the modern early-branching angiosperm Laurelia
novaezelandiae (pukatea), common in forest swamp
environments, it is more likely to be a fossil
Asparagales/Liliales pollen type. These pollen types
occur in most Late Cretaceous to Neogene samples,
but their palaeoenvironmental significance is not
known. They can occur in significant numbers in
Miocene samples.
The Late Cretaceous palynomorph L. kaitangataensis Couper (Fig. 2M) has been compared with pollen
from Lilium L., but that affinity is unlikely. The
Palaeogene species L. perforatus Pocknall (Fig. 2N)
has been compared with pollen from the Asparagales/
Liliales, particularly Amaryllidaceae, but also Arecaceae (Pocknall, 1982).
ASPARAGACEAE: LOMANDROIDEAE
ASPARAGALES
AMARYLLIDACEAE?
Monogemmites gemmatus (Couper, 1960) Krutzsch
1970
Botanical affinity is possibly with Hymenocallis declinata Salisb., as described by Erdtman (1952: 45).
New Zealand age range is late Palaeogene to
Neogene.
Hymenocallis is a genus of American open subtropical or tropical environments, including wetlands.
Monogemmites gemmatus (Fig. 2T) is a gemmate,
monosulcate pollen type that is clearly associated
with lowland rainforest vegetation in the New
Zealand Cenozoic. However, this placement in Amaryllidaceae is not considered definitive, and there are
Liliacidites intermedius Couper (1953)
Botanical affinity is with Arthropodium R.Br.
New Zealand age range is Cretaceous to Neogene.
This palynomorph (Fig. 2L) has been compared
with pollen of the rengarenga lily (Arthropodium), a
widespread and common plant of shady lowland
forest floors and coastal situations in modern New
Zealand (Moore & Edgar, 1970).
ASTELIACEAE
Liliacidites sp. cf. Collospermum Skottsb.
Botanical affinity is with Collospermum.
New Zealand age range is Palaeogene and younger.
A reticulate, monosulcate grain, similar to the
pollen of Collospermum, has been recorded from late
Oligocene sediments in which it was preserved in a
subtropical setting (Mildenhall et al., 2014).
There are two modern Collospermum spp., growing
primarily as epiphytes in mainly warm temperate,
coastal lowland forest environments north of about
42°S (Allan Herbarium, 2000). Most identifications of
Collospermum pollen are from Pleistocene sediments
deposited in lowland settings.
Monogemmites sp. Krutzsch (1970)
Botanical affinity is with Astelia Banks & Sol. ex
R.Br.
New Zealand age range is Palaeogene and younger.
Krutzsch (1970) gave the form name Monogemmites
to a fossil Astelia-like pollen grain illustrated by
Couper (1960). Small, monosulcate, apiculate pollen
grains attributed to the modern genus Astelia have
been recorded from late Eocene sediments (Couper,
1960), but are more frequent in latest Miocene and
younger sediments. Maciunas et al. (2011) described
Astelia macrofossils from a late Oligocene–early
Miocene, 23-Mya deposit, the first definitive Astelia
macrofossils identified, although Pole (2007) reported
Miocene-aged dispersed cuticle from southern South
Island drill cores. Astelia is a widespread Southern
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NEW ZEALAND FOSSIL MONOCOT POLLEN
Hemisphere epiphytic and ground-dwelling genus
(Rudall et al., 1998) and, in New Zealand today, may
be found growing in lowland coastal rainforest and
alpine areas, especially in bogs (Moore & Edgar,
1970). Therefore, Astelia pollen can be expected to
occur in sediments from a wide range of environments. Nevertheless, the Astelia/Collospermum
lineage is thought to have radiated near the Eocene–
Oligocene boundary, and Birch, Keeley & Morden
(2012) suggested that current distributions might, in
part, result from long-distance dispersal; with the
modern alpine New Zealand taxa in the complex
evolving in the late Miocene to Pliocene.
ORCHIDACEAE
Fossil pollinia and tetrads attributed to Orchidaceae
are rare and have only been positively identified in
occasional Quaternary samples. In contrast, macrofossils of Dendrobium Sw. and Earina Lindl. are
known from the early Miocene Foulden Maar
(Conran, Bannister & Lee, 2009a), demonstrating a
much longer presence in New Zealand for the family
than the pollen records currently suggest.
XANTHORRHOEACEAE (NOW ASPHODELACEAE;
APPLEQUIST, 2014): HEMEROCALLIDOIDEAE
Luminidites phormoides (Stover & Partridge, 1982)
Pocknall & Mildenhall (1984) (Fig. 2P)
Botanical affinity is with the genus Phormium
J.R.Forst. & G.Forst and possibly other Hemerocallidoideae, including Pasithea D.Don (Macphail &
Cantrill, 2006).
New Zealand age range is Neogene (one possible
Palaeogene record).
Luminidites phormoides pollen is also known from
the Palaeogene of Australia (Macphail, 1999) and
Miocene of Heard Island (Macphail & Cantrill, 2006).
However, the affinities of this palynomorph require
detailed comparison with other hemerocallid genera
in the phormioid and johnsonioid clades of Wurdack &
Dorr (2009), nearly all of which bear pollen superficially similar to this palynomorph (Furness et al.,
2014).
Luminidites reticulatus (Couper, 1960) Pocknall &
Mildenhall (1984) (Fig. 2Q)
Botanical affinity is possibly with Phormium and certainly with Hemerocallidoideae.
New Zealand age range is Palaeogene to Neogene.
Two Phormium spp. occur in New Zealand today,
but both produce pollen that differs from the two
trichotomosulcate fossil species (Luminidites phormoides and L. reticulatus). One species grows mainly
in lowland swamps and the other is less common and
427
grows on coastal cliffs and mountain slopes (Moore &
Edgar, 1970). If the fossil taxa occurred in the same
environment, they represent a swampland or periodically flooded lowland environment in warm to cool
temperate conditions. Luminidites reticulatus pollen
is never abundant, but is often found in Palaeogene
and Neogene sedimentary sequences. If L. phormoides, a much rarer pollen type than L. reticulatus,
is related to Pasithea, this pollen type might indicate
drier, open, sunny environments. A pollen grain, identified from late Eocene to late Oligocene sediments as
Phormium and Luminidites sp. and similar, but not
identical to, the present-day lowland Phormium tenax
J.R.Forst. & G.Forst. was illustrated by Pocknall
(1985). The climate at the time was regarded as cool
to warm temperate and humid. However, not necessarily all records of this palynomorph represent Phormium and some may be from Pasithea, Dianella or
other members of the phormioid clade (Furness et al.,
2014), and require further investigation.
ARECALES
Palms have a long fossil record going back well into
the Cretaceous, with palm-like pollen well known
from a wide range of sites around the world (e.g.
Harley & Baker, 2001; Pan et al., 2006; Dransfield
et al., 2008). The family appears to have had
maximum diversity in the Palaeogene, when many
groups were much more widely distributed than in
the present day (Baker & Couvreur, 2013a, b).
Similarly, although there is only a single living
palm in New Zealand, the nı̄kau palm (Rhopalostylis
sapida H.Wendl. & Drude) from the warmer areas of
both main islands, the fossil record is much more
diverse, both for macrofossils (Conran et al., 2015,
this issue) and, particularly, for palm-associated
microfossils (Raine et al., 2011; Mildenhall & Raine,
2012; GNS Science, 2014).
There are pollen records of Arecipites Wodehouse,
considered to be similar to nı̄kau, from sites in the
Palaeogene to Neogene, but, in addition, there are at
least six other Arecipites (Fig. 2A–C), one Cocos L.?,
two calamoid Dicolpopollis Pflanzl, one Palmidites
Couper and three nypoid Spinizonocolpites J.Muller
palynomorphs (Appendix; Raine et al., 2011; J. I.
Raine pers. comm.).
Arecipites otagoensis (Couper, 1960) Mildenhall &
Pocknall (1989)
Botanical affinity is with Arecaceae.
New Zealand age range is Palaeogene to Neogene.
Arecipites otagoensis (Fig. 2B) is one of a number of
monosulcate pollen types with similar smooth or relatively smooth exines identified from New Zealand
Late Cretaceous and Cenozoic deposits. Arecipites
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J. CONRAN ET AL.
spp. can be the dominant pollen type, especially in
Miocene samples where there is evidence of rapid
climatic changes, particularly in the thick coalbearing, lacustrine sediments of central Otago
(Mildenhall & Pocknall, 1989). Their palaeoenvironmental significance is uncertain, but most palms
today live in warm temperate to tropical environments. One exception to this is Rhopalostylis sapida,
which extends to the temperate coastal forests of
north-western South Island.
Dicolpopollis sp. cf. D. metroxylonoides Khan (1976)
Botanical affinity is with the sago palm Metroxylon
Rottb. or the rattan cane Calamus L.
New Zealand age range is Neogene.
This dicolpate pollen type (Fig. 2G), illustrated
from New Zealand by Mildenhall (1989) and
Mildenhall & Pocknall (1989), was compared with
Metroxylon amicarum (Wendl.) Becc. by Khan (1976)
and with Calamus by Truswell, Sluiter & Harris
(1985). Calamus and Metroxylon are both tropical to
subtropical genera, and may indicate similar conditions in New Zealand where this dicolpate pollen type
is rarely found.
Spinizonocolpites prominatus (McIntyre, 1965)
Stover & Evans (1973)
Botanical affinity is with the nipa palm, Nypa fruticans Wurmb. (Kumaran, Punekar & Limaye, 2011).
New Zealand age range is Palaeogene.
This spiny, monosulcate pollen type (Fig. 2A,B) was
regarded as a fossil Nypa Wurmb. by Kumaran et al.
(2011) and is largely restricted to Eocene sites, where
it occurs sporadically in samples from coastal or nearcoastal palaeoenvironments, although Pocknall (1989)
recorded it from marginal marine sediments of early
Oligocene age. The lack of Nypa pollen and pollen
from later mangrove communities suggests that the
Miocene New Zealand climate was never truly tropical, although Conran et al. (2014b) reported a possible
mangrove community in apparently subtropical late
Oligocene sediments from Cosy Dell in southern New
Zealand, based on the presence of warm-water mangrove snails.
POALES
RESTIONACEAE S.L. (INCLUDING
CENTROLEPIDACEAE), ANARTHRIACEAE (INCLUDING
HOPKINSIACEAE AND LYGINIACEAE) OR, POSSIBLY,
FLAGELLARIACEAE
Milfordia Erdtman emend. Stover & A.D.Partr. is a
widespread palynomorph which is known from the
Late Cretaceous of the Northern Hemisphere, where
it was taken traditionally as evidence of Restionaceae. However, Linder, Briggs & Johnson (1998)
noted that this palynomorph lacked diagnostic features for Restionaceae and might also represent
Flagellariaceae, Joinvilleaceae or even Poaceae. Similarly, although Cenozoic Southern Hemisphere Milfordia affinities to Restionaceae were accepted by
Linder, Eldenas & Briggs (2003), Smith et al. (2010)
noted that detailed ultrastructural study of Milfordia
and related palynomorphs is needed to better assess
their affinities.
Milfordia homeopunctata (McIntyre, 1965) Partridge
in Stover & Partridge (1973)
Botanical affinity is with the restiids: Restionaceae
s.l. or Anarthriaceae (including Lyginiaceae and Hopkinsiaceae) (Briggs et al., 2014), especially Lyginia
R.Br. (Macphail, 1999) and Hopkinsia W.Fitzg.
(Macphail & Stone, 2004). However, this pollen type
has also been related to Flagellariaceae (Macphail &
Stone, 2004).
New Zealand age range is Palaeogene to Neogene.
This punctate, monoporate pollen type (Fig. 2R)
occurs in late Palaeogene and Neogene sediments
becoming extinct at the end of the Miocene. Lyginia
grows in dry sandy areas in south-western Australia
and Hopkinsia grows in similar environments, but
more usually along watercourses. Neither genus
occurs naturally in New Zealand. Although the uncertainty of the botanical affinity of M. homeopunctata
negates its usefulness as an environmental indicator,
it mainly occurs in samples representing wetland and
coastal lagoon environments from cool temperate to
warm temperate or subtropical climatic conditions. It
has also been recorded in some numbers in a late
Oligocene site (25–24 Mya) interpreted as a subtropical, coastal, swampy estuarine palaeoenvironment
(Conran et al., 2014b). Milfordia homeopunctata is
not comparable with any present-day New Zealand
pollen type.
Milfordia hypolaenoides Erdtman (1960) (Fig. 2S)
Botanical affinity is with Restionaceae s.l., especially
Empodisma minus (J.D.Hook.) L.A.S.Johnson &
D.F.Cutler and related taxa.
New Zealand age range is Neogene and younger.
Empodisma minus [= Calorophus minor J.D.Hook.
and Hypolaena lateriflora (R.Br.) Benth.] grows in
lowland swamps and mountain bogs in New Zealand
(Moore & Edgar, 1970) and similar habitats in southeastern Australia (Meney & Pate, 1999), particularly
in cool, moist, often elevated environments (Wagstaff
& Clarkson, 2012). Pollen of this type first appears in
the Miocene, but is only common in the Pleistocene.
Similarly, using fossil-calibrated molecular analyses,
Wagstaff & Clarkson (2012) dated the divergence of
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NEW ZEALAND FOSSIL MONOCOT POLLEN
Empodisma L.A.S.Johnson & D.F.Cutler from its
sister genus Winifredia L.A.S.Johnson & B.G.Briggs
to the mid-Oligocene or early Miocene (28–16 Mya).
TYPHACEAE (INCLUDING SPARGANIACEAE)
Sparganiaceaepollenites barungensis Harris (1972)
Sparganiaceaepollenites sphericus (Couper, 1960)
Mildenhall in Mildenhall & Crosbie (1979)
Botanical affinities are with Sparganium L.
New Zealand age range is Palaeogene to Neogene.
Spherical, reticulate, monoporate pollen grains
(Fig. 2Y–AA) are common throughout the late Palaeogene and Neogene, but are rarely abundant. Several
different types have been identified. They are commonly found in samples from wetland environments;
they appear to occur in a range of climatic conditions
from subtropical and cool temperate. At present, the
burr weed Sparganium grows as an emergent macrophyte in freshwater swamps and wetlands on both
North and South Islands (Moore & Edgar, 1970).
Sparganiaceaepollenites sparganioides (Meyer, 1956)
Krutzsch (1970)
Botanical affinity is with Typhaceae (Typha orientalis
C.Presl.).
New Zealand age range is Neogene and younger.
Spherical, monoporate, reticulate pollen grains with
an affinity to extant raupō (T. orientalis; synonym
T. muelleri Rohrb.) occur in the Neogene, particularly
the late Neogene and younger sediments. Modern
Typha L. grows as an emergent macrophyte in freshwater wetland situations, and most of the older fossil
samples tend to come from similar palaeoenvironments, as do the relatively common New Zealand
Neogene Typha macrofossil and dispersed cuticle
records (Pole, 2007; Conran et al., 2015, this issue).
CYPERACEAE
Cyperaceaepollis neogenicus Krutzsch (1970)
Botanical affinity is with Cyperaceae.
New Zealand age range is Neogene and younger.
All Cyperaceae pollen older than Pleistocene is
placed into this form taxon. This periporate pollen
type only becomes prominent in the latest Miocene
and younger sediments and, as such, is usually only
indicative of swampy conditions and wetlands.
However, identifications of older Cyperaceae pollen
need to be treated with caution, as many could represent poorly preserved specimens of other taxa (D. C.
Mildenhall, unpubl. observ.)
POACEAE
Graminidites media Cookson (1947)
Botanical affinity is with the grass family Poaceae.
New Zealand age range is Palaeogene and younger.
429
All Poaceae pollen older than Pleistocene is placed
into this form taxon and, at present, cannot be placed
into subfamilial or lower taxonomic affinities. This
monoporate pollen type only becomes prominent in
the latest Miocene and younger sediments and, as
such, is usually only indicative of drier conditions in
grass- or scrublands, but is also common in wetlands.
Little evidence occurs in New Zealand for savannahlike conditions, although Mildenhall (1993) recorded a
late Pliocene(?) pollen assemblage from samples collected from the base of a 61.6-m drill hole at Wainuiomata near Wellington containing Apiaceae (7%),
Casuarina L. (31–44%) and Poaceae (28–41%). This
assemblage was interpreted as representing a cool
(but more likely dry) climate, in which a scattered
Casuarina-dominated open woodland grew in a predominantly grassland setting. Nevertheless, the
prominence of Eucalyptus L’Hérit. and Casuarinaceae, together with other sclerophyllous taxa, led
Pole (2003) to suggest that the late early Miocene was
seasonally dry with frequent fires, developing into
fully dry climate vegetation by the mid Miocene.
OTHER
MONOCOT TAXA REPORTED FROM
NEW ZEALAND
CENOZOIC
SEDIMENTS
A number of additional taxa not mentioned above,
and currently most without form generic names, have
been listed by their botanical affinity in various publications and largely summarized in Mildenhall
(1980) and Mildenhall & Raine (2012). These include
?Cocos zeylandica Berry (in Couper, 1953), Cordyline
Comm. ex R.Br., Dianella Lam., Juncaceae, Libertia
Spreng., Palmidites maximus Couper (1953), Potamogeton L. and ?Ruppia L. (Appendix).
Large, monosulcate, thin-walled pollen grains were
identified by Couper (1953) as ?Cocos zeylandica, a
name given by Berry (1926) to fossil coconut-like palm
fruits from the Miocene of New Zealand. Although
these pollen grains come from coconut-bearing beds,
they have not been identified positively by other
palynologists and so are regarded as dubious, pending
further study. Similar, but larger grains identified by
Couper (1953) as Palmidites maximus (Fig. 2X) have
also not been identified confidently, and both palynomorphs may not be from palms (or even pollen), but
algal cysts instead (D. C. Mildenhall, unpubl. observ.)
Nupharipollis mortonensis Pocknall & Mildenh.
(Fig. 2U–W) is a monocolpate–monosulcate, shortly
echinate pollen grain which is widespread in Palaeogene to Neogene New Zealand pollen samples.
However, this palynomorph has been variously allied
with Amaryllidaceae, Araceae, Arecaceae, Liliaceae
and/or Nymphaeaceae (Nuphar L.), making phylogenetic, palaeoclimatic or palaeoecological comparisons
difficult.
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430
J. CONRAN ET AL.
Most of the taxa listed above have been identified
occasionally from Cenozoic sediments, but require
more study before their affinities can be determined
with certainty. Others, such as Assamiapollenites
incognitus Pocknall & Mildenh. (Fig. 2E), from Palaeogene to Neogene sediments, and the Neogene
palynomorph A. inanis Pocknall & Mildenh.
(Fig. 2D), although apparently monocotyledonous,
have yet to be affiliated with any living or fossil
family.
DISCUSSION
LATE CRETACEOUS
About 17 palynomorph taxa, including monosulcate
and reticulate lilioid and echinate and gemmate
arecoid palynomorphs, are recognized to date from
the Late Cretaceous, but most are of uncertain affinity and some may not be monocots, as similar pollen
types occur in some early-branching angiosperms of
the Amborellaceae, Nymphaeaceae, Austrobaileyaceae (ANA) grade and of Atherospermataceae and
Chloranthaceae (e.g. Friis, Pedersen & Crane, 2000;
Sampson, 2007).
NEOGENE
In the early Miocene, the climate of New Zealand was
still more or less subtropical to warm temperate, with
rainforests similar in structure and diversity to those
of south-east Queensland (Bannister, Lee & Conran,
2012; Lee et al., 2012), but with significant elements
now restricted to New Caledonia and South America.
Many modern (largely Australasian) monocot groups
were present, together with many of the more conspicuous modern broad-leaved forest and aquatic New
Zealand monocots, such as Asteliaceae, Cordyline
(Fig. 3I), Luzuriaga (Fig. 3E), Phormium (Fig. 4C)
and Potamogetonaceae, and the first appearance of
Asphodelaceae subfamily Asphodeloideae: Bulbinella
Kunth (Fig. 4B). There is also palynological evidence
for locally diverse habitats, despite uniform topographic conditions, possibly reflecting responses to
variations in soil structure and fertility (Mildenhall
et al., 2014). Similarly, Pole (2003, 2014) suggested
that there was both cooling and drying from the early
Miocene, although this is contradicted by Reichgelt
et al. (2013, 2014), who found that Miocene precipitation rates were high, but with a significant difference
between the dry and wet seasons, becoming cooler,
but still relatively moist, towards the middle Miocene.
PALAEOGENE
QUATERNARY
Monocot pollen diversity increased during the Eocene,
possibly, in part, as a response to climate warming
(Harrington, Kemp & Koch, 2004; Sluijs et al., 2011;
Pross et al., 2012), and the climate of New Zealand by
the middle to late Eocene is thought to have been
subtropical to possibly marginally tropical (Hartwich
et al., 2010; Lee et al., 2012). At this time, members of
Arecaceae (Fig. 4D) were diverse and included the
tropical mangrove palm (Nypa), but there were also
asparagoid lineages (including Monogemmites gemmatus), Pandanaceae (Fig. 3D) and Poales, such as
Cyperaceae (Fig. 4I), Restionaceae and Typhaceae
(Fig. 4G), and the enigmatic Nupharipollis mortonensis, present.
By the Oligocene (34–23 Mya), Zealandia was
largely submerged, leading to ‘The Drowning’
hypothesis (Campbell & Hutching, 2007; Trewick,
Paterson & Campbell, 2007; Landis et al., 2008).
However, the New Zealand pollen record generally
and the various monocot lineages show no clear
break in pollen records to support this (Mildenhall
et al., 2013), with many of the same pollen taxa
present before and after the supposed complete submergence in the latest Oligocene or earliest Miocene
(c. 23 Mya). Similarly, there is evidence for a
minimum land area of c. 25 000 km2, comparable
with that of modern New Caledonia (Graham, 2008;
Lee et al., 2013).
Climatic deterioration (cooling) was well underway by
the late Miocene (c. 12 Mya), and most of the changes
seen in the monocot pollen floras at this time were
caused by the loss of tropical or warmer growing taxa
in the late Miocene to early Pleistocene and the
arrival of modern elements in the Quaternary. The
New Zealand Quaternary monocot pollen records
(Moar, Wilmshurst & McGlone, 2011, see also Appendix) suggest a recent and significant influx for many
extant taxa (Fig. 1B, C). These apparently include
members of the Alismatales: Juncaginaceae (Fig. 3C)
and Ruppiaceae (Fig. 3A), Arales: Araceae: Lemna
(Fig. 3B), Asparagales: Iridaceae (Fig. 4A), Orchidaceae (Fig. 3G) and Poales: Juncaceae and Restionaceae subfamily Centrolepidoideae (Fig. 4F), together
with a significant expansion of Cyperaceae and
Poaceae.
The presence of Orchidaceae pollen in Quaternary
sediments contrasts with the occurrence of early
Miocene macrofossils (Conran et al., 2009a), and is
probably at least partly a reflection of the specialized
insect pollination syndrome with the pollen grains
aggregated into pollinia (Faegri & van der Pijl, 1979),
making widespread dispersal of individual grains less
likely. Nevertheless, many of the extant New Zealand
terrestrial orchids are conspecific with, or show close
affinities to, Australian species and represent Quaternary arrivals (Johns & Molloy, 1983).
© 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 421–440
NEW ZEALAND FOSSIL MONOCOT POLLEN
431
Figure 3. Representative extant taxa from New Zealand families or subfamilies present as pollen in Neogene or earlier
fossil deposits, suggesting an ancient historical presence in New Zealand, or some which are only known from Quaternary
deposits. A, Alismatales: Ruppiaceae (Ruppia polycarpa R.Mason) – recent expansion? B, Alismatales: Araceae (Lemna
disperma Hegelm.) – recent expansion. C, Alismatales: Juncaginaceae (Triglochin striata Ruíz & Pav.) – recent expansion.
D, Pandanales: Pandanaceae (Freycinetia banksii A.Cunn.). E, Liliales: Alstroemeriaceae subfamily Luzuriagoideae
[Luzuriaga parviflora (J.D.Hook.) Kunth]. F, Liliales: Ripogonaceae (Ripogonum scandens). G, Asparagales: Orchidaceae
(Thelymitra pulchella J.D.Hook.) – recent expansion? H, Asparagales: Asparagaceae subfamily Lomandroideae [Arthropodium cirratum (G.Forst.) R.Br.]. I, Asparagales: Asparagaceae subfamily Lomandroideae [Cordyline australis (G.Forst.)
Endl.]. Photographs: A, C–I, J. G. Conran; B, Jeremy Rolfe.
Similarly, in Restionaceae, subfamily Centrolepidoideae seems to be a recent arrival, whereas Sporadanthoideae (Fig. 4E) and Leptocarpoideae both
apparently have Neogene or older pollen records
(Mildenhall, 1980; Raine et al., 2011), suggesting possible Miocene or older origins for some of these lineages (Wagstaff & Clarkson, 2012).
As with other elements of the modern New Zealand
flora, the picture seems to show a complex combina-
tion of histories and origins. There is evidence in
some taxa for long-distance dispersal (Pole, 1994,
2001), particularly involving interchange with the
northern Zealandian islands including New
Caledonia (Ladiges & Cantrill, 2007; Pole, 2010), and
vicariance scenarios, or at least long-term occupancy
for several lineages (Lee et al., 2001, 2012). Other
lines also show recent expansion in response to
climate change and tectonic events, creating new
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432
J. CONRAN ET AL.
Figure 4. Representative extant taxa from New Zealand families or subfamilies present as pollen in Neogene or earlier
fossil deposits, suggesting an ancient historical presence in New Zealand, or some which are only known from Quaternary
deposits. A, Asparagales: Iridaceae (Libertia peregrinans Cockayne & Allan). B, Asparagales: Asphodelaceae subfamily
Asphodeloideae (Bulbinella gibbsii Cockayne var. balanifera L.B.Moore). C, Asparagales: Asphodelaceae subfamily
Hemerocallidoideae (Phormium tenax). D, Arecales: Arecaceae (Rhopalostylis sapida). E, Poales: Restionaceae subfamily
Sporadanthoideae (Sporadanthus ferrugineus de Lange, Heenan & B.D.Clarkson). F, Poales: Restionaceae subfamily
Centrolepidoideae [Centrolepis strigosa (R.Br.) Roem. & Schult.] – recent expansion. G, Poales: Juncaceae (Luzula rufa
Edgar var. rufa) – recent expansion. H, Poales: Typhaceae (Typha orientalis). I, Poales: Poaceae [Austroderia richardii
(Endl.) N.P.Barker & H.P.Linder] – recent expansion. Photographs: A–D, F–I, J. G. Conran; E, Jeremy Rolfe.
environments that allowed speciation and/or successful invasion (Pirie et al., 2010; Birch et al., 2012;
Wagstaff & Clarkson, 2012; Linder et al., 2013).
CONCLUSIONS
Monocot pollen has formed a small, but significant,
part of the New Zealand pollen flora from the Late
Cretaceous to present. Throughout the Palaeogene
and Neogene, monocots were highly diverse and climatic conditions ranged from tropical or, more likely,
subtropical to warm temperate with possible periods
of drought, particularly in the Miocene (Mildenhall &
Pocknall, 1989). Throughout this time, some monocot
taxa have persisted and some became extinct as
climate conditions deteriorated towards the end of the
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NEW ZEALAND FOSSIL MONOCOT POLLEN
Neogene, and tropical to subtropical conditions gradually became cool to warm temperate.
From the late Miocene (c. 12 Mya), tectonic activity
and cooler climatic conditions prevailed, with the
result that, through the Neogene into the Pleistocene,
there was a gradual change in diversity with subtropical and rare tropical elements becoming extinct
and a gradual increase in diversity of monocot taxa
adapted to cool temperate environments as new
niches appeared (Mildenhall, 1980).
Many of the taxa described above have botanical
affinities which suggest tropical to subtropical conditions with high humidity. Many are wetland taxa,
which is not surprising as terrestrial sediments tend to
be preserved in such situations, especially where tectonic activity assisted in preserving thick sequences.
The full range of diversity is yet to be elucidated as
many fossil monocot pollen types have yet to be
described and some of those described have no known
modern botanical affinity.
Quaternary sediments hold the most diverse range
of monocot pollen types, but only because of the
widespread nature of terrestrial sediments of this age
and the ease with which the fossilized pollen can be
given modern botanical affinities. Although many
taxa seem to appear at this time, it is more likely that
they gradually appeared over the previous 20 Myr,
evolving and spreading as new ecological niches
appeared in response to orogenesis, habitat fragmentation, climate change and cooling.
ACKNOWLEDGEMENTS
The authors thank GNS Science for permission and
funding to produce this short summary of monocot
pollen history. The School of Earth & Environmental
Sciences, The University of Adelaide and the Departments of Geology and Botany at the University of
Otago are thanked for the provision of resources to
undertake this study. Additional funding was provided by grants from the Royal Society of New
Zealand Marsden Fund, Australian Research Council
and a University of Otago Research Grant. Jennifer
Bannister is thanked for the extraction and preparation of pollen samples from various Neogene and
Palaeogene sites in South Island.
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NEW ZEALAND FOSSIL MONOCOT POLLEN
437
APPENDIX
List of monocotyledonous palynomorphs reported from New Zealand and their modern affinities, where known.
Data from Raine et al. (2011), the New Zealand Fossil Record Electronic Database (FRED), Moar et al. (2011)
and J. I. Raine pers. comm. Family affiliations follow APG III (2009). *Indicates taxon present in modern New
Zealand.
Family affiliation
Taxon
NZ age range
Notes
Liliacidites contortus Mildenhall &
Bannister in Conran et al. 2014a
*Luzuriaga Ruiz & Pav.
*Luzuriaga parviflora (J.D.Hook.)
Kunth
Miocene
Foulden: Luzuriaga
Alstroemeriaceae
subfamily
Luzuriagoideae
Neogene–Quaternary
Quaternary
Recorded also as Luzuriaga marginata
Benth. & J.D.Hook., now recognized
to be a distinct South American
species
Amaryllidaceae?
Monogemmites gemmatus (Couper,
1960) Krutzsch 1970
Late Palaeogene to
Neogene
Foulden: cf. Hymenocallis declinata
Salisb. Ref. Erdtman (1952)
*Lemna L.
Nupharipollis mortonensis Pocknall &
Mildenhall 1984
Holocene
Palaeogene to Neogene
Arecipites ‘magnificus’ Pocknall ms.
Arecipites minutiscabratus (McIntyre
1968) Milne 1988
Arecipites otagoensis (Couper, 1960)
Mildenhall & Pocknall 1989
Arecipites subverrucatus (Pocknall)
Mildenhall & Pocknall 1989
Arecipites waitakiensis (McIntyre 1968)
Mildenhall & Pocknall 1989
Arecipites cf. Rhopalostylis sapida
H.Wendl. & Drude 1878
?Cocos zeylandica Berry 1926
Palaeogene
Palaeogene
?Arecaceae
Palaeogene to Neogene
Foulden
Palmidites maximus Couper 1953
Spinizonocolpites echinatus Muller 1968
Spinizonocolpites prominatus (McIntyre,
1965) Stover & Evans 1973
Spinizonocolpites sp. A Raine 1982
Dicolpopollis cf. D. metroxylonoides
Khan 1976
Dicolpopollis bungonensis Truswell &
Owen 1988
Neogene
Palaeogene
Palaeogene
*Arthropodium R.Br.
Pliocene–Quaternary
*Arthropodium candidum Raoul
*Arthropodium cirratum (G.Forst.) R.Br.
*Cordyline Comm. ex R.Br.
*Cordyline australis (G.Forst.) Endl.
Liliacidites intermedius Couper 1953
Holocene
Quaternary
Neogene–Quaternary
Holocene
Cretaceous to Neogene
*Astelia chathamica (Skottsb.)
L.B.Moore
*Astelia trinervia Kirk
*Collospermum Skottsb.
Quaternary
Araceae
?Araceae, Amaryllidaceae, Arecaceae,
Liliales/Asparagales or Nymphaeaceae
(Nuphar L.)
Arecaceae
Asparagaceae
subfamily
Lomandroideae
Palaeogene
Palaeogene to Neogene
Palaeogene to Quaternary
Rhopalostylis sapida
Neogene
Pollen from C. zeylandica macrofossil
locality in Couper (1953)
?Arecaceae
Nypa
Nypa
Palaeogene
Miocene
Eocene
Not illustrated by Raine (1982)
Metroxylon or Calamus: single NZ
record
Metroxylon or Calamus: single NZ
record
Also recorded as Liliacidites arthroides,
a manuscript name
?Arthropodium
Asteliaceae
Quaternary
Palaeogene –Quaternary
Recorded also as Liliacidites sp. cf.
Collospermum
© 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 421–440
438
J. CONRAN ET AL.
APPENDIX Continued
Family affiliation
Taxon
NZ age range
Notes
Monogemmites Krutzsch 1970
Palaeogene –Quaternary
Includes records of Astelia Banks & Sol.
ex R.Br.; Foulden.
*Baumea type Moar et al. 2011
Holocene
Includes Baumea Gaudich.,
Lepidosperma Labill., Machaerina
Vahl and Tetraria P.Beauv.
*Carex L.
*Carpha alpina R.Br.
Cyperaceaepollis neogenicus Krutzsch
1970
Quaternary
Holocene
?Cretaceous to Quaternary
*Cyperus ustulatus A.Rich.
*Eleocharis R.Br.
*Eleocharis sphacelata R.Br.
*Fimbristylis squarrosa Vahl
*Gahnia type Moar et al. 2011
Holocene
Quaternary
Holocene
Holocene
Quaternary
*Isolepis type Moar et al. 2011
Holocene
*Schoenus type Moar et al. 2011
Holocene
*Oreobolus R.Br.
Holocene
Milfordia homeopunctata (McIntyre,
1965) Partridge in Stover & Partridge
1973
Palaeogene to Neogene
*Libertia Spreng.
Quaternary
*Juncus L.
*Luzula DC.
Quaternary
Quaternary
*Triglochin L.
Quaternary
Liliacidites Couper 1953
Late Cretaceous to
Quaternary
Liliacidites aviemorensis McIntyre 1968
Liliacidites bainii Stover 1973
Palaeogene to Neogene
Palaeogene to Neogene
Liliacidites kaitangataensis Couper
1953
Liliacidites lanceolatus Stover 1973
Liliacidites perforatus Pocknall 1982
Late Cretaceous
Miocene
Palaeogene to Neogene
Liliacidites variegatus Couper 1953
Cretaceous to Neogene
*Orchidaceae pollen indet.
*Thelymitra J.R.Forst. & G.Forst.
?Neogene to Quaternary
Holocene
Dryptopollenites semilunatus Stover in
Stover & Partridge 1973
*Freycinetia banksii A.Cunn.
Palaeogene
Cyperaceae
Recorded by Moar et al. (2011)
Includes records of Cyperaceae Juss.;
records prior to Neogene need to be
re-examined. Foulden.
Recorded by Moar et al. (2011).
Possible late Miocene record
Recorded by Moar et al. (2011).
Recorded by Moar et al. (2011).
Includes Gahnia J.R. & G. Forst. and
Morelotia Gaudich.
Includes Isolepis R.Br. and Uncinia
Pers.
Includes Bolboschoenus (Asch.) Palla,
Desmoschoenus Hook.f.,
Schoenoplectus (Rchb.) Palla and
Schoenus L.
Recorded by Moar et al. (2011).
Flagellariaceae/
Anarthriaceae
Anarthriaceae: Hopkinsia-type or
Flagellariaceae (Macphail & Stone,
2004)
Iridaceae
Juncaceae
Juncaginaceae
Liliaceae
Includes several manuscript species.
Possibly Liliales/Asparagales s.l.,
Monimiaceae(?), Atherospermataceae,
etc.
Foulden
Tentative identifications of Australian
species
Liliales/Asparagales (?Lilium)
Single record of this Australian species
Liliales/Asparagales: Amaryllidaceae, or
Arecaceae. Foulden.
Liliales/Asparagales;
Atherospermataceae (cf. Laurelia
novaezelandiae A. Cunn.)
Orchidaceae
Pandanaceae
Holocene
© 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 421–440
NEW ZEALAND FOSSIL MONOCOT POLLEN
439
APPENDIX Continued
Family affiliation
Taxon
NZ age range
Notes
Lateropora glabra Pocknall &
Mildenhall 1984
Palaeogene to Quaternary
Includes records of Freycinetia Gaud.;
Foulden.
Graminidites media Cookson 1947
*Chionochloa type Moar et al. 2011
*Austroderia (= Cortaderia) type Moar
et al. 2011
*Poa type Moar et al. 2011
Palaeogene to Quaternary
Holocene
Holocene
Includes records of Poaceae; Foulden.
Quaternary
One Pleistocene locality – Solander
Island Volcanics, distinguished from
undifferentiated Poaceae in same
sample
Poaceae
Potamogetonaceae
*Potamogeton L.
Restionaceae
subfamily
*Centrolepis type Moar et al. 2011
Centrolepidoideae
subfamily
*Empodisma type Moar et al. 2011
Leptocaryoideae
Milfordia hypolaenoides Erdtman 1960
*Sporadanthus F.Muell.
Ripogonaceae
*Ripogonum scandens J.R.Forst. &
G.Forst.
Ruppiaceae
*Ruppia L.
Typhaceae
Sparganiaceaepollenites barungensis
Harris 1972
Sparganiaceaepollenites polygonalis
Thiergart 1938
Sparganiaceaepollenites robustisporis
Martin 1973
Sparganiaceaepollenites sparganioides
(Meyer, 1956) Krutzsch 1970
Sparganiaceaepollenites sphericus
(Couper, 1960) Mild. in Mild. &
Crosbie 1979
*Sparganium subglobosum Morong
*Typha orientalis C.Presl.
Xanthorrhoeaceae
(now
Asphodelaceae)
*Bulbinella angustifolia (Cockayne &
Laing) L.B.Moore
*Bulbinella gibbsii Cockayne
*Bulbinella hookeri (Hook.) Cheesem.
Neogene to Quaternary
Quaternary
Quaternary
Neogene
Neogene to Quaternary
Includes Centrolepis Labill. and
Gaimardia Gaudich.
Includes Empodisma L.A.S.Johnson &
D.F.Cutler and Leptocarpus R.Br.
Similar to Empodisma type
Includes Lepyrodia traversii R.Br.
Neogene to Quaternary
Holocene
Palaeogene to Neogene
Sparganium
Quaternary
Sparganium
Neogene
Neogene to Quaternary
Includes records of Typha L.
Palaeogene to Quaternary
Includes records of Sparganium L.
Quaternary
Quaternary
Late Quaternary
Late Quaternary
Neogene to Quaternary
*Herpolirion type Moar et al. 2011
Quaternary
*Dianella intermedia Endl.
*Dianella nigra Colenso
Quaternary
?Neogene to Quaternary
According to Moar et al. (2011),
morphological discrimination of pollen
of Bulbinella species (except B. rossii)
is difficult; therefore the ranges of
individual species given herein should
be regarded as tentative
Includes Dianella Lam. ex Juss. and
Herpolirion J.D.Hook.
The rare, uncertain Neogene records
need re-examination
© 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 421–440
440
J. CONRAN ET AL.
APPENDIX Continued
Family affiliation
Taxon
NZ age range
Notes
Luminidites phormoides (Stover &
Partridge, 1982) Pocknall &
Mildenhall 1984
?Palaeogene to Neogene
Luminidites reticulatus (Couper, 1960)
Pocknall & Mildenhall 1984
Luminidites sp. Pocknall 1985
*Phormium cookianum Le Jol.
Palaeogene to Neogene
Phormium J.R.Forst. & G.Forst and
possibly other Xanthorrhoeaceae:
Hemerocallidoideae, especially
Pasithea D.Don.; includes
pre-Quaternary records of Phormium
J.RForst. & G.Forst. (e.g. by Couper,
1953, 1960)
Xanthorrhoeaceae (Phormium)
Palaeogene
?Neogene to Quaternary
*Phormium tenax
?Neogene to Quaternary
Assamiapollenites incognitus Pocknall &
Mildenhall 1984
Assamiapollenites inanis Pocknall &
Mildenhall 1984
Asteropollis asteroides Hedlund &
Norris 1968
Couperipollis perspinosus (Couper,
1953) Venkatachala & Kar 1969
Neogene
Monosulcites Cookson ex Couper 1953
(pars)
Monosulcites granulatus Couper 1960
Late Cretaceous to
Neogene
Palaeogene
Monosulcites palisadus Couper 1953
Late Cretaceous
Retimonocolpites peroreticulatus
(Brenner 1963) Doyle 1975
Retimonocolpites textus (Norris 1967)
Singh 1983
Trichotomosulcites waronuiensis Couper
1953
Monoporopollenites fossulatus McIntyre
1968
Mid-Cretaceous
Xanthorrhoeaceae (Phormium)
Neogene records need critical
re-examination
Neogene records need critical
re-examination
Presumed monocot,
unknown family
affiliation
Foulden
Palaeogene to Neogene
Mid-Cretaceous
?May be Chloranthaceae
Neogene
Nomen dubium = Monosulcites
perspinosus; not relocated in type
material, possibly not monosulcate
Nomen dubium, possibly = Dilwynites
granulatus (Araucariaceae)
Nomen dubium – morphology poorly
known
Mid-Cretaceous
Late Cretaceous
Nomen dubium – morphology poorly
known
Palaeogene to Neogene
© 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178, 421–440