3/21/2017 PhyllocladusfossilsfromtheMioceneofNewZealand,andCretaceousProtophyllocladusMikePole

MikePole

March 4, 2017 / comment 0 / Edit

PhyllocladusfossilsfromtheMioceneofNewZealand,and
CretaceousProtophyllocladus

A rareplant fossil in the Miocene Manuherikia Group of New Zealand, is Phyllocladus(the Celery
Pine). This is a strange conifer which, instead of leaves, the adult plant has multi-veined attened
branches that are called phylloclades. With thesephylloclades, the average person would scarcely
believe Phyllocladus is a kind of conifer (very broadly, a pine). But it is common in some parts of New
Zealand today (typically in difcult places, cold, or with poor soils) and also in Tasmania and
Malaysia.

Fossil pollen suggestsPhyllocladus has been in New Zealand since the Eocene (Mildenhall 1980).
There are somewhat similar, but older pollen records, for example from New Zealands Late
Cretaceous (Couper, 1953; Cookson & Pike, 1954) but these are likely to represent something
distinct. Despite conifers being verydiversein the Miocene St Bathans Paleovalley sediments of the
Manuherikia Group, Phyllocladus appears to have been absent there (Pole 1992a, 1997, 2007). The
climate may have been so warm, and the soils rich enough, to preclude it (Pole 2014).
However,Phyllocladus is present in the Miocene Nevis Oil Shale (Pole 1992, see gured image), which
was deposited in very different conditions (Douglas 1986). There are hints from the Nevis, including
large fossil legumes (Pole et al. 1989, 1992b) that the climate may have had a distinct dry season. This
may have favouredPhyllocladus. DNA evidence suggest the extant New Zealand species only radiated
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in the Late Cenozoic (Wagstaff, 2004), and there are certainly Phyllocladus fossils from this time (e.g.
From Mataora, in the Coromandel. Pole and Moore, 2011).

Fossil Phyllocladus from Mataora,

New Zealand.

Phylloclades appears to be a strategy to increase light-intercepting area beyond that of typical,

simple, needle-like leaves ( e.g. Brodribb 2011). In essence, Phyllocladusistrying to become more like
the broad leaves of typical owing plants, in order to compete with them. Separate from that
hypothesis,there seems to be an unstated assumption that the worlds forests became darker,
shadier places, with the evolution of broad-leaved owering plants. But did they? Conifer forests can
be damned dark under the canopy. An individual pine needle doesnt create much shade, but a whole
lot of them do.

In one of the places I lived in for a few years in Brisbane, my neighbour on the opposite side of the
street was Ray Specht. There was a big Eucalyptus tree growing between his house and mine. Ray
happened to be one of the most inuential people in Australian botany. For example, the various
schemes of Australian vegetation classication mostly stem from him. On visiting Ray, the
conversation would usually quickly go to the 1948 American-Australian Scientic Expedition to
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Arnhem Land, which was clearly a highlight of his life. Ray would expound on the expedition until his
wife would look up from her book and go Oh Ray shut up! Then, fascinating as the Expedition
was, I could get him back to what I really wanted him to talk about his views on howvegetation
structure could be predicted from basic environmental properties.

One of Spechts understandings was that the amount of shade under the canopy of a forest was
fundamentally determined by the environment. Basically, closed canopy forests (like rainforests) are
more open underneath the canopy, whereas open canopy forests (like Eucalyptus forests) have a
denser zone of shrubs. Down near the ground, things even out. There are no doubt plenty of
botanists who might take issue with the details, but I nd it a valuable insight. At least in Spechts
view, given the right environmental conditions, forests would have been just as dark and shady
before owering plants evolved, as after.

Ive wondered if any other fossils might be helpful. In fact, there is a fossil plant with the intriguing
name Protophyllocladus has long been known from Cretaceous rocks right across the Northern
Hemisphere. In terms of gross morphology, it has phylloclades uncannily like Phyllocladus. However,
their detailed structure, and its afnities, have remained unclear up until very recently. But in 2014
the Russian scientists Natalya Nosova and Lena Golovneva from St Petersburg made a detailed study
of the fossil epidermis of Protophyllocladus and compared it with Phyllocladus (Nosova and Golovneva
2014). They concludedProtophyllocladus was a conifer, and likely in the same family asPhyllocladus
(the Podocarpaceae). Protophyllocladus is much older thanPhyllocladus, but it still comes from a time
when owering plant forests had spread across the Northern Hemisphere (see g. 7 in Pole et al.
2016). In that respect, it is consistent with the idea that phylloclades evolved in response to the
spread of owering plants. If fossils ofconifers with phylloclades turn up from pre-angiosperm times,
then Spechts ideas should be looked at closely.

Extant Phyllocladus trichomanoides

However, my feeling is that the epidermal morphology of Protophyllocladus is distinct from

Phyllocladus or any of the extant related plants (the family Podocarpaceae). For instance the

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epidermal cells of Protophyllocladus do not often occur in les. That is, I think the phylloclades of
Protophyllocladus probably evolved quite independently of Phyllocladus.

Lightening, so it is said, never strikes the same place twice. But of course, we know this is wrong
lightening does hit some places multiple times and even the odd person has been struck more than
once. I think of evolutionary developments as a bit like lightening striking a particular place (a new
morphological feature evolves), but just occasionally, it strikes there again (and a similar feature
evolves in another lineage. The technical term for this is convergent evolution. I suspect the
phylloclades ofProtophyllocladus andPhyllocladus are convergent evolution.

And that bigEucalyptus between my place and Ray Spechts ? According to him it had been hit by
lightening at least three times.

References

Links will take you to a site to download pdfs of the papers.

Brodribb, T.J., 2011. A functional analysis of podocarp ecology, in: Turner, B.L., Cernusak, L.A. (Eds.),
Ecology of the Podocarpaceae in Tropical Forests, Smithsonian Contributions to Botany, No. 95. .
Smithsonian Institution Scholarly Press, Washington, D.C., pp. 165-173.

Douglas, B. J. 1986. Lignite resources of Central Otago. New Zealand Energy Research and
Development Committee Publication P104: Volume one, Volume 2.

Mildenhall, D.C., 1980. New Zealand Late Cretaceous and Cenozoic plant biogeography: a
contribution. Palaeogeography, Palaeoclimatology, Palaeoecology 31, 197-233.

Nosova, N., Golovneva, L., 2014. The Mesozoic genus Protophyllocladus Berry (Pinopsida). Review of
Palaeobotany and Palynology 210, 7788.

Pole, M.S., 1992a. Early Miocene ora of the Manuherikia Group, New Zealand. 2. Conifers. Journal
of the Royal Society of New Zealand 22, 287-302.

Pole, M.S., 1997. Miocene conifers from the Manuherikia Group, New Zealand. Journal of the Royal
Society of New Zealand 27, 355-370.

Pole, M., 2007. Conifer and cycad distribution in the Miocene of southernNew Zealand. Australian
Journal of Botany 55, 143-164.

Pole, M., Moore, P.R., 2011. A late Miocene leaf assemblage from Mataora, Coromandel, New
Zealand and its climatic implications. Alcheringa 35, 103-121.
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Pole, M.S., 1992. Fossils of Leguminosae from the Miocene Manuherikia Group of New Zealand, in:
Herendeen, P.S., Dilcher, D.L. (Eds.), Advances in Legume Systematics: Part 4. The Fossil Record. The
Royal Botanic Gardens, Kew, pp. 251-258.

Pole, M.S., Holden, A.M., Campbell, J.D., 1989. Fossil legumes from the Manuherikia Group (Miocene),
Central Otago, New Zealand. Journal of the Royal Society of New Zealand 19, 225-228.

Pole, M., Wang, Y., Bugdaeva, E.V., Dong, C., Tian, N., Li, L., Zhou, N., 2016. The rise and demise of
Podozamites in east AsiaAn extinct conifer life style, . Palaeogeography, Palaeoclimatology,
Palaeoecology.

Wagstaff, S.J., 2004. Evolution and biogeography of the austral genus Phyllocladus (Podocarpaceae).
Journal of Biogeography 31, 15691577.

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