Phyllocladus

Superregnum: Eukaryota
Regnum: Plantae
Divisio: Tracheophyta
Divisio: Pinophyta
Classis: Pinopsida
Ordo: Pinales

Familia: Podocarpaceae
Genus: Phyllocladus
Species: P. alpinus – P. aspleniifolius – P. hypophyllus – P. toatoa – P. trichomanoides

Paleospecies: †P. aberensis – †P. annulatus – †P. elongatus – †P. lobatus – †P. morwellensis – †P. palmerii
Name

Phyllocladus Rich. ex Mirb., Mém. Mus. Hist. Nat. 13: 48 (1825), nom. cons.

Type species: Phyllocladus aspleniifolius (Labill.) Hook.f., London J. Bot. 4: 151 (1845) designated as synonym Phyllocladus billardierei Mirb., Mém. Mus. Hist. Nat. 13: 76 (1825), nom. superfl.

Synonyms

Heterotypic
Brownetera Rich. ex Tratt., Ann. Mus. Hist. Nat. 16: 299 (1810)
Thalamia Spreng., Anleit. Kenntn. Gew., ed. 2, 2: 218 (1817), nom. superfl.

Note: Sometimes placed in its own monotypic family Phyllocladaceae Bessey Nebraska Univ. Stud., 7: 325. (1907), but see Sudianto et al. (2017). New Zealand botanists always favour Phyllocladus alpinus as a separate species not a variety of Phyllocladus trichomanoides (Wassilieff, 2021 cf. Govaerts et al., 2021).
References
Primary references

Richard, L.C.M. 1825. Mémoires du Muséum d'Histoire Naturelle 13: 48. BHL

Additional references

Jordan, G.J., Carpenter, R.J.B., Bannister, J.M.C., Lee, D.E.D, Mildenhall, D.C.E. & Hill, R.S. 2011. High conifer diversity in Oligo-Miocene New Zealand. Australian Systematic Botany, 24(2): 121–136. DOI: 10.1071/SB11004 Reference page.
Pole, M.; Moore, P.R. 2011. A late Miocene leaf assemblage from Coromandel Peninsula, New Zealand, and its climatic implications. Alcheringa 35(1): 103–121. DOI: 10.1080/03115518.2010.481829
Sudianto, E., Wu, C.S., Leonhard, L., Martin, W.F. & Chaw, S.M. 2019. Enlarged and highly repetitive plastome of Lagarostrobos and plastid phylogenomics of Podocarpaceae. Molecular Phylogenetics and Evolution 133: 24-32. DOI: 10.1016/j.ympev.2018.12.012 Paywall PDF Reference page.
Wagstaff, S.J. 2004. Evolution and biogeography of the austral genus Phyllocladus (Podocarpaceae). Journal of Biogeography 31(10): 1569-1577. DOI: 10.1111/j.1365-2699.2004.01066.x Paywall ResearchGate Reference page.
Wassilieff, M. 2021 Conifers - Celery pines Te Ara - the Encyclopedia of New Zealand, [1] Accessed 12 March 2021

Links

Govaerts, R. et al. 2016. Phyllocladus in World Checklist of Selected Plant Families. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2016 Jan. 1. Reference page.
International Plant Names Index. 2016. Phyllocladus. Published online. Accessed: Jan. 1 2016.
Tropicos.org 2016. Phyllocladus. Missouri Botanical Garden. Published online. Accessed: 1 Jan. 2016.

Vernacular names
English: Celery Pines
suomi: Lapakkapuut

Phyllocladus, the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.[1] Species occur mainly in New Zealand, Tasmania, and Malesia in the Southern Hemisphere, though P. hypophyllus ranges into the Philippines, a short way north of the equator.

Characteristics

They are small to medium-sized trees, reaching 10–30 m tall, or sometimes small shrubs. The main structural shoots are green for 2–3 years, then turn brown as the bark thickens. The leaves are sparse, tiny, scale-like, 2–3 mm long, and only green (photosynthetic) for a short time, soon turning brown. Most photosynthesis is performed by highly modified, leaf-like short shoots called phylloclades; these develop in the axils of the scale leaves, and are simple or compound (depending on species). Simple phylloclades are rhombic, 2–5 cm long, and compound phylloclades are up to 20 cm long and subdivided into five to 15 leaflet-like phylloclades 1–3 cm long. The seed cones are berry-like, similar to those of several other Podocarpaceae genera, notably Halocarpus and Prumnopitys, with a fleshy white aril; the seeds are dispersed by birds, which digest the soft, fleshy aril as they pass the hard seeds in their droppings.

Phylloclades of P. trichomanoides

A seedling of P. aspleniifolius with needle-like juvenile leaves

Pollen cones of P. alpinus

Seed cones of P. aspleniifolius

Taxonomy

Phyllocladus is sufficiently morphologically distinct from the other Podocarpaceae genera that some botanists have treated them separately in their own family, Phyllocladaceae.[2] However, while at least one molecular phylogenetic analysis found Phyllocladus to be sister to Podocarpus sensu stricto [3] and another was equivocal on its position relative to Podocarpaceae s.s.,[4] two more recent molecular phylogenetic analyses have placed Phyllocladus within Podocarpaceae as a sister taxon to Lepidothamnus.[5][6] Morphological analysis supports this placement, and therefore it has been suggested that the distinctive phylloclades in the genus are a synapomorphy.[6]

The five species are genetically distinct, and probably arose between 5 and 7 million years ago.[4]
Species
The phylloclades of P. aspleniifolius vary in shape between lobed and pinnate.

Phyllocladus alpinus (P. trichomanoides var. alpinus) - mountain toatoa (New Zealand)
Phyllocladus aspleniifolius - celery-top pine (Tasmania)
Phyllocladus hypophyllus - Malesian celery-pine (New Guinea to Borneo and Philippines)
Phyllocladus toatoa - toatoa (New Zealand)
Phyllocladus trichomanoides - tanekaha (New Zealand)

References

James E. Eckenwalder. 2009. Conifers of the World. Timber Press: Portland, OR, USA. ISBN 978-0-88192-974-4.
Christopher N. Page. 1990. "Phyllocladaceae" pages 317-319. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) The Families and Genera of Vascular Plants volume I. Springer-Verlag: Berlin;Heidelberg, Germany. ISBN 978-0-387-51794-0
William T. Sinclair, R. R. Mill, M. F. Gardner, P. Woltz, T. Jaffré, J. Preston, M. L. Hollingsworth, A. Ponge, and M. Möller. 2002. "Evolutionary relationships of the New Caledonian heterotrophic conifer, Parasitaxis usta (Podocarpaceae), inferred from chloroplast trnL-F intron/spacer and nuclear rDNA ITS2 sequences". Plant Systematics and Evolution 233(1-2):79-104. doi:10.1007/s00606-002-0199-8
Steven J. Wagstaff. 2004. "Evolution and biogeography of the austral genus Phyllocladus (Podocarpaceae)". Journal of Biogeography 31(10):1569-1577.
Ed Biffin, Timothy J. Brodribb, Robert S. Hill, Philip Thomas and Andrew J. Lowe. 2012. "Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation". Proceedings of the Royal Society B Biological Sciences 279:341-348. doi: 10.1098/rspb.2011.0559.

Patrick Knopf, Christian Schulz, Damon P. Little, Thomas Stutzel and Dennis W. Stevenson. 2012. "Relationships within Podocarpaceae based on DNA sequence, anatomical, morphological, and biogeographical data". Cladistics 28: 271–299. doi:10.1111/j.1096-0031.2011.00381.x

Sources

Quinn, C. J. & Price, R. A. 2003. Phylogeny of the Southern Hemisphere Conifers. Proc. Fourth International Conifer Conference: 129-136.