Podocarpaceae

Kahikatea Dacrycarpus dacrydioides, from here.

Belongs within: Cupressophyta.

The Podocarpaceae are a family of more than 150 living species of mostly Southern Hemisphere conifers. Some are dominant forest species within their range and are used for timber. Many species produce fleshy 'pseudo-fruits' (really an enlarged seed scale) with seeds that are animal-dispersed as with many true fruits. In Podocarpus and Dacrydium, the ovule is exposed atop the seed scale; in Prumnopitys, the seed scale encloses the ovule. Members of the genus Phyllocladus, the celery pines, are distinctive in having the leaves largely replaced by broad, flattened, leaf-like photosynthesising stems. The genus Nageia, found in tropical and subtropical parts of south-east Asia and Australasia, has broad flat leaves more similar to those of Agathis (kauri) species than other Podocarpaceae.

Characters (from Eckenwalder 2009): Evergreen trees or shrubs with scale-, claw- or flattened, needlelike or broader leaves, usually with single undivided midvein. Leaves attached individually in spiral arrangement or in pairs radiating around twig or distichously in more or less flat rows on either side of predominantly horizontal branchlets. Branching typically with extra weaker branches along trunk between main tiers of three to five major branches. Pollen cones small to moderate, single at tips of branchlets or single to clustered in axils of foliage leaves or bracts on otherwise ordinary branchlets or on specialised reproductive shoots, often with one or a few rows of distinctive bracts at base. Pollen scales usually numerous to very numerous, attached spirally or apparently so by a (usually) short stalk, with two (or four) round to slightly elongated pollen sacs attached to stalk and to lower, outer edge of upturned more or less triangular blade. Pollen grains usually with two (or three) variously large to small air bladders. Seed cones single or clustered and often upright on short or long specialised reproductive branchlets in axils of leaves or at tips of otherwise ordinary shoots, almost always with some kind of fleshy structure(s). Seed scales typically few, often just a single fertile scale, with only a little fusion between the sometimes fleshy lower bract portion (podocarpium if fleshy) and the more frequently fleshy upper ovuliferous scale portion (epimatium). Seeds almost always single, folded within epimatium, small to large, plump and wingless, usually animal-dispersed themselves or as part of intact seed cone, with fleshy seed coat, aril, epimatium or podocarpium or combination thereof. Cotyledons normally two, each with two veins. Chromosome base numbers x = 9-19.

Podocarpaceae
    |--Phyllocladus L.C. & Rich. 1826 MM12, A61 [Phyllocladinae]
    |    |--P. alpinus Hook. f. 1853 [=P. trichomanoides var. alpina Parl. in DC. 1868] A61
    |    |--P. aspleniifolius J00 [=Podocarpus aspleniifolius BR00]
    |    |--P. asplenioides Ettingshausen 1886 F71
    |    |--P. glaucus Carr. 1855 [=P. trichomanoides var. glauca Parl. in DC. 1868] A61
    |    |--P. hypophyllus P88
    |    `--P. trichomanoides Don in Lamb. 1832 [incl. P. rhomboidalis Rich. 1832 non L.C. & Rich. 1826] A61
    `--+--Acmopyle MM12
       |    |--A. pancheri MM12
       |    `--A. sahniana MM12
       `--+--Dacrycarpus MM12
          |    |--D. compactus C78
          |    |--D. cumingii H03
          |    |--D. dacrydioides WH02
          |    |--D. imbricatus MM12
          |    |--D. kinabaluensis H03
          |    `--D. steupii H03
          `--+--Nageia nagi MM12
             `--Podocarpus L’Heritier 1807 MM12, A61
                  |--P. acutifolius Kirk 1884 A61
                  |--P. aristulatus J87
                  |--P. atjehensis H03
                  |--P. chinensis MM12
                  |--P. dacrydioides Rich. 1832 (see below for synonymy) A61
                  |--P. drouynianus GK00
                  |--P. falcatus K09
                  |--P. ferrugineus Benn. ex Don in Lamb. 1832 A61
                  |--P. hallii Kirk 1889 (see below for synonymy) A61
                  |--P. imbricatus P88
                  |--P. latifolius MM12
                  |--P. lawrencei MM04
                  |--P. × loderi Ckn. 1932 [P. hallii × P. totara] A61
                  |--P. macrophyllus (Thunberg) Sweet 1818 CD07 [=P. macrophylla LO98]
                  |--P. micropedunculatus H03
                  |--P. nagi MH98
                  |--P. nivalis Hook. 1843 [incl. P. montanus Col. 1895] A61
                  |    |--P. n. var. nivalis A61
                  |    `--P. n. var. erectus Ckn. 1916 A61
                  |--P. novae-caledoniae MM12
                  |--P. nubigena D03
                  |--P. oleifolius R96
                  |--P. praecupressina Ettingshausen 1886 F71
                  |--P. rotundus S03
                  |--P. rumphii H03
                  |--P. spathoides H03
                  |--P. spicatus Br. ex Mirbel 1825 (see below for synonymy) A61
                  |--P. totara Benn. ex Don in Lamb. 1832 A61
                  `--P. urbanii M93

Podocarpaceae incertae sedis:
  Eromangia Pole 2000 P00
    `--*E. douglasii Pole 2000 P00
  Thargomindia Pole 2000 P00
    `--*T. wordsmithiana Pole 2000 P00
  Prumnopitys MM04
    |--P. ferruginea MM04
    `--P. taxifolia WH02
  Dacrydium Sol. ex Forst. f. 1786 A61
    |--D. beccarii P88
    |--D. bidwillii Hook. f. ex Kirk 1878 (see below for synonymy) A61
    |--D. biforme (Hook.) Pilger 1903 (see below for synonymy) A61
    |--D. colensoi Hook. 1843 [incl. D. westlandicum Kirk ex Hook. f. 1877] A61
    |--D. comosum P88
    |--D. cupressinoides Ettingshausen 1886 F71
    |--D. cupressinum Lamb. 1807 [=Thalamia cupressina Spreng. 1826] A61
    |--D. dacrydioides BKS98
    |--D. elatum P88
    |--D. intermedium Kirk 1878 [incl. D. intermedium var. gracilis Kirk 1885] A61
    |--D. kirkii Muell ex Parl. in DC 1868 A61
    |--D. laxifolium Hook. f. 1845 A61
    |--D. magnum H03
    |--D. medium H03
    |--D. pectinatum P88
    `--D. pendulum A27

Dacrydium bidwillii Hook. f. ex Kirk 1878 [incl. D. bidwillii var. erecta Kirk 1889, D. bidwillii var. reclinata Kirk 1889] A61

Dacrydium biforme (Hook.) Pilger 1903 [=Podocarpus biformis Hook. 1843; incl. D. colensoi Hook. f. 1853 non Hook. 1843] A61

Podocarpus dacrydioides Rich. 1832 [incl. Dacrydium excelsum Don in Lamb. 1832, Nageia excela Kuntze 1891, Podocarpus excelsus (Don) Druce 1917, Dacrydium ferrugineum Houtte ex Gord. 1858 A61, P. thujoides Br. in Benn. 1838, D. thuioides Banks & Sol. ex Carr. 1855] A61

Podocarpus hallii Kirk 1889 [=P. totara var. hallii (Kirk) Pilger 1903; incl. P. cunninghamii Colenso 1884] A61

Podocarpus spicatus Br. ex Mirbel 1825 [=Prumnopitys spicata Kent in Veitch 1900; incl. Dacrydium mai Cunn. 1838, Podocarpus matai Lamb. ex Hook. f. 1867, D. mayi Houtte ex Gord. 1858, D. taxifolium Banks & Sol. ex Don in Lamb. 1824 non Po. taxifolia Kunth 1817] A61

*Type species of generic name indicated

REFERENCES

[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[A27] Andersen, J. C. 1927. Popular names of New Zealand plants. Transactions and Proceedings of the New Zealand Institute 57: 905-977.

[BKS98] Baars, R., D. Kelly & A. D. Sparrow. 1998. Liane distribution within native forest remnants in two regions of the South Island, New Zealand. New Zealand Journal of Ecology 22 (1): 71-85.

[BR00] Barnes, R. W., & A. C. Rozefelds. 2000. Comparative morphology of Anodopetalum (Cunoniaceae). Australian Systematic Botany 13 (2): 267-282.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[C78] Clunie, N. M. U. 1978. The vegetation. In Handbooks of the Flora of Papua New Guinea vol. 1 (J. S. Womersley, ed.) pp. 1-11. Melbourne University Press: Carlton South (Australia).

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In Reports of the Princeton University Expeditions to Patagonia, 1896-1899, vol. 8—Botany (W. B. Scott, ed.) pp. 1-34. The University: Princeton (New Jersey).

Eckenwalder, J. E. 2009. Conifers of the World: the complete reference. Timber Press.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[GK00] Gibson, N., & G. J. Keighery. 2000. Flora and vegetation of the Byenup-Muir reserve system, south-west Western Australia. CALMScience 3 (3): 323-402.

[H03] Heads, M. 2003. Ericaceae in Malesia: Vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311-449.

[J00] Johnston, P. R. 2000. Rhytismatales of Australia: The genus Coccomyces. Australian Systematic Botany 13 (2): 199-243.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1-136.

[K09] Kelbessa, E. 2009. Three new species of Acanthaceae from Ethiopia. Kew Bulletin 64 (1): 57-65.

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263-276.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793-7798.

[MM04] Martin, N. A., & L. A. Mound. 2004. Host plants for some New Zealand thrips (Thysanoptera: Terebrantia). New Zealand Entomologist 27: 119-123.

[M93] Mockford, E. L. 1993. North American Psocoptera (Insecta). Sandhill Crane Press, Inc.

[MH98] Morikawa, H., A. Higaki, M. Nohno, M. Takahashi, M. Kamada, M. Nakata, G. Toyohara, Y. Okamura, K. Matsui, S. Kitani, K. Fujita, K. Irifune & N. Goshima. 1998. More than a 600-fold variation in nitrogen dioxide assimilation among 217 plant taxa. Plant, Cell and Environment 21: 180-190.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153-197.

[P88] Polunin, I. 1988. Plants and Flowers of Malaysia. Times Editions: Singapore.

[R96] Righi, G. 1996. Colombian earthworms. Studies on Tropical Andean Ecosystems 4: 485-607.

[S03] Skog, J. E. 2003. Report of the Committee for Fossil Plants: 4. Taxon 52: 341.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).