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Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber Press. . It contains 19 genera if ''
Phyllocladus ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...
'' is included and ''
Manoao ''Manoao'' is a monotypic genus in the family ''Podocarpaceae.'' The single species, ''M. colensoi'', known as manoao ( Māori), silver pine, Westland pine, or white silver pine, is endemic to New Zealand. Before 1996 it was classified in gen ...
'' and ''
Sundacarpus ''Sundacarpus'' is a monotypic genus of conifers containing a single species ''Sundacarpus amarus'', belonging to the family Podocarpaceae. ''Sundacarpus'' was designated a genus by C.N.Page in 1989; formerly it had been classified variously as ...
'' are recognized. The family is a classic member of the Antarctic flora, with its main centres of
diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce * Diversity marketing, marketing communication targeting diverse customers ...
in
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologi ...
, particularly New Caledonia, Tasmania, and New Zealand, and to a slightly lesser extent Malesia and South America (primarily in the Andes Mountains). Several genera extend north of the equator into Indochina and the Philippines. '' Podocarpus'' reaches as far north as southern Japan and southern China in Asia, and Mexico in the Americas, and ''
Nageia ''Nageia'' is a genus of conifers belonging to the podocarp family Podocarpaceae.Christopher N. Page. 1990. "Podocarpaceae" pages 332-346. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and ...
'' into southern China and southern India. Two genera also occur in sub-Saharan Africa, the widespread ''Podocarpus'' and the endemic ''
Afrocarpus ''Afrocarpus'' is a genus of conifers of the family Podocarpaceae. Two to six species are recognized. They are evergreen trees native to Africa. ''Afrocarpus'' was designated a genus in 1989, when several species formerly classified in ''Podocar ...
''. ''
Parasitaxus usta ''Parasitaxus usta'' is a rare species of conifer of the family Podocarpaceae, and the sole species of the genus ''Parasitaxus''. It is a woody shrub up to 1.8 m found only in the remote, densely forested areas of New Caledonia, first discovered ...
'' is unique as the only known
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
gymnosperm. It occurs on New Caledonia, where it is parasitic on another member of the Podocarpaceae, '' Falcatifolium taxoides''.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. The genus ''Phyllocladus'' is sister to the Podocarpaceae ''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
''. It is treated by some botanists in its own family, the
Phyllocladaceae ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...
.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. Berlin, Heidelberg: Springer-Verlag.


Taxonomy

The Podocarpaceae show great diversity, both morphologically and ecologically. Members occur mainly in the Southern Hemisphere, with most genetic variety taking place in New Caledonia, New Zealand, and Tasmania. Species diversity of ''Podocarpus'' is found mainly in South America and the Indonesian islands, the latter also being rich in ''
Dacrydium ''Dacrydium'' is a genus of conifers belonging to the podocarp family Podocarpaceae. Sixteen species of evergreen dioecious trees and shrubs are presently recognized. The genus was first described by Solander in 1786, and formerly included many m ...
'' and ''
Dacrycarpus ''Dacrycarpus'' is a genus of conifers belonging to the family Podocarpaceae.Christopher N. Page. 1990. "Podocarpaceae" pages 332-346. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and Gene ...
'' species. ''Podocarpus'' (with 82 to 100 species)Christopher N. Page. 1990. "Podocarpaceae" pages 332–346. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and Genera of Vascular Plants'' volume I. Berlin, Heidelberg: Springer-Verlag. and ''Dacrydium'' (with 21 species) are the largest genera. A few genera are common to New Zealand and South America, supporting the view that podocarps had an extensive distribution over southern
Gondwanaland Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
. The breaking up of Gondwanaland led to large-scale
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
of the Podocarpaceae. Until 1970, only seven Podocarpaceae genera were recognized: ''Podocarpus'', ''Dacrydium'', ''Phyllocladus'', ''Acmopyle'', ''Microcachrys'', ''Saxegothaea'', and ''Pherosphaera''. All four of the African species fell under ''Podocarpus'' – ''P. falcatus'', ''P. elongatus'', ''P. henkelii'', and ''P. latifolius''. Taxonomists divided ''Podocarpus'' species into eight species groups based on leaf anatomy: ''Afrocarpus'' J.Buchholz & N.E.Gray, ''Dacrycarpus'' Endl., ''Eupodocarpus'' Endl., ''Microcarpus'' Pilg., ''Nageia'' ( Gaertn.)
Endl. Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian botanist, numismatist and Sinologist. He was a director of the Botanical Garden of Vienna. Bi ...
, ''Polypodiopsis'' C.E.Bertrand (non ''Polypodiopsis'' Carriére nom. rej. prop. 6), ''Stachycarpus'' Endl. and ''Sundacarpus'' J.Buchholz and N.E.Gray. Studies of embryology, gametophyte development, female cone structure, and cytology led to the belief that the eight categories probably deserved generic status. Researchers agreed on the need to recognize "fairly natural groupings which prove to have good geographic and probably evolutionary cohesion" and took the necessary steps to raise each section to generic status.Barker, N. P.; Muller, E. M.; and Mill, R. R. (2004)
"A yellowwood by any other name: molecular systematics and the taxonomy of ''Podocarpus'' and the Podocarpaceae in southern Africa"
. ''South African Journal of Science'', 100: 629–632.
In 1990, a treatment of the Podocarpaceae recognized 17 genera, excluding ''
Phyllocladus ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...
'' from the family, while recognizing ''
Sundacarpus ''Sundacarpus'' is a monotypic genus of conifers containing a single species ''Sundacarpus amarus'', belonging to the family Podocarpaceae. ''Sundacarpus'' was designated a genus by C.N.Page in 1989; formerly it had been classified variously as ...
'', but not ''Manoao''. In 1995, ''Manoao'' was segregated from '' Lagarostrobus'', based on morphological characteristics. Brian P. J. Molloy. 1995. "''Manoao'' (Podocarpaceae), a new monotypic conifer genus endemic to New Zealand". ''New Zealand Journal of Botany'' 33 (2): 183–201. In 2002, a
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study showed ''Sundacarpus'' is embedded in ''
Prumnopitys ''Prumnopitys'' is a genus of conifers belonging to the family Podocarpaceae. The nine recognized species of ''Prumnopitys'' are densely branched, dioecious evergreen trees up to 40 metres in height. Etymology The name ''Prumnopitys'' comes fr ...
'' and the monophyly of ''Lagarostrobos'' is doubtful if ''Manoao'' is included within it. More recent treatments of the family have recognized ''Manoao'', but not ''Sundacarpus''.Aljos Farjon. 2008. ''A Natural History of Conifers''. Portland, Oregon: Timber Press.


Evolution

Molecular evidence supports Podocarpaceae being the sister group to the
Araucariaceae Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the Jurassic and Cretaceous periods and the early Cenozoic, when it was distributed almost world ...
, and having diverged from it during the late Permian. The oldest known members of the family are known from the Lopingian (Late Permian), likely
Changhsingian In the geologic time scale, the Changhsingian or Changxingian is the latest age or uppermost stage of the Permian. It is also the upper or latest of two subdivisions of the Lopingian Epoch or Series. The Changhsingian lasted from to 251.902 mi ...
-aged sediments of the Umm Irna Formation in Jordan.


Genera

Studies based on anatomical, biogeographical, morphological, and DNA evidence suggest these relationships:


List of genera


References


Further reading

* Christopher J. Quinn and Robert A. Price. 2003. "Phylogeny of the Southern Hemisphere Conifers". ''Proceedings of the Fourth International Conifer Conference'': 129–136. DOI
10.17660/ActaHortic.2003.615.10


External links



A
The Gymnosperm Database
{{Taxonbar, from=Q157160 Pinales families Araucariales