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Podocarpaceae is a large
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of mainly Southern Hemisphere
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s, 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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
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 taxon, monotypic genus in the family ''Podocarpaceae.'' The single species, ''M. colensoi'', known as manoao (Māori language, Māori), silver pine, Westland, New Zealand, Westland pine, or white silver pine, is endem ...
'' 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 Antarctic flora are a distinct community of vascular plants which evolved millions of years ago on the supercontinent of Gondwana. Presently, species of Antarctica flora reside on several now separated areas of the Southern Hemisphere, includin ...
, 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 ecologica ...
, particularly
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
,
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
, and New Zealand, and to a slightly lesser extent
Malesia Malesia is a biogeographical region straddling the Equator and the boundaries of the Indomalayan and Australasian realms, and also a phytogeographical floristic region in the Paleotropical Kingdom. It has been given different definitions. The ...
and South America (primarily in the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
Mountains). Several genera extend north of the equator into
Indochina Mainland Southeast Asia, also known as the Indochinese Peninsula or Indochina, is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to the west an ...
and the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. ''
Podocarpus ''Podocarpus'' () is a genus of conifers, the most numerous and widely distributed of the podocarp family, the Podocarpaceae. The name comes from Greek πούς (poús, “foot”) + καρπός (karpós, “fruit”). ''Podocarpus'' species ...
'' 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 Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the African co ...
Africa, the widespread ''Podocarpus'' and the endemic '' Afrocarpus''. ''
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 c ...
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
. It occurs on New Caledonia, where it is parasitic on another member of the Podocarpaceae, ''
Falcatifolium taxoides ''Falcatifolium taxoides'' is a species of conifer in the family Podocarpaceae. It is found only in New Caledonia, and is the only known host of its non-photosynthetic, possibly parasitic relative, '' Parasitaxus usta''. Etymology ''Falcatifoliu ...
''.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 A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a family, familial relationship, it is sometimes used endearingly to r ...
to the Podocarpaceae '' sensu stricto''. 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
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
n 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. Joseph Gaertner (12 March 1732 – 14 July 1791) was a German botanist, best known for his work on seeds, ''De Fructibus et Seminibus Plantarum'' (1788-1792). Biography He was born in Calw, and studied in Göttingen under Albrecht von Haller ...
)
Endl. Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Pressburg, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian Empire, Austrian botanist, numismatist and Sinologist. He was a director of the Botan ...
, ''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 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 ...
'' and the
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
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 worldw ...
, and having diverged from it during the late
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
. The oldest known members of the family are known from the
Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...
(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 mill ...
-aged sediments of the
Umm Irna Formation The Umm Irna Formation is a geological formation in Jordan. It is found in several outcrops in Jordan in the area around the eastern shore of the Dead Sea. It is Late Permian (likely Changhsingian) in age, and is the oldest unit in the succession, ...
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