Podocarpaceae is a large

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

of mainly Southern Hemisphere

conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...

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'' 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 * ...

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 ecologic ...

, 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 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'' 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 ''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 ''Podocarp ...

''. ''

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 ha ...

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, γυμν ...

. 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 ''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 c ...

''. 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 Gui ...

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. 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 withi ...

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., ''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 ''

'' from the family, while recognizing '' Sundacarpus'', 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'' 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 ...

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 In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

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 Paleo ...

. 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 ...

-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

External links

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