Podocarpaceae is a large family 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 sin ...

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 20

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

if '' Phyllocladus'' is included and '' Manoao'' and '' Sundacarpus'' are recognized. The family achieved its maximum diversity in the

Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

, making the Podocarpaceae family one of the most diverse in the southern hemisphere. The family is a classic member of the Antarctic flora, with its main centres of diversity in

Australasia Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...

, particularly

New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...

Tasmania Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...

, 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. It is a phytogeographical floristic region in the Paleotropical kingdom. It was first recognized as a distinct region ...

and South America (primarily in the

Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...

Mountains). Several genera extend north of the equator into

Indochina Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) 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 th ...

and the

Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...

. ''

Podocarpus ''Podocarpus'' () is a genus of conifers, the most numerous and widely distributed of the podocarp family, the Podocarpaceae. ''Podocarpus'' species are evergreen shrubs or trees, usually from tall, known to reach at times. The cones have ...

'' 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 Africa, the widespread ''Podocarpus'' and the endemic '' Afrocarpus''. '' Parasitaxus usta'' is unique as the only known parasitic

gymnosperm The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...

. 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 A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...

to the Podocarpaceae '' sensu stricto''. It is treated by some botanists in its own family, the Phyllocladaceae. 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'' and ''

Dacrycarpus ''Dacrycarpus'' is a genus of conifers belonging to the Family (biology), 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 ...

'' 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 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., ''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'' 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...

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 is a Family (biology), family of conifers with three living Genus, genera, ''Araucaria'', ''Agathis'', and ''Wollemia''. While the family's native distribution is now largely confined to the Southern Hemisphere, except for a few spe ...

, and having diverged from it during the late

Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...

. While some fossils attributed to the family have been reported from the Late Permian and Triassic, like '' Rissikia'', these cannot be unambiguously assigned to the family. The oldest unambiguous members of the family are known from the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

period, found across both hemispheres, such as '' Scarburgia'' and '' Harrisiocarpus'' from the Middle Jurassic of England, as well as unnamed species from the Middle-Late Jurassic of Patagonia. Modern genera of the family first appeared during the Early Cretaceous, with the family probably reaching an apex of diversity during the early

Genera

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

List of genera

References

External links

Podocarpaceae
A
The Gymnosperm Database
{{Authority control Pinales families Araucariales