Podocarpaceae

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Podocarpaceae
Temporal range: Jurassic–present
Podocarpus macrophyllus inumaki.JPG
Podocarpus macrophyllus foliage and mature seed cones
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Pinophyta
Class: Pinopsida
Order: Araucariales
Family: Podocarpaceae
Endl.
Genera

See text

Synonyms
  • Acmopylaceae Melikian & Bobrov 1997
  • Bracteocarpaceae Melikian & Bobrov 2000
  • Dacrycarpaceae Melikian & Bobrov 2000
  • Dacrydiaceae Bobrov & Melikian 2000
  • Falcatifoliaceae Melikian & Bobrov 2000
  • Halocarpaceae Melikian & Bobrov 2000
  • Lepidothamnaceae Melikian & Bobrov 2000
  • Microcachrydaceae Doweld & Reveal 1999
  • Microstrobaceae Doweld & Reveal 2001
  • Nageiaceae Fu 1992
  • Parasitaxaceae Melikian & Bobrov 2000
  • Pherosphaeraceae Nakai 1938
  • Phyllocladaceae Bessey 1907
  • Prumnopityaceae Melikian & Bobrov 2000
  • Saxegothaeaceae Gaussen ex Doweld & Reveal 1999

Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs. [1] It contains 19 genera if Phyllocladus is included and Manoao and Sundacarpus are recognized.

Contents

The family is a classic member of the Antarctic flora, with its main centres of diversity in Australasia, 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 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. It occurs on New Caledonia, where it is parasitic on another member of the Podocarpaceae, Falcatifolium taxoides . [2]

The genus Phyllocladus is sister to the Podocarpaceae sensu stricto . [2] It is treated by some botanists in its own family, the Phyllocladaceae. [3]

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 species.

Podocarpus (with 82 to 100 species) [1] [4] 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 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 PodocarpusP. 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. [5]

In 1990, a treatment of the Podocarpaceae recognized 17 genera, excluding Phyllocladus from the family, while recognizing Sundacarpus , but not Manoao. [4] In 1995, Manoao was segregated from Lagarostrobus , based on morphological characteristics. [6] In 2002, a molecular phylogenetic study showed Sundacarpus is embedded in Prumnopitys and the monophyly of Lagarostrobos is doubtful if Manoao is included within it. [2] More recent treatments of the family have recognized Manoao, but not Sundacarpus. [7]

Evolution

Molecular evidence supports Podocarpaceae being the sister group to the Araucariaceae, and having diverged from it during the late Permian. [8] 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 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 Cenozoic. [9]

Genera

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

Knopf 2012 [10] Leslie et al. 2018 [11] [12]
Podocarpaceae

Saxegothaea

Halocarpus

Lepidothamnus

Lagarostrobos

Manoao

Phyllocladus

Prumnopitys

Microcachrys

Pherosphaera

Acmopyle

Dacrycarpus

Falcatifolium

Dacrydium

Retrophyllum

Nageia

Afrocarpus

Podocarpus

Phyllocladoideae
Phyllocladeae

Lepidothamnus

Phyllocladus

Prumnopityeae

Halocarpus

Parasitaxus

Lagarostrobos

Manoao

Pectinopitys

Sundacarpus

Prumnopitys

Podocarpoideae
Saxegothaeeae

Saxegothaea

Microcachrydeae

Microcachrys

Pherosphaereae

Pherosphaera

Acmopyleae

Acmopyle

Dacrydieae

Dacrycarpus

Falcatifolium

Dacrydium

Podocarpeae

Retrophyllum

Afrocarpus

Nageia

Podocarpus

List of genera

Related Research Articles

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms 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 Greek: γυμνόσπερμος, literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, Ginkgo. Gymnosperm lifecycles involve alternation of generations. They have a dominant diploid sporophyte phase and a reduced haploid gametophyte phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

<i>Phyllocladus</i> Genus of plants

Phyllocladus, the celery pines, is a small genus of conifers, now usually placed in the family Podocarpaceae.Species occur mainly in New Zealand, Tasmania, and Malesia in the Southern Hemisphere, though P. hypophyllus ranges into the Philippines, a short way north of the equator.

<i>Podocarpus</i> Genus of conifers in the family Podocarpaceae

Podocarpus is a genus of conifers, the most numerous and widely distributed of the podocarp family, the Podocarpaceae. The name comes from Greek πούς + καρπός. Podocarpus species are evergreen shrubs or trees, usually from 1 to 25 m tall, known to reach 40 m (130 ft) at times. The cones have two to five fused cone scales, which form a fleshy, berry-like, brightly coloured receptacle at maturity. The fleshy cones attract birds, which then eat the cones and disperse the seeds in their droppings. About 97 to 107 species are placed in the genus depending on the circumscription of the species.

<i>Nageia</i> Genus of conifers in the family Podocarpaceae

Nageia is a genus of conifers belonging to the podocarp family Podocarpaceae. Nageia includes evergreen shrubs and trees, from one to 54 meters in height. A 2009 treatment of the genus recognized five species. Some authors consider Nageia formosensis to be a separate species from Nageia nagi, thus recognizing six species. The podocarp genera have been reshuffled by various botanists. Most recently, several species formerly classed as Nageia were moved to the new genus Retrophyllum, while Nageia falcata and Nageia mannii were moved to the new genus Afrocarpus.

<i>Afrocarpus</i> Genus of conifers

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 Podocarpus and Nageia were reclassified.

<i>Prumnopitys</i> Genus of conifers

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.

<i>Dacrydium</i> Genus of conifers

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 more species, which were divided into sections A, B, and C by Florin in 1931. The revisions of de Laubenfels and Quinn, reclassified the former section A as the new genus Falcatifolium, divided Section C into new genera Lepidothamnus, Lagarostrobos and Halocarpus, and retained Section B as genus Dacrydium.

<i>Sundacarpus</i> Single-species genus of conifers

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 a species of Podocarpus or of Prumnopitys.

<i>Dacrycarpus</i> Genus of conifers

Dacrycarpus is a genus of conifers belonging to the family Podocarpaceae. The genus includes nine species of dioecious evergreen trees and shrubs to 55–60 metres (180–197 ft) in height.

<i>Manoao</i> Genus of conifers

ManoaoMAH-noh-ow 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 genus Dacrydium or Lagarostrobos, but has recently been recognised as a distinct genus; some botanists still treat it in Lagarostrobos on the basis that it is not phylogenetically distinct from that genus. In molecular phylogenetic analyses Manoao was found to be related to Parasitaxus and Lagarostrobos, but their exact relationships are unresolved.

<i>Dacrycarpus dacrydioides</i> Coniferous tree endemic to New Zealand

Dacrycarpus dacrydioides, commonly known as kahikatea and white pine, is a coniferous tree endemic to New Zealand. A podocarp, it is New Zealand's tallest tree, gaining heights of 60 m over a life span of 600 years. It was first described botanically by the French botanist Achille Richard in 1832 as Podocarpusdacrydioides, and was given its current binomial name Dacrycarpus dacrydioides in 1969 by the American botanist David de Laubenfels. Analysis of DNA has confirmed its evolutionary relationship with other species in the genera Dacrycarpus and Dacrydium.

<span class="mw-page-title-main">New Caledonia rain forests</span>

The New Caledonia rain forests are a terrestrial ecoregion, located in New Caledonia in the South Pacific. It is a tropical moist broadleaf forest ecoregion, part of the Australasian realm.

<i>Microcachrys</i> Genus of conifers

Microcachrys tetragona, the creeping pine or creeping strawberry pine, is a species of dioecious conifer belonging to the podocarp family (Podocarpaceae). It is the sole species of the genus Microcachrys. The plant is endemic to western Tasmania, where it is a low shrub growing to 1 m tall at high altitudes. Its leaves are scale-like, arranged in opposite decussate pairs, superficially resembling those of the unrelated Diselma archeri (Cupressaceae). It shares the common name Creeping pine with several other plants. Females produce tiny, red, edible berries in summer.

<i>Parasitaxus</i> Species of parasitic conifer from New Caledonia

Parasitaxus usta, also known in French as cèdre rabougri, is a rare species of conifer of the family Podocarpaceae, and the sole species of the genus Parasitaxus.

<i>Retrophyllum</i> Genus of conifers

Retrophyllum is a genus of conifers in the family Podocarpaceae. It contains five generally recognized extant species with a disjunct distribution in the Southern Hemisphere, found in Papuasia and also in South America. Retrophyllum are evergreen trees typically occurring in tropical rainforests and cloud forests.

Dacrydium gracile is a species of conifer in the family Podocarpaceae. It is found only in Malaysian Borneo.

<i>Nageia wallichiana</i> Species of conifer

Nageia wallichiana is a species of conifer in the family Podocarpaceae. It is a tree 10–54 m high, found in Brunei, Cambodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Thailand, and Vietnam. Nageia wallichiana is the most widely distributed species among the seven species in the genus Nageia. If the land areas of China and Japan are excluded, its distribution nearly coincides with that of the genus and includes both the western outliers in India and the easternmost part on Normanby Island. It is one of the most extensive conifer ranges recognized and is similar to Dacrycarpus imbricatus and Podocarpus neriifolius.

Podocarpus borneensis is a species of conifer in the family Podocarpaceae. It native to the island of Borneo, which is divided between Indonesia and Malaysia.

Podocarpus laubenfelsii is a species of conifer in the family Podocarpaceae. It is native to Borneo.

References

  1. 1 2 James E. Eckenwalder. 2009. Conifers of the World. Portland, Oregon: Timber Press. ISBN   978-0-88192-974-4.
  2. 1 2 3 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 Evolution233 (1–2): 79–104. doi : 10.1007/s00606-002-0199-8
  3. 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. ISBN   978-0-387-51794-0
  4. 1 2 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. ISBN   978-0-387-51794-0
  5. 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" Archived 2008-04-08 at the Wayback Machine . South African Journal of Science, 100: 629–632.
  6. Brian P. J. Molloy. 1995. "Manoao (Podocarpaceae), a new monotypic conifer genus endemic to New Zealand". New Zealand Journal of Botany33 (2): 183–201.
  7. Aljos Farjon. 2008. A Natural History of Conifers. Portland, Oregon: Timber Press. ISBN   978-0-88192-869-3
  8. Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (July 19, 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. ISSN   2055-0278. PMID   34282286. S2CID   236141481.
  9. Andruchow-Colombo, Ana; Escapa, Ignacio H; Aagesen, Lone; Matsunaga, Kelly K S (2023-08-04). "In search of lost time: tracing the fossil diversity of Podocarpaceae through the ages". Botanical Journal of the Linnean Society. doi:10.1093/botlinnean/boad027. ISSN   0024-4074.
  10. Knopf; Schulze; Little; Stützel; Stevenson (2012). "Relationships within Podocarpaceae based on DNA sequence, anatomical, morphological, and biogeographical data". Cladistics. 28 (3): 271–299. doi: 10.1111/j.1096-0031.2011.00381.x . PMID   34872191. S2CID   86581015.
  11. Leslie, Andrew B.; Beaulieu, Jeremy; Holman, Garth; Campbell, Christopher S.; Mei, Wenbin; Raubeson, Linda R.; Mathews, Sarah; et al. (2018). "An overview of extant conifer evolution from the perspective of the fossil record". American Journal of Botany. 105 (9): 1531–1544. doi: 10.1002/ajb2.1143 . PMID   30157290.
  12. Leslie, Andrew B.; et al. (2018). "ajb21143-sup-0004-AppendixS4" (PDF). American Journal of Botany. 105 (9): 1531–1544. doi:10.1002/ajb2.1143. PMID   30157290. S2CID   52120430.

Further reading