{{Short description|Podocarp family of conifers}} {{Automatic taxobox | image = Podocarpus elatus habit.jpg | image_caption = ''Podocarpus elatus'' Illawarra Plum | fossil_range = {{fossilrange|Middle Jurassic|present}} | taxon = Podocarpaceae | authority = Endl. | subdivision_ranks = Genera | subdivision = See text | synonyms = *Acmopyleaceae {{small|(Pilg.) Melikyan & A.V.Bobrov}} *Microcachrydaceae {{small|Doweld & Reveal}} *Microstrobaceae {{small|Doweld & Reveal}} *Nageiaceae {{small|D.Z.Fu}} *Phyllocladaceae {{small|Bessey}} *Phyllocladaceae {{small|Core ex H.Keng}} *Phyllolcadaceae {{small|Bessey}} *Prumnopityaceae {{small|A.V.Bobrov & Melikyan}} *Saxegothaeaceae {{small|Gaussen ex Doweld & Reveal}} |synonyms_ref = {{R|WFO|COL}} }}
'''Podocarpaceae''' is a large family of mainly southern hemisphere conifers, known in English as '''podocarps''', comprising about 201 species of evergreen trees and shrubs.<ref>{{Cite journal |last=Khan |first=Raees |last2=Hill |first2=Robert S. |last3=Liu |first3=Jie |last4=Biffin |first4=Ed |date=2023-03-03 |title=Diversity, Distribution, Systematics and Conservation Status of Podocarpaceae |url=https://www.mdpi.com/2223-7747/12/5/1171 |journal=Plants |language=en |volume=12 |issue=5 |pages=1171 |doi=10.3390/plants12051171 |doi-access=free|issn=2223-7747 |pmc=10005643 |pmid=36904033}}</ref> It contains 20 genera if ''Phyllocladus'' is included and ''Manoao'' and ''Sundacarpus'' are accepted. The family achieved its maximum diversity in the Cenozoic, 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, 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''.<ref name="sinclair2002">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. {{doi|10.1007/s00606-002-0199-8}}</ref>
Except for the two genera Pherosphaera and Saxegothaea, all members of Podocarpaceae have fleshy seed cones, which appears to be an ancestral trait.<ref>[https://academic.oup.com/aob/article/130/5/637/6652287 Detailed seed cone morpho-anatomy of the Prumnopityoid clade: an insight into the origin and evolution of Podocarpaceae seed cones]</ref>
The genus ''Phyllocladus'' is sister to the Podocarpaceae ''sensu stricto''.<ref name="sinclair2002"/> It is treated by some botanists in its own family, the Phyllocladaceae.<ref name="pagephyllocladaceae">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}}</ref>
==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)<ref name="eckenwalder2009">James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber Press. {{ISBN|978-0-88192-974-4}}.</ref><ref name="pagepodocarpaceae">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}}</ref> 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 accepted: ''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.<ref name="Barker et al. 2004">Barker, N. P.; Muller, E. M.; and Mill, R. R. (2004). [http://eprints.ru.ac.za/126/01/sajsci_v100_n11_a26%5B1%5D.pdf "A yellowwood by any other name: molecular systematics and the taxonomy of ''Podocarpus'' and the Podocarpaceae in southern Africa"] {{webarchive|url=https://web.archive.org/web/20080408221809/http://eprints.ru.ac.za/126/01/sajsci_v100_n11_a26%5B1%5D.pdf |date=2008-04-08 }}. ''South African Journal of Science'', 100: 629–632.</ref>
In 1990, a treatment of the Podocarpaceae accepted 17 genera, excluding ''Phyllocladus'' from the family, while recognizing ''Sundacarpus'', but not ''Manoao''.<ref name="pagepodocarpaceae"/> In 1995, ''Manoao'' was segregated from ''Lagarostrobus'', based on morphological characteristics.<ref name="molloy1995">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.</ref> 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.<ref name="sinclair2002"/> More recent treatments of the family have accepted both ''Manoao'' and ''Sundacarpus'', resolving the monophyly of ''Prumnopitys'' with respect to ''Sundacarpus'' by splitting some species out into a new genus ''Pectinopitys''.<ref name="POWO">{{cite web| title=Podocarpaceae Endl. | website=Plants of the World Online | date=2020-05-27 | url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:77126768-1 | access-date=2025-12-29}}</ref>
==Evolution== Molecular evidence supports Podocarpaceae being the sister group to the Araucariaceae, and having diverged from it during the late Permian.<ref>{{Cite journal|last1=Stull|first1=Gregory W.|last2=Qu|first2=Xiao-Jian|last3=Parins-Fukuchi|first3=Caroline|last4=Yang|first4=Ying-Ying|last5=Yang|first5=Jun-Bo|last6=Yang|first6=Zhi-Yun|last7=Hu|first7=Yi|last8=Ma|first8=Hong|last9=Soltis|first9=Pamela S.|last10=Soltis|first10=Douglas E.|last11=Li|first11=De-Zhu|date=July 19, 2021|title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms|url=https://www.nature.com/articles/s41477-021-00964-4|journal=Nature Plants|language=en|volume=7|issue=8|pages=1015–1025|doi=10.1038/s41477-021-00964-4|pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |issn=2055-0278|url-access=subscription}}</ref> 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.<ref>{{Cite journal|last1=Andruchow-Colombo |first1=Ana |last2=Escapa |first2=Ignacio H |last3=Aagesen |first3=Lone |last4=Matsunaga |first4=Kelly K S |date=2023-08-04 |title=In search of lost time: tracing the fossil diversity of Podocarpaceae through the ages |url=https://academic.oup.com/botlinnean/advance-article/doi/10.1093/botlinnean/boad027/7237351 |journal=Botanical Journal of the Linnean Society |volume=203 |issue=4 |pages=315–336 |language=en |doi=10.1093/botlinnean/boad027 |issn=0024-4074|hdl=11336/227952 |hdl-access=free }}</ref>
==Genera== Studies based on anatomical, biogeographical, morphological, and DNA evidence suggest these relationships:
{| class="wikitable" |- ! colspan=1 | Knopf 2012<ref name = knopf2012podocarpaceaerelationships/> ! colspan=1 | Leslie et al. 2018<ref>{{cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |display-authors=et al. |year=2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 | doi=10.1002/ajb2.1143 |pmid= 30157290|pmc= |bibcode= |doi-access=free}}</ref><ref>{{cite journal |last1=Leslie |first1=Andrew B. |display-authors=et al. |year=2018 |title=ajb21143-sup-0004-AppendixS4 |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430 |url=https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf |doi-access=}}</ref> |- | style="vertical-align:top| {{Clade | style=font-size:90%;line-height:90% |label1=Podocarpaceae |1= {{clade |1=''Saxegothaea'' |2={{clade |1={{clade |1=''Halocarpus'' |2={{clade |1=''Lepidothamnus'' |2={{clade |1=''Lagarostrobos'' |2=''Manoao'' }} |3=''Phyllocladus'' }} |3=''Prumnopitys'' }} |2={{clade |1={{clade |1=''Microcachrys'' |2=''Pherosphaera'' }} |2={{clade |1=''Acmopyle'' |2={{clade |1={{clade |1=''Dacrycarpus'' |2={{clade |1=''Falcatifolium'' |2=''Dacrydium'' }} }} |2={{clade |1={{clade |1=''Retrophyllum'' |2={{clade |1=''Nageia'' |2=''Afrocarpus'' }} }} |2=''Podocarpus'' }} }} }} }} }} }} }} | {{Clade | style=font-size:90%;line-height:80% |1={{clade |label1=Phyllocladoideae |1={{Clade |label1=Phyllocladeae |1={{Clade |1=''Lepidothamnus'' |2=''Phyllocladus'' }} |label2=Prumnopityeae |2={{Clade |1=''Halocarpus'' |2={{Clade |1={{Clade |1=''Parasitaxus'' |2={{Clade |1=''Lagarostrobos'' |2=''Manoao'' }} }} |2={{Clade |1=''Pectinopitys'' |2={{Clade |1=''Sundacarpus'' |2=''Prumnopitys'' }} }} }} }} }} |label2=Podocarpoideae |2={{Clade |label1=Saxegothaeeae |1=''Saxegothaea'' |2={{Clade |label1=Microcachrydeae |1=''Microcachrys'' |2={{Clade |label1=Pherosphaereae |1=''Pherosphaera'' |2={{Clade |label1=Acmopyleae |1=''Acmopyle'' |2={{Clade |label1=Dacrydieae |1={{Clade |1=''Dacrycarpus'' |2={{Clade |1=''Falcatifolium'' |2=''Dacrydium'' }} }} |label2=Podocarpeae |2={{Clade |1={{Clade |1=''Retrophyllum'' |2={{Clade |1=''Afrocarpus'' |2=''Nageia'' }} }} |2=''Podocarpus'' }} }} }} }} }} }} }} }} |}
===List of extant genera=== {{Columns-list| * ''Acmopyle'' * ''Afrocarpus'' * ''Dacrycarpus'' * ''Dacrydium'' * ''Falcatifolium'' * ''Halocarpus'' * ''Lagarostrobos'' * ''Lepidothamnus'' * ''Manoao'' * ''Microcachrys'' * ''Nageia'' * ''Parasitaxus'' * ''Pectinopitys'' * ''Pherosphaera'' (syn. ''Microstrobos'') * ''Phyllocladus'' * ''Podocarpus'' * ''Prumnopitys'' * ''Retrophyllum'' * ''Saxegothaea'' * ''Sundacarpus'' }}
== Extinct genera == {{Unreferenced section|date=September 2025}} Genera that have been moved into a new subfamily are tagged with ±. Genera that are wood{{explain|date=September 2025}} are tagged with #. * †''Podocarpoxylon ± #'' * †''Protophyllocladoxylon #'' * †''Dacrycarpites'' * †''Dacrydiumites'' * †''Lygistepollenites''? * †''Gamerroites''? Moved to Coniferae incertae sedis * †''Conites''? * †''Microcachryidites'' * †''Microcachryoxylon'' # * †''Phyllocladidites'' * †''Podocarpidites'' * †''Podosporites'' * †''Metapodocarpoxylon #'' * †''Protopodocarpoxylon #'' * †''Phyllocladoxylon ± #'' * †''Rissikia'' * †''Scarburgia'' * †''Harrisiocarpus''
== References == {{Reflist|refs = <ref name = knopf2012podocarpaceaerelationships>{{cite journal|title = Relationships within Podocarpaceae based on DNA sequence, anatomical, morphological, and biogeographical data|last1 = Knopf|last2 = Schulze|last3 = Little|last4 = Stützel|last5 = Stevenson|journal = Cladistics|volume = 28|date = 2012|issue = 3|pages = 271–299|doi = 10.1111/j.1096-0031.2011.00381.x| pmid=34872191 | s2cid=86581015 |doi-access = free}}</ref>
<ref name="COL">{{cite web |last1=Bánki |first1=Olaf |last2=Roskov |first2=Yury |last3=Döring |first3=Markus |last4=Ower |first4=Geoff |last5=Hernández Robles |first5=Diana Raquel |last6=Plata Corredor |first6=Camila Andrea |last7=Stjernegaard Jeppesen |first7=Thomas |last8=Örn |first8=Ari |last9=Pape |first9=Thomas |last10=Hobern |first10=Donald |last11=Garnett |first11=Stephen |last12=Little |first12=Holly |last13=DeWalt |first13=R. Edward |last14=Ma |first14=Keping |last15=Miller |first15=Joe |last16=Orrell |first16=Thomas |last17=Aalbu |first17=Rolf |last18=Abbott |first18=John |last19=Adlard |first19=Robert |last20=Aedo |first20=Carlos |last21=Aescht |first21=E. |last22=Alexander |first22=Sara |last23=Alonso-Zarazaga |first23=Miguel A. |last24=Alvarez |first24=Belinda |last25=Andrella |first25=Giovani Carlos |last26=Antonietto |first26=Lucas Silveira |last27=Arango |first27=Claudia |last28=Artois |first28=Tom |last29=Atahuachi Burgos |first29=Margoth |last30=Atkinson |first30=Stephen |display-authors=1 |title=''Podocarpaceae'' Endl. |url=https://www.catalogueoflife.org/data/taxon/C7ZN4 |website=Catalogue of Life |publisher=Species 2000 & ITIS |date=2025 |access-date=12 June 2025 |doi=10.48580/dgqdn }}</ref>
<ref name="WFO">{{cite web |url=https://www.worldfloraonline.org/taxon/wfo-7000000484#synonyms |title=''Podocarpaceae'' Endl. |website=World Flora Online |publisher=World Flora Online Consortium |year=2025 |access-date=12 June 2025}}</ref> }}
==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== {{Commons category}} {{Wikispecies}} * [http://www.conifers.org/po/index.htm Podocarpaceae] At [http://www.conifers.org The Gymnosperm Database]
{{Pinophyta}} {{Taxonbar|from=Q157160}} {{Authority control}}
Category:Podocarpaceae Category:Pinales families Category:Araucariales