Podocarpaceae

The podocarps, a family of about 201 evergreen trees and shrubs, are primarily Southern Hemisphere conifers. If the genera *Phyllocladus*, *Manoao*, and *Sundacarpus* are all counted, the family contains 20 genera. Their peak diversity occurred during the Cenozoic, making them one of the most varied conifer families in the Southern Hemisphere.

This family is a classic part of the Antarctic flora. Its main centers of diversity are Australasia—especially New Caledonia, Tasmania, and New Zealand—followed by Malesia and the Andes of South America. Some genera stretch north of the equator into Indochina and the Philippines. *Podocarpus* reaches as far north as southern Japan and China in Asia and Mexico in the Americas, while *Nageia* extends into southern China and India. Two genera appear in sub-Saharan Africa: the widespread *Podocarpus* and the endemic *Afrocarpus*.

The only known parasitic gymnosperm is *Parasitaxus usta*, found in New Caledonia, where it lives on another podocarp, *Falcatifolium taxoides*. Except for *Pherosphaera* and *Saxegothaea*, all podocarps have fleshy seed cones, which seems to be an ancestral trait. The genus *Phyllocladus* is the sister group to the rest of the family (Podocarpaceae *sensu stricto*); some botanists place it in its own family, Phyllocladaceae.

**Taxonomy** Podocarps are morphologically and ecologically diverse. Most genetic variety occurs in New Caledonia, New Zealand, and Tasmania. *Podocarpus* species are most diverse in South America and the Indonesian islands, while the latter also hosts many *Dacrydium* and *Dacrycarpus* species. *Podocarpus* (82–100 species) and *Dacrydium* (21 species) are the largest genera. A few genera are shared between New Zealand and South America, suggesting podocarps were widespread across southern Gondwanaland; its breakup drove large-scale speciation.

Until 1970, only seven genera were accepted: *Podocarpus*, *Dacrydium*, *Phyllocladus*, *Acmopyle*, *Microcachrys*, *Saxegothaea*, and *Pherosphaera*. All four African species were placed in *Podocarpus*: *P. falcatus*, *P. elongatus*, *P. henkelii*, and *P. latifolius*. Taxonomists later split *Podocarpus* into eight species groups based on leaf anatomy: *Afrocarpus*, *Dacrycarpus*, *Eupodocarpus*, *Microcarpus*, *Nageia*, *Polypodiopsis*, *Stachycarpus*, and *Sundacarpus*. Studies of embryology, gametophyte development, female cone structure, and cytology suggested these groups deserved generic status, and researchers agreed to recognize "fairly natural groupings which prove to have good geographic and probably evolutionary cohesion," raising each section to a genus.

In 1990, a treatment accepted 17 genera, excluding *Phyllocladus* from the family while recognizing *Sundacarpus* but not *Manoao*. In 1995, *Manoao* was separated from *Lagarostrobus* based on morphology. A 2002 molecular study showed *Sundacarpus* is embedded within *Prumnopitys* and that *Lagarostrobos* is not monophyletic if *Manoao* is included. More recent treatments accept both *Manoao* and *Sundacarpus*, resolving *Prumnopitys* monophyly by splitting some species into a new genus, *Pectinopitys*.

**Evolution** Molecular evidence places Podocarpaceae as the sister group to Araucariaceae, diverging during the late Permian. Some fossils from the late Permian and Triassic (e.g., *Rissikia*) have been attributed to the family but cannot be unambiguously assigned. The oldest clear podocarp fossils come from the Jurassic, found in both hemispheres—*Scarburgia* and *Harrisiocarpus* from the Middle Jurassic of England, plus unnamed species from the Middle–Late Jurassic of Patagonia. Modern genera first appeared in the Early Cretaceous, and the family likely reached its diversity peak in the early Cenozoic.

**Genera** Relationships among genera are based on anatomical, biogeographical, morphological, and DNA evidence. The following lists include extant and extinct genera.