Diversity, Distribution, Systematics and Conservation Status of Podocarpaceae

Abstract

Among conifer families, Podocarpaceae is the second largest, with amazing diversity and functional traits, and it is the dominant Southern Hemisphere conifer family. However, comprehensive studies on diversity, distribution, systematic and ecophysiological aspects of the Podocarpaceae are sparse. We aim to outline and evaluate the current and past diversity, distribution, systematics, ecophysiological adaptations, endemism, and conservation status of podocarps. We analyzed data on the diversity and distribution of living and extinct macrofossil taxa and combined it with genetic data to reconstruct an updated phylogeny and understand historical biogeography. Podocarpaceae today contains 20 genera and approximately 219 taxa (201 species, 2 subspecies, 14 varieties and 2 hybrids) placed in three clades, plus a paraphyletic group/grade of four distinct genera. Macrofossil records show the presence of more than 100 podocarp taxa globally, dominantly from the Eocene–Miocene. Australasia (New Caledonia, Tasmania, New Zealand, and Malesia) is the hotspot of living podocarps diversity. Podocarps also show remarkable adaptations from broad to scale leaves, fleshy seed cones, animal dispersal, shrubs to large trees, from lowland to alpine regions and rheophyte to a parasite (including the only parasitic gymnosperm—Parasitaxus) and a complex pattern of seed and leaf functional trait evolution.

1. Introduction

Conifers are economically and ecologically important, form extensive forests in both Hemispheres and are currently the most diverse gymnosperms. There are seven conifer families (Araucariaceae, Cupressaceae, Pinaceae, Podocarpaceae, Sciadopityaceae, Cephalotaxaceae and Taxaceae), including 72 genera and approximately 702 species [1]. They are estimated to have evolved in the late Devonian from progymnosperms, and then dominated the Mesozoic Era [2,3,4]. Leslie et al. [5] investigated the evolutionary dynamics of conifers on a hemispheric scale based on molecular studies of 489 species and concluded that extant conifers have diverged in the Neogene with older splits in the Southern Hemisphere. Pinaceae and Cupressaceae have their main distribution in the temperate and subtropical regions of the Northern Hemisphere, while the Southern Hemisphere conifers are dominated by the Araucariaceae, Podocarpaceae and the Callitroideae of the Cupressaceae. The podocarps are monoecious and dioecious evergreen trees, shrubs, and subshrubs with mostly spirally arranged leaves and fleshy cones [6]. They are morphologically highly diverse [7,8]. Although the ecological and environmental variation (mostly rainforest and wet montane) is restricted, the morphological variation in leaves and seed cones is very high [9,10]. The extant and extinct taxa present in Australasia and South America show the wider distribution of the family across Gondwana in the past. Phylogenetically, the Podocarpaceae are closest to the Araucariaceae, Sciadopityaceae and Taxaceae [11,12]. Podocarps are significant both ecologically and economically and are a vital component of global forests and biodiversity [4,13]. The Podocarpaceae are very important from an evolutionary and systematic point of view due to their remarkable eco-physiological adaptations as compared to other conifer families, and they can provide us with valuable information on evolution and response to climate change [4,14]. Podocarps provide an exceptional opportunity for the understanding of comparative diversification processes, the evolution of different functional traits and ecophysiological adaptations. However, comprehensive studies are lacking on the different taxonomic, phylogenetic, ecological, biogeographic, and evolutionary aspects of most Podocarpaceae due to fragmented data on living species, sparse fossil records, difficulty in sampling and ecological data collection and less attraction compared to other conifer families. Many species described from Papua New Guinea, Malaysia, Indonesia, and New Caledonia are based on single records, and these areas remain under-explored. To evaluate these different aspects and several other key features, updated checklists for living and extinct taxa, phylogenetic analysis and ecophysiological adaptation research is required.

In this study, we evaluate the diversity, distribution, taxonomy, phylogeny, ecophysiology, and ecology of podocarps with the following aims: 1. To tabulate updated podocarp checklists for both extant and extinct taxa. 2. To reconstruct a new dated phylogeny of Podocarpaceae using relevant macrofossil records. 3. To assess the historical overview of taxonomic classifications of podocarps. 4. To discuss the diversity and historical biogeography. 5. To consider ecophysiological adaptations and threats.

2. Material and Methods

2.1. Phylogenetic Studies

A new, dated phylogenetic tree was produced for the Podocarpaceae. For phylogenetic analysis, DNA sequences of Podocarpaceae and Araucariaceae were sourced from GenBank (https://www.ncbi.nlm.nih.gov/genbank/, accessed on 25 January 2021). The sequences were cleaned and six markers, rbcL, matK, trnL-trnF, NEEDLY, PHYP and ITS species, were selected based on alignment confidence and availability of sequences. The final concatenated alignment consists of 190 taxa with 14 taxa as an out-group. The data were analyzed with BEAST version 2.6.3 at the CIPRES Science Gateway [15], set to run an uncorrelated lognormal relaxed clock and a GTR+I+G substitution model [16]. For calibration of the tree, we used the 20 oldest unequivocal macrofossil records (see supplementary file Table S1). Sixteen fossil constraints were assigned to the Podocarpaceae, two were assigned to the Araucariaceae and two were assigned to other conifers. The Fossil Birth–Death (FBD) model [17] was used as the tree prior, which imposes a time structure on the tree, while accounting for uncertainty in the placement of the fossil data by allowing all plausible placements for the fossil taxon on the extant tree [18].

2.2. Updated Checklist, Distribution, and IUCN Conservation Status of Podocarps

An updated checklist of all podocarp species was compiled based on the available literature [4], herbarium specimen observations and online databases, e.g., The Gymnosperm Database [19]; Global Biodiversity Information Facility-GBIF [20]; plants of the world online [21]; Australasian Virtual Herbarium-AVH [22]; Flora of China [23].

2.3. Distribution Data Analysis

The distribution of the species was analyzed in PC-ORD [24]. The Cluster Analysis (CA) and the Two-Way Cluster Analyses (TWCA) were used to identify significant and species-rich countries using Sorensen measures, based on presence/absence data.

2.4. Updated Checklist for Macrofossils of Extant Genera of Podocarpaceae

A checklist of fossil podocarps species belonging to extant genera was compiled using published literature and an online database Fossilworks [25].

3. Results

3.1. Phylogenetic Relationships

The fossil-calibrated phylogenetic tree under the FBD model indicates that the Podocarpaceae and Araucariaceae diverged around the early–mid Permian and the extant podocarp clades split during the mid–late Triassic. The extant podocarp genera have an estimated divergence time of the early Jurassic. The extant podocarp species predominantly show recent diversification from the Oligocene onwards. The phylogeny of the Podocarpaceae shows three major clades, I. Podocarpoid, II. Dacrydioid, III. Prumnopityoid, as well as a distinctive paraphyletic group/grade (Figure 1).

Figure 1.

The phylogenetic relationships and divergence time of the 20 extant podocarp genera within Podocarpaceae. Blue node bars indicate the 95% highest posterior density divergence time estimates for the corresponding node. Branch lengths are proportional to time (Ma, millions of years).

I. Podocarpoid clade—four genera, i.e., Afrocarpus, Nageia, Podocarpus and Retrophyllum. The suggested crown age for the Podocarpoid clade is approximately 75 Ma (54–85 Ma). The phylogeny also supports the split of Podocarpus into two subgenera i.e., Foliolatus and Podocarpus.

II. Dacrydioid clade—three genera, i.e., Dacrydium, Dacrycarpus and Falcatifolium. The suggested crown age for the Dacrydioid clade is approximately 75 Ma (54–95 Ma).

III. Prumnopityoid clade—nine genera, i.e., Lepidothamnus, Phyllocladus, Manoao, Lagarostrobos, Parasitaxus, Halocarpus, Sundacarpus, Pectinopitys and Prumnopitys. The newly dated phylogeny shows that the crown age for the Prumnopityoid clade is approximately 175 Ma (150–210 Ma).

IV. Paraphyletic group/grade—four genera, i.e., Acmopyle, Pherosphaera, Microcachrys and Saxegothaea.

3.2. Diversity at Genus Level and Distribution

Currently, in the Podocarpaceae, 20 genera and approximately 219 taxa (201 species, 2 subspecies, 14 varieties and 2 hybrids) are recognized (Table 1). Podocarpus is the most speciose genus with approximately 120 species distributed in approximately 70 countries. Two-way cluster analysis of Podocarpaceae species distribution shows five major groups, I. New Caledonian group, II. New Zealand group, III. Malesian group, IV. Southeast Asian group and V. Podocarpian group, widely distributed across several countries (Figure 2). Some of the widely distributed species are Afrocarpus gracilior (7 countries) and A. falcatus (5 countries), Dacrycarpus imbricatus (11 countries), Dacrydium elatum (7 countries), Dacrydium pectinatum (5 countries), Nageia wallichiana (11 countries), Podocarpus coriaceus (7 countries), P. guatemalensis (9 countries), P. milanjianus (15 countries), P. neriifolius (16 countries), P. oleifolius (11 countries), P. pilgeri (9 countries), P. polystachyus (7 countries) and Sundacarpus amarus (6 countries). The current species diversity and distribution is listed in Table 1 and summarized here:

Table 1.

An updated checklist of living podocarp taxa.

S#	Name	Synonyms	Common Name	Distribution in World	IUCN Status
1	Acmopyle pancheri (Brongn. and Gris) Pilg.	Acmopyle alba, Dacrydium pancheri, Nageia pancheri, Podocarpus pectinatus	New Caledonian acmopyle, Pancher’s acmopyle	New Caledonia	NT
2	Acmopyle sahniana Buchholz and N.E. Gray	Parasitaxus vodonaivalui	Fijian acmopyle	Fiji (Namosi and near Mt Tomanivi.)	CR
3	Afrocarpus dawei (Stapf) C.N.Page	Afrocarpus mannii subsp. dawei, Nageia mannii var. dawei, Podocarpus dawei, Podocarpus usambarensis var. dawei	-	Congo, Tanzania (Kagara and Mara provinces), Uganda	NT
4	Afrocarpus falcatus (Thunb.) C.N.Page	Afrocarpus falcatus subsp. gaussenii, Afrocarpus gaussenii, Decussocarpus falcatus, Nageia falcata, Nageia falcata var. gaussenii, Nageia meyeriana, Podocarpus falcatus, Podocarpus falcatus, Podocarpus gaussenii, Podocarpus gracillimus, Podocarpus meyerianus, Taxus falcata	Outeniqua yellowwood, Bastard Yellow wood	Malawi, Mozambique, South Africa (Eastern Cape Province, KwaZulu-Natal, Limpopo Province, Mpumalanga, Western Cape), Swaziland, Madagascar	LC
5	Afrocarpus gracilior (Pilg.) C.N.Page	Afrocarpus falcatus subsp. gracilior, Decussocarpus gracilior, Nageia falcata var. gracilior, Podocarpus gracilior	East African Yellow wood	Ethiopia, Kenya, Tanzania, Congo, Rwanda, South Sudan, Uganda	LC
6	Afrocarpus mannii (Hook.f.) C.N.Page	Decussocarpus mannii, Nageia mannii, Podocarpus mannii	-	Sao Tomé and Principe	VU
7	Afrocarpus usambarensis (Pilg.) C.N.Page	Afrocarpus mannii subsp. usambarensis, Nageia mannii var. usambarensis, Podocarpus usambarensis	African Yellowwood	Tanzania, Kenya (Kyulu Hills, Taita Taveta District)	EN
8	Dacrycarpus cinctus (Pilg.) de Laub.	Bracteocarpus cinctus, Bracteocarpus dacrydiifolius, Dacrycarpus dacrydiifolius, Podocarpus cinctus, Podocarpus dacrydiifolius	-	Indonesia (Maluku, Papua, Sulawesi), Malaysia (Sarawak), Papua New Guinea	LC
9	Dacrycarpus compactus (Wasscher) de Laub.	Bracteocarpus compactus, Podocarpus compactus	Binban Kadzinam, Kaibigl, Kaipik, Pau, Pawa, Uba, Umba, Umbwa	Indonesia and Papua New Guinea	LC
10	Dacrycarpus cumingii (Parl.) de Laub.	Bracteocarpus cumingii, Nageia cumingii, Podocarpus cumingii, Podocarpus imbricata var. cumingii, Podocarpus imbricatus var. cumingii	-	Indonesia (Sumatera), Malaysia (Sarawak), Philippines	LC
11	Dacrycarpus dacrydioides (A.Rich.) de Laub.	Dacrydium ferrugineum, Nageia dacrydioides, Nageia excesla, Podocarpus dacrydioides, Podocarpus thujoides	Kahikatea (Maori), White Pine	New Zealand	LC
12	Dacrycarpus expansus de Laub.	Bracteocarpus expansus	-	Indonesia (Papua), Papua New Guinea	LC
13	Dacrycarpus imbricatus (Blume) de Laub.	Bracteocarpus imbricatus, Bracteocarpus kawaii, Dacrycarpus imbricatus var. imbricatus, Dacrycarpus imbricatus var. patulus, Nageia cupressina, Podocarpus cupressinus, Podocarpus imbricatus, Podocarpus javanicus, Podocarpus kawaii, Thuja javanica	-	Cambodia, China (Guangxi, Hainan, Yunnan), Fiji, Indonesia (Jawa, Lesser Sunda Is., Papua, Sulawesi, Sumatera), Lao People’s Democratic Republic, Malaysia, Papua New Guinea (Bismarck Archipelago), Philippines, Thailand, Vanuatu, Vietnam	LC
14	Dacrycarpus imbricatus var. curvulus (Miq.) de Laub.	Podocarpus cupressina var. curvula, Podocarpus imbricatus var. curvula	Tjamarah	Indonesia (Sumatra and Java)	LC
15	Dacrycarpus imbricatus var. robustus de Laub.	Podocarpus papuanus, Podocarpus javanica, Podocarpus cupressina, Dacrycarpus papuana	Pierur, tupi, daru and umba	Papua New Guinea, Indonesia (Moluccas), Malaysia (Sarawak) and Philippines (Luzon, Mindanao)	LC
16	Dacrycarpus kinabaluensis (Wasscher) de Laub.	Bracteocarpus kinabaluensis, Podocarpus imbricatus var. kinabaluensis	-	Endemic to Mt. Kinabalu in Sabah (Borneo), Malaysia	LC
17	Dacrycarpus steupii (Wasscher) de Laub.	Bracteocarpus steupii, Podocarpus steupii	-	Indonesia (Kalimantan, Papua, Sulawesi), Papua New Guinea, Philippines	NT
18	Dacrycarpus vieillardii (Parl.) de Laub.	Dacrydium elatum var. compactum, Dacrydium elatum var. tenuifolium, Nageia tenuifolia, Nageia vieillardii, Podocarpus taxodioides var. tenuifolius, Podocarpus tenuifolius, Podocarpus vieillardii	-	New Caledonia	LC
19	Dacrydium araucarioides Brongn. and Gris	Athrotaxis araucarioides, Dacrydium arthrotaxoides, Metadacrydium araucarioides, Podocarpus araucarioides	-	New Caledonia	LC
20	Dacrydium balansae Brongn. and Gris	Metadacrydium balansae	-	New Caledonia	LC
21	Dacrydium beccarii Parl.	Nageia beccarii	-	Indonesia (Maluku, Papua, Sulawesi, Sumatera), Malaysia (Peninsular Malaysia, Sarawak), Papua New Guinea (Bismarck Archipelago), Philippines, Solomon Islands	LC
22	Dacrydium comosum Corner	Corneria comosa	-	Malaysia (Genting Highlands, Gunung Hulu Kali, Negeri Pahang)	EN
23	Dacrydium cornwallianum de Laub.	Corneria cornwalliana, Dacrydium nidulum var. araucarioides	-	Indonesia (Papua), Papua New Guinea	LC
24	Dacrydium cupressinum Sol. ex G.Forst.	Dacrydium cupressiforme, Thalamia cupressina	Rimu, Red pine	New Zealand	LC
25	Dacrydium elatum (Roxb.) Wall. ex Hook.	Corneria elata, Corneria pierrei, Dacrydium beccarii var. subelatum, Dacrydium junghuhnii, Dacrydium pierrei, Juniperus elata, Juniperus elatus, Juniperus philippsiana	Sempilor	Cambodia, Indonesia (Sumatera), Lao People’s Democratic Republic, Malaysia (Peninsular Malaysia, Sabah, Sarawak), Philippines, Thailand, Viet Nam	LC
26	Dacrydium ericoides de Laub.	Corneria ericoides	Sempilor, Bintulu	Malaysia (Borneo)	NA (IUCN) VU
27	Dacrydium gibbsiae Stapf	Corneria gibbsiae, Dacrydium beccarii var. kinabaluense	-	Malaysia (Mount Kinabalu in Sabah)	LC
28	Dacrydium gracile de Laub.	Corneria gracilis	-	Malaysia (Sabah, Sarawak)	NT
29	Dacrydium guillauminii J.Buchholz	Gaussenia guillauminii	-	New Caledonia (river Madeleine (Riviére des Lacs) and along the riverbanks of Lac en Huit)	CR
30	Dacrydium leptophyllum (Wasscher) de Laub.	Bracteocarpus leptophyllus, Corneria lepto. phylla, Dacrycarpus leptophyllus, Dacrydium leptophyllum, Podocarpus leptophylla, Podocarpus leptophyllus	-	Indonesia (Mt. Goliath, Papua)	VU
31	Dacrydium lycopodioides Brongn. and Gris	Gaussenia lycopodioides	Mou	New Caledonia (southern massif, from Mont Nekandi to Mont Dzumac and Mont Mou)	NT
32	Dacrydium magnum de Laub.	Corneria magna, Dacrydium beccarii var. rudens	-	Indonesia (Maluku), Papua New Guinea (Tagula Island, Normanby Island)	NT
33	Dacrydium medium de Laub.	Corneria media	Sangu, Gajo	Indonesia (Sumatera), Malaysia (Peninsular Malaysia)	VU
34	Dacrydium nausoriense de Laub.	Corneria nausoriensis	-	Fiji (Nausori Highlands, Sarava)	EN
35	Dacrydium nidulum de Laub.	Corneria nidula, Dacrydium nidulum var. vitiensis	-	Fiji, Indonesia (Lesser Sunda Is., Maluku, Papua, Sulawesi), Papua New Guinea	LC
36	Dacrydium novoguineense Gibbs	Corneria novoguineensis	-	Indonesia (Papua, Sulawesi), Papua New Guinea	LC
37	Dacrydium pectinatum de Laub.	Corneria pectinata, Dacrydium pectinatum var. robustum	-	Brunei Darussalam, China (Hainan), Indonesia (Kalimantan, Sumatera), Malaysia (Sabah, Sarawak), Philippines	EN
38	Dacrydium spathoides de Laub.	Corneria spathoides	-	Papua New Guinea (Irian Jaya), Indonesia	NT
39	Dacrydium × suprinii Nimsch	-	-	New Caledonia	NA
40	Dacrydium xanthandrum Pilg.	Corneria xanthandra	Kerapui	Indonesia (Kalimantan, Maluku, Papua, Sulawesi, Sumatera), Malaysia (Sabah, Sarawak), Papua New Guinea (Bismarck Archipelago, North Solomons), Philippines	LC
41	Falcatifolium angustum de Laub.	-	-	Malaysia (Sarawak)	EN
42	Falcatifolium falciforme (Parl.) de Laub.	Podocarpus falciformis, Nageia falciformis, Falcatifolium usan-apuensis, Falcatifolium falciforme var. usan-apuensis, Falcatifolium falciforme var. kinabaluensis, Falcatifolium falciforme var. kinabaluensis	-	Brunei Darussalam; Indonesia (Kalimantan); Malaysia (Peninsular Malaysia, Sabah, Sarawak)	NT
43	Falcatifolium gruezoi de Laub.	-	-	Indonesia (Maluku, Sulawesi), Philippines	NT
44	Falcatifolium papuanum de Laub.	Dacrydium papuanum	-	Papua New Guinea (Morobe)	LC
45	Falcatifolium sleumeri de Laub. and Silba	-	-	Indonesia (Papua)	NT
46	Falcatifolium taxoides (Brongn. and Gris) de Laub.	Dacrydium taxoides, Nageia taxoides, Pinus falciformis, Podocarpus taxodioides, Podocarpus taxodioides var. gracilis	-	New Caledonia	LC
47	Halocarpus bidwillii (Hook. f. ex Kirk) C.J.Quinn	Dacrydium bidwillii, Dacrydium bidwillii var. erectum, Dacrydium bidwillii var. relinatum	-	New Zealand (North Island, South Island, Stewart Island)	LC
48	Halocarpus biformis (Hook.) C.J.Quinn	Dacrydium biforme, Podocarpus biformis	Yellow pine, Pink pine, Bog Pine, Mountain Pine, Tarwood	New Zealand (North Island, South Island and Stewart Island)	LC
49	Halocarpus kirkii (F.Muell. ex Parl.) C.J.Quinn	Dacrydium kirkii	Monoao	New Zealand (Hokianga, Manukau Harbor and Coromandel Peninsula)	NT
50	Lagarostrobos franklinii (Hook.f.) Quinn	Dacrydium franklinii, Lepidothamnus franklinii	Huon Pine	Australia (Tasmania,)	LC
51	Lepidothamnus fonkii Phil.	Dacrydium fonckii, Dacrydium fonckii	Chilean Rimu	Argentina (Chubut, Santa Cruz); Chile (Aisén, Los Lagos, Magellanes)	LC
52	Lepidothamnus intermedius (Kirk) Quinn	Dacrydium intermedium	Yellow silver pine, Mountain pine	New Zealand (South Island, Stewart Island and North Island)	LC
53	Lepidothamnus laxifolius (Hook.f.) Quinn	Dacrydium laxifolium, Dacrydium laxifolium var. compactum, Dacrydium laxifolium var. debile	Mountain rimu, Pigmy pine, Pygmy pine	New Zealand (Tongariro National Park in the North Island southwards to Stewart Island)	LC
54	Manoao colensoi (Hook.) Molloy	Dacrydium colensoi, Dacrydium westlandicum, Lagarostrobos colensoi	Manoao, Silver pine, Westland pine, White silver pine	New Zealand (Lake Te Anau, Central Volcanic Plateau and Auckland)	LC
55	Microcachrys tetragona (Hook.) Hook.f.	Dacrydium tetragonu, Athrotaxis tetragona	Strawberry pine, Creeping Pine,	Australia (Tasmania)	LC
56	Nageia fleuryi (Hickel) de Laub.	Decussocarpus fleuryi, Podocarpus fleuryi	Kim giao	China (Guangdong, Guangxi, Yunnan), Laos, Vietnam	NT
57	Nageia formosensis (Dummer) C.N.Page	Decussocarpus nagi var. formosensis, Nageia nagi var. formosensis, Nageia nagi var. koshuensis, Nageia nankoensis, Podocarpus formosensis, Podocarpus formosensis var. koshuensis, Podocarpus koshunensis, Podocarpus nagi var. koshuensis	-	Taiwan	NA
58	Nageia maxima (de Laub.) de Laub.	Decussocarpus maximus, Podocarpus maxima, Podocarpus maximus	Landin paya	Malaysia (Sarawak)	EN
59	Nageia motleyi (Parl.) de Laub.	Agathis motleyi, Dammara motleyi, Decussocarpus motleyi, Nageia baccarii, Podocarpus beccarii, Podocarpus motleyi	Podo kebal musang, Kayu bawa, Setebal, Medang buloh	Brunei Darussalam, Indonesia (Kalimantan, Sumatera), Malaysia (Peninsular Malaysia, Sabah, Sarawak), Thailand	VU
60	Nageia nagi (Thunb.) Kuntze	Myrica nagi, Agathis veitchii, Dammara veitchii, Decussocarpus nagi, Nageia caesia, Nageia cuspidata, Nageia grandifolia, Nageia ovata, Podocarpus caesius, Podocarpus cuspidatus, Podocarpus grandifolius, Podocarpus nageia, Podocarpus nageia var. angustifolius, Podocarpus nageia var. rotundifolius, Podocarpus nagi, Podocarpus nagi var. angustifolius, Podocarpus nagi var. caesius, Podocarpus nagi var. ovatus, Podocarpus nagi var. rotundifolius, Podocarpus ovata, Podocarpus ovatus	Broad-leaved podocarp	Introduced China (Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Sichuan, Zhejiang), Japan (Honshu, Kyushu, Nansei-shoto, Shikoku), Taiwan (introduced), Vietnam (Lang Son)	NT
61	Nageia wallichiana (C.Presl) Kuntze	Decussocarpus wallichianus, Nageia blumei, Podocarpus blumei, Podocarpus wallichianus	Mala almaciga	Brunei Darussalam, Cambodia, China (Yunnan), India (Assam, Kerala, Nicobar Island), Indonesia (Jawa, Kalimantan, Maluku, Papua, Sulawesi, Sumatera), Laos, Malaysia (Peninsular Malaysia, Sabah, Sarawak), Myanmar, Papua New Guinea, Philippines, Thailand, Vietnam	LC
62	Parasitaxus usta (Vieill.) de Laub.	Dacrydium ustum, Nageia usta, Parasitaxus usta, Podocarpus ustus	Corail, Cèdre rabougri	New Caledonia [Grand Terre in the mountains of the far south (Dzumac, Montagne des Sources), central west (Paéoua and Tchingou) and far northeast (Colnett/Panie)]	VU
63	Pherosphaera fitzgeraldii (F.Muell.) Hook.f.	Dacrydium fitzgeraldii, Microstrobos fitzgeraldii	Dwarf mountain pine, Blue Mountain dwarf pine	Australia (New South Wales)	CR
64	Pherosphaera hookeriana W.Archer bis	Dacrydium hookerianum (W.Archer bis) Eichler, Microstrobos niphophilus J.Garden and L.A.S.Johnson, Pherosphaera niphophila (J.Garden and L.A.S.Johnson) Florin	Drooping pine, Mount Mawson Pine	Australia (Tasmania)	NT
65	Phyllocladus aspleniifolius (Labill.) Hook.f.	Brownetera aspleniifolia (Labill.) Tratt., Phyllocladus glaucus Carrière, Phyllocladus rhomboidalis Rich., Phyllocladus serratifolius Nois. ex Henkel and Hochst., Phyllocladus trichomanoides var. glaucus (Carrière) Parl., Podocarpus aspleniifolius Labill., Thalamia asplenifolia (Labill.) Spreng.	Celery top pine	Australia (Tasmania)	LC
66	Phyllocladus hypophyllus Hook.f.	Phyllocladus hypophyllus var. protractus Warb., Phyllocladus major Pilg., Phyllocladus protractus (Warb.) Pilg., Podocarpus hypophyllus (Hook.f.) Kuntze	Celery top pine	Philippines, Brunei, Malaysia (Celebes, Moluccas, Sulawesi), Papua New Guinea, Indonesia (Maluku)	LC
67	Phyllocladus toatoa Molloy	Phyllocladus glaucus Kirk	Toatoa (Maori), Blue celery pine	New Zealand (North Island)	LC
68	Phyllocladus trichomanoides D.Don	Phyllocladus cunninghamii, Podocarpus trichomanoides	Tanekaha (Maori), Celery pine	New Zealand (North Island and South Island)	LC
69	Phyllocladus trichomanoides var. alpinus (Hook.f.) Parl.	Phyllocladus alpinus, Phyllocladus aspleniifolius var. alpinus	Mountain toatoa	New Zealand	LC
70	Podocarpus acuminatus de Laub.	-	-	Brazil (Serra da Neblina in the state of Amazonas), Venezuela (Bolívar)	NT
71	Podocarpus acutifolius Kirk	Nageia acutifolia	Needle-leaved totara, Westland totara	New Zealand (South Island from Marlborough Sounds to S Westland)	LC
72	Podocarpus affinis Seem.	Nageia affinis	Kuasi	Fiji (Higher ridges on Viti Levu)	NT
73	Podocarpus angustifolius Griseb.	Podocarpus victorinianus, Podocarpus leonii, Podocarpus aristulatus	Sabina cimarrona	Cuba Cienfuegos (Sierra de Trinidad), Guantánamo (Sierra Maestra), Sancti Spíritus (Sierra de Sancti Spíritu), Holguín and Santiago de Cuba (Sierra de Nipe, Sierra del Cristal, and the Baracoa Ranges).	VU
74	Podocarpus annamiensis N.E.Gray	-	-	Myanmar, Vietnam, China	TH
75	Podocarpus aracensis de Laub. and Silba	-	-	Brazil [Amazonas (Serra Araca, Cerro Neblina] and Venezuela [Amazonas (Cerro Yaví)]	LC
76	Podocarpus archboldii N.E. Gray	Margbensonia archboldi, Podocarpus crassigemma	Baula, Jamekang, juba, Kaibigltuga, Morumba, Puling, Yamekange, Mu, Soa, Sarau	Indonesia (Papua), Papua New Guinea (Morobe District0	VU
77	Podocarpus aristulatus Parl.	Nageia aristulata, Podocarpus angustifolius var. wrightii		Cuba, Dominican Republic,	TH
78	Podocarpus atjehensis (Wasscher) de Laub.	Podocarpus neriifolius var. atjehensis, Margbensonia atjehensis, Margbensonia atjehense, Podocarpus neriifolius var. membranaceaus	Atjeh	Malesia [Sumatera (Aceh, Gajo Lands, Kemiri and Bandahara), New Guinea (Papua, Wissel Lakes).	NT
79	Podocarpus barretoi de Laub. and Silba	-	-	Brazil (Minas Gerais)	NT
80	Podocarpus beecherae de Laub.	-	-	New Caledonia	TH
81	Podocarpus borneensis de Laub.	Podocarpus polystachyus var. rigidus	Bisit, Bubung, Buloh	Indonesia (Kalimantan), Malaysia (Sabah, Sarawak)	LC
82	Podocarpus bracteatus Blume	Nageia bracteata, Podocarpus bracteatus var. brevipes, Podocarpus neriifolius var. bracteatus, Podocarpus neriifolius var. brevipes	Kayu unung unung, Toba Batak, Bima, Kimarak, Kipantjar, Ki putri	Indonesia (Jawa, Lesser Sunda Is., Sumatera)	LC
83	Podocarpus brasiliensis de Laub.	Podocarpus barretoi	-	Brazil (Bahia, Brasília Distrito Federal, Goiás, Mato Grosso, Minas Gerais, Roraima), Venezuela	LC
84	Podocarpus brassii Pilg.	-	Baugwa, Baula, Chuga, Kaibigltuga, Kaipil, Karbuku, Maja, Mbagua, Tsula	Indonesia (Papua), Papua New Guinea	LC
85	Podocarpus brassii var. humilis de Laub.	Podocarpus brassii subsp. humilis	-	Papua New Guinea	LC
86	Podocarpus brevifolius (Stapf) Foxw.	Podocarpus neriifolius	-	Malaysia (Sabah)	NT
87	Podocarpus buchholzii de Laub.	Podocarpus buchholzii var. neblinensis Silba, Podocarpus buchholzii subsp. neblinensis (Silba) Silba	-	Venezuela (Guyana Highlands)	LC
88	Podocarpus buchii Urb.	Podocarpus aristulatus var. buchii, Podocarpus angustifolius Griseb. ssp. buchii	Tachuela, Chicharrón, Palo de Cruz	Dominican Republic (Southeast Haiti)	EN
89	Podocarpus capuronii de Laub.	Podocarpus capuronii var. capuronii, Podocarpus woltzii	-	Madagascar (Itremo Massif, Manandona)	EN
90	Podocarpus celatus de Laub.	-	Cinqui-mase	Bolivia (Potosi), Brazil (Goiás, Mato Grosso), Colombia, Ecuador, Peru (Junin, Loreto, Montana, Puno), Venezuela (Amazonas, Bolivar, Tachira)	LC
91	Podocarpus chinensis Wall. ex J.Forbes	Podocarpus macrophyllus var. maki, Podocarpus japonicus, Podocarpus makoyi, Podocarpus appressus, Podocarpus macrophyllus subsp. maki, Podocarpus maki, Nageia appressa, Nageia japonica, Nageia chinensis, Nageia macrophylla var. maki, Myrica esquirolii	-	China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Yunnan, and Zhejiang), Myanmar, Japan	LC
92	Podocarpus chinensis var. wardii de Laub. and Silba	Podocarpus chinensis subsp. wardii	-	Myanmar (N’mai Hka Valley)	LC
93	Podocarpus chingianus S.Y.Hu	Podocarpus macrophyllus var. chingiiI, Margbensonia chingiana	Zhu guan luo han song	China (Jiangsu, Zhejiang, Sichuan)	DD
94	Podocarpus confertus de Laub.	Podocarpus neriifolius var. penibukanensis	-	Malaysia (Sabah, Sarawak)	EN
95	Podocarpus coriaceus Rich. and A.Rich.	Nageia coriacea, Taxus lancifolia, Podocarpus coriaceus var. sulcatus	Yucca plum pine, Resinier moutaigue, Caoba del país	Dominican Republic, Guadeloupe, Martinique, Montserrat, Puerto Rico, Saint Kitts and Nevis, Saint Lucia, Trinidad, and Tobago	LC
96	Podocarpus costalis C.Presl	Nageia costalis, Podocarpus costalis var. taiwanensis	Lan yu luo han song, Arius	Philippines, Taiwan, China	EN
97	Podocarpus costaricensis de Laub.	-	Cipresillo	Costa Rica (San José)	CR (VU)
98	Podocarpus crassigemma de Laub.	Podocarpus crassigemmis	A-pul, Kaboga, Morumba, Baula, Iamuka, Jamekang, Juba, Kamga, Puling, Kabor, Kabiltugl, Kaibelparu, Kkaibig, Kaip, Nonofan, Ronohanini, Sula	Indonesia (Papua), Papua New Guinea (Bismarck Archipelago, Central highlands)	LC
99	Podocarpus decipiens N.E.Gray	-	-	Fiji (Viti Levu)	NA
100	Podocarpus decumbens N.E.Gray	-	-	New Caledonia (Grande Terre, Southern mountains)	CR
101	Podocarpus deflexus Ridl.	-	-	Indonesia (Sumatera), Malaysia (Peninsular Malaysia)	EN
102	Podocarpus degeneri (N.E.Gray) de Laub.	Margbensonia degeneri, Podocarpus neriifolius var. degeneri	-	Fiji	LC
103	Podocarpus dispermus C.T.White	Margbensonia disperma	Broad-leaved brown pine	Australia (Queensland)	LC
104	Podocarpus drouynianus F.Muell.	Nageia drouyniana, Margbensonia drouyniana, Podocarpus brownii	Emu Berry	Australia (Western Australia)	LC
105	Podocarpus ekmanii Urb.	-	Sabina Cimarrona	Cuba (Sierra del Cristal, Sierra de Moa and Sierra de Nipe)	LC
106	Podocarpus elatus R.Br. ex Endl	Margbensonia elata, Nageia elata	Illawarra plum, Brown pine, Plum pine, Turpentine pine, Yellow pine, Australian plum, White plum, Goongum, Native deal, Pencil cedar	Australia (New South Wales, Queensland)	LC
107	Podocarpus elongatus (Aiton) L’Hér. ex Pers.	Taxus elongatus, Taxus elongata, Taxus capensis, Nageia elongata, Podocarpus thunbergii var. angustifolia	Breede river yellowwood	Malawi, South Africa (Northern Cape Province, Western Cape), Zambia, Zimbabwe	LC
108	Podocarpus fasciculus de Laub.	Podocarpus macrophyllus var. liukiuensis, Podocarpus macrophyllus f. grandifolia	-	Japan (Nansei-shoto), Taiwan	VU
109	Podocarpus forrestii Craib and W.W.Sm.	Margbensonia forrestii, Podocarpus macrophyllus subsp. forrestii	-	China	VU
110	Podocarpus gibbsiae N.E.Gray	-	-	Malaysia (Endemic to Mt. Kinabalu in Sabah)	VU
111	Podocarpus glaucus Foxw.	-	Nipa	Indonesia (Maluku, Papua, Sulawesi), Papua New Guinea (Bismarck Archipelago), Philippines, Solomon Islands	LC
112	Podocarpus globulus de Laub.	-	Sapiro	Malaysia (Sabah, Sarawak)	EN
113	Podocarpus glomeratus D.Don	Nageia glomerata, Podocarpus rigidus, Podocarpus cardenasii	Pino de Monte, Intimpa, Huampo	Bolivia, Ecuador, Peru	NT
114	Podocarpus gnidioides Carrière	Nageia gnidioides, Podocarpus alpinus var. arborescens, Podocarpus alpinus var. caespitosus, Podocarpus caespitosus, Podocarpus gnidioides subsp. caespitosus	-	New Caledonia	NT
115	Podocarpus grayae de Laub.	-	Brown pine, Northern brown pine; Brown pine; Weeping brown pine	Australia (Northern Territory, Queensland)	LC
116	Podocarpus guatemalensis Standl.	Podocarpus allenii, Podocarpus guatemalensis var. allenii, Podocarpus guatemalensis subsp. allenii, Podocarpus guatemalensis subsp. pinetorum, Podocarpus guatemalensis var. pinetorum, Podocarpus pinetorum	Ocotillo de Llano, Alfajillo, Ciprecillo Amarillo, Ciprecillo Blanco, Cipresillo, Cypress de Montaña, Palo de Oro, Pinillo, Piño de Montaña	Belize, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico (Oaxaca, Veracruz), Nicaragua, Panama	LC
117	Podocarpus henkelii Stapf ex Dallim. and B.D.Jacks.	Podocarpus ensiculus, Podocarpus henkelii subsp. Ensiculus, Podocarpus thunbergii var. falcata	Henkel’s yellowwood, Falcate yellowwood, East griqualand yellowwood, Natal yellowwood, bastergeelhout, Nanjula	Malawi, South Africa (Eastern Cape Province, KwaZulu-Natal), Tanzania, Zimbabwe	CR
118	Podocarpus hispaniolensis de Laub.	-	-	Dominican Republic (Cordillera Central, San José de Ocoa, Cordillera Septentriona, Province Puerto Plata)	EN
119	Podocarpus hookeri de Laub.	Podocarpus neriifolius var. linearis, Podocarpus neriifolius var. staintonii	-	India (Sikkim), Indonesia (Sumatra, Java, Borneo), Philippines	LC
120	Podocarpus humbertii de Laub.	-	-	Madagascar (Mont Anjanaharibe, Mont Tsaratanana and Mont Marojezy)	EN
121	Podocarpus insularis de Laub.	-	Dala, tunum, ida-ayebo	Papua New Guinea (Bismarck Archipelago), Solomon Islands, Vanuatu	LC
122	Podocarpus idenburgensis N.E.Gray	-	-	Papua New Guinea, Fiji	NE
123	Podocarpus laetus Hooibr. ex Endl.	Podocarpus cunninghamii, Podocarpus hallii, Nageia hallii, Podocarpus totara var. hallii	Montane totara, Thin-bark totara, Hall’s totara	New Zealand (North Island, Tongariro National Park, South Island and Stewart Island)	LC
124	Podocarpus lambertii Klotzsch ex Endl.	Nageia lambertii, Podocarpus lambertii subsp. horsmanii, Podocarpus lambertii var. horsmanii, Podocarpus lambertii subsp. tigreensis, Podocarpus lambertii var. tigreensis	Pinheiro bravo	Argentina (Misiones), Brazil (Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo)	NT
125	Podocarpus laminaris de Laub.	Podocarpus rubens subsp. pabinamaensis, Podocarpus rubens var. pabinamaensis	-	Papua New Guinea	NA
126	Podocarpus latifolius (Thunb.) R.Br. ex Mirb.	Nageia latifolia, Nageia thunbergii, Podocarpus latifolius var. latior, Podocarpus latifolius subsp. latior, Podocarpus latior, Podocarpus nobilis, Podocarpus pinnata, Podocarpus thunbergii, Taxus latifolia	Broad-leaved yellowwood, Real yellowwood, True yellowwood, Upright yellowwood	Eswatini, South Africa (Eastern Cape Province, Free State, KwaZulu-Natal, Limpopo Province, Mpumalanga, Northern Cape Province, Western Cape)	LC
127	Podocarpus laubenfelsii Tiong	-	-	Indonesia (Kalimantan), Malaysia (Sabah, Sarawak)	EN
128	Podocarpus lawrencei Hook.f.	Podocarpus alpinus var. lawrencei, Podocarpus lawrencei subsp. errinundraensis	Mountain plum pine, Plum pine	Australia (Australian Capital Territory, New South Wales, Tasmania, Victoria)	LC
129	Podocarpus ledermannii Pilg.	Podocarpus idenburgensis	-	Indonesia (Papua), Papua New Guinea (Bismarck Archipelago)	LC
130	Podocarpus ledermannii Pilg. var. expansus de Laub.	-	-	Indonesia	LC
131	Podocarpus lenticularis de Laub.	-	-	Assam (India), Laos	NA
132	Podocarpus linearis de Laub.	-	-	Papua New Guinea	DD
133	Podocarpus levis de Laub.	-	Marisa, Sanru, Kayu tjina, Wasiwarare	Indonesia (Kalimantan, Maluku, Papua, Sulawesi)	LC
134	Podocarpus × loderi Cockayne	-	-	New Zealand	NA
135	Podocarpus longifoliolatus Pilg.	Podocarpus longifoliolatus	-	New Caledonia (Grande Terre)	EN
136	Podocarpus lophatus de Laub.	-	-	Philippines (Mt. Tapulao in Luzon and Mt. Halcon in Mindoro)	VU
137	Podocarpus lucienii de Laub.	-	-	New Caledonia (Massif du Colnett and the Massif du Tchingou)	LC
138	Podocarpus macrocarpus de Laub.	-	Malakawayan	Philippines (Luzon)	EN
139	Podocarpus macrophyllus (Thunb.) Sweet	Margbensonia forrestii, Margbensonia macrophylla, Nageia macrophylla, Nageia macrophylla, Podocarpus forrestii, Podocarpus macrophyllus var. angustifolius, Podocarpus macrophyllus subsp. angustifolius, Podocarpus macrophyllus f. angustifolius, Podocarpus macrophyllus subsp. forrestii, Podocarpus macrophyllus var. macrophyllus, Podocarpus macrophyllus var. rubra, Podocarpus verticillatus, Taxus macrophylla, Taxus makoya	Southern yew, Yew podocarp, Long-leaved podocarp, Buddhist pine, Kusamaki, Inumaki, luo han song	China (Anhui, Chongqing, Fujian, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Yunnan, Zhejiang), Hong Kong, Japan (Honshu, Kyushu, Shikoku), Taiwan, Malaysia, Singapore	LC
140	Podocarpus macrophyllus var. piliramulus Zhi X. Chen and Zhen Q. Li	-	-	China (Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Yunnan, Zhejiang); Hong Kong; Japan (Honshu, Kyushu, Shikoku); Myanmar; Taiwan	NT
141	Podocarpus madagascariensis Baker	Nageia madagascariensis, Podocarpus madagascariensis var. madagascariensis	-	Madagascar	NT
142	Podocarpus madagascariensis var. procerus de Laub.	Podocarpus madagascariensis subsp. procerus	-	Madagascar (Tolanaro [Fort Dauphin] and Massif de Bekolosy).	EN
143	Podocarpus madagascariensis var. rotundus L.Laurent	Podocarpus madagascariensis subsp. rotundus (L. Laurent) Silba	-	Madagascar (Massif de Bekolosy and the Massif du Manongarivo)	DD
144	Podocarpus magnifolius J.Buchholz and N.E. Gray	-	Cinqui-masé	Bolivia (La Paz), Colombia, Panama, Peru (Pasco, Oxapampa), Venezuela (States of Bolívar, Amazonas, Aragua, Yaracuy)	LC
145	Podocarpus marginalis de Laub.	-	-	Papua New Guinea	DD
146	Podocarpus matudae Lundell	Podocarpus reichei, Podocarpus matudae var. reichei, Podocarpus matudae var. macrocarpus, Podocarpus matudae var. jaliscanus, Podocarpus matudae subsp. jaliscanus, Podocarpus matudae subsp. macrocarpus, Podocarpus matudae subsp. reichei	Sabino	Mexico (Chiapas, Jalisco, Michoacán, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tamaulipas, Veracruz), El Salvador, Guatemala (Huehuetenango), Honduras	CR
147	Podocarpus matudae subsp. jaliscanus (de Laub. and Silba) Silba	Podocarpus matudae Lundell var. jaliscanus	-	Mexico (Jalisco)	VU
148	Podocarpus micropedunculatus de Laub.	-	Kayu china, kayu tjina	Brunei Darussalam, Indonesia, Malaysia (Sabah, Sarawak)	NT
149	Podocarpus milanjianus Rendle	Podocarpus ulugurensis	Lusamina	Angola, Burundi, Cameroon, Congo, Congo, Kenya, Malawi, Mozambique, Nigeria, Rwanda, Sudan, Tanzania, Uganda, Zambia, Zaire, Zimbabwe	LC
150	Podocarpus nakaii Hayata	Podocarpus macrophyllus var. nakaii	Nakai podocarp, Nakai yellowwood	Taiwan (Chianghua Co., Nantou Co., Taichung Co.)	EN
151	Podocarpus neriifolius D.Don	Nageia discolor, Nageia leptostachya, Nageia neriifolia, Nageia neglecta, Nageia decipiens, Nageia polyantha, Nageia annamiensis, Podocarpus discolor, Podocarpus leptostachya, Podocarpus annamiensis, Podocarpus epiphyticus, Podocarpus neglecta, Podocarpus junghuhniana, Podocarpus thailandensis, Podocarpus neriifolius var. polyanthus	Brown pine, Podo bukit, Ambai Ayam, Hatang, Hai nan luo han song, Thông tre, Thông lông gà	Brunei Darussalam, Cambodia, China (Guangxi, Yunnan), Fiji, India (Assam, West Bengal), Bangladesh, Indonesia (Bali, Jawa, Kalimantan, Lesser Sunda Is., Maluku, Papua, Sulawesi, Sumatera), Lao People’s Democratic Republic, Malaysia (Peninsular Malaysia, Sabah, Sarawak); Myanmar, Nepal, Papua New Guinea (Bismarck Archipelago), Philippines, Solomon Islands, Thailand, Vietnam	LC
152	Podocarpus neriifolius D.Don var. degeneri N.E.Gray	-	-	Fiji (Vanua Leva, Viti Levu)	LC
153	Podocarpus nivalis Hook.	Nageia nivalis, Podocarpus nivalis var. erectus, Podocarpus montanus	Alpine totara, Snow totara	New Zealand (North Island, South Island)	LC
154	Podocarpus novae-caledoniae Vieill. ex Brongn. and Gris	Nageia novae-caledoniae, Podocarpus beecherae, Podocarpus rivularis	-	New Caledonia (Grande Terre, Iledes Pins)	LC
155	Podocarpus neglectus Blume	Podocarpus discolor, Podocarpus leptostachyus, Podocarpus thailandensis, Nageia neglecta, Podocarpus discolor, Podocarpus junghuhnianus	-	Thailand, Indonesia, China (Hainan), Indonesia (Borneo, Java, Sumatra), Malaysia	NA
156	Podocarpus novoguineensis de Laub.	-	-	Papua New Guinea	NA
157	Podocarpus nubigenus Lindl.	Nageia nubigena, Saxegothaea gracilis	Huililahuani, Mañio, Mañío de Hojas Punzantes, Mañio Hembra, Mañio Macho, Mañiu de la Costa, Pino Amarillo	Argentina (Neuquén, Santa Cruz), Chile (Aisén, La Araucania, Los Lagos, Magellanes)	NT
158	Podocarpus oblongus de Laub.	Podocarpus rumphii subsp. Aruensis, Podocarpus rumphii var. aruensis	-	Papua New Guinea (Vogelkop)	DD
159	Podocarpus oleifolius D.Don	Nageia macrostachya, Nageia oleifolia, Podocarpus ballivianensis, Podocarpus ingensis, Podocarpus macrostachys, Podocarpus oleifolius subsp. equadorensis, Podocarpus oleifolius var. equadorensis, Podocarpus oleifolius var. macrostachys, Podocarpus oleifolius var. trujillensis, Podocarpus oleifolius subsp. trujillensis	Pino de pasto, Pinete	Bolivia, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico (Chiapas, Guerrero, Michoacán, Oaxaca, Puebla, Veracruz), Panama, Peru, Venezuela	LC
160	Podocarpus oleifolius subsp. costaricensis (J.Buchholz and N.E.Gray) Silba	-	-	Costa Rica, El Salvador, Guatemala, Honduras, Mexico Gulf, Mexico Southeast, Mexico Southwest, Nicaragua, Panamá	LC
161	Podocarpus orarius R.R.Mill and M.Whiting	Podocarpus spathoides subsp. solomonensis, Podocarpus spathoides var. solomonensis	-	Papua New Guinea (Solomon Island), Vanuatu	NT
162	Podocarpus palawanensis de Laub. and Silba	-	-	Philippines (Palawan)	CR
163	Podocarpus pallidus N.E.Gray	-	Uhiuhi	Tonga (island of Eua and islands of Vava’u)	VU
164	Podocarpus parlatorei Pilg.	Podocarpus angustifolius, Nageia angustifolia	Pino Blanco, Pino del Cerro	Argentina (Catamarca, Corrientes, Jujuy, Salta, Tucumán), Bolivia (Chuquisaca, Cochabamba, Potosí, Santa Cruz, Tarija, La Paz), Peru (Sierra de Chaglla)	NT
165	Podocarpus pendulifolius J.Buchholz and N.E.Gray	-	Pino Carbón, Pino Hayuco	Venezuela (Andes, Cordillera do Merida, Edo Lara, Merida, Tachira and Trujillo)	EN
166	Podocarpus perrieri Gaussen and Woltz	Podocarpus rostratus var. perrieri, Podocarpus rostratus subsp. perrieri	-	Madagascar (Andringitra Massif, Toamasina, Forêt d’Andasibé)	CR
167	Podocarpus pilgeri Foxw.	Podocarpus celebicus, Podocarpus pilgeri var. thailandensis, Podocarpus pilgeri subsp. wangii, Podocarpus schlechteri, Podocarpus wangii, Podocarpus tixieri	-	Cambodia (Kampuchea), China (Guangdong, Guangxi, Hainan, Yunnan), Indonesia (Maluku, Papua, Sulawesi), Laos, Malaysia (Sarawak), Papua New Guinea (Bismarck Archipelago); Philippines, Thailand, Vietnam	LC
168	Podocarpus polyspermus de Laub.	-	Mé Maoya podocarp	New Caledonia (Grande Terre)	EN
169	Podocarpus polystachyus R.Br. ex Endl.	Margbensonia polystachya, Nageia polystachya	Jati bukit, Kayu karamat, Podo laut, Mayu serai, Landin, Kandabang, kayu china, Saumah	Brunei Darussalam, Indonesia (Kalimantan, Maluku, Papua, Sumatera), Malaysia (Peninsular Malaysia, Sabah, Sarawak), Papua New Guinea, Philippines, Singapore, Thailand	VU
170	Podocarpus pseudobracteatus de Laub.	Podocarpus archboldii var. crassiramosus	Kaip, Kebu, Puling	Indonesia (Papua), Papua New Guinea	LC
171	Podocarpus pseudobracteatus de Laub. var. sicaris de Laub.	-	-	Papua New Guinea	LC
172	Podocarpus purdieanus Hook.	Nageia purdieana	Yacca, St. Ann Yacca	Jamaica (Claredon, St. Catherine, St. Ann, Trelawny, Sanders Hill, Mt. Diablo)	EN
173	Podocarpus ramosii R.R.Mill	Podocarpus rotundus	-	Indonesia (Kalimantan Timur), Philippines (Mt. Banahao in Luzon)	DD
174	Podocarpus ridleyi (Wasscher) N.E. Gray	Margbensonia ridleyi, Podocarpus neriifolius var. ridleyi	-	Malaysia (Peninsular Malaysia)	VU
175	Podocarpus roraimae Pilg.	Podocarpus buchholzii, Podocarpus buchholzii var. neblinensis, Podocarpus buchholzii subsp. neblinensis	Ai-yek	Guyana (Region of Cuyuni-Mazaruni on Mt. Roraima), Venezuela (Amazonas, Bolívar)	LC
176	Podocarpus rostratus J.Laurent	-	-	Madagascar (Antsiranana, Fianarantsoa, Mahajanga and Toamasina Provinces)	EN
177	Podocarpus rubens de Laub.	Podocarpus indonesiensis, Podocarpus rubens subsp. sumatranus, Podocarpus rubens var. sumatranu, Podocarpus rubens var. pabinamaensis, Podocarpus neriifolius var. timorensis	Bebi-è, Ungpop, Bin, Kaip, Nelil, Sukou	Indonesia (Maluku, Flores, Borneo, Sulawesi, Sumatera), Malaysia (Sabah), Papua New Guinea (Bismarck Archipelago, Papuasia, New Britain), Timor-Leste	LC
178	Podocarpus rumphii Blume	Cerbera nereifolia, Margbensonia koordersii, Margbensonia philippinensis, Margbensonia rumphii, Nageia rumphii, Podocarpus koordersii, Podocarpus philippinensis, Podocarpus rumphii subsp. arbainii, Podocarpus rumphii var. arbainii, Podocarpus sprengelii, Podocarpus rumphii var. aruensis	Nimsal	China (Hainan), Indonesia (Jawa, Lesser Sunda Island, Maluku, Papua, Sulawesi), Malaysia (Peninsular Malaysia, Sabah), Papua New Guinea (Bismarck Archipelago), Philippines (Luzon)	NT
179	Podocarpus rusbyi J.Buchholz and N.E.Gray	-	Pino Blanco, Pino del Monte	Bolivia (Cochabamba, La Paz, Santa Cruz), Peru (Cusco and near Machu Pichu)	VU
180	Podocarpus salicifolius Klotzsch and H.Karst. ex Endl.	Nageia salicifolia, Podocarpus pittieri	Pinabete	Venezuela, Brazil, Bolivia, Colombia, Peru	LC
181	Podocarpus salignus D. Don	Nageia chilina, Podocarpus chilinus, Podocarpus chilinus var. glaucus	Willow-leaf podocarp, Mañio, Mañío de Hojas Largas, Mañio de Hojas Punzantes, Mañio Hembra, Mañio Macho, Manique, Pino Amarillo	Chile (Biobío, La Araucania, Los Lagos, Maule)	VU
182	Podocarpus salomoniensis Wasscher	-	Dengali tolo	Solomon Islands (San Cristobal Island and San Jorge Island)	EN
183	Podocarpus sellowii Klotzsch ex Endl.	Nageia sellowii	-	Brazil (Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo)	EN
184	Podocarpus sellowii Klotzch ex Endl. var. angustifolius	-	-	Brazil (Rio de Janeiro)	CR
185	Podocarpus smithii de Laub.	-	Smith’s pine, Brown pine	Australia (Queensland)	LC
186	Podocarpus spathoides de Laub.	Podocarpus spathoides de Laub. var. solomonensis	-	Malaysia (Peninsular Malaysia, Sarawak, Maluku), Papua New Guinea (Solomon Islands)	DD
187	Podocarpus spinulosus (Sm.) R.Br. ex Mirb.	Margbensonia spinulosa, Nageia ensifolia, Nageia laeta, Nageia spinulosa, Podocarpus bidwillii, Podocarpus ensifolius, Podocarpus laetus, Podocarpus pungens, Taxus spinulosa	-	Australia (New South Wales, Queensland)	LC
188	Podocarpus sprucei Parl.	Nageia sprucei	Guabisay, Romerillo	Ecuador, Peru (Piura)	EN
189	Podocarpus steyermarkii J.Buchholz and N.E.Gray	-	-	Guyana (Pakaraima Mountains), Venezuela [Bolivar (Carrao-tepui, Uaipan-tepui, Cerro Jaua), Amazonas (Neblina Massif)]	LC
190	Podocarpus subtropicalis de Laub.	Podocarpus subtropicalis var. medogensis, Podocarpus subtropicalis subsp. medogensis	-	China (Sichuan, Yunnan)	DD
191	Podocarpus sylvestris J.Buchholz	Podocarpus colliculatus, Podocarpus novae-caledoniae var. colliculatus	-	New Caledonia (Grande Terre, Ile des Pins)	LC
192	Podocarpus tepuiensis J.Buchholz and N.E.Gray	-	-	Ecuador (Cordillera del Condo), Venezuela (Bolivar, Amazonas)	LC
193	Podocarpus teysmannii Miq.	Nageia teysmannii, Podocarpus epiphyticus, Podocarpus neriifolius var. polyanthus	Kalek rotan, Sikuju	Myanmar, Indonesia (Sumatera), Malaysia (Peninsular Malaysia, Sabah), Brunei Darussalam	NT
194	Podocarpus thevetiifolius Zipp. ex Blume	Margbensonia thevetiifolia, Nageia thevetiifolia, Podocarpus polystachyus subsp. thevetiifolius, Podocarpus polystachyus var. thevetiifolius	-	Papua New Guinea	NA
195	Podocarpus totara G.Benn. ex D.Don	Nageia totara, Podocarpus totara var. waihoensis, Podocarpus totara subsp. waihoensis	Totara	New Zealand (North Island and South Island)	LC
196	Podocarpus totara var. waihoensis Wardle	-	Totara, Westland totara	New Zealand (West Coast of the South Island)	NT
197	Podocarpus transiens (Pilg.) de Laub.	Podocarpus lambertii var. transiens, Podocarpus transiens var. harleyi, Podocarpus transiens subsp. harleyi	-	Brazil (Bahia, Goiás, Minas Gerais, Paraná, Santa Catarina)	EN
198	Podocarpus trinitensis J.Buchholz and N.E.Gray	-	-	Trinidad and Tobago (El Tucuche)	LC
199	Podocarpus urbanii Pilg.	-	Blue mountain yacca, Yacca	Jamaica (St. Andrew, Portland and St. Thomas within the Blue and John Crow Mountains)	CR
200	Podocarpus vanuatuensis de Laub.	-	-	Vanuatu	DD
201	Podocarpus victorinianus Carabia	Podocarpus leonii	-	Cuba	NE
202	Prumnopitys andina (Poepp. ex Endl.) de Laub.	Nageia andina, Nageia valdiviana, Podocarpus andinus, Podocarpus spicatus, Podocarpus valdivianus, Prumnopitys andina subsp. blijdensteinii, Prumnopitys elegans, Prumnopitys spicata, Stachycarpus andinus	Lleuque, Llleuqui, Uva de la Cordillera	Chile (Biobío, La Araucania, Maule), Argentina (Neuquen)	VU
203	Prumnopitys taxifolia (Sol. ex D.Don) de Laub.	Dacrydium mai, Dacrydium taxifolium, Nageia spicata, Podocarpus spicatus, Stachycarpus spicatus	Matai, Black pine	New Zealand (North Island and South Island)	VU
204	Prumnopitys montana (Humb. and Bonpl. ex Willd.) de Laub.	Botryopitys densifolia, Botryopitys meridensis, Botryopitys montana, Dacrydium distichum, Nageia montana, Podocarpus curvifolius, Podocarpus humboldtii, Podocarpus montanus var. densifolius, Podocarpus montanus var. diversifolius, Podocarpus montanus var. meridensis, Podocarpus taxifolius, Podocarpus taxifolius var. communis, Podocarpus taxifolius var. densifolius, Stachycarpus meridensis, Stachycarpus taxifolius, Taxus montana, Torreya montana	-	Bolivia (Cochabamba), Colombia (Belmira, San Andres, Arabuca, Villa de Leiva, Pensilvania, Cauca, Cesar, La Guajira, Magdalena, Quindío, Risarald, Tolima), Ecuado (Azuay, Cañar, Loja, Morona-Santiago, Zamora-Chinchipe), Peru (Cajamarca, Junín, Pasco, San Martín), Venezuela (Lara, Tachira, Zulia)	VU
205	Pectinopitys exigua (De Laub.) C.N.Page	-	Pino colorado, Jatun pino, Pino castilla, Pino negro	Bolivia (Cochabamba, Chuquisaca, Santa Cruz)	NT
206	Pectinopitys ferruginea (G.Benn. ex D.Don in Lamb.) C.N.Page	Nageia ferruginea, Podocarpus ferrugineus, Stachycarpus ferrugineus, Stachypitys ferruginea	Miro, Brown pine	New Zealand (North Island, South Island and Stewart Island)	LC
207	Pectinopitys ferruginoides (R.H.Compton) C.N.Page	Podocarpus distichus, Podocarpus distichus var. maialis, Podocarpus ferruginoides, Stachycarpus distichus, Stachycarpus ferruginoides, Stachypitys disticha, Stachypitys ferruginoides	-	New Caledonia	LC
208	Pectinopitys harmsiana (Pilg.) C.N.Page	Podocarpus harmsianus, Podocarpus utilior, Prumnopitys utilior	Uncumanu, Yellow miro	Bolivia (Abel Iturralde, Franz Tamayo, Sud Yungas), Colombia (Cauca, Quindío, Risaralda, Tolima, Sierra Nevada de Santa Marta), Ecuador (Loja), Peru (Ayacucho, Cajamarca, Cusco, Junín, Pasco, Piura, San Martín), Venezuela (Vargas, Miranda, Aragua, Yaracuy)	NT
209	Pectinopitys ladei (F.M.Bailey) C.N.Page	Stachypitys ladei, Podocarpus ladei, Stachycarpus ladei	Mount spurgeon black pine or Mount spurgeon brown pine	Australia (Queensland)	NT
210	Pectinopitys standleyi (J. Buchholz and N.E.Gray) C.N.Page	Podocarpus standleyi, Stachycarpus standleyi	Cipresillo, Ciprecillo, Ciprés lorito	Costa Rica (Alajuela, Cartago, Heredia, San José)	EN
211	Retrophyllum comptonii (J.Buchholz) C.N.Page	Decussocarpus comptonii, Nageia comptonii, Podocarpus comptonii	-	New Caledonia (Port Boise to Mt Ignambi)	LC
212	Retrophyllum filicifolium (N.E.Gray) R.R.Mill	Podocarpus filicifolius	Lehil, moegò	Indonesia (Maluku), Papua New Guinea (Bismarck Archipelago)	LC
213	Retrophyllum minus (Carrière) C.N.Page	Nageia minor, Podocarpus minor, Podocarpus palustris, Decussocarpus minor, Retrophyllum minor	Bois bouchon	New Caledonia (Grande Terre, Province Sud: Prony, Baie du Sud, Lac en Huit, Rivière des Lacs, Plaine des Lacs)	EN
214	Retrophyllum piresii (Silba) C.N.Page	Decussocarpus piresii, Nageia piresii	-	Brazil (Rondônia), Bolivia, Peru	DD
215	Retrophyllum rospigliosii (Pilg.) C.N.Page	Decussocarpus rospigliosii, Nageia rospigliosii, Podocarpus rospigliosii, Torreya bogotensis	Pino hayuelo, Diablo fuerte, Pino de monte, Pino real, Pino romero, Romerillo fino, Romerillo rojo, Saucecillo	Bolivia (Lap Paz), Colombia (Antioquia, Cundinamarca, Magdalena, Norte de Santander, Santander del Sur), Ecuador (Sucumbíos, Zamora-Chinchip), Peru (Junín, Pasco), Venezuela (Táchira, Mérida, Trujillo)	VU
216	Retrophyllum vitiense (Seem.) C.N.Page	Decussocarpus vitiensis, Nageia vitiensis, Podocarpus vitiensis	Ailumu, Dakua salusalu, Kau solo, Mungo	Fiji, Indonesia (Maluku, Papua), Papua New Guinea (Bismarck Archipelago), Solomon Islands (Santa Cruz Island), Vanuatu (Torba)	LC
217	Saxegothaea conspicua Lindl.	Squamataxus albertiana	Prince albert’s yew, Mañío, Mañío de hojas cortas, Mañío hembra, Mañío macho, Maniú	Argentina (Chubut, Neuquén, Rio Negro), Chile (Aisén, Biobío, La Araucania, Los Lagos, Maule)	NT
218	Sundacarpus amarus (Blume) C.N.Page	Nageia amara, Nageia eurhyncha, Podocarpus amara, Podocarpus dulcamara, Podocarpus eurhyncha, Podocarpus pedunculatus, Prumnopitys amara	Black pine, Choopoola	Australia (Queensland), Indonesia (Jawa, Maluku, Papua, Sulawesi, Sumatera), Malaysia (Sabah), Papua New Guinea (Bismarck Archipelago), Philippines, Timor-Leste	LC

Figure 2.

Diagram of two-way cluster analysis. The vertical matrix axis consists of more than 200 podocarp species coded by scientific name and the horizontal axis is the distributed countries (74 countries). The matrix was constructed depending on the presence (black) and absence (white). The species were grouped into five main clusters, I. New Caledonian group, II. New Zealand group and III. Malesian group, IV. Southeast Asian group and V. Podocarpian group.

1. Acmopyle has two species in New Caledonia (A. pancheri) and Fiji (A. sahniana).

2. Afrocarpus has five species distributed across Africa.

3. Dacrycarpus has nine species and two varieties distributed in New Caledonia, New Zealand, some Pacific Islands and Southeast Asia (Figure 3A).

4. Dacrydium has 20 species and two hybrids distributed in New Caledonia, New Zealand, some Pacific Islands and Southeast Asia.

5. Falcatifolium has six species distributed in New Caledonia, Papua New Guinea, Indonesia, Malaysia, the Philippines and Brunei Darussalam.

6. Halocarpus has three species endemic to New Zealand.

7. Manoao has one species distributed in New Zealand.

8. Pherosphaera has two species endemic to Australia.

9. Lagarostrobos has one species endemic to Australia.

10. Microcachrys has one species endemic to Australia (Figure 3B).

11. Lepidothamnus has three species distributed in Argentina, Chile and New Zealand.

12. Nageia has six species distributed in Southeast Asia to India and Myanmar, Papua New Guinea, the Philippines and Japan.

13. Parasitaxus has one parasitic species endemic to New Caledonia.

14. Phyllocladus has four species and one variety distributed in Papua New Guinea, Indonesia, Malaysia, the Philippines, New Zealand and Australia.

15. Podocarpus is the largest genus, with 120 species, 4 subspecies, 9 varieties and one hybrid and has a wide distribution, occurring in all continents (approximately 70 countries) except Europe and Antarctica.

16. Prumnopitys has three species distributed in New Zealand and South America.

17. Pectinopitys has six species distributed in New Zealand, Australia, New Caledonia and South America.

18. Retrophyllum has six species distributed in New Caledonia, South America and the Pacific Islands.

19. Saxegothaea has one species distributed in South America.

20. Sundacarpus has one species distributed in Australia, Indonesia, Malaysia, Papua New Guinea (Timor-Leste) and the Philippines.

Figure 3.

(A) Dacrycarpus dacrydioides (White pine, Kahikatea) tree in the rainforest of Wellington Kaitoke Regional Park, New Zealand. (B) Microcachrys tetragona (Strawberry pine) is a creeping shrub in the alpine region of cradle mountain summit, Tasmania.

The Podocarpaceae has its highest diversity of genera in New Zealand (nine) with fewer in other regions New Caledonia and Malesia (eight), Australia (seven), South America (four) and Africa and Asia (two). Of the 20 genera, three are endemic to Australia and two are endemic to New Zealand. Countries with a rich diversity of podocarps include Indonesia (51 species), Papua New Guinea (43 species), Malaysia (39 species), the Philippines (23 species), New Caledonia (20 species), New Zealand (19 species), China (17 species), Venezuela (15 species), Australia (14 species), Brazil, Peru and Bolivia (12 species each), Fiji (11 species), Ecuador (9 species) and Madagascar, Thailand, Brunei Darussalam and Colombia (8 species each).

3.3. Podocarpaceae in Space and Time

This checklist consists of macrofossils assigned to extant podocarp genera and includes more than one hundred taxa from the Cretaceous to the Pleistocene (Table 2). The macrofossils are predominantly recorded from Eocene–Miocene deposits. Australian and New Zealand macrofossil records dominate. Most of the macrofossils are foliage but well-preserved reproductive parts (seed and pollen cones) are also recorded for Lepidothamnus, Lagarostrobos, Dacrycarpus, Phyllocladus, Podocarpus and Nageia. Extant podocarps dominate in the Southern Hemisphere and analysis of extinct taxa assigned to living podocarp genera supports their past importance in the Southern Hemisphere (Table 2). A number of extinct species assigned to Podocarpaceae genera have been reported from Australia, New Zealand and South America [26]. An analysis shows that:

Table 2.

An updated checklist of podocarps fossil sp.ecies of extant genera.

S#	Fossil Taxa	Reported by	Part	Era/Period	Age Range Ma	Type Specimen	Distribution
1	Acmopyle antarctica	Florin, 1940 [27]	Leaf axis	Eocene	99.7 to 55.8	S132020-Swedish Museum of Natural History	Seymour Island (Antarctic Peninsula)
2	Acmopyle compactus	Pole, 1992 [28]	Leaf (Cuticle)	Middle–Late Eocene	48.6 to 33.9	S049-University of Tasmania	Hasties (Tasmania, Australia)
3	Acmopyle engelhardti	Florin, 1940 [27]	Leaf axis	Eocene–Miocene	55.8 to 48.6	PBDB 8673	(9 records) Rio Negro, Argentina
4	Acmopyle florinii	Hill and Carpenter, 1991 [29]	Sterile axis (Cuticle)	Late Paleocene	58.7 to 55.8	LB-063-University of Tasmania	Lake Bungarby (New South Wales, Australia)
5	Acmopyle glabra	Hill and Carpenter, 1991 [29]	Sterile axis (Cuticle)	Oligocene–Eocene	55.8 to 28.4	RPE-006-University of Tasmania	(9 records) Cethana—Oligocene, Regatta Point—Eocene (Tasmania, Australia)
6	Acmopyle masonii	Pole, 1997 [30]	Sterile axis (Cuticle)	Miocene	-	SB 1149-University of Tasmania.	Manuherikia Group, Central Otago, New Zealand
7	Acmopyle setiger	Hill and Carpenter, 1991; Carpenter and Pole, 1995; Townrow, 1965 [29,31,32]	Leaf (Cuticle)	Early Eocene	48.6 to 33.9	B-001-University of Tasmania	Lake Lefroy, Buckland and Monitor Bores (Tasmania and Western Australia)
8	Acmopyle tasmanica	Hill and Carpenter, 1991 [29]	Leaf (Cuticle)	Eocene	48.6 to 33.9	LA-060-University of Tasmania	Loch Aber (Tasmania, Australia)
9	Dacrycarpus acutifolius	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene to Early Miocene	28.4 to 15.97	M-235- University of Tasmania	Monpeelyata (Tasmania, Australia)
10	Dacrycarpus arcuatus	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene to Early Miocene	28.4 to 15.97	LRR1-243-University of Tasmania	Little Rapid River (Tasmania, Australia)
11	Dacrycarpus carpenterii	Jordan, 1995 [34]	Leaf	Early Pleistocene	2.588 to 0.126	RPU 525-University of Tasmania	Regatta Point (Tasmania, Australia)
12	Dacrycarpus chilensis	Wilf, 2012 [35]	Leaf (Cuticle)	Eocene		MMG PB SAT	Chile
13	Dacrycarpus crenulatus	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene to Miocene	28.4 to 15.97	P-221-University of Tasmania	Pioneer—tin mine (Tasmania, Australia)
14	Dacrycarpus cupressiformis	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Early Oligocene	33.9 to 23.0	LRR2-023-University of Tasmania	Little Rapid River 2 (Tasmania, Australia)
15	Dacrycarpus dacrydoides	Pole, 1992; 1997 [28,30]	Leaf (Cuticle)	Miocene	-	SB 1149-University of Tasmania.	Manuherikia Group, Central Otago, New Zealand
16	Dacrycarpus geminus	Pole, 1992 [28]	Leaf (Cuticle)	Eocene	56 to 33.9	S115-University of Tasmania	Hasties (Tasmania, Australia)
17	Dacrycarpus guipingensis	Wu et al., 2021 [36]	Seed cone+ Sterile axis	Miocene	-	GP109-Museum of Biology, Sun Yat-sen University	Guangxi, South China
18	Dacrycarpus involutus	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene to Miocene	28.4 to 15.97	M2023-University of Tasmania	Monpeelyata (Tasmania, Australia)
19	Dacrycarpus elandensis	Hill and Whang, 2000 [37]	Pollen cone, leaf	Miocene	-	ELD-005-University of Adelaide	Elands, New South Wales
20	Dacrycarpus falcatus	Carpenter, 1991 [38]	Sterile axis (Cuticle)	Oligocene	35	C-052, 203, 619-University of Tasmania	Cethana (Tasmania, Australia)
21	Dacrycarpus lanceolatus	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene to Miocene	28.4 to 15.97	M-1186-University of Tasmania	Monpeelyata (Tasmania, Australia)
22	Dacrycarpus latrobensis	Hill and Carpenter, 1991 [29]	Sterile axis (Cuticle)	Oligocene to Miocene	28.4 to 23.03	P 15714- Museum of Victoria, Melbourne	Southeastern Australia (Yallourn and Bacchus-3 locations)
23	Dacrycarpus linearis	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene	28.4 to 23.03	LRR2-051-University of Tasmania	Little Rapid River 2 (Tasmania, Australia)
24	Dacrycarpus linifolius	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Eocene to Oligocene	55.8 to 28.4	LRR1-851-University of Tasmania	Little Rapid River 1 (Tasmania, Australia) and Regatta Point (Tasmania, Australia)
25	Dacrycarpus microfolius	Jordan et al., 2011 [39]	Cuticle	Oligocene–Miocene	28.4 to 15.97	OU33024-Geology Museum (OU), University of Otago	F45/f0394, middle Gore Lignite Measure (Newvale Mine, New Zealand)
26	Dacrycarpus mucronatus	Wells and Hill, 1989; Carpenter, 1991; Lewis and Drinnan, 2013 [33,38,40]	Seed cone+ Sterile axis (Cuticle)	Eocene–Oligocene–Miocene	48.6 to 15.97	LRR2-044-University of Tasmania, (RPE-060-62, and RPE-4620	Little Rapid River 2 (Tasmania); Regatta Point (Tasmania) Cethana (Tasmania); Lochaber (Naracoorte, South Australia)
27	Dacrycarpus patulus	Hill and Merrifield, 1993 [41]	Leaf	Eocene–Oligocene	48.6 to 23.03	WAM P.84.34-Western Australian Museum	West Dale (Western Australia, Australia)
28	Dacrycarpus praecupressinus	Greenwood, 1987; Mill and Hill, 2004 [42,43]	Sterile axis (Leaves)	Eocene	37.2 to 33.9	F 51245-Geological Survey of New South Wales	Vegetable Creek—Witherdens Tunnel (NSW, Australia)
29	Dacrycarpus puertae	Wilf, 2012 [35]	Seed cone+ Sterile axis	Eocene	52–77.9	MPEF-Pb 4983-	Patagonia, Argentina
30	Dacrycarpus sp.	Pole et al., 1993 [44]	Leaf (Cuticle)	Late Oligocene–Early Miocene	-	SB282	Berwick Quarry, Victoria (Australia)
31	Dacrycarpus sp.	Carpenter and Pole, 1995 [32]	Leaf (Cuticle)	Middle Eocene	-	CD2999-University of Tasmania	Lefroy and Cowan Paleodrainages, Western Australia (Australia)
32	Dacrycarpus sp.	Carpenter, 1991 [38]	Foliage (Cuticle)	Oligocene	35	-	Cethana (Tasmania, Australia)
33	Dacrycarpus sp.	Carpenter et al., 1994 [45]	Foliage (Cuticle)	Oligocene–Early Miocene	-	R. J. Carpenter and R. S. Hill (unpublished data)	Lea River (Tasmania, Australia)
34	Dacrydium aciculare	Wells and Hill, 1989 [33]	Leaf (Cuticle)	Oligocene	33.9 to 28.4	LRR1-441-University of Tasmania	Little Rapid River 1 (Tasmania, Australia)
35	Dacrydium fimbriatus	Hill and Christophel, 2001 [46]	Sterile axis (Cuticle)	Middle Eocene	48.6 to 37.2	NC-004-University of Adelaide	Nelly Creek (South Australia, Australia)
36	Dacrydium microphyllum	Jordan et al., 2011 [39]	Cuticle	Oligocene–Miocene	28.4 to 15.97	OU33026-Geology Museum (OU), University of Otago	New Vale Mine, Waimumu (Coalfield, Southland, New Zealand)
37	Dacrydium mucronatus	Hill and Christophel, 2001 [46]	Fertile axis (Cuticle)	Eocene	48.6 to 37.2	NC-002- University of Adelaide	Nelly Creek (South Australia, Australia)
38	Dacrydium rhomboideum	Cookson and Pike, 1953; Blackburn, 1985 [47,48]	Foliage + Seeds	Oligocene–Miocene	28.4 to 15.97	P 209942	Morwell and Yallourm, Victoria (Australia)
39	Dacrydium sinuosum	Wells and Hill, 1989 [33]	Leaf (Cuticle)	Oligocene–Miocene	28.4 to 15.97	P-631-University of Tasmania	Pioneer- tin mine (Tasmania, Australia)
40	Dacrydium tasmanicum	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene	33.9 to 28.4	LRR1-1031-University of Tasmania	Little Rapid River 1 (Tasmania, Australia)
41	Dacrydium waimumuensis	Jordan et al., 2011 [39]	Cuticle	Oligocene–Miocene	28.4 to 15.97	OU33025-Geology Museum (OU), University of Otago	F45/f0394, middle Gore Lignite Measure (Newvale Mine, New Zealand)
42	Dacrydium Sp1	Carpenter, 1991 [38]	Foliage	Early Oligocene	35	C-202, 471-University of Tasmania	Cethana (Tasmania, Australia)
43	Dacrydium Sp2	Carpenter, 1991 [38]	Foliage	Early Oligocene	35	C-517, 519-University of Tasmania	Cethana (Tasmania, Australia)
44	Dacrydium Sp1	Carpenter and Pole, 1995 [32]	Foliage (disp.ersed cuticles)	Middle Eocene	-	CD2999 and DWT495-University of Tasmania	Lefroy and cowan paleodrainage, Western Australia
45	Dacrydium Sp2	Carpenter and Pole, 1995 [32]	Foliage (disp.ersed cuticles)	Middle Eocene	-	CD2999-University of Tasmania	Lefroy and cowan paleodrainage, Western Australia
46	Dacrydium Sp	Blackburn, 1985 [48]	Foliage	Oligocene–Miocene	-	-	Morwell, Victoria (Australia)
47	Dacrydium Sp	Blackburn, 1985 [48]	Foliage	Oligocene–Miocene	-	-	Morwell, Victoria (Australia)
48	Falcatifolium eocenica	Hill and Scriven, 1999 [49]	Sterile axis (Cuticle)	Middle Eocene	37.2 to 33.9	2351 and 2350-State Herbarium of South Australia.	ALCOA Anglesea Site II coal mine (Victoria, Australia)
49	Halocarpus highstedii	Jordan et al., 2011 [39]	Cuticle	Oligocene–Miocene	28.4 to 15.97	OU32899-Geology Museum (OU), University of Otago	F45/f0394, middle Gore Lignite Measure (Newvale Mine, New Zealand)
50	Lagarostrobos franklinii	Wells and Hill, 1989; Hill and Macphail, 1985; Carpenter et al., 1994; Jordan, 1995; Jordan et al., 2011 [33,34,39,45,50]	Seed cones and foliage	Late Pliocene–Early Pleistocene	2.588 to 0.126	RPU-190-University of Tasmania	Regatta Point (Tasmania, Australia)
51	Lagarostrobos marginatus	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene	33.9 to 28.4	LRR1-701-University of Tasmania	Little Rapid River 1 (Tasmania, Australia)
52	Lagarostrobos Sp	Peter, 1985 [51]	Sterile axis	Middle Cretaceous	145-100.5	-	Winton, Queensland
53	Lagarostrobos colensoi (correct name—Manoao colensoi)	Carpenter, 1991 [38]	Leaf (Cuticle)	Oligocene	35	-	Cethana (Tasmania, Australia)
54	Lepidothamnus intermedius	Pole, 1997 [30]	Sterile axis (Cuticle)	Miocene	-	S-632-University of Tasmania.	Manuherikia Group, Central Otago, New Zealand
55	Lepidothamnus diemenensis	Pole, 1992 [28]	Sterile axis (Cuticle)	Eocene	48.6 to 33.9	S014-University of Tasmania	Hasties (Tasmania, Australia)
56	Lepidothamnus	Peter, 1985 [51]	Seed cones and foliage	Middle Cretaceous	145-100.5	-	Winton, Queensland
57	Microcachrys tetragona	Jordan, 1995 [34]	Seed and sterile axes	Early Pleistocene		RPU2-University of Tasmania	Regatta Point (Tasmania, Australia)
58	Microcachrys novaezelandiae	Carpenter et al., 2011 [52]	Leaf (Cuticle)	Oligocene–Miocene	28.4 to 15.97	OU32896-Geology Museum (OU), University of Otago	F45/f0394, middle Gore Lignite Measure (Newvale Mine, New Zealand)
59	Nageia hainanensis	Jin et al., 2010 [53]	Leaf (Cuticle)	Eocene	-	CC-1200 a, b-The Museum of Biology of Sun Yat-sen University, Guangzhou, China	Changchang Basin, Hainan Island, south China
60	Nageia maomingensis	Liu et al., 2015 [54]	Leaf (Cuticle)	Late Eocene	-	MMJ1-001-The Museum of Biology, Sun Yat-sen University, Guangzhou, China.	Maoming Basin, Jintang, Maoming, Guangdong Province, South China.
61	(Podocarpus) Nageia ryosekiensis	Kimura et al., 1988 [55]	Leafy branches, connected seed, detached leaves	Lower Cretaceous	-	Makino Botanical Garden, Kochi	Southwest Japan
62	(Podocarpus) Nageia sujfunensis	Krassilov, 1965 [56]	Leaf (Cuticle)	Early Cretaceous	-	27/71-Far East Geological Institute, USSR Academy of Sciences	Far East Russia
63	Pherosp.haera sommervillae (Name correction Microstrobos sommervillae)	Townrow, 1965 [31]		Early Eocene	-	-	Buckland sediments in southeastern Tasmania
64	Pherosp.haera microfolius (Name correction Microstrobos microfolius)	Wells and Hill, 1989 [33]	Sterile axis (Cuticle)	Oligocene–Miocene	28.4 to 23.03	M-1155-University of Tasmania	Mudstone lens cutting Monpeelyata canal (Tasmania, Australia)
65	Phyllocladus aberensis	Hill, 1989 [57]	Leaf	Oligocene (Middle–Late Eocene)	28.4 to 23.03	LRR1-951-University of Tasmania	Little Rapid River 2 (Tasmania, Australia)
66	Phyllocladus annulatus	Hill, 1989 [57]	Leaf (Cuticle)	Oligocene	33.9 to 23.03	P-742-University of Tasmania	Pioneer (Tasmania, Australia)
67	Phyllocladus aspleniifolius	Ettingshausen, 1887, 1888; Cookson and Pike, 1954; Hill and Macphail, 1985; Hill, 1989, 1988; Pole, 1992 [28,50,57,58,59,60]	Leaf (Cuticle)	Late Eocene (Quaternary, Eocene and Cretaceous)	56-33.9 (84.9 to 0.012)	S 106-University of Tasmania	Deep leads NSW; Regatta point; Hasties, Tasmania (Australia) and Antarctica
68	Phyllocladus elongatus	Jordan et al., 2011 [39]	Leaf	Oligocene–Miocene	28.4 to 15.97	OU32901-Geology Museum (OU), University of Otago	F45/f0394, middle Gore Lignite Measure (Newvale Mine, New Zealand)
69	Phyllocladus lobatus	Hill, 1989 [57]	Cuticle	Oligocene	28.4 to 23.03	LRR1-1649- University of Tasmania	Little Rapid River 2 (Tasmania, Australia)
70	Phyllocladus morwellensis	Deane, 1925; Cookson and Pike, 1954; Hill, 1989 [57,60,61]	Leaf	Oligocene	28.4 to 15.97	P-15873-Museum of Victoria	Near Morwell, (Victoria, Australia)
71	Phyllocladus palmeri	Pole and Moore, 2011 [62]	Leaf (Cuticle)	Late Miocene	6 to 6.5	AU P340a and b-School of geography, geology and environmental science, University of Auckland	Near Matuora, (Coromandel Peninsula, New Zealand)
72	Phyllocladus Sp (P. lobatus)	Carpenter, 1991 [38]	Leaf (Cuticle)	Oligocene	-	-	Cethana, (Tasmania, Australia)
73	Phyllocladus Sp2 (P. hypophyllus)	Carpenter, 1991 [38]	Leaf (Cuticle)	Oligocene	-	-	Cethana, (Tasmania, Australia)
74	Phyllocladus Sp	Mc Loughlin and Hill, 1996; Mc Loughlin et al., 2001 [63,64]	Leaf (Cuticle)	Late Eocene	-	-	Kojonup, Western Australia
75	Phyllocladus Sp	Pole, 1992 [28]	Leaf (Cuticle)	Late Miocene	-	-	Near Cromwell, (South Island, New Zealand)
76	Phyllocladus Sp	Pole, 1992 [28]	Leaf (Cuticle)	Miocene	-	OU30068-Department of Geology, University of Otago.	Manuherikia Group, Central Otago, New Zealand
77	Phyllocladus Sp	Liz Kennedy, 2020 (unpublished)	Seed cones (Phyllocladus toatoa)	Miocene	23 to 5.3	-	Coromandel, North Island, New Zealand
78	Podocarpus andiniformis (Subgenus Foliatus)	Wilf et al., 2017/Berry, 1922 [65,66]	Leaf (Cuticle)	Late Triassic and Early Eocene	-	-	Patagonia, Argentina
79	Podocarpus alwyniae (Subgenus Podocarpus)	Pole, 1992 [28]	Sterile axis (Cuticle)	Miocene	-	OU29708, H411fD45-Department of Geology, University of Otago.	Manuherikia Group, Central Otago, New Zealand
80	Podocarpus oligocenicus	Awasthi et al., 1992 [67]	Leaf (Cuticle)	Oligocene	-	-	Mizoram, India; Manipur, India
81	Podocarpus araucoensis (as Decussocarpus araucoensis)	Berry, 1922; Mill and Hill, 2004 [43,66]	Foliage (Cuticle)	Eocene	55.8 to 33.9	-	Chile
82	Podocarpus brownei (probably Retrophyllum or Falcatifolium)	Greenwood, 1987 (Decussocarpus brownei); Mill and Hill, 2004 [42,43]	Foliage (Cuticle)	Eocene	37.2 to 33.9	2345-State Herbarium of South Australia	ALCOA Anglesea Site II coal mine (Victoria, Australia)
83	Podocarpus fildesensis	Zhou and Li, 1994 [68]	Foliage	Cretaceous	84.9 to 66.043	-	Half Three Point assemblage (Antarctica)
84	Podocarpus inopinatus	Florin, 1940 [27]	Foliage	Paleogene, Eocene–Miocene	55.8 to 33.9	-	Chile
85	Podocarpus platyphyllum	Greenwood, 1987 [42]	Leaf (Cuticle)	Middle to Late Eocene	37.2 to 33.9	1816-State Herbarium of South Australia	ALCOA Anglesea Site II coal mine (Victoria, Australia)
86	Podocarpus sinuatus	Pole, 1992 [28]	Leaf (Cuticle)	Eocene	48.6 to 33.9	S112-University of Tasmania	Hasties (Tasmania, Australia)
87	Podocarpus pliomacrophyllus (Subgenus Foliatus)	Chen et al., 2019; Wu et al., 2021 [69,70]	Leaf (Cuticle)	Lower Pliocene	-	MBU-16395- Institute of Palaeontology and Stratigraphy, Lanzhou University, Gansu Province, China.	Mannong Village (western Yunnan, China); Tuantian Town, Yunnan Province, southwestern China
88	Podocarpus travisiae (Subgenus Podocarpus)	Pole, 1993 [71]	Leaf	Miocene	23.03 to 15.97	0U30780- Department of Geology, University of Otago	Foulden Hills (New Zealand)
89	Podocarpus yunnanensis (Subgenus Foliatus)	Wu et al., 2021 [70]	Leaf (Cuticle)	Early Pliocene	-	MBU-19122301-Institute of Palaeontology and Stratigraphy, Lanzhou University	Tuantian Town, Yunnan Province, southwestern China
90	Podocarpus forrestii (Subgenus Foliatus)	Wu et al., 2021 [70]	Leaf (Cuticle)	Early Pliocene	-	MBU-20191221-Institute of Palaeontology and Stratigraphy, Lanzhou University	Tuantian Town, Yunnan Province, southwestern China
91	Podocarpus tasmanicus (Subgenus Podocarpus)	Townrow, 1965 [31]	Leaf (Cuticle)	Eocene	-	81905-University of Tasmania	Bed of Tea Tree Rivulet, Buckland, Tasmania, Australia
92	Podocarpus strzeleckianus (Subgenus Podocarpus)	Townrow, 1965 [31]	Leaf (Cuticle)	Eocene	-	81917-University of Tasmania	Bed of Tea Tree Rivulet, Buckland, Tasmania, Australia
93	Podocarpus witherdenensis (Subgenus Podocarpus)	Hill and Carpenter, 1991 [29]	Fertile axis+ seed cones (Cuticle)	Eocene	37.2 to 33.9	MMF 1201-Geological Survey of New South Wales, Sydney	Vegetable Creek—Witherdens Tunnel (NSW, Australia)
94	Podocarpus Sp	Carpenter et al., 1994 [45]	Foliage (Cuticle)	Oligocene–Early Miocene	-	R. J. Carpenter and R. S. Hill (unpublished data)	Lea River (Tasmania, Australia)
95	Podocarpus Sp1	Carpenter, 1991 [38]	Foliage (Cuticle)	Oligocene	-	University of Tasmania	Cethana, (Tasmania, Australia)
96	Podocarpus Sp2	Carpenter, 1991 [38]	Foliage (Cuticle)	Oligocene	-	C-251, 274, 275, 342, 492- University of Tasmania	Cethana, (Tasmania, Australia)
97	Podocarpus Sp	Carpenter et al., 1994 [45]	Foliage (Cuticle)	Oligocene–Early Miocene	-	R. S. Hill (unpublished data)	Little Rapid River (Tasmania, Australia)
98	Podocarpus Sp	Hill and Macphail, 1985 [50]	Foliage (Cuticle)	Late Pliocene–Early Pleistocene	-	University of Tasmania	Regatta Point (Tasmania, Australia)
99	Podocarpus Sp (Subgenus Podocarpus)	Jordan et al., 2011 [39]	Foliage (Cuticle)	Late Oligocene–Early Miocene	-	OU33027- Geology Museum, University of Otago	Newvale site, (South Island, New Zealand)
100	Podocarpus Sp	Pole, 1997 [72]	Foliage (Cuticle)	Miocene	-	H41/f74, S788-University of Tasmania.	Manuherikia Group, Central Otago, New Zealand
101	Podocarpus Sp	He and Wang, 2021 [73]	Leaf (Cuticle)	Miocene	17 to 14	-	Guangchang County, Jiangxi Province, southeastern China
102	Retrophyllum australe	Hill and Merrifield, 1993 [41]	Sterile axis	Eocene–Oligocene	48.6 to 23.03	WAM P.88.96-Western Australian Museum	West Dale (Western Australia, Australia)
103	Retrophyllum superstes	Wilf et al., 2017 [65]	Sterile axis (Leafy twig)	Cretaceous–Paleocene	70.6 to 61.7	MPEF-Pb 8910- Patagonia, Argentina	LefE (Chubut, Argentina)
104	Retrophyllum oxyphyllum (Retrophyllum sp.iralifolium)	Wilf, 2020 [74]	Sterile axis, cuticle, leaves, fertile axis	Eocene	52	MLP-4234 and MPEF–Pb 8915a Museo Paleontológico Egidio Feruglio	Trelew, Argentina
105	Retrophyllum vulcanense	Pole, 1992 [28]	Sterile axis (Cuticle)	Miocene	-	OU29857-Department of Geology, University of Otago.	Manuherikia Group, Central Otago, New Zealand
106	Prumnopitys tasmanica	Mill and Hill, 2004; Greenwood, 1987 [42,43]	Sterile axis (Cuticle)	Eocene	-	81905-University of Tasmania	Alcoa Anglesea, Victoria, Australia
107	Prumnopitys montana	Pole, 1992 [28]	Cuticle	Eocene	48.6 to 33.9	S 110.-University of Tasmania	Hasties (Tasmania, Australia)
108	Prumnopitys opihiensis	Pole, 1997 [72]	Cuticle	Cretaceous/Eocene	99.7 to 48.6	OU30932- University of Otago	Taratu Formation (New Zealand)
109	Prumnopitys portensis	Pole, 1992 [28]	Leaf	Eocene	48.6 to 33.9	S056-University of Tasmania	Hasties (Tasmania, Australia)
110	Prumnopitys taxifolia (Leaf morphology is similar to that of Sundacarpus)	Pole, 1997 [30]	Leaf	Miocene	-	SB 1154--Department of Geology, University of Otago.	Manuherikia Group, Central Otago, New Zealand
111	Sundacarpus anglica	Page, 2019 [75]	Leaf (Cuticle)	Eocene	48.6 to 33.9	V.46883- Natural History Museum, BM	Bandulska (Bournemouth, England)
112	Sundacarpus tzagajanicus	Page, 2019 [75]	Leaf (Cuticle)	Uppermost Cretaceous (Earliest Paleocene—65.5–61.7 Ma)	65.5–61.7	575-126-Far Eastern Scientific Centre, Vladivostok	Bureya River (Russia)

1. Acmopyle has eight fossil species recorded (A. antarctica, A. compactus, A. engelhardti, A. florinii, A. glabra, A. masonii, A. setiger and A. tasmanica) from the Eocene–Oligocene of the Antarctic Peninsula, Australia, New Zealand and Argentina [27,28,29,30,31,32]. The fossil record shows a wider past distribution of Acmopyle compared to its current occurrence in New Caledonia and Fiji.

2. Dacrycarpus has approximately 25 fossil taxa reported (D. acutifolius, D. arcuatus, D. carpenterii, D. chilensis, D. crenulatus, D. cupressiformis, D. dacrydoides, D. geminus, D. guipingensis, D. involutus, D. elandensis, D. falcatus, D. lanceolatus, D. latrobensis, D. linearis, D. linifolius, D. microfolius, D. mucronatus, D. patulus, D. praecupressinus, D. puertae, Dacrycarpus sp. (four separate records)) from the Eocene–Early Pleistocene of Australia, New Zealand, China, Argentina [28,30,32,33,34,35,36,37,38,39,40,41,42,43,45,71,76]. Although Dacrycarpus currently has no living species in Australia or South America, the fossil record shows its extensive distribution in both those landmasses in the past.

3. Dacrydium has approximately 14 fossil taxa reported from Australia (D. aciculare, D. fimbriatus, D. mucronatus, D. sinuosum, D. rhomboideum, D. tasmanicum and six taxa described to genus level) and two species from New Zealand (D. microphyllum and D. waimumuensis) [32,33,38,39,47,48]. These suggest an Australasian origin in the Late Cretaceous [77].

4. Falcatifolium has only one fossil extinct species (F. eocenica) from the Middle Eocene of Victoria, Australia [49]. Currently, Australia has no living species of Falcatifolium.

5. Nageia has four extinct species (N. hainanensis, N. maomingensis, N. ryosekiensis, N. sujfunensis), from the Early Cretaceous–Eocene in China, Southwest Japan, and Far East Russia [53,54,55,56,78]. This demonstrates that Nageia had a wider past distribution and occurred in Japan and Russia.

6. Retrophyllum has four extinct species (R. australe, R. oxyphyllum, R. superstes, R. vulcanense) recorded from the Cretaceous–Miocene in Australia, Argentina, and New Zealand [28,41,65,74]. Currently, Australia and New Zealand have no living species of Retrophyllum.

7. Podocarpus has at least 16 extinct species reported and approximately seven taxa identified at the genus level. These are Podocarpus platyphyllum, P. sinuatus, P. strzeleckianus, P. tasmanicus and P. witherdenensis (Eocene) from Australia; P. travisiae and P. alwyniae (Miocene) from New Zealand; P. andiniformis, P. araucoensis and P. inopinatus (Eocene–Miocene) from South America; and P. pliomacrophyllus, P. yunnanensis and P. forrestii (lower Pliocene) from China [27,28,38,42,65,69,70,71,79].

8. Halocarpus has a single occurrence of one extinct species (Halocarpus highstedii) reported from the Oligocene–Miocene of New Zealand [39].

9. Manoao colensoi has a fossil record from the Oligocene of Cethana, Tasmania (Australia) (reported as Lagarostrobos colensoi) [38], showing that this current New Zealand endemic genus was once also distributed in Australia.

10. Lepidothamnus has three fossil records: L. intermedius from the Miocene in New Zealand [30], L. diemenensis from the Eocene of Hasties, Tasmania [28] and an undescribed extinct species from the middle Cretaceous of Winton, Queensland [51]. This indicates a wider distribution of Lepidothamnus in the Late Mesozoic across the Southern Gondwana regions [51].

11. Lagarostrobos has two fossil records, e.g., L. marginatus from the Early Oligocene and the extant L. franklinii from the Early Pleistocene in Tasmania, Australia [33,34,39,45,50]. Current and macrofossil records suggest a narrow distribution and endemism to Australia for this genus.

12. Phyllocladus has seven fossil species described, including records of the extant P. aspleniifolius (the fossil species are P. aberensis, P. annulatus, P. elongatus, P. lobatus, P. morwellensis, P. palmeri) and six at genus level from Late Eocene–Pliocene in Australia, New Zealand and Antarctica (Cretaceous) [38,39,47,50,57,58,59,61,62,63,64]. Protophyllocladus Berry [80], an extinct genus that resembles the foliage morphology of Phyllocladus, is recorded from the Jurassic and Cretaceous of the United States and southwestern Canada, Western Greenland, Serbia, Romania, Portugal, Kazakhstan, Japan, northeastern Russia, Germany [81]. Dörken et al. [82] considered that it is unlikely that Protophyllocladus is closely related to Phyllocladus. Wagstaff [83] suggested that extant species are the remnants of one of the recently diverged lineages of Phyllocladus, but there is no unequivocal fossil evidence to support this.

13. Prumnopitys has two extinct species (P. portensis and P. tasmanica) and a fossil record of one extant species (P. montana) from the Eocene in Australia and one extinct species (P. opihiensis) and a fossil record of one extant species (P. taxifolia) from the Cretaceous–Miocene in New Zealand [28,30,42,43,72].

14. Sundacarpus has two extinct species: S. anglica and S. tzagajanicus, from the Eocene–Oligocene in Australia and Cretaceous–Paleocene in Argentina, respectively [75].

15. Pherosphaera has two extinct species: P. sommervillae (Microstrobos sommervillae) from the Early Eocene and P. microfolius (Microstrobos microfolius) from the Oligocene–Miocene of Tasmania [31,33].

16. Microcachrys has one extinct species (Microcachrys novae-zelandiae) from the Oligo-Miocene of New Zealand and fossils of the extant Microcachrys tetragona from the Early Pleistocene of western Tasmania [34,45,52].

3.4. Chromosomal Number

The chromosomal number in the Podocarpaceae varies from x = 9 to x = 19 (Acmopyle x = 10, Afrocarpus x = 12, Dacrycarpus x = 10, Dacrydium x = 10, Falcatifolium x = 10, Halocarpus x = 9, 11, 12, Lagarostrobos x = 15, Lepidothamnus x = 14, 15, Manoao x = 10, Microcachrys x = 15, Nageia x = 18, Parasitaxus x = 18, Pectinopitys x = 19, Pherosphaera x = 13, Phyllocladus x = 9, Podocarpus x = 10, 11, 17, 18, 19, Prumnopitys x = 18, Retrophyllum x = 10, Saxegothaea x = 12 and Sundacarpus x = 12) [84].

4. Discussion

4.1. Phylogenetic History of the Podocarpaceae

Molecular studies suggest Araucariaceae as the sister family of Podocarpaceae, although these families are morpho-anatomically divergent [4,11,12,13], which was also supported by our results. Previous molecular and fossil records suggest that podocarps originated in the Triassic–Jurassic in Gondwana [12,85], or the Early Cretaceous [10], or Late Triassic [13], but recent podocarp fossils from Jordan push back the origin of the Podocarpaceae to the Permian (Figure 1) and show that they survived the “great dying” at the end of Permian [86,87]. Our results suggest that Lepidothamnus and Phyllocladus diverged in the Late Jurassic, when incorporating the oldest Lepidothamnus fossil record [51,88], which is earlier than the previous estimate of mid-Cretaceous–early Paleogene [10,12] and Early Jurassic [13]. Our studies recognized the presence of three major Prumnopityoid, Dacrydioid and Podocarpoid clades and a paraphyletic group similar to Chen et al. [13].

Several studies based on both morphological [7,89,90,91,92] and molecular [7,92,93,94,95,96] studies have been published evaluating the phylogenetic relationship among different genera of the Podocarpaceae. Based on morphology and 18S rDNA, Kelch [7] concluded that the Podocarpaceae are monophyletic except for Podocarpus (paraphyletic) and Dacrydium (polyphyletic). Conran et al. [93], based on molecular analysis (plastid rbcL), reported that Podocarpaceae are polyphyletic and supported the separation of Afrocarpus from Podocarpus and its placement as sister to Retrophyllum instead of Nageia, and also suggested that Podocarpus is monophyletic, a conclusion supported by Sinclair et al. [94]. Biffin et al. [85], based on their molecular studies of 94 Podocarpaceae species, reported that Podocarpus is closely related to Nageia, Afrocarpus and Retrophyllum. Knopf et al. [92] investigated the phylogeny of 145 species (including 77 species of Podocarpus) of Podocarpaceae based on morphological, anatomical and DNA sequences (rbcL, nrITS1 and NEEDLY). Their most significant findings were the support of subgenera in Podocarpus, the transfer of Sundacarpus amarus to Prumnopitys and the incorporation of the Phyllocladaceae into the Podocarpaceae as Phyllocladus. Lu et al. [11] reported two monophyletic sister groups: the Dacrydioid group (Dacrycarpus, Dacrydium and Falcatifolium) and the Podocarpoid group (Retrophyllum–Nageia subclade and the Afrocarpus–Podocarpus subclade). Little et al. [95] used DNA barcoding (matK, rbcL and nrITS2 DNA barcodes) for the identification of Podocarpaceae (18 genera and 145 species) and to construct a phylogenetic tree. Quiroga et al. [97], based on molecular and fossil data, reported that Podocarpus originated in late Cretaceous–early Paleogene (~63 Ma) and supported the two subgenera in Podocarpus. Leslie et al. [96], using more comprehensive sampling and markers, recognized 19 genera and supported the division of Podocarpus into two subgenera. Recently, Page [75] again split the genus Prumnopitys into three genera (Prumnopitys, Sundacarpus and the new genus Pectinopitys) based on morphological and molecular data. The current phylogeny supports the division of the 20 genera of podocarps into main three clades and a paraphyletic grade (Figure 1). Similarly, the current phylogeny also recognizes and supports the division of Podocarpus into two subgenera, i.e., Podocarpus and Foliolatus [12,13,92,97].

4.2. Historical Taxonomic Treatment

The most extensive taxonomic studies on the Podocarpaceae have been by de Laubenfels, Buchholz, Gray and Page, with many other contributions, which are summarized in Table 3.

Table 3.

A brief historical overview of major taxonomic classifications of Podocarpaceae (Type genus Podocarpus elongatus).

Taxonomist	Taxonomic Treatment
Endlicher, 1847 [98]	He classified Podocarpaceae into three genera i. Podocarpus (with four sections i. Eupodocarpus, ii. Stachycarpus, iii. Nageia and iv. Dacrycarpus), 2. Dacrydium Sol. ex G. Forst, 3. Microcachrys Hook. f.
Pilger, 1926 [99]	He considered Podocarpaceae as subfamilies Podocarpoideae with Subgenus I. Protopodocarpus (with section i. Eupodocarpus, ii. Dacrycarpus), II. Stachycarpus with section B. i. Nageia ii. Saxegothaea iii. Microcachrys iv. Pherosphaera v. Acmopyle vi. Dacrydium, vii. section A, viii. Microcarpus and Phyllocladoideae with i. Phyllocladus
Buchholz and Gray, 1948 [100,101]	Classified Podocarpus into nine sections (P. sect. Eupodocarpus, P. sect. Nageia, P. sect. Afrocarpus, P. sect. Polypodiopsis, P. sect. Microcarpus, P. sect. Dacrycarpus, P. sect. Sundacarpus, P. sect. Stachycarpus)
Keng, 1973 [102]	Divided into two families, i.e., Podocarpaceae and Phyllocladaceae
Gaussen, 1974 [103]	Raised this group into suborder Podocarpineae and divided into three families, i.e., Podocarpaceae, Phyllocladaceae and Saxegothaeaceae.
de Laubenfels, 1985 [104]	Classified Podocarpus into two subgenera and 18 sections (subgenus Podocarpus: sect. Podocarpus, sect. Scytopodium, sect. capitulatis, sect. Australis, sect. Crassiforms, sect. Pratensis, sect. Lanceolatis, sect. Pumilis, sect. Nemoralis, subgenus Foliolatus: sect. Globulus, sect. Foliolatus, sect. Acuminatis, sect. Longifoliolatus, sect. Gracilis, sect. Macrostachyus, sect. Spinulosus, sect. Rumphius, sect. Polystachyus.)
Quinn, 1987 [105]	Placed back Phyllocladus in Podocarpaceae
Hart, 1987 [90]	Recognized 15 genera Lagarostrobos, Microstrobos (Pherosphaera), Microcachrys, Lepidothamnus, Halocarpus, Parasitaxus, Dacrycarpus, Falcatifolium, Dacrydium, Acmopyle, Nageia, Saxegothaea, Phyllocladus, Prumnopitys and Podocarpus
Page, 1988 [106]	Recognized eight genera in s.l. Podocarpus and five in Dacrydium
Page, 1990 [107]	Classified Podocarpaceae into Acmopyle, Falcatifolium, Dacrydium, Halocarpus, Lagarostrobos, Lepidothamnus, Microcachrys, Microstrobos (Pherosphaera), Phyllocladus and Podocarpus (P. subg. Podocarpus and P. subg. Foliolatus) Nageia (N. sect. Nageia, N. sect. Afrocarpus, N. sect. Polypodiopsis), Dacrycarpus, Parasitaxus, Prumnopitys, Sundacarpus, Saxegothaea
Dezhi, 1992 [108]	Placed Nageia into a new family Nageiaceae
Kelch, 1998 [7]	Produced the phylogeny of Podocarpaceae-based molecular markers (18S RNA) of 10 genera in the following sequences: Podocarpus, Dacrycarpus, Pherosphaera, Microcachrys, Afrocarpus, Saxegothaea, Dacrydium, Parasitaxus, Lagarostrobos and Phyllocladus.
Conran et al., 2000 [93]	Produced the phylogeny of Podocarpaceae-based molecular markers (rbcL) of 16 genera in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Acmopyle, Pherosphaera, Microcachrys, Lagarostrobos, Manoao, Prumnopitys, Halocarpus, Phyllocladus, Lepidothamnus and Saxegothaea.
Kelch, 2002 [14]	Produced the phylogeny of Podocarpaceae-based molecular markers (18S RNA) of 16 genera in the following sequences: Dacrydium, Falcatifolium, Dacrycarpus, Pherosphaera, Microcachrys, Saxegothaea, Acmopyle, Nageia, Afrocarpus, Podocarpus, Lagarostrobos, Halocarpus, Parasitaxus, Phyllocladus, Lepidothamnus and Prumnopitys.
Sinclair et al., 2002 [94]	Constructed the phylogeny of 18 genera-based molecular markers (trnL-trnF+ITS2) in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Acmopyle, Pherosphaera, Microcachrys, Saxegothaea, Lagarostrobos, Manoao, Parasitaxus, Halocarpus, Prumnopitys, Lepidothamnus and Phyllocladus.
Wagstaff, 2004 [83]	Constructed the phylogeny of 9 genera-based molecular markers (rbcL+matK) in the following sequences: Afrocarpus, Podocarpus, Dacrydium, Saxegothaea, Halocarpus, Lepidothamnus, Prumnopitys and Phyllocladus.
Biffin et al., 2012 [8]	Constructed the phylogeny of 18 genera based molecular markers (matK+ trnL-trnF+ITS2) in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Acmopyle, Pherosphaera, Microcachrys, Saxegothaea, Lagarostrobos, Manoao, Parasitaxus, Halocarpus, Prumnopitys, Lepidothamnus and Phyllocladus.
Knopf et al., 2012 [92]	Constructed the phylogeny of 18 genera-based molecular markers (ITS1+NEEDLY intron 2+ anatomy and morphology) in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Pherosphaera, Microcachrys, Halocarpus, Lepidothamnus, Lagarostrobos, Manoao, Phyllocladus, Prumnopitys and Saxegothaea.
Little et al., 2013 [95]	Used DNA barcoding (matK, rbcL and nrITS2 DNA barcodes) for the identification of Podocarpaceae (18 genera and 145 species) and to construct the phylogenetic tree
Lu et al., 2014 [11]	Constructed the phylogeny of 18 genera-based molecular markers (LEAFY+NEEDLY CDS+ introns) in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Acmopyle, Pherosphaera, Saxegothaea, Microcachrys, Lagarostrobos, Manoao, Parasitaxus, Phyllocladus, Lepidothamnus, Halocarpus and Prumnopitys.
Contreras et al., 2017 [109]	Constructed the phylogeny of 18 genera-based molecular markers in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrydium, Falcatifolium, Dacrycarpus, Acmopyle, Pherosphaera, Microcachrys, Saxegothaea, Halocarpus, Phyllocladus, Lepidothamnus, Prumnopitys, Lagarostrobos, Manoao and Parasitaxus.
Leslie et al., 2018 [12]	Recently constructed the phylogeny of 19 genera-based molecular markers (18S, rbcL and matK) in the following sequences: Podocarpus, Afrocarpus, Nageia, Retrophyllum, Falcatifolium, Dacrydium, Dacrycarpus, Acmopyle, Pherosphaera, Microcachrys, Saxegothaea, Prumnopitys, Sundacarpus, Manoao, Lagarostrobos, Parasitaxus, Halocarpus, Phyllocladus and Lepidothamnus.
Sudianto et al., 2019 [110]	Constructed the phylogeny tree of 12 genera based on Plastome in the following sequences: Afrocarpus, Nageia, Retrophyllum, Podocarpus, Dacrycarpus, Dacrydium, Pherosphaera, Saxegothaea, Phyllocladus, Lagarostrobos, Lepidothamnus and Prumnopitys.
Page, 2019 [75]	Recently divided the genus Prumnopitys into two genera, Prumnopitys (Subgenus Prumnopitys and Subgenus Botryopitys) and Pectinopitys.
Khan et al., 2023 [current classification]	Dacrycarpus, Halocarpus, Lepidothamnus, Manoao, Dacrydium, Lagarostrobos, Microcachrys, Pherosphaera, Parasitaxus, Acmopyle, Falcatifolium, Phyllocladus, Retrophyllum, Prumnopitys, Pectinopitys, Afrocarpus, Nageia, Podocarpus, Sundacarpus and Saxegothaea.

Initially, podocarps were placed in two genera, Podocarpus and Dacrydium, mainly based on leaf morphology [98]. Several early taxonomists including Gordon [111] and Philippi [112] recognized variation in Podocarpus and Dacrydium and classified them into several sections, subgenera, and subgroups. From the 1960s onwards, Podocarpus was then divided into eight genera and Dacrydium into five. Based on leaf morphology and anatomy, Podocarpus was initially divided into eight sections (Afrocarpus, Dacrycarpus, Eupodocarpus, Microcarpus, Nageia, Polypodiopsis, Stachycarpus and Sundacarpus). After a more detailed examination, de Laubenfels [113] raised section Dacrycarpus to the genus Dacrycarpus. Quinn [114] suggested raising the eight sections of Podocarpus to generic level and de Laubenfels [115] raised the section Microcarpus to generic level as Parasitaxus. Later, de Laubenfels [104] revised the genus Podocarpus into 18 sections and described 94 species. Page [106] raised section Sundacarpus into the genus Sundacarpus, section Polypodiopsis to the genus Retrophyllum, section Nageia to the genus Nageia and section Afrocarpus to the genus Afrocarpus. Some taxonomists reject these changes of status [116,117,118]. Page [107] divided the Podocarpaceae into 17 genera (Phyllocladus was excluded and Sundacarpus was included). Biffin et al. [85] recognized three major clades, i.e., the Podocarpoid clade (Afrocarpus, Nageia, Podocarpus, Retrophyllum), the Dacrydioid clade (Dacrydium, Dacrycarpus, Falcatifolium) and the Prumnopityoid clade (Halocarpus, Lagarostrobos, Manoao, Parasitaxus, Prumnopitys).

The concept of the separate family Phyllocladaceae has been supported in several different studies [93,107,119], and while this is no longer regarded as valid, its status as the genus Phyllocladus has been well supported by other taxonomists and recent phylogenetic studies [12,13,83,92,94,95,120,121,122,123,124].

Our studies recognize 20 extant genera (Acmopyle, Afrocarpus, Dacrycarpus, Dacrydium, Falcatifolium, Halocarpus, Lagarostrobos, Lepidothamnus, Manoao, Microcachrys, Nageia, Parasitaxus, Pectinopitys, Pherosphaera, Phyllocladus, Podocarpus, Prumnopitys, Retrophyllum, Saxegothaea and Sundacarpus), two subgenera in both Podocarpus (Podocarpus and Foliolatus) and Prumnopitys (Prumnopitys and Botryopitys) in Podocarpaceae, similar to Page [75] and Yang et al. [1], Chen et al. [13] proposes the splitting of Prumnopitys into Prumnopitys and Pectinopitys but reported 19 genera for Podocarpaceae. Our checklist enlists 201 living species for Podocarpaceae than previously reported 181 species by Yang et al. [1], which will increase the total number of conifer species from 702 to 722.

4.3. Current Diversity and Distribution of Podocarpaceae

Podocarps occur mainly in the Southern Hemisphere, although some genera extend northward, i.e., subtropical China and Japan and to Mexico and the Caribbean [125]. The living species of Podocarpaceae are a small representation of a once highly diverse group [55,126]. Today, several genera have low species representation (e.g., monospecific in Manoao, Lagarostrobos, Microcachrys, Parasitaxus and Saxegothaea and two in Acmopyle and Pherosphaera), although the fossil record suggests a more extensive diversity for at least some of these genera in the past. The center of diversity for the Podocarpaceae is Australasia (New Caledonia, Tasmania, New Zealand and Malesia), South America (Andes mountains), Indo-China and the Philippines [125,127].

Podocarps favor mostly cool and wet climates but usually do not tolerate extreme cold like some Northern Hemisphere conifers [128]. However, some temperate Podocarpaceae species occur as shrubs and prostrate woody plants above the tree line in the alpine ecosystems of Tasmania, Victoria, and New Zealand (Figure 4).

Figure 4.

(A) Podocarpus lawrencei in the alpine region, Mount McKay Falls Creek, Australia. (B) Pherosphaera hookeriana and Microcachrys tetragona populations in the alpine region of Cradle Mountain summit, Tasmania.

The tropical podocarps are mostly confined to mountain forests and heathlands and nutrient-poor habitats in the lowlands, although some also grow in forest understories. Temperate podocarps are good competitors in nutrient-poor soils probably because the light is more easily available within the incomplete canopies, but they are outcompeted in nutrient-rich soil as the canopy and forest floor is occupied by angiosperms and the growth of new individuals is slow due to shading. Such conditions favor broad-leaved podocarps (Nageia and broad-leaved Podocarpus species are shade-tolerant) and exclude imbricate-leaved podocarps due to competition [128]. This is supported by Adie and Lawes [129], who concluded that African podocarps are not lowland rainforest survivors but are temperate forest relicts.

4.4. Historical Biogeography

The historical reconstruction of Podocarpaceae confirms that it is a Southern Hemisphere family that was initially centered in Gondwana [130]. Leslie et al. [12] suggest that Podocarpaceae diversified in the Cretaceous and earliest Cenozoic after its appearance in the Triassic of Gondwana. Klaus and Matzke [10], based on the reconstruction of ancestral ranges, suggested that podocarps originated in the Early Jurassic in what is today Central–South America, Australia, and New Zealand. The family subsequently dominated Australasia and Southern America and later (and through to the present) in Malesia [77]. However, the discovery of macrofossils of podocarps from the early Permian of Jordan [86,87] will require a re-assessment of the early history of the family [88].

Klaus and Matzke [10] used living taxa to reconstruct the ancestral ranges and suggested that the ancestral area for the Podocarpoid clade is the Australia–New Guinea–Malesian region; for the Dacrydioid clade it is New Caledonia; for the Prumnopityoid clade it is New Zealand and for the paraphyletic group/grade, South America to Australia. Macrofossil evidence and the historical biogeographic reconstruction by Klaus and Matzke [10] support an Australian origin of Podocarpus and multiple dispersals to South America, Asia, New Zealand, Malesia, and New Caledonia. Morley [77] concluded that Podocarpus dispersed into South Asia in the Late Eocene, either by dispersal from India or by multiple long-distance dispersal events from Australia. Similarly, he concluded that Podocarpus was possibly present in Africa during the mid-Cenozoic but its dispersal to West Africa occurred by island-hopping in the late Pliocene. According to Nieto-Blázquez et al. [131], Podocarpus species in the Caribbean are the result of colonization from the Andes during the Eocene to Oligocene (c. 45–31 Ma). Fossil records and living species distributions of Nageia support an Asian origin [10,54]. The living species of Afrocarpus strongly support an African origin for that genus. The living taxa and fossil record suggest a Gondwanan origin of Retrophyllum, with it evolving by the early Eocene [65,74]. Although the historical biogeographic reconstruction produced by Klaus and Matzke [10] suggests the origin of Dacrydium in New Caledonia, the macrofossil record strongly suggests an Australasian origin [32,39]. Morley [77] also concluded that Dacrydium originated in Australasia in the Late Cretaceous and dispersed to Southeast Asia in the Early Oligocene, probably by island-hopping (e.g., it dispersed to the Ninety East Ridge by the Paleocene and to India by the Early Eocene and later expanded to Japan during the Middle Miocene climatic optimum). According to Wu et al. [36,76], Dacrycarpus also had an Australasian origin during the Late Cretaceous. Dacrycarpus was present by the Eocene in Patagonia, supporting biogeographic connections during the warm Eocene from Patagonia to Australasia across Antarctica [35]. According to Morley [77], Dacrycarpus dispersed to New Guinea from Australia by the late Miocene and then during the mid-Pliocene, it island-hopped to Borneo, and during the Pleistocene, to Sumatra and the Malay Peninsula. However, the Dacrycarpus megafossil from the Miocene of South China shows its earlier arrival to Asia from the Southern Hemisphere and China during Late Miocene [36]. Paleoclimatic studies also support the existence of Dacrycarpus in high-precipitation areas and explain its possible extinction in Australia as that continent dried [36,85]. Dacrycarpus possibly became extinct around the Paleogene–Neogene transition from both South America and Antarctica and during the Neogene from Australia [36]. Klaus and Matzke’s [10] historical biogeographic reconstruction suggests that Falcatifolium originated in the Fiji–New Guinea region around the Late Eocene. However, the fossil record of Falcatifolium from the Middle Eocene of Australia suggests an Australian origin [49], Falcatifolium probably dispersed later to New Caledonia and Papua New Guinea [84].

Klaus and Matzke [10] also concluded that the Prumnopityoid clade originated in New Zealand around the mid-Cretaceous. However, a recent phylogeny of the podocarps suggests an Early to Mid-Jurassic origin for this clade (Figure 1). Leslie et al. [5] and Wang and Ran [84] reported that the phylogenetic divergence of Podocarpaceae shows that the three genera (Lepidothamnus, Podocarpus and Prumnopitys) were dispersed from Australia to South America through Antarctica. A Lepidothamnus macrofossil from the middle Cretaceous of Winton, Queensland [51,88] also supports its Australian origin. The living and macrofossil records of Phyllocladus indicates a Gondwanan origin and wider distribution. Phyllocladus dispersed to New Guinea by the late Miocene and then, during the mid-Pliocene, it island-hopped to Borneo [77]. The extant and extinct species (Halocarpus highstedii from the Oligocene–Miocene) are endemic to New Zealand [39]. Today, Manoao is a monotypic endemic genus to New Zealand but one fossil specimen from the Oligocene (35 Ma) of Cethana, Tasmania (Australia) is similar to that of Manoao colensoi (reported as Lagarostrobos colensoi), showing this genus was once present in Australia [38]. Parasitaxus is a monotypic endemic genus to New Caledonia with no macrofossil records. Lagarostrobos is also a monotypic endemic genus in Tasmania and the macrofossil records from Early Oligocene to Early Pleistocene are also restricted to Tasmania [33,34].

Prumnopitys has three living species distributed in New Zealand and South America. The macrofossil records (Cretaceous–Miocene) demonstrate a Gondwanan origin and wider distribution [43,75]. Although Sundacarpus is now a monotypic genus, the macrofossil records (S. anglica from England and S. tzagajanicus from Russia) from the Uppermost Cretaceous and Eocene show a wider past distribution [75]. Pectinopitys is widely distributed in New Zealand, Australia, New Caledonia, and South America, but with no macrofossil record.

Klaus and Matzke [10] conclude that Acmopyle originated in New Caledonia, but macrofossils from the Eocene–Oligocene suggest a Gondwanan origin [27,28,29,30,31,32]. Microcachrys is now endemic to Australia but is also present in the Oligocene–Miocene of New Zealand [52]. Saxegothaea is the oldest genus in the family and is part of an ancient lineage endemic to South America. Pherosphaera has two living species and two macrofossils from Australia [33].

4.5. Eco-Physiological Adaptations

Most podocarps have evolved flattened leaves and fleshy seed cones, which enable them to survive in low-light conditions beneath the tree canopy and disperse their seeds biotically [85,88,132,133]. Podocarps mature as trees or shrubs. Some of the most significant ecophysiological adaptations and strategies are discussed here.

4.5.1. Seed Cone Morpho-Anatomy

The Podocarpaceae have evolved distinct seed cone morphotypes and display marked variation in functional traits across the 20 genera [88,133,134]. Most podocarp genera produce fleshy seed cones utilizing the epimatium, aril, bracts, receptaculum or a combination of these [109]. Podocarpus is the largest genus in the Podocarpaceae and has a cone composed of one or two seeds covered mostly by a papery and sometimes a fleshy epimatium [10,109]. Several podocarp genera have cones with a brightly colored, fleshy receptaculum [10,88,134].

4.5.2. Leaf Morpho-Anatomy

The Podocarpaceae is prominent in many mixed conifer/broadleaf vegetation types in the Southern Hemisphere, and they exhibit great variation in leaf morphology across the 20 genera [135]. The diversity in leaf morphology of Podocarpaceae is remarkable, ranging from uni-veined needle and scale-like leaves to multi-veined broad leaves. Podocarpaceae foliage can be divided into two main types, imbricate (Dacrycarpus, Dacrydium, Halocarpus, Manoao, Lagarostrobos, Lepidothamnus, Microcachrys, Pherosphaera and Parasitaxus) and broad (flattened) leaved (Acmopyle, Nageia, Afrocarpus, Falcatifolium, Phyllocladus, Podocarpus, Retrophyllum, Pectinopitys, Sundacarpus, Prumnopitys and Saxegothaea). These genera have leaves either spirally arranged or in opposite pairs. Most Podocarpaceae species possess flattened or composite leaves (in 11 genera and more than 140 species) and this may be an adaptation to light requirements, as most of these species grow in the understory of forests with a low-light environment and are unable to reach the canopy level and high sunlight [9] unless a canopy gap occurs. Nageia is characterized by having leaves with multiple parallel veins [55]. All Phyllocladus species have evolved multi-veined phylloclades (Supplementary Figure S1), probably to compete with angiosperms for light [9,82,136]. Acmopyle, Dacrycarpus and Falcatifolium have bilaterally flattened leaves, lacking a true petiole. Leaf dimorphism is present in many genera of Podocarpaceae (Supplementary Figure S2). All other broad-leaved species have bifacially flattened broad leaves [135].

4.5.3. Pollen Morphology

All conifer species are wind-pollinated and those in the Podocarpaceae (except Saxegothaea) and Pinaceae have developed special wing-like structures called sacci [2]. The Podocarpaceae usually have saccate pollen with a tectate exine but usually with a smaller grain than the Pinaceae [137]. Pollen of all genera are bi-saccate except Microcachrys, Pherosphaera and Dacrycarpus, which are tri-saccate, and Saxegothaea which does not have sacci [91,138,139]. Because of this, Erdtman [138] suggested shifting Saxegothaea to the Araucariaceae, while Gaussen [103] and Woltz [140] suggested promoting it to the new family Saxegothaeaceae. The fossil pollen record of the Podocarpaceae is not considered here but is in need of revision, with much important data currently difficult to assess without expert comment on the validity of published interpretations.

4.6. Dispersal Biology

The Podocarpaceae are predominantly zoochorous as their main seed dispersal mechanism, although some genera have other dispersal strategies [141]. The zoochorous mode of dispersal is reported in Dacrycarpus, Halocarpus, Dacrydium, Microcachrys, Afrocarpus, Nageia¸ Podocarpus, Lepidothamnus, Phyllocladus, Parasitaxus, Manoao, Sundacarpus, Falcatifolium, Retrophyllum, Prumnopitys and Pectinopitys [88,134]. Klaus and Matzke [10] reported that 11 genera of Podocarpaceae show endozoochory, two (Prumnopitys and Afrocarpus) epizoochory and seven genera are not ornithochorous. Barochory is present in Pherosphaera and Saxegothaea. Hydrochory and zoochory are reported in Retrophyllum comptonii, R. minor and Lagarostrobos [109].

Leslie et al. [96] reported that cone morphology and seed size are co-evolved in a correlated pattern in animal-dispersed conifers and animal-dispersed species have a relatively larger seed size to attract animals. Similarly, climate change (higher temperatures or water stress in drier conditions) can affect the evolution of cone shape. Interpreting the cone morphology and animal dispersal in Podocarpaceae is difficult because animal-dispersed seeds (fleshy cones) evolved many times in the deep past (from the Cretaceous or even earlier, based on ancestral reconstruction) [88,96,134]. Podocarpus can be interpreted as zoochorous and mainly bird-dispersed due to their colorful fleshy receptacle and epimatium. Bird and bat dispersal have been reported from South African podocarps [142]. The Emu (Dromaius novaehollandiae) is a large bird with a wide distribution range in Australia and it is the main disperser of Podocarpus drouynianus in southwestern Australia, keeping the seeds for up to 50 h in the digestive tract and dispersing them several kilometers [143].

4.7. Ecology of Podocarpaceae

The major Southern Hemisphere conifer family Podocarpaceae is different in morphology, functional physiology, and ecology from the Northern Hemisphere’s major conifer family Pinaceae. Pinaceae are successful in Northern Hemisphere forests, where angiosperms are outcompeted during freezing temperatures, and also occur in low-rainfall areas. Podocarp species are more abundant and compete more successfully with broadleaf angiosperms in the tropical montane forest through multiple morphological and anatomical adaptations but in most cases avoid low-rainfall areas [144]. Ecologically, podocarps have a highly conserved association with the conifer families Araucariaceae and Cupressaceae and with the angiosperm families Nothofagaceae, Winteraceae and Cunoniaceae [9,136]. However, ecological data are lacking for most of the species in these families [4].

Podocarps are unable to bear extreme cold temperate but can tolerate moderate frosts [128] and some exist as alpine shrubs in relatively cold climates (e.g., alpine Tasmania) where permanent snow is uncommon (Figure 4). They possess broad to scale leaves, phylloclades and fleshy cones and they are adapted to a range of conditions from alpine to lowland, understory environments beneath a dense canopy, semi-aquatic (Retrophyllum minus), drought-and fire-prone conditions (Podocarpus drouynianus). The only parasitic gymnosperm (Parasitaxus usta) grows on the roots of another podocarp species (Falcatifolium taxoides). The occurrence of extant species of Podocarpaceae in angiosperm-dominated humid forests is of great interest to ecologists and paleontologists. The Podocarpaceae have preferred wet climates throughout their history [77] and nutrients are a stronger limiting factor for their distribution than the temperature [145], with Coomes and Bellingham [128] reporting that within temperate and tropical rainforests with few exceptions, podocarps are well adapted to nutrient-poor soils.

Coomes and Bellingham [128] evaluated the ecological similarities and differences of temperate and tropical podocarps. They concluded that angiosperm diversification and expansion during the Late Cretaceous was responsible for driving conifers from the lowland tropics and mesic temperate regions due to inferior reproductive competitiveness. However, Bond [146] and Midgley and Bond [147] challenged this view and hypothesized that the physiological traits of conifers (slow seedling establishment and later growth) put them at a disadvantage in competitive regeneration in changing climates (increasing cold and droughts) and habitats (nutrient-poor soil, poorly drained soil, and low light). Podocarps are predominantly slow-growing with low photosynthetic capacity per unit leaf mass and per unit leaf area compared with angiosperms with the same leaf are [128]. The studies that evaluated the growth of podocarps in different habitats lead to the conclusion that podocarp growth is slow compared to other conifers and to angiosperms (e.g., in lowland cool temperate forest, the growth rate is half that of angiosperms [148], and in subalpine shrublands, podocarps grow more slowly than several angiosperm species [149]. In the nutrient-rich soil of southern New Zealand, even tree ferns grow faster than podocarps [150,151].

Brodribb [144] argues that drought is one of the major agents that prevents podocarp success at high altitudes in the Southern Hemisphere. The Late Cenozoic was a major drying period in the temperate region and resulted in the contraction and extinction of Australian and other southern podocarps [152]. The cool and wet conditions (on the continental margins of Gondwana) necessary for the diversification of the Podocarpaceae favor the theory of the drought sensitivity of Podocarps [135,153]. High wood density (that lowers hydraulic efficiency) and leaf sclereids (that collapse under water tension, which results in a loss of hydraulic and photosynthetic function in the leaf) are also present in the broad-leaved tropical podocarps and may be the cause of poorer drought performance and weak competition in drier forests but favor cool, shady, and wet regions of the Southern Hemisphere for podocarp persistence [135,144,154]. In contrast, the Pinaceae have tough and waxy needle-like leaves, lower wood density, fewer sclereids and a high photosynthetic rate, making them resistant and adaptable to drought and freezing temperatures that are common in parts of the Northern Hemisphere [144,155]. This also provides a possible insight into why podocarps are today almost absent from the Northern Hemisphere, despite their potential for long-distance dispersal. A few podocarps are tolerant of drier regions, e.g., Afrocarpus falcatus (southern Africa), Podocarpus drouynianus (Western Australia) and Halocarpus bidwillii, Phyllocladus alpinus and Podocarpus laetus (dry lowland forests of New Zealand) [134]. Podocarp morphology is unusual compared to other conifers, since, despite possessing thick tracheid walls that are vulnerable to embolism at low tensions [154]. (Pittermann et al., 2006b), they also have high hydraulic resistance across pit membranes [156] and that makes the implosion of sclereids in podocarp leaves under tension a real possibility [157].

4.8. IUCN Conservation Status and Threats

The analysis of the available data on the IUCN conservation status of Podocarpaceae shows that 8 species (1 variety) are Critically Endangered (CR), 27 species (2 varieties) are Endangered (EN), 23 species (one subspecies) are Vulnerable (VU), 3 species are Threatened (TH), 33 species (2 varieties) are Near Threatened (NT), 89 species (8 varieties and one subspecies) are Least Concern (LC), 10 species are Data Deficient (DD) and 7 species (2 hybrids) are Not Evaluated (NE) for IUCN status (Figure 5). The Critically Endangered (CR) species are Acmopyle sahniana (Fiji), Pherosphaera fitzgeraldii (Australia), Dacrydium guillauminii (New Caledonia), Podocarpus urbanii (Jamaica), P. costaricensis (Costa Rica and Panama), P. decumbens (New Caledonia), P. palawanensis (the Philippines), P. perrieri (Madagascar) and P. sellowii var. angustifolius (Brazil). The IUCN conservation status for tropical podocarps states that 5 species are considered critically endangered, 18 species are endangered, and 16 species are vulnerable (Cernusak et al., 2011). The New Caledonian podocarp species are facing serious conservation threats due to their restricted populations (Enright and Jaffré, 2011); i.e., Retrophyllum minus (endangered), Podocarpus decumbens (critically endangered) P. longefolaliatus (endangered), Dacrydium guillauminii (critically endangered), Acmopyle pancheri (nearly threatened) and Parasitaxus usta (vulnerable).

Figure 5.

The proportion of current IUCN conservation status of Podocarpaceae species. The conservation status is evaluated according to IUCN Red List categories and criteria, version 3.1 (IUCN Council, Geneva, Switzerland).

Deforestation associated with mining, expansion of tropical agricultural activities and other anthropogenic activities poses a serious threat to tropical podocarps [158]. Deforestation and climate change are also posing a serious threat to montane endemic podocarps [159]. Similarly, more extreme dry seasons are also damaging for tropical podocarps because they are drought and fire intolerant [158]. Wildfire is posing a huge threat to Australian podocarps (Figure 6) and in some areas, the podocarp population has been driven to extinction by these fires [160]. The harvest of podocarp timber has been an important industry, but their slow growth makes it detrimental and unsustainable for the species involved [161]. Mill [162] reported habitat loss, climate change and deforestation as major threats causing the extinction of Podocarpus species. Failure of regeneration and aging of the current populations are two major threats for at least some podocarp species [128,163,164].

Figure 6.

A wildfire in 2020 burnt the Tahune rainforest, Tasmania. This photo is of a burnt Celery-top Pine (Phyllocladus aspleniifolius) tree.

4.9. Current Gaps and Future Perspectives

Some clear gaps still exist that need to be filled in order for us to gain a better understanding of the Podocarpaceae and include some of the following aspects:

1. Descriptions and taxonomic treatments of several species from less explored/remote areas such as Papua New Guinea, Malaysia, Indonesia, and New Caledonia are based only on collections of one or a few specimens. Additionally, some of these areas are not well explored and may contain undescribed species.

2. Field-and laboratory-based studies on pollination biology, the reproductive cycle and anatomical structures are not well developed for most podocarps and require further detailed evaluation.

3. Extensive research is required to understand why Podocarpaceae have such remarkable seed cone and leaf morphology.

4. Very few studies report the dispersal biology of podocarp seeds and comprehensive assessments are required to understand the dispersal biology and ecology of podocarps.

5. Despite the several high-quality publications on the leaf cuticle morphology of various genera, a good quality publication is necessary that describes the taxonomic and phylogenetic authenticity of these foliar cuticular diagnostic characters. Similarly, studies are required to assess the infraspecific variation in the leaf morphology for different populations.

6. Phylogenomic and population-based studies are available only for a few Podocarpus species (P. matudae, P. nubigenus, P. parlatorei, P. salignus, P. latifolius, P. guatemalensis and P. oleifolius), and with fairly limited geographic scope (the Americas). With the availability of modern NGS techniques and bioinformatic tools, more comprehensive studies are required to unveil their phylogeny, historical biogeography, speciation, and population history.

7. Only a few studies are available on the historical biogeography of Podocarpaceae and the discovery of new podocarp fossils from the Early Permian (Paleozoic) of Jordan [86,87] questions the Gondwanan origin of the Podocarpaceae. The inclusion of well-placed podocarp fossils will help in better understanding the reconstruction of historical biogeography.

8. Comparative studies of the three Southern Hemisphere conifer families (Araucariaceae, Cupressaceae and Podocarpaceae) to evaluate the impact of these families on the habitats they occupy and their relationships with the rest of the Southern Hemisphere biota.

9. Evolution of photosynthetic units in these three families in response to the closed forests that predated the rise to dominance of the angiosperms and angiosperm-dominated rainforests and then the major aridification of the Southern Hemisphere.

10. A better understanding of the response of podocarp foliage to drought stress and the adaptations that have evolved to deal with the constraint of most podocarps in having only a single vein per leaf is required to better understand the distribution and ecology of the family.

11. Use of species distribution modelling to predict the possible ecological niche and the effect of climate change on species range dynamic.

12. A better understanding of the evolutionary history and biology, ecology and life history are important in conservation efforts, given that so many species are threatened.

5. Conclusions

The current study provides a comprehensive overview on the systematics, diversity, hotspots, evolutionary adaptations, and conservation status of podocarps. Podocarps are morphologically more diverse compared to other conifer families and the updated phylogeny based on more extensive macrofossil records broadens our understanding of the evolutionary history of Podocarpaceae. Most podocarp genera currently exhibit low species richness and high endemism and often have disjunct distributions. Today, the Malesian region is the diversity hotspot for living podocarp taxa. However, the fossil record demonstrates wider distributions in the past. Podocarpus, Dacrydium and Dacrycarpus are the most dominant genera (approximately 75% of living podocarps) and have acquired particular morpho-anatomical adaptations that help them to survive in tropical forests. Podocarps demonstrate a remarkable seed cone and leaf diversity compared with other conifers. The genera with fleshy seed cones predominantly rely on bird dispersal. Podocarps are facing serious threats from deforestation, climate change, drought and wildfire, and the need for further targeted research is urgent. Among the conifers, podocarps are less well known and receive less attention than their counterparts that dominate the Northern Hemisphere, despite their remarkable morphological diversity and long evolutionary history.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/plants12051171/s1. Figure S1. Phyllocladus aspleniifolius (with phylloclades) found in rainforest Tasmania. Figure S2. Leaf dimorphism in Dacrycarpus dacrydioides. Table S1. Fossil taxa used for calibration of the phylogeny [165,166].

Author Contributions

Conceptualization, R.K. and R.S.H.; methodology, E.B. and R.K.; software, E.B. and R.K.; validation, R.K., E.B. and R.S.H.; formal analysis, R.K., E.B. and R.S.H.; writing—original draft preparation, R.K.; writing—review and editing, E.B., J.L. and R.S.H.; supervision, R.S.H. All authors have read and agreed to the published version of the manuscript.

Data Availability Statement

Data available in article supplementary material. Additional supporting information may be found in the online version of the article at the publisher’s website.

Conflicts of Interest

The authors declare no conflict of interest.

Funding Statement

This research was funded by the Key Research Program of Frontier Sciences, CAS (ZDBS-LY-7001), National Natural Science Foundation of China (41971071, 42211540718), CAS “Light of West China” Program, and Top-notch Young Talents Project of Yunnan Provincial “Ten Thousand Talents Program” (YNWR-QNBJ-2018-146).