Fossil/Lineage Geographic origin

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Table S5. Fossils integrated into Ancestral Area Reconstruction (AAR) analyses for the Cupressaceae. Fossils are listed from older to younger and they were subdivided into 27 groups (at the first row of each group listed their names) by thick horizontal lines according to their closest living lineage/phylogenetic positions and region where they have been found. Fossil groups 2 to 27 and the three fossils within group 1 (marked with asterisks) were integrated into AAR analyses as 29 independent extinct lineages. For absolute age of each geological stratum, we relied on the latest geologic time scale (1). An asterisk (*) marks lineages named after fossil names and the dagger symbol (†) indicates that the fossil was not included in AAR analyses. Ma: million years ago.


Geographic origin



Type of fossil

Sister to

Assigned ages (Ma)

1. Fossil taxa assigned to basal grade of Cupressaceae

*Austrohamia minuta

S America

(NW Chubut, Argentina)


Leafy shoots, seed cones and pollen cones of this fossil share morphological characters with living Cunninghamia and Taiwania (2). The fossil comes from the Cerro Bayo locality, originally dated to the Middle to Late Jurassic, but recent evidence suggests an Early Jurassic age (2). We accepted the upper boundary of the Bajocian (167.7±3.5 Ma) (1) as the age of this taxon (2). Austrohamia minuta is the type species based on which the genus was first erected (see below under Hughmillerites for further justification of its placement).

A. acanthobractea in all three scenarios


*Austrohamia acanthobractea

Asia (Inner Mongolia, China)

Middle to Late Jurassic

Shared vegetative and reproductive traits between this fossil taxon and the type species Austrohamia minuta justify its assignment to the genus (3) (see also Hughmillerites, below). The fossil comes from the Tiaojishan Formation, Inner Mongolia, China, dated to 152-168 Ma based on stratigraphy and radioisotopes (3).

A. minuta in all three scenarios


*Sewardiodendron laxum

Asia (W Henan, China)

Middle Jurassic

This fossil taxon was reconstructed based on leafy shoots and female and male cones, and has its closest affinities with a group of Cunninghamia-like conifers (4) (see Hughmillerites below). The fossils Sewardiodendron laxum sensu Florin (5) and Elatides williamsonii (6-8), both from England, are of fragmentary nature and we therefore did not use them in our AAR analyses. Austrohamia minuta, A. acanthobractea and Sewardiodendron were placed as relatives to the extant Cunninghamia in our molecular phylogenetic analyses (“total evidence” analyses, see SI text, Fig. S5h). Since the AAR approaches that we used require fully dichotomous tree, we tested the three plausible phylogenetic positions of these fossils and performed likelihood and parsimony AARs for each scenario (see M & M section, Fig. 2c-e). Scenario 1: (((Sewardiodendron, Cunninghamioideae), Austrohamia), all other Cupressaceae); Scenario 2: (((Sewardiodendron, Austrohamia), Cunninghamioideae), all other Cupressaceae); Scenario 3: (Cunninghamioideae, ((Sewardiodendron, Austrohamia), all other Cupressaceae).

Scenario 1: Cunninghamioideae;

Scenario 2: Austrohamia; Scenario 3: Austrohamia


Hughmillerites juddii

Europe (Black Isle, NE Scotland, U.K.)

Late Jurassic

This fossil taxon was described based on a group of seed cones from the Lower Kimmeridgian (9). A cladistic analysis based on morphological and anatomical characters supports its placement among the basal grade of Cupressaceae (9). Our “total evidence” analysis, which included representatives from all living genera of Cupressaceae and Hughmillerites juddii (see SI text, Fig. S5b), showed this taxon as a member of basal Cupressaceae but without support. We therefore excluded it from AAR analyses.



2. Cunninghamia related fossils from Europe

Cunninghamites lignitum

Europe (Czech Republic)


Leaves of this fossil taxon are acicular, flattened, amphistomatic and helically arranged, and its ovuliferous cones consist of helically arranged peltate cone scales, each bearing four seeds (10). Among living Cupressaceae, this fossil is most similar to Cunninghamia konishii and C. lanceolata (10-11).



3. Cunninghamia related fossils from Asia

Cunninghamiostrobus yubariensis

Asia (Hokkaido, Japan)


The ovulate cone of this conifer bears numerous cone scales in a close helix, each with three ovules per scale, resembling extant Cunninghamia female cones (12).



4. Cunninghamia N American fossils

Cunninghamia hornbyensis

N America (British Columbia, Canada)


This conifer was reconstructed from leafy twigs. Its leaves are most like those of extant Cunninghamia in size, shape, and anatomy including the structure of vascular bundle and transfusion tissue (11).



Cunninghamia spec.

N America (Alberta, Canada)


Morphological as well as anatomical characters of leaves, seed cones and wood allow an assignment to the living genus Cunninghamia (13). The fossils have helically arranged leaves; resin canals were centrally located with a single adaxial vascular bundle and two smaller lateral canals (13). The bract margins are finely denticulate like those in extant C. konishii and C. lanceolata (13).



5. Taiwania N American fossils N America

Taiwania cf. T. cryptomerioides

N America (Alaska, U.S.A.)

Early Cretaceous


The leaves and seed cones of this fossil taxon are indistinguishable from those of the living genus Taiwania (14).



6. Taiwania Asian fossils

Taiwania cretacea

Asia (Arkagala River, Russia)

Late Cretaceous


Comparisons of leaf-bearing branches, seed cones and cuticle micromorphology with fossil and extant species of Taiwania suggest that this taxon represents a new fossil species of Taiwania (15).



7. Taiwania European fossils Europe

Taiwania schaeferi

Europe (Svalbard, Norway)


This fossil seed cone and associated leaves are similar to the extant Taiwania cryptomerioides (16) (and T. cryptomerioides from the Pliocene of Japan) (17).



8. Athrotaxis related fossils

*Athrotaxites berryi

N America (Kootenai Formation, Montana, U.S.A.)


Although leafy twigs and female and male cones of this conifer have Athrotaxis-like characters, they still differ from the living genus (18). Our “total evidence” analysis suggested that this taxon is a sister lineage of the extant Athrotaxis (SI text, Fig. S5h).



9. Athrotaxis S American fossils

Athrotaxis ungeri

S America (Santa Cruz, Argentina)

Early Cretaceous (early Albian)

This fossil taxon has helically arranged ovate and scale-like leaves and ovulate cones with a distinctive cone scale, similar to the extant A. cupressoides (19).



10. Metasequoia Northern Hemisphere fossils

Metasequoia occidentalis

Northern Hemisphere

Late Cretaceous to Pliocene

Metasequoia is unique in its decussately arranged cone scales and deciduous foliage shoots. Leafy twigs and female and male cones of this taxon are common throughout the Northern Hemisphere (20-23).



Metasequoia foxii

N America (central Alberta, Canada)


This whole-plant reconstruction (including seedlings) was made from compression remains. The fossil taxon shares many characters with the living species M. glyptostroboides (24).



Metasequoia milleri

N America (Princeton, B.C., Canada)


Leafy shoots and pollen cones of this taxon differ from the living species in a few traits, while seed cones cannot be distinguished from those of living species (25-27).



11. Sequoia and Sequoiadendron related fossils

Sequoia-like cone

Asia (Japan)

Late Cretaceous

This seed cone most closely resembles Sequoia and Sequoiadendron, but differs in having an abaxially concave reniform vascular cylinder traces to the bract-scale complex (28).

Sequoia and Sequoiadendron


Quasisequoia florinii

Europe (Scania, Sweden)

Late Cretaceous

While leaves of this fossil are most similar to Sequoiadendron, its seed cones share characters with Sequoia and Sequoiadendron (29).

Sequoia and Sequoiadendron


Sequoia reichenbachii

Northern Hemisphere

Late Cretaceous

These falcate leaves, which are tetragonal in cross sections with three canals, were widespread in the Northern Hemisphere during the Cretaceous (30-32).

Sequoia and Sequoiadendron


12. Austrosequoia

Austrosequoia wintonensis

Australia (Winton, Queensland, Australia)

Late Cretaceous


This conifer was reconstructed from a fertile branchlet. Its cone has 29-49 cone scales arranged helically around the axis. Each scale has four to seven ovules arranged in a single row. Reproductive shoot leaves are rhomboidal with an incurved apex and a distinct keel. Comparison with extant “taxodiaceous” genera (Cupressaceae genera except Cupressoideae and Callitroideae) reveals a strong similarity to extant monotypic Sequoia and Sequoiadendron (33).

Sequoia and Sequoiadendron


13. Cryptomeria European fossils

Cryptomeria rhenana

Europe (W Germany)


Reconstructed from compressed leafy shoots and seed cones, this fossil differs from the living C. japonica in several characters but clearly represents the genus Cryptomeria (30, 34).



14. Glyptostrobus N American fossils

Glyptostrobus spec.

N America (British Columbia, Canada)

Late Cretaceous (Cenomanian)

A cone of this fossil taxon from the Dunvegan Formation is obovate and was borne terminally on leafy shoots that resemble G. pensilis. It possesses a prominent reflexed mucro (plural: mucri) near the middle of the bract-scale complex and ovuliferous scales that exceed the length of the bracts (35). Its ovuliferous scales are composed of multiple lobes, with each lobe bearing a small apical hook along the outer margin that contributes to the prominence to the lobes (35). These features support its assignment to Glyptostrobus.

Glyptostrobus and Taxodium


15. Glyptostrobus European fossils

Glyptostrobus dunoyeri

Europe (Scotland, U.K.)


This fossil taxon was reconstructed based on leaves and seed cones, and both macro-morphology (of leaves and cones) and micro-morphology (of cuticles) are similar to the living G. pensilis (36).



16. Taxodium N American fossils

Taxodium wallissii

N America (Alberta, Canada)

Late Cretaceous (Maastrichtian)

This conifer was reconstructed from three-dimensionally preserved branches bearing dimorphic leaves, pollen cones, pollen, seed cones, and seeds. Comparisons with extant representatives of the subfamily Taxodioideae indicate that it is most similar to Taxodium, but also resembles Glyptostrobus and Cryptomeria in some features (35).



17. Taxodium European fossils

Taxodium balticum

Europe (Leipzig, Germany)


Reconstructed from foliage shoot, a female cone, and a male inflorescence, this conifer is assignable to Taxodium but differs from fossil and extant congeneric species in the cone scale complex (37).



18. Papuacedrus S American fossils

Papuacedrus prechilensis

S America (Patagonia, Argentina)


Nearly all features of foliage and seed cones of this fossil match extant Papuacedrus (Moluccas and New Guinea), but the fossil comes from S America (38).



19. Austrocedrus Australian fossils

Austrocedrus australis

Australia (Tasmania)

Early Oligocene

Seed cones of this fossil are similar to those of A. chilensis. Their bract-scale pairs are similar in shape, the bracts are large, and the bract projections are short and close to the apex of the outer scales. The fossil taxon’s tapered bases of the ultimate bract-scale pair and the position of the penultimate bracts beneath this tapered section are almost identical to those of A. chilensis. The only major difference between A. chilensis and the fossil is the small size of the latter (39).



20. Fitzroya Australian fossils

Fitzroya acutifolia

Australia (Tasmania, Australia)

Early Oligocene

This conifer has a terminal ovulate cone with bract-scale complexes in two whorls of three, and foliage consisting of alternate whorls or near-whorls of three scale-like, lanceolate leaves, each with a prominent midrib (40). These traits place it in the extant genus Fitzroya.



21. Thuja N American fossils

Thuja polaris

N America (Nunavut, Canada)

Middle Paleocene

Cones and leaves of this fossil taxon are similar to the extant Thuja plicata except that its cones have twice as many cone scale complexes (41).



22. Thuja European fossils

Thuja ehrenswaerdi

Europe (Greenland)


This conifer has cones similar to those of the extant Thuja sutchuenensis but its leaves are more similar to the extant T. plicata (42).



23. Chamaecyparis N American fossils

Chamaecyparis corpulenta

N America (Vancouver Island, B.C., Canada)

Late Cretaceous (Santonian)

Although leafy shoots of this fossil taxon are similar to those of Thuja and Chamaecyparis, its seed cones are woody and four-scaled and were borne at the end of leafy branches similar to those of Chamaecyparis (43). Because Cupressaceae seed cone provide a more secure identification than vegetative organs (30), this fossil here placed as sister to the extant Chamaecyparis -Fokienia clade.

Chamaecyparis -Fokienia lineage


24. Chamaecyparis European fossils

Chamaecyparis salinarum



Seed cones of this fossil taxon closely resemble the extant C. obtusa, which occurs in Japan and Taiwan (44). This fossil is common throughout Central European Middle Miocene to Pliocene strata (45).

Chamaecyparis obtusa


25. Fokienia N American fossils

Fokienia ravenscragensis

N America (SW Saskatchewan, Canada)

Early Paleocene

The cones of this fossil taxon are borne in opposite pairs, with 8-10 woody, decussate, peltate scales. Its foliage branches were borne oppositely and bear thin, flattened leaves in whorls of four. Although branching pattern and leaf morphology of this conifer differ from those of extant Fokienia, similarity in seed cone structure allows assignment to this genus (46-47).



26. Calocedrus European fossils

Calocedrus suleticensis

Europe (Czech Republic)


Seed cones of this fossil taxon differ from those of the extant Calocedrus species by more a pronounced basal pair of cone scales with a subterminal mucro (48). This taxon is considered as an extinct sister species to Asian Calocedrus species (rather than N American species due to biogeographic consideration) (49-51).



27. Tetraclinis N American and European fossils

Tetraclinis salicornioides

N America (Western U.S.A.) & Europe

Eocene to Pliocene

Although cones and seeds of this fossil taxon are closely similar to those of the extant T. articulata, its foliage is more “spreading” and composed of flattened segments with fused facial and lateral leaves (52).




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