Environmentally driven extinction and opportunistic origination explain fern

diversification patterns

Samuli Lehtonen*1,2†, Daniele Silvestro*3,4,5,6†, Dirk Nikolaus Karger2,7, Christopher Scotese8, Hanna
Tuomisto2, Michael Kessler7, Carlos Peña2, Niklas Wahlberg2,9, Alexandre Antonelli3,4,10

1
Herbarium, Biodiversity Unit, University of Turku, 20014 Turku, Finland
2
Department of Biology, University of Turku, 20014 Turku, Finland
3
Department of Biological and Environmental Sciences, University of Gothenburg, Carl Skottsbergs
gata 22B, Gothenburg 413 19, Sweden
4
Gothenburg Global Biodiversity Centre, Box 461, SE-405 30 Göteborg, Sweden
5
Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland
6
Swiss Institute of Bioinformatics, Quartier Sorge, 1015 Lausanne, Switzerland
7
Department of Systematic and Evolutionary Botany, University of Zurich, 8008 Zurich, Switzerland
8
Earth and Planetary Sciences, Northwestern University, Evanston, IL, USA
9
Department of Biology, Lund University, Lund, Sweden
10
Gothenburg Botanical Garden, Carl Skottsbergs gata 22A, Gothenburg 413 19, Sweden

†
These authors contributed equally to this work.

*Correspondence to: [email protected] or [email protected]

1
Supplementary Information

Supplementary Materials and Methods

PaleoDEMs

The paleogeographic digital elevation models1 were built on the basis of digital topographic and

bathymetric datasets of the modern world2–4. These data were combined into a global dataset with 6-

minute resolution. The individual grid cells were rotated to their paleopositions using the PALEOMAP

Global Plate Tectonic Model5, and the modern digital topographic and bathymetric values were

corrected and modified using the lithofacies and paleoenvironmental data. These data were constructed

using the database of Paleogeographic Atlas Project6–8 supplemented by additional lithological and

paleoenvironmental records for the Permian and Jurassic9,10 in combination with numerous regional

and global paleogeographic atlases. The plate tectonic reconstructions11 were used to model the

expected changes in topography and bathymetry caused by plate tectonic events, such as sea floor

spreading, continental rifting, subduction, and continental collision, as well as other isostatic events

such as glacial rebound12.

Fossil calibration priors for molecular dating

1. Ferns

Hard minimum age constraint: 359 Ma. Archaeocalamites from Upper Devonian to Lower Permian13,14

is considered as a member of the horsetail lineage15 and used in several studies to constrain the first

divergence within ferns16–18. We follow this practice to constrain the minimum age of crown group

ferns.

Soft maximum age constraint: 407 Ma. Oldest known Euphyllophytina are from Lower Devonian19,

and we used this age as the soft maximum age for the ferns.

2
2. Equisetum

Hard minimum age constraint: 359 Ma. Archaeocalamites is considered a member of the horsetail

lineage15 and we used their first appearance to constrain the minimum age of the Equisetum stem

lineage.

Soft maximum age constraint: 407 Ma. We used the oldest known Euphyllophytina19 as the soft

maximum for the stem lineage.

3. Tmesipteris

Hard minimum age constraint: 23 Ma. The Oligocene fossil Tmesipteris tasmanica is lacking

reproductive structures, but represents the distinctive vegetative characteristics of Tmesipteris20. We

assigned the constraint for the crown group Tmesipteris because the fossil is associated with extant

species having indefinite growth type20.

Soft maximum age constraint: 145.5 Ma. Whisk ferns have extremely poor fossil record (Collinson

1996), but are among the first diverging fern lineages15,17,21. In the lack of better fossil constraint we

applied the beginning of Cretaceous as the soft maximum age for this node.

4. Botrychium

Hard minimum age constraint: 55.8 Ma. Fossil trophophores, sporophores and spores were assigned to

the genus Botrychium with high confidence22. The fossil was considered remarkably similar with extant

B. virginianum22, a species that was resolved together with B. strictum as the sister lineage of all other

living Botrychium21. A detailed cladistic analysis incorporating morphological characters would be

needed to verify whether B. wrightonii represents a crown group Botrychium, or a stem group. We took

a conservative approach and allowed the fossil to represent the stem group.

Soft maximum age constraint: 201.6 Ma. Ophioglossales have extremely poor fossil record22, but are

among the first diverging fern lineages21. The split between Botrychium and Ophioglossum have

3
previously been estimated to have occurred ca. 162 Mya17. Without a better judgement we applied the

Triassic-Jurassic boundary as the soft maximum age for the stem lineage.

5. Marattiaceae

Hard minimum age constraint: 318 Ma. Marattioid fossils are abundant from Pennsylvanian onwards,

but poorly preserved specimens are known already from the Mississippian13. We assigned the

constraint to the Marattiaceae stem lineage.

Soft maximum age constraint: 407 Ma. We used the oldest known Euphyllophytina19 as the soft

maximum for the stem lineage, as the ancestors of marattioids remain unknown23.

6. Ptisana

Hard minimum age constraint: 176 Ma. Marattia asiatica is here considered as a member of segregate

genus Ptisana24. The species has sessile, deeply cut bivalve synangia with thick outer walls25,

characters typical of Ptisana24. These characters may represent plesiomorphies rather than

apomorphies26. However, Middle Jurassic Angiopteris blackii has free synangia and trilete spores27,

which are considered to be apomorphic states26. Therefore, it appears that Ptisana and Angiopteris were

separated by the end of the Early Jurassic. We constrained the Ptisana stem lineage to have originated

by then.

Soft maximum age constraint: 318 Ma. A cladistic analysis based on morphological data suggested that

the Early Permian Millaya may belong to crown group marattioids23. We applied the beginning of

Pennsylvanian as the soft maximum age for the stem lineage, as marattioids became common during

this period, but modern lineages apparently had not evolved yet23.

7. Angiopteris

Hard minimum age constraint: 172 Ma. We use the Middle Jurassic Angiopteris blackii27 to constrain

the age of the stem lineage.

4
Soft maximum age constraint: 318 Ma. We applied the same soft maximum age constraint for

Angiopteris as for Ptisana.

8. Leptosporangiate ferns

Hard minimum age constraint: 345 Ma. Tournaisian sporangia of Senftenbergia-type are placed in

Tedeleaceae28, a group of ferns resolved as stem group leptosporangiates15. We assigned the constraint

to the leptosporangiate stem lineage.

Soft maximum age constraint: 407 Ma. Oldest known Euphyllophytina are from Lower Devonian19,

and we used this age as the soft maximum age for leptosporangiate stem lineage.

9. Osmundaceae

Hard minimum age constraint: 271 Ma. Early Permian Grammatopteris is generally considered as an

early member of osmundaceous ferns16,17,29–31. We assigned the constraint to the Osmundaceae stem

lineage.

Soft maximum age constraint: 345 Ma. The hard minimum age of leptosporangiates was used as the

soft maximum age for Osmundaceae stem lineage.

10. Osmunda

Hard minimum age constraint: 201.6 Ma. The Triassic fossil Osmunda claytoniites was described as

remarkably similar to modern Osmunda32. The fossil shows medially positioned hemidimorphic leaves

which are characteristic to Osmunda subgenus Claytosmunda, thus providing a minimum age for the

genus32,33. We used this fossil to constrain the crown group age.

Soft maximum age constraint: 299 Ma. Members of Osmundaceae appeared first time in the

Permian34,35.

5
11. Osmundastrum

Hard minimum age constraint: 65.5 Ma. Fossils indistinguishable from the modern Osmundastrum

cinnamomeum are known from the Upper Cretaceous36. We assigned the constraint to the stem lineage

of the clade representing two O. cinnamomeum varieties in our analysis.

Soft maximum age constraint: 299 Ma. The soft maximum age of Osmunda was applied to the stem

lineage of the subgenus.

12. Osmunda subg. Plenasium

Hard minimum age constraint: 55.8 Ma. The Paleocene Osmunda arnoldii is very similar to extant

species of subgenus Plenasium34,37. We assigned the constraint to the stem lineage of subgenus

Plenasium.

Soft maximum age constraint: 299 Ma. The soft maximum age of Osmunda was applied to the stem

lineage of the subgenus.

13. Todea

Hard minimum age constraint: 136 Ma. Phylogenetic analysis combining morphological data and rbcL

sequences resolved the fossil species Todea tidwellii as a member of Todea35. Todea and Leptopteris

are morphologically readily distinguishable and strongly supported as each others closest relatives in

molecular phylogeny33, and we used this fossil to constrain the Todea-Leptopteris node excluding the

stem.

Soft maximum age constraint: 299 Ma. Members of Osmundaceae appeared first time in the

Permian34,35.

14. Hymenophyllaceae

Hard minimum age constraint: 228 Ma. Hopetedia praetermissa is the oldest known member of

Hymenophyllaceae based on its general frond morphology, indirect evidence of a filmy habit, and soral

6
position and morphology38. This fossil has been used to constrain filmy fern origin previously as well16.

We assigned the constraint to the Hymenophyllaceae stem lineage.

Soft maximum age constraint: 318 Ma. Hymenophyllites quadridactylites from the Upper

Carboniferous is the oldest known putative Hymenophyllaceae38, and the beginning of Pennsylvanian

is used here as the soft maximum age for the stem lineage.

15. Matoniaceae

Hard minimum age constraint: 235 Ma. Structurally preserved sori and sporangia of Tomaniopteris

katonii has a set of characters unique to Matoniaceae39. This constraint has been applied also in

previous studies31. We assigned the constraint to the Matoniaceae stem lineage.

Soft maximum age constraint: 299 Ma. Oligocarpia and Szea from Permian to Triassic have been

considered as possible stem group members of Gleicheniaceae, Matoniaceae, Dipteridaceae, or

Hymenophyllaceae16. The Carboniferous-Permian boundary was used here as soft maximum age for

the stem lineage.

16. Gleicheniaceae

Hard minimum age constraint: 99.6 Ma. Morphology based cladistic analysis resolved Gleichenia

chaloneri as the sister of extant Diplopterygium glaucum (=Gleichenia glauca)40. However, that

analysis suffered from poor taxon sampling and failed to cover the extant diversity within

Gleicheniaceae. As well, their results are different from those of a previous study41 and neither one of

these studies is supported by the molecular results21. Better sampled morphological data matrices

would be needed to verify the exact position of Gleichenia chaloneri and another fossil taxa,

Boodlepteris turoniana41, within the family. However, it seems apparent that both of these fossil taxa

represent members of crown group Gleicheniaceae, and following a previous study31 we used the older

one of them (G. chaloneri) to constrain the Gleicheniaceae crown group.

Soft maximum age constraint: 299 Ma. Justification as for the Matoniaceae.

7
17. Stromatopteris

Hard minimum age constraint: 89.3 Ma. Morphology based cladistic analyses40,41 resolved

Boodlepteris as the sister of extant Stromatopteris. Both of these studies suffered from poor taxon

sampling and failed to cover the extant diversity within Gleicheniaceae and their topology is not

supported by the molecular results. However, Boodlepteris and Stromatopteris share anatomical

apomorphies not observed in other members of the family, and we follow a previous study31 in using

this calibration point. We applied the constraint for the Stromatopteris moniliformis–Gleichenella

pectinata node.

Soft maximum age constraint: 299 Ma. The soft maximum age of the family was applied to this node.

18. Lygodiaceae

Hard minimum age constraint: 168 Ma. It has been suggested that mid-Jurassic Stachypteris provides a

minimum age for the divergence between Lygodium and Anemia-Schizaea42. Similarities in fertile

structures suggests affinity with modern Lygodium, despite the highly different frond morphology and

habit42. We assigned the constraint to the Lygodiaceae stem lineage following a previous study16.

Soft maximum age constraint: 299 Ma. Fossil evidence suggests that the second filicalean radiation,

into which Lygodiaceae belongs, began in Permian time43. Carboniferous-Permian boundary is applied

here as the soft maximum age for the stem lineage.

19. Schizaea

Hard minimum age constraint: 112 Ma. Fossil Schizaeopsis is very similar to living Schizaea in frond

morphology, but differs in its spore morphology42,44,45. Schizaeopsis has been considered as the sister

lineage of Schizaea42. The oldest known fossils are from the Lower Cretaceous45. We assigned the

constraint to the Schizaea stem lineage.

8
Soft maximum age constraint: 251 Ma. Spores with a possible affinity with Schizaea first appear in the

Triassic42 and consequently we applied the beginning of Triassic as the soft maximum age for the stem

lineage.

20. Anemia

Hard minimum age constraint: 136 Ma. The anatomically preserved fossils were assigned to the genus

Anemia, and compared most closely to subgenus Anemiorrhiza46. Anemiorrhiza is the sister group to a

clade comprising Mohria and the rest of the Anemia42. Because the fossil differed from Mohria and

presented characters suggesting affinity with Anemiorrhiza46, we used it to constrain the basal split of

Anemiorrhiza from the remaining clade. Further evidence to support this calibration point is provided

by Cretaceous Pelletixia, Ruffordia and Anemia fossils which have been variously resolved within the

modern Anemia subgroups in phylogenetic analyses42,47,48. This node was constrained as Valanginian

also in a previous study31. We assigned the constraint to the Anemia stem lineage.

Soft maximum age constraint: 251 Ma. Anemia and Schizaea are sister groups42 and we apply the same

soft maximum age for both of them.

21. Marsileaceae

Hard minimum age constraint: 140 Ma. Late Jurassic-Early Cretaceous Regnellites was resolved as a

member of Marsileaceae in a cladistic analysis of morphological data49. This calibration has also been

used before31. We assigned the constraint to the Marsileaceae stem lineage.

Soft maximum age constraint: 201.6 Ma. Oldest fossils possible related to Marsileaceae are Late

Jurassic microspores50,51. We therefore applied Triassic-Jurassic boundary as the soft maximum age for

the stem lineage.

22. Marsilea

Hard minimum age constraint: 99.6 Ma. Marsileaceaephyllum mahisensis was nested within Marsilea

in cladistic analysis of morphological data51. Taxon sampling was poor in that analysis, but the

9
relationships within Marsilea were in accordance with the molecular hypothesis52. We assigned the

constraint to the Marsilea stem lineage.

Soft maximum age constraint: 145.5 Ma. Based on fossil record it has been estimated that modern

Marsileaceae genera started to differentiate at mid-Cretaceous50. We used the beginning of Cretaceous

as the soft maximum age for the stem lineage.

23. Regnellidium

Hard minimum age constraint: 83.5 Ma. Fossils from the Late Cretaceous were assigned to

Regnellidium based on preserved sporocarp, megaspore and microspore characters50. This calibration

point has been used also in a previous study31. We applied this constraint to the Regnellidium

diphyllum–Pilularia globulifera node.

Soft maximum age constraint: 145.5 Ma. Based of fossil record it has been estimated that modern

Marsileaceae genera started to differentiate at mid-Cretaceous50 and we assigned the beginning of

Cretaceous as the soft maximum age for this node.

24. Azolla

Hard minimum age constraint: 83.5 Ma. Earliest fossil Azolla megaspores appear in the Santonian53.

We applied this constraint for the Azolla stem lineage.

Soft maximum age constraint: 201.6 Ma. The soft maximum age of Marsileaceae (sister family of

Salviniaceae, in which Azolla belongs) was applied as the maximum age for Azolla stem lineage.

25. Azolla section Azolla

Hard minimum age constraint: 13.8 Ma. The minimum age of this section was constrained following a

previous study54. The age constraint was applied to the section including the stem.

Soft maximum age constraint: 33.9 Ma. All modern Azolla share a collared megaspore, a character that

is observed in the fossil record first time in the Oligocene54. The age constraint was applied to the stem

lineage.

10
26. Loxsomataceae

Hard minimum age constraint: 112 Ma. Relationship of the Aptian Loxsomopteris anasilla rhizome

with Loxsomataceae was suggested by similarity of the hairs covering the rhizome55. We assigned the

constraint to the Loxsomataceae stem lineage.

Soft maximum age constraint: 251 Ma. Triassic fossils have often been associated with tree ferns56,

although it is possible that they represent extinct stem lineages of tree ferns or tree ferns and

Polypodiales17. Molecular clock suggested that the divergence between Polypodiales and tree ferns

occured in Late Triassic16. We apply the beginning of Triassic as the soft maximum age for the tree

ferns.

27. Lophosoria

Hard minimum age constraint: 112 Ma. Lophosoria cupulata from the Aptian of Antarctica is placed in

the extant genus57 and was used here to constrain the split between Lophosoria and Dicksonia.

Soft maximum age constraint: 251 Ma. We apply the beginning of Triassic as the soft maximum age

for the tree ferns.

28. Cyatheaceae

Hard minimum age constraint: 125 Ma. Permineralized fertiled structures have diagnostic characters of

modern Cyathea, including abaxially borne sori with globose indusia, annulate sporangia with

multiseriate stalks and vertical annuli that are not interrupted by the stalk, and 64 trilete spores58. The

Barremian Cyathea cranhamii represents the oldest unequivocal member of modern Cyatheaceae58 and

is used here as the minimum age constraint, although the permineralized stem genus Cyathocaulis

appears to be slightly older (Hauterivian)59 and was resolved as a member of Cyatheaceae in a cladistic

analysis60. We assigned the constraint to the Cyatheaceae stem lineage.

Soft maximum age constraint: 251 Ma. We apply the beginning of Triassic as the soft maximum age

for the tree ferns.

11
29. Lindsaeaceae

Hard minimum age constraint: 99.6 Ma. Cretaceous fossil roots display sclerenchymatous outer cortex,

parenchymatous inner cortex, and an innermost layer of six large cells61. This root anatomy is only

known in the family Lindsaeaceae61. We assigned the constraint to the Lindsaeaceae stem lineage.

Soft maximum age constraint: 201.6 Ma. Some Jurassic fossils have been associated with

Lindsaeaceae, although their affinity is highly uncertain61,62. Oldest fossils possibly associated with

Polypodiales are also Jurassic17,56,62 and we used the beginning of the Jurassic as the soft maximum age

for the stem lineage.

30. Pteridaceae

Hard minimum age constraint: 93.5 Ma. The shape of the pinnae and the presence of pseudoindusia in

a Cenomanian fossil63 closely resemble pteridoid ferns, and this fossil has been used to constrain the

origin of pteridoid lineage16,17,31. We assigned the constraint to the Pteridaceae stem lineage.

Soft maximum age constraint: 201.6 Ma. Oldest known putative members of Polypodiales are

Jurassic17,56,62 and therefore, we used the beginning of Jurassic as the soft maximum age constrain for

the stem lineage.

31. Acrostichum-Ceratopteris –clade

Hard minimum age constraint: 65.5 Ma. Macrofossils from Maastrichtian were assigned to

Acrostichum64, but more conservative views consider this fossil as putative stem group member17,31.

This more conservative approach is also followed here.

Soft maximum age constraint: 201.6 Ma. The soft maximum age of Pteridaceae was applied to the stem

lineage.

12
32. Ceratopteris

Hard minimum age constraint: 37.2 Ma. Diagnostic Ceratopteris spores are known from the

Bartonian47. This calibration has also been used in previous studies17,31. We assigned the constraint to

the Ceratopteris stem lineage.

Soft maximum age constraint: 145.5 Ma. If the Late Cretaceous Acrostichum64 indeed represents crown

group Acrostichum instead of stem group, the divergence between Acrostichum and Ceratopteris must

have occured earlier during the Cretaceous if not before. We apply the beginning of Cretaceous as the

soft maximum age for the stem lineage.

33. Dennstaedtia-Microlepia –clade

Hard minimum age constraint: 70.6 Ma. The rhizome and stipe anatomy of the genus Microlepiopsis

from Campanian are characteristic for the clade containing living species of Microlepia and

Dennstaedtia65. This calibration was also applied in a previous study31. We assigned the constraint to

the stem lineage of this clade.

Soft maximum age constraint: 201.6 Ma. It remains uncertain whether Dennstaedtiaceae or Pteridaceae

diverged earlier. For this early divergence within Dennstaedtiaceae we apply the same soft maximum

as for the family Pteridaceae.

34. Pteridium

Hard minimum age constraint: 23 Ma. Fossil evidence indicates worldwide distribution of Pteridium by

Oligocene66. This constraint was assigned to the Pteridium stem lineage.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the stem lineage in lack of good fossil record, but it has been estimated that Pteridium likely

originated during Cretaceous-Paleogene67.

13
35. Drynaria

Hard minimum age constraint: 33.9 Ma. Phylogenetic analysis resolved Protodrynaria takhtajanii from

the Eocene-Oligocene boundary as a member of Drynaria68. The constraint was assigned to the

Drynaria stem lineage.

Soft maximum age constraint: 145.5 Ma. Lower Cretaceous deposits of North America and China have

yielded fossils with eupolypod characteristics69–71. Molecular studies have also suggested Cretaceous

origin for the main eupolypod groups16,17,31. We apply the beginning of Cretaceous as the soft

maximum age for the eupolypod groups.

36. Drynaria mollis

Hard minimum age constraint: 2.6 Ma. The morphology-based and total-evidence analyses resolved

Pliocene D. callispora as sister to the extant species D. mollis, and these two formed sister lineage to D.

sinica68. Although the morphological results somewhat differed from the molecular results (e.g.

Drynaria monophyletic vs. paraphyletic), the nodes critical to place D. callispora in the phylogeny

remained the same and we constrained the D. mollis–D. sinica split to have occured before 2.6 Ma.

Soft maximum age constraint: 65.5 Ma. We applied the beginning of Paleogene as the soft maximum

age for this phylogenetically recent split.

37. Cyclosoroids

Hard minimum age constraint: 33.9 Ma. Eocene fossils can be assigned as members of cyclosoroids72,

which form a clade within Thelypteridaceae73. This calibration has also been used in previous

studies17,31. We assigned the constraint to the cyclosoroid stem lineage.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the eupolypod groups (see above).

14
38. Onoclea sensibilis

Hard minimum age constraint: 55.8 Ma. A large number of relatively complete Paleocene fossils

conform to the living species in all recognizable features74. This calibration has been accepted in

previous studies17,31, although in these more sparsely sampled analyses the calibration was assigned at

deeper phylogenetic level. We assigned the constraint to stem lineage of the clade of two Onoclea

sensibilis varieties.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the eupolypod groups (see above).

39. Stenochlaena

Hard minimum age constraint: 23 Ma. Distinctive spores of Stenochlaenidites are essentially identical

with the spores of modern Stenochlaena and first appear in the late Oligocene72. The constraint was

assigned to the Stenochlaena stem lineage.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the eupolypod groups (see above).

40. Woodwardia

Hard minimum age constraint: 55.8 Ma. Several fossil Woodwardia species are known from the

Paleocene onwards72, including W. bureiensis from Wuyun Formation of China75. This calibration has

been used previously as well31. The constraint was assigned to the Woodwardia stem lineage.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the eupolypod groups (see above).

41. Woodwardia virginica

Hard minimum age constraint: 15.4 Ma. Combination of vegetative pinnules, rhizome and stipe

anatomy, and fertile pinnules with sori and sporangia from the Middle Miocene were described to be

15
similar with the extant Woodwardia virginica76. The fossil bed was dated 15.6 ± 0.2 Ma by Ar/Ar

dating technique76,77. We used the lower estimate as the hard minimum age to constrain the crown

group Woodwardia.

Soft maximum age constraint: 145.5 Ma. We apply the soft maximum age of the genus as the soft

maximum age for this early diverging species.

42. Athyriaceae

Hard minimum age constraint: 37.2 Ma. The Middle Eocene Makopteris princetonensis is the oldest

fossil assigned to the family78 and has been used to calibrate Athyriaceae in other studies as well17,31.

The constraint was assigned to the Athyriaceae stem lineage.

Soft maximum age constraint: 145.5 Ma. We apply the beginning of Cretaceous as the soft maximum

age for the eupolypod groups (see above).

Coding of fern life-form

Taxa non-epiphytic (0) / epiphytic (1)

Abrodictyum caudatum 1

Abrodictyum elongatum 0

Abrodictyum rigidum 0

Acrostichum aureum 0

Acrostichum danaeifolium 0

Actiniopteris dimorpha 0

Actinostachys pennula 0

Acystopteris japonica 0

Acystopteris taiwaniana 0

16
Adenophorus abietinus 1

Adenophorus hymenophylloides 1

Adenophorus montanus 1

Adenophorus oahuensis 1

Adenophorus periens 1

Adenophorus pinnatifidus 1

Adenophorus tamariscinus 1

Adenophorus tenellus 1

Adenophorus tripinnatifidus 1

Adiantopsis radiata 0

Adiantum adiantoides 0

Adiantum aethiopicum 0

Adiantum argutum 0

Adiantum capillus-veneris 0

Adiantum diaphanum 0

Adiantum formosum 0

Adiantum hispidulum 0

Adiantum malesianum 0

Adiantum pedatum 0

Adiantum peruvianum 0

Adiantum raddianum 0

Adiantum tenerum 0

Adiantum tetraphyllum 0

17
Adiantum viridescens 0

Alansmia cultrata 1

Alansmia glandulifera 1

Alansmia lanigera 1

Alansmia senilis 1

Allosorus argenteus 0

Allosorus farinosus 0

Allosorus nitidulus 0

Alsophila bryophila 0

Alsophila cuspidata 0

Alsophila dregei 0

Alsophila foersteri 0

Alsophila hooglandii 0

Alsophila stelligera 0

Anemia adiantifolia 0

Anemia mexicana 0

Anemia phyllitidis 0

Anemia rotundifolia 0

Anemia tomentosa 0

Angiopteris evecta 0

Angiopteris lygodiifolia 0

Anogramma guatemalensis 0

Anogramma leptophylla 0

18
Antrophym latifolium 1

Arachniodes amoena 0

Arachniodes aristata 0

Arachniodes blinii 0

Arachniodes caudata 0

Arachniodes cavaleri 0

Arachniodes centrochinensis 0

Arachniodes chinensis 0

Arachniodes denticulata 0

Arachniodes festina 0

Arachniodes miqueliana 0

Arachniodes quadripinnata 0

Arachniodes rhomboides 0

Arachniodes simplicior 0

Arachniodes speciosa 0

Arachniodes standishii 0

Arachniodes superba 0

Argyrochosma fendleri 0

Argyrochosma incana 0

Argyrochosma jonesii 0

Argyrochosma limitanea 0

Arthromeris lehmannii 1

Arthromeris wallichiana 1

19
Arthropteris palisotii 0

Arthropteris parallela 0

Arthropteris paucivenia 1

Ascogrammitis dilatata 1

Ascogrammitis anfractuosa 1

Ascogrammitis david-smithii 1

Ascogrammitis nana 1

Ascogrammitis pichinchae 1

Ascogrammitis pichinchensis 1

Ascogrammitis sp.1 1

Ascogrammitis sp.2 1

Aspidotis californica 0

Aspidotis densa 0

Asplenium abscissum 0

Asplenium adiantum-nigrum 0

Asplenium affine 1

Asplenium alatum 0

Asplenium angustum 1

Asplenium anisophyllum 1

Asplenium aureum 0

Asplenium auritum 1

Asplenium australasicum 1

Asplenium bulbiferum 1

20
Asplenium bullatum 0

Asplenium contiguum 1

Asplenium cordatum 0

Asplenium cristatum 1

Asplenium cuneatiforme 1

Asplenium cuneifolium 0

Asplenium cuspidatum 1

Asplenium dielmannii 0

Asplenium difforme 0

Asplenium elliottii 1

Asplenium erectum 0

Asplenium feei 1

Asplenium fissum 0

Asplenium flabellifolium 1

Asplenium fontanum 0

Asplenium forisiense 0

Asplenium formosae 0

Asplenium formosum 0

Asplenium fragile 0

Asplenium gemmiferum 0

Asplenium hallbergii 0

Asplenium harpeodes 1

Asplenium hemionitis 0

21
Asplenium heterochroum 0

Asplenium hispanicum 0

Asplenium hobdyi 0

Asplenium hookerianum 0

Asplenium incisum 0

Asplenium jahandiezii 0

Asplenium juglandifolium 1

Asplenium laciniatum 0

Asplenium lamprophyllum 0

Asplenium lucidum 0

Asplenium lunulatum 0

Asplenium marinum 0

Asplenium monanthes 1

Asplenium myriophyllum 0

Asplenium nidus 1

Asplenium normale 1

Asplenium obtusatum 0

Asplenium planicaule 0

Asplenium platyneuron 0

Asplenium polyodon 1

Asplenium praemorsum 1

Asplenium prolongatum 0

Asplenium protensum 0

22
Asplenium pteropus 1

Asplenium resiliens 0

Asplenium rhizophyllum 0

Asplenium rigidum 0

Asplenium ritoense 0

Asplenium ruta-muraria 0

Asplenium sandersonii 1

Asplenium scolopendrium 0

Asplenium shuttleworthianum 0

Asplenium simplicifrons 1

Asplenium surrogatum 1

Asplenium tenerum 1

Asplenium theciferum 1

Asplenium tricholepis 1

Asplenium trichomanes 0

Asplenium varians 0

Asplenium vieillardii 0

Asplenium viride 0

Asplenium wrightii 0

Asplenium yunnanense 0

Athyrium decurrentialatum 0

Athyrium filix-femina 0

Athyrium nipponicum 0

23
Athyrium otophorum 0

Athyrium yokoscense 0

Azolla filiculoides 0

Azolla nilotica 0

Azolla pinnata 0

Azolla rubra 0

Blechnum brasiliense 0

Blechnum cyatheoides 0

Blechnum insigne 0

Blechnum medium 0

Blechnum neglectum 0

Blechnum nom.nov. 0

Blechnum occidentale 0

Blechnum orientale 0

Blechnum polypodioides 0

Blechnum schomburgkii 0

Blechnum spicant 0

Blotiella pubescens 0

Bolbitis appendiculata 0

Bolbitis auriculata 0

Bolbitis heteroclita 0

Bommeria ehrenbergiana 0

Bommeria hispida 0

24
Botrychium dissectum 0

Botrychium japonicum 0

Botrychium lanceolatum 0

Botrychium lunaria 0

Botrychium lunarioides 0

Botrychium simplex 0

Botrychium strictum 0

Botrychium virginianum 0

Calciphilopteris ludens 0

Callistopteris apiifolia 0

Calochlaena straminea 0

Calochlaena villosa 0

Calymmodon gracilis 1

Calymmodon luerssenianus 1

Campyloneurum angustifolium 1

Campyloneurum aphanophlebium 1

Campyloneurum asplundii 1

Campyloneurum brevifolium 1

Campyloneurum chlorolepis 1

Campyloneurum latum 1

Campyloneurum xalapense 1

Cephalomanes javanicum 0

Ceradenia fucoides 1

25
Ceradenia jungermannioides 1

Ceradenia kalbreyeri 1

Ceradenia pilipes 1

Ceradenia spixiana 1

Ceratopteris richardii 0

Ceratopteris thalictroides 0

Cheilanthes distans 0

Cheilanthes lasiophylla 0

Cheilanthes sieberi 0

Cheiloplecton rigidum 0

Cheiropleuria integrifolia 0

Christensenia aesculifola 0

Chrysogrammitis musgraviana 1

Cibotium barometz 0

Cibotium glaucum 0

Cibotium schiedei 0

Cochlidium punctatum 1

Cochlidium rostratum 1

Cochlidium seminudum 1

Cochlidium serrulatum 1

Coniogramme fraxinea 0

Coniogramme japonica 0

Crepidomanes bipunctatum 1

26
Crepidomanes minutum 1

Crepidomanes thysanostomum 0

Cryptogramma crispa 0

Ctenitis decurrentipinnata 0

Ctenitis eatonii 0

Ctenitis refulgens 0

Ctenitis rhodolepis 0

Ctenitis sloanei 0

Ctenitis submarginalis 0

Ctenopteris lasiostipes 1

Ctenopteris nutans 1

Ctenopteris repandula 1

Culcita coniifolia 0

Culcita macrocarpa 0

Cyathea alata 0

Cyathea arborea 0

Cyathea dejecta 0

Cyathea horrida 0

Cyathea howeana 0

Cyathea multiflora 0

Cyathea parvula 0

Cyathea poeppigii 0

Cyathea senilis 0

27
Cyclopeltis crenata 0

Cyclopeltis semicordata 0

Cyrtomium falcatum 0

Cyrtomium fortunei 0

Cyrtomium macrophyllum 0

Cyrtomium nephrolepioides 0

Cyrtomium urophyllum 0

Cystodium sorbifolium 0

Cystopteris fragilis 0

Danaea alata 0

Danaea falcata 0

Danaea nodosa 0

Danaea simplicifolia 0

Danaea sp. 0

Danaea urbanii 0

Danaea wendlandii 0

Davallia burbidgei 1

Davallia clarkei 1

Davallia corniculata 1

Davallia divaricata 1

Davallia epiphylla 1

Davallia fejeensis 1

Davallia griffithiana 1

28
Davallia hirsuta 1

Davallia hymenophylloides 1

Davallia mariesii 1

Davallia melanophlebia 1

Davallia parvula 1

Davallia perdurans 1

Davallia plumosa 1

Davallia polypodioides 1

Davallia pulchra 1

Davallia pyxidata 1

Davallia repens 1

Davallia solida 1

Davallia tasmanii 0

Davallia trichomanoides 1

Davallia triphylla 1

Davallia yunnanensis 1

Dennstaedtia punctilobula 0

Dennstaedtia scabra 0

Deparia bonincola 0

Deparia lancea 0

Deparia petersenii 0

Deparia unifurcata 0

Dicksonia antarctica 0

29
Dicksonia thyrsopteroides 0

Dicranopteris linearis 0

Dictymia brownii 1

Dictymia mckeei 1

Didymochlaena truncatula 0

Didymoglossum cuspidatum 1

Didymoglossum ekmanii 1

Didymoglossum hildebrandtii 1

Didymoglossum krausii 1

Didymoglossum membranaceum 1

Diplaziopsis javanica 0

Diplazium bombonasae 0

Diplazium centripetale 0

Diplazium cristatum 0

Diplazium dilatatum 0

Diplazium hachijoense 0

Diplazium legalloi 0

Diplazium okudairai 0

Diplazium plantaginifolium 0

Diplazium proliferum 0

Diplazium virescens 0

Diplazium wichurae 0

Diplopterygium bancroftii 0

30
Diplopterygium glaucum 0

Diplopterygium laevissimum 0

Dipteris conjugata 0

Doryopteris gleichenioides 0

Doryopteris nobilis 0

Doryopteris rediviva 0

Doryopteris sagittifolia 0

Doryopteris viridis 0

Dracoglossum plantagineum 0

Drynaria acuminata 1

Drynaria bonii 1

Drynaria coronans 1

Drynaria drynarioides 1

Drynaria laurentii 1

Drynaria meyeniana 1

Drynaria mollis 1

Drynaria nom.nov. 1

Drynaria novoguineensis 1

Drynaria pilosa 1

Drynaria rigidula 1

Drynaria sagitta 1

Drynaria sinica 1

Drynaria sp. 1

31
Drynaria sparsisora 1

Drynaria splendens 1

Drynaria volkensii 1

Dryopteris aemula 0

Dryopteris annamensis 0

Dryopteris bissetiana 0

Dryopteris crassirhizoma 0

Dryopteris diffracta 0

Dryopteris erythrosora 0

Dryopteris expansa 0

Dryopteris filix-mas 0

Dryopteris goldiana 0

Dryopteris hasseltii 0

Dryopteris hendersonii 0

Dryopteris heterolaena 0

Dryopteris hookeriana 0

Dryopteris labordei 0

Dryopteris marginalis 0

Dryopteris mariformis 0

Dryopteris maximowiczii 0

Dryopteris neolacera 0

Dryopteris nodosa 0

Dryopteris pacifica 0

32
Dryopteris sacrosancta 0

Dryopteris sieboldii 0

Dryopteris sparsa 0

Dryopteris squamiseta 0

Dryopteris uniformis 0

Dryopteris wallichiana 0

Elaphoglossum amygdalifolium 0

Elaphoglossum andicola 1

Elaphoglossum aubertii 0

Elaphoglossum burchellii 0

Elaphoglossum callifolium 0

Elaphoglossum crassifolium 1

Elaphoglossum crinitum 1

Elaphoglossum deltoides 0

Elaphoglossum erinaceum 1

Elaphoglossum flaccidum 1

Elaphoglossum glabellum 1

Elaphoglossum heterolepis 1

Elaphoglossum hybridum 0

Elaphoglossum lechlerianum 1

Elaphoglossum lingua 1

Elaphoglossum lonchophyllum 1

Elaphoglossum minutum 0

33
Elaphoglossum paleaceum 1

Elaphoglossum papillosum 1

Elaphoglossum peltatum 1

Elaphoglossum piloselloides 1

Elaphoglossum samoense 1

Elaphoglossum tripartitum 0

Enterosora parietina 1

Enterosora percrassa 1

Enterosora trifurcata 1

Equisetum arvense 0

Equisetum bogotense 0

Equisetum diffusum 0

Equisetum ferrissii 0

Equisetum giganteum 0

Equisetum myriochaetum 0

Equisetum pratense 0

Equisetum ramosissimum 0

Equisetum scirpoides 0

Equisetum sylvaticum 0

Equisetum telmateia 0

Equisetum variegatum 0

Eupodium laeve 0

Gaga arizonica 0

34
Galactodenia subscabra 1

Gleichenella pectinata 0

Gleichenia dicarpa 0

Goniophlebium amoenum 1

Goniophlebium argutum 1

Goniophlebium chinense 1

Goniophlebium formosanum 1

Goniophlebium manmeiense 1

Goniophlebium mehibitense 1

Goniophlebium microrhizoma 1

Goniophlebium percussum 1

Goniophlebium persicifolium 1

Goniophlebium pseudoconnatum 1

Goniophlebium subauriculatum 1

Grammitis bryophila 1

Grammitis deplanchei 0

Grammitis kyimbilensis 1

Grammitis melanoloma 1

Gymnocarpium dryopteris 0

Gymnocarpium oyamense 0

Gymnocarpium remotepinnatum 0

Gymnosphaera capensis 0

Gymnosphaera klossii 0

35
Gymnosphaera podophylla 0

Gymnosphaera ramispina 0

Gymnosphaera salvinii 0

Haplopteris elongata 1

Haplopteris graminea 1

Haplopteris taeniophylla 1

Haplopteris volkensii 1

Hecistopteris pumila 1

Helminthostachys zeylanica 0

Hemidictyum marginatum 0

Hemionitis arifolia 0

Hemionitis palmata 0

Histiopteris incisa 0

Homalosorus pycnocarpos 0

Hymenasplenium cheilosorum 0

Hymenasplenium excisum 1

Hymenasplenium laetum 0

Hymenasplenium unilaterale 1

Hymenophyllum acanthoides 1

Hymenophyllum apiculatum 1

Hymenophyllum armstrongii 1

Hymenophyllum atrovirens 0

Hymenophyllum badium 1

36
Hymenophyllum baileyanum 1

Hymenophyllum barbatum 1

Hymenophyllum bivalve 1

Hymenophyllum caespitosum 1

Hymenophyllum caudiculatum 1

Hymenophyllum cruentum 1

Hymenophyllum cupressiforme 1

Hymenophyllum demissum 1

Hymenophyllum dentatum 1

Hymenophyllum deplanchei 1

Hymenophyllum dicranotrichum 1

Hymenophyllum digitatum 1

Hymenophyllum dilatatum 1

Hymenophyllum exsertum 1

Hymenophyllum flexuosum 1

Hymenophyllum frankliniae 1

Hymenophyllum fuciforme 1

Hymenophyllum fucoides 1

Hymenophyllum heimii 1

Hymenophyllum hirsutum 1

Hymenophyllum hygrometricum 1

Hymenophyllum imbricatum 1

Hymenophyllum inaequale 1

37
Hymenophyllum javanicum 1

Hymenophyllum kuhnii 1

Hymenophyllum lanceolatum 1

Hymenophyllum leratii 0

Hymenophyllum lyallii 1

Hymenophyllum malingii 1

Hymenophyllum marginatum 1

Hymenophyllum minimum 1

Hymenophyllum mnioides 1

Hymenophyllum multifidum 0

Hymenophyllum pallidum 0

Hymenophyllum paniense 1

Hymenophyllum pectinatum 1

Hymenophyllum peltatum 1

Hymenophyllum perrieri 1

Hymenophyllum plicatum 1

Hymenophyllum polyanthos 1

Hymenophyllum poolii 1

Hymenophyllum pulcherrimum 1

Hymenophyllum rolandi-principis 1

Hymenophyllum rufescens 1

Hymenophyllum rugosum 1

Hymenophyllum scabrum 1

38
Hymenophyllum secundum 1

Hymenophyllum serrulatum 1

Hymenophyllum sibthorpioides 1

Hymenophyllum sp. 1

Hymenophyllum subdimidiatum 1

Hymenophyllum tenellum 1

Hymenophyllum tunbrigense 1

Hymenophyllum undulatum 0

Hymenophyllum villosum 1

Hymenophyllum wrightii 1

Hypodematium crenatum 0

Hypolepis tenuifolia 0

Jamesonia canescens 0

Jamesonia cheilanthoides 0

Jamesonia flexuosa 0

Jamesonia verticalis 0

Lastreopsis hispida 0

Lastreopsis subrecedens 0

Lellingeria apiculata 1

Lellingeria brevistipes 1

Lellingeria depressa 1

Lellingeria dissimulans 1

Lellingeria flagellipinnata 1

39
Lellingeria hirsuta 1

Lellingeria isidrensis 1

Lellingeria itatimensis 0

Lellingeria kaieteura 1

Lellingeria melanotrichia 1

Lellingeria phlegmaria 1

Lellingeria pseudocapillaris 1

Lellingeria randallii 1

Lellingeria simacensis 1

Lellingeria sp.1 1

Lellingeria sp.2 1

Lellingeria subsessilis 1

Lellingeria suspensa 1

Lellingeria tenuicula 1

Lemmaphyllum carnosum 1

Lemmaphyllum microphyllum 1

Lemmaphyllum rostratum 1

Lemmaphyllum squamatum 0

Lepidomicrosorium buergerianum 0

Lepidomicrosorium subhemionitideum 1

Lepidomicrosorium superficiale 1

Lepisorus accedens 1

Lepisorus annamensis 1

40
Lepisorus annuifrons 1

Lepisorus boninensis 1

Lepisorus excavatus 1

Lepisorus henryi 1

Lepisorus kawakamii 1

Lepisorus lewisii 0

Lepisorus marginatus 1

Lepisorus miyoshianus 1

Lepisorus monilisorus 1

Lepisorus mucronatus 1

Lepisorus nudus 1

Lepisorus platyrhynchos 1

Lepisorus pseudonudus 1

Lepisorus pseudoussuriensis 1

Lepisorus scolopendrius 1

Lepisorus sinensis 1

Lepisorus spicatus 1

Lepisorus thaipaiensis 1

Lepisorus thunbergianus 1

Lepisorus uchiyamae 0

Leptochilus cantoniensis 0

Leptochilus decurrens 1

Leptochilus digitatus 0

41
Leptochilus ellipticus 0

Leptochilus hemionitideus 0

Leptochilus henryi 0

Leptochilus simplicifrons 0

Leptochilus triphyllus 0

Leptochilus wrightii 1

Leptochilus xhemitomus 0

Leptolepia nom.nov. 0

Leptolepia novae-zelandiae 0

Leptopteris superba 0

Leptopteris wilkesiana 0

Leucostegia immersa 1

Leucostegia pallida 1

Leucotrichium mitchelliae 1

Leucotrichium mortonii 1

Leucotrichium organense 1

Leucotrichium pseudomitchelliae 1

Leucotrichium schenckii 1

Lindsaea blotiana 0

Lindsaea botrychioides 0

Lindsaea ensifolia 0

Lindsaea imrayana 0

Lindsaea multisora 0

42
Lindsaea parasitica 1

Lindsaea plicata 0

Lindsaea quadrangularis 0

Lindsaea rufa 0

Lindsaea stricta 0

Llavea cordifolia 0

Lomagramma matthewii 0

Lomariopsis japurensis 0

Lomariopsis pollicina 0

Lomariopsis sorbifolia 0

Lomariopsis sp. 0

Lomariopsis spectabilis 0

Lonchitis hirsuta 0

Lophosoria quadripinnata 0

Loxogramme abyssinica 1

Loxogramme avenia 1

Loxogramme dictyopteris 1

Loxogramme grammitoides 1

Loxogramme salicifolia 1

Loxsoma cunninghamii 0

Loxsomopsis pearcei 0

Lygodium japonicum 0

Lygodium lanceolatum 0

43
Lygodium radiatum 0

Lygodium reticulatum 0

Macrothelypteris torresiana 0

Marattia alata 0

Marsilea crenata 0

Marsilea crotophora 0

Marsilea drummondii 0

Marsilea gibba 0

Marsilea mutica 0

Marsilea nubica 0

Marsilea vera 0

Matonia pectinata 0

Matteuccia struthiopteris 0

Maxonia apiifolia 0

Megalastrum subincisum 0

Melpomene allosuroides 1

Melpomene gracilis 1

Melpomene melanosticta 1

Melpomene moniliformis 0

Metaxya rostrata 0

Mickelia guianensis 0

Mickelia nicotianifolia 0

Microgramma baldwinii 1

44
Microgramma bifrons 1

Microgramma fuscopunctata 1

Microgramma lycopodioides 1

Microgramma mauritiana 1

Microgramma megalophylla 1

Microgramma nitida 1

Microgramma percussa 1

Microgramma sp. 1

Microgramma squamulosa 1

Microgramma tecta 1

Microgramma vacciniifolia 1

Microlepia platyphylla 0

Microlepia speluncae 0

Microlepia szechuanica 0

Microsorum carnosum 1

Microsorum commutatum 0

Microsorum crustaceum 1

Microsorum cuspidatum 1

Microsorum grossum 0

Microsorum hainanense 1

Microsorum insigne 0

Microsorum lastii 1

Microsorum linguiforme 1

45
Microsorum luzonense 1

Microsorum membranaceum 1

Microsorum mirabile 1

Microsorum musifolium 1

Microsorum novae-zealandiae 1

Microsorum papuanum 0

Microsorum pteropus 0

Microsorum punctatum 1

Microsorum pustulatum 1

Microsorum sarcopus 1

Microsorum scandens 1

Microsorum sinuosum 1

Microsorum spectrum 1

Microsorum thailandicum 0

Microsorum varians 1

Microsorum vieillardii 1

Microsorum whiteheadii 0

Mildella intramarginalis 0

Monachosorum henryi 0

Moranopteris achilleifolia 1

Moranopteris hyalina 1

Moranopteris longisetosa 1

Moranopteris taenifolia 1

46
Moranopteris zurquina 1

Mycopteris alsopteris 1

Mycopteris semihirsuta 1

Mycopteris sp.1 1

Mycopteris sp.2 1

Mycopteris subtilis 1

Mycopteris taxifolia 1

Myriopteris alabamensis 0

Myriopteris aurea 0

Myriopteris covillei 0

Myriopteris lanosa 0

Myriopteris lendigera 0

Myriopteris myriophylla 0

Myriopteris newberryi 0

Myriopteris pringlei 0

Myriopteris rufa 0

Myriopteris wrightii 0

Neocheiropteris palmatopedata 1

Neolepisorus ensatus 1

Neolepisorus fortunei 1

Neolepisorus phyllomanes 0

Neolepisorus zippelii 1

Nephrolepis abrupta 0

47
Nephrolepis cordifolia 0

Nephrolepis davalliae 1

Nephrolepis davallioides 1

Nephrolepis exaltata 1

Nephrolepis falcata 1

Nephrolepis hirsutula 0

Nephrolepis lauterbachii 1

Nephrolepis rivularis 1

Nephrolepis undulata 0

Nesolindsaea kirkii 0

Niphidium crassifolium 1

Notholaena aschenborniana 0

Notholaena aurantiaca 0

Notholaena aurea 0

Notholaena brachypus 0

Notholaena bryopoda 0

Notholaena californica 0

Notholaena candida 0

Notholaena grayi 0

Notholaena greggii 0

Notholaena lemmonii 0

Notholaena leucopoda 0

Notholaena neglecta 0

48
Notholaena rigida 0

Notholaena rosei 0

Notholaena schaffneri 0

Notholaena standleyi 0

Notholaena sulphurea 0

Notholaena trichomanoides 0

Notogrammitis billardierei 1

Notogrammitis ciliata 0

Notogrammitis crassior 0

Notogrammitis heterophylla 1

Notogrammitis pseudociliata 1

Odontosoria aculeata 0

Odontosoria biflora 0

Odontosoria chinensis 0

Oleandra articulata 1

Oleandra pistillaris 1

Onoclea sensibilis 0

Onoclea sensibilis var. interrupta 0

Onocleopsis hintonii 0

Onychium japonicum 0

Ophioglossum pendulum 1

Ophioglossum reticulatum 0

Oreogrammitis congener 1

49
Oreogrammitis dolichosora 0

Oreogrammitis forbesiana 1

Oreogrammitis hookeri 1

Oreogrammitis knutsfordiana 1

Oreogrammitis padangensis 1

Oreogrammitis reinwardtioides 1

Osmolindsaea japonica 0

Osmolindsaea odorata 0

Osmunda banksiifolia 0

Osmunda claytoniana 0

Osmunda javanica 0

Osmunda regalis 0

Osmundastrum cinnamomeum 0

Osmundastrum cinnamomeum var. fokiense 0

Paesia scaberula 0

Paragramma longifolia 1

Parapolystichum glabellum 0

Parapolystichum glabellumeffusum 0

Pecluma alfredii 1

Pecluma chnoophora 1

Pecluma dulcis 1

Pecluma eurybasis 1

Pecluma hartwegiana 1

50
Pecluma longepinnulata 1

Pecluma longepinnulata 1

Pecluma ptilodon 1

Pecluma rhachipterygia 0

Pellaea andromedifolia 0

Pellaea atropurpurea 0

Pellaea bipinnata 0

Pellaea breweri 0

Pellaea cochisensis 0

Pellaea intermedia 0

Pellaea marantae 0

Pellaea rotundifolia 0

Pellaea sargentii 0

Pellaea sinuata 0

Pellaea truncata 0

Pellaea vestita 0

Pentagramma triangularis 0

Phanerophlebia nobilis 0

Phanerosorus sarmentosus 0

Phegopteris connectilis 0

Phegopteris decursivepinnata 0

Phegopteris hexagonoptera 0

Phlebodium areolatum 1

51
Phlebodium decumanum 1

Phlebodium pseudoaureum 1

Pilularia globulifera 0

Pilularia novae-hollandiae 0

Pityrogramma austroamericana 0

Pityrogramma calomelanos 0

Pityrogramma chaerophylla 0

Plagiogyria euphlebia 0

Plagiogyria japonica 0

Plagiogyria matsumureana 0

Plagiogyria semicordata 0

Platycerium alcicorne 1

Platycerium andinum 1

Platycerium bifurcatum 1

Platycerium coronarium 1

Platycerium elephantotis 1

Platycerium ellisii 1

Platycerium grande 1

Platycerium hillii 1

Platycerium holttumii 1

Platycerium madagascariense 1

Platycerium quadridichotomum 1

Platycerium ridleyi 1

52
Platycerium stemaria 1

Platycerium veitchii 0

Platycerium wallichii 1

Platycerium wandae 1

Platycerium willinckii 1

Pleocnemia rufinervis 0

Pleopeltis angusta 1

Pleopeltis astrolepis 1

Pleopeltis bradeorum 1

Pleopeltis buchtienii 1

Pleopeltis christensenii 1

Pleopeltis crassinervata 1

Pleopeltis desvauxii 1

Pleopeltis disjuncta 1

Pleopeltis ensiformis 1

Pleopeltis fraseri 1

Pleopeltis friedrichsthalianum 1

Pleopeltis fructuosa 1

Pleopeltis furfuraceum 1

Pleopeltis intermedia 1

Pleopeltis macrocarpa 1

Pleopeltis marginata 1

Pleopeltis monosora 1

53
Pleopeltis muenchii 1

Pleopeltis myriolepis 1

Pleopeltis platylepis 1

Pleopeltis pleopeltifolia 1

Pleopeltis polypodioides 1

Pleopeltis pycnocarpa 1

Pleopeltis rosei 1

Pleopeltis rzedowskianum 1

Pleopeltis sanctae-rosae 1

Pleopeltis thyssanolepis 0

Pleopeltis wiesbaurii 1

Pleurosoriopsis makinoi 1

Polybotrya alfredii 0

Polybotrya caudata 0

Polybotrya cervina 0

Polybotrya guianensis 0

Polybotrya sessilisora 0

Polybotrya trianae 0

Polyphlebium angustatum 1

Polyphlebium borbonicum 1

Polyphlebium endlicherianum 1

Polypodium adelphum 1

Polypodium arcanum 1

54
Polypodium chrysolepis 1

Polypodium glycyrrhiza 0

Polypodium hesperium 0

Polypodium macaronesicum 0

Polypodium martensii 1

Polypodium pellucidum 0

Polypodium plesiosorum 1

Polypodium rhodopleuron 1

Polypodium scouleri 0

Polypodium subpetiolatum 1

Polypodium vulgare 1

Polystichopsis chaerophylloides 0

Polystichum acutidens 0

Polystichum caducum 0

Polystichum capillipes 0

Polystichum conjunctum 0

Polystichum craspedosorum 0

Polystichum eximium 0

Polystichum fraxinellum 0

Polystichum glaciale 0

Polystichum hillebrandii 0

Polystichum hookerianum 0

Polystichum lemmonii 0

55
Polystichum luctuosum 0

Polystichum munitum 0

Polystichum neolobatum 0

Polystichum omeiense 0

Polystichum setiferum 0

Polystichum tonkinense 0

Polystichum transkeiense 0

Polystichum tripteron 0

Polystichum yunnanense 0

Polytaenium cajenense 1

Prosaptia alata 1

Prosaptia contigua 1

Prosaptia obliquata 1

Prosaptia rhodocarpa 1

Pseudophegopteris aurita 0

Pseudophegopteris cruciata 0

Psilotum nudum 1

Pteridium aquilinum 0

Pteridium esculentum 0

Pteridrys lofouensis 0

Pteris arborea 0

Pteris comans 0

Pteris cretica 0

56
Pteris ensiformis 0

Pteris fauriei 0

Pteris multifida 0

Pteris pallens 0

Pteris platyzomopsis 0

Pteris praestantissima 0

Pteris propinqua 0

Pteris quadriaurita 0

Pteris tremula 0

Pteris vittata 0

Pterozonium brevifrons 0

Ptisana attenuata 0

Ptisana fraxinea 0

Ptisana oreades 0

Ptisana purpurascens 0

Ptisana salicina 0

Pyrrosia angustata 1

Pyrrosia assimilis 1

Pyrrosia christii 1

Pyrrosia foveolata 1

Pyrrosia linearifolia 1

Pyrrosia lingua 1

Pyrrosia longifolia 1

57
Pyrrosia niphoboloides 1

Pyrrosia nuda 1

Pyrrosia piloselloides 1

Pyrrosia polydactyla 1

Pyrrosia rasamalae 1

Pyrrosia rupestris 1

Pyrrosia serpens 1

Pyrrosia subfurfuracea 1

Radiogrammitis havilandii 1

Radiogrammitis hirtelloides 1

Radiogrammitis holttumii 1

Radiogrammitis jagoriana 1

Radiogrammitis parva 1

Radiovittaria gardneriana 1

Regnellidium diphyllum 0

Rhachidosorus pulcher 0

Rheopteris cheesmaniae 1

Rumohra adiantiformis 1

Saccoloma brasiliense 0

Saccoloma elegans 0

Saccoloma inaequale 0

Salpichlaena hookeriana 0

Salpichlaena thalassica 0

58
Salpichlaena volubilis 0

Salvinia cucullata 0

Salvinia minima 0

Salvinia molesta 0

Salvinia natans 0

Salvinia oblongifolia 0

Schizaea dichotoma 0

Schizaea sp. 0

Scleroglossum sulcata 0

Selliguea brachypodia 1

Selliguea dareiformis 1

Selliguea enervis 1

Selliguea hellwigii 1

Selliguea heterocarpa 1

Selliguea laciniata 1

Selliguea lanceolata 1

Selliguea plantaginea 0

Selliguea triloba 1

Serpocaulon adnatum 1

Serpocaulon attenuatum 1

Serpocaulon catharinae 1

Serpocaulon dissimile 1

Serpocaulon falcaria 1

59
Serpocaulon fraxinifolium 1

Serpocaulon funckii 1

Serpocaulon lasiopus 1

Serpocaulon loriciforme 1

Serpocaulon meniscifolium 1

Serpocaulon ptilorhizon 1

Serpocaulon sessilifolium 1

Serpocaulon silvulae 1

Serpocaulon subandinum 1

Serpocaulon triseriale 1

Serpocaulon wagneri 1

Serpocaulon wiesbaueri 1

Sphaeropteris capitata 0

Sphaeropteris celebica 0

Sphaeropteris horrida 0

Sphaeropteris medullaris 0

Sphaeropteris robusta 0

Sphenomeris clavata 0

Stenochlaena milnei 0

Stenochlaena tenuifolia 0

Stenogrammitis hartii 1

Stenogrammitis hildebrandtii 1

Stenogrammitis limula 1

60
Stenogrammitis myosuroides 1

Stenogrammitis oosora 1

Stenogrammitis prionodes 1

Stenogrammitis pumila 1

Stenogrammitis saffordii 1

Stenogrammitis subcoriacea 1

Stenogrammitis wittigiana 1

Sticherus laevigatus 0

Sticherus palmatus 0

Stigmatopteris lechleri 0

Stigmatopteris longicaudata 0

Stromatopteris moniliformis 0

Synammia feuillei 1

Synammia intermedia 1

Tapeinidium luzonicum 0

Tapeinidium moorei 0

Tapeinidium pinnatum 0

Tectaria apiifolia 0

Tectaria decurrens 0

Tectaria devexa 0

Tectaria fimbriata 0

Tectaria fuscipes 0

Tectaria harlandii 0

61
Tectaria incisa 0

Tectaria nayarii 0

Tectaria prolifera 0

Tectaria subtriphylla 0

Tectaria trifoliata 0

Tectaria variolosa 0

Tectaria zeilanica 0

Telmatoblechnum serrulatum 0

Teratophyllum wilkesianum 0

Terpsichore eggersii 1

Terpsichore hanekeana 1

Terpsichore lehmanniana 1

Thelypteris abrupta 0

Thelypteris affinis 0

Thelypteris ciliata 0

Thelypteris clypeolutata 0

Thelypteris consanguinea 0

Thelypteris dayi 0

Thelypteris dentata 0

Thelypteris ecallosa 0

Thelypteris erubescens 0

Thelypteris esquirolii 0

Thelypteris gemmulifera 0

62
Thelypteris glandulosa 0

Thelypteris griffithii 0

Thelypteris interrupta 0

Thelypteris limbosperma 0

Thelypteris longissima 0

Thelypteris noveboracensis 0

Thelypteris oligocarpa 0

Thelypteris opulenta 0

Thelypteris ovata 0

Thelypteris palustris 0

Thelypteris poiteana 0

Thelypteris pozoi 0

Thelypteris reticulata 0

Thelypteris seemannii 0

Thelypteris simplex 0

Thelypteris taiwanensis 0

Thelypteris uraiensis 0

Thelypteris xylodes 0

Themelium conjunctisorum 1

Thylacopteris papillosa 1

Thyrsopteris elegans 0

Tmesipteris tannensis 1

Todea barbara 0

63
Tricholepidium maculosum 1

Trichomanes alatum 0

Trichomanes ankersii 0

Trichomanes arbuscula 0

Trichomanes crispum 0

Trichomanes elegans 0

Trichomanes hostmannianum 0

Trichomanes lucens 1

Trichomanes osmundoides 0

Trichomanes pinnatum 0

Trichomanes scandens 0

Triplophyllum funestum 0

Tryonia myriophylla 0

Vandenboschia davallioides 0

Vandenboschia radicans 0

Vittaria graminifolia 1

Vittaria lineata 1

Woodsia ilvensis 0

Woodsia manchuriensis 0

Woodsia obtusa 0

Woodsia polystichoides 0

Woodwardia areolata 0

Woodwardia fimbriata 0

64
Woodwardia japonica 0

Woodwardia kempii 0

Woodwardia prolifera 0

Woodwardia unigemmata 0

Woodwardia virginica 0

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Supplementary Figure 1. Paleodistribution of the fossil data used in this study. Maps show

continental positions at a, 50 Ma, b, 180 Ma and c, 320 Ma5, and green dots show locations of fern

fossils with mean age estimate falling between a, 99.6 and 0 Ma, b, 251 and 99.6 Ma and c, 388 and

251 Ma. Maps are based on PALEOMAP Global Plate Tectonic Model5 and were created using Adobe

Photoshop CC 2015, PointTracker79 and QGIS v. 2.0.1 (http://www.qgis.org).

74
Supplementary Figure 2. Paleoenvironmental trajectories used in this study. Diversity of a, ferns,

b, lycophytes etc. (free-sporing vascular plants excluding ferns), c, gymnosperms80, d, angiosperms80.

Extent of the biomes: e, wet tropical, f, arid, g, warm temperate, h, cool temperate, i, polar. j, Global

75
mean temperature81. k, Atmospheric CO282. l, Atmospheric O283. m, Eustatic sealevel84. n, Magmatic

activity85,86. o, Continental fragmentation87. p, Mountain area. q, Selenium concentration in marine

sediments88,89.

Supplementary Figure 3. Dynamics of fern generic diversity through time. a, The range-through

diversity trajectory of sampled genera based on the estimated times of origination and extinction. b,

The amount of turnover calculated as the fraction of new genera originating in a 5 Ma time bin out of

the total number of genera recorded within the time bin. Global mass extinction events are indicated by

dotted red lines.

76
Supplementary Figure 4. Rates of climatic niche evolution in ferns. The three rate shift

configurations in the 95% credible set of shift configurations with their overall frequencies for the

evolution of climatic niche across the fern phylogeny as estimated with BAMM90,91. No rate changes

are supported by the most frequent configuration.

77
Supplementary Figure 5. Climatic niche components. a, Bio1 (annual mean temperature). b, Bio6

(min temperature of coldest month). c, Bio12 (annual precipitation). d, Bio15 (precipitation

78
seasonality). e, interannual precipitation variation. f, interannual temperature variation. g, The first

principal component of the four climatic layers. Maps were created using the statistical programming

language R92.

Supplementary Table 1. Origination and extinction times of

fern genera as estimated from the fossil record.
Origination (Ma) Extinction (Ma)
Taxon
Mean 95% CI Mean 95% CI

Pseudoborniales
Pseudobornia 380.00 362.36 398.57 363.64 354.79 381.90
Sphenophyllales
Bowmanites 355.29 337.24 370.08 270.04 255.15 279.11
Cheirostrobus 346.63 336.67 366.44 321.23 286.22 339.06
Eviostachya 369.36 364.78 374.94 360.55 358.99 363.61
Gondwanophyton 299.73 272.76 345.26 267.47 245.05 288.86
Hamatophyton 382.99 372.41 393.02 363.52 359.01 374.80
Lilpopia 316.00 299.22 350.83 269.95 245.56 296.91
Parasphenophyllum 317.15 304.02 358.17 292.42 252.87 305.93
Peltastrobus 319.78 300.04 359.79 294.69 252.76 317.22
Rotafolia 369.52 359.07 379.25 362.70 353.10 373.93
Sentistrobus 314.56 299.03 355.03 289.63 251.82 305.85
Sphenophyllostachys 346.86 315.60 370.82 291.06 257.32 303.08
Sphenophyllum 367.70 364.81 370.39 251.55 250.65 252.63
Xihuphyllum 374.49 359.15 388.91 368.15 358.98 384.61
Equisetales
Annularia 349.23 328.28 365.23 247.83 243.79 252.15
Archaeocalamites 359.91 352.90 366.27 281.60 274.52 289.31
Arthropitys 334.59 317.83 357.60 246.31 241.83 250.73
Asterophyllites 358.83 356.59 360.29 257.64 251.75 264.38
Austroannularia 288.81 272.65 319.47 250.69 240.64 259.59
Bengaliaraniganjensis 301.34 271.34 345.38 270.10 245.32 297.51
Benlightfootia 296.36 273.21 327.53 251.38 242.08 259.08
Calamites 359.00 357.16 360.46 173.09 171.04 175.31
Calamostachys 357.77 349.00 365.23 231.18 225.50 236.80
Cruciatheca 300.25 285.72 318.53 279.83 253.83 292.78
Equisetinostachys 286.16 274.79 308.03 252.17 243.88 258.73
Equisetites 325.86 299.92 354.71 59.55 58.70 60.84

79
Equisetocaulis 229.06 204.62 245.73 202.34 145.52 227.34
Equisetum 342.03 324.72 359.39 extant
Giridia 296.87 284.59 318.41 273.34 250.20 286.63
Gondwanites 320.88 299.09 359.93 296.00 254.36 317.70
Gondwanostachys 298.97 272.53 343.53 266.91 245.26 289.62
Lelstotheca 262.15 251.04 293.50 252.37 243.76 259.99
Neocalamites 297.50 295.68 299.21 130.56 102.80 153.92
Notocalamites 303.19 284.04 347.16 271.15 246.20 287.94
Paracalamites 300.91 295.24 309.70 252.37 245.24 259.39
Paracalamitina 287.57 260.21 336.05 253.87 239.68 268.68
Peltotheca 311.41 297.03 351.53 283.35 249.70 298.97
Phyllotheca 302.35 294.47 312.28 226.36 215.63 234.65
Raniganjia 351.98 337.13 370.49 247.32 235.66 255.82
Sakoarota 302.49 280.66 333.87 269.33 245.19 293.15
Schizoneura 295.13 291.03 299.6 153.73 150.67 157.05
Spaciinodum 245.29 241.07 253.36 239.52 229.87 244.91
Tchernovia 300.61 251.15 349.32 274.60 245.14 316.36
Umbellaphyllites 292.75 287.88 300.16 248.67 245.92 251.57
Ophioglossales
Botrychium 78.10 57.87 110.55 extant
Ophioglossum 24.34 16.41 36.39 extant
Psilotales
Psilotum 41.78 16.09 87.40 extant
Tmesipteris 37.84 28.57 51.13 extant
Marattiales
Acitheca 322.31 311.79 332.95 252.93 246.29 259.87
Angiopteris 231.95 208.94 247.17 extant
Asterotheca 340.20 310.45 361.07 207.11 201.62 214.98
Boreopteris 252.64 248.47 257.13 246.82 242.6 250.82
Caulopteris 407.87 391.63 423.28 245.74 239.32 251.05
Danaea 69.67 35.45 128.24 extant
Danaeites 355.11 322.02 375.08 79.51 44.59 105.27
Danaeopsis 260.31 254.33 265.51 86.58 84.30 88.55
Dizeugotheca 290.48 274.75 307.23 250.77 244.16 256.76
Drepanozamites 239.28 230.97 250.70 187.45 144.05 203.98
Eoangiopteris 324.64 307.06 360.02 249.16 237.78 259.29
Gemellitheca 287.12 251.09 336.57 261.10 243.47 298.00
Goolandia 115.95 93.60 165.89 87.91 58.77 104.90
Grandeuryella 317.16 304.81 356.31 292.88 254.47 305.87
Herbstopteris 240.50 235.07 246.94 234.13 218.90 240.89

80
Lobatopteris 321.19 314.69 328.21 302.77 298.82 305.43
Marantoidea 240.70 235.05 247.81 233.75 218.43 240.95
Marattia 228.47 214.15 242.05 extant
Marattiopsis 245.88 241.07 253.56 82.82 72.85 88.45
Megaphyton 353.05 314.32 377.35 253.39 240.49 262.19
Mesozoisynangia 110.88 94.09 138.99 90.41 69.52 103.65
Myriotheca 348.48 313.21 371.27 294.61 258.85 305.77
Ogmos 242.83 235.03 252.32 236.29 221.17 244.98
Pecopteris 378.55 368.32 385.03 41.54 32.49 51.52
Pectinangium 302.34 274.38 335.26 269.14 245.00 292.47
Protoblechnum 261.64 251.01 294.38 251.76 243.04 259.90
Psaronius 393.40 363.64 419.69 248.94 241.91 254.60
Ptychocarpus 326.72 302.56 364.38 248.67 236.41 259.35
Qasimia 293.26 253.67 335.02 249.14 238.59 259.31
Radstockia 320.63 300.39 360.43 296.26 254.99 317.41
Rajahia 260.48 252.99 268.69 250.87 246.00 256.26
Rhinipteris 229.41 202.90 247.22 205.41 151.62 234.75
Rienitsia 233.46 213.42 246.76 201.73 169.62 225.36
Scolecopteris 321.76 315.02 332.04 239.94 234.40 244.78
Sydneia 321.49 307.09 359.96 296.79 254.39 311.34
Symopteris 241.75 236.77 247.12 199.59 190.53 203.94
Taiyuanitheca 261.77 251.06 294.50 251.83 243.12 259.72
Tietea 297.71 256.59 341.00 257.45 241.64 292.34
Tranquillia 228.03 203.22 245.61 202.70 148.73 227.51
Tuvichapteris 293.22 251.83 335.95 246.70 236.17 253.72
Zhutheca 261.15 251.09 290.75 251.91 243.16 259.88
Paleozoic leptosporangiate
ferns
Anachoropteris 348.10 322.39 369.16 251.80 245.24 259.02
Ankyropteris 351.10 325.32 367.07 274.69 253.76 292.26
Apotropteris 314.78 299.20 356.91 291.00 253.87 305.94
Botryopteris 357.63 346.36 367.34 268.08 246.85 292.83
Catenopteris 317.41 299.03 357.87 292.51 253.05 311.56
Doneggia 314.81 299.23 356.16 289.62 252.51 305.87
Kaplanopteris 314.77 299.11 354.98 289.25 251.67 305.92
Norwoodia 324.94 311.55 361.26 300.54 256.93 314.56
Psalixochlaena 354.30 331.07 373.65 298.84 270.71 312.77
Rhabdoxylon 322.99 312.58 344.14 298.27 273.70 311.73
Senftenbergia 351.26 329.16 369.00 260.79 245.92 280.79
Sermaya 319.97 306.24 350.05 292.30 257.92 305.52
Skaaripteris 261.21 251.06 291.41 251.90 243.46 259.81

81
Tubicaulis 320.77 312.56 332.04 251.57 244.21 259.64
Osmundales
Anomopteris 253.70 251.70 257.11 219.22 201.61 233.76
Ashicaulis 249.93 241.04 264.82 70.99 41.36 98.30
Aurealcaulis 69.80 57.91 85.64 30.80 23.01 38.03
Bathypteris 261.15 251.14 292.83 251.61 243.2 259.54
Birmoltia 236.91 228.37 243.99 224.84 209.36 234.20
Birtodites 238.41 228.36 247.76 220.56 196.21 234.99
Cacumen 161.54 136.96 213.62 88.47 58.16 111.07
Chasmatopteris 261.43 251.01 289.26 252.32 243.94 259.86
Cladophlebis 291.58 286.99 298.91 40.06 33.95 50.55
Donwelliacaulis 242.93 235.34 251.45 236.68 222.62 244.77
Elantodites 236.02 231.34 240.10 227.2 220.63 231.33
Grammatopteris 347.99 326.93 368.65 252.84 238.96 274.67
Guairea 300.34 251.94 352.63 194.38 92.30 292.80
Iegosigopteris 260.99 251.02 287.52 251.64 243.11 259.60
Itopsidema 231.12 204.47 247.95 206.26 149.68 234.94
Leptopteris 36.76 28.78 47.36 extant
Lunea 192.32 148.49 241.07 149.44 86.73 199.26
Millerocaulis 234.38 214.3 245.72 106.11 85.44 122.41
Nymbopteron 241.88 235.31 251.24 223.61 206.49 234.9
Osmunda 220.75 203.94 236.93 extant
Osmundacaulis 249.66 241.03 262.25 70.54 36.56 96.96
Osmundastrum 73.97 40.67 133.15 extant
Osmundopsis 243.60 236.03 254.55 126.55 91.31 144.93
Paleosmunda 260.97 251.07 290.48 251.40 242.90 259.54
Parsorophyllum 248.19 241.22 258.28 222.55 204.50 234.72
Phyllopteroides 148.01 138.68 158.76 91.72 81.27 99.21
Rastropteris 307.33 284.19 350.20 276.43 247.74 298.55
Rooitodites 236.32 231.58 241.81 226.91 219.24 232.51
Shuichengella 261.70 252.05 281.37 251.18 243.09 258.49
Sphenopteris 326.42 321.78 331.68 33.65 28.29 39.43
Stormbergia 240.57 235.07 247.08 225.42 213.05 233.95
Thamnopteris 287.00 273.48 315.65 265.83 247.72 278.23
Todea 192.13 172.31 235.79 extant
Todites 257.66 252.88 260.78 119.43 99.77 142.16
Zalesskya 262.02 251.09 294.81 251.94 242.93 259.65
Zhongmingella 257.58 252.21 275.22 249.82 242.32 254.19
Hymenophyllales
Eogonocormus 146.99 105.53 195.28 101.73 68.98 144.47

82
Hopetedia 238.42 228.23 248.23 221.20 198.08 234.95
Hymenophyllites 362.55 345.11 377.92 97.41 61.54 134.10
Hymenophyllum 94.55 85.95 106.37 extant
Trichomanes 86.13 66.77 116.20 extant
Gleicheniales
Aninopteris 204.05 192.26 218.82 176.19 156.41 192.15
Antarctipteris 245.78 241.05 254.05 239.77 230.70 244.91
Boodlepteris 101.36 89.33 128.17 76.31 57.19 85.73
Camptopteris 247.52 235.22 256.42 195.14 185.86 201.71
Chansitheca 301.81 280.55 331.85 248.06 238.47 253.93
Clathropteris 248.34 240.03 255.08 149.32 140.42 161.02
Delosorus 139.37 113.30 194.89 104.41 67.64 128.76
Dicranopteris 87.57 81.37 97.97 extant
Dictyophyllum 249.47 244.37 253.75 145.58 142.89 149.15
Gleichenia 178.50 174.42 182.67 extant
Gleicheniopsis 133.83 100.89 177.89 48.29 25.13 63.97
Gleichenipteris 243.15 235.54 252.17 237.00 223.75 244.98
Gleichenites 245.21 242.50 247.53 49.82 34.90 58.51
Goeppertella 227.44 203.85 245.00 149.84 120.73 167.02
Hausmannia 241.30 238.51 245.11 60.16 58.98 61.49
Kenderlykia 209.05 179.17 244.91 165.28 95.06 203.87
Konijnenburgia 101.53 94.62 111.44 91.80 82.10 97.12
Laccopteris 243.31 226.00 255.84 50.40 33.03 59.52
Matonia 207.65 179.88 242.22 extant
Matonidium 183.66 171.83 207.34 76.78 60.48 85.24
Microphyllopteris 258.69 247.22 289.83 85.43 58.55 106.02
Nathorstia 191.14 150.08 240.50 64.48 31.32 91.29
Oligocarpia 352.79 329.76 369.55 210.01 196.69 227.39
Phlebopteris 239.86 235.58 244.26 102.71 92.64 115.24
Piazopteris 207.38 179.55 242.18 97.80 61.92 133.49
Selenocarpus 211.78 191.51 241.76 154.49 116.56 175.35
Soloropteris 245.90 241.08 254.34 239.96 231.13 244.99
Sticherus 87.17 66.10 117.16 extant
Szea 295.05 271.18 340.55 262.34 243.81 283.46
Tasmanopteris 194.31 151.39 241.77 151.19 86.89 198.73
Thaumatopteris 244.82 236.24 255.95 178.59 159.46 195.16
Tomaniopteris 242.34 235.16 251.02 236.05 222.22 244.95
Weichselia 187.45 175.05 200.19 79.12 69.67 84.40
Wingatea 231.16 209.17 245.87 202.8 170.56 220.49
Schizaeales

83
Anemia 140.01 131.13 146.54 extant
Baieropsis 141.90 111.66 170.41 76.76 52.93 104.44
Cynepteris 249.18 237.73 258.57 202.79 191.24 215.84
Eleganopteris 252.98 247.78 258.91 245.17 238.68 250.74
Klukia 189.73 178.76 201.77 108.71 99.32 115.14
Klukiopsis 189.65 161.88 239.50 143.11 83.54 175.66
Lygodium 111.02 81.89 144.86 extant
Mohriopsis 166.87 146.9 202.18 138.88 100.00 158.86
Norimbergia 229.92 201.67 245.48 205.88 149.7 234.21
Paralygodium 51.80 34.51 77.00 32.56 22.90 46.53
Pekinopteris 228.94 202.24 246.63 203.43 148.02 234.20
Pelletixia 144.79 136.16 159.41 108.20 89.51 124.08
Ruffordia 161.49 140.68 201.24 30.80 8.78 47.79
Schizaea 64.01 56.05 75.90 extant
Schizaeangium 151.53 104.21 210.47 115.43 70.12 158.99
Schizaeopsis 140.01 114.60 180.89 71.35 39.46 98.86
Schizaeopteris 95.03 65.69 139.40 73.00 42.85 99.24
Stachypteris 200.59 177.29 238.65 152.28 117.96 171.69
Salviniales
Azinia 95.39 66.22 137.16 74.01 45.31 99.52
Azolla 126.74 99.47 145.32 extant
Hydropteris 71.04 68.59 73.55 65.60 62.82 67.84
Marsilea 194.75 178.21 210.06 extant
Marsileaceaephyllum 112.47 108.98 116.65 99.15 95.04 102.81
Mendozaphyllum 87.15 65.53 130.59 65.49 34.65 83.49
Nammouria 111.84 93.52 160.61 86.06 58.36 99.47
Parazolla 80.41 65.57 119.45 60.04 31.46 70.51
Pilularia 89.70 66.22 123.21 extant
Regnellidium 102.2 83.83 142.19 extant
Regnellites 169.00 140.18 233.49 131.84 81.24 159.96
Rodeites 78.11 61.05 116.81 56.59 29.08 68.91
Salvinia 83.34 69.34 98.97 extant
Cyatheales
Acanthopteris 164.27 156.67 177.18 105.93 87.17 129.92
Alienopteris 137.65 112.86 194.44 101.83 65.19 124.72
Alsophila 16.13 2.64 53.15 extant
Alsophilites 192.54 149.33 240.23 121.24 84.69 139.78
Alsophilocaulis 40.00 23.08 75.20 23.39 2.37 33.32
Cibotiocaulis 124.05 65.69 190.28 94.08 54.82 143.67
Cibotium 101.11 66.95 143.19 extant

84
Conantiopteris 137.12 112.1 195.24 101.36 63.33 124.53
Coniopteris 329.43 303.57 358.08 37.15 28.36 46.31
Culcitites 172.75 146.01 235.63 134.36 82.61 160.57
Cyathea 222.57 203.47 239.62 extant
Cyathocarpus 314.82 298.99 355.06 290.13 252.63 306.7
Cyathocaulis 171.81 145.92 210.31 64.03 30.42 89.69
Cyathodendron 55.26 53.27 56.87 33.52 32.38 34.28
Dendropteridium 27.26 6.09 53.60 5.44 0.00 17.17
Dicksonia 203.66 192.1 215.19 extant
Eboracia 206.92 197.96 221.27 116.64 90.29 142.2
Eocyathea 136.60 112.27 190.14 102.27 66.99 124.82
Gonatosorus 214.70 193.5 244.62 102.64 67.59 142.75
Haydenia 177.66 156.13 235.69 139.52 87.14 160.94
Kwazulupteris 115.76 91.69 148.52 87.62 63.30 107.81
Kylikipteris 207.44 181.45 242.14 138.71 98.49 157.19
Lophosoria 148.45 112.7 222.62 extant
Loxsomopteris 137.07 112.35 192.68 102.59 64.99 124.91
Natalipteris 115.72 93.81 149.88 87.60 61.18 106.55
Nishidacaulis 142.65 101.55 203.43 107.72 67.69 143.9
Oguracaulis 152.13 102.68 214.99 116.22 71.30 160.39
Onychiopsis 182.03 168.74 203.01 83.03 80.02 85.25
Paracyathocaulis 112.87 65.78 168.63 85.44 50.00 124.32
Protopteris 147.38 127.13 180.55 81.23 59.89 93.28
Rickwoodopteris 88.89 70.61 127.25 68.02 39.70 83.42
Sergioa 138.32 112.31 187.66 74.22 46.73 88.66
Solenostelopteris 154.11 130.21 214.40 117.06 71.92 139.79
Thyrsopteris 340.63 304.41 372.07 extant
Polypodiales
Acrostichum 71.42 67.84 75.78 extant
Adiantopteris 239.34 228.11 253.20 94.54 63.54 125.83
Adiantum 126.72 114.15 139.20 extant
Allantodiopsis 89.92 79.48 100.83 33.22 30.87 35.00
Anogramma 6.10 2.65 15.90 extant
Antrophyum 25.69 19.82 35.64 extant
Arthropteris 10.27 1.83 42.68 extant
Aspidistes 185.45 173.02 205.28 127.26 113.45 139.18
Asplenium 236.24 228.85 242.56 extant
Astralopteris 132.17 100.12 174.07 87.06 66.15 97.26
Athyrium 155.79 127.85 199.84 extant
Blechnum 151.01 141.00 163.04 extant

85
Ceratopteris 58.71 34.59 94.73 extant
Coniogramme 36.10 5.56 83.58 extant
Cretacifilix 138.47 99.82 197.11 104.69 65.11 142.01
Cryptogrammites 42.12 28.47 74.43 26.28 6.44 33.82
Ctenitis 7.68 2.07 23.17 extant
Cuyenopteris 39.08 23.02 73.30 23.13 2.40 32.97
Davallia 56.83 37.12 72.57 extant
Dennastra 77.66 61.72 118.94 54.53 27.64 66.38
Dennstaedtia 175.48 161.69 190.77 extant
Dennsteadtiopsis 50.71 40.74 70.04 30.38 23.03 36.42
Deparia 19.37 5.36 47.31 extant
Diplazium 71.37 34.81 134.93 extant
Drynaria 13.73 6.26 25.78 extant
Dryopteris 173.58 164.55 180.63 extant
Elaphoglossum 40.49 15.51 88.12 extant
Eogymnocarpium 139.66 99.85 199.93 104.38 63.47 145.14
Grammitis 24.20 15.32 39.13 extant
Hemionitis 81.67 57.77 123.27 extant
Histiopteris 36.40 28.79 46.81 extant
Hypolepis 23.14 16.15 33.99 extant
Krameropteris 121.81 98.03 177.85 91.22 60.57 111.25
Lastreopsis 16.47 2.64 54.64 extant
Lepisorus 38.04 28.70 52.09 extant
Leptolepia 8.11 0.06 36.88 extant
Lindsaea 40.64 34.07 50.81 extant
Makotopteris 59.12 40.52 101.36 35.54 18.91 48.40
Microlepiopsis 86.43 71.78 109.24 68.57 50.79 82.72
Microsorum 47.09 37.70 57.46 extant
Midlandia 88.43 70.68 128.09 67.59 38.66 83.38
Nephrolepis 10.64 5.39 17.68 extant
Oleandra 134.53 119.57 152.44 extant
Onoclea 112.70 94.05 141.68 extant
Paesia 40.39 34.17 50.31 extant
Pellaea 10.04 1.81 40.94 extant
Pleopeltis 41.29 16.11 91.51 extant
Polypodium 52.24 34.18 72.69 extant
Polystichum 79.86 65.63 108.17 extant
Protodrynaria 43.78 28.56 78.04 27.19 8.37 37.15
Pteridium 37.05 33.98 40.91 extant
Pteris 99.83 95.61 104.76 extant

86
Pyrrosia 5.60 3.64 9.20 extant
Rumohra 49.74 34.46 66.21 extant
Saccoloma 102.9 94.76 114.88 extant
Salpichlaena 74.39 68.98 84.41 extant
Stramineopteris 153.12 130.22 209.92 116.95 71.89 139.9
Tecaropteris 55.63 23.09 97.41 35.14 9.63 64.25
Tectaria 135.30 123.91 151.61 extant
Thelypteridaceophyllum 7.82 1.83 31.06 1.71 0.08 3.17
Thelypteris 76.00 66.42 85.21 extant
Trawetsia 55.73 37.34 92.61 34.09 16.55 48.46
Wessiea 97.48 70.75 149.79 5.63 0.01 14.40
Woodwardia 121.92 97.58 145.22 extant
Incertae cedis
Acrostichopteris 228.12 209.77 242.32 101.06 86.50 114.80
Adiantitophyllum 124.06 100.12 176.83 92.15 60.37 111.88
Asplenites 239.70 229.49 251.35 61.83 29.46 82.22
Boweria 355.30 329.70 376.35 311.2 271.78 349.97
Crenulopteris 318.51 314.97 323.44 302.42 297.86 305.14
Daeniopteris 254.78 245.32 274.99 203.07 163.95 233.80
Dictyonymba 242.78 235.11 251.93 235.73 220.34 244.91
Diplazites 316.65 304.08 354.62 292.00 252.83 305.95
Discopteris 320.49 300.39 360.80 296.08 254.98 317.60
Displinites 237.19 229.57 244.98 224.82 209.63 234.00
Dryopterites 147.38 121.65 177.66 97.73 68.36 117.11
Fremouwa 245.82 241.07 253.97 240.12 231.80 244.89
Gordonopteris 247.85 242.24 255.60 234.45 223.93 240.96
Gouldiopteris 242.82 235.22 251.83 236.40 221.21 244.98
Holmesopteris 317.07 304.36 340.51 299.23 273.46 312.57
Katasiopteris 253.92 251.34 258.17 245.67 239.89 250.73
Korallipteris 100.63 86.41 144.57 78.04 50.75 88.62
Leconama 242.81 235.02 251.65 236.63 222.96 244.87
Micronymbopteris 242.44 235.13 251.26 235.99 220.91 244.93
Molteniella 238.09 228.22 247.67 220.67 195.86 234.77
Neuropteridium 244.16 242.33 245.50 234.96 234.46 235.42
Nunatakia 124.27 99.96 177.83 92.33 60.22 111.95
Nymbofelicia 242.54 235.06 251.27 236.25 221.52 245.00
Nymboidiantum 239.78 235.60 244.62 226.89 218.88 232.09
Nymbophlebis 243.07 235.06 252.54 236.91 222.34 244.95
Nymborhipteris 243.10 235.14 252.04 236.40 221.32 244.83
Paratrizygia 297.04 273.34 329.02 267.49 245.56 287.35

87
Pterisorus 154.11 130.05 216.68 117.31 72.19 139.79
Ptilotonymba 242.97 235.05 251.58 236.50 222.40 244.96
Raphaelia 225.15 210.45 237.50 127.38 101.08 144.06
Renaultia 336.03 322.20 360.50 305.38 298.30 309.38
Schleporia 246.14 241.17 255.00 240.07 231.23 244.92
Schopfiopteris 245.83 241.22 254.44 239.79 231.03 244.96
Scolopendrites 253.71 251.71 257.12 238.26 233.28 243.61
Selleyopteris 172.97 151.03 235.61 134.13 83.26 155.67
Sendersonia 262.39 251.14 297.38 252.13 243.40 259.96
Speirsopteris 74.64 56.86 117.84 48.27 25.00 60.17
Tempskya 143.78 136.94 149.54 69.65 61.94 83.69
Trizygia 294.32 272.95 311.63 249.17 241.68 255.64
Yavanna 136.98 112.11 192.36 101.83 65.81 124.91
Yulebacaulis 142.60 100.23 205.12 107.04 65.22 143.72

Supplementary Table 2. Molecular age estimates.

crown age
Clade stem age (Ma) 95% HPD 95% HPD
(Ma)
MONILOPHYTES n.a. n.a. 421.3 379.16-465.97
EQUISETIDAE 421.3 379.16-465.97 80.1 50.87-113.95
EQUISETALES 421.3 379.16-465.97 80.1 50.87-113.95
Equisetaceae 421.3 379.16-465.97 80.1 50.87-113.95
Equisetum 421.3 379.16-465.97 80.1 50.87-113.95
OPHIOGLOSSIDAE 358.29 318.00-401.48 244.2 186.64-306.29
OPHIOGLOSSALES 244.2 186.64-306.29 148.94 108.17-187.78
Ophioglossaceae 244.2 186.64-306.29 148.94 108.17-187.78
Botrychium 29.63 14.51-48.37 16.1 6.00-28.52
Botrypus 54.61 27.25-86.01 22.44 14.89-45.08
Cheiroglossa not included
Helminthostachys 119.79 84.78-159.10 n.a. n.a.
Japanobotrychium not included
Mankyua not included
Ophioderma 69.8 41.38-101.05 n.a. n.a.
Ophioglossum 69.8 41.38-101.05 n.a. n.a.
Rhizoglossum not included
PSILOTALES 244.2 186.64-306.29 55.82 123.40-91.76
Psilotaceae 244.2 186.64-306.29 55.82 123.40-91.76
Psilotum 55.82 23.40-91.76 n.a. n.a.

88
Tmesipteris 55.82 23.40-91.76 n.a. n.a.
MARATTIIDAE 358.29 318.00-401.48 202.71 185.29-224.20
MARATTIALES 358.29 318.00-401.48 202.71 185.29-224.20
Marattiaceae 358.29 318.00-401.48 202.71 185.29-224.20
Angiopteris 182.69 172.00-198.04 1.78 0.06-4.69
Christensenia 202.71 185.29-224.20 n.a. n.a.
Danaea 188.8 177.71-203.20 30.43 17.29-45.96
Eupodium 181.39 176.19-191.92 n.a. n.a.
Marattia 182.69 172.00-198.04 n.a. n.a.
Ptisana 181.39 176.00-191.92 17.91 8.06-29.84
POLYPODIIDAE 414.34 374.15-456.27 377.19 339.23-414.21
OSMUNDALES 377.19 339.23-414.21 241.26 217.19-270.38
Osmundaceae 377.19 339.23-414.21 241.26 217.19-270.38
Claytosmunda embedded in Osmunda
Leptopteris 155.27 136.00-191.72 12.1 2.44-26.16
Osmunda 223.92 208.04-243.87 206.75 201.60-217.65
Osmundastrum 241.26 217.19-270.38 12 1.12-30.70
Plenasium embedded in Osmunda
Todea 155.27 136.00-191.72 n.a. n.a.
HYMENOPHYLLALES 315.27 282.11-348.57 216.09 175.76-255.40
Hymenophyllaceae 315.27 282.11-348.57 216.09 175.76-255.40
TRICHOMANOIDEAE 216.09 175.76-255.40 175.15 142.88-206.42
Abrodictyum 97.82 58.14-139.30 56.71 27.70-93.68
Callistopteris 128.45 84.72-169.36 n.a. n.a.
Cephalomanes 97.82 58.14-139.30 n.a. n.a.
Crepidomanes 130.04 103.42-157.17 88.5 63.90-114.91
Didymoglossum 140.78 114.95-167.68 103.56 79.33-127.81
Polyphlebium 151.65 123.66-178.90 62.46 30.86-98.15
Trichomanes 152.15 118.79-185.83 99.47 73.34-128.01
Vandenboschia 130.04 103.42-157.17 27.11 5.77-55.76
HYMENOPHYLLOIDEAE 216.09 175.76-255.40 47.37 35.92-60.01
Hymenophyllum 216.09 175.76-255.40 47.37 35.92-60.01
GLEICHENIALES 280.44 261.59-320.01 134.09 110.23-161.65
Gleicheniaceae 280.44 261.59-320.01 134.09 110.23-161.65
Dicranopteris 32.42 14.24-54.56 n.a. n.a.
Diplopterygium 91.74 55.83-127.57 28.83 12.00-50.02
Gleichenella 32.42 14.24-54.56 n.a. n.a.
Gleichenia 95.45 89.30-107.47 n.a. n.a.
Sticherus 110.35 95.11-129.08 76.99 37.80-110.62
Stromatopteris 95.45 89.30-107.47 n.a. n.a.

89
Dipteridaceae 242.48 235.00-256.62 50.38 22.57-87.25
Cheiropleuria 50.38 22.57-87.25 n.a. n.a.
Dipteris 50.38 22.57-87.25 n.a. n.a.
Matoniaceae 242.48 235.00-256.62 96.6 49.83-149.11
Matonia 96.6 49.83-149.11 n.a. n.a.
Phanerosorus 96.6 49.83-149.11 n.a. n.a.
SCHIZAEALES 322.79 290.55-354.72 250.76 213.60-287.39
Lygodiaceae 250.76 213.60-287.39 45.58 23.31-72.61
Lygodium 250.76 213.60-287.39 45.58 23.31-72.61
Schizaeaceae 190.36 161.39-222.54 91.32 51.37-133.42
Actinostachys 91.32 51.37-133.42 n.a. n.a.
Schizaea 91.32 51.37-133.42 24.01 15.59-48.87
Anemiaceae 190.36 161.39-222.54 145.26 136.00-162.15
Anemia 190.36 161.39-222.54 145.26 136.00-162.15
SALVINIALES 290.3 260.52-320.25 195.55 155.00-235.65
Marsileaceae 195.55 155.00-235.65 109.29 99.60-125.22
Marsilea 109.29 99.60-125.22 31.5 18.17-46.60
Pilularia 90.26 83.50-101.25 9.44 2.39-20.26
Regnellidium 90.26 83.50-101.25 n.a. n.a.
Salviniaceae 195.55 155.00-235.65 105.13 83.50-130.34
Azolla 105.13 83.50-130.34 52.39 34.91-70.97
Salvinia 105.13 83.50-130.34 49.73 29.85-71.40
CYATHEALES 279 249.60-307.72 150.76 133.22-171.67
Thyrsopteridaceae 140.81 123.85-160.72 n.a. n.a.
Thyrsopteris 140.81 123.85-160.72 n.a. n.a.
Loxsomataceae 121.24 112.00-137.26 23.5 8.66-42.22
Loxsoma 23.5 8.66-42.22 n.a. n.a.
Loxsomopsis 23.5 8.66-42.22 n.a. n.a.
Culcitaceae 62.89 34.77-95.55 14.54 4.21-28.85
Culcita 62.89 34.77-95.55 14.54 4.21-28.85
Plagiogyriaceae 62.89 34.77-95.55 31.89 15.99-50.50
Plagiogyria 62.89 34.77-95.55 31.89 15.99-50.50
Cibotiaceae 128.9 120.34-140.29 17.48 8.67-28.21
Cibotium 128.9 120.34-140.29 17.48 8.67-28.21
Cyatheaceae 132.06 125.00-142.80 30.28 21.65-41.12
Alsophila 24.18 17.38-31.72 16.29 10.91-22.26
Cyathea 24.18 17.38-31.72 14.43 9.63-19.49
Gymnosphaera 26.47 19.07-34.80 16.61 9.30-24.25
Sphaeropteris 30.28 21.65-41.12 27.15 17.69-37.01
Dicksoniaceae 127.43 118.31-138.30 118.77 112.72-126.46

90
Calochlaena 118.77 112.72-126.46 26.66 9.83-50.65
Dicksonia 115.94 112.00-122.86 10.1 2.95-20.13
Lophosoria 115.94 112.00-122.86 n.a. n.a.
Metaxyaceae 127.43 118.31-138.30 n.a. n.a.
Metaxya 127.43 118.31-138.30 n.a. n.a.
POLYPODIALES 279 249.60-307.72 248.81 221.72-273.80
Lonchitidaceae 182.43 155.69-211.48 n.a. n.a.
Lonchitis 182.43 155.69-211.48 n.a. n.a.
Saccolomataceae 235.21 208.09-262.00 15.16 5.77-27.39
Saccoloma 235.21 208.09-262.00 15.16 5.77-27.39
Cystodiaceae 221.07 194.15-248.43 n.a. n.a.
Cystodium 221.07 194.15-248.43 n.a. n.a.
Lindsaeaceae 182.43 155.69-211.48 127.49 106.54-150.87
Lindsaea 65.51 52.99-79.27 44.22 32.28-56.23
Nesolindsaea 40.98 29.24-52.95 n.a. n.a.
Odontosoria Old World 127.49 106.54-150.87 15.76 3.55-35.78
Odontosoria New World 79.91 64.93-95.64 n.a. n.a.
Osmolindsaea 48.57 36.61-61.18 10.44 2.73-20.53
Sphenomeris 71.25 57.63-86.04 n.a. n.a.
Tapeinidium 40.98 29.24-52.95 20.32 11.50-29.69
Xyropteris not included
Pteridaceae 223.92 199.06-247.93 177.61 155.14-200.82
CRYPTOGRAMMOIDEAE 161.03 136.24-186.88 79.99 49.15-115.07
Coniogramme 58.7 33.00-88.65 21.72 7.73-38.82
Cryptogramma 58.7 33.00-88.65 n.a. n.a.
Llavea 79.99 49.15-115.07 n.a. n.a.
PARKERIOIDEAE 161.03 136.24-186.88 144.2 119.48-169.89
Acrostichum 80.44 56.56-106.23 38.85 18.04-61.85
Ceratopteris 80.44 56.56-106.23 2.07 0.01-6.10
PTERIDOIDEAE 144.2 119.48-169.89 103.34 82.40-123.89
Actiniopteris 41.39 17.48-70.26 n.a. n.a.
Anogramma 17.8 3.28-37.25 n.a. n.a.
Austrogramme not included
Cerosora not included
Cosentinia not included
Jamesonia 57.67 42.02-73.47 25.86 14.69-38.39
Onychium 41.39 17.48-70.26 n.a. n.a.
Pityrogramma 40.46 23.01-58.89 8.83 1.83-18.71
Pteris 103.34 82.40-123.89 89.01 61.35-114.47
Pterozonium 29.94 11.97-50.65 n.a. n.a.

91
Syngramma not included
Taenitis not included
Tryonia 57.37 11.97-50.65 n.a. n.a.
CHEILANTHOIDEAE 156.11 135.56-178.76 135.21 102.05-166.43
Adiantopsis 31.99 23.36-40.92 n.a. n.a
Aleuritopteris 35 21.52-48.05 25.71 12.22-39.18
Allosorus 35 21.52-48.05 n.a. n.a.
Argyrochosma 48.06 34.35-62.37 29.35 15.31-44.23
Aspidotis 32.37 19.99-44.96 14.71 4.86-26.22
Astrolepis not included
Bommeria 93.74 71.05-119.26 26.98 7.24-53.85
Calciphilopteris 135.21 102.05-166.43 n.a. n.a.
Cheilanthes not monophyletic
Cheiloplecton 33.88 23.37-45.22 n.a. n.a.
Doryopteris 24.88 16.36-33.81 12.33 5.58-19.92
Gaga 32.37 19.99-44.96 n.a. n.a.
Hemionitis 23.07 13.35-32.59 n.a. n.a.
Lytoneuron not included
Mildella 52.09 36.00-68.56 n.a. n.a.
Myriopteris 52.09 36.00-68.56 37.9 25.15-51.68
Notholaena not monophyletic
Ormopteris 24.88 16.36-33.81 n.a. n.a.
Paragymnopteris embedded in Pellaea
Parahemionitis 23.07 13.35-32.59 n.a. n.a.
Pellaea 48.06 34.35-62.37 36.43 25.24-49.03
Pentagramma 48.79 38.11-60.28 n.a. n.a.
Trachypteris not included
VITTARIOIDEAE 156.11 135.56-178.76 141.16 121.06-161.49
Adiantum 141.16 121.06-161.49 115.94 87.40-143.20
Ananthacorus not included
Antrophyum 78.53 63.03-94.54 n.a. n.a.
Haplopteris 67.61 51.84-85.58 43.23 28.36-58.41
Hecistopteris 44.58 28.92-60.78 n.a. n.a.
Polytaenium 56.25 41.73-70.48 34.31 21.63-47.95
Radiovittaria 44.58 28.92-60.78 n.a. n.a.
Rheopteris 103.51 85.63-122.69 n.a. n.a.
Scoliosorus not included
Vaginularia not included
Vittaria 56.25 41.73-70.48 23.71 10.38-37.92
Dennstaedtiaceae 213.89 189.42-237.99 125.06 93.62-160.85

92
Blotiella 37.57 20.76-55.51 n.a. n.a.
Dennstaedtia 56.08 37.47-74.75 28.86 11.85-47.75
Histiopteris 37.57 20.76-55.51 n.a. n.a.
Hypolepis 56 36.87-77.27 n.a. n.a.
Leptolepia 83.55 70.60-104.22 30.25 11.24-53.26
Microlepia 56.08 37.47-74.75 16.36 8.30-26.14
Monachosorum 102.19 70.61-136.78 n.a. n.a.
Oenotrichia not included
Paesia 50.38 31.78-71.75 n.a. n.a.
Pteridium 61.69 40.42-84.38 11.33 3.91-20.91
Cystopteridaceae 91.53 53.17-133.49 62.29 35.35-93.75
Acystopteris 34.72 16.16-57.57 12.89 1.26-30.67
Cystopteris 34.72 16.16-57.57 n.a. n.a.
Gymnocarpium 62.29 35.35-93.75 18.36 4.49-36.92
Hemidictyaceae 137.04 114.28-161.35 n.a. n.a.
Hemidictyum 137.04 114.28-161.35 n.a. n.a.
Aspleniaceae 137.04 114.28-161.35 104.15 85.96-123.03
Asplenium 104.15 85.96-123.03 82.59 68.93-97.47
Hymenasplenium 104.15 85.96-123.03 56.09 34.04-80.00
Diplaziopsidaceae 91.53 53.17-133.49 40.34 17.19-70.02
Diplaziopsis 40.34 17.19-70.02 n.a. n.a.
Homalosorus 40.34 17.19-70.02 n.a. n.a.
Desmophlebiaceae not included
Desmophlebium not included
Thelypteridaceae 119.5 96.77-143.87 89.91 70.73-110.68
PHEGOPTERIDOIDEAE 89.91 70.43-110.68 59.96 34.57-85.89
Macrothelypteris 59.96 34.57-85.89 n.a. n.a.
Phegopteris 29.7 16.50-45.38 16.51 6.62-28.10
Pseudophegopteris 29.7 16.50-45.38 15.41 3.64-29.79
THELYPTERIDOIDEAE 89.91 70.43-110.68 77.4 60.94-94.55
Amauropelta embedded in Thelypteris
Amblovenatum embedded in Thelypteris
Ampelopteris not included
Chingia embedded in Thelypteris
Christella embedded in Thelypteris
Coryphopteris embedded in Thelypteris
Cyclogramma not included
Cyclosorus embedded in Thelypteris
Glaphyropteridopsis embedded in Thelypteris
Goniopteris embedded in Thelypteris

93
Meniscium embedded in Thelypteris
Menisorus not included
Mesophlebion not included
Mesopteris not included
Metathelypteris embedded in Thelypteris
Nannothelypteris not included
Oreopteris embedded in Thelypteris
Parathelypteris not included
Plesioneuron not included
Pneumatopteris embedded in Thelypteris
Pronephrium embedded in Thelypteris
Pseudocyclosorus embedded in Thelypteris
Sphaerostephanos embedded in Thelypteris
Stenogramma embedded in Thelypteris
Steiropteris embedded in Thelypteris
Thelypteris 89.91 70.43-110.68 77.4 60.94-94.55
Trigonospora embedded in Thelypteris
Woodsiaceae 110.9 90.92-132.19 62.55 38.33-89.01
Woodsia 110.9 90.92-132.19 62.55 38.33-89.01
Rhachidosoraceae 129.76 102.96-158.67 n.a. n.a.
Rhachidosorus 129.76 102.96-158.67 n.a. n.a.
Onocleaceae 91.75 76.40-107.94 66.46 55.80-79.43
Onoclea 91.75 76.40-107.94 66.46 55.80-79.43
Onocleopsis 42.24 16.84-63.74 n.a. n.a.
Matteuccia 42.24 16.84-63.74 n.a. n.a.
Blechnaceae 91.75 76.40-107.94 71.62 58.81-84.97
STENOCHLAENOIDEAE 67.84 54.58-80.61 53.34 38.72-68.22
Salpichlaena 53.34 38.72-68.22 13.1 4.09-21.51
Stenochlaena 41.85 25.66-58.02 2.45 0.00-9.30
Telmatoblechnum 41.85 25.66-58.02 n.a. n.a.
WOODWARDIOIDEAE 71.62 58.81-84.97 46.72 29.81-63.02
Anchistea embedded in Woodwardia
Lorinseria embedded in Woodwardia
Woodwardia 71.62 58.81-84.97 46.72 29.81-63.02
BLECHNOIDEAE 67.84 54.58-80.61 57.93 44.85-71.66
Austroblechnum embedded in Blechnum
Blechnidium embedded in Blechnum
Blechnopsis embedded in Blechnum
Blechnum 67.84 54.58-80.61 57.93 44.85-71.66
Brainea embedded in Blechnum

94
Cleistoblechnum embedded in Blechnum
Cranfilia embedded in Blechnum
Diploblechnum embedded in Blechnum
Doodia embedded in Blechnum
Icarus embedded in Blechnum
Lomaria embedded in Blechnum
Lomaridium embedded in Blechnum
Lomariocycas embedded in Blechnum
Neoblechnum embedded in Blechnum
Oceaniopteris embedded in Blechnum
Parablechnum embedded in Blechnum
Sadleria embedded in Blechnum
Struthiopteris embedded in Blechnum
Athyriaceae 105.23 87.24-124.38 88.85 67.04-111.17
Athyrium 74.29 54.43-95.29 43.13 24.77-64.09
Deparia 88.85 67.04-111.17 27.5 11.64-46.84
Diplazium 74.29 54.43-95.29 48.17 33.27-65.02
Didymochlaenaceae 144.01 99.99-175.90 n.a. n.a.
Didymochlaena 144.01 99.99-175.90 n.a. n.a.
Hypodematiaceae 144.01 99.99-175.90 72.68 30.37-123.07
Hypodematium 72.68 30.37-123.07 n.a. n.a.
Leucostegia 72.68 30.37-123.07 15.67 3.87-33.46
Dryopteridaceae 152.29 131.84-172.99 122.02 99.29-144.76
Aenigmopteris not included
Dryopolystichum not included
DRYOPTERIDOIDEAE 111.58 86.61-135.14 98.81 75.55-122.53
Arachniodes 77.2 56.74-97.75 50.99 35.80-67.38
Ctenitis 76.9 56.42-99.00 37.49 22.18-54.19
Cyrtomium 39.56 29.09-50.57 15.66 6.95-25.23
Dryopteris 77.2 56.74-97.75 45.82 33.49-59.02
Phanerophlebia 48.63 35.48-63.12 n.a. n.a.
Polystichum 39.56 29.09-50.57 34.95 25.49-44.86
POLYBOTRYOIDEA 111.58 86.61-135.14 95.57 62.98-127.00
Cyclodium embedded in Polybotrya
Maxonia 31.83 19.62-46.00 n.a. n.a.
Olfersia embedded in Polybotrya
Polybotrya 31.83 19.62-46.00 27.73 17.26-39.57
Polystichopsis 73.34 43.70-105.09 n.a. n.a.
Stigmatopteris 95.57 62.98-127.00 14.58 4.38-29.09
Trichoneuron not included

95
ELAPHOGLOSSOIDEAE 122.02 99.29-144.76 98.53 78.53-119.71
Arthrobotrya not included
Bolbitis 70 54.80-85.44 42.82 24.93-61.84
Elaphoglossum 58.83 45.95-72.27 46.67 35.72-58.67
Lastreopsis 49.08 25.96-73.53 32.27 11.60-56.05
Lomagramma 40.74 19.56-61.27 n.a. n.a.
Megalastrum 36.81 17.28-59.41 n.a. n.a.
Mickelia 58.83 45.95-72.27 45.34 29.16-61.80
Parapolystichum 80.18 48.75-108.96 23.38 7.50-45.52
Pleocnemia 91.02 72.02-111.37 n.a. n.a.
Rumohra 36.81 17.28-59.41 n.a. n.a.
Teratophyllum 40.74 19.56-61.27 n.a. n.a.
Lomariopsidaceae 134.71 114.80-155.38 99.13 73.79-124.14
Cyclopeltis 99.13 73.79-124.14 26.39 8.92-50.50
Dracoglossum 78.4 55.19-102.75 n.a. n.a.
Lomariopsis 78.4 55.19-102.75 29.03 17.45-41.59
Thysanosoria not included
Nephrolepidaceae 134.71 114.80-155.38 61.62 32.40-95.12
Nephrolepis 134.71 114.80-155.38 61.62 32.40-95.12
Tectariaceae 119.39 103.22-135.80 97.99 69.33-122.48
Arthropteris 97.99 69.33-122.48 39.17 15.68-67.16
Draconopteris not included
Hypoderris not included
Malaifilix not included
Pteridrys 83.43 58.37-108.82 n.a. n.a.
Tectaria 66.78 46.21-89.61 44.48 31.31-60.06
Triplophyllum 66.78 46.21-89.61 n.a. n.a.
Oleandraceae 114.36 98.64-129.78 32.05 5.63-80.22
Oleandra 114.36 98.64-129.78 32.05 5.63-80.22
Davalliaceae 108.91 94.53-123.99 63.11 41.19-88.53
Davallia 108.91 94.53-123.99 63.11 41.19-88.53
Polypodiaceae 108.91 94.53-123.99 101.47 87.48-114.93
LOXOGRAMMOIDEAE 101.47 87.48-114.93 73.77 55.16-91.40
Dictymia 73.77 55.16-91.40 27.43 8.73-50.81
Loxogramme 73.77 55.16-91.40 45.55 29.41-63.48
DRYNARIOIDEAE 90.1 77.99-102.13 67.17 39.70-93.39
Aglaomorpha 59.65 37.55-86.72 40.17 25.30-57.70
Arthromeris 28.11 17.42-39.52 12.07 2.97-23.37
Gymnogrammitis embedded in Selliguea
Paraselliguea not included

96
Phymatopteris not included
Polypodiopteris embedded in Selliguea
Selliguea 28.11 17.42-39.52 25.96 16.27-36.49
PLATYCERIOIDEAE 84.17 72.04-96.46 71.5 58.82-84.72
Platycerium 71.5 58.82-84.72 46.68 35.46-61.96
Pyrrosia 71.5 58.82-84.72 49.04 35.86-62.84
MICROSOROIDEAE 84.17 72.04-96.46 72.8 60.90-85.12
Dendroconche not included
Goniophlebium 66.26 54.85-77.38 43.18 26.75-59.79
Kaulinia not included
Kontumia not included
Lecanopteris embedded in Microsorum
Lemmaphyllum 32.23 22.83-41.64 17.55 8.56-27.34
Lepidomicrosorium 17.76 9.59-26.80 10.85 4.76-17.69
Lepisorus 42.48 33.41-51.66 37.92 28.61-47.18
Leptochilus 37.47 27.51-48.14 30.43 21.33-39.76
Microsorum 66.26 54.85-77.38 64.25 53.52-75.21
Neocheiropteris 17.76 9.59-26.80 n.a. n.a.
Neolepisorus 27.64 18.86-36.80 19.71 11.23-29.08
Paragramma 46.4 5.69-21.29 n.a. n.a.
Podosorus not included
Thylacopteris 72.8 60.90-85.12 n.a. n.a.
Tricholepidium 22.67 13.51-32.20 n.a. n.a.
POLYPODIOIDEAE 89.49 77.57-101.49 87.08 75.59-98.92
Acrosorus not included
Adenophorus 45.35 34.19-57.09 31.38 20.92-43.10
Alansmia 49.18 36.23-61.96 22.39 9.46-36.99
Archigrammitis not included
Ascogrammitis 52.05 43.68-60.77 35.52 24.10-47.22
Calymmodon 39.4 29.36-49.51 23.36 12.35-35.16
Campyloneurum 68.11 47.82-86.77 37.55 20.87-57.80
Ceradenia 42.27 29.73-54.62 32.35 20.86-44.57
Chrysogrammitis 47.29 37.25-57.04 n.a. n.a.
Cochlidium 36.37 25.25-48.10 25.4 15.34-36.66
Ctenopterella not included
Ctenopteris not monophyletic
Dasygrammitis not included
Enterosora not monophyletic
Galactodenia 42.77 34.52-50.98 n.a. n.a.
Grammitis 36.37 25.25-48.10 28.03 15.89-40.49

97
Lellingeria 38.02 30.26-45.64 32.57 24.91-40.58
Leucostrichum 49.18 36.23-61.96 31.11 18.85-44.47
Lomaphlebia not included
Luisma not included
Melpomene 36.15 28.14-43.80 26.95 15.53-37.78
Microgramma 77.37 60.78-92.21 45.68 30.42-62.98
Micropolypodium not included
Moranopteris 47.29 37.25-57.04 25.7 15.93-36.35
Mycopteris 45.03 36.66-53.27 30.06 18.11-42.49
Niphidium 68.11 47.82-86.77 n.a. n.a.
Notogrammitis 32.39 18.76-40.54 21.53 11.84-32.06
Oreogrammitis not monophyletic
Pecluma 62.91 44.95-80.11 43.26 29.29-58.73
Phlebodium 62.91 44.95-80.11 18.69 3.77-41.45
Pleopeltis 72.88 53.68-88.90 35.65 24.37-49.00
Pleurosoriopsis 49.08 27.17-75.18 n.a. n.a.
Polypodium 49.08 27.17-75.18 36.47 20.41-54.17
Prosaptia 20.86 12.92-29.40 9.99 4.81-15.90
Radiogrammitis not monophyletic
Scleroglossum 31.78 19.91-43.99 n.a. n.a.
Serpocaulon 69.32 47.88-87.53 44.5 29.26-60.95
Stenogrammitis 36.15 28.14-43.80 24 16.59-31.69
Synammia 67.17 39.70-93.39 10.55 1.28-24.98
Terpsichore 67.77 58.32-77.15 37 19.86-54.17
Themelium 16.07 5.33-28.40 n.a. n.a.
Tomophyllum embedded in Grammitis
1
Unnamed genus 69.32 47.88-87.53 n.a. n.a.
Xiphopterella not included
Zygophlebia not included
Ma=millions of years
HPD=highest porterior density
1
Polypodium chrysolepis Hook. is not resolved as a member of any named genus, but as a sister lineage
to Serpocaulon.

Supplementary Table 3. Minimum divergence

incongruence metrics.
Fossil estimate Molecular min-
Taxon MDI
95% CI max estimate
Acrostichum 67.84 75.78 18.04 106.23 0
Adiantum 114.15 139.20 87.40 161.49 0
Alsophila 2.64 53.15 10.91 31.72 0

98
Anemia 131.13 146.54 136.00 222.54 0
Angiopteris 208.94 247.17 0.06 198.04 -10.9
Anogramma 2.65 15.90 <3.28 37.25 0
Antrophyum 19.82 35.64 <63.03 94.54 n.a.
Arthropteris 1.83 42.68 15.68 122.48 0
Asplenium 228.85 242.56 68.93 123.03 -105.82
Athyrium 127.85 199.84 24.77 95.29 -32.56
Azolla 99.47 145.32 34.91 130.34 0
Blechnum 141.00 163.04 44.85 80.61 -60.39
Botrychium 57.87 110.55 27.25 187.78 0
Ceratopteris 34.59 94.73 0.01 106.23 0
Cibotium 66.95 143.19 8.67 140.29 0
Coniogramme 5.56 83.58 7.73 88.65 0
Ctenitis 2.70 23.17 22.18 99.00 0
Cyathea 203.47 239.62 9.63 31.72 -171.75
Danaea 35.45 128.24 17.29 203.20 0
Davallia 37.12 72.57 26.26 53.00 0
Dennstaedtia 161.69 190.77 11.85 74.75 -86.94
Deparia 5.36 47.31 11.64 111.17 0
Dicksonia 192.10 215.19 2.95 122.86 -69.24
Dicranopteris 81.37 97.97 <14.24 54.56 -26.81
Diplazium 34.81 134.93 33.27 95.29 0
Drynaria 6.26 25.78 25.3 86.72 0
Dryopteris 164.55 180.63 33.49 97.75 -66.8
Elaphoglossum 15.51 88.12 35.72 72.27 0
Equisetum 324.72 359.39 50.87 465.97 0
Gleichenia 174.42 182.67 <89.30 107.47 -66.95
Grammitis 15.32 39.13 15.89 48.10 0
Hemionitis 57.77 123.27 13.35 45.17 -12.6
Histiopteris 28.79 46.81 <20.76 55.51 0
Hymenophyllum 85.95 106.37 35.92 255.40 0
Hypolepis 16.15 33.99 <36.87 77.27 0
Lastreopsis 2.64 54.64 11.6 73.53 0
Lepisorus 28.70 52.09 28.61 51.66 0
Leptolepia 0.06 36.88 11.24 104.22 0
Leptopteris 28.78 47.36 2.44 191.72 0
Lindsaea 34.07 50.81 32.28 79.27 0
Lophosoria 112.7 222.62 <112.00 122.86 0
Lygodium 81.89 144.86 23.31 287.39 0
Marattia 214.15 242.05 <172.00 198.04 -16.11

99
Marsilea 178.21 210.06 18.17 125.22 -52.99
Matonia 179.88 242.22 <49.83 149.11 -30.77
Microsorum 37.70 57.46 53.52 77.38 0
Nephrolepis 5.39 17.68 32.40 155.38 14.72
Oleandra 119.57 152.44 5.63 129.78 0
Onoclea 94.05 141.68 55.8 107.94 0
Ophioglossum 16.41 36.39 41.38 159.10 4.99
Osmunda 203.94 236.93 201.60 243.87 0
Osmundastrum 40.67 133.15 1.12 270.38 0
Paesia 34.17 50.31 <31.78 71.75 0
Pellaea 1.81 40.94 25.24 62.37 0
Pilularia 66.22 123.21 2.39 101.25 0
Pleopeltis 16.11 91.51 24.37 88.90 0
Polypodium 34.18 72.69 20.41 75.18 0
Polystichum 65.63 108.17 35.48 99.00 0
Psilotum 16.09 87.40 <23.40 91.76 0
Pteridium 33.98 40.91 3.91 84.38 0
Pteris 95.61 104.76 61.35 123.89 0
Pyrrosia 3.64 9.20 35.86 84.72 26.66
Regnellidium 83.83 142.19 <83.50 101.25 0
Rumohra 34.46 66.21 <17.28 59.41 0
Saccoloma 94.76 114.88 5.77 262.00 0
Salpichlaena 74.39 84.41 4.09 68.22 -6.17
Salvinia 69.34 98.97 29.85 130.34 0
Schizaea 56.05 75.90 15.59 133.42 0
Sticherus 66.10 117.16 37.80 129.08 0
Tectaria 123.91 151.61 31.31 89.61 -34.3
Thelypteris 66.42 85.21 60.94 110.68 0
Thyrsopteris 304.41 372.07 <123.85 160.72 -143.69
Tmesipteris 28.57 51.13 <23.40 91.76 0
Todea 172.31 235.79 <136.00 191.72 0
Trichomanes 66.77 116.20 73.34 185.83 0
Woodwardia 97.58 145.22 29.81 84.97 -12.61

Origination and extinction times as estimated from

fossil record with PyRate compared with molecular
estimates (min=upper 95% CI estimate of the crown
group, max=lower 95% CI estimate of the stem
group) and their MDI incogruence metric.

100
Supplementary Table 4. Baseline origination and extinction
rates (λ0 and µ0) and correlation parameters.
Linear model Exponential model
Parameter Mean 95% HPD Mean 95% HPD
Baseline rates λ0 0.019 0.001 0.043 0.103 0.005 0.276
µ0 0.015 0.002 0.032 0.014 0 0.053
Correlation G_0 -1.759 -5.264 0.714 -2.414 -4.35 0.041
parameters G_1 -0.057 -1.62 1.299 0.012 -1.008 1.002
(origination) G_2 -0.609 -2.856 1.05 0.143 -0.467 0.891
G_3 0.919 -0.789 3.816 -0.084 -1.102 0.825
G_4 -0.986 -4.603 0.982 -0.41 -1.648 0.47
G_5 2.005 -1.1 8.799 1.822 -0.053 3.635
G_6 -0.653 -3.428 0.931 0.549 -0.575 2.237
G_7 0.299 -1.001 2.055 -0.646 -1.969 0.364
G_8 1.468 -0.919 5.006 0.248 -0.548 1.384
G_9 0.656 -0.992 3.029 0.068 -0.896 1.149
G_10 -0.048 -1.829 1.804 -0.568 -1.843 0.39
G_11 0.257 -1.59 2.54 0.007 -1.334 1.359
G_12 -0.201 -1.803 1.102 -0.814 -2.663 0.458
G_13 -0.202 -1.969 1.146 0.28 -0.771 1.722
G_14 3.855 -1.061 15.837 -0.398 -1.605 0.523
G_15 1.583 -1.038 7.377 -0.141 -1.69 1.032
G_16 0.419 -0.708 2.203 0.809 -0.383 2.392
Correlation H_0 1.29 -0.91 5.709 1.174 -0.531 3.661
parameters H_1 -0.536 -2.307 0.6 -0.379 -2.111 0.779
(extinction) H_2 -0.038 -1.22 1.025 -1.088 -2.391 0.133
H_3 -0.399 -2.443 1.062 -0.025 -1.145 1.055
H_4 -0.221 -1.769 1.145 -1.365 -3.253 0.265
H_5 0.5 -1.363 2.936 1.535 -0.268 3.474
H_6 -4.193 -10.495 -0.592 1.586 -0.228 3.283
H_7 -1.02 -3.506 0.628 0.282 -0.775 1.738
H_8 2.365 -0.535 6.292 0.216 -0.984 1.779
H_9 2.15 -0.226 6.422 0.605 -0.624 2.305
H_10 -1.234 -4.776 0.588 -1.004 -2.572 0.187
H_11 0.458 -0.609 2.179 2.739 0.007 4.859
H_12 0.316 -1.4 2.468 -3.307 -5.895 -0.814
H_13 -1.407 -3.834 0.255 0.194 -1.286 2.005
H_14 -0.682 -2.533 0.775 0.822 -0.528 2.95

101
 H_15 3.123 -0.302 9.244 -0.702 -2.818 0.617
H_16 0.472 -0.937 2.29 -0.304 -1.898 0.917

The curves were numbered as follows: 0 fern diversity; 1 selenium

concentration in marine sediments; 2 wet tropical biome extent; 3
cool temperate biome extent; 4 magmatic activity; 5 athmospheric
CO2; 6 gymnosperm diversity; 7 angiosperm diversity; 8 warm
temperate biome extent; 9 polar biome extent; 10 continental
fragmentation; 11 arid biome extent; 12 athmospheric O2; 13
eustatic sea level; 14 diversity of lycophytes etc. (free sporing
vascular plants excluding ferns); 15 global mean temperature; 16
mountain area.

Supplementary Table 3. Shrinkage weights, based on local and

global shrinkage parameters, and global shrinkage.
Linear model Exponential model
Parameter Mean 95% HPD Mean 95% HPD
Shrinkage weights w(G_0) 0.618 0.035 1 0.762 0.299 1
(origination) w(G_1) 0.289 0 0.923 0.269 0 0.864
w(G_2) 0.378 0 0.957 0.252 0 0.842
w(G_3) 0.452 0 0.971 0.28 0 0.881
w(G_4) 0.412 0 0.976 0.332 0 0.899
w(G_5) 0.515 0 0.992 0.68 0.175 1
w(G_6) 0.382 0 0.962 0.375 0 0.938
w(G_7) 0.295 0 0.93 0.416 0 0.931
w(G_8) 0.57 0.026 1 0.297 0 0.877
w(G_9) 0.4 0 0.959 0.286 0 0.879
w(G_10) 0.306 0 0.934 0.392 0 0.927
w(G_11) 0.33 0 0.95 0.31 0 0.898
w(G_12) 0.302 0 0.927 0.454 0 0.952
w(G_13) 0.309 0 0.931 0.319 0 0.897
w(G_14) 0.607 0.008 1 0.326 0 0.91
w(G_15) 0.456 0 0.988 0.313 0 0.905
w(G_16) 0.326 0 0.942 0.446 0 0.95
Shrinkage weights w(H_0) 0.456 0 0.983 0.518 0 0.972
(extinction) w(H_1) 0.353 0 0.946 0.351 0 0.923
w(H_2) 0.282 0 0.91 0.537 0 0.961
w(H_3) 0.348 0 0.946 0.284 0 0.875
w(H_4) 0.319 0 0.927 0.577 0 0.973
w(H_5) 0.375 0 0.958 0.611 0.033 1
w(H_6) 0.834 0.427 1 0.633 0.039 1

102
 w(H_7) 0.477 0 0.97 0.315 0 0.903
w(H_8) 0.69 0.102 1 0.322 0 0.905
w(H_9) 0.636 0.064 1 0.395 0 0.944
w(H_10) 0.488 0 0.981 0.511 0 0.961
w(H_11) 0.336 0 0.938 0.782 0.25 1
w(H_12) 0.364 0 0.943 0.84 0.487 1
w(H_13) 0.563 0.044 1 0.337 0 0.921
w(H_14) 0.426 0 0.954 0.448 0 0.959
w(H_15) 0.694 0.053 1 0.426 0 0.947
w(H_16) 0.352 0 0.947 0.346 0 0.92
Global shrinkage τ 0.999 0.138 2.48 0.839 0.306 1.491

Shrinkage weights greater than 0.5 (highlighted in bold) indicate

significant evidence for correlation (positive or negative depending on
the respective G or H value). The curves were numbered as follows: 0
fern diversity; 1 selenium concentration in marine sediments; 2 extent
of wet tropical biome; 3 extent of cool temperate biome; 4 magmatic
activity; 5 atmospheric CO2; 6 gymnosperm diversity; 7 angiosperm
diversity; 8 extent of warm temperate biome; 9 extent of polar biome;
10 continental fragmentation; 11 extent of arid biome; 12 atmospheric
O2; 13 eustatic sea level; 14 diversity of lycophytes etc. (free sporing
vascular plants excluding ferns); 15 global mean temperature; 16
mountain area.

103