ONLINE APPENDIX 2
Supplementary Text
I. Climatic Vegetation Maps (Figure 6):
To draw the climatic-vegetation maps presented in Figure 6, we used paleobotanical
evidence, as well as vegetation simulation models (see below). The close-canopy tropical
forest in the Southern Hemisphere is represented by light green color. The northern
vegetation belt in dark green color represents alternatively the boreotropical forest (Fig. 6a),
and its successors: the mixed-mesophytic forest (Fig. 6b,c) and the temperate forest (Fig. 6d).
The boreal forest belt in the northern regions of Eurasia and North America (Fig. 6c,d) is
represented by a darker shade of green. The background area is represented in yellow.
a) The limits of the boreotropical forest during the Early Tertiary were approximated
based on the location of paleontological sites with boreotropical elements, following Wolfe
(1975) and Tiffney (1985a,b). This extension is in accordance with the vegetation maps of
Parrish (1982) and Fine & Ree (2006), although see Morley (2007, 2003) for a reduced
distribution of the megathermal close canopy forest. Dashed lines show uncertainties in the
distribution of the boreotropical forest.
b) The extension of the mixed-mesophytic forest during the Oligocene–Late Miocene
follows Wolfe (1975), Tiffney (1985a,b), Wen & Ickert-Bond (2009), and Beerling &
Woodward (2001). It extends in the south into North Africa (Wen & Ickert-Bond, 2009) and
South East Asia (Tiffney 1985a,b; Wen 1999; Wolfe 1975; Xiang et al. 2000). The
palaeoflora of Egypt during the Eocene/Oligocene was similar to the floras of Europe
(Tiffney 1985b), and this was also the case for the mesothermal sclerophyllous vegetation in
Taiwan (Wolfe 1975). During the Early Tertiary continental South East Asia (South China,
Indochina, northwest Thailand, and West Malaysia) was covered by the same boreotropical
1
flora as Japan and China (Tiffney 1985b). The northern limit of the mixed-mesophytic forest
is justified by the presence of paleontological sites in northern America at 45 degrees of
actual latitude (Wolfe 1975).
c) The extension of the temperate and boreal forest during the Pliocene is based on the
climatic simulations of Salzmann et al. (2008), as well as on available paleobotanical
evidence (Schneck et al. 2012; Tiffney 1985a,b; Wen 1999; Willis and McElwain 2002;
Wolfe 1971, 1975). Information about the vegetation of Australia across time periods comes
from Morley (2003), Kemp (1978), and Frakes & Kemp (1972).
d) Position of the southern refuges for temperate forest taxa during the Last Glacial
Maximum comes from Fine & Ree (2006) and Woillez et al. (2011). Milne (2006), however,
excluded Central America and the eastern highlands of Mexico as potential refuges for the
distribution of the temperate forest.
II. Dating analysis
To estimate divergence times we used two calibration points, the Late Eocene fossil
H. antiquum (Meseguer and Sanmartín 2012) to constrain the crown node of Hypericum
(=Hypericeae; see Material and Methods for a discussion on the phylogenetic position of the
fossil in our phylogeny), using a lognormal prior, offset=33.9 Myr (the uppermost limit of the
Priabonian time interval as a minimum hard bound), and a standard deviation “SD” of 0.7
that includes the entire geological interval (33.9–37.2 Myr; after Walker and Geissman,
2009). We used a secondary calibration point for the root of the tree (crown-node family
Hypericaceae) from the calibrated clusioid clade phylogeny of Ruhfel (2011). This matrix
contains different plastid (matK, ndhF, and rbcL) and mitochondrial (matR) markers and
represents 71 of the 94 currently recognized clusioid genera. In this work, Ruhfel (2011) used
2
two fossil calibrations: the Upper Cretaceous macrofossil Palaeoclusia chevalieri and the
Eocene pollen Pachydermites diederexii and conducted two independent analyses with
different positions for Paleoclusia: 1/ as the stem age of the clusioid clade (MRCA of
Ochnaceae s.l. “O” and the clusioid clade “C”), thereafter named as the “OC” position, and 2/
as the stem node of the Clusiaceae family (MRCA of Bonnetiaceae “B” and Clusiacae s.s.
“C”), named as the “BC” position. They obtained a crown age for Hypericaceae between 58.9
and 71.5 Ma (OC and BC, respectively) depending on the position of Palaeoclusia. We
integrate the uncertainty on the position of Paleoclusia in our analysis by assigning a normal
prior to the crown age of Hypericaceae, with mean 65.2 Ma (the mean of the BC and OC
ages) and a SD of 11 to span the entire confidence interval (47.9–86.4 Ma) obtained by
Ruhfel (2011).
Results of our analysis are shown in Figure S1 in online Appendix 3. In general,
confidence intervals are small, except for some early divergent lineages, such as the root
node or the split of tribe Vismieae–Hypericeae. We estimated a crown age for Hypericaceae
of 53.8 Ma, confidence interval (CI 43.8 – 66.5 Ma), which is in agreement with the results
obtained in previous studies (Ruhfel 2011; 47.9–(58.9)–71.5 for the OC and 56.0–(71.5)–86.4
for the BC placement). Hypericum crown-group is dated 34.9 Ma (CI 34 – 37 Ma) in
agreement with the fossil record of the group (Meseguer and Sanmartín 2012).
III. Biogeographic Analysis
3.1 Delimitation of Biogeographic Regions
We split the world into seven operational areas. Delimitation was based on patterns of
endemism within Hypericum, i.e., shared distribution patterns among species, but also on the
palaeogeographic history of the continents with the aim to maximize congruence with other
3
continental-scale biogeographic studies (Buerki et al. 2011; Donoghue and Smith 2004;
Sanmartín et al. 2001). Several areas coincided with the floristic regions of Takhtajan (1986),
which were based on patterns of vegetation endemism, and therefore likely reflect
geographical and eco-physiographical (climatic) features.
Western Palaearctic region (“WP”): Eurasia west of the Ural Mountains (former Turgai
Sea). Eastern Palaearctic (“EP”): non-tropical Asia east of the Ural Mountains.
Irano-Turanian-Himalayan (“ITH”): the region formed by the collision of the Indian
and Arabian plates with Eurasia, extending from the Zagros Mountains and the Caucasus in
the west through the Iranian plateau, the Hindu Kush, Tien Shan and Kunlun Shan Mountains
to the Altay Mountains in the northeast, and the Tibetan Plateau and the Himalayan Mountain
range in the southeast
Nearctic region (“NE”): North America and Central America down to the Mexican
lowlands (i.e., excluding tropical Mexican distributions); the Caribbean region was included
here because there are very few endemics and most widespread species are shared with
eastern North America.
Oceania (“OC”): comprising the tropical regions of southwest and southern China and
eastern parts of Bangladesh, Burma, Thailand, Indochina, the Malay Peninsula and South
East islands of Borneo, Sumatra, Java, and the Inner Banda Arc, the Polynesian region, and
Hawaii, as well as the Indian subcontinent (all Indostan excluding the tropical foothills of
Himalayas in the north) and the former Gondwanan landmasses of Australia, New Guinea,
New Caledonia, and New Zealand.
Africa (“AF”): excluding the region north of the Saharan belt, which is considered as
part of the Western Palearctic region.
Neotropics (“NT”): South America and tropical Central America up to central Mexico.
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3.2. Biogeographic Models
We set up four different analyses (M1 to M4) in order of increasing complexity to
analyse the influence of fossils and ENM reconstructions (past and present environmental
tolerances) in biogeographic inference.
a) Model M1 (“Unconstrained”, Fig. S4 Appendix 3): We used only present data from
current distributions.
b) Model M2 (“Stratified”, Fig. S5): We used present occurrences but also
incorporated a palaeogeographic model to reflect changes in continental connectivity over
time. The evolution of Hypericum during the last 55 million years has been accompanied by
dramatic changes in continental configuration: tectonic plates moved apart and collided with
each other (e.g. Africa and Eurasia), but also the sea level changed producing the emergence
and sinking of land corridors (e.g. Beringia), whereas climatic oscillations created ecological
barriers to dispersal, e.g. the ice sheet covering part of the Northern Hemisphere in the
Pleistocene. These changes probably had an effect on the probability of Hypericum lineages
to disperse between areas over time.
To incorporate this information into our analysis, we stratified the phylogeny into four
time slices (TS), with boundaries selected to reflect major tectonic or climatic events: TSI
(65 to 35 Ma): this period is characterized by warm climates at northern latitudes and the
geographical proximity of northern landmasses, allowing the spread of a continuous forest
belt, the boreotropical forest, across the Holarctic (Wolfe 1975). It ends with the Terminal
Eocene Event (TEE). TSII (35 to 10 Ma): this period is marked by the major Antarctic
glaciations and the Terminal Eocene Event (TEE; Zachos et al. 2008), a global climatic
deterioration that produced a diversity decline at the beginning of the Oligocene (Morley
5
2007). During this period the boreotropical forest was replaced by the mixed-mesophytic
forest, a mix of angiosperm deciduous and conifer vegetation (Tiffney 1985a,b; Wolfe 1975)
that persisted until the Late Miocene (Wen 1999; Xiang et al. 2000). TSIII (10 to 3.5 Ma)
begins with the Late Miocene cooling event (LCM, ca. 10-8 Ma), a period when important
changes in vegetation occurred (Schneck 2012): a boreal forest expanded across the northern
regions of Eurasia and North America across Beringia, whereas the mixed-mesophytic
elements retreated to temperate regions in the south. TSIV (3.5-0 Ma) spans the period from
the opening of the Bering Strait (3.5 Ma), which interrupted biotic connections between
Palearctic and Nearctic regions for temperate elements to the glacial cycles that characterized
the end of the Pliocene and the Pleistocene period.
Dispersal rates in the biogeographic model (Q transition matrix; Buerki et al. 2011)
were scaled to reflect the changing continental configuration for each TS as follows: a)
dispersal rates between areas sharing an edge (adjacent plates) or connected by a land
corridor were not downscaled, i.e., assigned a scalar of 1, to reflect the facility of lineages to
migrate across this type of connection; for example, the NALB between the Western
Palearctic and North America in the Early Eocene. b) Dispersal rates between neighboring
areas not connected by a land bridge, such as the Western and Eastern Palearctic landmasses
before the closing of the Turgai Strait, were downscaled by a factor of 0.5 to reflect the
reduced probability of movement. c) Dispersal between non-neighboring areas or areas
separated by large ocean barriers was downscaled by a factor of 0.1; this also applies when
dispersal implies an intermediate area, for example, the possibility of dispersal between the
Eastern Palearctic and Africa through the Indian Plate in the Eocene. Besides
paleogeographic barriers, other abiotic factors such as marine or wind currents (e.g., the
Antarctic Cicum-Polar Current), or climatic “barriers” such as ice sheets, were considered in
6
modelling dispersal rates over time. Below follows a more detailed description of the
dispersal scalars used in our paleostratigraphic model:
Dispersal between Nearctic — Western Palearctic (NE — WP): During the Paleogene
(55-35 Ma), North America and Europe became connected by the North Atlantic Land
Bridge (NLAB). The first NLAB, the Thulean Bridge, connected the British Isles with the
Hudson Bay in North America and allowed movement of warm-temperate boreotropical
elements between 55 and 50 Ma, when it foundered. A later bridge, the De Geer Bridge,
lasted until ca. 35 Ma, but, because of its northern position, was more appropriate for coldtemperate plants (Sanmartín et al. 2001; Tiffney 1985ab). Our model reflects this connection
as a dispersal probability of 1 for TSI, which decreases to 0.5 after the break up of the NLAB
connection in the Late Eocene (TSII, 35 Ma). Finally, a low probability of 0.1 in the last two
time slices, TSIII and TSIV, reflects the widening of the North Atlantic Ocean and increasing
geographical distance between Nearctic and Palearctic regions after 20 Ma.
Nearctic — Eastern Palearctic (NE — EP): Land dispersal between North America
and eastern Asia was possible across the Beringian Bridge throughout the Early-Mid Tertiary
(Donoghue et al. 2001; Sanmartín et al. 2001; Tiffney 1985ab; Wen 1999), as reflected in our
model (dispersal probability of 1 in TSI and TSII). As the climate got increasingly cooler
from the Late Miocene onwards, the Beringian Bridge was covered by a boreal taiga forest
that extended across northern Eurasia and northern North America (Sanmartín et al. 2001).
This might have reduced the probability of dispersal between these two regions for temperate
taxa (scaling factor 0.5 in TSIII). Finally, the opening of the Bering Strait 3.5 Ma broke all
final connections between the Nearctic and Palearctic regions for boreo-temperate elements
(TSIV, scaling factor 0.1), although tundra-arctic elements could still migrate across Beringia
during the Pleistocene glaciations (Sanmartín et al. 2001).
7
Nearctic — Neotropic (NE — NT) were intermittently connected during the Tertiary,
first by the Late Eocene Proto-Greater Antilles land bridge, and later by GAARlandia
(Greater-Antilles Aves-Ridge) during the Eocene-Oligocene (33-35 Ma) boundary (Briggs
1994). This is reflected in our palaeostratigraphic model by a dispersal-scaling factor of 0.5
in TSI and TSII. This land bridge sank and the connection was interrupted or reduced until
the Late Miocene (ca. 10 Ma), when new mountain uplift in Central America and South
America created a new migration route: the Late Miocene Central America Land Bridge and
northern Andes orogeny (Antonelli et al. 2009). Finally, after the uplift of the Panama
Isthmus (3.5 Ma), land dispersal has been possible across the New World (scaling factor of 1
in TSIII and TSIV).
Western Palearctic — Eastern Palearctic (WP — EP): These two regions were
separated during the Early Tertiary by the Turgai Strait, although connections along the
coasts of the Tethys Seaway were possible (Sanmartín et al. 2001; Tiffney 1985b). Thus,
dispersal between these two regions was scaled by a factor of 0.5 in TSI. The Turgai Strait
closed at the Eocene-Oligocene boundary (30 Ma) with the formation of the Ural Mountains,
and from this moment on direct land dispersal was allowed in our biogeographic model
(scaling factor of 1 from TSII to TSIV).
Irano-Turanian-Himalayan (ITH) to all other regions. This region is formed by the
newly uplifted landmasses and mountain ranges that resulted from the successive collision
against Eurasia of the Indian plate in the Eocene and the Arabian plate in the Middle Miocene
(Wang et al. 2009). Before the Tertiary, this area formed the eastern arm of the Tethys
Seaway, and therefore dispersal to this region by the other landmasses was disallowed until
TSIII, i.e., by assigning it a probability of 0). Dispersal between the Irano-TuranianHimalayan region and Eurasia (ITH — WP+EP) might have been possible after the onset of
8
the Equatorial Ocean Currents in the Late Tertiary, involving dispersal along the coasts of
India, which forms part of area E (Sanmartín and Ronquist 2004). This is reflected in our
model by a dispersal rate of 0.5 in TSIII and TSIV.
Africa — Eurasia (AF — WP+EP): At the beginning of the Paleocene, 60 Ma, the
north-eastward drift of the African plate brought this continent into contact with the western
half of Eurasia, the Iberian Peninsula. The Gibraltar Strait was intermittently opened and
closed during the Tertiary (Meulenkampt & Sissingh 2003; Sanmartín 2003), but the African
and Western Palearctic plates have always been geographically close and this proximity is
reflected in our model by allowing dispersal between these two areas across all time slices
(scale factor of 1 from TSI to TSIV). In contrast, direct land dispersal between Africa (AF)
and the Eastern Palearctic (EP) only became possible in the Mid Miocene (16 Ma, TSII),
when the Arabian plate collided with Eurasia along the Anatolian fault zone, closing the
eastern arm of the Tethys Sea (Sanmartín 2003; although alternative reconstructions suggest
an earlier collision, 30 Ma, Allen & Amstrong 2008). The Arabian bridge acted as a dispersal
corridor between Africa and central-western Asia until the Late Miocene-Pliocene, when
progressive aridification of the African continent led to the formation of the Saharan (7 Ma)
and Arabian Deserts (3.5 Ma). These wilder lands probably constituted a barrier to dispersal,
reflected in our model by downscaling dispersal to 0.5 in TSIII and IV.
Neotropics — Oceania (NT — OC): South America and Australia were connected as
part of Eastern Gondwana until the opening of the Drake Passage between South America
and Antarctica in the Early Oligocene (32 Ma). Therefore, a scaling factor of 1 was assigned
to TSI to reflect the possibility of land dispersal. The opening of the Drake Passage isolated
these regions by opening new ocean barriers and initiated the Antarctic Circum-Polar
Current, which brought about the first Antarctic glaciation (Morley 2003; Sanmartín 2005).
9
This is reflected in our model by a low probability of dispersal from TSII to TSIV (scalar
0.1).
Eastern Palearctic — Oceania (EP — OC): During the Early Tertiary, these
landmasses were separated by the Tethyan Gulf (scalar 0.1 in TSI). Following its separation
from Antarctica in the Late Eocene-Early Oligocene, Australia began to drift rapidly towards
Asia. The collision of the Australian Plate with the Pacific and Eurasian Plates resulted in the
formation of the South West Pacific and South East Asian archipelagos (Metcalfe 1998),
which might have acted as a route of dispersal. This is reflected in our model by allowing
dispersal between areas B and E, albeit with a low rate (0.5) in TSII. Dispersal, however, was
freely allowed in TSIII and TSIV, after the final collision of the Australian plate with
Eurasia.
c) Model M3 (“Fossil-stratified”, Fig. S6): We used present occurrences but also
integrated information from fossil ranges and paleogeographic scenarios into the
reconstruction. In particular, the crown node of Hypericum was constrained to include the
Eastern Palearctic region (EP) as part of the ancestral range of this node. This is the area of
appearance of the fossil, H. antiquum, which was used to calibrate the age of the crown-node
in the phylogenetic dating analysis. In our approach, fossil constraints placed on nodes are
encoded as ancestral presence in the area containing the geographic locality of the fossil.
Lagrange then calculates the likelihood of the data conditional on that area being included in
the ancestral range at the node in question. The paleogeographic model in model M2 was also
used to constrain area connectivity through time.
d) Model M4 (“Integrative”, Fig. S7): We integrated fossil constraints and fossil-based
ENM predictions (Fig. 2) in the biogeographic analysis in two ways. First, by running
Lagrange with the crown node of Hypericum constrained to include the Eastern Palearctic
10
and Nearctic regions; these regions were predicted as potential distribution areas for
Hypericum lineages during the Eocene-Oligocene by the Eocene 560 ppm projections (Fig.
2). Second, we modified the paleogeographic dispersal matrix in Model 2 to reflect the
existence of climatic/ecological barriers or transient dispersal corridors that connected
regions, e.g., areas that are now outside the climatic envelope of the organism but were in the
past within its ecological tolerance. Therefore, this new model reflects both the geographical
and “ecological” connectivity between regions in Hypericum (see below). Dispersal rate
scalars as defined above were increased or decreased in accordance to the adequacy of the
world at different time periods to the present and past tolerances of Hypericum species (Fig
2). Areas were considered “ecologically connected” when there were regions/pixels within
the ecological tolerance of the group in a given time period, i.e., with Mahalanobis distances
lower than ten. Specifically:
TSII: The dispersal rate scalar between Africa — Eastern Palearctic (AF — EP) was
increased to 0.5 because ENM models favored this Miocene dispersal corridor for Hypericum
(Fig 2c).
TSIII: The dispersal rate scalar between Eastern Palearctic — Nearctic (EP — NE)
was increased to 0.7. Several studies suggest that during the Late Miocene-Pliocene the
Beringian corridor was only favorable for cold-adapted (boreal) taxa (Sanmartín et al. 2001),
but our ENM models suggest that it was still within the environmental tolerance of
Hypericum at least until the Pliocene (Fig. 2e). On the other hand, despite continuous
geographic connection between the Western Palearctic – Africa (AF – WP) after the
Paleogene, the dispersal rate between these two areas was downscaled to 0.7 in this time
slice, because our ENM models suggest that the aridification trend that affected North Africa
11
from the Late Miocene onwards (7-5 Ma) probably created an ecological barrier for the
dispersal of Hypericum lineages (Fig. 2d)
TSIV: Again, the ENM models predict a decrease in connectivity between Africa and
Western Palearctic in the Pliocene (Fig. 2e), related to the aridification episode that gave rise
to the Saharan and Arabian deserts, so dispersal rates between these two regions was
downscaled to 0.5 in this time slice.
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