Vol. 6/1,2, pp. 520 Urban & Fischer Verlag, 2003 http://www.urbanfischer.

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Perspectives in Plant Ecology, Evolution and Systematics

Interplate dispersal paths for megathermal angiosperms

Robert J. Morley
School of Geography and Environmental Science, Monash University, Victoria, Australia Department of Geology, Royal Holloway University, Egham, Surrey, UK
Received: 29 January 2003 Revised version accepted: 29 March 2003

Abstract
The dispersal of megathermal angiosperms between tectonic plates is reviewed on the basis of fossil evidence for the Cretaceous and Tertiary periods, since the radiation of the angiosperms, and the period of break-up of Gondwana. The combination of tectonic plate disassembly and redistribution, coupled with phases of global warming followed by pronounced cooling, has resulted in the formation of intermittent dispersal opportunities for frost-intolerant plants, and has been a major factor in determining the direction of angiosperm diversification. The Early Cretaceous radiation of angiosperms seems to show little relationship to the formation of Tethys. However, for the Late Cretaceous and Tertiary nine relevant dispersal routes can be differentiated that can be divided into two distinct categories: routes which formed following the break-up of Gondwana during the Late Cretaceous and Earlier Tertiary, when warm climates encouraged dispersal of megathermal elements globally, and routes which formed since the Middle Eocene, following phases of plate collision, as global climates were cooling down, inhibiting such dispersal. Most inter-plate dispersal of megathermal angiosperms took place in the Late Cretaceous and Early Tertiary at a time when global climates were markedly different from those of today, and the global area of megathermal vegetation several times greater than at present. Under such a scenario, it is likely than opportunities for speciation were much higher than for present-day megathermal plants. Key words: angiosperms, dispersal routes, megathermal, plate tectonics

Introduction
Angiosperms underwent their major phase of radiation and dispersal in unison with the break-up of Gondwana. At the same time, the global climate changed dramatically from an essentially greenhouse world during the Later Mesozoic and Earlier Tertiary, to the icehouse world of the Quaternary. The combination of tectonic plate disassembly and redistribution, coupled with phases of global warming followed

by pronounced cooling, resulted in the formation of intermittent dispersal opportunities for frost-intolerant (megathermal) angiosperms at different times, both within the low and mid latitudes and between low and mid latitudes, and has been a major factor in determining the direction of angiosperm diversification. Plate tectonic controls and patterns of global climate change also account for many of the present-day disjunctions seen in various groups of tropical flowering plants (Morley 2000).

Corresponding author: Robert J. Morley, School of Geography and Environmental Science, Monash University, Victoria 3800, Australia; and Dept of Geology, Royal Holloway University, Egham, Surrey TW20 0EX, UK. Mailing address: Palynova/PT Eksindo Pratama, Vila Indah Pajajaran, Jl Kertarajasa No 12A, Bogor, Indonesia 16153; e-mail: pollenpower@indo.net.id

1433-8319/03/6/01-02-005 $ 15.00/0

R. J. Morley

The process of inter-plate plant dispersal remains poorly understood; for instance, vicariance biogeographers who view the time of separation of African and S American plates at about 100 Ma as the time of cessation of plant dispersal between these regions may be making an unwarranted generalisation since pollen data from the two sides of the South Atlantic show that many dispersals continued well beyond that time. Reason suggests that dispersals are likely to continue between two tectonic plates well after the time of separation via wind, water and avian vectors. Also, the details of land connections during times of initial plate separation are equally poorly appreciated, and island chains associated with mantle plume hotspots between diverging plates are likely to have provided opportunities for filter dispersal well after the time of plate separation. In this paper shortcomings in our understanding of both dispersal processes and the nature of detailed palaeogeographies during times of plate separation or convergence are circumvented through the examination of the pattern of appearance of fossil (principally angiosperm) pollen on adjacent plates in addition to data on plant macrofossils and faunal migrations. The appearance of the same pollen types, or succession of types, on adjacent plates provides strong evidence for

interplate dispersal, and allows judgement to be made as to the presence of a dispersal corridor or enabling filter. Such judgements can be made whether or not the parent taxa of the pollen types are known. Emphasis has been placed on the fossil record of families considered megathermal, or megathermal pro majore parte by van Steenis (1962), but to clarify some dispersal routes, and to emphasise climatic barriers to dispersal of megathermal taxa during the Late Neogene and Quaternary, records of some mesothermal and microthermal taxa are also discussed. In several instances, dispersal paths are identified using the record of pollen types for which the parent plants are unknown; in such cases affinity with megathermal families cannot be demonstrated, it may be inferred on the basis of palaeobiogeography. The classification used here follows Takhtajan (1969).

Angiosperm origins and initial radiation

Only a few years ago, the relationship between angiosperms and other seed plants seemed more or less clear-cut based on the Anthophyte theory, with angiosperms being a sister group to Gnetales and extinct groups, such as the Bennettitales, and presumably ini-

Fig. 1. Diagram showing the palaeolatitudinal relationship of first occurrences of angiosperm pollen. Open circles and lighter shading, monosulcate grains, reflecting a grade of early dicotyledons and monocotyledons; closed circles and darker shading, triaperturate grains, reflecting eudicots. Geographic areas: (1) New Zealand, (2) Australia, (3) Patagonia, (4) Congo, (5) Brazil, (6) Peru, (7) Ivory Coast, (8) Israel, (9) Gabon, (10) Portugal, (11) Oklahoma, (12) Potomac Group, (13) Maryland (Virginia and S England), (14) SW Siberia, (15) not known, (16) Denver Basin, (17) E England, (18) Saskatchewan, (19) E USA, (20) Wyoming, (21) W Central Siberia, (22) SE Alberta and SW Manitoba, (23) SE Siberia, (24) Central Canada, (25) Central Alberta, (26) Greenland, (27) Ellesmere Island and NE Siberia, (28) Yukon, (29) Kuk River (Alaska), and (30) Umiat Region (Alaska) (after Hickey & Doyle 1977).

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Interplate dispersal paths for megathermal angiosperms

tially appearing sometime in the Triassic, some 230 Ma years ago (Crane et al. 1995). Recent molecular work contests this view, suggesting that although unambiguously supported hypotheses of phylogenetic relationships among seed plants have not yet been obtained, angiosperms form a sister clade to all gymnosperms (Bowe et al. 2000; Chaw et al. 2000; Madallon & Sanderson 2002), and hence point to an even older origin, in the Mid Palaeozoic. The oldest definitive angiosperm fossil is from the earliest Cretaceous of China (Sun et al. 2000), but this does not give a clue to the place of origin, since the deposits in which it was found provide only one of the best settings for plant and animal fossil preservation from this period (Zhou et al. 2003); more localities would be needed before such judgements could be made. It is not until well into the Early Cretaceous, within the Hauterivian (135132 Ma) or Barremian (132 124 Ma) stages, that angiosperm fossils become sufficiently widespread to make judgements regarding patterns of radiation. The appearance of monosulcate tectate pollen referable to the pollen genus Clavatipollenites, identified with the magnoliid family Chloranthaceae (Muller 1981; Walker & Walker 1984) is critically important. From the palynological perspective, the absence of angiospermid fossils prior to the Barremian/Hauterivian is easily explained if we look at the wall structure of the oldest pollen grains confidently identified with angiosperms, and compare this with

the succession of pollen wall structures seen in extant primitive angiosperm families, such as Anonaceae, we find that the wall structures of the most primitive angiosperm pollen types would have been indistinguishable from those of many gymnosperms (Walker & Walker 1984), so the appearance of angiosperm pollen in the Early Cretaceous indicates the time of evolution of pollen exines with a relatively complex tectate wall structure. Until this time there is no clear way of determining whether pollen was produced by angiosperms or gymnosperms unless it is found in situ in an angiosperm flower, e.g. in the Magnoliid Lesomasites fossulatus (Ward et al. 1989). The global radiation of the first angiosperms that produced tectate pollen is illustrated by the fossil record of two contrasting pollen groups (Fig. 1). The record of tectate monosulcate pollen reflects radiation of monocots and Chloranthoid dicots, whereas the record of triaperturate pollen reflects the appearance and radiation of non-magnoliid dicots or eudicots (Crane et al. 1995). The oldest records for tectate monosulcate pollen appear at about the same time in widely separated localities, such as Gabon, Brazil and Israel, in Western Gondwana, and England, Portugal and eastern USA in Western Laurasia (Crane 1987). Their pattern of appearance shows no relationship to the separation of Laurasia and Gondwana by the Tethyan Ocean, which was already well established in the Mid-Jurassic, at about 160 Ma, or to climatic

Fig. 2. Early Cretaceous, Albian palaeogeography and climatic zones, according to Vakhrameev (1991), together with the approximate locations and ages (in My) of the appearance of triaperturate pollen shown in Fig. 1.

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zones, suggesting that neither Tethys nor humidity-related climatic belts formed a barrier to dispersal for these early angiosperms. At the time of the first angiosperm radiation terranes eventually to form S Turkey, the Balkans and Italy were arranged as a lineament between Africa and Europe (Scotese 2001), and this may have facilitated this initial dispersal phase. Angiosperms appear, however, to have arrived some 10 Ma later on the Australian Plate (Burger 1991; Dettmann 1994), where the greatest concentration of primitive angiosperms occurs today, and in India. We can therefore hardly visualise Australia as their area of origin, or their origin on a Gondwanan shard or terrane that separated from Australasia in the Mesozoic (as proposed by Tahktajan 1987) but their area of most persistent survival. The second wave of angiosperm radiation, of early eudicots, shown by the record of triaperturate and triaperturate-derived pollen, begins in the Aptian (113108 Ma) of Gabon and Brazil (Doyle et al. 1977), during which time low-latitude climates were markedly subhumid, and probably much hotter than present day equatorial climates. This group subsequently dispersed poleward during the Albian (10896 Ma) into zones of mesic subtropical (still megathermal) climate and into areas with more mesothermal climates poleward of 60 N and S in the Cenomanian (9692 Ma)

(Fig. 2). As in the case of the earlier monosulcatepollen-producing angiosperms, the initial eudicot wave seems to have been little affected by the oceanic barrier of Tethys possibly for similar reasons (see above). By the Cenomanian/Turonian (9690 Ma), angiosperms became dominant components of vegetation in most areas, with major centres of radiation across the northern mid latitudes (Boreotropical province), southern mid latitudes (Gondwanan megathermal province) and the equatorial region (Palmae province; Fig. 3). It is from this time onward that the angiosperm pollen record can be used readily to identify periods of inter-plate dispersal (Morley 2000). Dispersal routes fall into two distinct categories. The first category consists of routes which formed during and following phases of Gondwanan break-up, in the Late Cretaceous and Early Tertiary, culminating with the Late Paleocene/Early Eocene thermal maximum (6049 Ma), when warm climates encouraged dispersal of megathermal elements globally. The second category is of dispersal routes that formed between the Middle Eocene and present day when dispersals relate primarily to phases of plate collision, and during which time global climates have gone through a long phase of stepwise cooling which has inhibited the dispersal of megathermal plants.

Fig. 3. Late Cretaceous, Turonian plate tectonic reconstruction and palaeogeography according to Smith et al. (1994), showing the three latitudinal belts within which moist megathermal angiosperm taxa first evolved. Noteworthy dispersals of megathermal taxa and of Normapolles are indicated for the Turonian and Santonian/Coniacian, suggested by the palynological record (from Morley 2000, with modifications). Normapolles province boundary from Herngreen et al. (1996).

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Interplate dispersal paths for megathermal angiosperms

Late Cretaceous to Early Eocene, fragmentation of Gondwana during period of greenhouse climates
During the Late Cretaceous to Early Eocene (5449 Ma), the fragmentation of Gondwana was well under way, with both the South Atlantic and Indian Oceans widening rapidly (Figs. 3, 4 and 5). Six main interplate dispersal paths for megathermal angiosperms need consideration during this period: (1) a transatlantic path between Europe and N America, (2) a route from Europe to Africa, (3) a land bridge between N and S America, (4) a transatlantic path between Africa and S America, (5) routes between Africa and India, and (6) a land bridge between S America and E Gondwana (Antarctica/Australasia). During this time global climates were warm, with temperature maxima during the Turonian (9290 Ma) and Late Paleocene/Early Eocene. A land connection also existed across Beringia throughout this time, but was probably too far north (75) for dispersal of megathermal plant taxa, although it formed a route for several microthermal Boreotropical taxa, reviewed by Manchester (1999).

Dispersal route from N America to Europe

Many megathermal angiosperm families originated in the North American and Eurasian Boreotropical

province during the Late Cretaceous, when virtually frost-free climates extended well into the mid-latitudes. Eurasia and N America would have been in tectonic contact via Greenland through the Late Cretaceous and Early Tertiary, but except for very high latitudes, would have been separated by a narrow seaway until the Late Paleocene (6054 Ma) and Early Eocene when a land connection was established across S Greenland at latitudes of 4550 N (Tiffney 1985a; Parrish 1987). Prior to the formation of this land connection, the timing of possible dispersal routes is suggested from the simultaneous appearance of pollen types in both regions. Pollen of the Normapolles group (affinity obscure but derived in part from early Juglandales), which characterizes Late Cretaceous pollen floras from the eastern USA to the Turgai Straits (Fig. 3) shows rapid diversification at this time (Tschudy 1981), with most similarities between the two areas during the Cenomanian and Santonian/Coniacian (88-84 Ma), suggesting the presence of a land connection or dispersal filter. The parent plants of the Normapolles pollen group were not necessarily megathermal, but their prominence in areas such as the Mississippi Embayment, considered to bear tropical and paratropical vegetation at this time by Upchurch & Wolfe (1987), suggests that strictly megathermal Late Cretaceous taxa which may have been characterized by less distinctive pollen could have followed the same route.

Boreotropical.
Ilex Anacolosa Sapindaceae

Meliosma Platycarya

Tropical rain forests

Land areas
u Evaporite deposits

Symplocos Bombacaceae Aquilapollenites

Palmae Restionaceae Ctenolophon Aquilapollenites Anacolosa Proteaceae

u u

u u Nypoidae*
Ctenolophon*

Palmae, SE Asian sub-province

Palmae
Walvis Ridge Cunoniaceae Didymelaceae Mytraceae

Casuarina Anacolosa

Southern Megathermal.

Fig. 4. Early Tertiary, Paleocene, plate tectonic reconstruction and palaeogeography according to Smith et al. (1994), with occurrences of evaporites from Parrish et al. (1982) with additions, and closed-canopy megathermal rain forests according to Morley (2000). Noteworthy Maastrichtian and Earlier Paleocene dispersals of megathermal taxa (prior to the Late Paleocene/Early Eocene thermal maximum) are indicated, as suggested by the palynological and macrofossil record.

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The development of a short-lived land connection across S Greenland during the latest Paleocene to Early Eocene at about 4550 N at the precise time of the Late Paleocene/Early Eocene thermal maximum provided a clear route for the dispersal of megathermal elements between the two areas for this short time period. Megathermal and mesothermal taxa that would have made this crossing, summarized from macrofossil data by Manchester (1999) and pollen by Morley (2000) include Mastixia and Toricella (Cornaceae), Gordonia (Theaceae), Symplocos (Symplocaceae), Alangium (Alangiaceae), Tapiscia (Staphyleaceae), Bombacaceae and Sapotaceae (Fig. 5). There are also many examples of mammalian dispersals between Europe and N America at this time (Simpson 1946; McKenna 1973). The Late Paleocene/Early Eocene thermal maximum allowed megathermal angiosperms to extend their ranges further poleward than at any time during the period in which they were dominant, as emphasised by records of Nypa macrofossils the Early Eocene from the London Clay (41 N palaeolatitude) (Reid & Chandler 1933; Collinson 1983), and also Tasmania (58 S palaeolatitude; Pole & MacPhail 1996). The most remarkable aspect of global palaeogeography at this time was the presence of land bridges between Europe and N America and between S America and E Gondwana before the termination of the C American connection, allowing the possibility of

unhindered dispersals over remarkably wide areas. For instance, at this time members of Bombacaceae and the mesothermal Alangium were able to disperse between Europe and N America on the one hand, and to Australia on the other (Fig. 5).

Dispersal route from Europe to Africa

Opportunities for Late Cretaceous trans-Tethyan dispersal from Europe of Africa is indicated by the common occurrence of the Laurasian Normapolles group in North Africa (Herngreen et al. 1996), delimiting the southern boundary of the Normapolles province (Fig. 3). With the northward drift of Tethyan terranes in the earliest Tertiary this dispersal route was probably severed, and it was not until the Late Eocene that dispersals of megathermal taxa between Europe and Africa are recorded (Cavagnetto & Anadon 1995), with Mimosaceae, Amanoa, Alchornea, Caesalpinia, Crudia and Thespesia dispersing into the Iberian Peninsula, only to disappear following Mid Tertiary climate deterioration.

Dispersal route from N to S America

The tectonics of the Caribbean region are complex and controversial, but the plate-kinematic model of Pindall et al. (1988) appears to be standing the test of time. With the western drift of both the N and S American

Sapotaceae, Mastixia, Platycarya Symplocos Toricella, Tapiscia, Alangium

Boreotropical.
u u u uu u

Bombacaceae

u u u u uu u u

u u

u u

Proto-Indian
Durio, Beauprea Ctenolophon Gonystylus Iguanurinae Sapindaceae Restionaceae (in M. Eocene)

Tropical rain forests Land areas u Evaporite deposits

(from M. Eocene)

African Neotropical.
Bombacaceae Polygonaceae Restionaceae Sapindaceae
u u

Amanoa Crudia C. dorogensis

u

Walvis Ridge

Southern Megathermal.

Fig. 5. Early Tertiary, Early Eocene plate tectonic reconstruction and palaeogeography according to Smith et al. (1994), with occurrences of evaporites from Parrish et al. (1982) with additions, and closed-canopy megathermal rain forests according to Morley (2000). Noteworthy dispersals of megathermal plants relating the thermal maximum are indicated, as well as Middle Eocene dispersals into SE Asia, relating to the collision of the Indian and Asian Plates, as suggested by the palynological and macrofossil record.

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Interplate dispersal paths for megathermal angiosperms

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plates since the Mid Cretaceous, the intervening oceanic Caribbean Plate, lying directly to the west, developed island arcs along both its leading and trailing margins. During the Late Cretaceous, the leading arc formed an island arc, or land connection, between Yucatan and Colombia (Fig. 6), which with further drift eastward subsequently separated from Yucatan after the Middle Eocene (4939 Ma), resulting in the formation of a marine trough permitting circulation of oceanic waters from the Atlantic to the Pacific. The arc subsequently drifted yet further east, but maintained a connection with the S American Plate, and during the Eocene-Oligocene is thought to have formed a land mass, recently termed Gaarlandia (Iturralde-Vincent & MacPhee 1999), prior to fragmenting into the present-day Caribbean islands. Evidence for this dispersal route is forthcoming from vertebrate fossils (Bonaparte 1984; Rage 1988; Hallam 1994), with dispersals of hadrosaurian and ceratopian dinosaurs from N America to S America in the Campanian, and of various snakes, lizards and titanosaurid dinosaurs in the northward direction. Exchanges began in the Campanian and increased in the Maastrichtian and Paleocene. The pollen record also provides evidence for dispersals over this period. Late Cretaceous dispersals included the parent plant of Aquilapollenites pollen (thought to be derived from an extinct Loranthalean (Santalalean) group) from north to south, and Gunnera (Gunneraceae) in the reverse direction (Fig. 3),

whereas in the Paleocene, Bombacaceae and Symplocos (based on macrofossils) dispersed into S America from the north, and Ilex (Aquifoliaceae), Anacolosa (Olacaceae) and members of Sapindaceae dispersed into N America from the south (Fig. 4).

Dispersal route from S America to Africa

Direct land connections between the African and S American Plates were severed at the end of the Albian (96 Ma), fitting approximately with faunal evidence (Parrish 1987). Up until that time there was a clear dispersal corridor for megathermal plants between the two plates, but the frequency of plant dispersals across the newly formed ocean after this date as shown from fossil pollen suggests that some form of land connection, or series of islands acting as stepping stones, remained throughout the Late Cretaceous and into the earliest Tertiary, possibly in the region of the Walvis Ridge/Rio-Grande Rise and Sierra Leone Ridges. The Walvis Ridge, stretching from Angola to Brazil, consists of a seamount lineament (volcanic lineament formed on spreading oceanic crust above a stationary mantle plume) with individual mounts dating from 82 Ma adjacent to Angola to 37 Ma within the Mid Atlantic (MacDougal & Douglas 1988). For the Rio Grande Rise on the other hand (which is less well dated), Theide (1977) conducted subsidence estimations, and suggested that the rise may have been above sea level until the Oligocene. No such study has been

Fig. 6. Campanian reconstruction of Middle America (Pindall et al. 1988). The Greater Antilles and Aves Ridge, created by subduction of the Proto-Caribbean Sea by the Caribbean Plate, formed a land bridge between the Americas during the Late Cretaceous and Paleocene. The Caribbean Plate drifted eastward relative to the Americas in the Mid-Tertiary, breaking this connection during the Eocene.

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performed on the Walvis Ridge, but clearly, this seamount ridge is a strong contender for Late Cretaceous and Palaeogene transatlantic dispersals. During the Late Cretaceous, well after the time of separation, numerous major dispersals can be demonstrated from similarities of the pollen record in West Africa and northern S America. The most important pollen taxa appearing virtually simultaneously on either side of the S Atlantic, summarised on Figs. 3 and 4, are: Triorites africaensis (ancestral Proteaceae) in the Cenomanian; Monocolpopollenites sphaeroidites (ancestral Palmae) and Droseridites senonicus (possibly Droseraceae) in the Turonian; Auriculiidites spp., Cupanieidites spp. (Sapindaceae), and the Contantinisporis group (possibly Myrtales) in the Santonian/Coniacian; Buttinia andreevii (unknown), Aquilapollenites spp. (Loranthales/Santalales), and Periretisyncolpites spp. (possibly Illiciaceae) in the Campanian (8474 Ma) and numerous Proteaceae and Palmae, Ctenolophon (Ctenolophonaceae), Anacolosa and Restionaceae in the Maastrichtian (7466 Ma). The first evidence for differences, which might reflect Late Cretaceous provincialism is suggested from Gnetalean pollen, which is more diverse in S America than in Africa from the Turonian onward (De Lima 1980), but obvious provincialism, based on the angiosperm pollen

record appears to have developed mainly in the Maastrichtian. There are also some Eocene (5436 Ma) dispersal events, with pollen of the Amanoa type (Euphorbiaceae), Crudia (Leguminosae), Pelliciera rhizophorae (Theaceae) and members of Malpighiaceae and spores of a Schizaeaceous fern (Cicatricosisporites dorogensis) appearing at the same time on both continents (Fig. 5), and even some Oligo-Miocene (365 Ma) crossings, such as the mangrove genus Rhizophora (Rhizophoraceae), a Sonneratioid or Lythraceous mangrove indicated by the pollen species Verrutricolporites rotundiporus (Morley 2000), the swamp-tolerant tree Symphonia globulifera (Guttiferae) and the climbing fern Lygodium scandens (Schizaeaceae) which were probably instances of sweepstake (dispersal by rare or chance events across a major barrier) dispersal (Fig. 7). The Walvis Ridge may also have acted as a dispersal path for Southern Hemisphere elements, such as Casuarina (sl) (Casuarinaceae), which appears in both southern Africa (Scholtz 1985) and southernmost S America (Romero 1993) in the Paleocene (Fig. 4).

Dispersal route between S America and E Gondwana

Clear dispersal events occurred between S America and Antarctica through to New Zealand and Australia

Fig. 7. Early Tertiary, Oligocene plate tectonic reconstruction and palaeogeography according to Smith et al. (1994), occurrences of evaporites from Parrish et al. (1982) with additions, and closed-canopy megathermal rain forests according to Morley (2000). Noteworthy dispersals are indicated for (a) Miocene sweepstakes dispersals across the S Atlantic; and (b) Oligo-Miocene dispersals between the Australian and Asian Plates. The black southward-pointing arrows reflect the proportions of Boreotropical taxa finding refuge in each of the three tropical rain forest blocks. Numbers refer to the number of Boreotropical genera which are represented in each rain forest block (from Tiffney 1985b).

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via the South Sandwich Islands and South Georgia, which would have formed a continuous connection during the Late Cretaceous and Paleocene. This path was followed by primitive angiosperms, such as winteraceous Bubbia/Bellobium and Drimys (Dettmann 1994), in the Campanian. Other taxa that followed this route during the Late Cretaceous and Early Paleocene were Gunnera, ancestral Proteaceae, members of Cunoniaceae, Didymelaceae and Myrtaceae (Figs. 3 and 4). Taxa that dispersed to the north were Ilex, which has its oldest records in the Turonian of Australia (Martin 1977), Casuarina (sl) and Anacolosa. It is likely that the S AmericaAntarctic connection involved a direct land bridge to facilitate dispersal of taxa such as the microthermal Nothofagus (Fagaceae), although due to its near-polar position, this bridge would strictly have acted as a filter for megathermal plants, inhibiting their dispersal. Climates were sufficiently warm, however, at the time of the Late Paleocene/Early Eocene thermal maximum, for a floristic interchange with S American megathermal and mesothermal elements with members of Bombacaceae, Restionaceae, Sapindaceae and Polygonaceae dispersing to Australia (Fig. 5).

Dispersal route between Africa and India

At the time of the initial radiation of angiosperms, the Indian Plate was located too far south to include a significant angiosperm element, or to bear vegetation of tropical aspect. The Indian Plate separated from Gondwana in the Aptian, and drifted rapidly northward during the Mid Cretaceous, during which time it bore a Gondwanan flora similar to that of Australia. By Cenomanian to Turonian times, it lay in close proximity to Madagascar, and at this time, many plant taxa were able to disperse from Africa, via Madagascar and its associated islands, to India. One of the first of these was a winteraceous plant that produced Afropollis pollen (IEDS 1996); it became extinct in the Mid Cenomanian. This pollen type was succeeded in the Turonian, or Early Senonian (9074 Ma), as India drifted across the southern hemisphere high-pressure zone, by the Constantinisporis group of pollen, which is triporate with subequatorial pores and widely misidentified with pollen of the palm Sclerosperma but more likely of Myrtalean affinity (Harley 1996). The Constantinisporis group, reported from the Late Cretaceous of W Africa (Belsky & Boltenhagen 1963) and of S America (Muller et al. 1987), is recorded from Madagascar by Chen (1978), and widely from the Late Cretaceous of India (Venkatachala 1974; Nandi 1991). Other taxa that are likely to have followed the same route (Figs. 3 and 4) are members of Sapindaceae, Palmae (including nypoid palms and the Lon-

gapertites group), Myrtaceae, Ctenolophon (Srivastava 1987/88) and Normapolles (Nandi 1984, 1991). Dispersals ceased during the Maastrichtian, after the Indian Plate separated from Madagascar and began to drift rapidly northward toward Asia. The dispersal path from Africa to India was likely to be a filter, as many African taxa failed to make the crossing (Morley 2000). Some lines of evidence suggest that the Cenomanian/Turonian dispersal path to India also reached Australia. The elater-bearing ephedroid pollen type Elateropollenites africaensis provides the best evidence for this dispersal route, being widespread throughout the Cenomanian and Turonian of Africa and S America, but appearing suddenly in the Turonian of Irian Jaya (Morley 2000). It is possible that some angiosperms followed the same route, perhaps following the Kohistan Arc, with which the Indian Plate collided during its northward drift from Gondwana (Treloar et al. 1989; Treloar & Coward 1991). A further dispersal opportunity to the Australian Plate may have been created as India moved northward, since a string of islands formed along its southern trailing edge (Scotese 2001). These islands form the NinetyEast Ridge and were generated at the Kerguelen mantle plume hotspot. Although there is no direct evidence, they may have facilitated dispersal between India/Madagascar and the Australian Plate in the Early Tertiary. Paleocene and Oligocene sediments from the NinetyEast Ridge have yielded pollen of Australian affinity with one Madagascan endemic (Didymeles, Didymelaceae) and are thought to reflect long distance dispersal (Kemp & Harris 1975).

Middle Eocene to present, phase of plate collision coupled with global cooling
From the Middle Eocene to present day, three tectonic events have had an impact on trans-oceanic angiosperm dispersal routes as follows: (1) the collision of the Indian Plate with Asia, (2) the complex collision of the Australian Plate with the Philippine and Asian Plates, and (3) the formation of the Panamanian Isthmus. During the period of these collisions, global climates underwent stepwise cooling, strongly inhibiting interplate dispersal of megathermal plants outside the tropical zone.

Collision of Indian Plate with Asia

At the time of the collision of the Indian Plate with Asia, during the Middle Eocene (5039 Ma), both India and SE Asia lay at similar latitudes and within the same

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moist climatic belt (Fig. 5), as evidenced by the common occurrence of coals within the sedimentary record of both areas. At this time, SE Asian palynofloras show a sudden appearance of pollen of taxa that were characteristic of the Paleocene and Early Eocene of India, or were of Gondwanan affinity, such as Durio type (Bombacaceae), Gonystylus (Thymelaeaceae), Iguanuroid palms, aff Beauprea (Proteaceae), Restionaceae and Mischocarpus type (Sapindaceae). At the same time, most of those SE Asian pollen types that cannot be referred to modern taxa disappear from the record. This is thought to reflect the establishment of a moist corridor between India and SE Asia, with the dispersal of a somewhat aggressive Indian flora into SE Asia and the extinction of many local elements (Morley 1998, 2000). Middle and Late Eocene (3935 Ma) palynofloras from Java are very diverse (Lelono 2000), suggesting that shortly after this phase of immigration, under the influence of a warm and moist climate, the SE Asian flora had become notably quite species-rich. Dipterocarpaceae are also likely to have dispersed to SE Asia from Africa via the Indian Plate, since they currently have representatives in both Africa, S America, Seychelles and Sri Lanka (Ashton & Gunatilleke 1987) and fossil woods referable to the SE Asian subfamily Dipterocarpoidae are reported from the Tertiary of E Africa (Bancroft 1935). In SE Asia, their earliest fossil record is from the Middle Eocene of Myanmar (Curiale et al. 1994) in the form on the geochemical biomarker bicadenane (thought to be derived from

dipterocarp resins; van Aarssen et al. 1990) and is consistent with dispersal via the Indian Plate. One reason why it has taken so long to realise the significance of the Indian Plate as a vector for rain forest plants from Africa to SE Asia is that there are currently very few wet climate refugia within the Indian subcontinent. The ones there have mostly disappeared as a result of the development of monsoonal climates following the uplift of the Himalayas and the drift of the Indian Plate into the northern sub-tropical high pressure zone. A recent molecular evolution study by Conti et al. (2002) on Crypteroniaceae and its allies confirm the importance of this dispersal route and of the Out of India hypothesis.

Collision of the Australian Plate with the Philippine and Asian Plates
To explain the intermingling of Australasian and SE Asian floras following the collision of the Australian and Asian/Philippine Plates provides a major challenge. Examination of present-day Sundanian floras show clear examples of taxa of Australasian affinity, especially within the kerangas (heath forest) floras of nutrient-poor podzolic soils, and in the mountains, whereas New Guinea and its associated islands in Eastern Indonesia and the Pacific, with its predominantly Gondwanan substrate, has a mainly SE Asian flora (Good 1962; Wiffin 2002). The rain forests of Eastern Australia also have some elements of SE Asian

Fig. 8. Noteworthy instances Pliocene and Pleistocene dispersals of microthermal taxa into the low latitudes. Distribution of closed canopy tropical rain forests during the last glacial maximum (Morley 2000).

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affinity. This scenario for vascular plants must be viewed against that for the fauna of the region, on which Wallaces Line was drawn, and which closely follows geological substrates, with faunas of predominantly Australasian affinity in New Guinea and to the east, and faunas of Asian affinity in Borneo, with a mixture of affinities in intervening Wallacea (George 1981). The concentration of SE Asian elements in the floras of Eastern Indonesia is thought to be relate to the tectonic history of the south-western arm of Sulawesi, which was attached to south-eastern margin of Borneo in the Middle Eocene prior to the formation of the Makassar Straits (Hall 1996, 2002), and has yielded Asian affinity palynomorph assemblages of Middle Eocene age similar to those recorded through the Eocene of Java and Kalimantan (Morley 1998). Following uplift of New Guinea and its associated islands in the Miocene, it is thought that this flora was able to disperse to the east without the need to cross Wallaces Line and the Makassar Straits, which has been a biogeographic barrier during the Younger Tertiary. The antiquity of this flora explains why Wallaces Line acts as a major dispersal barrier to the regions fauna, but has not been such an effective barrier to its flora. Dispersals from Australasia to Sunda fall into three groups: firstly, well-dispersed taxa which were probably able to island hop following the initial phase of collision of the Australian Plate with the Philippine Plate (Halmahera) close to the Oligo-Miocene boundary; secondly, chance dispersals across the Makassar Straits; and thirdly, dispersals of montane taxa following the Middle to Late Miocene uplift of New Guinea and other islands. The first group of dispersals, relating to the initial phase of the collision of the Australian Plate with the Philippine Plate, is based on pollen records of the Australasian taxa Phormium or Dianella (Hemerocallidaceae), Casuarina (sl) and Dacrydium (Podocarpaceae) which suddenly appear in Java in the MidTertiary (Fig. 7). This dispersal event was originally thought to be at about 21 Ma in the Early Miocene (Morley 1998, 2000), but recent unpublished work suggests that it is likely to be a little earlier, within the Late Oligocene. Following on this dispersal event, subsequent dispersals occur at irregular time intervals, with Myrtaceae pollen showing a sudden increase in abundance at about 17 Ma, interpreted by Muller (1972) as due to dispersal from the east, followed by the appearance of pollen of the mangrove tree Camptostemon (Bombacaceae) at about 14 Ma, and spores of the climbing fern Stenochlaena milnei (Blechnaceae) at 9 Ma. A further unpublished record suggesting dispersal from Australasia is of proteaceous pollen, recorded from the

Malay Basin by Jaizan Md Jais (1997). These dispersals are thought to represent sweepstake dispersals across the Makassar Straits. The third group of dispersals concerns mountain plants. The podocarp Dacrycarpus dispersed to New Guinea from Australia in the Middle Miocene, and subsequently to Borneo in the Mid Pliocene, at about 3.5 Ma, but recent unpublished work suggests that it did not disperse to Sumatra until 1.6 Ma, from where it quickly spread to Indochina. The second was Phyllocladus, which dispersed westward only to Borneo at about 1.6 Ma. These taxa dispersed from New Guinea at a time when there was considerable uplift in the southern Philippines (van der Kaas 1991). The presence of islands in this area probably allowed the birddispersed Dacrycarpus to island hop to the Sunda region. It is noteworthy that Nothofagus (Fagaceae) accompanied Dacrycarpus to New Guinea, where it first appears during the Middle Miocene (Morley 2000), but since Nothofagus is more poorly dispersed, it was never able to spread further west in the manner of Dacrycarpus. The montane dispersal route via New Guinea may therefore be viewed as a filter for montane elements with good dispersal mechanisms. The Australian pollen record provides relatively few data that might reflect dispersals from the SE Asian area following Neogene plate collision (Truswell et al. 1987). Possible examples of dispersal from the north, based on Mid-Tertiary pollen appearances in Australia, are the climbing fern Stenochlaena palustris (Blechnaceae), Acacia, Caesalpinia and Crudia (Leguminosae) and Merremia (Convolvulaceae). One surprising possible example is of Symplocos, which was an element of the northern hemisphere Boreotropical flora in the Palaeogene, but according to Martin (1994) reached Australia only in the Middle Miocene.

Formation of the Panama Isthmus

With further westward dispersal of the N and S American Plates, and eastward movement of the Caribbean Plate, the island arc that had formed along the trailing edge of the Caribbean Plate collided with the southern end of the N American Plate during the Mid Tertiary, and with Colombia in the Early Miocene (Pindall et al. 1988). With subsequent uplift it formed a continuous land bridge between N and S America during the Pliocene, which was followed by the faunal migrations of the Great American Interchange (Marshall et al. 1979, 1982). The most noteworthy plant taxa for which there is a fossil record indicating times of dispersal are the microtherm genera Juglans (Juglandaceae), Alnus (Betulaceae) and Quercus (Fagaceae), which dispersed into

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Colombia at about 2.2 Ma, 1.0 Ma and 300,000 Ka respectively. Alnus and Quercus have subsequently become major components of Andean forests in Colombia (van der Hammen & Gonzalez 1964; Andriessen et al. 1993; Hoogheimstra 1984; Hoogheimstra & Ran 1994). An additional possible emigration from N America during the Late Neogene is suggested by the sudden appearance of pollen of the montane forest tree Hedyosmum (Chloranthaceae) at about 4 Ma in Colombia and the Late Miocene of Guyana (Wymstra 1971). This genus has been shown by Colinvaux et al. (1996) to have been a component even of lowland vegetation during glacial periods. It is also likely that relics of the Palaeogene Boreotropical forests (see below) followed this route, such as Trigonobalanus, Meliosma and Saurauia, all of which have Mid-Tertiary fossil records in Europe and N America, with present distributions including the Colombian Andes.

Demise of the Boreotropical flora

One of the consequences of Mid-Tertiary global cooling was the replacement of the northern mid-latitude Boreotropical forests with frost-tolerant temperate vegetation (Fig. 7, 8). Frost-sensitive Boreotropical taxa had to disperse to the low latitudes, or face extinction. A large number of these taxa found refuge in rain forests of S China and Sunda, dispersing southward along the permanent land connection between the equatorial zone and northern mid-latitudes. Plant taxa were able to disperse between these two latitudinal zones in East Asia unhindered by oceanic barriers. With respect to the European Boreotropical flora, a combination of latitudinal barriers, such as the Mediterranean Sea, the Sahara desert and the uplift of the Alps prevented southward dispersal into the African low latitudes. For this reason there are hardly any Boreotropical elements finding refuge in African rain forests today. Tiffney (1985b) estimated that only eight Boreotropical genera are represented in the present African flora. Within the Americas, Boreotropical elements were probably able to find refuge to some degree along the southern margin of the N American Plate, but could not disperse to equatorial latitudes until the formation of the Isthmus of Panama during the Late Miocene and Pliocene. Those elements of the Boreotropical Province that were eventually able to find refuge in C and S America, typified by Ampelopsis (Vitaceae), Gordonia, Meliosma, Saurauia, Symplocos and Trigonobalanus are now preserved as the amphi-Pacific element of van Steenis (1962). Tiffney (1985b) recognised 22 Boreotropical genera in the present day Neotropics.

In the Far East, there are many more representatives of Boreotropical forests, with, in addition to amphi-Pacific elements, Alangium, Cinammomum and Endiandra (Lauraceae), Dracontomelon and Lannea (Anacardiaceae), Mastixia (Cornaceae), and the palms Livistona, Nypa and Oncosperma. Tiffney (1985b) identified 34 Boreotropical genera in Far Eastern forests. The relative representation of Boreotropical elements in the Far East, Africa and Neotropics (Fig. 7) thus reflects the different opportunities for southward dispersal to the tropical regions during the Late Tertiary (Morley 2001) and explains the frequent references in the literature to the relationship between Laurasian Early Tertiary floras with Malesia since the similarity was first noticed by Reid & Chandler (1933). The possibility of dispersals of mesic Boreotropical elements during the Late Cretaceous to Middle Eocene should also be given consideration since appropriate land bridges would have been in place to allow Boreotropical elements to disperse to both S America and SE Asia. Such taxa should also be poorly represented in Africa, which was isolated from the northern mid latitude land masses from the Paleocene to Middle Miocene. Symplocos might provide an example of this distribution type (Figs. 4, 7).

Discussion
The dispersal routes for megathermal plants as indicated from the fossil record correspond remarkably closely with the land bridges proposed by van Steenis (1962) with a single exception (Table 1). He proposed

Table 1. Comparison of the ages of interplate dispersal routes as suggested from plate tectonics with those of van Steenis (1962) in his Land Bridge theory (, no information). Steenis Land Bridge 1 Transatlantic 2 Transpacific 3 Madagascar Ceylon 4 Subantarctic 5 Bering 6 N Atlantic 7 C American 8 EuropeAfrica 9 AustraliaSE Asia Age (van Steenis 1962) JurassicMid Cretaceous JurassicCretaceous MidLate Cretaceous MidLate Cretaceous Age based on plate tectonics EarlyLate Cretaceous Not needed Late CretaceousEocene Mid CretaceousPaleocene

Late CretaceousTertiary Not applicable to tropical plants Late CretaceousEocene Late CretaceousEocene + Pliocene Late Cretaceous Eocene Neogene

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a transpacific route for his amphi-Pacific element, but the fossil record clearly shows that virtually all of the taxa that he explained through a trans-Pacific land bridge were actually elements of the Boreotropical forests which spread across the northern mid-latitudes during the Early Tertiary, their absence from Africa reflecting greater obstacles to dispersal from Laurasia to the African Plate than to SE Asia or S America. Outside areas of Neogene tectonic plate collision, such as Wallacea and the Panama Isthmus, which are current areas of active dispersal of megathermal plants, most interplate dispersal took place in the Late Cretaceous and Early Tertiary at a time when global climates were markedly different from those of today and the global area of megathermal vegetation several times greater than at present. Under such a scenario, it is likely than opportunities for speciation were much higher that for present day megathermal plants. It is also noteworthy that most higher rank biogeographical patterns reflect these relatively ancient dispersal patterns. The contrast between Early Tertiary and later dispersals is highlighted by the nature of Late Pliocene and Quaternary dispersal events, for in each of the three tropical rain forest blocks, the only dispersal events reflected by the pollen record are of microtherm taxa dispersing into the equatorial zone, with Alnus, Quercus and Juglans in the Neotropics, Dacrycarpus and Phyllocladus in the eastern tropics, and Podocarpus milanjianus in West Africa (Fig. 8). The fossil record also suggests that plants are unlikely to disperse from one plate to another unless not only an appropriate land connection is available, but also the climate is appropriate. Clearly dispersals are more likely within the same latitudinal climatic zone than between climatic zones. This may explain why the dispersals from N America to Europe during the Late Paleocene/Early Eocene and from India to SE Asia in the Middle Eocene were so successful, because at the times of connection, the two floras were within the same climatic zone, and similar climates are likely to have occurred in the corridor between both regions, facilitating dispersal. The relationship between plants and climate may also explain why the initial radiation of Chloranthoid angiosperms paid so little attention to either climatic or geological boundaries. If these plants became established initially in edaphically defined settings, such as along river courses, they would have been less dependent on rainfall. Conditions prevailing in gallery forests in regions of warm subhumid climate may well have been similar to those in riverine swamps in areas of warm ever-wet climate. Understanding the geological history of angiosperms in general, and the temporal framework for

rain forest expansion and contraction, is important for all taxonomic and molecular studies involving tropical plants, even for those without a fossil record. Knowledge of the availability and timing of appropriate dispersal pathways to a taxon limits the number of dispersal opportunities open to it and allows the taxonomist to make better judgements regarding phylogeny, even without the availability of a fossil record.

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