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Systematics and Phytogeography |
2Department of Biology, University of Missouri-St. Louis, St. Louis, Missouri 63121 USA; and The Missouri Botanical Garden, St. Louis, Missouri 63166 USA 3Institut für Spezielle Botanik, Universität Mainz, D-55099 Mainz, Germany
Received for publication August 22, 2000. Accepted for publication January 11, 2001.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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3000 species in the neotropics, 1000 in tropical Asia, 240 in Africa, and 225 in Madagascar in 150–166 genera, and the taxa sampled come from throughout this geographic range. Based on fossils, ranges of closest relatives, tree topology, and calibrated molecular divergences, Melastomataceae initially diversified in Paloecene/Eocene times in tropical forest north of the Tethys. Their earliest (Eocene) fossils are from northeastern North America, and during the Oligocene and Miocene melastomes occurred in North America as well as throughout Eurasia. They also entered South America, with earliest (Oligocene) South American fossils representing Merianieae. One clade (Melastomeae) reached Africa from the neotropics 14–12 million years ago and from there spread to Madagascar, India, and Indochina. Basalmost Melastomataceae (Kibessieae, Astronieae) are species-poor lineages restricted to Southeast Asia. However, a more derived Asian clade (Sonerileae/Dissochaeteae) repeatedly reached Madagascar and Africa during the Miocene and Pliocene. Contradicting earlier hypotheses, the current distribution of Melastomataceae is thus best explained by Neogene long-distance dispersal, not Gondwana fragmentation.
Key Words: biogeography • fossil calibration • long-distance dispersal • Melastomataceae • Memecylaceae • molecular clock • ndhF.
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; Gentry, 1982, 1993
; Barlow, 1990
; Romero, 1993
; Burnham and Graham, 1999
, their Table 2). However, this implies ages of 100–90 million years (Ma) for divergences among neotropical-African sister clades and even older divergences for taxa also found in Indochina and Southeast Asia. However, fossils supporting a Gondwanan range are available for few of the many pantropical eudicot families. Other workers have taken the absence of Cretaceous fossils at face value and attributed most tropical range disjunctions to long-distance dispersal (Smith, 1973
; Thorne, 1973
; Brenan, 1978
). In the absence of a dense fossil record from key areas, such as South America (Burnham and Graham, 1999
), the controversy pitting the break up of Gondwana against more recent long-distance dispersal as major factors shaping today's geographic ranges could not be resolved.
To resolve the problem, absolute age estimates for sister groups are needed. These can come from nucleotide substitution rates if a single rate across a phylogenetic tree can reliably be estimated and calibrated. Much discussion has surrounded both tasks. The assumption of similar substitution accumulation among descendents of a common ancestor must be justified prior to drawing inferences about either rates or times. And fossil or geological evidence must constrain the age of at least one calibration point on the tree in order for rates to be transformed into absolute ages. Doyle and Donoghue (1993
; see also Sanderson, 1998
) have characterized the kind of time constraint that can be inferred from fossils. To fix the age of a calibration node A, a fossil must exhibit not only the apomorphy of clade A, but also some additional synapomorphy of a subclade of A. Together, these traits indicate that the fossil is indeed part of clade A (thus proving its minimum age), rather than perhaps representing the sister group of A.
We here estimate absolute ages for key divergence events in the history of Melastomataceae from substitution rates estimated for their NADH dehydrogenase subunit F gene (ndhF) calibrated with different fossils. Melastomataceae are an important component of today's tropical floras and comprise 3000 species in the neotropics, 1000 in tropical Asia, 240 in Africa, and 225 in Madagascar in together 150–166 genera (Renner [1993
] for all genera's ranges and species numbers; a few genera have been synonymized since then). Their sister group, Memecylaceae, include 360 species in four genera in Africa, Madagascar, and Southeast Asia, and 90 species in two genera in tropical America. Melastomataceae fossil leaves and seeds are known from Eurasia and North America, and these fossils show morphological details that permit their assignment to particular nodes in a phylogeny (below). Memecylaceae wood is known from the Oligocene of Germany (Gottwald, 1992
). The families' closest relatives are species-poor lineages with restricted geographic ranges and poor fossil records, making it difficult to constrain the age of the entire clade. However, Myrtaceae, which are more distant relatives of Melastomataceae and Memecylaceae, have a fossil record that goes back to the Upper Cretaceous (Raven and Axelrod, 1974
; Manchester, 1999
).
Melastomataceae have been seen as having originated either in West Gondwana (South America + Africa) (Raven and Axelrod, 1974
; Gentry, 1982
: Table 5, 1993; Jacques-Félix, 1994
) or East Gondwana (Madagascar, India, Australia, Antarctica) (Renner, 1993
). Earliest fossils are Eocene leaves from northwestern North America (Hickey, 1977
; Wehr and Hopkins, 1994
), an Oligocene leaf from Andean Colombia (Huertas, 1977
), and Miocene seeds from throughout Eurasia (Dorofeev, 1960, 1963, 1988
; Collinson and Pingen, 1992
; Dyjor et al., 1992
; Fairon-Demaret, 1994
; Mai, 1995, 2000
). Pollen records are much younger. These fossils suggest that the family is too young to have reached its present pantropical distribution via overland dispersal. Nevertheless, the above-cited biogeographic analyses preferred the assumption of "missing older fossils" over the assumption of long-distance dispersal between South America and Africa, and Africa, Madagascar, and tropical Asia.
As will be presented, branch lengths in the ndhF tree can be calibrated by using two fossil-constrained calibration points. The resulting estimated times for the spread of Melastomataceae lineages across the tropics imply repeated long-distance dispersal events during the Neogene.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; primer sets used are specified in Clausing and Renner, 2001a
). For purification, PCR products were run on a low melting point agarose gel and then recovered with the help of QIAquick gel extraction kits (QIAGEN) or by using the QIAquick purification columns directly. Cycle sequencing of the amplified double-stranded products used the ABI Prism Dye Terminator ready reaction kit (Perkin Elmer, Norwalk, Connecticut, USA), the dye was removed by ethanol precipitation, and samples were run on an ABI 373 automated sequencer at the University of Mainz. Consensus sequences were constructed using Sequencher version 3.1 (GeneCodes Corporation, Ann Arbor, Michigan, USA) and SeqPup version 0.9 (D. Gilbert, Indiana University, Bloomington).
Taxon sampling, phylogenetic analysis, and rate and age estimation
A phylogenetic hypothesis for 45 representatives of Melastomataceae and Memecylaceae is available based on sequences from two chloroplast DNA genes and one intron (Clausing and Renner, 2001a
). For a biogeographic analysis, denser taxon sampling was desirable to substantiate unexpected groupings, such as the nesting of African, Madagascan, and Southeast Asian Melastomeae within a clade of neotropical Melastomeae and the nesting of Madagascan and African Sonerileae among Southeast Asian Dissochaeteae/Sonerileae. Second, to assess possible substitution rate constancy between sister groups, it was necessary to concentrate on one genome region. This is because the evolution of combined gene and intron sequences, the latter with large insertions and deletions, is less likely to be captured by a model of nucleotide substitution. Of the molecular data sets begun, the ndhF data appeared most suited for a molecular clock approach because of intermediate levels of divergence. We sequenced ndhF for 102 species from 59 genera of Melastomataceae and Memecylaceae of which 91 are included here. (A few congenerics differed little from each other and were therefore excluded.) Table 1 lists all species with their geographical ranges, vouchers, and GenBank accession numbers. Trees were rooted with representatives of Alzateaceae, Crypteroniaceae, Oliniaceae, Penaeaceae, Rhynchocalycaceae, Myrtaceae, Onagraceae, and Lythraceae. While ndhF alone is used for age estimation, the biogeographic discussion is restricted to clades that have strong bootstrap support in the two genes/one intron analysis with the exception of the Madagascan Medinilla clade of which but one species had been sampled.
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).
Maximum likelihood analyses were performed using the general time-reversible model (Yang, 1994
), which estimates independent probabilities for the six possible nucleotide substitutions in a symmetric rate matrix and also allows unequal base frequencies. This model had been found to fit the data significantly better than less parameter-rich models (data not shown). To compensate for among-site rate heterogeneity some proportion of nucleotide sites (Pinv) was modeled as completely resistant to change while substitution in the remainder was assumed to follow a gamma distribution with shape parameter 
. The rate matrix, Pinv, and were estimated simultaneously, using the discrete approximation of Yang (1994
; implemented in PAUP*) with four rate categories and empirically observed base frequencies. All searches were heuristic, with the "multiple trees" and "steepest descent" options in effect, and tree-bisection-reconnection (TBR) swapping. The initial search for model estimation used a minimum evolution (ME) starting tree (Rzhetsky and Nei, 1992a, b
; implemented in PAUP*) and log-determinant genetic distances (Lockhart et al., 1994
). The dependence of model parameters on tree topology is minor as long as strongly supported groups are maintained (Yang and Kumar, 1996
; Sullivan, Swofford, and Naylor, 1999
), and we therefore aborted parameter estimation after 400 TBR rearrangements (36 h). The estimated parameters were used in searches with 20 000 TBR rearrangements that used different starting trees, namely an ME tree, a neighbor-joining tree, and a randomly chosen parsimony tree. Nonparametric bootstrap frequencies were obtained with PAUP's "fast bootstrap" option under the ME optimality criterion and using ML genetic distances with the same ML settings as used before.
Likelihood ratio testing was used to assess whether substitution rates in sister groups can justifiably be modeled as clock-like. Likelihood ratio tests are extremely powerful at detecting differences in rates (Sanderson, 1998
), and in large data sets they usually reject the null hypothesis of rate constancy between sisters. This is the case here. The number of branches significantly shortened or lengthened in the clock tree compared to the nonclock tree was assessed by a Student's t test. Since there were few differences in topology and branch lengths between the two trees (Results), genetic distances (branch lengths) in the clock tree were calibrated by assigning fossils to two calibration points. We used binomial probability theory to estimate the standard deviation (SD) of the distance from a fixed calibration node to the tips and then used this value to obtain the SDs of the estimated ages. The number of nucleotide substitutions (S) is equal to the product of the total number of nucleotides in a sequence (N) times the proportion of nucleotides substituted (p). Thus, S = Np. The SD of this value is the square root of Np(1 – p), or SD(S) = sqrt(Np[1 – p]). The SD of the number of nucleotides substituted divided by the total number of nucleotides is the SD of the proportion of nucleotide substitutions. Thus, SD(p) = sqrt(p[1 – p]/N).
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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was 1.30, indicating that most sites have intermediate substitution rates, while few sites have very high or low rates (Yang and Kumar, 1996
).
The midpoint-rooted highest likelihood tree (Fig. 1) obtained under the GTR + Pinv + 
model shows Melastomataceae, Memecylaceae, and the Southeast Asian genus Pternandra (Kibessieae) as a strongly supported (100% bootstrap support) clade that is sister to a clade comprising Crypteroniaceae, Alzateaceae, Rhynchocalycaceae, Oliniaceae, and Penaeaceae. Within Melastomataceae, eight major groups (marked in Fig. 1) can be discerned that traditionally have been recognized at the tribal level (Cogniaux, 1891
; Clausing and Renner, 2001a
). They are (1) Astronia (Astronieae), (2) Merianieae, (3) Miconieae, (4) Blakeeae, (5) a complex formed by all Dissochaeteae and Sonerileae, (6) Bertolonieae, (7) Melastomeae, and (8) Microlicieae.
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2 = 2(10 072.09 – 9886.45) = 371.28; P < 0.001; 89 df). The highest likelihood topology found under the nonclock model (after 21 864 rearrangements when TBR swapping on two trees was aborted) and the topology found under the clock model (after 1400 rearrangements when swapping on two trees was aborted) differed in minor rearrangements (compare Figs. 1 and 2). (Searches were run for the same time but searching under the clock model took much longer.) Thus, relationships within Memecylaceae and within neotropical Melastomeae differ. A Student's t test showed that of 163 matched branches in the two trees, 25 (15%) significantly changed in length (P < 0.05) when the data were modeled under the clock. (Four of the 168 branches in the nonclock tree were so short that standard errors on them were undefined; one branch was not seen in the clock tree and could therefore not be matched; and 11 branches involved polytomies.) Note that 5% of branch lengths are expected to differ by chance alone. Since most branch lengths remained virtually unchanged (compare Figs. 1 and 2), including the ones of interest for the biogeographic analysis (nodes A through E), the use of branch length as a proxy for age is justified.
Branch length was calibrated in two ways, once with fossil seeds, once with leaves. Melastome seeds are known from Miocene deposits in Siberia, the Tambov region, Belorussia, Poland, several sites in Germany, and Belgium (Dorofeev, 1960, 1963, 1988
; Collinson and Pingen, 1992
; Dyjor et al., 1992
; Fairon-Demaret, 1994
; Mai, 1995, 2000
). They are 0.8–1.5 mm long and strongly curved (cochleate) with a round operculum (Fig. 2, inset). The testa is tuberculate and carries a secondary ornamentation. Such seeds characterize Melastomeae (including Rhexieae) (Whiffin and Tomb, 1972
; Renner, 1989, 1994
; F. Almeda, California Academy of Science, unpublished data). All other Melastomataceae have straight or cuneate seeds that lack round operculi and have different testas. The seed fossils thus constrain the age of node B in Figs. 1 and 2, making it minimally 23 Ma old (Berggren et al., 1995
). Fixing node B at 23 Ma yields a substitution rate of 0.0012 substitutions·site–1·Ma–1 (the distance from node B to the tips divided by 22 is 0.0012). Table 2 shows age estimates and standard deviations obtained for divergence events C, D, and E under this rate.
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), the Early Middle Eocene of Washington State (50–48 Ma; Wehr and Hopkins, 1994
), and the Oligocene of northern Colombia (30 Ma; Huertas, 1977
). Mid-Miocene leaves are known from the Latah formation near Spokane (18–16 Ma; J. Wolfe, personal communication, March 2000), Late Miocene leaves from Sumatra (Kräusel, 1929
), and Pliocene leaves from Brazil (Duarte, 1956
). A presumed Paleocene leaf from Antarctica (Dusén, 1908
) does not belong to the family (Birkenmajer and Zastawniak, 1989
) nor do Paleocene leaves from the Fort Union formation in Montana and the Ferris formation in Wyoming (Brown, 1962) whose leaf surfaces have glandular dots. The Early Middle Eocene leaves from Washington State (of which 20 were received on loan by S. S. Renner and one is depicted in Fig. 2) show numerous densely spaced pairs of tertiary veins running in strictly parallel rows between the midrib and pairs of secondary veins that run from the base to the leaf tip. A further faint pair of secondaries can sometimes be discerned very close to the margin. The leaves are entire-margined, have weakly developed drip tips, and are of medium size (between 5–9 cm long and 2.5–5 cm wide). The single Early Eocene leaf from North Dakota is very similar in shape and size, but shows the tertiary venation less clearly. While these fossils document that Melastomataceae with leaves resembling those of modern Miconieae and Merianieae were present in North America 53 Ma ago, it is difficult to assign them to a node in the phylogeny. Most conservatively, they constrain the age of node A in Fig. 2 because all Melastomataceae and Kibessieae (Pternandra) have the basic kind of venation seen in the fossils, while none of the outgroups do.
Fixing node A at 53 Ma yields a rate of 0.0011 substitutions·site–1·Ma–1 (the distance from node A to the tips divided by 53). Table 2 shows the age estimates obtained under this second rate for events C, D, and E.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; Gentry, 1982
: Table 5, 1993; Renner, 1993
; Jacques-Félix, 1994
) are rejected by the ndhF-based age estimates. While the fossil record of Melastomatacaeae and their relatives may be biased, undiscovered fossils would have to be nine times older than current oldest fossils to push back the estimates for entry into Africa and Madagascar to 90 or 100 Ma. Also, while unaccounted-for rate heterogeneity among sister groups must introduce error into the age estimates, the magnitude of such error in this data set may be small, judging from visual comparison of the clock and nonclock trees. Further error could stem from the tenuous links between calibration nodes and fossils assigned to them and from uncertainties in the dating of fossils. However, all three types of error are likely random with respect to under- or overestimation of rates and ages. Reduction of estimation error—and rejection of wrong biogeographic hypotheses—may come from additional fossils that can add calibration points or prove hypothesized ancient ranges.
The color-coding in Fig. 2 indicates current geographic ranges occupied by the respective taxa (with very limited back-extrapolation), while the tree's backbone is left black to indicate past ranges. From the Eocene through the Miocene both Melastomataceae and Memecylaceae occurred in tropical Laurasia, and the tree's black spine could be seen as symbolizing that region. Of their sister families, Crypteroniaceae (3 genera, 10 species) are endemic in Southeast Asia, where they are considered a Laurasian element (Raven and Axelrod, 1974
; their range is mapped in Zhu, 1997
). Alzateaceae (1 species) are neotropical (see Table 1 for detailed ranges), and Rhynchocalycaceae (1 species), Oliniaceae (8 species), and Penaeaceae (7 genera, 20 species) South African. Memecylaceae themselves have their greatest morphological diversity in tropical East Africa (Jacques-Félix, 1978a, b, 1994
), and two of the African genera extend into Malaysia (Bremer, 1982
). More data are needed to resolve whether the neotropical genera of Memecylaceae are nested among paleotropical memecylons, which is indicated by preliminary combined gene and intron data (S. S. Renner and G. Clausing, unpublished data). The family's only fossil record is Oligocene (40 Ma) Memecylon wood from Germany (Gottwald, 1992
).
First-branching Melastomataceae occur in Southeast Asia. This refers to Astronieae and Kibessieae, which based on morphological and sequence data from three genome regions belong in Melastomataceae (Clausing and Renner, 2001a
). Astronieae are endemic in Southeast Asia, with most of their 150 species occurring in Borneo; Kibessieae comprise 15 species in the Malay Archipelago, with three reaching Thailand (Renner et al., in press
). Since basal Melastomataceae and all Memecylaceae are rain forest trees or treelets and since their earliest fossils are from the Eocene and Oligocene of tropical Laurasia, their common ancestor likely lived under humid tropical conditions. In combination, three lines of evidence thus point to an origin of Melastomataceae in the tropical belt that bordered the northern Tethys margin: (1) earliest fossil record restricted to Laurasia; (2) occurrence in Southeast Asia of oldest surviving lineages; and (3) likely humid forest habitat of stem lineage. If melastomes indeed originated somewhere to the north of the Tethys, they could readily have spread into North America (explaining their Early Eocene presence in Dakota) as well as tectonically more labile Southeast Asia (Hall, 1996
). Going back further in time, the reconstruction of ancestral ranges based purely on topology (without fossils) becomes dubious.
Following initial radiation (viz. the divergence of Kibessieae and Astronieae), higher Melastomataceae spread into Eurasia and North America as documented by fossils. By the end of the Early Eocene, temperatures reached a global high, and many warm-humid-adapted groups reached North America via Beringia and/or the North Atlantic land bridge (Manchester, 1999
; Tiffney, 2000
). Malaysian genera become prominent among Alaskan fossils at that time (Graham, 1993
), and melastomes with leaves similar to the relatively basal Miconieae and Merianieae reached northern Dakota and northeastern Washington. The younger seed fossils from Eurasia represent the phylogenetically younger Melastomeae. However, carpofloras in North America are limited to a few sites, and with the exception of the Brandon Lignite, are not the same kind of environment as the more thoroughly studied Oligocene/Miocene Browncoals of Europe (S. Manchester, personal communication, April 2000). While this may explain the absence of melastome seeds from North America, the absence of leaves from Eurasia may have to do with the disintegration during sieving of samples being prepared for paleocarpological investigations (F. Velichkevich, personal communication, September 2000).
Merianieae/Miconieae spread from North America into South America sometime in the Oligocene, as indicated by an Oligocene Meriania-like leaf from northwestern Colombia (Huertas, 1977
). This is the oldest securely identified melastome fossil from South America. In North America, melastomes persisted at least until the Mid-Miocene based on fossil leaves found near the Washington/Idaho border (J. Wolfe, unpublished data, personal communication, March 2000). Among their most widespread groups may have been the open habitat-adapted Melastomeae whose seeds are known from the Miocene of Siberia, the Tambov region, Belorussia, Poland, Belgium, and several sites in Germany. Modern Melastomeae are the most cold-tolerant Melastomataceae, occurring at high altitudes in the Andes and the Himalayas, and the tribe also includes the family's only temperate zone endemic, the Eastern North American Rhexia with 11 species. The earliest South American records of Melastomeae are Pliocene leaves from Brazil (Duarte, 1956
).
Paleotropical Melastomeae form a clade (node C in Figs. 1 and 2) that is deeply embedded in neotropical Melastomeae and between 14 and 12 Ma old (see Table 2 for standard deviations on all estimates). West African genera branch off first, which together with the estimated age of this divergence implies that West Africa was reached from the neotropics via long-distance dispersal. The only two Asian genera of Melastomeae, Osbeckia and Melastoma, likely derive from African or Madagascan ancestors (Figs. 1, 2). In another analysis, Melastoma is sampled more densely (Renner and Meyer, in press), and those data suggest that the radiation of Melastoma began during the Pleistocene.
Junction of the Australian continent with the Philippine Sea plate arc in the Miocene (Hall, 1996
) and the resulting tectonic reconfiguration of Southeast Asia provided the opportunity for diversification of another large clade, Dissochaeteae/Sonerileae (Figs. 1, 2). Based on the clock tree (Fig. 2), Dissochaeteae/Sonerileae began to diversify during the Miocene. They are confined to rain forest habitat and comprise woody climbers, herbs, and terrestrial and epiphytic shrubs (Clausing and Renner, 2001b
). Two of their sublineages reached Madagascar, one wind dispersed, the other bird dispersed. The wind-dispersed group (node D in Figs. 1 and 2) may have arrived in the Late Miocene, 11 Ma ago (Table 2). Today, it has 107 species in one genus in Madagascar (Perrier de la Bâthie, 1951
) and 50 species in seven genera in tropical Africa (Jacques-Félix, 1994
). The later-arriving bird-dispersed genus Medinilla (node E in Figs. 1 and 2) today comprises 70 species in Madagascar and three in Africa.
This and other studies of the history of major components of tropical biota (Lavin and Luckow, 1993
; Chanderbali, van der Werff, and Renner, 2001
) have implications for our understanding of the build-up of tropical floristic diversity in general. Based on molecular phylogenies and Laurasian fossil floras, it appears that the ancestral floras shared directly among Gondwanan fragments may be smaller than sometimes assumed. Entry of boreotropical taxa into southern continents, beginning in the Oligocene and Miocene, must be seriously considered as an additional means of establishing pantropical distribution patterns. Tertiary floras document tropical forests at high latitudes in Laurasia during the Eocene worldwide climatic optimum, with connections via Beringia or the North Atlantic land bridge between the floras of Indochina, North America, and to a limited extent South America. Examples of families today more or less restricted to Indomalesia, tropical Africa, and/or the New World tropics but that were widespread in Eocene and Oligocene Laurasia include Actinidiaceae, Bignoniaceae, Lauraceae, Menispermaceae, Musaceae, groups of Myrtaceae, Rubiaceae, palms, and Zingiberaceae (Mai, 1995
; Kubitzki and Krutzsch, 1996
; Manchester, 1999
).
Our analysis also shows that species accumulation in one of the largest families in the three regions of today's tropics occurred over vastly different lengths of time. Especially illustrative is the case of Madagascar, which harbors 225 species of Melastomataceae (Perrier de la Bâthie, 1951
) of which 21% stem from successive arrivals of African members of an ultimately neotropical lineage (Melastomeae), 48% from a capsular-fruited Asian immigrant (Sonerileae), and 31% from a berry-fruited Asian immigrant (Medinilla). The paucity of African Melastomataceae (240 species) to some extent may reflect the family's relatively recent (14–12 Ma) arrival on that continent from the west and from the east. Other lineages, such as Osbeckia and Melastoma, dispersed from Africa and/or Madagascar to Indochina and Malesia, adding to the many ties between these floras (Thorne, 1973
) and increasing species diversity in Southeast Asia. Like other taxa in the neotropics that derive from Laurasian ancestors (Gentry, 1982
), Melastomataceae are most species rich in the Andes, while their morphological diversity (wood, placentation, anther morphology, etc.) is greatest in Indochina and Malesia, regions where oldest and youngest lineages now coexist.
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FOOTNOTES |
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4 Author for reprint requests.
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LITERATURE CITED |
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Berggren W. A. D. V. Kent J. D. Obradovitch C. C. Swisher III 1995 Toward a revised Paleogene geochronology. In D. R. Prothero and W. A. Berggren [eds.], Eocene–Oligocene climatic and biotic evolution, 29–45. Princeton University Press, Princeton, New Jersey, USA
Birkenmajer K. E. Zastawniak 1989 Late Cretaceous–Early Neogene vegetation history of the Antarctic Peninsula sector: Gondwana break-up and Tertiary glaciations. Bulletin of the Polish Academy of Sciences, Earth Sciences 37: 63-88
Bremer K. 1982 Lijndenia, a re-established paleotropical genus of the Melastomataceae-Memecyleae. Nordic Journal of Botany 2: 121-124
Brenan J. P. M. 1978 Some aspects of the phytogeography of tropical Africa. Annals of the Missouri Botanical Garden 65: 437-478[CrossRef][Web of Science]
Brown R. W. 1992 Paleocene flora of the Rocky Mountains and Great Plains. Geological Survey Professional Paper 375. U.S. Government Printing Office, Washington, D.C., USA
Burnham R. J. A. Graham 1999 The history of neotropical vegetation: new developments and status. Annals of the Missouri Botanical Garden 86: 546-589[CrossRef][Web of Science]
Chanderbali A. S. H. van der Werff S. S. Renner 2001 The relationships and historical biogeography of Lauraceae: evidence from the chloroplast and nuclear genomes. Annals of the Missouri Botanical Garden 88: 104–134
Clausing G. S. S. Renner 2001a Molecular phylogenetics of Melastomataceae and Memecylaceae: implications for character evolution. American Journal of Botany 88: 486-498
———, and ———. 2001b Evolution of growth form in epiphytic Dissochaeteae (Melastomataceae). Organisms, Diversity, and Evolution 1: 45-60
Cogniaux C. A. 1891 Melastomaceae (Monographiae Phanerogamarum). In A. and C. de Candolle [eds.], Monographiae Phanerogamarum, vol. 7, 1–1256
Collinson M. E. M. Pingen 1992 Seeds of the Melastomataceae from the Miocene of Central Europe. In J. Kovar-Eder [ed.], Palaeovegetational development in Europe, 129–139. Museum of Natural History, Vienna, Austria
Dorofeev P. I. 1960 On the Tertiary flora of Belorussia. Botanichesky Zhurnal 45: 1418-1434 (in Russian)
———. 1963 The Tertiary floras of western Siberia. Izdatelstvo Akademii Nauka SSSR, Moskva-Leningrad, Russia
———. 1988 Miocene floras of the Tambov district. Akademii Nauka, Leningrad, Russia [posthumous work ed. by F. Y. Velichkevich, in Russian]
Doyle J. A. M. J. Donoghue 1993 Phylogenies and angiosperm diversification. Paleobiology 19: 141-167[Abstract]
Duarte L. 1956 Melastomatáceas fósseis da Basia Tertiária de Fonseca, Minas Gerais. Divisio de Geologia e Minerologia Boletim 161: 8-32
Dusén P. 1908 Über die tertiäre Flora der Seymour-Insel. In Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition 1901–1903, vol. 3(3), 1–27. Lithographisches Institut, Stockholm, Sweden
Dyjor S. Z. Kvacek M. Lanucka-Srodoniowa W. Pyszynski A. Sadowska E. Zasrawniak 1992 The Younger Tertiary deposits in the Gozdnica region (SW Poland) in the light of recent palaeobotanical research. Polish Botanical Studies 3: 1-129
Fairon-Demaret M. 1994 [1996] Les fruits et graines du Miocene de Bioul (Entre-Samre-et-Meuse, Belgique). Etude qualitative, quantitative et considerations paleoecologiques. Annales de la Societé Geologique Belgique 117: 277-309
Gentry A. H. 1982 Neotropical floristic diversity: phytogeographical connections between Central and South America, Pleistocene climatic fluctuations, or an accident of the Andean orogeny?. Annals of the Missouri Botanical Garden 69: 557-593[CrossRef][Web of Science]
———. 1993 Diversity and floristic composition of lowland tropical forest in Africa and South America. In P. Goldblatt [ed.], Biological relationships between Africa and South America, 500–547. Yale University Press, New Haven, Connecticut, USA
Gottwald H. 1992 Hölzer aus marinen Sanden des Oberen Eozän von Helmstedt (Niedersachsen). Palaeontographica Abt. B 225: 27-103
Graham A. 1993 History of the vegetation: Cretaceous (Maastrichtian)-Tertiary. Flora of North America, vol. 1, 57–70. Oxford University Press, New York, New York, USA
Hall R. 1996 Reconstructing cenozoic SE Asia. Tectonic evolution of Southeast Asia. Geological Society Special Publications 106: 153-184
Hickey L. J. 1977 Stratigraphy and paleobotany of the Golden Valley formation (Early Tertiary) of western North Dakota. Geological Society of America, Memoir 150. Geological Society of America, Boulder, Colorado, USA
Huertas G. 1977 Una Melastomatácea fósil del Terciario carboneifero de Antioquia (Eoceno). Caldasia 12: 35-39
Jacques-Félix H. 1978a Les subdivisions du genre Memecylon (Melastomataceae) en Afrique. Adansonia 17: 415-424[Web of Science]
———. 1978b Les genres de Memecyleae (Melastomataceae) en Afrique, Madagascar et Mascareignes. Adansonia 18: 221-235
———. 1994 [1995] Histoire des Melastomataceae d'Afrique. Bulletin du Museum National d’ Histoire Naturelle, Paris, 4e sér., 16, sect. B, Adansonia 2–4: 235–311
Kräusel R. 1929 Fossile Pflanzen aus dem Tertiär von Süd-Sumatra. Verhandlingen der Geologie en Mijnbouw Genootschap voor Nederland en Kolonien, Geologie Serie, S'Gravenhage
Kubitzki K. W. Krutzsch 1996 Origins of East and South Asian plant diversity. In Z. Aoluo and W. Sugong [eds.], Floristic characteristics and diversity of East Asian plants, 65–70. Springer, Berlin, Germany
Lavin M. M. Luckow 1993 Origins and relationships of tropical North America in the context of the boreotropics hypothesis. American Journal of Botany 80: 1-14
Lockhart P. J. M. A. Steel M. D. Hendy D. Penny 1994 Recovering evolutionary tress under a more realistic method of sequence evolution. Molecular Biology and Evolution 11: 605-612[Web of Science]
Mai D. H. 1995 Tertiäre Vegetationsgeschichte Europas. G. Fischer, Jena, Germany
———. 2000 Die untermiozänen Floren aus der Spremberger Folge und dem II. Flözhorizont der Lausitz. Teil III. Dialypetalae und Sympetalae. Palaeontographica, B 253: 1-106
Manchester S. R. 1999 Biogeographical relationships of North American Tertiary floras. Annals of the Missouri Botanical Garden 86: 472-522[CrossRef][Web of Science]
Meyer K. In press Revision of the Southeast Asian genus Melastoma (Melastomataceae). Blumea 0: 000-000
Olmstead R. G. J. A. Sweere 1994 Combining data in phylogenetic systematics: an empirical approach using three molecular data sets in the Solanaceae. Systematic Biology 43: 467-481[CrossRef][Web of Science]
Perrier de la Bâthie H. 1951 Mélastomatacées. In H. Humbert [ed.], Flore de Madagascar et des Comores, 1–326. Firmin-Didot, Paris, France
Raven P. H. D. I. Axelrod 1974 Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden 61: 39-637
Renner S. S. 1989 Systematic studies in the Melastomataceae: Bellucia, Loreya, and Macairea. Memoirs of the New York Botanical Garden 50: 1-112
———. 1993 Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic Journal of Botany 13: 519-540[Web of Science]
———. 1994 A revision of Pterolepis (Melastomataceae: Melastomeae). Nordic Journal of Botany 14: 73-104[Web of Science]
———,G. Clausing N. Cellinese K. Meyer In press Melastomataceae. In K. Larsen and I. Nielsen [eds.], Flora of Thailand. The Forest Herbarium, Royal Forest Department, Bangkok, Thailand
———, and K. Meyer In press Melastomeae come full circle: biogeographic reconstruction and molecular clock dating. Evolution 55:
Romero E. J. 1993 South American paleofloras. In P. Goldblatt [ed.], Biological relationships between Africa and South America, 62–85. Yale University Press, New Haven, Connecticut, USA
Rzhetsky A. M. Nei 1992a A simple method for estimating and testing minimum-evolution trees. Molecular Biology and Evolution 9: 945-967[Web of Science]
———, and ———. 1992b Statistical properties of the ordinary least-squares, generalized least-squares, and minimum-evolution methods of phylogenetic inferences. Journal of Molecular Evolution 35: 367-375[CrossRef][Web of Science][Medline]
Sanderson M. J. 1998 Estimating rate and time in molecular phylogenies: beyond the molecular clock?. In D. E. Soltis, P. S. Soltis, and J. J. Doyle [eds.], Molecular systematics of plants, 2nd ed., 242–264. Kluwer, Boston, Massachusetts, USA
Smith A. C. 1973 Angiosperm evolution and the relationship of the floras of Africa and America. In B. J. Meggers and W. D. Duckworth [eds.], Tropical forest ecosystems in Africa and South America: a comparative review, 49–61. Smithsonian Institution Press, Washington, D.C., USA
Sullivan J. D. L. Swofford G. J. P. Naylor 1999 The effect of taxon sampling on estimating rate heterogeneity parameters of maximum-likelihood models. Molecular Biology and Evolution 16: 1347-1356[Web of Science]
Swofford D. L. 2000 PAUP*. Phylogenetic analysis using parsimony (*and other methods), version 4. Sinauer, Sunderland, Massachusetts, USA
Thorne R. 1973 Floristic relationships between tropical Africa and tropical America. In B. J. Meggers and W. D. Duckworth [eds.], Tropical forest ecosystems in Africa and South America: a comparative review, 27–47. Smithsonian Institution Press, Washington, D.C., USA
Tiffney B. H. 2000 Geographic and climatic influences on the Cretaceous and Tertiary history of Euramerican floristic similarity. Acta Universitatis Carolinae-Geologica, Biologica, Prague 44: 5-16
Wehr W. C. D. Q. Hopkins 1994 The Eocene orchards and gardens of Republic, Washington. Washington Geology 22: 27-34
Whiffin T. A. S. Tomb 1972 The systematic significance of seed morphology in the neotropical capsular-fruited Melastomataceae. American Journal of Botany 59: 411-422[CrossRef][Web of Science]
Yang Z. 1994 Maximum likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods. Journal of Molecular Evolution 39: 306-314[CrossRef][Web of Science][Medline]
———, and S. Kumar 1996 Approximate methods for estimating the pattern of nucleotide substitution and the variation of substitution rates among sites. Molecular Biology and Evolution 13: 650-659[Abstract]
Zhu H. 1997 Ecological and biogeographical studies on the tropical rain forest of south Yunnan, SW China with a special reference to its relation with rain forests of tropical Asia. Journal of Biogeography 24: 647-662[Web of Science]
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