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TREE vol. 5, no. 7, January

1990

Monospecific Dominance in Tropical Forests Rain
TereseB. Hart
Old-growth rain forests that are dominated monodominant rain forests are genby a single canopy speciesoccur throughout erally, though not always, members the tropics, though they account for a lim- of the legume family Caesalited proportion of the total ruin forest urea. piniaceae’ . Mora excelsa, also of These forests have been considered anom- the Caesalpiniaceae, forms monoalies in which development of a more diverse dominant forests in parts of eastern community is deflected by harsh conditions. South America and Trinidad. In Asia, Very poor soils or un otherwise extreme members of the Dipterocarpaceae, environment mug promote monodomi- Lauraceae and Guttiferae all occur nunce by excluding potentially competing as the dominant species of large stands2m5. In these species, 6ut it is now apparent that mono- monodominant dominant tropical forests also develop under forests the dominant species is more benign conditions. Field studies have present in all size classes and the shown that a single species may dominate forests have been considered cliof maintaining on undisturbed sites where the soils are max (i.e. capable similar to those of adjacent old-growth, approximately the same species mixed forests. In these situations the domi- composition in the absence of nant is a superior competitor and/or is severe disturbances). Dominance, by definition, affects particularly tolerant to stresses such us shade. Assertion of dominance 6g a single the evenness or relative represenspecies in an old-growth forest appears most tation of species. Therefore, it local species diversity, likely in areas where the species pool con- affects tains few late-succession species with simi- although total species number may not be altered over large areas. lar life history traits. Early studies from all three tropical continents reported monodominant forests having the same overall Tropical rain forests (Box I I are species composition as adjacent generally more diverse in species old-growth mixed forests5. The canthan temperate forests2,3. Richards, opy species associated with the dominant were all characteristic of in his early and extensive disthe neighboring forests and the cussion of this phenomenon4, also commented on the apparently species of these latter forests did occurrence not appear to be excluded from the anomalous of oldmonodominant stands. As a single growth tropical rain forests domispecies assumes an ever greater nated by a single canopy tree percentage of the basal area, howspecies. Such forests, he noted, ocever, other species lose represencurred on well-drained terrain and tation in the canopy and rare often abutted more diverse forests. species become rarer. The result is Richards described in detail only relatively small discrete stands of that any small plot in monodomiforest dominated by a single nant forest will contain fewer species, but he called attention to species than a plot of the same size the existence of more extensive forin mixed forest (Fig. I I. Monodominant forests have remations in various tropical regions. renewed attention during Examples of forested areas where a ceived the last decade, and our undersingle canopy tree species contribstanding of the possible causes of utes more than 80% to the total monodominance has advanced conbasal area (hereafter, monodomisiderably. Here, I review factors renant forests), and that are reputedly ported to control species diversity old-growth stands on well-drained in tropical rain forests on wellsoil, occur throughout the tropics (Table 1 )5,6 over areas of hundreds drained soils, and summarize of square kilometers. See Box 2 for examples from extra-tropical and an example. waterlogged forests that can conIn Africa, the dominant species of tribute to our understanding of monodominance. I also discuss how may be related to secTerese Hart is at Wildlife Conservation Inter- dominance succession in national, The New York Zoological Society, Bronx, ondary tropical NY 10460,USA. forests. 6

Abiotic factors It seems reasonable to expect that environmental factors thought to promote high species diversity in tropical forests might also control dominance by a single species. In some tropical areas, patterns of forest diversity were interpreted as showing that increased rainfall3 or soil moisture9 promote increased species richness. In an Asian study, floristic patterns were construed as showing a direct association between levels of specific soil nutrients and overall floristic richnessiO. The occurrence, in some cases, of species-poor tropical forests on particularly soils impoverished suggested that extremely low fertility could lead to low diversity”. In most tropical forest studies, however, correlations between particular species associations and soil factors have given only equivocal results2, and the high diversity of tropical forests cannot be related directly to edaphic diversity12. In certain instances, a single species may form monodominant stands over a wide variety of soil types5. Most of the large monodominant formations listed in Table 1, as well as some smaller stands, coexist with adjacent, more diverse mixed formations across similar soil conditions5,‘ 3. Biotic factors In tropical forests with high species diversity, there is some evidence for overall enhanced growth and recruitment of rare tree species relative to common species14. The in‘ compensatory mechanisms’ volved might include concentration of natural enemies on common species, intraspecific competition for resources, or interference between abundant conspecifics as, for instance, through allelopathy. Differential mortality of seeds and seedlings such that the greatest loss occurs near the parent trees has been shown to occur in at least some tropical forest species15,16. Such mortality, whether provoked by predation or other causes, could promote a more even distribution of tree species and hence greater species diversity. Treefalls have been attributed an important role in maintaining tropical forest diversity, in that the gaps they create present a rich and varied resource for tree germination

no. can confer advantage to a particular species by excluding other species. In Central America some freshwater swamps are nearly pure stands of a single species. 5. and Avicennia germinans does so where winter temperatures are particularly low26. In the La Selva forest (Costa Rica) there is a dramatic reduction in stem density and species richness with increasingly poor drainage. Soil salinity. there is a clear succession and one species eventually asserts dominance2*. however. if treetails created a level of heterogeneity that guaranteed not only the izermination and establishment. Some of the better-known tropical and subtropical monodominant forests occur on waterlogged or periodically flooded substrates.:ation of tropical tree species to this -esource gradient may maintain an elevated level of diversity. such as tolerance to full sunlight. Sumatra Borneo. fire-resistant species can maintain nearly single-species canopies in the presence of recurring fires20. or unable to compete under. the severe conditions. 8. additional environmental stresses compound the effect of waterlogging.g. Table 1. The result is a successional sequence of species as the relative competitive values of these traits change27. Where fire is long absent. The frequency. slow-growing shade- of timber Lauraceae Caesalpiniaceae Caesalpiniaceae Caesalpiniaceae ‘ Pure stands’ 85-95% canopy of the >75% of the canopy >90% of the canopy Gilbertiodendron dewevrei from Ref. such as shade or a build-up of leachates. or improves its ability to withstand stresses of conspecific origin. In mangrove forests. are often the inverse of traits favored in late succession. I. January 7990 #and growth17. as in the case of secondary succession. Prioria copaifera or Campnosperma panamensis23. In many ecosystems it appears that fire is important in determining species composition. in California chaparral communities the recurrence of fire preserves diversity. In the old-growth tropical monodominant stands where the canopy ‘ dominant is abundantly represen:ed in the understory. Absences from the forest canopy may be temporary. would promote monodominance. e. mono‘ dominance appears to be selfoerpetuating. Sumatra Trinidad East Africa Central Africa Adapted. dominance may be suppressed. therefore. the equalizing effect of seed predation may be suppressed in monodominant stands through predator satiation during synchronous fruit production’ 9. tidal flows and erosional species recurrent fires that continue episodes reduce growth rates and eliminate species intolerant of. tolerant are suppressed by to favor dominance by the shade-intolerant Nothofagus cunninghamii2’ (which in some cases comprises 95% of the basal area). In the pine flatwoods of Florida. rapid growth and efficient dispersion of small seeds. Fires may both promote and prevent dominance by a single species. that flooding. It may be that once nonodominance is well established. Dominant species Dryobalanops aromatica Eusideroxylon zwageri Mora excelsa Cynometra alexandri Family Dipterocarpaceae Description dominance 60-90% trees of Extra-tropical and waterlogged sites of examples of Examination single-species dominance in subtropical communities or in forests on poorly drained substrates may clarify the processes leading to dominance (Table 2). On the other hand. for example. When waterlogged forests are drained. 5. of a wide variety of tree . like fire. Old-growth monodominant tropical forests of well-drained soils that occur in stands of more than 100 km* Location Malaya. It has been suggested that tropical tree species that form species-specific ectomycorrhizal associations may have an enhanced ability to form monodominant stands that will be self-replacing and able to invade neighboring mixed forest$. Rhizophora mang/e forms self-replacing monocultures where tidal energy is high or ocean water deep.~lso the continued growth into the canopy. but . with permission. 7 . compensatory mechanisms 3re no longer effective in generating greater evenness. Ectomycorrhizae are common within at least two of the families containing species that achieve monodominance in tropical rain forest environments: the Dipterocarpaceae6 and the Caesalpiniaceae5. Monodominanceand succession The perturbations leading to dominance in a community need not be so stressful or continuous as to eliminate species permanently. depth and duration of flooding seem to be determinants of species composition and dominance. The differential adap. such as tolerance to shade and production of few seeds with large energy reserves. For example. hardwood species have invaded drained swamps and cypress has lost its dominance25. In Tasmanian temperate rain forests. The life history traits favored early in succession.‘ Any biotic factor that promotes the interspecific competitive advantage of the dominant tree species. This could only be true. this is a result of the exclusion of species intolerant of waterlogged soi124. in the cypress stands of Florida.. It seems.pecies.‘ TREE vol.

the slow-growing shade-tolerant will dominate an inspecies creasingly large proportion of the canopy. dewevrei as percentage of total stems 28. dewevrei. does not reproduce in its own shade and monodominance lasts only a single generation. Seedlings may persist for years in the shaded understory with little further growth. The best competitor. Wellknown forest communities that fit this category include monodominant stands of Musanga cecropioides or Trema guineensis in Africa and stands of Octomeles sumatranus Asia’ . if a forest remains undisturbed. a long-lived species believed to have grown up after major wind destruction in 18802. for instance. a single tree species can become established and dominate the canopy of rain forest more than a century later.5 92. dewevrei in the understory is evident in the profile diagram of a 100 m x 10 m plot near Yangambi. will steadily increase in abundances’ Accordingly. adapted from Ref.5 cm. locally known as mbau forest. Hurricane Joan of 1988. Nothofagus obfiqua dominates a Chilean forest but does not regenerate in its own shade: it has been suggested that the individuals now forming the canopy became established subsequent to a disturbance approximately 250 years ago30. such as Cecropia obtusifolia and Ochroma lagopus29. 1. These examples illustrate that. The first seedling leaves rapidly become coriaceous and resistant to herbivory. Lang and Knight32 describe a section of the forest on Barro Colorado Island (Panama) that exemplifies some of these mid-successional characteristics. According to the ‘ intermediate disturbance’ hypothesis3’ . At 60 years of age. these individuals are potentially self-replacing. Large-scale disturbances would have to occur only once every several centuries to reinitiate or rejuvenate a successional sequence in a tropical forest.1 72. diameter at 1. Large-scale storms occur regularly within the cyclone belt between IO” and 20” north and south of the equator34. covers hundreds of square kilometers in northeastern Zaire.6 16. Monodominance is common during early successions in tropical environments28. rapidly growing species. They drop beneath the parent and sprout within 3-5 days. For example. dewevrei stems per ha 8000 307 138 62 51 Mean total no. Size class? Seedlings Saplings Poles Trees Mean no. covered by only a thin seed coat. may have destroyed 7% . disturbances occurring at shorter intervals than the life span of the longest-lived trees in the systems2’ would have to be typical throughout much of the tropical rain forest zone. given fortuitous regeneration conditions following a major disturbance. or the species most resistant to prevailing stresses. shade-tolerant seedlings33 (Table 2). no. 7. in having been the first species to establish. Most tropical forests characterized by high species diversity may be in some middle stage of succession. in each case. For this to be the case. Able to persist in their own shade. dewevrei in the understory indicates that mbau forest is an old-growth forest in which canopy composition is stable. 5. The representation of G. Sapling data collected from 12 plots of 225 m* and seedling data from 12 plots of 25 m2. of dispersal is sacrificed in favor of large. producing first-year seedlings that are 0. The canopy dominant. of stems per ha 28 000 1911 191 77 55 G. heavy and often poorly dispersed seeds that produce robust. 45% of the basal area was accounted for by four species. the greater diversity of this stage derives from the persistence of mature individuals of early successional species as later successional species grow into the canopy. saplings: dbh > 2. Similar in species occur in tropical America. Data from Ref. 40 r -30 E -20 g 2 -10 -0 The homogeneity of the canopy contributes to the shade of the understory. of G. In mid succession. where the shade tolerance of the dominant species permits it to persist in all subcanopy size classes. three of which were clearly early successional. highly vagile. Zaire. as shown in the table below. are not dispersed by animals.4. Likewise. Exampleof an old-growthmonodominant tropical rain forest The monodominant rain forest dominated by Gilbertiodendron dewevrei.3 80. 5.5 cm and 5 10 Species replacement occurs relatively rapidly in early succession and there is a distinct advantage in being the first species to become established.TREE vol. Ten years later the same species were still present in the canopy although their relative representation had changed as individuals died and were replaced by more shadetolerant species.45 m high. January 1990 Box 2. Some monodominant forests do not have a history of recent or recurrent disturbance and yet the dominant species is characterized by certain early successional traits. The abundant representation of G. Dominance is likely to result from the fortuitous and massive establishment of a single. The large heavy seeds of G.X-0. efficiency .7 Large trees Data collected in the lturi Forest of Zaire from 24 plots of 625 mz each for all stems > 10 cm dbh. at this stage.3 m (dbhl 4 2. monodominance is less likely because the slower growth rates prolong the period of coexistence of numerous species27. dewevrei. There is little advantage. all darkened crowns are G. “Seedlings defined as height > 0.5 m. Diverse observations from various tropical regions suggest that such disasters may take place with this frequency and across significant areas of rain forest. the ‘ storm forest of Kelantan’ is dominated by Shorea parvifIora. in Asia.

Chile Past major disturbance Shade-intolerant 21. pers.30 tolerant to fire and 25 Southeast USA Flooding and fire Shade-intolerant. well-drained 4-6 9 . although fires might have escaped into uncut forest during drier periods. Similar extensive charcoal horizons have also been found under old-growth Africa5. may be more frequent in continental tropical forests than is commonly recognized. It also appears that large tracts of forest previously thought to have been untouched by human populations may actually have undergone extensive anthropogenic alteration recently enough to be relevant to the current successional status of the forest communities.i a “J. January 1990 of Nicaragua’ s total forest cover. Putz. The size of the species pool is likely to contribute to dominance in old-growth forests.). variable tolerance to fire and flooding 20 Tropicalforests without pronounceddry season Swamp forests Storm forests Mangrove Evergreen forest rain Central and South America ‘ Hurricane Coasts Throughout tropics belt’ Flooding. In late succession the prolonged coexistence of species is related to the number of tree species present in the system with near-equivalent life history traitss7. Accordingly. Characteristics monodominant of forests in extra-tropical and wet tropical sites Forest type Extra-tropical forests Location Environmental to dominance factor pertinent Traits of dominant species Refs Nothofagus forest Cypress swamps Pine fiatwoods Tasmania. dispersal shade-tolerant. Physiognomically mature and floristically diverse forest in Venezuela occurs over buried charcoal horizons whose radiocarbon dates suggest that the forest has been subjected to two separate fires in the last 500 year+. Plot size Fig. Exceptional droughts can also give rise to major disturbance. I. flooding Florida. resulting from tornadoes. Under prevailing humid conditions large fires would have been likely to occur only as intentional phases of agricultural cycles. waterlogging Tolerant to stress 23 2 26 Past disturbance Disturbance. I. Major environmental changes affect the species pool by causing local extinctions of rainforest trees. Aerial photographs and satellite imagery revealed numerous tornado tracks through sparsely populated forests of Paraguay. southwestern Brazil and northeastern Argentina35. Extensive damage. not necessariiv the exclusion of particular mixed forest species from monodominant &nds. and (2) major environmental change that occurs so rarely or at such distant intervals as to be on a time-scale other than that of secondary succession. 5. The movement of tree species northward after the last glaciation has been documented by pollen cores.TREE vol. On islands covered with rain forest at the time of their formation. Replenishment of the species pool after suitable growing conditions are re-established depends upon species mobility. poor Variable. Species-area curves illustrate that small plots in diverse forest support more species than plots of equal size in adjacent monodominant forest. or the number of species interacting during succession. forests in central t History: disturbance and the speciespool There are two further interrelated factors of possible significance to monodominance in tropical forest. monodominance is favored on isolated islands where the species pool is poor due to the difficult dispersal of propagules across large bodies of water. as illustrated by the 1982-1983 fire that ravaged 30 000 km* of tropical lowland forest in East Kaiimantan (Indonesia). USA Fire and flooding Shade-intolerant. The monodominance of old-growth forests on the Hawaiian Islands may be partly explained in this mannerY8. commun. Range expansion varied between 100 and 400 m per Table 2. no. These are ( I ) the size of the species pool. This is the result of increased rarity of nondominant species. This appears to be occurring on the smaller f 2 . islands isolated by the formation of the Panama canal (F. and salinity stress of tides soils Shade-intolerant Tolerant to stress. rapid colonization and growth Slow growth. there is also a trend towards monodominance as species are lost through an increased extinction rate relative to the colonization rate.

C.613-633 6 Connell. M. the local forest canopies would continue to be composed of less shade-tolerant.C. short life span Frequent. 54. Tracey. where the impact of Pleistocene climate change on the composition of vegetation has been the subject of much recent research. Unescol AETFAT/UNSO 2 Whitmore. (1989) Am. 81-101 I3 Swaine..W.B. 355-363. D C. PG. II9841 Am. In Africa. I. few. J. pp.K.H. (19871 Annu. January 1990 -. 119791 Am. ed.A. Nat Iin press) 7 Louis. and Murphy. (19771 Ann. M. Biof. Cambridge University Press 5 Hart. I.B.H. Nat.1. Conclusion Dominance in early rainforest succession may be established rapidly by a single species subsequent to its fortuitous massive establishment.H. 775-788 20 Abrahamson. H. P. or by local extinctions that reduce the species pool. II981 1 Ecology 62. Table 3. D. I 1983) Vegetation Map of Africa of the Sahara (2nd edn). The presence of numerous species with similar life history traits mitigates against dominance by any single tree species. The number of co-occurring species may be reduced by environmental stresses that preclude survival of some species in a given system. 124.S.S. Nat. T. F. D.- year for common wind.(l9801 Ecofogybl. late-succession species continues to reflect the impact of local extinctions after populations of more vagile species have converged. Monodominance is least likely during the mid-succession period.694-705 II Ianzen. References I White. C. pp. more vagile species until they were invaded by slowdispersing. J. ( 19841 Ecology 65. The absence of such species delays progress towards monodominance. and Clark. D. Rev.C.43.or animaldispersed forest species of eastern North America39. (19741 Biotropica 6. large. 18.. 431-451 I8 Boucher. slowgrowing rainforest species often have relatively restricted distributions4*. 69-103 12 Huston. but the results may help to clarify the processes that have led to the variable diversity and dominance of tropical rain forests. The extensive preColumbian agriculture on forest lands may have drastically reduced the propagule sources of maturephase trees. Accordingly. (I981 1 in The Biological Aspects of Rare Plant Conservation (Synge. tolerates direct sunlight. tolerates shade. At a late stage of succession. local composition of the maturephase forest that re-established during wet periods would have been dependent on which tree species survived intervening dry periods in small pockets maintaining suitable climatic conditions. Human disturbance is not the only imaginable cause of local extinctions in the tropics. W. small. 1169-1184 9 Hall. as on other tropical continents. M.TREE vol. The methods for such studies are only now being worked out for moist tropical areas.A.4’ The . long life span Disturbance Species pool regime intermediate Many species with similar life history traits Infrequent. as has been previously proposed by the ‘ intermediate disturbance’ hypothesis12. and Swaine. (1947) in Comptes Rendus de la South 19471. In order to construct a complete context in which to examine the dynamics of a community of long-lived organisms. In temperate regions. M. Hart.H. it is likely that the monodominant stands they form will expand only in regions undisturbed over long periods. Semaine Agricole fevrier au 5 mars de Yangambi Idu 26 64. The tree species that form monodominant stands in mature tropical rain forest have heavier seeds and many are without animal dispersal agents. T. Bot. Presumably. 5. the expression of the slow expansion of such a species from remnant populations. 141-164 I5 Augspurger. there have been marked changes in the total extent of rain forest area over the last 15 000 years.. The extent of monodominance is.D. welldispersed seeds. (1982) Evol. small Variable. in 10 . 902-915. D.B. more shade-tolerant Whitmore species. I. 133. poorly dispersed seeds. Inst.31. Publ. L. &of. Monogr.H. suggested that Central America might be in a stage of arrested succession. reflect its dispersal potential relative to the time elapsed since re- Wiley & Sons I4 Connell. large Mid succession: even coexistence Wide spectrum represented Late succession: monodominance Slow growth rate. Syst. Because late-succession species often have localized distributions and slow invasion rates. and Lowman. The impoverished assemblage of poorly dispersed. 15.I (1984) &co/. I I9841 Tropical Rain Forests of the Far East (2nd edn 1. advantage to first species to establish Single competitively superior species ~_-_. sufficient historical information must be available to allow a perspective that encompasses events pertinent to succession within the community and movement of slowly dispersing organisms in and out of the community. monodominance is again possible if the life history traits of a single species render it the competitive dominant (see Table 31.B. These variations have been associated with wet and dry climatic cycles40.D. then the composition of old-growth forest in the area where it occurs is determined by its unique presence in the species pool. and Webb. numerous. Their range expansion would accordingly be slower. thus. no. A. I-84 4 Richards. J.C and Hartnett. Nat1 Etude Agron. The more shade-tolerant. the importance of propagule availability subsequent to climate shifts has been well documented39. C. If such a species is a self-replacing competitive dominant. Card. 409-4 I 4 I9 Augspurger. Junk IO Ashton. I. Am.D. Congo (Hors S&iel 8 Lorimer. 1.K. such as rainforest trees. II9521 The Tropical Rain Forest. j. Nat. I I?. Communitycharacteristicsof tropical rain forests at different stagesof succession Early succession: monodominance Life history traits of canopy species Rapid growth rate. I1981 1 Distribution and Ecology of Vascular Plants in a Tropical Rain Forest: Forest Vegetation in Ghana. MO. Subsequent to a regional extinction. ( I98 I t Oecologia 49. Clarendon Press 3 Gentry. The size of the area over which a species occurs and the extent of the stands that it forms may lease from conditions unfavorable for dense forest growth. welldispersed mid-successional species of similar life history traits remained in the region. 769-788 I7 Denslow. P. 1705-1712 I6 Clark. and Hall.

D. 29v627-640 26 Lugo. NY 11794. edsl.studies of the sort that have clar$ed analogous issues such as heterostyly. the relative order in which the correlated traits originated in a lineage is also important: DonoghueZO discusses this and other issues with particular reference to the correlation between dioecy and fleshy fruits.E. Future correlational studies nzust explicitly consider phylogeny.L. and Zimmerman. 6.S.g. and (2) the conformity of the proposed mechanisms to empirical data.H. 505-516 25 Marois.B. None of the other correlations discussed below have received this level of scrutiny. and Futyma. 639-655.C.D. and Zimmerman. 129-162.S.). II9791 1. M. the statistical significance of phylogenetic correlations is hard to assess because of nonindependence of taxa within lineages. (19881 Photogramm. and Schlegel. Muenchow” and Charlesworth’ discuss this general statistical problem with particular respect to dioecy arguments. the woody habit. T. ( 1980) Biotropica (SuppI. R. pp. Blackwell Scientific Publications 23 Hartshorn. 303-306 38 Mueller-Dombois. T. ed. (1988) 1.D. (19801 Biotropica I2 (SuppI. I. When that change is one that Darwin confessed to finding ‘ much difficulty in understanding”. Pagel and HarveyI review several recent proposals for solving this difficulty. 558-584 22 Walker. F. 93-103 Over the last decade.. Clark.S. D. (1978) Science 199. 53-55 37 Fagerstr6m. but most have been described only recently. G. Shugart. M.. a correlation and a mechanism. pp. K.. 34-39 12 29 Hartshorn. Academic Press 42 Ewel. edsl. although a full interpretation of the correlations must include genetic constraints. Evol. P. We consider several pairs of correlations and mechanisms. assessing both the validity of the correlation and the adequacy of the mechanism. such broad studies should also 6e supplemented by detailed studies of particular lransitions to dioecy (e. K. covariation among the ‘ driving’ characters was usually ignored.P. The mechanisms typically invoke ecological circumstances that favor dioecy in ways specified by theoretical models of sexual selection and sex allocation theory. pp. and Ewel. We do not dwell on ‘ pathways to dioecy’ or on population genetic models12-16. Some of these correlations date to Darwin.S. Ideally. M. Sadarriaga. Nat.l. is properly counted as only one datum for a correlation analysis. Starting about ten years ago. however. M.C. F. a number of alternative hypotheses have been presented5-‘ I. M.H. 365-390. For testing a particular mechanism. ( I9801 Biotropica I2 (Suppl. Academic Press (in press) 21 Read. J. not five. 65-72 27 Huston. Hypotheses the Evolution for of Dioecy Seed in Plants JamesD. E. A. 2. State University of New York. Remote Sensing 54. we discuss the current status of these proposals and the progress made over the last decade. (19821 in The Plant Community as a Working Mechanism (Newman. 1302-1310 32 Lang. 73. 329-340 31 Connell. Most of the selective mechanisms invoked have been suggested and supported by phylogetietic correlations. D. 27-43.H. (1983) Forest Sci.). Here we review ( I) the validity of the correlations (especially in light of recent critiques of the comparative method]. eds). 76. Woods. ( 198 I ) in forest Succession: Concepts and Application (West.T.B.TREE vol.. The evolution of dioecy from hermaphrodite ancestors is such a problem.. no. ( 1978) in Tropical Trees as Living Systems (Tomlinson. W. and Peralta.C.. S.. more importantly. 168-198 28 Kellman. Webb. and Herrera. because most authors were typically concerned with only a single relationship. R. R. 119871 Am.L.P. 1429-1435 36 Sanford. all dioecious and all woody. 00 .B. but the strength of their support varies. D.L. An obvious consequence of separate sexes is the impossibility of selfing.I.M. and Hill. (19881 in North American Terrestrial Vegetation (Barbour. and Billings.C. eds). Thus. Cambridge University Press 24 Lieberman. II9801 Biotropica I2 ISuppl. M.B. January 1990 Ecosystems of Florida (Myers.S. The correlations are phylogenetic: dioecious taxa are more likely to exhibit fleshy ‘ fruits’ (sensu late). and Botkin. ( I9801 faleobiology 6. G. 5. R. Biogeogr. D.H. First. ( 1987) Trends Ecol. G.. A. (19851 Science 227. pp. (19781 in Tropical Trees as Living Systems ITomlinson.. Ecol. Uhl. G. I. 1075-1080 33 Ng.). ( 19861 Vegetatio 67.). Evol. Importantly. ( 19881 Trends Eco/. Here. Cambridge University Press 34 Whitmore. D.Thomsonand JohanneBrunet by Darwin’ ). 1. 130. M. Eng..D. D.A.USA (C 1990 Elsev~er Scercp Publishers L k? IlJKi 0169~5347. Ecol. Stony Brook. and Knight. M. I. or heterostyly. 1. 243-244 41 Hamilton.90:$02 General difficulties with correlations Recent critiques of the comparative method have established standards of rigor matched by few of the early papers concerning dioecy17. Ashton. pp. and Smith. 132-l 53.B.. T. Cambridge University Press 35 Dyer.H. R. Thus a genus containing five species. testing for correlation of traits requires mapping those traits 11 Any evolutionary change that has occurred independently in numerous lineages is likely to offer insight into fundamental mechanisms of evolution. (19821 Environmental History of East Africa: A Study of the Quaternary. the ‘ sample size’ should reflect the number of evolutionary events rather than taxa. 216-220 39 Davis. Hartshorn. 3. 23-30 30 Veblen. Springer-Verlag 40 Livingstone. D. C. with the risk that spurious or indirect correlations might have suggested incorrect causal relations. the avoidance of inbreeding has traditionally been invoked as an important selective force in the evolution of dioecy2-4 (despite an early caution lames Thomson and lohanne Brunet are at the Dept of Ecology and Evolution. within genera) . These hypotheses usually have two parts. None of the hypotheses can be flatly rejected on existing evidence. T. small and inconspicuous flowers.18.G. R. Lieberman. ( 1985) /. C.H. eds). 2-7 43 Davis. H..E. island habitats. P. it gains the stature of a central problem in biology.I. Second.. (19831 Ecology 64.E.. K. and Ewe\. new hypotheses have been proposed for the evolution of dioecy iM plants. K.. Ir.

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