Status of Miconia Calvescens (Melastomataceae), a Dominant Invasive Tree in the Society Islands (French Polynesia)!

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Status of Miconia calvescens (Melastomataceae), a Dominant Invasive Tree in the Society Islands (French Polynesia)!

2 JEAN-YVES MEYER ,3

ABSTRACT: Since its introduction to Tahiti in 1937 as an ornamental, Mico nia calvescens DC. (Melastomataceae) has become the most important plant pest in the Society Islands. Its ecological characteristics allow it to thrive and spread in a wide range ofhabitats, including native forest, where it forms dense monotypic stands. Miconia calvescens now dominates over two-thirds of Tahiti and has spread to the surrounding islands of Moorea and Raiatea. This species represents an immediate threat to the native flora of all the high islands of French Polynesia and a potential danger to many tropical oceanic islands.

NUMEROUS CASES OF biological invasions have tribution, and effects of M. calvescens in the been described in native vegetation of oce Society Islands, as well as some of the bio anic islands, including the Hawaiian Islands logical characteristics that contribute to the (Smith 1985, Stone et al. 1992), Galapagos success of this invasive tree and make it a Islands (Schofield 1989), Guam (Lee 1974), potential danger to the forest of many moist, Mauritius (Lorence and Sussman 1986), and tropical oceanic islands. La Reunion (Macdonald et al. 1991). High susceptibility ofisland ecosystems to invasion Study Area may be largely the result of relative impover ishment of flora and fauna (low species num The Society Islands, the largest group of ber, taxonomic disharmony) and of evolu high islands in French Polynesia, form one of tion in long isolation from outside influences the most isolated archipelagoes in the world, (Loope and Mueller-Dombois 1989). Another lying at 15-18°S, 148-154°W in the South factor in many instances is that aggressive Pacific Ocean, 5000-6000 km from the near invaders have been introduced without the est continents. They include nine high islands specific natural predators that have evolved (Bora Bora, Huahine, Maiao, Maupiti, Me in their native range. hetia, Moorea, Raiatea, Tahaa, and Tahiti) The case of Miconia calvescens De Can of volcanic origin, less than 4.5 myr old, and dolle in Tahiti provides an extremely dra five coral atolls (Manuae, Maupihaa, Motu matic example of the effect of a single in One, Tetiaroa, and Tupai). The archipelago vasive species on the biological diversity of is divided into the Windward Group and the an oceanic island. Introduced to Tahiti as a Leeward Group based on the position of the garden ornamental in 1937, it has spread to islands relative to the dominant southeast form monotypic stands, replacing former na trade winds (Figure 1). The climate is trop- tive forest, and now dominates the forest ical oceanic with two seasons: a warm and over 65% (ca: qO,OOO ha) ofthe island~ - ---- humid-season with torrential-rains from De This paper is an attempt to synthesize what cember to February and a cooler, drier sea- is known about the introduction, spread, dis- son from March to November; mean annual temperature is 26°C (absolute maximum 34°C in January, absolute minimum 15°C in Au 1 Manuscript accepted 27 March 1995. gust); relative humidity ranges from 76 to 2 Laboratoire d'Ecologie Vegetale, Centre ORSTOM de Tahiti, B.P. 529, Papeete, Tahiti. 80%; annual rainfall averages 1700 mm/yr 3 Current address: Delegation al'Environnement, RP. at sea level (pasturel 1993). Precipitation in 4562, Papeete, Tahiti, French Polynesia. creases rapidly orographically more on wind- 66 Status of Miconia calvescens in the Society Islands-MEYER 67

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FIGURE I. Distribution of Miconia calvescens in the Society Islands of French Polynesia (invaded islands are shown in black), ward sides (sometimes up to 10,000 mmjyr) Merril (Convolvulaceae), Mimosa invisa Mar than on leeward sides. tius ex Colla (Fabaceae), Psidium guajava L. Floristic richness of islands in the tropical and Psidium cattleianum Sabine (Myrtaceae), Pacific depends on their physiographic and and Rubus rosifolius Sm. (Rosaceae) have ecological diversity. The Society Islands, with become locally dominant in secondary vege their high, complex islands, have 623 native tation and are considered as "plant pests." species (including 273 endemics) of the 959 Miconia calvescens, although also a pest in vascular plant species native to French Poly disturbed sites, is unique in its extension in nesia (Florence 1987). Colonizing Polynesians apparently undisturbed native ecosystems and brought with them, 2000-2500 yr ago, an ad its efficiency in eliminating other species. ditional 80 plant species (1. Florence, pers. comm.) and several animal species. Over 1500 Description ofthe Study Species introduced plant species have been collected by J. Florence in the Society· Islands (pers. In the Society Islands, Miconia calvescens comm.). They were introduced either inten (Melastomataceae: Miconieae) is a small tree tionally (for agricultural, construction, medic up to 15 m high. Most often, mature trees are inal, or ornamental use) or inadvertently since between 6 and 12 m tall, with slender, stiffver European contact in the eighteenth century. tical stems. Because of its large, dark green, Many ofthem are naturalized, and some, such handsome leaves (up to 1 m long), which as Cyperus rotundus L. (Cyperaceae), Lantana have three prominent, pale green nerves above camara L. (Verbenaceae), Melinis minutiflora and are more or less purple-blue underneath P. Beauv. (poaceae), Merremia peltata (L.) (Figure 2), M. calvescens (also called M 68 PACIFIC SCIENCE, Volume 50, January 1996

Natural Distribution To check the native range of M calves cens, I observed the specimens available in some major herbaria (Royal Botanical Gar den of Kew [K], Museum national d'Histoire naturelle de Paris [p], Institut fUr Botanik der Universitat Wien [wu], U.S. National Arboretum of Washington [NA], Bishop Mu seum of Honolulu [BISH]). I collected other information from V. Sosa (Instituto de Eco logia, A. c., Mexico), M. G. Peiia (Herbario Nacional de Mexico, Mexico), S. C. Chiea (Instituto de Botanica, Brazil), F. Almeda (California Academy of Sciences, USA), and R. Burkhart (State Department of Agricul ture, Hawai'i, USA). The identification of most herbarium specimens was checked by J. J. Wurdack. The native range of M calvescens extends over more than 40° of latitude, from about 18°N in southern Mexico to about 26°S in southern Brazil and northern Argentina (Fig ure 3). Specimens ofthe bicolorous form with purple leaf undersides have only been col lected in southern Mexico (Chiapas, Oaxaca), northern Guatemala, Belize, and Costa Rica. FIGURE 2. Because ofits large, attractive leaves, more Information on herbarium labels suggests or less purple underneath, Miconia calvescens is culti that the species is found from lowland to vated in European greenhouses and tropical botanic gar montane tropical forest (up to 1800 m in dens and is still considered one of the most magnificent foliage plants (Graf 1986). Ecuador [Wurdack 1980]), sometimes in dense shade of primary forest, but most often in more open vegetation (old pastures, forest magnifica Triana in horticulture) was intro edges, river banks, trailsides and roadsides, duced to European botanical gardens and disturbed areas). Like most of the 1000 other cultivated in greenhouses for its ornamental species of the genus Miconia, M. calvescens value (Wurdack 1971) . It was then consid seems to be a shade-tolerant understory tree ered as "one of the best and most striking that behaves like a pioneer tree in forest gaps. of all conservatory foliage subjects" (Bailey In its native range, this species "is never 1900: 1012). common at anyone site" (F. Almeda, pers. The leaves are opposite, elliptic to obovate, comm.). entire or undulate, rounded or subcordate at the base; shortly acuminate or sometimes ob Ecology and Dispersal tuse to rounded at the apex with a 4- to 10 em-long petiole, glabrous above, and puber Many of the biological characteristics of ulous with minute stellate hairs underneath. M. calvescens make it a potential pest outside Flowers are small, white, more or less sessile, its native range. It is a fast-growing tn::e (up bisexual, 5-merous, and slightly fragrant, to 1 mjyr for juvenile plants under optimum arranged in large panicles. Younger branches conditions [Meyer 1994]) that can flower and and base of the panicle are also thinly cine fruit after 4-5 yr from seed. There are at least reous with minute stellate hairs. three peaks of flowering and fruiting per year 60 40 .... / ...... ~~~~:~lcL I

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FIGURE 3. Native range of Miconia calvescens according to herbarium specimens. Bicolorous form is indicated by closed circles and form with green leaves by open circles. 70 PACIFIC SCIENCE, Volume 50, January 1996 within populations (Meyer 1994), but with terrestrial bird, occupying all ecological hab some inter- and intraindividual variability itats on the island. The Red-vented Bulbul, (Gaubert 1992). Panicles of fleshy berries (up first noticed in Tahiti in 1979, has become to 500 fruits per panicle) are produced, with abundant in the coastal and lowland zones. each fruit 0.6-0.7 cm in diameter, blue-black Small rodents may play an important role in when ripe, containing an average of 140-230 Miconia seed dispersal: I have found numer seeds, each ca. 0.7 by 0.5 mm long (Meyer ous viable seeds in droppings of the quite 1994). With two panicles typical for the first abundant and frugivorous Polynesian rat, season of production, a young fruiting tree Rattus exulans Peale (Meyer 1994). Finally, produces ca. 200,000 seeds in its first season. both intentional and unintentional actions by A large fruiting tree with over 50 panicles can humans are believed to be important in the produce over 5 million seeds a year. In the short- and long-distance transport of M cal field, the large size of the soil seed bank vescens. Seeds are undoubtedly picked up (>50,000 seeds per m2 in the most invaded with mud and carried on shoes and vehicle area of Tahiti [Gaubert 1992]) and its lon wheels. They are known to have been carried gevity (>2 yr in soil samples [Meyer 1994]) with the soil of potted cultivated plants (see give M. calvescens a formidable reservoir of below). regenerative capacity even if all plants are removed from an area. In the laboratory, Introduction and Distribution in the Society some seeds germinate rapidly (15-20 days) Islands when exposed to light and moisture, but others remain dormant for up to 6 months; TAHITI. Tahiti, the largest (1045 km2) and over 90% eventually germinate under optimal highest (2241 m) island in the Society Archi conditions. Germination occurs even in pelago, is formed from two distinct ancient dense shade (at 0.02% of full sun [Meyer volcanoes, Tahiti Nui ("large," 30 kID in di 1994]) and on poor substrate (on rocks or ameter) and Tahiti Iti ("small," 22 kID by 13 barks of trees). Vegetative layering and re kID), connected by the low and narrow isth sprouting also occur. mus of Taravao (Figure 4). Ecosystems become completely trans M. calvescens was introduced in the pri formed in species composition as M. calves vate botanical garden of Motu Ovini (now cens gains dominance. Proliferation of the called Papeari Botanical Garden), on the broad leaves and overlapping architecture of southwestern coast of Tahiti, in 1937 by this species result in a dense shade that no Harrison Willard Smith, a retired American native species can tolerate. Meanwhile, the physics professor and exotic plant fancier. superficial, tentacular root system of M. cal The original label of the plant can still be vescens may not hold the soil well: M. cal seen at the botanical garden. Smith obtained vescens dense stands are suspected of con seeds of M. calvescens from the Royal Bo tributing to landslides in Tahiti (S.E.D.E.P. tanic Gardens of Peradeniya in Sri Lanka 1988). (then Ceylon), where it had been introduced Passive dispersal occurs by gravity (most from Mexico in 1888 (A. H. M. Jayasuriya, fruits fall beneath the trees), wind (seeds are pers. comm.). M. calvescens was apparently minute and light), and water (seeds germi planted later (date unknown) on the grounds nated from fruits I had left-for 3 months in -of the Agriculture Research Station at Tara water). Frugivorous birds have been ob vao by Jean Boubee (Raynal 1979), a French served in Tahiti feeding on Miconia fruits. agronomica1 engineer who was a colleague Among them, two introduced birds, the Sil and friend of Harrison Smith. vereye (Zosterops lateralis Latham) and the There are no records of the spread of M. Red-vented Bulbul (Pycnonotus cafer), are calvescens in Tahiti until the 1970s. In 1971, believed to be important dispersers (Gaubert botanist F. R. Fosberg of the Smithsonian 1992). Introduced to Tahiti in 1937, the Si1 Institution of Washington, D.C., saw this vereye has since become the most abundant species and was dismayed by its invasiveness. Status of Miconia calvescens in the Society Islands-MEYER 71

Jfitiaa t N

limit or cxtensive distribution * Sites of illtrodllL'tioll 6km

FIGURE 4. Distribution of Miconia calvescens on the island of Tahiti (after Birnbaum 1991).

He warned authorities in Hawai'i to prohibit An analysis of 1978 aerial photographs taken its introduction there (Loope and Medeiros on the plateau of Taravao showed that only 1994). Botanist Jean Rayna1 of the Museum an area of 100-200 ha of M calvescens was national d'Histoire naturelle of Paris pro visible in the forest canopy (Birnbaum 1991). claimed M. calvescens the "number one en However, this is an underestimate of the real emy of Tahitian vegetation" (Raynal 1973: spread of this plant pest, which was also 11), after noting that it already formed pure present in canopy and understory in other stands on the top of the plateau above valleys of Tahiti Hi, but not noticeable on Taravao and that it had been reported at aerial photographs (J. Florence, pers. comm.). 600-650 m elevation at the Belvedere above Furthermore, Raynal (1979: 149) noted that " Papeete~ more tban""30 km from fhenearesl "there are pure stands from Taravao to the"" known site of introduction. Whether M. cal Vaitepiha, the main valley of Tahiti Hi." In vescens was planted as an ornamental at the 1979, professor of biology B. Le Vot noted Belvedere or whether it had already expanded that M calvescens had invaded the totality of across the island is unknown. In 1976, ethno Tahiti Hi south of Tahiti Nui and entered botanist M. A. Martin found some individuals into the caldera (Le Vot 1979). In 1981, M in the lower part of the Papenoo Valley and calvescens had "supplanted all other vegeta wrote that"M calvescens would be more fre tion on certain slopes on the south side ofthe quent higher in the valley" (Martin 1976: 5). island, is spreading everywhere in the moist 72 PACIFIC SCIENCE, Volume 50, January 1996

valleys, and is proving to be serious com In less than 50 yr since its introduction, petition for the native vegetation" (Fosberg M. calvescens has spread widely throughout and Sachet 1981: 36). During an expedition much ofthe Tahitian forests, as shown by the to Lake Vaihiria in May 1982, Jacques distribution map made by P. Birnbaum in Florence (1982: 3-4), botanist of the Centre 1989 (Birnbaum 1991) (Figure 4). ORSTOM de Tahiti, noted that "the pres ence of Miconia is particularly impressive on MOOREA. Moorea is a small island (142 2 the sides of the lake where it has already km ) with steep relief: a large number of invaded the Cyathea-Pandanus forest and peaks, many between 700 and 900 m eleva makes the landscape look like a biological tion, form a mountainous chain that en desert." It has been suggested that a dramatic circles the Domaine d'Opunohu, a low-lying increase of Miconia followed the cyclones of agricultural and forestry zone (Figure 5). December 1982 and April 1983 (Birnbaum Two primary centers of establishment of 1991), but no conclusive evidence is avail M. calvescens are located on the summit and able. It is more likely that M calvescens was slopes of Mount Rotui (899 m) and Mount already present in the understory and that Mouaputa (860 m). Establishment of a small canopy openings caused by the cyclones population was first noted in the early 1970s favored growth and flowering (J. Florence, on Mount Mouaputa by an inhabitant of pers. comm.). Colonization of new open Afareiatu, the main village of the island (R. areas created by treefalls and landslides Shigetomi, pers. comm.). The remote moun seems less likely. taintop location of both the Mouaputa and

t _ limit of extensive distribution N • individual sightings o

FIGURE 5. Distribution of Miconia calvescens on the island of Moorea. Status of Miconia calvescens in the Society Islands-MEYER 73

Rotui populations suggests possible dispersal RAIATEA. Raiatea, the largest (171 km2) of from Tahiti (only 18 km distant) by wind, the Leeward Islands, is located 180 km WNW birds, or recreational hikers. Currently, occur of Tahiti. Unlike Moorea, Raiatea has rela rences ranging from individual trees to dense tively smooth relief dominated by the two localized populations have been reported in plateaus of Temehani at 500-800 m eleva many locations within and on steep slopes tion (Figure 6). above the Domaine d'Opunohu (Schwartz Small populations of M. calvescens cover 1994). ing a total of ca. 240 ha were reported in

.~;i""===,------

- limit of extensive distribution t • individual sightings N 3km

FIGURE 6. Distribution of Miconia calvescens on the island of Raiatea. 74 PACIFIC SCIENCE, Volume 50, January 1996

three separate valleys (Uturaerae, Tetooroa, light regime and is suspected of causing and Faaroa) in 1988 by the forestry section landslides; it alters, therefore, "the basic rules of the Service de l'Economie Rurale (J.-P. ofexistence ofall organisms" (Vitousek 1990 : Malet, pers. comm.). The Uturaerae popula 8). Its invasion of native forest is progres tion originated from an ornamental planting sively eroding native biological diversity: half in 1955 that had become naturalized by 1969 of all the endemic species of Tahiti are di 1970 (R. Amiot, pers. comm.); by 1989, there rectly endangered by M. calvescens (J. Flor were dense, monotypic stands in the valley ence, pers. comm.). The success of M. cal bottom (Florence and Birnbaum 1989). In the vescens in the Society Islands is apparently a moist (mean annual rainfall >5000 mmjyr) result of the ecological characteristics of the valley of Tetooroa, seeds of M calvescens species that allow it to thrive and spread are believed to have been introduced with (rapid growth rate under optimal conditions, soil of coffee plants from Tahiti in the 1970s. prolific seed production, efficient dispersal, Introduction of the Faaroa Valley popula persistence in the soil seed bank, shade toler tion probably dates back to 1981 with the ance) under suitable environmental condi beginning of the forestry industry (Swietenia tions; the absence in the Society Islands ofthe macrophylla King plantations) and the intro specific natural enemies that are present in its duction of "contaminated soil" from Tahiti native range in Central and South America (Malet 1992). (R. Burkhart, pers. comm.), where the species Dispersal from the three Raiatea popula is not abundant; and a presumed inherent tions ofM calvescens has been limited largely vulnerability of island forests to invasion. by surrounding plantations of pine forest on Because similar factors are present in high ridgetops. This fortuitous circumstance has oceanic islands throughout the Tropics, the allowed promising control efforts by the for possibility of destructive invasion of other estry section of the Service de l'Economie archipelagoes by M calvescens is obvious. Rurale. A total of ca. 25,000 plants (includ Stringent measures are needed to prevent the ing 500 large, seed-producing trees) was spread and escape of this aggressive species hand-removed in June 1992, with assistance to other locations. The striking appearance of 200 school volunteers. Follow-up in June of the plant, with its large green and purple July 1993, with the help of 100 soldiers from leaves, makes it attractive to plant fanciers the French Army, removed more than 77,000 (Graf 1986). plants. Seedlings and saplings were uprooted, Miconia calvescens recently was recognized and trees too large to be uprooted were cut as an unwelcome addition to the nearly 100 and stumps treated with herbicide (2,4-D) to dangerously invasive plant species of the Ha prevent resprouting. Although follow-up will waiian Islands (Smith 1985, Gagne et al. 1992); be necessary for several years to allow the unfortunately, by the time serious control exhaustion of the seed bank, there seems to measures were instituted, it had been present be a good chance of eradicating M. calves for over 20 yr and spread to multiple loci on cens from Raiatea. two islands (L. L. Loope and A. C. Medeiros, pers. comm.). It also occurs in such locations as the Philippines (R. Petocz, pers. comm.), Grenada (P. Cazin, pers. comm.), and New CONCLUSIONS Caledonia (R. Lavoix, pers. comm.) as an Miconia calvescens has already attained ornamental, not yet naturalized. dominance over most of the island of Tahiti. Dense, monotypic stands of this species oc cupy mesic and wet environments (mean an ACKNOWLEDGMENTS nual rainfall >2000 mm) at 10-1300 m ele vation. It not only occupies roadsides, forest I thank Jacques Florence (Centre ORS edges, and abandoned pastures, but also has TOM de Tahiti) for the help so generously invaded primary native forests. It changes the provided during my 2 yr field study in French Status of Miconia calvescens in the Society Islands-MEYER 75

Polynesia, John D. Thompson (CEFEj and invader plant species on the island of CNRS, Montpellier) and Paula Baylet for Guam. Biotropica 6(3): 158-164. revising the English on the first draft, and LE VOT, B. 1979. La vegetation aTahiti: In two anonymous reviewers for suggestions on troduction a une etude ecologique. Bull. the manuscript. I am also particularly grate Nat. Polynesie Fr. 6: 1-23. ful to Lloyd Loope (HaleakaUi National LOOPE, L. L., and A. C. MEDEIROS. 1994. Park, Maui) and Finn Kjellberg (CEFEj Interagency efforts to control the sprsad of CNRS, Montpellier) for their numerous cor a highly invasive tree (Miconia calvescens) rections, comments, and recommendations. on Maui, Hawaii (abstract). Bull. Ecol. Soc. Am. 75: (in press). LOOPE, L. L., and D. MUELLER-DoMBOIS. 1989. Characteristics of invaded islands, LITERATURE CITED with special references to Hawaii. Pages BAILEY, L. H. 1900. Cyclopedia of Ameri 257-280 in J. A. Drake, H. A. Mooney, F. can horticulture, vol. 3. Pages 1011-1012. di Castri, R. H. Groves, F. J. Kruger, M. Macmillan and Co., London. Rejmanek, and M. Williamson, eds. Bio BIRNBAUM, P. 1991. Comment une plante in logical invasion: A global perspective. troduite se transfome-t'elle en une plante John Wiley & Sons Ltd., Chichester. envahissante? Le cas de Miconia calves LORENCE, D. H., and R. W. SUSSMAN. 1986. cens a Tahiti (Polynesie Franyaise). Rap Exotic species invasion into Mauritius wet port de D.E.A., Universite des Sciences et forest. J. Trop. Ecol. 2: 147-162. Techniques du Languedoc, Montpellier. MACDONALD, I. A. W., C. THEBAUD, W. A. FLORENCE, J. 1982. Rapport de mission au STRAHM, and D. STRASBERG. 1991. Effects Lac de Vaihiria (Tahiti) du 17 au 19 mai of alien plants on native vegetation rem 1982. Centre ORSTOM de Tahiti, section nants on La Reunion (Mascarene Islands, botanique, no. 82-06. Indian Ocean). Environ. Conserv. 18(1): ---. 1987. Endemisme et evolution de la 51-61. flore en Polynesie Franyaise. Bull. Soc. MALET, J.-P. 1992. Point sur les operations Zool. Fr. 112(3-4): 370-380. d'eradication du Miconia sur 1'11e de Raia FLORENCE, J., and P. BIRNBAUM. 1989. Eval tea. Service de l'Economie Rurale, 2eme uation de la situation de Miconia calves Secteur Agricole. cens aRaiatea. Rapport de mission, Cen MARTIN, M. A. 1976. Compte-rendu de la tre ORSTOM de Tahiti. mission ethnobotanique effectuee a Ta FOSBERG, F. R., and M. H. SACHET. 1981. hiti, dans la vallee de la Papenoo (Juilletj Miconia calvescens de CandolIe. Poly Aout 1976). C.N.R.S., RCP no. 259: Pro nesian plant studies. Smithson. Contrib. gramme Papenoo. Bot. 47: 36-37. MEYER, J.-Y. 1994. Mecanismes d'invasion GAGNE, B. H., L. L. LOOPE, A. C. MEDEIROS, de Miconia calvescens DC. en Polynesie and S. J. ANDERSON. 1992. Miconia cal Franyaise. Ph.D. diss., Universite des vescens: A threat to native forests of the Sciences et Techniques du Languedoc, Hawaiian Islands. Pac. Sci. 46: 390-391 Montpellier. (abstract). PASTUREL, J. 1993. La climatologie des iles. GAUBERT, H. 1992. Les invasions biologiques Pages 42-43 in Atlas de la Polynesie fran en milieu insulaire: Le cas de Miconia cal yaise. Editions de l'ORSTOM, Paris. vescens aTahiti. Rapport de VAT, Centre RAYNAL, J. 1973. Les introductions mal ORSTOM de Tahiti. encontreuses d'especes etrangeres. Pages GRAF, A. B. 1986. Tropica, color cyclopedia 10-12 in Rapport du Museum national of exotic plants and trees, 3rd ed. Page d'Histoire naturelle, Paris. 645. Roehrs Co. Publishers, East Ruther ---. 1979. Three examples of endangered ford, New Jersey. nature in the Pacific Ocean. Pages 149 LEE, M. A. B. 1974. Distribution of native 150 in I. Hedberg, ed. Systematic botany, 76 PACIFIC SCIENCE, Volume 50, January 1996

plant utilisation and biosphere conserva operative National Park Resources Stud tion. Almquist & Wiksell, Uppsala. ies Unit, Honolulu. SCHOFIELD, E. K. 1989. Effects of introduced STONE, C. P., C. W. SMITH, and J. T. TUNI plants and animals on island vegetation: SON, EDS. 1992. Alien plant invasions in Examples from the Galapagos Archi native ecosystems of Hawaii: Manage pelago. Conserv. BioI. 3: 227-238. ment and research. University of Hawai'i SCHWARTZ, J. 1994. Distribution of an inva Press for University of Hawai'i Coopera sive introduced plant Miconia calvescens tive National Park Resources Studies on the island ofMoorea, French Polynesia. Unit, Honolulu. Department of Conservation and Re VITOUSEK, P. M. 1990. Biological invasions source Studies, University of California and ecosystem processes: Towards an in at Berkeley. tegration of population biology and eco S.E.D.E.P. 1988. Amenagement hydro-elec system studies. Oikos 57: 7-13. trique de la haute Papenoo. Etude d'im WURDACK, J. J. 1971. Notes on some culti pact, Papeete. vated species of Miconia (Melastomata SMITH, C. W. 1985. Impact of alien plants on ceae). Baileya 18: 17-19. Hawaiian native biota. Pages 180-250 in ---. 1980. Miconia calvescens DC. Pages C. P. Stone and J. M. Scott, eds. Hawai'i 171-172 in G. Harling and B. Sparre, terrestrial ecosystems preservation and eds. Flora of Equator, no. 13. University management. University of Hawai'i Co- Goteborg and Riksmuseum, Stockholm.