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Tmd’a Woody list for l@tlgua and Barbuda: Past and Present

John Francis, Cados Riira, and Julio Figurema S_

Durina;thel7thandl8~centuriea,theielandofAntiguawas~~ foragri~.viuauyalloftheremaining foresbwemcutover,grazed,and bumed.Sincethe aba&mentofaugamne~vati~xnuchofthelandhas retunledtQsecondaryf&.BarbudasufferedmeiIllyfromovercuttiIlgand overgrazing begiming in the 17th century. Very heavy grazing preamre con- tinues in ISarbuda. l&spit23 great d&W, a mqjority of the native qe45ee remainrmbothislande.Lista0fnativeandexoticepecieaarep~~bere.A prote&i~plansbauldbe~hedforsevleralareasmboth~ Tk~ward a List for and Barbuda: Past and Present John Francis, Carlos Rivera, and Julio Figureroa

INTRODUCTION AND HISTORY hectares of Antigua to grow up as secondary forest while sparing the pre-existing secondary forest from are two small islands with a further disturbance. total area of 440 km2 located in the in English attempts to settle Barbuda began in 1628 the eastern . Although not differing greatly and continued intermittently with attacks of the in size and separated by only 50 km, the origin and Caribs and French until early in the 18th century. The physiography of the islands are quite different. Antigua small island community subsisted for almost three cen- was formed by vulcanism more than 40 million years turies by raising livestock. During this period, vir- ago during the Eocene. The rock foundation of Bar- tually every large enough to use for construction buda is entirely of Pleistocene limestone. A land bridge materials was felled. Grazing must have been intense, apparently connected the two islands during the and fuelwood cutting heavy, at least near Codrington, Pleistocene (Schuchert 1935). From the time the the only town. Tourism, government services, and agri- islands emerged, a flora began to assemble through culture are the primary sources of income today for the chance introductions, mainly transported by birds. island’s 1,252 residents. Cement blocks and imported Intermittent human habitation of both Antigua and lumber are the current building materials. Barbuda has been dated from about 3,000 B.C. The Today, grazing may be as intense as ever in Bar- aboriginal groups, archaic Amerindians, agricultural buda’s history. Domestic goats and , feral donkeys Amerindians, and Island Caribs never had large popu- and pigs,. and introduced deer have left the ground lations and probably did not clear a great deal of land under the dry forest canopy nearly barren. All the for agriculture. However, they apparently introduced a and within reach, except for a few toxic or number of plant such as Annona muricdu L. unpalatable species, are being consumed or cropped and bijugatus Jacq. severely. Christopher Columbus visited Antigua and named it With the emergence of tourism as the driving force after Santa Maria della Antigua during his second voy- of the economy and the decline of agriculture, concern age in 1493. Because of the scarcity of fresh water and for the environment has grown. The protection of natu- the presence of threatening &ribs, no attempt was ral areas and the setting aside of areas to be used as made to colonize the island until a group of English parks are being suggested. This study was undertaken settlers arrived in 1632. The first export crop, tobacco, by the International Institute of Tropical Forestry was soon replaced by sugar cane. The sugar enterprise (IITF) in response to a request to identify local plant became profitable with the importation of slave labor, species and classify them as native or exotic. This basic and a number of large plantations were established. By data will be important for future management of natu- the early 1700’s, most of the island had been cleared for ral reserves. sugar production. Subsistence farming extended to the top of Boggy Peak, the highest mountain. Even the rocky hills in the drier parts were cut over periodically METHODS for firewood and charcoal. Goats, cattle, and horses grazed and browsed all the uncultivated remnants of Lists of trees, shrubs, and woody , including land. The phasing out of the sugar industry between ornamentals, were assembled from field observations 1960 and 1972 led to extensive abandonment of cultiva- (tables 1 and 2). The survey consisted of two visits to tion. At the same time, the emergence of the tourist Antigua and one to Barbuda. In the spring of 1991, a industry and urbanization has moved hundreds of peo- botanist and a research forester spent 5 days on the ple off subsistance farms and has allowed thousands of island of Antigua. Six months later, they returned with

John Francis is a research forester and Carlos Rivera is a botanist at the International Institute of lkopical Forestry, U.S. Department of Agriculture, Forest Service, Rio Pi&as, PR 00928-2500; Julio Figueroa is a graduate student at Oxford University, Oxford, UK. In cooperation with: University of , Rio Piedras, PR 009364984. a dendrologist and spent 3 days on Antigua and 2 days ralized species are only rarely found growing outside surveying Barbuda. Guided by local experts, the group cultivation in waste places near planted specimens. A visited as much of each island as possible. handful of species, notably nilotica (L.) Delile, Because Antigua is thoroughly interlaced with roads, lebbeck (L.) Benth., Azadirachta indica Adr. the survey was conducted by driving through all por- Juss., obliqua Willd., Haematoxylon camp- tions of the island. Frequent walks were taken a few echianum L., Leucaena leucocephala (Lam.) de Wit, hundred meters into forest stands. The names of Phoenix dactylifera L., juliflora (SW.) DC., positively identified species and the locations of their Thespesia populnea (L.) Sol., and Ziziphus mauritiana sightings were noted. Samples of species that were not Lam., compete successfully with native species in sec- readily identifiable were collected for checking against ondary forests on dry sites. A total of 127 woody species references. Those specimens that could not be identi- was observed on Barbuda, of which 65 were believed to fied were brought to the University of Puerto Rico be native to the island. The only exotic woody plant for examination by resident taxonomists. species competing significantly with native vegetation As a result, all but two of the observed species (repre- is H. campechianum. sented by foliage only) were identified. All scientific As an exercise to evaluate the rate of tree species loss names were checked for currency and spelling using as a result of the clearing, cutting, grazing, and fire Howard’s “Flora of the Lesser Antilles” (1979, 1988, associated with settlement, in table 5 of this survey is 1989a, 1989b). Other references consulted during the used to compare the canopy tree species (excluding process of identification and correlation of species were naturalized ornamentals and trees) that were Fournet (1978), Grisebach (1963), Liogier and Mar- observed with those collected by botanists from the torell (1982), Little and others (19741, and Little and 18th century onward (Howard 1979, 1988, 1989a, Wadsworth (1964). 198913). For Antigua, 39 species in common with the herbarium collections were listed. Howard lists six spe- cies that were not found, and two species not repre- RESULTS sented in the herbarium collections were found. Of those not found in the IITF survey, DacryocEes excelsa Plant lists derived from different surveys in Antigua Vahl has certainly been lost (it is unknown to Anti- and Barbuda are compared in tables 3 and 4. It should guans today). Beilschmiedia pendulu (SW.) Hemsley, be noted that the lists by Howard (1979, 1988, 1989a, Cecropia schreberiana Miq., Sapium caribaeum 198913) were compiled from ranges documented by Urban, and Vita divuricutu SW. probably have disap- herbarium specimens deposited in world museums and peared from Antigua. All are present on other Antillian that these specimens were collected from the early 18th islands. The two new species noted were Cordia century onward, many before the destruction and dis- Zaevigatu Lam. and Ochromapyrumidczle (Cav.) Urban. turbance of Antiguan forests were complete. The former probably was missed in earlier surveys; the A total of 286 woody species was observed on Anti- latter may be a new natural introduction. gua, of which 153 were determined to be native. A For Barbuda, the IITF survey and the species was concluded to be native if its documented had 15 canopy species in common. Howard lists three native range included Antigua or Barbuda and if it species that were not observed: pubescens L., grew in uncultivated conditions. A few technically saponaria L., and salicifolium native species, mainly food producers such as Annona (L.) Lam., which must be either rare or extinct on squamosa L. (sugar-apple) and Munihot esculenta Barbuda. Coccoloba diversifolia Jacq. was observed in Crantz (), were apparently introduced in pre- this survey as well as by Beard and Harris (1949,1960X Columbian times and today are rarely found growing It was undoubtedly overlooked in other surveys. outside cultivated areas. oleracea O.F. Cook is listed as exotic by Howard (1979), but may be native to Antigua. Antigua has established populations in moist secondary forests, DISCUSSION and large R. oleracea trees may be seen in the oldest photos of Antiguan plantations. ‘Iwo other islands with In a single survey of a flora, one cannot be sure of native populations of this species, Monserrat and finding every species. Field time in this survey was , lie within sight of Antigua. Because the somewhat limited; an additional 10 to 25 native species are bird-dispersed and the are useful to on Antigua and a few on Barbuda may have been found humans for roofing material, it would be remarkable if if a thorough search of all parts of both islands had the species had not become established before the been completed. This survey, combined with previous arrival of Columbus. efforts, adds a significant contribution to the compila- Nearly all the exotic species are or were cultivated as tion of a checklist of the woody flora of Antigua and ornamentals, shade, or crop . Most of the natu- Barbuda. The most satisfactory next step would be for

2 a resident botanist to conduct extensive field checking Bourreriu succulenta Jacq. and winterunu (L.) and eventually compile checklists and descriptive Gaertn. Rocky hills that may have passed the settle- floras. ment era as open woodland pasture are dominated Because of the individual experience of the partici- today by large Bucidu bucerus L. and Burseru sim- pants, the IITF survey was probably most reliable in urubu trees. With some variation reflecting soil condi- identifying forest trees and introduced trees and tions, the dry hills and plains of Antigua would have shrubs (ornament& and fruit species). Participants been dominated in pre-Columbian times by Bucida were less familiar and less attuned to the shrubs and bucerus, Burseru simaruba, Guuiacum ofiinale L., woody vines, although an effort was made to collect and Pimenta racemosa (Miller) J. Moore, Pisoniu subcor- identify every species seen, Early botanists contribut- data, and Zunthoxylum flwum Vahl. ing to the collections used by Howard (1979, 1988, Streams and nonsaline floodplains are still domi- 1989a, 1989b) to compile his flora of the Lesser nated by Annonu glabru L., pentundru (L.1 Antilles apparently missed a few species as well. There Gaertn., Hippomane mancinellu L., and Hura cre- are undoubtedly errors of identification in this survey’s pituns L. The swamps also still contain the lists and even in those from herbarium collections. original species, Avicennia germinuns CL.> L., Lugun- Careful rechecking by future researchers is suggested. culuriu rucemOsa (L.) Gaertn. f., and Rhizophoru mun- gle L. Barbuda was divided into four broad site types (fig. Present and Presettlement Forests 2): the highlands, clay slopes and plains, sandy coastal Today, a large portion of Antigua is covered with plains and beach strands, and mangrove swamps. The secondary forests. Most of these forests are less than 30 highlands are dominated today by CuneZZu winteranu, years old, although a few areas, such as the Wallings Haematoxylon campechianum, and Pilosocereus royeni watershed, have been largely undisturbed for more on the plateau and C. winterana, F. citrifoliu, Krugio- than 50 years. No primary forest remains, and, exclud- dendron ferreum (M. Vahl) Urban, and Pisonia s&or- ing the , only a few small areas were never data on the slopes. In presettlement times, the cleared for agriculture. Consequently, tree diameters highlands were probably dominated by 2. flmurn with tend to be relatively small. associates C. winteranu, F. citrifoliu, G. officinule, K. Four broad forest types that were probably more distinct in their pristine states than they are today were recognized (fig. 1). The upper portion of Boggy Peak is the wettest and coolest area of the island. It is HIGH ELEVATION MOIST currently dominated by Ingu Zaurina (SW.) Willd. Other MOIST important canopy species in the area are Clusia major m L., Cordia sulcutu DC., citrifoliu Mill., Lone- u DRY hocurpus benthaniunus (Jacq.) DC., and Pisonia sub- FLOODPLAIN, 8 MANGROVE SWAMPS corduta SW. The original forest was probably dominated by Vahl, other important forest dominants being C. major, I. Zaurinu, Simaroubu umuru Aubl., and Zunthaxylum murtinicense (Lam.) DC. The moist uplands of Antigua today have a mix of dominant Andiru inermis (Wright) Kunth ex DC., Bur- seru simarubu (L.) Sarg., Hymenaeu courbaril L., I. Zuurinu, Mungiferu indicu L., R. oleraceu, mombin L., and ‘Ihbebuiu heterophyllu (DC.) Britt. The primordial forest was probably dominated by CedreZa o&rata L., I. Zuurina, T. heterophylla, Z. murtinicense, and a host of other species. The dry hills and coastal plains support several for- est types depending on the history of the site. The most recently abandoned or most abused sites are currently dominated by Acacia spp. and Leucaenu Zeucocephala

(Lam.) de Wit. They are succeeded by Haematoxylon 0 IO mi cumpechianum and Prosopis juliflora. More advanced sites support Capparis spp., Pisoniu spp., PiZosocereus royeni (L.) Byles & Rowley, and Ziziphus muuritianu. Those that have developed further also support Figure 1. -Major forest types on Antigua. not surprising that extinctions and rarefaction CLAY SLOPES, 8. PLAINS occurred. In this survey, evaluation of tree loss indi- HIGHLANDS cates that about 13 percent of the native canopy tree SANDY COASTAL PLAINS, B BEACH STRANDS species have disappeared from Antigua since the late MANGROVE SWAMPS 18th century. odorata, once cut for its valuable , no longer grows in the wild. Guaiacum oflicinale and Z. flavum also produce very valuable wood and have become extremely rare in the wild. The rate of extinction of small trees, shrubs, and woody vines may be even more severe. Smaller plant forms are more short-lived, are more subject to browsing by goats and other livestock, and are more easily cut down than large trees. On the other hand, given the thorough destruction and disturbance of the forests of Antigua, it is remarkable that such a large percentage of species has survived. The Barbudan forests suffered in different ways, and severe disturbances continue. Most trees large enough to use for construction materials were felled long ago, resulting in the near disappearance of several species. Grazing over the centuries and very heavy browsing by goats, donkeys, cattle, deer, and feral hogs in recent times have made reproduction difficult or impossible for all but toxic or very unpalatable species. The very survival of several species is threatened. barbadensis and Thrinax morrisii H. Wendl., both palms, are cases in point. A hundred or more large C. barbadensis still grace the skyline in the northern part of the island; not a single plant less than 2 m high was Figure 2. -Major forest types on Barbuda. found in this survey. Thrinax morrisii seems to have disappeared entirely. The heavy grazing has allowed G. officinale and Canella winterana, both untouched by grazing, to reproduce in large numbers. ferreum, and l? subcordata. The clay slopes and plains The loss of biodiversity is somewhat offset by the are dominated today by Acacia farnesiana, C. win- naturalization of a number of exotic species. Notable on terana, Capparis spp., G. offtcinale, H. campechianum, Antigua are Acacia nilotica, Azadirachta indica Adr. and Pilosocereus royeni. In former times, the site type Juss., Calotropisprocera (Ait.) Ait. f., Cocos nucifera L., was probably dominated by Canella winterana, Cap- C. obligua, Haematoxylon campechianum, Leucaena paris Coccoloba diversifolia Coccothrinax spp., Jacq., leucocephala, Mangifera indica, Melicoccus bzjugatus barbadensis (Lodd. ex Mart.) Bacc., G. officinate, and Jacq., Parkinsonia aculeata L., P dactylifera, P. Pisonia subcordata. The sandy coastal plains and juliflora, Ricinus communis L., R. oleraceae, Solanum beach-strand areas are dominated today by Bucida torvum SW., mahagoni (L.) Jacq., Tam- buceras, Canella winterana, Chrysobalanus icaco L., arindus indica L., Terminalia catappa L., T. populnea, Hippomane mancinella, Jaquinia armillaris Jacq., and and Ziziphus mauritiana. Pilosocereus royeni. The dominant species were proba- Barbuda supports the naturalized exotics Caparia bly much the same in former times. Mangrove vegeta- biflora L., C. obliqua, Jatropha curcas L., S. mahagoni, tion, although it was heavily exploited for fuel before and Tamarindus indica. Because of its isolation, Bar- electricity and fossil fuels were available, is now much buda has escaped many introductions. Noticeably the same as it was in ancient times. absent are P. juliflora, Calotropis procera, R. com- munis, and T. populnea. Disturbance and Extinction

The degree of disturbance to the terrestrial habitat Plea for Conservation of Antigua was very severe. Excluding the mangroves, probably 95 to 99 percent of the forest was cleared and The natural vegetation of the island of Antigua has converted to agricultural uses at least once in the 300- passed through an era of neglect and abuse with sur- year history of its development. All the remaining for- prisingly few species lost. The current trends of est was heavily disturbed by cutting and grazing. It is and succession toward stand composi-

4 tions more like the original forest should be aided protected. The highlands are certainly the most unique through formalized protection of at least unique vege- geological and biological area of the island and, for that tative areas such as Boggy Peak, most of the remaining reason, deserve complete protection. Any management mangroves, the Wallings watershed, and representative scheme for Barbuda must address the problem of open- dry forest sites. For these sites, statutory protection range grazing of domestic animals and control of the and some minimal physical protection are a minimum populations of feral animals. Without it, the highland requirement. Likewise, in Barbuda, much of the man- thicket ecosystem will continue to deteriorate, and sev- groves, the salt ponds, and the dune areas should be eral species of plants will disappear from Barbuda.

Table 1. -Dee and species identified during field studies in Antigua

Family and species Where first seen Life form* Native range

Cycadaceae Cycas circinalis L. St. John’s t and +

Araucariaceae Aroucaria heterophylla (Salisb.) Franc0 St. John’s T Norfolk Island’

Cupressaceae Cupressus sempervirens L. Belleview Heights t South ’ Thuja orientalis L. St. John’s t West Asia+

Camarinaceae Casuarina equisetifolia L. Dickinson Bay T Oceania+

Piperaceae aduncum L. Figtree Native*

Ulmaceae Edema micrantha CL.1 Blume Body Ponds Native

Moraceae Artocarpus altilis 6. Park) Fosb. St. John’s Oceania+ A. heterophyllus Lam. Buckleys + Ficus benjamina L. St. John’s Malaya+ F. citrifolia Mill. St. John’s Native I? elastica Boxb. ex Hornem. St. John’s Tropical Asia+ I? microcarpa L. f. St. John’s + R trigonata L. St. John’s Tropical America Dophis racerrwsa CL.1 Urban Jolly Beach Native

Proteaceae Grevillea robusta A. Cunn. Botanic Garden T +

Olacacea Schoepfia artmaria Urban & Britt. Boggy Peak S Probably native S. Schreberi Gmel. Boggy Peak S Native

Polygonaceae Antigonon leptopus Hook. & Arn. English Harbour + Coccolobo diversifolia Jacq. Shirley Heights Native C. pubescens L. Boggy Peak Native C. uvifera CL.1 L. Runaway Bay Native C. wnosa L. Figtree Native

Nyctaginaceae Bougainvillea spectobilis Willd. Cedar Grove V + Pisonia fragrans DumXours. Cedar Grove t Native P. subcordata SW. Cedar Grove T Native lhble 1. - !lhee and shrub species identi@d during field studies in Antigua - Continued

Family and species Where first seen Life form* Native range

Annonaceae Anmna glnbra L. Collins t Native A. montana Macfad. Wallings t Native A. muricata L. St. John’s t Probably native A. reticulata L. St. John’s t Probably native A. squamosa L. St. John’s t Tropical America+

Lauraceae Licaria paruifolia (Lam.) Kostermans t Native Ocotea cernua (Nees) Mez Boggy Peak t Native 0. coriocea (SW.) Britt. Figtree t Native Persea americana Mill. St. John’s T Mexico+

Capparaceae Capparis flenuosa (L.) L. Cedar Grove t Native C. baducca L. Figtree t Native C. hastota Jacq. Mercers Creek t Native C. in&a (I~.) Druce Cedar Grove t Native

Moringaceae Moringa oleifera Lam. English Harbour t +

R.esaceae Rosa spp. St. John’s S Asia+

Leguminosae- Acacia farnesiana (L.) Wad. Cedar Grove t Native A. macracantha Humb. & Bonpl. Collins t Native A. n&&a CL.1 Delile Mercers Creek t Africa, Asia A. tortuosa CL.1 Willd. Cedar Grove t Native Albizia lebbek (L.) Benth. Cedar Grove T South Asia virgatus (L.) Willd. Runaway Bay Native cyclocarpum (Jacq.) Griseb. Botanic Garden ; Tropical America+ laurina (SW.) Willd. Boggy Peak T Native Leucaena leucocephulu (Lam.) de Wit Ailp0l-t t IYopical America ceratonia L. Body Ponds S Native Pitheceltiium unguis-coti CL.1 Benth. St. John’s 5 Native Prosopis ju&lora (SW.) DC. Cedar Grove t saman (Jacq.) Merr. Dickinson Bay T Tropical America+

Leguminosae- Bauhinia monandm Kurz Runaway Bay t Southeast Asia+ B. multineruia (Kunth.) DC. Figtree t Probably native Caesalpinin bonduc (L.) Roxb. Shirley Heights V Native C. pulcherrima CL.1 SW. Dickinson Bay South America+ Cassia fistula L. Golden Grove ; Asia+ C. grandis LX St. John’s T Tropical America+ C. jooanica L. Freemans t Southeast Asia+ Chnmaecrista glandulosa CL.) Greene var swartzii (W&s.) Irwin & Barneby Shirley Heights t Native Delonti regia (Bojer ex Hook.) Raf. St. John’s T + Haematoxylom campechianum L. Wallings t Hymenuea courbaril L. Boggy Peak T Native Parkinsonia oculeata L. Bethesda t Tropical America pterocarpum (DC.) Back. & Heyne Dickinson Bay T Southern Asia+ Sennu al&a CL.) Roxb. Greencastle s Tropical America+ 5’. atomaria (L.) Irwin & Barneby Shirley Heights 8 Native 5’. bicapsularis (L.) Roxb. var. bicapsularis Dark Valley Native S. siamea Irwin & Barneby St. John’s ; Southeast Asia+ lllzmarindus indica L. Body Ponds T Africa

Leguminosae-Papilionoideae inermis (Wright) Kunth Wallings T Native Cajanus cajun (L.) Huth Scotts Hill s Africa+ berterwann Urban St. John’s t Central America+ E. variegata L. St. John’s t India to Oceania+

6 Table 1. -7bee and shrub species identified during field studies in Antigua - Continued

Family and species Where first seen Life form* Native range

Gliricidia sepium (Jacq.) Kunth Runaway Bay t Tropical America+ Zndigofera suffruticosa Mill. Shirley Heights S Native Lonchocarpus violoceus (Jacq.) DC. Boggy Peak t Native Machaerium lunatum (L.f.1 Ducke Figtree t Probably native Piscidia carthagenensis Jacq. Shirley Heights t Native Sesbania gmndiflora CL.1 Pers. Body Ponds t South Asia+

Oxalidaceae Aver&a carambola L. Greencastle t Tropical Asia+

Erythroxylaceae Erythroxylon havanense Jacq. Shirley Heights t Native

Zygophyllaceae Guaiocum ofiinole L. Nelson’s Dockyard t Native

Rutaceae Citrus aurantifolia (Christm. & Panzer) Swingle St. John’s t East Indies+ C. limon CL.1 Burm.f. St. John’s t Southeast Asia+ C. maxima (J. Burm.) Merr. Greencastle t East Indies+ C. pamdisi Macf. Freemans t + C. reticulata Blanc0 Cades Bay t East Indies+ C. sinensis CL.1 Osbeck Christian Valley t + Murraya paniculata CL.1 Jack Mercers Creek S Southeast Aam+ Diphasia trifolia (Burm. f.) P. Wils. Mercers Creek Southeast Asia flavum Vahl Boggy Peak ; Native 2. martinicense (Lam.1 DC. Boggy Peak T Native 2. monophyllum (Lam.) P. Wils. Boggy Peak t Native 2. spinifex (Jacq.1 DC. Freetown t Native

Simaroubaceae Castela erecta %-pin Indian Town Point 8 Native Picmmnia pentandm Sw. Wallings t Native excelsa (SW.) Planch. Wallings t Native Simarwuba amara Aublet Boggy Peak T Native

Surianaceae Suriana maritima L. Cades Bay 8 Native

Burseraceae simaruba CL.1 Sarg. Jennings T Native

Meliaceae Amdirachta in&a Adr. Juss. Dickinson Bay T South Asia Cedrela oo?omta L. St. John’s T Native+ L. Cedar Grove t South Asia CL.) Jacq. St. John’s T Greater Antilles Wchilia hirta L. Figtree t Native

Malpighiaceae Bunchosia glandulosa (Cav.) L.C. Rich. Flashes 8 Native Byrsonima lucida (Mill.) DC. Greencastle t Native B. spicata (Cav.) DC. Boggy Peak t Native Malpighia emarginata DC. Runaway Bay t West Indies M. linearis Jacq. Nelson’s Dockyard 8 Probably native

Euphorbiaceae Acalypha amentocea Roxb. ssp. wilkesiana (Muell. Arg.) Fosberg St. John’s S Oceania+ Cod&urn variegatum CL.1 Blume Dickinson Bay S Sunda Islands+ Croton astroites Dryand. Cedar Grove 8 Native continifolia L. St. John’s t Tropical America+ E. lactea Haw. Dickinson Bay t Old World+ E. neriifolia L. Liberta t East Indies+ E. pulcherrima Willd. St. John’s S IYopicsl America+ E. tirucalli L. Runaway Bay 8 North Africa+

7 ‘Ibble 1. - !&ee and shrub species identified during f?eld studies in Antigua - Continued Family and species Where first seen Life form* Native range

Gymnanthes lucida SW. Shirley Heights t Native Hippomane mancinella L. Dickinson Bay T Native Hum crepitans L. Bethesda T Probably native Jatropha curcas L. St. John’s 9 Central America J. gossypiifdia L. Cades Bay S Probably native J. multifida L. St. John’s 8 Tropical America+ Manihot esculenta Crantz Sea View Farm 8 South America+ acidus (L.) Skeels Cedar Grove t Tropical Asia+ F! epiphyllanthus L. Shirley Heights t Native Ricinus communis L. Sea View Farm S Tropical Africa

Anacardiaceae Anacardium occidentate L. Hamiltons t Probably native+ Conwclodiu dodonoea CL.1 Urban Shirley Heights Native Ma&fern indica L. Hodges Bay ; Tropical Asia Spondias c@wea Sonn. English Harbour T Society Islands+ S. mombin L. St. John’s T Native S. purpurea L. St. John’s t Tropical America+

Celaetrace@s Cassine xylocarpo Vent. English Harbour t Native Cmssopetalum rhacoma Crantz Freetown 5 Native frutescens Jaq. Shirley Heights S Native

Sapindaceae Allophylus mcemosus SW. Boggy Peak t Native Melicoccus bgugatus Jacq. St. John’s T South America Sapindus sapon4wio L. Wallings t Native Schleichera oleosa (Low.1 Oken Dark Valley T South Asia+

Rhamnf8ceae CoZz&rinu arborescens (Mill.) Sarg. Crabs Mill t Native Krugiodendmn ferreum (M. Vahl) Urban St. Phillips t Native Ziziphus mauritiana Lam. Cedar Grove t Southeast Asia

TiIiaceae !&iumfetta grandifira M. Vahl Body Ponds 8 Probably native

Mahceae Hibiscus pernambucensis Arntda St. John’s t South America+ H. rosa-sinensis L. St. John’s t Tropical Asia+ H. schizopetalus (M.T. Mast.) HookX St. John’s S Africa+ Maluastrum coromand-elianum CL.) Garcke Wailings 9 Tropical America Malvaviscus arboreus Cav. St. John’s S Mexico-Brazil+ Thespesia populnea CL.) Sol. Dickinson Bay t Old World

Bombacaceme Ceiba pentandm (L.) Gaertn. Body Ponds T Native Ochmma pymmidale (Cav.) Urban Figtree T Native

Sterculiaceae Guaxumn ulmifdia Lam. Wallillgs t Native foetida L. Botanic Garden T Tropical Asia+ cacao L. John Hugh’s t Neotropicst

Ochnaceae Ouratea guildingii (Planch.) Urban Ail Saints S Native

Clusiaceae Calophyllum calaba L. Figtree T Native Clusia major L. Boggy Peak T Native Mammea americana L. Greencastle T West Indies+ lhmaricaceae !&maria sp. Old Road t Old World+

8 Table 1. - lhce and shrub species identified during field studies in Antigua - Continued Family and species Where first seen Life form* Native range

Cocblospermaceae Cochlospermum vitifolium (willd.1 Spreng. All Saints T Tropical America+

Canellaceae Canella wintemna (L.) Gaertn. Mercers Creek t Native

Flacourtiaceae Casearia okcandra Jacq. Walhngs t Native Samyda o!o&candra Jacq. Flashes 9 Native

Caricaceae Carica papaya L. St. John’s t Probably native

Cactaceae Opuntia cochenillifera (L.) Mill. Mercers Creek 8 + 0. rubescens Sslm-Dyck St. Phillips S Puerto Rico+ Pilosocereus royeni (L.) Byles & G. Rowley Shirley Heights t Native

Thymelaeaceae Daphnopsis americana (Mill.) J. Johnston ssp. americana (Griseb.) Nevl. Boggy Peak t Native

Lythraceae Lagerstmemia indica L. St. John’s 8 China+ Lawsonia inermis L. Body Ponds S Africa to Asia

Punicaceae Punica granatum L. St. John’s S South Asia+

Lecythidaceae Barringtonia asiotica (L.) Kurz St. Lukes T East Indies+

Rhizophoraceae L. Runaway Bay t Native

Combretaceae Bucida bucems L. Runaway Bay T Native erectus L. Runaway Bay 6 Native racemosa (L.) Gaertn. f. Flashes t Native !lkminalia catappa L. Body Ponds T East Indies

Myrtaceae Eucalyptus camaldulensis Dehnh. St. John’s T Australia+ Eugenia biflora CL.) DC. Boggy Peak t Native E. confusa DC. Boggy Peak 8 Native E. cordata SW. DC. var. sintenisii (Kiaersk.) Krug & Urban Boggy Peak 9 Native E. pseudopsidium Jacq. Boggy Peak 8 Native E. rhombea Krug & Urban Boggy Peak 9 Native Myrcia platyclada DC. Boggy Peak t Native Pimenta racemosa Will.) J. Moore Mercers Creek t Native Psidium guajava L. Wailings t Probably native Syzygium jumbos (L.) Al&on Figtree t Malaya S. malaccense CL.) Merr. & Perry Greencastle T Malaya+

Melastomataceae Miconia laevigata (L.) D. Don Boggy Peak S Native M. mirabilis (Aubl.) L.O. Wms. Body Ponds 9 Native M. pmsina (SW.) DC. Boggy Peak S Native !lktrazygia angustifolia (SW.) DC. Boggy Peak 8 Native

Araliaceae Brassaia actinophylla Endl. Greencastle 8 Australia+ guilfoylei (Bull) L.H. Bailey St. John’s 8 +

9 ‘Ibble 1. - !llce and shrub specks identifkd during field studies in Antigua - Continued Family and species Where first seen Life form* Native range

Sapotaceae Chrysophyllum argenteum Jacq. Boggy Peak Native C. cainito L. St. John’s Greater Antilles+ Manilkara zapota (L.) P. Royen Greencastle Central America+ Sidcmxylon foetidissimum Jacq. Botanic Garden Tropical America S. salicifdium (L.) Lam. Hamiltons Native

Oleaceae Chiananthus compacta SW. Boggy Peak t Native

Apocynaceae Alkmmda cathartica L. V South America 2i.p: Nerium oleander L. 8 Mediterranean+ alba L. St. John’s t Native l?rubaL. St. John’s t Central America+ Rauvolfia viridis Wtid. Jolly Bay Native %bernaemantana citrifdia L. Boggy Peak Native Thevetia peruviana Schum. Bolans S Tropical America

Adepiadaceae Calotropis procem (Ait.) Ait. E Dickinson Bay 8 Old World

COllVOIVUlaCe~ Ipomaea carnea Jacq. ssp. fistulosa (C. Mart.) D. Austin Runaway Bay 8 Tropical America+

Bomglnaceae Bourreria succuknta Jacq. Mercers Creek t Native Co&a alliodom (Ruiz & Pav..) Oken Boggy Peak T Native c. cdlowcco L. Boggy Peek Native C. cumswica (Jacq.) Roem. & Schult. Dark V&y Native C. dent&a Poiret Betty’s Hope Native C. nesophila I.M. Johnston Boggy Peak Native c. lo&g& Lam. Dark Valley Native C. obliqua Wad. Dickinson Bay India C. sebestena L. St. John’s Greater Antilles C. sulcata DC. Boggy Peak Native Heliatrvpium ternatum Vahl Freetown Native

Avicenniaceae CL.) L. Runaway Bay t Native A. schaueriana Stapf. & Leech. ex Mold. Runaway Bay S Native

Verbenaceae Citharezylum spinasum L. Boggy Peak Native Clerodendrum acukatum CL.) Schlecht. Shirley Heights Native Cornutia pymmidata L. Boggy Peak Native Dumnta repens L. Boggy Peak Native Lontano iruducrata L. Cedar Grove Native micnnnem Schauer var. helkrii (Britt.) Mold. Fh?etown Native !lbctona gmndis L.f. Greencastle Southeast Asia+

SoIanaceae Cestrum macmphyllum Vent. Boggy Peak Native Solanum racemasum Jacq. Shirley Heights Native S. mgosum Dunal Bogw Peak Probably native S. torvum SW. Figtree Pantropical

Scrophulariaceae Capraria biflora L. Runaway Bay Tropical America

Bignoniaceae Crescentia cujete L. St. John’s t Probably native Jacamnda mimasifolia D. Don St. John’s t South America+ Kiglia africana &am.) Benth. Botanic Garden T Africa+

10 Table 1. - lhee and shrub species identified during field studies in Antigua - Continued Family and species Where first seen Life form* Native range Spa&&a campanulnta Beauv. St. John’s Central Africa+ Zlzbebuia chrysantha (Jacq.) Nichols. Greencastle -Trinidad+ T. heterophylkz (DC.) Britt. Cedar Grove Native T. rosea (Bertol.) DC. Botanic Garden South America+ Tecoma stuns CL.) Juss. Valley Church Probably native

Rubiaceae Catesboca melanocarpa Urban Shirley Heights 8 Native alba (L.) Hitchc. Boggy Peak S Native ambica L. Body Ponds t Africa+ Erithalis fruticosa L. Shirley Heights t Native Exostema caribaeum (Jacq.) Roem. & Schult. Freetown 8 Native Fammea occidentalis (L.) A. Rich. Boggy Peak 8 Native Gardenia augusta CL.1 Merrill St. John’s 8 Asia+ Guettarda crispiflora Vahl Boggy Peak t Native G. oualifolia Urban Walljngs t Native G. parviflom Vahl Shirley Heights t Native G. scabra (L.) Vent. Boggy Peak t Native G. patens Jacq. St. John’s 8 Tropical America+ Zrora coccinea L. St. John’s 8 East Indies+ I. thwaitesii Hook E St. John’s 8 + Palicourea crocea (SW.) Roem. & Schult. Boggy Peak S Native Psychotria neruOsa SW. Boggy Peak S Native P tenuifolia Sw. Body Ponds 8 Native Randin oculeata L. Boggy Peak Native

Compositae Pluchea camlinensis (Jacq.) G. Don Cedar Grove 8 Native

PandanaCe~ utilis Bory St. Lukes East Indies+

Gramineae Bambusa sp. Collins Dam t Asia+ B. vulgaris Schrad. Swetes t Tropical Asia+

Palmae Acrocomia acukata (Jacq.) Lodd Body Ponds t Native Chrysalidocarpus lutescens (Borg) H. Wendl. St. John’s t Madagascar Cocos nucifem L. Cedar Grove T East Indies E&is guineensis N.J. Jacquin Figtree T Africa Hyophorbe verschuffelti Wendl. St. John’s t Mascarenet Phoenix doctylifem L. Airport T Africa, Asia Z+itchnrdia pacifica Seem. & Wendl. St. John’s t + Roystonea okracea O.F. Cook St. John’s T Probably native R. regia (HBK.) O.F. Cook Botanic Garden T and + Sobal causiarum (O.F. Cook) Becc. Old Road T Puerto Rico Thrinax morrisii H. Wendl. St. John’s Native Veitchia merrillii (Bacc.) H.E. Moore St. John’s t +

Agavaceae Agave kamtto Mill. Johnsons Point 8 Native

Liliaceae Yucca aloifolia L. Runaway Bay t + Dmcaena fragmns CL.1 Ker-Gawl. St. John’s S East Africa+

*t = small tree (sometimes shrub or medium tree), T = medium to large tree, s = shrub (sometimes a small tree). +Species that were observed only under cultivation (not naturalized). *Native means present on Antigua (and usuaIly in other areas) before European settlement.

11 Table 2. -Dee and shrub species observed in Barbuda Family and species Where first seen Life form* Native range

CaeuarinaCeae Casuarina equisetifolia L. Codrington T oceania+

Moraceae Artocarpus altilis (S. Park) Fosb. Codrington T Oceania+ &us benjamina L. Codrington Malaya l? citrifoliu Mill. Codrington ; Native F. elnstica Boxb. Codrington T Tropical Asia+

OIaeaeeae Schoepfia schreberi Gmel. Martello Tower S Native

Pol~ollaCWLe Coccoldm diversifdiu Jm. South of Codrington t Native C. krugii Lindau South of Codrington t Native C. uuifera (L.) L. South of Codrington t Native

Nyctaginaceae Bougainvillin spectabilis Wild. Codrington V Brazil+ Guapira discolor (Spreng.) Little Highlands Native Pisonia subcoraUa Sw. South of Codrington I Native

Ann- Annona muricaia L. Codrington t West Indies+ A. squamosa L. Codrington t West Indies

Lauraceae &o&a coriacea (SW.) Britt. Welsh Point t Native Persea americana Mill. Codrington T Mexico+

Capparaceae Capparis cynophallop~ra L. Welsh Point t Native C. in&a (L.) Bruce Pelican point t Native

Cbryaobalanaceae Chrysobalanus icaco L. Mart&o Tower 8 Native

Easaeeae Rosa spp. Codrington 8 India+

Legumhome-Mimosoideae Acacia farnesiana (L.) Wad. Codrington t Native L.eucaena leucocephalu (Lam.) de Wit South of Codrington t Probably nstive PitheceJlobium unguis-cati (L.) Benth. South of Codrington Native (Jacq.) Merr. Codrington ; Tropical Americs+

Leguminoeae-Caemlpinioideae Czesalpinia bonduc CL.1 Boxb. Martello Tower Native Dekmix &a (Bojer) Raf. Codrington ; Madagascar+ Haemutazylon campechianum L. South of Codrington t Mexico Parkinsonia aculeata L. Codrington t lbxas to Mexico+ sinmea Irwin & Barneby Codrington T Southeast Asia+ lbmarindus indica L. Msrtello Tower T Africa

Leguminosae-Papilionoideae Cajanus cajan CL.1 Huth Codrington S Africa+ Erythrina berteroana Urban Codrington t Tropical Americs+ Gliricidia sepium (Jscq.) Kunth Codrington t Central America+ Pictetia a&e&a (V&l) Urban South of Codrington S Native Sesbania grandiflora CL.1 Pers. Codrington t Tropical Asia+

ErythroxyIaoeae Erytmzylum sp. South of Codrington t Native

12 Tsble 2. - Ibee and shrub species observed in Barbuda- Continued

Where Life Family and species fast seen form* Native range

Zygophyllaceae Guaiacum officinale L. Codrington 8 Native

Rutaceae elemifera L. Highlands t Native Citrus aumntifolia (Christm. 8z Panzer) Swingle Codrington t East Indies+ C. reticulata Blsnco Codrington t South Asia+ C. sinensis (L.) Osbeck Codrington t South Asia+ Wphasia trifolia (Burmf.) P. Wils. Codrington Java+ Zanttiylum flwum Vahl South of Codrington ! Native Z. martinicense (Lam.) DC. Codrington T West Indies+

Simaroubaceae Castela erecta Bnpin South of Codringtan 8 Native

Surianaceae Suriana maritima L. South of Codrington 8 Native

Burseraceae Bursem simaruba (L.) Sarg. Codrington T Native

Meliaceae Azadirachta indica Adr. Juss. Codrington T India Melia azedarach L. Cockington t South Asia’ Swietenia mahagoni (L.) Jacq. South of Codringtan T Antilles

Malpighiaceae Byrsonima lucida Will.) DC. Codrington t Native Malpighia linearis Jscq. salt Pond 8 Native

Euphorbiaceae Codiaeum variegatum (L.) Blume CodrhlgtXXl 8 Sunda Islands+ Croton astroites Drysnd. Spanish Wells S Native Euphorbia &tea Haw. Codrington t Old World+ E. pulcherrima Wad. Codringtdn Mexico+ Hippomane mancinella L. Msrtello Tower f Native Jatropha curcas L. Codrington S central America+ J. gossypiifdia L. Msrtello Tower 8 Native J. multif?da L. Cadrington 8 lkopical America+ Manihot esculenta Grants Codringtan 8 South America+ Phyllanthus epiphyllanthus L. South of Cadriqton 8 Native

Anacardiaceae Comacladia a&fana.ea (L.) Urban South of Codri@on Nat& Mangifem indica L. Codringtan I lkopical Asia+ Spondias purpurea L. Codringtan t Tropical Anu?rica+

CeIastraceae Caasine xylocarpa Vent. Martello Tower t Native Cmssopetalum rhacoma Crantz Highlands a Nativs

SapincIaneae Doo!onaea viscosa CL.1 Jaq. Msrtello Tower Natiws Melicoccus bi,iugatus Jacq. Codringtcm ;

Rhamnaceae Colubrina arborescens (Mill.) Sarg. South of Codrin@n t Nativs Krugiodendron firmum (M. Vshl) Urban Highlands t Native Ziziphus mauritiana Lam. Cadrinmn t Sauth Asia Z. reticuhzta (M. Vahl) DC. South of Codri@an t Native

Malvaceae L. Welsh Point 8 Prabahly native Hibiscus rosa-sinensis L. vsr. rosa-sinensis Cadringtan t IYopical Asia+ Malvwiscus arboreus Cav. Codrington 8 ‘Itopical

13 Table 2. -Ike and shrub species observed in Barbuda- Continued Where Life Family and species first seen form* Native range

Bombacaceae Ceiba pentandra (L.) Gaertn. Codrington T ‘propical America+

Sterculiaceae Melochiu tomentosa L. Highlands 8 Probably native

CancIIaceae Can&o winterana (L.) Gaertn. Codrington t Native

Caricaceae Carica papaya L. Codrington t Probably native

Cactaceae Opuntia rubescens Salm-Dyck Codrington 8 Puerto Rico+ pilosocereus myeni (L.) Byles & G. Rowley Martello Tower t Native

Lythraceae Lagerstmemia in&a L. Codringkm 8 South Asia+

Punicaceac Punica granatum L. Codrington 8 Mediterranean+

Rhizophoraceae Rhizophom mangle L. Welsh Point t Native

Combretaceae Bucidu bucems L. South of Codrington T Native Conocarpus erectus L. South of Codrington 8 Native Laguncularin racenwsa (L.) Gaertn. f. ‘Iwo Foot Bay t Native !i%rminaliu catappa L. South of Codrington T +

Myrtaceae Eugeniu bifira (L.) DC. South of Codrington 8 Native E. cord&a SW. DC. var. sintenisii South of Codrington 8 Native E. rhombea Krug & Urban Highlands t Native Psidium guqjwa L. Codrillgtcn t Tropical America+

Araliaceae Polyscias guilfoylei (Bull.) L.H. Bailey Codrington 8 New Caledonia+

TheophrastaCe~ Jacquinia arborea Vahl Martello Tower 8 Native

Sapotaceae Sidemzylon obovatum Lam. Martello Tower t Native

Apocynal%W Allamandn cathartica L. Codrington V South America+ Nerium oleander L. Codrington 8 Old World+ Plumeriu alba L. Codrington t Native Raumlfiu viridis Wtid. South of Codrington t Native

CcmvcIvuIaceac Zponwea carnea Jacq. sap. fistulosa (C. Mart.) D. Austin Codrington 8 Brazil+

J3craginaceae Argusia gnapholodss CL.1 Heine Welsh Point 8 Native Bourreria succulenta Jaoq. Salt Pond t Native Cordio obliqua Willd. Codrington t India

Avicenniaceae Avicennia germinuns CL.1 L. South of Codrington t Native

14 ‘Ibble 2. - !lYee and shmb species obserwd in Barbuda- Continued

Where Life Family end species fast seen form* Native rsnge

Verbenaceae Citharerylum spinosum L. Codrington t West Indies Clerookndrum aculeatum CL.1 Schlecht. South of Codrington 8 Native Lantana involucrata L. South of Codrington 8 Native

Solanaceae Solanum racenwsum Jacq. Codrington 8 Native.

Scrophulariaceae Capraria biflora L. Rubbish Bay t Old World

Bignoniaceae Crescentia cujete L. Codrington t Tnpical America+ Jacaranda mimosifoliu D. Don Codrington South America+ lhbebuia heterophylla (DC.) Britt. South of Codrington I Native lkcoma stuns (L.) Juss. Codrington 8 West Indies+

Acanthaceae Gmptophyllum pi&urn (L.) Griff. Codrington 8 +

Rubiaceae Erithalis fruticosa L. ltK0 Foot Bay N&h Ernodea littomlis SW. Mertello Tower Native Gardenia augusta (L.) Merrill Codrington Asia+ Zoom coccinea L. Codrington East Indies+ aculeata L. Spanish Well Point Native

Compositae Eupatorium corymbosum Aubl. Highlands S Native Pluchea symphytifoZin (Mill.) Gillis Codrington 8 lYopical America

Palmae Coccothrinar barbadensis (Lodd. ex Mart.1 Becc. Msrtello Tower t Native Cocos nucifem L. Codrington T East Indies Sabal causiarum (O.F. Cook) Becc. Codrington T Puerto Rica+

Agavaeeae Agaue kamtto Miller Highlands 8 Native

l T = medium to large tree, t = small tree (sometimes shrub or medium tree), s = shrub (sometimes a small tree). +Species that were observed only in cultivation (not naturalized).

15 Table 3. -Native and naturalized trees and shrubs observed or collected in Antigua

Suecies Beard Harris M&H IITF Howard

CktSUarillaCeae Casuarina equisetifoliu L. - - - X X

Piperaceae Lepianthes pelt&a CL.) Fkf - - X Piper aduncum L. - - X E! am&go L. - - - X P dilatatum Rich. - - - X l? dussii C. DC. - - X &? reticulaturn L. - X

ulmaceae C&is iguaruxa (Jacq.) Sarg. - - - X lkemn micrantha (L.) Blume - X

Moraceae Cecropiu schreberianu Miq. r - - X Ficus americana Aublet - - - - X l? citrifdia Mill. - X X I? nymphaeifolia Mill. - - - X l? trigonata L. X X X X tiphis mcenwsa CL.) Urban - - - X

Loranthaceae Dendropenwn caribaeus Krug. & Urban - - - X

Olacaceae Wwepfiu arenaria Urban & Britt. - - X - S. schreberi Gmel. - - X X

Polygonaceae Coccobba dioersifolia Jacq. X - X X C. krugii Lindau - - - X C. pubescens L. X X X X X C. szuartzii Meisn. - X - X C. uvifem CL.) L. X X X X C. wwsa L. - X X

Amaranthaceae Irvsine angustifolia Euphrasen - - - X

Nyctaginaceae pisoni5 aculeata L. - - - - X I? fragmns Dum.-Cours. X X X X X p. subcordata SW. X X X X

Phytolaccaceae !&i&stigma octandrum (L.) H. Walter - - - X X

Meniqmmaceae Hyperboena domingensis (DC.) Benth. - - - X

Annonaceae Annona gl5bm L. X X - X X A. montana Macfad. - - - X X A. muricata L. - X X X A. reticulata L. - - X X A. squamosa L. - X X

16 ‘Ihble 3. -Notion and noturalizcd trees and shrubs observed or colkcted in Antigua - Continued source* Snecies Beard Harris M&H IITF Howard

Laumeeae Beikchmkdiu pendulu (SW.) Hemsley - X Licaria parvifdiu (Lam.) Kostermans - X X - Ocotea cernua (Nees) Mez - - - X X 0. coriacea (SW.) Britt. - X X 0. floribanda (SW.) Mez - - X X 0. kucoxylon (SW.) De Laness - - X X 0. membmnacea (SW.) R. Howard X X X X 0. patens (SW.) Nees - - X X capparaceae . Capparis baducca L. - X X c. cynopha8ophora L. X - X C. flexuosa CL.1 L. - - X X C. hnstata Jacq. - X X C. in&a CL.1 Druce X - X X Morisonia americann L. - - X

Moringaaeae Moringa okifem Lam. - X X c-m Chrysobalanus icaco L. - X - X

Leguminoaae-Mimosoideae Acacia farnesiann (L.) Wad. X X X X X A. mucracanthn Humb. & Bonpl. X X X X A. murk&a CL.1 Wdld. - - X A. nilotica CL.1 Delile X X X X A. polyacantha Willd. - X A. tortuosa (L.) Wtid. X X - X X AZbizia kbbeck CL.1 Benth. X X X X Calliandm purpurea CL.1 Benth. - - X Desmanthus virgatus CL.1 Willd. - X X Znga Zaurina (SW.) Wad. X X X X X Lcacacna kucocephulu (Lam.) de wit X X X X X Mimosa cemtonia L. - X X Pithe&obium unguis-cati CL.1 Benth. X - X X Pmsopis juliflom (SW.) DC. X X X X Samanea saman (Jacq.) Merr. - X X

Leguminosae-Caesalpinioideae Bauhinia acukatn L. - - X B. multinervin (Kunth.) DC. - - X X Caesalpiniu bonduc (L.) Boxb. - X X C. cilia&z (Berglus ex Wikstroml Urban - - X C. coriaria (Jacq.1 Willd. - - X C. pukherrima (L.1 Sw. - X X glandulosa var. swartzii GViitrom) Irwin & B=eby X X X D&nix regia @ojer) Raf. - X X Haemaiaxylum campechianum L. X X X X X Hymennea courbaril L. X X X X X Parkinsonia acukata L. - X X Senna atomaria CL.1 Irwin & Barneby - X X S. bocilknis (LX.1 Irwin & Bsrneby var. bacillnris - - X

17 ‘lkble 3. -Native and natbralized trees and shrubs observed or collected in Antigua - Continued source* Species Beard Harris M&H IITF Howard

S. bicapsularis (L.) Roxb. var. bicapsularis - - X X lkmarindus indica L. - - X X

Leguminosae-Papilionoideae Andim inermis (wright) Kunth X X - X X A. sapindoides (DC.) Benth. - - - X Dalbergia ecastaphyllum (L.) Taubert - - X Erythrina velutina Willd. - - - X - Flemingia stmbilifem (L.) Ait. f. - - - X Zndigofem suffruticosa Mill. - - - X X I. tinctoria L. - - - X Lonchocarpus pentaphyllus (Pair.) DC. X X - - - L. violaceus (Jacq.) DC. - - - X X Machaerium lunatum (LX.) Ducke - - X - Piscidia carthagenensis Jacq. X - - X X

Erythroxylaceae Erythroxylon brevipes DC. X - - - E. havanense Jacq. - - X X

Zygophylhceae Guaiacum ofiinale L. X X X

Rutaceae Amyris elemifera L. - - X 7kiphasia trifolia (Burm. f.) P. Wils. - - X X Zanthazylum flavum V&l - - X X Z. martinicense (Lam.) DC. X - X X X Z. monophyllum (Lam.) P. Wils. X - - X X Z. punctatum Vahl - - X Z. spinifa (Jacq.) DC. - X X

Simaroubaceae CasteZa erecta Turpin - - X X Picmmnia pentandm SW. - X - Z’icmsma excelsa (SW.) Planch. - X X Simarvuba amam Aubl. - X X

Surianaceae Suriana maritima L. - - X X

Burseraceae Bursem simaruba (L.) Sarg. X X X X X Dacryodes ercelsa Vahl - - X

Meliaceae Azadirachta indica Adr. Juss. - X X Cedm!a odorata L. X X X X X Guarea glabra Vahl - - X Meh azedamch L. X X Swietenia mahagoni CL.) Jacq. X X X X !Dichilia hirta L. X X

Malpighiaceae Bun&o& glandulosa Ov..) L.C. Rich. X X Byrsonima lucida (Mill.) DC. X X B. spicata V&v.) DC. X - X X B. trinitensis A&. Juss. - - X Malpighia emarginata DC. - X X M. linearis Jacq. - - X X M. martinicensis Jacq. - X

18 Table 3. -Native and naturalized trees and shrubs observed OF collected in Antigua - Continued Source* Species Beard Harris M&H IITF Howard

Krameriaceac Kramerin ixine L. - - - - X

Euphorbiaceae Actinostemon caribaeus Griseb. - - - - X Argythamnia polygama (Jacq.) Kuntze - - X Bernardia corensis (Jacq.) Muell. Arg. - X Chamaesyce articulata (Aubl.) Britt. - X C. mesembrianthemif~ia (Jacq.) Dugand - - X C&on astroites Dryand. - X X C. frowns L. X - - X Drypetes sermta (Maycock) Krug 8~ Urban - - - X Oymnanthes lucio!u Sw. - - X X Hippomane mancinelln L. X X - X X Hum crepitans L. X - X X Jatropha curcas L. - X - J. gossypiifdia L. - - - X X Margaritaria rwbilis LX - - X Pedilanthus tithymaloides CL.) Poit. - - - - X Phyllanthus epiphyllanthus L. - - X X Rich&a gmndis M. Vahl - X Ricinus communis L. - - - X X Sapium caribaeum Urban - - X

Anacardiaceae Ariacardium occidentale L. - X X (L.) Urban - - - X X Mangifem indica L. X X X X X L. X - - X X

Celastraceae Cassine xylocarpa Vent. X X Cmssopetalum rhacoma Crantz - - X X Gyminda latifolia (SW.) Urban - - X Schaefferia frutescens Jacq. - - X X

Sapimlaceae Allophylus racemosus SW. - - X X Dodonaea viscosa CL.) Jacq. - - X Melicoccus bijugatus Jacq. X - - X X Sapindus saponaria L. X X X X

Rhamnaceae Colubrina arborescens (Mill.) Sarg. - X X Krugiodendmn ferreum (M. V&l) Urban - - - X X Ziziphus muuritiuna Lam. - X X 2. reticuluta (M. Vahl) DC. X

Tiliaceae Wumfetta gmndifira Vahl. - - - X - T. rhomboioka Jacq. - - - X T. semitriloba Jacq. X

Malvaceae Abut&m guineense @chum.) E.G. Baker & Exell. X A. umbellatum CL.) Sweet - - - X

19 Table 3. -Native and naturalized trees and shrubs observed or collected in Antigua - Continued Source* Species Beard Harris M&H IITF Howard

Bastardia viscosa (L.) Kunth. var. viscosa - - - X Gossypium barbadense L. - - - - X Malvastrum coromandelianum (L.) Garcke - - X X Sida acuta Burman f. - - - - X S. spinosa L. - - - X Thespesia populnea (L.) Sol. - - - X X Urena lobata L. - - - - X

Bombacaceae Ceiba pentandra CL.) Gaertn. X X X X X Ochroma pyramidale (Cav.) Urban - - - X

Sterculiaceae Guasuma ulmifolia Lam. X - X X X Melochia tomentosa L. X - - X X indica L. - - X

Ochuaceae Ouratea guildingii (Planch.) Urban - - - X X

Clusiaceae Calophyllum calaba L. - - - X X Clusia major L. - - X X X

Can*eae Canella wintemna (L.) Gaertn. - X X X X

Flacourtiaceae Casearia o!ecandra Jacq. - - X X C. guianensis (Aubl.) Urban X - - - Pmckia crucis P. Browne ex L. - - - X Samyda o!odecandm Jacq. - - X X

Caricaceae Carica papaya L. - X X X X

ThymeIeaceae Daphnopsis americana (Mill.) J. Johnston sap. caribaea (Griseb.) Navl. X - X X X

LythraCeae Lawsonia inermis L. - - X X

Rhizophoraceae Rhiwphora mangle L. X X - X X

Combretaceae B&da bucems L. X X X X X Conocarpus erectus L. X X - X X Laguncularia racemosa (L.) Gaertn. E X X X X

MyrtaCeae Eugenia axillaris (SW.) Willd. - - X E. bifZom CL.) DC. - - - X X E. confusa DC. - - X X E. com!ata (SW.) DC. var. sintenisii (Kiaersk.) Krug & Urban - - X X E. monticola (SW.) DC. - - - X E. pseudopsidium Jacq. - - X

20 Table 3. -Native and naturalized trees and shrubs observed or collected in Antigua - Continued Source* Species Beard Harris M&H IITF Howard

E. rhombea Krug & Urban - - X X E. uniflora L. - X Myrcia citrifolia (Aubl.) Urban var. imrayana Greseb. X - - X M. o!efZexa (Pair.) DC. - - X M. splendens (SW.) DC. - - X M. platyclada DC. - - X - Myrcianthes fragrans (SW.) McVaugh - - X Myrciaria floribunda (West) 0. Berg. - - X Pimenta mcemosa (Mill.) J. Moore - - X X Psidium guajava L. X X X X P longipes (O.Berg) McVaugh var. orbiculare (O.Berg.) McVaugh - - - X Syzygium jambos CL.) AIston - X X lkrminalia catappa L. X X X

Melastomataceae Clidemia hirta (L.) D. Don - - - X Miconia laevigata (L.) D. Don - - X X M. mirabilis (Aubl.) L.O. Wms. - - X X M. prasina (SW.) DC. - X M. striata (M. Vahl) Cog-n. - - - X M. trichotoma (Dew.) DC. - - X Mouriri domingensis (lkss.) Spach - X i’ktrazygia angustifolia (SW.) DC. X - X X T. discolor (L.) DC. - - X

Myrsinaceae Ardisia obouata Ham. - X

Plumbaginaceae Plumbago auriculata Lam. - - - X P: scandens L. - - _ - X

Sapotaceae Chrysophyllum argenteum Jacq. X - X X Micropholis guyanensis (A. DC.) Pierre - - _ - X Sideraxylon foetidissimum Jacq. X - X X S. obovatum Lam. - - - - X S. salicifol’ium (L.) Lam. - X X

Oleaceae Chionanthus compacta SW. X - x x X Forestiera segregata (Jacq.) Krug. & Urban - - - - X

Apocynaceae Allamanda cathartica L. - - X X Plumeria alba L. - X X Rauvolfia viridis Willd. - X X lkbernaemontana citrifolia L. X - x x X Thevetia peruviana (Pers.) Schum. - - X X

Aeclepiadaceae Calotropis procem (Ait.) Ait. f. - X X

Boraginaceae Argusia gnaphalodes CL.) Heine - - X Bourreria succulenta Jacq. X - x x X Cordia alliodora (Ruiz & Pav.) Oken X - X X C. collococca L. - - X X

21 Table 3. -Native and naturalized trees and shrubs observed or collected in Antigua - Continued Source* Species Beard Harris M&H IITF Howard

C. curasavica (Jacq.) Boem. & Schult. - - - X X C. o!entata Poir. - - - X X C. globosa (Jacq.) Kunth - - - X C. laevigata Lam. - - - X - C. nesophila I.M. Johnston - - - X X C. obliqua Willd. - - - X X C. sebestena L. - - - X X C. sulcata DC. X - X X X Heliotmpium micmphyllum SW. ox Witstrom - - - - X H. ternatum Vahl - X - Rochefortia acanthophora (DC.1 Griseb. - - - - X R. spinosa (Jacq.1 Urban - X lburnefortia f&flora Griseb. - - - - X T. volubilis L. - - X

Avicenniaceae Avicennia germinans CL.1 L. X X - X X A. shaueriana Stapf. & Leech. ex Mold. - - X

Verbenaceae Aegiphila martinicensis Jacq. X - - - X Citharerylum spinosum L. X - X X Clemdendrum aculeotum CL.1 Schlecht. - - X X Cornutia pyramidata L. - - X X Duranta repens L. - - X X Lantana camara L. - X - - - L. involucrata L. X - X X L. urticifolia Mill. - - - X Lippia alba (Mill.) N. E. Br. - - X X Ocimum gratissimum L. - - - - X Stachytarpheta cayennensis (L.C. Rich.) Vahl - - X VitQ divaricata SW. - - - - X

Solanaceae Acnistus arborescens CL.1 Schlecht. - - - - X suaveolens (Willd.1 Berth. & Presl. - - - X Brunfelsia americana L. - - - X Cestrum laurifolium L’lterit. - - - - X C. macrvphyllum Vent. - - X Datum mete1 L. - - - X Lycium americanurn Jacq. - - - X Solanum racemosum Jacq. X - - X X S. rugosum Dunal - - X - S. torvum SW. - - X X

Bignoniaceae Amphitecna latifolia (Mill.) A. Gentry X - X - Cmscentia cujete L. - - - X X lhbebuia hetervphylla (DC.) Britt. X X X X X !&coma starts (L.) Juss. X - X X

Cactaceae Pilosocereus royenii CL.1 Byles 8z G. Rowley X X - X X

Acanthaceae Justicia carthaginensis Jacq. - - X X

22 Teble 3. -Native and naturalized trees and shrubs observed or collected in Antigua - Continued Source* Species Beard Harris M&H IITF Howard

Justicia carthaginensis Jacq. X X J. eustachiana Jecq. - - - X X Oobntonema nit&m (Jecq.) Kuntze - - X Oplonia microphylla (Lam.) Steern - - X

Myoporaceae Bontia daphnaides L. - - - X

Rubiaceae Antirhea acutata (DC.) Urban - - X melanocarpa Urban - X X Chiococca al&z (L.) Hitchc. - - - X X C. parvif&a Wiillschl. ex Griseb - - X Erithalis fruticosa L. X X E. odorifem Jacq. - - - X Ernodea littomlis SW. - X Exostema caribaeum (Jecq.) Roem. & Schult. X X X X X Fammea occidentalis (L.) A. Rich. X X Gonzalagunia hirsuta (Jecq.) Schum. - - - X Guettarda crispifim VahI - - - X G. odor&a (Jacq.) Lam. - - - X G. ovalifdia Urban X G. parvifim Vehl - - - X G. scabra (L.) Vent. X X X X X Zxam ferrea (Jacq.) Benth. - X Palicourea crvcea (SW.) Roem. & Schult. - - X X Psychotria obmingensis Jacq. - X l? micrvdan (DC.) Urban - - - X l? nervosa SW. - - X X Z? tenuifolia SW. - - - X X Randia aculeata L. X X X Spermacoce bahamensis (Britt.) Howard - - - X Strumpfia maritima Jacq. X

Compositae Eupatorium corymbosum Aubl. - X E. odamtum L. - - X E. sinuatum Lam. - - - - X Pluckea carolinensis (Jecq.) G. Don - X X Verrwnia alkicaulis Pers. - - - - X WeoMia calycina Rich. - X

Palmae Acmcomia aculeata (Jacq.) Lodd. X - - X X Coccothrinax barbadensis (Lodd.) Becc. X X X X Cocos nucifera L. - X X X Elaeis guineensis N.J. Jacquin - - - X Phoenix dactylifem L. X Roystonea oleracea O.F. Cook X - X Sabal causiarum (O.F. Cook) Becc. X X

Agavaceae Agate kamtto Mill. X X X X

*Beard = Beard 1949; Harris = Harris 1960; M&H = Mackler end Hannah 1988; IITF = International Institute of Tropical Forestry survey; Howard = Howard 1987; 1988; 1989a, 1989b.

23 lhble &-Native and naturalized tree and shrub specks observed or collected in Barbuda Source* Family and species Beard Harris IITF Howard

Ulmaceae Celtis iguanaea (Jacq.) Sarg. - - X Moraceae Ficus citrifolia Mill. X X X Olacaceae Schoepfia schreberi Gmel. - - X X Polygonaceae Coccoloba dioersifdia Jacq. X X X X C. krugii Lindau X - X X C. pubescens L. - - - X C. uuifera (L.) L. X - X X Nyctaginaceae Pisonia discolor Spreng. - - X I? fmgrans Dum.-Cours. X X X E! subcordata SW. X X X X PhYtolaccaceae Rchostigma octandrum (L.) H. Walter - - X Annonaceae Annonu gkbm L. X X - X A. squamosa L. - - X X Lauraceae Ocotea coriacea (SW.) Britt. - X X Capparaceae Capparis baducca L. - - X C. cynophallophora L. X - X X C. flezuosa (L.) L. X - - - C. in&a (L.) Druce - X X Cbqsobalanaceae Chrysobalanus kaco L. X X X Leguminoaae- Mimosoideae Acacia farnesiana (L.) Willd. - - X X A. macmcantha Humb. & Bonpl. - - X A. nilotica (L.) Delile - - X A. tortuosa (L.) Willd. - X - X Leucaenn kucocephala (Lam.) de Wit - - X X unguis-cati CL.) Benth. X - X X Rusopis juliflom (SW.) DC. - - X Leguminosae-Caeaalpinioideae Caesalpinia bonduc (L.) Roxb. - X C. pukherrima CL.) Sw. - - X Chamaecrista glandulosa var, swartzii (W&&ram) Irwin & Bameby X - - X Haematoxylon campechianum L. X X X X Senna bicapsularis (L.f.) Rnxb. ~81: bicapsularis - X Thmarindus indka L. X X X Leguminosae-Papilionoideae Indigofera tinctoria L. - - X L.on&ocarpus pentaphyllus (Pair.) DC. X - Pictetiu acukata (V&l) Urban - X Sophom tomentosa L. - X ZYeophyllace~ Guaiacum ofiinak L. - X X X Rutaceae Amyris ekmifera L. X X X X Zanthoxylum flavum Vabl X X X 2. spinifer (Jacq.) DC. - X SilUaroubaCe~ Castela erecta Turpin - X X SUrianaCe~ Suriana maritima L. X - X X Burseraceae Bursem simaruba CL.1 Sarg. X X X X

24 Table I.-Native and naturalized tree and shrub species observed or collected in Barbuda - Continued

SOUlTd Family and species Beard Harris IITF Howard Meliaceae Azadirachta indica Adr. Juss. - X - Swietenia mahagoni CL.) Jacq. - - X X Malpighiaceae Bunchosia glnndulosa (Cav.) L.C. Rich. - - - X Byrsonima lucida (Mill.) DC. X - X X Malpighiu linearis Jacq. X X X X Euphorbiaceae Chamnesyce articulata (Aubl.) Britt. X - - X C. mesembrianthemifoliu (Jacq.) Dugand - - X C&on astrvites Dryand. - X X c. plvem L. X X - X lucida Sw. - X Hippomane mancinella L. - X X Hum crepitans L. - - X Jatropha gossypiifdia L. - X X Phyllunthus angustifolius (SW.) SW. X - l? epiphyllanthus L. X X Ricinus communis L. - - X Anacardiaceae Comocludia dodonaea (L.) Urban X X X X Cassine xylocaqa L. - - X Cnxsopetalum rhacoma Crantz - X X Gyminda &if&a (SW.) Urban X X Dodonaea elueagrwides Rudolphi - - X D. viscosa CL.) Jacq. X X X X Melicoccus bijugatus Jacq. - - X X Sapindus saponaria L. - - X Rhamnaceae CoZubrina arborescens (Mill.) Sarg. - X X Krugiodendmn ferreum (M. Vahl) Urban - X X Ziziphus mauritinna Lam. X - Z. reticulata (M. Vahl) DC. - - X X Malvaceae Gossypium barbadense L. - X X Sidu spinosa L. - X Sterculiaceae Melochia tomentosa L. - X X CanelIaceae Can&z wintemna CL.) Gaertn. X X X X Caricaceae Carica papaya L. - X X Cactaceae Opuntia cochenillifem CL.) Mill. - - X Pilosocereus royeni CL.) Byles & G. Rowley X X X Rhizophoraceae Rhiwphom mangle L. X - X X Combretaceae Bucida buceras L. X X X X Comarpus erectus L. X X X Laguncularia racemosa (L.) Gaertn. f. X X X MyrtaCeae Eugenia axillaris (SW.) Wad. X - X E. bifim (L.) DC. - X - E. cord&a (SW.) DC. var. sintenisii (Kiaersk.) Krug & Urban X X X E. rhombea Krug 19, Urban - X X Pimenta mcemosa Nil.) J. Moore X X

25 l?&le 4.-Native and naturalized tree and shrub species observed or collected in Barbuda - Continued Source* Family and species Beard Harris IITF Howard Melastomataceae !llzmonea baxiana (Mold.) Howard - - X Theophrastaceae Jacquinia armillaris Jacq. X - X X Plumbaginaceae Plumbago scandens L. - - X Sapotaceae Sidenxylon obovatum Lam. X X X X S. salicifolium (L.) Lam. X X - X Oleaceae Fore&era segregata (Jacq.) Krng. & Urban - - - X Apocynaceae Plumeria alba L. X X X X Rauvolfia viridis Willd. - - X X Asclepiadaceae Calotropis procera (Ait.) Ait. f. - - - X Argusia gnaphalodes (L.) Heine - X X Bourreria succulenta Jacq. X - X X Cordia o!entata Poir. - - - X C. globosa (Jacq.) Kunth - - - X C. mawostachya Ftoem. & Schult. X - - - C. nesophila I.M. Johnston X C. obliqua Wtid. - - X - C. sebestena L. - - - X Heliotrvpium micmphyllum SW. ex Wiksttim - - - X Rochefortia acanthophom (DC.) Griseb. X !lbumefortia volubilis L. - - - X Avicenuiaceae Avicennia germinans CL.) L. - - X X Verbenaceae Cithamylum spinosum L. - - X X Clenxdendrum aculeatum (L.) Schlecht. X X X X Lantana camara L. X - X L. iruolucrata L. X X X X Solanaceae Solanum racemosum Jaca, X X Bignoniaceae !Ihbebuia hetervphylla (DC.) Britt. X X X X T. lepidota (HBK.) Britt. - X Acanthaceae Justicia carthaginensis Jacq. - - X J. eustachiana Jacq. - - - X Rubiaceae Catesbaea melnrwcarpa Urban - - X Chiococca parvifdia Wiillechl ex Griseb - - - X Erithalis fruticosa L. X - X X Ernoaka littoralis SW. X X X Guettarda o&rata (Jacq.) Lam. - X G. scabm CL.) Vent. X X Hamelia patens Jacq. - - X Randia aculeata L. X X Spermacoee bahamensis (Britt.) Howard - X Strumpfia maritima Jacq. X X Goodeniaceae Scaewlu Plumieri (L.) Vahl X X Compositae Baccharis dioica Vahl - X arborescens CL.) DC. - X Eupatorium corymbosum Aubl. - - X

26 lhble 4.-N&& and naturalized tree and shrub specks observed or collected in Barbuda - Continued Source* Femilv end species Beerd Harris IITF Howard Gundlachia corymbosa (Urban) Britt. - - X Pluchea carolinensis (Jecq.) G. Don - - X X We&la calycina Rich. - - X PalmIlt? Coccothrinax barbadensis &odd.) Becc. x - x X Thrinax morrisii H. WendI. - - X Agavaeeae Agaue kamtto MiII. x x x X *Beard = Beerd 1949; Harris = Harris 1960; IITF = the InternationaI Institute of Tropical Forestry survey; Howard = Howard 1987; 1988; 1989a, 1989b.

Teble 5.-Native canopy tree species listed in Howard (1979; 1988; 1989a, 1989bJ identified in the International Institute of Tropical Forestry (ZZTF) survey of Antigua and Barbuda, and appearing in both surveys Both Location end species name Howard IITF Surveys

Antigua Acrvcomia acukata (Jacq.) Lodd. X Andim inermis (Wright) Kunth X Avicennia germinans (L.) L. - X Beikchmiedia pendula (SW.) Hem&y Bourreria succuknta Jecq. X Bucida bucems L. X (L.1 Serg. - X Byrsonima spicata (Cav.) DC. X Cakphyllum calaba L. X Canella winterana (L.) Geertn. X Cecropia schmberiana Miq. Cedrela &rata L. X Ceiba pentandra CL.) Gaertn. X Clusia major L. - X Coccoloba dioersifdia Jecq. X C. pubescens L. X Coccothrinax barbadensis &odd.) Bacc. X Co&a alliodora (Ruis & Pav.) Oken X C. collococca L. - X C. laevigata Lam. X C. sukata DC. X Dacryodes exceka V&I - Ficus citrifdia MiII. X F. trigonata L. - X Guaiacum ofjicinak L. - X ulmifolia Lam. - X Hippomane mancinella L. X Hum crepitans L. X L. X Inga kurina &v.) Wtid. X Laguncularia racemosa CL.) Geertn. f. X Lmchocarpus violaceus (Jecq.1 DC. X Melicoccus buugatus Jacq. X Ochroma pyramidale (Cav.) Urban X Ocotea membmnacea (SW.) R. Howard X Pimenta mcemosa (MiII.) J. Moore X Pisonia subco?wka SW. X Rhiwphora mangle L. X Sapindus saponaria L. Sapium caribaeum Urban Siokmzylon foetidissimum Jacq. X Simarouba amam AubIet X

27 Table 5.-Native canopy tree species listed in Howard (1979; 1988; 1989a, 1989b), identified i,n the International Institute of lkopical Forestry (IITF) survey of Antigua and Barbuda, and appearing in both surveys-Continued

Both Location and species name Howard IITF Surveys

Spondias mombin L. _ X ZMebuia heterophylla (DC.) Britt. - X Vitex divaricata SW. x - - Zanthoxylum flwum Vahl - X 2. martinicense (Lam.) DC. - X

Barbuda Avicennia germinans (L.) L. - X Bourreria succulenta Jacq. - X Bucida buceras L. - X Bursera simaruba (L.) Sarg. - X Canella winterana (L.) Gaertn. - X Coccoloba dioersifolia Jacq. - X C. pubescens L. x - - Ficus citrifolia Mill. - X Guaiocum officinale L. - X Hippomane mancinella L. - X Krugiodendron ferreum (M. Vahl) Urban - X Laguncularia racemosa (L.) Gaertn. f. - X Ocotea coriacea (SW.) Britt. - X Pisonia subcordata SW. - X Rhizophora mangle L. - X Sapindus saponaria L. x - - Sideroxylon salicifolium (L.) Lam. x - - Rzbebuia heterophylla (DC.) Britt. X Zantharylum fhwum Vahl - X

LITERATURE CITED oneae. Jamaica Plain, MA: Arnold Arboretum, Har- vard University. 604 p. Vol. 5. Beard, J.S. 1949. The natural vegetation of the Wind- Howard, Richard A. 198913. Flora of the Lesser Antilles: ward and Leeward Islands. Oxford For. Mem. 21. Leeward and Windward Islands. Part 3: Dicotyle- Oxford, UK: Oxford University. 192 p. oneae. Jamaica Plain, MA: Arnold Arboretum, Har- Fournet, Jacques. 1978. Flore illustree des phan- vard University. 658 p. Vol. 6. erogames de Guadeloupe et de . Paris: Liogier, Henri AIain; Martorell, Luis F. 1982. Flora of Institut National de la Recherche Agronomique. Puerto Rico and adjacent islands: a systematic 1,654 p. synopsis. Rio Piedras, PR: Editorial de la Univer- Grisebach, A.H.R. 1963. Flora of the British West sidad de Puerto Rico. 342 p. Indian islands. New York: J. Cramer-Weinheim. 789 Little, Elbert L., Jr.; Wadsworth, Frank H. 1964. Com- P* mon trees of Puerto Rico and the . Harris, David R. 1960. The vegetation of Antigua and Agric. Handb. 249. Washington, DC: U.S. Depart- Barbuda, Leeward Islands, The West Indies. Berke- ment of Agriculture. 548 p. ley, CA: Department of Geography, University of Little, Elbert L., Jr.; Woodbury, Roy 0.; Wadsworth, California, preliminary report of field work. 88 p. Frank H. 1974. Trees of Puerto Rico and the Virgin Howard, Richard A. 1979. Flora of the Lesser Antilles: Islands. Second volume. Agric. Handb. 449. Wash- Leeward and Windward Islands. Monocotyledoneae. ington, DC: U.S. Department of Agriculture. 1,024 Jamaica Plain, MA: Arnold Arboretum, Harvard Pm University. 586 p. Vol. 3. Mackler, R.; Hannah, P. 1988. The forest at Wallings Howard, Richard A. 1988. Flora of the Lesser Antilles: Reservoir, Antigua, West Indies, 44 years after J.S. Leeward and Windward Islands. Part 1: Dicotyle- Beard’s study: conjecture on the patterns of change. oneae. Jamaica Plain, MAz Arnold Arboretum, Har- Commonwealth Forest Review. 67(3): 269-278. vard University. 673 p. Vol. 4. Schuchert, Charles. 1935. Historical geology of the Howard, Richard A. 1989a. Flora of the Lesser Antilles: Antillean-Caribbean region. New York: John Wiley & Leeward and Windward Islands. Part 2: Dicotyle- Sons, Inc. 811 p.

28 *U.S.GPO:1994-565-016/00009 Francis, John; Bivera, Carlos; Figueroa, Julio. 1994. Toward a woody plant list for Antigua and Barbuda: past and present. Gen. Tech. Rep. SO-102. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 28 p. Beginning in the 17th century, the forest lands of Antigua and Bar- buda were cleared for agriculture, burned, cut over, and grazed severely. A survey by personnel of the USDA Forest Service, Interna- tional Institute of Tropical Forestry and previous surveys were used to assemble lists of native and exotic woody plants. A large majority of the original woody flora still grows on both islands. Keywords: Botanical survey, extinction, Lesser Antilles native spe- cies, woody flora.

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