origin and distribution of adventive alien flowering
TRANSCRIPT
ORIGIN AND DISTRIBUTION OFADVENT1VE ALIEN FLOWERING PLANTS
IN HAWAI`I
Lyndon Wester
ABSTRACT
Hawai`i has more than 800 known species of alien plants that arereproducing without direct human assistance. During this century, new plantspecies have been arriving at the rate of about five per year. Patterns ofplant collecting in time and space suggest that other species await discoveryand that data on the ranges of known species are far from satisfactory. TheNew and Old World tropics have supplied most of the adventive alien plants inHawai`i, although other places, particularly those with which Hawai`i hasclose economic and cultural ties, have contributed significant numbers inrecent years.
INTRODUCTION
The nature and derivation of native floras have long attracted theattention of biologists, but the rearrangement and wholesale homogenizationof the world's biota caused by humans in the last 10,000 years has arousedcomparatively little interest until recently. Contributing reasons forthis are that alien plant invasions are generally short-lived, and thatpopulations depend upon continued disturbance, are unstable, and aredetermined to a large extent on chance dispersal. Furthermore, there is abuilt-in preference for what is natural and rare and a prejudice againstthe commonplace and alien among biologists, except perhaps where economicfactors are involved.
To protect natural ecosystems, agriculture, and even urban areas fromundesirable invasion from alien plants, reliable data about the alienspecies present are needed. Information about how plants were introducedis helpful to plan means to limit further additions to the alien flora.Understanding generalized patterns of establishment and spread is alsouseful in managing problems that already exist.
99
Wester \ Origin and Distribution of Alien Plants 100
DEFINITIONS AND DATA SOURCES
Considerable confusion exists about terminology applied to introducedplants and weeds (Frenkel 1984; Smith 1985), so it is important from theoutset to define, in exact terms, the subject of this study. The focus ofthe work here will be plants introduced to Hawai`i that have shown theability to reproduce themselves, either sexually or vegetatively, and tospread without direct human assistance. Excluded, therefore, are theintroduced plants cultivated as crops or ornamentals, although it must berecognized that new recruits for the naturalized alien flora often comefrom the plants cultivated intentionally by humans.
Weeds may be categorized according to the degree of human disturbancethey require for survival (Harlan and de Wet 1965; Hart 1976). Slightdisturbance, such as a small opening in the canopy of a forest, may besufficient to ensure the establishment of some species. However, ruderals,the plants of waste places, or the weeds of crops depend upon continualdisturbance. Some species may depend for their survival on humans tosupply water, to act as local dispersal agents, to set or extinguish fires,or to eliminate competitors by mowing, trampling, or spraying. All alienplants that are reproducing and actively spreading, or that have done so inthe past, were considered in this survey. The degree of direct or indirectdependence on humans for the survival of the populations was not taken intoaccount. Excluded, however, were plants confined to cultivation orpersisting without showing any tendency to spread after abandonment.Opinions as to whether species are disliked, welcomed as valued additionsto the flora, or simply tolerated were not taken into consideration.
There is no infallible way to determine whether a plant is indigenous oralien by its external characteristics. However, if a plant was notrecorded in early botanical surveys, has no closely related indigenousrelatives, seems incapable of dispersing over great stretches of open oceanwithout human assistance, and favors disturbed habitats created by humans,it seems reasonably safe to assume the species is an alien. There issubstantial agreement of opinion about the provenance of most species inHawai`i; however, there is some disagreement about a small number ofspecies that might be indigenous, Polynesian introductions, or very earlyEuropean introductions. Because many of these species have spread wellbeyond cultivation, this question must be addressed in more detail.
Exact and reliable information on the time and circumstances of anintroduction is rarely available. Furthermore, many adventive aliens werefirst introduced as crop or ornamental species and only later did theyescape from cultivation. It is rare for escapes to be recorded. In fact,a plant may spread for many years and build up a large population before itis noticed. For example, Oxyspora paniculata was observed growing as aplanted species in a forest reserve in 1958 when a collector commented,"this plant does not spread, does not seem to be a pest" (B.P. BishopMuseum herbarium specimen 58128). Only in 1984 was a serious infestationfound in the same area (Obata 1985). In the course of the present study,the earliest record of each alien species was sought, regardless of thesituation where it was first found. Sources for this information included
Wester \ Origin and Distribution of Alien Plants 101
published floras, checklists, other written accounts, accession records ofarboreta or botanical gardens, and botanical specimens. The majority offirst records (72%) are from herbarium specimens. The B.P. Bishop Museumherbarium was the most important source. However, other herbariacontaining the collections of 19th century botanists were also searched forplants believed to be early introductions to Hawai`i. The collectionsconsulted were U.S. National Herbarium, Gray Herbarium of HarvardUniversity, British Museum of Natural History, Royal Botanical Gardens atKew, Museum National d'Histoire Naturelle, Paris, UniversitatisFlorentinae, Instituto Botanico, Academy of Sciences of the U.S.S.R. inLeningrad, Harold L. Lyon Arboretum at University of Hawai`i, and Departmentof Botany, University of Hawai`i.
EARLY ALIEN INTRODUCTIONS
Most of the main food and fiber crops used by the Hawai`ians lackmechanisms of dispersal that would enable them to reach Hawai`i by naturalprocesses. Handy and Handy (1972) believed that only 14 species wereactually introduced by the Polynesians (Table 1), whereas St. John (1973,1978) and Nagata (1985) have a rather longer list of candidates. Includedare weedy species, which may have been introduced accidentally, and strandtrees of widespread distribution in the Indo-Pacific (Merrill 1946). Thestatus of several other plants, conventionally considered aliens introducedby Europeans, must be reconsidered in the light of the rediscovery ofcollections made by the first botanists to visit Hawai`i. As there isconsiderable controversy surrounding these species, each will be discussedseparately.
`Ahu`awa, (Mariscus javanicus)The first record of ahu`awa in Hawai`i comes from the collection by Lay
and Collie, botanists on the Beechey voyage, in 1827 (Hooker and Arnott1965). The plant is commonly found on sandy beaches, in taro (Colocasiaesculenta) patches, and in marshes and may be one of the indigenous, pan-tropical wetland species whose seeds were widely dispersed by waterbirds;or it could have been carried as an accidental passenger on taro corms.
Kili`o`opu (Kyllinga nemoralis)Mann (1866) was apparently the first botanist to collect kili`o`opu,
which was regarded as indigenous by Hillebrand (1965) and "longestablished" by St. John (1973). It was reported in Tahiti in 1769 byBanks and Solander (Merrill 1954) and, like Mariscus javanicus, mayeither be indigenous or one of those species spread accidentally by thePolynesians.
Kukaepua`a (Digitaria setigera)The botanist David Nelson, who accompanied Cook to Hawai`i, was the
first to collect botanical specimens of this plant in 1779 (St. John1978). Since it is frequently found growing around pig pens and itsHawai`ian name, kukaepua`a, implies an association with pigs, it mayhave been carried along accidentally when the Polynesians brought the pigto Hawai`i (Degener 1962).
Wester \ Origin and Distribution of Alien Plants 102
Table 1. Plants associated with Hawai`ian culture and land use.
Handy & St. John St. John NagataHandy 1972 1973 1978 1985
MAIN ECONOMIC PLANTSAleurites moluccanaAlocasia macrorrhizaArtocarpus altilisBroussonetia papyriferaCocos nuciferaColocasia esculentaCordyline fruticosaCurcuma longaDioscorea alataDioscorea bulbiferaDioscorea pentaphyllaIpomoea batatasLagenaria sicerariaMusa spp.Nasturtium sarmentosumPiper methysticumSaccharum officinarumSchizostachyum glaucifoliumSyzygium malaccenseTacca leontopetaloidesTephrosia purpureaZingiber zerumbet
STRAND TREES AND SHRUBSCalophyllum inophyllumCordia subcordataMorinda citrifoliaThespesia populnea
ACCIDENTAL INTRODUCTIONSDigitaria setigeraIndigofera suffruticosaLudwigia octovalvisMerremia aegyptiaOxalis comiculataUrena lobataWaltheria indica
****9*
*
*
*
*
*
* *
* *
*
*
* *
* *
* *
*
* *
*
*
* *
* *
* *
* *
*
*
*
*
*
OTHER POSSIBLE POLYNESIAN INTRODUCTIONSKyllinga nemoralisLycopersicon pimpmettifoliumLythrum maritimumMariscus javanicusSolanum americanumVigtia adenantha
Wester \ Origin and Distribution of Alien Plants 103
`Iniko (Indigofera suffruticosa)Hillebrand (1965) noted that Iniko was one of the most common weeds in
the Islands but believed that it had been introduced in 1836 for theproduction of indigo. Nelson, the botanist with Cook, listed"Indigofera, Indigo" in a report, which Beaglehole's informant assumedwas Indigofera suffruticosa (St. John 1978; Beaglehole 1967). Noherbarium specimen has been found to substantiate this observation. Norecord of the use of the plant by Hawai`ians has been found, and it is hardto imagine that such a useful dye plant would have escaped their attentionhad it been present in prehistoric times. It therefore seems likely thatthe plant was not a Polynesian introduction and that Nelson was mistaken inincluding it on his list.
Vigna adenanthaThis plant is probably a native of South America but was reported by
early botanists in many Polynesian islands as well as in easternAustralia. Plant specimens from Hawai`i bearing this name, including thosecollected by Menzies in 1794 (British Museum of Natural History), Macrae in1825 (British Museum of Natural History), Nuttall in 1835 (British Museumof Natural History), and Remy sometime between 1851 and 1855 (GrayHerbarium), can be found in a number of herbaria. Mann (1866) believed thespecies to be a Polynesian introduction, and Hillebrand (1965) reportedthat the Wilkes Expedition, which visited Hawai`i in 1840, found it on"Diamond Hill." A search for this specimen in the U.S. National Herbarium,where the collections from the expedition are said to be preserved, did notlocate it. No more recent reports based on voucher specimens have beenfound, so the extant specimens should perhaps be reexamined to ensure thatthe earlier botanists did not confuse this species with other taxa.
Yellow wood sorrel (Oxalis corniculata)Yellow wood sorrel or `ihi`ai is a pantropical weed collected by Banks
and Solander in Tahiti and was observed but not collected by Nelson inHawai`i in 1779 (St. John 1978). The fact that seeds have been found inprehistoric archaeological sites makes it clear that the plant was notintroduced by Europeans (Kirch 1985) but was either indigenous or anintroduction, probably accidental, by the Polynesians.
Hau (Hibiscus tiliaceus)Hau was a valuable plant for the Hawai`ians and was widely cultivated.
For this reason, it has been considered a likely Polynesian introduction(Krauss 1974). However, it is a widespread strand species whose seeds areadapted to long-distance dispersal over the ocean (Merrill 1946). Theplant is common along sheltered coasts and in wet upland gulches in Hawai`itoday. It is quite likely that it reached Hawai`i without humanassistance.
Milo (Thespesia populnea)The Polynesians held this tree in high regard as a shade tree and also
used it for tannin, dye, medicines, and wood (Krauss 1974). It was widelyplanted around houses and may be found persisting and occasionallyestablishing along the coast today. As a plant of the coastal strand, it
Wester \ Origin and Distribution of Alien Plants 104
is adapted to transoceanic dispersal and hence may be indigenous, althoughSt. John (1973) and Nagata (1985) judge it to be a Polynesian introduction.
Aramina (Urena lobata)Nelson observed "Urena, Indian Mallow" in Hawai`i in 1779 but did
not take a specimen of the plant. This was assumed to be the native opuhe,Urera sandwicensis, in the family Urticaceae; however St. John (1978)argued that Nelson, a competent botanist, would not have confused plants insuch distinctively different families. Merrill (1954) stated that Banksand Solander found Urena in Tahiti. Seeds of this plant could havebeen carried by birds to Hawai`i, or it might have been a Polynesianintroduction.
Kamani (Calophyllum inophyllum)This tree was cultivated by Hawai`ians and had many uses. It is not
found naturalized to any great extent in Hawai`i today but may have been soin the past. It is generally regarded as a Polynesian introduction, butsince its natural habitat is the coastal strand and its seeds are adaptedto ocean dispersal, the possibility that it is indigenous should not beentirely discounted.
Carrot (Daucus pusillus)Hillebrand (1965) believed the American carrot was undoubtedly in the
Islands before European discovery. A collection made by Menzies (BritishMuseum of Natural History) provides support for this conclusion. It mightpossibly have been brought to the Islands by migrating birds and thus be anindigenous species.
`Uhaloa (Waltheria indica)`Uhaloa has weedy properties and is considered by most to be
indigenous. It was collected by Nelson (St. John 1978) and has small seedsthat might adhere to birds; it may have been transported to Hawai`i by thismeans.
Pukamole (Lythrum maritimum)Today pukamole has been generally regarded as a European introduction in
Hawai`i. It was collected by many of the early botanists, includingMenzies in 1794. This would suggest that it is probably an indigenousspecies.
Primrose willow or Kamole (Ludwigia octovalvis)Nelson observed kamole in Hawai`i in 1779, and Menzies collected it in
1794 (St. John 1978). It is most abundant in wetland habitats, and itssmall seeds may have been transported to Hawai`i by waterbirds; or, as itis a frequent weed in taro patches, it might possibly have been introducedaccidentally by the Polynesians with taro.
Kou (Cordia subcordata)The wood of the tree kou was one of the most prized by the Hawai`ians and
was used in prehistoric times (Krauss 1974). It is a plant of the littoralzone, and its seeds are dispersed on the ocean, so it may be an indigenousplant. However, it is not found in the wild in Hawai`i, possibly
Wester \ Origin and Distribution of Alien Plants 105
indicating either that it was brought and cultivated by the Polynesians orthat its original habitat has been destroyed.
Hairy merremia or Kuahulu (Merremia aegyptia)Most botanists regarded kuahulu as a weed introduced in the 19th century
until it was realized that Nelson had collected it in Hawai`i in 1779 (St.John 1978; British Museum of Natural History). Thus, it may be regarded asindigenous or introduced.
Currant tomato (Lycopersicon pimpinellifolium)The tomato is found as a volunteer in waste places and has been regarded
as an alien introduced by Europeans. The fact that there is a specimenlabelled as a collection by Menzies from Hawai`i (British Museum of NaturalHistory) now casts doubt on this conclusion. It is conceivable that seedsmight have been transported to Hawai`i stuck to migrating birds; but, if itwere indigenous, it is surprising that Hawai`ians did not make moreconspicuous use of it. Furthermore, the fact that the plant was notcollected or mentioned by other botanists until Hillebrand suggests thatthe Menzies specimen may have been collected elsewhere and is mislabeled.This is believed to have occurred with some other Menzies specimens.
Glossy nightshade or Popolo (Solanum americanum)Although popolo was not recorded in Hawai`i until Nuttall made a
collection of it in 1835 (British Museum of Natural History), it was one ofthe most important medicinal plants for the Hawai`ians (Gutmanis 1979).Furthermore, its Hawai`ian name, popolo, comes from a proto-Polynesianroot word that is widely applied, with small variations, to the same plantthroughout Polynesia (Biggs 1979). It would appear that it is eitherindigenous or was introduced, perhaps on purpose, by Polynesians.
Noni (Morinda citrifolia)The noni was cultivated by Hawai`ians in prehistoric times and was a
valuable medicinal and dye plant. It has a very wide distribution in thetropical Pacific, and its buoyant seeds disperse by flotation. Although
generally regarded as a Polynesian introduction it may have reached Hawai`iy means of natural dispersal.
The status of the above species as indigenes or aliens could be asubject of endless debate. However, it is unlikely that any of thedisputes can be satisfactorily solved unless pollen or macrofossil evidenceis found in sediments that unmistakably date from a period before humansettlement.
ALIEN INTRODUCTIONSAFTER EUROPEAN CONTACT
Rate of IntroductionIn the last 200 years, 813 species of alien plants have naturalized in
Hawai`i (Appendix). (The nomenclature used in this study follows that ofthe Manual of the Flowering Plants of Hawai`i (Wagner et al. 1990)except in a few cases, where taxa used by earlier authorities are retained
Wester \ Origin and Distribution of Alien Plants 106
in order to minimize information loss in the present analysis. Instancesof retained older nomenclature are noted in the Appendix.) The rate ofintroduction increased by the end of the 19th century to the presentaverage rate of about five species per year (Fig. 1). A similar patternwas observed by Healy (1947) in New Zealand, where, in the hundred yearsprior to 1946, alien plants arrived at a more or less constant rate of 10per year.
Figure 1. Accumulated number of adventive alien plant species in Hawai i.
Bias in Herbarium CollectionsHerbarium specimens may be an important source of information to
establish the time of arrival of aliens in Hawai`i; thus, it is worthwhileto consider in detail the nature of the information preserved incollections. This approach is useful in interpreting the establishment andspread of colonizing alien plants and may also provide information usefulin managing alien invasions. Extensive use has been made of herbariummaterial for mapping the ranges of taxa (Hastings et al. 1972) and for
Wester \ Origin and Distribution of Alien Plants 107
phenology (Holland 1975), as well as for reconstructing the introductionand colonization of species (Strother and Smith 1970). However, it iswidely recognized that many biases exist in herbarium collections (Stearns1951; Davis and Heywood 1965). For example, cultivated plants, as well asthose difficult to preserve or store, tend to be poorly represented incollections. Furthermore, collecting activity tends to be concentrated incertain geographical areas and at certain times. This has specialimplications for alien species and will be discussed in more detail.
Temporal Biases. Sudden increases in accumulated species in Hawai`iherbaria during the 20th century reflect the intermittent nature ofbotanical surveys. In the 20th century, survey and systematic collectingof the flora were more frequent after the B.P. Bishop Museum and theUniversity of Hawai`i were founded and individuals with botanical trainingwere employed permanently by private organizations or government agencies.As a result, newly introduced species were probably noticed and recorded ona more regular basis. Fluctuations in numbers of species accumulated inherbaria can be noted, for example, in the 1930s. This might beinterpreted as a wave of invasion caused by the conscious attempt torevegetate degraded watershed with introduced plants. However, thehistorical records used to compile these data are not free from biases, andthe observed fluctuations may well be the result of these instead.
The dates of collection of a randomized sample of approximately 2,000sheets from the B.P. Bishop Museum herbarium reveal three main periods ofintensive collecting activity in the 20th century (Fig. 2):
a. 1908-1920: This was the period soon after the founding of the B.P.Bishop Museum, when C.N. Forbes and others were building arepresentative collection.
b. 1930-1940: During this time several enthusiastic collectors worked inHawai`i, including O. Degener, F.E. Egler, F.R. Fosberg, andH. St. John.
c. 1968-present: The last 18 years have been a period a renewed interestin ecological and taxonomic research.
Significant increases in the number of "first records" of alien plantscorrespond closely with these periods of intense collecting activity.Therefore, small fluctuations in the curve of accumulated numbers ofspecies (Fig. 1) should not be given much significance. However, it isreasonable to assume that still more searching would reveal additionalspecies, even though the period from 1968 to the present has been one ofintensive collecting activity, because new aliens continue to colonize.
Geographic Biases. Considerable difference in intensity of botanicalreconnaissance from place to place is obvious when examining the herbariumspecimens preserved in the B.P. Bishop Museum herbarium. A randomizedsample collected from O`ahu shows that many specimens were obtained fromonly a few favored areas such as the highest peaks of the Ko`olau andWai`anae ranges, offshore islands, and in and around Honolulu (Fig. 3).
Wester \ Origin and Distribution of Alien Plants 108
Figure 2. Number of herbarium specimens collected in Hawai`i that are preserved in theB.P. Bishop Museum herbarium.
The University of Hawai`i campus at Manoa and the B.P. Bishop Museumthemselves figure prominently as sites of intense collecting activity.However, on the whole, most collecting, and presumably other botanicalsurveying, is concentrated in essentially natural areas where aliens areleast common, not in the disturbed habitats where newly introduced plantsare most likely to become established first. This is unfortunate becausecontrol or eradication of aliens is simplest when plants are recognized assoon as possible after arrival in an area, when populations are small andgeographically confined.
If a policy of early detection and eradication were to be seriouslyadopted as a management technique, searches should be directed to sitesother than those typically surveyed in the course of botanical research.For example, we might expect newly introduced aliens to establish nearports of entry such as airports, or perhaps more likely, near harbors whereused automobiles are unloaded, as Wace (1979) demonstrated that cars carrya very large number of weed propagules. On the other hand, if manyadventive plants in Hawai`i were first introduced as ornamental or crop
INTENSITY OF COLLECTINGBOTANICAL SPECIMENS
ON OAHU(NON-CULTIVATED SPECIES)
Figure 3. Number of herbarium specimens collected on O`ahu in the B.P. Bishop Museum Herbarium.
Wester \ Origin and Distribution of Alien Plants 110
species, they may have first appeared in or near arboreta, botanicalgardens, nurseries, or experimental plantings. In point of fact, manyfirst records do come from these sites. Ornamental plants are alsointroduced and grown by hobbyists. Those that spread aggressively maycolonize from these foci or be removed and dumped at official refuse sites,along roads, or in wasteland. Areas such as these should be searched forpotential new invaders.
Roadsides not only provide a disturbed substrate for many alien species,but also provide corridors for dispersal through undisturbed landscapesotherwise inhospitable to aliens (Smith 1982; Wester and Juvik 1983).Roads allow aliens to achieve wide distribution rapidly.
Further sampling of B.P. Bishop Museum herbarium specimen sheets wasdone to determine relative numbers of collections from each island. Theresults show that the largest number of specimens come from O`ahu; with theexception of Ni`ihau, the small islands are represented by a large numberof specimens in proportion to their area (Table 2). On the other hand,fewer specimens have been taken from Hawai`i Island than from O`ahu, whichis only one-eighth its size.
A high correlation exists between the number of herbarium specimenstaken from each island and the number of species known from that island,higher in fact than with other factors, which are often taken to determinethe richness of an island flora (Table 3). It has been similarly noted inthe Galapagos Archipelago that the number of botanical collecting trips toeach of the islands is a better predictor of numbers of species than arearea, elevation, or isolation (Conner and Simberloff 1978). Hence, itseems likely that our knowledge of the ranges of alien plants in theHawai`ian Islands, based on herbarium material, is inadequate; efforts canbe most profitably be spent to correct this by concentrating on Hawai`i,Maui, Kaua`i, and Ni`ihau.
EXTINCTION OF ALIEN SPECIES
Invading species owe part of their success to an ability to increasepopulations rapidly. However, they are also subject to population crashescaused by environmental factors, such as drought or hurricanes, or tocompetition from other species. They often inbreed, forming homozygouspopulations vulnerable to attack by pests. As Baker (1965) pointed out,weeds are well adapted to rapid colonization and spread, but they have alimited future because they are often poorly equipped genetically to adjustto inevitable ecological and environmental changes. Although this does notseem to be the case in benign and isolated tropical areas, it is thereforeof interest to consider whether abundant new arrivals are at least partlybalanced by extinctions.
It is not possible to prove that a species has become extinct, butspecies once reported as aliens in Hawai`i that have not been collected atleast since 1959 are listed in Table 4. The recent period of intensivecollecting activity occurred after 1959, and it might usually be that a
Wester \ Origin and Distribution of Alien Plants 111
Table 2. Alien species per unit area (mi ) on major islands.
NW Hawn. Ids.Lana`iKaho`olaweNi`ihauO`ahuMoloka iKaua`iMauiHawai i
TOTAL
Number ofalien species
762176577
571232345379566
813
Area(mi2 )
51394573
608261553729
4,038
6,451
Number ofspeciesper mi
14.621.561.441.050.940.890.620.520.14
0.126
Number ofspecimensper mi
20.771.251.560.622.600.730.580.450.27
0.60
Table 3. Factors likely to influence the number of known adventive alien plant specieson each island.
Hawai`iMauiO` ahuKaua`iMoloka`iLana`iNi`ihauKaho`olaweNW Hawn. Ids.
Correlationcoefficient
Area(mi2)
4,03872960855326113973455
0.685*
MaximumElevation
(feet)
13,79610,0234,0205,2434,9703,3701,2811,477
874
0.757*
Island Age(m. years)
0.501.633.805.721.841.46n.a.1.03
11.30
-0.312
Numberof AlienSpecies
566379571345232217776576
Number ofHerbariumSpecimens
1,082327
1,5783191901744570
108
0.888**
*p < 0.05
**p < 0.01
Wester \ Origin and Distribution of Alien Plants 112
Table 4. Alien species not collected or observed since 1959.*
Species
Bidens laevisBlumea laciniataBlumea sessilifloraBothriochloa intermediaDigitaria longiflora**Echinochloa phyllopogon * * *Geranium dissectumLaportea interruptaPotamogeton nodosusPotamogeton pectinatusSpergula arvensisStachytarpheta x intercedensStachytarpheta x trimeniTypha domingensis
Date ofFirstRecord
18551923192019161916192819101819182519471888193919281939
Date ofLastRecord
19121937193719401938193019481959192019491958194019511942
Number ofHerbariumSpecimens
27125241525222
*Nomenclature according to herbarium labels; a number of commonly cultivated speciesand those observed as naturalized at only one location on one occasion have not beenincluded in this list.
**Not listed in Wagner et al. 1990; listed in St. John 1973.
***Not listed in Wagner et al. 1990 or St. John 1973.
u
Sparingly naturalized, according to Wagner et al. 1990.
species would be collected if populations persisted. Among the species notcollected and possibly extinct are a number that once had populationsestablished on several islands and that were collected in considerablenumbers. This would suggest that aliens may be suppressed, or possiblyexterminated, without purposeful eradication programs, and that largepopulations do not always survive. The present regression of many koahaole (Leucaena leucocephala) populations in the Hawai`ian Islands andelsewhere is a vivid illustration of the fact that, once introduced, alienplant populations may undergo dramatic fluctuation without direct humaninterference. This observation should not be a surprise: most aliens areopportunist species with general ecological roles. Their strategy forsurvival is effective dispersal and rapid population increase, rather thanspecialized adaptation to local environments. Successful competitionagainst all other plants over an extended period of time is not favored bythe generalist strategy. Disturbance, however, does favor suchgeneralists.
Wester \ Origin and Distribution of Alien Plants 113
The number of species that seem to disappear from herbarium records hassteadily increased during the 20th century; during the decade of the 1950s,this disappearance occurred at an average rate of less than one per year.Preliminary evidence thus suggests that turnover in the weed flora mayoccur, but the number of arrivals of new alien species is far greater thanthe number being exterminated.
Place of OriginThe immediate origin of a new alien is usually not known because the
colonizing propagule may not come directly from its native home. However,except in a few cases of pantropical species, it is possible to determinethe provenance of aliens. It appears that the New World supplied a largeproportion of aliens in the early European period of contact (Fig. 4).This is not surprising, since sailing ships coming to Hawai`i typicallystopped in South America and hence had the opportunity to pick up tropicalplants, on purpose or by accident, suited to the Hawai`ian environment. Inlater times, the Old World made a larger contribution to the Hawai`ian alienflora, perhaps as contacts with Asia became more frequent and as thecircumtropical weed flora became more homogenized. A significant number ofspecies arrived from North America in the 20th century, and theseintroductions reflect the close cultural and economic ties with thatregion.
Mode of IntroductionDirect evidence of the means of introduction of alien plants can only be
found for a small proportion of the species. These are mostly plantspurposefully introduced as crops, reforestation species, or ornamentals andobserved to establish from the original plantings. If we assume thatintroduction was purposeful for species that are commonly crop orornamental species, and that introduction was accidental for species withno redeeming social value, it appears that about half the introductionswere on purpose and the other half accidental (Table 5).
CONCLUSIONS
The native flora of the Hawai`ian Islands has been known in more detail,and for a longer period of time, than that of most other oceanic islands.However, the alien flora is not nearly so well known despite the fact thatit represents a large component of the present-day flora; additionally,many aliens are now ecological dominants over much of the land area ofHawai`i. A clearer knowledge of the nature and development of the alienflora is desirable so that further introductions can be minimized and thespread of aliens already present can be monitored and controlled to agreater extent. Competition that aliens exert on the diminishing nativeecosystems can thus be reduced.
Wester \ Origin and Distribution of Alien Plants 114
Figure 4. Region of origin of adventive alien plants in Hawai`i.
Table 5. Mode of introduction of alien species.
Mode of Number PercentIntroduction of Species of Species
PurposefulCropOrnamental
Accidental
199121
236
21.835.9
42.4
Wester \ Origin and Distribution of Alien Plants 115
Appendix. A Preliminary List of Alien Adventive Seed Plantsin the Hawai`ian Islands.
The following abbreviations are used in this list:
Life form Origin (Place of Origin)
B - tuber, bulb AA - Asia, AustraliaH-herb AF -AfricaS - shrub AS - AsiaT - tree ASP - Asia, PacificU - succulent AUP- Australia, PacificV - vine AUS - Australia
COS- cosmopolitanDIS - disjunctEAS - EurasiaEUR- EuropeHOR- horticultureMED- MediterraneanNAM- North AmericaNH - Northern HemisphereNW -New WorldOW -Old WorldP - PacificSAM- South AmericaTAM- Tropical AmericaTOW- Tropical Old WorldUSM- United States Mainland
Mode of IntroductionA - accidentalC - cropO - ornamental
Record (date of reports of species in the Hawai`ian Islands)first - earliest reportrecent - most recent record
Info. Source (information source for earliest report of species in Hawai`i)P - published referenceH - herbarium specimen
No. spms. (Number of herbarium specimens of species collected in Hawai`i and preserved in B.P.Bishop Museum Herbarium)
Island (Name of island where presence of species can be verified by an herbarium specimen.Note: many species occur more widely than indicated here.)
Hi - Hawai`iMa - MauiMo - Moloka`iLn - Lana`iKh - Kaho`olaweOh - O`ahuKa - Kaua`iNi- Ni`ihauNW - Northwest Hawai`ian Islands
Appendix, continued.
Appendix, continued.
Taxon
GYMNOSPERMSPINACEAE (Pine family)
Pinus elliottiiPinuspatulaPinus pinaster
ANGIOSPERMSMONOCOTYLEDONS
LifeForm
TTT
Origin
USMNAMEUR
Mode ofIntro.
CC
Record in Hi.First
197619441976
Recent
197619761976
Info.Source
HHH
No. Spms.
121
ALISMATACEAE (Water plantain family)* Echinodorus rostratus
Sagittaria latifoliaAGAVACEAE (Agave family)
Agave sisalanaCordyline fruticosaFurcraea foetidaPhormium tenax
ARACEAE (Taro family)Alocasia macrorrhizaColocasia esculentaPistia stratiotes
CANNACEAE (Canna family)Canna edulisCanna indica
HH
USUS
BBH
BB
NWEAS
TAMASPSAMP
ASPASOW
TAMTAM
AO
CCCO
CCO
CC
19391916
1893100018701933
100010001932
18981817
19391986
1977198519461949
197219861985
19291986
HH
PPPH
PPP
PH
113
468104
643
121
COMMELINACEAE (Spiderwort family)Callisia fragransCommelina benghalensisCommelina diffusa
HHH
TAMOWAS
OAA
192819091835
197519821983
HHH
81630
IslandHi
++++
+
+
Ma
++
+
+
+
++
Mo
++
+
+
Ln
++
+
+
Kh Oh
++
+++
+++
++
+++
Ka
+
+
+
+
+
+++
Ni
+
NW
+116
Taxon
Commelina erectaTradescantia geniculataTradescantia spathaceaTradescantia zebrina
CYPERACEAE (Sedge family)Bulbostylis capillarisCyperus altemifoliusCyperus compressusCyperus difformisCyperus esculentusCyperus gradlisCyperus halpanCyperus pilosus
* Cyperus rivularisCyperus rotundusCyperus trine/visCyperus virensEleocharis geniculataEleocharis obtusaEleocharis radicansFimbristylis aestivalisFimbristylis schoenoidesKyllinga brevifoliaKyllinga nemoralisSchoenoplectus
californicusTorulinium odoratum
LifeForm
HHHH
HHHHHHHHHHHHHHHHHHH
HH
Origin
NAMTAMTAMTAM
NAMAFCOSOWCOSAUSEASAS-cosAUSTAMTAMNAMTAM-OWOW-
NWAS
Mode ofIntro.
OOOO
AOAACOAAAAAAAAAAAAA
AA
Record in Hi.First
1925188819371926
1911192119351895195618931957191619491850192619761909185219171977198418651929
19121825
Recent
1963197319761976
1985198519821984195619851985191619751983196619781984198519821983198519841985
19861982
Info.Source
PPHH
HHHHHPHHHPHHHHHHHHH
HH
No. Spms.
0069
31206152121423213413646426616
213
IslandHi
+
+++
+++
++++
+
++
+
Ma
+
+
++
+
+
+
Mo
++
+
Ln
+
+
Kh Oh
+
+
+
+
+
+
+++
++
+
Ka
+
+
+
+++
+
++
+
++
+
Ni
+
+
+
NW
+
+
Appendix, continued. 117
Appendix, continued.
Taxon
DIOSCOREACEAE (Yam family)Dioscorea bulbiferaDioscorea pentaphylla
LifeForm
BB
Origin
ASASP
Mode ofIntro.
CC
Record in Hi.First Recent
10001000
19841984
Info.Source
PP
No. Spms.
4026
HYDROCHARITACEAE (Frog's-bits family)Egeria densa
IRIDACEAE (Iris family)Belamcanda chinensisCrocosmia x crocosmiifloraSisyrinchium rosulatumSisyrinchium strictumTrimezia martinicensis
JUNCACEAE (Rush family)Juncus acuminatusJuncus bufoniusJuncus effususJuncus ensifoliusJuncus planifoliusJuncus pofyanthemosJuncus tenuis
LEMNACEAE (Duckweed family)LemnaperpusillaSpirodela pofyrhiza
LILIACEAE (Lily family)Hippeastrum puniceum
LIMNOCHARITACEAE* *** Hydrocleys nymphoides
H
BBHHH
HHHHHHH
HH
B
H
SAM
ASAFTAMUSMTAM
NAMCOScos~
AUPAUSNH
COSCOS
TAM
SAM
O
OOAAO
AAAAAAA
O
O
O
1937
18881932194419381943
1935191019001911193019531915
18951918
1928
1939
1971
19761985194519381984
1935197119751980198419671974
19611976
1986
1986
H
PHHHH
HHPHHHH
HH
P
H
4
79412
1171041428
105
3
4
IslandHi
+
+
+++
+++++++
Ma
+
+
+
+
Mo
++
++
+
+
+
Ln Kh Oh
++
+
++
+
+
+
+
+
Ka
++
+
+
++
+
Ni NW
00
Taxon
MUSACEAE (Banana family)Musa troglodytarumMusa x paradisiac^
ORCHIDACEAE (Orchid family)Arundina graminifoliaEpidendrum x obrianianumPhaius tankarvilleaeSpathoglottis plicata
POACEAE (Grass family)Agrostis avenaceaAgrostis semiverticillataAgrostis stoloniferaAira caryophylleaAmmophila arenariaAndropogon virginicusAnthoxanthum odoratumArthraxon hispidusAvena barbataAvenafatuaAxonopus fissifoliusBothriochloa barbinodisBothriochloa intermediaBothriochloa pertusaBrachiaria muticaBriza maximaBriza minor
LifeForm
HH
HHHH
HHHHHHHHHHHHHHHHH
Origin
PASP
ASTAMASPAS
AUPEUREUREUREURUSMEURASEUREURUSMTAMAUSTOWAFEUREUR
Mode ofIntro.
CC
OOO
ACCAACC-ACCCCCCCC
Record in Hi.First
19601000
1950194019311929
18881864191219161913192419071972191618881912191019161916190219171840
Recent
19601957
1983198419851975
19821972196919851962197319841972196119431984196719401980198419841984
Info.Source
HP
HHHH
HPHHPHHHPPPHHPPHP
No. Spms.
34
2251734
631113102
3233145169217277
30
IslandHi
+
++
++++
+++
++
++++
Ma
+
+
+
++
++
++++
Mo
+
+
++
+
+++
+
Ln
+
+
+
+
++
+
Kh Oh
++
++++
+
+
+
+++++++
+
Ka
++++
+++
++
+++
Ni
+
NW
+
+
Appendix, continued. 119
Appendix, continued.
Taxon
Bromus mollisBromus rigidusBromus rubensBromus tectorumBromus willdenowiiCenchrus ciliarisCenchrus echinatusChloris barbataChloris divaricataChloris gayanaChloris petraeaChloris radiataChloris truncataChloris virgataChrysopogon aciculatusCoix lachryma-jobiCymbopogon refractusCynodon dactylonDactylis glomerataDactyloctenium aegyptiumDanthonia pilosaDanthonia semiannularisDichanthium annulatumDichanthium aristatumDichanthium sericeumDigitaria ciliaris
LifeForm
HHHHHHHHHHHHHHHHHHHHHHHHHH
Origin
EUREUREUREUR-
AFTAMTAMAUSAFTAMTAMAUSTAMTAMAS
AUSOW
EURTOWAUSAUSASASAUS-
Mode ofIntro.
CCCC-CAACC-CCCACACCACC---C
Record in Hi.First
19091910192018881909193218671902192419131968185119151903186518951929188819111909191019031927191119121912
Recent
19781984196719691982198219841984198019851980197619801982197219841982198419841983198519751963197619801982
Info.Source
HHPPHHPHHHHPPHHPHHHHPPPHPH
No. Spms.
101933
30751701219230228342510935133623181144
IslandHi
+++
++++
+
+
+++++++++
++
Ma
++
++
++++
+
+
+
+++++
+
+
Mo
+++
++
++++
++
+
Ln
+
+
+
+
++
+
+++
Kh
+++
++
+
+
+
Oh
+
++++++
+
+++++++
++++
Ka
++
++
+
++++++
+
Ni
++++
+++
NW
+
++
+
+
+
Taxon
Digitaria fuscescensDigitaria insularis
* Digitaria longifloraDigitaria pentziiDigitaria sanguinalisDigitaria setigeraDigitaria violascensEchinochloa colonaEchinochloa crus-galli
* Echinochloa cruspavonisEchinochloa glabrescens
** Echinochloa phyllopogonEhrharta stipoidesEleusine indicaEragrostis browneiEragrostis cilianensisEragrostis ciliarisEragrostis elongataEragrostis pectinaceaEragrostis pilosaEragrostis tenellaEragrostis unioloidesEustachys petraeaFestuca rubraGamotia acutiglumaGastridium ventricosum
LifeForm
HHHHHHHHHHHHHHHHHHHHHHHHHH
Origin
SAMCOSASAFEURASASASASSAMAS-
AUPAS
AUSEURCOSASNAMEURTOWASTAMEUR-
EUR
Mode ofIntro.
C-CCAAAACC--AACC-CCCAC-C-C
Record in Hi.First
18521948191619501864182619161864184619161895192819161940191618641976194919161913189519581978191218881909
Recent
19691957193819781984198319851982198319661966193019831984197319851976198319771983198419581978198219721982
Info.Source
HHHPPPHPHHHHHPHPHPHPHHHPHH
No. Spms.
38252756410546221321662113614161
40115913
IslandHi
+
+++++++++
++++
++
++
+++
Ma
+
+++++
+
+++
+
+
+++
Mo
++
++
++
+++
+
Ln
+
++
++
+
+
Kh
++
+
+
+
Oh
+++
++++++++++
++
+
+
Ka
+
++
++
+
+
+
+
+
Ni
+
+
+
+
NW
+
+
+
+
Appendix, continued. 121
Appendix, continued.
Taxon
Holcus lanatusHordeum leporinumHordeum vulgareHyparrhenia hirtaHyparrhenia rufaLolium multiflorumMelinis minutifloraMuhlenbergia microspermaOplismenus compositusOplismenus hirtellusPanicum maximumPanicum miliaceumPanicum repens
* Paspalidium radiatumPaspalum conjugatumPaspalum dilatatumPaspalum distichumPaspalum fimbnatumPaspalum paniculatumPaspalum scrobiculatumPaspalum urvilleiPaspalum vaginatumPennisetum clandestinumPennisetum macrostachyumPennisetum polystachionPennisetum purpureum
LifeForm
HHHHHHHHHHHHHHHHHHHHHHHHHH
Origin
EURMEDOWOWTOWEUREURNW
ASTAMAFHASOW
AUSTAMSAMTAMTAMTAMNWSAMTAMAFASTOWAF
Mode ofIntro.
A-CCCCCCCACCCCACCCC-CCCC-C
Record in Hi.First
19091909193519621933191019131930196618411888192319161939184019111916191619801852191419361924191619291912
Recent
19851938198419811982198019831974196619831984198119791939198319841978198319801982198419741984196719851976
Info.Source
HHHHPHPHHPHPHHPHPPHHHHHPHP
No. Spms.
351264161835111
84147121
55413211
49268883018
IslandHi
+++
+++
++++
++
++++++++
Ma
+++++++
+
++
++
++
+
++
Mo
+
+++
+
++
++
++
Ln
+
++
+
+
+
++
+
+
++
Kh
+
+
+
Oh
+
+
+++++++
++++
+++++++
Ka
+
+
+++
++
++++
+++
+++
Ni
++
NW
+
+
122
Taxon
Pennisetum setaceumPoa annuaPoapratensisPolypogon interruptusPofypogon mompeliensisRhynchelytrum repensSacciolepis indicaSchizachyrium condensationSchizostachyum
glaucifoliumSetaria glaucaSetaria gradlisSetaria palmifoliaSetaria verticillataSorghum halpense
* Sorghum sudanenseSporobolus africanusSporobolus dianderSporobolus elongatusSporobolus indicusSporobolus pyramidatusStenotaphrum secundatumStipa cernuaThemeda villosaTragus berteronianus
# Tricachne insularis
LifeForm
HHHHHHHH
HHHHHHHHHHHHHHHHH
Origin
AFEUREURSAMEURAFASNAM
POWTAMOWOW
MEDAFASAFAUSTAMTAMUSMUSMASOW
TAM
Mode ofIntro.
CCCCCCCC
CCCCACCCCCC-CCCAC
Record in Hi.First
19141841191118711909189519081961
10001910185519031895190919141911191119091912197818401957192419351913
Recent
19831983197819701967198419841974
19801983198419831984198319291984198319361983197919841982197919841984
Info.Source
HPHPHPHH
PPHHHHHHHHPHPHHPH
No. Spms.
2735161425634315
371645309724411391715436811843
IslandHi
++++++++
++++++
+
+
+
+
+
+
+
+
Ma
++++++
+
++++
+
++
++
Mo
++
+
+
+
+
+
+
++
Ln
++
+
++++
++
++
+
+
Kh
+
+
+
Oh
+++++++
+++++++++++++
+
+
Ka
++
+
++
+++
++
+
+
+
+
Ni
+
+
NW
+
++
+
+
++
+
Appendix, continued. 123
Appendix, continued.
Taxon
Vulpia bromoidesVulpia myuros
* Vulpia octoflora
LifeForm
HHH
Origin
EUREURUSM
Mode ofIntro.
.--
Record in Hi.First Recent
190119111966
197619851966
Info.Source
PHH
No. Spms.
39532
PONTEDERIACEAE (Pickerel weed family)Eichhomia crassipesMonochoria vaginalis
HH
TAMAS
0A
19001935
19761946
PH
54
POTAMOGETONACEAE (Pondweed family)Potamogeton foliosusPotamogeton nodosusPotamogeton pectinatus
TACCACEAE (Tacca family)Tacca leontopetaloides
TYPHACEAE (Cattail family)Typha domingensisTypha latifolia
HHH
B
HH
NAMCOSCOS
AS
OWCOS
AAA
C
OA
181718251947
1000
19391977
198419201949
1977
19421981
HHH
P
HP
1552
40
23
XYRIDACEAE (YeUow-eyed grass family)Xyris complanataXyris platylepis
ZINGIBERACEAE (Ginger family)Alpinia purpurataAlpinia zerumbetHedychium coronariumHedychium flavescensHedychium gardnerianumZingiber zerumbet
HH
BBBBBB
AUSUSM
PASASASASAS
AA
OOOOOC
19731951
192518881888191319541000
19851962
197919761986197219751983
PH
HPPHHP
26
16121216335
IslandHi
+++
+
+
+
++
+++++
Ma
++
+
++
+
++
+
Mo
+
+
+
++
+
Ln
++
++
+
Kh Oh
++
+
+
+
++
++++++
Ka
++
+
+
+
+
+
+
Ni
+
NW
+
Taxon
DICOTYLEDONSACANTHACEAE (Acanthus family)* Aphelandra aurantiaca
Asystasia gangeticaBafleria cristataDidiptera chinensisHemigraphis reptansJusticia betonicaJustida spidgera
** Odontonema cuspidatumRuellia brittonianaRuellia graedzansRuellia prostrataThunbergia alataThunbergia fragrans
AIZOACEAE (Fig-marigold family)Tetragonia tetragonioides
LifeForm
SHHHHHSSSSHVV
H
Origin
TAMTOWASASASTOW--
TAMSAMASAFAS
DIS
Mode ofIntro.
OOOOOOOOOOOOO
C
Record in Hi.First
1958192519291942195719431934193719301925194418651916
1909
Recent
1975197619741985197919841983197319811893198019461984
1976
Info.Source
HHHHHHHHHHHHH
H
No. Spms.
6188135143711105612
10AMARANTHACEAE (Amaranth family)
Achyranthes asperaAlternanthera brasilianaAltemanthera caracasanaAlternanthera pungensAlternanthera sessilisAlternanthera tenellaAmaranthus dubiusAmaranthus hybridus
HHHHHHHH
ASTAMAFTAMCOSTAMTOWNW
AAAAAAAO
18551973192519591935193619301930
19831973198519851983194419771982
HHHHHHPH
2611498244
IslandHi
+
+
+
+
+
+
+
+
Ma
+
+
+
+
Mo
+
++
+
+++
Ln
+
+
+
+
Kh Oh
+++++++++++++
+
+
++++++
Ka
++
+
+
+
Ni NW
+
+
Appendix, continued. 125
Appendix, continued.
Taxon
Amaranthus lividusAmaranthus spinosusAmaranthus viridisGomphrena globosa
ANACARDIACEAE (Mango family)Anacardium occidentalSchinus terebinthifolius
ANNONACEAE** Annona cherimolaAPIACEAE (Parsley family)
Anethum graveolensCentella asiaticaCiclospermum leptophyllumCoriandrum sativumCryptotaenia canadensisDaucus pusillusEryngium foetidumFoeniculum vulgareHydrocotyle bowlesioidesHydrocotyle sibthorpioidesHydrocotyle verticillataPetroselinum crispumTorilis nodosa
APOCYNACEAE (Dogbane family)Catharanthus roseus
LifeForm
HHHH
TT
T
HHHHHHHHHHHHH
H
Origin
COSTAMCOSTAM
TAMSAM
SAM
EASASNWEASNAMNAMTAMEUR-ASOW
MEDOW
TAM
Mode ofIntro.
AAAO
CO
C
OAACCAACAAACA
O
Record in Hi.First
1971191118191929
19151909
1791
1888188819091926192517941944186519311932181719101855
1865
Recent
1985198419841948
19831972
1975
1953197219831952197919811976198419841983196619841911
1983
Info.Source
HPPH
HH
P
PPHHHHHPHHHHH
H
No. Spms.
312254
923
7
3232652214215653183
22
IslandHi
+++
+
+
+++++
++++++
+
Ma
++
+
+
+++
+
Mo
+
+
++
+
+
+
Ln
+
+
+
+
+
+
Kh
++
Oh
++++
++
++++++++
+++
+
Ka
+
++
+
++
+
Ni
+
+
NW
++
+
+
Taxon
AQUIFOLIACEAE (Holly family)Ilex cassineIlex paraguariensis
ARALIACEAE (Ginseng family)Schefflera actinophylla
ASCLEPIADACEAE (Milkweed family)Asclepias curassavicaAsclepias physocarpaStapelia gigantea
ASTERACEAE (Sunflower family)Acanthospermum australeAcanthospermum hispidumAchillea millefoliumAgeratina adenophoraAgeratina ripariaAgeratwn conyzoidesAgeratwn houstonianumAmbrosia artemisiifoliaAnthemis cotulaArctium lappaArtemisia ponticaArtemisia vulgarisAster subulatusBaltimora rectaBidens albaBidens cynapiifolia
LifeForm
TT
T
HHH
HHHHHHHHHHSHHHHH
Origin
USMSAM
AUS
TAMAFAF
TAMTAMNHTAMTAMTAMTAMNAMEURHASEUREURTAMTAM-
TAM
Mode ofIntro.
OC
O
OOO
AAAAAOOAAOAAOAAA
Record in Hi.First
19491934
1900
185019191942
1840195119111909192518251909184019091910195919091826198419581929
Recent
19851984
1976
197619741984
1974196619751967197619741977198419761960195919591977198419831976
Info.Source
HH
P
PHH
PHHHPPHHHHHHHPHH
No. Spms.
157
6
44225
2821063135215401471141005
20
IslandHi
+
++
+
++++++++
++
++
Ma
+
+
+
+
+
Mo
+
++
+
+
+
+
Ln
++
+
+
+
+++
+
Kh
+
+
+
Oh
++
-f
+++
+
++++++
-h++++++
Ka
+
+
+
++
+
++
Ni
+
+
NW
+
Appendix, continued. 127
Appendix, continued.
Taxon
Bidens gdrdneriBidens laevis
##Bidens lanceolataBidens pilosaBlumea laciniataBlumea sessilifloraCalyptocarpus vialisCentaurea maculosaCentaurea melitensisChrysanthemum leucanthemum
* Chrysanthemum maximumCirsium vulgareConyza bonariensisConyza canadensisvar. canadensis
Conyza canadensisvar.pusilla
Coreopsis lanceolataCosmos caudatusCotula australisCrassocephalum crepidioidesCrepis capillarisEclipta albaElephantopus mollisElephantopus spicatusEmilia coccinea
LifeForm
HHHHHHHHHHHHH
H
HHHHHHHHHH
Origin
-NW-
TAMASASTAMEUREUREASEUREURSAM
NW
NWUSMTAMAUPAFEURCOSTAMTAMAS
Mode ofIntro.
AAAAAAAAAAOAA
A
AOOAAAAAAA
Record in Hi.First
1983185519631819192319201971196518551911191719091888
1911
1817193818651943192919341865192619351909
Recent
1984191219631975193719371985196519761975195919831985
1984
1984194619531983198419751974197419751963
Info.Source
HPHHHHHHHHHHP
H
HHHHHHPHHH
No. Spms.
121
5671111
31441953
6
574431681616611
IslandHi
++
++++++
+
++++
++
+
Ma
+
++
++
++
+
+
+
+
Mo
++
+
+
++
+
+
Ln
+
+
+
++
+
+
Kh
++
Oh
+
++
+
+
++
+++++++
++
Ka
+
+
+
++
++
++
Ni
+
+
++
NW
Taxon
Emilia fosbergiiEmilia sonchifolia
var. javanicusEmilia sonchifolia
var. sonchifoliaEncelia farinosaErechtites hieracifoliaErechtites valerianifoliaErigeron annuusErigeron bellioidesErigeron karvinskianusFilago gallicaFlaveria trinerviaGaillardia pulchellaGalinsoga parvifloraGalinsoga quadriradiataGnaphalium japonicumGnaphalium purpureumHelichrysum foetidumHeterotheca grandifloraHypochoeris glabraHypochoeris radicata
** Kalimeris pinnatifidaLactuca serriolaLapsana communisMadia sativa
LifeForm
H
H
HHHHHHHHHHHHHHHHHHHHHH
Origin
_
AS
ASUSMNW
TAMCOSTAMTAMEURNAMUSMTAMTAMASNAMNWUSMEURMED-EUREURSAM
Mode ofIntro.
A
A
AAAAAAOAAOAAAAOAAAAAAA
Record in Hi.First
1920
1927
1931190919091916193619841911196619371908190819641909188819491909190918951983192819091927
Recent
1984
1976
1983198319841984193619851983196719781984197619761981198419821975197419751983198519841937
Info.Source
H
H
HHHHHHHHHHHHHPHHHPHHHH
No. Spms.
92
4
9101534131738122743569933113916244
IslandHi
+
+++
++
++++++++
++
Ma
+
+
+
+
++
+
+
+++
Mo
+
++
+++
+
Ln
+
++
+
+++
Kh
+
+
+
Oh
+
+
+
++
++
++++++
++
Ka
+
+
+
+
+
+
+
+
++
Ni
+
NW
+
+
Appendix, continued. 129
Taxon
Matricaria chamomillaMontanoa hibiscifoliaOsteospermwn calendulaceumPalafoxia callosaParthenium hysterophorusPicris hieracioidesPluchea indicaPluchea symphytifoliaPluchea xfosbergiiReichardia picroidesRudbeckia laciniataSenecio mikanioidesSenecio sylvaticusSenecio vulgarisSigesbeckia orientalisSolidago canadensisSonchus asperSonchus oleraceusSynedrella nodifoliaTagetes minutaTanacetum vulgareTaraxacum laevigatumTaraxacum officinaleTithonia diversifoliaTragopogon porrifoliusTridax procumbens
LifeForm
HSHHHHSSSHHVHHHHHHHHHHHSHH
Origin
EASTAM-
USMUSMEASASTAM-OWNAMAFEUREUROWUSMCOSEURTAMSAMEUR-EURTAMEURTAM
Mode ofIntro.
AOAAAAAAAAOOAAAOAAAAAAAOOA
Record in Hi.First
19091919198319711960193019091931193419091958190919101932188819261975181719261932195919811909191719381922
Recent
19481975198319711985197419721974198019551958197519851982197619771980197219831974195919811976197519821976
Info.Source
HPHHHHHHHHHHHHPPHHHHHHHHHH
No. Spms.
4181179414421421142441632
3814141118113
26
IslandHi
+++
++
+
+
+++++++++
++++
Ma
+
+
+++
+++
++
++
+
Mo
++
++
+
+
+
Ln
+
+
+
+
+
+
Kh
+
+
Oh
+
+++++
+
++
+++++++
+
Ka
+
+
+++
++
++
+
Ni
++
+
NW
+
130
Appendix, continued.
Taxon
Verbesina encelioidesVemonia cinereaWedelia trilobataXanthium strumariumYoungia japonicaZinnia palmeriZinnia peruviana
LifeForm
HHHHHHH
Origin
NAMTOWTAMNAMASTAMTAM
Mode ofIntro.
AAOAAOO
Record in Hi.First
1888187019651840186519801909
Recent
1976196719751983197619801983
Info.Source
HPPPHHH
No. Spms.
26301342441
25BALSAMINACEAE (Touch-me-not family)
Impatiens oliveriImpatiens wallerana
BASELLACEAE (Basella family)Anredera cordifoliaBasella alba
BATACEAE (Saltwort family)Batis maritima
BEGONIACEAE (Begonia family)Begonia foliosa v miniataBegonia hirtellaBegonia reniformis
BIGNONIACEAE (Bignonia family)Spathodea campanulata
BIXACEAE (Arnotto family)Bixa orellana
BORAGINACEAE (Borage family)Amsinckia intermediaBothriospermum tenellum
HH
VV
H
HHH
T
T
HH
AF-
SAMOW
TAM
SAMSAM-
AF
TAM
USMAS
OO
OC
A
OOO
O
C
AA
19561939
19401865
1859
191719521976
1890
1830
19441854
19681984
19731945
1985
193719841985
1986
1986
19441975
HH
HH
P
HHH
H
P
HH
210
83
47
362
22
29
17
IslandHi
++++
+
+
+
+++
+
++
Ma
+
+++
+
+
+
+
Mo
+
++
+
+
+
Ln
++
++
+
+
Kh
+
+
+
Oh
++++++
+
++
+
4
+
+
+
Ka
+++++
+
+
+
+
Ni
+
+
+
NW
+
Appendix, continued. 131
Appendix, continued.
Taxon
Cordia subcordataCynoglossum amabileHeliotropium atnplexicauleHeliotropium procumbensMytosis discolorToumefortia argentea
BRASSICACEAE (Mustard family)Brassica campestrisBrassica junceaBrassica nigraCapsella rubellaCardamine flexuosaCardamine hirsutaCoronopus didymusLepidium bonarienseLepidium densiflorumLepidium hyssopifoliumLepidium oblongumLepidium virginicumLobularia maritimaNasturtium microphyllumNasturtium sarmentosumRaphanus raphanistrumRaphanus sativusSisymbrium altissimumSisymbrium irioSisymbrium officinale
LifeForm
THHHHT
HHHHHHHHHHHHHHHHHHHH
Origin
TOWASSAMTAM-AS
EURASEUREUREUREUROW
SAMNAM--
NAMMEDEURP
EUREUREASEUROW
Mode ofIntro.
COOA-O
ACAAAcAAAAAAOCCCCACA
Record in Hi.First
100019301871197519811888
19091942188819091840184018401975193519751914187219391888191219631888192719631909
Recent
197319761983198319811974
19801970198219851984197919761975193519751980198519801932198419841980198119821984
Info.Source
PHPHHP
HHPHHHHHHHHHHPHHPHHH
No. Spms.
311120101
21
1348
20433441111437157205974
25
IslandHi
+++
+
+++
+
+
++
+
+
+
+
+
++
+
+
+
+
+
Ma
+
++
+
+++
+
++++
+
+
Mo
+
+
+
+
++
+
+
Ln
+
+
+
+
+
+
+
Kh
+
++
Oh
++++
+
+++++
+
+++++++
Ka
+
+
+
++
+++++
+
Ni
+
+
+
+
NW
+
++
TaxonLifeForm Origin
Mode ofIntro.
Record in Hi.First Recent
Info.Source No. Spms.
BUDDLEIACEAE (Butterfly bush family)Buddleja asiaticaBuddleja madagascariensis
CACTACEAE (Cactus family)Cereus uruguayanusHarrisia martiniiHylocereus undatusOpuntia cochenilliferaOpuntia ficus-indicaOpuntia vulgaris
SS
UUUUUU
ASAF
SAMSAMTAMTAMNWSAM
OO
OOOOOO
19081931
193019591830188819851932
19841984
195919591985198519851944
HH
HPPPHP
587
409141
CAMPANULACEAE (Bellflower family)Hippobroma longifloraLobelia erinusTriodanis bifloraWahlenbergia gracilis
CAPPARACEAE (Caper family)Cleome gynandra
HHHH
H
TAMAFNWCOS
AF
AOAO
A
1865193119451928
1857
1983193119451985
1984
HHHH
P
241212
38CAPRIFOLIACEAE (Honeysuckle family)
Lonicera japonicaSambucus mexicana
CARICACEAE (Papaya family)Carica papaya
CARYOPHYLLACEAE (Pink family)Arenaria serpyllifoliaCerastium fontanumDianthus armeria
VT
T
HHH
ASTAM
SAM
EUREUREUR
OO
C
AAA
18821909
1823
193518651964
19651974
1984
198319851982
PP
P
HPH
1112
13
9322
IslandHi
++
+
++++
+
++
+
+++
Ma
+
+
++
Mo
+
+
+
+
Ln
+
+
Kh Oh
++
+
++++
+
+
++
+
+
Ka
+
+
+
+
+
Ni NW
Appendix, continued. 133
Appendix, continued.
Taxon
Drymaria cordataPetrorhagia velutinaPolycarpon tetraphyllumSagina decumbensSagina japonicaSilene gallicaSpergula arvensisSpergularia marinaStellaria media
CASUARINACEAE (She-oak family)Casuarina equisetifoliaCasuarina glauca
CECROPIACEAE (Cecropia family)Cecropia obtusifolia
CERATOPHYLLACEAE (HornwortCeratophyllum demersum
LifeForm
HHHHHHHHH
TT
Tfamily)
H
Origin
COSMEDEURUSMASEUREURNHEAS
ASPAUS
TAM
COS
Mode ofIntro.
AAAAAAAAA
CC
C
O
Record in Hi.First
189519381909186519851865188819091930
18951916
1926
1934
Recent
198419621982197419851984195819801982
19851985
1976
1985
Info.Source
HHHHHHPHH
HH
H
H
No. Spms.
3552231
5553517
5927
14
2CHENOPODIACEAE (Goosefoot family)
Atriplex eardleyaeAtriplex lentiformisAtriplex semibaccataAtriplex suberectaBassia hyssopifoliaChenopodium ambrosioidesChenopodium carinatumChenopodium murale
** Euchylaena tomentosa
HSSHHHHHS
AUSUSMAUSAUSEASNAMAUSEUR-
ACCCAAAA-
198419151895192319831858192218651985
198419741977198419831975198419841985
HHPHHHHHH
11034411
3524431
IslandHi
++++
++
+
+
+
+
++
+++
Ma
+
+
+
++
+
+++++++
Mo
+
++
+
+
+
+++
++
Ln
+
+
+
+
++
+
++
Kh
+
++
+
++
Oh
+
+++++
++
+
+
++
+
+
Ka
+
+
++
++
+
+
+
Ni
+
NW
+
+
+
Taxon
Salicornia virginicaSalsola kali
LifeForm
HH
Origin
NAMEUR
Mode ofIntro.
AA
Record in Hi.First
19681947
Recent
19681975
Info.Source
HP
No. Spms.
13
CLUSIACEAE (Mangosteen family)* Calophyllum antillanum
Calophyllum inophyllumClusia roseaHypericum canarienseHypericum gramineumHypericum mutilumHypericum parvulumHypericum perforatum
TTTSHHHS
TAMASPTAM-P
NAM-EUR
CCOAAAAA
19681000193419851911191119321961
19681986198519851984198319741964
HPHHHPHH
12111158112
COMBRETACEAE (Indian almond family)Terminalia catappa T AS O 1888 1986 P 16
CONVOLVULACEAE (Morning glory family)
*
*
Convolvulus arvensisDichondra repensIpomoea albaIpomoea aquaticaIpomoea batatasIpomoea cairica
var. hederaceaIpomoea hederifoliaIpomoea obscuraIpomoea ochraceaIpomoea purpureaIpomoea triloba
VHVVV
VVVVVV
EURP
COSTOWTAM
cosNWAFAFTAMTAM
AOOCC
AAAOOA
19181967181918881000
184019151926191918261943
19741967197619691984
198319851983198519611985
HHHPP
HHHHPH
51
298
106
805228218
IslandHi
+
++
++++
+
+
+
++++
Ma
+
+
+
+++++
+
+
Mo
+
+
+
+
+
Ln
+
++
Kh
+
Oh
+++
+
+
+++
+
++++
Ka
++
+
+
+
+
+
Ni
+
+
NW
+
+
++
Appendix, continued. 135
Appendix, continued.
Taxon
Ipomoea violacea* Merremia peltata
Merremia tuberosaMerremia umbellataStictocardia tiliifolia
LifeForm
VVVVV
Origin
_
ASTAMTAMAS
Mode ofIntro.
AOOOO
Record in Hi.First
19101911193219111837
Recent
19841911197619851955
Info.Source
HHHHH
No. Spms.
92101024
CORYNOCARPACEAE (Karakanut family)Corynocarpus laevigatus
CRASSULACEAE (Orpine family)Kalanchoe pinnataKalanchoe tubiflora
CUCURBITACEAE (Gourd family)Coccinia grandisCucumis dipsaceusLagenaria sicerariaMomordica charantiaSechium edule
CUSCUTACEAE (Dodder family)Cuscuta campestris
ELAEAGNACEAE (Oleaster family)Elaeagnus umbellata
EUPHORBIACEAE (Spurge family)Acafypha indicaAleurites moluccanaChamaesyce albomarginataChamaesyce hirtaChamaesyce hypericifolia
T
HH
VVVVV
H
S
STHHH
P
AFAF
OWASTOWOWTAM
NAM
AS
ASASPNAMCOSTAM
C
OO
OCCCC
A
O
OCAAA
1891
18881930
19691903100019091965
1955
1963
18881000194718171913
1976
19841983
19861984198619861986
1955
1984
18881973198419831983
P
PP
PHPHP
H
H
PPHHH
27
186
2396
364
1
5
0035316
IslandHi
+
+++
++
+++
+
+
+
Ma
+
+
++
+
+
+
Mo
+
+
+
+
+
++
Ln
++
+
+
+
Kh
+
Oh
+
+++
+
++
++++
+
+
+
+
Ka
++
+
+
+
+
++
+++
Ni
+
+
+
NW
++
Taxon
Chamaesyce hyssopifoliaChamaesyce maculataChamaesyce prostrataChamaesyce thymifolia
* Euphorbia cotinifoliaEuphorbia cyathophoraEuphorbia gramineaEuphorbia heterophyllaEuphorbia peplusJatropha curcasJatropha gossypiifoliaMacaranga mappaPhyllanthus debilisPhyllanthus tenellusRicinis communis
FABACEAE (Pea family)Abrus precatoriusAcacia confusaAcacia famesianaAcacia mearnsiiAlbizia lebbeckAfysicarpus vaginalisCaesalpinia bonducCaesalpinia decapetalaCaesalpinia major
* Caesalpinia pulcherrima
LifeForm
HHHHSHHHHTSTHHS
VTSTTHSSSS
Origin
NWCOSNWOW
TAMNW-TAMHASTAMTAMASASAFAF
ASASTAM-ASASCOSASCOSTAM
Mode ofIntro.
AAAAOOAOAOOCAAC
OCCCCCOOAO
Record in Hi.First
195919801909188819591917197818951855193719501927186519631819
1888191518651911186519091794188818651828
Recent
198319801983197819851980197819841984196319581971197819831974
1986198619761985198319851972198419601985
Info.Source
HHHPHHHHHHHHPHP
PPHPHHHPHP
No. Spms.
1111853131
41181241019620
8292113171837192420
IslandHi
+
++
+++
++
+
++
+
+++
Ma
+
+++
t
++++
++
Mo
+
+++
++
Ln
+
+
+
+
+
Kh
+
++
Oh
+
+++++++++++++
++++
+++++
Ka
+
+
+
+
+
+
+
+
+
+
+
+
Ni
+
+
+
+
+
+
NW
++
+
+
+
+
Appendix, continued. 137
Appendix, continued.
Taxon
Cajanus cajanCanavalia catharticaCanavalia sericeaChamaecrista nictitansClitoria ternataCrotalaria assamicaCrotalaria berteroanaCrotalaria incanaCrotalaria longirostrataCrotalaria micansCrotalaria pallidaCrotalaria retusaCrotalaria spectabilis
* Crotalaria verrucosa* Cytisus monspessulanus
Cytisus palmensisCytisus scopariusDesmanthus virgatusDesmodium cajanifoliumDesmodium heterocarponDesmodium incanumDesmodium intortumDesmodium sandwicenseDesmodium tortuosumDesmodium triflorumDioclea wilsonii
LifeForm
SVVSSHHHHHHHSHSSSSSHHHHHHV
Origin
ASOWP
COSCOSASASTAMTAMSAMOWASASAAMEDAFHASNAMTAMASTAMTAM-
TAMOW
TAM
Mode ofIntro.
COOAOCCCCOCAAOOOOA
CC-AAO
RecordFirst
18651908193018701865186518951895186519111965193118651926195619271909190019331916191619381847191318551825
in Hi.Recent
19861972198519761945197419851976198519831983197519611944195619851909198019841917198319821986198619851974
Info.Source
HHHPHHHHPHHHPHHHHPHHHHHHPH
No. Spms.
1554121057133
2969353832932273323
62183410
IslandHi
+
+
+
+
++
+++
++++++
Ma
+++
+++
+
+
+
+++++
Mo
++
+
++
+
Ln
+
+
+
+
+
Kh
+
+
+++
Oh
++++++++++++++
++++
++++
Ka
++
+
+
+
+
+
++++
Ni
+
+
+
NW
+
Taxon
* Flemingia strobiliferaHaematoxylum campechianumIndigofera spicataIndigofera suffruticosaLablab purpureusLathyrus latifoliusLathyrus tingitanusLeucaena leucocephalaLotus subbiflorusMacroptilium lathyroides
* Medicago intertextaMedicago lupulinaMedicago polymorphaMedicago sativaMelilotus indicaMelilotus officinalisMimosa pudicaMucuna urensParaserianthes falcatariaParkinsonia aculeataPithecellobium dulceProsopis pallidaRhynchosia minimaSamanea samanSenna didymobotryaSenna obtusifolia
LifeForm
STHHVHHSHHHHHHHHSVTTTTVTSS
Origin
ASPTAMTOWTAMASEUR-
TAMOW
TAMEUROWOW
HASHASEASSAMSAMASNW
TAMSAMTAMTAMAF-
Mode ofIntro.
OC
O
OCC
CCAO
OOCOCCA
Record in Hi.First
19101917197718361840198519851837193718651895188818821929190919201855181919301975187018281965191919301945
Recent
19791962197719861971198519851985198319851979198419841982198519841985196119761976198519741965198519861960
Info.Source
HHHPPHHHHHPPPHHPPHHHPPHHHH
No. Spms.
471
482511
244
446193011810208112
2029115133
IslandHi
++
++
++++++
+
++
+
+
Ma
+
+
++
+++++
++
+++
+
+
Mo
+
+
+
Ln
+
+
++
++
+
+
+
Kh
+
+
+
+
Oh
+++++
+
+
+++++++++
+++
Xa
++
+
+++
+
+
+
+
+
Ni
+
NW
+++
Appendix, continued. 139
Appendix, continued.
Taxon
Senna occidentalisSenna pendula
var. advenaSenna septemtrionalisSenna surattensisSesbania sesbanSpartium junceum
* Tamarindus indicaTephrosia purpureaTrifolium atvenseTrifolium dubiumTrifolium pratenseTrifolium procumbensTrifolium repensUlex europaeusVicia angustifolia
LifeForm
S
SSSSSTHHHHHHSH
FRANKENIACEAE (Frankenia family)Frankenia salina
GENTIANACEAE (Gentian family)Centaurium erythraea
GERANIACEAE (Geranium family)Erodium cicutariumGeranium dissectumGeranium homeanumGeranium pusillumGeranium retrorsumPelargonium zonale
H
H
HHHHHS
Origin
TAM
-AATOWEURTOWASPEUREUREUREUREASEUR-
USM
OW
MEDAA-EUR-
AF
Mode ofIntro.
O
OOOCOCCCCCCCC-
A
A
AAAAAO
Record in Hi.First
1836
19081913188819091956179710001932198519321966190919151909
1968
1909
188819101909190918881888
Recent
1983
19791984198519821972198519841984198519821966198419831985
1968
1984
197519481984190919801940
Info.Source
H
HHPHHPPHHHHHHH
H
H
PHHHHP
No. Spms.
26
20202415216661414191013
1
75
15425181
IslandHi
+
++
+++
+++++
+
+++++
Ma
+
+++
+++++
+++
+
+++
+
Mo
+
++
+
+
Ln
++
+
+
+
+
Kh
+
Oh
+
++++
++
+
+
Ka
+
+++++
+
+
+
Ni
+
+
NW
+
+
TaxonLifeForm Origin
Mode ofIntro.
Record in Hi.First Rejcent
Info.Source No. Spms.
HALORAGACEAE (Water milfoil family)Gonocarpus chinensisMyriophyllum aquaticum
LAMIACEAE (Mint family)Hyptis pectinataLeonotis leonurusLeonotis nepetifoliaLeonurus sibiricusMarrubium vulgareMentha pulegiumMentha spicataMentha x villosaOcimum basilicumOcimum gratissimumPlectranthus scutellarioidesPrunella vulgarisSalvia coccineaSalvia occidentalisStachys arvensis
LAURACEAE (Laurel family)Cinnamomum burmanniiPersea americana
LINACEAE (Rax family)Linum bienneLinum trigynum
HH
HHHHHHHHHHHHHHH
TT
HH
-SAM
TAMAFAFASHASOW
EUR
TOWTOW-EASUSMTAMEUR
ASTAM
EUR
-O
AOOOC
C
A
A
C
-A
19661919
193019291938190919131920193619531912192418901909188818351865
19341825
19821912
19831982
198319291985197119781948195919591984198419721975197519761976
19841954
19821985
HH
PHHHHHHHHHPHPHH
HP
HH
27
151
1319822311221315314340
187
13
IslandHi Ma
++
+
+++++++++++++
+
Mo
+
+
+
+++++++
+
Ln
++
++
+
Kh
+
+++
Oh
+
Ka
+
+
++
++++
+++
++
+
Ni
++
++
NW
++
Appendix, continued.
Appendix, continued.
Taxon
LYTHRACEAE (Loosestrife family)Ammannia auriculataAmmannia coccineaCuphea carthagenensisCuphea hyssopifoliaCuphea igrieaLythrum maritimum
MALVACEAE (Mallow family)Abelmoschus moschatusAbutilon grandifoliumGossypium barbardenseGossypium hirsutumHibiscus macrophyllusMalachra alceifoliaMalva parvifloraMalvastrum coromandelianumModiola carolinianaSida acutaSida rhombifoliaSida spinosaThespesia populnea
LifeForm
HHHSSS
HSSSTHHHHSSST
Origin
--
TAMNAMNAMSAM
ASCOSTAMTAMASTAMEURTAMNWTAMCOSTAMAS
Mode ofIntro.
A-AAOA
CACCOAAAAAAAC
Record in Hi.First
193619771855190919151794
1895190318171914191719401825184019111985183718881000
Recent
197419771975198219331975
1941198519851916193619711982198419851985198519811984
Info.Source
HHHHHH
HHPHPHPHHHHHP
No. Spms.
31
418558
2612027732513010438
26MELASTOMATACEAE (Melastome family)
Arthrostema ciliatumClidemia hirtaDissotis rotundifoliaHeterocentron
subtriplinervium
SSH
S
SAMTAMTAM
TAM
OAO
O
193919411958
1926
198519781975
1973
HPH
P
10174
20
IslandHi
++++
++
+++++++
+
+
Ma
+
+
+
+
+++
+
Mo
+
+
+
+
++
+
Ln
+
+++
++
+
Kh
+
++
Oh
+
++++
++++++++
++++
+++
+
Ka
++
+
++
+++
+
+
Ni
++
++
+
NW
+
++
Taxon
Medinilla venosaMelastoma candidumMelastoma sanguineumMiconia calvescensOxyspora paniculataPterolepis glomerataTetrazygia bicolorTibouchina herbaceaTibouchina wvilleanaTrembleya phylogiformis
MELIACEAE (Mahogany family)Melia azedarachToona ciliata
LifeForm
SSSTSSSSSH
TT
Origin
_
---ASTAMTAM-
SAM-
ASAA
Mode ofIntro.
O
OOA
OO
OC
Record in Hi.First
1957191619571985195419491932197919101947
18391929
Recent
1973198219791985197419831985197919831947
19851983
Info.Source
HPHPHHHHPH
PH
No. Spms.
2222051464291
3019
MOLLUGINACEAE (Carpetweed family)Mollugo cerviana
MORACEAE (Mulberry family)Artocarpus altilisBroussonetia papyriferaFicus microcarpaMows alba
MYRICACEAE (Bayberry family)Myrica faya
MYRSINACEAE (Myrsine family)Ardisia crenataArdisia elliptica
H
TTTT
T
SS
-
PASASAS
EUR
ASAS
A
C
OC
C
OO
1975
1000100019431920
1900
19301888
1982
1976198419761984
1976
19851986
H
HPHH
P
HP
3
1671413
38
1630
IslandHi
++
+++
++
+
+
+
+
++
Ma
+
+
++
++++
+
+
Mo
+
++
Ln
++
+
+
Kh Oh
+
+++
+
++
++++
+
++
Ka
+
+
++
++
++++
+
+
Ni
+
++
NW
Appendix, continued.
Appendix, continued.
TaxonLifeForm Origin
Mode ofIntro.
Record in Hi.First Recent
Info.Source No. Spms.
MYRTACEAE (Myrtle family)Eugenia unifloraLeptospermum flavescensLeptospermum laevigatumLeptospermum scopariumMelaleuca quinquenerviaPsidium cattleianumPsidium guajavaRhodomyrtus tomentosaSyzygium cuminiSyzygium jambosSyzygium malaccense
TSSSTTTSTTT
NYCTAGINACEAE (Four-o'clock family)Boerhavia cocdneaMirabilis jalapa
HH
TAMAUSAUSAUPAUPTAMTAMASASAS
ASP
-TAM
CCCCCC
CCC
A
18881927193019271920190818251920188818251000
19741888
19861983198519861985198319861986197819721986
19851985
PHPHPHPPPPP
HP
1581
4431494219271640
1018
OLEACEAE (Olive family)* Chionanthus ligustrina
Fraxinus uhdeiOlea europaeasubsp. africana
Olea europaeasubsp. europaea
TT
T
T
TAMNAM
AF
MED
AC
C
19731926
1932
1910
19731985
1973
1982
PH
H
H
114
7
5ONAGRACEAE (Evening primrose family)
Epilobium billardierianumEpilobium ciliatumFuchsia magellanica
SHS
ASPNAMSAM
AAO
190919291922
198419671983
HPH
37210
IslandHi
++
+++++++
++
+
+
+
+++
Ma
+++
+++
++
+
+
Mo
+
+++
+
+
Ln
++
+++
+
+
+
Kh Oh
++
++++++++
++
+
+
+
+
Ka
+
++++++++
++
Ni NW
+
+
Taxon
Fuchsia paniculataLudwigia octovalvisLudwigia palustrisOenothera affinisOenothera laciniataOenothera stricta
OXALIDACEAE (Wood sorrel family)Oxalis comiculataOxalis corymbosa
PAPAVERACEAE (Poppy family)Argemone mexicanaBocconia frutescensHunnemannia fumariifolia
LifeForm
SHHHHH
HH
HSH
Origin
TAMOW
EURSAMNAMSAM
EURSAM
TAMTAMTAM
Mode ofIntro.
OAAOOO
AO
OOO
Record in Hi.First
193310001934191519331911
10001840
193419201920
Recent
198019751982196119821984
19761985
198419831983
Info.Source
HPHHHH
PP
HHP
No. Spms.
340641420
4515
7169
PASSIFLORACEAE (Passion flower family)Passiflora x caeruleaPassiflora edulisPassiflora foetidaPassiflora laurifoliaPassiflora ligularisPassiflora mollissimaPassiflora pulchellaPassiflora quadrangularisPassiflora suberosaPassiflora subpeltata
VVVVVVVVVV
SAMSAMSAMTAMTAMTAMTAMTAMTAMSAM
O
OCCOOCOC
1865188818881888191019261924188818881914
1960198519841985197719831985194519861986
HPPPHHHPPH
6355914211464
3924
PHYTOLACCACEAE (Pokeweed family)Phytolacca octandra H TAM C 1910 1986 H 52
IslandHi
++++++
++
+
++++++++
Ma
+
++
+
+++
++
++
Mo
+
+
+
+
Ln
+
++
+
+
+
Kh
+
Oh
++
++
+
+
+++++
++++
+
Ka
++
++
+
+
++++
+
+
Ni
+
+
NW
+
+
Appendix, continued.
Appendix, continued.
Taxon
Rivina humilisPIPERACEAE (Pepper family)
Peperomia pellucidaPiper methysticum
LifeForm
H
HS
Origin
TAM
TAMP
Mode ofIntro.
C
OC
Record in Hi.First
1888
19181000
Recent
1985
19861984
Info.Source
P
PP
No. Spms.
15
348
PITTOSPORACEAE (Pittosporum family)Pittosporum undulatumPittosporum viridiflorum
PLANTAGINACEAE (Plantain family)Plantago aristataPlantago australisPlantago debilisPlantago lanceolataPlantago major
POLEMONIACEAE (Phlox family)Gilia capitata
POLYGALACEAE (Milkwort family)Pofygala paniculata
POLYGONACEAE (Buckwheat family)Antigonon leptopusEmex spinosa
* Fagopyrum esculentumMuehlenbeckia axillarisPolygonum argyrocoleonPotygonum avicularePolygonum capitatumPolygonum chinense
T
HHHHH
H
H
VHHSHHHH
TAF
NAM--
HASEAS
NAM
SAM
NAMMEDEASP
ASEURASAS
AUS
AAAAA
A
OACAAAA
O1954
19411908193418951850
1929
1974
19091928192519371937193219601966
18751978
19851985198519851985
1982
1984
19671983192519631937198219831983
1982H
HHHHH
H
H
HHHHHHHP
P5
32495845
5
6
14172312153
IslandHi
+
+
11+
++
++
+
+
+
+++
Ma
+
+
++
+
+
+
+++
Mo
+
+
++
+
Ln
+
++
+
+
Kh Oh
+
++
++
+++
++
+
Ka
+
+
++
Ni NW
++
Taxon
Polygonum convolvulusPolygonum glabrumPolygonum punctatumRumex acetosellaRumex browneiRumex crispusRumex obtusifolius
PORTULACACEAE (Purslane family)Portulaca oleraceaTalinum paniculatumTalinum triangulare
PRIMULACEAE (Primrose family)Anagallis arvensis
PROTEACEAE (Protea family)Grevillea banksiiGrevillea robusta
LifeForm
HHHHHHH
HHH
H
TT
Origin
COSASNW
HASAUSHASOW
COSNW
TAM
EUR
AUSAUS
Mode ofIntro.
AAAAAAA
AOO
A
C
Record in Hi.First
1981181919091895195819101917
181719311979
1895
19091909
Recent
1981198119831975197819781984
198519841979
1985
19821984
Info.Source
HHHHHHH
HHH
P
HH
No. Spms.
143102821616
104101
61
1616
RANUNCULACEAE (Buttercup family)Anenome hupehensisRanunculus muricatusRanunculus parviflorusRanunculus plebiusRanunculus repens
HHHHH
ASEURMEDAUSEUR
AAAA
19421939191119491922
19831943193219701951
HHHPP
82304
RHIZOPHORACEAE (Mangrove family)Bruguiera gymnorrhizaRhizophora mangle
ROSACEAE (Rose family)Eriobotrya japonica
TT
T
ASTAM
AS
C
C
19221902
1915
19461984
1967
PP
H
1041
6
IslandHi
++++++
+++
+
++
+++++
+
+
Ma
++
+
+
+
+
+
Mo
+
+
+
+
Ln
+
+
+
+
+
Kh
+
Oh
+
+
++
+
++
++
+
Ka
+
+
++
+
+
++
Ni
+
++
NW
+
+
Appendix, continued.
Appendix, continued.
Taxon
Fragaria vesca* Prunus x cerasifera
Pyracantha angustifolia* Pyracantha crenati-serrata
Rosa sp.Rubus argutusRubus discolorRubus ellipticusRubus niveusRubus rosifoliusRubus sieboldii
RUBIACEAE (Coffee family)Coffea arabicaGalium divaricatumHedyotis corymbosaMitracarpus hirtusMorinda citrifoliaPaederia scandensRichardia brasiliensisSherardia arvensisSpermacoce assurgensSpermacoce mauritiana
SAPINDACEAE (Soapberry family)Cardiospermum grandiflorumCardiospcrmum halicacabum
LifeForm
HTSSSSSSSSS
SHHHSVHHHH
VV
Origin
EURAFASAS-USM-AS-ASASP
AF-
SAMTAMASPASTAMEURAS-
COSCOS
Mode ofIntro.
CCOOOCCCCCC
CAAACAAAAA
OO
Record in Hi.First
18291933193719451910189419831961196518801971
1823198119651982100018541855190919291973
19511817
Recent
19831975198319841983198319831984196519671976
1985198119861982198519851983198419861983
19851984
Info.Source
PPHHHPHHHPH
PHHHPPPHHH
HH
No. Spms.
206742453251
443
27181
60393619234
527
SCROPHULARIACEAE (Figwort family)Antirrhinum orontium H MED O 1950 1960 H 4
IslandHi
++++++
+++
+
++++++++
+
Ma
+
+++
+
+++
+
++
+
Mo
+
+
+
+
Ln
++
+
+
+
Kh Oh
++
++
+
+
+
++
+
++
+
Ka
+++
+
+
+
+++
+
+
Ni
+
+
NW
Taxon
Castilleja arvensisDopatrium junceumLinaria canadensis
var. texanaLindemia CrustaceaLophospermum erubescensMaurandya antirrhinifloraOrthocarpus purpurascensParentucellia viscosaTorenia asiaticaVerbascum blattariaVefbascum thapsusVerbascum virgatumVeronica arvensisVeronica plebeiaVeronica serpyllifolia
SOLANACEAE (Nightshade family)Brugrnansia CandidaCapsicum annuumCestrum diumumCestrum noctumumDatura stramoniumLycopersicon esculentumLycopersiconpimpinellifolium
Nicandra physalodes
LifeForm
HH
HHHHHHHHHHHHH
SHSSHH
HH
Origin
TAM-
NWTOWNAMNAMUSMEURASHASHASEASEURAUSNH
TAMTAMTAMTAMASSAM
SAMSAM
Mode ofIntro.
AA
AAOOA-OAAAAAA
OCOOACCO
Record in Hi.First
19681977
1904195419171943194519711975195519321943194319111926
189018151888192718371888
18951850
Recent
19831977
1979198419821943194519711983195519751982198519741983
197619851984198519851983
19861985
Info.Source
HH
PPHHHHHHHHHHH
HPPHHP
HP
No. Spms.
61
1571411141
142103120
23162018201
4024
IslandHi
+
+++
+++++++++
+
++
++
Ma
+
+
++
+
++
Mo
+
+
+
+
Ln
+
+
Kh Oh
+
++
+
++++++
++
Ka
+
+
++++
++
Ni
+
NW
Appendix, continued. 149
Appendix, continued.
Taxon
Nicotiana glaucaNicotiana tabacumPhysalis angulataPhy satis peruviana
* Salpichroa rhomboideaSolatium aviculareSolatium capsicoidesSolatium elaeagnifoliumSolanum linnaeanumSolatium mauritianumSolanum pseudocapsicumSolanum robustumSolanum seaforthianumSolanum torvumSolanum tuberosumSolanum viride
* Streptosolen jamesoniSTERCULIACEAE (Cacao family)
Melochia umbellataTHEACEAE (Tea family)
Camellia sinensisTILIACEAE (Linden family)
Heliocarpus popayanensisTriumfetta rhomboideaTriumfetta semitriloba
LifeForm
SHHHHSSHHHSSVSBHH
T
S
THH
Origin
SAMSAMTAMSAMSAMOW
TAMUSMMEDSAMOW-
TAM-
SAM-
TAM
AS
AS
TAMAFCOS
Mode ofIntro.
AC-OOAAAAOOAO-C-O
O
OAA
Record in Hi.First
18651812197618251931195518371958189519091888197719161954181119371932
1929
1932
194118881910
Recent
19821986198319851985196419851979198319851986197719851965191919371974
1984
1977
196819761983
Info.Source
HPHPHHHPHHPHHHPHH
H
H
PHH
No. Spms.
314226145
3433329241143217
15
5
62720
IslandHi
+
+
+
+
+
+
+
+++
Ma
++
+
+
+
+++++
+
+
+
Mo
+
+
+++
+
+
+
Ln
++
+
++
+
+
Kh
+
+
Oh
++
++
+++++
+
+
+
+
+++
Ka
++++
+
+
+
++
+
Ni
+
NW
+
TaxonLifeForm Origin
Mode ofIntro.
Record in Hi.First Recent
Info.Source No. Spms.
TROPAEOLACEAE (Nasturtium family)Tropaeolum majus
TURNERACEAE (Turnera family)Tumera ulmifolia
ULMACEAE (Elm family)Trema orientalis
URTICACEAE (Nettle family)Boehmeria niveaLaportea interruptaPilea microphyllaUrtica urens
VALERIANACEAE (Valerian family)Centranthus ruber
VERBENACEAE (Verbena family)Citharexylum caudatumCitharexylum spinosumClerodendrum philippinumLantana camaraStachytarpheta dichotomaStachytarpheta jamaicensisStachytarpheta mutabilisStachytarpheta urticifoliaStachytarpheta x
intercedensStachytarpheta x trimeniVerbena bonariensis
V
S
T
HHHH
H
TTSSHHHH
HHH
SAM
TAM
AS
ASASPTAMOW
HAS
TAMTAMASTAMTAMTAMTAMCOS
ASHORSAM
O
C
C
OAOA
O
OOOOAAOA
AAA
1840
1888
1870
1928181919261909
1965
19291937183918581910189518881895
193919281914
1975
1984
1986
1976195919851975
1982
19831985197819761978198319801973
194019511918
P
P
H
HPHH
P
PHHPHHPH
HHH
4
10
50
12128
1
17817504623719
224
Hi
+
+
++
+
++++++
Ma
+
+
+
+
+
++
+
Mo
+
+
+
+
+
Ln
++++
+
IslaKh
adOh
+
+
+++
++++++++
+
Ka
+
+
+++
++
+
Ni
+
NW
+
Appendix, continued. 151
Wester \ Origin and Distribution of Alien Plants 153
Literature Cited
Baker, H.G. 1965. Characteristics and modes of origin of weeds. In The genetics ofcolonizing species, ed. H.G. Baker and G.L. Stebbins, 147-172. New York: Acad. Pr.
Beaglehole, J.C. 1967. The journals of Captain James Cook. The voyage of Resolutionand Discovery 1776-1780. Vol.3. Cambridge, England: Cambridge Univ. Pr.
Biggs, B. 1979. Pro-Polynesian word list II. Working papers in anthropology,archaeology, linguistics and Maori studies, No. 53. Anthropology Dep., Univ.Auckland, New Zealand.
Conner, E.F., and D. Simberloff. 1978. Species number and compositional similarity of theGalapagos flora and avifauna. Ecol Monogr. 48:219-248.
Davis, P.H., and V.H. Heywood. 1965. Principles of angiosperm taxonomy. Princeton,New Jersey: Van Nostrand.
Degener, O. 1962. Flora hawaiiensis: Digitariapruriens. Privately published.
Frenkel, R.E. 1984. Weeds and aliens: a conceptual and terminological jungle. Ann.Meet., Assn. Amer. Geographers, Washington D.C. April 22-25. Mimeo.
Gutmanis, J. 1979. Kahuna la`au lapa`au. Honolulu: Island Heritage.
Handy, E.S.C., and E.G. Handy. 1972. Native planters of old Hawai`i: their life, lore,and environment. B.P. Bishop Museum Bull. 233. Honolulu: Bishop Mus. Pr.
Harlan, J.R., and J.M.J. de Wet. 1965. Some thoughts about weeds. Econ. Bot.19:16-24.
Hart, R. 1976. An index for comparing weediness in plants. Taxon 25:245-247.
Hastings, J.R., R.M. Turner, and D.K. Warren. 1972. An atlas of some plantdistributions in the Sonoran Desert. Meteorology and Climatology Arid Regions Tech.Rep. 21. Tucson, Ariz.: Univ. Arizona Inst. Atmospheric Physics.
Healy, A.J. 1947. Evolution of the alien flora of New Zealand. Proc. N.Z. Sci.Congress, 160-162.
Hillebrand, W.F. [1888] 1965. Flora of the Hawai`ian Islands. New York and London:Hafner.
Holland, P.G. 1975. The use of herbarium records in biogeography. ProfessionalGeographer 27:475-479.
Hooker, WJ., and GA. Arnott. [1830] 1965. The botany of Captain Beechey's voyage:Sandwich Islands. Weinheim: Historiae Naturalis Classica 39:78-111.
Kirch, P. 1985. Feather gods and fishhooks. Honolulu: Univ. Hawai`i Pr.
Appendix, continued.
Taxon
Verbena litoralisVerbena rigida
VITACEAE (Grape family)Cissus nodosa
LifeForm
HH
VZYGOPHYLLACEAE (Creosote bush family)
Tribulus terrestris V
Origin
TAMSAM
AS
OW
Mode ofIntro.
AO
O
A
Record in Hi.First
18351932
1975
1947
Recent
19751938
1975
1984
Info.Source
HH
H
H
No. Spms.
613
1
5
IslandHi
++
+
+
Ma
+
Mo
+
Ln
+
Kh Oh
+
Ka
+
+
Ni
+
NW
* Taxon is not discussed in Wagner et al. 1990.** Taxon is not discussed in Wagner et al. 1990 or St. John 1973.# Tricachne insularis = Digitaria insularis; taxon is retained to avoid loss of data.
## Bidens lanceolata = Coreopsis lanceolata; taxon is retained to avoid loss of data.
152
Wester \ Origin and Distribution of Alien Plants 154
Kornas, J. 1971. Changements recents de la flore polonaise. Biol. Conserv. 4:43-47.
Krauss, B.H., comp. 1974. Ethnobotany of Hawai`i. Botany Dep. Honolulu: Univ.Hawai`i.
Mann, H. 1866. Flora of the Hawai`ian Islands. Communications Essex Institute5:113-248,6:105-111.
Mann, H. 1868. Enumeration of Hawai`ian plants. Proc. Amer. Acad. Arts and Sci.7:143-235.
Merrill, E.D. 1946. Man's influence on the vegetation of Polynesia, with specialreference to introduced species. Chronica Botanica 10:334-45.
Merrill, E.D. 1954. The botany of Cook's voyages. Chronica Botanica. 14:164-384.
Nagata, K.M. 1985. Early plant introductions to Hawai`i. Hawn. J. History 19:35-61.
Obata, J.K. 1985. Another noxious melastome? Oxyspora paniculata. Newsletter,Hawn. Bot. Soc. 24:25-26.
St. John, H. 1973. List and summary of the flowering plants in the Hawai`ian Islands.Mem. 1. Lawai, Kauai, Hawai`i: Pac. Trop. Botan. Gdn.
St. John, H. 1978. The first collection of Hawai`ian plants by David Nelson in 1779.Hawai`ian Plant Studies 55. Pac. Sci. 32:315-324.
Smith, C.W. 1985. Impact of alien plants on Hawai`i's native biota. In Hawai`i'sterrestrial ecosystems: preservation and management, ed. C.P. Stone and J.M. Scott,180-250. Univ. Hawai`i Coop. Natl. Park Resour. Stud. Unit. Honolulu: Univ. Hawai`i Pr.
Smith, J.B.M. 1982. Establishment of exotic woody plants along roadsides in New England(New South Wales). Research Ser. Applied Geogr., Geography Dep. Armidale,Australia: Univ. New England.
Stearns, W.T. 1951. Mapping the distribution of species. In The study of thedistribution of British plants, ed. J.E. Lousey, 48-64. Oxford: Botan. Soc. BritishIsles.
Strother, J.L., and A.R. Smith. 1970. Chorology, collection dates and taxonomicresponsibility. Taxon 19:871-874.
Wace, N.M. 1979. Assessment of dispersal of plant species ~ The car-borne flora in Canberra. Proc. Ecol. Soc. Australia 10:167-186.
Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 1990. Manual of the flowering plants ofHawai`i. Bishop Mus. Spec. Pub. 83. Honolulu: Univ. Hawai`i and Bishop Mus. Pr.
Wester, L., and J.O. Juvik. 1983. Roadside plant communities on Mauna Loa, Hawai`i. /.Biogeography 10:307-316.