Introduction to Parasitic Flowering Plants

Updated 2010

Nickrent, D.L. and Musselman, L.J. 2004. Introduction to Parasitic Flowering Plants. The Pl

Daniel L. Nickrent and Lytton J. Musselman

1. Department of Plant Biology, Southern Illinois University 2. Department of Biological Sciences, Old Dominion University The authors gratefully acknowledge information and ideas provided by Drs. Frank Tainter and Robert Eplee in the early stages of this publication.

INTRODUCTION

Fungi, nematodes, bacteria, and viruses are probably the first things th organisms certainly do cause damage to plants of economic importanc plants are also important pathogens. The purpose of this chapter is to relationships of these fascinating and unusual plants and also to focus

Most plants are autotrophs and produce their own carbon sources thr pipe (Monotropa) (Figure 1) lack chlorophyll and appear to be parasitic, and, hence, only indirectly parasitize the trees on which the mycorrhiza angiosperm (flowering plant) that directly attaches to another plant via morphological and physiological link between the parasite and host (Figu terms "parasite" and "pathogen." Parasite is from the Greek para (bes food". If a plant also induces disease symptoms in a host, then it is a p parasites and mycotrophs that derive carbon from sources other than t means "different feeding."

Figure 1.

Types of Parasitic Plants

Two main types of parasitic plants can be distinguished: stem parasite and pathogenic members include some mistletoes (see below) and do common and are found in diverse taxonomic groups. Some of the mos family, Orobanchaceae. Parasitic plants may also be classified as holo parasites (Figure 3). Facultative parasites contain chlorophyll and can the other hand, require a host for maturation.Hemiparasites contain c water, with its dissolved nutrients, by connecting to the host xylem via nonphotosynthetic) and must rely totally on the contents of the host xy Although these definitions imply absolute and discrete categories, som holoparasitic condition, e.g.Cuscuta (dodder).

Figure 3.

Morphological Features

In some stem parasites, such as Cassytha (Figure 4) and Cuscuta (dod and scale leaves. In contrast, many of the photosynthetic root hemipar casual observer because they are green with fully formed leaves. As th evolution from hemiparasitism to holoparasitism), profound changes oc holoparasites tend to have leaves reduced to scales (or absent in Hyd (derived from the seedling radicle). The best examples of the evolution various representatives of the broomrape family (Orobanchaceae, see

Figure 4.

The sandalwood order (Santalales), has no true holoparasites (excludi branching three families in this order are autotrophic (lacking haustoria hemiparasites.

Another family within this order, Loranthaceae, contains stem parasitic Saxon word Misteltan derives from the Old German word "mist" for dun is an apropos name given that birds defecate the seeds of mistletoes o with brittle stems and leaves. However, three genera of Loranthaceae most primitive in the family. Adult plants of dwarf mistletoe (Arceuthobi their hosts for carbohydrates. Despite this, their seedlings are photosy

Figure 6.

The range of flower sexual conditions (e.g. bisexual and unisexual) in separate male and female plants) is more frequent among heterotroph autotrophic plants. Of the 959 dioecious plant genera, at least 43 (4%) explained.

Evolutionary Development of Parasitism

What evolutionary conditions might favor the development of parasitism especially for annual plants. When the parasite seedling forms a haust "assuming" the root system of its host plant. Does it "cost less" for the rather than produce one of its own? Resource allocation studies of par hemiparasites have transpiration rates higher than their hosts, and so have dense groundcover vegetation and competition for resources in t the evolution of root parasitism.

The stem parasitic mistletoes also exceed their hosts' transpiration rat abundant in areas where access to sunlight is not limited, such as sav avoided. Recent work has shown that diversification in Loranthaceae o deciduous woodlands and grasslands were displacing tropical biomes. evolution of aerial parasitism (mistletoes) can be traced to root parasiti represented an unexploited niche that offered opportunities for coloniz

Host Interactions

The breadth of the modes of host selection and specialization of paras and Cuscuta (dodder) (Figure 8) can parasitize hundreds of different ho drops) (Figure 9) occurs only on Fagus grandifolia(beech). The same ge species are generalists and others specialists. Which strategy do you t evolutionary time?

Figure 7.

Figure 8.

The terms host range versus host preference describe different aspe number of different species that can be parasitized. For example,Seym but in pots a variety of angiosperms and gymnosperms can be parasiti desirable host for optimal growth, typically is much narrower. Cuscuta and can even be attached to many different hosts at once. But in natur often be located by first finding the preferred host.

Figure 10.

There has been considerable research on the chemical signals betwee indicates that chemical signaling is also common in root parasites, esp

most vulnerable part of the life cycle because life-sustaining attachmen (Figure 11) with minuscule food reserves, this period is especially critica attachment. Parasites with larger seed reserves can survive longer. In mm long, require a host stimulant to germinate whereas larger seeds d stimulants that may counteract or compete with the root stimulants.

Figure 11.

The stem parasitic dodders (Cuscuta) provide intriguing models for ho response to a host stimulus) that allow them to "forage" for (or move to that Cuscuta europaea selects shoots with the highest nutritional statu

We have very little information about host resistance mechanisms, exc fascinating avenue for further research would be to examine the anato Although the host-parasite relationship defines parasitic plants, we still selected, why some plants are favored, and why some are avoided en

Figure 12.

Distribution

Most root-hemiparasitic plants occur in temperate regions, particularly sunny habitats. For example, Orobanchaceae are most specialized in Southeastern United States (open, fire-maintained communities), and seems reasonable that many could potentially pose problems to crops southeastern United States. Thesium (Thesiaceae, formerly Santalace and about half of these occur in South Africa. One Eurasian species, T in the USA and Alberta, Canada. This species favors open or disturbed and pasturelands.

Dispersal

Like many weeds, several pathogenic parasitic plants have been effec North American dodder (Cuscuta campestris = C. pentagona) (Figure 1 most widespread parasitic weed in the world, and was first described f frequently intercepted by port inspectors, confirming that ports are foci implicated in the spread of Orobanche minor (Figure 14). Another sourc improved cultivated varieties of crop plants. The parasite seeds enter a have been identified as foci for the spread of Striga (witchweed) seed areas where improved crops have not been introduced, the likelihood o where subsistence grains contain little exotic germplasm, Striga is a lim

Figure 13.

While the spread of some root parasites and Cuscuta by humans is we mistletoe. Around 1900 in California, Luther Burbank introduced Viscu for sale at Christmas. An evaluation of its subsequent spread shows th miles) per year. It is interesting that the dwarf mistletoes (Arceuthobium have not spread to any pine species in the southeastern U.S. This pro and host resistance.

Figure 15.

Occasionally, indigenous plant parasites become serious pests when h examples in the American South: Seymeria cassioides (Figure 10), an a pubera (Figure 16), a shrub in Santalaceae that is endemic to the Appal of West Virginia have been attacked by P. pubera (Figure 17), thus redu of Seymeria in the late 1960s when slash pine (Pinus elliottii) was dam cassioides are exacerbated by drought conditions.

Figure 16.

A review of alien plants in California lists 151 alien species not include about 25 years before. Of these new aliens, only one is a plant parasit relatively benign root parasite. This suggests that parasitic plants spre USDA APHIS PPQ requires permits to import or move any parasitic pl five genera of parasitic plants that are considered to be the most serio and Striga.

Figure 18.

ECONOMICALLY IMPORTANT PARASITIC PLANTS Cassytha, Laurel Dodder (Lauraceae-Laurel Family) Cuscuta, Dodder (Convolvulaceae-Morning Glory Family) Orobanchaceae (Broomrape Family) Striga (Witchweeds-Broomrape Family)

Orobanche (Broomrapes-Broomrape Family) Sandalwood Parasites Loranthaceae (Showy Mistletoe Family) Santalaceae (Sandalwood Family) Viscaceae (Christmas Mistletoe Family)

Cassytha, Laurel Dodder (Lauraceae-Laurel Family)

Cassytha is a high climbing, parasitic vine with green or orange stems to Cuscuta (dodder). Cassytha is regularly mistaken for dodder, even b two unrelated genera is one of the most striking parallelisms among th both Cuscuta and Cassytha are hard seeds that require scarification (b and simultaneous attachment of a single parasite to several diverse ho

Cassytha is the only parasitic member of the otherwise autotrophic Lau its vigor (Figure 19) and wide distribution, it isremarkable that it is only o in Australia, but C. filiformis is pantropical and is also abundant in sout introduced species could easily be overlooked.

Figure 19.

Cuscuta, dodder (Convolvulaceae-Morning Glory Fam

Species of Cuscuta, commonly known as dodder, are among the best dodders was reviewed in Dawson et al. (1994). Dodders have a broad The genus Cuscuta contains three subgenera. Members of the subgen

trees, while species in the subgenus Cuscuta are more delicate in stru entirely New World subgenus, Grammica.

Figure 13.

Dodders may be the most important parasitic weeds of legumes in tem campestris (Figure 5) on alfalfa (Medicago sativa). The alfalfa and dodd the host. The wide range of hosts attacked by dodders is reviewed in D seed sanitation. Because the surface of dodder seeds is minutely roug pass over. Dawson et al. (1994) also reviewed several herbicide treatm dodder.

Figure 5.

Orobanchaceae (Broomrape Family)

This family includes the largest number of genera (90) and species (ca past, the hemiparasitic members of this family were classified as part o members were included in Orobanchaceae (the broomrape family). Re and holoparasites show that Orobanchaceae is monophyletic (i.e. all m

line is descended from nonparasitic Scrophulariaceae. The two most e and Orobanche (broomrape), are discussed below, with particular atte

Striga (witchweeds)

Witchweeds (Striga spp.) have a greater impact on humans worldwide subsistence crops grown widely in Africa and Asia. Such crops include sugar cane and legume crops such as cowpea and ground nut. The na on their host in which most damage occurs before the parasite is visibl in the Latin name, which means "hag" or "witch."

Striga is an obligate hemiparasite that reaches its greatest diversity in Far East, and Australia. Two species, S. asiatica (Figure 12) and S. her worldwide. Most Striga species have complex life cycles. Several discr conditioning, haustorial induction, attachment, penetration, seedling de summary is based on reviews in Sauerborn (1991), Parker and Riches

Figure 12.

Striga produces thousands of "dust" seeds (Figure 11) per capsule that season. Movement of seeds in nature is usually attributed to wind and are dormant for several months before they will respond to chemicals e ripening." We know virtually nothing about what controls after ripening

process. Conditioning occurs when two environmental factors are pres and adequate moisture. The moisture content of the soil must be near after conditioning, it is able to respond to chemical signals (Figure 21) fr and the distance to the host. Recent work has shown that host roots e mycorrhizal fungi colonization as well as regulate branch root formatio chemicals as germination stimulants. Germination in obligate root parasites is usually cryptocotylar, that is, t radicle emerges. After germination, Striga causes the release of chem formation. One such chemical is 2, 6-dimethoxybenzoquinone, a degra tip begins to produce structures that superficially resemble root hairs (F 23). Similar hairs are produced by facultatively parasitic members of O suitable, the haustorium penetrates and forms a link with the host vasc

Figure 11.

Figure 22.

As the parasite becomes established and matures, the distinctive seed specialized part of the life cycle is subterranean, lacks chlorophyll, pos

roots from which additional (lateral) haustoria can arise. The seedling e of the host by altering hormone balances and stimulating root producti stage, emergence from the soil, is critical for the parasite but poorly un for emergence? Does the condition of the host control emergence? Or develops, flowers form, and the life cycle is completed when the seeds

Figure 24.

Why have so few Striga species become serious pests? Mohamed et a S. hermonthica and S. asiatica are true agrestals, that is, plants assoc pathogens are spread with their hosts, are more abundant in crops tha Two observations support this. First, S. hermonthica and S. asiatica ar where there has been little introduction of improved grain varieties, Str situation exists in Togo where S. asiatica became a serious problem o grown.

When Striga was identified in the USA in 1956, federal and state action facilities were established to conduct basic and applied research to ga methodology to eradicate the infestation. Early research revealed that volatile form of the selective phenoxy herbicide 2,4-D. This method sto damage to the crop. Motorized high clearance sprayers and backpack (thousands of hectares) called for more effective methods. Moreover, t

on weedy grasses in broadleaf crops such as cotton, soybeans and ve herbicide.

A major problem in witchweed control is the persistence of the microsc identify the seed germination stimulant. A breakthrough occurred when germination. New equipment and application methodologies to introdu proper soil conditions, the injection of 1.5 kilograms per hectare of ethy viable Striga seed bank in a growing season. Through implementation the USA has been virtually eradicated. There still remain, however, a f monitoring and further treatment.

Figure 26.

Orobanche (Broomrapes-Broomrape Family)

The genus Orobanche, as currently classified, contains ca. 150 specie Worlds. These plants are known by the English name broomrape beca brooms (the common name for the legume Cytisus). The genus reache Western Asia. Most of the economically important pathogens are Old W not monophyletic, thus other generic concepts may eventually be appl

Major crop hosts for Orobanche are legumes, solanaceous crops (egg umbels (carrot, parsley, celery), cole crops (cabbage, cauliflower), lettu the soil (more than five decades), small seed size (less than the width and a subterranean phase (seeds germinate beneath the soil and para Broomrapes have their greatest impact in the Balkans, the Nile Valley, level of infestation, and total crop failures have occurred in some case

There have been numerous studies of the host range of Orobanche sp can parasitize plants from 11 different dicot families, in fact, more diffe important hosts include solanaceous crops, cabbage, cauliflower, hem aegyptiacacauses especially severe damage to melons in Central Asia the melons, but also induces the production of a toxin within the melon

Figure 27.

Evaluation of host range for parasitic plants is complicated by the fact field. For example, there are no reports of field parasitism of soybean b crenata in pots. Nonetheless, pot experiments are valuable because p parasites can be determined.

Germination requirements for Orobanche are different from those of S so germination temperatures range from 10 to 20C (50 to 68F), alter explain why O. ramosa is a problem in the Nile Valley of Sudan only in tobacco in the winter in India, but is not a problem on sunflower (Helia UnlikeStriga, Orobanche seedlings produce roots that are geotropically gravity. Orobanche also differs from Striga in the way the host root sys its host. An exception is O. cernua, which often forms a large, single h important for witchweed germination and a useful control for that paras

The longest documented viability of Orobanche seeds in the soil was a Tobacco Research Institute near Plovdiv were replanted in grapes due removed and tobacco replanted. Large numbers of O. ramosa emerge last time tobacco had been grown!

Orobanche ramosa has been introduced to California where it has bee effort to eradicate it, the parasite still persists. It is likely that seeds of t contaminated plants or contaminated tomato seeds. The more recent i we can expect O. ramosa to establish itself wherever suitable hosts ar

Figure 29.

Orobanche cernua (Figure 30) is widespread in Eastern Europe and the scattered occurrences in North Africa, China, and southern Europe. Its Sunflower is the most important oil crop in parts of eastern Europe, an Bulgaria where sunflower oil is the national favorite (Figure 31). Infected and poorer oil quality. Between 1947 and 1950, the broomrape problem continued existence of sunflower cultivation. New, resistant varieties w apparently selected for virulence in broomrape, with a resultant rapid lo documented in Bulgaria. Tobacco can also be severely damaged by O cernua, even in fields adjacent to heavily infested sunflower fields. The damaged and sunflower has not been attacked. It has been shown tha

and O. cernua from sunflower will not attack solanaceous crops. For m development of races in O. cernua. Their data should be re-examined pathogen is needed, considering the acreage devoted to sunflower in t

Figure 30.

The last of the major broomrapes to consider is O. crenata (Figure 32). carrots, parsley, and celery. It is restricted to Europe, the Middle East a the cold hardiness of this species, which has the northernmost range o this seems the least likely to be introduced and to become established major crops there. But damage is severe and commodities from infecte

Figure 32.

Two other genera of the Orobanchaceae, Aeginetia and Christisonia, m hosts for most broomrapes. Aeginetia indica (Figure 33) can be damagi

wightii damaging sugarcane in the Philippines.

Figure 33.

Sandalwood Parasites

The Sandalwood order (Santalales), as traditionally defined, contains p autotrophs, root hemiparasites, and stem hemiparasites. The mistletoe mistletoe refers to the plant habit (stem parasite), not just one taxonom order contains 160 genera and ca. 2300 species distributed worldwide human activity, four families deserve mention: Loranthaceae, Thesiace

Loranthaceae (Showy Mistletoe Family)

This large, mainly tropical, family is composed of 74 genera and ca. 90 members have large showy flowers (Figures 34 and 35) that are bird-poll reached a high level in this family, as evidenced also by the seed dispe genus Dicaeum have tongues that are specifically modified to sip nect that pass the viscid seeds in a remarkably short period of time. Many o has proposed that mistletoes function as keystone resources in many that positively affect diversity in these habitats.

Figure 34.

At least 30 genera of mistletoes in the family Loranthaceae occur on in reported to be particularly damaging: Tapinanthus bangwensis (Figure pentandra (Figure 36) on kapok (Ceiba pentandra) in Java, Phthirusa sp and Oryctanthus sp.(Figure 38) on cacao in Costa Rica. Four genera of andTapinanthus) are particularly damaging to the shea butter or karit the 16,000 trees examined in one study, 95% were parasitized by one mistletoes generally involves pruning the mistletoe and/or the host bra

Figure 35.

Figure 37.

A discussion of a typical member of Loranthaceae ("loranth" here) follo flowered Old and New World species, whereas insects are the pollen v

bisexual flowers, whereas others are dioecious or monoecious with un angiosperms in having an "aggressive" embryo sac which actually gro some species. After fertilization, the embryo and seed begin to form w single-seeded berry (or drupe, but without a stony endocarp), and the viscin, composed of cellulosic strands surrounded by mucilaginous pec dispersal. Unlike Orobanchaceae, loranth seeds do not require a host however, establishment only occurs on living hosts. Emerging from the phototropic and thus grows towards a dark surface (often the host bran a holdfast (Figure 42), and the cotyledons either remain within the endo and expand (e.g. Psittacanthus) (Figure 43). The haustorium forms from first aerial shoots of the mistletoe form from the epicotyl and, in some s haustorium along the host branch. These epicortical roots form new ha host branches (Figure 45). When in contact with the host cambium, the that enlarges in fluted columns, eventually forming a placenta-like sadd mistletoe haustorial tissue (composed mainly of parenchyma) decays, objects (Figure 47).

Figure 39.

Figure 40

Figure 42.

Figure 43

Figure 45.

Figure 46.

Sandalwood Family

The root-hemiparasitic sandalwood family or Santalaceae has recently a plant pathology perspective, few members of Santalaceae have a ne true for Santalum album, which is extensively cultivated because it is t perfumes, cosmetics, and medicine. Two families that have pathogenic Although Thesium contains weedy species, only a few cause significan can be a pathogen of trees under some circumstances in the eastern U aubrevillei of West Africa, also a member of this family, can inflict seve

Figure 16.

Viscaceae (Christmas Mistletoe Family)

The mistletoe family Viscaceae is small in terms of the number of gene Most of these species occur in two genera: Viscum (of the Old World) derives from a feature of the seeds - the viscin - that forms a sticky lay the seeds to the host branch. These are the mistletoes most often reco the green leafy shoots with white berries (Figure 50) often festoon doorw associated with mistletoe, see the Feature Story by Tainter (Decembe

Figure 48.

Figure 49.

Mistletoes in Viscaceae impact both positively and negatively on huma album (Figure 15) is used medicinally, for example to treat various form is questionable, there is growing scientific evidence of therapeutic acti used to treat ovarian cancer. The other major compounds extracted fro only have immunomodulatory effects, but are also strong cytotoxins. T

pose a safety risk for small children who may ingest them.

Figure 15.

Certainly the greatest economic impact on human activity is caused by "dwarf mistletoes" because some species are diminutive, their damagi America are enormous: 11.3 million cubic meters of wood is lost annua such asPhoradendron and Viscum can also cause damage to host tre scale as Arceuthobium. The leafy mistletoes occur on both hardwood a from conifer hosts (families Pinaceae and Cupressaceae).

Figure 6.

The life cycle of viscaceous mistletoes is similar to that described abov generally effected by insects and wind, and the flowers in this family ar unisexual flowers on spikes or cymes (Figure 51) (Figure 52) (Figure 53). I haustorium of Viscaceae never forms epicortical roots, but instead form structure is composed of portions that run parallel to the host branch a Other tissues called sinkers descend perpendicularly from the cortical are water hemiparasites, hence they manufacture at least some of the

documented that leafy mistletoes such as Phoradendron actually trans when host leaves are absent. In contrast, Arceuthobium (dwarf mistlet

Figure 51.

Figure 53.

Complex types of haustoria are found in Arceuthobium. In some cases the host is a smaller branch distal to the swollen infection. In other cas mistletoe haustorium that is composed of a dense group of host branc (Figure 55) where the mistletoe endophyte does not enter the witches' b present, sometimes even in the host branch apical meristem. When th also divide in a coordinated fashion. Systemic witches' brooms, as see the extremely intimate cellular relationship that has evolved between h

Figure 55.

In Viscum, Phoradendron, and most of the other genera of Viscaceae, of Arceuthobium, birds are not the primary seed vectors. Instead the fr The fruit walls are very elastic, and turgor pressure develops as the fru begins to weaken and, with the slightest disturbance, the fruit separate mechanism explosively expells the single seed at a rate of 27 meters ( to 16 meters (17.5 yards) before sticking to a host surface (Figure 59). M swell, and the seed may then slide to the base of the needle fascicle w are wasted in this process, one plant may produce hundreds of fruits a tree branches. The seeds can only penetrate younger host tissue beca

Figure 57.

Figure 59.

As with loranths, the seedling radicle forms a holdfast when it encount penetrates the host from within the holdfast. Unlike loranths and most mistletoes) are not formed from an epicotyl; once the endophyte is insi endosperm whither and die. The endophyte then develops within the h adventitiously from the cortical strands (Figure 61). In nature, various dw go from seed to seed.

Figure 61.

Controlling Dwarf Mistletoes - a Continuing Challenge

Dwarf mistletoes are often a serious threat in extensively managed for Although there is no relationship between presence of dwarf mistletoe trees growing on a poor site are invariably more seriously affected than form of control is to determine site quality before dwarf mistletoe has b infected trees during pre-commercial and commercial thinnings might b heavy dwarf mistletoe infestation might dictate early harvest or even a value stands such as seed orchards or recreation areas, severely infec

infected branches can be effective if more than half the crown will rem Forest Service has developed a computer program that is used by fore

Natural wildfires effectively eradicated dwarf mistletoe infections in the as a possible control agent. Fire has been most effective withArceutho species spreads locally to form discrete infection centers with a distinc fire has been less effective in western North American forests because centers. Instead, economic losses attributed to dwarf mistletoes are ef stand site quality and then reduce the incidence of infected trees by th planted to replace a susceptible species presently on the site. For exa hardwoods such as aspen or cottonwood.

Chemical control through the use of herbicides has limited success, m agents that affect only the mistletoe and not the host. An environmenta mistletoe in some situations. When applied in summer, the dwarf mistl produced. The chemical will not affect the endophytic system, so the in new dwarf mistletoe shoots appear. This technique is only useful wher

Various means of biological control have been investigated, including i with Arceuthobium, hence it is unlikely they can be used to dramaticall native to North America, Asia, Europe, and Africa - the only mistletoe t plants have been coevolving with their hosts for millions of years, so it dwarf mistletoe witches'brooms provide roosts and nesting sites for ma thus there is a growing tendency to take an enlightened view of the rol (Watson 2001), and to modify management practices with this view in

CONCLUSION

Although 270 genera and 4400 species of flowering plants are parasiti cultivated by humans and are thus considered pathogens (Table 1). Am pests: Striga (witchweed), Orobanche (broomrape), Cuscuta(dodder), most felt by farmers growing subsistence crops such as maize, sorghu The broomrapes are pests worldwide but are particularly damaging to East. Striga and Orobanche produce thousands of tiny seeds that pers mistletoes represent one of the major disease organisms affecting com of the ecosystem that have coevolved with their hosts for millions of ye approach. Understanding the complex and fascinating biology of paras engaged in both basic and applied scientific disciplines.

Table 1 Selected Pathogenic* Parasi Parasite Convolvulaceae dodder Cuscuta campestris Is the most widely distributed of about ten species that attack crops. Lauraceae Cassytha filiformis Orobanchaceae Aeginetia indica Alectra vogellii A few other species have been reported as pests of sunflower and tobacco. Christisonia wightii Broomrapes Orobanche ramosa O. crenata O. minor

diverse crops,

Woody plants

sugarcane (Sa

cowpea (Vign

sugarcane (Sa

Various crops

Mainly legum

legumes, toba

crops Rhamphicarpa fistulosa Seymeria cassioides Witchweeds Striga hermonthica S. asiatica S. gesnerioides Loranthaceae Agelanthus spp. Amyema spp. Tapinanthus bangwensis Dendrophthoe spp. Phthirusa spp. Psittacanthus spp. Struthanthus spp. Santalaceae Acanthosyris pauloalvimii Exocarpos spp. Osyris alba Pyrularia pubera Thesium spp. Viscaceae dwarf mistletoes Arceuthobium spp. "Christmas" mistletoes Dendrophthora poeppigii Phoradendron spp. Viscum spp. shea butter or Eucalyptus grains grains legumes

peanut (Arach

pine (Pinus sp

cacao (Theobr

kapok (Ceiba)

rubber (Hevea Citrus

Coffea, Citrus

cacao (Theobr Eucalyptus grape (Vitis)

fir (Abies fras

sugarcane (Sa (Hordeum), et

Pinaceae (New & Cupressace

rubber (Hevea various trees various trees

* Pathogenic defined as negatively impacting a host plant that is cultivated by humans

Scientific Name: Cassytha filiformisCommon Name: Chemar Hantu,Doddler-LaurelClassification: FlowerLocation: TampinesDate Taken: 03/10/2003

Scientific Name: Cassytha filiformisCommon Name: Chemar Hantu,Doddler-LaurelClassification: FlowerLocation: TampinesDate Taken: 03/10/2003