Transfer of pyrrolizidine and quinolizidine alkaloids toCastilleja (Scrophulariaceae) hemiparasites from composite and legume host plants

Castilleja (Scrophulariaceae) species of the western United States contain pyrrolizidine and quinolizidine alkaloids. TheCastilleja obtain the alkaloids by root parasitism on host plants, withSenecio atratus andS. triangularis (Asteraceae) furnishing the pyrrolizidines, and quinolizidines being obtained fromLupinus species andThermopsis montana (Leguminosae). Individual plants within a givenCastilleja species population may be devoid of alkaloids while others have high alkaloid content. No populations have been found which are made up of both pyrrolizidine- and quinolizidine-containing individuals. These results have important implications forCastilleja ecology, for the study of insect herbivores which areCastilleja specialists, and in the development of systems for the investigation of the role of alkaloids as plant defenses.Paper 10 in the series Chemistry of the Scrophulariaceae. Paper 9: D.R. Gardner, J. Narum, D. Zook, and F.R. Stermitz.J. Nat. Prod. 50: In press.     

A flavonoid glucoside,phellamurin, regulates differential oviposition on a rutaceous plant,Phellodendron amurense,by two sympatric swallowtail butterflies,Papilio protenor andP. xuthus: The front line of a coevolutionary arms race?

We studied the chemical basis for the differential acceptance of a rutaceous plant.Phellodendron amurense, by ovipositing females of two sympatricPapilio butterflies,P. protenor andP. xuthus, that have a very similar host range.P. amurense, on which larvae of both species perform well, was rejected byP. protenor females but was marginally accepted byP. xuthus, even though water-soluble fraction of this plant elicited significant ovipositional responses from the two species. A prenylated dihydroflavonol glucoside, 3,5,7,4-tetrahydroxy-8-(3-methylbut-2-enyl)flavanone-7-O--glucoside (phellamurin), was identified as a dominant active substance that deters oviposition.P. protenor responded highly sensitively to phellamurin, whereasP. xuthus was less susceptible and was affected conspicuously only at higher concentrations than that found in living plants, thereby accounting for their differential responses toP. amurense. InP. protenor, the deterrent effect of phellamurin was considered to outweigh the activity of coexisting stimulant(s) at the concentrations actually present in the foliage, resulting in avoidance of this plant. The phellamurin content in the foliage amounted to as much as 1.8% of the fresh weight, which is far higher than the average levels of flavonoids present in other major host plants ofPapilio. The dose-response experiments demonstrated that unacceptability ofP. amurense forP. protenor was attributable largely to such a high concentration of phellamurin. These results appear to provide information on possible coevolutionary interactions between herbivores and plants and also on the potential for shifts in host affiliation.     

Euphydryas anicia (Lepidoptera: Nymphalidae) utilization of iridoid glycosides fromCastilleja andBesseya (Scrophulariaceae) host plants

Iridoid glycosides were found to be sequestered by natural populations ofEuphydryas anicia after ingestion from the host plantsBesseya alpina, B. plantaginea, andCastilleja integra. Both major iridoids ofB. alpina, cataipol and aucubin, were found in butterfly populations where this was the only host plant. The catalpol-aucubin ratio was higher in the butterflies than in the host plant. AnE. anicia population which uses bothB. plantaginea andC. integra as host plants was found to sequester cataipol as well as another iridoid, macfadienoside. Macfadienoside was the major iridoid ofC. integra, while catalpol esters were the major iridoids ofB. plantaginea. Although it was a major sequestered iridoid, catalpol was a minor constituent in both host plants. The macfadienoside-catalpol ratio in the butterflies from this population was highly variable, and there appeared to be both sex and individual variation in host plant and/or iridoid glucoside utilization byE. anicia. Although other iridoids were present in the host plants, none was sequestered in more than trace amounts.This work was supported by grant CHE-8213714 to FRS, in part by grant DEB-06961 to PRE from the National Science Foundation, and a grant from the Koret Foundation of San Francisco to PRE. Paper 5 in the series Chemistry of the Scrophulariaceae. Paper 4: Roby, M.R. and Stermitz, F.R. 1984.J. Nat. Prod. 47:854–857.     

Ovipositional response of tobacco budworm moths (Lepidoptera: Noctuidae) to cuticular labdanes and sucrose esters from the green leaves ofNicotiana glutinosa L. (Solanaceae)

Field plots of three accessions ofNicotiana glutinosa L. (Nicotiana species accessions 24, 24A, and 24B) at Oxford, North Carolina and Tifton, Georgia were heavily damaged by natural populations of tobacco budworms,Heliothis virescens (F.), during 1985–1989. Experiments in outdoor screen cages demonstrated that all accessions ofN. glutinosa were as prone to oviposition byH. virescens moths as was NC 2326, a commercial cultivar of flue-cured tobacco,N. tabacum L. However, in greenhouse experiments, tobacco budworm larvae did not survive or grow as well when placed on plants ofN. glutinosa as they did when placed on plants of NC 2326. Four labdane diterpenes (manool, 2-hydroxymanool, a mixture of sclareols, and labda-13-ene-8,15-diol [labdenediol]) and two sucrose ester fractions (2,3,4-tri-O-acyl-3-O-acetyl-sucrose [G-SE-I] and 2,3,4,-tri-O-acyl-sucrose [G-SE-II]) were isolated from green leaves of the three accessions ofN. glutinosa. These components were bioassayed for their effects on the ovipositional behavior of tobacco budworm moths using small screen cages in a greenhouse at Oxford, North Carolina. Labdenediol, manool, and both sucrose ester fractions stimulated tobacco budworm moths to oviposit on a tobacco budworm-resistant Tobacco Introduction, TI 1112 (PI 124166), when these materials were sprayed onto a leaf.This article reports the results of research only. Mention of a proprietary product does not constitute endorsement or recommendation for its use by USDA, ARS, or by the North Carolina Agricultural Research Service.     

The “Raison D'être” of pyrrolizidine alkaloids inCynoglossum officinale: Deterrent effects against generalist herbivores

In this study we tested whether pyrrolizidine alkaloids (PAs) ofCynoglossum officinale serve as antifeedants against herbivores. Total PA N-oxide extracts of the leaves significantly deterred feeding by generalist herbivores. Specialist herbivores did not discriminate between food with high and low PA levels. Three PAs fromC. officinale, heliosupine, echinatine, and 3-acetylechinatine, equally deterred feeding by the polyphagous larvae ofSpodoptera exigua. Although the plants mainly contain PAs in their N-oxide form, reduced PAs deterred feeding byS. exigua more efficiently than PA N-oxides. On rosette plants, the monophagous weevilMogulones cruciger significantly consumed more of the youngest leaves, which had the highest PA level and the highest nitrogen percentage. Larvae ofEthmia bipunctella, which are oligophagous within the Boraginaceae, did not discriminate between leaves. All generalist herbivores tested significantly avoided the youngest leaves with the highest PA levels. In the field, the oldest leaves also were relatively more damaged by herbivores than the youngest leaves. It is hypothesized that the skewed distribution of PAs over the leaves of rosette plants reflects optimal defense distribution within the plant.     

Fungal endophyte-infected grasses: Alkaloid accumulation and aphid response

The occurrence of the alkaloidsN-formyl andN-acetyl loline, peramine, lolitrem B, and ergovaline and the response of aphids to plants containing these compounds were determined in species and cultivars ofFestuca,Lolium, and other grass genera infected with fungal endophytes (Acremonium spp., andEpichloe typhina). Twenty-nine of 34 host-fungus associations produced one or more of the alkaloids, most frequently peramine or ergovaline. Three alkaloids (lolines, peramine, and ergovaline) were found in tall fescue and in perennial ryegrass infected withA. coenophialum, while peramine, lolitrem B, and ergovaline were present in perennial ryegrass and in tall fescue infected withA. lolii and inF. longifolia infected withE. typhina. WhileA. coenophialum andA. lolii produced similar patterns of alkaloids regardless of the species or cultivar of grass they infected, isolates ofE. typhina produced either no alkaloids or only one or two different alkaloids in the grasses tested. Aphid bioassays indicated thatRhopalosiphum padi andSchizaphis graminum did not survive on grasses containing loline alkaloids and thatS. graminum did not survive on peramine-containing grasses. Ergovaline-containing grasses did not affect either aphid.     

Influence of cnicin,a sesquiterpene lactone ofCentaurea maculosa (Asteraceae), on specialist and generalist insect herbivores

The sesquiterpene lactone cnicin was extracted fromCentaurea maculosa andCentaurea vallesiaca. We examined its effects on the ovipositional response and larval development of generalist and specialist insect herbivores associated withC. maculosa. For the oviposition trials, three plant species (C. maculosa, Achillea millefolium, andCichorium intybus), half of which were sprayed with 3% of cnicin, were exposed to the specialist mothsStenodes straminea, Agapeta zoegana, andPterolonche inspersa in field cages. All three species significantly preferredC. maculosa to other plants andP. inspersa significantly preferred cnicin-sprayed plants to untreated plants for oviposition. Tested over all species, cnicin significantly increased the number of eggs laid on a given plant. A larval diet test examined the toxicity of cnicin for larvae of the generalist noctuid mothSpodoptera littoralis. Cnicin concentrations of 3% and 6% were lethal and 1% and 0.5% seriously inhibited growth and development. The larvae of theC. maculosa specialistStenodes straminea survived at 6% cnicin, but none of the pupae hatched.Agapeta zoegana was able to survive at 1% and 3% cnicin. Both specialists had difficulties with the artificial diet, but weight increase and survival was not further reduced when cnicin was present compared with on the control diet. In conclusion, cnicin influenced host recognition by the specialist species, and larvae of the generalist did not survive on natural levels of cnicin. Growth and survival of the specialist were not influenced by cnicin but were considerably hampered on artificial diet.     

Olfactory responses of a specialist and a generalist grasshopper to volatiles ofArtemisia ludoviciana nutt. (Asteraceae)

Hypochlora alba is a specialist grasshopper that lives and feeds almost exclusively onArtemisia ludoviciana, a plant that produces large amounts of allelochemics including a variety of monoterpenes. This plant is not a host for generalist grasshoppers such asMelanoplus sanguinipes. The role of olfaction in the grasshopper-plant relationship was investigated by comparing electroantennograms (EAGs) of males and females of both species generated by solvent-extracted volatiles from plant leaves and by major individual components. Volatiles ofA. ludoviciana were identified by gas chromatography-mass spectrometry. The major components were 1,8-cineole, camphor, camphene, and borneol, while minor identified compounds were - and -pinene, -thujene, myrcene,p-cymene, Artemisia ketone, -thujone, and bornyl acetate. The EAGs (mV) ofH. alba males to a range of concentrations of individual volatiles or the total plant extract were nearly double those of conspecific females or both sexes ofM. sanguinipes. However, both sexes ofM. sanguinipes were more sensitive than either sex ofH, alba to geraniol, a monoterpene that commonly occurs in many plant species but is absent or is present in only trace amounts inA. ludoviciana. The increased sensitivity ofH. alba males to the odor components of their host plant appears to be related to the greater number of certain olfactory chemoreceptors on male versus female antennae. The significance of this phenomenon is discussed.     

Iridoid glycoside metabolism and sequestration byPoladryas minuta (Lepidoptera: Nymphalidae) feeding onPenstemon virgatus (Scrophulariaceae)

A bivoltine checkerspot butterfly,Poladryas minuta, is aPenstemon specialist, not known to utilize any other plant genus for oviposition and larval feeding. At several intermontane plains sites of central Colorado, the butterfly utilizesPenstemon virgatus as its sole host plant. Analysis of the host plant showed it to contain three cinnamyl-type catalpol esters (scutellarioside-II, globularin, globularicisin) and catalpol. The host plant contained an average of 10% dry weight iridoids, but some variation among individual plants and leaves within plants was noted. Field-collected butterflies contained 2.1–8.7% dry weight catalpol, but no other iridoids. Adults from larvae fedP. virgatus in the lab contained 4.2–9.0% dry weight catalpol and excreted large amounts of catalpol in the meconium. No catalpol was found in the larval frass. Larvae did not consume three alternate iridoid-containing host-plant species, and most eventually died rather than feed on the alternate plants. Larvae did consume small amounts of artificial diets containing the alternate species andP. virgatus, but most went into diapause and some died. Survival was good on artificial diet containing 10% dry weight of the iridoid esters fromP. virgatus. Only catalpol was found in pupae and adults, but it was absent from the larval frass. The cinnamic-type acids expected from larval hydrolysis of the esters were not found in larval frass, pupae, or adults. These results are contrasted with those found for another checkerspot,Euphydryas anicia, which consumes a different host-plant species but was present at one of the same sites withPoladryas minuta.Paper 15 in the series Chemistry of the Scrophulariaceae. Paper 14 Boros, C.M., Stermitz, F.R., and Harris, G.H. 1990.J. Nat. Prod. 5372–80.     

Importance of quinolizidine alkaloids in the relationship between larvae ofUresiphita reversalis (Lepidoptera: Pyralidae) and a host plant,Genista monspessulana

Larvae ofUresiphita reversalis feed almost exclusively on legumes in the tribe Genisteae, which characteristically contain a variety of quinolizidine alkaloids. The larvae are aposematic, and onGenista monspessulana, a major host in California, they feed on the youngest leaves, at the periphery of the plant. These leaves, which were preferred over older foliage in choice tests, contained four to five times the level of alkaloid found in older leaves. The major alkaloids detected in these plants were dehydroaphylline andN-methylcytisine, together accounting for 74% of the total. Preliminary analyses showed the alkaloid profile of exuviae from larvae feeding on these plants was very similar to that of the plants. Two alkaloids, sparteine and cytisine, which are known components of some hosts ofU. reversalis, were phagostimulants for fifth-instar larvae when added to sucrose-impregnated glass-fiber disks. In addition, when sparteine was added to foliage ofG. monspessulana, effectively doubling the percent dry weight of alkaloid, the growth rate of late-instar larvae was positively affected. Cytisine added to plants had no discernible effect on growth of larvae. Alkaloid levels in larvae and in their frass were proportional to levels in the plants on which they fed. Although the majority of alkaloid was excreted, that which was sequestered by the insect was found entirely in the integument, possibly confering some protection from predators.     

Defensive role ofAllium sulfur compounds for leek mothAcrolepiopsis assectella Z. (Lepidoptera) against generalist predators

It has been shown previously that sulfur volatiles produced byAllium plants affect the behavior of their specialist phytophages and of their specialist entomophages. The action of these compounds in protecting the leek mothAcrolepiopsis assectella against generalist entomophages was studied in comparison to the proposed original defensive role of these compounds against generalist herbivorous insects. Two ants species,Formica selysi andF. fusca, were used as generalist predators. Six behavioral criteria of the predatory behavior of the ants were studied in presence of the last-instar caterpillars (C₅). C₅ reared on artificial diets with or without leek components were tested, as well as C₅ soaked in frass of leek-reared caterpillars or disulfide solutions. In addition, the response of the ants to pure chemicals found in leek was studied using honey solutions with or without sulfur compounds. The sulfur allelochemicals ofAllium plants have a negative action on predatory ants. Interestingly, the nonvolatile precursors of sulfur volatiles ofAllium plants seem to have a protective role for their phytophagous insects against generalist entomophages.     

Sequestration of pyrrolizidine alkaloids in several arctiid moths (Lepidoptera: Arctiidae)

Sequestration of dietary pyrrolizidine alkaloids (PA) by larvae and adults of six European arctiid moth species (Spilosoma lubricipeda, Arctia caja, Phragmatobia fuliginosa, Callimorpha dominula, Diacrisia sannio, andTyria jacobaeae) was investigated for comparison with the well-studied Asian arctiidCreatonotos transiens. Larvae of all species metabolized free PA bases into the respectiveN-oxides. Only adults ofA. caja, P. fuliginosa, andS. lubricipeda, but not their larvae, converted dietary 7(S)-heliotrine to 7(R)-heliotrine, a direct precursor of a male pheromone in some arctiids, 7(R)-hydroxydanaidal. The larval integument figures as the main storage site for resorbed alkaloids; only minor amounts were found in other tissues. In addition, a significant amount of alkaloid is deposited in the cocoon ofArctia caja; only traces of alkaloids could be found in the meconium and the exuviae of this species. A substantial part of the alkaloids fed was degraded to unknown, nonalkaloidal products.     

Sequestration ofVeratrum alkaloids by specialistRhadinoceraea nodicornis konow (Hymenoptera,Tenthredinidae) and its ecoethological implications

The larvae of the specialist sawflyRhadinoceraea nodicornis Konow (Hymenoptera, Tenthredinidae) store in their hemolymph ceveratrum alkaloids originating from the host plantVeratrum album L. (Liliales, Melanthiaceae). The major alkaloid found in the hemolymph is 3-acetyl-zygadenine. Qualitative and quantitative data showed that the plant alkaloid 3-angeloylzygadenine is most probably metabolized in the larval gut to zygadenine and then acetylated. A still unidentified alkaloid with a molecular weight of 591 Da was detected in plant leaves as well as in the gut, hemolymph, and excrement of larvae. Protoveratrine A and B, on the other hand, seem to be degraded by the larvae. These findings indicate that the pathway of ceveratrum alkaloids inR. nodicornis larvae is fourfold: direct sequestration, metabolism followed by sequestration, excretion of intact alkaloids, and degradation. In contrast, no ceveratrum alkaloids were detected in the hemolymph and excrement of larvae of the generalist sawflyAglaostigma sp. fed withV. album leaves. Bioassays with the antMyrmica rubra L. proved that the hemolymph ofR. nodicornis larvae is highly deterrent and toxic. In bioassays evaluating defensive efficiency against predators (ants, spiders, and bushcrickets), no larvae were eaten. Ceveratrum alkaloids were also detected in the hibernating prepupae ofR. nodicornis. In feeding bioassays, the shrewCrocidura russula Hermann rarely fed upon prepupae, suggesting that this stage is also protected from predation to some degree. In field surveys, the only parasitoids recorded were two ichneumonid species that are believed to be specialized onR. nodicornis. Bioassays and field observations enable us to suppose thatR. nodicornis and its enemies produce a food web of ion connectance.     

Mandibular gland secretions of the male beewolvesPhilanthus crabroniformis,P. barbatus,andP. pulcher (Hymenoptera: Sphecidae)

The composition of the territorial marking pheromones from mandibular glands of males of the beewolvesPhilanthus crabroniformis, P. barbatus, andP. pulcher have been determined. The structures of the components were elucidated by gas chromatography-mass spectrometry and gas chromatography-Fourier transform infrared spectroscopy. The major compound ofP. crabroniformis is isopropyl tetradecanoate, with somewhat lesser amounts of 2-tridecanone, 3-methyl-3-butenyl tetradecanoate, and 928 (Z)(E)-11-eicosen-1-ol. The major compounds ofP. barbatus are ethyl tetradecanoate and hexadecanal, which are present in approximately a 6040 ratio. These two compounds comprise over 95% of the neutral lipids. Also present in lesser amounts are ethyl dodecanoate, tetradecanal, hexadecan-1-ol, a ^x -octadecen-1-ol, and octadecan-1-ol. The major compounds ofP. pulcher are ethyl (Z)-7-hexadecenoate and geranylgeraniol acetate, which comprise nearly 90% of the neutral lipid fraction, with smaller amounts of tetradecanal, pentadecanal, and ethyl hexadecanoate; trace amounts of ^x hexadecenal, hexadecanal, and octadecanal are also present.     

Fate of quinolizidine alkaloids through three trophic levels:Laburnum anagyroides (Leguminosae) and associated organisms

The quinolizidine alkaloids (QA) of golden rain,Laburnum anagyroides, and those of phytophagous insects associated with the plant, as well as of parasitoids of the latter, were analyzed by capillary GLC and GLC-MS. The alkaloid content in samples of vegetative plant parts was high at the beginning of the season, then decreased, while that of reproductive organs was high throughout flowering, pod formation, and maturation. The analyses showed that the QA of the plant passed through two higher trophic levels (herbivorous insects and their parasitoids) and that the alkaloid pattern changed little during the passage. The alkaloids were present in two phytophagous insect species associated with golden rain: the predispersal seed predator,Bruchidius villosus [5–13g/g fresh weight (fw)], andAphis cytisorum (182–1012g/g fw), an aphid that feeds on shoots, leaves, and inflorescences. Braconid and chalcidoid parasitoids emerging from the bruchid host also contained alkaloids (1.3–3g/g fw), as did three foraging ant species,Lasius niger, Formica rufibarbis, andF. cunicularia (45g/g fw), that visited the aphid colonies or honeydew-covered leaves of aphid-infested plants. The hypothesis that developing bruchid larvae and/or the plant manipulate QA supply to infested seeds was not supported, because QA content of leftover endosperm in seeds after bruchid development was similar to that of uninfested seeds. The frass of developing bruchid larvae was rich in QA (31g/ g dry weight). While aphids sequestered, the bruchid larvae took up and eliminated QA with the frass without chemical transformation.     

Iridoid glycosides as oviposition stimulants for the buckeye butterfly,Junonia coenia (Nymphalidae)

Females ofJunonia coenia (Nymphalidae), a specialist on plants that contain iridoid glycosides, were found to use aucubin and catalpol, iridoid glycosides typical of a host plant,Plantago lanceolata (Plantaginaceae), as oviposition cues. Incorporating dried ground leaf material or pure iridoid glycosides into agar disks proved to be a very effective method of testing. In no-choice tests and choice tests, females laid more eggs on disks withP. lanceolata leaf material or iridoid glycosides, compared to agar controls. There was variation among individual females in preference for disks withP. lanceolata leaf material versus disks with iridoid glycosides. Females given a choice of three different concentrations of iridoid glycoside (0.2, 0.5, 1.0%) in the agar disks and a control laid more eggs on the disk with the highest concentration of iridoid glycoside.     

Iridoid glycoside sequestration byThessalia leanira (Lepidoptera: Nymphalidae) feeding onCastilleja integra (Scrophulariaceae)

A small population of a polyvoltine checkerspot butterfly,Thessalia leanira fulvia (also known asChlosyne leanira ssp.fulvia), was found to useCastilleja integra as a larval food plant at a localized site (Burnt Mill) southwest of Pueblo, Colorado. Field-captured adult butterflies contained the major iridoid glycosides (catalpol and macfadienoside) of theCastilleja. The content of a third iridoid glycoside, methyl shanzhiside, was also relatively high in the collected butterflies even though most individualCastilleja plants at Burnt Mill contained little or no methyl shanzhiside. Only a few plants, restricted to a small area, did contain appreciable methyl shanzhiside. Most of the plants that lacked the ester methyl shanzhiside contained shanzhiside, the corresponding free carboxylic acid.Thessalia larvae did not normally methylate the acid to produce methyl shanzhiside. Larvae that stopped feeding at an early instar, but yet survived several weeks, did contain major amounts of methyl shanzhiside. It is suggested that only larvae that overwinter or otherwise enter diapause convert shanzhiside to methyl shanzhiside. TheCastilleja food plant also contained iridoids other than catalpol and macfadienoside, sometimes in major amounts, but these were never found in larvae, pupae, or butterflies.Paper 27 in the series Chemistry of the Scrophulariaceae. Paper 26: Stermitz, F.R., Foderaro, T.A., and Li, Y.-X., 1993.Phytochemistry 32:1151     

Transfer of bipiperidyl and quinolizidine alkaloids toViscum cruciatum Sieber (Loranthaceae) hemiparasitic onRetama sphaerocarpa boissier (Leguminosae)

Plant material ofViscum cruciatum Sieber contains bipiperidyl (ammodendrine) and quinolizidine alkaloids (lupanine, 5,6-dehydrolupanine, retamine, cytisine,N-methylcytisine). This plant obtains the alkaloids by root parasitism onRetama sphaerocarpa Boissier (host plant). These results have important implications forViscum ecology, for the study of herbivores that areViscum specialists, and in the development of systems for the investigation of the role of alkaloids as plant defenses.     

Mating and oviposition byCotesia (=Apanteles) marginiventris (Hymenoptera: Braconidae) in presence of synthetic pheromone ofSpodoptera frugiperda (Lepidoptera: Noctuidae)

Two blends of fall armyworm,Spodoptera frugiperda (J.E. Smith), pheromonal components were evaluated in the laboratory for possible close-range effects on mating and oviposition byCotesia marginiventris (Cresson). Neither a two- nor a five-component blend had any significant effect on mating or parasitization ofS. frugiperda larvae by this parasitoid. Dosages of 200, 400, and 800 g were tested.     

Comparative study of allelopathy as exhibited byProsopis Juliflora swartz andProsopis cineraria (L) druce

The allelopathic effects ofProsopis juliflora were studied both in the laboratory and in nature and compared with that ofProsopis cineraria to understand the chemical nature of allelochemics. Both species occupy the same habitats butP. cineraria does not appear to have any toxic effect on other plants under its canopy.P. juliflora is highly allelopathic and does not allow the growth of any other species. Leaf extracts and leaf leachates ofP. juliflora were inhibitory. Decaying leaves were also inhibitory at early stages of decomposition. Live roots were not found to be inhibitory in cogermination and interplanting of seeds. Chemical investigation of the extracts showed the allelopathic compounds to be phenolic in nature in both the species. Slow decomposition and heavy accumulation of leaf litter belowP. juliflora may possibly result in accumulation of toxic substances in soil layers, inhibiting growth of other species.