Research ArticleSpecies Composition and Visiting Frequencies ofFlower Visitors of Chromolaena odorata in a Dry Zone ForestPatch of Sri Lanka

D G R M M Kaushalya Rathnayake and W M G Asanga S T B Wijetunga

Department of Biological Sciences Faculty of Applied Sciences Rajarata University of Sri Lanka Mihintale Sri Lanka

Correspondence should be addressed to D G R M M Kaushalya Rathnayake kaushdxyahoocom

Received 18 August 2016 Revised 27 October 2016 Accepted 1 December 2016

Academic Editor G Wilson Fernandes

Copyright copy 2016 D G R M M K Rathnayake and W M G A S T B Wijetunga This is an open access article distributedunder the Creative Commons Attribution License which permits unrestricted use distribution and reproduction in any mediumprovided the original work is properly cited

Plant-animal interaction has been a major theme in ecology as it has helped ecologists to rule out different patterns they observedin the surrounding environments Chromolaena odorata is another plant species that is studied extensively as it has become amajortroublesome weed in many parts of the tropics But handful of studies are available on pollination of this invasive plant species indry forests and its function as a pollinator sink in these environments The current study was carried out in a dry zone secondaryforest patch in North-Central Sri Lanka to assess the diversity abundance and pollination strength of flower visitors associatedwith the C odorata The results suggest that the diversity of Hymenoptera and Lepidoptera is higher than the other insect ordersthat visited C odorata but all species exerted equal pollination strength on the plant The attraction of large numbers of insects isconcluded to as one of the factors that contribute to the reproductive success of C odorata in dry zone forests

1 Introduction

Transfer of pollen grains to the stigma of the flower gynoe-cium (Pollination) is the crucial event in sexual reproductionprocess of flowering plants that ensures their long last survival[1ndash3] Therefore self-incompatible flowering plants in theworld heavily rely on animals for their pollination [1 3 4]Thus plants benefit from these mutualistic interactions toensure effective cross pollination and to maintain viablepopulation in their habitats

Disturbances that alter pollinator communities in habi-tats where self-incompatibale flowering plants are abundantmay reduce the pollination services Additionally loss orlow abundance of pollinating partners will lead ecologicallyspecialized plants to edge of local extinctions These dis-trupting factors can be (1) human alterations to environ-ments reducing pollinator abundance (agriculture pesticideseffect) (2) the effect of invasive ineffective pollinator speciesthat exclude effective native pollinators from floral resources(pollinator competition) and (3) introduced exotic floweringplants with higer attractability than native flowering plants

[5] However most of the pollinator communities consist ofhigher number of invertebrate generalists and few number ofspecialists who visit flower for floral resources [6ndash8] Thustropical pollinator networks have developed an immunityagainst the decline of specialized pollinators but it gives nogurantee that the fidelity between interaction partners will bekept after alterations to the system occur

Importance of pollination has been well documentedBut handful of studies are available which explore insectvisitors in exotic plants while studies on controlling invasiveplants are being largely available According to our under-standing studies on plant-insect mutualism in certain weedsare extremely important as they might reveal several under-lining causes to their inherent invasiveness [9] in the habitatsthey are invading Among different adaptive characteristicsseen in exotic invasive plant species competing with nativespecies for resources available in particular environmentincluding pollinators is commonly observed [10] This maybring a detrimental effect on native existing mutualismsGrowing as considerably large stands and mass floweringwith higher amount of reward for pollinators these exotic

Hindawi Publishing CorporationPsycheVolume 2016 Article ID 8746251 7 pageshttpdxdoiorg10115520168746251

2 Psyche

species attract pollen dependent insects that visit other nativeflowering plants [9]This abundance in exotic floral resourcesis known to alter the spatial distribution of floral resourcesand by that they compete for pollinators with native self-incompatible flowering plants [5] One of the resons thatthe exotic plants gets more dispersed is having high successin pollination and thereby high seed production to extendtheir dispersal range [11] In contrast several authors arguethat abundance of exotic plants with high amount of floralresourses is beneficial to neighboring native plants becausethey increase pollinators in the area by acting as a sink [12]

A native of Central and South America Chromolaenaodorata (L) King and Robinson the Siam weed has spreadthroughout the tropical and subtropical areas of the worldand is now a major weed in Central and Western AfricaSouth and Southeast Asia [13] This neotropical weed isbeing flagged as one of the most troublesome weeds in manyhumid tropical countries [14] Being perennial and having theability to withstand dry climate fires [15 16] and stressedenvironmental conditions [17 18] C odorata has becomemore abundant weed in the above regions while showingpotential extension of its distribution

This study was conducted to assess pollinators that visitwidely dispersed C odorata in tropical secondary dry forestpatch inNorth-Central Sri Lanka Visitation frequencieswereutilized as a surrogate tomeasure the importance of floweringplant to potential insect pollinators as a forage source

2 Materials and Methods

21 Chromolaena odorata C odorata formerly known asEupatorium odoratum L is a weedy pioneering shrub [13 1619] This invasive plant has spread over South Asian regionfrom mid-nineteenth century and currently stated as themost troublesome weed in the region [13 14 19]

C odorata is a scrambling perennial shrub with straightpithy and brittle stems usually grown up to 2-3m in heightBut with support from other vegetation and environmentalconditions it can reach up to 5ndash10m in height [15 19]Capitula are located in panicles at the ends of the branchesand are with only disk florets Each floral head usuallyconsisted of 21 to 28 tiny tubular florets These flower headsare about 10mm long and 3mmwide pale pink or palemauvein color bisexual and actinomorphic The ligule is tubularwith five teeth at the tip Five epipetalous stamens arisefrom the base of the corolla filaments are free but anthersare united syngenesious The anthers are dithecous and thepollen grains are very small round and ornamented Thestyles of florets are stretched beyond anthers and its branchesare receptive to pollen deposited by flower visitors [9] Inthe end of the first growth season the plants undergo sexualreproduction cycle producing approximately 8400 seedsm2[20] and reproduction becomes majorly apomictic in laterperiods of the life cycle Seeds aremainly wind-dispersed andthey also exhibit the tendancy to stick to fur feather andclothes

22 Study Site The study was conducted in Mihintale Sanc-tuary where dense vegetation of C odorata is found The site

is located in the dry zone of Sri Lanka between 8∘ 181015840ndash8∘ 231015840 Nand 80∘ 271015840ndash80∘ 351015840 E and extends over 9996 ha (2470 acres)area Extent of the sanctuary has been defined in the gazettenumber 8370 of Sri Lankan government in 1938 yet no properdemarcated boundaries are documented [21]

Mihintale sanctuary receives an annual rainfall between1000ndash1500mmyear from Northeast monsoon (NovemberndashJanuary) and intermonsoons (March-April September-October) The temperature fluctuates between 19∘C and 38∘Cthroughout the year with mean annual temperature of thestudy area being 273∘C From February to September is thedry period with August being the warmest month of theyear The Southwest monsoon winds blow across Mihintalebetween May and September creating higher evaporationrate

Mihintale sanctuary consists of different kinds of habi-tats dry zone semievergreen undisturbed secondary forestpatches semievergreen disturbed secondary forests scrub-lands aquatic habitats and highly degraded forest patchesCodoratawas observed as dense stands in around these habitatsas well as the open forest margins

23 Data Collection A field survey was conducted to doc-ument potential pollinators in C odorata shrubs in thearea of 05 km2 semievergreen forest patch from the endof August 2014 to January 2015 which coincided with oneflowering season of the weed [9 16 22] Five (5) C odoratainhabiting sites were selected to study flower visitors inMihinthale Sanctuary Vegetation density was calculatedusing the quadrate sampling method Altogether thirty (30)5m times 5m quadrats were layed randomly over stands in fivestudy sites and number of C odorata plants were countedin each quadrat Counts were averaged and the density ofC odorata was calculated Flowering density for 1m2 wascalculated by counting and averaging the number of flowerheads fell within randomly layed another thirty (30) 1mtimes 1mquadrates on C odorata strands [23]

Direct observation method was followed to encounterpotential insect visitors to C odorata stands in sunny dayswith gentle breeze [24 25] Rainy days with heavy windswere deliberately avoided as pollinators are disturbed by theseunsuitable climatic conditions Prior to insect observationsflowers were observed carefully to confirm the readiness offlowers for pollinationMature flowers from standswere sam-pled and observed to confirm that pollination has not beencarried out [4 26] This was easily determined by observingwhether the stamens were disturbed andor pollens wererobbed by flower visitors Fifteen (15) 1m times 1m quadrats wereselected randomly to be observed for insect visits Obser-vations and sampling insects were done from 0700 to 1800for 7 days during the flowering season with the assistanceof two observers Visitation frequencies of each insect werenoted during 30-minute period per hour Insect samplingswere done carefully using an insect net causing minimumas possible disturbance to the visiting insect communitiesTherefore five hours per each day was effectively utilizedfor insect survey Interactions were confirmed if an insecttouches anther or stigma of the flower [27] and visits wererecorded irrespective to previous visits of the insect to the

Psyche 3

same plant 30-minute settling period was given to the flowerstrands after each 30-minute capturing session to reduce themagnitude of disturbance created by the observers whencapturing insects Captured insect assemblages were takento the laboratory to identify using available keys [28] Somespecimens were identified to the genus by referring to theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment of Zoology University of Peradeniya At thelaboratory insects were checked for adhering pollens underthe optical microscope to ensure whether the insects act asa potential pollinator or not The insects that do not carrypollens on their bodies were excluded from the counts Theinsect collections were deposited with the laboratory insectcollection of the Rajarata University of Sri Lanka for furtherstudies

24 Data Analysis The data gathered during the field surveywas tabulated as insects in rows and visitation frequencies incolumns This contingency table was used to calculate dif-ferent indices for describing the topography of the observedflower-insect interaction system

Richness of insect visitor community was calculatedusing an abundance-based coverage estimator and richnessof each insect order was calculated using Margalef rsquos Index(M) Diversity of flower visiting insects was calculated usingMaximum Likelihood Estimator of Shannon Diversity Index[29] Diversity indices were calculated using the programSPADE developed by Chao and Shen [30]

Mean visitation frequencies were calculated for eachinsect observed in all 30 quadratsThe strength of interaction(T) which is indicative of the impact that each insectvisitor exerts on flowers was calculated followingmethods inVazquez et al [31] Interaction strengths were checked for sig-nificant difference among flower visitor families and amongflower visitor orders using a one-way ANOVA with residualanalysis for normality and homoscedasticity of variance Allstatistical analysis on interaction strength and visualizationwas done usingMINITAB version16 (Minitab Inc CoventryUK)

3 Results

Flowering occurred from October to December Mean Codorata vegetation density was 516 plusmn 514 while meanflowering density was 31253plusmn832 across five sites A total of2271 flower visitations were recorded during 35 hours within7 days of observation to C odorata strands Seventy-seven(77) insect flower visitors were recorded belonging to fivemajor insect orders and twenty-five families (Table 1) Speciesrichness of theC odorataflower visitor communitywas 775plusmn080 and species diversity was 4183 plusmn 002

Lepidopteran Hymenopteran and Dipteran diversitywas higher than the diversity of Hemipterans and Coleopter-ans (Table 2) Altogether 32 Lepidopterans belonging to fivefamilies 30 Hymenopterans belonging to eight families 10Dipterans in seven families three Hemipterans in threefamilies and twoColeopterans in two families were identifiedas potential pollinators in the study system

The interaction frequency of the pollinator species ishighly correlated with the species strength (ie impactexerted on plant by its interacting partners) [31] Thusthe most frequent flower visitors were thought to havea potentially high contribution for pollination of flowersand by that increasing their reproductive fitness or havehigher amount of the pollinator reward from the flowerOrders Hymenoptera (0014plusmn0001) Diptera (0013plusmn0001)and Lepidoptera (0012 plusmn 0001) showed higher interactionstrengths respectively while the order Coleoptera (0009 plusmn0001) had the least strength

When comparing interaction strengths of flower visitorsfamily Apidae and Halictidae in the order Hymenopterafamily Syrphidae and Calliphoridae in order Diptera andfamilyNymphalidae and Lycaenidae in the order Lepidopteraexhibited higher values in interaction strength scale How-ever we did not detect any statistically significant differencesbetween means of interaction strengths of flower visitorfamilies as determined by one-way ANOVA [119865 (24 52) =069 119901 ge 0841] These results suggest that each insect visitorhas equal importance in pollinating this invasive weed in dryforests

Though there is no significant difference in pollinatorstrengths among insect visitors top contributors were identi-fied by ranking them in descending order using their impactexerted on the plant Apis cerana Apis dorsata Ceratinabinghami Lasioglossum amblypygus Antepipora sp of orderHymenopteraEpisyrphus nectarinus of the orderDiptera andCepora nerissa Junonia hierta Melanitis leda and Castalliusrosimon of order Lepidoptera exhibited higher strength inpollination

Observers identified two types of insect visitors exhibitingtwo different types of behaviors namely foragers on flowersand hovers All insects who had shown high species strengthover flowers were observed always on flowers They all wereobserved to forage on closely arranged capitula of C odoratapassively loading sticky pollen grains on insects body Thusthey were expected to facilitate passive transport of pollengrains more than sudden visitors who visit flower only forsecond or two

4 Discussion

C odorata is a weedy plant attracting a rich diversity ofinsects Being available during major flowering season of thedry forest as well as the dry seasons it provides an importantsource of pollen nectar and foraging grounds to insects whoshare the same habitat that they grow in In general insectcommunities associated with cosmopolitan invasive weedsdiffer from one habitat to another depending on availabilityof particular species and essentially on availability of othernectar and pollen sources [5]Thus the interactions are prod-uct of relative abundance of plant and animal componentsand environmental alterations But some authors argue thatirrespective of the species living in the habitat exotic plantspecies reconstruct the plant-animal interactions by alteringthe resource availability to pollen or nectar gatherers [5]In that context researchers can deduct that being an exoticplant in dry zone secondary forests C odorata shares the

4 Psyche

Table 1 List of flower visitors in different orders and families observed in C odorata stands in Mihintale Sanctuary and the frequency ofvisits Rates presented are population means per 1 times 1m flower strands per hour (119899 = 15 35 hours)

Order Family Species Frequency SD (plusmn)

Hymenoptera

Apidae

Amegilla comberi 233 153Apis cerana 2267 115Apis dorsata 1900 200Apis florea 1400 265Braunsapis sp 733 416Ceratina binghami 1567 153Thyreus insignis 500 100Trigona iridipennis 900 100Xylocopa ruficornis 1133 058Xylocopa tenuiscapa 400 100

Halictidae

Curvinomia iridiscens 1300 100Hoplonomia westwoodi 1200 173Lasioglossum amblypygus 1833 208Leuconomia sp 1233 153Pachynomia sp 800 100

Megachilidae

Euaspis edentata 1200 100Lithurgus atratus 700 100Megachile lanata 1267 208Megachile vigilans 433 058

Chrysididae Chrysis oculata 433 058Stilbum cyanurum splendidum 1000 100

EumenidaeAntepipona sp 1500 100Delta emarginatum 700 100Subancistrocerus sichelii 200 100

Sphecidae

Ammophila atripes 267 058Sp 1 (H1) 1367 153Sp 2 (H2) 1267 208Sphex sp 700 100

Scelionidae Scolia jurinei Sauss 300 100Vespidae Sp 1 (H8) 367 252

Coleoptera Meloidae Mylabris phalerata 867 058Coccinellidae Coccinellidae 900 400

Diptera

Syrphidae Episyrphus nectarinus 1667 058

Asilidae Cophinopoda chinensis 1367 252Sp 2 (D1) 633 208

Ulidiidae Sp 1 (D2) 533 252Rhagionidae Sp 1 (D3) 433 058Bombyliidae Sp 1 (D4) 800 100

CalliphoridaeSp 1 (D5) 1000 265Sp 2 (D6) 1400 214Sp 3 (D7) 1233 112

Tephritidae Sp 1 (D8) 733 153

HemipteraReduviidae Ectomocoris cordigera 933 058

Pyrrhocoridae Sp 1 (He1) 633 208Scutelleridae Rhynchium brunneum 1200 200

Psyche 5

Table 1 Continued

Order Family Species Frequency SD (plusmn)

Lepidoptera

Papilionidae

Pachliopta hector Crimson Rose 733 153Papilio demoleus Lime butterfly 733 058Papilio polymnestor Blue Mormon 367 208Papilio polytes Common Mormon 1400 436

Pieridae

Appias albina Common Albatros 1233 153Appeals Galen Lesser Albatros 333 153Appias libythea Stripped Albatros 267 208Catopsilia pyrantheMottled Emigrant 833 252Cepora nerissa Common Gull 1733 289Delias eucharis Common Jezebel 467 115Eurema blandaThree-Spot Grass Yellow 1033 321Eurema hecabe Common Grass Yellow 233 153

Nymphalidae

Acraea violae Tawny Coster 800 100Cirrochroa thais Tamil Yeoman 1400 100Danaus genutia Common Tiger 1100 100Euploea core Common Crow 1467 208Junonia hierta Yellow Pansy 1733 115Junonia iphita Chocolate Soldier 1433 153Melanitis leda Common Evening Brown 2100 265Neptis hylas Common Sailer 567 208Parantica aglea Glassy Tiger 567 153Ypthima ceylonicaWhite Four Ring 767 115

Lycaenidae

Castalius rosimon Common Pierrot 1500 361Catochrysops strabo Forget-me-not 300 100Jamides celeno Common Cerulean 833 058Pachliopta hector Crimson Rose 800 436Rapala manea Slate Flash 1200 100

Hesperiidae

Ampittia dioscorides Bush Hopper 500 100Badamia exclamationis Brown Awl 733 306Iambrix salsala Chestnut Bob 833 153Spialia galba Indian Skipper 467 208Telicota colon Pale Palm Dart 533 252

Table 2 Diversity and richness of flower visitor orders in Codorata stands inMihintale Sanctuary (MLEMaximum LikelihoodEstimator of Shannon Weiner Diversity Index M Margalef rsquos Indexfor measuring species richness)

Order Number of species Diversity (MLE) Richness (M)Coleoptera 2 0679 plusmn 0023 0351 plusmn 0025

Diptera 10 2227 plusmn 0012 1963 plusmn 0022

Hemiptera 3 1049 plusmn 0053 0602 plusmn 0004

Hymenoptera 30 3259 plusmn 0049 5113 plusmn 0058

Lepidoptera 32 3267 plusmn 0050 5469 plusmn 0062

same strength to alter topography of native interaction websin its associated habitats Thus the current study bears animportance in determining the relative importance of theplant to the insect community and the strength it exerts onpotential pollinators residing in the habitat Also being aplant with higher number of associated potential pollinators

C odorata exhibits its ability to act as pollinator sink in thegrowing environment

Shihan and Kabir [32] argue that the attraction of theButterflies in higher level may be due to presence of hexosesugars and amino acid rich nectar in the nectaries available inflowersThe flower bears minute amount of nectar more thanmany other flowers used by nectar gatherers But bees tendto forage on C odorata as much as on any other nectar plant[33ndash36]The reasonmay be the availability of large quantity ofpollens and nectar collectively in one flower standThereforethis weedy plant has been used as an important honey plantin commercial apiculture all over the world [33 35 36] Bylooking at our insect visitor checklist compiled for dry zoneforests shown above it is concluded that wild bees are alsoattracted to the plant as much as the domesticated bees suchas Apis cerana Apis dorsata or Trigona iridipennis

In addition to the nutrients and the reward for thepollinators the shape of the flowers was also observed to havewell adapted to facilitate the visitors Floral heads create a fine

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

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2 Psyche

species attract pollen dependent insects that visit other nativeflowering plants [9]This abundance in exotic floral resourcesis known to alter the spatial distribution of floral resourcesand by that they compete for pollinators with native self-incompatible flowering plants [5] One of the resons thatthe exotic plants gets more dispersed is having high successin pollination and thereby high seed production to extendtheir dispersal range [11] In contrast several authors arguethat abundance of exotic plants with high amount of floralresourses is beneficial to neighboring native plants becausethey increase pollinators in the area by acting as a sink [12]

A native of Central and South America Chromolaenaodorata (L) King and Robinson the Siam weed has spreadthroughout the tropical and subtropical areas of the worldand is now a major weed in Central and Western AfricaSouth and Southeast Asia [13] This neotropical weed isbeing flagged as one of the most troublesome weeds in manyhumid tropical countries [14] Being perennial and having theability to withstand dry climate fires [15 16] and stressedenvironmental conditions [17 18] C odorata has becomemore abundant weed in the above regions while showingpotential extension of its distribution

This study was conducted to assess pollinators that visitwidely dispersed C odorata in tropical secondary dry forestpatch inNorth-Central Sri Lanka Visitation frequencieswereutilized as a surrogate tomeasure the importance of floweringplant to potential insect pollinators as a forage source

2 Materials and Methods

21 Chromolaena odorata C odorata formerly known asEupatorium odoratum L is a weedy pioneering shrub [13 1619] This invasive plant has spread over South Asian regionfrom mid-nineteenth century and currently stated as themost troublesome weed in the region [13 14 19]

C odorata is a scrambling perennial shrub with straightpithy and brittle stems usually grown up to 2-3m in heightBut with support from other vegetation and environmentalconditions it can reach up to 5ndash10m in height [15 19]Capitula are located in panicles at the ends of the branchesand are with only disk florets Each floral head usuallyconsisted of 21 to 28 tiny tubular florets These flower headsare about 10mm long and 3mmwide pale pink or palemauvein color bisexual and actinomorphic The ligule is tubularwith five teeth at the tip Five epipetalous stamens arisefrom the base of the corolla filaments are free but anthersare united syngenesious The anthers are dithecous and thepollen grains are very small round and ornamented Thestyles of florets are stretched beyond anthers and its branchesare receptive to pollen deposited by flower visitors [9] Inthe end of the first growth season the plants undergo sexualreproduction cycle producing approximately 8400 seedsm2[20] and reproduction becomes majorly apomictic in laterperiods of the life cycle Seeds aremainly wind-dispersed andthey also exhibit the tendancy to stick to fur feather andclothes

22 Study Site The study was conducted in Mihintale Sanc-tuary where dense vegetation of C odorata is found The site

is located in the dry zone of Sri Lanka between 8∘ 181015840ndash8∘ 231015840 Nand 80∘ 271015840ndash80∘ 351015840 E and extends over 9996 ha (2470 acres)area Extent of the sanctuary has been defined in the gazettenumber 8370 of Sri Lankan government in 1938 yet no properdemarcated boundaries are documented [21]

Mihintale sanctuary receives an annual rainfall between1000ndash1500mmyear from Northeast monsoon (NovemberndashJanuary) and intermonsoons (March-April September-October) The temperature fluctuates between 19∘C and 38∘Cthroughout the year with mean annual temperature of thestudy area being 273∘C From February to September is thedry period with August being the warmest month of theyear The Southwest monsoon winds blow across Mihintalebetween May and September creating higher evaporationrate

Mihintale sanctuary consists of different kinds of habi-tats dry zone semievergreen undisturbed secondary forestpatches semievergreen disturbed secondary forests scrub-lands aquatic habitats and highly degraded forest patchesCodoratawas observed as dense stands in around these habitatsas well as the open forest margins

23 Data Collection A field survey was conducted to doc-ument potential pollinators in C odorata shrubs in thearea of 05 km2 semievergreen forest patch from the endof August 2014 to January 2015 which coincided with oneflowering season of the weed [9 16 22] Five (5) C odoratainhabiting sites were selected to study flower visitors inMihinthale Sanctuary Vegetation density was calculatedusing the quadrate sampling method Altogether thirty (30)5m times 5m quadrats were layed randomly over stands in fivestudy sites and number of C odorata plants were countedin each quadrat Counts were averaged and the density ofC odorata was calculated Flowering density for 1m2 wascalculated by counting and averaging the number of flowerheads fell within randomly layed another thirty (30) 1mtimes 1mquadrates on C odorata strands [23]

Direct observation method was followed to encounterpotential insect visitors to C odorata stands in sunny dayswith gentle breeze [24 25] Rainy days with heavy windswere deliberately avoided as pollinators are disturbed by theseunsuitable climatic conditions Prior to insect observationsflowers were observed carefully to confirm the readiness offlowers for pollinationMature flowers from standswere sam-pled and observed to confirm that pollination has not beencarried out [4 26] This was easily determined by observingwhether the stamens were disturbed andor pollens wererobbed by flower visitors Fifteen (15) 1m times 1m quadrats wereselected randomly to be observed for insect visits Obser-vations and sampling insects were done from 0700 to 1800for 7 days during the flowering season with the assistanceof two observers Visitation frequencies of each insect werenoted during 30-minute period per hour Insect samplingswere done carefully using an insect net causing minimumas possible disturbance to the visiting insect communitiesTherefore five hours per each day was effectively utilizedfor insect survey Interactions were confirmed if an insecttouches anther or stigma of the flower [27] and visits wererecorded irrespective to previous visits of the insect to the

Psyche 3

same plant 30-minute settling period was given to the flowerstrands after each 30-minute capturing session to reduce themagnitude of disturbance created by the observers whencapturing insects Captured insect assemblages were takento the laboratory to identify using available keys [28] Somespecimens were identified to the genus by referring to theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment of Zoology University of Peradeniya At thelaboratory insects were checked for adhering pollens underthe optical microscope to ensure whether the insects act asa potential pollinator or not The insects that do not carrypollens on their bodies were excluded from the counts Theinsect collections were deposited with the laboratory insectcollection of the Rajarata University of Sri Lanka for furtherstudies

24 Data Analysis The data gathered during the field surveywas tabulated as insects in rows and visitation frequencies incolumns This contingency table was used to calculate dif-ferent indices for describing the topography of the observedflower-insect interaction system

Richness of insect visitor community was calculatedusing an abundance-based coverage estimator and richnessof each insect order was calculated using Margalef rsquos Index(M) Diversity of flower visiting insects was calculated usingMaximum Likelihood Estimator of Shannon Diversity Index[29] Diversity indices were calculated using the programSPADE developed by Chao and Shen [30]

Mean visitation frequencies were calculated for eachinsect observed in all 30 quadratsThe strength of interaction(T) which is indicative of the impact that each insectvisitor exerts on flowers was calculated followingmethods inVazquez et al [31] Interaction strengths were checked for sig-nificant difference among flower visitor families and amongflower visitor orders using a one-way ANOVA with residualanalysis for normality and homoscedasticity of variance Allstatistical analysis on interaction strength and visualizationwas done usingMINITAB version16 (Minitab Inc CoventryUK)

3 Results

Flowering occurred from October to December Mean Codorata vegetation density was 516 plusmn 514 while meanflowering density was 31253plusmn832 across five sites A total of2271 flower visitations were recorded during 35 hours within7 days of observation to C odorata strands Seventy-seven(77) insect flower visitors were recorded belonging to fivemajor insect orders and twenty-five families (Table 1) Speciesrichness of theC odorataflower visitor communitywas 775plusmn080 and species diversity was 4183 plusmn 002

Lepidopteran Hymenopteran and Dipteran diversitywas higher than the diversity of Hemipterans and Coleopter-ans (Table 2) Altogether 32 Lepidopterans belonging to fivefamilies 30 Hymenopterans belonging to eight families 10Dipterans in seven families three Hemipterans in threefamilies and twoColeopterans in two families were identifiedas potential pollinators in the study system

The interaction frequency of the pollinator species ishighly correlated with the species strength (ie impactexerted on plant by its interacting partners) [31] Thusthe most frequent flower visitors were thought to havea potentially high contribution for pollination of flowersand by that increasing their reproductive fitness or havehigher amount of the pollinator reward from the flowerOrders Hymenoptera (0014plusmn0001) Diptera (0013plusmn0001)and Lepidoptera (0012 plusmn 0001) showed higher interactionstrengths respectively while the order Coleoptera (0009 plusmn0001) had the least strength

When comparing interaction strengths of flower visitorsfamily Apidae and Halictidae in the order Hymenopterafamily Syrphidae and Calliphoridae in order Diptera andfamilyNymphalidae and Lycaenidae in the order Lepidopteraexhibited higher values in interaction strength scale How-ever we did not detect any statistically significant differencesbetween means of interaction strengths of flower visitorfamilies as determined by one-way ANOVA [119865 (24 52) =069 119901 ge 0841] These results suggest that each insect visitorhas equal importance in pollinating this invasive weed in dryforests

Though there is no significant difference in pollinatorstrengths among insect visitors top contributors were identi-fied by ranking them in descending order using their impactexerted on the plant Apis cerana Apis dorsata Ceratinabinghami Lasioglossum amblypygus Antepipora sp of orderHymenopteraEpisyrphus nectarinus of the orderDiptera andCepora nerissa Junonia hierta Melanitis leda and Castalliusrosimon of order Lepidoptera exhibited higher strength inpollination

Observers identified two types of insect visitors exhibitingtwo different types of behaviors namely foragers on flowersand hovers All insects who had shown high species strengthover flowers were observed always on flowers They all wereobserved to forage on closely arranged capitula of C odoratapassively loading sticky pollen grains on insects body Thusthey were expected to facilitate passive transport of pollengrains more than sudden visitors who visit flower only forsecond or two

4 Discussion

C odorata is a weedy plant attracting a rich diversity ofinsects Being available during major flowering season of thedry forest as well as the dry seasons it provides an importantsource of pollen nectar and foraging grounds to insects whoshare the same habitat that they grow in In general insectcommunities associated with cosmopolitan invasive weedsdiffer from one habitat to another depending on availabilityof particular species and essentially on availability of othernectar and pollen sources [5]Thus the interactions are prod-uct of relative abundance of plant and animal componentsand environmental alterations But some authors argue thatirrespective of the species living in the habitat exotic plantspecies reconstruct the plant-animal interactions by alteringthe resource availability to pollen or nectar gatherers [5]In that context researchers can deduct that being an exoticplant in dry zone secondary forests C odorata shares the

4 Psyche

Table 1 List of flower visitors in different orders and families observed in C odorata stands in Mihintale Sanctuary and the frequency ofvisits Rates presented are population means per 1 times 1m flower strands per hour (119899 = 15 35 hours)

Order Family Species Frequency SD (plusmn)

Hymenoptera

Apidae

Amegilla comberi 233 153Apis cerana 2267 115Apis dorsata 1900 200Apis florea 1400 265Braunsapis sp 733 416Ceratina binghami 1567 153Thyreus insignis 500 100Trigona iridipennis 900 100Xylocopa ruficornis 1133 058Xylocopa tenuiscapa 400 100

Halictidae

Curvinomia iridiscens 1300 100Hoplonomia westwoodi 1200 173Lasioglossum amblypygus 1833 208Leuconomia sp 1233 153Pachynomia sp 800 100

Megachilidae

Euaspis edentata 1200 100Lithurgus atratus 700 100Megachile lanata 1267 208Megachile vigilans 433 058

Chrysididae Chrysis oculata 433 058Stilbum cyanurum splendidum 1000 100

EumenidaeAntepipona sp 1500 100Delta emarginatum 700 100Subancistrocerus sichelii 200 100

Sphecidae

Ammophila atripes 267 058Sp 1 (H1) 1367 153Sp 2 (H2) 1267 208Sphex sp 700 100

Scelionidae Scolia jurinei Sauss 300 100Vespidae Sp 1 (H8) 367 252

Coleoptera Meloidae Mylabris phalerata 867 058Coccinellidae Coccinellidae 900 400

Diptera

Syrphidae Episyrphus nectarinus 1667 058

Asilidae Cophinopoda chinensis 1367 252Sp 2 (D1) 633 208

Ulidiidae Sp 1 (D2) 533 252Rhagionidae Sp 1 (D3) 433 058Bombyliidae Sp 1 (D4) 800 100

CalliphoridaeSp 1 (D5) 1000 265Sp 2 (D6) 1400 214Sp 3 (D7) 1233 112

Tephritidae Sp 1 (D8) 733 153

HemipteraReduviidae Ectomocoris cordigera 933 058

Pyrrhocoridae Sp 1 (He1) 633 208Scutelleridae Rhynchium brunneum 1200 200

Psyche 5

Table 1 Continued

Order Family Species Frequency SD (plusmn)

Lepidoptera

Papilionidae

Pachliopta hector Crimson Rose 733 153Papilio demoleus Lime butterfly 733 058Papilio polymnestor Blue Mormon 367 208Papilio polytes Common Mormon 1400 436

Pieridae

Appias albina Common Albatros 1233 153Appeals Galen Lesser Albatros 333 153Appias libythea Stripped Albatros 267 208Catopsilia pyrantheMottled Emigrant 833 252Cepora nerissa Common Gull 1733 289Delias eucharis Common Jezebel 467 115Eurema blandaThree-Spot Grass Yellow 1033 321Eurema hecabe Common Grass Yellow 233 153

Nymphalidae

Acraea violae Tawny Coster 800 100Cirrochroa thais Tamil Yeoman 1400 100Danaus genutia Common Tiger 1100 100Euploea core Common Crow 1467 208Junonia hierta Yellow Pansy 1733 115Junonia iphita Chocolate Soldier 1433 153Melanitis leda Common Evening Brown 2100 265Neptis hylas Common Sailer 567 208Parantica aglea Glassy Tiger 567 153Ypthima ceylonicaWhite Four Ring 767 115

Lycaenidae

Castalius rosimon Common Pierrot 1500 361Catochrysops strabo Forget-me-not 300 100Jamides celeno Common Cerulean 833 058Pachliopta hector Crimson Rose 800 436Rapala manea Slate Flash 1200 100

Hesperiidae

Ampittia dioscorides Bush Hopper 500 100Badamia exclamationis Brown Awl 733 306Iambrix salsala Chestnut Bob 833 153Spialia galba Indian Skipper 467 208Telicota colon Pale Palm Dart 533 252

Table 2 Diversity and richness of flower visitor orders in Codorata stands inMihintale Sanctuary (MLEMaximum LikelihoodEstimator of Shannon Weiner Diversity Index M Margalef rsquos Indexfor measuring species richness)

Order Number of species Diversity (MLE) Richness (M)Coleoptera 2 0679 plusmn 0023 0351 plusmn 0025

Diptera 10 2227 plusmn 0012 1963 plusmn 0022

Hemiptera 3 1049 plusmn 0053 0602 plusmn 0004

Hymenoptera 30 3259 plusmn 0049 5113 plusmn 0058

Lepidoptera 32 3267 plusmn 0050 5469 plusmn 0062

same strength to alter topography of native interaction websin its associated habitats Thus the current study bears animportance in determining the relative importance of theplant to the insect community and the strength it exerts onpotential pollinators residing in the habitat Also being aplant with higher number of associated potential pollinators

C odorata exhibits its ability to act as pollinator sink in thegrowing environment

Shihan and Kabir [32] argue that the attraction of theButterflies in higher level may be due to presence of hexosesugars and amino acid rich nectar in the nectaries available inflowersThe flower bears minute amount of nectar more thanmany other flowers used by nectar gatherers But bees tendto forage on C odorata as much as on any other nectar plant[33ndash36]The reasonmay be the availability of large quantity ofpollens and nectar collectively in one flower standThereforethis weedy plant has been used as an important honey plantin commercial apiculture all over the world [33 35 36] Bylooking at our insect visitor checklist compiled for dry zoneforests shown above it is concluded that wild bees are alsoattracted to the plant as much as the domesticated bees suchas Apis cerana Apis dorsata or Trigona iridipennis

In addition to the nutrients and the reward for thepollinators the shape of the flowers was also observed to havewell adapted to facilitate the visitors Floral heads create a fine

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

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Signal TransductionJournal of

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BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

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Nucleic AcidsJournal of

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 3

same plant 30-minute settling period was given to the flowerstrands after each 30-minute capturing session to reduce themagnitude of disturbance created by the observers whencapturing insects Captured insect assemblages were takento the laboratory to identify using available keys [28] Somespecimens were identified to the genus by referring to theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment of Zoology University of Peradeniya At thelaboratory insects were checked for adhering pollens underthe optical microscope to ensure whether the insects act asa potential pollinator or not The insects that do not carrypollens on their bodies were excluded from the counts Theinsect collections were deposited with the laboratory insectcollection of the Rajarata University of Sri Lanka for furtherstudies

24 Data Analysis The data gathered during the field surveywas tabulated as insects in rows and visitation frequencies incolumns This contingency table was used to calculate dif-ferent indices for describing the topography of the observedflower-insect interaction system

Richness of insect visitor community was calculatedusing an abundance-based coverage estimator and richnessof each insect order was calculated using Margalef rsquos Index(M) Diversity of flower visiting insects was calculated usingMaximum Likelihood Estimator of Shannon Diversity Index[29] Diversity indices were calculated using the programSPADE developed by Chao and Shen [30]

Mean visitation frequencies were calculated for eachinsect observed in all 30 quadratsThe strength of interaction(T) which is indicative of the impact that each insectvisitor exerts on flowers was calculated followingmethods inVazquez et al [31] Interaction strengths were checked for sig-nificant difference among flower visitor families and amongflower visitor orders using a one-way ANOVA with residualanalysis for normality and homoscedasticity of variance Allstatistical analysis on interaction strength and visualizationwas done usingMINITAB version16 (Minitab Inc CoventryUK)

3 Results

Flowering occurred from October to December Mean Codorata vegetation density was 516 plusmn 514 while meanflowering density was 31253plusmn832 across five sites A total of2271 flower visitations were recorded during 35 hours within7 days of observation to C odorata strands Seventy-seven(77) insect flower visitors were recorded belonging to fivemajor insect orders and twenty-five families (Table 1) Speciesrichness of theC odorataflower visitor communitywas 775plusmn080 and species diversity was 4183 plusmn 002

Lepidopteran Hymenopteran and Dipteran diversitywas higher than the diversity of Hemipterans and Coleopter-ans (Table 2) Altogether 32 Lepidopterans belonging to fivefamilies 30 Hymenopterans belonging to eight families 10Dipterans in seven families three Hemipterans in threefamilies and twoColeopterans in two families were identifiedas potential pollinators in the study system

The interaction frequency of the pollinator species ishighly correlated with the species strength (ie impactexerted on plant by its interacting partners) [31] Thusthe most frequent flower visitors were thought to havea potentially high contribution for pollination of flowersand by that increasing their reproductive fitness or havehigher amount of the pollinator reward from the flowerOrders Hymenoptera (0014plusmn0001) Diptera (0013plusmn0001)and Lepidoptera (0012 plusmn 0001) showed higher interactionstrengths respectively while the order Coleoptera (0009 plusmn0001) had the least strength

When comparing interaction strengths of flower visitorsfamily Apidae and Halictidae in the order Hymenopterafamily Syrphidae and Calliphoridae in order Diptera andfamilyNymphalidae and Lycaenidae in the order Lepidopteraexhibited higher values in interaction strength scale How-ever we did not detect any statistically significant differencesbetween means of interaction strengths of flower visitorfamilies as determined by one-way ANOVA [119865 (24 52) =069 119901 ge 0841] These results suggest that each insect visitorhas equal importance in pollinating this invasive weed in dryforests

Though there is no significant difference in pollinatorstrengths among insect visitors top contributors were identi-fied by ranking them in descending order using their impactexerted on the plant Apis cerana Apis dorsata Ceratinabinghami Lasioglossum amblypygus Antepipora sp of orderHymenopteraEpisyrphus nectarinus of the orderDiptera andCepora nerissa Junonia hierta Melanitis leda and Castalliusrosimon of order Lepidoptera exhibited higher strength inpollination

Observers identified two types of insect visitors exhibitingtwo different types of behaviors namely foragers on flowersand hovers All insects who had shown high species strengthover flowers were observed always on flowers They all wereobserved to forage on closely arranged capitula of C odoratapassively loading sticky pollen grains on insects body Thusthey were expected to facilitate passive transport of pollengrains more than sudden visitors who visit flower only forsecond or two

4 Discussion

C odorata is a weedy plant attracting a rich diversity ofinsects Being available during major flowering season of thedry forest as well as the dry seasons it provides an importantsource of pollen nectar and foraging grounds to insects whoshare the same habitat that they grow in In general insectcommunities associated with cosmopolitan invasive weedsdiffer from one habitat to another depending on availabilityof particular species and essentially on availability of othernectar and pollen sources [5]Thus the interactions are prod-uct of relative abundance of plant and animal componentsand environmental alterations But some authors argue thatirrespective of the species living in the habitat exotic plantspecies reconstruct the plant-animal interactions by alteringthe resource availability to pollen or nectar gatherers [5]In that context researchers can deduct that being an exoticplant in dry zone secondary forests C odorata shares the

4 Psyche

Table 1 List of flower visitors in different orders and families observed in C odorata stands in Mihintale Sanctuary and the frequency ofvisits Rates presented are population means per 1 times 1m flower strands per hour (119899 = 15 35 hours)

Order Family Species Frequency SD (plusmn)

Hymenoptera

Apidae

Amegilla comberi 233 153Apis cerana 2267 115Apis dorsata 1900 200Apis florea 1400 265Braunsapis sp 733 416Ceratina binghami 1567 153Thyreus insignis 500 100Trigona iridipennis 900 100Xylocopa ruficornis 1133 058Xylocopa tenuiscapa 400 100

Halictidae

Curvinomia iridiscens 1300 100Hoplonomia westwoodi 1200 173Lasioglossum amblypygus 1833 208Leuconomia sp 1233 153Pachynomia sp 800 100

Megachilidae

Euaspis edentata 1200 100Lithurgus atratus 700 100Megachile lanata 1267 208Megachile vigilans 433 058

Chrysididae Chrysis oculata 433 058Stilbum cyanurum splendidum 1000 100

EumenidaeAntepipona sp 1500 100Delta emarginatum 700 100Subancistrocerus sichelii 200 100

Sphecidae

Ammophila atripes 267 058Sp 1 (H1) 1367 153Sp 2 (H2) 1267 208Sphex sp 700 100

Scelionidae Scolia jurinei Sauss 300 100Vespidae Sp 1 (H8) 367 252

Coleoptera Meloidae Mylabris phalerata 867 058Coccinellidae Coccinellidae 900 400

Diptera

Syrphidae Episyrphus nectarinus 1667 058

Asilidae Cophinopoda chinensis 1367 252Sp 2 (D1) 633 208

Ulidiidae Sp 1 (D2) 533 252Rhagionidae Sp 1 (D3) 433 058Bombyliidae Sp 1 (D4) 800 100

CalliphoridaeSp 1 (D5) 1000 265Sp 2 (D6) 1400 214Sp 3 (D7) 1233 112

Tephritidae Sp 1 (D8) 733 153

HemipteraReduviidae Ectomocoris cordigera 933 058

Pyrrhocoridae Sp 1 (He1) 633 208Scutelleridae Rhynchium brunneum 1200 200

Psyche 5

Table 1 Continued

Order Family Species Frequency SD (plusmn)

Lepidoptera

Papilionidae

Pachliopta hector Crimson Rose 733 153Papilio demoleus Lime butterfly 733 058Papilio polymnestor Blue Mormon 367 208Papilio polytes Common Mormon 1400 436

Pieridae

Appias albina Common Albatros 1233 153Appeals Galen Lesser Albatros 333 153Appias libythea Stripped Albatros 267 208Catopsilia pyrantheMottled Emigrant 833 252Cepora nerissa Common Gull 1733 289Delias eucharis Common Jezebel 467 115Eurema blandaThree-Spot Grass Yellow 1033 321Eurema hecabe Common Grass Yellow 233 153

Nymphalidae

Acraea violae Tawny Coster 800 100Cirrochroa thais Tamil Yeoman 1400 100Danaus genutia Common Tiger 1100 100Euploea core Common Crow 1467 208Junonia hierta Yellow Pansy 1733 115Junonia iphita Chocolate Soldier 1433 153Melanitis leda Common Evening Brown 2100 265Neptis hylas Common Sailer 567 208Parantica aglea Glassy Tiger 567 153Ypthima ceylonicaWhite Four Ring 767 115

Lycaenidae

Castalius rosimon Common Pierrot 1500 361Catochrysops strabo Forget-me-not 300 100Jamides celeno Common Cerulean 833 058Pachliopta hector Crimson Rose 800 436Rapala manea Slate Flash 1200 100

Hesperiidae

Ampittia dioscorides Bush Hopper 500 100Badamia exclamationis Brown Awl 733 306Iambrix salsala Chestnut Bob 833 153Spialia galba Indian Skipper 467 208Telicota colon Pale Palm Dart 533 252

Table 2 Diversity and richness of flower visitor orders in Codorata stands inMihintale Sanctuary (MLEMaximum LikelihoodEstimator of Shannon Weiner Diversity Index M Margalef rsquos Indexfor measuring species richness)

Order Number of species Diversity (MLE) Richness (M)Coleoptera 2 0679 plusmn 0023 0351 plusmn 0025

Diptera 10 2227 plusmn 0012 1963 plusmn 0022

Hemiptera 3 1049 plusmn 0053 0602 plusmn 0004

Hymenoptera 30 3259 plusmn 0049 5113 plusmn 0058

Lepidoptera 32 3267 plusmn 0050 5469 plusmn 0062

same strength to alter topography of native interaction websin its associated habitats Thus the current study bears animportance in determining the relative importance of theplant to the insect community and the strength it exerts onpotential pollinators residing in the habitat Also being aplant with higher number of associated potential pollinators

C odorata exhibits its ability to act as pollinator sink in thegrowing environment

Shihan and Kabir [32] argue that the attraction of theButterflies in higher level may be due to presence of hexosesugars and amino acid rich nectar in the nectaries available inflowersThe flower bears minute amount of nectar more thanmany other flowers used by nectar gatherers But bees tendto forage on C odorata as much as on any other nectar plant[33ndash36]The reasonmay be the availability of large quantity ofpollens and nectar collectively in one flower standThereforethis weedy plant has been used as an important honey plantin commercial apiculture all over the world [33 35 36] Bylooking at our insect visitor checklist compiled for dry zoneforests shown above it is concluded that wild bees are alsoattracted to the plant as much as the domesticated bees suchas Apis cerana Apis dorsata or Trigona iridipennis

In addition to the nutrients and the reward for thepollinators the shape of the flowers was also observed to havewell adapted to facilitate the visitors Floral heads create a fine

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

4 Psyche

Table 1 List of flower visitors in different orders and families observed in C odorata stands in Mihintale Sanctuary and the frequency ofvisits Rates presented are population means per 1 times 1m flower strands per hour (119899 = 15 35 hours)

Order Family Species Frequency SD (plusmn)

Hymenoptera

Apidae

Amegilla comberi 233 153Apis cerana 2267 115Apis dorsata 1900 200Apis florea 1400 265Braunsapis sp 733 416Ceratina binghami 1567 153Thyreus insignis 500 100Trigona iridipennis 900 100Xylocopa ruficornis 1133 058Xylocopa tenuiscapa 400 100

Halictidae

Curvinomia iridiscens 1300 100Hoplonomia westwoodi 1200 173Lasioglossum amblypygus 1833 208Leuconomia sp 1233 153Pachynomia sp 800 100

Megachilidae

Euaspis edentata 1200 100Lithurgus atratus 700 100Megachile lanata 1267 208Megachile vigilans 433 058

Chrysididae Chrysis oculata 433 058Stilbum cyanurum splendidum 1000 100

EumenidaeAntepipona sp 1500 100Delta emarginatum 700 100Subancistrocerus sichelii 200 100

Sphecidae

Ammophila atripes 267 058Sp 1 (H1) 1367 153Sp 2 (H2) 1267 208Sphex sp 700 100

Scelionidae Scolia jurinei Sauss 300 100Vespidae Sp 1 (H8) 367 252

Coleoptera Meloidae Mylabris phalerata 867 058Coccinellidae Coccinellidae 900 400

Diptera

Syrphidae Episyrphus nectarinus 1667 058

Asilidae Cophinopoda chinensis 1367 252Sp 2 (D1) 633 208

Ulidiidae Sp 1 (D2) 533 252Rhagionidae Sp 1 (D3) 433 058Bombyliidae Sp 1 (D4) 800 100

CalliphoridaeSp 1 (D5) 1000 265Sp 2 (D6) 1400 214Sp 3 (D7) 1233 112

Tephritidae Sp 1 (D8) 733 153

HemipteraReduviidae Ectomocoris cordigera 933 058

Pyrrhocoridae Sp 1 (He1) 633 208Scutelleridae Rhynchium brunneum 1200 200

Psyche 5

Table 1 Continued

Order Family Species Frequency SD (plusmn)

Lepidoptera

Papilionidae

Pachliopta hector Crimson Rose 733 153Papilio demoleus Lime butterfly 733 058Papilio polymnestor Blue Mormon 367 208Papilio polytes Common Mormon 1400 436

Pieridae

Appias albina Common Albatros 1233 153Appeals Galen Lesser Albatros 333 153Appias libythea Stripped Albatros 267 208Catopsilia pyrantheMottled Emigrant 833 252Cepora nerissa Common Gull 1733 289Delias eucharis Common Jezebel 467 115Eurema blandaThree-Spot Grass Yellow 1033 321Eurema hecabe Common Grass Yellow 233 153

Nymphalidae

Acraea violae Tawny Coster 800 100Cirrochroa thais Tamil Yeoman 1400 100Danaus genutia Common Tiger 1100 100Euploea core Common Crow 1467 208Junonia hierta Yellow Pansy 1733 115Junonia iphita Chocolate Soldier 1433 153Melanitis leda Common Evening Brown 2100 265Neptis hylas Common Sailer 567 208Parantica aglea Glassy Tiger 567 153Ypthima ceylonicaWhite Four Ring 767 115

Lycaenidae

Castalius rosimon Common Pierrot 1500 361Catochrysops strabo Forget-me-not 300 100Jamides celeno Common Cerulean 833 058Pachliopta hector Crimson Rose 800 436Rapala manea Slate Flash 1200 100

Hesperiidae

Ampittia dioscorides Bush Hopper 500 100Badamia exclamationis Brown Awl 733 306Iambrix salsala Chestnut Bob 833 153Spialia galba Indian Skipper 467 208Telicota colon Pale Palm Dart 533 252

Table 2 Diversity and richness of flower visitor orders in Codorata stands inMihintale Sanctuary (MLEMaximum LikelihoodEstimator of Shannon Weiner Diversity Index M Margalef rsquos Indexfor measuring species richness)

Order Number of species Diversity (MLE) Richness (M)Coleoptera 2 0679 plusmn 0023 0351 plusmn 0025

Diptera 10 2227 plusmn 0012 1963 plusmn 0022

Hemiptera 3 1049 plusmn 0053 0602 plusmn 0004

Hymenoptera 30 3259 plusmn 0049 5113 plusmn 0058

Lepidoptera 32 3267 plusmn 0050 5469 plusmn 0062

same strength to alter topography of native interaction websin its associated habitats Thus the current study bears animportance in determining the relative importance of theplant to the insect community and the strength it exerts onpotential pollinators residing in the habitat Also being aplant with higher number of associated potential pollinators

C odorata exhibits its ability to act as pollinator sink in thegrowing environment

Shihan and Kabir [32] argue that the attraction of theButterflies in higher level may be due to presence of hexosesugars and amino acid rich nectar in the nectaries available inflowersThe flower bears minute amount of nectar more thanmany other flowers used by nectar gatherers But bees tendto forage on C odorata as much as on any other nectar plant[33ndash36]The reasonmay be the availability of large quantity ofpollens and nectar collectively in one flower standThereforethis weedy plant has been used as an important honey plantin commercial apiculture all over the world [33 35 36] Bylooking at our insect visitor checklist compiled for dry zoneforests shown above it is concluded that wild bees are alsoattracted to the plant as much as the domesticated bees suchas Apis cerana Apis dorsata or Trigona iridipennis

In addition to the nutrients and the reward for thepollinators the shape of the flowers was also observed to havewell adapted to facilitate the visitors Floral heads create a fine

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 5

Table 1 Continued

Order Family Species Frequency SD (plusmn)

Lepidoptera

Papilionidae

Pachliopta hector Crimson Rose 733 153Papilio demoleus Lime butterfly 733 058Papilio polymnestor Blue Mormon 367 208Papilio polytes Common Mormon 1400 436

Pieridae

Appias albina Common Albatros 1233 153Appeals Galen Lesser Albatros 333 153Appias libythea Stripped Albatros 267 208Catopsilia pyrantheMottled Emigrant 833 252Cepora nerissa Common Gull 1733 289Delias eucharis Common Jezebel 467 115Eurema blandaThree-Spot Grass Yellow 1033 321Eurema hecabe Common Grass Yellow 233 153

Nymphalidae

Acraea violae Tawny Coster 800 100Cirrochroa thais Tamil Yeoman 1400 100Danaus genutia Common Tiger 1100 100Euploea core Common Crow 1467 208Junonia hierta Yellow Pansy 1733 115Junonia iphita Chocolate Soldier 1433 153Melanitis leda Common Evening Brown 2100 265Neptis hylas Common Sailer 567 208Parantica aglea Glassy Tiger 567 153Ypthima ceylonicaWhite Four Ring 767 115

Lycaenidae

Castalius rosimon Common Pierrot 1500 361Catochrysops strabo Forget-me-not 300 100Jamides celeno Common Cerulean 833 058Pachliopta hector Crimson Rose 800 436Rapala manea Slate Flash 1200 100

Hesperiidae

Ampittia dioscorides Bush Hopper 500 100Badamia exclamationis Brown Awl 733 306Iambrix salsala Chestnut Bob 833 153Spialia galba Indian Skipper 467 208Telicota colon Pale Palm Dart 533 252

Table 2 Diversity and richness of flower visitor orders in Codorata stands inMihintale Sanctuary (MLEMaximum LikelihoodEstimator of Shannon Weiner Diversity Index M Margalef rsquos Indexfor measuring species richness)

Order Number of species Diversity (MLE) Richness (M)Coleoptera 2 0679 plusmn 0023 0351 plusmn 0025

Diptera 10 2227 plusmn 0012 1963 plusmn 0022

Hemiptera 3 1049 plusmn 0053 0602 plusmn 0004

Hymenoptera 30 3259 plusmn 0049 5113 plusmn 0058

Lepidoptera 32 3267 plusmn 0050 5469 plusmn 0062

same strength to alter topography of native interaction websin its associated habitats Thus the current study bears animportance in determining the relative importance of theplant to the insect community and the strength it exerts onpotential pollinators residing in the habitat Also being aplant with higher number of associated potential pollinators

C odorata exhibits its ability to act as pollinator sink in thegrowing environment

Shihan and Kabir [32] argue that the attraction of theButterflies in higher level may be due to presence of hexosesugars and amino acid rich nectar in the nectaries available inflowersThe flower bears minute amount of nectar more thanmany other flowers used by nectar gatherers But bees tendto forage on C odorata as much as on any other nectar plant[33ndash36]The reasonmay be the availability of large quantity ofpollens and nectar collectively in one flower standThereforethis weedy plant has been used as an important honey plantin commercial apiculture all over the world [33 35 36] Bylooking at our insect visitor checklist compiled for dry zoneforests shown above it is concluded that wild bees are alsoattracted to the plant as much as the domesticated bees suchas Apis cerana Apis dorsata or Trigona iridipennis

In addition to the nutrients and the reward for thepollinators the shape of the flowers was also observed to havewell adapted to facilitate the visitors Floral heads create a fine

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

6 Psyche

landing stage for many flower visitors and closely arrangedflowers in the stands facilitate movement of insects fromflower to flower easily or efficiently

The current study does not suggest very strong interactionbetween flower visitors and the plant C odorata Howeverthe top potential contributors were identified includingbutterflies and adding bees and flies to the global checklistof interacting species with C odorata Since the dry forestsgenerally experience a scarcity of flowering plants during thedry spell and the heavy rains generalist insect foragers mayhave to use whatever floral resources available in the systemto ensure their survival and continuation of generationsTherefore almost equal visitation frequencies and interactionstrengths can occur due to this dependence of the floralvisitors on the available resources Accordingly this plantcan be viewed as a refuge for insects when it is flowering ininvaded habitats where no other plants are available to giveout pollen andor nectar in the habitat

Competing Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Authorsrsquo Contributions

D G R M M Kaushalya Rathnayake and W M G AsangaS T B Wijetunga contributed equally to this work

Acknowledgments

The authors are also grateful to the respective Heads of theDepartment of Biology Faculty of Applied Sciences RajarataUniversity of Sri Lanka Mihintale and the Departmentof Zoology University of Peradeniya for their support inconducting the study and Dr Inoka Karunarathne of theDepartment of Zoology Faculty of Science University ofPeradeniya for the invaluable support given for the iden-tification of pollinators and granting permission to use theInvertebrate Systematic and Diversity Facility (ISDF) of theDepartment Professor Tom Chapman Travis Heckford andMaria Esther Neito Blazquez of the Memorial University ofNewfoundland Canada are highly acknowledge for theirsupport in producing the manuscript

References

[1] D P Abrol Pollination Biology Biodiversity Conservation andAgricultural Production Springer 2011

[2] L Pijl The Principles of Pollination Ecology Pergamon PressOxford UK 1979

[3] L Van der Pijl Principles of Dispersal Springer Berlin Ger-many 1982

[4] K Faegri and L Van der Pijl The Principles of PollinationEcology William Clowes amp Sons Limited London UK 3rdedition 1979

[5] J Ghazoul ldquoAlien abduction disruption of native plant-pollinator interactions by invasive speciesrdquo Biotropica vol 36no 2 pp 156ndash164 2004

[6] MH AllsoppW J De Lange and R Veldtman ldquoValuing insectpollination services with cost of replacementrdquo PLoS ONE vol3 no 9 Article ID e3128 2008

[7] J Bascompte and P Jordano ldquoPlant-animal mutualistic net-works the architecture of biodiversityrdquo Annual Review ofEcology Evolution and Systematics vol 38 pp 567ndash593 2007

[8] J Memmott N M Waser and M V Price ldquoTolerance ofpollination networks to species extinctionsrdquo Proceedings of theRoyal Society of London B Biological Sciences vol 271 no 1557pp 2605ndash2611 2004

[9] P V Lakshmi A J S Raju D J Ram and K V RamanaldquoFloral biology psychophily anemochory and zoochory inChromolaena odorata (L) King and the Robins (Asteraceae)rdquoPakistan Journal of Scientific and Industrial Research vol 54 pp1ndash8 2011

[10] C C Daehler ldquoPerformance comparisons of co-occurringnative and alien invasive plants implications for conservationand restorationrdquo Annual Review of Ecology Evolution andSystematics vol 34 pp 183ndash211 2003

[11] A Traveset and D M Richardson ldquoBiological invasions asdisruptors of plant reproductive mutualismsrdquo Trends in Ecologyamp Evolution vol 21 no 4 pp 208ndash216 2006

[12] E Moragues and A Traveset ldquoEffect of Carpobrotus spp onthe pollination success of native plant species of the BalearicIslandsrdquo Biological Conservation vol 122 no 4 pp 611ndash6192005

[13] L Gautier ldquoTaxonomy and distribution of a tropical weedChromolaena odorata (L) R King amp H Robinsonrdquo Candolleavol 47 pp 645ndash662 1992

[14] A De Rouw ldquoThe invasion of Chromolaena odorata (L) Kingamp Robinson (ex Eupatorium odoratum) and competition withthe native flora in a rain forest zone south-west Cote drsquoIvoirerdquoJournal of Biogeography vol 18 no 1 pp 13ndash23 1991

[15] R C McFadyen and B Skarratt ldquoPotential distribution ofChromolaena odorata (siam weed) in Australia Africa andOceaniardquo Agriculture Ecosystems amp Environment vol 59 no1-2 pp 89ndash96 1996

[16] R Muniappan and M Marutani ldquoEcology and distribution ofchromolaena odorata in Asia and the pacificrdquo in Proceedingsof the 1st International Workshop on Biological Control of Chro-molaena Odorata pp 21ndash24 Agricultural Experiment StationUniversity of Guam Bangkok Thailand 1988

[17] D J Kriticos T Yonow and R E McFadyen ldquoThe potentialdistribution of Chromolaena odorata (siam weed) in relation toclimaterdquoWeed Research vol 45 no 4 pp 246ndash254 2005

[18] S Inderjit Invasive Plants Ecological and Agricultural AspectsSpringer 2006

[19] R Muniappan G Reddy and P-Y Lai ldquoDistribution andbiological control of chromolaena odoratardquo in Invasive PlantsEcological and Agricultural Aspects pp 223ndash233 BirkhauserBasel Switzerland 2005

[20] G A Epp ldquoThe seed bank of Eupatorium odoratum along asuccessional gradient in a tropical rain forest in Ghanardquo Journalof Tropical Ecology vol 3 no 2 pp 139ndash149 1987

[21] S M W Ranwala and P G I Thushari ldquoCurrent statusand management options for invasive plants at the MihintaleWildlife sanctuaryrdquo Journal of the National Science Foundationof Sri Lanka vol 40 no 1 pp 67ndash76 2012

[22] C Zachariades M Day R Muniappan and G Reddy ldquoChro-molaena odorata (L) king and robinson (Asteraceae)rdquo inBiological Control of Tropical Weeds Using Arthropods pp 130ndash160 Cambridge University Press Cambridge Mass USA 2009

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 7

[23] M Stang P G L Klinkhamer and E Van der MeijdenldquoAsymmetric specialization and extinction risk in plant-flowervisitor webs amatter ofmorphology or abundancerdquoOecologiavol 151 no 3 pp 442ndash453 2007

[24] A M Basilio D Medan J P Torretta and N J Bartoloni ldquoAyear-long plant-pollinator networkrdquoAustral Ecology vol 31 no8 pp 975ndash983 2006

[25] K Jędrzejewska-Szmek andMZych ldquoFlower-visitor and pollentransport networks in a large city structure and propertiesrdquoArthropod-Plant Interactions vol 7 no 5 pp 503ndash516 2013

[26] K Faegri and L Van der Pijl Principles of Pollination EcologyElsevier 2013

[27] Y L Dupont and J M Olesen ldquoEcological modules and rolesof species in heathland plant-insect flower visitor networksrdquoJournal of Animal Ecology vol 78 no 2 pp 346ndash353 2009

[28] S G Potts A Dafni and G Nersquoeman ldquoPollination of a coreflowering shrub species in Mediterranean phrygana variationin pollinator diversity abundance and effectiveness in responseto firerdquo Oikos vol 92 no 1 pp 71ndash80 2001

[29] A Chao and S-M Lee ldquoEstimating the number of classes viasample coveragerdquo Journal of theAmerican Statistical Associationvol 87 pp 210ndash217 1992

[30] A Chao and T-J Shen ldquoNonparametric estimation of Shan-nonrsquos index of diversity when there are unseen species insamplerdquo Environmental and Ecological Statistics vol 10 no 4pp 429ndash443 2003

[31] D P Vazquez S B Lomascolo M B Maldonado et al ldquoThestrength of plant-pollinator interactionsrdquo Ecology vol 93 no 4pp 719ndash725 2012

[32] T R Shihan and N Kabir ldquoButterfly diversity in relation toChromolaena odorata (L) King and HE Robins as a nectarplant from two selected regions of Bangladeshrdquo Journal ofEntomology and Zoology Studies vol 3 no 3 pp 258ndash264 2015

[33] P Akratanakul Beekeeping in Asia Food ampAgriculture Organi-zation 1986

[34] P Latham ldquoManuals for bee forage in Tanzania and Congordquo BeeWorld vol 82 no 2 pp 57ndash59 2001

[35] A F Hill ldquoAmerican honey plants together with those whichare of special value to the beekeeper as sources of pollen FrankC Pellettrdquo The Quarterly Review of Biology vol 23 no 4 pp354ndash354 1948

[36] RThapa and SWongsiri ldquoEupatorium odoratum a honey plantfor beekeepers inThailandrdquoBeeWorld vol 78 pp 175ndash178 1997

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology