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Journal of Threatened Taxa Building evidence for conservaton globally www.threatenedtaxa.org ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) Communication Pollination ecology of nodiflora (L.) Gaertn. ()

B. Usharani & A.J. Solomon Raju

26 October 2018 | Vol. 10 | No. 11 | Pages: 12538-12551 10.11609/jot.4008.10.11.12538-12551

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Threatened Taxa Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551

Pollination ecology of Synedrella nodiflora (L.) Gaertn.

Communication (Asteraceae)

ISSN 0974-7907 (Online) 1 2 ISSN 0974-7893 (Print) B. Usharani & A.J. Solomon Raju

1,2 Department of Environmental Sciences, Andhra University, Visakhapatnam, Andhra Pradesh 530003, OPEN ACCESS 1 [email protected], 2 [email protected] (corresponding author)

Abstract: Synedrella nodifora (L.) Gaertn grows almost throughout the year if the soil is damp. It produces heterogamous capitula with female ray forets anthesing on the frst day and bisexual disc forets anthesing on the next three consecutve days. Disc forets are dichogamous, herkogamous, self-compatble, self-pollinatng (vector-mediated) and display secondary pollen presentaton through an intermediate form of brush mechanism. Ray and disc forets exhibit facultatve xenogamy. Buterfies are principal pollinators while bees, wasps and fies are supplementary pollinators. Thrips Microcephalothrips abdominalis uses the forets as breeding and feeding sites; the feeding actvity efects pollinaton. The ray and disc forets produce cypselas; the cypselas produced by ray forets are heavier, elliptcal, membranous with upwardly-pointng teeth along the margins and two short terminal awns while those produced by disc forets are lighter, cylindrical and tangentally compressed with 2 or 3 stf terminal divaricate awns. Seed dispersal is polychorous and represented by anemochory, anthropochory, zoochory and ombrohydrochory. Cypselas of ray forets disperse to short distances and germinate under specifc germinaton conditons either at parental sites or in similar habitats while those of disc forets disperse farther away from parental sites and germinate readily under a wide range of conditons. Therefore, bimorphic cypselas with diferent germinaton abilites enable the to grow as a widespread weed but not as an invasive weed.

Keywords: Bimorphiccypselas, entomophily, polychory, secondary pollen presentaton, Synedrella nodifora, thripsophily.

DOI: htps://doi.org/10.11609/jot.4008.10.11.12538-12551

Editor: Kannan C.S. Warrier, Insttute of Forest Genetcs and Tree Breeding, Coimbatore, India. Date of publicaton: 26 October 2018 (online & print)

Manuscript details: Ms # 4008 | Received 13 January 2018 | Final received 14 September 2018 | Finally accepted 26 September 2018

Citaton: Usharani, B. & A.J.S. Raju (2018). Pollinaton ecology of Synedrella nodifora (L.) Gaertn. (Asteraceae). Journal of Threatened Taxa 10(11): 12538–12551; htps://doi.org/10.11609/jot.4008.10.11.12538-12551

Copyright: © Usharani & Raju 2018. Creatve Commons Atributon 4.0 Internatonal License. JoTT allows unrestricted use of this artcle in any medium, reproducton and distributon by providing adequate credit to the authors and the source of publicaton.

Funding: Self-funded.

Competng interests: The authors declare no competng interests.

Author Details: Ms. B. Usharani is a PhD student in the Department of Environmental Sciences, Andhra University. Recently, she has been awarded PhD under the guidance of Prof. A.J. Solomon Raju. She has published two research papers on pollinaton ecology. Prof. A.J. Solomon Raju is working in the Department of Environmental Sciences, Andhra University, Visakhapatnam. He is the recipient of several natonal and internatonal awards. He has more than 300 research papers in internatonal and natonal Journals. He is on the editorial board of several internatonal journals. He visited USA, Canada, UK, Spain, Brazil, Paraguay, Mexico, Italy, , , Ethiopia, Tanzania, Hong Kong, China and .

Author Contributon: Both the authors contributed equally.

Acknowledgements: We thank the Andhra University, Visakhapatnam, for providing physical facilites to carry out this research work.

12538 Pollinaton ecology of Synedrella nodifora Usharani & Raju

INTRODUCTION MATERIALS AND METHODS

Synedrella is a monotypic with only a single Populatons of Synedrella nodifora growing in , S. nodifora. The generic name is derived from Visakhapatnam region (17.6860N & 83.2180E ) in Andhra the Greek word “synedros” meaning small fowers seated Pradesh State, India were used for the study from together while the species name indicates the presence 2015–2017. Observatons regarding the organizaton of fower clusters around the nodes in the upper parts of inforescences, the spatal positoning of fowers, of the plant (Davidse et al. 2015). It is natve to tropical and their positon on the plant were made since America and from there it has become widespread these features are regarded as important for efectng throughout the warmer regions of the world (Holm et pollinaton by foragers. The life tme of individuals of al. 1997). It is spread throughout the Southeast Asian two foret types was recorded by marking twenty just region, found in the plains of India, in the Andamans open forets each and following them untl fall of. and West . It is also found in Bangladesh, Japan, Anthesis was initally recorded by observing ten marked Spain, China and England (Chauhan & Johnson 2009). mature capitula in the feld. Later, the observatons The plant is used as feed for livestock and in traditonal were repeated fve tmes on diferent days; on each medicine to treat various health problems in Ghana, day we observed twenty marked mature capitula in Nigeria, Malaysia and (Burkill 1985; Idu & order to provide an accurate anthesis schedule. Twenty Onyibe 2007). This plant displays foral features which mature disc forets were followed for recording the are characteristc of thripsophily (Ananthakrishnan tme of anther dehiscence. The presentaton patern of 1993). Further, it is reported that this plant with small pollen was also investgated by recording how anthers and less atractve heterogamous heads is inhabited dehisced and confrmed by observing the anthers under only by one terebrantan thrip, Microcephalotrhips a 10x hand lens. The details of fower morphology such abdominalis which is able to move freely within the as fower sex, shape, size, colour, odour, sepals, petals, capitulum and also fy to other capitula of the same plant stamens and ovary were described. or other nearby . As a result, they efect both self Twenty mature but undehisced anthers from disc and cross-pollinaton (Ananthakrishnan et al. 1981). forets were collected from fve randomly chosen plants Kissmann & Groth (1992) reported that Synedrella and placed in a petri dish. The pollen output per anther/ nodifora produces dimorphic cypselas. Rocha (1996) disc foret and pollen-ovule rato was calculated using stated that the morphological diferences between the protocol given by Cruden (1977). Individual volumes these cypselas infuence dispersion strategies directly. of nectar were recorded for 25 ray and 25 disc forets Bradford (1990) stated that both the cypselas of S. and then the average volume of nectar per each foret nodifora exhibit the same patern of imbibiton rates type was determined and expressed in µl. The capitula indicatng no relatonship to the germinaton rate or used for this purpose were bagged at the mature bud diferences in the fruit coat permeability. Brandel (2004) stage, opened afer anthesis and squeezed the nectar reported that the germinaton responses between from each foret into a micropipete to measure the these cypsela types are not so marked because of a volume of nectar. Based on nectar volume in individual lack of deep innate dormancy. However, they show ray and disc forets, the total volume of nectar secreted some diferences in germinaton rates depending on in a capitulum was calculated. Similarly, the nectar sugar their environment percepton. The state of informaton concentraton at the capitulum level was determined available on pollinaton and seed ecology is insufcient to using a Hand Sugar Refractometer (Erma, Japan). understand the pollinaton and propagaton abilites of S. Nectar analysis for sugar types was done as per the nodifora as a widespread weed across tropical lattudes. Paper Chromatography method described in Dafni et In this context, the present study was contemplated al. (2005). The sugar content/foret is expressed as the to investgate the following objectves: fowering product of nectar volume and sugar concentraton per phenology, foral biology, pollinaton mechanism, unit volume, mg/µl. This was done by frst notng the pollinators, sexual system and seed dispersal ecology. conversion value for the recorded sugar concentraton This informaton presented in this paper enables us to on the refractometer scale and then by multplying it understand the studied aspects of S. nodifora. with the volume of nectar/foret. Table 5.6 given in Dafni et al. (2005) was followed for recording the conversion value to mg of sugars present in one µl of nectar. Dinitrosalicylic acid method was followed for the frst

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12539 Pollinaton ecology of Synedrella nodifora Usharani & Raju two sugar types while Resorcinol method was followed afer they are dispersed or not. Field observatons on for the last sugar type. The caloric reward of nectar/ seed germinaton and seedling formaton were made foret/day was measured as per the formula given in to record the approximate number of generatons Heinrich (1975). He assumed that 1 mg of sugar yields produced during the rainy season. 16.74 joules or 4 calories of energy and accordingly he used the formula for calculatng the caloric reward of the nectar. RESULTS Nectar volume (µl) X Concentraton of nectar (%) ------x 16.74 Phenology 100 The plant is an erect, branched, ephemeral herb Paper chromatography method described in Dafni with long internodes and swollen nodes (Image 1a). et al. (2005) was followed for identfying the amino The lower parts of the stems root at the nodes in damp acid types in the nectar of ray and disc forets; the soil conditons. The shallow tap root system is strongly amino acid types were the same in both ray and disc branched. It grows in humid places, shaded and nutrient forets. The fower visitors were collected and identfed rich soils, crop felds, wastelands, roadsides, lawns and with the representatve specimens available with disturbed areas. The stems are ribbed, dichotomously the Department of Environmental Sciences, Andhra branched, sub-angular, smooth to sparsely hairy. The University, Visakhapatnam. All buterfies were further leaves occur in opposite pairs, ovate to elliptc, fnely confrmed by consultng the books of Kunte (2007) and hairy with short petoles and joined by a ridge across Gunathilagaraj et al. (1998) while other , some the stem. It propagates only by seed. Individual plants to species level while a few others to genus level only. complete their life cycle within 4–5 months. The seeds Thrips were identfed using the key provided by Bhat germinate and produce new plants as soon as they are (1980) for Indian thrips. The insects were observed dispersed from the mother plants and in efect, the carefully for 10 hours a day for 15 days in diferent plants show vegetatve, fowering (Image 1b) and fruitng months each year during the profuse fowering period. phases simultaneously in diferent habitats. However, The hourly foraging visits of each species were recorded the plant shows prolifc growth during the rainy season on ten diferent days for which thirty capitula were and displays peak fowering from October-November. selected. The data obtained was used to calculate the percentage of foraging visits made by each species per Flower morphology day and also to calculate the percentage of foraging visits A plant produces several capitula and their numbers of each category of insects per day. Simultaneously, the vary depending upon the age, number of branches insects were observed for their foraging behavior such and nutrient conditon of the soil. But, it produces an as mode of approach, landing, probing behaviour, the average of 22.6±4.6 capitula. The capitula are sessile type of forage they collected, contact with essental and borne either singly or as groups of 2–8 to nodes in organs to result in pollinaton, and inter-plant foraging leaf axils; they are covered by two green herbaceous actvity. The insects were captured from the capitula involucral bracts. Each capitulum consists of odorless during 10:00–12:00 hr on fve diferent days for pollen outer ray forets and central disc forets and each foret analysis in the laboratory. For each species, 10 type is enclosed by 3–5 mm long erect bracts. Ray specimens were captured and the proboscides were forets vary from 5–8 (7.3±1.31) while disc forets vary separated for examinaton to record whether they from 11–15 (13.06±1.50); they are produced in the rato carried pollen grains or not. They were washed frst in of 1:2. The calyx is reduced to pappus of 2–3 barbed ethyl alcohol and the contents stained with aniline-blue and strong bristles in both types of forets. The corolla on a glass slide and then observed under a microscope is tubate, zygomorphic, yellow and tpped with one to count the number of pollen grains present. ligulate lobe in ray forets and fve actnomorphic lobes A sample of capitula was tagged and followed for in disc forets. The ray foret is 8.1±0.7 mm long and fruit set rate of ray and disc forets in open-pollinatons 2.0±0.3 mm wide while disc foret is 8.2±0.6 mm long and the same sample was used for notng the duraton and 1.2±0.3 mm wide. The ray forets lack stamens but of fruit maturaton. Seed characteristcs of ray and disc have well developed pistl with ovary and style which is forets were carefully examined to note their special terminated into two smooth stylar arms (Image 1h,i). adaptatons for dispersal modes. Field visits were made The disc forets have four stamens with yellow flaments to record whether the seeds germinate immediately and dark brown anthers. The stamens are epipetalous

12540 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 Pollinaton ecology of Synedrella nodifora Usharani & Raju

a b c

d e f g h i

j k l m n o

Image 1. Synedrella nodifora a. Habit, b. Plant in fowering phase, c. Capitulum, d. Anthesing capitulum, e–g. Diferent stages of anthesis in capitulum, h. Ray foret, i. Ray foret with bifd stgma, j. Disc forets, k. Disc foret with unreceptve stgma, l. Disc foret with receptve stgma, m. Syngenecious anthers, n. Relatve positons of stamens and stgma, o. Pollen grain. © A.J. Solomon Raju

and anthers are dithecous, fertle, united and cohered Floral biology forming a hollow cylinder representng syngenesious The ray and disc forets open during early morning conditon (Image 1m). The anther has its connectve from 07:00-09:00 hr on clear sunny days(Image 1c– prolonged into a hood and tend to separate when the g) (Table 1). The forets open completely on sunny fower withers. The pistl is well developed with ovary days while they are partally open on rainy days. In a and long style which is terminated into two linear- capitulum, the ray forets open frst and all of them open lanceolate stylar arms covered abaxially at the base with simultaneously; the disc forets open concentrically sweeping hairs. A nectariferous disc is present at the inwards from the next day for three successive days with base of the style inside the corolla tube. The style with 38% opening on day 1 and 31% each on day 2 and day its aligned arms extend beyond the height of anthers 3 (Table 1). Individual ray and disc forets take about (Image 1j,k,n); the stylar arms diverge and curve inwards three hours to open from mature bud phase. The disc exposing the hidden stgmatc surfaces and fnally over- forets are protandrous with anther dehiscence taking arching the forets (Image 1l). In both foret types, the place during mature bud stage by longitudinal slits. The ovary is bicarpellary, syncarpous and unilocular with narrow anthers are united to form a hollow space into one anatropous ovule on basal placentaton. The foral which pollen is liberated. At the mature bud stage the features indicate that ray forets are female and disc style with its aligned stylar arms lies below the anthers, forets are bisexual. its forked arms are converged at this stage. During and immediately afer anthesis, the style grows, elongates

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12541 Pollinaton ecology of Synedrella nodifora Usharani & Raju

Table 1. Anthesis of disc forets as a functon of tme in Synedrella are present in that order of dominance. The ray and nodifora disc forets wither away on the 3rd day and fall of on the th Time No. of disc forets anthesed 4 day. The tubate corolla in ray forets and the tubate (h) Day 1 % Day 2 % Day 3 % Total corolla and stamens in disc forets gradually fall of

06:00 ------following fertlizaton and initaton of fruit formaton.

07:00 2 15 1 8 1 8 31 Pollinaton mechanism 08:00 2 15 2 15 2 15 45 The ray forets devoid of stamens act as female 09:00 1 8 1 8 1 8 24 and they expose the stgmatc region prominently by First, all ray forets open at once on the frst day of anthesis of capitulum. The next day onwards disc forets open for three consecutve days. Total No. of disc unfolding stylar arms immediately afer anthesis against forets per capitulum 13 the ligulate petal to receive pollen from the foragers on par with disc forets. The disc forets present the stamens and stylar arms at diferent positons. The and passes through the anther tube brushing the anthers dehisce inwardly and discharge pollen grains pollen with its sweeping hairs. At this stage, the inner into the anther tube during mature bud stage. At this stgmatc surfaces are un-receptve and not exposed, the stage, the style lies below the basal part of the anthers. functonal situaton of which prevents the occurrence of During and immediately afer anthesis, the style with autogamy. Such a form of pollen presentaton is referred its aligned stylar arms elongates within the anther tube to as “secondary pollen presentaton mechanism” and brushes the pollen out of the anther tube by stylar which ensures the pollen availability to insects visitng hairs called “sweeping hairs” presentng the pollen the capitula on a daily basis. The style with its aligned only on the abaxial surface at the base of stylar arms branches gradually diverge in the early hours of the (sterile portons). Such a pollen presentaton patern is second day; then the inner stgmatc surfaces atain indicatve of secondary pollen presentaton functonal receptvity and remain so untl the end of that day. The through an intermediate mechanism representatve stylar arms curve downwards completely exposing the of brush mechanism. The stylar arms are in closed receptve stgmatc surfaces. Ray forets display stgma state on day 1 and the inside stgmatc surfaces are un- receptvity by diverging the stylar arms day-long on the receptve, there is no possibility for the occurrence of day of anthesis only. The pollen grains are bright yellow, autogamy. The staminate phase ends at 1700 h and the spheroidal, tricolporate, 35.96±0.04 µm in size and sterile porton of the stylar arms has residual pollen at echinate. The pollen grains are 134.13±21.16 per anther, that tme. This is followed by the retracton of flaments 536.53±84.66 per foret and 6,975 per capitulum (Image and the consequent partal retracton of anthers into 1o). The pollen-ovule rato is 955:1. The ray and disc the corolla tube. The pistllate phase starts in the early forets are nectariferous. A ray or disc foret produces hours of the second day, the stgmatc surfaces atain 1.3μl of nectar which rises up as it accumulates in the receptvity, gradually diverge and curve downwards foret due to the narrow corolla tube. The beginning within three hours totally exposing the inner stgmatc of nectar secreton coincided with anthesis in ray surfaces; the stgmatc surfaces remain receptve untl forets and with anther dehiscence in disc forets; its the end of that day. The brush type pollen presentaton secreton ceased by the evening of day 1 in ray forets mechanism, staminate phase on day 1 and pistllate while it ceased by the evening of day 2 coinciding with phase on day 2 appear to have evolved to prevent cessaton of stgma receptvity. A capitulum produces an autogamy and promote cross-pollinaton. However, average volume of 22.4μl of nectar during its lifespan; the anthesis of disc forets for three consecutve the nectar sugar concentraton is 27±1.6 % with 0.4mg days in the same and diferent capitula on the same of sugar containing 1.6 calories of energy at foret level plant facilitates the occurrence of vector-mediated and 6.67mg sugar containing 26.68 calories of energy self-pollinaton. Therefore, the secondary pollen at capitulum level. The sugar types present in the presentaton mechanism and the sexual system functon nectar include sucrose, glucose and fructose; they are do not insulate completely from the occurrence of self- present in that order of dominance. The nectar contains pollinaton and hence the fowers set fruit and seed fve essental amino acids (arginine, histdine, lysine, through self- as well as cross-pollinaton. tryptophan and threonine) and six non-essental amino acids (cysteine, glycine, serine, aspartc acid, glutamic Thrips breeding, feeding and pollinaton acid, hydroxyproline, alanine, cystne and proline); they Thrips species, Microcephalothrips abdominalis

12542 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 Pollinaton ecology of Synedrella nodifora Usharani & Raju

a b c d e f

g h i j k Image 2. Synedrella nodifora a. Trigona iridipennis collectng nectar, b. Trigona iridipennis collectng pollen, c. Xylocopa latpes collectng nectar, d. Ceratna sp. collectng nectar, e. Ceratna smaragdula collectng nectar, f. Nomia sp. collectng nectar, g. Lasioglossum sp. collectng nectar, h. Rhynchium sp. collectng nectar, i. Ammophila sp. collectng nectar, j. Sarcophaga sp. collectng nectar, k. Eristalinus sp. collectng nectar. © A.J. Solomon Raju.

Table 2. List of insect foragers on Synedrella nodifora

Order Family Genus Species Common name Forage sought

Hymenoptera Apidae Trigona iridipennis Smith Stngless Honey Bee Pollen + Nectar

Ceratna sp. Small Carpenter Bee Pollen + Nectar

Ceratna smaragdula F. Small Carpenter Bee Pollen + Nectar

Anthophoridae Xylocopa latpes L . Large Carpenter Bee Nectar

Halictdae Nomia sp. Alkali Bee Pollen + Nectar

Lasioglossum sp. Sweat Bee Pollen + Nectar

Vespidae Rhynchium sp. Black Poter Wasp Nectar

Sphecidae Ammophila sp. Digger Wasp Nectar

Diptera Sarcophagidae Sarcophaga sp. Flesh Fly Nectar

Syrphidae Eristalinus sp. Hover Fly Nectar

Lepidoptera Pieridae Eurema hecabe L. Common Grass Yellow Nectar

Leptosia nina F. Psyche Nectar

Lycaenidae Zizula hylax F. Tiny Grass Blue Nectar

Pseudozizeeria maha Kollar Pale Grass Blue Nectar

Zizeeria karsandra Moore Dark Grass Blue Nectar

Everes lacturnus Godart Indian Cupid Nectar

Hesperiidae Pelopidas mathias F. Small Branded Swif Nectar

(Thysanoptera: Thripidae) oviposited during early bud nectar, especially from disc forets. They were dusted stage of forets of capitula. The larvae emerge from the with pollen in their upward and downward movements eggs in synchrony with anthesis and nectar producton in within the corolla tube of disc forets. The echinate both ray and disc forets. The centripetal development pollen surface facilitated the thrips to carry 180 to 345 of the capitulum was found to provide contnuous pollen grains on their body setae, wings and legs. The availability of pollen and nectar for three days for their thrips dispersed the pollen on the stgmatc region of the growth. The thrips were found feeding on pollen and stylar arms of both ray and disc forets due to their actve

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12543 Pollinaton ecology of Synedrella nodifora Usharani & Raju movements, rubbing the abdomen on the stgmatc visits (Fig. 3). The body washings of insects collected surface, cleansing of their body parts with their hind legs from the fowers during peak foraging period revealed and also by their wing combing mechanism. Since the that all insects carry pollen but bees carry the highest disc foret is staminate on day 1 and pistllate on day 2, number of pollen grains. Further, the mean number of the foraging actvity of thrips within that foret does not pollen grains varied with each insect species (Table 3). contribute to self-pollinaton but may contribute to self- pollinaton between disc forets of the same or diferent Fruitng ecology and seed dispersal capitula of the same plant. Further, self-pollinaton may The fertlized disc forets produce single-seeded also take place in ray forets due to depositon of pollen cypselas within two weeks. Natural cypsela set is 86% of disc forets of the same capitulum by thrips. As the in ray forets and 98% in disc forets (Table 4). In ray plant occurs as small or large populatons, thrips could forets, the cypsela is 3–5 mm long, 2.5–3.1 mm wide, fy to migrate to the capitula of other closely spaced pale brown, oval, tangentally fatened with upwardly- plants and efect cross-pollinaton. pointng teeth along the marginal wing and 2 short terminal awns (Image 3g). In disc forets, the cypsela is Insect foraging actvity 4–5 mm long, 1.8–2.5 mm wide, cylindrical, thickened, The capitulum is the unit of atracton for insect 4-sided and tangentally compressed with 2 or rarely 3 foragers. Within the capitulum, the yellow ligulate petal terminal stf divaricate awns (Image 3h). The cypselas of ray forets acts as chief atractant. The ray and disc of disc forets detach earlier than those produced from forets were foraged by bees, wasps, fies and buterfies. ray forets. The stf echinate structures of cypselas of The buterfies were the consistent and regular foragers both types of forets enable them to stck readily to hair, while all others were inconsistent foragers and also they fur, clothing and skin for dispersal; the cypselas forage especially during the peak fowering season. The of disc forets being light in weight disperse easily to long bees were Trigona iridipennis (Image 2a,b), Ceratna sp. distances than those of ray forets which are comparably (Image 2d), Ceratna smaragdula (Image 2e), Xylocopa heavy. Wind dispersed the cypselas efciently when latpes (Image 2c), Nomia sp. (Image 2f) and Lasioglossum ambient air is dry. Water also acted as an efcient sp. (Image 2g). The wasps were Rhynchium sp. (Image dispersal agent for the dispersal of cypselas during the 2h) and Ammophila sp.(Image 2i). The fies were Sarcophaga sp. (Image 2j) and Eristalinus sp. (Image 2k). The buterfies included Eurema hecabe (Image 3a), Table 3. Pollen recorded in the body washings of insect foragers on Synedrella nodifora Leptosia nina (Image 3b) (Pieridae), Zizula hylax (Image 3c), (Image 3d), karsandra, Insect species Number of pollen grains Sample Everes lacturnus (Image 3e) () and Pelopidas size Range Mean S.D mathias (Image 3f) (Hesperiidae) (Table 2). Of these, all (N) Trigona iridipennis 10 56-134 106.7 21.08 bees except Xylocopa sp. foraged for both pollen and nectar while Xylocopa sp. and all other foragers foraged Ceratna sp. 10 25-56 37.6 8.54 for nectar only. All these insects approached the fowers Ceratna smaragdula 10 21-63 48.5 11.78 in upright positon, landed on the fat-topped capitulum Xylocopa latpes 10 17-50 29.8 9.47 and then probed ray and disc forets for nectar. They Nomia sp. 10 20-53 35.7 10.13 foraged several forets in a single visit and made Lasioglossum sp. 10 15-49 34.4 8.79 multple visits to several capitula on the same plant in Rhynchium sp. 10 11-42 28.2 8.31 quest of forage. They made frequent visits to capitula Ammophila sp. 10 8-25 15.9 5.27 of diferent closely and distantly spaced plants to collect Sarcophaga sp. 10 10-31 20.5 6.00 forage. Such a foraging behavior was considered to be Eristalinus sp. 10 11-26 18.4 4.08 promotng both self- and cross-pollinaton. The foraging Eurema hecabe 10 21-54 37.9 8.64 actvity patern of insects showed a defnite patern with Leptosia nina 10 8-27 20.2 5.61 reference to foraging schedule. They foraged fowers during 08:00-16:00/17:00 hr with peak foraging during Zizula hylax 10 5 -24 21.7 6.86 10:00-12:00 hr (bees, wasps and fies) and during 10:00– Pseudozizeeria maha 10 11-28 21.5 4.60 11:00 hr (buterfies) coinciding well with the standing Zizeeria karsandra 10 9-36 26.4 7.47 crop of nectar by that tme (Figs. 1,2). Bees made 42%, Everes lacturnus 10 10-32 23.2 5.97 wasps 13%, fies 11% and buterfies 34% of total foraging Pelopidas mathias 10 7-24 15.7 4.37

12544 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 Pollinaton ecology of Synedrella nodifora Usharani & Raju

Figure 1. Hourly foraging actvity of bees, wasps and fies on Synedrella nodifora

Figure 2. Hourly foraging actvity of buterfies on Synedrella nodifora

Table 4. Natural cypsela set in Synedrella nodifora Ray forets Disc forets

No. of forets No. of forets set Cypsela set (%) No. of forets No. of forets set Cypsela set (%) sampled cypsela sampled cypsela 98* 84 86 258** 253 98

*Sample size: 14; **Sample size: 20

rainy season. Therefore, zoochory, anthropochory, anemochory and ombrohydrochory are the functonal forms of seed dispersal in this plant. The cypselas of both ray and disc forets are viable and germinate within a week afer their dispersal from the parental plants if the soil is damp and fertle (Image 3i). Cypselas of both foret types sowed in diferent pots flled with fertle soil produced seedlings within a week; germinaton rate is 60% in ray foret cypselas and 80% in disc foret cypselas. The seedlings have epigeal Figure 3. Percentage of foraging visits of diferent categories of germinaton. The hypocotyl is 10–18 mm long, purplish insects on Synedrella nodifora and hairless. The cotyledons are elliptc, 5–9 mm long, purplish and shortly stalked. The paired juvenile leaves are similar to adult leaves but smaller. Afer two weeks, observatons indicated that both types of cypselas they were transplanted to natural soil and followed produce identcal individuals and produce both types of for their life cycle, which is nearly four months. The cypselas.

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12545 Pollinaton ecology of Synedrella nodifora Usharani & Raju

a b c

d e f

g h i

Image 3. Synedrella nodifora a & b. Pierids - a. Eurema hecabe, b. Leptosia nina, c-e. Lycaenids ; c. Zizula hylax, d. Pseudozizeeria maha, e. Everes lacturnus, f. Hesperiid, Pelopidas mathias, g. Ray foret seeds, h. Disc foret seeds, i. Emergence of seedlings during rainy season. © A.J. Solomon Raju.

DISCUSSION agricultural systems. As a result, it displays vegetatve, fowering and fruitng phases simultaneously in diferent Synedrella nodifora is a herbaceous weed and grows habitats throughout the year. But, the rainy season in soils with sufcient moisture for its rapid germinaton, is the best period for its prolifc growth in any habitat growth, fowering and seed set. It grows in a very wide because the soil is damp and contains nutrients due range of habitats and tolerates most forms of cultvaton to decompositon of organic mater at this tme. Peak because of its short life cycle (Holm et al. 1997). As an fowering occurs during October-November in diferent exotc species, it is widely distributed in both natural populatons. Therefore, it can be said that soil moisture habitats and cultvated felds in India (Ansari et al. and nutrient content greatly infuence the growth and 2016). It is one of the widespread weeds endangering populaton size of this weed in diferent habitats. the natve fora (Singh et al. 2002) and causing a menace Jefrey (2009) stated that Asteraceae members in cultvated felds due to its invasiveness (Singh et al. produce capitula consistng of peripherally located 2010). Despite this situaton created by this weed, there ray forets and centrally located disc forets. The ray are no studies on this species as to how it is able to grow forets are highly specialized in pollinator atracton and invade varied habitats. The present study shows while disc forets assume the reproductve functon and that S. nodifora is a therophyte and produces new sets hence improve the chances for reproductve success of populatons contnuously in damp soils of forest and and facilitate a more fexible basis for breeding system

12546 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 Pollinaton ecology of Synedrella nodifora Usharani & Raju evoluton than does a single fower. Further, this author this study falls in the range of pollen/ovule rato (244.7- stated that a partcularly diverse trait in the capitulum is 2,588) for facultatve xenogamy provided by Cruden the perianth symmetry exhibited by peripheral ray and (1977) and hence this plant has facultatve xenogamous central disc forets; the ray forets display zygomorphic breeding system with low self-incompatbility. In this symmetry while disc forets display actnomorphic breeding system, xenogamy enables plants to increase symmetry. The present study shows that S. nodifora genetc heterogeneity which favors their establishment uses nectar-less ray forets for pollinator atracton and in heterogeneous and variable environment (Hsu 2006) reproductve functon, and nectariferous disc forets while geitonogamy facilitates to increase their populaton exclusively for reproductve functon. The positon of in the currently growing sites or other sites with similar several forets in a capitulum as a unit enables the plant environment. Therefore, the widespread and invasive to maximize reproductve success. Torices et al. (2011) character of S. nodifora could be atributable to low stated that the expression of foral sexuality is associated self-incompatbility and facultatve xenogamy. with changes in symmetry, which has important Floret opening in the morning is a common feature consequences for the evoluton of reproductve biology among the Asteraceae (Proctor & Yeo 1978), and it in Asteraceae. This generalizaton is absolutely true in S. usually occurs before 08:00hr (Mani & Saravanan 1999). nodifora because the peripheral zygomorphic ray forets In S. nodifora also, anthesis of both ray and disc forets are female while the actnomorphic disc forets are occurs in the morning from 07:00–0900 hr. Within a bisexual. Chapman & Abbot (2009) reported that the capitulum, ray forets open frst exposing simultaneously presence of peripheral ray forets shows a marked efect the white ligulate petal and the receptve stgmatc region on atractveness to pollinators, cross-pollinaton rate by unfolding the stylar arms to receive pollen from other and ftness for the plant. In S. nodifora, the highest fruit capitula of the same or diferent plants. These forets set recorded in both ray and disc forets indicate that ray cease stgma receptvity by the end of the same day and forets atract pollinator fauna and enhance both self and become non-functonal to receive pollen from the next cross-pollinaton rate in both types of forets enabling day onwards. The disc forets open concentrically on the plant to increase ftness as an invasive weed. three consecutve days but each foret shows staminate Allen et al. (2011) reported that dichogamous and pistllate phases on two consecutve days to avoid protandry and self-incompatbility are functonal in spontaneous autogamy and minimize geitonogamy. In Asteraceae; self-incompatbility occurs at the stgmatc Asteraceae, secondary pollen presentaton mechanism surface either by failure of germinaton of self-pollen is an important characteristc associated with protandry grains or by the arrest of pollen tube growth. Harder (Howell et al. 1993) but it is an ancestral feature in et al. (2000) noted that dichogamous protandry reduces this family (Jefrey 2009). This pollen presentaton rates of self-fertlizaton and enhances out-crossing. mechanism has been considered to be a strategy to Netancourt (2001) stated that self-incompatbility is improve accuracy in pollen removal and depositon in another evolutonary strategy to avoid self-fertlizaton order to enhance male and female ftness (Ladd 1994). and inbreeding. The present study shows that S. In the present study, the disc forets of S. nodifora nodifora is dichogamous, protandrous, self-compatble display this form of pollen presentaton. In these forets, and self-pollinatng. But, disc forets have a mechanism the pollen shed from the anthers is brushed by the sterile to prevent spontaneous self-pollinaton but not vector- sweeping hairs present at the base of the abaxial side mediated self-pollinaton. In disc forets, the staminate of stylar arms when the style branches are joined and phase on day 1 and pistllate phase on day 2 characterize inner stgmatc surfaces are not receptve. This pollen temporal dioecy. These forets are also herkogamous presentaton mechanism is an intermediate mechanism as they have a physical barrier between the site of representatve of brush mechanism sensu Leins & Erbar pollen depositon and stgmatc region; herkogamy (2006). As the style grows out of the anther tube, the favors xenogamy, and pollinaton occurs during pistllate outer sweeping hairs of the style arms present pollen phase. The occurrence of self-pollinaton, however, for pollinaton. The receptve papillate stgmatc surface is possible due to the centripetal opening of forets is hidden between the two appressed style arms, in the capitulum on consecutve days. In this study, preventng self pollinaton. During the functonally fruit set standing at 86% in ray forets and 98% in disc female phase of the foret which occurs on day 2, the forets in open-pollinatons indicate the functon of style arms separate exposing the receptve papillae for self-compatbility and self-pollinaton in this plant. The the receipt of pollen. The style arms serve as secondary pollen/ovule rato recorded for S. nodifora (955:1) in pollen presenters in the staminate phase and expose

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12547 Pollinaton ecology of Synedrella nodifora Usharani & Raju receptve stgmatc surfaces for pollen during pistllate the body. Further, they use stylar arms for take-of and phase. This type of actve pollen presentaton is typical landing during which the stgmatc area is dusted with of disc forets of Asteraceae (Ladd 1994). The secondary pollen. The feeding actvites of larvae and adult thrips pollen presentaton system functonal in S. nodifora within the capitulum contribute to self-pollinaton. As appears to have evolved to enhance the efciency and there is a contnuous emergence of thrips in synchrony accuracy of pollen exportaton and/or pollen recepton, with sequental anthesis within the capitulum, the thus increasing male and/or female ftness of the plant available forage becomes insufcient to meet their (Yeo 1993). food requirement and in efect they migrate to other Varatharajan & Daniel (1984) reported that thrips have capitula on the same plant or nearby plants in search an intmate associaton with the capitula of Asteraceae. of forage due to which, chances of cross-pollinaton are Laughlin (1977) reported that in Asteraceae, the duraton enhanced. Ananthakrishnan et al. (1981) stated that the of growth and development of thrips synchronizes well thrips living in the heterogamous capitula of Asteraceae with the centripetal disc foral development whereby with solitary inforescences spend more energy for their thrips, which are mostly pollen feeders, efciently use visits to other fowers where the food is plenty. In S. the capitulum for their growth and survival. Kirk (1997) nodifora, the solitary as well as grouped capitula borne noted that thrips puncture the pollen coat and drain the in leaf axils enable thrips to optmize energy expenditure grains; individual thrips consume more than 800 pollen to visit and acquire more energy from the forage they grains per day. Abrol (2012) noted that thrips carry pollen collect from diferent capitula within the plant. Such an ranging from 129–180 pollen grains but the pollen loads interacton between S. nodifora and M. abdominalis carried vary with the larvae and adults with the later benefts both partners, the former in pollinaton and the carrying an increased load due to greater surface area later in breeding and feeding. such as wing fringes, abdominal setae, as well as the In S. nodifora, the peripheral nectar-less yellow ray antenna. Grimaldi & Engel (2005) stated that the higher forets atract foragers. In ray forets, the the stylar arms the pollen producton the greater the level of pollen with exposed receptve stgmatc region standing above carrying capacity. Ananthakrishnan (1982) reported that the tubular porton of the corolla is an adaptaton for the pollen carrying capacity could also depend upon the easy pollen depositon and pollinaton by the foragers pollen surface and architecture. Pollen grains atached which visit the disc forets for pollen and/or nectar. In to the body setae, wings and legs of thrips are dispersed the central yellow disc forets, the narrow tubular corolla on the stgma by way of their actve movement, rubbing containing nectar, pollen accessibility to foragers within the abdomen on the stgmatc surface, cleansing of their and outside the corolla tube, and pollen characteristcs body parts with their hind legs and also by their wing- such as spheroidal shape, tri-colporate apertures and combing mechanism. Kirk (1997) reported that thrips echinate exine are adaptatons for insect-pollinaton. use the stgma for landing and take-of. During this Wodehouse (1935) stated that the echinate trait of the process, thrips place the pollen directly on the stgma. pollen grains is a special adaptaton for adherence to The impressive petal colour and corolla tube of a small insect vectors. Therefore, the characteristcs of both fower form is an ideal site to atract thrips for ovipositon, ray and disc forets suggest that this plant is adapted for enabling the emerging larvae to become dusted with insect-pollinaton. pollen in their upward and downward movement Faegri & Pijl (1979) described the foral characters eventually leading to pollinaton. Ananthakrishnan of buterfy-pollinaton. They usually possess large, et al. (1981) reported that heterogamous capitula of white, pink, red, yellow or blue, narrow, tubular fowers Asteraceae facilitate free movement of both larvae and with deep nectaries and nectar guides. Baker & Baker adults in between the individual forets, and adults carry (1982; 1983) categorized two categories of fowers a maximum pollen load on the body. In the present study, with reference to fower-buterfy relatonships. The Microcephalothrips abdominalis is the only thrips species frst category is “true buterfy fowers” which are which uses S. nodifora capitula for its breeding. The characterized by deep, narrow corolla tubes with larvae emerge in synchrony with the tming of anthesis relatvely copious sucrose-rich nectar. The second of capitulum which occurs for four consecutve days. category is “bee and buterfy fowers” which are The larvae and adults move freely up and down within characterized by short-tubed corolla with hexose- and between ray and disc forets in search of pollen and rich nectar. In S. nodifora, the nectar of disc forets is nectar; the larvae carry less pollen while the adults carry sucrose-rich with moderate sugar concentraton and more pollen because of variaton in the surface area of sugar content, and has some essental and non-essental

12548 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 Pollinaton ecology of Synedrella nodifora Usharani & Raju amino acids. These characteristcs of disc forets All these insects also probe the ray forets for nectar conform to “true buterfy pollinaton syndrome”. The without any discriminaton although there is no nectar; tubular corolla of ray forets lacking nectar and stamens their visits to ray forets result in pollinaton. All these and having functonal pistl indicates that it is adapted insects carry pollen on their body, transfer and efect self- for buterfy-pollinaton by decepton. and cross-pollinaton. Since they show foraging actvity In the present study, buterfies are the consistent mainly during the peak fowering period, they serve as and regular foragers throughout the year for S. supplementary pollinators. Nectar feeding by the larvae nodifora. The capitula atract buterfies belonging to and adults of M. abdominalis leads to reduced levels pieridae, lycaenidae and hesperiidae but lycaenids are of nectar and make all nectar feeding insect species to the principal pollinators. It is interestng to note that increase foraging actvity across populatons to meet papilionid buterfies never visited the capitula. The their nectar requirement and in efect, both self- and small, narrow tubate ray and disc forets aggregated into cross-pollinaton rates are enhanced. capitula are appropriate for pollinaton by buterfies Diferent authors reported that Synedrella nodifora which efciently handle several forets to collect produces two types of fruits (Banerji & Pal 1959; nectar successfully in a single visit. In such an act, Kissmann & Groth 1992; Rocha 1996). The present their proboscis gains contact with the dehisced anthers study also found that S. nodifora produces two types situated inside the corolla tube and also with the pollen of cypselas within a capitulum. The cypselas produced adhered to sweeping hairs during staminate phase in from ray forets are heavier, elliptcal, membranous with disc forets. The proboscis is the carrier of pollen and upwardly-pointng teeth along the margins and two the number of pollen grains carried by them vary with short terminal awns while those from disc forets are the length of proboscis and the tme of nectar collecton. lighter, cylindrical and tangentally compressed with 2 In pistllate phase of disc forets, the buterfies with or 3 stf terminal divaricate awns. Chauhan & Johnson their proboscis contact the exposed stgmatc regions (2009) stated that the heavier cypselas produced from and in efect transfer and deposit pollen at this region ray forets could be related to the inclusion of additonal efectng pollinaton. Further, the buterfies with pollen- carbohydrate resources by the plant to help in the plant laden proboscis pollinate the ray forets while probing emergence while Brandel (2007) atributed the presence their tubular corolla lacking nectar. In both disc and of superior mass in the cypselas produced from ray ray forets, self-pollinaton occurs due to simultaneous forets to the pericarp wing which is developed as an occurrence of staminate and pistllate phases in diferent alternatve structure for dispersion. The present study disc forets, and only pistllate phase in ray forets at shows that the diferences in morphological features capitula and plant level. The standing crop of nectar of these two types of cypselas favour spatal dispersion at plant or populaton level is commensurate with the having diferent strategies which promote spreading in requirement of the buterfies as sufcient volume of various sites. In a capitulum, the cypselas of disc forets nectar is available at capitulum level. The buterfies detach frst and disperse to farther sites while those frequent the capitula of the same and/or diferent of ray forets disperse later in the vicinity of mother closely or distantly spaced individuals and promote plants. The morphological features such as marginal and cross-pollinaton. Their foraging schedule and the peak terminal teeth in ray foret cypselas and terminal teeth foraging actvity period coincide well with the availability in disc foret cypselas enable them to be dispersed by levels of standing crop of nectar. The consistent foraging and humans. Further, wind during dry season actvity of buterfies on this plant suggests that they use and water during rainy season also disperse them this plant as an important nectar source. Therefore, S. farther away. Therefore, the morphological features nodifora is principally psychophilous. of cypselas of S. nodifora are adapted for dispersal In the present study, it is found that bees, wasps and by multple vectors, the situaton of which is referred fies also use the disc forets of S. nodifora as pollen to as “polychory” involving zoochory, anthropochory, and/or nectar sources, especially during peak fowering anemochory and ombrohydrochory. season. The foraging behavior of the bees indicated that The present study shows that the cypselas of they are successful mostly as pollen collectors due to a both ray and disc forets of S. nodifora are viable slight mismatch between the length of their tongue and and germinate within a week afer their dispersal the length of the corolla tube to collect the deeply seated from the mother plants and produce new plants. This nectar. In case of wasps and fies, their proboscis length situaton indicates that both types of cypselas do not easily facilitates nectar collecton from the disc forets. have dormancy. Similarly, Bradford (1990) reported

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12549 Pollinaton ecology of Synedrella nodifora Usharani & Raju that both cypsela types in this plant do not have any aob/mcr147 diference in the fruit coat permeability and hence Ananthakrishnan, T.N. (1982). Thrips and pollinaton biology. Current Science 51: 168–172. germinaton responses between them are almost the Ananthakrishnan, T.N. (1993). Bionomics of thrips. Annual Review same; both cypselas types germinate immediately afer of Entomology 38: 71–92; htps://doi.org/10.1146/annurev. en.38.010193.000443 dispersal. On the contrary, Brandel (2004) reported Ananthakrishnan, T.N., R. Varatharajan & K. Gopinathan (1981). some diferences in germinaton rates of the two cypsela Seasonal periodicity of thrips infestng some Compositae in relaton types in S. nodifora and related these diferences to the to pollinaton. 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The dormant for several months and germinate when Biology of Nectaries. Columbia University Press, New York. favourable conditons exist. The present study indicates Banerji, I. & S. Pal (1959). A contributon to the life history of Synedrella nodifora Gaertn. The Journal of Linnean Society Botany that heterocarpy in S. nodifora is a mixed strategy in 55: 810–817. which disc foret cypselas successfully germinate under Bhat, J.S. (1980). Species of the genus Thrips from India (Thysanoptera). Systematc Entomology 5: 109–166. diferent conditons in tme and space and ray foret Bradford, K.J. (1990). A water relatons analysis of seed germinaton cypselas germinate in restricted environment, usually rates. Plant Physiology 94: 840–849; htps://doi.org/10.1104/ near mother plants or the same habitats. The ability of pp.94.2.840 Brandel, M. (2004). Dormancy and germinaton of heteromorphic disc foret cypselas ofers the chance to colonize new achenes of Bidens frondosa. Flora 199: 228–233. sites, free from sibling competton or other local sources Brandel, M. (2007). Ecology of achene dimorphism in of stress, whereas the remainder of the ofspring stays in Leontodonsaxatlis. Annals of Botany 100: 1189–1197; htps://doi. org/10.1093/aob/mcm214 the same habitat (Imbert & Ronce 2001). Interestngly, Burkill, H.M. (1985). The Useful Plants of West Tropical Africa. Royal both types of cypselas produce identcal plants and also Botanical Gardens, Kew, 981pp. produce bimorphic cypselas within the capitulum during Chapman, M.A. & R.J. Abbot (2009). Introgression of ftness genes across a ploidy barrier. New Phytologist 186: 63–71; htps://doi. fowering period suggestng that they are genotypically org/10.1111/j.1469-8137.2009.03091.x identcal but diferent only by morphology which is Chauhan, B.S. & D.E. Johnson (2009). Seed germinaton and seedling confgured by environmental situatons. The study emergence of Synedrella (Synedrella nodifora) in a tropical environment. 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Threatened Taxa

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12538–12551 12551 OPEN ACCESS The Journal of Threatened Taxa is dedicated to building evidence for conservaton globally by publishing peer-reviewed artcles online every month at a reasonably rapid rate at www.threatenedtaxa.org. All artcles published in JoTT are registered under Creatve Commons Atributon 4.0 Internatonal License unless otherwise mentoned. JoTT allows unrestricted use of artcles in any medium, reproducton, and distributon by providing adequate credit to the authors and the source of publicaton.

ISSN 0974-7907 (Online); ISSN 0974-7893 (Print)

October 2018 | Vol. 10 | No. 11 | Pages: 12443–12618 Date of Publicaton: 26 October 2018 (Online & Print) www.threatenedtaxa.org DOI: 10.11609/jot.2018.10.11.12443-12618

Artcle A winter roost count of the Short-eared Owl Asio fammeus (Aves: Strigiformes: Strigidae) at Porbandar, Gujarat, India The terrestrial life of sea kraits: insights from a long-term study on two -- Dhavalkumar Varagiya & Anita Chakraborty, Pp. 12566–12570 Latcauda species (Reptlia: Squamata: Elapidae) in the Andaman Islands, India -- Zoya Tyabji, Nitya Prakash Mohanty, Erina Young & Tasneem Khan, Pp. 12443– Crocodiles of river Krishna: impact on agriculture, economy, and the sociology of 12450 human populaton in Sangli, Maharashtra, India -- Rajaram Hindurao Atgre, Pp. 12571–12576 Communicatons A new report on the clasper movements of a captve Sand Tiger Shark Fishing Cat Prionailurus viverrinus Bennet, 1833 (Carnivora: Felidae) distributon Carcharias taurus (Lamniformes: Odontaspididae) and a possible reason for and habitat characteristcs in Chitwan Natonal Park, Nepal the behaviour -- Rama Mishra, Khadga Basnet, Rajan Amin & Babu Ram Lamichhane, Pp. 12451– -- Helen Colbachini, Rafael Caprioli Guterrez, Cristane Schilbach Pizzuto & 12458 Oto Bismarck Fazzano Gadig, Pp. 12577–12581

Status distributon and feeding habit of Wild Boar Sus scrofa (Mammalia: New species of termite Pericapritermes travancorensis sp. nov. (Isoptera: Artodactyla: Suidae) in Pench Tiger Reserve, Madhya Pradesh, India Termitdae: Termitnae) from India -- Shaheer Khan & Orus Ilyas, Pp. 12459–12463 -- Jobin Mathew & Chinnu Ipe, Pp. 12582–12588

The compositon and status of waterbirds of Perur Lake in Tamil Nadu, India A checklist of vascular epiphytes of El Cometa Lagoon, Pantanos de Centla -- G. Parameswaran & R. Sivashankar, Pp. 12464–12488 Biosphere Reserve, Mexico -- Derio Antonio Jiménez-López, Candelario Peralta-Carreta, Jonathan V. Solórzano, The herpetofauna of Jigme Singye Wangchuck Natonal Park in central Bhutan: Gerardo Luis Cervantes-Jiménez & Marco Antonio Domínguez-Vázquez, status, distributon and new records Pp. 12589–12597 -- Sangay Tshewang & Letro Letro, Pp. 12489–12498 Notes The Odonata (Insecta) of Dhofar, southern -- Elaine M. Cowan & Peter J. Cowan, Pp. 12499–12514 Two moth species of Lasiocampidae (: Lasiocampoidea) recorded for the frst tme in Bhutan New kissing bug (Hemiptera: Reduviidae: Triatominae) records from Napo and Jatshwor Singh Irungbam­ & Meenakshi Jatshwor Irungbam, Pp. 12598–12601 Morona-Santago provinces with distributon updates in Ecuador -- Ana Soto-Vivas, Sandra Enríquez, Ernesto Villacrés, Jazzmin Arrivillaga, New nymphalid buterfy records from Jammu & Kashmir, India Martn Hinojosa & Jonathan Liria, Pp. 12515–12522 -- Shakha Sharma & Neeraj Sharma, Pp. 12602–12606

Orchid diversity in two community forests of Makawanpur District, central Nepal Argostemma khasianum C.B. Clarke (Rubiaceae): a new record of a genus and -- Bijaya Pant, Mukt Ram Paudel, Mukesh Babu Chand, Shreet Pradhan, species of fowering plant for the state of Arunachal Pradesh (India) and its Bijaya Bahadur Malla & Bhakta Bahadur Raskot, Pp. 12523–12530 lectotypifcaton -- Umeshkumar L. Tiwari & V.K. Rawat, Pp. 12607–12609 Habitat distributon modeling for reintroducton and conservaton of Aristolochia indica L. - a threatened medicinal plant in Assam, India Amorphophallus longiconnectvus and A. margaritfer: additonal aroids from -- Bhaskar Sarma, Prantk Sharma Baruah & Bhaben Tant, Pp. 12531–12537 Maharashtra with notes on the foral variatons -- Avinash R. Gholave, Ravikiran S. Govekar, Vasanta I. Kahalkar, Milind M. Sardesai Pollinaton ecology of Synedrella nodifora (L.) Gaertn. (Asteraceae) & Shrirang R. Yadav, Pp. 12610–12613 -- B. Usharani & A.J. Solomon Raju, Pp. 12538–12551 Cultvaton of the Himalayan seasoning Allium in a remote village of Review Utarakhand, India -- Chandra Prakash Kuniyal & Bir Singh Negi, Pp. 12614–12617 Status of studies on zooplankton fauna of Arunachal Pradesh, India -- Bikramjit Sinha, Pp. 12552–12560 Miscellaneous

Short Communicatons Natonal Biodiversity Authority

First record of the endangered Arabian Tahr Arabitragus jayakari (Thomas, 1894) in the Hata Mountain Conservaton Area, Dubai, Member -- Jeruel Cabadonga Aguhob, Junid N. Shah, Esmat Elfaki Mohammed Elhassan, Aisha Almurr Al Muhery, Mohamed Mustafa Eltayeb Mohamed, Juma Abdulla Saeed Mohammad Al Omairi, Hamad Hashim Mohammed Khalaf Albedwawi, Obaid Mohammed Salem Mohammed Al Bedwawi, Hassan Zain Alsharif & Afra Mahmood Mohammad Ali Haji, Pp. 12561–12565

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