OPEN ACCESS The Journaf of Threafened Taxa fs dedfcafed fo buffdfng evfdence for conservafon gfobaffy by pubffshfng peer-revfewed arfcfes onffne every monfh af a reasonabfy rapfd rafe af www.fhreafenedfaxa.org . Aff arfcfes pubffshed fn JoTT are regfsfered under Creafve Commons Afrfbufon 4.0 Infernafonaf Lfcense unfess ofherwfse menfoned. JoTT affows unresfrfcfed use of arfcfes fn any medfum, reproducfon, and dfsfrfbufon by provfdfng adequafe credff fo fhe aufhors and fhe source of pubffcafon.

Journaf of Threafened Taxa Buffdfng evfdence for conservafon gfobaffy www.fhreafenedfaxa.org ISSN 0974-7907 (Onffne) | ISSN 0974-7893 (Prfnf) Arffcfe Pofffnaffon ecofogy of Rhynchosfa cana (Wfffd.) DC. (), an erecf sub-shrub, fn penfnsufar Indfa

A.J. Sofomon Raju & K. Venkafa Ramana

26 Ocfober 2017 | Vof. 9| No. 10 | Pp. 10757–10770 10.11609/jof. 3172 .9. 10. 10757–10770

For Focus, Scope, Afms, Poffcfes and Gufdeffnes vfsff hfp://fhreafenedfaxa.org/Abouf_JoTT For Arfcfe Submfssfon Gufdeffnes vfsff hfp://fhreafenedfaxa.org/Submfssfon_Gufdeffnes For Poffcfes agafnsf Scfenffc Mfsconducf vfsff hfp://fhreafenedfaxa.org/JoTT_Poffcy_agafnsf_Scfenffc_Mfsconducf For reprfnfs confacf

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Threafened Taxa

Journaf of Threafened Taxa | www.fhreafenedfaxa.org | 26 Ocfober 2017 | 9(10): 10757–10770 ARTICLE Pofffnaffon ecofogy of Rhynchosfa cana (Wfffd.) DC. (Fabaceae), an erecf sub-shrub, fn penfnsufar Indfa

A.J. Sofomon Raju 1 & K. Venkafa Ramana 2 ISSN 0974-7907 (Onffne) ISSN 0974-7893 (Prfnf) 1 Deparfmenf of Envfronmenfaf Scfences; 2 Deparfmenf of Bofany, Andhra Unfversffy, Waffafr Juncfon, Vfsakhapafnam, Andhra Pradesh 530003, Indfa OPEN ACCESS 1 [email protected] (correspondfng aufhor); 2 [email protected]

Absfracf: Rhynchosfa cana fs a perennfaf erecf sub-shrub. If fowers durfng November-January wffh peak fowerfng fn December. The fowers are hermaphrodffc, necfarfferous, seff-compafbfe and dfspfay expfosfve pofffnafon mechanfsm adapfed for pofffnafon by bees. They do nof fruff fhrough aufonomous seffng, buf rafher fhrough manfpufafed seffng, geffonogamy and xenogamy medfafed prfncfpaffy by bees and occasfonaffy by fycaenfd buferffes. In fhe focafffes of fhfs herb, weeds such as Hypfs suaveofens (Lamfaceae) and Chromofaena odorafa (Asferaceae) show fuxurfanf growfh and fower sfmuffaneousfy, and fhefr fnfense fowerfng and hfgh fower densffy ensures fhaf mosf fower-vfsffng fnsecfs vfsff fhefr fowers. In fhfs sffuafon onfy fwo bee specfes, Nomfa and Anfhfdfum exhfbffed fdefffy fo R. cana fowers. Un-frfpped fowers faff of whffe frfpped ones sef fruff. In open-pofffnafons, fruff sef was 81% and seed sef was 54%. Seed dfspersaf occurs by expfosfve pod dehfscence. Perennfaf roof sfock resurrecfs back fo fffe and resfarfs ffs reproducfve cycfe durfng fhe rafny season. Seeds afso germfnafe af fhe same fme buf fhefr confnued growfh fs subjecf fo fhe avaffabffffy of soff mofsfure confenf. Thfs sfudy suggesfs fhaf R. cana fs unabfe fo compefe wffh fhe co-fowerfng weed specfes for pofffnafors, and afso has regenerafon consfrafnfs due fo nufrfenf-defcfenf rocky habffafs wffh profffc growfh of weeds.

Keywords: Enfomophffy, expfosfve pofffnafon mechanfsm, expfosfve seed dfspersaf, hermaphrodffsm, Rhynchosfa cana .

DOI : hfp://dof.org/10.11609/jof.3 172 .9. 10 .10757- 10770

Edffor: Cfeofas Cervancfa , Unfversffy of Phfffppfnes Los Baños Coffege, Laguna, Phfffppfnes. Dafe of pubffcafon: 26 Ocfober 2017 (onffne & prfnf)

Manuscrfpf defaffs: Ms # 3172 | Recefved 27 November 2016 | Ffnaf recefved 01 Augusf 2017 | Ffnaffy accepfed 18 Sepfember 2017

Cffafon: Raju, A.J.S. & K.V. Ramana (2017). Pofffnafon ecofogy of Rhynchosfa cana (Wfffd.) DC. (Fabaceae), an erecf sub-shrub, fn penfnsufar Indfa . Journaf of Threafened Taxa 9(10): 10757–10770 ; hfp://dof.org/10.11609/jof.3172.9.10. 10757-10770

Copyrfghf: © Raju & Ramana 2017 . Creafve Commons Afrfbufon 4.0 Infernafonaf Lfcense. JoTT affows unresfrfcfed use of fhfs arfcfe fn any medfum, reproduc - fon and dfsfrfbufon by provfdfng adequafe credff fo fhe aufhors and fhe source of pubffcafon.

Fundfng: The work reporfed was carrfed ouf from seff-fundfng sources.

Compefng fnferesfs: The aufhors decfare no compefng fnferesfs.

Aufhor Defaffs: Prof. A.J. Sofomon Raju fs fhe Chafrman, Board of Sfudfes fn fhe Deparfmenf of Envfronmenfaf Scfences and he has pubffshed more fhan 300 research papers fn fnfernafonaf and nafonaf Journafs. He fs presenffy workfng on keysfone and endemfc specfes of Easfern Ghafs wffh fnancfaf supporf from DST and UGC.. Dr. K. Venkafa Ramana fs DST Posf-Docforaf Research Feffow (Young Scfenfsf Scheme) workfng fn Andhra Unfversffy. He has pubffshed more fhan fwenfy papers fn fnfernafonaf and nafonaf journafs. He fs presenffy workfng on keysfone and endemfc specfes of Easfern Ghafs wffh fnancfaf supporf from DST and UGC.

Aufhor Confrfbufon: Bofh fhe aufhors confrfbufed fo a sfmffar exfenf overaff.

Acknowfedgemenfs: We fhank fhe Andhra Unfversffy, Vfsakhapafnam, for provfdfng physfcaf facffffes.

LOGOs

10757 Pollination ecology ofRhynchosia cana Raju & Ramana

INTRODUCTION India at 13.68333056 N & 79.31666667 E. This area constitutes the forest cover of Tirumala Hills in the is a genus of the legume family Fabaceae, Seshachalam Hill Range in Chittoor District, which is subfamily , tribe , and subtribe primarily deciduous forest. The site is characterized by Cajaninae (Lackey 1981; Jayasuriya 2014). It consists a combination of rocky, undulating and steep terrain. In of approximately 200 and occurs in both the this area Rhynchosia cana grows in small populations eastern and western hemisphere in warm temperate or as scattered individuals in open areas, intermingled and tropical regions (Grear 1978). In the Eastern prominently with two prolific weeds, Hyptis suaveolens Ghats, 12 species of this genus have been reported to and Chromolaena odorata. be occurring almost in one region, Seshachalam Hills of southern Eastern Ghats of Andhra Pradesh. They include Flowering and floral biology R. beddomei, R. rufescens, R. suaveolens, R. cana, R. The flowering season was defined via regular field albiflora, R. capitata, R. courtollensis, R. densiflora, R. trips. Anthesis was initially recorded by observing 25 heynei, R. minima, R. rothii, R. rufescens, R. suaveolens marked mature buds in the field. These observations and R. viscosa. These species are either climbers or were repeated five times on different days in order shrubs (Madhavachetty et al. 2008). Of these,R. cana is to determine an accurate anthesis schedule for each distributed only in India and Sri Lanka. species. Similarly, the mature buds were monitored to Franco (1995) provided floral details of a species of record the time of anther dehiscence. The presentation Rhynchosia in Brazil. He reported that it is autogamous, pattern of pollen was also investigated by recording limited by spatial segregation between stigma and how anthers dehisced and confirmed by observing anthers. Levels of out-crossing are maintained by the anthers under a 10x hand lens. Details of flower retention of a pollination mechanism.Hypanthidium sp. morphology such as flower sex, shape, size, colour, and Centris sp. are the primary pollinators, and pollen is odour, sepals, petals, stamens and ovary were described deposited on the ventral part of their abdomen when the based on 25 flowers randomly collected from five . flower is probed. Craufurd & Prins (1979) reported that Careful observations were made regarding the position Rhynchosia sublobata is self-compatible and pollinated and spatial relationships of stamens and stigma in by Xylocopa bees. Etcheverry et al. (2011) reported mature buds, at anthesis and during flower lifespan with that Rhynchosia edulis and R. senna var. texana display reference to self and/or cross-pollination. valvular pollination mechanism; the former is facultative xenogamous while the latter is obligately xenogamous. Pollen output There is no other information on flowering phenology, Thirty mature but un-dehisced anthers from five breeding systems, pollen presentation mechanisms, different plants were collected and placed in a petri pollination mechanisms, pollinators and fruiting ecology dish. Single anthers were removed, placed on a clean of any species of Rhynchosia. There is one detailed microscope slide (75x25 mm) and dabbed with a study on the biochemical and nutritional aspects of R. drop of lactophenol-aniline-blue. Anther tissue was cana, R. filipes, R. rufescens and R. suaveolens, which observed under the microscope for pollen; if pollen suggests that their seeds have significant protein content was present the pollen mass was drawn into a band although they are under-utilized legumes (Kalidass & and the total number of grains was counted under a Mohan 2012). Keeping this in view, the present study compound microscope (40x objective, 10x eye piece). of the floral biology, sexual system, breeding systems, This procedure was followed for counting the number pollinators and fruiting ecology ofR. cana was conducted. of pollen grains in each anther collected. Based on This information is useful to take up breeding programs these counts, the mean number of pollen produced per aimed at improving the nutritional quality of legumes anther was determined. The mean pollen output per involving Rhynchosia species and Cajanus cajan. anther was multiplied by the number of anthers in the flower for obtaining the mean number of pollen grains per flower. The characteristics of pollen grains were also MATERIALS AND METHODS recorded.

Study site Pollen-ovule ratio The study region is located in the southern The pollen-ovule ratio was determined by dividing Eastern Ghats of Andhra Pradesh in peninsular the average number of pollen grains per flower by the

10758 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pollination ecology ofRhynchosia cana Raju & Ramana number of ovules per flower. The value thus obtained 2. The stigmas were pollinated with the pollen of was taken as pollen-ovule ratio (Cruden 1977). the same flower manually just five minutes after anthesis at 11:30hr by using a brush; they were bagged and Nectar characters followed to observe fruit set in manipulated autogamy. The presence of nectar was determined by observing 3. The stigma in each emasculated flower was mature buds and open flowers. Ten flowers were hand-pollinated with the pollen of two anthers of a used to determine the average volume of nectar per different flower on the same at 11:30hr by using flower expressed in µL. Flowers used for this purpose a brush; they were bagged and followed for fruit set in were bagged at the mature bud stage, opened after geitonogamy. cessation of nectar secretion and nectar was squeezed 4. The stigma in each emasculated flower was into micropipette to measure the volume. Nectar hand-pollinated with the pollen of two anthers of a sugar concentration was determined using a Hand different plant at 11:30hr by using a brush; they were Sugar Refractometer (Erma, Japan). Ten samples were bagged and followed for fruit set in xenogamy. used for examining the range of sugar concentration in All these categories of flower pollinations were the nectar. For the analysis of sugar types, the paper followed for fruit set for three weeks. If fruit set is there, chromatography method described by Harborne (1973) the percentage of fruit set was calculated for each mode. was used. Nectar was placed on Whatman No. 1 filter paper alongside standard samples of glucose, fructose Flower visitors and sucrose. The paper was run in ascending order for Flower foragers included bees and butterflies. The 24 hours with a solvent system of n-butanol-acetone- hourly foraging visits of each species were recorded on water (4:5:1), sprayed with aniline oxalate spray reagent three or four occasions, and the data were tabulated for and dried at 1200C in an electric oven for 20 minutes for further analysis. Plants in full bloom were selected to the development of spots from the nectar and standard record the foraging visits of insects. The data obtained sugars. Sugar types present and the most dominant type was used to calculate the percentage of foraging visits were recorded based on the area and colour intensity made by each species per day, and also to calculate the of spots. The sugar content/flower is expressed as the percentage of foraging visits of each species per day in product of nectar volume and sugar concentration per order to understand the relative importance of each unit volume, mg/µL. This is done by first noting the species. Their foraging behaviour was observed on a conversion value for the recorded sugar concentration number of occasions for the mode of approach, landing, on the refractometer scale and then by multiplying probing method, the type of forage collected, contact it with the volume of nectar/flower. Table 5.6 given with essential organs to result in pollination, inter-plant in Dafni et al. (2005) was followed for recording the foraging activity in terms of cross-pollination. conversion value to mg of sugars present in 1µl of nectar. Determination of pollen carryover efficiency of insects Stigma receptivity Ten specimens of each species were captured from In visual method, the stigma physical state (wet or flowers and brought to the laboratory. Each specimen dry) was considered to record the commencement was washed first in ethyl alcohol and the contents of receptivity. H2O2 test as given in Dafni et al. (2005) stained with aniline-blue on a glass slide and observed was followed for the confirmation of stigma receptivity under microscope to count the number of pollen grains period. present. From this, the average number of pollen grains carried by each species was calculated to know its pollen Breeding systems carryover efficiency. Mature flower buds of five inflorescences on five individuals were tagged and enclosed in paper bags. Natural fruit set, seed set, seed dispersal and seedling The breeding systems were repeated second time with ecology the same number of inflorescences and individual trees. Flowers on 50 plants were tagged prior to anthesis They were tested in the following way, and the number and followed for fruit set rate in open-pollinations. The of flower buds used for each mode of pollination is given same fruits were used to record seed set rate. Fruit in Table 1. maturation period, fruit dehiscence and seed dispersal 1. The flowers were fine-mesh bagged without aspects were observed to the extent possible. Field hand pollination for autonomous autogamy. observations were also made on fruit and seed dispersal

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 10759 Pollination ecology ofRhynchosia cana Raju & Ramana modes, seed germination and seedling establishment to the extent possible.

Photography Plant habitat, flowering inflorescences, and flower and fruit details were photographed with Nikon D40X digital SLR (10.1 pixel) and TZ240 stereo zoom microscope with SP-350 Olympus digital camera (8.1 pixel). Olympus binoculars (PX35 DPSR Model) was also used to make field observations. Magnus compound microscope - 5x, 10x, 40x and 100x magnification was a used for studying the pollen characteristics.

RESULTS

Phenology It is a perennial erect sub-shrub with slender stem that grows in open, rocky areas. The branchlets are glandular, sticky and velvet-hairy. The plant re-grows from below ground perennial root stock and from the seed during June-October (Image 4c). The plants show almost complete vegetative growth by the end of October b c (Image 1b). Hyptis suaveolens, a weedy labiate also Image 1. Rhynchosia cana: a - Habitat showing individual plants shows profuse growth simultaneously and its individuals mixed with the dense population ofHyptis suaveolens; are intermingled with the individuals of R. cana in many b - Vegetative phase; c - Flowering initiation. © A.J. Solomon Raju places (Image 1a). In this situation,H. suaveolens stands out prominently while R. cana with comparably low height is not prominent in appearance. In R. cana, the Flower morphology leaves are trifoliate with reticulate venation. The leaflets The flowers are pedicellate, small (9.2±0.7 mm long are petiolate, ovate, pointed, pubescent, especially and 7.6±0.5 mm wide), yellow, odorless, papilionaceous, beneath. The flowering occurs during November- zygomorphic and bisexual. The calyx is green with January with peak flowering in December (Image 1c). yellow tinge and consists of five free, lanceolate, hairy, The flowers are yellow, stalked (5mm) and borne in 5–6 mm long sepals. The corolla is bright yellow, pairs in leaf-axils. The plants wither and disappear in pubescent, specialized and consists of upper standard late March/early April. In H. suaveolens, the flowering petal, two wing petals and two keel petals. The standard occurs during November–February with peak flowering petal is large (8.3±0.6 mm long and 6.7±0.4 mm wide), during November–January. It produces multi-flowered yellow streaked with purple veins outside and inside cymes with violet-blue flowers and attracted most of but prominent at the bottom of the inside mid-region the local insect foragers, which exhibit fidelity to this which serves as nectar guide; the petal base is clawed floral source. In some areas Chromolaena odorata and consists of two inflexed fingernail auricles. The and R. cana were in full bloom simultaneously, and the standard petal envelops the rest of the petals in bud former species deprived the latter of insect foragers. H. but reflexes when the flower blooms. The two adjacent suaveolens and C. odorata with their profuse flowering petals (7.4±0.5 mm long and 3.7±0.4 mm wide), called outcompete R. cana when all the three occur in the wing petals surround the two bottom petals, called keel same habitat, hence R. cana with paired flowers in leaf petals (7.2±0.4 mm long and 3.4±0.5 mm wide). The axils attracted few insects even during its peak flowering keel petals form a proximal cylindrical part and a distal phase. In areas where H. suaveolens and C. odorata part consisting of a pressed angular pouch, with an acute flower simultaneously with R. cana, seeds produced by porate tip in which the stamens and stigma are housed. the latter species appear to be mainly derived from self The keel and the wing petals are attached by means of fertilization via autogamy and geitonogamy. two notched folds. The wing petals serve as alighting

10760 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pollination ecology ofRhynchosia cana Raju & Ramana platform for insects visiting the flowers. The stamens Table 1. Anthesis as a function of time inRhynchosia cana are 10, 5.4±0.5 mm long, diadelphous; nine filaments Time (h) No. of flowers anthesed Percentage of Anthesis are fused by the basal part into a sheath open along the 0930 0 0 upper side while the tenth filament is free and lies on the 1030 5 8 others (Image 2f). The distal parts of the filaments are 1130 15 25 free and contain 1mm long uniform dithecous anthers. 1230 26 43 The ovary is sessile, green, villous, 3.4±0.5 mm long and 1330 9 15 lies in the sheath of the filaments along the cylindrical part of the keel (Image 2h). It is monocarpellary and 1430 4 7 monolocular with two ovules or rarely three ovules 1530 1 2 arranged on marginal placentation (Image 2j,k). It has 1630 0 0 a long glabrous style with a capitate wet shiny stigma No. of mature buds tagged: 60 (Image 2i), both together accounts for a length of 4.3±0.4 mm. The stigma is situated at the height of the anthers. The distal portion of free filaments and style and stigma Pollination mechanism are incurved and clamped into the keel petals. The reproductive column is held under pressure within the keel part in open flowers, and it is exposed Floral biology when the pollinator presses against the wing and the Mature buds open during 1030–1530 h with peak keel petals. When bees land on the wing petals the anthesis during 1130–1230 h (Table 1; Image 2a-d; l-o). keel petals release the reproductive column explosively. Unfolding of the standard petal and wing petals indicates Consequently, the reproductive column snaps forward flowering opening. The keel petals do not unfold and against the standard petal, causing most of the pollen to remain in their original position as in mature bud stage be instantly released. The pollen thus released comes (Image 2e). All the 10 anthers in a flower dehisce at the into contact with the ventral side of the bee body. Since same time by longitudinal slits in mature bud stage. The the incurved stigma is situated above the height of the number of pollen grains per anther is 867±68.06 and anthers, it strikes the bee’s body first due to which cross- per flower is 8670. The pollen-ovule ratio is 4,335:1. pollination occurs if the bee visited the other flowers The pollen grains are monads, spheroidal, 31.52±3.80 previously and carried pollen on its ventral side and also µm in size, powdery and tricolporate, angulaperturate then the pollen ejected from the anthers powders the with reticulate exine (Image 2g). A nectariferous disc ventral side of the bee instantly. If it is the first visit for is present at the base of the ovary. The initiation of the bee to the flower, then it effects self-pollination upon nectar secretion occurs during mature bud stage and explosive release of reproductive column from the keel its cessation occurs an hour after anthesis. Individual boat. With the departure of the bee from the flower, the flowers produce 2.3±0.16 µl of nectar with 0.66mg of reproductive column does not return back to its former sugar. The nectar sugar concentration is 26% (Range 25– position but the keel moves forward partly covering the 28 %) consisting of sucrose, glucose and fructose with stamens and stigma. The downward movement of keel the first as dominant. Nectar is deeply concealed and it petals occurs in each subsequent foraging visit by bees. is open through two windows between the joined and Lycaenid butterflies were found to visit the flowers for the free filaments at the flower base. These windows nectar but they were ineffective to trip the keel petals allow access to the nectar. The stigma attains receptivity to cause explosive pollen release. If the flower is during anthesis and remains receptive for about three untouched or the keel is not tripped, the reproductive hours. After three hours of anthesis, the standard, column is never exposed and remains enclosed in the wing and keel petals gradually move close to each other keel boat. Such flowers wither and fall off. enclosing the reproductive organs (Image 2p–r). The closed flowers remain so even during most part of the Breeding systems fruit development. The calyx initially encloses the ovary In mature buds, anthers dehisce but autonomous and subsequently turns light brown and discloses the autogamy does not occur. Fruit set is absent in un- ovary since the latter gradually bulges and develops into manipulated autogamy, 18% in hand-pollinated a seeded pod. autogamy, 46% in geitonogamy, 90% in xenogamy and 81% in open-pollination (Table 2). In open-pollination, seed set rate is 54% (Table 3).

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 10761 Pollination ecology ofRhynchosia cana Raju & Ramana

a b c d e

j

f g h i k

l m n o

p q r Image 2. Rhynchosia cana: a–d - Different stages of anthesis from bud to flower; e - Keel and wing petals with stamens and stigma inside; f - Diadelphous condition of stamens; g - Pollen grain; h - Ovary; i - Capitate stigma, j–k - Ovules, l–o - Different stages of anthesis; p–r - Different stages of flower closure. © A.J. Solomon Raju.

10762 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pofffnafon ecofogy of Rhynchosfa cana Raju & Ramana

Tabfe 2. Resuffs of breedfng sysfems fn Rhynchosfa cana Nomfa sp. No. of Anfhfdfum sp. No. of fruffs Pofffnafon mode fowers Fruff sef (%) formed Freyerfa frochyfus pofffnafed Lampfdes boefcus Aufogamy (un- 50 0 0 manfpufafed and bagged) Chffades pandava Aufogamy (hand- 50 9 18 pofffnafed and bagged)

Geffonogamy 50 25 vfsffs 46 foragfng of No.

Xenogamy 50 44 90

Open-pofffnafon 1000 807 81 Tfme (h) Ffgure 1. Hourfy foragfng acfvffy of fnsecfs on Rhynchosfa cana

Insecf foragfng acfvffy and pofffnafon The fowers were foraged by bees for bofh necfar Thefr vfsffs fo un-frfpped fowers dfd nof confrfbufe fo and poffen, and by buferffes for necfar (Tabfe 4). The any pofffnafon. However, fhefr vfsffs fo frfpped fowers bees showed foragfng acfvffy from 1000h onwards confrfbufed fo pofffnafon due fo confacf befween when fhe mafure buds are sfff fn fhe process of anfhesfs fhefr proboscfs/forehead and fhe exposed sfamens and sfopped vfsffng fowers af 1700h (Ffg. 1). The bees and sfgma; as fhe fowers were afready frfpped, necfar fncfuded Apfdae and Megachfffdae of Hymenopfera. wfndows were exposed af fhe base of fhe sfandard pefaf The bees fncfuded Nomfa sp. (Apfdae) (Image 3a) and and hence buferffes were abfe fo coffecf necfar. Bofh Anfhfdfum sp. (Megachfffdae) (Image 3b–d); fhey bees and buferffes were found fo make fnfer-pfanf were afmosf consfsfenf foragers durfng fhe fowerfng ffghfs frequenffy due fo producfon of a few fowers per season. Buferffes were nof consfsfenf foragers and pfanf and fn fhfs process fndfvfduafs fowers appeared fo fhey showed foragfng acfvffy durfng 1100–1600 h wffh be vfsffed more fhan one or severaf vfsffs enhancfng fhe peak foragfng acfvffy 1200–1300 h (Ffg. 1). The foragfng pofffnafon rafe. acfvffy pafern of fnsecfs fndfcafed fhaf fhefr fower- vfsffng rafe fs dfrecffy refafed fo fhe avaffabffffy of Fruffng behavfor fowers; fhe peak anfhesfs fme fs confned fo noonfme Fruff growfh and devefopmenf begfns fmmedfafefy hours when humfdffy fs usuaffy fow and ffghf fnfensffy fs afer pofffnafon and ferfffzafon. The fruffs mafure hfgh. Bees accounfed for 68% and fycaenfd buferffes wffhfn fhree weeks (Image 3g–f). The sepafs encfose fhe 32% of fofaf foragfng vfsffs (Ffg. 2). Among bees, Nomfa growfng fruff fnffaffy and fhe fruff emerges ouf of fhe sp . made 35% whffe Anfhfdfum sp. made 33% of fofaf sepafs graduaffy. Fruff fs green fnffaffy and brown when foragfng vfsffs. The body washfngs of foragfng bees rfpe and dry. If fs a non-feshy, hafry, obfong, 15.7±0.6 and buferffes showed varfafon fn fhe poffen carryfng mm fong, 6.7±0.5 mm wfde, compressed befween capacffy; fhe average poffen recorded on Nomfa sp . seeds, rounded and apfcufafed pod. The pods produced was 168.2, Anfhfdfum sp. 125.4, F. frochyfus 32.8 , L. 1 or 2 or 3 seeds or none; 1-seeded pod sef fs 91.57%, boefcus 40.7 and C. pandava 45.1 (Tabfe 5; Image 3e,f). 2-seeded pod sef 7.19%, 3-seeded pod sef 0.62% and The fowers were vfsffed severaf fmes by bees buf new pod sef wffhouf seeds 0.62% (Tabfe 3; Image 3j–p). vfsffs fasfed shorfer fhan fhe frsf one. Wffh respecf fo fhefr behavfor, fhe bees fanded on fhe wfng pefafs Seed ecofogy and fhe keef, wffh fhefr head near fhe sfandard. They Mafure and dry fruffs wffh bf-vafvafe confgurafon fhen exerfed a cerfafn pressure wffh fegs on fhe wfng dehfsce expfosfvefy fo ejecf seeds vfofenffy due fo whfch pefafs unff fhese and fhe keef benf downwards, and fhey faff weff away from fhe parenf pfanf (Image 4a,b). fhen proceeded fo coffecf necfar durfng whfch fhe bee’s The seed fs brownfsh bfack, renfform, fnefy pubescenf, abdomen appeared poffen smofhered (sfernofrfbfc 3.7±0.5 mm fong and 2.8±0.3 mm wfde and shfny poffen deposffon). The bees fook “U” furn afer necfar wffh cushfoned sfrophfofe (Ffg. 3q). Seeds germfnafe coffecfon and proceeded fo fhe sfamens fo coffecf durfng rafny season whfch sfarfs from June. Seedffngs poffen. In case of buferffes, fhey afempfed fo vfsff bofh grow confnuaffy buf fhefr growfh rafe fs subjecf fo fhe un-frfpped and frfpped fowers. They never frfpped keef avaffabffffy of mofsfure sfafus of fhe soff (Image 4d-h). pefafs due fo fhefr ffghf wefghf and afso coufd nof access necfar due fo profecfon of necfar by fhe sfandard pefaf.

Journaf of Threafened Taxa | www.fhreafenedfaxa.org | 26 Ocfober 2017 | 9(10): 10757–10770 10763 Pollination ecology ofRhynchosia cana Raju & Ramana

a b c

d e f

g h i j k l m

n o p q

Image 3. Rhynchosia cana: a - Nomia sp. collecting nectar; b–d - Anthidium sp. collecting pollen from un-anthesed flowers (b), collecting nectar from open flower (c), collecting pollen from open flower (d); e & f - Lycaenid, Chilades pandava collecting nectar; g–i - Different stages of fruit development before maturity; j - 1-seeded pod; k - 2-seeded pod; l - 3-seeded pod (not dehisced); m–p - Dehisced 3-seeded, 2-seeded and 1-seeded pods; q - Seeds. © A.J. Solomon Raju.

10764 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pofffnafon ecofogy of Rhynchosfa cana Raju & Ramana

Tabfe 3. Nafuraf fruff/seed sef rafe fn Rhynchosfa cana

Rafe of seed sef/fruff No. of No. of Fruff sef Seed sef No. of 1-seeded No. of 2-seeded No. of 3-seeded No. of Seedfess fowers fowers (%) (%) 1-seeded fruff sef 2-seeded fruff sef 3-seeded fruff sef seedfess fruffs sampfed sef fruff fruffs (%) fruffs (%) fruffs (%) fruffs (%) 1000 807 81 54 739 91.57 58 7.19 5 0.62 5 0.62 No. of ovufes/fower = Two; rarefy Three

Tabfe 4. Lfsf of fnsecf foragers on Rhynchosfa cana Order Famffy Sub-famffy Genus Specfes Common Name Foragfng Forage coffecfed schedufe Hymenopfera Apfdae Nomffnae Nomfa sp. Afkaff bee 1000–1700 Necfar + Poffen

Megachfffdae Megachfffnae Anfhfdfum sp. Pofer bee 1000–1700 Necfar + Poffen

Lepfdopfera Lycaenfdae Pofyommafnae Freyerfa Trochyfus Freyer Grass Jewef 1100–1600 Necfar

Lampfdes boefcus L. The Pea Bfue 1100–1600 Necfar

Chffades Pandava Horsfefd The Pfafns Cupfd 1100–1600 Necfar

Tabfe 5. Poffen recorded fn fhe body washfngs of bee foragers on Rhynchosfa cana

Sampfe Number of poffen grafns Bee specfes sfze (N) Range Mean S.D % of foragfng vfsffs foragfng of % Nomfa sp. 10 76-281 168.2 67.62

Anfhfdfum sp. 10 58-185 125.4 48.87 Freyerfa frochyfus 10 21-47 32.8 8.70 Ffgure 2. Refafve percenfage of foragfng vfsffs of bees and Lampfdes boefcus 10 23-56 40.7 11.50 buferffes on Rhynchosfa cana

Chffades pandava 10 31-59 45.1 9.26

specfes oufcompefe R. cana fn afracfng a wfde array of fnsecfs fo fhefr fowers, ffmffng pofffnafor avaffabffffy. DISCUSSION In R. cana , fhe hermaphrodffc sexuaf sysfem fs funcfonaf due fo producfon of ferffe poffen grafns and The presenf sfudy shows fhaf Rhynchosfa cana fs funcfonaf ovarfes. The fowers dfspfay near-synchronous a dry decfduous specfes and grows fn rocky areas wffh hermaphrodffsm or homogamy due fo fhe occurrence ffffe fffer confenf. In fhe absence of fnformafon on fhe of anfher dehfscence fn fhe mafure bud sfage, and habffafs of ffs occurrence, ff fs nof possfbfe fo evafuafe recepfvffy of sfgmas durfng anfhesfs. The enfre whefher ff has fhe abffffy fo grow fn dfverse habffafs or reproducfve cofumn sfays fnsfde fhe keef pefafs even nof. If fs an erecf sub-shrub, whfch shows growfh from afer anfhesfs; fn fhfs sffuafon, fhere fs a ffkeffhood of perennfaf roof sfock durfng fhe rafny season. If afso aufonomous aufogamy. Neverfhefess, hand pofffnafon produces new pfanfs from seed sfock af fhe same fme. fesfs fndfcafed fhaf aufonomous aufogamy does nof Fuff feaf fushfng fs compfefe by fhe end of Ocfober and occur despffe seff-compafbffffy, affhough ff remafns fhen onwards, foraf bud fnffafon fakes pface. The funcfonaf because fruffng occurred when sfgmas fowerfng perfod fs confned fo wfnfer season. Indfvfduaf were brushed wffh fhefr own poffen. Such a sffuafon pfanfs produce a smaff number of fowers durfng fhefr suggesfs fhaf fhe fowers are essenfaffy dependenf fffefme due fo producfon of onfy a pafr of fowers fn feaf on fower foragers for fruff sef fhrough seff- as weff as axffs. The pfanf durfng fowerfng phase afracfs onfy fwo cross-pofffnafon. If appears fhaf fhe sfgma, affhough bee specfes consfsfenffy and fhree specfes of buferffes recepfve, bfocks fhe germfnafon of fhe seff-poffen occasfonaffy due fo sfmuffaneous fowerfng of fhe weeds whffe ff fs fn keef pefafs and hence ff essenfaffy requfres Hypfs suaveofens and Chromofaena odorafa, whfch fhe rupfure of ffs surface by a pofffnafor fo affow fhe seff- produce farge popufafons wffh profffc fowerfng. These or cross poffen fo germfnafe. Such a sfgmafc regufafory

Journaf of Threafened Taxa | www.fhreafenedfaxa.org | 26 Ocfober 2017 | 9(10): 10757–10770 10765 Pollination ecology ofRhynchosia cana Raju & Ramana

a b c

d e f

g h

Image 4. Rhynchosia cana: a - Dry fruits; b - Explosive pod dehiscence; c - Emergence of new branches from perennial root stock; d–f - Growth of new plants; g & h - Established seedlings from seeds. © A.J. Solomon Raju. function appears to have evolved to discourage selfing pollinator permits the pollen to germinate in the flowers and promote out-crossing. Shivanna & Owens (1989) of Phaseoleae members with thick stigmatic cuticle. On stated that the rupture of the stigmatic surface by the contrary, Castro & Agullo (1998) reported that in

10766 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pollination ecology ofRhynchosia cana Raju & Ramana

Vigna, a member of the tribe Phaseoleae, autonomous nectar at the corolla base and hidden pollen in keel self-pollination may occur by spontaneous rupture of petals. the stigmatic membrane. Similar stigmatic surface that Within the sub-family Papilionoideae, primary and prevents self-fertilization has also been reported inVicia secondary pollen presentations have been reported. faba (tribe Vicieae) (Lord & Heslop-Harrison 1984) and In plants with primary pollen presentation, pollen is in scutellata (tribe ) (Kreitner & delivered directly from the anthers to the vector’s Sorensen 1985); however, in these species auto-fertile body. In plants with secondary pollen presentation, lines have been reported to have thin stigmatic cuticles pollen grains are delivered first on a floral part such as allowing spontaneous disruption and self-fertilization. the keel petals in Papilionoideae and then on the body In R. cana, the stigmatic surface appears to have thick of the vector implying an accurate delivery of pollen cuticle and does not have the mechanism of causing on the vector’s body (Howell et al. 1993). These two spontaneous rupture to facilitate autonomous self- pollen presentation patterns are associated with the pollination. In effect, the tripping of keel petals appears four types of basic pollination mechanisms - valvular, to be essential to cause rupture on the stigmatic surface pump, explosive and brush, all of them are associated by the tripping agent due to which there is more with a particular floral architecture and kinetics. In likelihood of the occurrence of either geitonogamy or the valvular type, pollen presentation is primary, xenogamy. The fruit set rates recorded in hand-pollinated whereas in the other three mechanisms, it is secondary geitonogamy and xenogamy also substantiate that the (Yeo 1993). In the explosive mechanism, commonly plant is facultative xenogamous, a breeding system that only one pollination event occurs and it has evolved is flexible and keeps the options open for both selfing independently in several tribes (Small 1988), while and out-crossing mediated by pollen vectors. in the other three mechanisms, repeated visitation is Schrire (1989) stated that the ecological and possible (Westerkamp 1997). In the present study, R. evolutionary success of Leguminosae has been related cana flowers have explosive pollination mechanism to biotic pollination mechanisms. The three sub- and deliver pollen directly from the anthers to the families within this family have achieved a characteristic bee’s body when keel petals are tripped by the foraging floral architecture, in which plants within the subfamily bee; this type pollen delivery is the representative of Papilionoideae have developed the most complex floral primary pollen presentation associated with explosive mechanisms. Plants within the Papilionoideae have pollination mechanism. In the flowers, the staminal zygomorphic flowers that are mainly bee-pollinated column is held under pressure within the keel, and when (Westerkamp 1997); although bird pollination and the tension is released by the forager, the same column bat pollination have also been recorded within the snaps forward against the standard petal causing all the subfamily (Ortega-Olivencia et al. 2005). In bee- pollen to be instantly released. The reproductive column pollinated flowers of Papilionoideae, it is assumed that remains exposed and does not return back to its original each part of the corolla is specialized for a particular role state but the keel petals return back partially covering in pollinator attraction and the success of pollination. the stamens and stigma. The efficiency of explosive The flag or standard petal attracts pollinators; the keel pollination mechanism depends on the ambient protects androecium and gynoecium and, together weather conditions, especially temperature and relative with the wings, provides a platform for the insects to humidity. Since R. cana flowers during winter season, land on. The wings also operate as levers that raise or it accordingly commences anthesis from late morning lower the keel (Stirton 1981). The flowers typical of onwards by which time the ambient air will be relatively pollination by the bee family Apidae are zygomorphic, dry and hence is conducive for the efficient functioning of bright yellow or blue with nectar guides, and frequently the explosive pollination mechanism. Further, the bees with hidden rewards such as those in the Lamiaceae, also commence their foraging activity at about the same Scrophulariaceae, Fabaceae and Orchidaceae (Faegri & time and continue forage collection until the flowers van der Pijl 1979). In the present study, the Fabaceae close back. The concealment of the stamens within member, R. cana has papilionaceous corolla with flag, the keel petals until it is tripped is an advantage for the wing and keel petals; the flag petal serves as a visual plant to secure pollen from unusual rains and ambient attractant, wing petals provide landing platform and moisture conditions during the flowering season of this keel petals protect the entire reproductive column. The plant (Peter et al. 2004). flowers are typical of pollination by bees since they are Percival (1961) stated that plants with deep-tubed zygomorphic, standard petal with nectar guide, hidden flowers tend to produce sucrose-rich nectar, whereas

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 10767 Pollination ecology ofRhynchosia cana Raju & Ramana those with open or shallow-tubed flowers tend to suaveolens and Chromolaena odorata with huge flower be hexose-rich. Baker & Baker (1983) stated that density diverted bee populations to their flowers; in this flowers with long corolla tube possess more sucrose situation, Nomia and Anthidium bees confined their in their nectar while those with short tubes possess foraging visits to R. cana flowers and effected pollination. more hexoses in their nectar. In the present study, R. In R. cana, the keel tripping process is not self-activated cana with short corolla tube presents sucrose-rich to effect pollination. The flowers depend on bees for nectar because the nectar is perfectly concealed and tripping of the keel petals to enable the working of hence is not exposed for the breakdown of sucrose explosive pollination mechanism. The flowers that were into hexoses. Concealment of nectar in this species is not tripped by external agents subsequently fall off. This adaptive to protect against microorganisms, particularly situation explains that the plant is obligately dependent yeasts, whose metabolic activities dramatically change on bees for pollination. The bees visiting the flowers nectar chemistry and the plant benefits from keeping seem to be efficient in tripping the flowers because the the nectar as sterile as possible to maintain control flower size and petal strength are commensurate with over its chemical composition in order to maximize the bee size and the force employed to depress the wing pollination rate by attracting appropriate pollinators petals to access the nectar. (Herrera et al. 2008). Bees prefer flowers with sucrose Cruden (1977) used the pollen-ovule (P/O) ratios as as chief constituent of nectar (Kevan 1995). The flowers indicators of breeding systems of plants. He provided pollinated by long-tongued bees produce sucrose-rich P/O ratios for different breeding systems: 168.5+22.1 nectar (Baker & Baker 1990). In line with this, R. cana for facultative autogamy, 798.6+87.7 for facultative with melittophilous pollination syndrome also produces xenogamy and 5859.2+936.5 for xenogamy. Several sucrose-rich nectar which is utilized principally by long- workers followed these P/O ratios to classify breeding tongued bees and occasionally by lycaenid butterflies. systems of the plant species studied by them. Arroyo Opler (1983) and Cruden et al. (1983) noted that bee- (1981) stated that the P/O varies according to the flowers tend to produce a small volume of nectar with pollination mechanism within Papilionoideae. These higher sugar concentration than the nectar of flowers authors suggested that the plants with explosive pollinated by other animals. Pyke & Waser (1981) mechanism have a low P/O because a single pollinator stated that the nectar sugar concentration of flowers visit is needed for efficient transference of pollen; this pollinated by bees is generally higher when compared low P/O is a consequence of the highly specialized, to those pollinated by butterflies and hummingbirds; irreversible pollination mechanism, which allows only bee-pollinated flowers tend to produce nectar with one effective exchange of pollen with pollinators. sugar concentration more than 35% while butterfly or Small (1988) stated that Medicago species of the tribe hummingbird pollinated flowers tend to produce nectar Trifolieae with explosive pollination mechanism displays with sugar concentration ranged between 20% and 25%. the lowest pollen-ovule ratios. Lopez et al. (1999) In the present study the flowers of R. cana produce a recorded explosive pollination mechanism with highest small volume of nectar with 26% sugar concentration. pollen-ovule ratios in certain genera of the Fabaceae such Further, the energy yield from the nectar appears to as Cytisus, Pterospartum, Teline, , Stauracanthus be in tune with the requirement of energy by bees. and Cytisophyllum. Etcheverry et al. (2011) stated that Therefore, R. cana flowers with explosive pollination the Fabaceae plants which they studied with explosive mechanism, primary pollen presentation, and hidden pollination mechanism had intermediate pollen-ovule nectar and pollen have evolved to discourage other ratios. These authors mentioned that Rhynchosia foragers from visiting the flowers and to ensure that edulis and R. senna var. texana have valvular pollination the bees get the floral rewards. Accordingly, bees mechanism with primary pollen presentation. Both the have been found to be the principal pollinators while species are classified as obligate xenogamous based on lycaenid butterflies to be the secondary pollinators. P/O ratio but R. edulis has been found to be facultative This observation has been substantiated by their pollen xenogamous in hand-pollination tests. Craufurd & Prins carrying capacity as realized in body washings after (1979) reported that R. sublobata is self-compatible and their visits to the flowers. The foraging activity pattern facultative xenogamous in hand-pollination tests; it is of insects suggests that their peak foraging activity is pollinated by Xylocopa bees. In the present study, R. correlated with peak anthesis time when humidity is cana shows highest P/O ratio when compared to that usually low and light intensity is high. In the habitats of facultative xenogamy used by Cruden (1977). The of R. cana, the extensive populations of weeds, Hyptis highest P/O ratio in this plant species appears to be a

10768 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770 Pollination ecology ofRhynchosia cana Raju & Ramana consequence of pollen collection activity by bees. It during rainy season but there is a deficit of rainfall due appears to be inevitable for R. cana to produce high to which the migration of seeds is profoundly affected. P/O to compensate the pollen loss caused by pollen Remanandan (1981) stated that Rhynchosia, being collectors and ensure the function of its vector- closely related to the genus Cajanus, some of its species dependent facultative xenogamous breeding system. can be used to provide substantial contributions towards Further, this is substantiated by natural fruit and seed crop improvement in pigeon pea. Furthermore, some set rates evidenced in the study. The study showed species of Rhynchosia have been experimented in India that seed set rate is only 54% although natural fruit set to provide physiological resistance against insect pests is 81% of which the production rate of 1-seeded fruits such as pod-borer and pod-fly in pigeon pea. In this exceed 90% despite the characteristic of production of study, the seeds of R. cana have not been infested by any two ovules per flower. This state of fruiting and seed pod-borer or pod-fly suggesting that it has physiological set rates could be attributable to pollen and pollinator resistance against insect pests. Therefore, intensive and limitation, and also to nutrient environment of the soil extensive research is suggested to identify and select where the plant grows. The study also showed that desirable genotypes of R. cana that give physiological the flowers of R. cana produce three ovules rarely and resistance against pod or seed pests in order to use them such flowers could produce fruits with three seeds if for crop improvement in pigeon pea. pollinated. Further, the study also noted the production of fruits without seeds but the percentage of such fruits is highly negligible. However, this situation in R. cana REFERENCES indicates that the plant initiates and produces fruit cover first and then the gradual production of seeds. Ali, H.H., A. Tanveer & Nadeem (2012). Evaluation of some seed dormancy breaking methods on germination ofRhynchosiacapitata The production of seedless fruits can be attributed to (Roth DC). Pakistan Journal of Weed Sciences Research 18: 423–432. nutrient-deficiency in the soil and to genetically inferior Arroyo, M.T.K. (1981). Breeding systems and pollination biology in fertilization arising from self-pollination. Leguminosae, pp. 723–769. In: Polhill, R.M. & P.H. Raven (eds.). Advances in Legume Systematics, Part 2, Royal Botanical Gardens, Tran & Cavanagh (1984) reported that in Leguminosae, Kew, London. seeds of many taxa exhibit physical dormancy due to the Baker, H.G. & I. Baker (1983). Floral nectar sugar constituents in presence of a water impermeable seed coat. With this relation to pollinator type, pp. 117–140. In: Jones, C.E. & R.J. Little (eds.). Handbook of Experimental Pollination Biology, Scientific and dormancy, they remain viable for long period of time. Academic Editions, New York. Ali et al. (2012) reported such physical dormancy in Baker, H.G. & I. Baker (1990). The predictive value of nectar chemistry Rhynchosia capitata to the recognition of pollinator types. Israel Journal of Botany 39: due to which this species is successful 157–166; http://doi.org/10.1080/0021213X.1990.10677140 as a weed. Shaukat & Burhan (2000) described seed Castro, M.A. & M.A. Agullo (1998). Anatomy of the stigma of characteristics and the factors regulating germination Vigna adenantha (G.F. Mayer) Marechal. Mascherpa and Stainer (Leguminosae, Papilionoideae). Biocell 22: 9–18. of R. minima in Pakistan; it exhibits differential success Craufurd, R.Q. & W.H. Prins (1979). Munkolo (Rhynchosia sublobata), in different habitats with different microclimates. Seed a promising pasture legume for Zambia. Tropical Grasslands 13: characteristics or dormancy studies on R. cana have not 45–52. Cruden, R.W. (1977). Pollen-ovule ratios: a conservative indicator of been made by previous workers. In the present study, it breeding systems in flowering plants. Evolution 31: 32–46; http:// is found that the seeds of R. cana respond to monsoonal doi.org/10.2307/2407542 rains, germinate and produce seedlings. During the Cruden, R.W., H.M. Hermann & S. Peterson (1983). Patterns of nectar production and plant-pollinator coevolution, pp. 80–125. study period, the rainy season is characterized by erratic In: Bentley, B. & T. Elias (eds.). The Biology of Nectaries. Columbia rainfall with long dry spells; in effect, the many growing University Press, New York. seedlings perished while a few continued to grow in Dafni, A., P.G. Kevan & B.C. Husband (2005). Practical Pollination Biology. Enviroquest Ltd., Cambrige, Ontario, 315pp. areas where water stress is not severe. In case of H. Etcheverry, A.V., M.M. Aleman, T. Figueroa-Fleming, D. Lopez- suaveolens and C. odorata, they showed quick growth Sphar, C.A. Gomez, C. Yanez, D.M. Figueroa-Castro & P. Ortega- and soon shaded or covered the growing seedlings of Baes (2011). Pollen: ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): an evaluation with R. cana due to which the latter could not grow further Argentine species. 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Threatened Taxa

10770 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2017 | 9(10): 10757–10770

OPEN ACCESS The Journal of Threatened Taxa is dedicated to building evidence for conservation globally by publishing peer-reviewed articles online every month at a reasonably rapid rate atwww.threatenedtaxa.org . All articles published in JoTT are registered under Creative Commons Attribution 4.0 International License unless otherwise mentioned. JoTT allows unrestricted use of articles in any medium, reproduction, and distribution by providing adequate credit to the authors and the source of publication. ISSN 0974-7907 (Online); ISSN 0974-7893 (Print) October 2017 | Vol. 9 | No. 10 | Pages: 10741–10864 Date of Publication: 26 October 2017 (Online & Print) DOI: 10.11609/jott.2017.9.10.10741-10864 www.threatenedtaxa.org

Articles New records of the subfamily Stratiomyinae (Diptera: Stratiomyidae) from Pakistan The relationship between artificial food supply and natural -- Muhammad Asghar Hassan, Imran Bodlah, Zafar Iqbal & food selection in two troops of commensal Hamadryas Rubina Jabeen, Pp. 10831–10834 Baboons Papio hamadryas (Mammalia: Primates: Cercopithecidae) in Saudi Arabia A new distribution report of the Critically Endangered -- Ahmed Boug, M. Zafar-ul Islam, Toshitaka Iwamoto, Amomum kingii Baker (Zingiberaceae) outside Sikkim, India Akio Mori, Akihiro Yamane & Amy L. Schreier, Pp. 10741– -- Sreetama Bhadra & Maumita Bandyopadhyay, Pp. 10835– 10756 10838

Pollination ecology ofRhynchosia cana (Willd.) DC. A note on the population status and threats on two endemic (Fabaceae), an erect sub-shrub, in peninsular India and Endangered species of Garcinia of Agasthyamalai -- A.J. Solomon Raju & K. Venkata Ramana, Pp. 10757–10770 Biosphere Reserve, Tamil Nadu, India -- G. Manikandan & R. Ramasubbu, Pp. 10839–10845

Communications Feasibility study on the vegetative propagation of four endemic rare balsams (Impatiens spp.) through stem Feeding habits and behaviour of Bagre bagre and Genidens cuttings for conservation and management in Idukki District, barbus, two ariid catfishes (Pisces: Siluriformes) from Kerala, India southeastern Brazil -- G. Prasad, P. Rajan & N. Bhavadas, Pp. 10846–10849 -- Maria Thereza Manhães Tavares & Ana Paula Madeira Di Beneditto, Pp. 10771–10775 Notes The Odonata (Insecta) of northern and central Oman -- Elaine M. Cowan & Peter J. Cowan, Pp. 10776–10791 Observation of Shaheen Falcon Falco peregrinus peregrinator (Aves: Falconiformes: Falconidae) in the Illustrated description and notes on biology of Nilgiris, Tamil Nadu, India Dicranocephalus lateralis (Signoret) (Coreoidea: -- Arockianathan Samson, Balasundaram Ramakrishnan, Stenocephalidae) from Maharashtra State, India Palanisamy Santhoshkumar & Sivaraj Karthick, Pp. 10850– -- Balasaheb V. Sarode, Nikhil U. Joshi, Swapnil S. Boyane, 10852 Subodh S. Gaikwad, Pratik P. Pansare & Hemant V. Ghate, Pp. 10792–10803 A first report of Redigobius oyensi (de Beaufort, 1913) (Teleostei: Gobionellinae) from Car Nicobar Island, India Floristic diversity of the Indian Cardamom Research Institute -- J. Praveenraj, R. Kiruba-Sankar, Lohith Kumar, campus, Myladumpara, Western Ghats, India J. Raymond Jani Angel & S. Dam Roy, Pp. 10853–10855 -- Anoop P. Balan & S. Harikrishnan, Pp. 10804–10822 New butterfly records from the Jammu Shiwaliks, Jammu & Kashmir, India Short Communications -- Shakha Sharma & Neeraj Sharma, Pp. 10856–10859

Mammals of Papikonda Hills, northern Eastern Ghats, India Range extension of the Indian Tortoiseshell -- Vikram Aditya & Thyagarajan Ganesh, Pp. 10823–10830 Aglais caschmirensis aesis (Fruhstorfer, 1912) (Lepidoptera: Nymphalidae) into the hills of Manipur, India -- Jatishwor Singh Irungbam, Harmenn Huidrom & Baleshwor Singh Soibam, Pp. 10860–10864

Threatened Taxa