the indian stingless bee or dammar bee , tetragonula iridipennis , is a species of bee belonging to the family apidae , subfamily apinae .

it was first described by frederick smith in 1854 who found the species in what is now the island of sri lanka .

many older references erroneously placed this species in melipona , an unrelated genus from the new world , and until recently it was placed in trigona , therefore still often mistakenly referred to as trigona iridipennis .

for centuries , colonies of t. iridipennis have been kept in objects such as clay pots so that their highly prized medicinal honey can be utilized .

tetragonula iridipennis

tetragonula iridipennis <EOT> indian stingless bee ( tetragonula iridipennis ) collecting nectar and pollen from a table rose plant .

supplementary figure 1 : map indicating the sampling sites of tetragonula iridipennis . ( jpeg 151 kb )

supplementary figure 2 : scatterplot of the two first axis of the canonical discriminant analysis of the indian populations of tetragonula iridipennis . ( jpeg 134 kb )

supplementary figure 4 : nest structure of tetragonula iridipennis bees sampled in assam state , with the brood cells constructed as layers . ( jpeg 371 kb )

supplementary figure 3 : nest structure of tetragonula iridipennis bees sampled in kerala state , with the brood cells constructed as a cluster . ( jpeg 494 kb )

kerala will host an indo - australian collaborative research project on deploying stingless bee ( tetragonula iridipennis ) as pollinators to increase the agricultural yield of fruits and vegetables .

floral sources for stingless bees ( tetragonula iridipennis ) in nellithurai village , tamilnadu , india doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra04

view from inside nest of tetragonula cf sarawakensis , you can see brood cells , food storage and queen . pemandangan dari dalam sarang tetragonula cf sarawakensis , anda boleh nampak sel telur , storan makanan dan ratu .

stingless bees are a large and diverse taxon and more than 500 species are recorded worldwide . based on the previous reports , totally seven species are reported from india . the present study reports male and female morphological characteristics of tetragonula iridipennis belonging to tetragonula subgenus in nellithurai village , tamil nadu , india . the species of tetragonula group are extremely similar in the external morphology of the workers . the glabrous bands on mesoscutum of female bees and structure and arrangement of gonostylus and the penis valve of male bees are used to separate t . iridipennis from other species in this group . the identity of t . iridipennis is made based on the male and female key morphological characteristics in the previous literature .

< p > stingless bees are a large and diverse taxon and more than 500 species are recorded worldwide . based on the previous reports , totally seven species are reported from india . the present study reports male and female morphological characteristics of < em > tetragonula iridipennis < / em > belonging to < em > tetragonula < / em > subgenus in nellithurai village , tamil nadu , india . the species of < em > tetragonula < / em > group are extremely similar in the external morphology of the workers . the glabrous bands on mesoscutum of female bees and structure and arrangement of gonostylus and the penis valve of male bees are used to separate < em > t . iridipennis < / em > from other species in this group . the identity of < em > t . iridipennis < / em > is made based on the male and female key morphological characteristics in the previous literature < strong > . < / strong > < / p >

[ pooja singh and m . s . khan ( 2015 ) ; assessment of comparative foraging activity in queen right and queen less colony of stingless bee , tetragonula iridipennis smith ( hymenoptera : apidae ) int . j . of adv . res . 3 ( 7 ) . 498 - 502 ] ( issn 2320 - 5407 ) . urltoken

danareddi , c . s . & s . viraktamath ( 2009 ) . morphometrical studies on the stingless bee , trigona iridipennis smith karnataka journal of agricultural sciences 22 : 796 - 797 .

furthermore , a case of collapse of a managed colony of a stingless bee , tetragonula iridipennis , was reported in india ( vijayakumar et al . , 2013 ) . although the mite illustrated in this paper belongs to the family cheyletidae , the yellowish carpet of dust on the bottom of the nest ( figs . 2a , b ) may represent an astigmatid mite ( carpoglyphus ) .

mohan , r . & s . devanesan ( 1999 ) . dammer bees , trigona iridipennis smith . ( apidae : meliponinae ) in kerala . insect environment 5 ( 2 ) : 79 .

sakagami , s . f . ( 1978 ) . tetragonula stingless bees of the continental asia and sri lanka ( hymenoptera : apidae ) . journal of the faculty of science , hokkaido university , series vi , zoology 21 : 165 - 247 .

rathor , v . s . , c . rasmussen & m . s . saini ( 2013 ) . new record of the stingless bee tetragonula gressitti from india ( hymenoptera : apidae : meliponini ) . journal of melittology 7 : 1 - 5 .

sakagami , s . f . & t . inoue ( 1987 ) . stingless bees of the genus trigona ( subgenus trigonella ) with notes on the reduction of spatha in male genitalia of the subgenus tetragonula ( hymenoptera , apidae ) . kontyû 55 : 610 - 627 .

vijayakumar , k . , m . muthuraman & r . jayaraj ( 2012 ) . predation of stingless bees ( trigona iridipennis : apidae , meliponinae ) by centipede ( scolopendra hardwicki : chilopoda : scolopendramorpha ) . international journal of advanced life sciences 5 ( 2 ) : 156 - 159 .

trigona ( tetragonula ) iridipennis smith is the common stingless bee found in south india . they are domesticated in mud pots , bamboo bits , wooden boxes or coconut shells . meliponiculture is being popularized in the rural homesteads for poverty alleviation and additional income generation . the chapter discusses the studies conducted as part of all india coordinated research project ( aicrp ) on honey bees and pollinators at vellayani centre of kerala agricultural university . among the various types of hives with different volumes , bamboo bits with 1500 cc showed better brood development and storage of honey . due to scarcity of bamboo nodes , a wooden box with 1960 cc volume with two equal halves was designed which helped for easy division and mass multiplication of colonies . the technologies were disseminated to the public by imparting trainers’ training in different districts of kerala in which 174 women and 322 men were trained . augmentation , conservation and management of t . iridipennis should be intensified for ensuring sustainable agriculture and the conservation of biological diversity resulting in food security for poverty eradication .

description of biodiversity is often cited as one of the most important actions necessary for conservation programs . there are more than 600 species of stingless bees spread over the tropical regions of the world ; though for various species , little is known about their biology and taxonomy . we sampled bees from feral colonies from various regions of india and compared them using wing morphology . the results of population analysis of the patterns of wing venation , using geometric morphometric techniques , suggested the existence of at least two phenotypic clusters within our samples of the so - called tetragonula iridipennis complex . these findings were supported by other features , including differences in nest architecture . this helps to explain the patterns of variability found in stingless bees in india and also will be valuable for conservation planning .

three social bees accounted for 99 % of flower visitors : apis dorsata , the giant asian honey bee ( 47 % of visitors ) , apis cerana ( 24 % ) , and tetragonula iridipennis ( 27 . 9 % ) . all these bees were observed to carry pollen and contact the stigma during flower visits . two ceratina and one xylocopa species were occasional visitors to coffee flowers ( 1 % of visits ) . as bee diversity was low , we evaluated pollination and fruit set on the basis of overall bee abundance , assuming that each pollinator species is equivalent in terms of pollination effectiveness . as there is no information available on the equivalency of pollinator effectiveness of these flower visitors , we feel the assumption of equivalence is preferable to the introduction of potential errors based on speculation .

stingless bees are good pollinator in tropical and sub - tropical regions in india and known to pollinate various plants of economic importance and generally better used in planned pollination . foraging performance by bees is a clear indicator of functioning and strength in colony . a various circumstances affect the normal foraging activity by bees . the queenless ness in the bee colony may be one of those reasons which alter the normal foraging activity in respect to foraging activity in a normal queen right bee colony . the present study was done with the aim to know how the absence of queen in a colony of stingless bee tetragonula iridipennis smith affects its foraging activity and to find out a comparative conclusion between foraging activity in a queen less and a queen right colony of t . iridipennis . the study revealed some important differences in foragging jobs in both types of colonies . number of outgoing forager bees and resin collector bees were remain almost similar in queen right ( 61 . 149 and 2 . 893 bees per 5 min ) and queenless colony ( 62 . 744 and 3 . 618 bees per 5 min ) , respectively , but pollen collector bees were found to be more active in queen less colony ( 20 . 560 bees per 5min ) then in queen right colony ( 1 . 814 bees per 5 min ) , respectively , and the activity of cleaner bees and nectar collecting bees were more in queen right colony ( 5 . 367 and 61 . 39 bees per 5 min ) than in queen less colony ( 2 . 569 and 44 . 957 bees per 5 min ) , respectively .

peeking into the entrance of a stingless bee hive - kuala koh , taman negara , peninsular malaysia .

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the authors are grateful to the department of biotechnology , ministry of science and technology , government of india , new delhi , for funding this study . acknowledgements are also due to dr . m . muthuraman , tamil nadu agricultural university , coimbatore , dr . a . j . solomon raju , andhra university , visakhapatnam , dr . k . v . lazar , calicut university , calicut , dr . s . devanesan , kerala agricultural university , thiruvananthapuram , dr . sushamachapalkar , vidyapratishtan’s school of biotechnology , baramati and dr . a . rahman , assam agricutural university , jorhat , for making the material available for the study . the authors are grateful to fapesp ( proc . 2011 / 07857 - 9 to tmf and proc . 2013 / 20358 - 7 to vb ) and nap biocomp ( university of são paulo ) for financial support and to thibaut de meulemeester and an anonymous reviewer for suggestions . dr . david de jong helped us to improve the english of our manuscript .

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lcc subject category : science : biology ( general ) : ecology | science : natural history ( general ) : general . including nature conservation , geographical distribution

k . vijayakumar ( kongu nadu arts and science college ) r . jeyaraaj ( kongu nadu arts and science college )

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department of zoology , kongu nadu arts and science college , g . n . mills , coimbatore , tamil nadu 641029 , india

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we are grateful to dr . deborah smith from university of kansas , usa for helping in identifying the stingless bee species and useful comments . i also express my sincere thanks to dr . m . muthuraman , for his encouragement and motivation to pursue the research work . i would also like to thank beekeepers mr . k . udayakumar , mr . g . k . thangavelu , mr . navaneethan from nellithurai village for their participation during sample collection and documentation .

stingless bees are a monophyletic group principally found in the tropical and subtropical areas of america , africa , australia and parts of asia ( roubik 1989 ) . the stingless bee species are taxonomically organized into

( michener 2013 ) . taxonomic identification of stingless bees remains unclear and requires experienced taxonomists . the total number of species is estimated to be about 500 described species worldwide ( michener 2013 ) and 43 species occur in the asian region ( michener 2000 ) .

is the single largest and most widespread genus in the indo - malayan regions , reported from india and extending to the solomon and caroline islands .

rasmussen 2013 ) and this species was originally described from ceylon by smith in the year 1854 .

in different parts of india are lacking . scarce literature is available on stingless bee species diversity in india . the distribution of

was reported in bengaluru , karnataka ( biesmeijer 1993 ) and in kerala ( mohan & devanesan 1999 ) .

the diversity of stingless bees in the indian subcontinent has been summarized by rasmussen ( 2013 ) .

) of stingless bees by using relative warp analysis of the forewings from india . the present study reports the morphological key characteristics of male and female bees of

most of the researchers in india concentrated only on stingless bee biology , morphometry , natural enemies and its pollination biology ( muthuraman & saravanan 2004 ; danareddi & viraktamath 2009 ; vijayakumar et al . 2012 ) . the studies on stingless bee species diversity in india are rare and our understanding of their identification is at an early stage .

53’6 . 24”e ) , tamil nadu , india ( image 1 ) and preserved in 90 % ethyl alcohol for further identification . taxonomic identification was based on the nest architecture and morphology according to sakagami et al . ( 1983 ) and schwarz ( 1939 ) . the preserved male and female stingless bees were dissected and the morphological characteristics of the worker and drone stingless bees were photographed by using a leica m165c stereo microscope with image analyzer . the male genitalia were cleared in a 0 . 5 % koh solution and examined in both wet and dry conditions . in addition , male sternum v and sternum vi were used as a key characteristic for distinguishing the

deborah smith , department of ecology and evolutionary , biology / entomology , the university of kansas , usa .

smith , depends strongly on the works of schwarz ( 1939 ) , sakagami ( 1978 ) and sakagami & inoue ( 1987 ) for the subgeneric characterization and for species recognition . these authors indicated that the size differences as well as male genitalia and sternal characters are important key morphological characteristics for identifying the species . the species descriptions are given below .

. this species has formerly been regarded as very widespread from india to solomon island . the “

species group is the largest and most widespread group in the indo - pacific areas .

the malar space is vestigial and the scutellum is extended backward . the mandible with two well developed teeth on the inner half of its apex . the basal sericeous area positioned more than half of the length of basitarsus .

the total body length ranges from 3 . 5–4 . 0 mm , and head width ranges from 1 . 50–1 . 68 mm ; forewing length includes tegula ranging from 3 . 2–3 . 9 mm , bifurcation between vein

ranges from 0 . 90–1 . 12 mm and hind tibial length ranges from 1 . 29–1 . 57 mm .

the entire body is black to blackish - brown . the clypeus , tegula , legs and metasoma are dark in colour ( image 2a ) . the frontal hairs are fulvous to whitish and plumose ( image 2b ) . the mesoscutal hairs well banded and fulvous to testaceous in colour ( image 2c ) . the mesoscutellar fringes are fulvous to testaceous . the erect hairs of the mesopleura are silvery to white in colour . the frontal hairs are mostly fulvous to whitish ( image 2d ) . the anterior veins and stigma of forewings are dark brown .

the hairs fringing on the hind tibiae posterior are plumose and the outer surface of hind tibia with dark brown stout setae ( image 2e ) . the hairs on upper surface of thorax is mostly light in colour and the hairs fringing the anterior contour of the hind tibiae are black ( image 2f ) . the basal and apical half of the wings are uniform in colour and the anterior veins and stigma dark brown ( image 2g ) . the number of hamuli on hind wing is constantly five per wing ( image 2h ) .

the structural characteristics are more similar to female bees ( images 3a–c ) . the posterior margin of basitarsus is imperceptibly angulate . the outer surface of the hind tibiae medially gently convex and apically slightly depressed ( image 3d ) .

the total body length ranges from 2 . 5–3 . 5 mm , head width ranges from 1 . 38–1 . 43 mm , forewing length including tegula ranges from 3 . 1–3 . 8 mm , bifurcation between m and cu ranges from 0 . 88–0 . 95 mm ; hind tibial length ranges from 0 . 96–1 . 23 mm .

the male stingless bees are similar to female bees in colour characteristics . the dense plumose hairs cover the posterior fringe of the mid tibia . the hairs on the outer surface of the hind tibiae are plumose and sparser ( image 3e , f ) . the sternum v ( s5 ) is small and the sternite vi ( s6 ) is antegladular area medially long and postgladular area short ( image 3g ) . the median depression of sternite vi with sparse hairs and the apex is narrowly and shallowly incised .

the male genitalia are one of the most important characteristics for taxonomic study . the gonostylus is long and slender , more or less sinuous with sparse hairs at apex , penis valve is very robust , tapering only at the apex , about as long as or slightly shorter than gonostylus .

stingless bees are a tropical group of over 500 species worldwide ( michener 2013 ) . recently , rathor et al . ( 2013 ) referred to seven stingless bee species in india . rasmussen ( 2013 ) listed eight species from the indian subcontinent and

is a wide spread species in india . he pointed out that the species of the

group are extremely similar in external morphology of the workers and a taxonomic revision of the species of india should include morphological characteristics of the male genitalia . the present study reports the key morphological characteristics of worker bees and male genitalia .

group is characterized by having a dark mesoscutum with four distinct hair bands separated by broad glabrous interspaces . the genital morphology and molecular data are needed for correctly describing the

group in nellithurai village , tamil nadu , india . the female bees were morphologically similar and are marked with minor differences in hairs on head and thorax , sereceous space and marginal cell in forewing . the male genitalia are the best diagnostic characteristics for differentiating within

is long and slender and sinuous with sparse hairs at apex . the penis valve is very robust and tapering only at the apex .

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the authors extend sincere thanks to dr . ( ms ) debjani dey , incharge , insect identification service , national pusa collection , division of entomology , iari , new delhi and dr . rajiv k . gupta , professor and former head , department of zoology , jai narain vyas university , jodhpur for confirming the identity of the bee specimens .

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pessarakli m ( 2016 ) handbook of cucurbits , 1st edn . crc press , boca raton , p 561

carpoglyphus robin , 1869 ( sometimes , the year is cited as 1860 , however , carpoglyphus robin , 1860 is not an available name ; robin , 1860 cites some collective characters for " carpoglyphus " and other genera of mites , which does not constitute a valid description , and , thus , the name carpoglyphus robin , 1860 ( nom . nud . ) is unavailable ( iczn art . 12 . 1 ) ) .

phoretic deutonymph : empodial claws arising from membranous ambulacra i - iv ( not from tarsal apices ) ( figs . 4 , 5 ) . leg iv with empodial claw present ( fig . 5 ) . leg iv generally similar in form to leg iii ( figs . 2 , 5 ) . posterior median ventral apodeme absent ( fig . 2 ) . ocelli present , widely separated on propodosoma ( fig . 3 ) .

adult : prodorsum with setae ve absent ( fig . 8 ) . prodorsal sclerite absent ( fig . 8 ) . setae vi situated about half - way between anterior edge of propodosoma and setae si ( fig . 8 ) . ocelli present ( fig . 8 ) . supracoxal gland opening not associated with a large sclerotized region ( fig . 8 ) . tarsi i - ii elongate ( fig . 11 ) . pretarsi similar on all legs ( fig . 11 ) . pretarsi with long , thin condylophores ( figs . 11 , 13 ) . empodial claws present ( figs . 11 , 13 ) . dorsal setae smooth , not heavily barbed ( fig . 6 ) . males without paranal suckers or sucker - like setae on tarsus iv ( fig . 13 ) . coxal apodemes i fused medially with coxal apodemes ii closing coxal fields i in both sexes ( fig . 9 ) . condylophores asymmetrical in male ( fig . 12 ) . male with genital setae ( g ) and coxal setae 4b present ( fig . 10 ) .

a dichotomous key to adults ( males and females ) is available in fain and rack , 1987 . this key can be used to identify the two species found in associations with bees : carpoglyphus lactis and carpoglyphus munroi . one non bee - associated species , carpoglyphus wardleorum , was described later ( clark , 2010 ) .

cosmopolitain . mites from honey bees have been reported from the holarctic , oriental , and australian regions .

feeding stages live in beehives of honey bees ( apis ) and probably in nests of stingless bees ( meliponini ) .

these mites live in a variety of habitats that often consist of sugary and fermenting materials . in honey bee hives , they feed on various components , including pollen stored in honeycombs ( bee bread or bee pollen ) , plant pollen , honey , and nest debris .

phoretic deutonymphs disperse on adult insect hosts to suitable habitats . phoresy is known on lepidoptera and coleoptera but not on bees .

carpoglyphus lactis is a relatively common species in beehives , occurring in the debris on the bottom boards of beehives , honeycombs , dead bees , honey , and especially on the bee bread ( bee pollen with added honey and bee secretions that is stored in brood cells ) . this mite can penetrate the brood cells and make burrows in the stored pollen , consuming it and causing the pollen and the debris to spill from the cells . the infested bee bread , mixed with large quantities of dead and live mites , turns to a golden - brown or yellow powdery material covering honeycombs and bottom boards of beehives . the mite damage is especially severe in stored overwintering nests . for example , 250 honeycombs were destroyed by these mites over one winter in a single storage area in germany ( zander , 1947 ) . a similar case was reported in the usa ( alabama ) , where stored honeycombs were found heavily infested after winter storage ( baker and delfinado , 1978 ) . weak bee colonies are more susceptible to mite attacks ( zander , 1947 ) , while healthy bee colonies usually can clean up the infested pollen ( baker and delfinado , 1978 ) .

aside from beehives , carpoglyphus lactis is also found in old honeycombs , wine barrels in cellars , dried sweet fruits , canned fruits , fruits preserved in sugar , fermenting pulp , dairy products ( milk and cheese ) , and stored honey . it often infests products following substantial development of yeast . in the field , this species is found in fermenting tree sap flows , burrows of moles , and as phoretic deutonymphs on butterflies , moths , and scarabaeid beetles ( e . g . , gnorimus ) .

when large numbers of c . lactis mites are ingested with infested food or beverages , they can cause dysentery . these mites also cause dermatitis to handlers of infested materials , such as dried plums ( o ' donovan , 1922 ) , and occupational allergies in the biological pesticide industry ( krop et al . , 2012 ) . however , they may be beneficial in the wine industry , as mite alarm pheromones enhance the aroma of pale and dry wines aged under flor yeasts ( marin et al . , 2009 ) .

the second mite species found in beehives , carpoglyphus munroi , is relatively rare and has been recorded from beehives from the czech republic .

fig . 5 . carpoglyphus lactis phoretic deutonymph legs iii - iv , dorsal view .

biotechnological innovations in aquaculture ( p . 01 - 06 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ga01

mangesh m . bhosale , r . r . mugale , pabitra barik , b . r . honnananda , h . k . vardia

four insectivorous birds in search of foraging niche in and around an agricultural ecosystem of nalgonda district of telangana , india ( p . 07 - 15 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra01

a virtual survey based debate on conservation strategies of indian giant flying squirrel ( petaurista p . philippensis ) ( p . 16 - 21 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra02

ectoparasites ( insecta and acari ) associated with bats in south and south - western caves of iran ( p . 22 - 28 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra03

orthoptera fauna , it’s habitat ecology and threats in barnawapara wildlife sanctuary , chhattisgarh , india ( p . 29 - 37 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra04

distribution and habitat preference of four - horned antelope ( chowsingha ) , tetracerus quadricornis in kumbhalgarh wildlife sanctuary , rajasthan ( p . 38 - 42 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra05

estimation of dusty days using the model of time series : a case study of hormozgan province ( p . 43 - 48 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . ra06

breeding biology of critically endangered long - billed vulture ( gyps indicus ) at a unique site in telangana state , india ( p . 49 - 51 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . nn01

hathipol : biodiversity of a tunnel ‘cave’ of chhattisgarh , india ( p . 52 - 54 ) doi : 10 . 21276 / ambi . 2016 . 03 . 1 . nn02

young ecologists talked and interacted in delhi ncr : culmination of yeti delhi 2016 / krem - puri : india’s longest sandstone cave explored ( p . 55 - 57 )

systematic numbers for monte carlo integration doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ga01

cloud computing : goals , issues , soa , integrated technologies and future - scope doi : 10 . 21276 / ambi . 2016 . 03 . 2 . rv01

ultrasound guidance in detection of pneumothorax and thoracentesis performance : a review doi : 10 . 21276 / ambi . 2016 . 03 . 2 . rv02

traditionally used medicinal plants in the treatment of kidney stone : a review on ethnobotanical studies in iran doi : 10 . 21276 / ambi . 2016 . 03 . 2 . rv03

best , useful and objective precisions for information retrieval of three search methods in pubmed and ipubmed doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ta02

preprocessing and optimization of smooth data - driven model for emergency conditions against air pollution doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ta01

a . ardalan , h . mohammadi , a . massah bavani , k . naddafi , m . t . talebian

one - dimensional transport simulation of pollutants in natural streams doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ta04

proposed suitable methods to detect transient regime switching to improve power quality with wavelet transform doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ta03

ahp - vikor bridge structural system selection in urban areas tehran : interchangescase study doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ta05

diversity and habitat association of birds in a vindhyan gorge of kekariya , rajasthan , india doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra01

cogitation on routine drowned cases in mazandaran province : a case study in between the year 2008 to 2013 doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra02

effect of glacial retreat on floral distribution and / or displacement in khersan glacier , central alborz mountain range , iran doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra05

fecal carriage of esbl types tem , shv , ctx producing genera proteus , morganella , providencia in patients of iran doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra06

diversity and habitat preference of odonata fauna ( insecta ) in kaziranga - karbi hills , central assam , northeast india doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra03

locating of rural health centers equipped with telehealth using gis : a case study on khorramabad city , iran doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra07

growth and phytoremediation potential of watercress nasturtium officinale r . br . in ammonium - rich wastewater doi : 10 . 21276 / ambi . 2016 . 03 . 2 . ra08

redescription of cadrema pallida var . bilineata ( de meijere , 1904 ) ( diptera : chloropidae ) and its role as pollinator and carrion feeder fromindiansunderbans doi : 10 . 21276 / ambi . 2016 . 03 . 2 . nn01

a report on resident , local migratory and migratory water fowl diversity in mayurbhanj district , odisha , eastern india doi : 10 . 21276 / ambi . 2016 . 03 . 2 . nn02

an analysis on views of iranian women about incentive policies on childbearing decision - making doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ga01

informationseeking behavior in blind people of iran : asurvey based onvariousexperiences faced by them doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ta01

assessment of human resources performance in high - tech aviation industries by dematel technique and fuzzy - analytic network process doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ta02

extracorporeal shock wave lithotripsy for the treatment of ureteral stones : a critiqueon success rate doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ta03

the role of cognitive dysfunction and behavioral activation system on life quality life of employees in iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ta04

cogitation on occupational stress and social support among hospitalnurses : acasestudyof zanjanprovince , iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra02

occurrence of cleft lip and palate in terms of maternal health , parents ' kinship , and neonateweight : a case study in the infants of southeastern part of iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra01

effect of mindfulness - based cognitive therapy on psychological sequels in hypertensive people ( hbp - patients ) doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra04

power of stressmanagement skill training on levels of depression , anxiety and stress in patients suffering from type - 2 diabetes doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra05

the role of motivational interviewing in depression and psychological well - being in mothers of students having learning disabilities doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra06

dental anxietyand pain perception related the appearance of dental injectors : a randomized clinical trial doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra07

womens psychological sexual disorder and hypertensive husbands doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra08

consequences of split shift work in indian traffic police personnel : day time sleepiness , stressors and psychological distress doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra09

pregnancy related anxiety questionnaire : reliability , validity and factor analysis doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra10

relation between emotion adjustment and perceived social support with quality of life of athletes with disability doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra11

personal and social predictors about safe sexual behavior inpatients with immune deficiency virus in ahwaz , iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra12

relationship between job characteristics model and learning organization : a case study of a ceramic and tile company of maybod city , iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . ra03

the relationship between lifestyle and pain in patients with spinal discherniation doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . nn01

a debate on haryngolaryngeal histopathologic examination of the cadaver suspected of pressure on vital elements of neck : death cause doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . nn02

occupational causes related male infertility : a case study in iran doi : 10 . 21276 / ambi . 2016 . 03 . sp1 . nn03

the effects of water crisis on food production in iran doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ga01

research and development on optimal expenditures evaluation and prediction : case study of irans agricultural sector doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ta01

anassessment of natural regeneration of non timber forest product ( ntfp ) species : a case study of mandla , india doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ta02

evaluating nitrogen efficiencies and accumulation in sugar beet ( beta vulgaris l . ) under tape - drip irrigation doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ra01

the assessment of sugar beet half - sib families based on some morphophysiological traits under drought stress conditions doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ra02

evaluation of ceres - rice model in simulation of rice growth under constraint irrigation and nitrogen fertilizer conditions doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ra03

phylogenetic analysis of medicagoid trigonella l . species based on its sequence data doi : 10 . 21276 / ambi . 2016 . 03 . sp2 . ra04

copyright © 2016 - 2018 national cave research and protection organization . all rights reserved

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virginie boreux , a , 1 cheppudira g . kushalappa , b philippe vaast , c and jaboury ghazoul a , 2

2 to whom correspondence should be addressed . e - mail : hc . zhte . vne @ luozahg . yruobaj .

edited by kenneth g . cassman , university of nebraska , lincoln , ne , and accepted by the editorial board september 18 , 2012 ( received for review june 22 , 2012 )

author contributions : v . b . , c . g . k . , and j . g . designed research ; v . b . performed research ; v . b . , p . v . , and j . g . analyzed data ; and v . b . and j . g . wrote the paper .

1 present address : institute of ecology , ecosystem functions , leuphana university , 21335 lüneburg , germany .

crop productivity is improved by ecosystem services , including pollination , but this should be set in the context of trade - offs among multiple management practices . we investigated the impact of pollination services on coffee production , considering variation in fertilization , irrigation , shade cover , and environmental variables such as rainfall ( which stimulates coffee flowering across all plantations ) , soil ph , and nitrogen availability . after accounting for management interventions , bee abundance improved coffee production ( number of berries harvested ) . some management interventions , such as irrigation , used once to trigger asynchronous flowering , dramatically increased bee abundance at coffee trees . others , such as the extent and type of tree cover , revealed interacting effects on pollination and , ultimately , crop production . the effects of management interventions , notably irrigation and addition of lime , had , however , far more substantial positive effects on coffee production than tree cover . these results suggest that pollination services matter , but managing the asynchrony of flowering was a more effective tool for securing good pollination than maintaining high shade tree densities as pollinator habitat . complex interactions across farm and landscape scales , including both management practices and environmental conditions , shape pollination outcomes . effective production systems therefore require the integrated consideration of management practices in the context of the surrounding habitat structure . this paper points toward a more strategic use of ecosystem services in agricultural systems , where ecosystem services are shaped by the coupling of management interventions and environmental variables .

ecosystem services are widely used as an economic argument for the conservation of natural habitats , including forests ( 1 ) . one such ecosystem service , crop pollination by animals , is thought to benefit 75 % of the major crops , representing 35 % of the world crop production ( 2 ) . pollination services have received particular attention in view of the declines in honey bees , a major crop pollinator ( 3 ) . a current concern is that a continued loss of honey bees ( and perhaps other bees ) may undermine crop pollination services and hence crop production worldwide ( 4 ) .

although insect pollinators have been shown to improve coffee crop productivity ( 5 ) , many studies have either limited the assessment of production to initial fruit set ( around 5 wk after pollination ) or overlooked the impact of environment and management system on production . assessment of fruit yield within a few weeks of flowering might be misleading as coffee has a 6 - to 10 - mo fruit maturation period . during this production cycle a range of management practices , including fertilization and pruning , are implemented that , together with environmental conditions , might play a major role in fruit development and early fruit loss . consequently , the early positive benefits of pollination for fruit set might have disappeared by fruit maturation and harvest ( 6 ) . in consequence , it is difficult to ascribe the degree to which pollination alone contributes to coffee production . at the same time , if pollinators do enhance initial fruit set ( as appears to be the case ) , then an improved understanding of management and environmental factors might allow pollination benefits to be retained through crop maturation . such information would facilitate the integration of pollination services within the management system , for instance through the provision of potential nesting sites and the management of alternative floral resources .

a further consideration for coffee farmers is whether interventions that maximize pollination service benefits ( e . g . , shade management ) incur trade - offs with other services or management priorities . it is thus conceivable that an intervention that increases pollination benefits might diminish production by , for example , reducing light or resource acquisition . a complete study of the potential benefits of pollination services to coffee production requires a more comprehensive evaluation of production at the time of fruit harvest , as well as an evaluation of the various trade - offs inherent to different management practices . ultimately , trade - offs should be presented in crop production or monetary terms , but here we seek to understand their ecological expression .

in this study , we investigated the potential role of pollinators in promoting coffee production in the context of soil and tree management practices as well as environmental variables such as soil fertility , shade cover , and rainfall .

agroforests in kodagu , south india , flowered between february 10 and march 20 , 2008 , with each agroforest flowering on a single day within this period . twenty - six farmers used irrigation to stimulate flowering ( irrigation is otherwise not used ) . two large postrainfall “mass flowering” events , covering nonoverlapping areas of 250 km 2 and 80 km 2 , occurred on february 19 and march 20 and included 75 and 16 agroforests , respectively . in the first region only four farmers had irrigated their crop before the february rainfall event . the intensity of the march flowering event was less than in february as many more agroforests had already been induced to flower by irrigation . all 19 agroforests that flowered between march 15 and march 31 received rain on the flowering day , which greatly reduced the number of insect visitors ( mean = 0 . 4 ± 0 . 3 se insects per observation as opposed to 6 . 4 ± 1 . 2 se insects per observation for agroforests that did not receive rain ) .

there was high variation in pollinator abundance , with 45 agroforests ( 40 % ) receiving no visitors at all during the observation periods . in such cases it was very obvious that there were almost no pollinators across the entire agroforest , and thus a “0” value is not simply an artifact of a limited observation time but is an accurate representation of extremely low bee visitation .

shade varied from 15 % in the most open agroforests to 76 % in the most shaded , with an average at 45 % . densities of grevillea robusta and native shade trees differed at plot and agroforest scales due to uneven distribution of trees within agroforests ( fig . s1 ) . on the basis of data collected from the 113 agroforests we estimated shade tree species richness across a range of agroforest areas ( fig . s2 ) .

widespread rainfall throughout the study region triggered simultaneous flowering across multiple agroforests , whereas irrigation allowed farmers to control the timing of coffee flowering , which was often at times when few other agroforests were flowering . bee abundance varied as a function of the number of agroforests flowering simultaneously (

) . agroforests flowering asynchronously via irrigation ( i . e . , only 1 , 2 or 3 agroforests flowering at any one time ) had significantly higher bee abundance ( 15 . 2 ± 3 . 0 bees ) than agroforests that flowered following rain ( 16 or 75 agroforests flowering concurrently ; 2 . 3 ± 0 . 7 bees ) ( kruskal–wallis test : χ

bee abundance as a function of the number of agroforests flowering on the same day . all agroforests in categories 1–3 were irrigated , and the number of each category signifies the number of agroforests in flower at that time . the numbers at top of the graph indicate how many agroforests flowered under such conditions : agroforests flowering alone ( category “1” ) numbered 13 , those flowering simultaneously with one other agroforest ( category “2” ) totaled 6 , and those flowering with two other agroforests ( category “3” ) totaled 9 . four of the 16 coflowering agroforests ( category “16” ) were irrigated . flowering in the 75 coflowering agroforests was stimulated by rainfall .

the “sign” column indicates the effect of the independent variable on bee abundance / presence .

the interaction between bee abundance and number of flowers had a positive impact on the number of coffee berries harvested . thus coffee production increased with increasing bee abundance , and this effect was amplified by an increase in the initial number of flowers (