in zoology , a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants .

nectar as a food source presents a number of benefits as well as challenges .

it is essentially a solution of ( as much as 80 % ) the simple sugars sucrose , glucose and fructose , which are easily ingested and digested , representing a rich and efficient source of nutrition .

this solution is often diluted either by the plant that produces it or by rain falling on a flower and many nectarivores possess adaptations to effectively rid themselves of any excess water ingested this way .

however , nectar is an incomplete source of nutrition .

while it does contain proteins and amino acids , these are found in low quantities , and it is severely deficient in minerals and vitamins .

very few organisms consume nectar exclusively over their whole life cycle , either supplementing it with other sources , particularly insects ( thus overlapping with insectivores ) or only consuming it exclusively for a set period .

many species are nectar robbers or nectar thieves , performing no pollination services to a plant while still consuming nectar .

nectar is produced by flowering plants to attract pollinators to visit the flowers and transport pollen between them .

flowers often have specialized structures that make the nectar accessible only for animals possessing appropriate morphological structures , and there are numerous examples of coevolution between nectarivores and the flowers they pollinate .

for example , hummingbirds and hawkmoths have long narrow beaks that can reach nectar at the bottom of long tubular flowers .

bats , meanwhile , visit open flowers where the nectar is not as deeply hidden .

nectarivore

nitrogen requirements of an old world nectarivore , the orange - tufted sunbird nectarinia osea .
nitrogen requirements of an old world nectarivore , the orange - tufted sunbird nectarinia osea . - pubmed - ncbi
nectarivore - ( from the greek work nektar , the drink of the gods ) a creature deriving its energy from the sugar rich nectar of flowering plants .
…that feed on nectar ( nectarivore s ) and have life spans longer than the flowering time of one plant species have mutualistic relationships with a succession of pollinating species in order to survive .
during the same time that a [ pis ] m [ ellifera ] adansonii encounters on flowers were registered , we noted the type of floral products collected by this bee . this parameter was measured to determine if a . m . adansonii is strictly a pollinivore , nectarivore or pollinivore and nectarivore . this could give us an idea of its implication as a cross - pollinator of g [ ossypium ] hirsutum .
citation : welch kc jr , allalou a , sehgal p , cheng j , ashok a ( 2013 ) glucose transporter expression in an avian nectarivore : the ruby - throated hummingbird ( archilochus colubris ) . plos one 8 ( 10 ) : e77003 . urltoken
colour me in . it ' s nectar of a new sort . nectarivore certifed organic ipa , a big hoppy , colourful cocktail of organic new zealand hops , pineapple sage and hibiscus . there ' s oodles of colour inside , but we ' ve left the outside to you . stay in the lines , or not - it ' s up to you . happy 25th birthday common sense organics .
another possible route for elimination of surplus water is through evaporation . evaporative water loss ( ewl ) of anna ' s hummingbird is high because of its high mass - specific metabolic rate , but high excretory losses mean that ewl still represents a far smaller proportion of daily water loss compared with other birds and mammals ( powers , 1992 ) . estimates calculated from the difference between water gain and cloacal fluid output have also yielded high ewl values for sunbirds and honeyeaters ( collins , 1981 ; fleming and nicolson , 2003 ; lotz and nicolson , 1999 ) . nothing is known of the partitioning of evaporation between cutaneous and respiratory routes , but cutaneous ewl is influenced by hydration state in other birds ( williams et al . , 2012 ) . unfortunately , the high excretory output of avian nectarivores complicates the direct measurement of ewl , and the pharmacokinetic method has proved unreliable for its estimation ( purchase et al . , 2013b ) . further research in this area is required to determine the role of evaporation in osmoregulation for the three avian nectarivore lineages .
sugar preferences and sucrase activity in nectar - feeding birds . ( a ) concentration - dependent sugar preferences of red wattlebirds , anthochaera carunculata ( fleming et al . , 2008 ) . birds were offered pairs of sucrose and hexose ( fructose + glucose ) solutions of varying concentrations from 0 . 075 to 2 mol l −1 sucrose equivalents ( se ) . diets where birds did not achieve energy balance are indicated with increasingly lighter shaded symbols . values are means ± 1 s . d . index of sugar preference ( i . e . hexose ingested as a proportion of total sugar ingested ) . asterisks indicate significant preferences for either hexose or sucrose diets ( * p < 0 . 05 ) and different letters indicate diets that were significantly different from each other in terms of the index of sugar preference ( anova and post hoc analysis ) . ( b ) relationship between degree of hexose preference ( i . e . minimum concentration where no sugar preference was indicated ) and standardized intestinal sucrase activity in 11 nectarivore species ( napier et al . , 2013 ) . data are average values for each species . white symbols denote diet generalists and grey symbols nectar specialists .
whole - kidney glomerular filtration rate ( gfr ; a measure of the rate at which the kidneys filter the plasma ) is much more responsive to hydration state in birds than in mammals , decreasing with dehydration or salt loading and increasing with water loading ( goldstein and skadhauge , 2000 ) . studies carried out in a range of bird species indicate that this is due to release of arginine vasotocin ( avt ) , which acts mainly on gfr , while having a relatively modest effect on the water permeability of the collecting ducts ( goldstein , 2006 ; nishimura and fan , 2003 ) . the reduction of gfr by avt occurs through a decrease in filtration by reptilian - type nephrons , with mammalian - type nephrons being less affected ( goldstein and skadhauge , 2000 ) . in nectarivores , plasma avt concentration increases with increasing sugar concentration , presumably reflecting a hormonal response to increased plasma osmotic concentration ( gray et al . , 2004 ) . surprisingly , in view of the high variation in water intake , gfr does not vary with water loading for the three avian nectarivore lineages ( goldstein and bradshaw , 1998 ; hartman bakken and sabat , 2006 ; purchase et al . , 2013b ) , although gfr is greatly reduced at night ( see below ) .
high water fluxes are a problem for salt conservation , and avian nectarivores are very efficient at minimizing electrolyte losses in the cloacal fluid : for example , whitebellied sunbirds feeding on 0 . 25 mol l −1 sucrose solutions void cloacal fluid with an osmolality of only 6 mosmol kg −1 ( fleming and nicolson , 2003 ) . despite similar abilities to produce copious volumes of dilute excreta across the three nectarivore lineages , the birds handle dilute diets differently . extremely dilute , sucrose diets devoid of electrolytes cause hummingbirds to stop feeding and go into torpor , while honeyeaters and sunbirds suffer decreased plasma sodium levels and are unable to maintain energy balance ( fleming et al . , 2004c ; goldstein and bradshaw , 1998 ; lotz and martínez del rio , 2004 ) . nectarivores thus appear to be constrained in their intake of dilute diets due to hyponatremia , i . e . low plasma sodium concentration ( fleming and nicolson , 2003 ) , and the addition of even small amounts of sodium ( 5–10 mmol l −1 ) to very dilute sucrose diets results in a marked decrease in plasma aldosterone concentration ( the active principle of the renin–angiotensin–aldosterone system , stimulating renal reabsorption of sodium ) ( fleming et al . , 2004a ) and enables whitebellied sunbirds and new holland honeyeaters to increase consumption , up to an extraordinary eight times their body mass daily ( purchase et al . , 2010 ) .
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parallel adaptations to nectarivory in parrots , key innovations and the diversification of the loriinae . - pubmed - ncbi
warning : the ncbi web site requires javascript to function . more . . .
parallel adaptations to nectarivory in parrots , key innovations and the diversification of the loriinae .
schweizer m 1 , güntert m 1 , seehausen o 2 , leuenberger c 3 , hertwig st 1 .
naturhistorisches museum der burgergemeinde bern bernastrasse 15 , ch 3005 , bern , switzerland .
aquatic ecology and macroevolution , institute of ecology & evolution , university of bern baltzerstrasse 6 , ch 3012 , bern , switzerland ; fish ecology and evolution , eawag seestrasse 79 , ch 6047 , kastanienbaum , switzerland .
department of quantitative economics , university of fribourg boulevard de pérolles 90 , ch 1700 , fribourg , switzerland .
specialization to nectarivory is associated with radiations within different bird groups , including parrots . one of them , the australasian lories , were shown to be unexpectedly species rich . their shift to nectarivory may have created an ecological opportunity promoting species proliferation . several morphological specializations of the feeding tract to nectarivory have been described for parrots . however , they have never been assessed in a quantitative framework considering phylogenetic nonindependence . using a phylogenetic comparative approach with broad taxon sampling and 15 continuous characters of the digestive tract , we demonstrate that nectarivorous parrots differ in several traits from the remaining parrots . these trait - changes indicate phenotype - environment correlations and parallel evolution , and may reflect adaptations to feed effectively on nectar . moreover , the diet shift was associated with significant trait shifts at the base of the radiation of the lories , as shown by an alternative statistical approach . their diet shift might be considered as an evolutionary key innovation which promoted significant non - adaptive lineage diversification through allopatric partitioning of the same new niche . the lack of increased rates of cladogenesis in other nectarivorous parrots indicates that evolutionary innovations need not be associated one - to - one with diversification events .
pmid : 25165525 pmcid : pmc4130445 doi : 10 . 1002 / ece3 . 1131
longitudinal section through the epithelium at the border ( arrow ) between the esophagus ( right ) and the proventriculus ( left ) of psittrichas fulgidus . the compound glands ( cg ) of the proventriculus can be clearly distinguished from the mucous glands ( mg ) of the lower part of the esophagus .
schematic illustration of a caudal view and a transverse section along the median plane between the tendineal centers ( right ) of the gizzard . the measurements taken in this study are indicated . modified from ziswiler ( ) .
schematic illustration of a transverse section through the gizzards along the median plane between the tendineal centres of a nectarivorous ( vini australis , left ) and a granivorous parrot ( neophema chrysostoma , right ) .
best - scoring maximum likelihood tree including bootstrap values above 70 % indicated at nodes . this tree was used for the formulation of the hypothesis of the variance - covariance matrix of the error terms in the regression analyses . nectarivorous species are shown in red .
natural - log transformed values of the different independent variables against natural - log transformed body masses including the regression lines of the best - fitting model . for all these traits , the data were best explained by an allometric relationship between the dependent and independent variables .
natural - log transformed values of the different independent variables against natural - log transformed body masses including the regression lines of the best - fitting models . for all these traits , a model including the nectarivorous diet as a covariate with different intercepts but the same slope was considered as the best - fitting model .
out of the 40 . 000 or more species of spiders found up to date , every single one of them is a predator , capturing their prey using a complicated web , hunting them directly , or using other creative methods ; all except one , that is . bagheera kiplingi ( odd choice for a veggie spider ) is the first species that primarily feasts on something else than meat .
the type of plants it enjoys is called beltian bodies , and it basically represents leaf - tip structures that are produced by acacia . what’s really interesting is that another animal that fancies the same food is the ant . some ants have an almost symbiotic relationship with the acacia shrubs , living in the hollow spines and eating an amount that also allows the acacia to thrive , and protecting the plants from other invaders . this relationship has been studied and described in numerous studies .
“this is really the first spider known to specifically ‘hunt’ plants ; it is also the first known to go after plants as a primary food source , ” said christopher meehan of villanova university , who took notice of the spiders during a field course in mexico .
even more , this species of spider is the first known to eat solid vegetative food at all . they do occasionally prey on small invertebrates , but in an overwhelming proportion , their food is vegetal .
“i’ve done the math several times , and even the most conservative estimates point to near - total vegetarianism , ” meehan said . “almost all of the prey that the spiders do eat are acacia - defending ant larvae . ”
“spiders aren’t really thought to be capable of eating solid food at all , ” meehan said . spiders digest their prey externally , he explained , and any matter bigger than about one micrometer gets filtered out of the vital juices in the spider’s pharynx . beltian bodies , however , are 80 percent structural fiber , and quite large by spider standards . “the acacia spiders consume these nutrient - enriched—but not particularly nutrient - rich—’vegetables’ completely , often in less than five minutes . ”
but eating the food of an ant colony… that’s definitely not something you should be eager to do , so how does the spider get away with it ? ? well , according to meehan , they have what can only be called “sheer wit” .
“jumping spiders in general possess incredibly advanced sensory - cognitive skills and eight - legged agility , and bagheera is no exception , ” he said . “individuals employ diverse , situation - specific strategies to evade ants , and the ants simply cannot catch them . ”
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avenues of water gain and loss in bees are examined here at two levels of organisation : the individual and the colony . compared with the majority of terrestrial insects , bees have a high water turnover . this is due to their nectar diet and , in larger species , substantial metabolic water production during flight , counteracted by high evaporative and excretory losses . water fluxes at the colony level can also be very high . when incoming nectar is dilute , honeybees need to remove large volumes of water by evaporation . on the other hand , water is not stored in the nest and must be collected for evaporative cooling and for feeding the brood . water regulation has many similarities at individual and colony levels . in particular , manipulation of nectar or water on the tongue is extensively used by bees to increase evaporation for either food - concentrating or cooling purposes .
bees are not normally model insects for studying water balance . bee physiology is characterised by endothermy in even small species ( stone and willmer , 1989 ) and is complicated by the social context . bees show the full spectrum of sociality : although most are solitary it is the social bumblebees and honeybees whose physiology and behaviour are most familiar to us ( willmer and stone , 2004 ) . in addition , the sophisticated thermal physiology of bees has received much more attention ( e . g . heinrich , 1985 ; kovac et al . , 2007 ) than their water regulation . here i briefly examine the water balance physiology of bees , considering the usual avenues of water gain and loss , at two levels of organisation : the individual and the colony , sometimes termed a ` superorganism ' ( moritz and southwick , 1992 ) . it is accepted that social homeostasis can be explained by the coordinated activity of bees as individuals , responding to their internal or external environment , without the need to invoke centralised control ( jones and oldroyd , 2007 ; seeley , 1995 ) .
water fluxes of honeybees can be very high at the colony level . for example , seeley estimated average annual requirements of 120 kg of nectar , 20 kg of pollen and 25 litres of water for a single wild colony ( seeley , 1995 ) . this applies to honeybees in cold temperate conditions , and the estimate for nectar includes substantial energy stores required for over - wintering . the seasonal cycles of african honeybees are limited by rainfall , not temperature , and better foraging weather means that in comparison massive honey stores are not needed ( hepburn and radloff , 1998 ) .
while male bees of most species take care of their own energetic needs , all female bees forage far beyond their individual needs , collecting pollen and nectar as provisions for the offspring they raise ; therefore activity patterns of the sexes may be very different ( willmer and stone , 2004 ) . where comparisons have been made , male xylocopa capitata and anthophora plumipes carry much smaller crop loads than do females ( louw and nicolson , 1983 ; stone , 1995 ) . the impermeable crop expands greatly for storage and transport of nectar and protects the haemolymph from osmotic shock . apis mellifera workers can carry close to their body mass in nectar , although crop loads tend to be much smaller ( schmid - hempel et al . , 1985 ) and are positively correlated with the nectar concentration ( hunt et al . , 1995 ) . this suggests that under natural conditions honeybees might carry less than during experiments in which they are fed concentrated sugar syrup ( e . g . park , 1932 ; waller , 1972 ) . their feeding system is designed to retain nectar in the crop for as long as possible . sugar leaves the crop at a rate dependent on the metabolic rate of the individual , but the fluid - emptying rate ( and the rate of rectal filling ) are inversely proportional to nectar concentration ( roces and blatt , 1999 ) .
female allodapula variegata concentrating the dilute ( 14 % ) nectar of aloe arborescens by evaporation . the regurgitated droplet , held under the tongue , is repeatedly sucked in and out and may be very large in relation to the size of the bee ( body length 7 mm ) . photo , michael ellis .
fresh pollen is relatively dehydrated after exposure at anthesis , but its water content increases after collection by bees due to the addition of nectar and glandular secretions ( human and nicolson , 2006 ) . the larval diets of bees vary greatly in water content ( roubik , 1989 ) . the water content of royal jelly is around 67 % ( wongchai and ratanavalachai , 2002 ) , but the larval food of stingless bees contains less water , a thicker consistency being necessary for larvae floating on top of mass provisions ( hartfelder and engels , 1989 ) . large carpenter bees provision their nests with semi - solid masses of pollen combined with nectar , giving a final water content of only 20 % in the provisions of x . capitata ( louw and nicolson , 1983 ) . in xylocopa mordax , nectar is pre - concentrated on the tongue for this purpose ( corbet and willmer , 1980 ) .
honeybees prefer sugar concentrations of 30–50 % ( sugar concentrations here are given as % w / w as in refractometer measurements ) under experimental conditions ( waller , 1972 ) , but in practice they collect from a much wider range of nectars . seeley measured 15–65 % in nectar loads being brought into a single colony ( seeley , 1986 ) , and hunt and colleagues recorded a similar range of concentrations in incoming loads ( hunt et al . , 1995 ) . the choice of nectar concentration depends on the ecological context , i . e . on the other food sources available at the time . this has long been a complicating factor in experimental studies of honeybee foraging behaviour . for example , lindauer found that the threshold sucrose concentration for eliciting recruitment behaviour declined from 55 % to 4 % as the german summer progressed ( lindauer , 1948 ) . empirical measurements of energy intake rate in bees show peak values at sucrose concentrations around 60 % in bumblebees , stingless bees and honeybees ( harder , 1986 ; roubik and buchmann , 1984 ) . note that for orchid bees ( euglossini ) , which use suction feeding rather than a lapping mechanism , optimal concentrations are lower and more dilute nectars are collected ( borrell , 2004 ) .
communal food storage requires that the osmolality of honey is high enough to inhibit microbial growth ( pusey , 1999 ) . in honeybees this is achieved first by hydrolysis of nectar sucrose to glucose and fructose , through the addition of hypopharyngeal gland enzymes , and then through evaporative processing by food - handling bees to reach a concentration of about 82 % . these bees evaporate nectar on their tongues before placing droplets in open cells for further evaporation , accelerated by fanning ( park , 1925 ) . among stingless bees , workers drink water condensed in the nest during honey ripening and regurgitate it outside the entrance ( roubik , 2006 ) . their ripened honey is around 70 % in concentration and tends to ferment ( cortopassi - laurino et al . , 2006 ; roubik , 2006 ) . note that uncapped honey is hygroscopic and absorbs water , so can be both a sink and source of water in the nest .
trophallaxis is the regurgitation of the crop contents of a donor bee for ingestion by receiver bees . extremely rapid distribution of incoming nectar was demonstrated by nixon and ribbands , who fed radiolabelled food to six foragers and were able to detect the label in 62 % of all foragers after only 4 h and in all large larvae in unsealed cells after 48 h ( nixon and ribbands , 1952 ) . trophallactic interactions ensure that homeostasis is achieved in the ` collective stomach ' of all workers , which is a nectar reserve for the colony ( schmickl and crailsheim , 2004 ) . similarly , colonies preparing to swarm store concentrated food in their crops , comprising 20–30 % of the mass of individuals and of the swarm ( combs , 1972 ) . in addition to its nutritional significance , liquid transfer between adults is a means of exchange of information about the quality and quantity of food reserves in the colony ( crailsheim , 1998 ) .
it is not clear whether solitary bees drink water for their own needs , as distinct from seeking dilute nectar . large aggregations of bees of various genera can be seen foraging at wet soil substrates in the tropics , then regurgitating and reimbibing fluid , and this may be a means of obtaining salts ( roubik , 1989 ) . although bumblebees are not expected to drink , marked bombus terrestris were observed drinking repeatedly from a water trough during warm conditions ( ferry and corbet , 1996 ) : it is unlikely that individual bumblebees would have a water deficit so this was probably for the benefit of the colony . when groups of 100 honeybees are confined in cages and provided with 67 % sugar , they drink about 10 μl of water daily at t a of 35°c and 40°c ( free and spencer - booth , 1958 ) .
honeybee colonies collect water for two reasons , related to different types of weather : for cooling of the brood area by evaporation on hot days , and for feeding the larval brood when foraging is limited on cool days ( lindauer , 1955 ; seeley , 1995 ) . the classic studies of lindauer showed how bees regulate the hive temperature in hot conditions ( lindauer , 1955 ) . water is collected by water foragers , then distributed around the hive and in cells containing eggs and larvae ; fanning accelerates its evaporation , as does regurgitation and evaporation on the tongue ( lindauer , 1955 ) . visscher and colleagues measured mean water loads of 44 mg in honeybees collecting water under desert conditions ( visscher et al . , 1996 ) . paper wasps and hornets also use water for cooling their nests , but the highly social stingless bees do not ( jones and oldroyd , 2007 ; roubik , 2006 ) .
the second need for water – for consumption by nurse bees when feeding the brood – is an aspect of water use by honeybees that tends to be underestimated ( johansson and johansson , 1978 ) . nurse bees feed young larvae a secretion from their hypopharyngeal glands ; for worker larvae after the third day this jelly is supplemented with honey and pollen ( crailsheim , 1998 ) . as already mentioned , the water content of royal jelly is high , so nurse bees have a great need for water when brood rearing is intensive ; this water cannot always be obtained from nectar .
the regulation of water collection in honeybees is discussed in detail by seeley ( seeley , 1995 ) . in essence , the rate of unloading of water foragers indicates the colony demand for water ( i . e . the feedback system is similar to that for nectar ) . in this way the balance between collection and consumption of water is maintained . importantly , water collection does not interfere with the collection of concentrated nectar by the colony ( kuhnholz and seeley , 1997 ) . the first bees to start water collection may be stimulated by the collective increase in crop sugar concentration of all bees in the nest , due to trophallaxis ( lindauer , 1955 ; seeley , 1995 ) , or possibly by the collective increase in haemolymph osmolality . apart from environmental factors , the tendency of honeybee foragers to collect water , nectar or pollen has a genetic component ( hunt et al . , 1995 ) . workers with the lowest sucrose response thresholds , i . e . those able to distinguish low sucrose concentrations from water in proboscis extension response tests , become water foragers ( pankiw and page , 2000 ) .
water foraging is regulated according to current demand and water is not stored in combs by temperate honeybee colonies : this is because nectar availability fluctuates widely and water sources usually do not ( seeley , 1995 ) . for african honeybees , occasional water storage has been recorded in wild bee nests , as after summer rain in the kalahari desert ( eksteen and johannsmeier , 1991 ) . park recorded temporary storage of water in the crops of ` reservoir bees ' ( park , 1923 ) .
studies on the mechanisms by which bees thermoregulate in flight have yielded data on the relative magnitudes of metabolic water gains and evaporative water losses for bees as individuals . metabolic water production by large flying bees ( xylocopa and bombus ) is substantial ( bertsch , 1984 ; nicolson and louw , 1982 ) , especially at low or moderate t a . honeybees ferrying large water loads to the hive at high t a produce enough metabolic water to offset their evaporative water losses ( louw and hadley , 1985 ) . however , it cannot be assumed that metabolic gains will balance evaporative losses in flight . because variation in metabolic heat production is used by bees ( apis and centris ) as a primary mechanism of thermoregulation in flight ( roberts and harrison , 1999 ; roberts et al . , 1998 ) , at high t a metabolic water production will decrease as evaporative water loss increases . for individual bees , the thermal environment is crucial to water balance in flight .
bees engaged in brood warming generate metabolic heat using their flight muscles . at low t a , the metabolic rates of incubating bumblebees are extremely high ( heinrich , 1974 ) . honeybee brood nest temperatures are maintained constant at about 35°c by bees that ` shiver ' on the comb surface or inside empty cells in the brood area ( kleinhenz et al . , 2003 ) , and high metabolic water production can be assumed during this energy - intensive heating activity . however , the general colony heat production to which all workers contribute does not require much increase in metabolism ( harrison , 1987 ) .
in flying a . mellifera and centris pallida there is a negative relationship between water balance and t a , the bees being in negative water balance at t a above 31°c ( roberts and harrison , 1999 ; roberts et al . , 1998 ) . although the cuticular permeability of both species increases with t a , neither cuticular nor respiratory water losses are sufficient to explain the very high evaporative water losses at high t a . these probably involve cooling mechanisms such as regurgitation of crop contents onto the proboscis ( heinrich , 1980 ) . this ` tongue lashing ' at high t a has been observed in a variety of bees ( roberts and harrison , 1998 ) and in honeybees has been shown to increase evaporative losses dramatically ( louw and hadley , 1985 ) . as a result , the thoracic temperatures of nectar and water foragers are significantly lower than those of pollen foragers ( cooper et al . , 1985 ; feuerbacher et al . , 2003 ) .
the amount of water that has to be evaporated from dilute nectar is enormous ( fig . 2 ) . in order to increase the sugar concentration from 20 % to 82 % , bees must evaporate 0 . 75 g water for every 1 g of nectar collected , and the mass of honey produced from a given mass of nectar is correspondingly reduced . recently we have shown that foragers of a . mellifera scutellata collecting dilute nectar of aloe greatheadii var . davyana in dry winter air begin to concentrate the nectar before returning to the hive ( nicolson and human , 2008 ) . because the crop is impermeable to both sugar and water , we can only explain the doubling of crop sugar concentration , from 20 % to 40 % , by evaporation on the tongue . this contradicts the conventional wisdom that the concentration of nectar is unchanged during its transport by bees between flowers and the hive ( park , 1932 ) . the advantage for the bees lies in reducing the water load that has to be carried and the amount of evaporation needed in the hive ( fig . 2 ) ; the cooling effect is less desirable for individual foragers but disturbances of heat or water balance can be corrected in the hive .
relative masses of sugar and water in nectars of various concentrations and in honey ( horizontal red line ) . the figure shows the nectar - processing advantage for apis mellifera scutellata workers concentrating the nectar of aloe greatheadii var . davyana from 20 % to 40 % on their tongues : two - thirds of the necessary evaporation is achieved before the bees return to the hive ( nicolson and human , 2008 ) .
the various processes involved in cooling the honeybee nest – collecting water , spreading it within the brood comb , and speeding its evaporation by fanning and regurgitation – have been mentioned above . this enables the temperature in the brood area to be precisely regulated at 35°c , but humidity in the hive is less constant ( human et al . , 2006 ) . air in the hive will generally be more humid than outside , as a result of transpiration of the inhabitants and evaporation during nectar flows . while high humidity is necessary for brood development , a dry atmosphere favours nectar ripening . we have measured absolute humidity ( thus excluding temperature effects ) in various regions of the hive , and found higher values in the brood area than in nectar stores , suggesting adjustments by the bees ( human et al . , 2006 ) . however , trade - offs with regulation of temperature and respiratory gases will disrupt the establishment of optimum humidity levels .
colony level respiration is important in social homeostasis . periodic synchronised fanning leads to tidal ventilation in honeybees and stingless bees when only one nest entrance is present ( moritz and crewe , 1988 ; southwick and moritz , 1987 ) . measurement of cyclic fluctuations in water vapour pressure and temperature at the nest entrance would enable estimation of evaporative water losses at the colony level .
the non - random disposal of the excreta of social insects , such as ejecting faeces late in larval development , is assumed to be for hygienic reasons ( weiss , 2006 ) . in honeybee larvae , the midgut–hindgut junction is occluded until the end of the larval stage , and defaecation coincides with cocoon formation , the excreta being incorporated into the structure of the cocoon ( jay , 1964 ) . healthy adult apis do not defaecate in the nest , even during overwintering in temperate climates ( but defaecation in or around the nest is a sign of infection with the midgut parasite nosema ) . defaecation flights occur when weather permits , and until then rectal fluid may be stored for prolonged periods , the distended rectum occupying much of the abdominal cavity ( fig . 3 ) . obviously an individual honeybee ' s water content fluctuates enormously depending on the volume of crop or rectal contents . an inverse relationship between crop and rectal volumes has been measured in honeybees confined for varying times after feeding ( roces and blatt , 1999 ) .
mass defaecation flights are conspicuous in tropical honeybees , and led to the ` yellow rain ' scare , due to the high proportion of pollen exines in the faeces ( mardan and kevan , 1989 ) . heat - shedding benefits have been suggested for the giant honeybee apis dorsata , which builds exposed combs and engages in mass flights involving half the colony , each bee jettisoning 20 % of its body mass ( mardan and kevan , 1989 ) . however , more comprehensive recordings for the same species show that mass flight activity is highest during maximum brood production , and timed so that brood temperature is minimally affected by the temporary absence of the protective curtain of bees ( woyke et al . , 2004 ) . water shedding may be a more important function of mass flights than heat shedding .
bees are less subject to desiccation than most terrestrial insects . this is because their nectar diet and high metabolic water production during flight frequently generate excess water . water fluxes in the honeybee colony are also high , due to honey ripening and periodic water demand for evaporative cooling and for feeding the brood . very importantly , the favourable microclimate created by the nest architecture and its densely aggregated inhabitants reduces evaporative water losses and provides a hydric and thermal refuge for returning foragers .
honeybee abdomen showing full crop ( a ) or full rectum ( b ) . adapted from plate 8 in dade ( dade , 1962 ) .
workers of apis mellifera do not forage for themselves and in social bees there is a blurring between the individual and colony in terms of water balance physiology . common to water regulation at both the individual and colony level is the regurgitation of nectar or water on the tongue for evaporative purposes . this is involved in water elimination from nectar both in the hive and during foraging , and water is evaporated in the same way to cool either the hive or the individual bee in flight . heinrich has previously drawn attention to the similarities in individual cooling , nest cooling and food storage behaviours ( heinrich , 1985 ) . perhaps these all originate in bubbling ( a term i prefer to tongue lashing ) , which appears to function as a nectar - concentrating mechanism in a variety of bees , and may have a profound influence on the water balance physiology of solitary bees in addition to social homeostasis . the high propensity of bees for regurgitation is important in both trophallaxis , which is not confined to highly social species ( kukuk and crozier , 1990 ) , and bubbling to evaporate water .
i thank julian dow for inviting me to contribute to this volume , connal eardley for identifying allodapine bees , and christian pirk for commenting on the manuscript . the paper is dedicated to simon maddrell and his enduring fascination with water in insects .
cooper , p . d . , schaffer , w . m . and buchmann , s . l .
) . pollination of the yellow passionfruit : nectar , pollen and carpenter bees .
cortopassi - laurino , m . , inmperatriz - fonseca , v . l . , roubik , d . w . , dollin , a . , heard , t . , aguilar , i . , venturieri , g . c . , eardley , c . and nogueira - neto , p .
feuerbacher , e . , fewell , j . h . , roberts , s . p . , smith , e . f . and harrison , j . f .
) . effects of nectar concentration and flower depth on flower handling efficiency of bumble bees .
) . the composition of larval food in stingless bees : evaluating nutritional balance by chemosystematic methods .
hendrichs , j . , cooley , s . s . and prokopy , r . j .
) . post - feeding bubbling behaviour in fluid - feeding diptera : concentration of crop contents by oral evaporation of excess water .
human , h . , nicolson , s . w . and dietemann , v .
hummon , a . b . , richmond , t . a . , verleyen , p . , baggerman , g . , huybrechts , j . , ewing , m . a . , vierstraete , e . , rodriguez - zas , s . l . , schoofs , l . , robinson , g . e . et al .
hunt , g . j . , page , r . e . , fondrk , m . k . and dullum , c . j .
johansson , t . s . k . and johansson , m . p .
kleinhenz , m . , bujok , b . , fuchs , s . and tautz , j .
kovac , h . , stabentheiner , a . , hetz , s . k . , petz , m . and crailsheim , k .
) . uber die einwirkung von duft - und geschmackstoffen sowie anderer faktoren auf die tänze der bienen .
moritz , r . f . a . and crewe , r . m .
moritz , r . f . a . and southwick , e . e .
) . the importance of osmosis in nectar secretion and its consumption by insects .
) . simultaneous measurement of evaporative water loss , oxygen consumption , and thoracic temperature during flight in a carpenter bee .
) . effect of nectar on microbial antagonists evaluated for use in control of fire blight of pome fruits .
roberts , s . p . , harrison , j . f . and hadley , n . f .
( hymenoptera : apidae ) and the ecology of nectar intake by bee colonies in a tropical forest .
) . inner nest homeostasis in a changing environment with special emphasis on honey bee brood nursing and pollen supply .
schmid - hempel , p . , kacelnik , a . and houston , a . i .
) . social foraging by honeybees : how colonies allocate foragers among patches of flowers .
southwick , e . e . and moritz , r . f . a .
) . warm - up rates and body temperatures in bees : the importance of body size , thermal regime and phylogeny .
visscher , p . k . , crailsheim , k . and sherman , g .
) . evaluating responses of honeybees to sugar solutions using an artificial - flower feeder .
) . behavioral , ecological , and physiological determinants of the activity patterns of bees .
woyke , j . , kruk , c . , wilde , j . and wilde , m .
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photo credit : t . - c . francis pan some oyster larvae grow faster than others , but now donal t . manahan and colleagues reveal that the fastest growers are marked out by their high protein synthesis rates .
“one of the underappreciated benefits of fellowships is the act of applying for them , because you have to write and articulate your ideas . ”
in our latest early - career interview , we chat to simon sponberg , assistant professor at the georgia institute of technology , usa . he shares the story of his career , beginning with how he combined a love of physics and biology by studying how geckos stick to walls .
a new review from craig p . mcgowan and clint e . collins looks at the ecology , biomechanics and evolution of bipedal hopping in mammals , with a focus on why bipedal hopping has arisen in multiple clades of mammals .
photo credit : ari friedlaender not all orca species prey on aquatic mammals , so how do delphinids know when they are at risk ? a new study shows that pilot whales and risso’s dolphins flee from a subset of orca calls that share acoustic characteristics with other mammal alarm calls , including human screams . this article was featured in science magazine .
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regulation of energy and water are by necessity closely linked in avian nectarivores , because the easily available sugars in nectar are accompanied by an excess of water but few electrolytes . in general , there is convergence in morphology and physiology between three main lineages of avian nectarivores that have evolved on different continents – the hummingbirds , sunbirds and honeyeaters . these birds show similar dependence of sugar preferences on nectar concentration , high intestinal sucrase activity and rapid absorption of hexoses via mediated and paracellular routes . there are differences , however , in how these lineages deal with energy challenges , as well as processing the large volumes of preformed water ingested in nectar . while hummingbirds rely on varying renal water reabsorption , the passerine nectarivores modulate intestinal water absorption during water loading , thus reducing the impact on the kidneys . hummingbirds do not generally cope with salt loading , and have renal morphology consistent with their ability to produce copious dilute urine ; by contrast , as well as being able to deal with dilute diets , honeyeaters and sunbirds are more than capable of dealing with moderately high levels of added electrolytes . and finally , in response to energy challenge , hummingbirds readily resort to torpor , while the passerines show renal and digestive responses that allow them to deal with short - term fasts and rapidly restore energy balance without using torpor . in conclusion , sunbirds and honeyeaters demonstrate a degree of physiological plasticity in dealing with digestive and renal challenges of their nectar diet , while hummingbirds appear to be more constrained by this diet .
nectar has been described as ‘the simplest food on earth’ ( martínez del rio et al . , 2001 ) , but nevertheless may pose significant physiological challenges for animals feeding on it . since we reviewed the topic a decade ago ( nicolson and fleming , 2003b ) , researchers in the americas , africa and australia have made substantial progress towards comparing the physiology of the three main lineages of specialized avian nectarivores : the hummingbirds ( trochilidae ) , sunbirds ( nectariniidae ) and honeyeaters ( meliphagidae ) ( fig . 1 ) . numerous other bird families show varying reliance on nectar ( nicolson and fleming , 2003b ; symes et al . , 2008 ) , but this commentary will focus on how the main radiations deal with the challenges of nectarivory , highlighting both convergence and differences in their responses .
the relatively low concentration of bird nectars [ compared with insect nectars ( pyke and waser , 1981 ) ] is the fundamental reason for the physiological problems associated with a nectar diet . flowers visited by hummingbirds and sunbirds produce nectar averaging 10–30 μl in volume and 15–25 % w / w ( 0 . 5–0 . 8 mol l −1 sucrose equivalents ) in concentration [ while flowers visited by generalist birds produce nectars that are even more copious and dilute ( johnson and nicolson , 2008 ) ] . nectar can also be diluted by rainfall events . the challenge of drinking dilute diets is that the birds must handle large volumes of preformed water , necessitating fast transit times ( i . e . time for food to pass through the gut , thus minimizing mass gain , which would interfere with flight ) , and yet maintain high assimilation efficiency , while also warming this large volume of fluid to body temperature . more concentrated nectars are likely to be less problematic , because switching between plants would enable birds to access more dilute nectar and thus balance water intake . the other major challenge is that nectar also contains very little by way of other solutes ; nectarivorous birds therefore have to deal with low electrolyte intake as well as low nitrogen , although their nitrogen requirements are relatively low ( tsahar et al . , 2006 ) .
it might be predicted that the physiological constraints of a ‘simple’ nectar diet would result in convergence in physiological responses of the different lineages of nectarivorous birds . however , recent findings indicate that this is not always the case and we now know that these birds show interesting differences in response to their nectar diets ( fig . 2 ) . in the first half of this commentary , we examine some common strategies for dealing with the challenges of a nectar diet . in the second half , we examine available data that indicate differences in how hummingbirds , sunbirds and honeyeaters deal with these challenges .
) , this field is receiving renewed attention , and not without controversy . rico - guevara and rubega (
) presented morphological measurements of hummingbird tongues and high - speed videos of drinking that showed that the tongue traps liquid as a result of surface tension ; this requires no energy expenditure by the bird and does not involve capillary action , which suggests that viscosity is not important . this finding was disputed by kim et al . (
) , who used a modeling approach to differentiate between different modes of nectar drinking , showing that the ‘capillary suction’ mechanism used by hummingbirds , sunbirds and honeyeaters is associated with optimal concentrations in the range 30–50 % ( 1 . 0–1 . 8 mol l
sucrose ; much higher than natural nectar concentrations ) . after further theoretical and experimental analysis of drinking in ruby - throated hummingbirds ,
the methods / techniques used to study the interactions between a substance and the body from administration to excretion ( absorption , distribution , metabolism ) .
in addition to concentration , nectar volumes are important for tongue loading . the volume of nectar determines its height in tubular floral corollas , and partial immersion of the tongue is common ( kim et al . , 2012 ) . when honeyeaters and hummingbirds feed from tubular artificial flowers , the amount of nectar loaded per lick increases with greater nectar volumes or shorter flowers , both of which increase the contact between tongue and nectar ( collins , 2008 ) . however , honeyeaters also commonly extract nectar from open , brush - type inflorescences and the effects of inflorescence shape on feeding rates have yet to be investigated .
specialized avian nectarivores from three lineages . ( a ) whitebellied sunbird , cinnyris talatala ( photo credit : rudi van aarde ) , ( b ) new holland honeyeater , phylidonyris novaehollandiae ( kathryn napier ) and ( c ) charming hummingbird , amazilia decora ( cole wolf ) .
some key physiological differences between the three specialist bird lineages . a representative hummingbird , honeyeater and sunbird ( note the differences in body size ) are shown feeding on nectar of an erythrina species , a major bird - pollinated genus that occurs in the natural range of all three lineages . all lineages show high rates of intestinal sucrase activity ( highest for the hummingbirds ; thicker yellow lines ) and rapid absorption of hexose sugars ( orange arrows ) , but the passerines are capable of shunting water directly through the gut ( blue dotted lines ) and therefore do not only rely on filtration by the kidneys ( blue solid lines ) . * hummingbirds show only a small medullary portion to their kidneys ( shaded green ) , which limits their abilities to produce hyperosmotic urine .
nectar - feeding birds are compensatory feeders over a wide range of nectar concentrations ( martínez del rio et al . , 2001 ; nicolson and fleming , 2003a ) . this means that they adjust their volumetric intake rapidly in response to changes in energy density ( fleming et al . , 2004b ; köhler et al . , 2008 ) . sugar concentration consequently has a marked effect on their sugar preferences . probably because of the rapid transit times and because hexose nectars do not require hydrolysis in order for the sugars to be assimilated , a hexose diet is preferred by both generalist and specialist nectar - feeding birds on dilute diets [ for references , see napier et al . ( napier et al . , 2013 ) ] .
interestingly , most species tested to date show a switch to no preference or even sucrose preference on concentrated diets ( e . g . data for the red wattlebird , anthochaera carunculata , are shown in fig . 3a ) . the concentration at which the switch occurs varies between species , and the minimum sugar concentration at which birds show no hexose preference is significantly correlated with intestinal sucrase activity for 11 bird species , even after phylogenetic correction ( fig . 3b ) ( napier et al . , 2013 ) . birds with no sucrase activity ( e . g . redwinged starlings , onychognathus morio ) or relatively low sucrase activity ( e . g . rainbow lorikeets , trichoglossus haematodus ) prefer hexoses at higher sugar concentrations , while birds with the greatest sucrase activity ( i . e . hummingbirds , but also sunbirds and honeyeaters ) either show no hexose preference or hexose preference on only the most dilute diets ( napier et al . , 2013 ) .


chalcosiinae is a subfamily of the zygaenidae , containing many species , mostly little known .

prominent sexual dimorphism , bright aposematic coloration and mimicry complexes are widespread .

chalcosiinae

no one has contributed data records for chalcosiinae yet . learn how to contribute .
chalcosiine day - flying moth caterpillar ( cyclosia midama , chalcosiinae , zygaenidae ) by sinobug ( itchydogimages ) on flickr . pu’er , yunnan , china see more chinese caterpillars on my flickr site here … . .
chalcosiine day - flying moth ( pidorus albifascia , chalcosiinae , zygaenidae ) by sinobug ( itchydogimages ) on flickr . pu ' er , yunnan , china see more chinese moths on my flickr site here … . .
chalcosiine day - flying moth ( retina sp . , chalcosiinae , zygaenidae ) by sinobug ( itchydogimages ) on flickr . pu ' er , yunnan , china see more chinese moths on my flickr site here … . .
chalcosiine day - flying moth , female ( soritia sp . , chalcosiinae , zygaenidae ) by sinobug ( itchydogimages ) on flickr . pu ' er , yunnan , china see more chinese moths on my flickr site here … . .
yen , s . - h . , g . s . robinson and d . l . j . quicke . 2005 . the phylogenetic relationships of chalcosiinae ( lepidoptera , zygaenoidea , zygaenidae ) . zoological journal of the linnean society 143 : 161341 .
yen , s . - h . , g . s . robinson and d . l . j . quicke . 2005 . phylogeny , systematics and evolution of mimetic wing patterns of eterusia moths ( lepidoptera : zygaenidae : chalcosiinae ) . systematic entomology 30 : 358397 .
lower ditrysian family groups : zygaenoidea s . l . ( especially chalcosiinae , procridinae and phaudidae of oriental / indo - australian regions ) , crambidae ( especially acentropinae ( = nymphulinae ) of asia and indo - pacifics ) , hyblaeidae , thyrididae ( strigulininae ) higher ditrysian family groups : uraniidae ( epipleminae ) , epicopeiidae , callidulidae , erebiidae ( syntomini ) . rhopalocera ( especially those are mimetic such as elymnias , penthema and dananinae )
now as ph . d . student of department of biology , imperial college of science , technology and medicine , university of london . ( supervisors : dr . donald quicke , imperial college and dr . gaden robinson , british museum ( natural history ) . thesis : phylogenetic reconstruction of chalcosiinae ( zygaenidae ) of the world , with systematic revision on tribal and generic levels . major interests in lepidoptera : phylogeny / evolution / biology / systematics of : primitive families : micropterigidae , paleosetidae , neopseustidaemonotrysian families : adelidae
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dried specimens and a small reverberation chamber ( fig . 1 ) were used to measure the random incidence absorption factors of the wings of each study species ( separately for male and female saturniids ) over the frequency range 20–100 khz ( fig . 2 ) . for each treatment n = 6 sets were used , with each set comprising the maximum number of non - overlapping moth wings of same - sex specimens of a species that could fit on the reverberation chamber floor . the number of wings of each set depended on species size and availability of dried specimens .
small reverberation chamber for the measurement of absorption factors in the ultrasonic spectrum . ( a ) experimental set - up . funnels direct the ultrasound emitted by the transducers into the chamber through a 7 mm opening . the microphone can move freely up and down inside the chamber , allowing for spatial averaging of the reverberation times . ( b ) overview of reverberation chamber showing the engraved diffusers . ( c ) absorption factor of empty reverberation chamber ( eqn 1 ) ; the error bars indicate 95 % confidence intervals ( cis ) ( n = 27 microphone positions ) .
absorption factors of moth wings measured with small reverberation chamber ( eqn 3 ) . ( a ) chalcosiines . ( b - d ) female and male saturniids . the absorption factors of a . mittrei and s . c . ricini differ significantly according to sex for the frequencies indicated by circles . ( e ) absorption factors grouped by preparation group . the absorption factors differ significantly from each other for frequencies within the range 20–40 khz . circles indicate significant difference between male and female saturniids , triangles between female saturniids and chalcosiines , and squares between male saturniids and chalcosiines . all pairwise comparisons were conducted with tukey - kramer test for each frequency separately . one , two , or three symbols correspond to 0 . 01 < p < 0 . 05 , 0 . 001 < p < 0 . 01 , and p < 0 . 001 , respectively . the error bars indicate the 95 % cis and n = 6 sets for each sample .
estimated marginal means of absorption factors of moth wings across the frequency range 20–100 khz . ( a ) estimated marginal mean of each preparation . there is a significant effect of preparation on estimated marginal mean ( anova , f 8 , 45 = 23 . 78 , p < 0 . 001 ) . ( b ) estimated marginal means grouped by preparation group . the estimated marginal means differ significantly according to preparation group ( anova , f 2 , 51 = 19 . 08 , p < 0 . 001 ) and also differ significantly from each other ( tukey - kramer test ) . the error bars show the 95 % cis [ n = 48 in a ( 6 sets×8 frequencies ) and n = 144 in b ( 3 preparations×6 sets×8 frequencies ) ] .
survival advantage of female s . c . ricini over males in terms of smaller detection distance by bats
the detection distance of a target by a bat depends on the transmission loss due to spherical spreading and atmospheric attenuation , and on the target strength ( møhl , 1988 ) . the detection distances of male and female s . c . ricini were compared because they have wings of similar size and shape ; consequently the comparison is feasible because their target strengths , and hence their detection distances , differ due to the different absorption factors of their wings alone . detection distances of moths typically range between 1–10 m ( surlykke et al . , 1999 ) . eqn 13 was used with three hypothetical detection distances within this range for male s . c . ricini in order to derive the respective detection distances of female s . c . ricini ( fig . 4 ) . at 25 khz , where the difference between the absorption factors of male and female s . c . ricini is maximized ( fig . 2 d ) , the detection distance of female s . c . ricini is 20–30 % smaller .
percentage difference in the detection distance of female s . c . ricini compared to three hypothetical detection distances of male s . c . ricini [ r ( 2 ) ] . the difference between the detection distances of male and female s . c . ricini is maximized at 25 khz , where the difference between the absorption factors of their wings is also maximized ( fig . 2 d ) . the detection distances of female s . c . ricini were calculated with eqn 13 for atmospheric attenuation at temperature 20°c and 70 % relative humidity ( surlykke , 1988 ) .
studies have reported bat echolocation calls with dominant frequencies ranging from 11 khz ( euderma maculatum ; fullard and dawson , 1997 ) to 212 khz ( cloeotis percivali ; fenton and bell , 1981 ) , though most insectivorous bats echolocate with dominant frequency 20–60 khz ( fenton et al . , 1998 ) . the ultrasound absorbance of the wings of this study ' s moth species peaks at the lower end of this range ( 20–25 khz ; fig . 2 ) . despite the similar patterns , there are significant differences not only between the nocturnal saturniids and the diurnal chalcosiines ( fig . 2 e ) , but also between male and female a . mittrei ( fig . 2 b ) and s . c . ricini ( fig . 2 d ) .
zeng et al . ( 2011 ) proposed that the scales on the wings are responsible for the ultrasound absorption . the scales have interstitial spaces between them that create a network of interconnected pores similar to that found in porous sound absorbers ( zeng et al . , 2011 ) ; when a sound wave propagates through this network , thermal and viscous effects cause the dissipation of its acoustic energy ( cox and d ' antonio , 2009a ) . in addition , the ultrastructure of the scales resembles a perforated panel backed by air ( zeng et al . , 2011 ) , which could act as a microperforated panel absorber ( cox and d ' antonio , 2009b ) .
male and female saturniid specimens ( a . io from usa , a . mittrei from madagascar , and s . c . ricini from china ) and male chalcosiine specimens ( a . a . analis and c . burmanus from thailand , and e . pulchera from burma ) were obtained from the lepidoptera breeders association ( bourne , uk ) .
the time period for which it takes the sound pressure level to drop 60 db after a sound stops is termed reverberation time and depends on the total absorption inside a reverberation chamber . consequently , by comparing the reverberation time before and after the introduction of an absorbent material ( e . g . moth wings ) inside the chamber , the random incidence absorption coefficient of the introduced material can be derived . the standard for the measurement of the random incidence absorption coefficient in a reverberation chamber essentially concerns frequencies below 20 khz ( iso , 2003 ) . for this frequency spectrum , the measurement procedure requires large chambers and 10 - 12 m 2 of absorbent material ( cox and d ' antonio , 2009c ) ; however , such a quantity of moth wings is practically infeasible . besides , the shorter wavelengths of the ultrasonic spectrum render feasible the measurement of the absorption coefficient with a smaller chamber and smaller quantities of absorbent material . still , since this study did not follow the standard faithfully , the measured absorption quantity has been called absorption factor . the same term was employed by the only other documented study of sound absorption in the ultrasonic spectrum ( zeng et al . , 2011 ) .
even with these measures , the reverberation time is spatially dependent within the reverberation chamber . to reduce the effect of non - diffuseness , two ultrasonic transducers equipped with funnels were utilised to direct the ultrasound into the chamber through 7 mm openings , scattering multi - directionally the acoustic wave that entered the chamber . in addition , the recording device , an ultrasonic microphone , could move freely up and down inside the chamber , allowing for spatial averaging of the measured reverberation times ( fig . 1 a ) .
the absorption factors of the empty chamber and of the moth wings were measured over nine 1 / 3 octave bands that have centre frequencies from 20 to 100 khz . the parameter
( s ) is the reverberation time after the introduction of the moth wings .
) . the wings of each set were placed on a white background of known surface area and an image was obtained . the image was smoothened with edge - preserving bilateral filter (
) . in the binary image , one class of pixels represents the wings and the other represents the background .
first , the recorded signal of main frequency f was filtered with a bandpass digital filter designed with a kaiser window . the filter passed frequencies between f l and f h in order to retain only frequencies within the 1 / 3 octave band , frequencies outside this range were attenuated 80 db ( fig . s2a ) .
the decay curve of the filtered signal has many fluctuations that render difficult the identification of the point where the response drops 60 db after the offset of sound ; therefore , the curve has to be smoothed . the first step of the smoothing process is to obtain the filtered signal ' s envelope by using the hilbert transform :
( s ) is the time point where the signal merges with the noise level . the parameter
is a maximally flat curve , meaning that its gradient is constantly negative . as a result , the curve crosses the −60 db point once . this property makes schroeder ' s integration method useful for the estimation of the reverberation time . however , in most cases l (
) > –60 [ db ] . this means that the signal merges with the noise level before reaching the −60 db point . therefore , the reverberation time has to be estimated using linear regression .
to compare the absorption factors among the nine preparations ( 3 chalcosiines and 3×2 male and female saturniids ; fig . 2 a - d and fig . 3 a ) , a repeated measures model , specifically a subject - by - treatment model ( doncaster and davey , 2007 ) , was fitted . in the model , the absorption factors are the responses , preparation is the between - subjects factor , which is used as the predictor variable , and frequency is the within - subject factor . anova was used to test if the absorption factors differ significantly according to preparation ( fig . 3 a ) , and repeated measures anova to test if there is a significant effect of frequency on absorption factor as well as significant preparation - frequency interaction . the p - values of the repeated measures anova were computed using a greenhouse - geisser correction ( ε = 0 . 79 ) because mauchly ' s test for sphericity indicated that that the sphericity , hence the compound symmetry assumption , does not hold ( χ 2 = 144 . 80 , p < 0 . 001 ) . to do pairwise comparisons between preparations , post hoc tukey - kramer tests were used ( fig . 2 a - d and fig . 3 a ) .
accordingly , a repeated measures model was used for the comparisons among the three preparation groups ( chalcosiines , male and female saturniids ; fig . 2 e and fig . 3 b ) , albeit with the preparation group as the between - subjects factor . the same tests were carried out as with the above model . again , the sphericity did not hold ( χ 2 = 142 . 61 , p < 0 . 001 ) , and a greenhouse - geisser correction ( ε = 0 . 81 ) was used for the calculation of the repeated measures anova p - values . all statistical analysis was conducted in matlab ( mathworks , uk ) , and a p - value of < 0 . 05 was considered statistically significant .
is the noise . the equation is in db form and all quantities are measured in db .
( m ) is the detection distance of the target ( e . g . moth ) , and
) is the atmospheric attenuation factor . the first term of the equation accounts for loss due to spherical spreading and the second one for loss due to atmospheric attenuation .
have wings of similar size and shape , their ratios of incident to returned sound intensity depend on the absorption factors of their wings alone . specifically , they are related as follows :
) , which in this study returns exactly one real solution . eqn 13 was used to compare the detection distances of female and male
we thank colleagues in the laboratories of the centre of ultrasonic engineering at the university of strathclyde for their advice and assistance during this work .
a . n . conceived the study , which was designed by a . n . , f . g . and j . f . c . w . experiments were carried out by a . n . and f . g . ; data analysis was performed by a . n . a . n . , f . g . and j . f . c . w . interpreted the findings . a . n . drafted the article . all authors read and approved the final version of the manuscript .
the research leading to these results has received funding from the european research council under the european union ' s seventh framework programme ( fp / 2007 - 2013 ) / erc grant agreement no . 615030 ; and was also supported by the engineering and physical sciences research council ( epsrc ) grant ep / l026511 / 1 .
all data created during this research are openly available from the university of strathclyde pure / knowledgebase at urltoken .
this is an open access article distributed under the terms of the creative commons attribution license ( urltoken ) , which permits unrestricted use , distribution and reproduction in any medium provided that the original work is properly attributed .
auditory influences on the flight behaviour of moths in a nearctic site . ii . flight times , heights , and erraticism
handbook of zoology / handbuch der zoologie . lepidoptera , moths and butterflies , morphology and physiology
auditory influences on the flight behaviour of moths in a nearctic site . i . flight tendency
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moths from yunnan , china click on and scroll through images for individual ids… . . by sinobug ( itchydogimages ) on flickr . pu ' er , yunnan , china see more chinese moths on my flickr site here …
the view from above - chinese caterpillars seen from the dorsal aspect click individual images for ids… . view all images in the the view from above series on my flickr here . by sinobug ( itchydogimages ) on flickr . pu’er , yunnan , china see more chinese caterpillars on my flickr site here … . .
this is a moth species that is quite common in hk . the species is distributed across se asia from thailand to china , but the hk population seems to be especially prominent .
safety in numbers - caterpillars ( click individual images for ids in captions ( where known ) ) by sinobug ( itchydogimages ) on flickr . pu’er , yunnan , china see more chinese caterpillars on my flickr site here … . .
red click on and scroll through individual images for ids in captions . by sinobug ( itchydogimages ) on flickr . pu ' er , yunnan , china see more chinese insects and spiders on my flickr site here ……
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the root of the current tree connects the organisms featured in this tree to their containing group and the rest of the tree of life . the basal branching point in the tree represents the ancestor of the other groups in the tree . this ancestor diversified over time into several descendent subgroups , which are represented as internal nodes and terminal taxa to the right .
you can click on the root to travel down the tree of life all the way to the root of all life , and you can click on the names of descendent subgroups to travel up the tree of life all the way to individual species .
for more information on tol tree formatting , please see interpreting the tree or classification . to learn more about phylogenetic trees , please visit our phylogenetic biology pages .
bode , w . and c . m . naumann . 1988 . little - known accessory glands in female zygaena moths ( lepidoptera , zygaenidae ) . zoological journal of the linnean society 92 : 2742 .
naumann , c . m . 1988 . the internal female genitalia of some zygaenidae ( insecta : lepidoptera ) : their morphology and remarks on their phylogenetic significance . systematic entomology 13 : 8599 .
naumann , c . m . , g . m . tarmann and w . g . tremewan . 1999 . the western palaearctic zygaenidae . apollo books , stenstrup .
niehuis , o . , c . m . naumann , and b . misof . 2006 . phylogenetic analysis of zygaenoidea small - subunit rrna structural variation implies initial oligophagy on cyanogenic host plants in larvae of the moth genus zygaena ( insecta : lepidoptera ) . zoological journal of the linnean society 147 : 367 - 381 .
niehuis , o . , s . - h . yen , c . m . naumann and b . misof . 2006 . higher phylogeny of zygaenid moths ( insecta : lepidoptera ) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence . molecular phylogenetics and evolution 39 ( 3 ) : 812 - 829 .
tarmann , g . 1994 . a preliminary review of the classification of the zygaenid subfamily procridinae ( lepidoptera ) . nota lepid . suppl . 5 : 115 - 123 .
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tree of life web project . 2006 . zygaenidae . burnet moths . version 10 august 2006 ( temporary ) .
each tol branch page provides a synopsis of the characteristics of a group of organisms representing a branch of the tree of life . the major distinction between a branch and a leaf of the tree of life is that each branch can be further subdivided into descendent branches , that is , subgroups representing distinct genetic lineages .
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no . 11 , alley 10 , lane 93 , shin - nan rd . , sect . 1 , chung - ho city , taipei hsien 235 , taiwan
born in november of 1971 in tainan city of taiwan . b . sc . of department of entomology , national chung - hsing university ( taichung , taiwan ) . m . sc . of institute of biological science , national sun yat - sen university ( kaohsiung , taiwan ) . thesis : phylogenetic analyses of the major lineages of nymphulinae and musotiminae , with taxonomic revision on taiwanese fauna ( lepidoptera , pyralidae s . l . )
awards : on 9th january 2004 dr . shen - horn yen received the 5th r . j . h . hintelmann wissenschaftspreis für zoologische systematik for his thesis
phylogenetic analyses of the major lineages of nymphulinae and musotiminae , with taxonomic revision on taiwanese fauna ( lepidoptera , pyralidae s . l . )
. the award was presented in a ceremonial act in zoologische staatssammlung münchen organized by the museum ' s management and the
freunde der zoologischen staatssammlung münchen e . v society
.
systematics , phylogenetics and biogeography of elymnias ( with mr . chia - hsuan wei and dr . david lohman )
evolution of mimetic wing patterns of the papilio memnon complex ( with dr . mathieu joron )
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marocaster is an extinct genus of sea stars in the family goniasteridae .

it existed in what is now morocco during the early cretaceous period .

it was described by daniel b. blake and roland reboul in 2011 , and the type species is m. coronatus .

marocaster

belongs to marocaster according to d . b . blake and r . reboul 2011
no one has contributed data records for marocaster coronatus yet . learn how to contribute .
sinopse : marocaster coronatus é um gênero de estrela - do - mar extinto . fonte : wikimedia commons - didier descouens . url : urltoken
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full reference : d . b . blake and r . reboul . 2011 . a new asteroid ( echinodermata ) faunule from the early cretaceous ( barremian ) of morocco . journal of paleontology 85 ( 6 ) : 1021 - 1034
type specimen : holotype mhnt . pal . 2010 . 2 . 2 , paratypes 2010 . 2 . 4a , 2010 . 2 . 5 , 2010 . 2 . 1h
parent taxon : goniasteridae according to d . b . blake and r . reboul 2011
eol content is automatically assembled from many different content providers . as a result , from time to time you may find pages on eol that are confusing .
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daniel b . blake & roland reboul ( 2011 ) .
a new asteroid ( echinodermata ) faunule from the early cretaceous ( barremian ) of morocco
.
this page was last modified on 20 april 2015 , at 22 : 23 .
this article ' s content derived from wikipedia , the free encyclopedia ( see original source ) .


eupithecia nigristriata is a moth in the geometridae family .

it is found in western china ( tibet , qinghai ) .

the wingspan is about 17 – 18 mm .

the fore - and hindwings are uniform pale brown .

eupithecia nigristriata

eupithecia lavicaria fuchs , 1902 ( syn : eupithecia lavicata prout , 1914 ) , described from norway .
ash pug ( angle - barred pug ) ( eupithecia innotata f . fraxinata )
eupithecia is a large genus of moths of the family geometridae . there are hundreds of described species , found in all parts of the world ( 45 in the british isles alone ) , and new species are discovered on a regular basis .
eupithecia species form the bulk of the group commonly known as pugs . they are generally small with muted colours and specific identification can be difficult . as a group they are easily identified by their narrow wings held flat at 90° to the body with the hindwings almost hidden behind the forewings .
the larvae of many species feed on the flowers and seeds of their food plants rather than the foliage . many species have a very specific food plant . some hawaiian eupithecia are predators of other insects ( e . orichloris , e . staurophragma , e . scoriodes ) . they mimic twigs but when sensitive hairs on their backs are triggered , they quickly grab the insects touching them . the defensive behavior of snapping may have pre - adapted hawaii ' s ancestral eupithecia for shifting to predation from feeding on pollen . also , insect predators that behave in this way are lacking in hawaii ' s fauna .
a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
the author ' s name is made up of a string of letters , with no blanks , and multiple authors ' names are separated by a comma . spelling of author ' s name is based on the original publication . if there are more than three authors , only the names of the first two authors are shown , followed by
, +
and the number of omitted authors .
attribute is enclosed in square brackets . this is rarely needed , but to differentiate homo - identifiers , this will contain the page , line or plate number of original publication .
all diacritic marks , hyphens , and apostrophes are eliminated , thus only the following characters are used : a to z , a to z , 0 to 9 , blank , comma , and opening and closing square brackets . although upper and lower cases are used for the convenience of human recognition , it is not case sensitive .
created by dicky sick ki yu 1997 - 2012 please send me information about errors and omissions ( contact information ) with supporting references , possibly with pdf or hard copy .
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to date 683 species of macro - moth along with 1159 species of micro - moth have been recorded in norfolk since records began in victorian times . this site aims to provide full details of all the species that occur ( or once occurred ) in norfolk , with photographs , descriptions , flight graphs , latest records , distribution maps and more !
if you have photos of any moths featured on this site , and would like them displayed along with your name and comments . . . please send them in ( any size . jpg images ) .
please consider helping with the running costs of norfolk moths . thank you : - )
underlying maps using digital map data © norfolk online lepidoptera archive - nola™ 2018 . © james wheeler - n o r f o l k m o t h s 2007 - 2018 . data © nola™ 2018
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chinery , michael ( 1986 ) . collins guide to the insects of britain and western europe ( reprinted 1991 ) .
skinner , bernard ( 1984 ) . colour identification guide to moths of the british isles .
this article was sourced from creative commons attribution - sharealike license ; additional terms may apply . world heritage encyclopedia content is assembled from numerous content providers , open access publishing , and in compliance with the fair access to science and technology research act ( fastr ) , wikimedia foundation , inc . , public library of science , the encyclopedia of life , open book publishers ( obp ) , pubmed , u . s . national library of medicine , national center for biotechnology information , u . s . national library of medicine , national institutes of health ( nih ) , u . s . department of health & human services , and urltoken , which sources content from all federal , state , local , tribal , and territorial government publication portals ( . gov , . mil , . edu ) . funding for urltoken and content contributors is made possible from the u . s . congress , e - government act of 2002 .
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. ” on his return to spain he found his old regiment about to march fo . . . . . . t imitate that life of mine when i in lonely sadness on the great rock
there to pine ; thou , . . . . . . what thou dost ; touch it not unless thou wouldst lay down thy life as the
of thy rashness . ” the car - rier gave no heed to these words ( and he . . . . . . be with you , and keep in mind what you have promised and sworn under those
that have been already declared to you . ” so saying , he gave rocin . . . . . . ut , as there might be some to be found among them that did not deserve the
of fire . “no , ” said the niece , “there is no reason for showing merc . . . . . . ou , master nicholas , i say let this and ‘amadis of gaul’ be remit - ted the
of fire , and as for all the rest , let them perish with - out further . . . . . . never slept a day under a roof , went to their graves as much maids as the
that bore them . i say , then , that in these and other respects our g . . . . . . ar , hatred nor love , should make them swerve from the path of truth , whose
is history , rival of time , storehouse of deeds , witness for the past . . . . . . ked plough had not dared to rend and pierce the tender bowels of our first
, belonging to a genus that feeds on feathers ; a beetle ( quedius ) and * . . . . . . brating so rapidly as to be scarcely visible , i was reminded of the sphinx
: their movements and habits are indeed in many respects very similar . . . . . . . than any other race of animals . i allude only to the butterflies ; for the
, contrary to what might have been ex - pected from the rankness of the . . . . . . ads . nothing could be more interest - ing than some of the family groups . a
with one or two daughters would often come to our rancho , mounted on . . . . . . manner in which his laws were enforced . one of these was , that no man , on
of being put into the stocks , should carry his knife on a sunday : t . . . . . . ate individual , but likewise used them , as old spain had done before for a
settlement . en - gland claimed her right an seized them . the english - . . . . . . yages of the adventure and beagle , is in lat . 46 degs . 50 ' , in the gulf of
. it is 15 miles long , and in one part 7 broad and descends to the sea . . . . . . an rafael . the posi - tion of the glaciers at this place and in the gulf of
may be put even in a more striking point of view , for they descend to . . . . . . in charge of this same fortress . after we left south america , he paid the
in the usual manner , by being con - quered , taken prisoner , and shot . . .
that 60 don quixote have been already declared to you . ” so sayin . . . . . . ut , as there might be some to be found among them that did not deserve the
copyright © world library foundation . all rights reserved . ebooks from project gutenberg are sponsored by the world library foundation , a 501c ( 4 ) member ' s support non - profit organization , and is not affiliated with any governmental agency or department .
here you will find one or more explanations in english for the word nigritaria . also in the bottom left of the page several parts of wikipedia pages related to the word nigritaria and , of course , nigritaria synonyms and on the right images related to the word nigritaria .
in turkey .
home of ichneumonoidea
. taxapad . * * * * y sick ki yu . 1997–2012 . retrieved . . .
this is the place for nigritaria definition . you find here nigritaria meaning , synonyms of nigritaria and images for nigritaria copyright 2017 © urltoken


polyrhachis gracilior is a species of ant found in the southwest and northeast india .

it is one of the few ants that build arboreal nests made of leaves stitched together using silk produced by their larvae .

originally described as a " race " of polyrhachis furcata , it was elevated to a full species by c t bingham who noted differences in the shape of the spines .

a species described from travancore as weberi by horace donisthorpe in 1943 was identified as being identical to gracilior by barry bolton .

polyrhachis gracilior

the above specimen data are provided by antweb . please see polyrhachis gracilior for further details
weberi . polyrhachis ( myrmhopla ) weberi donisthorpe , 1943b : 206 ( w . ) india . junior synonym of gracilior : bolton , 1974b : 174 .
polyrhachis gracilior is an endemic ant of the western ghats . they live in small colonies that nest between leaves stitched together with silk . the queen is not very different from the workers in appearance . this worker is cleaning her antennae .
{ author1 , author2 . . . } , ( n . d . ) . polyrhachis gracilior forel , 1893 . [ online ] india biodiversity portal , species page : { name of species field } available at : urltoken [ accessed date jul 10 , 2018 ] .
bolton , b . 1974b [ 1973 ] . new synonymy and a new name in the ant genus polyrhachis f . smith ( hym . , formicidae ) . entomol . mon . mag . 109 : 172 - 180 ( page 174 , senior synonym of weberi )
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forel , 1893e pdf : 25 ( diagnosis in key ) ( w . )
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from urltoken
in the figure caption . for websites , images must be clearly identified as coming from urltoken , with a backward link to the respective source page . see how to cite antweb .
antweb is funded from private donations and from grants from the national science foundation , deb - 0344731 , ef - 0431330 and deb - 0842395 . c : 0
dealate queen with brood within a disturbed nest . kerala , india . photo from kalesh sadasivan .
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
forel , 1893c : 25 ( diagnosis in key ) ( w . ) india . combination in
) : emery , 1925b : 193 . raised to species : bingham , 1903 : 388 . senior synonym of
karavaiev , v . 1927f . ameisen aus dem indo - australischen gebiet . iii . zb . prats zool . muz . 3 : 3 - 52 [ = tr . ukr . akad . nauk fiz . - mat . vidd . 7 : 3 - 52 ] . ( page 11 , worker , queen , male described )
santschi , f . 1928h . fourmis de sumatra , récoltées par mr . j . b . corporaal . tijdschr . entomol . 71 : 119 - 140 ( page 139 , replacement name : amboinae )
bingham , c . t . 1903 . the fauna of british india , including ceylon and burma . hymenoptera , vol . ii . ants and cuckoo - wasps . london : taylor and francis , 506 pp . ( page 388 , raised to species )
emery , c . 1925d . hymenoptera . fam . formicidae . subfam . formicinae . genera insectorum 183 : 1 - 302 ( page 193 , combination in p . ( myrmhopla ) )
forel , a . 1893e . les formicides de l ' empire des indes et de ceylan . part iii . j . bombay nat . hist . soc . 8 : 17 - 36 ( page 25 , ( diagnosis in key ) worker described )
this page was last modified on 6 july 2016 , at 02 : 20 .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
bharti h , guénard b , bharti m , economo ep ( 2016 ) an updated checklist of the ants of india with their specific distributions in indian states ( hymenoptera , formicidae ) . zookeys 551 : 1 - 83 . doi : 10 . 3897 / zookeys . 551 . 7052
enumerates geographic entities where the taxon lives . covers ranges , e . g . , a global range , or a narrower one ; may be biogeographical , political or other ( e . g . , managed areas like conservencies ) ; endemism ; native or exotic . does not include altitudinal distribution , which is covered under habitat .
if the taxon is in the ecological state of being unique to a defined geographic location , such as an island , nation or other defined zone , or habitat type , and found only there ; organisms that are indigenous to a place are not endemic to it if they are also found elsewhere .
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tiny ants living in acorn gutting a cricket - temnothorax cf . nylanderi ( 720p )
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .