peters ' elephant-nose fish ( gnathonemus petersii ; syn . gnathonemus brevicaudatus pellegrin , 1919 , mormyrus petersii günther , 1862 ) is an african freshwater elephantfish in the genus gnathonemus .

other names in english include elephantnose fish , long-nosed elephant fish , and ubangi mormyrid , after the ubangi river .

as the latin name petersii confirms it is named after someone called " peters " ( probably wilhelm peters ) , although the apostrophe is often misplaced and the common name given as " peter 's elephantnose fish " .

it uses electrolocation to find prey , and has the largest brain-to-body oxygen use ratio of all known vertebrates ( around 0.6 ) .

peters ' elephantnose fish

to find a peters’ elephantnose fish , you must lurk in muddy , slow - moving water . look closely , because the fish is brown and so is the background .
the retina of the peters’ elephantnose fish looks completely different . it is covered with cup - shaped depressions . around 30 cones sit inside each cup , and a few hundred rods are buried underneath .
peters’ elephantnose fish has no such limitations . its peculiar eyes allow it to use the two types of receptor at the same time . that could help it to spot predators as they approach through the murky water it calls home .
black or dark brown in colour with attractive white stripes across the tail , elephantnose fish are smooth - skinned . like many scaleless fish they can be
if your elephantnose fish slides its proboscis across your other fish from time to time , don ' t worry - it isn ' t trying to eat them .
however , the peters’ elephantnose fish were very good at spotting large moving objects against a cluttered background – essential for fish that live in dirty water . presented with a monitor displaying a black stimulus on a white background , they took as long to spot it as goldfish . but when a grey noise pattern – like an untuned tv – was superimposed , the elephantnose fish spotted the stimulus faster than the goldfish .
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the elephantnose fish , or peter ' s elephant , are very unique fish that not many hobbyists can say they have . elephantnose fish are thin and oblong , primarily dark brown or gray with white markings and a long trunk - like nose , thus the name , elephantnose . elephantnose fish are shy , sensitive fish , who need pristine water conditions to thrive . they are , in fact , so delicate , that they are used at government water departments in the usa and germany to test the quality of the water . whether this is humane or not is debatable ; the point is that an elephantnose needs to be added to a fully cycled , mature tank .
peters rc , denizot jp . miscellaneous features of electroreceptors in gnathonemus petersii ( günther , 1862 ) ( pisces , teleostei , mormyriformes )
what the bonn team found was that the elephantnose can switch between these two senses despite having a tiny brain .
peters’ elephantnose fish belongs to a large family called the elephantfish , all of which live in africa . they get their name from the trunk - like protrusions on the front of their heads . but whereas the trunks of elephants are extensions of their noses , the trunks of elephantfish are extensions of their mouths .
, peters ' elephantnose fish are second to none . their fascinating appearance , with the lower lip extended like a long snout , is accompanied by a high degree of intelligence and intriguing behaviour . although they can be shy , patience and care will encourage them to be friendly , active participants in the aquarium community .
according to bonn , statistical analyses showed identical performance as they swapped between vision and electrical impulses . also , they did it with a consistency that indicates that all elephantnose fish share this talent .
peters ' elephantnose fish ( gnathonemus petersii ) is common to the slower rivers of west africa , where it hunts for insect larvae just before dawn and after sunset . it ' s also one of a number of fish equipped with an electrical organ that allows it to generate a weak electrical field that allows it to sense its immediate environment like a living motion detector . it also has eyes that it uses to navigate as well as to track and catch its supper .
as to why the elephantnose can do this , the bonn team hasn ' t speculated , but it could be that the fish uses a specialized algorithm or processing system , the job of sense - swapping is less complex than believed , or fish are just really clever .
peters’ elephantnose fish are native to the rivers of west and central africa , in particular the lower niger river basin , the ogun river basin and in the upper chari river . it prefers muddy , slowly moving rivers and pools with cover such as submerged branches . it is a dark brown to black in colour , laterally compressed ( averaging 23–25 cm ) , with a rear dorsal fin and anal fin of the same length . its caudal or tail fin is forked . it has two stripes on its lower pendicular . its most striking feature , as its names suggest , is a trunk - like protrusion on the head . this is not actually a nose , but a sensitive extension of the mouth , that it uses for self - defense , communication , navigation , and finding worms and insects to eat . this organ is covered in electroreceptors , as is much of the rest of its body . the elephantnose fish has poor eyesight and uses a weak electric field , which it generates by muscular contractions , to find food , to navigate in dark or turbid waters , and to find a mate . peters’ elephantnose fish live to about 6 - 10 years , but there are reports of them living even longer .
peters’ elephantnose fish is probably the most commonly - available mormyrid in aquarium stores in the usa . in the aquarium ( which should be at least 200 liters ) it is timid , preferring a heavily planted environment with subdued lighting . ideally , a pipe or hollow log should be provided . the substrate should ideally be soft sand to allow the fish to sift through it with its delicate extended lip . it feeds on small worms ( bloodworms ) and aquatic invertebrates such as mosquito larvae , but in the aquarium will usually accept frozen or even flake food . how peaceful an elephantnose fish is can depend on the individual ; some are quite aggressive with other species , while others are retiring . they may be kept in a community aquarium with peaceful species who share their water preferences . however , unless kept in an aquarium of over 400 liters , it is unwise to keep more than one elephantnose fish as they can be territorial . the conditions suggested to keep them in an aquarium are as follows : ph of 6 . 8 to 7 . 2 , water temperature 26 to 28 degrees celsius , and water of medium hardness . the substrate should always be something that does not irritate the sensitive snout of the fish .
the weak electrical impulses generated by this fish can be made audible by placing two electrodes in the fish tank , which are then hooked up to an audio amplifier or a piezoelectric earbud . the sonar - like clicks that this fish emits can sound like a squeaky door when the fish is excited . – wikipedia
hello , i have an elephant nose fish just the one , they are fine in groups when young but as soon as they reach sexual maturity they will not tolerate there own species . in my experience these fish are great community fish best kept with medium to large species like the south - american cihlids , clown loaches , botia , larger tetra ' s and angel fish . however these fish are slow growing and nocturnal but these fish are very elegant and a very intelligent fish species . the elephant nose fish has an electric force - field depending on their mood so these fish can ' t be kept with any other fish with an electric force - field like the black ghost knife fish . elephant nose fish are not for the beginners of fish keeping , they are somewhat sensitive to water conditions and will be very unhappy and lethargic in anything above ph7 . i keep my elephant nose fish in ph 6 . 8 which is spot on in the middle as the ph for these fella ' s ranges form 6 . 5 - 7 . i have been keeping fish for nearly 14 years now , every year has been a joy . i love keeping these lovely elegant fella ' s and have don ' t for many years now , so good luck with your elephant nose ' s , hope all is good .
- medium / difficult . this fish needs perfect water and is somewhat intolerant to medication .
hopkins cd , bass ah . temporal coding of species recognition signals in an electric fish .
ciali s , gordon j , moller p . spectral sensitivity of the weakly discharging electric fish
maler l . the posterior lateral line lobe of a mormyrid fish - a golgi study .
. far gentler than that generated by a stingray or electric eel , this is used not for defence but for navigation , for finding food , and for finding a mate . male and female elephantnose fish have distinctly different electrical signatures but these can be distorted in the aquarium , which may be why they have never bred in captivity .
fish may be smarter than we thought . not only can some recognize human faces , but others can use their senses in a way that it was believed only humans and other mammals could manage . a team of zoologists at the university of bonn has discovered that , despite lacking a complex brain , the african elephantnose fish can swap between its electrical and visual senses in the same way a person can switch between sight and touch .
schwarz s , emde g von der . distance discrimination during active electrolocation in the weakly electric fish
kramer b , tautz j , markl h . the eod sound response in weakly electric fish .
arnegard me , carlson ba . electric organ discharge patterns during group hunting by a mormyrid fish .
so that you can supply them with an extra meal at dusk when other fish are less active .
. with other types of fish they tend to be shy , though individual temperament varies a lot .
emde g von der . discrimination of objects through electrolocation in the weakly electric fish , gnathonemus petersii .
grau hj , bastian j . neural correlates of novelty detection in pulse - type weakly electric fish .
szabo t , hagiwara s . a latency change mechanism involved in sensory coding of electric fish ( mormyrids )
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electric fish navigate and explore their dark and turbid environment with a specialised electric sense . this active electrolocation involves the generation and perception of an electric signal and fish have proven to be useful model systems for the investigation of sensory - motor interactions . a well studied example is the elephantnose fish , gnathonemus petersii , which has a characteristic and unique elongated chin covered with hundreds of electroreceptor organs . this highly moveable so - called schnauzenorgan constitutes the main fovea of the active electrosensory system . here we present first evidence for a sensory - motor loop relating active electrical sensing to active motor exploration of the environment .
people and animals can navigate a darkened room by touch , hearing , or even smell , and then switch immediately to sight when the lights come on without having to reassess their surroundings . this was thought to require the cerebral cortex of the mammalian brain , but the elephantnose fish doesn ' t have a cerebral cortex and seems to manage it like a zx - 80 running windows 10 .
the fish’s ability to see the wood for the trees probably helps it spot incoming predators like catfish . so reichenbach thinks its oddball visual system isn’t a mistake . “it’s the right type for this fish , ” he says .
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emde g von der . non - visual environmental imaging and object detection through active electrolocation in weakly electric fish .
emde g von der , zelick r . behavioral detection of electric signal waveform distortion in the weakly electric fish ,
the ability to recognize objects across different is senses is regarded as an evolutionary plus because it provides the animal with greater sensory flexibility in different environments . however , it ' s been thought that this ability was confined to certain highly developed mammals , including monkeys , dolphins , rats , and humans , because the information processing was thought to require a cerebral cortex . that is , until the elephantnose fish came along .
s ' , you can use this method from time to time to check on your fish ' s well being .
szabo t . sense organs of the lateral line system in some electric fish of the gymnotidae , mormyridae and gymnarchidae .
emde g von der , bleckmann h . finding food : senses involved in foraging for insect larvae in the electric fish ,
post n , emde g von der . the
novelty response
in an electric fish : response properties and habituation .
hall c , bell c , zelick r . behavioral evidence of a latency code for stimulus intensity in mormyrid electric fish .
engelmann j , pusch r , emde g von der . active sensing : pre - receptor mechanisms and behavior in electric fish .
also known as peter’s elephantnose , this fascinating species is currently the most common mormyrid in the hobby . sadly , its popularity means it is often kept in unsuitable conditions . to deny the fish a soft substrate really is cruel , as it uses its “trunk” to locate food hidden in the substrate . it also needs dim lighting , and will commonly become withdrawn and pine away if kept under bright lights . if you cannot provide suitable lighting , add food to the tank at lights out to allow the fish to feed , along with plenty of cover for them to hide in during tanklight hours .
like other mormyrids , it produces a weak electric field using specially - adapted muscle tissue located towards the tail . it also possesses electroreceptors which allow the fish to receive electrical signals . with these , the fish can sense the tiniest of movements as the field around it is disturbed , and navigate in total darkness , useful skills for a poorly - sighted fish when hunting prey or avoiding predators in the gloom . what is most fascinating about this adaptation is that the fish also use it to communicate with each other and find partners .
elephantnoses are a member of the mormyridae family , or electric fish . they emit small electrical pulses from an organ located inside their nose . the pulses will change with the mood of the fish . female elephantnoses have shorter pulses , males longer . they also have very poor eyesight , so this organ helps them navigate around the tank . this slight electricity should not hurt the other fish in the tank .
the fish were started in the experiments with the lights on , where they could use both senses , then in the dark . with the lights off , the team could see that the fish found their goal , but only at short range – indicating that it was using its electrical sense . the scientists then made the objects electrically invisible , so the fish could use only sight and repeated the experiment .
overall the color of the fish is a dark brown or black color . there are a pair of somewhat yellow vertical bands that extend from the back of the dorsal to the anal fin . not a stunning fish as far as color goes , but nice in its own right .
. they grow up to twenty four centimetres in length but can live happily alongside peaceful fish of anything down to a third of their size .
twenty - four fish ( gnathonemus petersii , günther 1862 ) , all purchased from aquarium glaser ( rodgau , germany ) , were used in this study . fish were housed in two 100 litre tanks of constant temperature ( 22 - 24°c ) and electrical conductivity ( 100 to 120 μs / cm ) .
elephantnoses are very timid and nocturnal . a very well planted tank with multiple hiding places in the form of driftwood , pots and pipes is a must , otherwise your elephant nose will be very stressed . soft gravel is also necessary , so they don ' t damage their sensitive noses . hobbyists have reported that these fish only swim in the open when all lights in the room are completely off , yet others say that their fish are very active , and will swim around their owners ' hands during water changes , so it ' s sort of hit or miss . the fish will be much more friendly , however , once comfortable in its surroundings . one tip for viewing your elephant nose during the day is to purchase some see - through plastic pipes , and put a few plants in front . that way the elephantnose can feel secure , and you can view them .
, though they can share that space with other types of fish . they are best kept either singly or in groups of three or more ; keeping two will
i have not been so lucky as everyone else . i have tried twice now to have the elephantnose in my 55g tank with no success . they have lasted 2 to 3 days and wind up dead in the bottom of my tank . all the water levels are where they should be and i have been feeding them at night ( frozen brine shrimp ) , plenty of cover . . . i see where some recommend 3
wöhl s , schuster s . the predictive start of hunting archer fish : a flexible and precise motor pattern performed with the kinematics of an escape c - start .
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the fish has paid a price for its electrical sensitivity . processing the signals takes brainpower , so it has an exceptionally large brain . as a result , 60 per cent of the oxygen taken in by the fish goes to its brain . even humans , with our whopping brains , only devote 20 per cent of our oxygen to them .
emde g von der , ronacher b . perception of electric properties of objects in electrolocating weakly electric fish : two - dimensional similarity scaling reveals a city - block metric .
hollmann m , engelmann j , emde g von der . distribution , density and morphology of electroreceptor organs in mormyrid weakly electric fish : anatomical investigations of a receptor mosaic .
to study this phenomenon , the zoologists took 10 fish and taught them how to recognize objects using both their visual and electrical senses . they did this using an aquarium separated into two chambers that held a sphere and a cuboid object . each fish was introduced to the aquarium and taught to select one or the other object with rewards of larvae .
the unusual appearance and novelty of the elephant nose has increased its popularity in recent years . if the proper care is taken they will be a rewarding member of your fish community .
these experiments were conducted in a separate tank ( 27 × 13 × 8 cm 3 , 100 μs cm - 1 ) with fish that were curarized and artificially respirated . fish initially were anaesthetised ( ms - 222 , 306 . 17 μmol l - 1 ) followed by an injection of 30 μl pancuronium ( solution 4 mg pancuroniumbromid per 2 ml , thinner 1 : 400 , organo teknika ) . afterwards the fish was rigidly placed in the tank where it was connected to an artificial respiration system . recording technique and animal handling was similar to that described elsewhere [ 51 ] .
this is one of the most interesting fishes . very exciting to watch . they are freindly with all . the nose bends around while searching for food . one of few fish that swims backwards often .
while our systematic mapping of the response was restricted to the head region of the fish , experiments with the objects placed at the trunk of the fish never evoked sors . in these experiments , the trunk was only partially covered . we therefore conclude that particularly the schnauzenorgan is the focal region for evoking sors , but can not exclude that sors might also be evoked form other regions in freely behaving animals .
i bought an elephantnose about two months ago . i built a cave for him out of some flat rocks that i found in lake ontario , but he refuses to come out of it unless it is completely dark in the room . at this moment in time he is about 13 cm long . they also have some electrical organs in their tails . it ' s too weak to affect any fish at all , they can ' t even feel it . i don ' t know why they are there , but i just thought it would be cool to know . they eat blood worms , it ' s best to feed them after you turn off the lights or they won ' t get the food and they ' ll starve to death .
hi ! i have 5 elephant nose fish in a 150 gal . tank . it shares its space with 5 cichlids , 1 black ghost knife fish , 2 chinese eaters , 1 pleco , and a bunch of different species of tetras . my elephant nose fish are doing great ! i feed them frozen ( and live ) brine shrimp , bloodworms , sometimes krill , sinking meaty pellets , and cyclop - eeze ( which is a crustacean ) , and chopped up night crawlers . my tank temperature is 79 degrees f . and ph is 6 . 8 . it is also well planted with 2 pieces of driftwood .
please remember that the following comments are personal experiences and may or may not apply to your setup . use them as guide to help better understand your fish , like us all individuals will behave differently under different circumstances .
weakly electric fish produce electric signals ( electric organ discharges , eods ) with a specialised electric organ creating an electric field around their body . objects within this field alter the eod - induced current at epidermal electroreceptor organs , which are distributed over almost the entire body surface . the detection , localisation and analysis of objects performed by monitoring self - produced electric signals is called active electrolocation . electric fish employ active electrolocation to detect objects that are less than 12 cm away and have electric properties that are different from those of the surrounding water . within this range , the mormyrid gnathonemus petersii can also perceive the distance of objects . depth perception is independent of object parameters such as size , shape and material . the mechanism for distance determination through electrolocation involves calculating the ratio between two parameters of the electric image that the object projects onto the fish ' s skin . electric fish can not only locate objects but can also analyse their electrical properties . fish are informed about object impedance by measuring local amplitude changes at their receptor organs evoked by an object . in addition , all electric fish studied so far can independently determine the capacitative and resistive components of objects that possess complex impedances . this ability allows the fish to discriminate between living and non - living matter , because capacitance is a property of living organisms . african mormyrids and south american gymnotiforms use different mechanisms for capacitance detection . mormyrids detect capacitance - evoked eod waveform distortions , whereas gymnotiforms perform time measurements . gymnotiforms measure the temporal phase shift of their eods induced at body parts close to the object relative to unaffected body parts further away .
i find that having more fish and a large group of elephant noses will bring them out during the day time . i currently have 13 elephant nose in a 140 gallon tank with tetras , plecos , rainbows and corys . there is probably about 80 different fish with the elephant nose . the elephant nose are so happy that they come up and feed out of my hand . there is plenty of driftwood , rocks and plants for them to hide in .
elephant noses are a great fish to have but i wouldn ' t recommend it to beginners as they are very sensitive to water quality . i use to have 2 of these fish but you are supposed to have 3 + because they are very territorial with each other so you need more than to so they share the bullying . they wont harm each other but will skillfully chase one another dodging in and out plants and décor with great precision . make sure you have places for them to hide as they are more active at night . one of mine use to live under a slightly raised piece of slate and the other in a hollow coconut . to keep them thriving do regular water changes maybe 1 every 2 weeks change 20 % of water . they will take all frozen foods mainly bloodworm and will be trained to eat from your hand . make sure you don ' t have large gravel for the bottom or it can harm their noses i recommend sand or fine gravel . they are very god with other fish they lived with maybe 19 other fish including 2 cichlids and 2 red eyed puffers in my 45 gallon tank . very enjoyable fish .
it finds its way through the murk using its trunk , which generates a weak electrical field that helps it sense its surroundings and even discriminate between different objects . the fish’s electric sense allows it to hunt insect larvae in pitch darkness .
in order to diminish the amplitude of the eod produced by the fish and thereby impair active electrolocation , the tail of the fish was wrapped in aluminium foil . this caused a short circuit of the electrical organ . in these experiments , the two dipole objects were positioned at a lateral distance of 3 mm at the middle of the so , and novelty responses and sors were evoked , investigated , and analysed in the same manner as described for the standard paradigm above .
, you can tune into it by placing two electrodes in the aquarium water and connecting them to an amplifier . the field will then manifest as a series of clicks , increasing in frequency hen the fish is distressed or excited . although you
pusch r , emde g von der , hollmann m , bacelo j , nobel s , grant k , engelmann j . active sensing in a mormyrid fish : electric images and peripheral modifications of the signal carrier give evidence of dual foveation .
pusch r , emde g von der , hollmann m , bacelo j , nöbel s , grant k , engelmann j . active sensing in a mormyrid fish - electric images and peripheral modifications of the signal carrier give evidence of dual foveation .
i bought 3 , within a couple of weeks two were dead , too shy to come out and eat . the survivor gets along fine with the other fish in the 110 gal . community tank , ( black ghost knife , red tail shark , bala sharks , angel fish , gouramis , plecos , clown loaches , tiger barbs and rasboras ) it likes to hang around with the black ghost knife the most . have had it over a year with no problems . especially likes blood worms .
it ' s an omnivore which prefers live foods , but will take frozen foods , and possibly flake . the ' trunk ' is used for hunting small food organisms , a soft substrate is necessary to prevent injury to the snout . these fish have a weak electric organ which they use for navigation . you must take great care if you ever need to treat these fish , or a tank containing them , with a medication . they are sensitive to several of the common treatments - read the product insert carefully .
because of the peculiar design of the fish’s retina , it was thought to be blind until about 10 years ago , says andreas reichenbach of the paul flechsig institute for brain research in leipzig , germany . reichenbach has now worked out what the cups are for .
there is no other fish that even closely resembles the elephant nose . the top and bottom profiles are almost identical , which come to an end in a forked shaped caudal fin ( tail . ) the small dorsal fin is located at the end of the body , it to is mirrored by the anal fin . the most distinguishing trait of the fish is its ' mouth . it is located at the eye level and the lower jaw section extends into a moveable trunk like extension . guess where the common name comes from .
- very difficult to breed in captivity . studies have shown that when introduced into aquaria , the electrical organ that is used to find food can get reversed from male to female , making it impossible for even the fish to tell the gender of their tank mates .
it cannot be sexed by external means ( some sources suggest the shape of the anal fin may be significant ) , although the sexes can be distinguished by examining the electric fields produced by the fish . obviously this approach is not an option for the general aquarist .
what the researchers found is that the fish could recognize objects electrically that they had only seen visually , in addition , they favored whichever sense was most reliable at the time with the electrical sense dominating when the object was close up , while vision dominated with distant objects .
) . switches were between a shunted and open - circuit object ( see standard stimulus in the material and methods section ) . motor - responses of the schnauzenorgan could be evoked in all fish by both , an impedance - switch from a low to a high impedance ( fig .
this is a nocturnal fish , so dim lighting is required to make it feel secure . plenty of cover should also be provided , particularly if you’re keeping a few together . smooth rocks , driftwood and plants that can survive under low lighting , such as anubias sp . , java fern and vallisneria can all be used . a sand substrate is absolutely critical to the well - being of this species , as it burrows into it with it’s proboscis - like lower lip . sharp - edged or coarse substrates can damage the mouthparts of the fish and prevent it from feeding naturally .
not thought to have been achieved in captivity . one scientific study suggests that being kept in the confines of an aquarium causes the sexually - defining patterns of electric impulses to become less clearly defined or even reversed . this means the fish cannot recognise members of the opposite sex , and inhibits spawning .
mormyrids also have relatively large brains , possessing the equivalent of a human brain in terms of its size . there are 3 different types of electroreceptors that carry information to different parts of this , amazingly allowing g . petersii to distinguish between different species of fish and ascertain the sex of conspecifics in total darkness !
before animals could be transferred to the experimental set - up , they were anaesthetised by immersion in ms - 222 ( tricaine methane sulfonate ; sigma , st . louis , mo , usa ; concentration 306 . 17 μmol l - 1 ) and transferred to a holder . here , the fish ' s trunk was lightly restrained . the animals head was not covered from the pectoral fin to the so . the holder was placed in a tank ( 30 × 19 . 5 × 18 . 5 cm 3 lxwxh ) with fresh water ( 123 μs cm - 1 ) , where the fish recovered from anaesthesia within a few minutes .
we have had two of these fish in a 96 let tank for over eighteen years . they have plenty of bog wood to hide under and usually rest at opposite ends of the tank . there is also some plants but not enough to make it too crowded as they do sometimes like to shoot around when the lights are off . they only like bloodworm which they are given once the lights are off . they have shared the tank with a variety of other community fish over the years , currently eight harlequins , three pandas , a corydoras myersi and a few cardinal tetras . 30 % water change every three ( ish ) weeks
it is well established that some electric fish can respond to changes in their environment with a change in the rate of their eods [ 16 , 19 , 22 , 26 , 30 , 31 ] . these novelty responses can be evoked by all sensory modalities . here we show that novelty responses caused by changes in the impedance of an object located near the schnauzenorgan can be accompanied by a previously un - described motor - response , the schnauzenorgan response . following a novelty response , fish show a quick movement of their schnauzenorgan towards or away from the object . both sor and novelty response probabilities scale with the change in the eod - amplitude .
it’s less easily kept with conspecifics . buying only 2 is not recommended , as the stronger will often bully the weaker to death . if you wish to keep more than one of these you’ll need a suitably large tank with lots of hiding places . buy at least 5 fish in order to disperse aggression within the group .
these are the coolest fish . i have one named elefunky . he is in a large tank with 2 kissing gouramis , 2 gold gouramis , 1 dwarf gourami , 2 small barbs , 3 kuhli loaches , 3 albino catfish and 1 bristle nose catty . he loves swimming amongst the plants , and spends the majority of his time there . he does however cruise around the tank , nose butting everyone else just to remind them of his dominence . he has it all over them , even the larger fish . saying that , he and the bristle nose have buddied up together . i highly recommend elefunkies , everyone should have one . be careful if treating the tank with medication , they have no scales and it can burn their soft skin .
sors were undirected single movements of the schnauzenorgan , but up to seven consecutive movements were observed . theses were reminiscent of the rhythmic scanning movements of the schnauzenorgan , which these fish perform during foraging [ 39 , 40 ] . irrespective whether a single or multiple sors were evoked , sors were quick , with angular velocities reaching 89 . 4 ± 34 . 3° / sec . this value is lower than the natural scanning movement observed in freely moving animals , where values of up to 500° / sec have been reported [ 28 ] . however , these data were based on regular video analysis at frame rates of 40 ms and an analysis of freely moving fish using high - speed videos ( 128 frame / sec ) indicates that natural scanning movements are comparable in their speed to evoked sors .
in order to analyse the latency of the sor and to compare it with the latency of the simultaneously recorded novelty response , we used high - speed video sequences ( 128 frames / sec ; 16 sors ) obtained from one fish . these data show that the peak of the novelty response occurred at a mean latency of about 40 ms after changing the electrical properties of the dipole - object . the latency of
the elephant nose are shy by nature and will require a well planted aquarium , with subdued lighting and a soft substrate . you must include hiding places in the form of clay pipes , rockwork or driftwood . they should be kept singly as they are somewhat aggressive to their own species . nocturnal in nature they will become more active at dusk and start looking for food , it is important to provide this to them after the lights come out . given time they will become active with the other fish with the lights on . feed them with tubifex worms , mosquito larva and bloodworms . they are a member of the electric fish family and emit a small electrical charges that change with their mood . they are very sensitive to water quality and i have read that some municipal water companies use them as a gauge of water purity .
like other scaleless fish , it is sensitive to many aquarium medications , including salt . it is also very sensitive to deteriorating water conditions and is actually used as an indicator of water quality in municipal supplies in parts of germany and the us . the frequency of its electrical discharges increases as the water becomes more polluted . when maintained correctly , however , it makes a superb addition to the african biotope aquarium , or a community of compatible species .
in this study we are mainly interested in possible links between ( motor ) behaviour and electrolocation . it has been shown that the mormyromast system is important for foraging and orientation [ 10 , 11 ] . fish can perceive a wealth of information form their ' electrical ' world , including parameters such as size and distance of objects and the differentiation of various object properties , like capacitive and resistive electrical properties [ for review see : [ 12 ] ] .
) . except for the tip of the schnauzenorgan , where the orientation of the dipole - object was along the rostro - caudal axis of the animal , two such dipole - objects were placed at equal distances at opposing sides of the animal . distances between dipole - object and the skin of the fish were 1 , 3 , 5 , 7 , 10 , 12 and 15 mm , and placement of the objects along the length of the animal is shown in figure
a quiet but territorial species . it is peaceful towards heterospecifics , although it should not be kept with very active or aggressive species , as it will be out - competed at feeding time . it also does not mix well with other mormyrids . suggested tankmates include other african species such as butterfly fish , smaller bichirs , congo tetras , synodontis catfish and ctenopoma species . g . petersii can also be kept successfully with peaceful cichlids such as satanoperca , some geophagus and angelfish .
in the course of evolution many different sensory systems and sensory receptors have developed . one of the rather unique sensory systems is that of active electrolocation and electro - communication found in mormyriform and gymnotiform weakly electric fishes from africa and south america , respectively . during active electrolocation mormyrids emit and simultaneously perceive electric signals , which enable them to detect and analyse nearby objects . this is considered as an adaptation enabling electric fish to extend their activity to the hours of darkness , since the dependence on vision is expected to be reduced .
capacitive switches evoked a sor in 45 % of all stimuli ( 70 switches , 32 sors ) . in comparison , purely resistive switches ( from a short to an isolator ) under otherwise identical conditions lead to sors in 55 out of 64 cases ( 86 % ; 64 switches in 4 fish ) . sors caused by capacitive switches were less frequent and also lower in amplitude compared to those caused by purely resistive switches ( mann - whitney u - test , z = - 2 . 5 , p < 0 . 011 ; fig .
probability of sor . a . probability of the sor as a function of the dipole - object ' s position along the body of the fish . black filling represents a single sor , medium grey two , and light grey bars three consecutive responses , respectively . the total probability of a sor is given by the white bars . numbers above the graphs show the absolute number of stimuli ( 100 % ) at each position . b . box - and - whisker plot of the amplitude ( peak amplitude ) of the sor as a function of distance . sor amplitude were almost constant at object distances of 1 to 12 mm , while amplitudes decreased at a distance of 15 mm c . box - and - whisker plot of the peak amplitude of the sor as a function of dipole position . sors declined in amplitude form rostral to caudal along the fish ' s body . in b and c horizontal lines above the plots summarise the results of the dunn post - hoc test ( p < 0 . 05 ) . data labelled far in b and c indicates the false alarm rate amplitudes .
i have read that the elephant noses are not for beginners but i am new to having a larger tank i have one in a 40 gallon tank and it is very happy and healthy so far . we have had it about 2 months now . we have lots of live plants and two driftwood pieces we put together to form a sort of cave / tunnel for it . we also have 2 fantail goldfish , 2 mickey mouse platys , 2 sunburst platys , 3 dalmation mollys , a peacock eel ( who swims constantly and never hides which i heard is not normal ) , a pleco and ghost shrimp . the elephant nose swims around day and night although more so at night . it doesn ' t really bother any of the other fish unless they go into one small part of the
cave
and even then it doesn ' t bite them , just kind of scares em off . he / she always has it ' s head sticking out , watching the other fish and never really hides completely . within 3 days it was eating the frozen blood worms out of my hand ( although i have to bring the food to it , it doesn ' t come and get it ) and when i clean the tank it swims around my hand and through my fingers . we have had a high nitrite ( or nitrate ? i always confuse the two ) problem and it did not seem to effect the fish at all . that problem is now under control . i have to honestly say i was not thrilled when my husband first got the fish , i thought it was incredibly ugly but it ' s temperament and the way it responds to me has changed my mind . i can defiantly see myself getting another in the future . the only issue we had was that my husband got 2 at the same time and they chased / harassed each other all the time . we gave one away and now they are both happy . the pet store didn ' t think to mention that they are territorial , especially with their own kind and they had had the two in a tank . . . go figure . thank goodness for the internet and sites like this one or i would never have known !
yet a different example of electrosensory foveation in the passive electrosensory system can be found in paddlefish ( polyodon spathula ) , which have the highest density of ampullary receptors on their rostrum [ 44 ] . these fish swim in an undulatory manner that imparts a lateral oscillating motion to their rostrum . these saccade - like motions of the rostrum are interpreted as an adaptation for prey detection , since they increase the width of the electrical scan field . in contrast to the sor , however , these movements are rather slow . if a potential prey is encountered , however , a fast and precise turning of the rostrum takes place .
a video camera ( sony dcr - hc 40e ) was placed above the tank and images of the animals were acquired and stored at 22 frames / s with a pc running viewer 2 ( biobserve , bonn , germany ) . a pair of carbon electrodes on the inside wall of the tank between the head and tail of the fish was used to record the electric organ discharges ( custom - built amplifier , band pass filtered 10 hz - 10 khz ) . eods were stored using a ced digitiser ( cambridge electronic design , micro 1401 12 bit , 200 khz , analogue - digital converter ) and spike2 software ( cambridge electronic design ltd ) .
my experience with elephant nose is they need worm - type food ( e . g frozen blood worm , live black worm ) diet due to the structure of the mouth . the mouth is the hole above the nose sensory . the elephant nose will refuse dry food because of they lack of ability to consume the food from the shape of their mouth . the elephant nose has poor sight and relies on the electric sensory organs . the requirement of cave fixtures whether from rocks or drift wood is a must to make them feel secure . caves preferred by the elephant nose is one with two doorways . elephant nose needs to be the dominant bottom dweller as it can easily be stressed or bullied by a more bossy species . this is the major reason i do not keep elephant noses anymore , because of my larger clown loaches were giving the fish a difficult time . other bottom dwelling fish to avoid is the pictus catfish , eeltails or tandanus catfish , large clown loaches and black ghost , due to their bossy nature towards the elephant nose . another thing is elephant nose are sensitive to poor water quality . you need to practice good aquarium management , such as regular water changes . although elephant nose is nocturnal and secretive , they can be trained in behaviour by food rewards to be more outgoing or to come out in the day .
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 .
vertebrates have repeatedly been noted for having remarkably constant ratios of brain to body o2 consumption , the brain using 2 - 8 % of resting body o2 consumption , suggesting that evolution has put strict limits on the energetic cost of brain function . only man , with a value of 20 % , is an exception to this rule . however , the results presented here suggest that , in the electric fish gnathonemus petersii , the brain is responsible for approximately 60 % of body o2 consumption , a figure three times higher than that for any other vertebrate studied , including man . the exceptionally high energetic cost of the g . petersii brain appears to be a consequence both of the brain being very large and of the fish being ectothermic . it was also found that g . petersii has a high ability to utilise o2 at low levels . thus , during falling [ o2 ] , this species was found to maintain both its o2 uptake and its electric discharge rate down to an ambient o2 level of 0 . 8 mg l - 1 ( at 26 & deg ; c ) , although it was unable to tolerate an [ o2 ] below 0 . 3 mg l - 1 . during severe hypoxia ( < 0 . 8 mg l - 1 ) , g . petersii attempted to gulp air from the water surface . these results establish a new record for the energetic cost of a vertebrate brain and they show that the species possessing such a brain has a high capacity for utilising o2 at very low ambient concentrations .


kallima , known as the oakleaf or oak leaf butterflies , is a genus of butterflies of the subfamily nymphalinae in the family nymphalidae .

they are found in east , south and southeast asia .

their common name is a reference to the lower surface of their wings , which is various shades of brown .

when the wings are held closed , this results in a remarkable similarity to a dead leaf , further emphasized by their wing shape .

kallima

kallima inachus boisduval , 1846 = inachus ( boisduval , 1846 ) = kallima formosana fruhstorfer .
kallima paralekta ( horsfield , [ 1829 ] ) = paphia paralekta horsfield , [ 1829 ] = kallima hewitsoni moore = kallima limborgii moore 1878 .
unwind before or after a holistic treatment in the idyllic surroundings of the kallima spa .
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frequency of feeding of kallima inachus at different fermented fruit juices . high quality figures are available online .
kallima alompra moore , 1879 ; trans . ent . soc . lond . 1879 ( 1 ) : 14
kallima buxtoni moore , 1879 ; trans . ent . soc . lond . 1879 ( 1 ) : 10
kallima spiridiva ; [ bow ] : pl . 147 , f . 1 ; [ bor ] , 287
each treatment at kallima is carried out using premium products by elemis , kallima ' s exclusive skincare partner . each product , treatment and package is specially designed to treat the individual needs of your body and skin .
c1s , c1s plus and c2s – the kallima® fsc® - certified coated cover collection delivers print performance and reliability .
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frequency of feeding of kallima inachus at different colored flowers supplemented with fermented pear juice . high quality figures are available online .
kallima photography is a south florida based photography and videography studio specializing in weddings in south florida , the bahamas , and beyond .
kallima albofasciata moore , 1877 ; proc . zool . soc . lond . 1877 ( 3 ) : 584 ; tl : s . andamans
kallima inachus ; [ bow ] : pl . 195 , f . 10 ; [ bor ] , 290 ; [ mrs ] , 563
kallima paralekta ; [ ebw ] ; [ bow ] : pl . 146 , f . 11 - 12 ; [ bor ] , 288
kallima inachis [ sic ] eucerca fruhstorfer , 1898 ; berl . ent . zs . 43 ( 1 / 2 ) : 191 ; tl : okinawa
kallima limborgii moore , 1878 ; proc . zool . soc . lond . 1878 ( 4 ) : 828 ; tl : moolai , 3000 - 6000ft
kallima inachis [ sic ] amplirufa fruhstorfer , 1898 ; berl . ent . zs . 43 ( 1 / 2 ) : 192 ; tl : malacca
kallima wellbeing centre is a spiritual goods shop that also provides holistic treatments , 1 - 2 - 1 spiritual readings , spiritual workshops and holistic courses .
at the kallima spa , you ' ll experience total relaxation and rejuvenation for mind , body and soul . choose from an exclusive selection of innovative beauty and relaxation treatments , all designed to deliver outstanding results . each luxurious treatment uses specially - designed premium products by comfort zone , kallima ' s exclusive skincare partner .
kallima buxtoni trebonia fruhstorfer , 1909 ; ent . zs . 23 ( 1 ) : [ 4 ] 231 , f . 18 ; tl : w . sumatra
kallima chinensis swinhoe , 1893 ; ann . mag . nat . hist . ( 6 ) 12 ( 70 ) : 255 ; tl : omei - shan , china
callima herrich - schäffer , [ 1858 ] ; samml . aussereurop . schmett . ( ii ) 1 : 54 ( unj . emend . kallima doubleday , [ 1849 ] )
make the earth friendly paper choice . fsc ® certified kallima coated cover meets the most rigorous responsible forestry standards in the world . let ' s keep print a sustainable medium .
kallima albofasciata ; wood - mason & de nicéville , 1881 , j . asiat . soc . bengal 49 pt . ii ( 4 ) : 227 ; [ bor ] , 288
kallima inachis [ sic ] alicia joicey & talbot , 1921 ; bull . hill mus . 1 ( 1 ) : 170 , pl . 21 , f . 9 ; tl : haina , five finger mtns
welcome to kallima club spa at hilton london syon park . experience total relaxation and rejuvenation for the mind , body and soul with a range of treatments , membership options and personalised fitness for a balanced wellness lifestyle .
kallima ' s bright , blue - white shade is perfect for creating beautiful , vivid print and top quality jobs . its consistent printability and reliable runnability on press ensure you get the job done efficiently and on - time .
fsc ® - certified kallima ® c1s , c1s plus and c2s are the ideal grades for all your coated cover printing needs . produce beautiful , vivid print while saving up to 20 % on paper cost and shipping of printed material .
kallima coated cover has a unique , low density , high bulk construction giving your print the stiffness and presence you expect , all while offering savings of up to 20 % over competing products . measure your savings potential today with our savings calculators .
kallima philarchus ; moore , [ 1880 ] , lepid . ceylon 1 ( 1 ) : 37 , ( 2 ) pl . 20 , f . 1 ; [ bow ] : pl . 146 , f . 13 ; [ bor ] , 288
at the kallima spa , you ' ll experience total relaxation and rejuvenation for mind , body and soul . choose from an exclusive selection of innovative beauty and relaxation treatments , all designed to deliver outstanding results . each luxurious treatment uses specially - designed premium products by
there’s nothing as memorable as a kallima spa gift certificate – the ultimate indulgence for the people you love the most . choose from a range of treatments , products or experiences , or give them the pleasure of choosing for themselves with any of our monetary value options .
the feeding responses of kallima inachus to different volatiles . a : single compounds at concentrations of 5 % . b : compound mixtures at a total concentration of 5 % . c : different doses of ethanol and ethyl acetate ( mean ± sd ) . different letters indicate significant differences with the lsd test ( p < 0 . 05 ) . high quality figures are available online .
no subspecies are listed under this species . it is also known as the south indian blue oakleaf .
this species has been referred to as
southern blue oakleaf
in some recent books . this is a slight misnomer since the species is endemic to the western ghats , it does not occur elsewhere in southern india .
kollar , 1844 – sahyadri blue oakleaf . kunte , k . , s . sondhi , and p . roy ( chief editors ) .
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copyright © 2018 , all rights reserved . national centre for biological sciences ( ncbs ) holds copyright for all the original material and compilations on this website , although contributing writers and photographers may hold copyright for their material as cited . material from this website can be used freely for educational , basic research and conservation purposes , provided that this website is acknowledged and properly cited as the source . contact us to obtain prior permission for any other use , including for large data downloads and collaborative research .
600x400 ( ~ 53kb ) upperside india : pune , 7 . 4 . 2007 © kedar tokekar
600x400 ( ~ 55kb ) underside india : pune , 7 . 4 . 2007 © kedar tokekar
w . china , c . china , japan , himalayas , assam , pachmarhi , e . ghats , s . bihar . see [ maps ]
1024x768 ( ~ 118kb ) upperside yonahadake , okinawa , ryukyu , japan , 7 - 93 , photo © s . shuichi haupt
1024x768 ( ~ 104kb ) underside oppadake , okinawa , ryukyu , japan , 7 - 93 , photo © s . shuichi haupt
600x400 ( ~ 96kb ) upperside india : subansiri , 1 . 11 . 2005 , photo © kedar tokekar
1024x768 ( ~ 107kb ) oppadake , okinawa , ryukyu , japan , 7 - 93 , photo © s . shuichi haupt
paphia paralekta horsfield , [ 1829 ] ; descr . cat . lep . ins . mus . east india coy : ( explic ) , pl . 6 , f . 4 ,
philarchus ( westwood , 1848 ) ( amathusia ) ; cabinet orient . ent . : 56
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
the butterflies of the malay peninsula . fourth edition revised by j . n . eliot with plates by bernard d ' abrera
the genera of diurnal lepidoptera , comprising their generic characters , a notice of their habitats and transformations , and a catalogue of the species of each genus ; illustrated with 86 plates by w . c . hewitson
( 1902 ) , : ( ragadia - argyronympha - hypocysta ) : 1 - 6 , pl . [ 1 ] ( 1895 ) , [ 2 ]
aufzählung und beschreibung der von freiherr c . v . hügel auf seiner reise durch kaschmir und das himaleygebirge gesammelten insekten in hügel ,
the butterflies of india , burmah and ceylon , a descriptive handbook of all the known species of rhopalocerous lepidoptera inhabiting that region , with notices of allied species occurring in the neighbouring countries along the border , with numerous illustrations . vol . 2
a list of the lepidopterous insects collected by mr . ossian limborg in upper tenasserim , with descriptions of new species
if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .
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you can rest assured that our paper meets the most rigorous responsible forestry standards in the world .
rayonier advanced materials is a manufacturer of forest products – lumber , paper pulp , paper and specialty cellulose pulp – and a global leader in sustainable forest management practices .
we are photojournalistic wedding photographers , who strive to provide our clients with amazing portraits as well .
we believe that every wedding has a unique story , each couple is different . we love that our couples come from all walks of life and backgrounds . each have their own passions about their wedding and what they hope for in a wedding photographer . it’s those unique personalities that make weddings fun for us , and we look forward to being a part of your day . here’s a snippet of some of the things clients are saying about us :
i have no words to express how amazing ben and beka are . we got married almost a year ago in the dominican republic and these 2 came along for the ride . it was hands down the best decision we made having them there with us . i think they became a part of the family in this whole process . the pictures we have from that weekend were incredible . we relive our wedding and those amazing memories every time we look at them . not to mention they are amazing human beings . probably would have invited them to the wedding regardless . i recommend them to everyone i know , eyes closed and without a doubt . miss you guys !
purchasing your gift certificates is simple , call us on 0208 380 1590 , email us on kallimaspa @ urltoken or simply pop in to the spa and talk to one member of the team .
includes a 30 minute vitality back , neck and shoulder massage . a 30 minute elemis revitalise express facial . full use of the spa facilities , & lunch .
includes 1 x 30 minute scrub , 1 x 30 minute massage , 1 x 30 minute facial . served with a glass of bubbly and afternoon tea .
relax this summer with two soothing treatments , a glass of prosecco and some delightful strawberries with cream .
includes bed and breakfast , champagne and chocolates in your room , one 25 - minute spa treatment each plus three - course dinner .
our signature treatments are designed by elemis to work in natural synergy with the skin , body and mind . every treatment is specifically designed to offer a unique experience , using powerful massage sequences and the most potent actives available in the world today . no matter which treatment you choose , our experts will look after you
a powerful treatment to stimulate every cell in the body , helping alleviate muscular pain and remove toxins . this is a revolutionary , mineral - charged experience of skin conditioning , metabolic balancing and energising wellness .
a survival treatment boosting sluggish complexions . stimulating the removal of impurities and dead skin cells ; then restores moisture for instantly clearer and visibly brighter skin .
the hard - working facial for ageing , dehydrated skin and tired eyes . it maximises cell regeneration , a multi - dynamic facial massage sequences boost circulation , whilst scalp and foot massage deeply relax .
we are proud partners with elemis which is naturally sourced , science led , results driven and always personally prescribed .
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we strive to help others on their spiritual path by providing the tools to help them reach their goals .
feel free to visit us at our centre to view products and take part in our workshops & courses or even to have a nice relaxing holistic treatment .
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rarely am i so disappointed with a place that i resort to leaving a negative review , but after . . .
i had two 25 minutes treatments - both back and neck massages . both therapists were friendly . . .
rarely am i so disappointed with a place that i resort to leaving a negative review , but after absolutely nothing being done to resolve my complaints on the day apart from being told a manager would email me , and then coming here to see that . . .
i had two 25 minutes treatments - both back and neck massages . both therapists were friendly , welcoming and professional . the first therapist gave me a robe and towel as i hadn ' t checked in yet , which was a nice touch . the massages were slightly different styles . . .
stayed there overnight for a relaxing spa day and i had a great time . the room was beautifull with a nice wiev on the fountain . the main highlight was of course the spa . fantastic design of the swimming pool facility really helped me relax . the . . .
the spa facilities themselves are lovely . we were a group of five and disappointingly 3 out of 5 of us were disappointed with the treatments . therapists felt unqualified , lacking in spa etiquette , ignored requests for a firmer massage , shortened treatment time , no ambient music . two . . .
afternoon tea was not very nice , spa facilities were lacking and service was confusing . a lot of staff coming and going and had to explain the same things to multiple people which was frustrating . the spa rooms were disturbed by loud school children coming in . . .
we booked a spa break and were treated with an upgraded room . the staff were very welcoming and the spa was just what we needed to unwind . dinner in the restaurant was included , the only note is the drinks were pretty pricey with dinner .
me and my sister took my mum as a b - day spa day treat .
this place is nice an calming . i booked 3 30 minute treatments which were body scrub , massage and a facial . this also included afternoon tea . i was unable to book all treatments together and later in the afternoon which wasn ' t a massive problem , just not . . .
we had a fabulous relaxing day here as a birthday celebration . staff were really friendly and showed us around making us feel v welcome , providing slippers and robes . we started our day with a swim - nice size pool and good size jacuzzi . there . . .
i have a monday to friday membership which is great value for money and offers full access to the gym , spa , sauna , steam and pool . . . you get discount on room rates and treatment rates as a member , staff are all very lovely and always willing . . .
what a wonderful day , as soon as you walk into the spa reception you are met with helpful staff and the ambience of the area is fresh and welcoming . my friend and i had our treatments first which we absolutely great i had ingrid give . . .
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the spa lunch consist of soup of the day and main course from the choice of salads , sandwiches , paninis or wraps . unfortunately we do not have packages with dinner . you are more than welcome to have dinner in the restaurant . . .
the spa lunch consist of soup of the day and main course from the choice of salads , sandwiches , paninis or wraps . unfortunately we do not have packages with dinner . you are more than welcome to have dinner in the restaurant after the spa .
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with special rates and complimentary breakfasts for two - from healthy to decadent - our breakfast included packages are the perfect way for you to relax and recharge .
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enjoy a 25 - minute treatment plus dinner at marco pierre white restaurant monday through thursday .
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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .
species catalog enables to sort by characteristics such as expansion , flight time , etc . .
[ 85 ] urltoken ( insecta . pro previous version / cтарая версия insecta . pro )
butterfly is famous as an example of protective coloration , as well as being in form ( with folded wings ) is strikingly similar dry leaf .
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> press w . h . , flannery b . p . , teukolsky s . a . and vetterling w . t . < br / > < strong > cyclic redundancy and other checksums < / strong > < br / > < a href =
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to maintain daily life activities and reproduction , adult insects must look for and find suitable nutrients to supplement their diets . insects are known to make use of a variety of sensory modalities in foraging , and the integration of visual , olfactory , and gustatory cues are usually involved in their orientation to and finding of food sources ( barth 1991 ) .
depending on the food type , adult butterflies can be classified into three guilds : nectar - feeding , fruit - feeding , and a combination of both ( gilbert 1972 ; devries 1988 ; devries et al . 1997 ; devries and walla 2001 ; molleman et al . 2005 ; ômura and honda 2009 ) . most butterfly adults are found in temperate regions and feed on nectar , while fruit - feeding butterflies are found mainly in tropical and subtropical regions and feed on rotting fruits , exuded tree sap , mud , carrion , and dung ( boggs and jackson 1991 ; ômura et al . 2000 ; devries and walla 2001 ; krenn 2008 ) .
the scientific literature clearly demonstrates that the flower - visiting behavior of adult butterflies is affected by the color and odor of flowers , which provide specific information for adults , and that colors are extremely important in flower recognition and preference in some species ( use and vaidya 1956 ; weiss 1997 ; andersson and dobson 2003 ; borges et al . 2003 ) . flower scent is an olfactory stimulus that influences butterfly visitation ( andersson 2006 ; ômura 2006 ) . for example , 2 - phenylethanol and phenylacetaldehyde stimulate the foraging behavior of pieris rapae ( honda et al . 1998 ; ômura et al . 1999a , b ; ômura and honda 2005 ) .
experiments were performed at the experimental station of the research institute of resource insects of the chinese academy of forestry in yuanjiang , yunnan ( 400 m . a . s . l . , 101 . 59°–101 . 00° e , 23 . 35°–23 . 36°n ) . a net house ( 12 m long , 8 m wide , and 5 m tall ) with evenly scattered sunlight was used as the experimental chamber .
four - day - old naive adults of k . inachus were obtained from artificial culture . larvae of k . inachus were reared on fresh leaves of strobilanthes cusia ( acanthaceae ) at 25°c and with a photoperiod of 14 : 10 l : d . adults were supplied with purified water and were not allowed to contact fruit or sap before tests .
when an adult k . inachus landed nearby the food , and then reached to the food and uncoiled the proboscis to probe and feed , it was recorded as a visit .
the cotton flowers were arranged the same as in the test of feeding responses to different colors of artificial flowers . then , we sprayed fermented pear juice evenly on the petals . the juice that couldn ' t be absorbed by the cotton petals flowed into the disk . the evening before the bioassay , about 200 adult butterflies ( sex ratio not controlled ) were released into the net house and allowed free - flight to acclimatize them to the experimental arena . the number of feedings during two 2 - hr periods ( 10 : 00–12 : 00 and 15 : 00–17 : 00 ) was recorded for each model . during the bioassay , the positions of the different juice disks were changed in a clockwise direction every hour , and the flowers and juice were renewed at the same times . the observations were repeated on a second day . the data were analyzed with a chi - square test .
based on the results of auto thermal - desorption gas chromatography / mass spectrometry , we used seven pure chemical compounds in behavioral and eag response tests . all seven chemicals were commercially purchased and were more than 98 % pure . these compounds were 3 - methyl - 1 - butanol , ethyl acetate , isoamyl acetate , α - pinene , the 2 - pentanone homologue butanone , ethanol , and acetic acid . the first five compounds were present in at least three of the six kinds of fermented fruits ; ethanol and acetic acid are the major volatiles of fermentation , but were not found in our fermented fruits .
in our long - term field observation , we found k . inachus habitually basks on light colors . in order to avoid color interference , white disks ( d = 20 cm ) were covered with deep - purple cloth , spaced above the floor in the net house , and filled with 25 ml of a solution , which was dissolved in deionized water as described below . deionized water was used as a control . three feeding experiments were conducted :
in the center of the net house , a 6 m × 6 m square was delimited , and the four corners of the square and the midpoint of the four sides were chosen as the placement points of the treatments . the feeding performances of adult butterflies were characterized by assaying seven single compounds at concentrations of 5 % v / v .
in the center of the net house , a 6 m × 3 m rectangle was delimited , and the four corners of the rectangle and the midpoint of the long sides were chosen as the placement points . the synergistic effects of different compound mixtures were evaluated . based on the results of the single - compound assays , we mixed ethanol , α - pinene , and ethyl acetate at equal volumes in the following combinations : ethanol + ethyl acetate + α - pinene , ethanol + ethyl acetate , ethanol + α - pinene , ethyl acetate + αpinene , and ethanol . each mixture had a total concentration of 5 % .
in the center of the net house , a 6 m × 5 m rectangle was delimited , and the four corners of the rectangle and the one - third point of the four sides were chosen as the placement points . to identify the most effective doses of ethanol and ethyl acetate , a concentration series ( 0 . 001 , 0 . 01 , 0 . 1 , 1 , or 10 % ) of each compound was bioassayed ; 0 . 5 % ethanol was also used .
before each of the three bioassays , about 100 adults ( sex ratio not controlled ) were released into the net house and allowed free - flight to acclimatize them to the experimental arena . the number of feedings during two 2 - hr periods ( 09 : 30–11 : 30 and 14 : 00–16 : 00 ) was recorded for the treatment and control models . every model was duplicated . during each bioassay , the positions of the models were rotated clockwise every 30 min , 40 min , or 20 min , respectively , and solutions were renewed to account for evaporation . tests were replicated for two days . significant differences between treatments and controls were determined using analysis of variance , followed by the lsd test .
the eag responses to the solutions at doses of 5 µl / ml were tested . a series of odorant compounds were applied to butterfly antennal preparations in the following order : ether control , standard stimulus ( see below ) , individual samples ( 3 - methyl - 1 - butanol , butanone , αpinene , acetic acid , ethyl acetate , isoamyl acetate ) in a random order , standard stimulus , and ether control . each sample was tested five times in each antenna .
the eag responses to the solutions at doses of 50 µl / ml were tested , with solutions and procedures as described above .
the eag responses to mixtures at the dose of 50 µl / ml were tested . mixtures mixed at equal volume were as described for the behavioral feeding experiments .
to identify the eag responses to different doses of ethanol and ethyl acetate , ethanol was tested at six doses ( 0 . 01 , 0 . 1 , 1 , 5 , 10 , or 100 µl / ml ) and ethyl acetate was tested at five doses ( 0 . 01 , 0 . 1 , 1 , 10 , or 100 µl / ml ) .
ethanol at 5 µl / ml was used as the standard stimulus in eag experiments 1 and 4 , and 50 µl / ml was used in experiments 2 and 3 . the responses were averaged across individual trials and expressed as percentages of the response relative to the standard . data were normally distributed ; sexual differences in eag intensities were analyzed with a t - test . the intensity of eag responses to different compounds within males and within females were analyzed using analysis of variance , followed by the lsd test .
k . inachus showed no attempted feeding or orientation responses to the four different single colors , suggesting that foraging adults were not innately sensitive to these colors , and that color is not a forging signal for k . inachus .
showed no visits to deionized water . the different fermented fruit juices were not significantly differently attractive to
showed the strongest response to fermented pear juice , accounting for 16 . 47 % of feedings , while ethanol had the weakest effect , attracting only 11 . 01 % of feedings (
< 0 . 01 ) . adults showed the strongest attraction to white , with 35 . 71 % feeding at that color , and weaker attractions to red , yellow , and purple ( 19 . 48 % , 20 . 56 % , and 24 . 24 % feeding , respectively ) . there were no differences in feeding among red , yellow , and purple flowers ( χ
) . the volatile compounds found in more than half of the fruits were as follows : α - pinene , limonene , isobutyl acetate , ethyl acetate , isoamyl acetate , 3 - methylbutyl butyrate , 3 - methyl - 1butanol , and 6 - methyl - 5 - hepten - 2 - one . half of these are esters , so it can be concluded that esters are commonly found in fruit volatiles . all of the fruits we tested contained terpenoids , except for banana . esters were the major volatiles in banana , apple , and watermelon , while terpenoids dominated the orange volatiles , which also contained very small amounts of aromatics , such as 1 - methyl - 4 - ( 1 - methylethenyl ) - benzene . persimmon contained a high relative concentration of 3 - methyl - 4 - oxo - pentanoic acid , about 49 . 36 % . in contrast , esters and nitrogen compounds were predominant in pears , which also contained a small amount of hydrocarbons , such as nonane and 1 - nonene .
chemical components of six kinds of fermented fruits and their relative contents ( % ) .
single - compound bioassays of seven chemicals at 5 % concentration were performed . the number of adult butterflies attracted to these compounds was in the following order , from most attractive to least : ethanol > 3 - methyl - 1butanol > isoamyl acetate > ethyl acetate > αpinene > butanone > acetic acid > deionized water (
the synergistic effects of different compound mixtures were also tested . adults were attracted to these mixtures in the order from most to least , as follows : ethanol + ethyl acetate > ethanol > ethanol + ethyl acetate + α - pinene > ethanol + α - pinene > ethyl acetate + α - pinene > deionized water (
) . ethanol + ethyl acetate was the most effective of all mixtures . overall , the mixtures containing ethanol were significantly more attractive than deionized water (
> 0 . 05 ) . the presence of α - pinene reduced the number of feedings .
the effective doses of ethanol and of ethyl acetate were evaluated . the number of responses was significantly positively correlated with concentration for both compounds ( r = 0 . 974 ,
< 0 . 05 ) , but there were no differences between 0 . 01 % , 0 . 001 % ethyl acetate , and deionized water (
> 0 . 05 ) . ethanol was more attractive than deionized water regardless of its concentration . overall , 10 % ethanol was the most effective lure , and concentrations of 10 % , 1 % , 0 . 5 % , and 0 . 01 % were significantly more attractive than deionized water (
< 0 . 05 ) , but there were no differences between concentrations of 0 . 1 % , 0 . 001 % , and deionized water (
the relative electroantennogram responses of males and females to different compounds , a : reponses to compunds at the dose of 5 µl / ml b : responses to compounds at the dose of 50 µl / ml c : responses to mixtures at the dose of 50 µl / ml d : responses to different doses of ethanol . e : responses to different doses of ethyl acetate ( mean ± se ) . significant differences by the paired t - test are indicated by * ( p < 0 . 05 ) and * * ( p < 0 . 01 ) . high quality figures are available online .
> 0 . 05 ) , indicating that the perception of different compoundswas varied , but the differences were not significant . ethanol , which was used as a standard , elicited the lowest response in both sexes . eag responses of both sexes to six chemicals at the dose of 50 µl / ml were weaker than at 5 µl / ml , except for female and male responses to acetic acid , male responses to butanone , and female responses to ethyl acetate , which were somewhat greater than at 5 µl / ml .
> 0 . 05 ) ( figure5d , e , respectively ) . the values of eag responses were not correlated to the doses of chemicals ( ethanol : females r = - 0 . 194 ,
> 0 . 05 ; ethyl acetate : females r = 0 . 071 ,
alcohols , ketones , and esters were found in all six fruit volatiles , either as the majority components or at lower levels . esters were the main volatiles in most of the fruits tested . ethanol and acetic acid , both of which are characteristic volatile products of fermentation ( ômura et al . 2000 ; ômura and honda 2003 , 2009 ) , were not found in our samples .
this result may be because the fruit was not thoroughly fermented . ripe fruits are high in aromatic volatiles , rather than producing decay odor ( molleman et al . 2005 ) . the six kinds of fruit were fermented under the same conditions , so the variations in volatile composition were most likely due to differences in the fruits themselves and to different microbial species associated with the juice extracts ( wood 1961 ; ômura et al . 2000 ) .
although there was no ethanol or acetic acid in the fruit volatiles , the fruits emitted a variety of different types of chemicals and were strongly attractive to the butterfly , indicating that k . inachus uses a variety of food chemical signals .
in behavioral tests , ethanol was the most attractive volatile , while acetic acid was the least attractive . in contrast , the perception of antennae to acetic acid in eag tests was the strongest , while ethanol was the weakest , regardless of whether the test dose was 5 or 50 µl / ml . this result showed acetic acid induces the strongest perception by antenna , but k . inachus showed the strongest preference for ethanol . thus , the compound that most strongly stimulated feeding behavior did not induce high eag responses , and eag responses did not totally reflect the food choices of adults foraging in the field ( honda et al . 1998 ; ômura and honda 2009 ) . this interesting result suggests that antennal perception of these olfactory cues was caused by a specific sensory system that selectively elicited feeding behavior ( ômura and honda 2009 ) . antennal perception is not only related to foraging behavior , but also involves a large number of other behavioral functions . therefore , the antennal perception of volatiles is not necessarily interpreted as a food signal ; some signals may relate to host recognition , to the identification of conspecific individuals , or to defense ( kreher et al 2008 ) .
another striking result of the eag experiments was the differences found between genders for a few chemicals , such as 5 µl / ml butanone , 50 µl / ml acetic acid , and 100 µl / ml ethanol . these differences may be due to males and females differing in their dietary needs because of different reproductive strategies ( dierks and fischer 2008 ) . however , these differences were not consistent , as they changed at different volatile concentrations . different concentrations of the same compounds may reflect the distance between adult butterflies and food , but the relative eag responses of males and females to different concentrations of ethanol and ethyl acetate showed no significant correlation . the phenomenon may be caused by the tested concentration , which could have been higher than the perception threshold of the antenna of k . inachus .
according to the results of our study , we conclude the following : 1 ) the foraging , adult k . inachus has no sensitivity to food color , but innately uses olfaction to detect and locate food ; 2 ) the adults feed on a wide range of fruits , so they also use a wide range of volatiles such as alcohols , esters , ketones , and terpenoids , and the coexistence of multiple chemical signals is most attractive ; 3 ) antennal perception is not only related to foraging behavior , but also involves a large number of other behavioral functions , such as hostrecognition and defense behavior , so feeding preference is not necessary correlated with eag responses .
we thank yong cao , chuang jiang , zhen chen , tichu li , and chunzi luo for their participation in field observations , fei chen for his help in the eag experiment , and cheng - zhu yang for his kindness in rearing butterflies . special thanks to ning liang and other staff of the yuanjiang experimental station of riricaf for their careful logistical services during our fieldwork . this study was supported by science and technology of social development projects in yunnan province and the special fund for basic research cafybb2007018 and riri200705z .
andersson s . floral scent and butterfly pollinators . in : dudareva n , pichersky e , editors .
balkenius a , rosén w , kelber a . the relative importance of olfaction and vision in a diurnal and a nocturnal hawkmoth .