the blister beetle genus meloe is a large , widespread group commonly referred to as oil beetles .

they are known as " oil beetles " because they release oily droplets of hemolymph from their joints when disturbed ; this contains cantharidin , a poisonous chemical causing blistering of the skin and painful swelling .

members of this genus are typically flightless , without functional wings , and shortened elytra .

as in other members of the family , they are hypermetamorphic , going through several larval stages , the first of which is typically a mobile triungulin that finds and attaches to a host in order to gain access to the host 's offspring .

in this genus , the host is a bee , and each species of meloe may attack only a single species or genus of bees ; while sometimes considered parasitoids , it appears that in general , the meloe larva consumes the bee larva along with its provisions , and can often survive on the provisions alone , thus they do not truly qualify ( see parasitoid for definition ) .

meloe

separating the sexes of the short - necked meloe brevicollis , rugged meloe rugosus and mediterranean oil beetles meloe mediterraneus requires examining the underside of the last segment of the abdomen . this is distinctly notched in males and rounded in females .
oil beetles can sometimes be difficult to identify particularly the black meloe proscarabaeus and violet oil beetles meloe violaceus . see buglife oil beetle guide and below are some extra notes and photographs .
meloe - 1 is a new antigen overexpressed in melanomas and involved in adoptive t cell transfer efficiency .
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meloe - 1 is a new antigen overexpressed in melanomas and involved in adoptive t cell transfer efficiency . - pubmed - ncbi
black meloe proscarabaeus and violet oil beetles meloe violaceus vary greatly in colour . some violet oil beetles are all black and some black oil beetles can be violet - blue . despite their names colour is not always the best way to separate them .
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it is fairly easy to tell the sexes of the _ black meloe proscarabaeus and violet oil beetles meloe violaceus apart as the males have a distinct kink in their antennae and the antennal segments are very broad at the kink . the females have slightly kinked antennae with all segments about the same width .
preferential expression of meloe cdna in melanoma cell lines measured by qpcr , and impact of meloe expression on specific ctl clone activation . ( a ) four melanoma , one breast cancer , two renal carcinoma , and one lung cancer cell lines were tested by qpcr for the expression of meloe . rplpo and β2 - microglobulin gene expression were used as internal controls . the relative expression of meloe was calculated after normalization on the efficiency of the pcr reaction and the mean expression of these two housekeeping genes , reported to its normalized expression in melanocytes . ( b ) tnf secretion by the m170 . 48 ctl clone in response to hla - a2 tumor cell lines nontransfected ( open bars ) or transfected with meloe ( hatched bars ) or meloe - 1 ( shaded bars ) expression plasmids . tumor cells were transiently transfected with 100 ng of each plasmid with a lipofectamine reagent kit . 10 4 ctls were added to 3 × 10 4 target cells , and the ctl clone reactivity was assessed by a tnf release assay . ( c ) meloe relative expression measured by qpcr in 16 human healthy tissues .
detection of meloe - 1 / a2–specific ctls in tils infused to relapse - free melanoma patients and analysis of their repertoire diversity . ( a ) hla - a2 til populations labeled with the a2 / meloe - 1 36 - 44 tetramer . ( top ) tils infused to relapse - free patients . ( bottom ) tils infused to patients who relapsed . tils were coincubated with meloe - 1 tetramer and anti - cd8 mab . values indicate the percentage of tetramer - positive cells among cd8 + tils . ( b ) repertoire diversity of multimer - sorted populations was evaluated by labeling with 25 anti - vβ mabs . insets illustrate the purity of each sorted til population , assessed by meloe - 1–specific tetramer labeling .
bland r . g . ( 1986 ) antennal and mouthpart sensilla of the blister beetle , meloe campanicollis ( coleoptera : meloidae ) . great lakes entomologist 19 ( 4 ) : 209–215 .
short - necked oil beetle meloe brevicollis – rectangular shaped thorax which is wider than long , shiny blue - black upperparts , short straight antennae which thicken towards the tip , up to 24 mm long .
the bionomics of blister beetles of the genus meloe and a classification of the new world species pinto j . d . , selander r . b . 1970 . ill . biol . monogr . 42 : 1 - 222 .
characterization of the cdna coding for the recognized antigen . ( a ) m170 . 48 tnf responses to cos - 7 cells ( e / t ratio = 1 : 3 ) transfected with the indicated plasmids . the t cell clone was added 2 d after the transfection , and the ctl clone reactivity was assessed by a tnf release assay . ( b ) comparison of the nucleotide sequences of meloe and bc008026 cdnas and the localization of this sequence on the hdac - 4 gene . the indicated nucleotides correspond to snps between the meloe sequence isolated from m134 and a498 tumor cell lines and the meloe sequence isolated from the m117 and sw480 cell lines , and the bc008026 cdna sequence .
rugged oil beetle meloe rugosus and mediterranean oil beetle meloe mediterraneus – are very similar in appearance , both have a rectangular shaped thorax which is wider than long , rough matt black upperparts , long straight antennae , up to 22 mm long ( rugosus ) and 36mm ( mediterraneus ) . the rugged oil beetle has a sulcus ( narrow slit ) running down the middle of the thorax this is lacking in the mediterranean oil beetle . the thorax is more densely punctured in the rugged oil beetle .
characterization of meloe - derived peptide recognized by the m170 . 48 t cell clone . ( a ) structure of meloe cdna . boxes illustrate orfs > 120 bp present along the meloe sequence , and black boxes correspond to orfs tested for recognition by the ctl clone . ( b ) m170 . 48 tnf responses to cos - 7 cells ( e / t ratio = 1 : 3 ) transfected with the indicated plasmids . the t cell clone was added 2 d after the transfection , and the ctl clone reactivity was assessed by a tnf release assay . ( c ) nucleotide and amino acid sequences of the orf 1 , 230–1 , 370 of meloe isolated from the m134 cdna library . the two candidate peptides are bolded . ( d ) cytotoxicity of the m170 . 48 ctl clone against peptide - pulsed t2 cells . target cells were 51 cr - labeled for 60 min and incubated for 30 min with a range of the indicated peptides . the m170 . 48 t cell clone was added ( e / t ratio = 10 : 1 ) , and chromium release was then measured after a 4 - h incubation period .
a cytotoxic t lymphocyte ( ctl ) clone was derived from a tumor - infiltrating lymphocyte ( til ) population infused to a melanoma patient who remained relapse free for 10 yr after this adoptive transfer . this clone recognized all melanoma cell lines tested and , to a lower extent , melanocytes , in the context of human histocompatibility leukocyte antigen a2 ( hla - a2 ) , but it did not recognize other tumor cell types . the gene coding for the antigen recognized by this clone was identified by the screening of a melanoma complementary dna expression library . this antigen is overexpressed in melanomas , compared with other cancer cell lines and healthy tissues , and was thus called melanoma - overexpressed antigen ( meloe ) . remarkably , the structure of meloe was unusual , with multiple short open reading frames ( orfs ) . the peptide recognized by the ctl clone was encoded by one of these orfs , called meloe - 1 . using a specific hla - a2 / peptide tetramer , we showed a correlation between the infusion of tils containing meloe - 1 - specific t cells and relapse prevention in hla - a2 patients . indeed , 5 out of 9 patients who did not relapse were infused with tils that contained meloe - 1 - specific t cells , whereas 0 out of the 21 patients who relapsed was infused with such til - containing lymphocytes . overall , our results suggest that this new antigen is involved in immunosurveillance and , thus , represents an attractive target for immunotherapy protocols of melanoma .
reactivity of meloe - 1 / a2–specific tils against hla - a2 tumor cell lines . ( a ) lysis of the m170 melanoma cell line ( closed circles ) and of the 1355 lung carcinoma cell line ( open circles ) by the m170 . 48 ctl clone and meloe - 1–specific til populations . 51 cr - labeled tumor cells were co - cultured with t cells at various e / t ratios . chromium release in the supernatants was measured after a 4 - h incubation period . ( b ) cytokine production by the m170 . 48 ctl clone and meloe - 1–specific til populations in response to m170 melanoma cells . effector and target cells were incubated at a 1 : 2 ratio in the presence of brefeldin a and stained with anti - tnf antibody ( open bars ) , anti– ifn - γ antibody ( hatched bars ) , or anti–il - 2 antibody ( closed bars ) , and 10 4 t cells were analyzed by flow cytometry .
the oil beetles are a family of beetles that share a fascinating life - cycle in which the larvae are parasites of certain bees or grasshoppers ( 2 ) . this species , meloe proscarabaeus is bluish black in colour with a long swollen abdomen ( 2 ) , which is particularly pronounced in females when they are producing eggs . females are usually much bigger than males ( 4 ) .
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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
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primarily holarctic ( mostly palaearctic ) , with meager representation in more southern areas ; throughout na ( to nw . colombia ; hispaniola )
in some species , triungulins aggregate and use chemical signals to attract male bees to which they attach themselves . this allows transport ( and transfer ) to a female bee who carries them back to her nest ( saul - gershenz & millar 2006 ) .
first - instar larvae climb to the top of a plant as a group , clump together in the shape of a female solitary ground bee , exude a scent imitating the female bee pheromone . when a male bee comes and tries to mate with the clump of larvae , some of these clamp onto his hairs and eventually get to female bees when he mates for real . impregnated female bees fly off and build nests in burrows ; triungulins move to the new nests and feed on honey and pollen stocked by the bee for her own young . - - jim mcclarin ' s comment
pinto j . , mayor a . ( 1986 ) size , mating success and courtship pattern in the meloidae ( coleoptera ) . ann . ent . soc . am . 79 : 597–604 .
saul - gershenz l . s . , millar j . g . ( 2006 ) phoretic nest parasites use sexual deception to obtain transport to their host ' s nest . pnas 103 : 14039 - 14044 ( abstract )
peterson field guides : beetles richard e . white . 1983 . houghton mifflin company .
american beetles , volume ii : polyphaga : scarabaeoidea through curculionoidea arnett , r . h . , jr . , m . c . thomas , p . e . skelley and j . h . frank . ( eds . ) . 2002 . crc press llc , boca raton , fl .
the new world genera of meloidae ( coleoptera ) : a key and synopsis pinto j . d . , bologna m . a . 1999 . j . nat . hist . 33 : 569 - 620 .
contributed by troy bartlett on 16 february , 2004 - 12 : 32pm additional contributions by mike boone , cotinis , beatriz moisset , mcclarinj , chuck entz , mike quinn , aaron schusteff , ceiseman , v belov , harsi s . parker last updated 30 september , 2015 - 9 : 11pm
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detailing the physical features , habits , territorial reach and other identifying qualities of the american oil beetle .
plain and small in stature , the seemingly simple american oil beetle has a few noteworthy tricks up its sleeve .
american oil beetles are a type of blister beetle . when threatened or squeezed under pressure , they emit a chemical called cantharidin that creates blisters and irritates human skin . these wounds will heal , but they are painful . this chemical defense can ward off predators and give the beetle time to escape . american oil beetles have a soft , yet stout abdomen with a shell covering that looks like a series of overlapping plates . the insect can appear as a dull black or , in some cases , a shiny black or dark blue . the surface texture is slightly bumpy , not slick and smooth . antennae are visible on the head . these particular beetles do not fly and are slow movers . adults can be found gingerly walking around plants they eat , such as buttercups , and in grass . they are active all year , but more so in the spring , when they are more likely to be seen . the larvae are somewhat devious . one will sit on flowers , waiting for a bee to land . it will latch onto the bee for a free ride back to the hive . once there , the beetle larva feeds on the same food as the bee larvae . it will pupate safely inside and emerge in the spring .
antennae : beetles have a pair of antennae on the head used as sensors .
head : the head is home to the insect ' s eyes , antennae , and mandibles ( jaws ) .
thorax : holds the three pairs of legs as well as vital internal organs .
elytron : one of two wing cases on a beetle that protects its wings ( plural : elytra ) .
legs : beetles have three pairs of legs located at the thorax , numbering six legs in all .
an insect ' s reach is not limited by lines drawn on a map and therefore species may appear in areas , regions and / or states beyond those listed below as they are driven by environmental factors ( such as climate change ) , available food supplies and mating patterns . grayed - out selections below indicate that the subject in question has not been reported in that particular territory . u . s . states and canadian provinces / territories are clickable to their respective bug listings .
the map below showcases ( in red ) the states and territories of north america where the american oil beetle may be found ( but is not limited to ) . this sort of data can be useful in seeing concentrations of a particular species over the continent as well as revealing possible migratory patterns over a species ' given lifespan . some species are naturally confined by environment , weather , mating habits , food resources and the like while others see widespread expansion across most , or all , of north america .
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( zool . ) a genus of beetles without wings , but having short oval elytra ; the oil beetles . these beetles are sometimes used instead of cantharides for raising blisters . see oil beetle , under oil .
proscarabaeus , and gooden ' s nomad bee nomada goodeniana - both found at cwm tydu , in ceredigion .
brevicollis , was found by entomologist bob beckford on trust land in south devon .
brevicollis - were found near salcombe , south devon - more than 200 miles from their last sighting in sussex nearly 60 years ago .
( 1990 , 73 ) : one should situate oneself within the practice that the object belongs to , and then investigate the object and its contribution to that practice .
americanus ( leach ) 1 05 - 05 monotomidae pycnotomina cavicolle ( horn ) 4 05 - 05 to 06 - 03 mordellidae mordella marginata melsheimer 3 06 - 16 to 08 - 17 mordellistena aspersa ( melsheimer ) 1 06 - 09 mordellistena bihamata ( melsheimer ) 1 07 - 14 mycetophagidae litargus balteatus leconte 4 06 - 09 to 11 - 04 litargus tetraspilotus leconte 1 06 - 09 mycetophagus melsheimeri leconte 1 05 - 05 oedemeridae asclera ruficollis ( say ) 3 04 - 28 to 05 - 19 passandridae catogenus rufus ( f .
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american labor - - a 50 - year chronology ,
monthly labor review , july 1950 , pp .
franciscanus marks the first recorded case of parasitic insects cooperating to mimic their target , report hafernik and his san francisco state colleague leslie saul - gershenz .
ah , my pretty , you ' re . . . # & ! a beetle pile !
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oil beetles have fascinating life - cycles . the larvae are parasites of a number of species of ground - nesting solitary bee . towards the end of spring , female oil beetles dig burrows in the ground close to colonies of host bees , into which they lay around 1000 eggs . these eggs usually hatch the following year in order to coincide with the emergence of the bees . the oil beetle larvae ( known as tringulins ) are very active , and climb up onto flowers where they wait for a host bee . they attach themselves to the bee , and if they are lucky and attach to the right type of species they will be flown to the host’s burrow , where the tringulin oil beetle turns into a grub - like larva , and develops , feeding upon the pollen stores and eggs of the host . the larva pupates and the resulting adult beetle spends the winter inside the host’s burrow before emerging the following spring ( 4 ) .
only three of the nine oil beetle species native to britain remain , and the number of sites supporting these species has declined drastically . this beetle was once common ( 3 ) , but is now restricted to the west of britain ( 4 ) . it is also found in europe ( 2 ) .
you can view distribution information for this species at the national biodiversity network atlas .
in europe , the black oil beetle shows a preference for low - lying flat terrain ( 2 ) . in britain it is found on heaths , coastal cliffs and moors ( 1 ) .
the reasons for the decline of this species may reflect declines in host bee populations .
buglife , the invertebrate conservation trust , is currently running the oil beetle project , which aims to establish the current range of britain’s remaining oil beetles and to carry out research into their life - cycles and ecology in order to guide conservation actions targeted at these beetles ( 4 ) .
there may be further information about this species available via the national biodiversity network atlas .
this information is awaiting authentication by a species expert , and will be updated as soon as possible . if you are able to help please contact : arkive @ urltoken
abdomen in arthropods ( crustaceans , insects and arachnids ) the abdomen is the hind region of the body , which is usually segmented to a degree ( but not visibly in most spiders ) . in crustacea ( e . g . crabs ) some of the limbs attach to the abdomen ; in insects the limbs are attached to the thorax ( the part of the body nearest to the head ) and not the abdomen . in vertebrates the abdomen is the part of the body that contains the internal organs ( except the heart and lungs ) . larvae stage in an animal ' s lifecycle after it hatches from the egg . larvae are typically very different in appearance to adults ; they are able to feed and move around but usually are unable to reproduce . parasites organisms that derives food from , and live in or on , another living organism at the host’s expense . pupates pupation is the the process of forming a pupa , the stage in an insect ' s development when huge changes occur that reorganise the larval form into the adult form . in butterflies the pupa is also called a chrysalis .
harde , k . w . ( 2000 ) a field guide in colour to beetles silverdale books , leicester .
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godet y 1 , moreau - aubry a , guilloux y , vignard v , khammari a , dreno b , jotereau f , labarriere n .
institut national de santé et de recherche médicale , unité mixte de recherche 892 , 44093 nantes , france .
pmid : 18936238 pmcid : pmc2571940 doi : 10 . 1084 / jem . 20081356
t cell clone selection and characterization . ( a ) percentage of tnf - producing t cells and of hla - a2 / melan - a a27l tetramer–positive t cells in the m170 til population in response to the autologous melanoma cell line . 10 5 tils and 2 × 10 5 melanoma cells were incubated for 5 h in the presence of brefeldin a , stained with hla - a2 / melan - a a27l tetramer , fixed , and stained with anti - tnf antibody in a permeabilization buffer . 10 4 t cells were then analyzed by flow cytometry . ( b ) tnf secretion by the m170 . 48 t cell clone in response to the autologous melanoma cell line . 10 4 ctls were added to 3 × 10 4 m170 melanoma cells in the presence of blocking antibodies directed against class i , a2 , and b / c hla , diluted to 1 : 50 ( shaded bars ) , 1 : 500 ( hatched bars ) , and 1 : 5 , 000 ( open bars ) . ctl clone reactivity was assessed by a tnf release assay . ( c ) tnf response of the m170 . 48 ctl clone to hla - a * 0201 tumor cell lines . the m6 cell line ( hla - a2 negative ) was used as a negative control . ( d ) ifn - γ response of the m170 . 48 ctl clone ( shaded bars ) and of a melan - a / a2–specific ctl clone ( hatched bars ) to hla - a * 0201 melanocytes . the m170 cell line was added as a positive control .
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any oil beetles over 2 . 5 cm long with kinked anteannae will be either black or violet oil beetles though there is great size variation in all species – see photos . newly emerged oil beetles have very short abdomens , they usually feed up quickly and the abdomen swells enormously . females generally have very large bloated looking abdomens but the males can have large abdomens as well .
the males of the black and violet oil beetles have very kinked antennae ( the antennae of the females is slightly kinked ) . the rugged , mediterranean and short - necked oil beetles have straight antennae in both sexes .
violet oil beetle – square shaped thorax , indented lower edge of thorax with depressed area at base , large tooth at base of thorax , fine punctures on thorax , up to 30 mm long .
black oil beetle – square shaped thorax , almost straight lower edge of thorax which is flat ( not with depressed area at base ) , small tooth at base of thorax , large coarse punctures on thorax , up to 30 mm long .
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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 .


myrmecia midas is an australian ant which belongs to the myrmecia genus .

this species is native to australia .

myrmecia midas is distributed mainly across the coastline of several eastern australian states .

it was first described by clark in 1951 .

workers are typically 13-15 millimetres long , with the queens bigger at 18-19 millimetres and males are smaller .

the head and thorax are a red colour , gaster black , and the mandibles , antennae , and the legs are brownish red .

myrmecia midas

the above specimen data are provided by antweb . please see myrmecia midas for further details
polarized light use in the nocturnal bull ant , myrmecia midas . - pubmed - ncbi
the study site for the nocturnal ant myrmecia midas . the field site was located on the macquarie university campus . the four nests were located within 100 m of each other .
freas ca , narendra a , lemesle c & cheng k . 2017 polarised light use in the nocturnal bull ant myrmecia midas . royal society open science 4 : 170598 [ pdf ]
freas ca . wystrach a , narendra a & cheng k . 2018 . the view from the trees : nocturnal bull ants , myrmecia midas , use the surrounding panorama while descending from trees . frontiers in psychology 9 : 16 . [ pdf ]
jayatilaka p , raderschall ca , narendra a & zeil j . 2014 . individual foraging patterns of the jack jumper ant , myrmecia croslandi . myrmecological news 19 : 75 - 83 .
ramirez - esquivel f , zeil j & narendra a . 2014 . the antennal sensory array of the nocturnal bull ant , myrmecia pyriformis . arthropod structure and development 43 : 543 - 558
raderschall ca , narendra a & zeil j . 2016 . head roll stabilisation in the nocturnal bull ant myrmecia pyriformis : implications for visual navigation . journal of experimental biology 219 : 1449 - 1457
narendra a & ribi wa . 2017 . ocellar structure is driven by the mode of locomotion and activity time in myrmecia ants . journal of experimental biology 220 : 4383 - 4390 . [ pdf ] featured inside jeb
narendra a , greiner b , ribi wa & zeil j . 2016 . light and dark adaptation mechanisms in the compound eyes of myrmecia ants that occupy discrete temporal niches . journal of experimental biology 219 : 2435 - 2442 . featured inside jeb
experiments were conducted from april 2015 to october 2016 on four m . midas nests located at the northern end of the macquarie university campus in sydney , australia ( 33°46′11″s , 151°06′40″e ) . all testing was conducted on nests that were located within a 100 m area ( fig . 1 ) . the area ' s vegetation consisted of stands of eucalyptus trees with largely barren understories interspersed with grassy areas . myrmecia midas nests were located at or close to the base of a tree ( 25 . 6±4 . 36 cm mean±s . d . , n = 20 ) , and a portion of the nest ' s nightly foragers ( ∼30 % ) travelled straight up the nest tree while the remaining foragers travelled to one of the surrounding trees ( observations : c . a . f . , three m . midas nests ) . we used headlamps with red filters to observe the ants at night , and forager behaviour suggests they were not affected by this light . working with this ant species requires no animal ethical approval within australia . all collection and testing procedures were non - invasive and we witnessed no adverse effects either in the tested individuals or in their nests during or after testing . all displacement collection and testing procedures occurred during the evening or morning twilight .
here , we investigated the compass cues used by a nocturnal bull ant , myrmecia midas clark 1951 . first , we determined their daily activity schedule . next , we tested whether foragers use a home vector to orient in the absence of familiar landmark information , by displacing foragers to unfamiliar locations . we then asked how ants weight their home vector and terrestrial cues by creating a conflict between the two cue sets . to explore the possibility that foragers navigate using cues beyond the surrounding landmarks , we displaced foragers locally near the nest with the surrounding landmark panorama obscured . finally , we compared the navigational knowledge of ants when they had to travel different distances ( 0 . 3–14 . 0 m ) from the nest to reach their foraging tree .
nightly foraging activity pattern of m . midas foragers . the observation period began on 2 april 2016 with sunset at 18 : 49 h . bars indicate outgoing foragers ( blue , n = 171 ) and incoming foragers ( red , n = 169 ) . the black line indicates the number of individuals actively foraging away from the nest site during this period . data are shown in 10 min bins .
myrmecia midas appear to detect and use both path integration and the surrounding terrestrial landmarks to navigate at low light levels during twilight . our results suggest that vector cue strength is weighted by vector length , as only at the longest observed vector lengths did foragers orient successfully in unfamiliar surroundings . the lack of orientation to smaller ( 2–5 m ) home vectors in the presence of unfamiliar terrestrial cues and the very weak orientation at the maximum observed natural vector ( 12 . 8–14 . 0 m ) coupled with the lack of orientation to all vector lengths ( 2–14 m ) when in the presence of conflicting familiar terrestrial cues indicates that vector cues may be weakly integrated during orientation and can be overridden by stronger terrestrial cues or that these foragers do not path integrate over short distances . additionally , terrestrial visual cues appear critical to successful homeward orientation , as blocking these cues in local environments results in foragers no longer being oriented to the nest direction . finally , our results show that a portion of m . midas ' foraging force lacks the ability to navigate back to the nest site using landmark cues over short distances , and that these ants may be unable to extrapolate beyond the nest site panorama even over short distances if they have only experienced the terrestrial panorama immediately surrounding the nest site .
circular distributions of individual m . midas foragers ' headings during the long - vector 1–2 m inbound condition . histograms show raw data of exit orientation under the filter with the individual ' s initial orientation and reorientation with the forager ' s exit orientation under the filter . the triangle denotes 45° in each distribution . the arrow denotes the length of the mean vector direction . n , number of individuals ; ø , mean vector ; r , length of the mean vector .
the overnight and morning inbound testing conditions were chosen so as to mirror the typical inbound activity patterns of this species , with foragers being released during the pre - dawn twilight when motivation to return to the nest should be high . our 15 min delay conditions were chosen as this holding period allows time for the forager to feed in the collection tube while still being tested during twilight , as the navigational abilities of nocturnal myrmecia have been shown to suffer after twilight ends ( narendra et al . , 2013a , b ) .
we carried out a 24 h observation at one m . midas nest on 2 april 2016 to establish the species ' activity pattern . we set up a 60 cm diameter reference marker around the nest entrance and filmed ants departing and entering the nest with an hd c920 webcam ( logitech international ) . a lamp covered with a red filter was placed 50 cm away from the nest entrance . filming was carried out for a 24 h period from 1 h before sunset . astronomical data were obtained from calculations in the astronomical almanac ( urltoken ) .
circular distributions of individual m . midas foragers ' headings during the inbound conditions . histograms show raw data of exit orientation under the filter with the individual ' s initial orientation and reorientation with the forager ' s exit orientation under the filter . the triangle denotes 45° in each distribution . the arrow denotes the length of the mean vector and mean direction . ( a ) orientations for nest 1 during the 4–6 m inbound condition . ( b ) orientations for the 1–2 m inbound condition . n , number of individuals ; ø , mean vector ; r , length of the mean vector .
solitary foraging ants have a navigational toolkit , which includes the use of both terrestrial and celestial visual cues , allowing individuals to successfully pilot between food sources and their nest . one such celestial cue is the polarization pattern in the overhead sky . here , we explore the use of polarized light during outbound and inbound journeys and with different home vectors in the nocturnal bull ant , myrmecia midas . we tested foragers on both portions of the foraging trip by rotating the overhead polarization pattern by ±45° . both outbound and inbound foragers responded to the polarized light change , but the extent to which they responded to the rotation varied . outbound ants , both close to and further from the nest , compensated for the change in the overhead e - vector by about half of the manipulation , suggesting that outbound ants choose a compromise heading between the celestial and terrestrial compass cues . however , ants returning home compensated for the change in the e - vector by about half of the manipulation when the remaining home vector was short ( 1 - 2 m ) and by more than half of the manipulation when the remaining vector was long ( more than 4 m ) . we report these findings and discuss why weighting on polarization cues change in different contexts .
observations of forager activity at the nest indicate that a portion ( ∼30 % ) of m . midas foragers do not travel farther than 30 cm to reach their foraging tree . to determine the navigational knowledge of these foragers , we displaced outbound full - vector ants 5 m away from the nest , either directed toward the nest entrance ( nest - side condition ) or directed toward the tree ( tree - side condition ; see fig . 2 b ) . foragers were collected on their outbound trip as they reached the base of the foraging tree . these individuals were fed , held for 15 min , and then released on the goniometer at one of the displacement sites . foragers ' orientations at 20 cm were determined at nests 1 , 2 and 3 . after testing , individuals were marked and returned to the nest .
circular distributions of individual m . midas foragers ' headings during outbound conditions . histograms show raw data of exit orientation under the filter and the reorientation after exiting the filter . the triangle denotes 45° in each distribution . the arrow denotes the length of the mean vector and the mean direction . ( a ) orientations for nest 1 during the 4–6 m outbound condition . ( b ) orientations for nest 2 during the 4–6 m outbound condition . ( c ) orientations for nest 1 during the 1–2 m outbound condition . closed circles indicate individuals that continued on to the forging tree after testing . open circles represent foragers that retreated once the filter was placed overhead and these individuals returned to within 30 cm of the nest entrance after testing . n , number of individuals ; ø , mean vector ; r , length of the mean vector .
circular histograms of initial headings of individual m . midas foragers at 20 cm in nest - tree experiments . nest location for both conditions was set at 0 deg . histograms show orientation data in 15 deg bins . the arrow denotes the length of the mean vector direction and the filled triangle designates the true nest direction . outbound foragers travelling up the nest - tree were collected at the tree base and displaced 5 m from the nest site either ( a ) on the nest side of the tree ( nest - side condition ; v - test at 0 deg ; v = −0 . 04 , p = 0 . 101 ) or ( b ) with the tree in between the nest site and the displacement site ( tree - side condition ; v - test at 0 deg ; v = 0 . 143 , p = 0 . 639 ) .
circular histograms of initial headings of individual m . midas foragers at 20 cm in unfamiliar location experiments . nest direction indicated by path integration was at 0 deg and the actual nest site direction was at 180 deg . histograms show orientation data in 15 deg bins . the arrow in each histogram denotes the length of the mean vector and the direction of the average orientation . the filled triangle in each histogram indicates the true nest direction and the open triangle indicates the vector direction . ( a ) foragers were collected during their morning inbound trip and displaced to a site 100 m away ( inbound 100 m condition ; v - test at 0 deg ; v = 0 . 085 , p = 0 . 225 ) . ( b ) foragers were collected during the outbound trip , at the base of the foraging tree , 2–5 m from the nest entrance , held for 15 min and then released 100 m from the nest site ( outbound 15 min delay 100 m condition ; v - test at 0 deg ; v = −0 . 139 , p = 0 . 906 ) . ( c ) foragers were collected during their outbound trip , held for 12 h , and displaced to a site 100 m away ( outbound overnight delay 100 m condition ; v - test at 0 deg ; v = 0 . 083 ; p = 0 . 240 ) . ( d ) foragers were collected during the outbound trip with the longest observed vectors ( 12 . 8–14 . 0 m ) , held for 15 min and displaced to a site 100 m away ( outbound long vector 15 min delay condition ; v - test at 0 deg ; v = 2 . 361 , p = 0 . 009 ) . n , number of individuals in each condition ; ø , mean vector ; r , length of mean vector .
circular histograms of initial headings of individual m . midas foragers at 20 cm in local displacement experiments . nest direction indicated by the home vector was at 270 deg and the actual nest site direction was at 0 deg . histograms show orientation data in 15 deg bins . the arrow denotes the length of the mean vector direction . in all off - route conditions , the filled triangle indicates the true nest direction and the open triangle indicates the vector direction . ( a ) outbound foragers were collected at the base of the foraging tree , held for 15 min and then released at the base of the tree on their foraging route ( outbound 15 min delay on - route condition ; v - test at 0 deg ; v = 0 . 351 , p < 0 . 001 ) . ( b ) inbound foragers were collected during their morning inbound trip and displaced to a site 5 m away ( inbound off - route condition ; v - test at 0 deg ; v = 0 . 703 , p < 0 . 001 ) . ( c ) outbound foragers were collected at the base of the foraging tree 2–5 m from the nest entrance , held for 15 min and then released at the 5 m displacement site ( outbound 15 min delay off - route condition ; v - test at 0 deg ; v = 0 . 580 , p < 0 . 001 ) . ( d ) foragers were collected during their outbound trip , held for 12 h and then displaced to a site 5 m away ( outbound overnight delay off - route condition ; v - test at 0 deg ; v = 0 . 554 , p < 0 . 001 ) . ( e ) outbound foragers were collected at the base of their foraging tree 14 m from the nest entrance , held for 15 min and then released at the 5 m displacement site ( outbound long vector 15 min delay off - route condition ; v - test at 0 deg ; v = 0 . 618 , p < 0 . 001 ) . nest direction indicated by path integration was at 270 deg and is marked with an open triangle . ( f ) outbound foragers were collected at the base of their foraging tree 2–5 m from the nest entrance , held for 15 min and then released at the 5 m displacement site with the surrounding landmarks blocked from the forager ' s view ( landmark blocking 15 min delay condition ; v - test at 0 deg ; v = −0 . 04 , p = 0 . 631 ) .
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holotype , worker , dorrigo , new south wales , australia , froggatt , w . w . , anic32 - 006507 , australian national insect collection .
paratype , 1 worker , tambourine mtn , queensland , australia , hacker , h . , anic32 - 006513 , australian national insect collection .
paratype , 4 workers , dorrigo , new south wales , australia , heron , w . , anic32 - 012805 , australian national insect collection .
clark , j . 1951 . the formicidae of australia . 1 . subfamily myrmeciinae : 230 pp . csiro , melbourne . [ ( 31 . xii ) . 1951 . ] pdf
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clark , 1951 pdf : 55 , figs . 34 , 35 ( w . q . )
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these vision - based navigational abilities have been widely studied in diurnal ants , which are active when visual cues are easy to distinguish ( wehner et al . , 1996 ; fukushi , 2001 ; beugnon et al . , 2005 ; cheng et al . , 2009 ; bühlmann et al . , 2011 ) . nocturnal foragers have the challenge of navigating during twilight or at night when ambient light levels drop significantly and visual cues become increasingly difficult to detect ( warrant , 2008 ; warrant and dacke , 2011 ) . yet , little is known about navigation in ant species that forage at low light intensities .
we displaced full - vector foragers in three conditions . in the first , we collected inbound homing ants at the base of the tree during the morning twilight as they travelled down the foraging tree ( inbound 100 m condition ) . in the second condition , we collected individuals during their nightly outbound trip at the base of the foraging tree , held them for 15 min with a small amount of honey and released them at the distant displacement site ( outbound 15 min delay 100 m condition ) . in the third condition , we collected outbound foragers during the evening twilight , offered them a small amount of honey and stored them overnight ( 11–12 h ) . these foragers were tested in the morning twilight ( outbound overnight delay 100 m condition ) . after testing and marking , foragers were returned to the nest site . foragers ' orientations did not differ between nests and results were pooled across all four nests .
additionally , we determined whether the length of the home vector itself affected the ability to orient using path integration . for this test , we studied a subset of ants from nest 1 and nest 3 that foraged on a tree 14 . 0 and 12 . 8 m from the nest entrance , respectively , which were the maximum observed foraging distances at any nest at this site , distances we term long vector . we again collected outbound foragers at the base of the foraging tree , but only during the evening twilight . foragers were held for 15 min , and given a small amount of honey , then released at the 100 m displacement site ( outbound long vector 15 min delay 100 m condition ) , where we recorded their initial orientations . tested foragers were marked and returned to the nest site . foragers ' orientations from the two nests were pooled .
next , we asked how ants weight path integration and landmark cues while returning home to the same four nests . we addressed this by capturing ants at the base of their foraging tree under identical collection conditions to the unfamiliar displacements and then displacing them locally . we first tested foragers in a control condition without creating a conflict between the home vector direction and the local landmarks . for this , we released outbound ants during the evening twilight after a 15 min holding period on the goniometer at the base of their foraging tree ( outbound 15 min delay on - route condition ) . we then tested a separate group of foragers with conflicting terrestrial landmarks and the home vector by displacing foragers away from the nest site and perpendicular to the foraging route . the displacement site was 90 deg clockwise from the individual ' s foraging tree with respect to the nest location and 5 m from the nest entrance ( fig . 2 a ) .
schematic diagram of the experimental setup . ( a ) diagram of collection and displacement conditions for the experiments on cue conflict . ( b ) diagram of collection and displacement conditions for the nest - tree foragers .
foragers were tested in three separate conditions , similar to the procedures for testing the use of path integration . in the first condition , incoming foragers were collected during the morning twilight at the base of their foraging tree ( inbound off - route condition ) and then tested . in the second and third conditions , foragers were collected on their outbound trip as they reached one of the foraging trees 2–5 m from the nest ; foragers were either held for 15 min ( outbound 15 min delay off - route condition ) and then tested or held until morning twilight ( outbound overnight delay off - route condition ) and then tested . at the testing site , under all conditions the forager ' s initial orientation was recorded using the goniometer at 20 cm . after testing , individuals were marked and returned to the nest entrance . we analysed foragers ' initial orientations toward both the true nest direction and the vector from path integration . orientations were pooled across all four nests .
again , we determined whether the length of the home vector itself affected the weighting of the home vector . for this test , we studied a subset of ants ( n = 30 ) from nest 1 that foraged on a tree 14 . 0 m from the nest ( outbound long vector 15 min delay off - route condition ) . we collected outbound foragers at the base of the foraging tree , but only during the evening twilight . foragers were held for 15 min , given a small amount of honey , and then released at a displacement site 90 deg clockwise from the foraging tree with respect to the nest and 5 m from the nest entrance ( fig . 2 a ) . at the release site , we recorded initial orientations , and then tested foragers were marked and returned to the nest site .
next , we asked whether foragers successfully orient to the nest direction after local displacement in the absence of familiar terrestrial landmark cues . for this test , we collected outbound foragers at the base of their foraging tree 2–5 m from nest 1 , nest 3 or nest 4 during the evening twilight ( landmark blocking 15 min delay ) . foragers were held for 15 min , offered a small amount of honey , and then released at a displacement site 90 deg clockwise from the foraging tree with respect to the nest and 5 m from the nest entrance ( fig . 2 a ) . at the release location , a 38 cm high plastic sheet was used to block the landmark cues below 50 deg . the plastic sheet was erected into a 55 cm diameter circle around the goniometer . this sheet was raised 5 cm off the ground to allow any potential scent cues to reach foragers at the centre of the goniometer . after releasing the forager onto the goniometer , we recorded the orientation of animals at 20 cm . after testing , individuals were marked and returned to the nest .
for statistical analysis , a minimum of 30 foragers were collected for each condition . in each testing condition , foragers were collected as they were discovered and assigned a condition within that experiment group randomly . data were analysed with circular statistics ( batschelet , 1981 ; zar , 1998 ) using the statistics package oriana version 4 ( urltoken ) . watson and wheeler f - tests were used to compare mean vectors between testing conditions . a rayleigh ' s test was conducted on each displacement condition , to test whether the data met the conditions of a uniform distribution ( p > 0 . 05 ) . we used a v - test , with α set at p = 0 . 05 , to determine whether the initial orientation of ants at the release site was significantly towards the nest direction or to the home vector . during displacement tests where foragers did not orient to the nest direction , v - tests were conducted across all 15 deg sections of the goniometer to test whether foragers were oriented to directions other than the nest . all data are available upon request .
the onset of foraging activity occurred just before sunset , with most foragers heading out from the nest in the evening twilight ( fig . 3 ) . a small number of foragers left the nest after twilight ended . foragers began returning during the evening twilight , with some individuals carrying insect prey . foragers returned throughout the night , and a burst of incoming foragers arrived at the nest during the morning twilight and just after sunrise . activity was greatly reduced after this burst , and activity ceased completely after 1–2 h from sunrise .
ants released on the goniometer would initially remain motionless while within the tube . after emerging from the tube and onto the wooden goniometer board , ants typically scanned the environment along the horizontal axis and then headed off in a chosen direction . the nest - tree distance varied between nests ( fig . 1 ; nest 1 , 2 m ; nest 2 , 2 . 5 m ; nest 3 , 4 m ; nest 4 , 5 m ) , and we found no difference between nests in forager orientation to a home vector .
ants with home vectors ranging from 2 to 5 m when displaced to distant locations did not successfully orient in the direction indicated by their path integrator under any collection condition . foragers ' initial orientations met conditions of a uniform distribution and they were not significantly oriented in the direction of their home vector at 0 deg ( fig . 4 a–c , table 1 ) . foragers from all three conditions were not oriented in any direction on the goniometer , even when using v - tests ( p > 0 . 05 , orientation data binned at 15 deg ) . further observations of individuals with 5 m vectors after distant displacement indicate that they do not continue in this initial direction but loop back to the release point , a behaviour indicative of systematic search .
in contrast , ants that travelled long distances to their nest - specific tree ( nest 1 and 3 ) and were then displaced to the distant site within 15 min of capture were significantly orientated toward their home vector and away from the true nest location when analysed using a v - test . yet , forager orientations also met the conditions of a uniform distribution when analysed using a rayleigh ' s test , indicating random initial headings and no orientation . as results from nest 1 and nest 3 did not differ ( v - test , p < 0 . 05 ; rayleigh ' s test , p > 0 . 05 ) and had similar mean vectors ( watson and wheeler f - test , p > 0 . 05 ) , they were pooled ( table 1 , fig . 4 d ) . at the end of each experiment , foragers were returned to the nest vicinity , and all foragers immediately searched for and entered the nest .
for local on - and off - route displacement tests , full - vector ants were displaced either back at their foraging tree or off - route ( fig . 2 a ) . outbound forager orientations when displaced back on - route after 15 min were non - uniform and were directed towards the nest ( outbound 15 min delay on - route condition ; fig . 5 a , table 1 ) . the orientation of full - vector ants , displaced ( 5 m off - route ) to create a conflict between the vector and local landmark cues , was non - uniform . these ants were significantly oriented toward the nest ( fig . 5 b–d ) and not toward the home vector at 270 deg ( inbound off - route condition , v - test , v = 0 . 776 , p = 0 . 22 ; outbound 15 min delay off - route condition , v - test , v = 0 . 844 , p = 0 . 34 ; outbound overnight delay off - route condition , v - test , v = −0 . 771 , p = 0 . 779 ) . in all local displacement conditions , mean vector did not significantly differ between conditions ( watson and wheeler f - test , p > 0 . 05 ) .
the orientation of long - vector foragers , accumulating 14 m vectors to their nest - specific tree ( nest 1 ) , then displaced locally with a 90 deg cue conflict within 15 min of collection ( outbound long vector 15 min delay off - route condition ) was also significantly non - uniform and oriented toward the true nest direction and not the vector at 270 deg ( table 1 , fig . 5 e ) . different holding durations did not have an effect on the heading directions in all off - route conditions ( watson and wheeler f - test ; p > 0 . 05 ) .
the initial orientation of foragers displaced locally without access to familiar landmark cues within 15 min of collection met the conditions of a uniform distribution , and ants were not significantly oriented in the direction of their nest at 0 deg ( table 1 , fig . 5 f ) .
full - vector ants , when displaced 5 m either side of the nest ( fig . 2 b ) , did not orient significantly toward the nest direction . their initial orientation had a uniform distribution at both displacement sites ( nest - side condition , fig . 6 a ; tree - side condition , fig . 6 b ; table 1 ) . foragers from both conditions were not oriented in any direction or toward any other tree ( v - tests , p > 0 . 05 ) . at the end of the experiment , foragers were returned to the nest vicinity , and all foragers immediately searched for and entered the nest .
in the current study , we show that nocturnal ants with short home vectors of ≤5 m when displaced to distant unfamiliar locations did not orient toward the fictive nest . but ants with longer home vectors of 12 . 8–14 . 0 m weakly oriented toward the fictive nest , suggesting vector length influences cue strength . additionally , when foragers were displaced locally with conflicting vector and landmark cues , they oriented exclusively using landmark cues and ignored any potential home vector regardless of vector length . when the landmark panorama ( up to 50 deg elevation ) around the displacement site was obscured , foragers could no longer orient to the nest direction , suggesting terrestrial cues are critical to homeward orientation . finally , we found that the nest - tree forager subset did not orient to local landmark cues when displaced , and these animals appear to have reduced navigational knowledge compared with their nest mates that travel away from the nest to forage .
conceived and designed experiments : c . a . f . , a . n . , k . c . collected data : c . a . f . drafted and revised paper : c . a . f . , a . n . , k . c .
this research was supported by the australian research council through a discovery grant to k . c . and a . n . ( dp150101172 ) and a future fellowship to a . n . ( ft140100221 ) .
homing strategies of the australian desert ant melophorus bagoti i . proportional path integration takes the ant half - way home
homing strategies of the australian desert ant melophorus bagoti ii . interaction of the path integrator with visual cue information
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r soc open sci . 2017 aug 30 ; 4 ( 8 ) : 170598 . doi : 10 . 1098 / rsos . 170598 . ecollection 2017 aug .
freas ca 1 , narendra a 1 , lemesle c 1 , cheng k 1 .
department of biological sciences , macquarie university , sydney , new south wales 2109 , australia .
pmid : 28879002 pmcid : pmc5579118 doi : 10 . 1098 / rsos . 170598
schematics of the polarization filter and experimental set - up . ( a ) diagram of the polarization filter . during the forager ' s outbound trip to the foraging tree , a polarization filter was placed over the forager with the polarization e - vector rotated ±45° of the ambient e - vector . this filter apparatus was used in a previous study [ ] . ( b ) diagram of measurements collected during polarization filter test . measurements were made using a compass application on a smartphone . initial orientation routes were measured from the nest entrance ( a ) to when the polarization filter was centred over the forager ( b ) . initial route directions ( a ° ) were calculated with the tree direction from the nest as 0° . the magnitude of angle a has been artificially enlarged in this diagram for clarity , with angle a averaging 4 . 42° across all conditions during testing . exit orientations were measured from the centre of the polarization filter ( b ) to the exit location of the ant on the filter ' s edge ( c ) . route directions under the filter ( b ° ) were calculated from the forager ' s initial route direction . reorientations were measured from the forager ' s exit location from the polarization filter ( c ) to the forager ' s path 50 cm after exiting the filter ( d ) . reorientation route directions ( c ° ) were calculated from the under - filter route direction .
r soc open sci . 2017 aug ; 4 ( 8 ) : 170598 .
i am interested in how animals navigate in their natural habitats , the information content in the environment they use , the sensory structures they have and the central processing units that allow them to memorise , integrate and recall information .
my research has largely focused on ants to take advantage of the inter - and intra - specific variation in size , in locomotion , in tasks and also in the different ecological and temporal niches they occupy .
i have a keen interest in photography and love to experiment with macro - photography when time permits . some of my pictures are here .
phd in biological sciences ; 1 book ; 47 publications ; 3 australian research council fellowships ; 10 research grants .
2016 . macquarie research and development grant ( with lead ci frances kamhi ) , macquarie university , australia . ( $ 50k )
palavalli - nettimi r and narendra a . 2018 . does size affect orientation usage celestial cues . insects sociaux . [ in press ]
palavalli - nettimi r and narendra a . 2018 . miniaturisation decreases visual navigational competence in ants . journal of experimental biology . doi : 10 . 1242 / jeb . 177238
ramirez - esquivel f , ribi wa and narendra a . 2017 . techniques to investigate the anatomy of the ant visual system . journal of visualized experiments 129 : e56339 . [ pdf ]
narendra a , kamhi jf & ogawa y . 2017 . moving in dim light : behavioural and visual adaptations in nocturnal ants . integrative and comparative biology 57 : 1104–1116 [ pdf ]
ramirez - esquivel f , leitner ne , zeil j & narendra a . 2017 . the sensory arrays of the ant , temnothorax rugatulus . arthropod structure and development 46 : 552 - 563 [ pdf ]
narendra a & ramirez - esquivel f . 2017 . subtle changes in the landmark panorama disrupts visual navigation in a nocturnal bull ant . philosophical transactions of the royal society b 372 : 20160068 . [ pdf ]
freas ca , narendra a & cheng k . 2017 . compass cues used by a nocturnal bull ant ,
card a , mcdermott c & narendra a . 2016 . multiple orientation cues in the trunk trail forming ant , iridomyrmex purpureus . australian journal of zoology 64 : 227 - 232 .
narendra a , ramirez - esquivel f & ribi wa . 2016 . compound eye and ocellar structure for walking and flying modes of locomotion in the australian ant , camponotus consobrinus . scientific reports 6 : 22331
schultheiss p , raderschall ca , narendra a . 2015 follower ants in a tandem pair are not always naïve . scientific reports 5 : 10747
ogawa y , falkowski m , narendra a , zeil j , hemmi jm . 2015 . three spectrally distinct photoreceptors in diurnal and nocturnal australian ants . proceedings of the royal society b 282 : 20150673
stürzl w , grixa i , mair e , narendra a & zeil j . 2015 . three - dimensional models of natural environments and the mapping of navigational information .
zeil j , ribi wa & narendra a . 2014 . polarization vision in ants , bees and wasps . in polarized light and polarization vision in animal sciences ( ed . horváth g ) . springer series in vision research ( eds . collin s & marshall j ) . details here .
cheung a , collett m , collett ts , dewar a , dyer f , graham p , mangan m , narendra a , philippides a , stürzl w , webb b , wystrach a & zeil j . 2014 . still no convincing evidence for cognitive map use by honeybees . proceedings of the national academy of sciences usa 111 : 4396–4397
zeil j , narendra a & stürzl w . 2014 . looking and homing : how displaced ants decide where to go . philosophical transactions of the royal society b 369 : 20130034