the rufous-tailed jacamar ( galbula ruficauda ) is a near-passerine bird which breeds in the tropical new world in southern mexico , central america and south america as far south as southern brazil and ecuador .

the jacamars are elegant brightly coloured birds with long bills and tails .

the rufous-tailed jacamar is typically 25 centimetres ( 10 in ) long with a 5 centimetres ( 2 in ) long black bill .

the subspecies g. r. brevirostris has , as its name implies , a shorter bill .

this bird is metallic green above , and the underparts are mainly orange , including the undertail , but there is a green breast band .

sexes differ in that the male has a white throat , and the female a buff throat ; she also tends to have paler underparts .

the race g. r. pallens has a copper-coloured back in both sexes .

this species is a resident breeder in a range of dry or moist woodlands and scrub .

the two to four rufous-spotted white eggs are laid in a burrow in a bank or termite mound .

this insectivore hunts from a perch , sitting with its bill tilted up , then flying out to catch flying insects .

one commonly preyed upon insect is the social wasp agelaia vicina .

further , the bird distinguishes between edible and unpalatable butterflies mainly according to body shape .

the rufous-tailed jacamar 's call is a sharp pee-op , and the song a high thin peeo-pee-peeo-pee-pe-pe , ending in a trill .

rufous - tailed jacamar

protection / threats / status : rufous - tailed jacamar is common and widespread in its range . the major threats for this species are deforestation and habitat loss . rufous - tailed jacamar populations are not threatened at this moment .
a lateral view of a perched male ' rufous - and - green jacamar ' .
the commonest species is the rufous - tailed jacamar ( galbula ruficauda ) , 25 cm ( 10 inches ) long , found from southern mexico to argentina .
the rufous - tailed jacamar inhabits a range from central america to south america as far as brazil and northern argentina . the beauty of its shimmering green upperparts and rufous underparts cannot be overlooked . while the male’s throat is white , the female’s is rufous .
range : rufous - tailed jacamar is widespread on a large range , from southern mexico to northern argentina , including mexico , costa rica , ecuador , colombia , brazil , argentina , and trinidad and tobago .
behaviour : rufous - tailed jacamar feeds mainly on insects caught on the wing , by flycatching , aerial sallies or pursuit in the air . when the prey is caught , the bird beats it against a branch or other hard surface , in order to kill it , and to remove the more resistant parts of the body and the wings . then , the prey is swallowed whole . this behaviour is observed only for relatively large preys . rufous - tailed jacamar hunts from an exposed perch .
habitat : rufous - tailed jacamar frequents several kinds of habitats , such as shrubby forest edges , gallery forests , second - growth thickets , streams and rivers’ banks , marshes and savannahs with scattered trees . it may be seen from lowlands up to 900 metres of elevation , rarely up to 1300 metres .
flight : rufous - tailed jacamar has swift flight , fairly brief when pursuing preys . if the bird needs to fly long distances , for escaping a danger or between two feeding areas , it flies with bursts of rapid wing beats and glides . at this moment , the flight seems to be undulating .
some might say that the rufous - tailed jacamar ( galbula ruficacauda ) is reminiscent of a green - and - rufous kingfisher for its similar color patterns and heavy bill whereas others might see this bird and think of it as an oversized hummingbird . whatever side you fall on it’s a species that you can’t seem to get enough of and is hard to turn away from for its brilliant colors and sassy attitude .
description : rufous - tailed jacamar is a brightly coloured bird . adult male has shining metallic coppery - green plumage on upperparts . on the wings , the primaries are blackish . the long , graduated tail shows two longer rectrices . the other tail feathers are cinnamon . on the underparts , chin and throat are white . breast shows a broad metallic coppery - green band . belly , vent and undertail are rufous - chestnut . underwing coverts are cinnamon and flight feathers are dark grey .
diet : rufous - tailed jacamar feeds mainly on insects caught by flycatching and sallies . the diet includes a wide variety of insects , even wasps . the bird perches 1 to 3 metres above the ground on relatively exposed perch . when the prey is caught , it beats it several times against the branch , in order to kill it and to remove the indigestible parts before swallowing .
earlier this month at springhill , a high - pitched call of pee - pee - pee alerted us to the presence of a rufous - tailed jacamar nearby . turning out to be just in front of springhill’s main house , employees and guests alike were delighted to spot trinidad and tobago’s only representative of the galbulidae ( jacamar ) family . jacamars are neotropical birds which are similar to old world bee - eaters . they look like oversized hummingbirds and share a few characteristics with ‘hummers’ . they have iridescent feathers , long , thin bills and a fullness of energy .
there are 15 species of jacamar in tropical america which belong in their own family galbulidae situated between the puffbirds and toucans . all belong to the order piciformes along with woodpeckers and honeyguides . there are two species found here in costa rica , the great jacamar ( jacamerops aurea ) found on the caribbean slope and the rufous - tailed jacamar found at low elevations on both caribbean and pacific slopes . they are fairly common here on the osa peninsula mainly found along forest edges , streams and the open understory of tall second growth forests . it is also known to frequent cocoa plantations . we recently saw one perched along a road near one of our avian monitoring points on friends of the osa property in an area that we know as arbolito . it is not as abundant as some species and so it is a special moment to come across one .
voice : sounds by xeno - canto rufous - tailed jacamar’s call is a sharp , thin “peeup” repeated 3 to 10 times by both sexes . it also utters a rapid trill . song is a rapid phrase of several notes ending in trill “peeo - peeo - peea - pee - pee - pe - pe - pe - pe - pe - illllll” . this prolonged song uttered by the male may vary , including melodious or buzzy parts , twittering , ascending trills and low “pee - pee - pee” .
rufous - tailed jacamars breed , in trinidad , from february to june and in tobago , from february to august , where i might add , they are quite common . both sexes share nesting duties and interestingly , during courtship males remove the insects’ wings to feed them to females . clutches of two to four white eggs with cinnamon spots , are laid .
rufous - tailed jacamar nests in burrows excavated by both sexes . it is situated in sandy banks or termitary , and even in higher roots of fallen trees , from the ground up to three metres high . the burrow is usually straight , and the nest - chamber is sometimes partially visible . if there is any obstruction such as roots , the angle may deviate as sideway or downwards . the entrance is usually rectangular with vertical sides . the burrow is about 20 to 50 cm deep , with terminal cavity for the nest - chamber which is unlined .
locally called the ‘king hummingbird’ , the rufous - tailed jacamar is common in humid lowlands – on forest edges , in clearings and in secondary forest . it often perches a few metres above ground alongside a road , narrow stream or other type of clearing . there , it waits for prey , mostly flying insects , for which it hawks then thrashes against a branch in order to de - wing . these lively birds are regularly seen dust bathing on gravel roads . they measure 26 cm in length and usually nest in short tunnels on earth banks or even in termite nests .
rufous - tailed jacamars dig long burrows ( 11 - 20” ) in a bank , the root mass of an upturned tree or a termitary in which to put their nest . parents feed their 2 – 4 young during a 3 week nestling period by regurgitating insect parts . by the time the nestlings are ready to fledge the nest can look like a tomb of glittery insect wings and chitinous body parts .
it is highly sedentary , only performing some short dispersal , but it does not migrate . rufous - tailed jacamar is very active , often seen in pairs in shrubs , hunting for insects . during breeding season , the male performs regular courtship - feeding . it catches insects for its mate , offered without wings or hard parts . both mates perch close to each other while the male bobs its head upwards , downwards and sideways , and swings the tail up and down . at this moment , the male utters low notes and female gives trills . at other moment , always perched side to side , both mates fan their tails and bow together .
vary slightly in the amounts of black on the chin and in the number of green central rectrices , but in general males are an iridescent coppery / golden green above with a white throat and cinnamon - rufous underparts . females are a slightly duller green and have a cinnamon - buff throat .
feed almost exclusively on flying insects , especially dragonflies , butterflies and moths . these birds forage from a perch on an exposed branch 1 to 3 meters from the ground , and sally out to catch insects on the wing . after the jacamar has caught an insect it beats it several times against a branch to stun it and remove the insect ' s wings before it swallows .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
sacc . 2005 and updates . a classification of the bird species of south america . available at : urltoken .
justification : this species has an extremely large range , and hence does not approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . despite the fact that the population trend appears to be decreasing , the decline is not believed to be sufficiently rapid to approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size is very large , and hence does not approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
partners in flight estimate the total population to number 500 , 000 - 4 , 999 , 999 individuals ( a . panjabi in litt . 2008 ) . trend justification : despite this species tolerance of degraded and man - made habitats declines have been reported in panama and costa rica owing to hunting pressure , human interference and habitat loss ( del hoyo et al . 2002 ) .
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cuvier 1816 ) is a beautiful inhabitant of forest edges and clearings of central and south america . it occurs in several disjunct populations : from eastern mexico south to western panama ; from eastern panama south to western ecuador , colombia and venezuela ; in guyana ; and from bolivia east to eastern brazil . the six recognized subspecies of
) , version 1 . 0 . in neotropical birds online ( t . s . schulenberg , editor ) . cornell lab of ornithology , ithaca , ny , usa .
hbw alive contains information on descriptive notes , voice , habitat , food and feeding , breeding , movements , status and conservation plus a list of bibliographical references for this species account .
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josep del hoyo , pieter de groot boersma , greg baker , carlos gussoni , david ascanio , lucas andrei campos - silva , keith blomerley , antonio silveira , max roth , alex garcía · videoaves , thore noernberg , peter nash , keith and lynn youngs , jacob . wijpkema , cedric prys - roberts , pere sugranyes , richard garrigues , manakincarmelo , daniêl jimenez , mauricio rueda , rigdon currie , yaiza aragón , yoël jimenez , eldert groenewoud , monicayhector , pedro h ramos , rodrigo y castro , sidnei sampaio , agustin carrasco .
margareta wieser , carlos gussoni , lars petersson , luis r figueroa , antonio silveira , lmarce , richardgreenhalgh031 , manakincarmelo , lsargentini , phillip edwards , steve garvie , sanjiv parasram , joe tobias , philgunson , stephen romany , josep del hoyo , kperezleon , jacqueserard , rolando chavez , lutz duerselen , ian barker , anselmo d affonseca , mikko pyhälä , miriam bauman , paul van giersbergen , róger rodríguez , netosevero , nimali digo and thilanka edirisinghe , delareina , hal and kirsten snyder , mauricio rueda , orlando jarquín g . , nigel lallsingh , eduardo freitez gassan , gustavo a . rodríguez , marco valentini , markus lilje , digitalbcon , sirroyalty , hector ceballos - lascurain , jacob . wijpkema , josé frade , luca boscain , nelsonlage , sam shaw , anderson rosado gomez , horacio luna , carlos ruiz . - guerra , marianhw , juanjaimemg , marinês eiterer , aisse gaertner , stu elsom , daniêl jimenez , dubi shapiro , pedro h ramos , frédéric pelsy , zanon , michaelp , andre zambolli , juan carlos marín , kevin b agar , bjørn kjellemyr , roger ahlman , samantha klein , erkki lehtovirta , holger teichmann , luis eduardo urueña , ken havard , mattias hofstede , dusan m . brinkhuizen , sebastian duque , sam , tomas grim , joselito nardy ribeiro , daniel avendaño , tadeusz stawarczyk , lindolfo souto , greg baker , josef widmer .
josep del hoyo , joe klaiber , peter boesman , noé eiterer , antonio silveira , dusan m . brinkhuizen .
male heard of forest edge of tropical dry forest . recording distance was 15 meters .
one individual seen and heard on forest edge of secondary vegetation at understory near a pasture . it was catching insects at fly . recording distance was 6 meters .
heard on forest interior at understory of lowland rain forest . recording distance was 10 meters or more .
probably a male at understory into forest interior of dry forest . different individual from xc382459 .
a single individual perched and flying for catching cicadas on forest edge of dry forest . recording distance was 8 meters .
recorded in the edge of vereda at if ' s school farm , in the company of ruan henrique silveira martins .
grabación hecha en el marco del proyecto
estudio técnico y participativo de la biodiversidad aviar del municipio de ibagué – tolima , como estrategia para la consolidación del ecoturismo de aves , la conservación de sus hábitats y el desarrollo de procesos de educación ambiental
universidad de ibagué - rednatur - colciencias .
singing while perched about 12 m from me . this was more than 2 km from my other recording of this species in this area ( xc402069 ) , so it is a different bird .
a pair , singing together . these three recordings were taken from about 1 - 1 / 2 hours of following these birds and attempting to get good recordings ; wind , some children , a photographer , and chickens caused me to not include many others .
original recording . habitat riparian strip of forest along arroyo de coloso at ecoparque ecoloso .
sonogram images © xeno - canto foundation . sonogram images share the same license terms as the recording they depict .
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these colours fit right in with christmastime , so we presume that the bird’s unusual appearance at asa wright was a fitting start to the holiday season .
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myavibase allows you to create and manage your own lifelists , and produce useful reports to help you plan your next birding excursion .
there are more than 12 , 000 regional checklists in avibase , offered in 9 different taxonomies , including synonyms more than 175 languages . each checklist can be viewed with photos shared by the birding community , and also printed as pdf checklists for field use .
there are a few ways by which you can help the development of this page , such as joining the flickr group for photos or providing translations of the site in addition languages .
howard and moore 4th edition ( incl . corrigenda vol . 1 - 2 ) :
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horse , ( equus caballus ) , a hoofed , herbivorous mammal of the family equidae . it comprises a single species , …
animal , ( kingdom animalia ) , any of a group of multicellular eukaryotic organisms ( i . e . , as distinct from…
female resembles male , but she has cinnamon - buff chin and throat . she is slightly paler and duller than male .
juvenile has shorter bill and tail than adults . the upperparts are duller , and the breast band is less regular and broader than in adults .
head is metallic green . the long , sharp black bill is about 4 to 5 cm length . eyes are dark brown with bare greyish eye - rings and lores . legs and feet are yellowish - brown .
we find six subspecies : g . r . melanogenia ; g . r . pallens ; g . r . brevirostris ; g . r . ruficauda ; g . r . rufoviridis ; g . r . heterogyna . they differ in varying amount of colours in plumage and bill size .
female lays 2 to 4 white eggs . incubation lasts about 19 to 23 days , shared by both parents , and female mainly at night . calls and displays occur at each change of incubating bird . at hatching , the chicks are covered with long whitish down . they are fed by both adults with insects caught around the nest - site , within 50 metres from the burrow . young fledge about 20 to 26 days after hatching . adults and youngs roost at night in the burrow after fledging , and during about 8 weeks . this species produces one brood per season , but probably two .
they measure in at about 9 inches ( 23 cm ) long and weight approximately 27 g . those long bills are used for capturing insects . they perch on horizontal branches swaying their bill back and forth in search of flying insects . they are known to capture bees and beetles as well as large brilliantly colored butterflies such as morpho and swallowtails . their long beak allows them to grab hold of butterfly bodies while keeping flapping wings as well as bee stings at bay .


eupithecia peregovitsi is a moth in the geometridae family that is endemic to vietnam .

the wingspan is about 25 – 26 millimetres ( 0.98 – 1.02 in ) .

the forewings are pale brown and the hindwings are white .

eupithecia peregovitsi

eupithecia lavicaria fuchs , 1902 ( syn : eupithecia lavicata prout , 1914 ) , described from norway .
ash pug ( angle - barred pug ) ( eupithecia innotata f . fraxinata )
montgomery , sl 1983 . carnivorous caterpillars : the behavior , biogeography and conservation of eupithecia ( lepidoptera : geometridae ) in the hawaiian islands . geojournal 7 ( 6 ) : 549 - 556 . doi : 10 . 1007 / bf00218529
eupithecia is a large genus of moths of the family geometridae . there are hundreds of described species , found in all parts of the world ( 45 in the british isles alone ) , and new species are discovered on a regular basis .
eupithecia species form the bulk of the group commonly known as pugs . they are generally small with muted colours and specific identification can be difficult . as a group they are easily identified by their narrow wings held flat at 90° to the body with the hindwings almost hidden behind the forewings .
the larvae of many species feed on the flowers and seeds of their food plants rather than the foliage . many species have a very specific food plant . some hawaiian eupithecia are predators of other insects ( e . orichloris , e . staurophragma , e . scoriodes ) . they mimic twigs but when sensitive hairs on their backs are triggered , they quickly grab the insects touching them . the defensive behavior of snapping may have pre - adapted hawaii ' s ancestral eupithecia for shifting to predation from feeding on pollen . also , insect predators that behave in this way are lacking in hawaii ' s fauna .
the larvae of many species feed on the flowers and seeds of their food plants rather than the foliage . many species have a very specific food plant . some hawai ' ian eupithecia are predators of other insects ( e . orichloris , e . staurophragma , e . scoriodes ) . they mimic twigs but when sensitive hairs on their backs are triggered , they quickly grab the insects touching them . the defensive behavior of snapping may have pre - adapted hawai ' i ' s ancestral eupithecia for shifting to predation from feeding on pollen . also , insect predators that behave in this way are lacking in hawai ' i ' s fauna .
a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
the author ' s name is made up of a string of letters , with no blanks , and multiple authors ' names are separated by a comma . spelling of author ' s name is based on the original publication . if there are more than three authors , only the names of the first two authors are shown , followed by
, +
and the number of omitted authors .
attribute is enclosed in square brackets . this is rarely needed , but to differentiate homo - identifiers , this will contain the page , line or plate number of original publication .
all diacritic marks , hyphens , and apostrophes are eliminated , thus only the following characters are used : a to z , a to z , 0 to 9 , blank , comma , and opening and closing square brackets . although upper and lower cases are used for the convenience of human recognition , it is not case sensitive .
created by dicky sick ki yu 1997 - 2012 please send me information about errors and omissions ( contact information ) with supporting references , possibly with pdf or hard copy .
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to date 683 species of macro - moth along with 1159 species of micro - moth have been recorded in norfolk since records began in victorian times . this site aims to provide full details of all the species that occur ( or once occurred ) in norfolk , with photographs , descriptions , flight graphs , latest records , distribution maps and more !
if you have photos of any moths featured on this site , and would like them displayed along with your name and comments . . . please send them in ( any size . jpg images ) .
please consider helping with the running costs of norfolk moths . thank you : - )
underlying maps using digital map data © norfolk online lepidoptera archive - nola™ 2018 . © james wheeler - n o r f o l k m o t h s 2007 - 2018 . data © nola™ 2018
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chinery , michael ( 1986 ) . collins guide to the insects of britain and western europe ( reprinted 1991 ) .
skinner , bernard ( 1984 ) . colour identification guide to moths of the british isles .
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biston mediolata ( n . jiang , d . y . xue & h . x . han , 2011 )
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it is endemic to mexico and only known from the area of its type locality near ayutla , oaxaca .

its common name is mustache false brook salamander or mustached false brook salamander .

the specific name refers to the whitish protuberances on the lips that resemble a mustache in the frontal view of the male holotype .

pseudoeurycea mystax

pseudoeurycea mystax is a species of salamander in the family plethodontidae . it is endemic to mexico .
pseudoeurycea mystax is a species of salamander in the plethodontidae family . it is endemic to mexico .
pseudoeurycea ( pseudoeurycea ) lineola — dubois and raffaëlli , 2012 , alytes , 28 : 77 - 161 .
iucn ssc amphibian specialist group 2016 . pseudoeurycea mystax . the iucn red list of threatened species 2016 : e . t59388a53983330 . urltoken
parra olea , g . & wake , d . 2004 . pseudoeurycea mystax . 2006 iucn red list of threatened species . downloaded on 23 july 2007 .
pseudoeurycea unguidentis is a species of salamander in the plethodontidae family . it is endemic to mexico .
new salamanders of the plethodontid genus pseudoeurycea from the sierra madre del sur of mexico . american museum novitates ; no . 2314
parra olea , g . & wake , d . 2004 . pseudoeurycea unguidentis . 2006 iucn red list of threatened species . downloaded on 23 july 2007 .
reviewed by tanner and dundee , 2000 , cat . am . amph . rept . , 705 : 1–5 ( as lineatriton lineolus prior to the naming of lineatriton orchimelas and lineatriton orchileucos ) . parra - olea and wake , 2001 , proc . natl . acad . sci . usa , 98 : 7888–7891 , suggested on the basis of molecular evidence that
lineatriton
lineolus is composed of at least two species , one more closely related to pseudoeurycea werleri and pseudoeurycea mystax and another , most closely related to pseudoeurycea leprosa . see comment under pseudoeurycea . see photograph , map , description of geographic range and habitat , and conservation status ( as lineatriton lineolus ) in stuart , hoffmann , chanson , cox , berridge , ramani , and young , 2008 , threatened amph . world : 577 . raffaëlli , 2013 , urodeles du monde , 2nd ed . : 270–271 , provided a brief account , photograph , and map .
reviewed by tanner and dundee , 2000 , cat . am . amph . rept . , 705 : 1 - 5 ( as lineatriton lineolus prior to the naming of lineatriton orchimelas and lineatriton orchileucos ) . parra - olea and wake , 2001 , proc . natl . acad . sci . usa , 98 : 7888 - 7891 , suggested on the basis of molecular evidence that
lineatriton
lineolus is composed of at least two species , one more closely related to pseudoeurycea werleri and pseudoeurycea mystax and another , most closely related to pseudoeurycea leprosa . see comment under pseudoeurycea . raffažlli , 2007 , les urodťles du monde : 223 - 224 , provided a brief account ( as lineatriton lineolus ) , photograph , and map . see photograph , map , description of geographic range and habitat , and conservation status ( as lineatriton lineolus ) in stuart , hoffmann , chanson , cox , berridge , ramani , and young , 2008 , threatened amph . world : 577 .
pseudoeurycea lineola — frost , grant , faivovich , bain , haas , haddad , de sá , channing , wilkinson , donnellan , raxworthy , campbell , blotto , moler , drewes , nussbaum , lynch , green , and wheeler , 2006 , bull . am . mus . nat . hist . , 297 : 359 .
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american museum novitates novitates ( latin for
new acquaintances
) , published continuously and numbered consecutively since 1921 , are short papers that contain descriptions of new forms and reports in zoology , paleontology , and geology . new numbers are published at irregular intervals .
department of library services american museum of natural history central park west at 79th st . , new york , ny 10024 © american museum of natural history , 2011
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
frost , d . r . 2013 . amphibian species of the world : an online reference . version 5 . 6 ( 9 january 2013 ) . electronic database . american museum of natural history , new york , usa . available at : urltoken .
justification : listed as critically endangered because its extent of occurrence ( eoo ) is 13 km 2 , it occurs in a single threat - defined location , and there is a continuing decline in the extent and quality of its habitat .
this species is known only from near ayutla , east - central oaxaca , mexico , at 2 , 100 m asl . it is only known from one threat - defined location and its extent of occurrence ( eoo ) is 13 km 2 .
it is an uncommon species , having last been seen in 1999 ( lamoreux et al . 2015 ) . due to ongoing decline in the extent and quality of habitat , the population is suspected to be decreasing .
this species is a terrestrial inhabitant of pine - oak forest and madroño ( arbutus forest ) . it can tolerate some habitat disturbance , as evidenced by the fact that a handful of small subpopulations survive in several tiny fragments of remaining habitat . it breeds by direct development and is not dependent upon water .
the major threat to this species is the extensive loss of habitat ( only small fragments of original habitat remain ) that has taken place due to agriculture ( including slash and burn practices ) , logging and human settlement .
conservation actions this species has not been recorded in any protected areas . it is listed as
threatened
( amenazada ) by mexican law . conservation needed there is an urgent need for the protection of forested areas near ayutla . research needed research is needed on its population size and distribution and to monitor its population trends .
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this species is known only from near ayutla , east - central oaxaca , mexico , at 2 , 100 m asl . it is only known from one threat - defined location and its extent of occurrence ( eoo ) is 13 km
. 2015 ) . due to ongoing decline in the extent and quality of habitat , the population is suspected to be decreasing .
this species has not been recorded in any protected areas . it is listed as
threatened
( amenazada ) by mexican law .
research is needed on its population size and distribution and to monitor its population trends .
, it occurs in a single threat - defined location , and there is a continuing decline in the extent and quality of its habitat .
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frost , darrel r . 2004 . amphibian species of the world : an online reference . version 3 . 0 ( 22 august , 2004 ) . electronic database accessible at urltoken american museum of natural history , new york , usa
amphibian species of the world : an online reference v5 . 3 , database ( version 5 . 3 )
frost , darrel r . 2009 . amphibian species of the world : an online reference . version 5 . 3 ( 12 february , 2009 ) . electronic database accessible at urltoken american museum of natural history , new york , usa
flores - villela , oscar / mccoy , c . j . , ed .
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 .
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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 .
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its natural habitat is subtropical or tropical moist montane forests . it is threatened by habitat loss .
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spelerpes lineolus cope , 1865 , proc . acad . nat . sci . philadelphia , 17 : 197 . holotype : not designated ; ansp 735 , according to dunn , 1926 , salamanders fam . plethodontidae : 423 , and malnate , 1971 , proc . acad . nat . sci . philadelphia , 123 : 348 . type locality :
table land , mexico
; probably eastern central veracruz , mexico , according to smith and taylor , 1948 , bull . u . s . natl . mus . , 194 : 21 . restricted to
metlac
, veracruz , mexico by smith and taylor , 1950 , univ . kansas sci . bull . , 33 : 349 .
opheobatrachus lineolus — cope , 1869 , proc . acad . nat . sci . philadelphia , 21 : 101 .
spelerpes ( oedipus ) infuscatus peters , 1879 , monatsber . preuss . akad . wiss . berlin , 1879 : 778 . holotype : zmb 6556 according to bauer , günther , and klipfel , 1995 , in bauer et al . ( eds . ) , herpetol . contr . w . c . h . peters : 40 . type locality :
hayti [ = haiti ] ; in error according to dunn , 1923 , proc . new england zool . club , 8 : 39 - 40 , who also made the synonymy ( but see wake , 1993 , herpetologica , 49 : 229 - 237 ) . type locality restricted to
metlac
, veracruz , mexico by smith and taylor , 1950 , univ . kansas sci . bull . , 33 : 349 .
geotriton lineolus — garman , 1884 , bull . essex inst . , 16 : 39 .
oedipina lineolus — cope , 1887 , bull . u . s . natl . mus . , 32 : 8 .
oedipina lineola — cope , 1896 , am . nat . , 30 : 1022 .
oedipus lineolus — fowler and dunn , 1917 , proc . acad . nat . sci . philadelphia , 69 : 21 . dunn , 1926 , salamanders fam . plethodontidae : 422 .
lineatriton lineola — tanner , 1950 , great basin nat . , 10 : 39 .
lineatriton lineolus — brodie , mendelson , and campbell , 2002 , herpetologica , 58 : 194 .
veracruz worm salamander ( liner , 1994 , herpetol . circ . , 23 : 12 ; frank and ramus , 1995 , compl . guide scient . common names amph . rept . world : 32 ; liner and casas - andreu , 2008 , herpetol . circ . , 38 : 33 ) .
mexican slender salamander ( tanner and dundee , 2000 , cat . am . amph . rept . , 705 : 1–4 ) .
known only from a small area of pine - oak forest in the sierra madre oriental of veracruz , mexico , in the vicinity of cuautlapan , nearby barranca metlac , and jalapa , 800–1250 m elevation .
please note : these links will take you to external websites not affiliated with the american museum of natural history . we are not responsible for their content .
for access to available specimen data for this species , from over 350 scientific collections , go to vertnet .
copyright © 1998 - 2018 , darrel frost and the american museum of natural history . all rights reserved .
this species occurs in cerro san felipe / cerro san luis ( the type locality ) in north - central oaxaca , mexico . elevation 2 , 600 - 2 , 800 m asl . taxonomic status of specimens provisionally assigned to this species found in llano de las flores and cerro machin remain to be validated .
paratype : taylor , e . h . 1941 . herpetologica . 2 ( 3 ) : 57 .
this taxon is found in the sierra madre de oaxaca pine - oak forests , an ecoregion of northern oaxaca , mexico exhibiting a large number of endangered species , so that the conservation value is outstanding in terms of uniqueness of the habitat . the sierra madre de oaxaca pine - oak forests is within the tropical and subtropical conifer forests biome , and the ecoregion is known for elevated plant endemism , especially within the sierra de juarez montane forests .
this ecoregion is located in northern oaxaca state , and is delineated by the sierra norte de oaxaca mountains , which have characteristically abrupt and rugged topography . its tallest peak is zempoaltepetl ( 3400 metres ) , and most of the terrain in this area is above 1000 metres . three mountain chains or sierras constitute the sierra madre de oaxaca : juarez , aloapaneca and zempoaltepec . the climate is temperate and humid with annual temperatures ranging from 16°c to 20°c . the annual mean precipitation varies greatly from 700 millimetres ( mm ) to as great as 4000 mm .
the sierra juarez spiny lizard ( sceloporus cryptus ) is endemic to the ecoregion , and limited in range to drier parts of the sierra de juarez , in northeastern oaxaca . there are a number of threatened reptilian taxa in the ecoregion including the ribbon graceful brown snake ( rhadinaea fulvivittis vu ) , a limited distribution snake endemic to southern mexico .
avian taxa found here include the dwarf jay ( cyanolyca nana en ) , bearded tree quail ( dendrortyx barbatus cr ) , tamaulipas pygmy - owl ( glaucidium sanchezi ) and grey - barred wren ( campylorhynchus megalopterus ) as restricted - range bird species , which includes this ecoregion . the oaxaca sparrow ( aimophila notosticta nt ) , golden - cheeked warbler ( dendroica chrysoparia en ) , russet nightingale - thrush ( catharus occidentalis ) , hooded yellowthroat ( geothlypis nelsoni ) , and collared towhee ( pipilo ocai ) are also species which thrive in the habitats offered by this mountainous ecoregion .
this ecoregion presents a mosaic of vegetatative associations , due to the varied climate and topography . these formations include tropical evergreen forest , montane cloud forest , pine forest , pine - oak forest , and oak forest . the pine forests , at elevations between 1600 and 2600 metres ( m ) , include trees that are 25 to 40 m tall . dominant pine species are mexican white pine ( pinus ayacahuite ) ; lawson ' s pine ( p . lawsonii ) , a mexican endemic ; chiapas white pine ( p . strobus var . chiapensischiapensis ) ; michoacan pine ( p . devoniana lr / lc ) and smooth - barked mexican pine ( p . pseudostrobus ) . these pine forests have a robust understory and an herbacious layer dominated by numerous species of the ericaceae family .
this is a semi - arboreal species that inhabits pine - oak and fir forests , and has commonly been found under logs . it can live in somewhat disturbed forest . it reproduces by direct development .
listed as critically endangered because its extent of occurrence is less than 100 km , all individuals are known with certainty only from a single location , and there is continuing decline in the extent and quality of its habitat , and in the number of mature individuals , in oaxaca , mexico .
formerly common , this species was not found in the 1960s and the early 1970s , and , although it was recorded briefly , there have been no records since 1976 at the type locality .
the cause for declines remain a mystery . it is probably being negatively impacted by agricultural expansion , human settlements , and logging , all of which are taking place extensively within its range . however , these threats do not explain the level of decline that has been observed , because the habitat is still in quite good condition in some places . possible explanations include climate change , and disease ( such as chytridiomycosis ) .
it occurs in parque nacional benito juarez . there is a need for further research to establish the cause of the declines observed in this species in suitable habitat . this species is listed as
threatened
( amenazada ) by the mexican government .
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smith and taylor , 1948 , bull . u . s . natl . mus . , 194
smith and taylor , 1950 , univ . kansas sci . bull . , 33
fowler and dunn , 1917 , proc . acad . nat . sci . philadelphia , 69
frost , grant , faivovich , bain , haas , haddad , de sš , channing , wilkinson , donnellan , raxworthy , campbell , blotto , moler , drewes , nussbaum , lynch , green , and wheeler , 2006 , bull . am . mus . nat . hist . , 297
mexican slender salamander ( tanner and dundee , 2000 , cat . am . amph . rept . , 705 : 1 - 4 ) .
known only from a small area of pine - oak forest in the sierra madre oriental of veracruz , mexico , in the vicinity of cuautlapan , nearby barranca metlac , and jalapa , 800 - 1250 m elevation .
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parra olea , g . & wake , d . 2004 . 2006 iucn red list of threatened species . downloaded on 23 july 2007 .
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temnothorax albipennis is a species of small ant in the subfamily myrmicinae .

it occurs in europe and builds simple nests in rock crevices .

temnothorax albipennis

the above specimen data are provided by antweb . please see temnothorax albipennis for further details
junior synonym of temnothorax unifasciatus : nylander , 1856b pdf : 92 ; of temnothorax nylanderi : smith , 1858a pdf : 120 ; of temnothorax tuberum : emery , 1924f pdf : 256 .
( 2013 ) nest - seeking rock ants ( temnothorax albipennis ) trade off sediment packing density and structural integrity for ease of cavity excavation .
until recently the ant temnothorax albipennis was known as leptothorax albipennis ( = tuberointerruptus ) and had been misidentified as temnothorax tuberum in britain . it appears that all mainland records of temnothorax tuberum refer to t . albipennis ( orledge , 1998 ) . whether the channel islands records for t . tuberum are similarly in error is unknown , although they are included here on the t . albipennis map . workers of t . albipennis are smaller than those of leptothorax acervorum , and generally distinguishable from workers of the three other british temnothorax species , by their combination of darkened antennal clubs , straight and relatively short propodeal spines , and rather poorly - defined darkening on their gasters .
image showing a marked temnothorax albipennis colony . colonies may be marked in this way so that individuals can be tracked in order to determine what behaviours they demonstrate .
revived from synonymy and senior synonym of temnothorax tuberointerruptus : orledge , 1998 : 31 .
combination in leptothorax : orledge , 1998 : 31 ; in temnothorax : bolton , 2003 pdf : 271 .
salata , s . & borowiec , l . 2013 . temnothorax albipennis ( curtis , 1854 ) in poland and identification of the t . tuberum species complex ( hymenoptera formicidae ) . genus ( wroclaw ) 24 , 403 - 413 .
temnothorax albipennis workers forage singly . they scavenge small arthropods and may also take small live invertebrates ( orledge , 1995 ) . no carbohydrate sources have been documented , but the author ' s observation of large numbers of workers on umbellifer flowers suggests that nectar is taken .
plateaux , l . & cagniant , h . 2012 : 436 : we conclude from all these observations that leptothorax tuberum unifasciatointerruptus is a hybrid of temnothorax unifasciatus and temnothorax albipennis . hybridization works in both directions , parents may be alternately of the two species , and males are from the parthenogenetic egg of the queen mother . note that both species exist also in places ( especially in luberon and ventoux ) where the where the form was found . we therefore ask : t . albipennis x t . unifasciatus = l . tuberum unifasciatointerruptus forel .
on continental europe t . albipennis has been recorded from the netherlands , southern france , northern spain , south and central germany and central italy .
bolton , b . 2003 . synopsis and classification of formicidae . mem . am . entomol . inst . 71 : 370pp ( page 271 , combination in temnothorax )
in britain , t . albipennis colonises warm , south - facing , open rocky areas , typically coastal cliffs , screes , undercliffs , slumped cliff slopes and old quarries .
wiezik , m . 2005a . first records of leptothorax albipennis and l . nadigi ( hymenoptera , formicidae , myrmicinae ) from slovakia . biológia ( bratisl . ) 60 : 170 ( page 170 , see also )
plateaux , l . & cagniant , h . 2012 . quelques synonymies dans le genre temnothorax mayr , 1855 ( hymenoptera , formicidae ) . bulletin de la société entomologique de france 117 ( 4 ) , 427 - 440 .
by contrast , in established temnothorax albipennis colonies , the aggregate is stable and is contained inside a nest rather than being exposed in the open . in t . albipennis , decisions to leave an established nest are not solely a function of group size [ 14 ] . even more than for a temporary aggregation , individual decisions to leave a stable collection of nest - mates may be affected by a wide variety of inter - related factors such as age or physical caste [ 15 ] , genetic predisposition [ 16 ] , physiological status [ 17 , 18 ] , reproductive status [ 19 ] , dominance interactions [ 20 , 21 ] , previous experience and spatial location [ 22 ] .
we tested the exit dynamics of t . albipennis ant colonies under the following three experimental conditions . note , all colony - size statistics were collected in a census immediately prior to the commencement of removal of workers as they left the nest .
orledge , g . m . 1998 . the identity of leptothorax albipennis ( curtis ) ( hymenoptera : formicidae ) and its presence in great britain . syst . entomol . 23 : 25 - 33 pdf ( page 31 , revived from synonymy , combination in leptothorax and senior synonym of tuberointerruptus )
we assumed that a similar process applies to individual ants within intact colonies , but in combination with social information . consistent with earlier experimental work on the collective decision making of temnothorax ants , we incorporated the use of quorums in our model ( 31 ) . when the population of ants at a nest site surpasses a threshold , commitment to that site increases significantly ( 32 , 33 ) . with this addition , the site acceptance probabilities now depend on the number of ants at the site as follows :
tuberointerruptus . leptothorax tuberointerruptus bondroit , 1918 : 126 ( w . q . ) switzerland . [ first available use of leptothorax tuberum var . tuberointerruptus forel , 1874 : 86 . nomen nudum . leptothorax tuberum r . interruptus var . tuberointerruptus forel , 1915d : 24 ( in key ) ; unavailable name . ] status as species : betrem , 1926 : 217 . subspecies of interruptus : novak & sadil , 1941 : 93 . revived status as species : seifert , 1994 : 22 . junior synonym of albipennis : orledge , 1998 : 31 .
i first determined whether ants assess nest population throughout the emigration or only until they begin to transport . i induced emigrations and followed recruiters until they switched from tandem runs to transports . i then emptied the new nest of ants and noted whether the recruiters continued to transport or reverted to tandem runs . two colonies of t . albipennis were housed in observation nests constructed of thin cardboard or wood partitions sandwiched between glass microscope slides ( 50 × 76 mm ) . these colonies and all others used in this study were collected at portland bill , dorset , u . k . ants were individually marked with four drops of paint on the head , thorax , and gaster . emigrations were induced by removing the roof slide from the old nest , forcing the ants to find a new home .
thirty - two colonies of t . rugatulus were used for the colony - level tests . half of them were small ( 20–80 workers ) , and half were large ( 150–250 workers ) . an additional eight medium - sized colonies ( 100–130 workers ) provided 16 worker ants ( 2 ants per colony ) for the individual tests . only a minority of workers in temnothorax colonies participate actively in nest - site scouting and transport of nestmates and brood items , whereas the rest of the colony waits at the home nest ( 24 , 33 ) . to ensure that we tested only these active ants , we placed brood items outside of the colony’s nest and selected workers that attempted to retrieve them . we reasoned that ants willing to leave the nest and retrieve brood were likely to carry out similar tasks during colony emigrations ( 34 ) .
the collective choice of a new home by emigrating colonies of the ant t . albipennis depends critically on the detection of a nest - mate quorum by the ants evaluating each candidate site . the quorum allows ants to condition their full commitment to a site—signaled by the switch from slow tandem runs to rapid transports—on the parallel decisions of their nest mates ( pratt et al . , 2002 ) . in this study , i show that quorum detection depends on direct contact between an ant and her nest mates at the site , probably acting through density - dependent encounter rates . i further show that ants cease monitoring the presence of a quorum once they have made the switch to transport , failing to revert to tandem runs even if the site ' s population falls well below the quorum . finally , my observations suggest that quorum detection is not instantaneous but requires ants to invest 1–2 min patrolling the site to gather information .
a markov chain model was used to describe the temnothorax house hunting process . in the model , each ant begins in the exploring state , from which she can either find the better nest , entering state a , or the worse nest , entering state b . if the ant fails to find either nest , she continues exploring . once at a site , she may also leave and find the other site , with some probability . with each subsequent nest visit , the ant enters state ca i or cb i , where i represents the number of comparisons she has made between nests . this number is incremented each time the ant goes from a to b , or from b to a . at each state the ant has a probability of accepting the nest she is currently assessing , modeled as the final states a and b . si appendix , table s2 provides a summary of model states ; si appendix , table s3 the model’s transition probabilities ; and si appendix , table s4 a description of the model parameters . the flowchart in si appendix , fig . s4 summarizes all possible state transitions .
from a previous dataset in which individually marked t . albipennis ants were allowed to freely leave and return to the nest [ 14 ] , we obtained the times that each ant left the nest over a 5 day period . from this , for each ant , we calculated the mean waiting time between successive exits . it should be emphasized that here the waiting time refers to the time elapsed between successive exits by the same ant . we then took the inverse of each ant ' s mean waiting time , giving the mean exit rate for each ant . thus , we obtained the distribution of the instantaneous decay probabilities ( exit rates ) within each of seven colonies . these rate distributions were well fitted by a log - logistic distribution . a variable that follows a log - logistic distribution is one in which the logarithm of the variable is logistically distributed ( electronic supplementary material , figure s2 ) . at the start of each run of the simulation , every ‘ant’ in the colony was assigned an individual exit rate from the distribution . in every time step , each ant decided whether to leave the nest by comparing its exit probability to a uniformly distributed random number between 0 and 1 . if the ant ' s exit probability was greater than this random number , the ant leaves the nest . we recorded the same statistics for these simulated ants , as for the real ants . for comparison , we also ran the model without any individual variation , by assigning the mean instantaneous decay probability from the log - logistic distribution to all the ants , which is a homogeneous poisson process .
these ants inhabit grasslands and light scrub , especially on lime subsoil and are sometimes common in dunes . they nests in rock crevices and rubble or in tree stumps and in dry fallen branches . colonies are reported as monogynous , with up to 200 workers , and may form temporary polydomous systems ( salata & borowiec , 2013 ) .
this species is a host for the endoparastic fungus myrmicinosporidium durum ( espadaler & santamaria , 2012 ) .
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
curtis , j . 1854 . on the genus myrmica and other indigenous ants . trans . linn . soc . lond . 21 : 211 - 220 ( page 218 , worker , male described )
espadaler , x . , santamaria , s . 2012 . ecto - and endoparasitic fungi on ants from the holarctic region . psyche article id 168478 , 10 pages ( doi : 10 . 1155 / 2012 / 168478 ) .
nylander , w . 1856b . synopsis des formicides de france et d ' algérie . ann . sci . nat . zool . ( 4 ) 5 : 51 - 109 ( page 92 , junior synonym of unifasciatus )
smith , f . 1858a . catalogue of hymenopterous insects in the collection of the british museum . part vi . formicidae . london : british museum , 216 pp . ( page 120 , junior synonym of nylanderi )
this page was last modified on 31 october 2016 , at 00 : 51 .
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0 times found in rocks ( rocky - calcareous grasslands ) , 4 times found in pinar , 0 times found in unknown , 3 times found in bosque claro de p . sylvaticus , 2 times found in robedal / castañar , 1 times found in agrícola , 0 times found in dry grassland , 1 times found in pinar ( p . sylvaticus ) cerrado , 1 times found in pinar en dunas , 1 times found in pinus sylvestris , . . .
1 times nido en tocón , 2 times nido en piedra , 1 times madera , 2 times forrajeando , 2 times bajo corteza ymusgo p sylvaticus , 1 times sous pierres , 1 times nido dentro de piedra , 1 times nido bajo piedra , 1 times nbp , 1 times bajo corteza pino , 1 times bajo corteza p sylvaticus , . . .
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british mainland records come only from southern england and from wales . most are from sites on or near the coast , a notable exception being the record from castle drogo , south devon ( alexander , 1983 ) . there are no confirmed records from ireland .
listed as a nationally notable ( na ) species ( falk 1991 ) . work for this atlas has shown that the species is currently known from 40 post - 1970 10 km squares on mainland britain and so the status should be revised .
alate males and females are produced during june and july and have left the nest by the middle of august ( from data in partridge ( 1993 ) ) . there are no field records of swarming , but laboratory observations suggest that mating flights occur during the two hours following sunrise ( plateaux 1984 ) .
nests are most frequently found between rock laminae , in rock cavities and crevices , on rock surfaces under mat - forming plants , or on soil under stones . between rock laminae the workers often surround the nest with a ' wall ' of soil particles and rock fragments . there are several records of nests within the dead stems of herbaceous and shrubby plants , in rotting wood and in old snail shells . near dungeness , kent , nests have also been found amongst pebbles and gorse roots , and in peat over pebbles ( felton , 1965 ) . colonies rarely exceed 350 workers , and have a single queen . adult workers are produced each year during the summer months .
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t . o . richardson , 1 , 2 , * k . christensen , 3 , 4 n . r . franks , 2 h . j . jensen , 3 , 5 and a . b . sendova - franks 1
* author for correspondence ( ku . ca . lotsirb @ nosdrahcir . mot ) .
many purely physical complex systems , in which there are both stochasticity and local interactions between the components , exhibit record dynamics . the temporal statistics of record dynamics is a poisson process operating on a logarithmic rather than a linear time scale ( i . e . a log - poisson process ) . record dynamics often drive substantial changes in complex systems when new high water marks in partially stochastic processes trigger new events . social insect colonies are exemplary complex biological systems in which many of the local interactions of the components have been moulded by natural selection for the common good . here , we combine experimental manipulation of ant colony demography with modelling to test the hypothesis that social interactions are the mechanism underlying the record dynamics . we found that compared with the control , log - poisson statistics were disrupted in colonies in which the pattern of interactions was modified by the removal of the brood , and disappeared completely in ‘callow’ colonies composed entirely of very young workers from the same age cohort . we conclude that a subtle interplay between the demography of the society and the pattern of the interactions between the ants is crucial for the emergence of record dynamics . this could help identify what makes an ant colony a cohesive society .
there may be safety in numbers [ 1 ] , so for many social animals , individuals are more likely to remain in a group the larger it is . hence , when deciding whether to leave the safety afforded by the group , individuals may often take the size of the group into account [ 2 – 4 ] .
complex systems in which events are governed by record dynamics exhibit rapidly , and crucially non - exponentially , declining event rates ( the rates are proportional to the inverse of time , hence events accumulate at a constant rate in logarithmic time ) . further , such systems always include strong interactions between the component parts . record dynamics have been shown to govern rapidly declining event rates in a diverse variety of non - equilibrium systems , for example declining extinction rates [ 27 – 29 ] , the ‘tangled nature’ model of macroevolution [ 25 , 30 ] , fluctuating commodity prices [ 31 ] , type - ii superconductors [ 32 ] , colloidal gels [ 33 ] and relaxation of the angle of repose in vibrated sandpiles [ 34 ] .
here , we use experimental manipulations of colony structure in combination with modelling to investigate the mechanisms underlying the log - poisson characteristics of nest exits when ants are experimentally prevented from returning home ( richardson et al . 2010 ) . our experiments compare ( i ) control colonies where worker and brood demographies have not been manipulated with ( ii ) ‘no - brood’ colonies where worker demography is not manipulated but the brood has been removed and ( iii ) colonies that have a full brood complement but all the workers belong to the same cohort of very young ‘callow’ workers . with our models , we compare statistics generated by two scenarios based on the poisson process against the well - known log - poisson statistics generated by a record dynamics process .
our second experimental manipulation was to construct callow colonies in which all the workers belonged to the same age - cohort of very young workers . both worker developmental stage and experience influence , and are influenced , by social interactions . hence , the motivation for this manipulation was to check whether a narrower distribution of probabilities for performing some action could still reproduce the log - poisson statistics found in unmanipulated colonies .
) with food and water provided ad libitum . all colonies had a single queen that showed normal behaviour for a fertilized queen , for example a strong tendency to take up a central position within the colony .
apparatus for automated removal of ants as they leave the nest . ( a ) microscope - slide nest ( 50 × 35 × 1 mm ) ; ( b ) high - resolution webcam ; ( c ) computer running automated motion - detection software ; ( d ) compressed air supply ; ( e ) access tube through which ants could access the exploration arena prior to the removal , and through which they left the nest during the removal . the access tube contained an inner tube and nozzle arrangement that delivered the pressurized air , hitting the ant from behind .
sixty - four hours prior to the beginning of the period when ants were removed as they left the nest , each colony was stimulated to emigrate to a new nest by removing the roof of their original nest . during this ‘settling - in’ period , no food or water was provided to the colonies , although they were allowed access to an exploration arena ( 100 × 100 mm ) , which was accessible via a tube attached to the front of the nest (
during the period when ants were removed as they left the nest , the exploration arena was disconnected . then , every time an ant walked to the glass tip of the access tube , its motion was detected by the high - resolution webcam ( logitech qc deluxe ) positioned immediately above (
) . this webcam was attached to a pc running automated motion detection software , thus sensing the exit of the ant . when an ant was detected leaving the nest , the computer opened a valve , releasing a short ( 0 . 2 s ) burst of pressurized air through the inner tube , hitting the ant from the rear , and resulting in the ant being blown out of the access tube , into a collection dish .
nine unmanipulated colonies with brood present ( median n adults = 90 , inter - quartile range ( iqr ) = 34 ; median n brood = 127 , iqr = 111 . 8 ) .
eight unmanipulated colonies with all brood removed ( median n adults = 118 , iqr = 24 ; median n brood = 0 . 5 , iqr = 4 . 8 ) . the brood was removed during the enforced emigration into the experimental nest , that is 64 h prior to commencement of experimental ant removal . in some colonies , the queen laid a few eggs after the removal of the brood , hence the median n brood is non - zero .
eight callow colonies were constructed ( median n adults = 101 , iqr = 67 , median n brood = 77 , iqr = 45 . 5 ) in the following way . the queen plus a retinue of 10 adults were removed from each of eight unmanipulated colonies , thus creating eight ‘proto - colonies’ . then , the 10 adult workers from each proto - colony were marked using pactra paint . these adult workers were used to perform the crucial task of helping the late - stage pupae eclose into callows , after which the adult workers were removed . we then collected brood items from 20 donor colonies , and sorted them according to their developmental stage ( eggs , larvae , pre - pupae and pupae ) . each proto - colony received a complement of brood containing equal proportions of eggs , larvae and pupae . we aimed to ensure there were equal numbers of workers and brood in each colony . the modal length of time from the point at which the majority of the pupae had eclosed , until the emigration into the experimental nest , was 12 days . newly eclosed callows are extremely sluggish , hence the delay to allow time for them to acquire some mobility . although not all the late - stage pupae eclosed at exactly the same time , the variation was relatively low , with the delay between the first and last eclosions of approximately 2 days .
which is the expectation from a homogeneous poisson process , such as radioactive decay .
( a ) the mean proportion of the colony remaining , normalized by the starting colony size , in logarithmic time ( log - linear axes ) . in a log - poisson process , n ( t ) grows , on average , at a constant rate in logarithmic time , hence 1 − ( n ( t ) / n t = 1 ) will decrease at a constant rate in logarithmic time . open circles , control ( n = 9 ) ; filled grey circles , no - brood ( n = 8 ) ; filled black circles , callow colonies ( n = 8 ) . ( b ) the proportion of the original colony remaining in the nest , as a function of time ( note linear - log axes ) . the proportion of the original colony remaining 1 − ( n ( t ) / n t = 1 ) , at the end for the experiment was : for the control , median = 0 . 39 , iqr = 0 . 47 ; no - brood , median = 0 . 37 , iqr = 0 . 27 ; callows , median = 0 . 50 , iqr = 0 . 12 . as in a homogeneous poisson process , the number remaining in the nest declines exponentially over time in the callow colonies ( λ = −2 ) but not in the control or the no - brood colonies . the linear fit is a guide to the eye only .
, that occurs simply due to there being fewer and fewer ants ‘available’ to leave the nest .
) is negative , this indicates social amplification , that is , the smaller the aggregate , the higher the exit rate and hence the shorter the residence times of those remaining . however , we found
> 0 , so the smaller the aggregate , the lower the exit rate and hence the longer the residence times , which we term social inhibition . the rate
exit rate as a function of the proportion remaining than did the no - brood colonies . we again found a qualitatively different pattern in the callow colonies , in which the
; electronic supplementary material , figure s4 ) . in other words , when colonies were constructed so as to minimize the variation in the ants ' exit probabilities , the
exit rate was stationary , and thus the absolute exit rate was dependent only upon the finite number of ants in the nest .
in the controls against those in the no - brood and callow colonies . we used glms of the form , response ∼ colony + experiment + covariate + experiment * covariate , where ‘ * ’ denotes interaction . this interaction term determines whether the coefficients ( i . e .
) are significantly different in the no - brood and callow colonies , compared with the control . the standardized residuals from the above glms were slightly more right skewed than a normal distribution ; however , the qualitative differences between the coefficients were the same when individual regressions were preformed on the ensemble average data depicted in
( a ) control , ( b ) no brood , and ( c ) callows . each data point represents the ensemble average , that is an average across the colonies . the curved lines are lowess regressions ( degree of smoothing , f = 0 . 5 , n steps = 2 ) . row 1 : testing for social amplification ( m < 0 ) or social inhibition ( m > 0 ) . rows 2–4 : testing for log - poisson statistics . logarithmic binning was used throughout for time plotted on the x - axis . row 2 : checking that the per capita exit rate behaves like r k / n t ∝ t − β . row 3 : variance - to - mean ratio , < vmr > , of the normalized accumulated number of events over time . if the timing of events obeys log - poisson statistics , the vmr will be constant over time . row 4 : open square boxes , the waiting time between exits , < t k − t k − 1 > , as a function of time . filled circles , the ratio , < x > , that is , the ratio of the waiting time between exits , t k − t k − 1 , to the time of occurrence of the previous ( k − 1 ) th event , t k − 1 , as a function of the time of occurrence of t k − 1 .
; electronic supplementary material , figure s3 ) . however , as the callow colonies showed a qualitatively different decrease in the event rate that was not well described by linear regressions of the form log
) , that is , the waiting time between exits increases more rapidly when the brood is present , compared with when the brood is absent .
testing for quantitative differences between the control and no - brood colonies . the callows were excluded from the comparison as they displayed qualitatively different temporal statistics . glms of the same form as in
rate and the waiting times were significantly different in the control versus no - brood colonies .
) of the accumulated number of events , that is , the vmr is equal to one , and does not vary over time . if the event rate is only
to a log - poisson variable . however , the same cannot be said for callow colonies , in which the < vmr > increases rapidly over time .
in the poisson process , the waiting time is stationary , exponentially distributed ( i . e .
) and there is no correlation between successive waiting times . similarly , in a log - poisson process , the logarithmic waiting times ( ln
) and without temporal correlation . however , here we wish to compensate for the falling colony size by calculating the logarithmic waiting times on a
) , and are not temporally correlated ( electronic supplementary material , figure s1 ) . however , in the callow colonies ,
are considerably more right - skewed , non - stationary and temporally correlated , while in the no - brood colonies these metrics are again intermediate between poisson and log - poisson .
the cumulative distribution of the per capita logarithmic waiting times , p ( τ / n t ) > x . ( a ) controls , ( b ) no - brood colonies , ( c ) callows . in a poisson process , the waiting times are exponentially distributed , ( p ( t k − t k −1 > x ) = e − λx ) , and similarly in a log - poisson process the logarithmic waiting times ( ln t k − ln t k −1 = ln ( t k / t k −1 ) = τ ) are exponentially distributed ( p ( τ > x ) = e − λx ) . each line represents a different colony .
the mechanism described above is a natural null model because it is so simple , and also because the poisson statistics it produces are well studied . however , the temporal statistics of a poisson process operating on a logarithmic time scale , that is ‘log - poisson’ statistics , characterizing the nest exit dynamics in unmanipulated colonies ( richardson et al . 2010 ) , suggest an alternative mechanism of record dynamics [ 25 , 26 ] derived from the statistical mechanics of complex systems . in this scenario , individual exits are triggered when a fluctuating variable , the ‘record signal’ , reaches a never - before - seen ( record ) level . if the record signal is uncorrelated in time , the increment between successive record values will rapidly decrease , and the interval between successive records will rapidly increase . clearly , there are strong interactions between ants [ 41 – 43 ] , and the interaction rate—measured by the collective activity—fluctuates non - periodically over time [ 44 – 46 ] .
here we assume that within each colony there is a distribution of intrinsic decay probabilities , and that a ‘decay’ event is equivalent to an exit . then , the declining exit rate will be because of the diminishing number of ants , but also because on average the more ‘unstable’ ants leave the nest before those with lower exit probabilities . this model is essentially a non - homogeneous poisson process that incorporates a finite - size effect .
here we present an alternative phenomenological explanation for the production of the observed log - poisson statistics . we assume that an event occurs ( an ant leaves the nest ) whenever a fluctuating record signal hits a new maximum ( or minimum ) value . then , the temporal stochasticity of the signal itself is responsible for regulating the exit rate , rather than the variation across individuals , as in the null model . time series created in this manner are known as ‘white noise’ . as each instance of the signal is randomly chosen , the value of the signal at time
the temporal sequence of events ( records ) generated by three realizations of the record dynamics model . records accumulate at a constant rate in logarithmic time . insert : the fluctuation of the underlying white - noise record signal for a single realization . here , the x - axis shows linear time . open circles , the underlying record signal and its associated records ( solid line ) .
when there was no individual variation in the exit rates , or rather , when the exits were organized by a homogeneous poisson process operating on a finite population , we found typical poisson statistics . specifically , the number of events did not accumulate at a constant rate in logarithmic time (
) . thus , this model qualitatively reproduces the statistics produced by the callow colonies .
results of the null model based on : ( a , c ) a homogeneous poisson process , instantaneous exit rate for all ants = 0 . 0003 s −1 , ( b , d ) a heterogenous poisson process ( log - logistic distribution , see electronic supplementary material , figure s2 ) mean instantaneous exit rate = 0 . 0003 s −1 . for both exit probability distributions , the starting colony size was identical , n t = 1 = 100 ants . the number of time steps in both models was of the same order of magnitude as in the experimental data . for further details of model parametrization , see text . ( a , b ) open circles , the mean proportion of the original colony remaining , 1− ( n ( t ) / n t = 1 ) ; filled circles , the vmr for n ( t ) . ( c , d ) open squares , the mean waiting times as a function of time , t k − t k −1 ∝ t k −1 ; filled squares , the ratio of the waiting times , x , as a function of the time of the previous event .
we do not give further evidence of the log - poisson nature of the event rate statistics generated by the record dynamics model as it has been demonstrated elsewhere [ 25 , 47 – 49 ] that a fluctuating record signal produces log - poisson statistics .
] . in the no - brood colonies , the absence of hungry brood may have removed stimuli that trigger foraging and thus contributed to the observed reduction in the overall exit rate .
it is clear from the null model that individual variation is not , by itself , sufficient to reproduce event rates that decelerate according to record dynamics . merely assigning individual nest - leaving rates ( derived from previous empirical observations of unimpeded nest leaving ) to the non - interacting ‘ants’ did not reproduce the log - poisson statistics (
) . therefore , the scenario in which the decision to leave the nest is a purely probabilistic consequence of the individual exit rate is inaccurate .
statistics in the control colonies demonstrates that the log - poisson deceleration is not merely an artefact of the declining colony size . colonies composed entirely of a single age - cohort of very young and inexperienced callows differed qualitatively from the log - poisson statistics described in the control and , to a lesser extent , in the no - brood conditions . in the callow colonies , the
) or stationary ( electronic supplementary material , figure s5 ) . instead of the ‘normal’ log - poisson temporal statistics found in unmanipulated colonies , the timing of nest exits in callow colonies was more like a homogeneous poisson process (
) . so , when the society lacks variation in age and task experience among the component parts , it displays none of the interesting dynamics exhibited by systems in which events are regulated by record dynamics .
the importance of the structuring of social interactions is supported further by the result that no - brood colonies show statistics that are intermediate between log poisson and poisson . we hypothesized that the removal of the brood would change the spatial distribution of the interactions between the workers within the colony , so causing a loss of log - poisson temporal statistics . indeed , the loss of the log - poisson behaviour suggests that the particular pattern of ant–ant interactions mediated by the presence of brood of various developmental stages sorted into an annular structure is important for the generation of log - poisson statistics . in summary , to explain the production of the log - poisson statistics , it is necessary but not sufficient to invoke only the natural variation in the ants ' nest - leaving tendencies or the nature of the interactions between the workers .
so what then would be sufficient ? it is reasonable to suppose that decreasing the density of ants within the nest might cause a reduction in the rate at which ants encounter one another . then , if individual exit probabilities were downregulated ( i . e . inhibited ) as the encounter rate decreases , the exit rate would decline more rapidly than if there were no density effects . clearly , in order to construct an individual - level model that reproduces the collective log - poisson statistics , we will need a much more detailed understanding of how ant–ant interactions within the nest influence subsequent decisions to leave it .
our results demonstrate how analysing the behaviour of social insect colonies using the tools of record dynamics brings new insights in their organization and opens up the prospect of developing a new theory to help identify what makes an ant colony a cohesive society . substantial and irreversible changes in complex systems can originate from the combination of temporal stochasticity and strong interactions between the components . in the physical sciences there is strong evidence for the role of such record dynamics in a wide range of condensed - matter systems . the application of record dynamics to complex systems in biology promises to be fruitful because both stochasticity and agent–agent interactions are ubiquitous at all levels of biological organization .
t . o . r . and a . b . s . - f . acknowledge epsrc grant ep / e061796 / 1 . k . c . acknowledges epsrc grant ep / e061761 / 1 . we are grateful to everyone in the ant laboratory at bristol for helpful advice .
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address correspondence to s . c . pratt . e - mail : spratt @ urltoken .
how do ants actually implement the quorum rule ? that is , what proximate cues cause them to change their recruitment behavior ? when we say that this change relies on attainment of a quorum , this reflects both the experimental observation that the number of nest mates at a site determines subsequent recruitment behavior as well as the likelihood that nest - mate number is the functionally important feature for decision - making performance at the colony level . the ants themselves , however , are not likely to assess nest - mate numbers directly . although evidence exists for a limited number sense in many animals , including honey bees ( chittka and geiger , 1995 ; dehaene et al . , 1998 ) , insects and simpler organisms likely rely instead on simple cues correlated with either the absolute numbers or the density of conspecifics .
alternatively , ants may respond to their rate of encounter with nest mates . during each visit to her candidate site , the scout wanders through it , approaching nest mates and touching them with her antennae . the rate of such encounters is likely to increase with the local density of ants ( gordon et al . , 1993 ) . one possibility is that an ant simply notes the time elapsed between her entry at the site and her first encounter . if this interval declines with increasing site population , it could serve as a simple cue of nest - mate density . similar time intervals play an important role in allocating labor in partitioned tasks , efficiently balancing the numbers of workers foraging for material with the numbers using this material at the nest ( jeanne , 1986 ; seeley , 1995 ) . a second possibility is that each ant integrates information from many encounters over the duration of her visit . use of multiple encounters could improve the precision of her estimate , as others have suggested for partitioned tasks ( ratnieks and anderson , 1999 ) . similar cues have already been implicated in labor allocation within harvester ant colonies , where an ant ' s probability of switching to a given task is positively related to her rate of encounter with nest mates already performing that task ( greene and gordon , 2003 ; gordon and mehdiabadi , 1999 ) .
in this study , i examined the potential roles of encounter rate and indirect pheromonal signals in detection of a nest - mate quorum . i began by determining when , during an emigration , scouts pay attention to quorum attainment , and by estimating the time they devote to its measurement . i then explicitly tested the importance of each cue by manipulating its availability to scouts and determining how this affected their subsequent recruitment decisions .
apparatus used to control access of ants to a new nest site . the dish containing the old nest is mounted on a rack - and - pinion mechanism that allows it to be joined with or separated from the dish containing the new nest .
the time required by ants to decide whether a quorum is present may help to distinguish among potential mechanisms . detection of volatile pheromones is potentially very rapid , while measurement of lag to the first encounter may be more time consuming , and integration of a series of encounters should take longer still . to determine whether time devoted to quorum detection changed as ants switched from tandem runs to transports , i measured the duration of each visit to the new site by individually marked recruiters . emigrations were observed in a large tray with fluon - coated walls . an inhabited nest was placed against one wall of the tray and a single empty nest against the opposite wall , 60–65 cm away . digital video cameras recorded the ants ' activity at each nest throughout the emigration . visit durations and subsequent recruitment behavior were measured from the videotapes . only data from ants that led tandem runs were analyzed . i excluded the visit before each ant ' s first recruitment to avoid confounding time devoted to quorum detection with time needed to assess the nest . also excluded were visits that followed searches in the arena , to avoid confounding quorum assessment with prolonged stays in the nest that may follow these searches . six emigrations were observed : three by colony a6 , two by colony 6 , and one by colony 1 .