the wrybill or ( in māori ) ngutuparore ( anarhynchus frontalis ) is a species of plover endemic to new zealand .

it is special since it is one of the two species of bird in the world with a beak that is bent sideways ( it is always to the right for wrybill , and the other species is loxia pytyopsittacus ) .

a 2015 study found it to actually be within the charadrius clade , with its closest relatives other plovers found in new zealand , the nearest the new zealand plover or new zealand dotterel ( charadrius obscurus ) , and then the double-banded plover or banded dotterel ( charadrius bicinctus ) .

it lays its eggs among the rocks along rivers and distracts intruders by pretending to be in distress and moving away from its " nest " .

wrybill

wrybill <EOT> miranda is an amazing shorebird location , and we spend time here looking for wrybill and other migrant shorebirds .

range : the wrybill is found in new zealand . it breeds in c south i and winters in n north i .

with it ' s bill curved to the right the wrybill is an uncommon visitor / resident of awarua bay , new zealand .

wrybill breed on large braided rivers in central south island from august - january . they prefer large dynamic rivers that will not become overgrown with weeds . once prevalent on smaller rivers , the wrybill’s range as contracted to about 60 % of its estimated original habitat (

wrybill ( anarhynchus frontalis ) feeding and walking head on showing the bent bill . manawatu estuary , manawatu , new zealand . september .

the wrybill became fully protected in new zealand in 1940 ( 2 ) . research is currently being undertaken into the impact of predation on wrybill population numbers ( 2 ) . long - term monitoring is vital in determining the overall trends in this threatened species ( 2 ) .

embed this arkive thumbnail link ( " portlet " ) by copying and pasting the code below . < a href = " urltoken " title = " arkive species - wrybill ( anarhynchus frontalis ) " > < img src = " urltoken " alt = " arkive species - wrybill ( anarhynchus frontalis ) " title = " arkive species - wrybill ( anarhynchus frontalis ) " border = " 0 " / > < / a >

wrybill . front view of adult male in breeding plumage . ohau river delta , mackenzie country , november 2006 . image © craig mckenzie by craig mckenzie

hay , j . r . 1984 . the behavioural ecology of the wrybill plover anarhynchus frontalis . unpublished phd thesis , university of auckland , auckland . urltoken

pierce , r . j . 1979 . foods and feeding of the wrybill ( anarhynchus frontalis ) on its riverbed breeding grounds . notornis 26 : 1 - 21 .

dowding , j . e . 2013 [ updated 2017 ] . wrybill . in miskelly , c . m . ( ed . ) new zealand birds online . urltoken

no other bird in the world has a bill like new zealand’s wrybill . its bill curves to the right , allowing it to probe for insects under river stones .

calls and songs : sounds by xeno - canto the wrybill gives short , high - pitched “tweeps” in flight , before landing . this bird does not call frequently .

davies , s . j . 1997 . population structure , morphometrics , moult , migration , and wintering of the wrybill ( anarhynchus frontalis ) . notornis 44 : 1 - 14 .

protection / threats / status : the wrybill has small population estimated at 3 , 000 / 3 , 300 mature individuals . it is threatened by deterioration of the habitat due to the encroachment of weeds with reduction of the water flow . predation by gulls , stoats and cats is probably substantial too . the increasing use of riverbeds for recreational purposes and floods are also important threats . the wrybill is currently listed as vulnerable .

riegen , a . c . ; dowding , j . e . 2003 . the wrybill anarhynchus frontalis : a brief review of status , threats and work in progress . wader study group bulletin 100 : 20 - 24 .

the wrybill or ngutuparore ( māori ) anarhynchus frontalis is a species of plover endemic to new zealand . it is unique in that it is the only species of bird in the world with an asymmetrically bent bill , which it uses to dig around river stones for freshwater invertebrates . measuring 20–21cm long and weighing between 43–71gm , the wrybill is slightly sexually dimorphic . the most distinctive feature of the bird is the long black bill , which is always curved to the right .

their normal call is a high–pitched staccato whistle resembling that of the banded dotterel , though it is rather more musical . the wrybill does not , however , call nearly so frequently as the dotterel , either when on the ground or in flight .

introduction : the wrybill is the only bird in the world with the bill tip curving to the right . this “tool” is adapted to the feeding behaviour of this species when the bird extracts insects and aquatic preys from rock crevices and under stones . the maori name is ngutuparore .

birds begin to leave the breeding grounds by late december ; this species is one of the first of the season to begin its migration ( 4 ) . using its specially adapted beak , the wrybill is able to forage under stones for invertebrates such as mayfly larvae ( 3 ) .

in those localities on the river - beds , where a diminishing stream leaves large shingle , there the bent bill is of some use . in such situations i have seen wrybill peck under a stone on its left , and then turn around to get the same stone on its right , and peck under it again . it must be admitted , however , that i have often seen banded dotterel act similarly , and , i daresay , just as often as i have seen wrybills do it , so that any advantage the wrybill has would seem to be very slight .

behaviour in the wild : the wrybill feeds on aquatic invertebrates during the breeding season such as worms , annelids and gastropods , and also mites , spiders , and eggs , larvae and pupae of numerous insects , including the adults . it also feeds on crustaceans , fish and their eggs .

the recently rediscovered new zealand storm - petrel , refound for the first time in 150 + years by brent stephenson and sav saville from wrybill birding tours , nz is a key target species . our hauraki gulf pelagic not only finds several other endemic breeding seabirds , but specifically targets this species .

the wrybill is migratory . it breeds in c south i and moves after the breeding season to the mudflats of estuaries and harbours on n north i . they migrate along the e coast of south i and the w coasts of north i . the adults show strong site - fidelity to their wintering grounds .

but on intertidal mudflats , the wrybill tilts the head to the left and performs sideways sweeps with the bill , usually from right to left , in order to sieve preys from water and mud . it often forages in shallow pools , in shallow water in shingle riverbed , on intertidal mudflats and along riverbanks .

the wrybill is an internal migrant . after breeding , almost the entire population migrates north to winter in the harbours of the northern north island , notably the firth of thames and manukau harbour . on their wintering grounds , wrybills form dense flocks at high - water roosts ; the highly - coordinated aerial manoeuvres of these flocks have been described as resembling a flung scarf .

in most writings on the wrybill , the greatest emphasis is laid on the importance of its peculiar bill , but , in my opinion , its colouration is far more important , for this harmonises so perfectly with that of rounded stones among which it breeds that if the bird remains stationary ( and it has a habit of doing so ) it becomes exceedingly difficult to detect .

habitat : the wrybill breeds inland on large , fast - flowing rivers running in bare beds of shingle and sand . occasionally , it may breeds in smaller , slower - running rivers . outside the breeding season , it frequents shallow estuaries and sheltered coasts with large mudflats and muddy lagoons . it may occasionally frequent ploughed areas and muddy shores of small lakes and ponds in mountains .

this site is maintained and copyrighted by wrybill birding tours , nz © 2016 . all photos ( unless otherwise stated ) were taken by brent stephenson @ eco - vista and are copyrighted . the use of any image without permission is not allowed . however , all photos on this site are for sale , please email brent for more information or check out eco - vista ' s website for details .

reproduction of this species : the laying starts in mid - september and continues throughout october . a second clutch may occur from november to late december . the wrybill’s nest is on the ground , in sand , among large , smooth , rounded stones . it is protected by a piece of wood or a larger stone . occasionally , small pebbles are added around the edges of the nest , as lining for the eggs .

wiersma , p . & kirwan , g . m . ( 2018 ) . wrybill ( anarhynchus frontalis ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .

as to the wrybill’s chances of survival , one would say that they are good , at any rate on those rivers whose beds among the hills above the gorges are sufficiently wide to be of the bird’s liking , for there the winter temperature seems to have prevented the yellow lupin from getting a foothold . but the open shingle must be a quarter of a mile wide before the wrybill would make their home on it . they were never numerous , and even thirty years ago i should think two to three pairs to the square mile on the rakaia was their maximum population , and , since the lupin has filled the river bed , below the bridge , their numbers have certainly been much reduced . their chief bird enemies are harriers and black–backed gulls , yet the toll that both these take would not equal that of stoats and weasels during the birds’ breeding season .

the wrybill is a small pale plover which breeds only in braided rivers of the south island . it is the only bird in the world with a laterally - curved bill ( always curved to the right ) , which it uses to reach insect larvae under rounded riverbed stones . wrybills are completely dependent on braided rivers for breeding ; all their life stages are predominantly grey , and highly cryptic among the greywacke shingle of the riverbeds .

the wrybill is a distinctive wading bird , which possesses a uniquely bent bill . the tip of the black bill is curved to the right , this adaptation allows these birds to forage under stones for insect larvae ( 3 ) . the plumage is ash - grey above with white underparts . during the breeding season , individuals have a black band across the upper chest and males also have a band on the forehead ( 2 ) .

wrybill birding tours , nz …… . started out as a couple of birding mates making the decision to set up their own independent bird - guiding business . we felt there was a need for real birders , who were passionate about birding , really knew their new zealand birds and birding sites inside out , and were based in new zealand , with access to up - to - date information on the new zealand birding scene . sure we know our natural history as well , but we are birders , the name says it all !

the wrybill is monogamous . nesting and feeding territories are usually strongly defended . the mating season starts when the birds return to south i . they are very active , chasing each other in the air and on the ground . if the pair is disturbed during the courtship displays by another bird , the intruder is driven away by the bird of the same sex . it runs after it with outstretched neck among the stones , while maintaining its balance by slightly opening the wings . the female shows great submission to the male once mated . they nest solitary in loose colonies with nests at least 400 metres apart , and never less than 40 metres .

in the wrybill , however , the bill , which is one and a quarter inches long , is bent to the right in the middle , the end being offset at an angle of 12 degrees to the base . this is claimed to be an adaptation to its existence on the shingle river - beds of canterbury , where it nests , the bent bill supposedly enabling it to get its food the more readily from under stones . and no doubt it does this , when the occasion demands , but there can be very few occasions when this peculiarity is of any decided benefit to its possessor , for over nearly all the river - beds on which the bird feeds , the stones are so much buried in sand as to make the bill quite unnecessary . moreover . the bird spends less than half the year on shingle river–beds , living for the remaining months on mud–flats , and sea beaches , where its abnormality can be of no benefit .

html public " - / / w3c / / dtd xhtml 1 . 0 strict / / en " " urltoken "

among new zealand’s many bird curiosities , writes edgar stead , the wry - billed plover is often not included , but it certainly ought to be , for it is the only bird in the world which has its bill bent sideways . the crossbills of the northern hemisphere might claim that they share in this distinction , but their bills are symmetrical , the upper mandible being bent one way , and the lower , the other .

it has to be taken into account also , that while the bill , bent as it is to the right , gives the bird some advantage when feeding under stones on its right , this structure is a very distinct disadvantage to the bird when it wishes to feed on its left . a bill with an upwards curve , one would have thought , would have been of greater use .

wrybills are in their full mating plumage when they return south , the black band across the chest being very conspicuous ; the narrow white frontal band , with the brownish band behind it , not showing up at all except at very close range . the whole of the upper parts are blue–grey , but with age the feathers lose a certain amount of their blue tint , and also , particularly in the case of scapulars , become much frayed at the edges . it has been stated by some writers that the black chest band is wider on one side than the other , but this is not the case , either in fact or in appearance .

wrybills nest on high shingle spits , choosing a place where the stones are rather large , and clear of all growths and drift . a mere scratching in the sand , occasionally with a few small pebbles added around its edges , serves as a receptacle for the two eggs , which are wonderfully like the stones among which they are laid .

both birds take their turn at incubation , and sit very close , so that they will often allow a person to pass within twenty yards of them without getting off the eggs . owing to the remarkable similarity of their colouring to that of the surroundings , wrybills are easily passed when they keep still , and nests would certainly remain undetected if the birds did not move . but if the bird does get off the nest , and run towards an intruder ( wrybills do not fly around intruders in the same way that banded dotterel do ) , the latter has only to stand still for a few moments , when the bird will , as a rule , run straight back , and sit on its nest , even though it be in full view , and not fifteen yards away .

laying commences about the middle of september and continues throughout october . i have found eggs in november , but i think it probable that they were second clutches , the first having been destroyed . the young when hatched are thickly covered with down , white on the breast and underparts , the whole of the upper surface being stone grey , faintly marked with smoky black . when they are hatched , and even as an embryo in the egg , they have the bend in the bill well defined . they leave the nest within a day of hatching , and follow their parents about in search of food . by the time the young are full grown , the old birds are moulting into their winter plumage , when they closely resemble the young — having no black chest band , and no frontal bands — and then old and young leave the river–beds . the earliest families have left by the end of december , and all of them have gone by the end of february . sometimes they stay awhile on the coastal lagoons , and i have several times seem them on the shores of lake ellesmere in january and february , but they are the first birds to go north , migrating nearly two months earlier than the main body of dotterel and stilts . they spend the winter on the sea coast of the north island , frequenting the mouths of rivers , and the mud–flats of the big harbours of the far north . so far as my information goes , it would seem they travel up the west coast of the north island in greater numbers than they do up the east .

wrybills always display when their eggs or young are handled , running around with the wing near the intruder trailing on the ground , the other lifted in the air ; the feathers of the rump are raised ; the tail spread fanwise , and depressed so that the tip is almost on the ground ; and the bird all the time makes a continuous purring noise .

it is sincerely to be hoped that this most interesting little bird does survive , for , on its nesting ground , it exhibits in all its stages — adults , eggs and young — the most amazingly perfect protective colouration that there is among new zealand birds .

20 cm . , 55 g . , pale grey upper parts , black bill , legs grey green , white forehead tinged with black in breeding plumage , breeding adult underparts white except for black band across the chest .

the main breeding rivers in the south island are the rakaia , rangitata , waimakariri , and upper waitaki . after breeding the head to the tidal harbours of northland , auckland , and south auckland , the firth of thames .

ibis , 1869 . buller , walter lawry , birds of new zealand , 1888 .

heather , b . , & robertson , h . , field guide to the birds of new zealand , 2000 . stead , edgar f . , life histories of new zealand birds , 1932 .

thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .

turbott , e . g . 1990 . checklist of the birds of new zealand . ornithological society of new zealand , wellington .

20 cm . stocky , pale grey plover , tip of black bill turned to right . ash - grey crown , nape , upperparts . white underparts . black band across upper chest thick in breeding male , thinner in breeding female , sometimes absent in non - breeding birds . black frontal band above white forehead in male , absent in female . juvenile breast - band absent , back feathers tinged with white .

this species is listed as vulnerable because it has a small population , which is undergoing a continuing decline owing to habitat degradation and the impacts of introduced predators .

. it is found on over 26 riverbeds , but is only common on 10 . it winters mainly north of 38°s in the north island . in the last 40 years , population counts have varied between 3 , 000 and 5 , 000 individuals ( sagar

, probably reflecting the difficulty in surveying the species ( j . e . dowding

, which is supported by preliminary results from a long - term demographic study ( j . e . dowding

the population is estimated to number 4 , 500 - 5 , 000 individuals , roughly equating to 3 , 000 - 3 , 300 mature individuals .

analysis of wintering flocks indicates a slow decline over the last 40 years ( veitch and habraken 1999 ) , which is supported by preliminary results from a long - term demographic study ( j . e . dowding

it breeds on braided riverbeds , and frequents sheltered estuaries and coasts over the non - breeding season . nests are built within 250 m of running water , and are usually hollows in bare shingle , lined with more than 100 small pebbles ( marchant and higgins 1993 , j . e . dowding

. it lays two eggs . young usually begin to breed at two or three years of age ( marchant and higgins 1993 )

, the average adult life expectancy is c . 5 . 4 years ( hay 1984 )

. 2012 ) . land intensification in the high country will have similar effects on reducing water flow , as well as increasing the concentration of nutrients in rivers , further encouraging weed growth ( rebergen 2011 ) . the extent of predation by stoats

. the recent illegal introduction of rabbit haemorrhagic disease has resulted in the localised switching of some predators to a diet containing proportionately more birds . predation by kelp gulls

may pose an increasing threat as they become more numerous in association with human activities ( j . e . dowding

. increasing use of riverbeds for recreational purposes and floods are also threats ( marchant and higgins 1993 , a . grant

1999 ) . water quality deterioration and disturbance pose threats at the species ' s wintering grounds in auckland and northland ( rebergen 2011 ) . further threats may include the conversion of coastal habitat for aquaculture , development of wind farms , and the spread of mangroves ( d . melville

. research on the impact of predation and prey - switching is being undertaken . predator control for black stilt

. project river recovery carries out habitat restoration and predator research in the mckenzie basin ( a . grant

. continue to monitor wintering aggregations . control introduced predators and invasive plants at important sites . control the recreational use of riverbeds , perhaps by delimiting areas where humans are excluded . identify and monitor key areas of habitat ( rebergen 2011 ) . increase awareness of river - dwelling birds as part of the current government policy formulation on freshwater resource management ( d . melville

to make use of this information , please check the < terms of use > .

wrybills are small , pale plovers that are much more approachable than most new zealand waders . their underparts are white , with a black upper breast band from mid - winter to the end of the breeding season . the upper parts and sides of the face are pale grey , and the forehead white . the bill is long and black , with the distal third curved 12 - 26° to the right . the legs are dark grey to black . the sexes are alike in eclipse plumage ; juveniles lack the black breast band . in breeding plumage , males are distinguishable by a black line above the forehead ; this is highly variable however , and difficult to see in some individuals .

voice : the most common call is a ' chip ' that appears to indicate alertness . rapid ' churring ' is used when chasing intruding banded dotterels or other wrybills , and very quiet ' grating ' call used to communicate with chicks . flocks in flight ( and sometimes when milling on the ground ) may indulge in excited high - pitched ' chattering ' .

similar species : wrybills are unlikely to be mistaken for other species in breeding plumage or if the bill is seen well . in eclipse plumage the banded dotterel is superficially similar , but has browner upper parts and short straight bill . confusion is possible with some rare transequatorial migrant waders , notably sanderling and terek sandpiper , but these lack the black breast band , and their bill shapes differ .

wrybills breed only in the south island , east of the main divide . the large majority of the population breeds in canterbury between the waimakariri river in the north , and the rivers of the upper waitaki ( mackenzie ) basin in the south . main strongholds are the rakaia , upper rangitata , and mackenzie basin . four rivers in otago ( hunter , makarora , matukituki , and dart ) and two in north canterbury ( ashley and waiau ) have small populations . wrybills previously bred in marlborough , but their breeding range has contracted southwards over the past 120 years .

from january to july , wrybills are present in harbours of the northern north island , mainly manukau and firth of thames , smaller flocks elsewhere . during migration , small flocks are often seen briefly at south island east coast lakes and estuaries , and flocks may settle at rivermouths in the southwestern north island .

wrybills breed exclusively on braided riverbeds . on their wintering grounds , they feed on inter - tidal mudflats in harbours and estuaries . high - water roosts are usually near foraging areas , commonly on local shellbanks and beaches ; occasionally on pasture . in the upper manukau harbour , birds regularly roost on roofs of large buildings ; occasionally on tarmac at auckland airport . on migration , flocks generally follow coastlines , but are not averse to flying overland . a high proportion of the population passes through lake ellesmere on both migrations .

counts on breeding grounds are impractical – wrybills are highly cryptic and widely dispersed . combined counts from wintering flocks suggest a total population of 5000 - 5500 . counts show high variability ( which obscures trends ) , but the population is thought to be declining slowly .

the main threats faced by wrybills are predation ( by introduced mammals and native birds ) , flooding of nests , and loss or degradation of breeding habitat . threats vary temporally and spatially , and their relative importance overall is unclear . actual and potential threats to habitat include loss to hydro - electric power schemes , abstraction of water for irrigation , weed growth , and disturbance caused by a wide range of recreational and commercial activities . on wintering grounds , predation of adults has been recorded , and some loss of roosting habitat is likely as mangroves encroach . wrybills appear to be prone to collision with man - made objects . occasional significant strikes have occurred at auckland airport , and collisions with fencelines and powerlines have been recorded .

a small proportion of the population is managed through predator control programmes . the largest such area covers the whole tasman river catchment , but benefits are equivocal to date . a few small areas are managed by community groups ( e . g . the lower ashley river ) .

wrybills breed in monogamous dispersed pairs . territories may overlap with those of other species ( e . g . banded dotterel , black - fronted tern , pied stilt ) , but are vigorously defended against other wrybills . the nest is a shallow scrape in the gravel , lined with many small stones . the normal clutch is 2 ; first clutches are laid in september or october . replacement clutches laid after loss may occur through to january . some pairs will double - brood if they are successful early . incubation is shared and is protracted ; accurate data are few , but values from 30 - 36 days recorded . chicks fledge after 35 - 40 days ; chicks are guarded by one or both parents during their first 3 weeks at least , often becoming increasingly independent before fledging .

wrybills usually allow a close approach . when incubating , they rely on camouflage to avoid detection , and flush from the nest very late . distraction displays are used to defend nests ; banded dotterels are aggressively chased if close to eggs or chicks . chicks freeze to escape detection ( especially when small ) , and swim well from a few days old . away from the breeding grounds , wrybills are highly gregarious and form dense flocks , normally also very approachable .

northward migration typically begins in late december , and peaks in january . southward migration is mainly in august and early september ( adults ) , with some first - year birds ( which will not breed ) following in october - november .

on the breeding grounds , wrybills consume a wide range of aquatic invertebrates , but predominantly mayfly and caddisfly larvae . on wintering grounds , a range of small marine and littoral invertebrates are taken ( including annelid and polychaete worms , small molluscs , and insects ) , and the occasional small fish .

dowding , j . e . ; moore , s . j . 2006 . habitat networks of indigenous shorebirds in new zealand . science for conservation 261 . department of conservation , wellington .

heather , b . d . ; robertson , h . a . 2005 . the field guide to the birds of new zealand . 2nd edition . penguin : rosedale , auckland .

marchant , s . ; higgins , p . j . ( eds ) 1993 . handbook of australian , new zealand and antarctic birds . vol . 2 , raptors to lapwings . oxford university press , melbourne .

incubation behaviour shared incubation length ( mean ) 30 - 36 days nestling type precocial nestling period ( mean ) leave nest within 24 hours of hatching of second egg age at fledging ( mean ) 35 - 40 days age at independence ( mean ) 30 - 50 days age at first breeding ( typical ) 2 plus years age at first breeding ( min ) 1 years maximum longevity 22 plus years maximum dispersal migrate annually 700 - 1000km each way .

a relatively small pale plover with a long black bill curving to the right , dark grey to black legs and , in breeding males , a black line above the white forehead . the underparts are white , with a black upper breast band from mid - winter to the end of the breeding season , and the upperparts and sides of the face are pale grey .

the laying season runs between september and october ; a clutch of two eggs is laid into a slight depression amongst the gravel . both parents take it in turn to incubate the eggs that are well camouflaged against the shingle , resembling the stones around them ( 4 ) . the parents also rely on camouflage to remain undetected with their ash - grey plumage barely visible amongst the stones ( 4 ) . the chicks are able to leave their nest within a day of hatching and follow their parents on foraging trips ( 4 ) .

endemic to new zealand , wrybills migrate annually from breeding grounds in canterbury and otago ( on south island ) , to spend the winter in northern areas of north island ( 2 ) .

breeding occurs on rivers where there are large amounts of bare shingle . the wintering sites to the north tend to consist of mudflats at the mouths of large rivers ( 4 ) .

classified as vulnerable ( vu ) on the iucn red list 2006 ( 1 ) .

large flocks of wrybills were recorded in the early 19th century but the species has since undergone a long - term decline , principally as a result of habitat loss at breeding sites ( 2 ) . the shingle beds that comprise the nesting habitat of this species have decreased in size due to the encroachment of weeds , and altered river flooding regimes caused by hydroelectric schemes ( 2 ) . it is also likely that predation by introduced stoats ( mustela erminea ) and cats has played a large part in the decline of this ground - nesting bird ( 2 ) .

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

endemic a species or taxonomic group that is only found in one particular country or geographic area . invertebrates animals with no backbone . 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 .

getty images 101 bayham street london nw1 0ag united kingdom tel : + 44 ( 0 ) 800 376 7981 sales @ urltoken http : / / www . urltoken

terms of use - the displayed portlet may be used as a link from your website to arkive ' s online content for private , scientific , conservation or educational purposes only . it may not be used within apps .

myarkive offers the scrapbook feature to signed - up members , allowing you to organize your favourite arkive images and videos and share them with friends .

team wild , an elite squadron of science superheroes , needs your help ! your mission : protect and conserve the planet’s species and habitats from destruction .

wildscreen is a registered charity in england and wales no . 299450 wildscreen usa is a registered 501 ( c ) ( 3 ) non - profit organisation in the usa

we also try and see as many marine mammal species as possible on our tours , with the tiny hector ' s dolphin highly likely to be seen .

for our pelagics we use small charter boats that are specifically certified for taking passengers . the operators we are use are people we know and trust and provide the best pelagic experienced based on working with us over many years .

the new zealand breeding endemic black ( parkinson ' s ) petrel is generally seen on several of the north island pelagics .

near okarito we will spend an evening in search of the rarest of the kiwi species - the okarito brown kiwi . we have a chance of seeing four species , plus great - spotted kiwi which is unlikely to be seen , but generally heard .

the new zealand forest can be extremely beautiful , but with all this moss in a south island nothofagus beech forest it means there can sometimes be rain .

although not specifically a photographic tour , our 21 - day birding tours allow excellent photographic opportunities .

we target three species of penguin during the tour , with the largest being the yellow - eyed penguin . we generally find this bird both at sea and near its breeding sites on land .

small group sizes and relaxed travel times generally allow us to stop and make the most of birding locations we are enjoying .

a courting pair of dusky dolphins ( lagenorhynchus obscurus ) leap from the water . kaikoura , canterbury , new zealand .

as large as the wandering albatross , the southern royal albatross is likely to be seen on at least a couple of our pelagic opportunities .

the ancient looking takahe is a flightless endemic rail , considered extinct for almost 50 years , and now found on several predator free offshore islands .

the south island is renowned for its spectacular scenery , and rightly so . this is franz josef glacier on the west coast of the south island and a place we stay near and stop at .

one of the worlds rarest shorebirds , the black stilt , is a key endemic we aim to see .

we use clean and tidy accomodation around the country . some such as this spot we use on our first night are in beautiful scenic locations and birding around the accomodations can often be good .

a male south island wren ( xenicus gilviventris ) perched on a rock in its alpine habitat . homer tunnel , fiordland national park , new zealand . january .

buller ' s albatross is surely one of the most beautiful of the albatrosses , and one on our target list .

the south island is renowned for its spectacular scenery , and rightly so , with this being part of the road through to the famous milford sound , which we visit if time allows .

the endemic blue duck can be difficult to find , but we visit the best location ( and several others ) for this species in the north island , and also try to find it in the south island .

southern brown kiwi ( apteryx australis ) feeding for crustaceans on a sandy beach . no flash used , just dim flashlight . ocean beach , stewart island , new zealand .

new zealand dotterel ( charadrius obscurus ) , sometimes known as red - breasted plover , feeding on the shoreline . waipu estuary , northland , new zealand .

the south island is renowned for its spectacular scenery , and rightly so , with views like this of mount cook ( nzs highest peak ) sometimes being possible .

it is not only about birding , but having a great time as well .

the alpine kea makes for a strange parrot , but will be seen in several places on the south island . an incredibly curious bird they provide much entertainment .

north island kokako ( callaeas wilsoni ) feeding on leaves in the sub - canopy . tiritiri matangi island , auckland , new zealand .

we generally grab lunch from a cafe or bakery as we travel , and then enjoy it in a place where we will maximise our birding or scenic opportunities .

we generally grab lunch from a cafe or bakery as we tarvel , and then enjoy it in a place where we will maximise our birding or scenic opportunities .

the largest flying bird in the world , the wandering albatross , can be seen almost at arms length off kaikoura and possibly also during our other pelagic opportunities .

adult male yellowhead ( mohoua ochrocephala ) peering into moss whilst foraging . haast pass , west coast , new zealand . january . endangered .

a cook ' s petrel ( pterodroma cookii ) in flight , showing the underwing pattern . hauraki gulf , auckland , new zealand .

generally we dine at cafes and resturants , choosing from the normal menu , however on some nights we take the opportunity to get a little more personal , with catered meals providing an introduction to real kiwi food .

new zealand falcon is our only endemic diurnal raptor , and we have excellent opportunities to see this species both in the north and south islands .

the white - capped albatross , one of the larger ' small ' abatrosses is frequently very common during our stewart island pelagic .

our pelagic out of kaikoura offers unrivalled close views of several species of albatross and other tube - nosed seabirds - not to be missed .

buller ' s shearwater is a new zealand endemic breeder , and will be seen commonly during several of our north island pelagics .

pelagic birding trips are just one of our specialties , with unrivalled knowledge within new zealand . our rediscovery of the supposedly extinct new zealand storm - petrel in 2003 , as we were just starting to operate our business , was an amazing sign that we were really doing the right thing . however , we also know our land - birds just as well and know you will enjoy your new zealand birding experience with us !

so here we are…… . sav saville and brent stephenson , and recently due to being so popular , and restricting our group sizes to just 8 people , we have had to enlist the help of three other great new zealand birders – phil hammond , matt jones , and neil robertson . the decision to hire other guides was not one we took lightly , but as their clients have told us , they are excellent and fit well within our team ! our list of services can be found here , with anything from pre - trip planning advice , to a full north and south island organised tour ( including stewart island ) , and anything in between . as sav , brent , phil , matt , and neil live in different parts of the country , we are also able to offer guided trips around our ‘neck - of - the - woods’ for one or more days , and make custom itineraries . trip reports from our pelagics , personal birding trips , and guided trips can be found here . if you are visiting new zealand , then email us to let us know how we can help .

we must be doing something right if pelagic experts and overseas field leaders think we do a good job ! check out endorsements from world class seabirders here and see our testimonials page here .

we only operate in new zealand , choosing to run tours on our home turf where we know things best . that doesn’t mean we don’t travel and bird overseas though !

our 21 - day tours are only run by birders we know and trust ! sav & brent have been running tours by themselves since 2002 , but have now enlisted the help of phil hammond , matt jones , and neil robertson . both phil , matt , and neil are excellent guides and birders and we have no hesitation bringing them both onboard as guides .

d’s conservation estate . this means some of your fee goes towards ensuring the conservation of the birds you will see . we also hold concessions for many of the

we love getting out and about birding , we are birders through and through – we even go birding on the weekend ! but , we are also keen on plants , invertebrates , and other new zealand wildlife . so a tour with us is not just hardcore birding , we take time to see the country and other things of interest during our 21 - day tours .

plus we love showing new zealand and its fantastic birds to overseas birders ! read client testimonials here .

we are pleased to announce that we will again be at the british birdwatching fair in 2018 ! we will be sharing a stand with our good friends from albatross encounter - marquee 2 , stands 46 & 47 . so come and see both brent and sav there during the weekend of 17 - 19 . . .

well our last places are filling on this upcoming seasons tours - 2018 - 2019 summer . we have just just three places left in five 21 - day tours confirmed in late 2018 , and just a couple of places left in the three confirmed tours in early 2019 . there have been recent . . .

well it is the start of the new year - happy 2018 every one ! we are gearing up for our early 2018 21 - day tours which kick off in a week or so . looking forward to some excellent birding . we have also just added dates for our pelagics in warkworth ( into the hauraki . . .

well our tours in november and december 2017 went really well , with both tours run getting almost all of the ' gettable ' endemics . low numbers of the less common arctic shorebirds accounted for lower numbers than usual on these tours , but these really are additional . . .

sav will be attending the american birding expo , which is being held this weekend - 29 september to 1 october 2017 , in philadelphia . it is being held at the greater philadelphia expo centre , and sav will be around all weekend . so if you are in the area and wanting . . .

wrybills are classified as a threatened species due to their low and declining numbers .

birds on braided rivers have evolved to feed in distinct ways . specialisation minimises competition for food between the bird species .

wrybills feed in shallow channels , riffles and the edges of pools . their bent bill is specially adapted to allow them to reach under stones for mayfly larvae .

when food is scarce , they move into stableside channels and pond areas to find food . in winter , wrybills migrate to north island harbours and feed in flocks on the mudflats .

each pair of wrybills , stilts , dotterels and oystercatchers defends a territory and nests alone . the chicks are active soon after hatching , following the parents as they forage for food . within hours , newly - hatched chicks can hunt for food and swim if necessary .

breeding on a riverbed is a risky business . many eggs and chicks do not survive . riverbed birds have adapted to cope with floods and are able to renest if eggs or chicks are lost .

birds with good nesting sites are more likely to raise chicks successfully . the best nest sites have :

swamp harriers / kāhu and black - backed gulls / karoro are natural predators of braided river birds . these avian predators have taken advantage of changes made by humans and their numbers have increased dramatically .

braided river birds have good camouflage and use distraction to cope with avian predators . wrybills , oystercatchers and dotterels often pretend to have a broken wing to lead predators away . terns , gulls and oystercatchers may dive - bomb and call loudly .

however these defences against avian predators are little use against introduced predators such as cats , stoats , ferrets , rats and hedgehogs . these are the main threats .

ensuring the survival of the birds ' natural open habitat is important in combating predation .

the fragile ecology of the braided river system is being destroyed by the invasion of weeds .

human activity including land development and recreational activities disturb nesting birds . birds may abandon their nests if disturbed .

call 0800 doc hot ( 0800 362 468 ) immediately if you see anyone catching , harming or killing native wildlife .

only take dogs to areas that allow them , and keep them under control .

don ' t drive on riverbeds , or keep to formed tracks if you have to .

braided rivers are important features of the upper waitaki basin . their distinctive wildlife and plant communities make them unique worldwide .

recommended citation birdlife international ( 2018 ) species factsheet : anarhynchus frontalis . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .

the male has a white forehead and pale grey crown , nape , back , wings and tail and a white throat , breast , belly and rump , with a thin black band across the breast . this band is thinner in the female , and much less distinct in both sexes in the non - breeding season . males have a small black bar between the white forehead and the grey crown . as with the breast band , this is reduced in the non - breeding season .

their eggs are blue - grey and lightly speckled , making them well camouflaged against river stones and pebbles , which generally make up the main structure of a very simple nest .

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 .

no flash player has been set up . please select a player to play flash videos .

a bird walking slowly to detect prey , catching a worm , cleaning and eating it .

a bird looking for food , catching a worm , cleaning it on water and swallowing it .

josep del hoyo , doug and denise norris , pieter de groot boersma , greg baker , nick talbot , mkennewell , mauriravasini , brooke clibbon .

nick talbot , glenda rees , pluvius , josep del hoyo , greg baker , georges olioso , marco valentini , rick and elis simpson , rémi bigonneau , alex berryman , frédéric pelsy , ken havard , tomas grim , martin främke .

new zealand : breeds in c south i ( canterbury and otago riverbeds ) ; winters mainly in n north i .

20–21 cm ; 43–68 g . rather plump , greyish plover with conspicuous long bill curved to the right ; male has black frontal bar and breastband . female has duller , . . .

breeds inland on large braided rivers , occupying large bare beds of shingle and sand , with . . .

during breeding season takes aquatic invertebrates , including flatworms , annelids , gastropods , mites , spiders , and eggs , larvae , pupae and . . .

lays late aug to late oct , with second clutch late oct to late dec . monogamous on long - term basis . solitary ; holds vigorously defended . . .

migratory . moves from breeding grounds in c south i to mudflats of estuaries and harbours on n end . . .

vulnerable . hunted prior to 1940 ; after hunting ceased , population increased , but stabilized by early 1960s , with c . 5000 birds . in 1994 , total of 5111 birds counted , with c . . .


harpalus anxius is a species of phytophagous and xerophilous ground beetle that is native to palearctic ( including europe ) and the near east .

in europe , it is only absent in the following countries or islands : andorra , the azores , the canary islands , the channel islands , crete , cyclades , cyprus , dodecanese , the faroe islands , franz josef land , gibraltar , iceland , madeira , malta , monaco , the north aegean islands , norway , novaya zemlya , portugal , san marino , the savage islands , sicily , svalbard and jan mayen , and vatican city .

its presence on the balearic islands and sardinia is doubtful .

harpalus anxius

harpalus anxius <EOT> no one has contributed data records for harpalus rubripes yet . learn how to contribute .

no one has contributed data records for harpalus atratus yet . learn how to contribute .

harpalus ( harpalus ) anxius ( duftschmid , 1812 ) family : carabidae size : 8 . 1 mm ( 6 . 1 to 8 . 6 mm ) distribution : europe to west siberia , west china ecology : inhabitants of steppes , loves warm and dry areas location : france , ardeche , les vans leg . w . veldkamp , 10 . vii . 1979 ; det . u . schmidt , 1986 photo : u . schmidt , 2017

irish status : like the majority of harpalus , this species is of restricted distribution due to the scarcity of free - draining soils , particularly in the west . its range appears to be entirely coastal except for a few inland sites on sandy or morainic soils .

seasonal dynamics of activity , as well as the age structure of the populations of harpalus rufipes from reedbeds along the river khara ( a ) and pseudotaphoxenus rufitarsis major from the lakeside salt - marsh ( b ) , in 2006 ( see figure 2 for further explanations ) .

seasonal variation in abundance curves and reproduction aspects in four resident carabid species coupled with abundance of a migrant - species harpalus rufipes from reedbeds , combined data for 2006 / 07 ( r and l right and left y axis , respectively ; n ( ex . ) – number of specimens ) .

as such , the contribution of migrants to the trophic structure of carabid communities is not apparent and requires further research . that a particular carabid species inhabits and breeds in , and even dominates a certain habitat , is only a hypothesis that needs corroboration each time . species with high abundance levels and high frequency of occurrence in a particular habitat can belong to both labile and stable components . thus , in the lake elton region , calosoma auropunctatum , dolichus halensis , amara aenea , harpalus calceatus , harpalus rufipes , harpalus distinguendus and anisodactylus signatus belong to the labile component in all the habitats where they occur ; cephalota elegans comprises the main element of the stable component in several azonal habitats ; while calathus ambiguus , cymindis lateralis and pseudotaphoxenus rufitarsis maior play the main role in the composition of the stable component in the majority of zonal habitats ( appendix ) . overall , 65–75 % of the species diversity of both individual habitats and the landscape as a whole comprised of non - residential species . it is important to note that almost half of the migrants ( 41 of 94 species ) failed to breed in any of the studied habitats . thus , the distances of their movements are substantially greater compared to the size of the site . so , the migrations of such species should be characterised at the landscape scale .

overall , we collected 34 , 108 individuals belonging to 98 carabid species . we recorded 65 species in rice field banks , 73 species in buffer strips and 78 in restored habitats . eight species were found only in rice field banks ( agonum sexpunctatum , amara nitida , cicindela campestris , clivina collaris , harpalus albanicus , harpalus pumilus , ophonus cribricollis , pterostichus macer ) , 6 species only in herbaceous buffer strips ( amara littorea , bembidion quadripustulatum , bradycellus verbasci , dolichus halensis , ophonus azureus , panagaeus cruxmajor ) , 2 species only in arboreal buffer strips ( badister sodalis , drypta dentata ) , 2 species only in herbaceous restored habitats ( acupalpus elegans , ophonus diffinis ) and 11 species only in arboreal restored habitats ( acinopus picipes , acupalpus flavicollis , acupalpus notatus , agonum versutum , agonum viduum , limodromus assimilis , limodromus krynickii , nebria brevicollis , ophonus parallelus , paranchus albipes , synuchus vivalis ) . poecilus cupreus and pseudoophonus rufipes consituted about 55 % of the capture with 18 753 individuals .

the most interesting aspect of this carabid coenosis is the presence of several species with southern distribution , quite common in clay soil on the right bank of the po river , and known only in few stations north of the po river . among these species , we list acinopus picipes , amblystomus niger , dinodes decipiens , harpalus cupreus , harpalus oblitus , parophonus mendax , parophonus planicollis , pterostichus macer . although a comparison with the past coenosis is not possible for the lack of similar surveys in the area , it could be hypothesized that these species are recent colonizers ( 7 - 10 years ) . they are not reported in the historical catalogue of magistretti ( 1965 ) , and are also not listed in several recent faunistic investigations carried out in the lombardy lowland , particularly along the ticino river ( pasquetto 1992 , bogliani et al . 2003 ) , adda river ( conti 1991 ) , po river ( pilon et al . 1991 , rancati and sciaky 1994 ) and in milan ( pilon et al . 2010 ) , where potentially suitable habitats were sampled . even in an intensive survey along the po river included in piedmont region , only some of these species have been collected ( allegro and sciaky 2001 ) . if so , we could assume a tendency to a northward shift in the distribution of these species , according to what has been observed for other zoological groups well studied and that have great mobility , such as birds ( chen et al . 2011 ) and dragonflies ( ott 2010 ) .

soil disturbance is recognised as an important restoration measure for conserving biodiversity in sandy soils . we used a soil disturbance ( ploughing ) experiment in a sandy grassland as well as a semi - natural disturbance ( slope erosion enhanced by cattle trampling ) gradient on a sandy slope to test the soil disturbance effects on the ground - living beetle community . both experimental disturbance and semi - natural disturbance favoured sandy grassland specialists , but there was no overall effect on beetle richness and abundance . amara lucida and harpalus spp . were favoured by disturbance while calathus melanocephalus was disfavoured . experimental ploughing significantly increased the proportion of red - listed species in disturbed plots compared to non - disturbed controls . in the semi - natural disturbance gradient we found that the beetle community on the disturbed slope differed from that of the flat areas , and there were tendencies for a higher proportion of red - listed species on the slope . we conclude that increasing the area of bare sand in sandy grasslands can have positive effects on many threatened species . soil disturbance should thus be included as a regular measure in sandy grasslands under conservation management and as a measure to restore high biodiversity in areas where bare sand is rare .

although the sampling effort was quite different between 1983–1984 and 2002 , we assume that around 11 records of the 16 new species during the second sampling period can also be due to the invasion by elymus . in fact , several continental species were discovered after the invasion in relatively high numbers ( i . e . , more than five individuals ) , both on the dyke and in the salt marsh . among them , most species are linked to high contents of organic matter and a more pronounced litter layer ( e . g . , agonum muelleri , bembidion obtusum and the polyphagous pterostichus versicolor ) or are even partly phytophagous ( amara spp . and harpalus spp . : dajoz 1988 , ikeda et al . in press ) . conversely , halophilic species discovered in 2002 are hardly related to the invasion . pogonus littoralis and dicheirotrichus obsoletus could have been misidentified earlier , as these species are very similar to pogonus chalceus and dicheirotrichus gustavii , respectively ( forel and leplat 2005 , dhuyvetter et al . 2007 ) . dicheirotrichus obsoletus could also have been missed in 1983–1984 ( the sampling stopped in september ) as more than 89 % of individuals were caught in october - november during 2002 . tachys scutellaris appears as a new species in 2002 , but was present in 1983–1984 , but at another station located below the mean sea level ( slikke habitat : fouillet 1986 ) . the ‘appearance’ of several species , sampled in low numbers in 2002 , can be due to differences in sampling effort and / or to random catches .

enter the name or part of a name you wish to search for . the asterisk character * can be used as wildcard . e . g . ' papilio * ' . keep in mind that the search is only based on the full taxon name .

we are still having problems with the search feature . unfortunately we cannot give a timeline when the advanced search will be fixed .

the id resolving service for stable taxon ids is currently under maintenance . meanwhile , please use the name search in order to find the taxon page .

museum für naturkunde leibniz - institut für evolutions - und biodiversitätsforschung invalidenstr . 43 10115 berlin germany e - mail : fauna - eu ( at ) mfn - berlin . de website : urltoken

html public " - / / w3c / / dtd xhtml 1 . 0 transitional / / en " " urltoken "

description : an 8 . 5 - 12mm long black beetle with colourful and varied metallic reflections . the male is usually metallic bronze , green or blue , the female usually unmetallic black . phytophagous . common in dry grassland and coastal dunes .

world distribution : a eurasian wide - temperate ( 65 ) species and found across europe except the extreme north , south to asia minor and iran , and east to the river lena in siberia . introduced in n . america .

ecology : fairly eurytopic in areas which suit it , on arable land , in quarries , on dry grassland and on sand dunes at or near the coast .

if you have images for this taxon that you would like to share with nbn atlas , please upload using the upload tools .

in continuing your browsing of this site , you accept the use of cookies to offer you suitable content and services and realize visits statistics . learn more about cookies .

to get the picture , please visit : bouyon hervé herve . bouyon @ urltoken

any reuse of one or more photographs on this site is subject to an authorization request from the author . link to the code of intellectual property ( legifrance )

thank you for your contribution to the improvement of the inpn . the information submitted has been sent to an expert for verification and correction .

corresponds to a report on the basis of at least one observation proved within a period of 10 years ( 20 years for little - known invertebrates ) preceding the year and no presumption of extinction since obtaining the last data nor doubt on reproductive and implemented nature of this population . for migratory species , the presence indicated concerns areas of reproduction .

the last reliable sighting is older than 10 years compared to the reference date , no recent specific research and no presumption of extinction from that date [ vertebrates , invertebrates and plants well studied ( rhopalocera , grasshoppers , dragonflies . . . ) ] ;

the last reliable observation being older than 20 years , no recent specific research and no presumption of extinction from that date [ poorly known taxa : fungus , many invertebrates . . . ] .

this point covers the absence , more difficult by nature to demonstrate than presence . this status is based on one or more of the following criteria :

this status must be assigned to a department in which the presence of the species is casual .

particular case of absence due to a proven extinction less than a half century ago ( older disappearances are treated as " no probable or definite " ) .

in the state of knowledge , we can not comment on the presence or absence in the current department . this is the default status when not comprised in one of the previous categories or whenever there is doubt .

warning : the data available reflects the progression status of knowledge or the availability of the inventories . it should never be considered as comprehensive .

national inventory of natural heritage , website : https : / / inpn . mnhn . fr .

description : a medium - sized ( 6 . 8 - 8 . 2mm ) black phytophagous beetle of sandy heaths and dunes . eastern and southern .

world distribution : this species has a eurasian wide - temperate distribution ( 65 ) and is found across europe , except the north , to asia minor and siberia .

irish status : restricted to the south and east coasts between meath and kerry .

ecology : a strongly xerophilous species in ireland restricted to loose sand habitats in coastal fore dunes .

paleartic . open habitats . macropterous , with summer larvae . small size . zoospermatophagous .

common in the study area ( n = 331 ) . recorded in all habitat categories .

eol content is automatically assembled from many different content providers . as a result , from time to time you may find pages on eol that are confusing .

to request an improvement , please leave a comment on the page . thank you !

html public " - / / w3c / / dtd html 4 . 0 transitional / / en "

checklist of beetles of the british isles ( andrew duff ; http : / / www . coleopterist . org . uk ) , version 1 ( recommended )

we use cookies to optimise your experience when using this site . view our cookie policy and our new privacy notice .

asiatic - european . open habitats . macropterous , with summer larvae . medium size . zoospermatophagous .

common in the study area ( n = 180 ) . recorded in all habitat categories .

europe ( except the north ) and the balkans , where it prefers foothills to alpine regions ( arndt et al . 2011 ) .

it is a mesoxerophilous and polyphagous species , which prefers to live in forested areas ( varvara and apostol 2008 ) . it feeds on a mixture of seeds from agricultural crops , weeds and shrub species ( lundgren 2009 ) . in this study , it was only found in the wheat field in the homogeneous area ( n = 1 ) .

copyright © 1999 - 2018 john wiley & sons , inc . all rights reserved

enter your email address below . if your address has been previously registered , you will receive an email with instructions on how to reset your password . if you don ' t receive an email , you should register as a new user

warning : the ncbi web site requires javascript to function . more . . .

corresponding author : andrey v . matalin ( ur . akhcot @ nilatam _ a ) .

this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .

pitfall trapping is one of the most commonly used techniques to quantify terrestrial arthropods ( barber 1931 ) . the simplicity of the method and the possibility of data standardization are the main advantages of their application in numerous entomological studies . pitfall trapping is easy , and as such arthropods can be captured in different places at the same time . this explains the extensive use of pitfall traps in ecological investigations of ground beetles ( scherney 1959 ; skuhravý 1959 ; novák 1964 ; kabacik - wasylik 1970 ; tietze 1973 ; den boer 1977 ; brandmayr and zetto brandmayr 1986 ; østbye and hägvar 1996 ; gryuntal 2008 ; makarov and matalin 2009 ) .

however , doubts concerning the reliability of the obtained results were already expressed during the first pitfall trap studies and have been discussed subsequently ( for example , see adis 1979 ) . numerous factors have been found to affect pitfall trap catches , such as , the size of a trap and its inlet ( luff 1975 ; waage 1985 ; work et al . 2002 ; koivula et al . 2003 ) , the colour of a trap ( buchholz et al . 2010 ) , the presence and type of preservative ( luff 1968 ; feoktistov 1980 ; gryuntal 1982 ; karpova and matalin 1992 ; weeks and mcintyre 1997 ) and the ways of setting traps across habitats ( greenslade 1964 ; perner and schueler 2004 ; korczycski and sienkiewicz 2006 ) . in addition , the mobility of beetles in relation to both their physiological condition and the environment vary widely in the course of a season and between seasons ( den boer 1977 ; van huizen 1977 , 1979 ; baars 1979 ; matalin 1994 , 1997 , 2003 ; desender 2000 ) .

towards the second half of the 20th century it became clear that pitfall trapping reflected not as much the abundance as the locomotor activity of beetles . numerous steps have been taken to increase the reliability of the results of catches : changes in trap construction ( reeves 1980 ; boucher 1981 ; kuschka et al . 1987 ; loreau 1987 ; dufrêne 1988 ; makarov and tshernyakhovskaya 1990 ; karpova and matalin 1992 ; kuschka 1998 ) and in the type of preservative used ( louda 1970 ; feoktistov 1980 ; gryuntal 1982 ; pekar 2002 ) , exhaustive catches from enclosed areas ( kudrin 1971 ; gryuntal 1981 ; desender and maelfait 1986 ) , the calculation of correction coefficients from the re - trapping of marked specimens ( holland and smith 1991 ; raworth and choi 2001 ) , and the comparisons of dynamic ( pitfall trapping ) and static ( standard soil fauna quadrate sampling ) population densities ( kudrin 1966 ; arnoldi et al . 1972 ; desender and segers 1985 ; spence and niemelä 1994 ) . in spite of these important advances , standard pitfall trapping has ‘de facto’ become a standard technique used in synecological investigations of carabidae .

at the same time , when pitfall - trapped data are interpreted , the beetles’ migratory capacities are often ignored . this is because there is no universal technique for quantitatively estimating beetle locomotion ( den boer 1977 ; prisnyi 1987 ) . interpretation of life cycles to evaluate the demographic structure of local populations can provide a new approach to solving this problem . for example , a significantly deficient demographic structure recently observed in some carabid species in agricultural or disturbed habitats shows that in many places the populations are represented only by certain ‘age groups’ ( borkowski and szyszko 1984 ; wallin 1989 ; makarov and tshernyakhovskaya 1989 ; tshernyakhovskaya 1990 ; khotuleva 1997 ) . according to data obtained by bokhovko ( 2006 ) , five of the 11 dominant carabid species from arable soils in the kuban region , southern russia , demonstrated high abundance levels , coupled with incomplete demographic spectra . for example , in semi - centennial forest belts as well as in alfalfa fields , about 80 % of the dominants completed their development . on the other hand , in corn fields and in a forest belt with robinia , about 75 % of the carabid beetles did not complete their full life cycle .

the last case clearly illustrates the probable scales of migration in carabidae , showing that populations are often incapable of reproducing in such environments . however , it still remains unclear whether this situation is general or not . we can assume that the proportion of species with incomplete demographic spectra represented in pitfall traps is higher in disturbed habitats , while in undisturbed or moderately disturbed habitats , the sex and age structures of the populations are more or less balanced .

in the present study , we highlight a key methodological problem that the actual community structure ( e . g . , the roles of individual species ) cannot be understood based on pitfall counts alone . we also demonstrate how demographic analysis can be used to address this problem .

ground beetle communities in the lake elton region , volgograd area , south - eastern russia ( 49o12 . 47’n , 46o39 . 75’e ) were studied in 2006–2007 . lake elton is situated within the botkul - bulukhta drainless desert depression , which belongs to the caspian lowland . a strongly pronounced salt - dome structure is characteristic of this region , and desert steppes are typical plant associations in most of the habitats present ( nekrutkina 2006 ; safronova 2006 ) . dense reedbeds occur in the river valleys , in gullies at lakesides there are trees and shrubs , while lakesides near the mouth of most large rivers are characterised by salt - marshes . near the village of elton , all desert steppes are fragmented or transformed into pastures .

pitfall trapping was conducted in 10 habitats : six zonal characteristic of this particular biogeographical area , and four azonal present in a variety of biogeographical areas ( walter 1973 ; chernov 1975 ) . three selected habitats were located near the village of elton , while seven were placed on the north - western shore of lake elton , on the right bank of the river khara ( for more details see makarov and matalin 2009 ) . zonal habitats were represented by sagebrush and sagebrush - grassland steppe with varying degrees of anthropogenic disturbances ( strong near elton village , moderate on the northern slope of mt . ulagan , and weak in the watershed of river khara ) . azonal habitats were chosen along salinity and solar irradiation gradients ( strong in the lakeside salt - marsh , moderate in the salina on the floodplain terrace of river khara , and weak in reedbeds along river khara ) .

plastic cups of 0 . 5 l capacity and 95 mm upper diameter containing 4 % formaldehyde solution as a preservative were used . in each habitat , 10 traps were arranged along transects at 10 m intervals . the traps were checked every ten days from 10 may to 31 october in 2006 and from 1 april to 10 may in 2007 .

all captured carabids were dissected . based on gonad condition ( gilbert 1956 , skuhravý 1959 , van heerdt et al . 1976 , wallin 1989 ) , as well as on the degree of wear - and - tear of the mandibles , claws and cuticle ( houston 1981 , brandmayr and zetto brandmayr 1986 , butterfield 1986 , davies 1987 ) , six physiological states in the adults of both sexes were distinguished .

recently emerged beetles with soft and pale cuticle ; mandibles and claws sharp . ovaries thin , white or translucent without any trace of developing oocytes ; corpora lutea absent ; lateral oviducts very thin . testes thin and dull or relatively large and white ; accessory glands always thin and poorly visible .

cuticle fully hardened and coloured ; mandibles and claws pointed . ovaries compact , opaque and white , with or without distinctly visible oocytes , but always without ripe eggs ; corpora lutea absent ; lateral oviducts long and thin . testes opaque and white ; accessory glands no longer than half of the abdominal length , occupying less than a third of the abdominal space .

cuticle slightly worn ; mandibles and claws hardly or distinctly dulled . ovaries with ripe eggs ; corpora lutea absent or yellowish , hardly visible ; lateral oviducts wide . testes large and white or cream - coloured ; accessory glands long and white or light - yellow , filling more than three - quarters of the abdominal space .

cuticle clearly worn ; mandibles and claws dull . ovaries with ripe eggs ; corpora lutea distinctly light or dark brown ; lateral oviducts wide . testes large and cream - coloured ; accessory glands long and cream - coloured or light - brown , filling more than three - quarters of the abdominal space .

cuticle clearly worn ; mandibles and claws as a rule distinctly dull . ovaries compactly opaque and cream - coloured , without ripe eggs ; corpora lutea clearly visible and dark brown , often deposited above last developing oocytes ; lateral oviducts wide . testes medium - sized or relatively small ( regressed ) , opaque and cream - coloured or yellow ; accessory glands thin opaque and yellow or light - brown , occupying less than a third of the abdominal space .

cuticle very worn ; mandibles and claws blunt . ovaries compactly opaque and cream - coloured or light - brown , without ripe eggs ; corpora lutea clearly visible and dark brown , as a rule deposited under the developing oocytes ; lateral oviducts wide . testes medium - sized or relatively small ( regressed ) , opaque and yellow or brown ; accessory glands thin opaque and yellow , yellow - orange or brown , occupying less than a third of the abdominal space .

the separation between parental and ancestral generations was somewhat subjective and should be interpreted with caution . however , in most cases this separation was not required for the reasonable interpretation of demographic structures of the studied populations .

detection of the chronology of the maximum activity of the above - mentioned groups of specimens in the key stages of their life cycles as a result of feeding , reproduction or preparation for hibernation , forms the basis of our analysis . in such an approach , the quantitative recording of eggs , larvae , and pupae is not required . moreover , we can evaluate the demographic spectra of a local population from small numbers ( several dozen ) of individuals .

) . during this sequence , the abundance of species can be high or low . for example , in the reedbeds along the river khara in early spring , peaks of abundance in the populations of

) . in spite of this , both species are characterised by a complete demographic spectrum .

chronology of changes in periods of activity of individual ‘age’ groups , characterised by female gonad condition , in ‘spring’ ( a ) and ‘autumn’ ( b ) breeding carabid beetles ( t – teneral , im – immature , m – mature , sp – spent beetles ) .

seasonal dynamics of activity , as well as the age structure of the populations of pogonus transfuga ( a ) and brachinus hamatus ( b ) from reedbeds along the river khara , combined data for 2006 / 07 ( t – teneral , im – immature , m – mature , sp – spent beetles ; solid lines below graphs parental generation , dashed lines below graphs – new generation ; n ( ex . ) – number of specimens ; 1 , 2 , 3 – first , second and third ten - day periods per month , respectively ) .

) , the same order of physiological conditions of the adults is observed , but without an aestivation parapause . as in the previous case , the abundance of species can vary widely . for example , in the grass - forb steppe , the abundance of

) , yet the sex and age structure in the populations of both species was complete .

seasonal dynamics of activity , as well as the age structure of the populations of calathus ambiguus from grass - forb steppe with amygdalus nana ( a ) and pseudotaphoxenus rufitarsis major from sagebrush - grassland desert steppe on the northern slope of ulagan mountain ( b ) , in 2006 ( breaks in the periods of activity of immature specimens correspond to the time of aestivation parapause ; see figure 2 for further explanations ) .

importantly , in all these cases there are clear changes in successive waves of activity of different adult ‘age’ groups . it should be noted that in populations of many carabid species , the individuals of ancestral generations ( which live and breed during two or more years ) are often represented . in these cases the pattern of change in the physiological conditions can be blurred because separate successive waves of activity overlap each other .

thus , it is not abundance , but rather a regular change in the physiological condition that allows for a reconstruction of the life cycle at the local population scale , and this must be regarded as the criterion for the successful existence and breeding of a population in a particular habitat . species that meet these demands are considered ‘residents’ and their habitats ‘residential’ .

an incomplete demographic spectrum of a population means that the probability of a complete life cycle in a particular habitat is low to zero . such a situation is often followed by extremely high abundance levels . in reedbeds from the end of june until the end of july ,

was by far the most numerous carabid beetle collected , with abundance levels of 1753 , 7047 , 3770 and 2830 for successive ten - day periods . without information on the physiological conditions of individuals , this species may be considered dominant in this habitat . however , mature females were completely absent from the demographic spectra in this local population of

) . in these cases a reproductive phase in the demographic spectra of the local populations was absent .

yet the presence of mature specimens is not necessarily evidence of successful breeding . for example , in lakeside salt - marshes , the demographic spectrum of

was mainly represented by mature specimens . the abundance of spent beetles was very low , while teneral and immature beetles were completely absent (

) . the lack of young specimens in the demographic spectrum of this species provides evidence of immigration of mature beetles . species with incomplete demographic spectra are here considered ‘migrants’ and their habitats as ‘transit’ .

the spatial distribution of carabid species is determined by the availability both of habitats and landscape suitable for the complete realization of their life cycle . so the same habitat can be residential for one species and transit for another . among the examples discussed above , reedbed is a residential habitat for

) . at the same time , various habitats offer different living conditions to the same species . the sagebrush - grassland desert steppe on the northern slope of the ulagan mountain is a residential habitat for

in summary , the demographic structures of 66 carabid species found in the lake elton region were analyzed . the other 109 carabid species were represented by only one or two individuals ( appendix ) . considering the differences in abundance and demographic structure of the populations , three groups of carabidae of the studied habitats can be distinguished :

residents with their life cycles completed in a given habitat . in such species , migration forms only a facultative part of the life cycle . the catches of different species vary widely and sometimes differ by two orders of magnitude .

migrants that are characterised by relatively high numbers , yet rarely dominant , but with an incomplete demographic structure in particular habitats . because their reproduction and development are observed in different habitats , their roles in specific assemblages would be minor . migration forms both facultative and obligatory parts of their life cycles .

sporadic species with very low numbers , probably not associated with a particular habitat , neither during migration nor reproduction .

without question , residents interact both with their prey and with each other in a particular habitat . sporadic species are hardly important to a carabid community because of their low abundance levels . the role of migrants in the local carabid community remains unknown , with possible interactions between the migrants and residents . first , even very high numbers of migrants in relatively small - sized habitats do not reflect the condition of the populations of other carabid species . for example , in reedbeds of an area of 1 km2 , more than 13 000 specimens of

were trapped . this equates to a population density of about six individuals per square meter . this is a very high value . for example , the pest threshold of

, is two - three individuals per square meter . hence , if the captured specimens of

fed in this habitat and interacted with other species , we would expect changes in the demographic parameters of residents during this period . however , this is not the case , because the dynamics of the demographic structure in the populations of resident carabid beetles failed to change during this period (

) . second , relatively high numbers and species diversity levels of migrants were recorded at some seemingly unsuitable sites . these sites included the lakeside salt - marsh with high salt concentrations , poor vegetation and soil , as well as occasional floods . under these conditions , only some specialist

habitat preferences of individual species and the composition of carabid assemblages with the labile component dominant and subdominant species ( combined data for 2006 / 07 ) .

habitat preferences of individual species and the composition of carabid assemblages without the labile component resident species only ( combined data for 2006 / 07 ) .

“stable” and “labile” components can be recognized in ground - beetles communities ( makarov and matalin 2009 ) . the former includes species whose life cycles are realized in certain habitats ( residents ) , while the latter comprises species that are not capable of breeding in particular habitats ( migrants and sporadic species ) .

the ratio of stable to labile components in the studied habitats varied strongly and was not always in favour of residents . resident species comprised only 6–35 % of the species list and 15–90 % of total abundance . in zonal habitats , residents formed the dominant part of the assemblage . more than 65 % of total abundance and 15–35 % of total species diversity consisted of resident species . in azonal habitats the labile component prevailed . these species accounted for about 75 % of the fauna and about 80 % of total abundance (

) . only in zonal habitats did results from pitfall trapping adequately reflect the state of the carabid community while azonal and apparently disturbed habitats are only transit sites for many species of ground beetles .

species diversity and the share of labile / stable components in particular habitats in the lake elton region , combined data for 2006 / 07 ( black bars – labile component , white bars – stable component , line – number of species ; n ( sp . ) – number of species ) .

according to our data , the capture in a pitfall trap indicates only the fact that the beetle has moved across the trap area , but do not reflect true abundances . in some cases , errors occurring from direct interpretations of pitfall trapping data can be severe , and statistical techniques can not compensate for this . this is evident from cases in which high numbers of some carabid species are collected from seemingly unsuitable locations , for example from city dumps ( budilov 2002 ; romankina et al . 2007 ) , urban quarters ( khotuleva 1997 ; sharova and kiselev 1999 ) , places with strong oil or chemical pollution ( avtaeva 2006 ) and along roads ( noordijk et al . 2008 ; solodovnikov 2008 ) . the varying contribution of the labile component substantially distorts our knowledge of species diversity in carabid communities . taking into account the contribution of the labile component can change conclusions based on pitfall trapping data considerably .

firstly , criteria for determining the most abundant , or dominant species inevitably vary . the abundance of migrants in some cases is one order of magnitude higher than that of residents . therefore , estimating the faunistic or community features based solely on abundant or dominant species , fail to solve the problem and can even worsen the situation . in reedbeds , for example , 36 migrant species made up about 83 % of the total abundance . the complex of dominants in this community , as identified by the usual criterion ( abundance exceeding 5 % ) while discarding the demography of individual species , contains only two polyzonal migrants

numbers of the 10 most abundantly collected carabid species in reedbeds with regards to migrants ( a ) and residents only ( b ) . dominant species are in bold text , combined data for 2006 / 07 ; n ( ex . ) – number of specimens ( after makarov and matalin 2009 ) .

secondly , common information regarding the habitat preferences of particular species , as well as indicator species , is considerably altered . in our case , all studied habitats belong to two contrasting groups : dry desert steppes and riparian , more or less halophilic habitats . as such , variation in carabid populations is expected . when analyzing the habitat distribution of all dominants - subdominants , we find more or less eurytopic species inhabiting both zonal dry steppes on floodplain terraces and azonal alluvial salt - marshes . the grouping of dry steppes is very poor and contains one or two species which occur in one to three habitats , as a rule . in contrast , the inhabitants of salt - marshes are very diverse and peculiar . interestingly , the woodland in the ‘biological’ ravine supports not only a native carabid beetle community , but also a peculiar species ,

) . results from an analysis of the habitat distribution based solely on residents are distinctly different . only one species ,

, can be labelled eurytopic because it reproduces in nine of the ten studied habitats . the communities of carabid beetles on floodplain terraces and in flood - plains are clearly isolated from each other . each of them includes the main body of oligotopic species and a few stenotopic ones . finally , the riverine woodland does not have a native carabid community and can be considered a transit habitat for practically all carabid species (

because we have only very few examples that illustrate more or less close relations between ground beetles and their habitats , we are unable to assess the commonality of the situation described in the present study . however , it is conceivable that migrants in a carabid beetle community contribute to diversity estimates . based on results from this study , some preliminary conclusions can be made .

a study of the demographic structure of local populations and an assessment of the migratory / residential status of particular carabid species are possible ways to increase the reliability of pitfall trapping information .

up to 65–75 % of species diversity , both of particular habitats and the landscape as a whole , can comprise of non - residential carabid species , i . e . migrants .

results from pitfall traps adequately reflect the state of carabid communities only in zonal habitats . azonal and apparently disturbed habitats are only transit sites for many species of ground beetles .

knowledge concerning the composition of carabid communities , as well as study techniques , need to be significantly updated . no statistical method is capable of correcting the errors inferred from direct interpretations of pitfall trapping results .

we extend our thanks to all colleagues who assisted in our work , especially to the directors of the elton natural park , mrs yulia nekrutkina ( volgograd , russia ) and viktor gerdt ( elton , russia ) , as well as to dr . artem zaitsev ( moscow state pedagogical university , moscow , russia ) . we also want to thank professor sergei golovach ( institute for problems of ecology and evolution , moscow , russia ) for a critical review of the text , to dr . gabor lövei ( university of aarhus , denmark ) for fruitful discussions on the subject of the present paper , as well as to mr . stephen venn ( university of helsinki , finland ) and professor andrei alyokhin ( university of maine , orono , usa ) , who kindly checked the english . this study was financially supported by the russian foundation for basic research ( projects nos 09 - 04 - 01311 and 09 - 04 - 08112 ) .

classification of the carabid species from the elton lake region in migrant ( m ) , resident ( r ) and sporadic ( s ) species , based on their abundance and demographic spectrum in each habitat type ( combined data for 2006 / 07 ) .

zuphium ( s . str . ) olens olens ( p . rossi , 1790 )

arnoldi kv , sharova ikh , klyukanova gn , butrina nn . ( 1972 )

ground - beetles ( carabidae , coleoptera ) of the streletskaya steppe near kursk and their seasonal activity dynamics .

, moscow state pedagogicla institute publisher , moscow , 215–230 [ in russian ] .

the influence of different types of soil pollution on the population structure of ground beetles ( coleoptera , carabidae ) in grozny ( chechen republic ) .

phd thesis , moscow , russia : moscow state pedagogical university ; [ in russian ] .

life - cycles of ground - beetles ( coleoptera , carabidae ) in an agro - landscape in the south of the kuban - cisazov lowland .

phd thesis , moscow , russia : moscow state university ; [ in russian ] .

number of eggs in the ovaries of some carabidae ( coleoptera ) species in various pine stands on fresh coniferous forest habitats .

phenology of ground beetles and its ecological significance in some of the main habitat types of southern europe .

in : den boer pj , mossakowski d , luff ml , weber f . ( eds )

the population of ground beetles ( coleoptera , carabidae ) on areas with human impact ( on the example of ureiskiy crushed stone quarry ) .

buchholz s , jess a - m , hertenstein f , schirmel j . ( 2010 )

effect of the colour of pitfall traps on their capture efficiency of carabid beetles ( coleoptera : carabidae ) , spiders ( aranea ) and other arthropods .

changes in life - cycle strategies of carabus problematicus over a range of altitudes in northern england .

long adult life , low reproduction and competition in two sub - antarctic carabid beetles .

dispersal power and survival . carabids in a cultivated countryside ( with a mathematical appendix by j . reddingius ) .

miscellaneous papers 14 , h veenman & zonen bv press , wageningen , 190 pp .

flight muscles development and dispersal in the life cycle of carabid beetles : patterns and processes .

pitfall trapping within enclosures : a method for estimating the relationship between the abundances of coexisting carabid species ( coleoptera : carabidae ) .

a simple device and technique for quantitative sampling of riparian beetle populations with some carabid and staphylinid abundance estimates in different riparian habitats ( coleotera ) .

the natural histories of four species of calathus ( coleoptera , carabidae ) living on sand dunes in anglesey , north wales .

to the method of quantitative evaluation of ground - beetles ( coleoptera , carabidae ) .

community organization in ground beetles ( coleoptera , carabidae ) of the forests of the east european ( russian ) plain .

sampling epigeal arthropods : an evaluation of fenced pitfall traps using mark - release - recapture and comparison to unfenced pitfall traps in arable crops .

the life cycles and age of carabus glabratus paykull and c . problematicus herbst ( col . : carabidae ) on moorland in northern england .

the structure of populations of ground beetles ( coleoptera , carabidae ) in urbanized landscapes of north of meschera lowland .

koivula m , kotze dj , hiisivuori l , rita h . ( 2003 )

the influence of the manner of pitfall traps setting in forest habitat on their catchability .

to the question of the use of soil traps for the study of the distribution and interactions of entomofauna elements on the soil surface .

about updating the quantity estimation applying the methods of exhaustive catches using of pitfall traps .

nĕkteré čisté chemické látky jako návnada v zemních pastech při studio střevlíkovitých ( carabidae ) .

ground - beetle comunities in the lake elton region , southern rusia : a case study of a local fauna ( coleoptera : carabidae ) . in : golovatch si , makarova ol , babenko ab , penev ld ( eds ) species and communities in extrem environments . festschrift towards the 75th anniversary and a laudatio in honour of academician yuri ivanovich chernov .

variability of seasonal dynamics of activity in ground beetle pterostichus melanarius ill . ( coleoptera , carabidae ) in different forest types .

, institute of biology of latvian academy of science publisher , riga , 55–56 [ in russian ] .

the phenology and population structure of loricera pilicornis ( f . ) ( coleoptera , carabidae ) in agrocoenosis .

in : desender k , dufrêne m , loreau m , luff ml , maelfait j - p . ( eds )

specific features of life cycle of pseudoophonus ( s . str . ) rufipes deg . ( coleoptera , carabidae ) in southwest moldova .

variations in flight ability with sex and age in ground beetles ( coleoptera , carabidae ) of south - western moldova .

diversity of ground beetles ( coleoptera : carabidae ) and spiders ( araneae ) in roadside verges with grey hair - grass vegetation .

synekologická studie sezónního výskytu střevlíkovitých na řepných polich hané ( col . carabidae ) .

pit - fall catches of surface - active arthropods in high mountain habitats at finse , south norway . iv . coleoptera .

differential effects of formaldehyde concentration and detergent on the catching efficiency of surface active arthropods by pitfall traps .

estimating the density of ground - dwelling arthropods with pitfall traps using a nested - cross array .

determining numbers of active carabid beetles per unit area from pitfall - trap data .

ecological and fauna structure of the population of coleoptera , carabidae in the forest belt in the city dump area .

dynamics of population structure of ground beetles ( coleoptera , carabidae ) in urbanized landscapes of saransk .

mordovian state pedagogical university publisher , saransk , 213 pp [ in rusian ] .

ground beetles ( coleoptera , carabidae ) of byelorussian lakeland with catalogue of species of byelorussia and adjacent lands .

uo p . m . masherov vgu , vitebsk , 325 pp [ in russian ] .

carabid beetles in their environments . a study on habitat selection by adaptations in physiology and behaviour .

springer - verlag , berlin – heidelberg - new - york , xvii + 369 pp .

zur ökologie , soziologie und phänologie der laufkäfer ( coleoptera - carabidae ) des grünlandes im süden der ddr . hercynia n . f .

seasonal dynamics of activity and population structure of pterostichus niger schaller in different habitats .

the reproductive cycle and age composition of a population of pteropstichus oblongopunctatus f . in the netherlands ( coleoptera : carabidae ) .

the significance of flight activity in the life cycle of amara plebeja gyll . ( coleoptera , carabidae ) .

in : den boer pj , thiele hu , weber f . ( eds )

trapping efficiency of carabid beetles in glass and plastic pitfall traps containing different solutions .

the influence of different age classes on the seasonal activity and reproduction of four medium - sized carabid species inhabiting cereal fields .

work tt , buddle cm , korinus lm , spence jr . ( 2002 )

pitfall trap size and capture of three taxa of litter - dwelling arthropods : implications for biodiversity studies .

we gratefully acknowledge financial support from the county administrative board in skåne and the swedish research council .

bascompte j , jordano p ( 2007 ) plant–animal mutualistic networks : the architecture of biodiversity . annu rev ecol syst 38 : 567–593

bertoncelj i , dolman pa ( 2013 ) conservation potential for heathland carabid beetle fauna of linear trackways within a plantation forest . insect conserv divers 6 : 300–308

cizek l , hauck d , pokluda p ( 2012 ) contrasting needs of grassland dwellers : habitat preferences of endangered steppe beetles ( coleoptera ) . j insect cons 16 : 281–293

dahlström a , cousins sao , eriksson o ( 2006 ) the history ( 1620–2003 ) of land use , people and livestock , and the relationship to present plant species diversity in a rural landscape in sweden . environ hist camb 12 : 191–212

desender k , bosmans r ( 1998 ) ground beetles ( coleoptera , carabidae ) on set - aside fields in the campine region and their importance for nature conservation in flanders ( belgium ) . biodiv cons 7 : 1485–1493

gärdenfors u ( ed ) ( 2010 ) the 2010 red list of swedish species . artdatabanken , slu , uppsala

gazenbeek a ( 2005 ) life , natura 2000 and the military . life focus , european commission

hansen v ( 1927 ) biller vii , bladbiller og bønnebiller ( chrysomelidae & laridae ) . danmarks fauna 31 , copenhagen , denmark

hobbs rj , huenneke lf ( 1992 ) disturbance diversity and invasion implications for conservation . cons biol 6 : 324–337

irmler u , hoernes u ( 2003 ) assignment and evaluation of ground beetle ( coleoptera : carabidae ) assemblages to sites on different scales in a grassland landscape . biodiv cons 12 : 1405–1419

jentsch a , friedrich s , steinlein t , beyschlag w , nezadal w ( 2009 ) assessing conservation action for substitution of missing dynamics on former military training areas in central europe . rest ecol 17 : 107–116

johansson lj , hall k , prentice h , ihse m , reitalu t , sykes m , kindström m ( 2008 ) semi - natural grassland continuity , long - term land - use changes and plant species richness in an agricultural landscape on öland , sweden . landsc urban plan 84 : 200–211

knisley cb ( 2011 ) anthropogenic disturbances and rare tiger beetle habitats : benefits , risks , and implications for conservation . terr arthropod rev 4 : 41–61

. final report to u . s . fish and wildlife service , ventura , california , usa . pp 34

landin bo ( 1957 ) svensk insektsfauna 9 : skalbaggar , bladhorningar : fam . scarabaeidae . entomologiska föreningen , stockholm , sweden

lin y - c , james r , dolman pm ( 2007 ) conservation of heathland ground beetles ( coleoptera , carabidae ) : the value of lowland coniferous plantations . biodiv cons 16 : 1337–1358

lindroth ch ( 1961 ) svensk insektfauna . 9 , skalbaggar , coleopetra , sandjägare och jordlöpare , fam . carabidae , 2nd ed . entomologiska föreningen , stockholm , sweden