the spanish imperial eagle ( aquila adalberti ) , also known as the iberian imperial eagle , spanish eagle , or adalbert 's eagle , is a threatened species of eagle native to the iberian peninsula .

the binomial commemorates prince adalbert of bavaria .

formerly , the spanish imperial eagle was considered to be a subspecies of the eastern imperial eagle , but is now widely recognised as a separate species due to differences in morphology , ecology , and molecular characteristics .

spanish imperial eagle

the spanish imperial eagle , also known as iberian imperial eagle or adalbert ' s eagle , ( aquila adalberti ) is closely related to the eastern imperial eagle ( aquila heliaca ) .
hello we have checked it and it opens a spanish imperial eagle video . arnau
long - term changes in nest defense intensity of the spanish imperial eagle aquila adalberti .
. today treated as a seperate species . forms a superspecies with the eastern imperial eagle . the closest relatives of the spanish and eastern imperial eagle are probably the steppe eagle
spanish imperial eagle ( aquila adalberti ) . presumed 5cy . 14 - 6 - 2011 . valladolid . photographer : juan sagardia
description : spanish imperial eagle is classified as endangered species . it has restricted habitat and numerous threats make of this eagle the rarest bird of prey in the world .
it closely resembles the spanish imperial eagle , but has far less white to the
shoulder
and it is slightly larger .
ferrer m , hiraldo f ( 1991 ) evaluation of management techniques for the spanish imperial eagle . wildlife soc b 19 : 436–442 .
ferrer m , hiraldo f ( 1992 ) man - induced sex - biased mortality in the spanish imperial eagle . biol conserv 60 : 57–60 .
[ grin 2009 ] global raptor information network . 2009 . species account : spanish imperial eagle aquila adalberti . downloaded from urltoken on 27 feb . 2009
meyburg , b . u . ( 1975 ) on the biology of the spanish imperial eagle . publicado en ardeda , 21 : 1 - 10 .
bird life international ( 2005 ) imperial eagle - birdlife species fact sheet . birdlife international , cambridge
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spanish imperial eagle ( aquila adalberti ) . central spain , autumn 2011 . photographer : juan pablo fuentes serrano a juvenile sie with notorious streaking on the upper breast .
spanish imperial eagle ( aquila adalberti ) . valladolid , june 2011 . photographer : juan sagardía a young adult female spanish imperial eagle . a stunning bird , for sure , particularly if seen this well ! adults are relatively straightforward to identify , but some juveniles , and especially immature birds , can be very difficult to tell apart from easterns .
the spanish imperial eagle has been discovered in 1860 by the german naturalist dr . reinaldo brehm ( renthendorf , 1830 – barcelona , 1891 ) . the first research about the spanish imperial eagle was financed by the prince albert of bavaria , great friend of the naturalist . that explains why dr . reinaldo brehm gave the name of prince’s albert eagle ( aquila adalberti ) as a tribute to his friend .
habitat : the eagle in its favourite habitat the spanish imperial eagle lives in cork - oak plantations , as in plains or hills . it can be seen in marshes , along the streams and in steppes with scattered isolated trees .
eastern imperial eagle ( aquila heliaca ) . juvenile . january 2011 . oman . photographer : daniel lópez - velasco
eastern imperial eagle ( aquila heliaca ) . juvenile . january 2011 . oman . photographer : daniel lópez - velasco
eastern imperial eagle ( aquila heliaca ) . 5cy . january 2011 . oman . photographer : daniel lópez - velasco
spanish imperial eagle ( aquila adalberti ) . juvenile . southern spain . december 2011 . photographer : josé arcadio . another example of an already bleached , pale sandy buff juvenile sie .
the spanish imperial eagle is resident in its range . it lives in territories of variable sizes , and according to the food resources , several pairs may nest in the same area .
gonzález lm , hiraldo f , delibes m , calderón j ( 1989a ) zoogeographic support to consider the spanish imperial eagle as a distinct species . bull br ornithol club 109 : 86–93
imperial eagle sp , probably 4cy eastern . sweden , july 2012 . photographer : lasse olsson . photo taken from surfbirds gallery
spanish imperial eagle ( aquila adalberti ) . presumed 5cy . photographer : jose luis rodriguez . the first white feathers on the leading edge have already appeared , which helps separate it from eastern .
land stewardship net created by seo / birdlife and the preceding flying high programme for the conservation of spanish imperial eagle spanned 2006 - 2012 ( s . cabezas - díaz , j . c . atienza
ferrer , m . and hiraldo , f . ( 1992 ) man - induced sex - biased mortality in the spanish imperial eagle . biological conservation , 60 ( 1 ) : 57 - 60 .
with this species , the name imperial eagle being used in both circumstances . however , the two are now regarded as separate species
i was recently asked about an interesting looking immature ( presumably a 4cy ) imperial eagle sp . seen in sweden , and i thought it was about time to share some photos , and comments , on variation within juvenile and immature spanish imperial eagles ( aquila adalberti ) .
the spanish imperial eagle relies mainly on rabbits as prey , and following drops in rabbit abundance due to shooting and disease , food supplies have been limited causing reduced breeding success ( 2 ) ( 11 ) .
gonzález lm , margalida a , mañosa s , sánchez r , oria r , et al . ( 2007 ) causes and spatio - temporal variations of non - natural mortality in the vulnerable spanish imperial eagle (
voice : sounds by xeno - canto the spanish imperial eagle utters series of bark - like calls “krao - krao” . this eagle calls mainly during the breeding season , and to defend its hunting area against other birds of prey . it is rather silent outside of breeding period .
feeding almost exclusively on rabbits , the spanish imperial eagle’s population density , range and reproductive performance are influenced by the abundance and distribution of prey in the area ( 2 ) . this eagle species is also known to eat greylag geese in southern spain in the winter ( 9 ) .
spanish imperial eagle ( aquila adalberti ) . juvenile . photo : jose luis rodriguez . a fairly pale bird , with obvious streaking on the upper breast . note however little streaking on the underwing coverts , unlike heliaca .
his preferred perch is the cliff top , among the vultures , often above them because the spanish imperial eagle does not like them . sometimes they may rest together , relatively close to each other on the rocks . but today , the eagle is upset and some aggressive chases can be observed .
spanish imperial eagle ( aquila adalberti ) . fresh juvenile . valladolid , july 2012 . photographer : juan sagardía . a typical juvenile sie , showing uniformly warm rufous colouration on the underparts , with very little streaking on the chest .
spanish imperial eagle ( aquila adalberti ) . juveniles . photographer : jose luis rodriguez . photo taken from jose luis rodriguez website . note the heavy upper breast streaking of the juvenile on the left , similar to that of eastern .
gonzáles , l . m . y margalida , a . ( 2008 ) . biología de la conservación del águila imperial ibérica ( aguila adalberti ) . conservation biology of the spanish imperial eagle ( aquila adalberti ) . organismo autónomo parques nacionales . ministerio de medio ambiente y medio marion y rural . madrid .
in europe , the eastern imperial eagle is threatened with extinction . it has nearly vanished from many areas of its former range , e . g .
the tracking of a young spanish imperial eagle ( aquila adalberti ) released with a transmitter in southern spain has resulted in an unexpected discovery . it was found dead in october 2015 under an electricity pylon near guelmim , in southern morocco , along with the remains of two other spanish imperial eagles , a golden eagle ( a . chrysaetos ) and a bonelli ' s eagle ( a . fasciata ) # r . up to four more bonelli ' s eagles were found dead along the same electricity line during a second visit there in early november .
spanish imperial eagle ( aquila adalberti ) . central spain , autumn 2011 . photographer : juan pablo fuentes serrano note the overall pale and cold background colour of this bird , and also the striking pale tips to all the wing coverts .
some examples of different age classes of spanish imperial below , showing variation . there are few published pics of these plumages , so they might be of interest .
mostly found in mediterranean forests but also in marshes like in the doñana national park . the spanish imperial eagle can also be found in
dehesas
, a typical wooded pastureland found in spain . enough prey , especially rabbits are important .
there are many eastern imperial eagle nests in the bulgaria / turkey section of the european green belt ( the uncultivated belt along the former iron curtain . )
the monarchy of austria - hungary once chose the imperial eagle to be its heraldic animal , but this did not help this bird . the eagle ' s preferred habitat is open country with small woods ; unlike many other species of eagle , it does not generally live in mountains , large forests or treeless steppes .
flight : the spanish imperial eagle performs beautiful flight displays . it flies low from the ground for hunting , and soars in order to detect preys . the broad wings allow the bird to soar and glide easily , with few wing - beats .
but i will try to offer some visual information on variation and id pitfalls of juvenile and immature spanish imperial eagles , which can perhaps be useful for future records .
left – spanish imperial eagle ( aquila adalberti ) . juvenile . avila . january 2010 . photographer : juan sagardia . right – eastern imperial eagle ( aquila heliaca ) . juvenile . oman . january 2011 . photographer : daniel lópez velasco note the similarities between these 2 birds , although the white tips of the upperwing gcs are broader in the eastern . a view of the underparts should id both , though .
[ gonzáles & margalida 2008 ] gonzáles , l . m . & margalida , a . ( 2008 ) . biología de la conservación del águila imperial ibérica ( aguila adalberti ) . conservation biology of the spanish imperial eagle ( aquila adalberti ) . organismo autónomo parques nacionales . ministerio de medio ambiente y medio marion y rural . madrid .
aquila heliaca , commonly called the imperial eagle , is a large bird of prey that are migratory and winter in northeastern africa , southern and eastern asia .
citation : lópez - lópez p , ferrer m , madero a , casado e , mcgrady m ( 2011 ) solving man - induced large - scale conservation problems : the spanish imperial eagle and power lines . plos one 6 ( 3 ) : e17196 . urltoken
2016 ) . several spanish imperial eagles have been found shot in the past decade , in both spain and portugal ( s . cabezas - díaz , j . c . atienza
imperial eagles are beneficial to farmers and ranchers because they feed on rabbits and other small mammals . by eating these animals , the imperial eagle reduces their numbers , and thus reduces the damage they can do . this eagle also feeds on venomous reptiles , many of which have been known to cause serious harm to humans . ( channing 2000 )
adult male is very similar to the eastern imperial eagle ( aquila heliaca ) , but the spanish imperial eagle shows conspicuous white leading edges of the wings on the lesser coverts . the plumage is reddish - brown overall , without any streaks , but we can see several irregular white patches on scapulars . the tail is greyish - brown with broad darker terminal band . on the brown underparts , underwing shows black wing coverts and paler flight feathers .
the spanish imperial eagle is soaring among several eurasian griffon vultures ( gyps fulvus ) , when suddenly , he dives onto the one below . he pursues the large raptor during one or two minutes , and performs some threat displays . he holds his legs towards the vulture with outstretched talons . and finally , the eagle turns and reaches the cliff top , as usual .
this large eagle is a gift for all the bird lovers watching for his silhouette soaring over the rocky cliffs of spanish mountains . the spanish imperial eagle appears as a survivor after numerous years of persecutions , poisoning , habitat destruction , power lines and reduction of numbers of his preferred preys , rabbits . today , thanks to several conservation and protection programs , this species is slowly recovering better numbers . but this eagle is still evaluated as vulnerable , with small but increasing populations of about 396 pairs , scattered throughout spain and 11 pairs in portugal ( census 2013 ) .
the spanish imperial eagle could be very similar to the golden eagle ( aquila chrysateos ) , but the white shoulder’s feathers allow a good identification . he is living only in spain and portugal , but formerly , his range probably included north africa and southern france . this species is sedentary , but the young birds may wander over some distance , sometimes to the former range .
martínez - cruz b , godoy ja ( 2007 ) genetic evidence for a recent divergence and subsequent gene flow between spanish and eastern imperial eagles . bmc evol biol 7 : 170 . doi :
the imperial eagle is legally protected in azerbaijan , bulgaria , croatia , georgia , greece , hungary , romania , slovakia , turkey and ukraine . the eastern imperial eagle working group was established in 1990 and the european action plan was published in 1996 . crucial steps in the conservation of this eagle are the improvement of forestry practices , the maintenance of large trees and the prevention of mortality via nest robbing , illegal trade , poisoning and power lines ( 6 ) .
2011 ) , with more than 50 eastern imperial eagles poisoned in hungary ( m . horvth
the spanish imperial eagle is smaller , 2 . 5–3 . 5 kg ( 5 . 5–7 . 7 lbs ) and 75–84 cm ( 30–33 in ) in length , and darker than its eastern cousin , and is a resident species ( a . heliaca migrates to the southeast during winter ) .
pain , d . j . , meharg , a . a . , ferrer , m . , taggart , m . and penteriani , v . ( 2005 ) lead concentrations in bones and feathers of the globally threatened spanish imperial eagle . biological conservation , 121 ( 4 ) : 603 - 610 .
the spanish imperial eagle is legally protected in spain , and 62 % of the breeding population occurs in 20 protected areas . a european action plan was published in 1996 , and national and regional governments have worked to implement a coordinated conservation plan . a reintroduction plan has seen 13 young released successfully ( 2 ) .
eastern imperial eagle ( aquila heliaca ) . 3cy . january 2011 . oman . photographer : daniel lopez - velasco . for comparison , a 3cy eastern from january . outer primaries still juvenile , thus very worn and ragged . note the heavier streaking on the breast and underwing coverts compared to the above 3cy spanish .
range : the spanish imperial eagle is resident in spain , mainly in extremadura and andalucía , in mountains such as sierras of guadarrama and gredos , along the tajo and tietar rivers , and in marshes in the guadalquivir area . this species is extinct in morocco , but it is slowly increasing in portugal after extinction .
spanish imperial eagle ( aquila adalberti ) . 3cy . 4 - 6 - 2011 . aragon . photographer : alberto bueno . another 3cy , more advanced in terms of primary moult , with no juvenile primaries left , than the above bird . note , again , obvious streaking on the chest , and relatively cold looking underparts and underwing coverts . see , however , the fairly warm rufous head of this individual , typical of spanish .
spanish imperial eagle ( aquila adalberti ) . presumed 5cy . 25 - 5 - 2011 . central spain . photographer : antonio cavadas . if no pure white feathers on the leading edge of the wing can be seen ( and beware of light effects ) , then the id of birds like this one can be very tricky .
we saw the spanish imperial eagle several times in extremadura , as well in the steppes searching for rabbits , as more often in monfragüe park . the cliffs are isolated and protected from the road by the meanders of the tagus river . numerous eurasian griffon vultures inhabit these rocky faces where their white downy chicks hatch every year .
although rare , imperial eagles have been known to attack humans who come too close to their nests . they may dive at a person who gets too close and strike at them with their talons . ( eagle 1988 )
and below , a 5cy eastern , from oman . note how scarily similar the bird is to the above 2 spanish .
unesa ( 2005 ) el sector eléctrico a través de unesa ( 1944 - 2004 ) . spanish association of electricity industry .
our results demonstrate that solving bird electrocution is an affordable problem if political interest is shown and financial investment is made . the combination of an adequate spatial planning with a sustainable development of human infrastructures will contribute positively to the conservation of the spanish imperial eagle and may underpin population growth and range expansion , with positive side effects on other endangered species .
in a study that examined the death of 267 spanish imperial eagles between 1989 and 2004 , 91 . 7 % of the causes of mortality ( where the cause of death could be determined ) were of human origin . electrocution and poisoning were by far the most common causes of death . 115 spanish imperial eagles were electrocuted and 74 were poisoned . other causes of death were shooting ( 16 cases ) and disease ( 13 cases ) [ gonzáles & margalida 2008 ] .
native forest has been lost to the forestry industry as trees are felled and replaced with introduced species , depriving the imperial eagle of nesting and feeding sites . nest robbing by humans is common , and trade in this species remains a problem . additionally , imperial eagles are shot and poisoned , and are electrocuted on power lines . shortages of prey species , particularly the suslik ( spermophilus citellus ) , have also contributed to the decline of this eagle ( 4 ) .
bisson , i . a . , ferrer , m . and bird , d . m . ( 2002 ) factors influencing nest - site selection by spanish imperial eagles . journal of field ornithology , 73 ( 3 ) : 298 - 302 .
margalida a , gonzález lm , sánchez r , oria j , prada l , caldera j , aranda a , molina ji ( 2007 ) a long - term scale study of the breeding biology of spanish imperial eagles . j ornithol 148 : 309–322
most fresh juvenile spanish imperial eagles show uniformly rich warm rofous , almost unstreaked , underparts . upperparts usually lack many distinct pale spots or streaks . thus , they are very different to juvenile eastern imperials . see 2 photos below of typical juvenile adalberti .
spanish imperial eagle ( aquila adalberti ) . 3cy . 31 - 7 - 2011 . navarra . photographer : josé ardaiz . note 2 moult waves on the primaries of this 3cy , and the 2 outermost juvenile primaries . the bird shows notorious dark and pale streaking on the upper breast , but the head , part of the underparts and underwing coverts are typically warm rufous .
through a recent observation , we saw again an altercation in the sky between the spanish imperial eagle and the eurasian griffon vulture . the eagle is perched on its usual dead branch in the vicinity of the nest - site used the last season . the raptor is within its territory . but this area is also occupied by a large colony of vultures . they must share the territory , and usually , they are able to live close to each other . their hunting behaviour is not the same . no competition for food !
horváth , m et al . ( 2010 ) .
spatial variation in prey composition and its possible effect on reproductive success in an expanding eastern imperial eagle ( aquila heliaca ) population
( pdf ) . acta zoologica academiae scientiarum hungaricae 56 : 187–200 .
since 1974 , a total of 158 spanish imperial eagles have been recorded dead in andalucía , 101 of them ( 63 . 92 % ) inside doñana national park . electrocution was the most frequent cause of death accounting for 39 . 87 % of the total mortality events . since 1974 when the first death by electrocution was recorded , 37 spanish imperial eagles have been found electrocuted in doñana ( 36 . 63 % of the mortality cases ) and 26 cases in andalucía ( 41 . 94 % of the records ) ( figure 2 ) .
[ gonzáles et al . 1989 ] gonzáles , l . m . & heredia , borja & gonzáles jose l . & alonso , juan c . ( 1989 ) . juvenile dispersal of spanish imperial eagles . journal of field ornithology 60 ( 3 ) 369 - 379 .
but one vulture joins him and perches close to the eagle . one minute later , a second vulture arrives , but the eagle is still upset and performs again threat displays and calls . then , he takes off and flies away , leaving the two perched vultures .
[ seo 2011 ] el marm , las ccaa y portugal analizan los avances conseguidos en la conservación del águila imperial ibérica . downloaded from
diet : the spanish imperial eagle feeds mainly on rabbits and hares , but it also captures birds such as greylag goose , coots , ducks , pigeons and magpies . it catches them often on the ground diving from a perch , or sometimes by soaring , taking the bird perched in open , at the tip of a branch . it also consumes reptiles and insects taken by walking on the ground , some fish and carrion .
the spanish imperial eagle is classified as vulnerable ( vu ) on the iucn red list ( 1 ) and is listed on appendices i and ii of both cites ( 3 ) and the convention on migratory species ( cms or bonn convention ) ( 4 ) . it is also listed on appendices ii and iii of the berne convention of european wildlife and natural habitats ( 5 ) and annex i of the ec birds directive ( 6 ) .
the imperial eagle is found from southern europe to southern russia , as well as northwest india and central siberia . in winter it migrates to the middle east , east africa as far south as tanzania , the arabian peninsula , the indian subcontinent and south and east asia ( 2 ) .
spanish imperial eagle ( aquila adalberti ) . presumed 4cy . central spain . photographer : jose luis rodriguez . photo taken from jose luis rodriguez website . this presumed 4cy bird ( not possible to confirm the ageing on this photo alone ) , still , apparently , with no white on the leading edge of the wing , is probably impossible , on actual knowledge , to separate with confidence from the corresponding plumage of eastern . id mainly based on location .
man - induced mortality of birds caused by electrocution with poorly - designed pylons and power lines has been reported to be an important mortality factor that could become a major cause of population decline of one of the world rarest raptors , the spanish imperial eagle ( aquila adalberti ) . consequently it has resulted in an increasing awareness of this problem amongst land managers and the public at large , as well as increased research into the distribution of electrocution events and likely mitigation measures .
the eagle is flying too in the same area , circling quietly in thermals . sometimes , one or two vultures are passing by… and suddenly , vulture and eagle are facing each other . the large vulture almost stands up with the wings widely open , while the eagle is flying on its back with legs and talons hold forwards in front of the vulture . during less than one second , both raptors are facing each other with the wings widely spread while flying .
hiraldo f , delibes m , calderón j ( 1976 ) sobre el status taxonómico del águila imperial ibérica . doñana acta vertebr 3 : 171 - 182
illegal poisoning must be stopped . public education and better law enforcement are necessary . powerlines should all be made safe for birds . not only in the breeding areas , but also in the dispersal areas where many juvenile and immature birds can be found after they leave their natal area . in spain , several organisations like wwf or seo / birdlife spain are working on conservation programs for the spanish imperial eagle . the national and some regional governments are also working to protect the species .
the spanish imperial eagle breeds in central and southwest spain , with populations found in the sierras of guadarrama and gredos , the plains of the tajo and tiétar rivers , montes de toledo , the central hills of extremadura , the alcudia valley , sorena morena and the guadalquivir marshes . it previously bred in portugal , algeria and morocco , but is now only found there as a vagrant ( 2 ) . it is also found in france and gibraltar as a vagrant ( 1 ) .
meyburg , b . u . , kirwan , g . m . & de juana , e . ( 2018 ) . spanish imperial eagle ( aquila adalberti ) . 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 ) .
meyburg , b . u . ( 1994 ) . eastern imperial eagle ( aquila heliaca ) . pp . 194 - 195 in : del hoyo , elliott & sargatal . eds . ( 1994 ) . handbook of the birds of the world , vol . 2 . isbn 84 - 87334 - 15 - 6
on of the most serious threats is electrocution . in 2008 , 33 electrocuted spanish imperial eagles were found [ gonzáles 2009 ] . despite work to make powerlines safe in many areas , there are still many eagles dying because of electrocution . the expanding population and dispersing juveniles visit new areas where powerlines have not yet been made safe .
castaño lópez , juan pablo ( 2005 ) . el águila imperial ibérica en castilla la mancha - estatus , ecología y conservación . graphitis impresores . madrid .
behaviour : the spanish imperial eagle is often seen perched in treetops or electric pylons or towers during the day . it hunts in open areas , flying and soaring relatively low from the ground . when a prey is detected , it dives and captures it with its strong talons . its preferred preys are rabbits and hares , but it also takes other mammals , birds , and reptiles such as lizards . sometimes , it may feed on fish and insects , and also carrion . reptiles and insects are taken while walking on the ground . birds are mainly taken on the ground or when perched in open place , at the tip of a branch . it also catches young birds at nest . the spanish imperial eagle cannot pursue birds in flight , due to its great size , making impossible to enter the tree branches or bushes where the scared birds hide . it is able to catch preys from 500 to 2500 grams . it hunts mainly in the morning and at the end of the afternoon until dusk .
a second video shows the eagle returning to his nest , but he also takes a long branch with leaves from the rim and flies away . this behaviour probably indicates a change of nest . the eagle prefers to build a new nest in order to protect his young against the vultures . these facts could explain the threat behaviour recently observed .
i am , by no means , an expert on the 2 imperial eagles , although i have seen quite a few easterns , mainly in oman , and of course plenty of spanish over here in spain ( mainly adults and juveniles of the latter though ; it’s not easy at all to see immature birds , as they wander around a lot ) .
eagles according to cytochrome b nucleotide sequences . in : meyburg bu , chancellor rd ( eds ) eagle studies . world working group on birds of prey , london , pp 1–15
from above , certain birds can be almost identical . see the 2 shots below , both taken in january , showing a juvenile spanish on the left , and a juvenile eastern on the right .
[ gonzáles 2009 ] . gonzáales , luis mariano ( 2009 ) . más de 250 parejas : continúa la recuperación del águila imperial . quercus . cuaderno ( issue ) 279 .
padilla , j . a . ; martinez - trancón , m . ; rabasco , a . ; fernández - garcía , j . l . ( 1999 ) .
the karyotype of the iberian imperial eagle ( aquila adalberti ) analyzed by classical and dna replication banding
. cytogenetics and cell genetics 84 : 61–66 . doi : 10 . 1159 / 000015216 .
the imperial eagle tends to live in forests , where it will build a huge , conspicuous nest in an isolated tall tree . old forests , and the forests in mountains , hills , and along rivers are all common habitats . it has , however , also been known to live in steppes , open landscapes , agricultural areas , and even semideserts . imperial eagles hunt in open fields or wetlands . during migration , a variety of habitats are used , though they seem to prefer to winter in wetlands . ( heredia 1996 , peterson 1993 )
the importance of human - induced mortality should also be considered in the context of population dynamics . demographic variables are not independent of one another when endangered small populations are analyzed , as the case of spanish imperial eagles [ 29 ] , [ 30 ] . in long - lived spatially structured populations , pre - adult and adult mortality play a key role in influencing population persistence [ 31 ] – [ 33 ] . therefore conservation measures should be focused on mitigating detrimental effects on these parameters [ 10 ] , [ 30 ] . in the case of spanish imperial eagles , adult survival is higher than pre - adult and floater survival and hence small electrocution rate of adult eagles leads to more severe consequences on population dynamics than the death of non - breeding individuals [ 34 ] .
spanish imperial eagle ( aquila adalberti ) . juvenile . ávila , february 2012 . photographer : juan sagardía . another typical juvenile , with no streaking on the chest , and plain rufous underparts and underwing coverts . the photo was taken in a mountain area of central spain , certainly less sunny in the winter than southern spain or morocco . therefore , the plumage state of the bird is still good , not worn or bleached at all being february . birds wintering in more southerly , sunnier , areas are more prone to bleaching , and by mid winter can be already very worn and whitish .
protection / threats / status : the spanish imperial eagle is an endangered species , living in restricted areas in spain . the species is threatened by poisoning , habitat loss , human disturbances , electrocution , trapping in rabbit - traps and the decline of rabbit’s populations . the species is slightly colonizing portugal again with 11 pairs . in spain , there are currently 396 pairs ( census 2013 ) . but it is still listed as vulnerable . the most important threat is the increasing use of poisoning . however , the small numbers of pairs are slowly increasing , thanks to important protection measures and increase of protected areas .
originally a lowland species , the imperial eagle has been pushed to higher elevations by habitat loss and hunting . in central and eastern europe it is found in forests up to 1 , 000 metres , as well as steppe and agricultural areas with large trees . in the caucasus , it is still found in lowland and riverine forests and semi - deserts . it winters in wetlands ( 5 ) .
the time for them to regain their balance , they continue their way until the next encounter in the sky . the vulture returns to the cliff and the eagle to its perch… see you soon ! ! !
this type of behaviour occurs regularly and lasts a few seconds . this is not a direct attack , but probably an intimidation attempt in order to remind the vulture to keep its distance from the eagle’s nest .
schaub m , aebischer a , gimenez o , berger s , arlettaz r ( 2010 ) massive immigration balances high anthropogenic mortality in a stable eagle owl population : lessons for conservation . biol conserv 143 : 1911–1918 .
funding : p . lópez - lópez was supported by fpu grant of the spanish ministerio de educación ( reference ap2005 - 0874 ) and wrote this paper during a pre - doctoral stay at doñana biological station ( csic ) in sevilla , spain . the funders of this study ( spanish ministry of education , natural research ltd . and junta de andalucía ) had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
reproduction : the spanish imperial eagle nests at tree top or sometimes on electric tower or pylon . the nest may be very large or smaller , and it is reused year after year . in average , this nest is about 150 cm of diameter and 60 cm of depth . but year after year , new materials are added , and these structures become huge , up to 2 metres of diameter , and can be seen at great distance . the nest is made with sticks , and the interior is lined with dried grasses and green twigs . nest materials are brought by both adults , but the female builds the nest . these works often last about three weeks .
soutullo a , lópez - lópez p , urios v ( 2008 ) incorporating spatial structure and stochasticity in endangered bonelli ' s eagle ' s population models : implications for conservation and management . biological conserv 141 : 1013–1020 .
another interesting observation show the large eagle harassed by the two iberian magpies ( cyanopica cooki ) . these birds also nest in cork - oaks , and the presence of the raptor close to their nest is a threat for the young magpies . so , they fly around him , very close , and dive bomb onto the eagle which remains quietly perched on his rock . but after several attacks , he takes off , pursued by the angry corvids !
cites appendix i and ii . cms appendix i and ii . it is legally protected in armenia , azerbaijan , bulgaria , croatia , georgia , greece , hungary , romania , slovakia , turkey and ukraine . the eastern imperial eagle working group was established in 1990 . a european action plan was published in 1996 and its implementation reviewed in 2010 ( barov and derh 2011 ) . regional action plans have been published for the balkan peninsula ( stoychev
future research should be focused on the development of new high - efficiency , low - cost devices that reduce electrocution risk of distribution power lines . these mechanisms should be easy to install . they should allow birds to perch and nest while protecting the lines to ensure that power supply is not disrupted due to bird electrocution events or short - circuiting by nest material . finally , conservation actions should include adequate spatial planning , avoiding the placement of power lines in areas of special conservation interest . the combination of an adequate spatial planning with a sustainable development of human infrastructures will contribute positively to the conservation of the spanish imperial eagle and may underpin population growth and range expansion , with positive side effects on other endangered species .
below , a darker , and more rufous , eastern , but with very heavy streaking on the breast and underwing coverts . such heavy streaking , especially on the lower breast and underwing coverts , shouldn´t be found even in the more streaked extremes of juvenile spanish .
in the 1960s , with just 30 pairs remaining , conservation efforts succeeded in increasing the population of the spanish imperial eagle by a considerable amount . however , since 1994 , the population has been allowed to decline again to just 160 pairs . there are several causes of this decline : habitat fragmentation due to deforestation for agricultural land and timber has disturbed breeding grounds , intentional poisoning on hunting reserves to reduce natural predators of game species , lead shot poisoning as a result of ingestion of game killed with bullets , and electrocution on power cables ( 2 ) . approximately 80 % of eagles killed on power lines are female , causing a greater impact on this monogamous species than if equal numbers of males and females were killed ( 10 ) .
adult spanish imperial eagles are sedentary birds and stay in their territory all year . juvenile birds disperse after becoming independent . some birds breed close to their birth site when they are old enough , others settle farther away from their birth site . this latter is called natal dispersal . in a study in spain that involved tracking 84 nestlings between 1990 and 2002 , the mean natal dispersal distance was 101 . 2 km with a maximum of 310 km ( 255 km for females ) [ gonzáles & margalida 2008 ] .
< p > a report about the electrocution of three spanish imperial eagles ( < em > aquila adalberti < / em > ) , seven bonelli’s eagles ( < em > aquila fasciata < / em > ) and one golden eagle ( < em > aquila chrysaetos < / em > ) in the guelmim region , morocco . the electrocuted eagles found in three days in autumn 2015 . < / p > < p > - additional photographs of the electrocuted eagles found on 22 october and 7 november are available in the link 1 below . < / p > < p > - photographs of the bonelli’s eagles found electrocuted on 5 december are available in the link 2 below . the pdf of the report updated on 17 december to include this new information . < / p >
large , dark eagle . generally dark brownish - black with prominent white “shoulders” on forewing and scapulars . pale golden - cream nape and pale grey basal area on uppertail . juvenile red - brown fading to pale buff with dark flight feathers and white fringes to coverts . in soaring and gliding flight wings held flat . similar spp . adult golden eagle a . chrysaetos lacks white “shoulders” and is less dark overall . immature has large white wing flashes and white base to tail . wings held in a flattened “v” shape . voice repeated barking owk .
the species is classified as vulnerable . threats include loss of habitat , human encroachment , collisions with pylons , and illegal poisoning . there has also been a decline in the spanish rabbit population , as a result of myxomatosis and other viral illnesses . the current population is estimated at less than 500 .
2009b ) . habitat alterations associated with agricultural expansion threaten historical and potential breeding sites in former range countries . hunting , poisoning , prey depletion and other mortality factors are also likely to pose threats along migration routes and in wintering areas . competition for nest sites with greater spotted eagle
. this network involved 112 municipalities , 22 , 000 ha of private land and 96 schools and focused on habitat management , species conservation , awareness and information activities across the whole spanish distribution ( cabezas - díaz and hernáez 2012 ) . in castilla - la mancha an eu life - funded projected ,
conservación de especies prioritarias en el monte mediterráneo en castilla - la mancha
is working for the conservation of this and other species related to mediterranean forests , while a counterpart project , life imperial , is focued on the conservation of the species in portugal ( r . alcazar
the most important prey is the rabbit . in a study in the montes de toledo ( castilla - la manche , spain ) the pellets of spanish imperial eagles were studied . in 93 % , remains of rabbits were found . other important prey was carrion ( found in 20 % of the pellets ) , pigeons ( 8 . 4 % ) , other birds ( 10 % ) , hare ( 2 % ) , reptiles ( 1 . 8 ) and partridge ( 0 . 6 % ) [ castaño lópez 2005 ] . that clearly shows how important the rabbit is for the species .
source : gonzález , l . m . ( 2012 ) . águila imperial ibérica – aquila adalberti . en : enciclopedia virtual de los vertebrados españoles . salvador , a . , morales , m . b . ( eds . ) . museo nacional de ciencias naturales , madrid . urltoken
immature spanish , mainly 3cy - 5cy , until they start to show pure white feathers on the leading edge of the wing , are in some cases , as already commented , very difficult to separate from eastern . at least , on actual knowledge . more work is needed to try to improve this in the future . .
separating immature ( mainly 3 - 5cy ) spanish from eastern , before the first obvious and diagnostic white feathers on the leading edge of the wing start to appear , usually by late 4cy - 5cy , can be very difficult , or even impossible in some cases . especially on birds lacking warm rufous immature - type feathers on the head , underparts and underwing coverts . and these birds exist . differences in structure ( spanish looking more “compact” , with proportionally shorter and broader wings , and a longer tail than eastern , and showing a heavier bill and more robust head / neck ) are somewhat variable , and probably of little use in the field for separating both species .
i do not know very well why the eagle threatens the vultures . but an observation by seo birdlife - spain , may give an explanation . thanks to a video , we can see a vulture coming to an abandoned eagle’s nest , maybe searching for chicks of eggs , or nest materials ? but later , a second vulture joins the first and the one in the nest is moving a stick with the bill . is it a pair ? we know that eurasian griffon vultures nest in cliff faces on rocky ledges , and maybe they need some sticks more to arrange the nest ? i do not have the answer .
female lays 2 or 3 eggs , between mid - february and late march , sometimes later . eggs usually are whitish , speckled with grey and brown . both sexes incubate , but mainly the female which is fed at nest by the male . the incubation lasts about 43 days . the chicks are covered with white down . usually , the smallest dies , but sometimes , all the chicks grow successfully . they are aggressive as soon as they hatch . they are fully feathered at 55 - 60 days of age , and they remain near the nest one month more . they return regularly at the nest - site and the members of the family may spend the winter together . the spanish imperial eagle may have 3 to 5 different nests , mainly in corn - oaks or pines , at about 10 to 25 metres above the ground . this species breeds between 300 and 900 metres of elevation , but sometimes up to 1500 metres in mountain areas .
adult spanish imperial eagles occupy territories in pairs all year - round ( 7 ) . they defend these territories from intrusion by other raptors , allowing only immature members of their own species to enter ( 8 ) . nests are built in tall oak trees , in small stands of trees far from human disturbance ( 7 ) . during the breeding season , both the male and female of the monogamous pair will participate in the incubation of the eggs and the care of the chicks . prey is torn into manageable pieces for the chicks until they are ready to hunt for themselves . parent eagles will even shelter the chicks from heavy rain ( 8 ) .
spp . ) , human disturbance of breeding sites , nest robbing and illegal trade , shooting , poisoning and electrocution by powerlines . an average of c . 450 eastern imperial eagles were killed by powerlines during the 2009 breeding season in the altai region 25 % of the total population of the region ( karyakin
the imperial eagle , like other birds of prey , is a carnivore . its main source of food is small mammals , such as rabbits and susliks ( ground squirrels in the genus spermophilus ) . it will also eat reptiles , including some poisonous snakes , and other birds , such as young magpies , waders , and ducks . it has been known to feed on some carrion as well , carrying off lambs that were already dead . it generally hunts by slowly flying around the open areas near the forest where it resides , staying just above the treetop level , and scanning for prey . its excellent eyesight makes spotting prey easy . when it sees a suitable prey item it will drop down on it , using its talons to kill and carry the prey . it will also , on a few occasions , perch high on a tree near the edge of the forest , and drop on its prey from there . ( channing 2000 , eagle 1988 )
i´m not completely sure of the id of the swedish bird . it is most likely an eastern , typically lacking any warm rufous colouration on the immature - type underparts and underwing coverts . however , based on my experience , it might be difficult to rule out with certainty a spanish . some of them can be scarily similar . see for yourself after checking some of the photos below…
the imperial eagle is the second largest eagle to reside in europe . it can grow to a length of about 0 . 92 meters . it can have a wingspan approaching 2 . 14 meters and have a mass of about 3 . 6 kilograms . adults are black - brown in color . they have a pale golden crown and nape , and a grey base that extends to the tail . they also have very apparent white
braces
on their scapulars . juveniles are paler and more variegated than the adults . they are yellow - brown with rump , wing , and tail patterns similar to the patterns of the lesser spotted , pale spotted , tawny and steppe eagles . they are distinguished from these eagles , however , by the bolder streaks on their under - wings and body , the lack of a pale band under the wing - coverts , and their larger and more majestic build . both the juveniles and adults have noticeably protruding head and long , parallel - edged wings . like other eagles , they have strong legs and feet . their feet include long , curved talons that they use to seize , kill and carry their prey . they also have large eyes that are located slightly to the side of the head . these eyes provide them with extremely keen eyesight , allowing them to spot prey from high in the air . ( peterson 1993 , wyss 1997 , eagle 1988 )
75 - 84 cm . large , dark eagle . generally dark brownish - black with prominent white
shoulders
on forewing and scapulars . pale golden - cream nape and pale grey basal area on uppertail . juvenile red - brown fading to pale buff with dark flight feathers and white fringes to coverts . in soaring and gliding flight wings held flat .
the action plan proposes the discouragement of intentional poisoning , as well as annual surveying of the breeding population , protection and management of all breeding sites , increasing the rabbit population and modifying dangerous power lines ( 2 ) . the modification of just 19 % of power poles in the range of the eagle could reduce total mortality by 52 % ( 12 ) .
apart from the previous pitfall , worn juvenile spanish imperials , especially those wintering in more southerly , sunny , latitudes , can be quite bleached in winter and spring , and can look very pale , with cold sandy - buff or even whitish upperparts , and also very pale , worn , upperwing coverts . they can be very similar to pale belisarius tawny eagles , and also to fulvescens gse .
eastern imperial eagles generally prefer to construct a nest in a tree which is not surrounded by other trees , so that the nest is visible from a considerable distance , and so that the occupants may observe the surroundings unobstructed . tree branches are taken in order to build the nest , which is upholstered with grass and feathers . very rarely it nests on cliffs or the ground .
this large eagle hunts for food , and his preferred prey is the rabbit , but partridges and other birds , mammals , some reptiles , fish and carrion are also taken according to the prey availability . he frequents plains and hills with cork - oaks or pines , and can be seen near streams and marshes . he may nest in mountains , from 300 - 900 metres up to 1500 metres of elevation .
the eagle nests in cork - oak or pine in the surroundings of the cliffs . sometimes , he perches on dead branch in tree , in the vicinity of the nest . his dark plumage contrasts against the green vegetation . but from the observatory , our eyes only see a small dark spot ! he is really the king of the park and we enjoy his presence when he deigns to appear above his domain .


phrynus longipes is a species of amblypygid found in the caribbean .

they are nocturnal predators that hide during the day in small retreats , and are sit-and-wait predators at night .

p. longipes can be found in subtropical and tropical forests , and caves within their range .

phrynus longipes

additional file 3 : video s2 : wettability of the cuticle of phrynus longipes some hours after moulting and between moultings ( playback speed : 50 % of real time ) . ( wmv 2236 kb )
ms . oh hell no . ( female phrynus parvulus whipscorpion ) by robin agarwal
owen , j . , & cokendolpher , j . ( 2006 ) . tailless whipscorpion ( phrynus longipes ) feeds on antillean crested hummingbird ( orthorhyncus cristatus ) . the wilson journal of ornithology , 118 ( 3 ) , 422 - 423 .
formation of the secretion coat in phrynus longipes . a . bare cuticle of a phrynus longipes 6 h after moulting , exhibiting tubercles ( tub ) and gland openings . asterisks indicate the major gland openings , arrowheads the minor gland openings . b . detail of minor gland opening in a specimen 24 h after moulting . note the thin homogeneous coat on the cuticle , except in the vicinity of the pore . note also the nano - crystals accumulating at the edge of the secretion coat ( arrowheads ) . c . detail of major gland opening in a specimen of p . longipes 24 h after moulting . note the continuous secretion film with evenly distribution of tiny droplets of a volatile substance ( arrowheads ) . d . detail of the margin of a major gland opening showing formation of globular granules from a continuous phase . ( jpg 719 kb )
guabá / tailless whipscorpion / phrynus marginemaculatus ( c . l . koch , 1841 ) ( juvenil / juvenile ) by carlos de soto molinari
additional file 2 : video s1 : phrynus decoratus showing super - hydrophobicity ( playback speed : 50 % of real time ) . ( wmv 1549 kb )
quintero , d . ( 1981 ) . the amblypygid genus phrynus in the americas ( amblypygi , phrynidae ) . the journal of arachnology , 9 , 117 - 166 .
phrynidae is a primarily neotropical family , ranging from southern florida and the caribbean to the amazon basin ( quintero , 1981 ) . phrynus , paraphrynus , and acanthophrynus are mostly represented by mesoamerican species , whereas heterophrynus contains typically south american species . one species of phrynid , phrynus exsul , is solely known from the flores and rinca islands of indonesia , making it the only known paleotropical member of its family . ( harvey , 2002 )
santer , r . , & hebets , e . ( 2009 ) . prey capture by the whip spider phrynus marginemaculatus c . l . koch . journal of arachnology , 37 ( 1 ) , 109 - 112 .
hebets , e . , & chapman , r . ( 2000 ) . electrophysiological studies of olfaction in the whip spider phrynus parvulus ( arachnida , amblypygi ) . journal of insect physiology , 46 ( 11 ) , 1441 - 1448 .
fowler - finn , k . , & hebets , e . ( 2006 ) . an examination of agonistic interactions in the whip spider phrynus marginemaculatus ( arachnida , amblypygi ) . journal of arachnology , 34 ( 1 ) , 62 - 76 .
peretti , a . ( 2002 ) . courtship and sperm transfer in the whip spider phrynus gervaisii ( amblypygi , phrynidae ) : a complement to weygoldt ' s 1977 paper . journal of arachnology , 30 ( 3 ) , 588 - 600 .
hebets , e . ( 2002 ) . relating the unique sensory system of amblypygids to the ecology and behavior of phrynus parvulus from costa rica ( arachnida , amblypygi ) . can . j . zool . , 80 ( 2 ) , 286 - 295 .
hebets , e . , & chapman , r . ( 2000 ) . surviving the flood : plastron respiration in the non - tracheate arthropod phrynus marginemaculatus ( amblypygi : arachnida ) . journal of insect physiology , 46 ( 1 ) , 13 - 19 .
phrynids are morphologically distinguished from other amblypygids by having 4 tibial segments on leg iv ( except for phrynus daimonoidaensis , which only has 3 ) and lacking pulvilli ( adhesive pads on the tarsi of many arthropods ) of their walking legs ( quintero , 1981 ) .
a possible inquiline relationship between phrynus gervaisii and the giant neotropical ant species paroponera clavata has been observed on barro colorado island ( leclere et al . , 1987 ) , and could extend to other species of phrynids across the bullet ant’s range . see “ behavior” section for details .
leclere , m . , mcclain , d . , black , h . , & jorgenson , c . ( 1987 ) . an inquiline relationship between the tailess whip - scorpion phrynus gervaisii and the giant tropical ant paraponera clavata . the journal of arachnology , 15 , 129 - 130 .
difference between the cuticle before and shortly after moulting in phrynus longipes . some hours after moulting ( left column ) the cuticle is highly wettable by water droplets ( arrowhead ) ( see also video additional file 3 : video s2 ) . the cuticle lacks the granules and is rather smooth on the microscale , except for small denticles ( den ) that help in arresting the later secretion layer . there are two different gland openings , the 2 - lipped major ( maj ) and the pore - like minor ( min ) one . between moultings ( right column ) , water droplets are completely repelled ( arrowhead ) by the cuticle . the secretion layer is visible , as white ‘powder’ ( arrowhead ) , composed of wrinkled granules ( gra ) , creating a micro - and nano - roughness
weygoldt , p . , rahmadi , c . , & huber , s . ( 2010 ) . notes on the reproductive biology of phrynus exsul harvey , 2002 ( arachnida : amblypygi : phrynidae ) . zoologischer anzeiger - a journal of comparative zoology , 249 ( 2 ) , 113 - 119 .
my boys and i found it under a rock just off a trail in the rancho ecologico sol de mayo in the sierra de la laguna . phrynus asperatipes , endemic to baja california sur . with those whip - like 1st legs and spined pedipalps , this is the most unusual arachnid i ' ve ever seen .
phrynidae consists of 4 extant genera : phrynus , paraphrynus , acanthophrynus , and heterophrynus ( quintero , 1981 ) . phrynids , due to their abundance throughout the neotropics , are some of the better - studied members of amblypygi , with much of what little we do know about whip spiders’ life history coming from these organisms ( hebets , 2002 ) .
effect and structure of the cuticular secretion in phrynus decoratus . water dripped from above onto the carapace forms a perfect droplet and rolls completely off ( see also video additional file 2 : video s1 ) . close inspection of the cuticle reveals a white ‘powder’ assembled on and between tubercles ( tub ) , consisting of globular particles , called granules ( gra ) ( from left to right ) . the granules arise from a colloidal secretion and may contain crystalline particles ( cry ) . the underlying cuticle ( bcl ) is smooth
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two of them fell off her and she never picked them up ( i don ' t know if amblypigid mothers even pick up fallen young ) , so i removed them . hopefully they survive without her .
so cute ! sorry about the two that fell off . unfortunately , in my experience ( with d . diadema ) , any babies that fall off that young aren ' t going to make it - but if they do pull through , i ' d love to hear how you did it !
so cute ! sorry about the two that fell off . unfortunately , in my experience ( with d . diadema ) , any babies that fall off that young aren ' t going to make it - but if they do pull through , i ' d love to hear how you did it ! congratulations on the rest of the babies !
thanks . i remember a thread from a while ago where someone raised a few babies that had fallen off ( or maybe their mother died ? i don ' t remember exactly ) and successfully got them to 2nd instar and beyond , so hopefully these two pull through .
cool , are you in the u . s . ? also , how large is the adult female ?
so the two that fell off while hatching into first instars just molted into second instars , at the same time as their siblings still with the mother . i will reintroduce these two to the others shortly .
awesome result ! nice to see more than once that it is possible to save babies that fell off the mother .
indeed . i didn ' t do anything special for them either , just carefully placed them on a piece of torn egg crate on some substrate .
glad to see some success in keeping the two younger ones alive ! keep us updated . i would love to see more cb amblies in the us hobby .
so the two that fell off while hatching into first instars just molted into second instars , at the same time as their siblings still with the mother . i will reintroduce these two to the others shortly . view attachment 223978
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senior curator & head , dept . of terrestrial zoology , western australian museum , locked bag 49 , welshpool dc , wa 6986 , australia
smaller arachnid orders catalogue ( saocat ) , database ( version oct . 2007 )
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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department of entomology , university of maryland , college park , maryland 20742 , u . s . a .
new terrestrial arachnids from the devonian of gilboa , new york ( arachnida , trigonotarbida ) .
mastigoproctus giganteus ( lucas ) ( arachnida : uropygi ) and its evolutionary significance . zoological journal of the linnean society
the feeding organs of the arachnida , including mites and ticks . smithsonian miscellaneous collections
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
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harvey , m . ( 2002 ) . the first old world species of phrynidae ( amblypygi ) : phyrynus exsul from indonesia . journal of arachnology , 30 ( 3 ) , 470 - 474 .
large spined pedipalps used in prey capture and intraspecific interactions are another shared feature of all amblypygids , with specific morphology of the weapons varying from species to species ( hebets , 2002 ) . some species of phrynids display sexual dimorphism in their pedipalps , with those of the males growing longer in proportion to their bodies than in females ( quintero , 1981 ; weygoldt et al . , 2010 ) .
little data exist on predation of amblypygids , but one individual was observed being eaten by a scorpion ( scorpiones ) ( fowler - finn & hebets , 2006 ) .
male phrynids have been observed wandering more than their relatively sedentary female counterparts , likely searching for mates . this observation likely applies to all amblypygid families ( fowler - finn & hebets , 2006 ) .
levi , h . , levi , l . , zim , h . , & strekalovsky , n . ( 1968 ) . a guide to spiders and their kin ( p . 117 ) . new york : golden press .
breeding in at least some phrynid species appears to be concentrated in the dry season , although low levels of breeding occurr year - round ( hebets , 2002 ) .
amblypygid courtship consists of the male orienting himself towards the female and vibrating his whips for several minutes , as in same - sex agonistic interactions . the male then deposits his spermatophore , a sperm - filled capsule attached to the substrate by a flexible stalk , and steps backwards to lure the female into taking up his sperm . ( weygolt et al . , 2010 )
phrynids are distiguished from other amblypygids in the scultped , triangular heads of their spermatophores ( peretti , 2002 ) , and a unique step in courtship where the male embraces his spermatophore for some minutes with his pedipalps before luring the female to it ; the purpose of this is unknown ( weygolt et al . , 2010 ) .
all amblypygid females carry their eggs in a sac on the ventral side of their opisthosoma ( the posterior segment of the chelicerate body ) . in phrynids , the eggs tend to develop for around 4 months ( weygolt et al . , 2010 ) . the hatched nymphs are then carried on the mother’s back for about 2 weeks ( weygolt et al . , 2010 ) , longer than average amblypygid period of 4 - 6 days ( levi et al . , 1968 ) .
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effects of graded hypophysectomy on the gonads of the male crested newt ( triturus cristatus carnifex laur . )
foraging ecology of the great crested grebe ( podiceps cristatus l . ) at a mesotrophic - eutrophic reservoir
this is a tailless whip - scorpion . best i can tell there is only one species in florida and it also occurs on some caribbean islands . for more information : urltoken
this remarkable arachnid belongs to the ancient order of whip spiders ( amblypygi ) and , as a such , differs from modern spiders in many ways . one pair of elongate legs is used like antennae and only three pairs for walking . whip spiders do not produce venom , but grab their prey with raptorial pedipalps . they do not produce silk , and the mating behaviour is more scorpion like : the male deposits a spermatophore on the ground and guides the female over it , which will pick it up .
es una araña no verdadera , que a diferencia de las arañas verdaderas esta posee y anda en seis patas y el otro par ha sido modificada a pinzas y que su obdomen posee segmentos como los alacranes y escorpiones , se alimenta de insectos , lagartos y sapos , es de vida nocturna basicamente . es como una mezcla entre anaña y escorpion , se le llama a veces escorpion sin cola o araña latigo por la forma de sus paras delanteras .
saltan , son rapidisimos e incluso algunos han atacado el lente de mi camara . es comun en todo el mundo , en r . d . hay descritas unas cinco especies .
nürnberg : in der c . h . zeh ' schen buchhandlung , 1831 - 1848 . .
an amblypygid , or whip spider . identified by
d _ b
at inaturalist . org .
taken at the seattle bug safari in 2011 . this photo is licensed creative commons so please use it for any purpose you see fit .
falch iawn cael hyn yr un dydd â phrynu a dechrau darllen ready player one . # airwolf by nic dafis
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fiona m . jones , jason a . dunlop , matt friedman , russell j . garwood ; trigonotarbus johnsoni pocock , 1911 , revealed by x - ray computed tomography , with a cladistic analysis of the extinct trigonotarbid arachnids , zoological journal of the linnean society , volume 172 , issue 1 , 1 september 2014 , pages 49–70 , urltoken
trigonotarbids ( arachnida : trigonotarbida ) are an extinct order of arachnids that ranged from the late silurian ( přídolí ; c . 420 mya ) through to the early permian ( sakmarian ; c . 290 mya ; dunlop , 1996a ; dunlop & rößler , 2013 ) . sixty - seven species are currently recognized in the literature , although this is probably an overestimate ( see comments in garwood & dunlop , 2011 ) . historically trigonotarbid genera and species were often diagnosed on taphonomic features , sexual dimorphism , and / or different growth stages . this has led to the creation of numerous synonyms , which revisions are gradually starting to address ( rößler , 1998 ; garwood & dunlop , 2011 ) .
reconstructions of representatives of nine trigonotarbid families , shown to scale . a , palaeocharinus rhyniensis ( palaeocharinidae ) . b , archaeomartus levis ( archaeomartidae ) . c , anthracomartus hindi ( anthracomartidae ) . d , anthracosiro woodwardi ( anthracosironidae ) . e , trigonotarbus johnsoni ( trigonotarbidae ) . f , lissomartus schucherti ( lissomartidae ) . g , aphantomartus pustulatus ( aphantomartidae ) . h , eophrynus prestvicii ( eophrynidae ) . i , kreischeria wiedei ( kreischeriidae ) . scale bar = 10 mm .
the name trigonotarbida stems from petrunkevitch ( 1949 ) who split anthracomarti into two , supposedly unrelated , groups . he retained anthracomartus karsch , 1882 , within anthracomarti , along with the other genera whose opisthosomal tergites were divided into five separate plates . anthracomarti and another extinct order , haptopoda , were placed by petrunkevitch in a subclass stethostomata : defined as arachnids with a broad prosoma–opisthosoma junction and downward - hanging mouthparts . the remaining fossils–including trigonotarbus pocock , 1911 –with tergites divided into three plates , formed the new order trigonotarbida . this was placed in its own suborder soluta : defined as arachnids with a prosoma–opisthosoma junction of variable width . this scheme was maintained by petrunkevitch ( 1953 ) , becoming the standard classification through its use in the treatise on invertebrate paleontology ( petrunkevitch , 1955a ) . here the order names were emended to anthracomartida and trigonotarbida to fit nomenclatural conventions .
based on a series of fossils from the famous coal measures site of coseley near dudley in the english west midlands ( see material and methods ) . these specimens originated from the egginton , johnson , and madeley collections . henry johnson ( 1823–1885 ) , after whom the species is named , was a civil and mining engineer and a fellow of the geological society who lived in dudley and was noted as an avid fossil collector (
is one of the smaller coal measures trigonotarbids , typically about 5–6 mm in body length , and has a rather simple body plan without strong ornamentation of the body . the carapace is notably triangular and is presumably the source of the genus name : a combination of the ancient greek
. the carapace has a raised central ridge bearing a pair of median eyes . the legs are fairly short and stubby . as noted above ,
trigonotarbus johnsoni is known from at least 13 specimens ( see systematic palaeontology ; fig . 2 ) , all housed in the natural history museum london ( nhmuk ) . the fossils are from the late carboniferous , similis–pulchra zone of the clay croft open - cast works at coseley near dudley , staffordshire , uk .
hand specimens of trigonotarbus johnsoni from the nhmuk collections . scale bars = 1 mm .
tomographic reconstruction of trigonotarbus johnsoni based on specimen nhmuk i . 15860 . a , dorsal view . b , fourth right leg , podomeres labelled . c , ventral view . d , ventrolateral view of the coxo - sternal region . abbreviations : 1–12 , segment number ; ce1−2 , coxal endites 1–2 ; ch , chelicerae ; cl , clypeus ; fe , femur ; l1−l4 , legs 1–4 ; me , median eye tubercle ; mt , metatarsus ; pa , patella ; pp , pedipalp ; pr , projection ; ta , tarsus ; ti , tibia ; tr , trochanter . scale bars : a , c = 2 mm , b = 1 mm , d = 0 . 5 mm .
an idealized reconstruction of trigonotarbus johnsoni based on the computed tomography scan of nhmuk i . 15860 and additional hand specimens . scale bar = 2 mm .
the results of the cladistic analysis . the four trees show , as labelled , the agreement subtree and strict consensus of both the equally weighted analysis and implied weights analyses with concavity constants ( k ) of 0 . 25 , 1 , 3 , and 10 .
1911 trigonotarbus johnsoni pocock , pp . 74–75 , fig . 38 , pl . 3 , fig . 4 .
1949 trigonotarbus johnsoni pocock ; petrunkevitch , pp . 250–255 , figs 104–109 , 111 , 250–257 .
1955a trigonotarbus johnsoni pocock ; petrunkevitch p . 112 , figs 77 ( 2 ) , 79 ( 1–2 ) .
1996a trigonotarbus johnsoni pocock ; dunlop , pp . 611–612 , text - fig . 4 .
nhmuk in 31239 . from coseley near dudley , staffordshire , uk . late carboniferous ( similis–pulchra zone , westphalian b in the european stratigraphical scheme and moscovian in an international stratigraphical framework ; wilson , 2005 ) .
nhmuk i . 7895 , i . 13911 , i . 13919 , i . 13921 , i . 15859 , i . 15860 , i . 15897 , in 31240–41 , in 31243 , and in 31254–55 . all from the same locality as the holotype .
small , stubby - limbed trigonotarbids with triangular carapace , lacking lateral lobes , but with a raised median ridge bearing the eyes and a deep , inverted t - shaped depression behind the eyes . rounded opisthosoma has laterally undivided tergite nine , apparently with fused sternites 9 and 10 bearing the pygidium at their anterior . ( emended from petrunkevitch , 1955b . )
trigonotarbus johnsoni was described in some detail by pocock ( 1911 ) and petrunkevitch ( 1949 ) . here , we redescribe it from restudy of the hand specimens ( fig . 2 ) and tomographic reconstruction of nhmuk i . 15860 ( fig . 3 , files s2 , s3 ) . except where noted , measurements derive from the latter . small trigonotarbid ; holotype 4 . 8 mm long ( carapace 2 . 0 mm in length , opisthosoma 2 . 8 mm long by 2 . 5 mm wide ) ; scanned specimen 6 . 3 mm in length ( carapace 2 . 8 mm in length , opisthosoma 3 . 5 mm long by 3 . 6 mm wide ) . carapace triangular with raised median area , increasing in width posteriorly ( fig . 3a ) . raised median area drawn into anteriorly pointing , rounded clypeus , surrounded by zones of lower relief . carapace with smooth lateral margins ( see , e . g . nhmuk in 32139 ) , i . e . no division into lobes , posteriorly demarcated by strong transverse groove ; some specimens ( e . g . nhmuk in 31241 ) reveal depression on median ridge behind the eyes and shaped like an inverted t that effectively merges into this groove . closely spaced median eyes situated on median ridge , towards front of carapace ( nhmuk in 31239 ; fig . 2 ; nhmuk i . 15860 , fig . 3 ) .
if an alternative scheme were to be chosen in which the first fully visible tergite is ( despite its short length ) a fused diplotergite 2 + 3 , then tergite nine must be both laterally and longitudinally divided . a comparable condition is observed in anthracomartidae ( garwood & dunlop , 2011 ) and some eophrynidae . this would then no longer require a fused sternite 9 + 10 , but would make the anterior ventral segmentation difficult to reconcile with either the hand specimens or the tomographic model . for this reason the former hypothesis of segmentation is preferred .
despite the increased morphological information provided in the current study , trigonotarbidae remain very difficult to place definitively . in some iw analyses ( k = 0 . 25 , 1 ; see supporting information file s4 for trees ) , the two included trigonotarbus species are recovered as sister taxa , whereas in other analyses they are variously a grade at the base of the non - anthracomartid / palaeocharinus clade ( k = 3 ) or widely separated ( k = 10 ) . this could be partly because of misinterpretation of the devonian species , trigonotarbus stoermeri , for which schultka ( 1991 ) confused dorsal and ventral features . a redescription of this species with a test of its generic affinities would be welcome . it is interesting to note that in several trees the trigonotarbus species group with palaeotarbus , the earliest known trigonotarbid , which appears superficially similar in appearance to the trigonotarbidae .
despite several problematic , highly mobile , taxa removing the signal in the strict consensus trees , a consistent result of our analysis is a split at the base of the trigonotarbids . one reliable ‘anthracomartid’ clade ( fig . 5 ) comprises ( palaeocharinus ( archaeomartidae + anthracomartidae ) ) , which is principally defined by the presence of a somewhat box - shaped carapace . our data thus offer cladistic support to the suggestion by poschmann & dunlop ( 2010 ) that the palaeocharinus species from the rhynie and windyfield cherts are closely related to anthracomartidae , with archeaomartidae as a putative intermediate taxon .
the resolved phylogeny also demonstrates that the split between these two major clades must have occurred by the early devonian at the latest . the earliest ( silurian ) trigonotarbid palaeotarbus is consistently recovered within the non - anthracomartid clade ( see supporting information file s4 ) , whereas palaeocharinus from the devonian rhynie chert is placed in the ‘anthracomartid’ clade . this implies either an early split as part of rapid diversification after terrestrialization within the trigonotarbids , or it could reflect early terrestrialization and then a long period from which we have no trigonotarbids prior to the earliest fossils , a possibility given the patchy fossil record of pre - carboniferous terrestrial arthropods . either scenario lends credence to the suggestion that the peak in richness seen in coal measures deposits is more likely to be taphonomic , or sampling bias , rather than representing a genuine interval of diversification within the trigonotarbids .
we thank : r . mounce for duplicating our cladistic results ; d . legg and r . sansom for advice on phylogenetic inference ; g . edgecombe for access to his microscope ; c . mellish for access to fossils in her care ; and two anonymous reviewers whose comments and suggestions improved the manuscript . r . garwood is an 1851 royal commission research fellow , a scientific associate at the natural history museum , london , and member of the interdisciplinary centre for ancient life ( umri ) . infrastructure for the processing of ct data at oxford was supported by the john fell fund of the university of oxford and a leverhulme prize to m . friedman . part of the paper was completed during a synthesys - funded visit to berlin awarded to r . garwood .
the bridgewater treatises on the power , wisdom and goodness of god as manifested in the creation . treatise iv . geology and mineralogy with reference to natural theology
the youngest trigonotarbid permotarbus schuberti n . gen , n . sp . from the permian petrified forest of chemnitz germany
contributions to our knowledge of the arthropod fauna of the west indies . –part i . scorpiones and pedipalpi ; with a supplementary note upon the freshwater decapoda of st . vincent
first palaeozoic arachnid from iberia : aphantomartus areolatus pocock ( basal stephanian ; prov . león , n . w . spain ) , with remarks on aphantomartid taxonomy
arthropods from the lower devonian ( lower emsian ) of alken an der mosel , germany . part 1 : arachnida
0 , flat ; 1 , subrectangular and box - like ; 2 , subtriangular ; 3 , rounded .
a distinctly box - like carapace is found in several groups including palaeocharinus and anthracomartidae , whereas a subtriangular carapace is typical of , for example , the eophrynid assemblage–often coupled with a spine - like , anteriorly projecting clypeus . a rounded carapace is unusual , but found in both some outgroups and trigonotarbids such as anthracosironidae . a flat carapace is found in some outgroup arachnids amongst the taxa scored here .
0 , with elongate depression ; 1 , unmodified ; 2 , raised into a ridge ; 3 , raised into a lobed ridge .
the median axis of the trigonotarbid carapace , although sometimes unmodified , is often marked by either a ridge ( with or without lobes , the latter being typical of the eophrynid assemblage ) , or a depression in some anthracomartids .
in eophrynidae , and some apparently associated taxa , the lateral margins of the carapace are divided into a prominent series of lateral lobes . we also score lobes as present in lissomartus where these divisions ( faintly ) appear .
the clypeus in trigonotarbid nomenclature refers to the presence of an anteriorly projecting region of the prosomal dorsal shield ( or carapace ) , in front of the median eyes .
0 , lobate ; 1 , projecting spine ; 2 , downward - pointing projection .
anthracosironidae and a number of other taxa have a rounded or lobate clypeus , whereas others , such as eophrynidae , possess a projecting anterior spine . anthracomartidae and palaeocharinus tend to have a ventrally projecting clypeus resembling a small nose . this character is inapplicable to the outgroups .
trigonotarbid genera such as palaeocharinus , archaeomartus , and gigantocharinus have small , tooth - like projections on the tip of the clypeus . this character is inapplicable to the outgroups .
in palaeocharinus and gigantocharinu s the clypeus bears slightly raised ridges , which run up towards the median eyes . this character is inapplicable to the outgroups .
in anthracomartidae , palaeocharinidae , and some outgroup taxa , the ventral carapace lateral margins possess a series of distinct indentations to accommodate the coxa−trochanter articulation .
anthracomartidae have a distinct transverse depression near the back of the carapace , but in front of the locking ridge . this is most pronounced in anthracomartus hindi .
some trigonotarbids ( and all outgroups ) posses a concave posterior prosomal boundary , whereas a number of other trigonotarbid taxa have instead a straight margin here .
in the outgroups palpigradi and amblypygi the sternum is composed of a series of separate sclerites . in araneae and all trigonotarbida where the sternum can be seen , the sternum is a single , undivided plate . it is not clear which of these conditions represents the plesiomorphic state . ricinuleids lack a sternum .
in anthracosiro woodwardi the median eyes appear to be ( uniquely ) quite far apart at the ends of an essentially t - shaped ridge ( e . g . pocock , 1911 : fig . 35 ) . although currently autapomorphic for the species in this analysis , we include the character for reference in case later finds are scored into our matrix .
0 , in front of leg i ; 1 , level with leg i ; 2 , level with leg ii ; 3 , level with leg iii .
this character essentially explores the relative position of the median eyes , namely the degree to which they are towards the front or middle of the carapace .
outgroups such as xiphosura and eurypterida imply that the arachnid ground pattern included a pair of lateral eye tubercles . lateral eyes were retained in araneae and amblypygi amongst the outgroups tested here , and these eyes–or at least their tubercles if individual lenses cannot be resolved–are retained in at least palaeocharinids , anthracomartids , and ( probably ) archaeomartus and anthracosiro amongst the trigonotarbids . in other trigonotarbids the lateral eyes have unequivocally been lost .
trigonotarbids possess a broad prosoma−opisthosoma boundary , a trait they share with the ricinuleids but not with the other arachnid outgroups in this analysis , which have a very narrow junction here .
the first tergite of most trigonotarbids is modified into a locking ridge that is tucked into a fold at the back of the carapace and presumably prevented movement at the prosoma−opisthosoma boundary . the same trait is found in ricinuleids , although the character was apparently reduced in some trigonotarbids .
trigonotarbids apparently lack a sternite associated with the first opisthosomal segment , an observation confirmed on the basis of rhynie and windyfield chert taxa . outgroups retain a first opisthosomal sternite ; thus its absence is potentially diagnostic for trigonotarbida .
0 , rounded to suboval ; 1 , distinctly longer than wide with straight , subparallel lateral margins .
anthracosironidae are unique amongst the trigonotarbids in having a long opisthosoma with almost - parallel lateral margins .
the posterior opisthosomal margin is highly variable in trigonotarbids , being variously straight , convex , or concave in nature .
this character captures the number of sclerites for each opisthosomal tergite–i . e . one in outgroups , three in most trigonotarbids , but five in anthracomartidae .
0 , greater than half of body width ; 1 ; less than half of body width .
the median row of opisthosomal sclerites are noticeably narrower in anthracomartidae and archaeomartidae compared with other trigonotarbid taxa , in which the median sclerites make up the majority of the opisthosomal width .
in some taxa there is a clear reduction in the width of the median sclerites going posteriorly .
many trigonotarbid taxa have tergites two and three fused into a so - called diplotergite . the plesiomorphic state of this character is unclear , but it could be part of the trigonotarbid ground pattern with reversals in some more derived species .
eophrynidae , and some other trigonotarbid taxa , have a median procurved , anteriorly directed diversion of the tergite boundary .
the majority of trigonotarbid taxa possess a laterally divided ninth tergite , but some trigonotarbids ( and the outgroups ) have tergite nine undivided .
anthracomartidae and pleophrynus have a transverse line dividing the ninth tergite into anterior and posterior plates .
trigonotarbus johnsoni uniquely possesses in our interpretation of segmentation a very narrow tergite nine . although currently phylogenetically uninformative , this is included for future additions to the matrix .
0 , absent ; 1 , incomplete and only in anterior segments ; 2 , complete and surrounding the whole ventral opisthosomal margin .
anthracomartidae and palaeocharinus possess a ventral suture line , which is incomplete in the latter ( devonian ) genus , but contiguous in carboniferous taxa , creating median and lateral sternal plates .
ventral sacs are enigmatic opisthosomal structures found in many trigonotarbids , with possibly homologous features known in both palpigrades and amblypygids amongst the outgroups . they are variable in form within the trigonotarbids , being variously found as a paired structure , with occasional transverse ridge ( s ) anterior to this .
0 , straight ; 1 , rounded ; 2 , angles and chevron - shaped .
sternite shape within the trigonotarbids is variable . some taxa have straight margins–based on outgroup comparison this is probably the plesiomorphic state . others have rounded sternites whose edges are posteriorly directed , whereas some have chevron - shaped sternites ( i . e . straight sternal margins meet to form a sharp v - shape at the median axis ) .
the trigonotarbid pygidium comprises segments eleven and twelve , and is not seen in the outgroups . ricinuleids have by contrast three terminal ring - like segments .
0 , intersects segment 9 only ; 1 , cuts segment 8 ; 2 , cuts segment 7 .
this character captures the position of the pygidium relative to the opisthosomal segmentation if a horizontal line were to be carried laterally from the pygidial position . it is impacted by the degree of curvature of the sternites .
although rarely preserved , the rhynie and windyfield chert trigonotarbids possess a terminal palpal chela ; a character they share with ricinuleids , but not with other outgroup taxa .
a laterigrade stance , with the legs twisted to have the prolateral surface more or less facing upwards , is observed in some trigonotarbids ; namely within anthracomartidae and anthracosironidae in which it is presumably an adaptation towards ambush predation . this is also present in the amblypygid outgroup .
a laterigrade stance is accompanied in some taxa , e . g . anthracosiro , by spination of the anterior limbs , which we score here as an adaptation for a raptorial limb function .
palaeocharinus and anthracomartidae possess median endites on the leg coxae of indeterminate function . the lip - like projection on the first leg coxae of tr . johnsoni are not considered homologous for this character as it is found on a single limb , and is anteriorly rather than medially projecting .
0 , leg iv less than or equal to total body length ; 1 , longer than total body length .
0 , noticeably shorter than tarsus ; 1 , as long as or longer than tarsus .
anthracomartidae and palaeocharinidae possess a metatarsus , which is noticeably shorter than the tarsus , whereas in other trigonotarbids ( and the outgroups ) the metatarsus is usually proportionally longer .
a pitted surface ornamentation can be seen in the cuticle of anthracomartidae and eophrynidae .
both eophrynus and pleophrynus are characterized by a unique grouping of four tubercles forming a cross - shape immediately behind the median eye tubercle .
in addition to large surface pits , anthracomartidae have a granular surface texture on their cuticle . granulation is also seen in tynecotarbus .
the eophrynid - like trigonotarbids can be recognized in part by a heavily ornamented , pustulate or tuberculate dorsal surface .
in eophrynidae and kreischeriidae the tubercles are found in a distinct pattern on each tergite , allowing the easy identification of individual segments through this repeating pattern .
in eophrynus the lateral margins of the tergites are punctuated by upward - pointing spines . this is only applicable to a single taxon in the present study ( and hard to detect in flattened material ) , but is included to allow later expansion of the matrix .
eophrynidae and kreischeriidae have backwards - directed , presumably defensive , spines on the posterior opisthosomal margin .
three trigonotarbids possess an ornament in the form of a row of small tubercles following the lateral margins of the opisthosoma .
this character reflects the ( rare ) presence of any form of tuberculation on the ventral opisthosoma , and is unknown in those taxa that only preserve the dorsal surface .
0 , absent ; 1 , paired ridges on the midline ; 2 , paired angled ridges tapering posteriorly .
a variety of ventral ridges are found amongst trigonotarbid species . if present , these can be observed as a parallel pair of ridges along the midline of the opisthosoma of pleophrynus , and an angled pair of ridges in anthracomartidae .
a few trigonotarbid species have limb articles with ornamentation in the form of tubercle rows , or continuous ridges or depressions forming longitudinal lines .
barranco & harvey , 2008 ; barranco & mayoral , 2007 ; broza , poliakov & conde , 1993 ; harvey , štáhlavský & theron 2006 ; van der hammen , 1969 .
file s1 . the matrix used for the current analysis in the form of a . tnt file . file can be opened in any text editor , in addition to tnt .
file s2 . an animation of the tomographic reconstruction of trigonotarbus johnsoni of specimen nhmuk i . 15860 .
file s3 . a vaxml model of the tomographic reconstruction of trigonotarbus johnsoni of specimen nhmuk i . 15860 . see sutton et al . ( 2012 ) for further information on the vaxml interchange format .
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of territorial behavior , which showed that residency effects were more profound in epigean interactions than in those of cave conspecifics . the high density of amblypygids in the cave population may have resulted in more tolerant , less aggressive behaviors relative to epigean conspecifics . thus , these findings fit the ecological and demographic conditions of each environment .
among both plants and arthropods , super - hydrophobic surfaces have evolved that enable self - cleaning , locomotion on water surfaces , or plastron respiration . super - hydrophobicity is achieved by a combination of non - polar substances and complex micro - and nano - structures , usually acquired by growing processes or the deposition of powder - like materials .
here we report on a multi - phasic secretion in whip spiders ( arachnida , amblypygi ) , which externally forms durable , hierarchical microstructures on the basically smooth cuticle . the solidified secretion crust makes the previously highly wettable cuticle super - hydrophobic . we describe the ultrastructure of secretory cells , and the maturation and secretion of the different products involved .
whip spiders represent intriguing objects of study for revealing the mechanisms of the formation of complex microstructures in non - living systems . understanding the physical and chemical processes involved may , further , be of interest for bio - inspired design of functional surface coatings .
] . however , further data on this ‘powder’ are lacking . it has been reported that whip spiders are able to survive submerged due to plastron respiration [
] , however the physical mechanism of plastron formation remained obscure . it was proposed that the wrinkled structure of the cuticle close to the book lungs is responsible for water - repellence and air - entrapment [
] . this may indicate the importance of secretions for waterproofing . we hypothesize that the granular coating reported by weygoldt [
] is a secretion product that is responsible for the entrapment of air and a repellence of water . in the present study we sought to test this hypothesis and to reveal the fine structure and origin of this substance .
teruel & armas 2005 from cienfuegos - cuba ( phrynidae ) . this covers all extant whip spider families , except paracharontidae , which are nearly unobtainable [
] . temperature was kept constant at 26–27°c and relative humidity varied between 65 and 75 % . animals were fed every seven days with cricket nymphs (
to test the wettability of the whip spider cuticle , 30 μl droplets of tap water were dripped on the carapace ( dorsal prosomal shield ) from a height of 1–5 cm ( depending on the size of the animal ) using a micro - pipette . tap water had the following characteristics : conductance 448 μs ; gh 14° ; kh 10 , 5 ; ph 8 , 0 nh 4 < 0 , 05 mg / l ; no 2 0 , 025 - 0 , 05 mg / l ; no 3 1 , 0 mg / l ; fe 0 , 02 - 0 , 05 mg / l ; po 4 < 0 , 02 mg / l ; sio 2 2 , 0 - 3 , 0 mg / l ; mg 10 , 0 ; k 2 , 0 mg / l . in a second test series , two days later , droplets were carefully placed on the carapace , and we observed whether they remained attached or rolled off when the animal moved . both nymphs and adults were tested , with 3–15 individuals per species .
freshly moulted and inter - moult specimens of c . acosta were treated as described above , including the complete dehydration step in acetone before being embedded into agar low viscosity resin ( lvr , agar scientific , stanstead , essex , uk ) . cured resin blocks were sectioned at 1 μm ( semithin ) or 60 nm ( ultrathin ) on a leica uc6 ultramicrotome ( leica microsystems , wetzlar , germany ) . semithin sections were stained with toluidine blue for few seconds on a heating plate at 60 °c , whereas ultrathin sections were contrasted with uranylacetate for 30 min and leadcitrate for 5–10 min . light microscopy sections were viewed with an olympus bx53 equipped with a dp73 microscope camera ( olympus , tokyo , japan ) . ultrathin sections were analyzed with a zeiss libra120 tem ( zeiss , oberkochen , germany ) .
we found that the whip spider integument is covered with a crust of a solidified secretion that forms globular microstructures ( granules ) with a diameter of 0 . 3–5 . 0 μm . granules may exhibit a coat of assembled crystal - like nano - particles ( figs .
) . the granules have a rough surface structure that is highly species - specific ( fig .
) . the secretion is present on all body parts , except regions situated close to and at joint membranes and segment borders , the distitarsi and pretarsi of legs , parts of the chelicerae and the eyes . it may also be absent on the tips of tubercles that widely cover the dorsal cuticle of the carapace and parts of the legs ( except in
water repellence and ultrastructure of granules in different species of whip spiders . 1 st row : tap water droplets , directly ejected on the carapace , form a spherical shape in most species . 2 nd row : the fine structure of granules is highly species - specific . 3 rd row : the ultrastructure of colloid particles differ between species
) . the bulk of a granule is formed by a homogeneous material . it is covered by a porous crystal - like layer of assembled nano - particles . the outermost layer is thin and irregular and composed of a loose granular substance . all these substances are rather electron - dense . sem observations of fractured cuticles in
) show that the inner material appears always homogeneous , and that species - specific differences in granule morphology are the result of the specific structure of the second crystal - like layer . the uppermost layer was not visible in sem images , and may be a fluid .
. histological section of the integument of a freshly moulted animal ( dorsal carapace ) . different epidermis cells are present between the cuticle ( cut ) and the musculature ( mus ) , containing either spherical strongly stained vesicles ( ves ) or large vacuoles ( vac ) . the 2 - lipped major gland openings ( maj ) are connected to a secretory cell ( sce ) , in which the content of different vesicles is apparently mixed ( see fig .
. an sem micrograph of the cuticle of a freshly moulted animal shows the absence of the granular secretion coat .
. histological section of the integument of an inter - moult animal ( same scale as in a . ) . the epidermal cells are reduced in size , the vacuoles shrink , and the stained vesicles are mostly absent . the solidified secretion layer is assembled on the cuticle and consists of a thin continuous layer and globular granules ( gra ) , both of which are highly stained like the vesicles in a . the different in the thickness of the cuticle ( cut ) if compared to a . is due to the difference in the animal instar .