kimberleyeleotris is a genus of sleeper gobies endemic to australia , where they are only known from rivers in the kimberley region of western australia .

kimberleyeleotris

drysdale gudgeon , kimberleyeleotris notata . source : jill ruse . license : all rights reserved
mitchell gudgeon , kimberleyeleotris hutchinsi . source : mark allen . license : all rights reserved
kimberleyeleotris hutchinsi voucher ams i . 33464 - 004 zic family member 1 ( zic1 ) gene , partial cds
kimberleyeleotris hutchinsi voucher ams i . 33464 - 004 cytochrome b ( cytb ) gene , complete cds ; mitochondrial
kimberleyeleotris hutchinsi voucher ams i . 33464 - 004 recombination activating protein 1 ( rag1 ) gene , partial cds
kimberleyeleotris hutchinsi voucher ams i . 33464 - 004 g protein - coupled receptor 85 ( gpr85 ) gene , partial cds
kimberleyeleotris hutchinsi voucher ams i . 33464 - 004 12s ribosomal rna gene , partial sequence ; trna - val gene , complete sequence ; and 16s ribosomal rna gene , partial sequence ; mitochondrial
wager , r . 1996 . kimberleyeleotris notata . in : iucn 2011 . iucn red list of threatened species . version 2011 . 2 . < urltoken > . downloaded on 10 january 2012 .
wager , r . 1996 . kimberleyeleotris hutchinsi . in : iucn 2011 . iucn red list of threatened species . version 2011 . 2 . < urltoken > . downloaded on 10 january 2012 .
kimberleyeleotris notata hoese & allen 1987 , mem . mus . vict . 48 ( 1 ) : 40 , figs 3 , 4 . type locality : drysdale river , 4 km above junction with forest creek , kimberley region , western australia .
kimberleyeleotris hutchinsi hoese & allen 1987 , mem . mus . vict . 48 ( 1 ) : 36 , figs 1 , 2 [ incorrectly spelled kimberleotris hutchinsi ] . type locality : tributary of mitchell river , kimberley region , western australia .
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, select family and click on ' identification by pictures ' to display all available pictures in fishbase for the family .
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cfm script by eagbayani , 30 . 11 . 04 , , php script by cmilitante , 05 / 11 / 2010 , last modified by cmilitante , 14 / 03 / 2013
known only from the drysdale river in the kimberley region of north western australia . the species inhabits clear , freshwater pools of slow - flowing streams over rocky and sandy bottoms .
the specific name notata is from the latin nota ( mark , sign ) , in reference to the pattern of markings on the side of this species .
allen , g . r . 1982 . inland fishes of western australia . perth : western australian museum 86 pp . 6 figs 20 pls [ p . 61 , pl . 13 ( 7 ) , as new genus and species b ]
allen , g . r . 1989 . freshwater fishes of australia . neptune , new jersey : t . f . h . publications 240 pp . , 63 pls
allen , g . r . , midgley , s . h . & allen , m . 2002 . field guide to the freshwater fishes of australia . perth : western australian museum 394 pp .
morgan , d . l . , allen , g . r . , pusey , b . j . & burrows , d . w . 2011 . a review of the freshwater fishes of the kimberley region of western australia . zootaxa 2816 : 1 - 64 .
morgan , d . l . , unmack , p . j . , beatty , s . j . , ebner , b . c . , allen , m . g . , keleher , j . j . , donaldson , j . a . & murphy , j . 2014 . an overview of the ' freshwater fishes ' of western australia . journal of the royal society of western australia 97 : 263 - 278 .
unmack , p . j . 2001 . biogeography of australian freshwater fishes . journal of biogeography 28 : 1053 - 1089 .
oceania : known only from the drysdale river system in the northern kimberley region of western australia .
maturity : l m ? range ? - ? cm max length : 4 . 0 cm tl male / unsexed ; ( ref . 5259 )
inhabits clear , freshwater pools of slow - flowing streams over rock and sand bottoms ( ref . 44894 ) .
allen , g . r . , 1989 . freshwater fishes of australia . t . f . h . publications , inc . , neptune city , new jersey . ( ref . 5259 )
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 7500 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01000 ( 0 . 00244 - 0 . 04107 ) , b = 3 . 04 ( 2 . 81 - 3 . 27 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 1 ±0 . 1 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 10 of 100 ) .
if you have images for this taxon that you would like to share with atlas of living australia , please upload using the upload tools .
yearsley , g . k . , last , p . r . & morris , g . b . 1997 ,
codes for australian aquatic biota ( caab ) : an upgraded and expanded species coding system for australian fisheries databases
, pp . 15 pp . + appendices
urn : lsid : biodiversity . org . au : afd . taxon : 4c50c7ca - a0a1 - 4e17 - b27c - 17d9fe569d8d
urn : lsid : biodiversity . org . au : afd . taxon : 2dd4c6c8 - c2ce - 4d3b - 83b2 - 77207e14f225
urn : lsid : biodiversity . org . au : afd . name : 450751
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oceania : known only from the mitchell river system in the northern kimberley region of western australia .
inhabits still or slow - flowing waters , usually in rocky pools . occurs in streams . usually found close to submerged sandstone boulders , but occasionally seen hovering in mid - water ( ref . 44894 ) . solitary , often hovering over submerged sandstone boulders , but sometimes forms loose aggregations in mid - water . males are more colorful and slightly larger than females . attractive species well suited for captivity , but is unknown to aquarists ( ref . 44894 ) .
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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 .
urn : lsid : biodiversity . org . au : afd . taxon : ad4ce6ca - 0e2f - 4d29 - 9617 - 26b23a9be30b
urn : lsid : biodiversity . org . au : afd . taxon : 48e069e4 - d923 - 45b1 - a8c0 - 107279c373f1
urn : lsid : biodiversity . org . au : afd . name : 434542
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format summary genbank genbank ( full ) fasta asn . 1 xml insdseq xml tinyseq xml feature table accession list gi list gff3
finds sub - sequences or patterns in the sequence and highlights the matching regions . the tool works with standard single letter nucleotide or protein codes including ambiguities and can match prosite patterns in protein sequences . more . . .
finds sub - sequence or patterns in the sequence and highlights the matching region . the tool works with standard single letter nucleotide or protein codes including ambiguities and can match prosite patterns in protein sequences . more . . .
a small pale greyish - brown gudgeon with a darker underside , with black anterior dorsal - fin spines and an orange patch on the fin , a large black blotch anteriorly on the base of the second dorsal fin and white spots on the fin , a reddish - orange anal fin , and a white margin on the dorsal , anal and pelvic fins . males are more densely covered in melanophores than females ; .
known only from the mitchell river in the kimberley region of western australia . the species inhabits still to moderately fast - flowing freshwater rocky streams and pools .
pale greyish - brown , pale purple ventrally on head , body darker ventrally ; scale pockets edged with melanophores and lines of melanophores on skin along margins of muscle segments ; males with dense melanophores scattered over body , often more so ventrally on caudal peduncle ; females with few scattered melanophores ; large median black blotch ventrally behind urogenital papilla connecting to thin black line on either side of anal fin base , lines joining behind anal fin to form thin median black stripe extending to anterior base of unsegmented caudal rays . first dorsal fin black before fourth spine , black extending onto and along distal tip of fin ; fin otherwise mostly clear ; second dorsal with large black blotch at base anteriorly , extending just anterior to third segmented ray , thin black distal margin ; often with broad , black stripe on distal upper third of fin ; rest of fin mostly clear ; anal fin mainly reddish orange with faint blotch near base anteriorly ; caudal fin pale grey with scattered melanophores . pectoral base with few scattered melanophores ; pectoral fin clear to grey . pelvic fins clear to white , with few elongate melanophores along lateral margins of rays in males .
the species is named for j . b . hutchins , former curator of fishes at the western australian museum , who collected the type specimens .
allen , g . r . 1982 . inland fishes of western australia . perth : western australian museum 86 pp . 6 figs 20 pls [ 61 , pl . 13 ( 7 ) , as new genus and species b ]
morgan , d . l . , allen , g . r . , pusey , b . j . & burrows , d . w . 2011 . a review of the freshwater fishes of the kimberley region of western australia . zootaxa 2816 : 1 - 64
allen , g . r . 1982 . inland fishes of western australia . perth : western australian museum 86 pp . 6 figs 20 pls [ 61 , pl . 13 ( 7 ) ] ( as new genus and species b )
allen , g . r . 1989 . freshwater fishes of australia . neptune , new jersey : t . f . h . publications 240 pp . , 63 pls [ 201 ]
allen , g . r . , midgley , s . h . & allen , m . 2002 . field guide to the freshwater fishes of australia . perth : western australian museum 394 pp . [ 302 ]
hoese , d . f . 2006 . eleotridae ( pp . 1596 - 1610 ) . in : beesley , p . l . & wells , a . ( eds ) 2006 . zoological catalogue of australia . volume 35 australia : abrs & csiro publishing parts 1 - 3 2178 pp .
urn : lsid : biodiversity . org . au : afd . taxon : b543ce97 - 963e - 48f2 - 82f7 - ea917623c872
urn : lsid : biodiversity . org . au : afd . taxon : 12f7654c - 734e - 4b95 - bc75 - 16fb722f293f
urn : lsid : biodiversity . org . au : afd . name : 328118
this is intended to provide a comprehensive up to date list of australian freshwater fishes . it includes : all species which are strictly freshwater , that is they never occur in estuaries or marine environments ; species which primarily occur in freshwater , but also occur in tidal reaches , or occasionally in estuaries , e . g . ,
ophisternon gutturale ( richardson 1845 ) multiple spp . ( 2 - 3 ? )


inglis drever ( 18 march 1999 – october 2009 ) was a champion racehorse trained in the north of england by howard johnson .

he was one of the most successful hurdle racing horses to date and won the world hurdle three times , bettered only by the former champion big buck 's .

he is seen by many as having been the greatest staying hurdler of all time .

inglis drever

inglis drever : ruled out for the remainder of this season . photograph : pa .
bbc sport | other sport . . . | horse racing | inglis drever eyes chepstow test
inglis drever’s owner graham wylie is now enjoying great success with ireland’s dominant force willie mullins .
but more than any , inglis drever was the horse who got me interested in racing .
triple world hurdle winner inglis drever was put down on friday after suffering a serious bout of colic .
racing is mourning a jumps legend after three - time world hurdle hero inglis drever died on friday morning .
inglis drever began his career on the flat for sir mark prescott , for whom he won four races .
and when inglis drever made it three , he joked :
i ' m a complete slapper .
three - time world hurdle winner , inglis drever , has been put down after a bout of colic .
inglis drever denied baracouda a third stayers ' crown with a famous success in the ladbrokes world hurdle at cheltenham today .
wylie ' s substantial and still growing investment in racing is one of inglis drever ' s legacies to the winter game .
3 times world hurdle winner inglis drever was sadly put to sleep this morning after suffering from a bad bout of colic .
inglis drever made history at the cheltenham festival last march when he became the first horse to win the world hurdle three times .
inglis drever will bid to maintain his status as champion stayer after being confirmed for the rescheduled totesport long walk hurdle on 27 december .
a new order is emerging in jump racing and it was suitably demonstrated as inglis drever saw off baracouda in the ladbrokes world hurdle .
massive repsect to inglis drever , a wonderful horse who won at the festival only to get beaten at aintree such was his way .
plans for inglis drever to return to cheltenham this march were scrapped after he failed to recover from an injury sustained at newbury last november .
inglis drever ' s trainer howard johnson said :
chepstow is a flat , left - handed track and that is what he likes .
inglis drever runs in to win the world hurdle for a second time in 2007 . photograph : tony marshall / empics sport / pa photos
but more than any horse - probably apart from lord transcend - inglis drever was the horse who got me seriously interested in racing .
johnson has had a number of top - class horses pass through his hands , but places inglis drever at the top of the tree .
it wasn’t his ability that we found so endearing . greater horses have come and gone that didn’t enjoy half the love and respect that inglis drever rightfully earned
very nice tribute bosranic . cant add much to that but to say inglis drever will be missed lots . condolences to all connected with this mighty racehorse .
staying power : thelaadbroke world hurdle ( stayers hurdle ) cheltenham : inglis drever and baracouda d . m . dent 15
x10
ltd ed 200 £59 giclee
but all the while lee was stoking up inglis drever ( 5 - 1 ) and the writing was on the wall for his rivals as he surged towards the last .
in the meantime , johnson hopes to send inglis drever back to the flat for the northumberland plate , a race with near - mythical status in the north - east .
but a disappointing workout at the weekend convinced them to abandon plans to send inglis drever back to the cheltenham festival , after which the decision was taken to retire him .
inglis drever made history at the cheltenham festival last year when he became the first horse to win the world hurdle three times , showing tremendous courage to overhaul kasbah bliss .
entries for the ladbrokes world hurdle at the cheltenham festival close this week but the list is set to be drawn up without howard johnson ' s triple winner inglis drever .
another endearing quality in the constitution of inglis drever was his tendency to hit a flat spot in his races - as evidenced by his world hurdle victories in 2005 and 2007 .
inglis drever was ridden at cheltenham by three different jockeys , the last of them being denis o’regan who said : “words can’t describe how much of a legend he truly was .
as inglis drever came to the last that day , finding himself in yet another battle , the outcome was inevitable . he was never going to come second in a fight , but still the cheltenham faithful , who cast aside their own selfish motivations for his success , roared inglis drever up the hill to a victory that would catapult him into deserved immortality .
the big players were held up towards the rear , however , with baracouda as usual looking to be cruising as the others , including inglis drever , started to come under pressure .
inglis drever made history at the festival last march when he became the first horse to win the world hurdle three times and wylie credits the horse with getting him seriously involved in racing .
on friday howard johnson’s yard , and racing in general , were mourning the death of inglis drever , the only horse to win the ladbrokes world hurdle at the cheltenham festival three times .
edward gillespie , the managing director of cheltenham , said yesterday that the course will take time to consider the best way to commemorate inglis drever . a statue , however , does not seem likely .
inglis drever was only six years old when he beat the great baracouda in this race two years ago , and was sidelined with an injury when my way de solzen took the prize 12 months ago .
really sad news his third ( ? ) win in the stayers hurdle at cheltenham with his devoted groom cheering him on was a magic moment . thanks for the memories inglis drever and rest in peace .
perhaps the most piquant aspect to the retirement of inglis drever , announced yesterday , is that he should so narrowly have missed the opportunity to measure the pretensions of his obvious heir . there is no doubting the regal bearing of punchestowns , whose accession as the new monarch of staying hurdlers could well be hastened at cheltenham on saturday . thus far , indeed , he has gone about things with rather more panache than inglis drever . but it would be unusual for any horse to dominate this exacting discipline without also disclosing a coarser grain , and inglis drever would never have surrendered his crown without an almighty scrap .
wylie does not like to name horses in advance of particular meetings or races -
it jinxes them ,
he maintains - so we won’t mention the name of possible champion hurdle entry inglis drever .
in a move away from the sadness that normally heralds the end of a career , trainer howard johnson described himself as
over the moon
after the retirement of triple world hurdle winner inglis drever today .
meanwhile , the pair at the top of the pecking order to take over the staying hurdle mantle from inglis drever are set to clash in the cleeve hurdle on festival trials day at cheltenham a week on saturday .
inglis drever , the only horse to win the staying hurdlers ' championship at cheltenham three times , will not get the chance to make it four , after he was retired from racing yesterday by graham wylie and howard johnson , his owner and trainer . ladbrokes , the sponsors of the world ( formerly stayers ' ) hurdle , said later that they will refund all ante - post bets on inglis drever for this year ' s race .
when he hears the roar of the crowd , though , inglis drever concentrates hard and has a finishing surge that is difficult to resist . paddy brennan powered him towards the lead on the run down the hill and had two lengths to spare at the final flight . mighty man then summoned a strong challenge as they galloped up the hill but inglis drever was never going to yield the advantage and held on by three - quarters of a length .
inglis drever , won 17 races from 35 starts earning nearly £800 , 000 in prize - money . thirteen of his victories came over hurdles after he was recruited from sir mark prescott’s flat stable for 110 , 000 guineas .
although he was to make his mark as a staying hurdler , inglis drever was kept to two miles for most of the 2004 / 05 season and even won a recognised trial for the champion hurdle , wincanton ' s kingwell hurdle . but there was a strong field for that year ' s champion , with hardy eustace , harchibald and brave inca in their prime , so inglis drever was stepped up in trip for the three - mile world hurdle .
racing : three - time world hurdle hero inglis drever has died . the howard johnson - trained 10 - year - old , who retired from racing in january , was put down this morning after suffering a fatal bout of colic .
inglis drever won 17 of 35 career outings , amassing almost £800 , 000 in prize money . trained on the flat by sir mark prescott , he won four handicaps but was only 11th behind landing light when fancied for the 2003 cesarewitch .
wylie , who owned three - time world hurdle winner inglis drever , admitted he knew that striking article had undergone a neurectomy but , like johnson , claimed to be unaware that running a horse after the treatment breached the rules of racing .
inglis drever , who racked up a hat - trick of victories in the world hurdle at the cheltenham festival , has been retired this morning . the 10 - year - old failed to recover from a leg injury sustained at newbury in november .
inglis drever , a horse who failed three veterinary inspections before being sold privately to graham and andrea wylie , galloped his way into festival history yesterday by winning the ladbrokes world hurdle a record third time to the wild cheering of an appreciative cheltenham crowd .
but johnson , best known for his handling of three - time world hurdle winner inglis drever and horses like direct route who was denied the 2000 queen mother champion chase by a whisker , has no intention of making what would be a dramatic comeback .
inglis drever was pulled up after suffering a hock injury on his seasonal debut at newbury last november , but graham wylie ' s gelding was given time to recover and a positive x - ray last week appeared to give hope of a fourth cheltenham festival victory .
the bookmakers ladbrokes and paddy power announced they would refund all single bets on inglis drever for this year ' s world hurdle , as a goodwill gesture . the horse had been as short as 4 - 1 in the ante - post market before his injury .
formal presentations of the season ' s order of merit also took place with john webb , owner of overall winner and top hurdler royal shakespeare receiving a cheque for £250 , 000 from andrea and graham wylie , owners of last year ' s inaugural winner inglis drever .
the 2005 , 2007 and 2008 world hurdle winner was pulled up in the long distance hurdle at newbury in november and while johnson doesn ' t want inglis drever to go out of the game with a whimper , he would find it hard to suffer his champion being heckled at his favourite racecourse .
the trainer has little else to regret in his supervision of a horse whose influence on his own career can scarcely be exaggerated . for it was inglis drever who convinced graham wylie that racing could become a conduit between his wealth and pleasure , and that johnson was the man to trust with the project .
hopes that inglis drever might bid for a fourth win at the festival were raised earlier this month , when a scan on a leg injury , sustained at newbury in november , appeared clear . however , the 10 - year - old worked poorly at the weekend and the decision to retire him followed soon afterwards .
in 14 runs over three miles , inglis drever failed to finish in the first three on only two occasions , when falling at chepstow and when pulled up at newbury on his final start . he returned with an injury on both occasions , the most recent of which has brought his fine career to an end .
but despite the dark clouds , tidal bay , the winner of tuesday ' s arkle trophy for rider denis o ' regan and the same owner - trainer combination , and inglis drever , who displayed typical courage for o ' regan in one of the most thrilling finishes of the week , both secured major prizes for their connections .
more than any horse , probably apart from lord transcend , inglis drever was the horse who got me seriously interested in racing ,
wylie said yesterday .
people might think cheltenham gave me my favourite memories of him , but it was actually a day at haydock in 2005 when both him and lord transcend won .
as it is , the substance of his achievements can be represented at the festival in march only by kasbah bliss , the french horse who made him work so hard for that unprecedented third success in the ladbrokes world hurdle last year . in the meantime , inglis drever will abide in the esteem of all and the affection of many .
inglis drever instilled in me such a passion ,
wylie said yesterday .
more than any horse – probably apart from lord transcend – he was the one who got me seriously interested in racing . people might think cheltenham gave me my favourite memories of him , but it was a day at haydock in 2005 when both he and lord transcend won .
inglis drever had a career record of 17 wins from 35 starts and his appetite for a battle , particularly on the climb to the line at cheltenham , made him hugely popular with punters . he was famous , too , for hitting a
flat spot
in a race when he appeared to be in trouble , only to run on strongly in the closing stages .
at around evens , however , this is surely one favourite that has to be taken on . it is time for a fresh face on the stayers ' scene , and inglis drever looks the one . at 10 , baracouda is not getting any better . iris ' s gift last year showed that the french horse is beatable , though that is not to denigrate his tremendous record .
since then , wylie’s colours have been carried by some of the best recruits to national hunt which racing has seen in many years . inglis drever , valley henry , royal rosa , arcalis , and chivalry are but five of the powerful string owned by the modest man whose quiet tones betray his upbringing in the north - east of england , but which also mask his true origins .
after hitting his usual flat spot in the middle stages of the race , inglis drever knuckled down to an intense battle with the francois doumen - trained kasbah bliss , which he won by calling on all stamina reserves . he jumped into the lead at the final flight and held off kasbah bliss by a length , with a gap of seven lengths back to the game kazal in third .
prevented by a tendon injury from defending his crown , inglis drever won it back in 2007 , outbattling mighty man by three - quarters of a length . his pattern of finishing strongly after appearing to struggle was now well established and he again made his supporters wait when landing his third world hurdle last year , getting past kasbah bliss only on the run - in for what turned out to be his final victory .
if there were a group of horses ahead of him turning in , as far as he was concerned they all had bullseyes on their backsides and the drever would pop his eyes out of their sockets and take aim .
black jack ketchum had a question to answer as he set off for the ladbrokes world hurdle here yesterday , and if the reply was not one that many punters wanted to hear , at least he did not keep them waiting . he barely took off at the third flight and suffered the first fall of his career , which left the way clear for inglis drever to win the stayers ' championship event for the second time in three years in his familiar , curmudgeonly style .
inglis drever can be a difficult horse to warm to , as punters tend to prefer their horses to look like potential winners at every stage of a race . howard johnson ' s runner , by contrast , often needs persuasion to keep him interested in the early part of a race , and he was on and off the bit throughout the first two miles yesterday . there were several sloppy jumps on the first circuit too , as if the eight - year - old could not quite be bothered .
devastating news . inglis was a proper nh legend and he will live long in the memory of all those who had the pleasure of seeing him race . huge condolences to connections and also his die - hard fans , of which he had many .
inglis drever introduced his owner as a man agreeably lacking airs , while johnson ' s bluff geordie cadences also enriched the tale . then there was the horse ' s groom , ginni wright , whose emotional celebrations were shared by many who learned of her battles with cancer . but the central character was always the horse . his habit of hitting a flat spot before gathering momentum implied a latent vulnerability – a pleasing sense that it did not come easily , but that he would always do his utmost .
inglis drever left the door ajar by apparently recovering from his newbury injury , but johnson explained :
we put shoes on him the other day , rode him out , and he didn ' t even want the jockey to get on him . he was telling me he ' d had enough . i think i ' ve done the right thing for the public . i didn ' t want to go to cheltenham against younger horses and be pulled up or get beat 20 lengths . one day we might find another like him . but we ' ll struggle .
graham wylie ' s personal favourite is lord transcend , the horse who managed to get him involved in the racing game , but inglis drever is now in the same bracket .
he ' s a street fighter who never knows when he is beaten and he loves the attention he gets in the winner ' s enclosure ,
wylie said .
maybe one day i ' ll realise what he ' s done , but it is all so emotional now . i just sat quietly in the stand and watched and watched , and he had so many traffic problems . but when he came over the last , he was too good for them .
connections of inglis drever ( 3 . 15 ) have always considered him a stayer in the making , and that certainly looked the case when he finished strongly to take second place behind fundamentalist in last season ' s sunalliance hurdle . nothing was in his favour in the kingwell hurdle at wincanton last time out - a right - hand course and having to make his own running - but he still won by five lengths . at odds of 7 - 1 he is the each - way value . rule supreme , who would have beaten baracouda and crystal d ' ainay in a few more strides at windsor , is another reason to be wary of taking a short price about the favourite .
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“they took him away yesterday and from what i can understand they couldn’t do anything with him , ” johnson said .
“graham ( wylie , owner ) rung me yesterday morning to say he was thrashing on the ground so they got the vets up there to look at him and they rushed him straight in ( for surgery ) .
“they put him on a drip overnight and they rang mr wylie back this morning to say his heart rate was over 70 and it should 33 , so they have had to put him down on humane grounds . ”
“without a doubt he’d be number one when it comes to horses i’ve trained , ” added johnson .
“he went to graham’s for a good retirement and i haven’t actually seen him since he left . it is just one of those things . ”
his indomitable spirit and appetite for a scrap made him one of the most popular national hunt horses in training .
the brilliant gelding signed off with 17 wins - 12 of which came at graded level - from 35 starts and amassed nearly €880 , 000 in prize - money .
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having his first try at this three - mile trip , the six - year - old surged to the front at the last and was far too strong for baracouda up the hill .
the win continued a great week for trainer howard johnson , owner graham wylie and jockey graham lee , who were landing their third prize following arcalis and no refuge .
baracouda ' s pacemaker knife edge took the field along at a fair old clip in the early stages , with emotional moment , westender and yogi also prominent .
westender led from some way out as knife edge gave way , and he was still there two out , with rule supreme hot on his heels and baracouda set to pounce .
baracouda , the 6 - 5 favourite , gave brave chase but was three lengths away at the line , with a further three - quarters of a length back to rule supreme .
he didn ' t travel great , he didn ' t jump great but he ' s got a big heart and in the end i think i got there a bit too soon - but i wasn ' t going to stop when i got him going ,
lee told channel 4 racing .
i was forced to track baracouda for a while because i wasn ' t travelling well , but i ' m so happy for the horse . to be narrowly beaten in the sunalliance last year and now this , i ' m so chuffed for him . he ' s tough .
johnson said :
i never usually shake , but watching that race today made me shake a little bit because i love this horse . he ' s a proper little jack russell terrier .
he jumped as straight as a die today . graham gave him a lovely ride . all credit to the staff at home and the girl who does him . it ' s great .
it ' s working out well with graham and andrea ( wylie ) . let ' s hope it lasts .
i knew he ' d come up the hill . we ' ve got one that ' s high and steep at home . when he ' s on form he goes up our hill great .
that ' s the secret about training our horses - going up steep hills . it seems to get their wind right and helps them be fit and healthy .
wylie added :
it ' s all down to howard and to graham . howard ' s spent a lot of time getting the horses ready . he ' s done a brilliant job .
he was really a splendid second ,
he said .
tony ( mccoy ) was impeccable , he gave him the perfect ride . we were just beaten by a better horse and we have to respect that .
he may go to aintree . we will certainly enter him , but it is too early to say whether he will go over fences or not .
maybe as he is getting older he does not have the speed he had in the past , but he has run a good race .
owner jp mcmanus added :
we were just beaten by a better horse . he ran his heart out . all credit to francois , he had him ready to run the race of his life .
rule supreme ' s trainer willie mullins said :
he ' s had a hard race . i ' m happy with the horse . there are no excuses really .
i am a bit disappointed as he was in the form of his life , but he was beaten fair and square , and perhaps that ' s as good as he is .
he ' ll definitely go to punchestown now and then we ' ll go for the french champion hurdle again .
earlier , thisthatandtother denied fondmort in a thrilling finish to the new daily telegraph festival trophy chase .
fondmort held a slight advantage after jumping the last , but the paul nicholls - trained thisthatandtother ( 9 - 2 ) , ridden by ruby walsh , battled on just the better up the hill and wore down nicky henderson ' s charge in the last 50 yards to win by half a length .
irish raider rathgar beau , who finished third , lost his winning chance with a mistake at the final fence when bang in contention .
and even earlier , king harald put up tremendous front - running performance under mattie batchelor to land the jewson novices ' handicap chase on the third day of the cheltenham festival .
the mark bradstock - trained seven - year - old had four lengths to spare at the line from lacdoudal after an incident - packed affair .
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staying hurdlers often endure much longer than the two - milers and if he stays sound , the 6 - 1 offered by ladbrokes for the race next year could prove to be a very fair price .
he tends to come on and off the bridle ,
johnson said ,
but i said to paddy that as soon as you come down that hill and get a bit of light , he ' ll pick up .
he will never go chasing now , as you don ' t need to send a dual winner of this race over fences . it ' s always been an ambition of mine to have a runner in the northumberland plate , so i may run him in that and then turn him out .
there was little to be said about black jack ketchum , though jonjo o ' neill , his trainer , confirmed that the horse seemed to have emerged unscathed .
life goes on ,
he said .
the horse is ok , and we ' ll just have to try again .
taranis was a tenacious winner of the ryanair chase after being left in front by the fall of crozan at the fourth - last , which was probably a couple of furlongs further from the finish than ruby walsh would have chosen . the line eventually arrived just in time as our vic staged an extraordinary late rally , failing only by a neck , having looked to be one of the first beaten three - out .
he ' s a horse that tends to idle in front and i was a bit concerned when crozan fell , but ruby has given him a great ride ,
paul nicholls , who was welcoming back his 100th winner of the season , said .
when they looked to be catching him at the finish it was more to do with him idling than the others making ground . he ' s only six and i think he may well get three miles next season .
previous form at the festival usually proves significant and idole first , who took the coral cup over hurdles two years ago , added a victory over fences in the racing post plate .
ferdy murphy , meanwhile , saddled his second winner of the week when l ' antartique took the jewson novice handicap chase at 20 - 1 , giving anyone who played up their winnings from the 50 - 1 chance joes edge on tuesday a 1070 - 1 double .
it ' s 27 years since anaglog ' s daughter won here [ in the arkle trophy ] ,
tony durkan , the winning owner , said ,
and she was beaten in the champion chase the next year . this is a lovely horse and we hope that he hope that he will go on to better things .
with hardy eustace successfully defending his crown , and moscow flyer regaining his , the pattern is set for a repeat performance by baracouda in this afternoon ' s ladbrokes world hurdle , the race he won two years in succession before relinquishing his title 12 months ago .
coming into last year ' s race baracouda had won the long walk hurdle and newbury ' s long distance hurdle by a total of 37 lengths , but this season he has won the same races only narrowly from crystal d ' ainay . admittedly he had crystal d ' ainay a long way behind him in third place here last year , but alan king ' s gelding faced a stiff task then as a five - year - old .
2 . 00 jewson h ' cap chase : one of the new races at this year ' s festival , this has been the target for liverpool echo , whose trainer , henry daly , has a fine record in handicaps here . liverpool echo was becoming somewhat expensive to follow until he beat copsale lad convincingly at kempton . that form is working out well .
2 . 35 festival trophy chase : our vic runs here rather than in the gold cup and this 2m5f is more his trip , because he seemed to get outstayed when third in last season ' s sun - alliance chase . though given time to get over his final - fence fall in the bonusprint chase here in december , he has yet to confirm his high home rating on the track . at 10 - 1 , fondmort makes more appeal . best when fresh , nicky henderson ' s nine - year - old has a fine record at cheltenham and last season ran a highly creditable third under top weight in the mildmay of flete . he now meets our vic and thisthatandtother - second in the bonusprint - on more favourable terms than when only seventh in that race , when henderson ' s horses were out of sorts .
4 . 00 mildmay of flete chase : martin pipe has won four of the last seven runnings of this handicap , and he looks to have prepared polar red for a second crack after his creditable sixth behind tikram 12 months ago . polar red , now 3lb lower , has been freshened up by a mid - season break and two recent spins over hurdles .
4 . 40 national hunt chase : stormez and celestial gold have finished second for the pipe stable here in the last two seasons . sixo could go one better , having run well against the useful novice distant thunder at newbury . by roselier , a strong influence for stamina , sixo should be in his element over this marathon trip .
5 . 20 pertemps hurdle final : with some solid course form to his name , the dark lord should be a leading player . he was staying on strongly at the finish of the sandown handicap hurdle last time and his stable is in better form now .
the decision was made following a disappointing workout at the weekend by the 10 - year - old , who notched up 17 wins from 35 starts , amassing almost £800 , 000 in prize money in the process .
but rather than wallowing in despair at the decision , johnson breathed a sigh of relief .
i ' m over the moon i ' ve retired him ,
he said .
we put shoes on him the other day , we rode him out and he didn ' t even want the jockey to get on him .
he was telling me he ' d had enough , so he ' s gone into retirement . i think i ' ve done the right thing for everybody .
i didn ' t want to go to cheltenham against younger horses and either be pulled up or get beat 20 lengths - that wasn ' t for me .
he said :
we decided on saturday that he wasn ' t interested any more . at the end of the day , he has won three world hurdles and has been one of the greatest horses .
howard says to let him chill out and he ' ll have a box next to one of our broodmares . i ' m sure he ' ll have a long and happy retirement .
the howard johnson - trained 10 - year - old , who retired from racing in january , had to be put down after suffering a fatal bout of colic .
johnson said :
they took him away yesterday and from what i can understand they couldn ' t do anything with him .
graham ( wylie , owner ) rung me yesterday morning to say he was thrashing on the ground so they got the vets up there to look at him and they rushed him straight in ( for surgery ) .
they put him on a drip overnight and they rang mr wylie back this morning to say his heart rate was over 70 and it should be 33 , so they have had to put him down on humane grounds .
johnson added :
without a doubt he ' d be number one when it comes to horses i ' ve trained .
he went to graham ' s for a good retirement - it is just one of those things .
the brilliant gelding signed off with 17 wins - 12 of which came at graded level - from 35 starts and amassed nearly £800 , 000 in prize - money .
he was transferred to johnson ' s yard in 2003 , when he made a winning debut for his new team in november .
although the star hurdler will be best remembered for his exploits at cheltenham , he also won the long distance hurdle at newbury three times ( 2005 - 07 ) - in addition to big - race triumphs at sandown , warwick , wincanton , haydock and wetherby .
johnson said :
he ' s been a fantastic servant to me and the yard and has kept me going since we bought him off sir mark .
he won three world hurdles and was the first horse to win the order of merit .
you could always tell when he was right and he was a peach to train .
he wouldn ' t hurt a butterfly and you always knew he would come up hills because as soon as he saw our hills at home he used to fly .
it ' s very upsetting and i ' ve said to mr wylie that we ' ll get him cremated and get him back into what i call the ' millionaire ' s field ' in front of our house .
he used to love it in there , and spent his summers in there every year since he arrived , so i think we should bury his ashes in there .
there was no sign of that trademark quirk at prestbury park last march , however , as he powered up the hill to record a no - nonsense verdict over kasbah bliss .
defeat at aintree - never his happiest hunting ground - followed last april , after which he was sent to newbury for his seasonal debut .
but denis o ' regan ' s mount suffered a hock injury and was pulled up after four out .
although there were hopes he had made a full recovery , johnson realised the game was up in january .
the jockey was in the saddle when the horse won his third ladbrokes world hurdle and described him as the best horse he will ever ride .
it ' s shocking news . i won the world hurdle on him and the long distance hurdle as well ,
he said .
he was a top - class hurdler and it ' s a shame he ' s died so young , but he ' s a horse that will stick in the memory for a long time .
he ' s the best horse i ' ve ridden and will probably be the best horse i ' ll ever ride .
he ' s brought my career to the fore and it ' s a sad ending to a great career for that horse .
graham lee ' s association with the horse lasted from november 2003 to december 2005 .
their seven victories included a first world hurdle in 2005 when the pair dethroned the french champion baracouda .
lee said :
when you are a jockey you want to have big rides in big races and he was a very , very good horse .
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the ten - year - old , who retired in january this year , underwent surgery on thursday after being found lying in a field at owner graham wylie’s stud at humshaugh in northumberland , but his condition worsened on friday morning .
at kelso on saturday after silk drum had made a winning start over fences for graham wyile , the owner said : “yesterday was very difficult . i walk around the house and there are photos everywhere , all his trophies and dvds .
“i have very happy memories , but on a day like today it’s very distressing . ”
johnson was also distressed saying : “i’ve not seen the lovely horse since he left here for his retirement ; i will never have one like this to win three world hurdle and an order of merit . he’s been my pride and joy and i will never get another to replace him . ”
“he had a fantastic racing career and will be remembered for a very long time . i was very lucky to ride him and his final world hurdle win was a wonderful performance . ”
staying on a sombre theme , barry keniry faces a lengthy spell on the sidelines due to a broken leg at kelso on saturday .
the leyburn - based jockey was riding the shy man for the george moore stable when he came crashing down from fences from the finish .
yesterday he underwent surgery on the injured leg in the boarders hospital , and it remains to be seen just how long the irishman will be out of action .
this week’s local racing kicks off today at pontefract where there is an eight race card due of at 2 . 10pm .
the highlight race at the west yorkshire comes up at 3 . 40pm , a listed race over the one mile trip .
among the 11 runners is hot prospect , who looks a very interesting colt . trained by michael jarvis , he was a very impressive winner at sandown last time out and with kieren fallon booked he could just turn out to be a very hot prospect .
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the race was abandoned at windsor last weekend and so switches to chepstow as part of the welsh national meeting .
he has come back from windsor in good order and i will definitely put him in - he won ' t mind the soft ground .
officials at the bhb have also decided to revert to the original entry system , which will allow four - times winner of the race baracouda to run , after he was initially missed out of the entry stage due to a clerical error .
however , francois doumen has not yet decided whether baracouda will take part in the rescheduled event .
he said :
i will need to think about it and will make a decision later in the week .
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the magnitude of this triumph can be viewed against the backdrop of depression that descended on howard johnson ' s yard in the build - up to jump racing ' s most important meeting when a collection of his stable stars were coughing and breaking blood vessels .
o ' regan declared :
what a horse . he just took off . howard said beforehand that he wouldn ' t want to get there before the last . i was a bit down at the last but he jumped well , and he did the rest himself .
johnson ' s comment that
this is near enough the greatest training performance of my life
is indeed accurate . the winner , who had been bought out of sir mark prescott ' s newmarket yard off the flat to win the northumberland plate , was purchased unvetted . a persistent leg injury had been his problem .
johnson said suggestions of retirement had been put on hold .
he ' s only nine , but i don ' t want anything to happen to him ,
johnson said .
he might go to aintree , he might come back here next year , but there are not many races left in him .
owner trevor hemmings landed the first two races , the peter o ' sullevan national hunt chase with the alan king - trained old benny , and the royal & sunalliance chase with the jonjo o ' neill - trained albertas run , who was a first winner at this year ' s festival for tony mccoy .
our vic , who chased home kauto star in the king george at kempton park on boxing day , finally enjoyed his moment of glory at the highest level when taking the ryanair chase , a race in which he finished second last year . first - time blinkers worked a treat as our vic blazed a trail in front .
trainer david pipe said :
the blinkers sharpened him up a bit and timmy got some fantastic jumps out of him .
we decided on saturday that he wasn ' t really interested in his racing any more ,
wylie said yesterday .
we always knew that he would tell us when he wanted to be retired . that day has now come .
obviously we would like to do something appropriate in due course ,
gillespie said .
at this stage we are just delighted that he has been retired fit and healthy .
we have just four statues [ of horses ] at the moment : arkle , dawn run , golden miller and , most recently , best mate . there are no hurdlers , though istabraq is one about whom the question has been asked , but it has just not been possible to achieve . we have plenty of room but statues are incredibly expensive .
cheltenham will stage its last meeting before the festival this saturday and , though the track lost a meeting to waterlogging in december , simon claisse , the clerk of the course , is confident that this card will proceed as planned .
the rain we are forecast is due to arrive between now and friday morning , after which it should be basically dry through to racing ,
he said .
it seems certain that great leighs racecourse in essex will remain closed for at least another week , however . the british horseracing authority said yesterday that it has yet to receive a formal application for a racecourse licence from deloitte , the administrator trying to rescue the track .
with entries for next thursday ' s meeting due to close tomorrow , that card now seems sure to be staged elsewhere .
his trainer , howard johnson , had indicated recently that an x - ray showed the injury to be clearing . johnson said at that stage that the horse might return to the racecourse but the horse ' s demeanour has convinced him otherewise .
we decided on saturday that he wasn ' t really interested in his racing any more ,
said the horse ' s owner , graham wylie .
we always knew that he would tell us when he wanted to be retired . that day has now come .
i ' m over the moon i ' ve retired him ,
said johnson .
i ' m happier in myself now . we put shoes on him the other day , we rode him out and he didn ' t even want the jockey to get on him . he was telling me he ' d had enough . we gave him a jog along the road and i wasn ' t happy , so he ' s gone into retirement .
the following month , he joined johnson ' s yard and made his hurdling debut at aintree , winning easily . he remained unbeaten over hurdles until that season ' s cheltenham festival , when he was edged out by fundamentalist in the race known then as the royal & sunalliance novices hurdle .
it proved the making of him . his performance that day was a revelation as , after appearing beaten half a mile from home , he bounded up the hill , finishing three lengths clear of baracouda , himself a dual winner of the race .
i just hope sir mark will find me another one - he ' s on the case ,
added johnson .
i think i ' ve done the right thing for the public and everybody . i don ' t want to go to cheltenham against younger horses and either be pulled up or get beat 20 lengths - that wasn ' t for me .
prescott added his own tribute , saying :
he ' s one of those lovely horses who ' s done everybody well all his life . he was a very nice yearling - any fool could have picked him . he was a very nice horse for us and made a jolly nice price for us and he gave mr wylie a marvellous introduction into racehorse ownership .
time has been called on the 10 - year - old ' s illustrious career after he failed to recover from an injury sustained at newbury in november .


orthotylus marginalis is a species of stinkbugs from miridae family that can be found throughout europe ( except for liechtenstein and various european islands ) .

orthotylus marginalis

in great britain and / or ireland : foodplant / feeds on orthotylus marginalis feeds on salix caprea foodplant / feeds on orthotylus marginalis feeds on salix cinerea ssp . oleifolia foodplant / feeds on orthotylus marginalis feeds on salix foodplant / feeds on orthotylus marginalis feeds on alnus glutinosa foodplant / feeds on orthotylus marginalis feeds on malus foodplant / feeds on orthotylus marginalis feeds on ribes foodplant / feeds on orthotylus marginalis feeds on prunus spinosa animal / predator orthotylus marginalis is predator of tetranychus urticae animal / predator orthotylus marginalis is predator of aphidoidea animal / predator orthotylus marginalis is predator of insecta
leaf beetles on willows : orthotylus marginalis
. department of ecology , swedish university of agricultural sciences .
orthotylus marginalis population density was estimated in 15 of the gray willow stands in june 1999–2011 and in the two other stands in 1999–2010 and 1999–2005 . all insects on the upper 35 cm of each current - year shoot were dislodged into a white plastic container . orthotylus marginalis nymphs were counted and then re - released . the number of shoots sampled was proportional to the size of the stand . for a detailed description of the sampling method see dalin ( 2006 ) .
the adult orthotylus flavosparsus is normally about 4 millimetres ( 0 . 16 in ) in length , and green in colour .
population densities related to leaf nitrogen status . predicted mean number of orthotylus marginalis individuals per 35 cm shoot ( based on data from 17 gray willow stands collected over 13 years ) related to measured spad values ( relative difference in leaf nitrogen concentration— lower horizontal scale ) and to the predicted corresponding leaf nitrogen concentration ( upper horizontal scale )
orthotylus flavosparsus is a species of plant - eating bug in the miridae family , which is found everywhere in europe except for albania and iceland . it was introduced to north america .
predation rate related to leaf nitrogen status . number of leaf beetle eggs consumed by orthotylus marginalis ( day × mg dry weight ) −1 related to leaf nitrogen status under three nitrogen treatment levels ( n = 37 ) . predation rates differed between all treatments at p < 0 . 01 . box plots show medians with the first and third quartile and 95 % confidence interval of the median
total numbers of o . marginalis nymphs surviving to the adult stage under the 1 . 4 , 8 . 4 and 15 . 4 mg n week −1 nitrogen treatments were : 6 in all cases with prey absent ; and 15 , 10 and 13 , respectively , with prey present . orthotylus marginalis survival was clearly higher on plants in the presence of prey ( f 1 , 72 = 22 . 41 , p < 0 . 001 ) , but there was no difference in survival on plants under different nitrogen treatments ( f 2 , 72 = 1 . 50 , p = 0 . 23 ) . however , these results should be treated cautiously as various mortality factors , such as parasitization and molting failure , were not necessarily related to leaf nitrogen status . mortality due to parasitization was 16 % for o . marginalis with no prey and 11 % with prey present .
performance related to leaf nitrogen status . dry weight ( mg ) of orthotylus marginalis adults in the a absence ( n = 18 ) and b presence of prey ( n = 38 ) on plants exposed to different nitrogen treatments . performance differed between all nitrogen treatments in the absence of prey at p < 0 . 01 , but there was no difference between any of the treatments in the presence of prey . box plots show medians with the first and third quartile and 95 % confidence interval of the median
population variability related to leaf nitrogen status . variability ( cv = coefficient of variation ) in population density of orthotylus marginalis related to measured spad values ( relative difference in leaf nitrogen concentration— lower horizontal scale ) and to the predicted corresponding leaf nitrogen concentration ( upper horizontal scale ) . solid and dotted lines indicate model predictions and standard error of predicted means ( r 2 = 0 . 53 ) , respectively . open circles are data points used in the model . data points indicated by crosses were not considered in the model . population variability was calculated from data collected in 15 stands over 10 years . a small cv represents high stability between years
two new records of the subfamily orthotylinae ( heteroptera : miridae : orthotylinae ) , zanchius tarasovi kerzhner , 1988 and orthotylus bilineatus ( fallén , 1807 ) , are reported for the first time to the korean fauna . the genus zanchius is also first recorded from the korean peninsula . redescriptions of genitalia , diagnoses of each species and genus , and biological notes are presented with the photographs and illustrations .
the effects of leaf nitrogen status on performance and predation rate in the greenhouse experiment were examined using two glmms . response variables in the two performance models were o . marginalis survival ( binomial distribution , logit link ) and adult dry weight ( µg ) ( poisson distribution , log link ) . the response variable in the predation model was total egg consumption by o . marginalis ( poisson distribution , log link ) using an offset for development time ( days ) and adult dry weight ( mg ) . over - dispersion was accounted for using quasi - models . fixed effects in both performance models were nitrogen treatment and the presence / absence of prey , and in the predation model nitrogen treatment . we incorporated a variance structure in the omnivore performance models to account for differences in residual spread between groups with and without prey . gray willow clone was included as a random effect in all models .
altogether 90 o . marginalis nymphs were collected for the greenhouse experiment from the same gray willow stand . all individuals were collected simultaneously , on the first day that first stage nymphs appeared in late may . the nymphs were randomly assigned to plants under different nitrogen treatments and plants with or without prey : 45 to plants with prey and 45 to plants without prey . perforated plastic bags ( 0 . 5 mm diameter perforations , baumann saatzuchtbedarf , germany ) sealed at the base of the pot , prevented mirids from escaping and stopped potential prey from accessing the plants .
the relationship between o . marginalis population density and gray willow leaf nitrogen status in the field study was examined using a generalized linear mixed model ( glmm ) ( poisson distribution , log link function ) with an offset for number of samples in each stand . the fixed effects were the spad values recorded in june , sampling year and stand area . by treating gray willow stand as a random effect , the model intercept was allowed to vary between stands and between years within stands . temporal autocorrelation was accounted for by introducing an autoregressive first - order structure , with observation year nested within willow stand .
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is one of the larger species and is found on willows and sallows . the upper surface is covered in dense pale hairs , and the membrane veins are green . the 1st antennal segment is brownish .
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the distribution in this summary table is based on all the information available . when several references are cited , they may give conflicting information on the status . further details may be available for individual references in the distribution table details section which can be selected by going to generate report .
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are 6 millimetres ( 0 . 24 in ) long , and are green coloured . their upper surface is covered with dense pale hairs , with brownish
kerzhner i . m . , josifov m . ( 1999 ) .
family miridae
. in aukema , berend and rieger , christian . catalogue of the heteroptera of the palaearctic region . 3 , cimicomorpha ii . amsterdam : netherlands entomological society . pp . 1–577 , pages 253 & 263 . isbn 978 - 90 - 71912 - 19 - 1 .
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european union funding : for a one - year period ( 2017 - 12 - 16 to 2018 - 12 - 15 ) , eppo has been awarded an eu grant for the further development of the eppo code system ( agreement nb : sante / 2017 / gs / eppo / s12 . 768842 ) . the eu commission is not responsible for any use that may be made of the information from this project subsequently included in the eppo global database .
adults are 6 millimetres ( 0 . 24 in ) long , and are green coloured . their upper surface is covered with dense pale hairs , with brownish antennas .
the species members feed on alder , apple trees , currant , sloe , sallow , and willows . adults feed on aphididae , carabidae , and psyllidae . in some cases , they also feed on plants of pear trees , causing the pears to have stoney pits , among other damages .
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the online version of this article ( doi : 10 . 1007 / s00442 - 016 - 3742 - y ) contains supplementary material , which is available to authorized users .
plant traits , such as nutrient status , morphology and secondary metabolites affect herbivore performance , long - term population dynamics and community structure and can even cascade across trophic levels to mediate trophic interactions ( price et al . 1980 ; underwood and rausher 2000 ; harvey et al . 2003 ; dalin and björkman 2003 ; kagata and ohgushi 2006 ; bukovinszky et al . 2008 ; carmona et al . 2011 ; volf et al . 2015 ) . the interactions between plant - feeding omnivores and their herbivore prey can be mediated by plant traits ( 1 ) via density effects and ( 2 ) by altering the omnivores’ trophic status ( behavioral effects ) ( agrawal et al . 1999 ; eubanks and denno 1999 , 2000 ; eubanks 2005 ) .
omnivory and trophic behavior is determined by evolutionary history , driven by resource abundance and quality to optimize nutritional needs ( agrawal et al . 1999 ; coll and guershon 2002 ; eubanks et al . 2003 ) . plant feeding allows for survival of small omnivore nymph stages and for persistence during periods of rapidly declining prey densities ( naranjo and gibson 1996 ; wheeler 2001 ; coll and guershon 2002 ; eubanks and styrsky 2005 ) . feeding on high - quality host plants can , thus , strengthen the buffering effect of this alternative resource , which should increase and stabilize omnivore population density and enhance the long - term predation pressure ( density effect ) . high plant quality can , however , also increase the omnivores’ relative consumption of plant resources and reduce the average per capita predation rate ( behavioral effect ) ( eubanks and denno 2000 ; coll and guershon 2002 ) .
nitrogen concentration is a characteristic of plant nutrient status with a potentially strong direct effect on omnivore performance in the absence of prey , which is also relevant for comparing the relative importance of resources at different trophic levels ( eubanks and denno 1999 , 2000 ; fagan et al . 2002 ; denno and fagan 2003 ; matsumura et al . 2004 ) . herbivore population density commonly increases on nitrogen enriched plants ( mattson 1980 ; awmack and leather 2002 ) and sap - feeding insects ( i . e . , many heteropteran omnivores ) may be especially responsive to enhanced plant nitrogen , since they feed selectively and often on tissue that does not contain nitrogen - based allelochemicals ( toxic secondary metabolites ) ( holopainen et al . 1992 ; wheeler 2001 ; huberty and denno 2004 ) . nitrogen concentration is substantially lower in plant than in herbivore biomass and the relative abundance of nitrogen in relation to growth conditions is , by orders of magnitude , more variable in plants than in herbivores ( mattson 1980 ; sterner and elser 2002 ; andersen et al . 2004 ) . the stoichiometric mismatch in nitrogen across trophic levels has also been critical for the evolution of omnivory in heteropteran insects ( eubanks et al . 2003 ) .
the gray willow ( salix cinerea l . ) grows in wet moderately nutrient - rich soils and often forms dense stands along small streams , ditches and pastures and at forest edges ( jonsell 2000 ) . stands generally consist of individuals of the same clone and range from a few square meters to several hectares in size ( although small stands are more common ) .
leaf nitrogen status in the field study and the greenhouse experiment was estimated using an optical chlorophyll meter ( model , spad - 502 , konica minolta sensing , japan ) . this is a non - destructive alternative to analytical methods that can be used to estimate leaf nitrogen content per unit area ; the method is commonly used in plant sciences and has been extensively evaluated for a number of hardwoods , including salix and populus species ( bonneville and fyles 2006 ; weih and rönnberg - wästjung 2007 ; bonosi et al . 2010 ) . spad values are , however , mainly useful for relative comparisons of nitrogen content in leaves , under similar environmental conditions ( chang and robison 2003 ) .
one - year - old shoots were cut from 15 of the gray willow stands and used to create a leaf nitrogen gradient in the greenhouse , similar to the one recorded in the field . the shoots were stored at −5 °c for 3 months and then divided into 90 ( 6 × 15 ) cuttings ( length 20 cm , diameter 0 . 9–1 . 7 cm ) . the cuttings were placed in water in the greenhouse for 2 weeks to initiate root development , before being planted in pots with 2 dm 3 sand ( grain size 0 . 2–1 mm ) . the insides of the pots were lined with cloth , so that water and air ( but not the sand ) could pass through the holes in the base of the pots . the sand was saturated with water at all times . this ensured that water availability would not influence access to nitrogen and simulated the conditions gray willows experience in the field ( growing in wet soils or more or less directly in water ) .
six cuttings from each of the 15 stands were randomly assigned to three different fertilization treatments in the greenhouse , corresponding to 1 . 4 , 8 . 4 or 15 . 4 mg n × week −1 . these fertilization levels were used in a preceding pilot study and therefore known to create nitrogen levels similar to those observed in gray willow stands in the field . to fertilize the plants we used a complete nutrient solution , ‘blomstra’ ( wallco , sweden ) ( n : p : k 5 : 1 : 4 . 3 , ph 7 . 8 ) . the temperature in the greenhouse ranged from 18 to 24 °c . leaf nitrogen status of plants in the greenhouse was recorded 30 days after the first fertilizer application to ensure that the selected treatments resulted in leaf nitrogen concentrations similar to those observed in the field . spad values were recorded at five different positions between the mid - rib and the leaf margin , on three leaves from the mid part of each plant .
the relationship between spad - 502 chlorophyll meter values and leaf nitrogen concentrations was determined using three leaf samples collected from each gray willow stand ( n = 51 ) in the field in june , 2011 , covering almost the full spectrum of values found in gray willow stands at that time of the year . in addition , three leaf samples were collected from clones of the same willows grown in the greenhouse ( n = 45 ) . the aim was to determine whether the relationship between spad values and nitrogen concentrations differed between greenhouse and field samples , for instance due to differences in light and hydrological conditions . mass - based leaf nitrogen concentrations of sampled leaves were determined by gas chromatography using a carlo erba na 1500 elemental analyzer ( carlo erba , milano , italy ) .
the relationship between spad values and mass - based leaf nitrogen concentrations in leaves sampled in the field and in the greenhouse experiment was validated using a linear model . leaf origin was treated as a fixed effect in the model .
the predicted difference in spad values between gray willow stands in the field study and nitrogen treatments in the greenhouse experiment was tested using two separate linear mixed models . potential correlation between data points due to repeated measures of spad values in the field was accounted for by incorporating a first - order autoregressive correlation structure for the time variable ( month ) . by treating stand as a random effect in the models , the intercept was allowed to vary among willow clones .
two measures were used to estimate population variability : the coefficient of variation ( cv ) and the inter - quartile ratio ( iqr ) . cv was calculated for each stand by dividing the standard deviation of density by the mean density . cv thus provides a standardized measure of variability which allows for comparisons regardless of the mean . however , cv is sensitive to zero counts and low means ( heath 2006 ) . the iqr equals the inter - quartile range divided by the median and provides an alternative measure of variability that is independent of both observation extremes and the mean . we fitted two separate linear models with the response variable population variability expressed either as cv or iqr and leaf nitrogen status and willow stand area as fixed effects . cv and iqr were estimated using a subset of the population density data ( for 15 stands , 1999–2008 ) . the selection of data was intended to optimize the number of stands sampled over the same years and the longest continuous time period . the model was restricted to spad values in the range 30–36 because of the low number of observations below this range ( spad < 30 ) . including or excluding the two observations in this lower range in the model did not change the overall results .
analyses were performed in r version 3 . 1 . 0 ( r development core team 2014 ) using the mass package glmmpql function for the poisson glmms ( venables and ripley 2002 ) and the nlme package lme function for the linear mixed model ( pinheiro et al . 2011 ) .
) . the relationship was best described by the model spad = 0 . 41n
spad - leaf nitrogen validation models plotted with field - and greenhouse - recorded leaf nitrogen gradients . predictions were based on linear models describing the relationship between spad values and leaf nitrogen concentrations ( mg × g −1 ) in gray willow leaves ( r 2 = 0 . 67 ) , collected in the greenhouse ( solid line ) and the field ( dotted line ) . fine dotted gray lines are standard errors of prediction estimates . open squares show mean spad values recorded in june for 17 gray willow ( salix cinerea ) stands in forest and open habitats ( squares with white and gray backgrounds , respectively ) . solid squares show mean greenhouse spad values recorded under the 1 . 4 , 8 . 4 and 15 . 4 mg n × week −1 nitrogen treatments . bars show standard errors
the recorded spad values ranged from 25 . 07 to 36 . 39 in may and from 20 . 06 to 36 . 72 in june ( fig .
< 0 . 001 ) . when spad values were translated into nitrogen concentrations using the spad–nitrogen validation for gray willow in the greenhouse , the range was similar to the field - recorded nitrogen concentrations ( fig .
) . population density increased by 195 % with a spad value increase from 30 . 75 to 36 . 49 , which is approximately equivalent to an average increase in leaf nitrogen from 26 to 40 mgn × g
= 0 . 44 ) . population variability , expressed as cv , decreased by 63 % with a spad value increase from 30 . 75 to 36 . 49 , which is approximately equivalent to an average increase in leaf nitrogen from 26 to 40 mgn × g
) . adults with access to both plant and prey exhibited , on average , 89 % higher dry weight , compared to individuals without access to prey . dry weight of individuals without prey increased by , on average , 69 % from the lowest ( 1 . 4 mg n week
< 0 . 01 . predation rate decreased by , on average , 28 % from the lowest ( 1 . 4 mg n week
we utilized a combination of data from long - term field studies and controlled greenhouse experiments to show that feeding on plant resources with a high nitrogen status increases and stabilizes omnivore population density , and enhances performance in the absence of prey . to our knowledge , this is the first empirical evidence under field conditions for this effect . moreover , we found that leaf nitrogen status can alter the omnivore’s trophic behavior : individuals on plants with higher nitrogen status consume less prey . both results are consistent with accumulating ( but scattered ) evidence linking the distribution , dispersal , performance , oviposition preference and trophic behavior of omnivorous insects to intra - specific variation in plant quality ( agrawal et al . 1999 ; eubanks and denno 1999 , 2000 ; eubanks and styrsky 2005 ; groenteman et al . 2006 ; jiménez et al . 2012 ) .
the long - term population level effect and the behavioral effect ( plant vs . prey feeding ) of leaf nitrogen status differed with respect to the consequence for omnivore prey suppression and the outcome of the plant–herbivore–omnivore interaction . the different environmental conditions ( field vs . greenhouse ) and spatial and temporal scales make it impossible to integrate the effects entirely and to predict the net outcome of leaf nitrogen status on prey suppression . the density effect associated with increased plant nutrient status seems , however , as expected , to be stronger than the per capita changes in trophic behavior—especially when longer time scales are considered ( eubanks and denno 2000 ) . consequently , we expect the long - term effect of high leaf nitrogen status ( population stability ) to outweigh the short - term behavioral effect on per capita predation rates—resulting in a net increase in population predation rates with increasing leaf nitrogen status .
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we thank martin weih for advice on spad - nitrogen validation , tomas grönqvist for gc analyses , marie melander and karin eklund for field assistance . support for this work was provided by energimyndigheten , future forests and the oscar and lili lamm foundation .
asl , pd and cb conceived and designed the experiments . asl and pd collected the data . asl analyzed the data and wrote the manuscript ; other authors provided editorial advice .
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sources results page - southwood , t . r . e . & leston , d . ( 1959 ) land and water bugs of the british isles 1 - 436 : 261
the influence of plant sex on the performance of a detrimental herbivore and two biocontrol agents in the dioecious grey willow , salix cinerea .
dioecious plants can be used as model systems for studying how variation in host plants affects plant - insect interactions . this thesis describes investigations into the impact of plant sex on plant - herbivore - predator interactions for the host plant salix cinerea . both herbivores and predators were found to prefer female plants , all other things being equal . as a result , top - down control had a more significant impact on the herbivore population on female plants than was the case for males . despite this , phratora beetles were more abundant on , and preferred , female plants . this may be due to bottom - up control , since female plants have longer leaves . both herbivores and biocontrol agents were more abundant on female plants in the field . because of predation by omnivores , survival rates for herbivore eggs on female plants were lower than those for male plants , and omnivore predation was the primary cause of death among herbivore eggs . these findings may provide novel opportunities for pest control in salix short rotation coppices .
( nl , nj ) > dept . of ecology ( s ) > dept . of ecology
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