the jamaican ibis , jamaican flightless ibis or clubbed-wing ibis ( xenicibis xympithecus ) is an extinct bird species of the ibis subfamily uniquely characterized by its club-like wings .

it is the only species in the genus xenicibis , and one of only two flightless ibis genera , the other being the genus apteribis endemic to hawaii .

jamaican ibis

the jamaican flightless ibis ( xenicibis xympithecus ) , a bird that became extinct approximately 10 , 000 years ago , possessed clublike wings .
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drawing of the wing bones of an american white ibis ( left ) and jamaican ibis ( right ) . bones are scaled such that the humeri are the same size to enable easier comparison of morphological changes .
nicholas r . longrich , and storrs l . olson ( 2010 ) . the bizarre wing of the jamaican flightless ibis xenicibis xympithecus : a unique vertebrate adaptation
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ibis young stay in the nest for a relatively long time ,
he said .
the wing of xenicibis ( top ) compared with that of a typical ibis eudocimus ( middle ) and the extinct , flightless ibis apteribis ( bottom ) . from longrich and olson , 2010 .
xenicibis xympithecus is an extinct flightless ibis from jamaica that had an incredibly rare adaptation — it used its wings as clubs .
a member of the ibis family , it was probably about the size of a chicken , but with an infinitely more robust armoury .
xenicibis is a large , extinct , flightless ibis . it was discovered by storrs olson storrs olson storrs olson from the smithsonian institution , who found some partial remains in a jamaican cave in 1977 . when olson eventually saw the bird’s wing bones , he was baffled . they were so “utterly strange” that he thought the animal must have been suffering from some inexplicable disease .
unlike most flightless birds whose wings turned to vestigial and useless stumps , the jamaican flightless ibis retained extremely long wing bones , but they adapted into use for attack . the wing had a long , curved , banana like structure , with a narrow base and heavy tip . it also had a special joint , which allowed it to be smashed downwards at high speed like a club .
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the jamaican ibis was endemic to jamaica . bones have been excavated from several caves , including the long mile cave , the swansea cave , the jackson ' s bay cave and the red hills fissure . bones from cuba claimed to be of this genus were later identified as those of a limpkin . jamaica and cuba have always been separated by the sea , so it is improbable that a flightless species could reach the other islands .
. . . a roughened area on the carpometacarpal extensor process of phorusrhacos led andrews ( 1901 ) to suggest the presence of a spur or knob used in intraspecific combat . the club - like , thick - walled carpometacarpi of the jamaican ibis xenicibis were interpreted by longrich & olson ( 2011 ) as weapons used in combat . such a role is supported by the presence of healed carpometacarpal and humeral injuries in two specimens . . . .
the initial description of xenicibis was followed two years later by the discovery of a humerus ( upper arm bone ) found in jamacia’s swansea cave . the unusual , robust nature of this bone led olson and steadman to believe that xenicibis was flightless that had evolved on jamacia . along with another fossil flightless ibis – apteribis glenos – described by olson and colleague a . wetmore in 1976 , xenicibis was one of the first flightless ibis to be recognized by science .
while doubtless used for defense , as the ibis evolved its flightlessness on an island filled with predators , it ' s thought the club wing was used against other members of the same species , as evidenced by the high number of fractured bones found in the skeletal remains .
chichester art i did not do any wildlife today , but jean and i had an interesting day of art in chichester . first , we went to see the john minton centenary exhibition at the pallant gallery . minton was a rather sad and tortured artist who committed suicide aged 39 . his art was not great , but he was a fine illustrator and did a large number of books and other things . his paintings included a huge mural called ' jamaican village ' .
. . . birds , however , have been largely ignored until now . hitherto , only about ten fossil bird species were known from the pleistocene of jamaica ( morgan , 1993 ; mcfarlane et al . , 2002 ) , one of which is xenicibis xympithecus , a now - extinct , flightless ibis ( olson and steadman , 1977 ; longrich and olson , 2011 ) . . . .
a very well written and presented article , tying in aspects of larmarckism whilst relating the importance of this new work on the genus xenicibis . the lack of native predators , ( excepting the presence of one or two birds of prey ) , leads me to suspect that the wings evolved for intraspecific combats , perhaps over nesting sites . after all , a number of extant ibis are extremely territorial .
the skeleton of the sacred ibis of the ancient egyptians , as found in mummified form in egyptian tombs , from a paper cuvier had published in 1804 . cuvier identified this bird to be comparable to modern species . however since it was distinctly different from the living bird commonly identified as the same species , it had been used ( by other naturalists ) as evidence for the gradual
transformation or evolution of animals over long periods of time ( from rudwick 1997 ) .
xenicibis might have used its wings to clobber enemies in defence . unlike its living cousins , this ibis couldn’t fly . many island birds lose the ability to fly because they aren’t threatened by any land predators . as a result , their wings become small and stunted , as in the kiwi or the flightless cormorant of the galapagos . but prehistoric jamaica had no shortage of predators , including a boa , an extinct monkey , and several birds of prey . defence would have been important .
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fortunately , there are a few indications that xenicibis truly was a fightin’ ibis . other than the fact that some modern birds fight with their wings , longrich and olson cite two broken arm bones which show signs of healing . one , a humerus , began to heal after being split in two , and the other , a carpometacarpus , has a large callus of bone which probably grew in response to a fracture . these findings are consistent with the idea that these birds were using their wings to strike each other .
what remains unknown is what individual xenicibis were fighting and why . the most plausible answer is “other xenicibis . ” since there is no indication that the anatomy of the weapon differed between males and females , longrich and olson propose that xenicibis fought each other over territory , particularly for good nesting spots . the wing clubs could have also been used to fight off predators , too , and this brings up an important point . we can generate ideas about the function of the bird’s wings by looking at their anatomy and come up with multiple possible uses , but identifying those uses does not tell us why those particular traits evolved in the first place . the clubs of xenicibis may have evolved as a result of ibis - on - ibis combat and could also be used to fight predators once they had already evolved , or perhaps the clubs are some sort of exaptation – a structure used for one function which became co - opted for something else . given the state of the evidence , it is likely that xenicibis used its clubs in combat with each other , but we should not be so careless as to say that the clubs evolved for that function .
while no living ibis shows adaptation of the wing for combat , ibises are highly territorial during nesting and feeding [ 6 , 18 ] , and disputes frequently escalate to fights . of particular note is the fact that plegadis has been seen to grasp its opponent with the bill and then strike with the wings [ 19 ] . in many species , intraspecific combat involves access to mates , but ibises are monogamous [ 5 ] , and there is no evidence that the club is dimorphic . it is therefore likely that intraspecific combat would have focused on securing territory , rather than mates .
it took nearly a century to recognize the strange character of this bird . sometime between 1910 and 1920 the zoologist harold elmer anthony collected a few bird bones from long mile cave in trelawny parish , jamaica . they sat , virtually unnoticed , in the american museum of natural history collections for decades , but in 1977 the ornithologists storrs olson and d . w . steadman used anthony’s fossils to describe the bird as a previously unknown fossil ibis . they named it xenicibis xympithecus . about the size of a chicken , this bird only recently became extinct ( around 2 , 200 years ago according to a table included in the new book holocene extinctions . )
dixon’s restorations are so delightful because they take creatures native to our own sliver of time and transform them into alien shapes . probable or not , they are wonderful because they are both recognizable and alien . the same can be said of actual creatures which existed just yesterday in geological terms . we live in the wake of the earth’s last great extinction . ( in fact , we are arguably still in the midst of it . ) the giant sloths , saber - toothed cats , glyptodonts , mammoths , giant hyenas , and other impressive pleistocene beasts only became extinct within the past 15 , 000 years or so . nor were all the strange creatures charismatic , mammalian megafauna . one odd ibis , first described over thirty years ago , was unlike any other bird to have ever evolved .
the main object of interest is the bird’s bizarre hand . the carpometacarpus – part of the bird’s “hand” you tear the meat off of while having chicken wings – is wider in diameter than the bird’s thigh bone ( femur ) and has extraordinarily - thickened walls . behind this bone , the radius – a lower arm bone in line with the carpometacarpus – was unusually thick compared to the relatively slender companion bone , the ulna . additionally , there was a modified groove – properly called the carpal trochlea – on the end of the carpometacarpus closest to the rest of the body which allowed the wrist to be quickly swung forward in a way different from other avians . this was not the arm of flying bird , but it also differed from the wings of many ground - dwelling birds . rather than becoming reduced or vestigial , the wings of xenicibis actually became heavier and more robust . ( see the comparison of ibis wing skeletons below . )
warblington shore birds peter milinets - raby was out this morning for a couple of hours visiting the shore at warblington ( 6am to 8 : 11am - tide coming in ) . main observations . warblington cemetery extension : 2 green woodpecker . ibis field : 2 chiffchaff , 1 jay , 1 swallow . hedgerow behind conigar point : 1 chiffchaff , 1 sedge warbler , 1 stock dove . conigar point : 1 grey plover , 2 common tern , 2 little egret , 1 whimbrel , 1 snipe flushed by kestrel , 9 dunlin , 7 ringed plover , 1 greenshank . off pook lane : 7 grey plover , 8 dunlin , 14 greenshank ( 5 with colour rings rg / / - + yy / / - & g / / r + yn / / - & g / / r + ll / / - & b / / r + lo / / - & gr / / - + yy / / - ) , 3 common tern . great spotted woodpecker flew over heading across the water to hayling , 7 swallow , 2 yellow wagtail - yellow adult over , then a dull , light brown juvenile landed on the shore for a couple of minutes before flying on - autumn has truly arrived now ! 1 sandwich tern , 9 black - tailed godwit .
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a few million years ago , jamaica was home to one of the strangest boxers in the animal kingdom – a flightless bird called xenicibis xympithecus that could batter its enemies with club - shaped wings .
since then , olson has found more remains including an almost complete skeleton . now , he and his partner nicholas longrich from yale university , have a very different view of the wing . they think it was a club . weapons like clubs and bats have large weighted ends to deliver heavy impacts , and long handles to increase the speed of the swing . that’s exactly what you see in xenicibis ’s wing .
its hand bone ( the metacarpal ) was massive , curved and inflated – perfect for inflicting strong blows . it sat at the end of a long ‘handle’ , made up of the wrist and the forearm – perfect for creating a fast swing . the metacarpal is also hollow , just like many baseball bats are , allowing it to produce a stronger blow without adding too much weight . and its joints allowed it to swing its wing out very quickly , and extend it as far as possible , giving it speed and reach .
the bones are telling , but did xenicibis really punch with its wing ? it’s hard to be sure , especially because there are few modern birds with similar bones to compare against . however , longrich and olson have found some compelling evidence that the bird struck heavy blows with its wings at least two specimens of xenicibis had arm bones that had broken and healed . the first had broken its upper arm ( humerus ) in two and the bones hadn’t knitted together properly . the second had fractured its hand , and a massive callus had grown over the front edge . these birds struck something with enough force to injure themselves .
alternatively , the bird could have boxed with its rivals . longrich and olson note that a couple of flightless birds have similar ( but far less extreme ) forearms , including the steamer duck and the extinct rodriguez island solitaire . and both of these species occasionally use ( or used ) their wings in to beat other individuals in fights .
in fact , many birds use their wings as weapons ( including some ibises ) . some even have special adaptations for combat . waterfowl in particular , such as geese , ducks and swans , have a wide variety of spurs , spikes and bony knobs on their wrists . they use these weapons in battle and conflicts can be very violent ( although there’s some debate about whether swans can actually break a person’s arm ) . xenicibis just expanded on a theme that’s common in the bird world and took it to an evolutionary extreme .
birds have repeatedly evolved flightlessness to exploit insular environments where mammalian predators and competitors are absent , and flightlessness is typically associated with reduction of the forelimbs and pectoral girdle [ 1 , 2 ] . flightlessness has evolved dozens of times in island habitats , and in many taxa , including waterfowl , megapodes , rails , pigeons , parrots [ 2 ] and ibises [ 3 , 4 ] .
later , a partial skeleton prepared from a block of cave breccia ( usnm 460349 ) included the radius , ulna and a carpometacarpus so utterly strange in morphology that it appeared to represent some inexplicable pathology . however , subsequent discoveries of additional material from numerous additional individuals , including almost all of the skeleton ( figure 1 ) , show that the bizarre morphology of the wing represents an adaptation unprecedented among the vertebrates , which we here describe and attempt to interpret .
skeletal reconstruction of xenicibis xympithecus based on usnm 460349 and fossils from the red hills fissure cave deposits , saint andrews parish , jamaica .
the highly modified manus is unlike that of any other bird , flightless or volant ( figure 2 a – d ) . the major metacarpal is massive , inflated , strongly bowed and expanded distally . its diameter is up to 12 . 8 mm dorsoventrally and 9 . 9 mm anteroposteriorly , exceeding the diameter of the femur . the bone is hollow , but the anterior wall is broad and thickened ( figure 2 e ) , being up to 3 mm thick in this area , whereas the cortex of the femur is under 2 mm thick .
wing skeleton of x . xympithecus compared with eudocimus albus and apteribis glenos . ( a ) left carpometacarpus ( usnm 543067 ) in cranial view , ( b ) dorsal view , ( c ) ventral view and ( d ) caudal view . ( e ) major metacarpal ( usnm 543074 ) sectioned at midlength ( note the greatly thickened anterior wall ) . ( f ) radiograph of carpometacarpus of eudocimus compared with ( g ) xenicibis ( usnm 543067 ) . ( h ) composite left wing of xenicibis ( usnm 543067 - 543073 ) compared with ( i ) eudocimus ( usnm 266467 ) and ( j ) apteribis .
the alular metacarpal has a reduced extensor process . the shaft of the minor metacarpal has a thick cortex and a subtriangular section , unlike the strap - like form seen in other birds , and is unusual in being shifted dorsally to lie directly behind the major metacarpal .
the carpal trochlea has a reduced ventral ridge and the articulation for the radiale is symmetrical , pulley - shaped and extended anteriorly . this joint allowed the wrist to swing forward in the plane of the antebrachium , without the complex pronation and supination that occurs in the wing of volant birds [ 10 ] , while the anterior extension of the carpal trochlea permitted hyperextension of the manus .
the proximal phalanx of the major digit is a stubby and block - like element ( figure 2 h ) . it has a subtriangular section and weakly developed articular surfaces .
the radius ( figure 2 h ) is expanded distally to give it a club shape , and it is unlike typical birds in that its diameter exceeds that of the ulna . unusually , the ulna is slender and nearly straight ( figure 2 h ) . its proximal articular surface is on the proximal end of the bone such that the elbow can be fully extended , placing the ulna ' s long axis in line with the humerus . distally , the shaft is dorsoventrally flattened and the articulation for the ulnare is reduced .
the humerus ( figure 2 h ) is typical of flightless birds [ 9 ] except that the shaft is elongate and bowed dorsally . the shaft is slender proximally but expanded distally . the distal end is twisted by approximately 30° such that the articular facets are directed ventrally .
unlike other flightless birds , particularly apteribis , the pectoral girdle is relatively well developed . the coracoids ( figure 3 e ) and furcula ( figure 3 f ) are large and robust . the sternal carina ( figure 3 g ) , while reduced compared with volant ibises , is large when compared with that of apteribis or other flightless birds .
selected skeletal elements of x . xympithecus from the red hills fissure cave deposits , representing multiple individuals . skull elements : ( a ) rostrum ( usnm 543075 ) and mandible ( usnm 543076 ) in lateral view and ( b ) dorsal view ; ( c ) cranial cap ( usnm 543077 ) in left lateral view and ( d ) dorsal view . pectoral girdle : ( e ) coracoid ( usnm 543078 ) , ( f ) clavicle ( usnm 543079 ) and ( g ) two fragments of sternal carina ( usnm 543081 and 54080 ) . hindlimb : ( h ) tibiotarsus ( usnm 543084 ) , ( i ) femur ( usnm 543083 ) and ( j ) tarsometatarsus ( usnm 543085 ) . pelvis ( usnm 543082 ) : ( k ) dorsal view and ( l ) lateral view .
other skeletal elements show modification , albeit less extreme . as in other ibises , the beak is long and decurved ( figure 3 a , b ) , but the tip is knob - like , as in apteribis , rather than spatulate . the parietals and frontals ( figure 3 d ) are thickened by a honeycomb of cancellous bone . the vertebrae and notarium are robust , but otherwise unremarkable . the hindlimb ( figure 3 h – j ) resembles that of apteribis in being massive , with a shortened tibia and tarsometatarsus . the anterior portion of the ilium is dorsally expanded to form a tall crest and the synsacrum is robust ( figure 3 k , l ) , again resembling apteribis .
adaptation for a terrestrial , flightless lifestyle is correlated with hypertrophy of the hindlimbs and pelvis , and reduction of the forelimbs and pectoral girdle [ 1 , 2 ] . in this context , the long forelimbs and well - developed pectoral girdle of xenicibis suggest that the wings were not functionless vestiges . the twisted humerus , short antebrachium and massive , bowed metacarpus make it extremely unlikely that its wing could have functioned in flight ; but the extreme modification of the forelimb argues that the wings had been adapted to function in some capacity .
attempts to identify avian analogues are complicated by the unique morphology of xenicibis . the distally expanded radius does find parallels among steamer ducks ( tachyeres spp . ; n . r . longrich 2010 , personal observation ) and the extinct rodriguez island solitaire ( pezophaps solitaria , columbidae [ 11 ] ) . in those species , the wings are ( or were ) used to deliver hammering blows to conspecifics [ 11 , 12 ] , suggesting that the wing of xenicibis may have functioned as a weapon . perhaps a better analogue is to be found among some of the mantis shrimps ( stomatopoda : gonodactyloidea ) ; these have a club - like , distally inflated dactyl that is used to strike prey and conspecifics [ 13 ] , again hinting that xenicibis may have used the club - shaped hands to deliver blows .
we therefore propose that the wing of xenicibis functioned as a club or flail . several features of the limb would have facilitated this function . kinetic energy is the product of mass and velocity squared ; accordingly , weapons such as clubs and flails have a long handle to increase the angular velocity of the club , and are heavily weighted to increase the mass accelerated by the swing , and the centre of mass is near the end of the club , where the angular velocity is highest . precisely this design is seen in the hand of xenicibis , where the end of the wing is massive , and the proximal metacarpus and long forelimb could act as a handle . the comparatively weak wrist joint does not preclude such a function , because during impact a club acts as a free body [ 14 ] ; the hollow metacarpal also allows the hand to achieve greater strength for a given amount of material , much like an aluminium baseball bat [ 14 ] .
several morphological adaptations would have further increased the wing ' s effectiveness as a weapon . reduction of the extensor process and elongation of the manus would decrease the mechanical advantage of the wing extensors , producing a more rapid wing extension . the retention of long wing bones allows the forelimb to be swung rapidly , while the ability to hyperextend at the elbow and wrist increase the wing ' s effective length , and therefore its angular velocity when swung .
this hypothesis can be tested by looking for direct evidence of agonistic behaviour in the form of traumatic injuries sustained from delivering or receiving blows . fractures are common in pugnacious birds such as the steamer ducks and solitaire [ 11 , 12 ] , and are therefore predicted to occur in xenicibis . two bones of xenicibis show evidence of healed fractures . the first is a humerus that was broken in two ( figure 4 a ) ; a fracture callus indicates healing although the bones failed to knit . the second specimen ( usnm 460349 ) is a carpometacarpus ( figure 4 b ) with a massive callus overgrowing the anterior surface of the major metacarpal and extending inside the bone as well ; the hand apparently suffered an impact that fractured the anterior wall of the metacarpal .
fossils of xenicibis showing evidence of trauma . ( a ) fractured left humerus ( usnm 543086 and 543087 ) showing fracture callus ( fc ) : ( i ) photograph ; ( ii ) radiograph . ( b ) carpometacarpus of xenicibis ( usnm 460349 ) with rugose fracture callus on anterior , dorsal and ventral surfaces of the major metacarpal : ( i ) photograph ; ( ii ) radiograph showing development of callus inside the medullary cavity .
we thank ross macphee for the opportunity to study the red hills fossils . discussions with helen f . james and richard l . zusi and guidance from b . rosemary grant were helpful in developing this project . n . r . l . received support from an nsf graduate research fellowship , an alberta ingenuity graduate studentship and the yale institute for biospheric studies . finally , we thank the reviewers for their helpful reviews of this paper . fossil specimens are deposited in the department of paleobiology , national museum of natural history ( usnm ) , smithsonian institution , washington , dc .
descriptions of thirty - two new species of birds from the hawaiian islands . part i . non - passeriformes
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this is the only known example of a vertebrate adapting a limb into club .
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ornithologists speculate that the wings were used as weapons , in the manner of a club or flail , similar to the adaptations found in some mantis shrimps ( stomatopoda : gonodactyloidea ) that possess a club - like distally inflated dactyl used to strike prey and other shrimps . however , among birds , this adaption appears to be unique . adaptations of the wing to fight in birds is an example of contingency in which various bird species find different solutions to the same problem based in chance .
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what comes next for evolution ? this seems like a simple question . every day we are learning more about the history of life on earth , and we would expect that , over 150 years since charles darwin published on the origin of species , the life of the past could be used to extrapolate the trajectory of evolution’s arrow . regarding our own species , especially , there is a pervasive desire to know what our distant descendants might be like .
but this question is flawed . even though there are evolutionary constraints – not everything we can imagine is possible – life is not following a tightly - restricted road of progress towards a pre - determined goal . natural selection , the primary engine of evolutionary change , generates both diversity and disparity – charles darwin’s “endless forms most beautiful and most wonderful” – as organisms are constantly subjected to changing conditions . there is no way to create an evolutionary law or take everything into account so that we can predict what will exist one thousand , one million , one billion years in the future . that is part of the wonder of evolutionary science . we can’t know how the vast array of lineages we see around us today are going to change in years to come , but we still feel compelled to ask what the future holds .
this isn’t to say that we can’t have a little fun with speculation . in 1981 the scottish geologist dougal dixon published one of the lasting cult favorites of paleontologists – after man : a zoology of the future . imagining the world 50 million years after our extinction – which , one takes it , is probably not all that far off in geological terms – dixon takes the reader on a tour of the world’s ecosystems . there is an odd familiarity to all of it , seeing the prospective descendants of modern creatures twisted into new and bizarre forms . in the seas , fully aquatic descendants of penguins ( the vortex and the porpin ) have replaced the long - extinct whales , the long - legged rabbucks stride across the grasslands , and my personal favorite , a terrestrial bat dixon calls the night stalker , “roams screeching and screaming” in search of prey under the cover of darkness .
( dixon also wrote two similar books : the new dinosaurs [ 1988 ] , chronicling the evolutionary history of still - living non - avian dinosaurs , and man after man [ 1991 ] , a grotesque gallery of future humans modified through bioengineering . after man was arguably his most successful outing , however ; it was turned into a documentary and cartoon in japan . )
the complete skeleton of xenicibis , based upon multiple individuals , as reconstructed by nicholas longrich . from longrich and olson , 2010 .
there must be a reason why xenicibis had such strange wings . the trouble is that no other bird had wings quite like it . spurs , spikes , and other wing weapons are seen among birds like steamer ducks – which use them in combat with members of their own species – but nothing as extreme as the banana wings of xenicibis . ( the closest thing , storrs and longrich suggest , might be the formidable claws of mantis shrimp , though this analogy breaks down in terms of anatomy and function – imagine xenicibis using its wings to trap and consume prey ! ) being that many birds do have a modified carpometacarpus used for combat , however , it is a reasonable hypothesis that the odd wings of xenicibis were used as some kind of club or flail . the bird could have swung its arms so that the wide , thick parts of the carpometacarpus near the tip of the wing struck its opponent .
the actual manner in which xenicibis fought is not discussed in the paper – i assume that will be the focus of further research – but longrich and olson propose that the unique weaponry of this bird underscores a grander theme of evolutionary change . of all the birds which have spurs , spikes , and clubs on their wings , none was quite like xenicibis . despite similar evolutionary pressures – fighting one another for territory , mates , or some other resource – multiple species of birds from various lineages have been adapted to have very different weaponry , with xenicibis having the oddest of all . even when selective pressures are the same , evolutionary quirks and contingencies can generate quite different outcomes .
as strange as it may sound , prehistoric creatures like xenicibis make dixon’s speculative beasts seem almost conservative in their anatomy . many of the creatures featured in after man are products of convergent evolution , testaments to the power of natural selection to funnel organisms down similar channels from disparate starting points . convergence is a real and significant part of life’s pattern , but chance and contingency can open up unique evolutionary possibilities that cannot be predicted or foreseen . the more we learn about the fossil record , the stranger life becomes , and though i am a little saddened by the fact that i will never know what life is going to be like 50 million years from now , i am left in awe by the fantastic creatures which inhabited this planet in ages past .
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an extinct flightless bird from jamaica fought rivals and predators using wings evolved into clubs , scientists suggest .
the boney bludgeons carried by xenicibis xympithecus are unlike anything else known in the bird world - or in mammals , reptiles or amphibians .
writing in the royal society journal proceedings b , the scientists report finding bones that had apparently been broken by another bird ' s club .
fossils show that the metacarpus - one of the
hand
bones - was elongated and much bigger than in related species , with very thick walls .
this allowed the wings to function
in combat as a jointed club or flail
, the researchers write .
we don ' t really know how they would have used these clubs , but we do know that modern ibises grab each other by the beak and pound away with their wings ,
said nicholas longrich , from yale university in the us .
and we analysed two bones that had been broken during fighting , including a humerus ( upper arm bone ) that had been snapped in half - it had started to re - heal , although the two ends hadn ' t knitted together ,
he told bbc news .
dr longrich ' s colleague in this research , storrs olson from the smithsonian institution , was one of the scientists who first identified xenicibis xympithecus back in the 1970s .
a number of other birds are known to fight by whacking each other with their wings - including swans , who will also protect their young this way .
some , including screamers , lapwings and and spur - winged goose , have evolved spurs to increase the damage they can wreak .
the extinct solitaire from the indian ocean island of rodrigues - a cousin of the dodo - had bony growths colloquially known as
musketballs
on their wings , which appear to have served the same purpose .
but among vertebrates - there ' s no animal of any sort that has anything like a limb modified as a club ,
noted dr longrich .
julian hume , an avian palaeontologist with london ' s natural history museum who was not involved in the research , noted that unlike most flightless birds , xenicibis retained long wings , possibly making its flailing more powerful .
so if they retained that feature , that suggests they needed it for defence against predators - and there were quite a few on jamaica .
however , ibises also tend to be intensely territorial , so the flailing clubs may also - or alternatively - have found employment in disputes between individuals , probably with both sexes involved .
the foreign secretary quits , telling theresa may
needless self - doubt
is leading to
semi - brexit
.
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. . . and todus todus . of these , x . xympithecus was previously known from the late pleistocene of jamaica and from the rhrc ' s sediment ( longrich and olson , 2011 ) , but t . todus , and corvus ? sp . . . .
. . . then , what function did dakotaraptor co - opt them for ? aside from brooding , examination of certain extant flightless birds suggests that they could have played a role in hunting strategy , mating display , aggressive territorial behavior , and / or shielding the young ( sensu cooper & harrison , 1994 ; hopp & orsen , 2004 ; longrich & olson , 2011 ) . implementing feathers in aggressive territorial behavior , as has been observed for extant avian taxa ( cooper & harrison , 1994 ; longrich & olson , 2011 ) , is a behavior that could easily have augmented the pack - hunting strategies already proposed for deinonychus , utahraptor , and other dromaeosaurs ( kirkland , burge , & gaston , 1993 ; maxwell & ostrom , 1995 ) . . . .
. . . aside from brooding , examination of certain extant flightless birds suggests that they could have played a role in hunting strategy , mating display , aggressive territorial behavior , and / or shielding the young ( sensu cooper & harrison , 1994 ; hopp & orsen , 2004 ; longrich & olson , 2011 ) . implementing feathers in aggressive territorial behavior , as has been observed for extant avian taxa ( cooper & harrison , 1994 ; longrich & olson , 2011 ) , is a behavior that could easily have augmented the pack - hunting strategies already proposed for deinonychus , utahraptor , and other dromaeosaurs ( kirkland , burge , & gaston , 1993 ; maxwell & ostrom , 1995 ) . dakotaraptor ' s remiges would have lengthened the outstretched arms , providing long and robust outstretched wings that would have been useful when implemented in such strategies . . . .
. . . scale bar = 10 mm . olson & steadman , 1977 ( threskiornithidae ) is unique in having a thick - walled elongate , inflated metacarpus ( longrich & olson , 2011 ) . within columbidae ( pigeons and doves ) , the extinct flightless natunaornis gigoura and the extant goura ground pigeons both have a small bony outgrowth on the processus extensorius of the carpometacarpus ( goodwin , 1983 ; worthy , 2001 ) ( fig . 8b ) , which closely resembles that of a female solitaire . . . .
. . . within columbidae ( pigeons and doves ) , the extinct flightless natunaornis gigoura and the extant goura ground pigeons both have a small bony outgrowth on the processus extensorius of the carpometacarpus ( goodwin , 1983 ; worthy , 2001 ) ( fig . 8b ) , which closely resembles that of a female solitaire . in our examination of carpal spurs and knobs , we note that the literature does not accurately reflect the number of taxa that exhibit some form of carpal growth , but without exception all of these structures are or were used as weapons ( jefferies , 1881 ; sclater , 1886 ; rand , 1954 ; goodwin , 1983 ; livezey & humphrey , 1985 ; worthy , 2001 ; longrich & olson , 2011 ) . our study confirms that the solitaire ' s carpal knob was also a weapon . . . .
fósiles del holoceno tardío provenientes de un depósito subterráneo en la península de portland ridge al sur de jamaica extienden el rango de buteogallus aequinoctialis hasta la cuenca del caribe , a más de 1700 km al oeste - noroeste de la distribución actual más occidental de esta especie . los fósiles se localizaron cerca de las áreas más extensas de manglares , el cual es el hábitat requerido . . . [ show full abstract ]
in the present paper we briefly reviewed and commented the terminology for bird topography , especially wing and tail . we named here newly required japanese terms , such as alula coverts and carpal covert , and also illustra - ted some of them . terms for bird topography are given , which including new japanese ones indicated by asterisk ( * ) in table 1 .
a new genus and species of unusual tern ( aves : laridae : anoinae ) from the middle miocene calvert for . . .
feducciavis loftini , new genus and species , is described from a single partial associated skeleton from the middle miocene calvert formation of virginia . this bird was evidently most closely related to the noddy terns ( anoinae , anous , procelsterna ) . compared with those genera , the tibiotarsus was much shorter and the ulna much longer in relation to the size of the humerus . the absolute size of . . . [ show full abstract ]
history , morphology , and fossil record of the extinct puerto rican parakeet psittacara maugei souanc . . .
the extinct puerto rican parakeet ( psittacara maugei ) has been known with certainty only from mona island and is usually regarded as a poorly defined subspecies of p . chloroptera of hispaniola . examination of skin specimens and comparison of skeletons with fossil and archeological material from puerto rico , show that psitticara maugei is a fully distinct species from p . chloroptera , differing . . . [ show full abstract ]
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this charming hotel enjoys a convenient location in a quiet part of amsterdam city centre , just a few steps from rembrandtplein , a major square in the centre of this city . amsterdam central station and the waterlooplein metro are just around the corner , providing easy access to amsterdam schiphol airport , the world trade centre amsterdam and rai . guests will find themselves near some of the most important tourist attractions in the area including anne frank house , madame tussaud & apos ; s , dam square , and the van gogh museum , all easily reach by metro . the rooms boast functional furniture and have been decorated in soothing tones to create a relaxing atmosphere in which to completely unwind . this pet - friendly establishment includes parking as well as wi - fi internet connection .
located in the centre of amsterdam , close to the waterlooplein and a short walk from central station .
20 kms to the nearest airport ( schiphol airport ) 2 minute walk to the nearest metro station ( waterloo plein ) 3 km to the nearest station ( amsterdam central railway stat ) 2 minute walk to the nearest bus stop 8 km to the nearest fair site ( rai exhibition and congress ce ) .
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make use of convenient amenities , which include complimentary wireless internet access and tour / ticket assistance .
our modern and cozy rooms are the perfect place to relax , with comfortable beds and well - appointed bathrooms - everything you need to enjoy a pleasant stay . there are 77 rooms .
public transport by taxi or train can take you from schiphol airport to your hotel in 30 minutes . the hotel is then a mere 3 minutes walk from amsterdam central station .
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excellent food , would definitely recommend ! ! ! visited restaurant on carnival . good hearty caribbean food available and for a good price .
rates of extinction vary widely . for example , during the last 100 , 000 years of the pleistocene epoch ( about 2 . 6 million to 11 , 700 years ago ) , some 40 percent of the existing genera of large mammals in africa and more than 70 percent in north america , south america , and australia went extinct . ecologists estimate that the present - day extinction rate is 1 , 000 to 10 , 000 times the background extinction rate ( between one and five species per year ) because of deforestation , habitat loss , overhunting , pollution , climate change , and other human activities—the sum total of which will likely result in the loss of between 30 and 50 percent of extant species by the middle of the 21st century .
according to the best estimates of the world’s environmental experts , human activities have driven species to extinction at rates perhaps 1 , 000 times the natural , or background , rate , and future rates of extinction will likely be higher . to…
although extinction is an ongoing feature of earth’s flora and fauna ( the vast majority of species ever to have lived are extinct ) , the fossil record reveals five unusually large extinctions , each involving the demise of vast numbers of species . these conspicuous declines in diversity are referred to as mass extinctions ; they are distinguished from the majority of extinctions , which occur continually and are referred to as background extinction . ranked in descending order of severity , they are :
the impact of a near - earth object 66 million years ago in what is today the caribbean region , as depicted in an artist ' s conception . many scientists believe that the collision of a large asteroid or comet nucleus with earth triggered the mass extinction of the dinosaurs and many other species near the end of the cretaceous period .
and many species of plants . although many scientists contend that this event was caused by one or more large
( 407 million to about 359 million years ago ) , which included 15–20 percent of marine families and 70–80 percent of all animal species . roughly 86 percent of marine brachiopod species perished , along with many corals , conodonts , and trilobites .
in essence , mass extinctions are unusual because of the large numbers of taxa that die out , the concentrated time frame , the widespread geographic area affected , and the many different kinds of animals and plants eliminated . in addition , the mechanisms of mass extinction are different from those of background extinctions .
the trilobite modocia typicalis lived during the middle cambrian period . as a group , trilobites were among the longest - lasting organisms , first evolving at the beginning of the cambrian period ( about 541 million years ago ) and dying out some 289 million years later during the permian extinction , which occurred near the end of the permian period ( roughly 252 million years ago ) .
many species have become extinct because of hunting and overharvesting , the conversion of wetlands and forests to croplands and urban areas , pollution , the introduction of invasive species , and other forms of human - caused destruction of their natural environments . indeed , current rates of human - induced extinctions are estimated to be about 1 , 000 times greater than past natural ( background ) rates of extinction , leading some scientists to call modern times the sixth mass extinction . this high extinction rate is largely due to the exponential growth in human numbers : from about 1 billion in 1850 , the world’s population reached 2 billion in 1930 and more than 6 billion in 2000 , and it is expected to reach about 10 billion by 2050 . as a result of increasing human populations , habitat loss is the greatest factor in current levels of extinction . for example , less than one - sixth of the land area of europe has remained unmodified by human activity , and more than half of all wildlife habitat has been eliminated in more than four - fifths of countries in the paleotropics ( the old world tropics that span africa , asia , and indonesia ) .


steinernema scapterisci , the mole cricket nematode , is a species of nematode in the order rhabditida .

it is a parasite of insects in the order orthoptera , the grasshoppers , crickets and their allies .

it has been introduced into florida in the united states in an effort to provide a biological pest control of mole crickets .

steinernema scapterisci

steinernema scapterisci is a close relative of the insect - parasitic nematode steinernema carpocapsae .
11 . a formulation as claimed in claim 10 in which the two or more species of steinernema are selected from the group comprising : steinernema feltiae ; stinernema scapterisci ; steinernema riobravis ; steinernema carpocapsae ; and steinernema krussei .
16 . a formulation as claimed in claims 14 or 15 in which the at least one species of steinernema is selected from the group comprising : steinernema feltiae ; stinernema scapterisci ; steinernema riobravis ; steinernema carpocapsae ; and steinernema krussei .
infection of sound - trapped mole crickets , scapteriscus spp . , by steinernema scapterisci
12 . a formulation as claimed in claim 10 in which the two species of steinernema are selected from the group comprising steinernema feltiae , steinernema krussei , and steinernema carpocapsae .
. vertical dispersal of steinernema scapterisci . journal of nematology , 22 : 574 - 578 .
17 . a formulation as claimed in claim 16 in which the at least one species of steinernema is selected from the group comprising : steinernema feltiae ; steinernema krussei and steinernema carpocapsae .
. bacterial symbionts of steinernema scapterisci . journal of invertebrate pathology , 62 : 68 - 72 .
. addendum to the morphology of steinernema scapterisci . journal of nematology , 24 : 478 - 481 .
. pathogenicity of steinernema scapterisci to selected invertebrates . journal of nematology , 23 : 7 - 11 .
. life cycle of steinernema scapterisci nguyen & smart , 1990 . journal of nematology , 24 : 160 - 169 .
. mode of entry and sites of development of steinernema scapterisci . journal of nematology , 23 : 267 - 268 .
. steinernema scapterisci n . sp . ( steinernematidae : nematoda ) . journal of nematology , 22 : 187 - 199 .
nguyen kb , smart gc jr . 1990b . vertical dispersal of steinernema scapterisci . journal of nematology 22 : 574 - 578 .
anguillera mm , smart gc jr . 1992 . bacterial symbionts of steinernema scapterisci . journal of invertebrate pathology 62 : 68 - 72 .
nguyen kb , smart gc jr . 1992b . addendum to the morphology of steinernema scapterisci . journal of nematology 24 : 478 - 481 .
nguyen kb , smart gc jr . 1991a . pathogenicity of steinernema scapterisci to selected invertebrates . journal of nematology 23 : 7 - 11 .
. preliminary studies on survival of steinernema scapterisci in soil . soil and crop science society of florida , proceedings , 49 : 230 - 233 .
since 1985 steinernema scapterisci has been detected in mole crickets collected from several other sites in alachua county , undoubtedly spread from our original release sites by infected mole crickets . by november 1998 , about 10 to 30 percent of mole crickets collected in alachua county , florida were infected with steinernema scapterisci .
figure 1 . mole cricket nematodes , steinernema scapterisci nguyen & smart , emerging from a mole cricket . photograph by k . nguyen , university of florida .
figure 8 . mole cricket nematodes , steinernema scapterisci nguyen & smart , emerging from a cockroach nymph . photograph by k . nguyen , university of florida .
nguyen kb , smart gc jr . 1992a . life cycle of steinernema scapterisci nguyen and smart , 1990 . journal of nematology 24 : 160 - 169 .
nguyen kb , smart gc jr . 1991b . mode of entry and sites of development of steinernema scapterisci . journal of nematology 23 : 267 - 268 .
nguyen kb , smart gc . pathogenicity of steinernema scapterisci to selected invertebrates . j nematol . 1991 ; 23 ( 1 ) : 7–11 . pmid : 19283088
nguyen kb , smart gc jr . 1990 . steinernema scapterisci n . sp . ( steinernematidae : nematoda ) . journal of nematology 22 : 187 - 199 .
. development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . journal of invertebrate pathology , 62 : 289 - 294 .
aguillera mm , hodge nc , stall re , smart gc . bacterial symbionts of steinernema scapterisci . j invertebr pathol . 1993 ; 62 ( 1 ) : 68–72 .
this project was initiated in june 1999 to re - establish commercial production of steinernema scapterisci for mole cricket control in florida . in about one year , we have :
nematicides were used on golf courses against plant - parasitic nematodes and doubtless would kill steinernema scapterisci . however , such chemicals no longer are labeled for that purpose by u . s . environmental protection agency . studies in the early 2000s showed that steinernema scapterisci is remarkably resistant to chemical insecticides . the effect of chemical herbicides on this nematode is unknown .
nguyen kb , smart gc . steinernema scapterisci n . sp . ( rhabditida , steinernematidae ) . j nematol . 1990 ; 22 ( 2 ) : 187–99 . pmid : 19287709
figure 3 . spicules ( top ) and gubernaculum ( bottom ) of the mole cricket nematode , steinernema scapterisci nguyen & smart . photographs by k . nguyen , university of florida .
nguyen kb , smart gc jr . 1990a . preliminary studies on survival of steinernema scapterisci in soil . soil and crop science society of florida , proceedings 49 : 230 - 233 .
nguyen kb , smart gc . life - cycle of steinernema - scapterisci nguyen and smart , 1990 . j nematol . 1992 ; 24 ( 1 ) : 160–9 . pmid : 19283218
. inoculative release of steinernema scapterisci ( rhabditida : steinernematidae ) to suppress pest mole crickets ( orthoptera : gryllotalpidae ) of golf courses . environmental entomology , 23 : 1331 - 1337 .
figure 6 . infective juvenile of the mole cricket nematode , steinernema scapterisci nguyen & smart , with the head showing labial raising disc . photograph by k . nguyen , university of florida .
anguillera mm , smart gc jr . 1992 . development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . journal of invertebrate pathology 62 : 289 - 294 .
aguillera mm , smart gc . development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . j invertebr pathol . 1993 ; 62 ( 3 ) : 289–94 .
citation : lu d , sepulveda c , dillman ar ( 2017 ) infective juveniles of the entomopathogenic nematode steinernema scapterisci are preferentially activated by cricket tissue . plos one 12 ( 1 ) : e0169410 . urltoken
nguyen , k . b . , smart , g . c . , jr . 1990 . steinernema scapterisci n . sp . ( rhabditida : steinernematidae ) . journal of nematology 22 : 187 - 199 .
steinernema scapterisci was successfully introduced by inoculative applications ( parkman et al . 1993 , 1994 ) in florida . since 1993 , this nematode has been commercialized in florida to control mole crickets in golf courses and pastures .
biological control , entomopathogenic nematode , mole cricket parasite , morphology , new species , neoaplectana , scapteriscus , steinernema scapterisci n . sp . , taxonomy , light microscopy ( lm ) , scanning electron microscopy ( sem )
. biological control of mole crickets in the genus scapteriscus with the nematode steinernema scapterisci nguyen & smart , 1990 . rencontres caraibes en lutte biologique , guadeloupe , 5 - 7 november 1990 . ed . inra , paris 1991 .
30 . use of a formulation according to claim 1 to treat one or more of the tawny mole cricket and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema scapterisci and two species of heterorhabditis .
parkman jp , frank jh , nguyen kb , smart gc jr . 1994 . inoculative release of steinernema scapterisci ( rhabditida : steinernematidae ) to suppress pest mole crickets ( orthoptera : gryllotalpidae ) of golf courses . environmental entomology 23 : 1331 - 1337 .
parkman jp , frank jh , nguyen kb , smart gc jr . 1993 . dispersal of steinernema scapterisci ( rhabditida : steinernematidae ) after inoculative applications for mole cricket ( orthoptera : gryllotalpidae ) control in pastures . biological control 3 : 226 - 232 .
24 . a method as claimed in claim 22 in which the target insect is one or more of the tawny mole cricket and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema scapterisci and two species of heterorhabditis .
steinernema biddulphi n . sp . can be distinguished from other steinernema species by means of a combination of morphological and morphometric characteristics of males and ijs . based on these data , steinernema biddulphi n . sp . belongs to the “ bicornutum ” clade within the steinernematidae family . molecular data show that within this clade , steinernema biddulphi n . sp . is sister to the pair of s . pakistanense and s . bifurcatum and this group is related to the pair of s . yirgalemense and s . abbasi .
figure 5 . a , b - face views of first generation females ; c - female tail ; d , e , f - double flapped epiptygma ; of the mole cricket nematode , steinernema scapterisci nguyen & smart . photograph by k . nguyen , university of florida
parkman jp , frank jh , nguyen kb , smart gc . inoculative release of steinernema - scapterisci ( rhabditida , steinernematidae ) to suppress pest mole crickets ( orthoptera , gryllotalpidae ) on golf - courses . environmental entomology . 1994 ; 23 ( 5 ) : 1331–7 .
a multigene approach for assessing evolutionary relationships of xenorhabdus spp . ( gamma - proteobacteria ) , the bacterial symbionts of entomopathogenic steinernema nematodes
in 1985 , a nematode parasite of mole crickets was brought from uruguay to florida . that nematode , which was described as a new species , steinernema scapterisci ( nguyen and smart 1990 ) , appears to be a major factor in limiting populations of mole crickets in south america .
comparative morphometrics ( in μm ) of infective juveniles of steinernema biddulphi n . sp . ; mean ± sd with ranges given in brackets .
some commercial companies may sell steinernema nematodes other than s . scapterisci as biopesticides for use against mole crickets . if the label does not tell you the species of nematode , then ask the supplier to give the name in writing . it is true than other steinernema species will kill mole crickets , at least to some extent , but do not expect them to give any residual activity because they are not known to reproduce in mole crickets .
comparative morphometrics ( in μm ) of first generation males of steinernema biddulphi n . sp . ; mean ± sd with ranges given in brackets .
phylogenetic relationships of the species from “ bicornutum ” group and other related species of steinernema based on analysis of its rdna regions . steinernema nepalense and s . scapterisci was used as outgroup taxon . the percentage of replicate trees in which the associated taxa clustered together in the bootstrap test ( 10 , 000 replicates ) are shown next to the branches . branch lengths indicate evolutionary distances and are expressed in the units of number of base differences per site .
smart gc jr , nguyen kb , parkman jp , frank jh . 1990 . biological control of mole crickets in the genus scapteriscus with the nematode steinernema scapterisci nguyen and smart , , 1990 . rencontres caraibes en lutte biologique , guadeloupe , 5 - 7 november 1990 . ed . inra , paris 1991 .
scapterisci ( rhabditida : steinernematidae ) after inoculative applications for mole cricket ( orthoptera : gryllotalpidae ) control in pastures . biological control , 3 : 226 - 232 .
meeting held with a company in georgia to rear s . scapterisci in vivo ( 11 / 17 / 99 ) , confidentiality agreement completed and business plan submitted .
meeting held with uf , otl ( 1 / 20 / 00 ) , agreed to assist in locating a company to produce s . scapterisci using in vitro methods .
phylogenetic relationships of the species from “ bicornutum ” group and other related species of steinernema based on analysis of d2 - d3 expansion segments of the 28s rdna . steinernema nepalense and s . scapterisci was used as outgroup taxon . the percentage of replicate trees in which the associated taxa clustered together in the bootstrap test ( 10 , 000 replicates ) are shown next to the branches . branch lengths indicate evolutionary distances and are expressed in the units of number of base differences per site .
foreign company and uf , otl drafted an exclusive license agreement for production of s . scapterisci for sale in the u . s . ( 6 / 19 / 00 ) .
steinernema scapterisci was more pathogenic to insects tested in the order orthoptera than to those in the orders lepidoptera or hymenoptera ; it was not pathogenic to earthworms . the nematode also infected and killed the mole crickets scapteriscus acletus and s . vicinus when released four successive times at 10 - day intervals in containers of soil infested with the nematode .
9 . a formulation as claimed in any preceding claim in which the first and second genus are selected from the group consisting of : steinernema ; and heterorhapditis .
furthermore , the first generation females of steinernema biddulphi n . sp . differ from the females of s . pakistanense and s . bifurcatum by having a postanal swelling .
10 . a formulation as claimed in claim 9 in which the first genus ( majority species ) is steinernema and the second genus ( minority species ) is heterorhapditis .
14 . a formulation as claimed in claim 9 in which the first genus ( majority species ) is heterorhapditis and the second genus ( minority species ) is steinernema .
the whole genome shotgun project for s . scapterisci has been deposited at ddbj / embl / genbank under the accession [ azbw00000000 ] . the version described in this paper is version azbw01000000 .
nguyen kb , maruniak j , adams bj . diagnostic and phylogenetic utility of the rdna internal transcribed spacer sequences of steinernema . j nematol . 2001 ; 33 : 73–82 .
morphometric characters ( in μm ) of steinernema biddulphi n . sp . based on the holotype and 20 paratypes of each generations with mean ± sd and ranges given in brackets .
parasitism is a major ecological niche for a variety of nematodes . multiple nematode lineages have specialized as pathogens , including deadly parasites of insects that are used in biological control . we have sequenced and analyzed the draft genomes and transcriptomes of the entomopathogenic nematode steinernema carpocapsae and four congeners ( s . scapterisci , s . monticolum , s . feltiae , and s . glaseri ) .
meeting held with richard gaskalla , fdacs / dpi ( 1 / 13 / 00 ) , determined that they would be receptive to rearing s . scapterisci if provided with the necessary funding and direction .
report with background information on mole cricket infestations , damage and efficacy of control by s . scapterisci prepared for fdacs ( 2 / 25 / 00 ) , report sent with justification for a federal appropriation .
hox cluster architecture in steinernema . comparisons of the hox clusters of c . elegans , s . carpocapsae , and s . scapterisci . each cluster is mapped at the same scale , with the colored boxes representing different putative genes between the lin - 39 and ceh - 13 orthologs . genes marked in blue are specific to steinernema , not having orthologs in c . elegans . gray genes have a c . elegans ortholog , though they are not syntenic in the nematodes compared . genes marked in brown are unique , not having obvious orthologs in the other nematodes in this comparison
32 . use of a formulation according to claim 1 to treat a root weevil , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis megidis , and two species of steinernema .
figure 7 . diagram of the life cycle of steinernema scapterisci . g1 = first - generation adults , g2 = second - generation adults , j1 = first - stage juvenile , j2 = second - stage juvenile that may be the preinfective or non - preinfective stage , j3 = third stage noninfective juvenile , pi = preinfective stage , ij = third - stage infective juvenile , j4 = fourth - stage juvenile .
26 . a method as claimed in claim 22 in which the target insect is a root weevil , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis megidis , and two species of steinernema .
meeting held with uf , office of technology licensing ( 10 / 20 / 99 , 2 month delay ) , agreed that up front costs would be minimal for a license to sell s . scapterisci for mole cricket control .
29 . use of a formulation according to claim 1 to treat one or more of sciarid larvae and fungus gnats , wherein the cohort of infective juvenile nematodes consists of steinernema felitae and two species of heterorhabditis .
the phylogenetic trees of the its and 28s genes were obtained by the minimum evolution method ( rzhetsky and nei , 1992 ) in mega 6 . 0 ( tamura et al . , 2013 ) . steinernema nepalense and steinernema scapterisci were used as outgroup . the minimum evolution tree was searched using the close - neighbor - interchange ( cni ) algorithm ( nei and kumar , 2000 ) . the neighbor - joining algorithm ( saitou and nei , 1987 ) was used to generate the initial tree . the evolutionary distances were computed using the p - distance method ( nei and kumar , 2000 ) and are expressed as the number of base differences per site .
sleinemema scapterisci nguyen et smart , 1990 is recorded for the first time in argentina . this entomopathogenic nematode was found parasitizing the mole cricket scapteriscus borelli giglio - tos at the locality of colon ( province of buenos aires ) , argentina .
nadler sa , bolotin e , stock sp . phylogenetic relationships of steinernema travassos , ( nematoda : cephalobina : steinernematidae ) based on nuclear , mitochondrial and morphological data . syst parasitol . 2006 ; 63 : 161–81 .
23 . a method as claimed in claim 22 in which the target insect is one or more of sciarid larvae and fungus gnats , and wherein the cohort of infective juvenile nematodes consists of steinernema felitae and two species of heterorhabditis .
34 . use of a formulation according to claim 1 to treat one or more of root weevils , wood borers , and scarabs , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis bacteriophora and two species of steinernema .
meeting held with uf , otl , and a foreign company to discuss an agreement to produce in vitro and market s . scapterisci in the u . s . ( 4 / 13 / 00 ) , the company requested a 60 - day non - compete agreement .
28 . a method as claimed in claim 22 in which the target insect is one or more of root weevils , wood borers , and scarabs , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis bacteriophora and two species of steinernema .
lee mm , stock sp . a multigene approach for assessing evolutionary relationships of xenorhabdus spp . ( gamma - proteobacteria ) , the bacterial symbionts of entomopathogenic steinernema nematodes . j invertebr pathol . 2010 ; 104 ( 2 ) : 67–74 . pmid : 20102721
meeting held with fdacs , dpi and the mole cricket task force to determine the status of state funds requested to produce s . scapterisci and initiate a mole cricket control program ( 7 / 21 / 00 ) , appropriation is in a trust fund and project is unable to proceed .
figure 2 . males of the mole cricket nematode , steinernema scapterisci nguyen & smart n . sp . a ) spicules of the first - generation male showing angular head , ribs , and gubernaculum with anterior end bent upward . b ) variation in the tail shape of the first - generation males . c ) tail of the second - generation male showing elongate spicule head . d ) variation in tail shape of second - generation males . e ) entire body of the first - generation male . drawing by k . nguyen , university of florida .
31 . use of a formulation according to claim 1 to treat one or more of root weevils , the tawny mole cricket , and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema riobravis , and two species of heterorhabditis .
25 . a method as claimed in claim 22 in which the target insect is one or more of root weevils , the tawny mole cricket , and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema riobravis , and two species of heterorhabditis .
33 . use of a formulation according to claim 1 to treat one or more of armyworms , cutworms , webworms , root weevils , wood borers , artichokes , and plume moths , and wherein the cohort of infective juvenile nematodes consists of steinernema carpocapsae and two species of heterorhabditis .
nadler sa , bolotin e , stock sp . phylogenetic relationships of steinernema travassos , 1927 ( nematoda : cephalobina : steinernematidae ) based on nuclear , mitochondrial and morphological data . systematic parasitology . 2006 ; 63 ( 3 ) : 161–81 . epub 2006 / 03 / 17 . pmid : 16541298
meeting held with fdacs , dpi to plan production and prepare a budget , $ 281 , 000 year 1 ( 2 / 17 / 00 ) , planned a realistic structure and budget for solid culture of s . scapterisci , and delivered it to dr . martha roberts ( fdacs ) in the commissioner ' s office .
27 . a method as claimed in claim 22 in which the target insect is one or more of armyworms , cutworms , webworms , root weevils , wood borers , artichokes , and plume moths , and wherein the cohort of infective juvenile nematodes consists of steinernema carpocapsae and two species of heterorhabditis .
a . house crickets were anesthetized at 4°c and were then put individually into the wells of a cryo storage box . each well was lined with a small piece of paper towel loaded with approximately 100 ijs of s . scapterisci suspended in 100 μl of tap water . b . about 3–4 infected crickets were transferred to white traps [ 31 ] .
we have identified a set of expanded gene families that are likely to be involved in parasitism . we have also identified a set of non - coding motifs associated with groups of orthologous genes in steinernema and caenorhabditis involved in neurogenesis and embryonic development that are likely part of conserved protein–dna relationships shared between these two genera .
comparing activation rates of ijs exposed to different insect homogenates , mole cricket homogenate induced significantly higher activation than house cricket homogenate at 10 % ( p < 0 . 0001 ) , and 50 % ( p = 0 . 009 ) concentrations . both cricket homogenates induced significantly higher activation in s . scapterisci ijs than waxworm homogenate ( p < 0 . 0001 ) .
steinernema biddulphi sp . n . was reared on last instar of g . mellonella as described by kaya and stock ( 1997 ) . the first and second generation adults were obtained from the dissection of 3 - and 5 - d - old infected g . mellonella larvae , respectively . infective juveniles were collected approximately 1 wk after emergence from the cadavers .
dillman ar , macchietto m , porter cf , rogers a , williams b , antoshechkin i , lee mm , goodwin z , lu x , lewis ee , goodrich - blair h , stock sp , adams bj , sternberg pw , mortazavi a . comparative genomics of steinernema reveals deeply conserved gene regulatory networks . genome biol , 2015 ; 16 ( ) : 200
protein gels showing the quantity and diversity of proteins secreted by activated ( 12h ) and non - activated ( na ) ijs . equal percentage of total volumes of concentrated proteins from non - activated and activated s . scapterisci ijs was loaded onto each gel . ijs were activated with 50 % cricket homogenate . a . a protein gel stained with colloidal coomassie blue . b . a protein gel with silver staining .
the effects of environmental factors ( such as temperature , moisture , aeration , and soil type [ esp . texture and chemistry ] ) and biotic factors ( species of epn , targeted insect , age of insect , soil fauna ) have been documented by numerous researchers ( gaugler and kaya , 1990 ; kaya and gaugler , 1993 ; shapiro - ilan et al . , 2012 ; grewal et al . , 2005a ; georgis et al . , 2006 ) . temperature range for survival and infectivity will depend on the species of epn and its native habitat and center of origin ( kaya , 1990 ) . for example , steinernema feltiae can be infective from 2 - 30°c , whereas some heterorhabditids can infect host insects from 7 to 35°c and steinernema carpocapsae is nearly inactive at 10°c ( kaya , 1990 ; georgis et al . , 2006 ; lacey et al . , 2006a ) .
the life cycle of s . biddulphi n . sp . is similar to other steinernema species . in an experimental infection of g . mellonella with a dose of 50 ijs per insect at a temperature of 22°c , the majority of insects were dead after 48 h , and fourth stage juveniles were present in the hosts . two amphimictic generations occur inside the host , and the first and second generation stages could be observed after 2 and 8 d of initial infection , respectively . infective juveniles appeared 10 d postinoculation .
s . scapterisci ijs were exposed to three different host homogenates ; scapteriscus borellii , acheta domesticus , and galleria mellonella . the homogenates were made in different concentrations ( 10 % , 25 % , 50 % and crude ) . we were unable to test crude waxworm homogenate for technical reasons ; the crude homogenate was too thick to be separated from nematodes after exposure but we were able to test 10 % , 25 % , and 50 % waxworm homogenate . each of these activation experiments was collected and observed after 18 hours of exposure to the homogenate . ij activation was then quantified as described above .
steinernema biddulphi n . sp . lm of infective juvenile ( ij ) , male and female . a , c . first generation female . a . tail with postanal swelling . c . vulval region . b , d . second generation female . b . tail with postanal swelling . d . vulval region . e . first generation male , tail with spicules and gubernaculum . f . second generation male , tail with spicules and gubernaculum . g , h . ij . g . anterior portion showing rounded head and excretory pore ( arrow ) . h . tail with anus and hyaline region .
steinernema biddulphi n . sp . a . first generation male , tail with spicules and gubernaculum , lateral . b . first generation female , vulval region . c . first generation female , tail . d . second generation female , vulval region . e . second generation female , conical tail . f , g . infective juvenile . f . anterior region showing excretory pore and nerve ring . g . tail showing anus and hyaline region . ( scale bar in micrometers : a = 58 ; b = 143 ; c = 68 ; d = 71 , e = 61 ; f = 91 ; g = 75 ) .
in recent years , several surveys have been conducted in south africa resulting in the recovery of eight new steinernematid species , namely steinernema khoisanae ( nguyen et al . , 2006 ) , s . citrae ( stokwe et al . , 2011 ) , s . sacchari ( nthenga et al . , 2014 ) , s . tophus ( cimen et al . , 2014 ) , s . innovationi ( cimen et al . , 2015 ) , s . jeffreyense ( malan et al . , 2015 ) , s . beitlechemi ( cimen et al . , 2016 ) , and s . fabii ( abate et al . , 2016 ) .
based on the blast search and phylogenetic analysis of the concatenated sequences , sequences of the 16s rdna and reca and gyrb genes ( data not shown ) , the symbiotic bacterium of s . biddulphi n . sp . ( bacterial strain sgi - 246 ) seems to be most closely related to xenorhabdus indica ( blast similarities 98 % for 16s rdna , and 97 % for reca and gyrb genes ) . this bacterium was found in association with other nematodes from the “ bicornutum ” group , namely steinernema abbasi ( somvanshi et al . , 2006 ) , s . yirgalamense ( ferreira et al . , 2016 ) , and s . bifurcatum ( shahina et al . , 2014 ) .
steinernema biddulphi n . sp . scanning electron microscopy of infective juvenile ( ij ) , male and female . a–c . ij . a . head region with horn - like structures . b . lateral field in mid - body ( ridges numbered 1 to 6 ) . c . lateral field in tail region . d . first generation female , tail , and four projections on tip of the tail ( arrow ) . e , f . second generation female . e . tail with postanal swelling . f . vulva . g . first generation male , tail with paired genital papillae ( numbered ) and single papilla ( s ) , lateral . h , i . second generation male . h . tail with paired genital papillae , single papilla ( s ) and mucron ( arrow ) . i . tail with a part of paired genital papilae ( numbered ) , mucron ( arrow ) , and phasmid opening ( p ) , ventro - lateral .
s . scapterisci activation was categorized based on 3 morphological features ( fig 2 ) — ( 1 ) the mouth being open or closed , ( 2 ) the state of the anterior gut , or the opening of the gut immediately posterior to the pharyngeal bulb , and ( 3 ) expansion of pharyngeal bulb and how pronounced it appeared . if an individual nematode had all three characteristics ( an open mouth , an open and expanded anterior gut , and an expanded and visible pharyngeal bulb ) they were considered fully activated ( fig 2c ) . if the nematode had a visible pharyngeal bulb , though had not proceeded to a further stage of activation as indicated by the gut and mouth opening , it was considered partially activated ( fig 2d ) . if the nematode had no visible pharyngeal bulb , it was considered non - activated ( fig 2b ) . non - activated nematodes were generally still ensheathed , though presence of the l2 cuticle sheath was not used to determine nematode activation . dead nematodes were not counted . any nematodes that had developed beyond l3 into l4 or even adulthood were counted as fully activated .
proteins were denatured by sds sample buffer ( final 1x buffer : 50 mm trishcl ph6 . 8 , 2 % sds , 2 . 5 % ficoll , 0 . 01 % bromophenol blue , and 0 . 1 m dtt ) and heating for 10 min in boiling water before being loaded to the 4–15 % mini - protean® tgx™ precast gels ( biorad , 456–1084 ) . protein molecular weights were marked by the precision plus protein dual color standards ( biorad , # 1610374 ) . the electrophoresis was run at 100 volts for 60–90 min and the gels were stained by colloidal coomassie blue [ 35 ] or by the pierce silver staining kit ( pierce , # 24600 ) . equal percentages of the total volumes of collected proteins from non - activated and activated s . scapterisci ijs were loaded onto each gel . 10 % of protein volumes of activated ( total 210 μl x 1 . 08 μg / μl ) and non - activated ( total 200 μl x 0 . 008 μg / μl ) ijs were loaded onto the gel for colloidal coomassie blue staining ; for the silver staining , the volumes were 1 % of the total collected proteins .
conserved non - coding networks in steinernema and caenorhabditis . a a hierarchically clustered heat map of 30 derived regulatory motifs and the go terms that the target genes of these motifs are enriched in . only motif - go term associations that are shared between s . carpocapsae and c . elegans are shown . p - values depicted are from c . elegans associations . colored arrows point to single go term or groups of go terms that belong to the four developmental categories shown . b a network of conserved s . carpocapsae and c . elegans motif - target gene associations related to neurogenesis go terms . only nodes for motifs and downstream genes with degrees ≥5 are shown in the network . c a network of conserved s . carpocapsae and c . elegans motif - target gene associations related to muscle go terms . only nodes for motifs and downstream genes with degrees ≥5 are shown in the network . d zag - 1 gene model in s . carpocapsae and c . elegans showing conserved motifs , and well as conserved regulatory modules ( clusters of conserved motifs ) . sequence conservation tracks are displayed below each gene model . associations between zag - 1 and motifs are highlighted in red in the neurogenesis network . go gene ontology
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male , second generation : a second generation male is similar morphologically to that of the first generation except that it is about two - thirds as long and one - half as wide and the spicules have an elongate head .
female , first generation : anteriorly , the female is similar to the male but much larger . lateral fields and phasmids not observed . six labial papillae and four cephalic papillae present . stoma with prominent cheilorhabdions unusually thickened , appearing as a circular or hexagonal ring in face view . esophagus typical of family . esophago - intestinal valve large . excretory pore located anteriorly to mid - metacorpus .
excretory duct unusually prominent forming an elliptically - shaped structure seemingly with a hole at the center . gonads didelphic , amphidelphic , reflexed . vulva appears as a transverse slit with a prominent double - flapped epiptygma . tail length shorter than anal body width , with a mucron at end .
females , second generation : second generation female similar morphologically to that of the first generation with the following exceptions : about one - half as long and two - thirds as wide , valve in basal bulb of esophagus more prominent , elliptically - shaped structure less prominent , tail , which tapers to a point bearing a mucron , longer than body width at anus .
infective juveniles ( third stage ) : the infective stage , when newly formed , is always enclosed in the cuticle of the second - stage juvenile as a sheath . however , the sheath is lost rather easily , even in storage , and thus may not always be present . body thin , head with a labial raising disc ; lip region not offset , oral aperture not observed ; six labial , four cephalic papillae and an elevated oral disc prominent . esophagus degenerate and thus not seen clearly , but its basal bulb is elongate and has a valve . lateral field with six incisures . tail tapers gradually dorsally but abruptly ventrally .
three years after the nematode was released , a few mole crickets collected in a farm 10 miles from the nearest release site were found infected with the nematode . by the fifth year , 25 to 65 percent of the mole crickets collected at this farm were infected ( smart et al . 1990 ) .
in 1990 and 1991 , the nematode was released in pastures in six different counties and on nearly 30 golf courses in florida . it has become established and has reduced populations of mole crickets in all of the pastures .
mcsorley r . ( july 1997 ) . soil - inhabiting nematodes . uf / ifas featured creatures . eeny - 12 . ( august 1999 ) .
nguyen kb . ( may 1999 ) . mole cricket control by entomopathogenic nematodes . uf / ifas . ( no longer available online )
nguyen kb . ( february 1999 ) . taxonomy of entomopathogenic nematodes . uf / ifas . ( no longer available online )
nguyen kb . ( february 1998 ) . symbiotic bacteria of entomopathogenic nematodes . uf / ifas . ( no longer available online )
smart gc jr , nguyen kb . 1995 . biological control of orthoptera pest insects . united state patent , patent number 5 , 466 , 448 , date of patent : november , 14 1995 .
publication date : june 1999 . latest revision : june 2014 . reviewed : april 2017 .
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copyright : © 2017 lu et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
data availability : all relevant data are within the paper and its supporting information files .
funding : this work was supported by the national institutes of health , national institute of allergy and infectious diseases . k22 career transition award ai119155 to ard . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
galleria mellonella waxworms used for activation experiments were purchased from crittergrub ( urltoken ) . once received , waxworms were immediately sorted to remove dead ones , and the healthy waxworms were frozen at - 80°c until used . acheta domesticus house crickets ( american cricket ranch , lakeside , ca ) were immediately sorted to remove dead ones , and the healthy crickets were frozen at - 80°c until used . scapteriscus borellii mole crickets used for activation experiments were collected from the rio hondo golf course ( downey , ca ) as previously described [ 33 ] . adult mole crickets were fed a diet of wheat germ ( 15 g agar , 166 g wheat germ , 900–1000 ml h 2 o ) and water ad libitum until used . healthy adult mole crickets were frozen at - 80°c prior to use in activation experiments .
to recover nematodes from the small - scale activation assays , tap water was added to soak the sponge pieces in the petri dish for 5 min and the nematode suspension was transferred to a 15 ml tube . the process was repeated two more times and all nematodes were combined in one 15 ml tube . the nematodes were pelleted by centrifugation at 700 rcf for 1 min and washed 3 times with tap water . the supernatant was removed leaving ~ 1 . 5 ml of nematode suspension . aliquots of the nematode suspension were visualized with a compound microscope at the 10 x 40 magnification and scored for activation . higher magnification was occasionally used when the morphology was unclear . 3–4 aliquots were quantified per treatment meaning that 150–600 nematodes were used for each replicate .
the mouth , pharynx , and anterior gut are marked in these images by red arrows and labeled m , p , and g respectively . a . schematic drawing illustrating the morphological features of an activated ij on the left and a non - activated ij on the right . in the non - activated ij the pharynx and anterior gut can be difficult to visualize . b . two pictures showing the opaque morphology of non - activated ijs . the mouth is closed , the anterior gut is closed , and the pharyngeal bulb is not visible . c . two pictures showing the visible morphology of activated ijs . these fully activated ijs have visibly open mouths and stoma . they have expanded pharyngeal bulbs and the anterior gut is open . d . this picture shows a nematode scored as partially active because the pharyngeal bulb is only partially expanded .
ijs were exposed to insect homogenate for different amounts of time : 6 , 12 , 18 , 24 , and 30 hours . though a 3 - hour exposure was initially tested , it was too difficult to reliably distinguish morphological characteristics . for the time course we used 50 % acheta domesticus homogenate , made as described above .
statistical analysis was performed using graphpad prism 6 . 04 . standard statistical tests were used for all experiments , as described in the figure legends . one - way anovas were conducted to compare experimental effects among different exposure times to 50 % house cricket homogenate and the effects of the percentage of host homogenate on activation and partial activation . unpaired t tests were used to compare the experimental effects between two treatments within an experiment ( e . g . differences between two time points of the time course activation or differences between two homogenate concentrations within an experiment ) .
comparing activation rates for each type of host homogenate we found that for house cricket homogenate , 25 % , 50 % , and crude homogenate all produced significantly higher activation than 10 % homogenate ( p < 0 . 0001 ) ( fig 5a and s2 file ) . 50 % and crude house cricket homogenate both induced significantly higher activation than 25 % homogenate ( p < 0 . 0001 ) , and crude homogenate induced significantly higher activation than 50 % homogenate ( p = 0 . 04 ) . using mole cricket homogenate , there was no significant difference in activation between 10 % and 25 % homogenate , but 50 % and crude homogenate both induced significantly higher activation than 10 % or 25 % homogenate ( p≤0 . 03 ) ( fig 5b and s2 file ) . there was no significant difference between 50 % and crude homogenate . using waxworm homogenate , 25 % and 50 % homogenate both induced significantly more activation than 10 % homogenate ( p≤0 . 002 ) ( fig 5c ) , but there was no significant difference between 25 % and 50 % waxworm homogenate .
to study ij activation we first identified quantifiable morphologic changes that could be used to differentiate between activated and non - activated ijs ( fig 2 ) . we found that by examining the state of the mouth , pharyngeal bulb , and anterior gut , we could reliably distinguish between non - activated , partially activated , and fully activated ijs ( fig 2 ) . some early - activated ijs developed to l4 and young adult stages when incubated for more than 18 hours in cricket homogenate and in this study we categorized those as fully activated ijs . in future studies it may be informative to distinguish between activated ijs and nematodes that have developed beyond the activated l3 stage .
s2 file . activation data using different insect homogenates and different concentrations of homogenate .
we thank john rodriguez and the rio hondo golf course grounds crew for access to the grounds for collecting mole crickets ; sudarshan aryal and dennis chang for collecting and maintaining mole crickets and for providing feedback on the manuscript ; tiffany baiocchi , edwin lewis , and byron adams for insightful comments on the manuscript .
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ijs in the wells of a plastic cryo storage box with 100 slots designed for 1 . 5 ml microtubes ( e . g . , vwr cat # 40000–322 ) , with the lid removed . a sheet of transparent plastic was cut to fit the top of the box and cover all the wells to prevent crickets from escaping ; small holes were punched in the plastic for aeration . single layers of paper towels ( ~ 4 cm x 4 cm ) were pushed into the wells of the microtube storage box using a pen to make infection chambers . about 100 ijs of
) . the box was covered by the plastic sheet described above , which was pushed against the wells using a slightly smaller freezer box ( e . g . , vwr cat # 82007–162 ) . the boxes were securely bound together by rubber bands or tape . the infection boxes were incubated for 2 days in the dark at 25°c . dead crickets , presumably infected , were transferred to white traps [
] in which a piece of 5 . 5 cm filter paper was raised by the lid of a 3 . 5 cm petri dish in a 10 cm petri dish with a thin layer of tap water ( about 2 mm deep ) (
) — ( 1 ) the mouth being open or closed , ( 2 ) the state of the anterior gut , or the opening of the gut immediately posterior to the pharyngeal bulb , and ( 3 ) expansion of pharyngeal bulb and how pronounced it appeared . if an individual nematode had all three characteristics ( an open mouth , an open and expanded anterior gut , and an expanded and visible pharyngeal bulb ) they were considered fully activated (