prioniodontida , also known as the " complex conodonts " , is a large clade of conodonts that includes two major evolutionary grades ; the prioniodinina and the ozarkodinina .

it includes many of the more famous conodonts , such as the giant ordovician promissum ( prioniodinina ) from the soom shale and the carboniferous specimens from the granton shrimp bed ( ozarkodinina ) .

they are euconodonts , in that their elements are composed of two layers ; the crown and the basal body , and are assumed to be a clade .

prioniodontida

this is a recent cladogram of the prioniodontida , simplified from donoghue et al . , ( 2008 ) .
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in earlier sections , we have disposed of the contentious and speculative subject of conodont soft - tissue anatomy . we now belatedly turn to the conodont apparatus itself - - hard tissues about which we have hard data . specifically , we will describe , albeit briefly and without much detail , the apparatus of ozarkodinid idiognathodus which is well known from the work of purnell & donoghue ( 1997 ) and purnell ( 1995 , 2001 , 2001a ) .
ozarkodinids , like most euconodonts , have three basic types of elements . these three basic types are the s , m , and p elements . the precise nomenclature has varied considerably over the years . to the extent it makes a difference , we follow the system described by purnell et al . ( 2000 ) . the ozarkodinid elements are better differentiated than the elements of many other groups , which makes them convenient to study .
the reader may note that we have made it a point to use certain terminology which is de rigueur in any discussion of conodonts . the point here is not to demonstrate our lamentably poor knowledge of these terms , nor to obscure the facts while purporting to explain them . to the contrary , the objective of our using this
stupid stuff
is , like that of houseman ' s mithradates , to inure the reader to these linguistic poisons by introducing them in small doses and under controlled conditions . [ 1 ]
s - elements : in the image of an idiognathodus natural assemblage at right , the s - elements are the long , rod - like bones with straight , transverse pectinate teeth and a sort of pick - axe at the anterior or rostral end . more generally , s - elements are frequently curved , as in the manticolepis model below . however , they are all relatively simple in shape , relatively gracile , and with strong longitudinal asymmetry , the base being located much closer to one end than the other . each conodont has 4 or 5 mirror - image pairs of s - elements , numbered s 1 to s 4 or s 5 in addition , each conodont possesses a symmetrical , unpaired , median s 0 element , shown as green on the figure on the left . the s 0 is an unpaired , bilaterally symmetrical , medial element which is effectively at least triramous . two of its branches create a stout , transverse , rastrate bar across the bottom of the mouth , denticles facing upward . the s - element numbering system is symmetrically arranged around this s 0 element , with the first element to the left of the s 0 designated s 1 s , the first to the right s 1 d , and so on ( with the superscript designations for sinister and dexter ) .
m - elements : conodonts have a single pair of m - elements . these are only loosely connected to the s - assemblage and appear to have been attached in the mouth cavity on its dorsal or lateral surface . their shapes are frequently complex , vaguely reminiscent of anything from a nunchuck to a dart . while the m - elements are structurally obscure , their functional role is clear . they are the fork which held the food item in place as it was sliced or scooped by the s - elements , which acted as a combination knife and spoon .
p - elements : the p - elements are the stout bones at the right of the right image , the left of the left image , and the bottom of the bottom ( manticolepis ) image . contrary to everyone ' s expectations , they appear to have been oriented vertically , with the two sets of denticles facing and , in fact , interdigitating as shown in the idiognathodus natural assemblage . purnell has produced convincing sem images showing regular wear facets . the clear implication is that , at least in idiognathodus , the teeth occluded in a regular and precise way . high resolution images of these facets may be found at wear on conodont elements .
the functional biology of the idiognathodus apparatus : purnell & donoghue reconstruct the functional biology of the idiognathodus along the following lines . the s - elements were attached to the medial surface of ( at least ) two strong lateral cartilaginous plates , by analogy to the oral apparatus of hagfishes and lampreys . the entire apparatus of s - and m - elements was everted over the anteroventral edge of the mouth . in this position , the distal ends of the s - elements would spread out like the ribs of a fan . the s - element denticles ( or keratinous
teeth
mounted on them ) would grasp and / or slice the food items . the oral plates were then drawn back into the mouth , forcing the s - elements closer together , gripping the food more tightly and forcing it into the oral cavity . the food material was then drawn past the p - elements which crushed , sliced , and sheared it before it was passed to the gut . atw020728 .
notes : [ 1 ] speaking of stupid stuff , this poem is slightly misquoted at the link , destroying the meter of the critical last two lines . the correct lines are : - - i tell the tale that i heard told . / mithradates , he died old . furthermore , contrary to various other web sources , the poem is by houseman , not by auden or yeats . the concluding lines are an ironic reference to houseman ' s own to an athlete dying young . the full story behind the poem is interesting , but strains even the elastic notions of relevance which constrain this web site .
phylogeny : euconodonta : : protopanderodontida + * : balognathidae + ( prioniodinida + ozarkodinida ) .
characters : s - elements with hairpin shape ; s - elements oriented with cusps rostral and denticles obliquely dorsal [ p + 00 ] ; p elements less massive & more angular ; 3 pairs of p elements ; apparatus probably eversible .
references : purnell et al . 2000 ) [ p + 00 ] . apw040316 .
note : according to a recent ( june , 2003 ) pander society newsletter , an analysis of the distomodus apparatus is nearing completion .
links : prioniodinida ; pan 51 . ( abstracts of dhanda and purnell & von bitter ) ; albe r iana t . apw040316 .
links : ozarkodinida ; muzeum geologiczne - konodont ; abstract ; pandd 97 . pdf ; . apw040316 .
bactrognathidae is actually only one of two families ( the other unnamed ) belonging to a superfamily that is as yet unnamed .
remarks on the evolution of ordovician conodonts . j dzik , acta palaeontologica polonica , 1976 , volume 21 , no 4 ( pdf , retrieved 30 april 2016 )
donoghue , p . c . j ; m . a purnell ; r . j aldridge ; s . zhang ( 2008 ) .
the interrelationships of ' complex ' conodonts ( vertebrata )
.
gabbott , s . e . ; r . j . aldridge ; j . n . theron ( 1995 ) .
a giant conodont with preserved muscle tissue from the upper ordovician of south africa
.
briggs , d . e . g ; e . n . k clarkson ; r . j aldridge ( 1983 ) .
the conodont animal
.
this article is issued from wikipedia - version of the 6 / 5 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .
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master - dr c . giles miller ; natural history museum ; palaeontology ; miller
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yang & zhang , in an et al . , 1983 | | ` - - †
albanesi , g . l . & barnes , c . r . , 2000 : subspeciation within a punctuated equilibrium evolutionary event : phylogenetic history of the lower - middle ordovician paroistodus originalis - p . horridus complex ( conodonta ) . –journal of paleontology : vol . 74 , # 3 , pp . 492 - 502
ethington , r . l . , lehnert , o . & repetski , j . e . , 2000 : stiptognathus new genus ( conodonta : ibexian , lower ordovician ) , and the apparatus of stiptognathus borealis ( repetski , 1982 ) . –journal of paleontology : vol . 74 , # 1 , pp . 92 - 100
gabbott , s . e . , aldridge , r . j . & theron , j . n . , 1995 : a giant conodont with preserved muscle tissue from the upper ordovician of south africa . –nature : vol . 374 , 27 april , pp . 800 - 803
janvier , p . , 1995 [ preview ] : conodonts join the club . –nature : vol . 374 , 27 april , pp . 761 - 762
janvier , p . , 1999 [ preview ] : catching the first fish . –nature : vol . 402 , 4 november , pp . 21 - 22
leslie , s . a . , 2000 : mohawkian ( upper ordovician ) conodonts of eastern north america and baltoscandia . –journal of paleontology : vol . 74 , # 6 , pp . 1122 - 1147
long , j . a . , 1995 : the rise of fishes : 500 million years of evolution . –johns hopkins university press , baltimore & london , pp . 1 - 223
purnell , m . a . , 1994 : microwear on conodont elements and macrophagy in the first elements . –nature : vol . 374 , 27 april , pp . 798 - 800
purnell , m . a . , donoghue , p . c . j . & aldridge , r . j . , 2000 : orientation and anatomical notation in conodonts . –journal of paleontology : vol . 74 , # 1 , pp . 113 - 122
sweet , w . c . , 2000 : conodonts and biostratigraphy of upper ordovician strata along a shelf to basin transect in central nevada . –journal of paleontology : vol . 74 , # 6 , pp . 1148 - 1160
sweet , w . c . & donoghue , p . c . j . , 2001 : conodonts : past , present and future . –journal of paleontology : vol . 75 , # 6 , pp . 1174 - 1184
zhang , s . & barnes , c . r . , 2000 : anticostiodus , a new multielement conodont genus from the lower silurian , anticosti island , quebeg . –journal of paleontology : vol . 74 , # 4 , pp . 662 - 669
zhen , y . y . , nicoll , r . s . , percival , i . g . , hamedi , m . a . & stewart , i . , 2001 : ordovician rhipidognathid conodonts from australia and iran . –journal of paleontology : vol . 75 , # 1 , pp . 186 - 207
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the following descriptions come from benton ( 2005 ) and nelson ( 2006 ) . the structure of the following system is based on sweet and donoghue ( 2001 ) . taxa in red are extinct .
although many genera are recognized , they are based on tooth morphology ; so , relationships between the taxa are artificial .
acodus , amorphognathus , anticostiodus , basselodus , belodina , carniodus , cornuodus , culumbodina , curtognathus , dapsilodus , decoriconus , diaphorodus , drepanodus , erismodus , erraticodon , fahraeusodus , gamachignathus , hamarodus , histiodella , istorinus , juanognathus , jumudontus , kallidontus , lenodus , microzarkodina , oepikodus , oistodus , oulodus , paroistodus , pedavis , phragmodus , polonodus , pseudobelodina , pseudooneotodus , pteracontiodus , pterospathodus , pygodus , scabbardella , scolopodus , scyphoidus , spinodus , staufferella , stereoconus , strachanognathus , streptognathus , taoqupognathus , tripodus , walliserodus , yaoxianognathus .
benton , m . j . 2005 . vertebrate paleontology . third edition . blackwell publishing , malden , ma . [ l ]
donoghue , p . c . j . , p . l . forey , and r . j . aldridge . 2000 . conodont affinity and chordate phylogeny . biological reviews . 75 : 191–251 .
janvier , p . 2008a . early jawless vertebrates and cyclostome origins . zoological science . 25 : 1045 - 1056 .
nelson , j . s . 2006 . fishes of the world . 4th edition . john wiley and sons , inc . new york .
sweet , w . c . and p . c . j . donoghue . 2001 . conodonts : past , present , future . journal of paleontology . 75 : 1174 - 1184 .


graphocephala coccinea is a meadow and woodland-dwelling species of brightly colored leafhopper native to north and central america , from canada south to panama .

common names include candy-striped leafhopper , red-banded leafhopper , scarlet-and-green leafhopper and red-and-blue leafhopper .

g. coccinea adults measure 6.7 – 8.4 mm in length and have vivid blue ( or green ) and red ( or orange-red ) stripes on their wings and the top of their thorax combined with bright yellow coloration on their head , legs , abdomen , and elsewhere .

leafhoppers feed on plant sap with the aid of specialized mouthparts .

graphocephala coccinea

at least three subspecies of this leafhopper have been named : g . coccinea confluenta , g . coccinea punctata , and g . coccinea sambuci .
deniz martinez marked
candy - striped leafhopper
as trusted on the
graphocephala coccinea
page .
deniz martinez added the english common name
candystriped leafhopper
to
graphocephala coccinea ( forster , 1771 )
.
deniz martinez added the english common name
red and blue leafhopper
to
graphocephala coccinea ( forster , 1771 )
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deniz martinez added the english common name
red - banded leafhopper
to
graphocephala coccinea ( forster , 1771 )
.
deniz martinez added the english common name
scarlet and green leahopper
to
graphocephala coccinea ( forster , 1771 )
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the rhododendron leafhopper { graphocephala fennahi } and the scarlet and green leafhopper { graphocephala coccinea } are not the same . . . . there is a difference betweeen them , as described in the text below .
hamilton k . g . a . ( 1985 ) the graphocephala coccinea complex in north america ( homoptera , auchenorryncha , cicadellidae ) . entomologische abhandlungen 49 ( 6 ) : 105 - 11 .
birds have nothing on insects when it comes to being colorful . one of the most outstanding examples of this is the candystriped leafhopper , graphocephala coccinea . it also goes by the names “red - banded leafhopper” and “scarlet - and - green leafhopper . ” there are two subspecies , g . c . coccinea being found only in coastal areas along the atlantic seaboard ; and the slightly larger g . c . quadrivittata , found inland .
g . picta and g . fennahi are more green than red ( at most , equally marked green and red ) whereas g . coccinea is usually dominantly red .
g . c . coccinea ( confined to the atlantic coast ) : ♂♂ 5 . 7 - 6 . 2 mm , ♀♀ 6 . 7 - 7 . 3 mm
the sharpshooters graphocephala versuta , g . coccinea , and oncometopia orbona commonly occur in oklahoma vineyards . of these , g . versuta is the most prevalent species . graphocephala hieroglyphica , cuerna costalis and paraulicizes irrorata also occur in oklahoma , but are rarely collected in vineyards . in research trials , individuals of g . versuta collected within vineyards and those nearby , demonstrated the ability to transmit x . fastidiosa to susceptible plants . however , all of these species are potential vectors of pierce’s disease in oklahoma because they are known carriers of x . fastidiosa .
. exact measurements of body length and sex of individual are essential in being 100 % certain which species you have , as g . teliformis is typically larger than g . coccinea .
photo by : kyle kittelberger , brian bockhahn , paul scharf new hanover co . comment : open woodlands , pine dominated ; male , 6 . 34 mm , g . c . coccinea
graphocephala coccinea – this species is called the candy - striped leafhopper , red - banded leafhopper or red - and - blue leafhopper due to the prominent , alternating red and blue stripes present on the forewings and upper thorax ( pronotum ) . adults measure 3 / 8 inch ( 8 mm ) long and have a yellow head with a black stripe running through the eyes along the margin of the head . the legs and abdomen are bright yellow .
graphocephala hieroglyphica – adults of this species measure 3 / 8 inch ( 8 mm ) long with black and white ( or light red ) stripes running along the forewings and pronotum . the head is white to light red with black markings .
there are a couple known subspecies of g . coccinea ( as mentioned in the description ) : g . c . coccinea , which is found along the atlantic coast and g . c . quadrivittata , which is found inland . there are however intermediates in size between the two subspecies , and one may not be able to determine which subspecies in some instances ( though in nc it should be much more distinctive ) . for more pictures of the two subspecies , see :
g . constricta , known only from ar , w . ms & e . tx , is similar to g . coccinea but is smaller ( 6 . 2 - 7 . 4 mm ) . [ from andy hamilton ' s comments ]
there is usually only one generation of graphocephala coccinea each year in the northern part of its range , but potentially two in southern climates . the winter is spent in the egg stage . eggs are inserted in between layers of leaf tissue by the adult females , with the aid of her blade - like ovipositor . oviposition results in mechanical injury to the leaf , but may also introduce diseases into the host plant . the nymphs that hatch from the eggs molt five times before reaching adulthood .
graphocephala coccinea is distributed throughout north america , occurring from ontario , canada south to florida and west to california and mexico . this species is even less common than o . orbona in oklahoma vineyards as it prefers to feed on blackberry , forsythia , pittosporum , rose , milkweed , goldenrod and other plants . recent sampling efforts in oklahoma vineyards yielded too few specimens to determine peaks in the abundance of this species ; however , it was collected from late may through august , with most individuals captured in june .
several other members of the genus graphocephala resemble the candystriped leafhopper , and there is great variation between individuals of each species . it pays to consult sources like urltoken to learn the differences . i’ll also share information about some of these lookalikes in future “true bug tuesday” posts .
the brilliantly colored candy - striped leafhopper ( graphocephala coccinea ) , is one of the most common leafhoppers of north america’s roughly 2500 leafhopper species . it is native to north and central america , extending from canada south to panama . it goes by several additional common names , including the red - banded leafhopper , scarlet - and - green leafhopper , and red - and - blue leafhopper . it has a particular fondness for blackberry and raspberry leaves , but may also be found on the leaves of a variety of ornamental plants including roses and rhododendrons . the eggs are laid in the leaves .
there are no reports of any of the known insect vectors of xf being present in the midwest , including homalodisca vitripennis ( syn . coagulata ) , draeculacephala minerva , xyphon fulgida , graphocephala coccinea , g . versuta , and g . atropunctata ( ifas , global invasive species database ) . however , the midwest is home to many related xylem feeding , hemipteran insects . many of the known vectors of xf increase their numbers and overwinter on lush grasslands , riparian buffer strips , and alfalfa fields , which also are prevalent in the midwest . of further concern is the observation that alfalfa may be an alternate host for xf ( 55 ) .
description : a medium sized graphocephala , with two subspecies : adult males of g . c . coccinea ( coastal ) are 5 . 7 - 6 . 2 mm long while females are 6 . 7 - 7 . 3 mm , and males of g . c . quadrivittata ( inland ) are 6 . 6 - 7 . 6 mm long while females are 7 . 2 - 8 . 4 mm . this is a brightly colored species with a reddish to orange scutellum . the color of the non - red stripes is variable among individuals , with colors ranging from blue ( as above ) to turquoise to greenish . the top of the head is yellowish to orange in color , with the underside of the body yellow , and there is a bold black line going around the side of the face between the eyes . ( bg )
graphocephala versuta – commonly referred to as the versute sharpshooter , adults measure 3 / 16 inch ( 5 mm ) long and have green forewings with several black , tooth - like markings at the wing tips . a light blue stripe runs diagonally along each forewing and extends to the front of the head . this blue stripe is bordered on both sides by an orange stripe in some individuals . the top of the head is light brown to orange and has a light blue stripe along the midline .
graphocephala versuta is common in the southern u . s . , ranging from as far west as texas , north to illinois . it is active in oklahoma vineyards from early june through late august with a distinct peak in abundance from late june through mid - july . adults overwinter in woody habitats and move to herbaceous perennials in march . females lay eggs on the lower leaf surface of ragweed , sunflower , okra and other species . this insect feeds on blackberry , grape , honeysuckle , privet , cherry and other deciduous trees . while this species occurs throughout most of the growing season , inoculative individuals ( i . e . , those capable of transmitting x . fastidiosa to susceptible plants ) are not present until mid to late june . there appears to be one generation per year in oklahoma .
in coastal california , the principal vector of pd was graphocephala atropunctata , known as the blue - green sharpshooter ( bgss ) ( 52 ) . bgss overwinters in riparian vegetation but readily feeds and multiplies on grapevines . these insects transmit xf in a persistent manner , with a short or even no latent period required between acquisition and transmission ( 44 ) . in the southeastern usa , another sharpshooter , homalodisca vitripennis germar ( fig . 2 ) ( 61 ) , was identified as a vector of xf . homalodisca vitripennis [ formerly homalodisca coagulata ( 60 ) ] , popularly known as the glassy - winged sharpshooter ( gwss ) , was introduced to california in the 1990s . bgss and gwss have similar feeding habits . however , bgss prefers to feed on young leaves and tissues while gwss , although it can exist on these tissues , is also found on woody parts of the plants ( 29 ) . at present , gwss is considered to be the most efficient vector of pd because it displays greater mobility , is capable of feeding on older woody stems , and has a wider host range than the native sharpshooters ( 33 ) . its establishment in california threatens the overall wine grape , table grape , and almond industries ( 48 ) .
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checklist of the hemiptera of canada and alaska maw , h . e . l . , r . g . foottit , k . g . a . hamilton and g . g . e . scudder . 2000 . nrc research press .
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pierce’s disease of grapes ( vitis spp . ) is caused by the bacterial plant pathogen , xylella fastidiosa subsp . fastidiosa . this disease is a serious concern for grape growers because once vines are infected , there is no cure and vines will eventually die . since 2009 , pierce’s disease has been detected in several oklahoma vineyards , and there is concern that oklahoma grape producers will experience heavy losses similar to california and texas . thus , pierce’s disease poses a threat to the vitality of viticulture in oklahoma , and measures must be implemented to reduce its impact in the state .
pierce’s disease can spread several ways . sharing cuttings with other growers may lead to the accidental introduction of infected vines to new vineyards . once pierce’s disease is established , there is potential for spread among vines via pruning shears . also , vine - to - vine transmission is aided by insect vectors . key vectors include several species of sharpshooters , a group of leafhoppers that feed on xylem tissue where x . fastidiosa resides . these sucking insects are the most common source of disease transmission . because they feed on a wide variety of plants that can harbor x . fastidiosa , weed management within the vineyard is an important strategy to reduce disease transmission . effective management of pierce’s disease depends on correct identification of key vector species , monitoring their presence in the vineyard , and applying broad - spectrum insecticides when peak numbers are detected .
oncometopia orbona – this species is known as the broad - headed sharpshooter , presumably for its wide head . adults measure 1 / 2 inch ( 12 mm ) and have an orange head , eyes , legs and abdomen . the forewings and pronotum have light blue and black speckles . occasionally , females have white patches called brochosomes on the sides of the forewings . brochosomes contain proteins that females smear onto their eggs to prevent desiccation .
paraulicizes irrorata – this species is among the largest encountered in vineyards , with adults measuring 9 / 16 inch ( 14 mm ) long . the common name , speckled sharpshooter , refers to the mottled appearance of the forewings and pronotum . the body and head are dark , with white to pale yellow markings scattered along the lateral margins . the front of the head may have parallel orange markings .
cuerna costalis – this colorful species measures 5 / 16 inch ( 10 mm ) long . the forewings have narrow , alternating black and red stripes with a thicker red stripe along the wing margins . the head and pronotum are black with white markings and a white stripe extends from the front of the head , through the eyes , to the end of the abdomen . the legs are red and black .
oncometopia orbona is distributed widely throughout the eastern u . s . from florida to maryland , and south from missouri and texas to northern mexico . it is found in oklahoma vineyards , but less frequently than g . versuta . this sharpshooter prefers feeding on sunflower , hollyhock , okra , lambsquarter , ash , oak , silktree , peach and crapemyrtle . based on data extrapolated from oklahoma tree nurseries where o . orbona is more abundant , this species is likely to be active in oklahoma vineyards from late may through late august with a distinct peak in abundance in late july . there appears to be one generation per year in oklahoma .
successful management of pierce’s disease depends on effective control of potential insect vectors and eliminating weeds that may serve as sources of future infections . while all of the sharpshooter species described in this publication have potential to carry and transmit x . fastidiosa to grapevines , monitoring efforts should focus on the versute sharpshooter because of its high prevalence in oklahoma vineyards and demonstrated ability to transmit x . fastidiosa to susceptible plants . based on several years of sampling data from oklahoma vineyards , monitoring for the versute sharpshooter should occur from early june through late august . the best monitoring tools for jumping and flying insects like sharpshooters are sticky cards . sticky cards can be purchased online from garden and greenhouse supply companies . purchase yellow sticky cards that measure approximately 3 inches by 5 inches and have adhesive on both sides . sticky cards should be attached to the upper vine support cable using a clip or clothespin and spaced evenly around the perimeter of the vineyard and on diagonals through the vineyard center . for most oklahoma vineyards , which are two acres or less , 12 to 16 sticky cards should be sufficient for monitoring versute sharpshooters . sticky cards should be checked at least twice per week , and changed every two weeks . due to the small size of these insects , use a hand lens with at least 15x magnification to identify specimens caught on sticky cards . if identification is in doubt , check with the local county extension office for assistance .
table 1 . inse cticide products available to control insect vectors of pierce’s disease in oklahoma vineyards .
do not apply more than 3 . 2 fl oz / a per 14 - day interval
† restricted use pesticide ; a pesticide applicators license is required to purchase and apply this product .
1 insecticide resistance action committee chemical subgroup ; rotate among products with different irac numbers to reduce likelihood of insecticide resistance developing in target pest population .
2 re - entry interval ; do not allow workers to enter treatment area of crop following application for the minimum amount of time indicated .
3 pre - harvest interval ; do not apply less than the number of days indicated prior to harvest .
oklahoma state university , in compliance with title vi and vii of the civil rights act of 1964 , executive order 11246 as amended , and title ix of the education amendments of 1972 ( higher education act ) , the americans with disabilities act of 1990 , and other federal and state laws and regulations , does not discriminate on the basis of race , color , national origin , genetic information , sex , age , sexual orientation , gender identity , religion , disability , or status as a veteran , in any of its policies , practices or procedures . this provision includes , but is not limited to admissions , employment , financial aid , and educational services . the director of equal opportunity , 408 whitehurst , osu , stillwater , ok 74078 - 1035 ; phone 405 - 744 - 5371 ; email : eeo @ urltoken has been designated to handle inquiries regarding non - discrimination policies . any person who believes that discriminatory practices have been engaged in based on gender may discuss his or her concerns and file informal or formal complaints of possible violations of title ix with osu’s title ix coordinator 405 - 744 - 9154 .
issued in furtherance of cooperative extension work , acts of may 8 and june 30 , 1914 , in cooperation with the u . s . department of agriculture , director of oklahoma cooperative extension service , oklahoma state university , stillwater , oklahoma . this publication is issued by oklahoma state university as authorized by the vice president , dean , and director of the division of agricultural sciences and natural resources .
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l . c . galvez , k . korus , j . fernandez , and j . l . behn , department of plant pathology , and n . banjara , department of agronomy , university of nebraska lincoln , lincoln , ne 68583 - 0722 galvez , l . c . , korus , k . , fernandez , j . , behn , j . l . , and banjara , n . 2010 . the threat of pierce’s disease to midwest wine and table grapes . online . apsnet features . doi : 10 . 1094 / apsnetfeature - 2010 - 1015 .
grape production in the united states is expanding rapidly , especially as new growers are starting vineyards in many regions of the country . commercial wine grape production in the midwest , along with production in california , oregon , and new york , has increased along with the demand for high - quality wine . there are more than 652 commercial wineries throughout the midwest region ( 11 , 17 ) , defined by the u . s . census bureau as the 12 states in the central and northeastern united states . the number of wineries in the united states has steadily increased over the past twenty years and currently exceeds 6000 ( 36 ) .
the wine industry has expanded considerably in recent years in the midwest ( 15 , 17 ) . this increase can be attributed to the growing demands for wine both in local and international markets . ohio has the most wineries of the midwest states ( table 1 ) . wineries are considered a new industry in minnesota , where there were 3 wineries in 1997 , expanding to 36 wineries at present ( 21 ) . nebraska and surrounding states are developing rapidly as table grape and wine producers . wisconsin had 24 wineries in the early 2000 and now has around 50 wineries ( 66 ) . kansas , south dakota , and north dakota have the least number of wineries , with 17 , 14 , and 5 , respectively ( 17 ) . this increase in wineries corresponds to the increase in vineyards to meet the demands for wine production .
table 1 . winery numbers in the midwest united states ( 14 , 16 ) .
grape cultivars grown in the midwest are selected on the basis of vine hardiness , fruit characteristics , ripening time , tolerance to diseases and insects , vine characteristics , market outlet , and requirements of the processor or consumer who will purchase the crop . the comparison of popular grape cultivars grown in the midwest to those of california reveals almost no overlap ( table 2 ) . some of these varieties show differences in relative susceptibility to powdery mildew ( erysiphe necator ) , botrytis bunch rot and blight ( botrytis cinerea ) , phomopsis cane and leaf spot ( phomopsis viticola ) , eutypa dieback ( eutypa lata ) , and crown gall ( agrobacterium vitis ) . only a few of the cultivars grown in the midwest have been tested for resistance to xylella fastidiosa , the causal agent of pierce’s disease [ ( 49 ) as cited by ( 63 ) ] .
table 2 . comparison of grape cultivars grown in california and the midwest us .
dechaunac concord esprit edelweiss frontenac ( mn 1047 ) jupiter lacrescent ( mn 1166 ) leon millot mars reliance st . croix swenson red vidal blanc vignoles
lacrosse leon millot limberger marechal foch melody niagara seyval blanc st . pepin traminette ventura vidal blanc vignoles reisling st . croix
xylella fastidiosa ( xf ) is a fastidious gram - negative , xylem - limited bacterium associated with a large number of diseases including pierce’s disease , phony peach disease , plum leaf scald , citrus variegated chlorosis , scorch of almond , oleander leaf scorch , and several other diseases ( 2 , 10 , 29 , 38 , 54 ) . this bacterium has a very wide host range and has been reported to cause diseases in over 100 plant species of both monocots and dicots ( 29 ) . different subspecies or strains of xf can cause disease in different plants , and in some cases the bacterium can inhabit the plant without causing symptoms . the fastidious bacterium resides in the plant xylem tissue or water conducting vessels and is spread exclusively by xylem - feeding insects known as sharpshooter leafhoppers ( 26 ) . pierce’s disease is known to be prevalent within the united states from florida to california , and in other countries throughout central and south america .
in 1892 , newton pierce was the first plant pathologist to describe the
mysterious vine disease
eventually known as pierce’s disease ( pd ) . since the 1880s , xf has been reported to cause considerable losses in southern california , destroying more than 35 , 000 acres of vineyards . as a result , nearly all viticulture was forced to move northward due to the inability to produce a quality grape in this area due to pd .
fig . 1 . symptoms of pierce ' s disease in grapevines caused by xylella fastidiosa ( courtesy a . c . goheen ; reproduced from compendium of grape diseases , 1988 , american phytopathological society , st . paul , mn ) .
the vectors . for quite some time , pd was thought to be caused by an insect - borne virus because of its transmissibility via xylem - feeding insect vectors and via grafting ( 24 ) . similarly , no fungi or bacteria were isolated from pd infected plants ( 25 ) . the reemergence of pd in the southern california vineyards enabled researchers to identify xylem - feeding vectors such as sharpshooter leafhoppers belonging to the subfamily cicadellinae ( 51 ) and spittlebugs from the family cercopidae ( 14 ) .
fig . 2 . the glassy - winged sharpshooter leafhopper vector of xylella fasidiosa ( courtesy b . bextine , fletcher and wayadande , 2002 , the plant health instructor ) .
pathogen - vector - host interaction . three events are essential for the successful transmission of the pathogen to the plant . first is the acquisition of the pathogen by a vector from a diseased host plant . acquisition efficiency is directly proportional to the population of the bacteria from the infected hosts ( 26 , 48 ) . similarly , efficiency of acquisition also depends upon the sharpshooter feeding site whether it be leaves , stems , or older woody branches , however , no report suggests a preferred feeding site by the vector ( 26 ) . the second event is the attachment or retention of the acquired bacteria into the vector foregut . de la fuente and co - workers ( 13 ) suggested that type i and type iv pili are involved in the attachment to the cuticle within an environment described as a
fast fluid flow
condition ( 13 ) . microscopy studies revealed that the bacteria form a biofilm in the foregut of the sharpshooters where they are polarly attached to the cuticle , with the cuticle being shed at each molt ( 43 ) . lastly , the pathogen must be detached and introduced into a new host during vector feeding . the lack of a latent period suggests that the population of bacteria within the vector is not a significant factor affecting the transmission efficiency ( 1 ) ( fig . 3 ) .
chatterjee and co - workers ( 7 ) used green flourescent protein ( gfp ) - tagging of xf to monitor the correlation between the colonized vessels and symptom expression . they found a five - fold increase in colonized vessels from symptomatic leaves compared to symptomless leaves ( 7 , 42 ) . it was also observed that vessels in symptomatic leaves contained large bacterial colonies while asymptomatic leaves from infected plants had vessels that contained very small colonies of bacterial cells . this finding suggested that vessel blockage determined symptom expression .
like many plant pathogenic bacteria , xf harbors a cell - to - cell signaling system which plays an important role in colonization , pathogenesis , and biofilm formation ( 62 ) . a set of genes known for regulation of pathogenic factor ( rpf ) is required for the synthesis of diffusible signaling factor ( dsf ) , involved in the formation of biofilm in the foregut of the vector ( 7 ) . rpf was also involved in colonization of both the vector and the host ( 43 ) and pathogenesis in the host ( 7 ) . genomic characterization of xf also revealed its richness in adhesins and hemaglutinin - encoding genes that are involved in the interaction with vectors and hosts , and cell - to - cell attachment in biofilm formation ( 8 ) . whether these virulence factors are secreted via type i or type ii secretion systems is not yet clear , but sequence analysis reveals that xf possesses genes for both type i and type ii secretion systems for the export of exoenzymes involved in plant cell wall degradation and allow bacterial colonization of plant xylem ( 30 , 53 ) .
the primary factor to consider with respect to the probability of pd development in the midwest is climate . in the years since the discovery of pd , reports of its presence have been restricted to areas with a sub - tropical climate . in the united states , its distribution has been mostly limited to the southernmost grape growing states and west coast , with the greatest distribution and greatest disease severity occurring in florida and california . the correlation between disease incidence and the concentration of vineyards may be coincidental . however , this pattern of disease seclusion to sub - tropical climates appears to be a result of the intolerance of xf and its respective vectors to cold temperatures . even though xf is favored by a sub - tropical climate , it also is sensitive to elevated temperatures , and growth above 34ºc does not occur ( 16 ) .
many xylem feeding insects found throughout the midwest are known to feed on the younger , greener tissues of their respective host plants . theoretically , if xf and a suitable vector were introduced into the midwest , the insect would likely also feed on the younger tissues . if xf is introduced to these parts of the grape vine , disease severity is expected to be low and overwintering inside the plant hindered because of vine pruning practices . the risk of disease in the midwest , aside from the effect of temperature , relies on two important parameters : the introduction of xf ; and the association between xf and a xylem - feeding insect pest of grapes .
little work has been done to correlate the effects of severe cold weather , such as seen in the midwest during a typical winter , and its effect on the survival of either h . vitripennis or xf . however , there is a relationship between low temperatures ( ~ 5ºc ) and the corresponding growth rate of xf . henneberger and stevenson ( 22 ) isolated xf from the roots and shoots of sycamore trees ( a natural alternate host ) at various times throughout the year at varying temperatures . they constructed a plot correlating the frequency of xf isolation with the respective temperature when the isolation was attempted which revealed a decrease in isolation success as the temperature approached 0ºc . these data suggest that survival of xf is minimal at temperatures below freezing . in a similar study , feil and purcell ( 16 ) found a similar correlation between temperature and the growth rate of xf . they were able to show that , in vitro , the optimal growth conditions for this bacterium ranged from 25 to 32ºc and that growth declined and was even suppressed when temperatures fell below 12ºc .
the above findings conflict with the discovery of a xf strain that causes mulberry leaf scorch ( 9 ) . based on restriction fragment length polymorphism ( rflp ) and random amplified polymorphic dna ( rapd ) molecular genome assays , it was found that this strain is closely related to the subspecies that causes pd . furthermore , the mulberry trees from which the bacterium was isolated were growing naturally in two different locations , namely massachusetts and nebraska , suggesting that this particular strain has adapted to the harsh temperature fluctuations of the midwest and northeast .
more recently , a report from oklahoma revealed that the table grape cultivar ‘concord’ displayed typical symptoms of pd . real - time pcr analysis of dna isolated from vine samples using available specific primers coupled with elisa confirmed the presence of xf ( 57 ) . this finding in oklahoma has caused concern among midwest wineries because of its geographical proximity . the recent identification of xf in grapes suggests the possibility that the strain of xf infecting woody trees was able to overcome previous grape cultivar resistance . currently there are no studies to investigate the ability of this strain to infect any of the widely grown grape cultivars throughout the midwest . until this is done , it will not be possible to extrapolate the possibility of pd on the midwestern grape and wine industry .
climate change adaptations . over the last 100 years average temperatures across the northern midwest have increased ca . 4ºf ( 2ºc ) , and average temperatures have decreased in the southern midwest by 1ºf ( 0 . 5ºc ) . these trends are projected to continue ( 39 ) . a shift to warmer temperatures throughout the midwest may enable establishment of organisms that would otherwise be unable to exist in these areas . an increase in the minimum temperature by 10ºf ( 12ºc ) might shift a particular region’s climate such that freezing would be avoided altogether . the pathogen and vector may also be able to adapt in the presence or absence of climate change . rising temperatures would also lengthen the growing season , and potentially increase drought conditions throughout the midwest . in combination , these elements could enable an increased proliferation of insect pests and pathogens . drought stress not only increases the probability of pd incidence , but may also increase the expression of disease symptoms on stressed plants due to compromised defense system . in fact , mcelrone et al . ( 34 ) revealed that occlusion of the xylem vessels by xf was exaggerated when host plants experienced drought stress ( 19 , 34 ) .
there are several disease management strategies used with some success in areas with severe pd . these include host resistance , vector control , and changes in cultural practices . used in combination as an integrated pest management strategy , they should prove to be fairly successful in preventing the infection and spread of xf in midwestern vineyards .
the most popular grape cultivars of european and american descent are susceptible to pd to some degree . however , there are a few native species found in the southeastern part of the usa that have shown resistance to xf . muscadine grape ( vitis rotundifolia ) cultivars have varying degrees of resistance to x . fastidiosa , but are not immune to the pathogen . although some muscadine grape cultivars may be highly resistant to pd , they lack the fruit quality that most popular cultivars offer . also , vitis vinifera cultivars have shown differences in pd resistance in california , with ‘petit sirah , ’ ‘chenin blanc , ’ and ‘sylvaner’ among the most resistant cultivars ( 29 ) . these cultivars may be useful for resistance gene introduction into popular wine and table grape cultivars .
potential transgenic grapes resistant to pd are highly controversial and are not currently considered within the grape industry ( 40 ) . wine - making , an
old world
industry based on historical european wine regions , carries with it original wine grape names representing different cultivars , tradition , and prestige . the general opposition to genetically altering the classic cultivars is that they will not be the same cultivar anymore , hence losing their status among the top wine - making grapes . plant breeders and the skeptical wine consumers are at a standstill , involving regulatory and marketing issues , with the growers left in the middle , scrambling for a better solution to pd . if resistance genes can be identified and introduced into a desirable but susceptible grape genome , new cultivars may be produced having varied levels of resistance to pd which may or may not be accepted in the trade .
with management of any mobile pest , such as the gwss , timing is crucial . targeting the first generation of gwss is most effective in controlling xf transmission ( 29 ) . these studies have shown that an early insecticide application before the appearance of the first generation of gwss helps to slow pd spread . insecticides may also need to be applied to surrounding vegetation containing hosts of gwss and other insect vectors of xf . although effective for shorter annual growth periods , it is less effective in areas with longer growing seasons and warmer climates , such as those experienced in florida ( 29 ) . because systemic insecticides last a few months inside plants , a second application may be needed in areas where the growing season is extended .
several insecticides ( i . e . , foliar - applied acetamiprid and soil - applied imidacloprid and thiamethoxam ) have been tested for control of gwss and other xf transmitting insects ( 3 ) . the most effective insecticides for insect control , inhibiting bacterial transmission , and pd spread are the neonicotinoids ( 1 ) . these insecticides are systemic and aid in disrupting the h . vitripennis - x . fastidiosa interface .
pierce’s disease of grapes has recently been reported in oklahoma , adjacent to the lower midwest region . while pd is unlikely to become a serious problem in the near future , this could change with adaptation of both the vector and the pathogen to colder climates and introduction of susceptible cultivars . with a warmer climate predicted for the midwest , it is also possible that climate would not be a limiting factor in the future . this review of the current status reveals that growers , consultants and investigators associated with grape and wine production are at some risk of pd and should plan accordingly . such plans should include periodic surveys , determination of susceptibility of current cultivars and potential forward breeding for resistance to this potentially devastating disease .
this review is one of the final requirements of the course plant path 867 . the authors were all enrolled in this course , and wish to acknowledge their professors drs . anne vidaver and gary yuen and drs . fred baxendale , robert harveson , paul read , for their valuable comments and suggestions on the manuscript .
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amiskwia is a genus of large , soft-bodied animals of unknown affinity known from fossils of the middle cambrian lagerstätten both in the burgess shale formation in british columbia and the maotianshan shales of yunnan province , china .

very few specimens of this organism have been found , only eighteen of the burgess shale species , a. sagittiformis , and a single specimen of the maotianshan species , a. sinica — which may be a reflection of its genuine rarity , but is more likely to be due to taphonomic ( preservational ) or behavioural factors .

the fossils reach 25 mm ( 1 in ) in length .

the head is rounded , tipped with two tentacles , and appears to contain a four-ganglion brain ; the body flattens out and broadens in the trunk , which appears to have been fairly muscular .

where the trunk meets the head there is a small tubular opening , which can be interpreted as the mouth ; the gut terminates where the trunk narrows and meets the tail , which is broad and paddle shaped .

the body morphology suggests a free energetic swimmer , which may be consistent with the dearth of fossils ( in other words , a free-swimming animal would be able to avoid being buried alive in a mudflow on a regular basis , compared to a benthic animal ) .

amiskwia was originally categorized by paleontologist charles walcott .

walcott thought he saw three buccal spines in the fossils , and therefore categorized amiskwia as a chaetognath worm ( arrow worm ) .

however , amiskwia appears to lack the characteristic grasping spines and teeth of other burgess fossil arrow worms .

later scientists suggested an affinity with the nemerteans ( ribbon worms ) , but the evidence for this was somewhat inadequate .

conway morris , on re-examining of the burgess shale fauna in the 1970s , described it as being the single known species in an otherwise unknown phylum , given that it has two tentacles near its mouth , rather than the characteristic single tentacle of true nemerteans .

( nemerteans do not have a single tentacle . however , a pair of antero-lateral tentacles is present in 2 of the many genera of pelagic nemerteans . nemerteans do have a single eversible — normally internal — proboscis , which when everted could resemble an anterior median tentacle if fossilized . whether retracted or everted , the proboscis is the only structure in pelagic nemerteans likely to fossilize , as it is the only structure with substantial connective tissue and muscle . the body wall has almost no muscle or connective tissue and is exceedingly unlikely to fossilize ; hence , a pelagic nemertean fossil would be only the proboscis ) .

butterfield implies from the appearance of the fossils that the organisms may have lacked a cuticle : while this is also true of the nemerteans , these organisms lack a coelom and are thus unlikely to fossilise .

he goes on to argue that the absence of cuticle is characteristic of the chaetognaths ; whilst teeth would be expected , a similar fossil , wiwaxia , shows such structures in only 10 % of the expected instances , and anomalocaridids are often found detached from their mouthparts , so the absence may be taphonomic rather than genuine .

the absence of spines could simply mean that the fossils represent young organisms — or that later chaetognath evolution involved paedomorphosis .

while more work on undescribed specimens is required before firm consensus is reached , the discovery of an undisputed chaetognath , protosagitta spinosa , which strongly resembles modern chaetognaths , in the similarly aged chengjiang biota places doubts on amiskwia 's position in the chaetognath crown group .

it has also been likened to the molluscs , and bears limited resemblance to unusual forms such as pterotrachea .

of course , many similarly enigmatic cambrian fossils probably represent stem groups to living taxa , and there is no reason to assume that this organism must fall into an extant crown group at all .

the scientific name amiskwia sagittiformis derives from the nearby amiskwi river , and its sagittiform shape .

" sinica , " of a. sinica , refers to that species ' origin from china .

amiskwia

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amiskwia sagittiformis ( ah - miss - kwi - ah sa - jih - tih - form - is ) . after amiskwia , a river west of mt . burgess + sagittiformis ( l . ) = arrow - shaped .
amiskwia – from the cree amiskwi , “beavertail , ” a name given to various topographical features in yoho national park .
amiskwia is a rare fossil because only 18 specimens have been found in the burgess shale . because of its hydrodynamic shape and its powerful tail , amiskwia looks like it could have been a fast swimmer . maybe the reason why amiskwia is rare could be that they were territorial and living very far away from each other . but that ' s just my hypothesis .
amiskwia looked like a fat worm with a head that had two antenna - like tentacles on it . on the side it had stubby fins , and on the back end it had a paddle - like tail possibly for locomotion . specimens of amiskwia are almost transparent so that the internal organs are visible .
note the rather conspicuous pair of bulb - like organs in the head of this amiskwia reconstruction : this is the artist ' s depiction of its cerebral ganglia .
both amiskwia and opabinia would have quivered in fear at the sight of a meter - long anomalocaris ! ! it just goes to show that size exists only by way of comparison .
amiskwia was a bizarre genus of cambrian animal . it was about one inch long and probably used its flat tail to swim . when charles doolittle walcott first discovered amiskwia in the burgess shale , he described it to be a chaetognath , sometimes called an arrow worm . then later , other scientists classified it as a ribbon worm . but now no one really knows what it was .
conway morris , s . 1977 . a redescription of the middle cambrian worm amiskwia sagittiformis walcott from the burgess shale of british columbia . palaontologische zeitschrift , 51 ( 3 ) : 271 - 287 .
owre , h . b . and f . m . bayer . 1962 . the systematic position of the middle cambrian fossil amiskwia walcott . journal of paleontology , 36 ( 6 ) : 1361 - 1363 .
amiskwia ‘s entire body is dorso - ventrally compressed . its head sports a pair of ventrally - situated tentacles , just behind and between which lies an ovular mouth . though the creature’s “trunk” is unsegmented , it does contain a pair of lateral fins which are unsupported by any sort of stiffening device : the same is true for the caudal fin which comprises the beast’s tail . the trunk and caudal regions of amiskwia ‘s anatomy appear to have been quite muscular : a piece of evidence which supports the hypothesis that amiskwia may have been a pelagic ( free - swimming ) organism , which would also help to explain its aforementioned rarity in the formerly muddy burgess basin .
the phylogenetic relationships of amiskwia are a matter of some debate . however , the general consensus amongst cambrian paleontologists is that while the genus exhibits characteristics akin to those of the chaetognatha and nemertea . it cannot be scientifically accomodated by either group .
the presence of fins demonstrates that amiskwia was well adapted for swimming . its rarity in the burgess shale suggests that it may have spent much of its time well above the sea bed , above the depth at which it could be caught in submarine mudslides .
amiskwia shows us three definite body segments : a head with two prominent tentacles , an unsegmented trunk with stubby side fins , and a flattened tail . the fins and tail suggest this was an active swimmer ( also suggested by its rare appearance in the burgess shale formation ) . walcott first described this fossil in 1911 and assigned it as a chaetognath ( arrow worm ) . however , amiskwia appears to lack the characteristic grasping spines and teeth of other burgess shale fossil arrow worms , so later scientists suggested it was more likely a nemertean ( ribbon worm ) . however , key characteristics of that group are also absent , so amiskwia remains another of the burgess shale weird wonders with no obvious relationship ( affinity ) to any other living or extinct group . fossil sizes up to one inch .
the soft bodied and rather delicately - built amiskwia sagittiformis is a superb example of the conflict which often arises when a fossil is subjected to exceptional preservation and a debatable amount of deformity . the soft composition and meager proportions of amiskwia and its kin prevent their fossil record from becoming remotely extensive . hence , we may never know the degree to which geologic forces have altered the appearance of this superficially worm - like animal . nonetheless , enough information has been gathered to make a few tentative educated guesses concerning its lifestyle and possible taxonomic affiliations .
amiskwia is a symmetrical , flattened worm . it bears a pair of lateral fins in addition to a paddle - like tail fin . a pair of small tentacles is situated on the bottom of its head , just in front of its mouth . the trace of a gut and other internal organs are preserved in the fossils .
opabinia was also a giant , at a mighty one inch long ! but amiskwia is one of the burgess critters who really strikes me as alien . maybe one day my museum could have a big walk through of the cambrian with the animals blown up to our scale . this guy would certainly get a place of honor .
described by walcott in 1911 , amiskwia was originally interpreted as an arrow - worm ( walcott , 1911 ) . originally popular , this interpretation fell into disrepute after further studies ( conway morris , 1977 ; owre and bayer , 1962 ) , but it has more recently been reconsidered as a possible arrow - worm ( butterfield , 1990 ) .
though the statement may sound comical , at a full inch in length , amiskwia was a comparatively large inhabitant of the burgess shale and the elder maotianshan shales . this considerable size appears to have come at the expense of abundance , with a mere eighteen specimens of this peculiar genus having been found in the former deposit as of this writing .
the phylogenetic position of amiskwia is uncertain . despite significant objections to its traditional interpretation as a chaetognath ( conway morris , 1977 ; owre and bayer , 1962 ) , some workers still hold this view ( butterfield , 1990 ) . nemertine ( owre and bayer , 1962 ) and molluscan ( chen and huang , 2002 ; chen et al . , 2005 ) affinities have also been suggested , but not substantiated .
amiskwia sagittiformis walcott wird auf der basis von fünf erhältlichen exemplaren neu beschrieben . der dorsoventralisch komprimierte körper besteht aus einem kopf mit einem paar hervorstehender fühler und einem rumpf mit seiten - und schwanzflossen . ein darm mit subterminalen dffnungen sowie gehirnganglien , nervenstränge , blutgefäße und muskeln konnten mit variierendem sicherheitsgrad identifiziert werden . eine aktive pelagische lebensform wird als wahrscheinlich angenommen . die abgrenzung dieses wurms von den pfeilwürmern ( chaetognatha ) , die in früheren arbeiten unternommen wurde , wird bestätigt , aber deren klassifizierung als nemertinen ( nemertea ) kann aufgrund fehlender kritischer daten nicht aufrechterhalten werden . die systematische einordnung der a . sagittiformis bleibt weiterhin ungelost .
amiskwia sagittiformis walcott is redescribed on the basis of the five available specimens . the dorsoventrally compressed body consists of a head bearing a pair of prominent tentacles and a trunk with lateral and caudal fins . a gut with subterminal openings , cerebral ganglia , nerve cords , blood vessels and muscles have been identified with varying degrees of confidence . an active pelagic mode of life is considered probable . the rejection by earlier workers of this worm from the arrow - worms ( chaetognatha ) is confirmed , but their identification of it as a nemertean ( nemertea ) cannot be supported owing to the absence of critical data . the systematic position of a . sagittiformis remains unresolved .
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paleobiology : a quarterly journal of the paleontological society . ( ejournal / emagazine , 1975 ) [ worldcat . org ]
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paläontologische zeitschrift : wissenschaftliche beiträge der paläontologischen gesellschaft . ( journal , magazine ) [ worldcat . org ]
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paläontologische zeitschrift : wissenschaftliche beiträge der paläontologischen gesellschaft . / paläontologische gesellschaft . ; ; stuttgart : e . schweizerbart ' sche .
an early cambrian problematic fossil : vetustovermis and its possible affinities . - pubmed - ncbi
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nanjing institute of geology & palaeontology , chinese academy of sciences , nanjing 210008 , china . chenjunyuan @ urltoken
the early cambrian problematic fossil vetustovermis ( glaessner 1979 alcheringa3 , 21 - 31 ) was described as an annelid or arthropod . anatomical analysis of 17 new specimens from the lower cambrian maotianshan shale at anning , kunming ( south china ) does not support its affinities with annelids or arthropods . anatomical features instead resemble other animal groups including modern flatworms , nemertines and molluscs . the presence of a pelagic slug - like form and ventral foot , as well as a head with eyes and tentacles indicates a possible affinity with molluscs , but these characters are not present only in molluscs ; some of them are shared with other animal groups , including flatworms and nemertines . for example , a ventral foot - like structure is found in nemertines , ' turbellarians ' , and some polychaete groups . the well differentiated head is seen in separate bilaterian groups , but among molluscs it did not occur before the evolutionary level of the conchifera . unlike the ctenia - gills in molluscs , the gills in vetustovermis are bar - like . all the characters displayed in this 525 million - year old soft - bodied animal fail to demonstrate clear affinity with molluscs or any other known extant or extinct animal groups , but argue for representing an independently evolved animal group , which flourished in early cambrian and possibly in middle cambrian time .
pmid : 16191609 pmcid : pmc1559895 doi : 10 . 1098 / rspb . 2005 . 3159
anatomical features of the early cambrian mollusc - like organism vetustovermis planus glaessner 1979 . ( a ) dorsal view ; ( b ) reconstructed cross section . abbreviations ( also used in and ) : a , anus ; b , buccal mass and stomach ; d , diverticula ; da , dorsal aorta ; e , eye ; f , foot ; fn , fin ; g , gills ; h , head ; i , intestine ; m , mantle ; o , outer margin of foot ; p , spine - like structure ; t , cephalic tentacles .
vetustovermis planus glaessner 1979 from the lower cambrian maotianshan shale , at shankou village , anning , kunming . ( a ) and ( b ) a nearly complete specimen ( coll . no . sk70001 ) with rock split along a plane near to the dorsal surface to separate it into a part ( a ) and a counterpart ( b ) , showing : a round small head ( a ) that carries a pair of cephalic tentacles ( b ) and stalked eyes ( a and b ) ; a buccal mass and stomach with anterior diverticula ( a ) ; lateral margins of a large ventral foot that lie near the outer margins of the trunk ( b ) ; the imprints of dorsal spine - like structures ; dorsal artery intestine ; and an anus . ( c ) a trunk , showing gills and a mud - filled intestine ( coll . no . sk70010 ) . ( d ) ventral view of an animal showing a large foot with its outer margins near margins of the trunk proper ( coll . no . sk70011 ) . ( e ) a sagittal plane near to dorsal surface ( coll . no . sk7002a ) showing gills , lateral fins , dorsal aorta and an intestine with an anus . ( f ) a sagittal plane near to dorsal surface ( coll . no : sk70007 ) showing cephalic tentacle , eye , lateral fins , and a dorsal artery . ( g ) enlargement of figure 2 a , showing conic ( red arrow ) and rounded ( white arrow ) imprints of spine - like structures . ( h ) head of an animal ( coll . no . sk70012 ) showing a round head , a three dimensionally preserved buccal mass and stomach with anterior diverticula . ( i ) a sagittal plane near ventral surface of a trunk ( coll . no . sk70009b ) showing ventral foot with its outer margins and the gills in lateral area of the trunk proper ; intestine ; and folded lateral fins . ( j ) a sagittal plane near dorsal surface of a trunk ( coll . no . sk70013 ) showing gills extended axially near to the centre and some of the gills bent ; the imprints of spine - like structures ; and a median dorsal aorta . ( k ) ventral view of a trunk ( coll . no . sk70003 ) showing ventral foot and its outer margins , and an anus near terminal end of the trunk . ( l ) a sagittal plane near dorsal surface of a trunk ( coll . no . sk70008a ) showing imprints of spine - like structures , lateral fins and an intestine . scale bars , all are 5mm except for ( g ) which is 1 mm .
interpretive drawing of vetustovermis planus glaessner 1979 . diagrams are the following figure parts : ( a ) a ; ( b ) b ; ( c ) e ; ( d ) f ; ( e ) l ; ( f ) j ; ( g ) .
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( 1968 ) : the free - living lower invertebrates . — 229 pp . ; new york .
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, a new organism from the middle cambrian burgess shale of british columbia . — n . jb . geol . palaont . mb . . ,
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und beschreibung einer neuen art . — veroff . oberseemus . bremen , reihe a ,
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, r . c . [ ed . ] : treatise on invertebrate paleontology , part w , miscellanea , w 144 - w 177 , lawrence ; kansas .
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— ( 1959 ) : the invertebrates . vol . 5 . smaller coelomate groups . — 783 pp . ; new york .
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( 4 ) : 115–118 ; falls church , virginia . [ agi transl . of paleont . zh . ]
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( 1839 ) : note on the annelida . — in : murchison , r . i . [ ed . ] : the silurian system : 699–702 ; london .
( 1974 ) stratigraphic relationships at the western edge of the middle cambrian carbonate facies belt , field , british columbia . — geol . surv . pap . canad . ,
from the northumbrian plankton , with a note on a trematode parasite . — proc . zool . soc . lond . ,
( 1842 ) : über einige neue fossile schalenlose cephalopoden und eine neue gattung ringelwürmer ( anneliden ) . — beitr . petrefactenkunde ,
( 1839 ) : the silurian system . — 768 pp . ; london .
( 1972 ) : sediments of the middle cambrian burgess shale , canada . — lethaia ,
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( 1952 ) : classe des chétognathes . in : piveteau , j . [ ed . ] : traité de paléontologie , 2 : 161 ; paris .
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sagittiformis – from the latin sagitta , “arrow , ” and formis , “shape , ” in reference to the general outline of the animal .
lectotype – unsm 57644 in the national museum of natural history , smithsonian institution , washington , dc , usa .
other deposits : a . sinica ( chen et al . , 2002 ) from the lower cambrian chengjiang deposits , yunnan , china .
middle cambrian , bathyuriscus - elrathina zone ( approximately 505 million years ago ) .
a . saggitiformis is known from only a couple dozen specimens from the walcott quarry , comprising only 0 . 025 % of the specimens counted ( caron and jackson , 2008 ) .
butterfield , n . j . 1990 . organic preservation of non - mineralizing organisms and the taphonomy of the burgess shale . paleobiology , 16 ( 3 ) : 272 - 286 .
caron , j . - b . and d . a . jackson . 2008 . paleoecology of the greater phyllopod bed community , burgess shale . palaeogeography , palaeoclimatology , palaeoecology , 258 : 222 - 256 .
chen , j . - y . and d . - y . huang . 2002 . a possible lower cambrian chaetognath ( arrow worm ) . science , 298 ( 5591 ) : 187 .
chen , j . - y . , d . - y . huang and d . j . bottjer . 2005 . an early cambrian problematic fossil : vetustovermis and its possible affinities . proceedings of the royal society b : biological sciences , 272 ( 1576 ) : 2003 - 2007 .
chen , l . , h . luo , s . hu , j . yin , z . jiang , z . wu , f . li and a . chen . 2002 . early cambrian chengjiang fauna in eastern yunnan , china . yunnan science and technology press , kunming , china , 199 p .
walcott , c . d . 1911 . middle cambrian annelids . smithsonian miscellaneous collections , 57 ( 2 ) : 109 - 144 .
note that in the fossil preparation the head shows a highly reflective area ( cerebral ganglia ? ) . ( these are also sketched in above in the artist ' s rendering . ) the broad light area running along the trunk is the gut , while the narrow linear structures along the trunk may be traces of blood vessels and a nerve cord .
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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 .
random thoughts on paleontology , evolution , and philosophy from a life - long eccentric .
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“note that in the fossil preparation the head shows a highly reflective area ( cerebral ganglia ? [ aka : the bilobed ‘brain’ of many modern worms and arthropods ] ) … the broad light area running along the trunk is the gut , while the narrow linear structures along the trunk may be traces of blood vessels and a nerve cord . ”
that exhibit sounds awesome ! have you ever seen the chicago field museum’s burgess display ? if not , i think you’d greatly appreciate it : the wall contains a perpetually - moving animated diorama depicting dozens of the shale’s residents interacting and roaming about .
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i finally started reading ' crucible of creation ' . ( my local library had to borrow a copy from miami ) i really liked ' wonderful life ' when it cam out , and this book is sort of a rebuttal to that . you should write a review of both of them ! if you want to read an interesting debate between stephen jay gould and simon conway morris go to : urltoken
i started this blog when i was seven years old and now i ' m thirteen . i retired this blog a few years ago , but i still check on it and you can still contact me . i might even post again !
art started writing this blog at age seven and completed the majority of the work before he turned nine . he did his own research and for a long time dictated the blog entries to me , his mother . i typed exactly what he said and did my best to spell everything correctly .
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parent taxon : problematica according to r . k . bambach et al . 2007