normannites is a strongly ribbed evolute middle jurassic ammonite included in the stephanoceratoid family stephanoceratidae .

normanites was previously included in the otoitidae by arkell et al. 1957 .

more recent classifications show it to belong to the family stephanoceratidae , along with genera like stephanoceras and stemmatoceras .

normannites

photo information for stephanoceras ( normannites ) sp . ( munier - chalmas 1892 )
intraspecific variation of phragmocone chamber volumes throughout ontogeny in the modern nautilid nautilus and the jurassic ammonite normannites .
photo information for stephanoceras ( normannites ) sp . ( munier - chalmas 1892 ) - the fossil forum
3d reconstructions of the two specimens of normannites mitis , modern nautilus pompilius ( specimen 17 ) , and their phragmocones .
intraspecific variation of phragmocone chamber volumes throughout ontogeny in the modern nautilid nautilus and the jurassic ammonite normannites . - pubmed - ncbi
volumes and widths of chambers plotted against chamber numbers in normannites mitis . squares and diamonds represent volumes and widths , respectively .
volumes plotted against chamber numbers in normannites mitis . the volumes prior to chamber 25 ( nm . 1 ) and 27 ( nm . 2 ) have not been measured .
( 1a ) 3d model of normannites mitis ( nm . 1 ) ; ( 1b ) 3d model of normannites mitis ( nm . 2 ) ; ( 1c ) extracted phragmocone of nm . 1 ( 1d ) ; extracted phragmocone of nm . 2 ; ( 2a , b ) 3d models of nautilus pompilius ( specimen 17 ) ; ( 2c ) extracted phragmocone of nautilus pompilius ( specimen 17 ) ; ( 2d ) backface of 3d model of nautilus pompilius ( specimen 17 ) . scale bars are 1 cm .
specimen count 1 site number 27579 record last modified 5 jul 2018 geological age mesozoic - jurassic - middle - bajocian stratigraphy snowshoe fm nmnh - paleobiology dept . taxonomy animalia mollusca cephalopoda ammonoidea collector dr . ralph w . imlay see more items in paleogeneral types : mollusca paleozoic and mesozoic mollusca mesozoic cephalopoda type paleobiology place harney county , oregon , united states collection date 1959 type citation imlay . 1973 . u . s . geol . survey prof . paper . ( n . 756 ) : 82 , pl . 41 , f . 9 . usnm number pal168581 published name normannites ( normannites ) orbignyi buckman
this is one of the finest normannites we have had for sale on fossils direct . both lappets are complete and the ammonite ' s centre is pin point immaculate . this specimen was colleted in 1989 and is from an old collection andy cowap has since spent many hours in preparing this stunning specimen . to get both lappets preserved is exceptionally rare . ammonites of this superior quality only rarely come on to the open market for sale . serious collectors of rare ammonites will really appreciate this unique fossil approximately 170 million years old .
specimen count 1 site number 19773 record last modified 5 jul 2018 geological age mesozoic - jurassic - middle - bajocian stratigraphy kialagvik fm nmnh - paleobiology dept . taxonomy animalia mollusca cephalopoda collector l . b . kellum see more items in paleogeneral types : mollusca paleozoic and mesozoic mollusca mesozoic cephalopoda type paleobiology place alaska , united states type status holotype type citation imlay . 1964 . u . s . geol . survey prof . paper . ( n . 418 - b ) : b43 , pl . 13 , f . 6 , 7 , 8 , 10 . usnm number pal131401 published name normannites kialagvikensis imlay
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tajika a 1 , morimoto n 2 , wani r 3 , naglik c 1 , klug c 1 .
laboratory of physical anthropology , graduate school of science , kyoto university , kyoto , japan .
faculty of environment and information sciences , yokohama national university , yokohama , japan .
pmid : 26500816 pmcid : pmc4614987 doi : 10 . 7717 / peerj . 1306
( a ) scatter plot of chamber numbers and individual chamber volumes ; ( b ) scatter plot of chamber numbers and cumulative phragmocone volumes .
( a ) scatter plot of chamber numbers and individual chamber volumes ; ( b ) scatter plot of chamber numbers and phragmocone volumes .
comparison between males and females . chamber volumes plotted against chamber numbers in nautilus pompilius .
squares and diamonds represent the female and male , respectively . ( a ) scatter plot of chamber numbers and individual volumes ; ( b ) semilog scatter plot of chamber numbers and individual volumes ; ( c ) scatter plot of chamber numbers and cumulative phragmocone volumes .
squares , diamonds , and triangles represent the female , male , and indeterminable sex , respectively . ( a ) scatter plot of maximum conch diameters and chamber numbers of a specimen ; ( b ) scatter plot of maximum conch diameters and the phragmocone volume .
squares and diamonds represent volumes and widths , respectively . ( a ) specimen 8 ; ( b ) specimen 7 ; ( c ) specimen 53 . specimens with different growth trajectories were analysed .
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thesis ( phd - - geology ) - - university of auckland , 2004 .
bajocian–bathonian ( middle jurassic ) ammonites from the polish jura . part 2 : families stephanoceratidae , perisphinctidae , parkinsoniidae , morphoceratidae and tulitidae - palaeontographica abteilung a band 292 lieferung 4 - 6 — schweizerbart science publishers
bajocian–bathonian ( middle jurassic ) ammonites from the polish jura . part 2 : families stephanoceratidae , perisphinctidae , parkinsoniidae , morphoceratidae and tulitidae
palaeontographica abteilung a band 292 lieferung 4 - 6 ( 2010 ) , p . 115 - 213
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. colonization of novel environments can alter selective pressures and act as a catalyst for rapid evolution in nature . theory and empirical studies suggest that the ability of a population to exhibit an adaptive evolutionary response to novel selec . . .
. dna typing offers a unique opportunity to identify individuals for medical and forensic purposes . probabilistic inference regarding the chance occurrence of a match between the dna type of an evidentiary sample and that of an accused suspect , howe . . .
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. isolated left ventricular non - compaction is a rare genetic disorder manifesting mainly with heart failure , ventricular arrhythmias and systemic embolism . isolated ventricular tachycardia originating from the right ventricular outflow tract is an a . . .
. the course of total gorwth and of total natural removal ( mortality caused by natural agencies ) in naturally normal stands of pinus sil - vestris , picea abies , and betula alba on various sites in n . and s . finland were compared . growth continuously e . . .
stevens , b . m . ; folts , c . j . ; cui , w . ; bardin , a . l . ; walter , k . ; carson - walter , e . ; vescovi , a . ; noble , m .
. we discovered that glioblastoma ( gbm ) cells use cool - 1 / β - pix to inhibit normal activation of the c - cbl ubiquitin ligase via the redox / fyn / c - cbl pathway and that c - cbl inhibition is critical for gbm cell function . restoring normal c - cbl activity b . . .
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. . . outside europe , data concerning middle jurassic belemnites often lack precise taxonomic and / or stratigraphic information . exceptions are , for the northern hemisphere , the aalenian - bajocian belemnite faunas described by sachs & nalnjaeva ( 1970 , 1975 ) from north siberia , ne russia and the russian far east , which have been stratigraphically reviewed by meledina et al . ( 2005 ) . their taxonomic composition is characterized by the presence of hastitidae ( hastites , sachsibelus ) , pseudodicoelitidae ( lenobelus , pseudodicoelites ) and megateuthididae ( orthobelus [ = rarobelus ] , mesoteuthis [ = megateuthis ] , paramegateuthis ; see also dzyuba & weis 2015 ) . . . .
. . . aberhan ( 1998 ) arrived at similar plate tectonic reconstructions based on the pattern of pliensbachian ammonite and bivalve distributions . also , the comparative palaebiogeographic analysis of bivalves and ammonites ( benthic and nekto - benthic ) in the jurassic of siberian basins showed a good agreement ( meledina et al . , 2005 ) . however , liu et al . ( 1998 suggested that the boundaries of provinces based on ammonites and bivalves do not always coincide and explained this by the differing mode of life of the two groups . . . .
. . . in west - central nevada , the ammonite diversity is basically not affected by eustatic changes at all , probably due to special conditions of local ecology , environment or preservation . in general , broad patterns of the ammonite diversity show good correlation with eustatic fluctuations ( e . g . , o ' dogherty et al . , 2000 ; meledina et al . , 2005 ) . the changes in diversity and faunal turnover at small scales ( zone level ) are usually the result of multiple factors , including eustatic changes , and therefore need to be interpreted on a case by case basis . . . .
. . . since the upper toarcian , only the genus pseudolioceras existed in the arctic seas , that brings some uncertainties to the understanding of the stratigraphic extent of the upper toarcian and the possibility of its subdivision . in the middle jurassic , there was a significant change in the ammonite composition in the arctic seas caused by the isolation of the arctic from the west european seas ( meledina et al . , 2005 ) . european genera or species are almost absent . . . .
. . . the belemnite distribution over the section is also nonuniform . they occur massively near the lower - middle jurassic boundary , and they are also abundant in the upper jurassic and lower half of the lower cretaceous ( dzyuba , 2013 ; meledina et al . , 2005 ; saks and nal ' nyaeva , 1979 ) . in the bajocian - oxfordian and hauterivian , the number and taxonomic diversity of belemnites in siberian sections are low . . . .
. . . increasing seawater temperatures produced a massive bloom of the diverse marine species and the immigration of abundant west european faunas ( saks , 1976 ) . this time was also marked by colonization of the siberian seas by a belemnite fauna ( meledina et al . , 2005 ; saks anf nalnyaeva , 1975 ) . . . .
. . . faunal interchange increased and decreased alternately . sometimes some groups colonized the vast expanses of boreal seas for a long time and evolved there ; this resulted in the wide spread of the endemic groups of the boreal fauna ( meledina et al . , 2005 ; zakharov et al . , 2002 ) . in early studies ( saks et al . , 1980 ) , the jurassic zonation was based on data on ammonite occurrence in the numerous sections of central siberia and northeastern russia . . . .
. . . knyazev ' s ammonite zonal scale for the upper toarcian correlates with the bivalve , foraminiferal , ostracod , and microphytofossil zones distinguished simultaneously in the same sections , this scale is still incorporated into the regional scale for northern siberia and into the boreal jurassic zonal standard ( resolution , 2004 ; shurygin et al . , 2000 ; zhamoida and petrov , 2008 ) . in the middle jurassic , there was a considerable change in the taxonomic composition of ammonites in the arctic seas because of their isolation from the western european ones ( meledina et al . , 2005 ) . here , european genera and species were nearly absent . . . .
. . . comparative analysis of taxonomic diversity of jurassic benthic associations in different facies areas of the boreal basins allows the definition of development phases of macro - and microfaunal associations and certain marker levels with the sharpest structural and taxonomic changes as well as their relation to the main abiotic factors ( nikitenko & mickey 2004 ; meledina et al . 2005 ; nikitenko 2008 ) ( fig . 5 ) . in the early jurassic and aalenian there was a stable connection between the biota of the arctic and palaeopacific , while the connection with the palaeoatlantic was only periodically intensified during the major transgressions , which were accompanied by climate warming . . . .
stages in development of mollusks , paleobiogeography of boreal seas in the early - middle jurassic and . . .
we compared stages of the taxonomic restructurings that occurred in communities of ammonites , belemnites , and bivalves in siberian sea basins in the second half of the early jurassic and at the beginning of the middle jurassic . in general , the main stages are similar in different groups of mollusks , but the restructuring borders often do not coincide . degree of endemism and portion of . . . [ show full abstract ]
cylindroteuthid belemnite correlation of the jurassic / cretaceous boundary strata in northern siberia . . .
in the northern hemisphere there are some sections of the jurassic / cretaceous ( j / k ) boundary strata where boreal and tethyan molluscan fauna can be found together . they are all located on the pacific margins , in northeast asia ( far east of russia , northeast china ) and on the western coast of north america ( western british columbia in canada , northern california and southwest oregon in the . . . [ show full abstract ]
comprehensive zonal subdivisions of siberian jurassic and their significance for circum - arctic corre . . .
we show the present state of the set of parallel zonal scales for the siberian jurassic , based on various fossil groups , and the principles of their construction . we discuss the significance of siberian biostratigraphic scales for the boreal zonal standard of the jurassic units . the stratotype region for this standard must have a typical boreal ( arctic rather than mixed ) fauna . a possible . . . [ show full abstract ]
the jurassic / cretaceous boundary in northern siberia and boreal - tethyan correlation of the boundary . . .
there is no international consensus regarding the gssp for the berriasian , the basal stage of the cretaceous system . any of the events discussed by the international expert community can be regarded as a marker of the jurassic / cretaceous boundary : a phylogenetic change of taxa , paleomagnetic reversal , or isotopic excursion . however , the problem of identification of this level in boreal . . . [ show full abstract ]
ammonite assemblages and biostratigraphy at the lower to upper bajocian boundary in the digne area ( se france ) . implications for the definition of the late … | marta zunino - urltoken
ammonite assemblages and biostratigraphy at the lower to upper bajocian boundary in the digne area ( se france ) . implications for the definition of the late …
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citation : zell p , stinnesbeck w ( 2016 ) paleobiology of the latest tithonian ( late jurassic ) ammonite salinites grossicostatum inferred from internal and external shell parameters . plos one 11 ( 1 ) : e0145865 . urltoken
copyright : © 2016 zell , stinnesbeck . 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
funding : the research ( including all scientific analyses ) was funded by the deutsche forschungsgemeinschaft to ws ( dfg sti 128 - 17 and sti 128 - 31 ) . the dfg had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
ammonite morphology and internal structures yield important information on both the systematic position of species as well as their paleobiology ( e . g . dimorphism , habitat preferences , migration during sexual maturity , responses to seasonality ) , but only a small number of taxa has been analyzed in detail by using a combination of external and internal sets of data ( e . g . [ 1 ] ) . in consequence , the complete shell ontogeny of ammonites from embryonic to late post - embryonic stages is little known to date . for instance , an analysis of septal spacing in combination with shell diameter and ornamentation data revealed a gradual decrease in septum distances in adults reaching maturity ( e . g . [ 2 ] ) , while septal crowding was regarded as rare in premature individuals [ 2 ] . in other taxa septal crowding was attributed to adverse ecological circumstances , or injuries [ 2 – 6 ] .
here we provide a detailed analysis of the ontogenetic development of the late jurassic ammonite salinites grossicostatum ( imlay 1939 ) [ 7 ] , combining data on external morphology with internal characters such as septal spacing and angles , suture and siphuncular position and size . our data allow us to differentiate sexual dimorphism and four stages of growth . we discuss the implications of these data for the interpretation of life span , habitat preferences , seasonal environmental changes and migration of mature macroconchs .
map of mexico with inset of northeastern mexico and the jurassic - cretaceous transition at puerto piñones .
salinites grossicostatum specimens described here originate from a shaly limestone of the latest tithonian “ kossmatia - durangites - salinites beds” ( cf . [ 12 ] ) , respectively the uppermost crassicollaria zone [ 8 ] . locality map after [ 13 ] . puerto piñones : n25°02 . 719’ / w101°03 . 396’ .
our collection of salinites grossicostatum consists of 169 specimens ( fig 2 ) preserved three - dimensionally ( cpc - 1214–1242 ; sample box cpc - 1243 , cpc - 1405–1413 ) . they were collected by wolfgang stinnesbeck between 1986 and 1994 . no permits were required to access the sampling site at puerto piñones . logistic support and export licenses were provided by the facultad de ciencias de la tierra of the universidad autónoma de nuevo león .
specimens were collected from a single layer of the uppermost tithonian la caja formation at puerto piñones . d = diameter .
salinites grossicostatum is well preserved and suturelines are locally visible . the body chamber is completely preserved in most specimens and a long apophysis is present in eight individuals . descriptive terms are those of arkell et al . [ 14 ] , verma & westermann [ 15 ] and wright et al . [ 16 ] , and the ontogenetic description follows neige ( [ 17 ] , and references therein ) . abbreviations for ammonites : d , diameter ; wh , whorl height ; ww , whorl width ; u , umbilical diameter ; u / d , umbilical ratio ; ww / wh , whorl height to width ratio . specimens analyzed here are deposited in the colección de paleontología de coahuila ( cpc - 1214–1242 ; sample box cpc - 1243 , cpc - 1405–1413 ) of the museo del desierto , carlos abendrop dávila no . 3745 , parque las maravillas , saltillo , coahuila c . p . 25015 , mexico ( urltoken ) .
the latest tithonian [ 8 ] hildoglochiceras grossicostatum was first described by imlay [ 7 ] and was assigned to the genus salinites by cantú - chapa ( [ 18 ] , p . 19 ) .
representing an ontogenetic series from juvenile ( a ) to mature ( i ) .
( a ) cpc - 1405 ; ( b ) cpc - 1406 ; ( c ) cpc - 1407 ; ( d ) cpc - 1408 ; ( e ) cpc - 1409 ; ( f ) cpc - 1410 ; ( g ) cpc - 1411 ; ( h ) cpc - 1412 ; ( i ) cpc - 1413 ; scale bars = 10 mm .
an explicit sexual dimorphism is indicated by differential growth rates and internal growth features as detailed below , indicating that the assemblage consists of roughly equal numbers of micro - and macroconchs . the largest specimens ( d = 53–63 mm ) are here interpreted to represent macroconchs , while no individuals with shell diameters between 45 and 53 mm were found . this bimodal distribution pattern ( fig 2 ) may indicate changing habitat preferences of female adults during sexual maturity .
salinites grossicostatum is endemic to the ancient gulf of mexico , where it is restricted to shallow marine , outer shelf environments at the transition between the continental platform and slope [ 7 ] – [ 8 ] , [ 18 – 24 ] .
the embryonic ammonoid , termed ammonitella [ 25 ] , consists of the protoconch ( initial chamber ) and approximately one spiral whorl initiating at the caecum and terminating at the primary constriction [ 6 ] , [ 17 ] , [ 25 ] . in salinites grossicostatum , the center of the protoconch ( sensu [ 17 ] ) was used to determine the diameter of protoconch and ammonitella ( fig 4a 2 ) . the mean minimum diameter of the protoconch is 0 . 18 mm and its mean maximum diameter is 0 . 265 mm ( fig 4 ) . this considerable variation ( 68 % ) is due to the subellipsoidal form of the protoconch . the mean diameter of the ammonitella is 0 . 58 mm ( fig 4 ) . the ratio between the diameters of ammonitella and maximum protoconch values averages 2 . 15 .
( a ) cpc - 1217 , with sketch to the right ( a 2 ) to illustrate internal shell structures ; ( b ) cpc - 1214 ; measurements are provided in mm .
apertural heights and their corresponding diameters were measured in four specimens from the protoconch to the shell opening at the end of the last whorl . results are illustrated in fig 5 as a logarithmic ratio between the apertural height and shell diameter . our measurements reveal a small but significant break in growth between the embryonic and post - embryonic stages , coincident with the primary constriction . our data also indicate that allometric growth was higher in the post - embryonic development than in the initial ontogenetic stage considered as embryonic .
apertural height ( ah ) versus shell diameter ( d ) for two micro - and two macroconchs ( cpc - 1214–1217 ) .
note that apertural height is illustrated logarithmically . the regression lines indicate a change in growth at the primary constriction and that allometric growth is higher during post - embryonic stages than before .
the siphuncular diameter was measured in five specimens ( cpc - 1214–1217 and 1228 ; fig 6 ) . septa are mostly straight but are slightly incised concavely at the position of the septal neck . the latter is retrochoanitic during the earliest ontogenetic stages ( fig 4a 2 ) and becomes prochoanitic ( fig 6b ) at d≈1 . 8 mm , after approximately 18 septa and in the first half of the second whorl ( in cpc - 1214 ) . the connecting ring is thin and concave . the siphuncle is initially located in a subcentral position but gradually shifts to a ventral position shortly after the primary constriction ( fig 4a 2 ) . the siphuncle resumes its final ventral position at d≈0 . 77 mm . due to recrystallization and partial internal fragmentation , the siphuncle is only partially preserved in cpc - 1214–1217 and 1228 ( fig 6b ) . however , it is well - preserved in the two specimens cpc - 1214 and 1228 , with a minimum siphuncular diameter of 0 . 05 mm at d = 0 . 56 mm ( cpc - 1228 ) . a maximum diameter of 0 . 9 mm was reached at the last septum ( end of the phragmocone ) of macroconch cpc - 1214 at d = 40 . 9 mm ( postembryonic stage ) . the siphuncular diameter gradually increases in size and reveals a uniform first stage up to d≈28 mm ( ah≈10 mm ) . in macroconchs this stage is followed by an abrupt increase in siphuncular diameter . in general , however , the increase of the siphuncular diameter is slower than that of the increase in shell diameter and apertural height , thus indicating a negative allometry .
the upper plot illustrates the ratio between siphuncular diameter [ ds ] versus shell diameter [ d ] ; the lower plot figures the siphuncular diameter versus apertural height at the plane of coiling [ ah ] . to the right , high - resolution scans illustrate the siphuncle ; ( a ) longitudinal median section of cpc - 1228 ; ( b ) transversal section through the plane of coiling in cpc - 1214 . black arrows indicate position of the siphuncle ; scale bars = 2 . 5 mm .
the complete number of septa was analyzed in four adult specimens ( cpc - 1214–1217 ) and one juvenile ( cpc - 1231 ; fig 7 ) . the number of septa in the four adult individuals reaches a maximum of 97 and shows no major variation ( maximum between 86 and 97 ) , even though the two adult microconchs ( cpc - 1216 and 1217 ) are considerably smaller ( d = 11 mm , cpc - 1217 ; d = 18 . 7 mm , cpc - 1216 ) than the macroconchs ( d = 42 . 8 mm , cpc - 1214 ; d = 37 . 6 , cpc - 1215 ) . the number of septa thus appears to be genetically confined to a maximum of about 100 in both micro - and macroconchs . fig 7 illustrates the correlation between the number of septa and the diameter of s . grossicostatum . the general shape of these graphs is similar in the five individuals analyzed and reveals a uniform first stage of ontogenetic development , up to septum numbers of 25 to 30 . subsequently , however , the growth rates of micro - and macroconchs differ substantially ( fig 7 ) . the end of growth in specimen cpc - 1215 is marked by an inflection .
sexual dimorphism is indicated between septum numbers 25 and 30 by a considerably higher growth increase in macroconchs .
the interseptal angle , which corresponds to interseptal spacing , varies during ontogeny ( fig 8 ) . three general phases are indicated in mature specimens ( cpc - 1214–1217 ) . specific septal “events” are detected here and are indicated , in fig 8 , by dashed lines .
ratio between septal angle versus septal number for four mature specimens and one juvenile ( cpc - 1231 ) .
specific correlative septal “events” are marked by dashed lines . d = diameter at the last septum .
each trajectory initiates with a stable stage of strong septal angle variations , with widest angles reaching 39 . 4° . this stage ends after 20 to 23 septa and is followed by a stable stage in macroconchs ( cpc - 1214 and 1215 ) , and a relatively unstable stage in microconchs ( cpc - 1216 , 1217 , 1231 ) , with the majority of angles of about 16 . 6° but also including extremely low angles of 5° . this stage ends after approximately 63 septa . the final growth stage ( 64 septa and more ) is characterized by septal crowding with angles ranging between 6° ( cpc - 1215 ) and 18 . 3° ( cpc - 1214 ) , with an average of about 12 . 7° .
suture lines are only fragmentarily visible in three specimens ( fig 9 ) . in the early post - embryonic stage ( d = 8 mm ; fig 9c ) , the largest visible saddle is the third lateral , while the first lateral is smaller . the largest lobe is the second lateral . the ventral lobe is simple and small , higher than wide . the ventral saddle becomes larger and wider towards late ontogenetic stages . at d = 20 . 9 mm , the ventral saddle is subrectangular and furcated , while lateral lobes and saddles are stronger furcated .
( a ) cpc - 1218 ; ( b ) cpc - 1219 ; ( c ) cpc - 1220 .
the ornamentation shows five ontogenetic stages ( fig 3 , fig 10 ; see also [ 7 ] , p . 27 ) . the earliest ontogenetic stages are smooth with the exception of fine growth lines , which are , however , not visible on internal molds . ventrolateral ribs appear at a diameter of 6 . 4 mm ; they gradually increase in strength . a smooth keel is first seen at diameters of about 8 . 6 mm , which agrees with the description of imlay [ 7 ] who noted a keel at diameters of about 10 mm . a spiral groove , situated at about mid - flank , first appears on internal molds at diameters of about 9 . 2 mm in microconchs and at about 13 . 1 mm in macroconchs . the keel becomes serrated at diameters of about 11 . 2 mm in microconchs and at about 15 . 5 mm in macroconchs . the serration gradually increases in strength and finally reaches the ventrolateral plain . an apophysis is first present at diameters of about 19 . 5 mm in microconchs and of about 25 . 3 mm in macroconchs ; it is here interpreted to represent the adult stage , in accordance with previous interpretations of this character ( e . g . [ 26 ] ) .
in macroconchs , the spiral groove , the serration of the keel and the apophysis appear at larger diameters .
the adult stage starts by the first appearance of an apophysis at d≈19 . 5 mm in microconchs and at d≈25 . 3 mm in macroconchs . it is characterized further by a distinct reduction of space between the septa , marking the final stage of their secretion ( fig 8 ) . a similar pattern was previously documented in other ammonite taxa [ 2 ] , [ 17 ] , [ 27 – 29 ] , among other authors . in the most completely preserved macroconchs ( e . g . cpc - 1411–1413 ) a slight decrease in the whorl overlap ratio is present at d > 38 mm and the apophysis is consequently projecting ventrolaterally ( fig 3g , fig 3h , fig 3i ) , thus expressing a change in shell geometry . a crowding of growth lines , an additional characteristic of mature ammonites [ 29 ] , is only visible in macroconch cpc - 1413 ( fig 3i ) at a diameter of 63 mm .
our analysis of the ontogeny of salinites grossicostatum is based on 169 specimens collected from a single limestone layer at puerto piñones , coahuila and assigned to the latest tithonian “ durangites beds” . results are summarized in fig 11 .
sexual dimorphism and four major stages are recognized : embryonic , neanic , juvenile and mature . the diameter indicates the first occurrence of characteristic modifications .
septal spaces during earliest ontogenetic stages agree in all four specimens analyzed here ( micro - and macroconchs ) and are marked by strong variations in septal spacing . septal crowding during similarly early ontogenetic stages in nautiloids , bactritoids and ammonoids were interpreted to be related to hatching ( e . g . [ 6 ] , [ 30 – 33 ] ) .
numerous authors considered ammonite hatchlings to be planktonic ( e . g . [ 40 – 45 ] ) . in that case , the highly variable septal angles observed in early ontogenetic stages of s . grossicostatum could reflect a change of life habit to the juvenile stage ( cf . [ 6 ] ) . during the neanic development and up to about 30 septa , the ratio between the number of the septa and the diameter remains uniform for all specimens analyzed here .
the shell remains unornamented until diameters of about 6 . 4 mm , where ventrolateral ribs first appear . a smooth keel develops at diameters of about 8 . 6 mm . a spiral groove was identified in microconchs at about 9 . 2 mm diameter and in macroconchs at about 13 . 1 mm . serration of the keel is first present at diameters of about 11 . 2 mm in microconchs and at about 15 . 5 mm in macroconchs .
the abundance of juveniles ( d≈10 mm ) at puerto piñones , but also at sierra de parras and sierra de jimulco originally described by imlay [ 7 ] , may indicate high mortality rates during early ontogenetic stages before the expression of sexual dimorphism , suggesting that salinites grossicostatum reproduced early , quickly , and in large numbers , to ensure that at least some offspring of the population survive ( r - selection ) . ammonites associated with s . grossicostatum at puerto piñones are mostly represented by single adult individuals .
based on our analysis of septal spacing we propose that seasonal environmental changes are evidenced in salinites grossicostatum . ten low septal angle events with almost constant amplitudes of fluctuation were identified during the ontogenetic development of this ammonite , all separated by eight to nine wide and regularly spaced septa . in nautilus , these fluctuations in septal spacing are related to variations in the overall rate of growth and to the shape of shell and soft body ( [ 39 ] , and references therein ) , even though they are also seen in nautilus kept in captivity [ 64 ] .
six ornamentational stages are differentiated . during early ontogeny , s . grossicostatum is unornamented . this phase is followed by the subsequent appearance of ventrolateral ribs , a smooth keel , a spiral groove , and by the serration of this keel . maturity is reached at approximately 19 . 5 mm diameter in microconchs and 25 . 3 mm in macroconchs , and is marked by the appearance of an apophysis and a reduction of septal spacing . additional mature characteristics are expressed in macroconchs , including smoothening of surface ornamentation , increase in siphuncular diameter , reduced whorl overlap ratio , and the crowding of growth lines on the apophysis .
mortality rates were high during early ontogenetic stages before the expression of sexual dimorphism , suggesting that salinites grossicostatum was an r - selective species . size distribution and a sudden increase in siphuncular diameter in macroconchs may indicate changing habitat preferences of macroconchs during an interval after reaching sexual maturity and before their end of growth . macroconchs may have left the mating area at puerto piñones to reach shallow water areas for egg deposition .
the authors acknowledge seija beckmann ( institut für geowissenschaften , heidelberg ) for photography . we thank the referees and editor of plos one for their helpful comments that greatly improved the quality of this work .
conceived and designed the experiments : pz ws . performed the experiments : pz . analyzed the data : pz . contributed reagents / materials / analysis tools : ws . wrote the paper : pz ws . collected samples : ws . designed the figures : pz .
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the froghoppers , or the superfamily cercopoidea , are a group of hemipteran insects , in the suborder auchenorrhyncha .

adults are capable of jumping many times their height and length , giving the group their common name , but they are best known for their plant-sucking nymphs which encase themselves in froth in springtime .

froghopper

froghopper is miniature bug that can reach around 0 . 25 inches in length .
) . morphology and action of the hind leg joints controlling jumping in froghopper insects .
mouth apparatus of froghopper is designed for stabbing and extraction of the sap from the plant tissue .
froghopper is herbivore ( plant - eater ) . it eats sap of various species of plants .
watch the steady stream of honeydew droplets excreted by this beautiful froghopper as it sucks plant sap .
name
froghopper
refers to the frog - shaped head of this insect and its ability to jump .
driggers bf ; pepper bb , 1935 . the spittle insect or froghopper . new jersey agricultural experiment station bulletin 593 .
an adult froghopper ( aphrophora alni ) reaches a body length of under a half inch and can jump to heights of 28 inches .
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froghopper can jump 27 inches into the air . even though it is heavier than flea , it can jump higher thanks to strong , well - developed muscles of the hind legs .
second , does a froghopper have to anticipate the possible need to jump and thus hold its hind legs in readiness ? this would explain the cocked position adopted by the hind legs during walking .
froghopper is an insect that belongs to the group of true bugs . froghopper can be found around the world . it inhabits densely vegetated areas such as meadows , parks and gardens . froghoppers are numerous and widespread in the wild . several species of froghoppers are classified as agricultural pest due to ability to decrease the yield of commercially important species of plants ( such as sugar cane ) .
hind legs of froghopper generate g - force of 400 gravities , when it prepares to jump . this force is 80 times greater than g - force generated during the launching of the rockets into the space .
burrows , whose primary research interest is in how animals use the individual cells in their brains to generate movement , stumbled upon the froghopper ' s leaping agility while looking for an insect model to clear the next hurdle in his work .
philaenus spumarius , commonly known as a froghopper or spittle bug , is a mere 0 . 2 inches ( 6 millimeters ) long , but employs a novel catapult mechanism to launch itself upwards of 28 inches ( 70 centimeters ) into the air .
life cycle of froghopper consists of three developmental stages : egg , nymph and adult insect . nymph is often green colored and it looks like miniature , wingless version of adult . it molts several times until it reaches the size of an adult insect .
froghopper has two pairs of wings and three pairs of legs . first pair of wings covers the body like a tent . wings of some froghoppers form false head at the end of the body .
two headed - body
is designed to confuse the predators .
when disturbed , the adults can jump as high as 70 cm with enormous force using their powerful back legs . recent research has shown that within a millisecond they can accelerate to over 14 km / h ! very few potential predators could catch the common froghopper once it has jumped .
the adult common froghopper is not often seen by the casual observer . although it is 6 mm long , it can move so quickly when disturbed that it seems just to ‘disappear . ’ it is also not distinctively coloured : indeed , its pattern of coloration is very variable , often being various shades of mottled pale and dark brown , but also ranging from pure sandy brown to dull black . however , the nymph of the common froghopper is well - known for the distinctive white frothy ‘cuckoo - spit’ it produces – and hides within – on the stems of its food plants .
a sallow bush at the edge of the wood is home to a great many variegated willow froghoppers ( aphrophora pectoralis ) . like their more familiar smaller relative , the common froghopper ( philaenus spumariu s ) , their soft young live in “cuckoo - spit” , though these feed only on willows .
the common froghopper is found in a variety of habitats , but it is perhaps most abundant on waste ground and road - side verges where its weedy herbaceous food plants , such as thistles and mugwort , are often plentiful . within the ‘cuckoo - spit’ , the nymph of this species feeds by sucking sap from the food plant .
which of the five species of froghoppers examined is the best jumper ? the answer lies in which aspects of jumping performance are considered and how they are related to body mass and volume . the five species of froghopper analysed have a tenfold range of body masses ( 3 . 2 mg in neophilaenus to 32 . 9 mg in cercopis ) , and vary in length from 4 . 0 mm in neophilaenus to 9 . 8 mm in aphrophora .
a brief report on the kinematics of the jumping movements of a froghopper , philaenus spumarius ( burrows , 2003 ) , has demonstrated its jumping prowess , and a mechanism for jumping has been proposed for cercopis vulnerata ( gorb , 2004 ) . this paper analyses the detailed jumping performance of froghoppers and shows that in a jump they are airborne in less than 1 ms from the first propulsive movement of the hind legs . the enormous acceleration needed to achieve take - off velocities of over 4 . 7 m s - 1 in this short time is equivalent to 550 g .
spittlebugs are first noticed in late spring within white frothy masses of bubbles on grasses , herbaceous plants , shrubs and conifers . small nymphs of the spittlebug are mostly greenish with conspicuous red eyes ; different species may have different coloration . in most landscape gardens , there may be some wrinkling of the leaf or stem where the young spittlebugs are feeding . froghopper adults are found later in spring and summer . they look like leafhopper adults , but froghoppers are shorter and wider . one diagnostic feature is that the hind legs of froghoppers lack spines . damage from adults is slight .
spittlebugs are first noticed in late spring within white frothy masses of bubbles on grasses , herbaceous plants , shrubs and conifers . small nymphs of the spittlebug are mostly greenish with conspicuous red eyes ; different species may have different coloration . in most landscape gardens , there may be some wrinkling of the leaf or stem where the young spittlebugs are feeding , but generally their damage to plants is insignificant . froghopper adults are found later in spring and summer . they look similar to leafhopper adults , but froghoppers are shorter and wider . one diagnostic feature is that the hind legs of froghoppers lack spines . damage from adults is slight .
first , at what distance and with what sense does a froghopper detect an approaching predator ? a vibratory sense could give advanced warning of an approaching danger by detecting footfalls or movements of the plant on which it is feeding . this would allow the necessary time for developing the forces needed to jump ( burrows , 2007 ) . related families of auchenorrhyncha use this sense to communicate with each other on the same plant ( claridge , 1985 ; cocroft et al . , 2000 ; cokl and virant - doberlet , 2003 ) and cercopis appears to signal by vibrating its wings while remaining stationary on a plant ( kehlmaier , 2000 ) .
these are the bugs that in the summer you see as a nymph on bushes surrounded by what looks like spit ( hence the name ) . this is the adult , quite boring looking and uninspiring at first glance , but this teeny insect has one of the fastest movements in the terrestrial animal kingdom . considering its small size a froghopper can move at 4 metres per second , and can jump 70 times there own length ! i tried to film the jump on my sony avchd camcorder , but even after slowing the footage down to a fraction of its speed the jump is invisible . it looks like the insect just vanishes into thin air ! i would need a camera capable of shooting 1000 frames per second to show this behaviour in full slow mo .
from the start of the first visible movements of the hind legs to a froghopper becoming airborne takes no more than 0 . 875 ms in philaenus and a maximum of 1 . 5 ms in the heavier cercopis or aphrophora . in this short time , the body is accelerated to a take - off velocity of 4 . 7 m s - 1 in the best jumps by philaenus . in the best jumps by the different species , the applied acceleration ranged from 2267 - 5400 m s - 2 . philaenus experiences the equivalent of 550 g at take - off and the others from 231 - 428 g . the best jumps by philaenus require an energy output of 136 μj , a power output of 155 mw and exert a force of 66 mn . these forces and accelerations generated in jumping could not be produced by direct contractions of the muscles and indicate that muscular force must be generated in advance of the movement , energy stored and then released rapidly .
the head of froghoppers is flattened dorso - ventrally and has short antennae . its dorsal cuticle , and that of the prothorax , has many small indentations and its ventral cuticle is ribbed . the mouthparts point backwards and in aphrophora extend to the coxae of the hind legs . the folded fore wings cover the body , extend beyond the abdomen posteriorly and cover most of the hind legs when viewed from the side . the five species of froghopper analysed have a tenfold range of body mass , from 3 . 2±0 . 08 mg ( n = 7 ) in neophilaenus to 32 . 9±1 . 0 mg ( n = 16 ) in cercopis ( table 1 ) . their body lengths have a 2 . 5 fold range from 4 . 0±0 . 03 mm ( n = 7 ) in neophilaenus to 9 . 8±0 . 24 mm ( n = 23 ) in aphrophora . philaenus is toward the middle of this range with a body mass of 12 . 3±0 . 41 mg ( n = 34 ) and a body length of 6 . 1±0 . 08 mm .
graphs of leg and body movements during a jump by philaenus captured at 8000 s - 1 . ( a ) five points on the body ( see cartoon inset ) are plotted against time . take - off is indicated by the right arrow and vertical yellow bar . the first movement of a hind leg occurred 0 . 875 ms before take - off ( left arrow and yellow bar ) . the tarsi of the front and middle legs left the ground 0 . 5 ms before take - off ( middle arrow and yellow bar ) . velocity , measured as the movement of the centre of an ellipse representing the overall shape of the body , is plotted as a two - point average of successive frames ( blue line ) . ( b ) sequential movements of the five points on the body as the insect moved through the field of view of the stationary camera , superimposed on an image of the froghopper in its starting position . the black arrowheads and the linking black lines show the position of these five points at take - off . the corresponding positions of these points at different times during the jump can be read frame - by - frame from these positions at take - off .
have you noticed spit on your plants ? did you immediately think it was one of your kids or your spouse who did that ? before you go blaming your kids or partner , check your plants for spittlebugs first .
spittlebug drinks about 300 times its body weight in plant fluids in one hour ' s time . as a result of all the drinking , they produce a lot of waste , and that’s what causes all the spit on your plants .
because of all the spit on your plants , you probably want to get rid of these spittlebugs , but spit aside , spittlebugs can also harm your precious plants ! they not only will feed on your plants , but your plants may undergo stunted development and can also lose some of its vitality .
but don ' t panic just yet ! after some thorough research and several readings , we’ve chosen 7 efficient ways to help you get rid of these nasty bugs .
look around your garden and keep your eyes peeled for spittlebug patches . when you find them , use your hand to physically remove them off of your plant . if you don ' t want to touch them , you can knock them off by spraying water directly on the spittlebug patch . this is one of the easiest methods of how to get rid of spittlebugs ; however , i would only recommend doing this for light and easily accessible infestations only .
another option that is becoming popular nowadays is using a bug zapper . a bug zapper is an insect control system , also called an electrocutor trap device , which kills bugs , mosquitoes , and other flying insects through electrical discharge . although using a bug zapper offers an easy way of controlling pests in your garden , including spittlebugs , this method is only suitable to mild insect infestation . for severe infestations , you already need the help of insect - killer chemicals .
spittlebug eggs will form in and around garden debris and other similar areas . you need to clean up your garden now and then to get rid of old plant matter . cleaning up your garden is another method of how to get rid of spittlebugs .
with religious cleaning , you’ll get rid of the eggs and will , thereby limit , the number of spittlebugs that will hatch .
row covers are made of lightweight fabric or plastic and can be used to protect your crops from spittlebugs and other pests . they are an effective and cheap method to get rid of spittlebugs .
unlike plastic row covers , fabric row covers have tiny holes in them which allow the rain to come in and heat to go out . i would definitely recommend the fabric variety over the plastic ones .
although there are a lot of commercial pesticide sprays available on the market , these are very harmful to you and your plants .
pesticide , when ingested or inhaled , is toxic and very hazardous to your health . similarly , putting these chemicals on your plants would get rid of the pests but will change your plants and taint the flowers and fruits .
i would recommend making a homemade pesticide spray as the way to go on how to get rid of spittlebugs .
i would suggest using garlic or something spicy as your base for your organic homemade pesticide . you can even mix garlic and peppers with water to make it twice as effective .
the spittlebug is usually a bright yellow color and tends to blend in with your leaves . [ via : flickr . com ]
cover your nose and mouth with the face mask to protect your airways from the strong - smelling fumes of the pepper and garlic .
pour 1 cup of water in the blender along with the garlic cloves and the peppers . add the cayenne pepper and blend this mixture on high until the mixture is pureed .
using the cheesecloth or coffee filter , strain the mixture into the large jar .
add the dish detergent or the castile soap little by little . gradually stir it completely in using the wooden spoon .
place the lid on the jar , then let it sit in a cool , dark place for 24 to 48 hours .
first , you ' ll want to wipe all the spit off of your plants then spray away . be sure to test your pesticide on a small portion of your plant to make sure that you don ' t do any serious harm to it .
also , only do this on gloomy and rainy days because your spicy pesticide could burn your plant and kill it .
praying mantises tend to prey on most insects and pests . you could try ordering a few and releasing them onto your pest - laden plants and monitor the results after a week .
this method is a more drastic means regarding how to get rid of spittlebugs and only try this when the others have failed .
plant - based essential oils are a great way to deter spittlebug feeding patterns and can help disrupt their normal activities . i would recommend trying neem oil or any citrus - based oil such as lemon or orange oil to control and prevent spittlebug infestation .
there are a lot of organic insecticidal soaps available in the market . these insecticidal soaps help control and rid your plants of pests like spittlebugs .
they contain potassium salts of fatty acids which can break down the pests ' outer shells and cause them to become dehydrated and lose body fluids .
spittlebugs are really a nuisance in any garden . not only do they spit on your plants , they will harm your plants too . don’t let spittlebugs ruin your plants ! plan accordingly and follow the 7 tips and tricks above detailing how to get rid of spittlebugs .
please let me know if you have any other comments or suggestions or your own tips and tricks for getting rid of spittlebugs . i’d love to hear your thoughts !
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in the best jumps by philaenus , take - off occurs within 0 . 875 ms of the start of movements of the hind legs at a peak velocity of 4 . 7 m s - 1 and involves an acceleration of 5400 m s - 2 , equivalent to 550 times gravity . this jumping performance requires an energy output of 136 μj , a power output of 155 mw and exerts a force of 66 mn .
insects have evolved many different mechanisms for jumping so that they may increase the speed of their locomotion , launch themselves into flight , or escape rapidly from a potential predator . the repeatable nature of these movements has enabled detailed analyses of the underlying neuronal mechanisms ( burrows , 1996 ) and determination of the mechanical and muscular solutions to these extreme locomotory demands . click beetles ( elateridae ) jack - knife a joint in their thorax ( evans , 1972 ; evans , 1973 ) , bristletails ( archaeognatha ) ( evans , 1975 ) , springtails ( collembola ) ( brackenbury and hunt , 1993 ) and the larvae of some flies ( maitland , 1992 ) use movements of their abdomens . particular ants ( baroni et al . , 1994 ; tautz et al . , 1994 ) and the stick insect sipyloidea sp . ( burrows and morris , 2002 ) combine forward movements of their abdomens with movements of their legs .
five species of froghoppers were analysed : aphrophora alni ( fallén 1805 ) , cercopis vulnerata ( rossi 1807 ) , lepyronia coleoptrata ( linnaeus 1758 ) , philaenus spumarius ( linnaeus 1758 ) , and neophilaenus exclamationis ( thunberg 1784 ) . neophilaenus were collected near wells - next - the - sea in norfolk , uk and lepyronia from the nanus region of slovenia and near ljubljana . the other species were collected near wells - next - the - sea and around cambridge , uk . observations on live insects were made on the same day of collection , or after they had been in the laboratory for no more than a few days feeding on live chrysanthemum plants .
sequential images of jumps were captured at rates of 1000 or 2000 s - 1 with a high speed camera ( redlake imaging , san diego , ca , usa ) and associated computer , or at 4000 - 8000 s - 1 with a photron fastcam 512 or 1024pci camera [ photron ( europe ) ltd , marlow , bucks . , uk ] and with exposure times of 0 . 05 - 0 . 25 ms . spontaneous jumps , or jumps encouraged by delicate mechanical stimulation with a fine paintbrush or a 100 μm silver wire , were performed in a chamber of optical quality glass 80 mm wide , 80 mm tall and 25 mm deep with a floor of high density foam . selected image files were analysed with motionscope camera software ( redlake imaging ) or with canvas ( acd systems of america ) . jumps were aligned by designating the point of take - off as time t = 0 ms .
drawing of the anterior part of philaenus , viewed from the side , to show the orientation of the proximal parts of its three pairs of legs . each leg is shown in its most anterior position ( black ) and in its most posterior position ( grey ) . the pivots of the coxae with the thorax are indicated by black dots and vertical arrows . the plane of movement is not orthogonal to the plane of the drawing .
higher temporal resolution of the movements of a hind leg of a restrained aphrophora was obtained by gluing a 0 . 2 mm disc of reflective tape to a hind femur close to the femoro - tibial joint . a modified single lens reflex camera with a concentric light around the lens was focussed on the disc and the light it reflected was captured by a photocell in the film plane of the camera ( hedwig , 2000 ) . this method recorded the movement of the femur and was combined with sequential images of the hind legs captured by a high speed camera .
seventy nine jumps by 19 aphrophora , 92 jumps by 19 philaenus , 47 jumps by 13 cercopis , 8 jumps by 5 neophilaenus and 16 jumps by 4 lepyronia were captured and analysed . data are given as means ± standard error of the mean ( s . e . m . ) . temperatures ranged from 24 - 30°c .
sequential images of a jump by philaenus captured at 7500 s - 1 and with an exposure time for each of 0 . 05 ms . the images are arranged vertically in two columns and are timed from the image at take - off ( 0 ms ) . the first movements of the right hind leg ( white arrow ) occurred 1 . 04 ms before take - off .
the hind legs are only just over half the length of the body , ranging from 52 . 3±1 . 24 % ( n = 23 ) of body length in aphrophora to 66 % in philaenus and neophilaenus ( table 1 ) . in all species the front and middle legs are of similar length , but the hind legs are about one and half times longer so that the ratio of leg lengths ranges from 1 ( front ) : 1 ( middle ) : 1 . 4 - 1 . 6 ( hind legs ) in the different species . the increased length of a hind leg is due to its longer tibia . by contrast , the femur of a hind leg is the same length and shape as those of the other legs . the mass of the two hind legs of aphrophora , including the trochanter and all the more distal segments , represents only 2 . 0±0 . 11 % ( n = 7 ) of the total body mass .
the coxae of the three pairs of legs articulate with the thorax at different angles ( fig . 1 ) . in its most forward position the coxa of a hind leg subtends an angle of 155° to the longitudinal axis of the body and rotates , as determined by imposed movements , backwards and upwards in one plane about its paired pivots by only a further 20 - 25° . movements of segments distal to the coxa are in this plane . by contrast , in their most forward positions the coxae of the front and middle legs subtend angles of 80 - 90° and can rotate backwards through an angle of about 40° , or almost twice the range of a hind leg .
the same rapid movements of the hind legs propelled jumping by all species but the following analysis focuses on philaenus , with information from other species illustrating particular features .
in preparation for a jump from a horizontal surface , the front of the body was raised or lowered by movements of the front and middle legs to give a mean attitude of the body relative to the ground at take - off of 28±1 . 9° ( n = 20 ) . after adjustment of the body attitude was complete , the hind legs then remained still for 1 - 2 s with only the distal tips of their tarsi in contact with the ground . a rapid and simultaneous depression of both hind legs then powered an explosive take - off . no differences could be detected in the timing of the movements of the two hind legs and both left the ground at the same time .
the first movement of a hind leg in a jump was a downward and backward thrust of the trochanter and femur ( their individual movements cannot be distinguished in these images viewed from the side ) which , as transmitted through the tibia , forced the whole ventral surface of the tarsus to the ground ( fig . 2 , fig . 3a ) . images captured at 8000 s - 1 showed that this first movement of a hind leg occurred only 0 . 875 ms ( 7 frames ) before the insect became airborne . the force from the continuing backwards movement of the hind legs began to lift the body because their tarsi were now directly applied to the ground ( fig . 2 , fig . 3a , b ) . the body continued to be raised as the hind femora were further depressed and the hind tibiae were extended so that the tarsi of the front and middle legs were raised from the ground before take - off . the velocity of the insect followed these movements of the legs . the first surge in velocity corresponded to the initial movement of the femur ( fig . 3a ) and after a short pause of 0 . 25 ms was followed by a rapid acceleration to a peak velocity of 4 . 7 m s - 1 at take - off .
jump by aphrophora viewed ventrally and captured at 5000 s - 1 with an exposure time of 0 . 05 ms . ( a ) sequence of four images from the jump . ( b ) drawings to show the joint positions of the hind legs before ( - 0 . 4 ms , fully levated ) and after ( 0 ms , fully depressed ) their rapid movements .
the same sequence of movements of the joints of a hind leg were also seen in philaenus jumping from a horizontal position toward the camera and therefore moving out of its focal plane . ( fig . 5a - c ) . the first movement of a hind leg was a downward movement of the femur resulting from a depression of the trochanter about the coxa , accompanied by an extension of the tibia . with the tarsus pushed fully to the ground , further depression of the femur and extension of the tibia resulted in an upward movement of the body . at take - off the coxo - trochanteral joint had been depressed through its full range at angular velocities of 75 500 deg . s - 1 and the femoro - tibial joint extended at an angular velocity of 105 000 deg . s - 1 .
( a ) sequential images captured at 5000 s - 1 and with an exposure of 0 . 05 ms of philaenus jumping toward and to the right of the camera to show the leg movements . movements of the right hind femur are indicated by the arrows . ( b ) graphs of changes in the angle between the femur and the longitudinal axis of the body and of the femoro - tibial joint ( see inset drawings ) in a jump by another philaenus captured at 4000 s - 1 . ( c ) movements of four points on the body ( see cartoon ) and of the angular changes of the femur and tibia ( coloured lines ) . the changes in the femoro - tibial angle at the times indicated ( ms ) are shown in detail on the right .
hind leg movements of a restrained aphrophora recorded simultaneously by a camera capturing images at 1000 s - 1 and with an exposure of 0 . 25 ms and by a photoelectric device ( see materials and methods ) . ( a ) two sequential images , the first showing the hind legs fully levated before the attempted jump and the second after the rapid movement . the inset drawings show the position of the hind legs in these two frames . note the small piece of reflective tape on the hind femur . the outputs of the photoelectric device during six jumps were captured at a sampling rate of 45 khz and low - pass filtered at 2 khz . one trace in blue shows the unfiltered recording . the leg movements were complete in 0 . 3 ms . ( b ) seven attempted jumps by a second aphrophora show that the movement was again complete in 0 . 3 ms .
further detail of the joint movements was obtained by fixing aphrophora ventral surface uppermost in plasticene™ with the hind legs free to move ( fig . 6 ) . rapid and simultaneous movements of both hind legs occasionally occurred spontaneously or could be evoked by gently tickling hairs on the abdomen with a fine paintbrush . no differences in the form of these attempted jump movements could be discerned compared with those in free jumping . they were , however , much faster and were completed in 0 . 3 ms because they did not lift the mass of the body . the key movement was again a simultaneous depression of both trochantera about the coxae which occurred at 267 000 deg . s - 1 , almost three times faster than in a real jump . the speed of these movements was consistent in 6 attempted jumps by one aphrophora and in 7 by a second ( fig . 6a , b ) .
in the smallest of the froghoppers , neophilaenus , take - off was also achieved within 1 ms of the first movements of the hind legs ( fig . 7a , b ) . the first and key movements of the hind legs were again a rapid depression of the trochanter , with an accompanying extension of the tibia . before take - off in some jumps , the tarsi of the front and middle legs had already lost contact with the ground ( fig . 7b ) .
in the heaviest of the froghoppers , cercopis , the body was accelerated for a longer period to achieve take - off , with the movements of the hind legs beginning 1 . 5 ms before take - off ( fig . 8a , b ) . the whole jumping sequence began with the front and middle legs adjusting the attitude of the body , which , in this example was only 16° . both front and middle pairs of legs were again off the ground before take - off ( fig . 8b , c ) . movements of the hind legs led to the head being raised while the posterior of the body was lowered , giving a take - off angle of 45° despite the initial shallow body attitude .
in some jumps when lepyronia took off almost vertically the middle legs were already off the ground and the front legs were fully depressed and extended even before the first movements of the hind legs began ( fig . 9 ) . at 1 ms before take - off , the front and middle legs were clear of the ground but the take - off velocity of 4 . 0 m s - 1 was , nevertheless , as great as that achieved at take - off angles closer to the mean of 45° when the front and middle legs remained in contact with the ground for a longer period .
images from two jumps by neophilaenus captured at 2000 s - 1 with an exposure of 0 . 25 s . ( a ) a jump in which the take - off occurred within 1 ms of the first movements of the hind legs : body angle , 36° ; take - off angle , 55° ; take - off velocity , 4 . 2m s - 1 . ( b ) a jump toward and to the right of the camera in which the body was raised by the front and middle legs to assume a higher take - off angle : body angle , 50° ; take - off angle , 72° ; take - off velocity , 4 . 0 m s - 1 ; body mass , 3 . 2 mg .
philaenus had a mean take - off angle of 46 . 8±2 . 0° ( range 18° to 90° , n = 50 ) and a mode of 45° ( fig . 10a ) . in the first few milliseconds after take - off the insect typically maintained a stable orientation , and in many jumps this continued until a landing feet - first on a vertical or horizontal surface . in other jumps , however , the body rotated about its long or transverse axes and occasionally about both axes ( fig . 10b ) . in the example shown , philaenus spun through four complete cycles during the first 50 ms after take - off . in a second jump , the abdomen started to rotate forward about the transverse axis 10 ms after take - off so that it rather than the head pointed forwards . in a third jump , the body began to rotate about its long axis after 10 ms and after 19 ms had rotated by 180° so that the legs were pointed upwards . the rotation was completed 28 ms after take - off and then the next cycle of rotation began . in a fourth jump , the body first started to rotate about its long axis and then some 5 ms later also began to rotate about its transverse axis .
in all species the wings remained folded during take - off , so that the movement was a pure jump powered by the hind legs and not assisted by active wing movements . occasionally , however , the wings were opened after take - off , and flapping flight was assumed though this did not always lead to sustained flight or even to maintaining the height attained by the initial jump ( fig . 11 ) .
jump by cercopis . ( a ) sequential frames from the jump viewed from the side and captured at 2000 s - 1 with an exposure of 0 . 25 ms . ( b ) graph of the movements of six points on the body ( see cartoon inset ) during this jump . ( c ) sequential movements of the same six points superimposed on an image of cercopis in its starting position to show their vertical and horizontal displacement . the yellow arrows show the direction of the initial movements of the femoro - tibial joint . body angle at take - off , 16° ; take - off angle , 45° ; take - off velocity , 3 . 8m s - 1 ; body mass , 41 . 9 mg .
the energy required to achieve this performance depends on body mass . in the heavier species such as cercopis the best jumps required 238μ j but in the much lighter neophilaenus this fell to 28 μj ; philaenus required 136 μj . the power output in a jump depends on the time during which the energy is expended . in the 0 . 875 ms that philaenus took to accelerate its body the power output was thus 155 mw . the force exerted during the best jumps by philaenus was 66 mn . for the heavier cercopis the force was highest at 83 mn and for the lighter neophilaenus was lowest at 13 mn .
jump by lepyronia . ( a ) two columns of sequential images from a jump captured at 4000 s - 1 and with an exposure of 0 . 25 ms . at the start , the body was raised at an angle of 61° to the ground so that only the tips of the tarsi of the front and middle legs were in contact with the ground . ( b ) movements of five points on the body ( see image in c ) against time . ( c ) movements of the same body points to show their vertical and horizontal displacement . the black curved arrows show the direction of the initial movements of the femoro - tibial joint . body angle at take - off , 61° ; take - off angle , 90° ; take - off velocity , 4 . 0 m s - 1 ; body mass , 17 mg ; temperature , 36°c .
in a laboratory chamber at a temperature of 25°c and in still air , the average height jumped by philaenus was 428±26 mm ( n = 17 insects ) with the highest jumps reaching 700 mm , or 115 times its body length . none of the other species bettered these performances ; for example , in the same conditions aphrophora reached an average height of 263±20 mm ( n = 13 ) . in a particular individual , jumping performance declined with increasing attempts to encourage jumping with the consequence that averaged values are likely to have underestimated the true jumping performance of these insects . the best indication of ability came by taking the maximal performance of particular individuals , which under laboratory conditions and temperatures may still be an underestimate .
the orientation of the hind legs , and their key role in powering jumping , raised the question of whether this compromised their ability to contribute to walking . the striking feature of horizontal walking was that the hind legs did not show rhythmic movements in the walking pattern and were not sequentially placed on the ground and then lifted ( fig . 12a ) . instead they were held in the cocked position with the trochantera fully levated about the coxae so that the tarsi did not contact the ground . the hind legs were , however , used when climbing on a vertical surface on which there was limited traction ( fig . 12b ) . they moved rhythmically and were placed on the ground in time with the walking pattern so that they might therefore be expected to contribute thrust to the movement .
( a ) trajectories of five jumps by the same philaenus . take - off angles are the average over the first 10 ms after take - off . ( b ) rotation of the body during a single jump by philaenus . a fixed point on the head in the vertical or in the horizontal plane was plotted against time . the body spins about its longitudinal ( y axis ) and transverse ( x ) axis undergoing five cycles of rotation in the first 70 ms that it is airborne . images were captured at 1000 s - 1 ; body mass , 12 mg .
a jump in which aphrophora flapped its wings after take - off . the selected images captured at 1000 s - 1 and with an exposure of 0 . 25 ms are arranged in two columns . at take - off ( time 0 ms ) the wings were not opened . 5 ms later it started to lose height and 8 ms after take - off opened its hind wings and flapped them .
the main thrust for jumping is provided by the hind legs . any contribution from the front and middle legs is limited as they are often lifted from the ground before movements of the hind legs begin . their primary responsibilities are therefore to provide a stable platform and to set the angle of the body and the take - off trajectory , by raising or lowering the anterior part of the body . the critical movement of the hind legs in generating the thrust for a jump is the depression of the trochantera about the coxae . before take - off , the hind legs are levated forwards at the coxo - trochanteral joints so that that the femora are tucked between the thorax and the middle legs and are apposed to the lateral and ventral surfaces of the coxae . the tibiae are also flexed about the femora . by contrast , the front and middle tibiae are held in their most forward positions at angles of 80 - 90° to the longitudinal axis of the body . these critical actions of the hind legs in jumping are at the expense of their ability to contribute to the propulsion of the body in horizontal walking .
in terms of the height jumped then philaenus comes out on top . its average height jumped was 428 mm with the best jumps attaining heights of 700 mm , or 115 times its body length . by contrast , in the same conditions aphrophora reached an average height of only 263 mm , or 27 times its body length . distance and height achieved will be determined by take - off velocity , take - off angle and by the drag . all the froghoppers achieve high take - off velocities ranging from 3 . 4 to 4 . 7 m s - 1 and average take - off angles are close to 45° . drag will , however , be different because of the different body sizes and masses ( bennet - clark and alder , 1979 ) . the distance lost due to drag by philaenus is estimated to be about 25 % ( vogel , 2005 ) based on my data . the smaller neophilaenus would be expected to experience greater drag while the larger froghoppers should experience less .
in terms of velocity , acceleration and force relative to body mass generated at take - off , then philaenus again comes out on top . it accelerates its body in less than 1 ms to achieve an average velocity over the first 3 ms of the jump of 4 . 7 m s - 1 . both neophilaenus and lepyronia approach but never better these velocities at take - off , but the heavier aphrophora and cercopis both take longer ( 1 . 5 ms ) to accelerate their bodies and achieve lower velocities .
contribution of the hind legs of philaenus in walking . images were captured at 1000 s - 1 with an exposure of 0 . 5 ms . selected images , viewed ventrally , are shown at the times indicated . ( a ) horizontal walking . the middle and front legs were lifted in a sequence to perform a swing phase of similar duration ( thick black bars ) and contact the ground during a stance phase of variable duration ( thin blue bars ) . the hind legs did not move . ( b ) vertical walking in which the hind legs did contribute .
fleas have been considered the best jumpers amongst the insects , accelerating their body within 1 ms to a take - off velocity of 1 m s - 1 ( bennet - clark and lucey , 1967 ; rothschild et al . , 1975 ; rothschild et al . , 1972 ) . froghoppers produce a substantially better jumping performance . philaenus accelerates its body to a take - off velocity that is more than 4 . 7 times faster than a flea despite having a body mass that is 27 times greater and a body that is four times longer . once airborne , however , the flea is likely to have its jumping distance reduced by 80 % due to drag compared to the 25 % reduction experienced by philaenus ( vogel , 2005 ) . heavier orthopteran insects such as locusts ( schistocerca gregaria ) with a mass of 1 - 2 g take 20 - 30 ms to extend their hind legs and accelerate their body ( brown , 1967 ) to a take - off velocity of 3 m s - 1 ( bennet - clark , 1975 ) while prosarthria teretrirostris with a mass of 280 mg takes 30 ms of acceleration to achieve a take - off velocity of 2 . 5 m s - 1 ( burrows and wolf , 2002 ) . the jumping distance of these larger insects is likely to be curtailed by only some 6 % due to drag ( vogel , 2005 ) . if jumping performance is expressed as the force exerted relative to body mass , then froghoppers again outperform other insects and other jumping animals . the force that froghoppers exert at take - off is more than 400 times their body weight and is , therefore , much higher than in other jumpers such as the flea ( ∼135 times ) ( bennet - clark and lucey , 1967 ) , locust ( ∼8 ) ( bennet - clark , 1975 ) and humans ( ∼2 - 3 ) ( dowling and vamos , 1993 ) .
potential predators of froghoppers are many and include birds , solitary wasps that provision their nests with froghoppers , and predatory social wasps or flies . parasitoids such as pipunculidae attack the pre - adult stages which are unable to jump . a further major danger may be unwitting predation by grazing mammals . they share this danger with all the other insects that live or feed on vegetation , but a rapid and long jump out of harm ' s way becomes advantageous . it may be that froghoppers are vulnerable while airborne and thus need to minimise exposure time in the air by jumping rapidly . this assessment of the value of jumping , however , poses further questions .
the structural specialisations of the joints and the sequence of muscle actions that enable this remarkable jumping performance is analysed in two subsequent papers ( burrows , 2006 ; burrows , 2007 ) and further papers will explore the evolution of the particular jumping mechanisms in other families of these plant sucking bugs .
i thank my cambridge colleagues for their help in collecting these bugs , for their constructive suggestions during the course of this work , and for their comments on the manuscript . this work was initiated at the wells field study centre , wells - next - the - sea , norfolk , england during an undergraduate field course , and i am most grateful to the warden christine marshall for the use of these facilities . some experiments were also carried out at the department of entomology , national institute of biology , ljubljana , slovenia and i am most grateful to dr meta virant and her colleagues for their support .


chloealtis aspasma is a species of grasshopper in the family acrididae .

it is native to northern california and southern oregon in the united states .

it is known by the common names siskiyou short-horned grasshopper and siskiyou chloealtis grasshopper .

chloealtis aspasma

rehn , j . a . g . & hebard . 1919 . trans . amer . entomol . soc . 45 : 82 > > chloealtis aspasma urn : lsid : orthoptera . speciesfile . org : taxonname : 61230
rehn , j . a . g . & hebard . 1919 . a new species of grasshopper of the genus chloealtis ( acrididae ) from the pacific coast . transactions of the american entomological society ( trans . amer . entomol . soc . ) 45 : 82
rehn , j . a . g . & hebard . 1919 . a new species of grasshopper of the genus chloealtis ( acrididae ) from the pacific coast . transactions of the american entomological society ( trans . amer . entomol . soc . ) 45 : 81 - 87
rehn , j . a . g . & hebard ( 1919 ) a new species of grasshopper of the genus chloealtis ( acrididae ) from the pacific coast : transactions of the american entomological society ( trans . amer . entomol . soc . ) 45 : 81 - 87
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gurney , strohecker & helfer . 1964 [ 1963 ] . trans . amer . entomol . soc . 89 : 130
type locality : northern america , northwestern u . s . a . , oregon , jackson county , siskiyou mountains
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bey - bienko . 1932 . orthoptera palaearctica critica xi . the group chrysochraontes ( acrid . ) . eos , revista española de entomología ( eos ) 8 : 88
gurney , strohecker & helfer . 1964 [ 1963 ] . a synopsis of north american acridine grasshoppers of the genus group chrysochraontes ( orthoptera : acrididae ) . transactions of the american entomological society ( trans . amer . entomol . soc . ) 89 : 130
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jago . 1971 . a review of the gomphocerinae of the world with a key to the genera ( orthoptera : acrididae ) . proceedings of the academy of natural sciences of philadelphia ( proc . acad . nat . sci . philad . ) 123 : 293
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