chrysotoxum cautum is a species of hoverfly .

it is found in southern britain and europe east into the palearctic but is normally encountered in small numbers .

the larvae are thought to feed on root aphids .

adults are usually found on the edges of woodland or scrub or along hedgerows where they visit a wide range of flowers .

chrysotoxum cautum

kari pihlaviita added the finnish common name
etelänsarvikirvari
to
chrysotoxum cautum ( harris , 1776 )
.
chrysotoxum cautum - large species , female with long dense hairs on thoracic dorsum . antennal segment 1 shorter than segment 2
tergites of investigated species , lateral view . a , chrysotoxum festivum a ; b , chrysotoxum festivum b ; c , chrysotoxum elegans ; d , chrysotoxum elegans –intermediate form ; e , chrysotoxum octomaculatum .
investigated localities . a , chrysotoxum festivum a ; b , chrysotoxum festivum b .
thin - plate - spline deformation grids shape differences . a , males of chrysotoxum festivum a and chrysotoxum festivum b ; b , females of chrysotoxum festivum a and chrysotoxum festivum b ; c , males of chrysotoxum festivum a and chrysotoxum elegans ; d , females of chrysotoxum festivum a and chrysotoxum elegans ; e , males of chrysotoxum festivum b and chrysotoxum elegans ; f , females of chrysotoxum festivum b and chrysotoxum elegans . differences between the species were exaggerated five - fold to make them more visible .
chrysotoxum tomentosum giglio - tos , 1890 : 159 stat . nov . corresponds to chrysotoxum festivum b
fig . 5 . superimposed outline drawings showing wing shape differences between analysed species . a , chrysotoxum vernale and chrysotoxum montanum sp . nov . ; b , chrysotoxum orthostylum sp . nov . and chrysotoxum montanum sp . nov . ; c , chrysotoxum orthostylum sp . nov . and chrysotoxum vernale .
pilosity on mesoscutum , lateral view . a , chrysotoxum festivum a ; b , chrysotoxum festivum b .
it just doesn ' t quite seem to have markings which match cautum or verralli when i check ' google ' .
fig . 10 . male genitalia . a , chrysotoxum orthostylum sp . nov . , lateral view ; b , chrysotoxum orthostylum sp . nov . , dorsal view ; c , chrysotoxum festivum , lateral view ; d , chrysotoxum festivum , dorsal view ; e , chrysotoxum vernale , lateral view ; f , chrysotoxum vernale , dorsal view .
fig . 11 . abdomen , dorsal view . a , chrysotoxum montanum sp . nov . ; b , chrysotoxum orthostylum sp . nov . ; c , chrysotoxum vernale .
figure 1 . adult female hover fly , chrysotoxum cautum ( harris ) , showing resemblence of some hover fly species to yellowjackets . photograph by david a . iliff , cheltenham , england .
fig . 6 . superimposed outline drawings showing surstylus shape differences between chrysotoxum montanum sp . nov . and chrysotoxum vernale .
fig . 14 . distribution maps of the material examined . a , chrysotoxum montanum sp . nov . ; b , chrysotoxum orthostylum sp . nov . ; c , chrysotoxum vernale .
fig . 13 . right surstylus of the male genitalia , lateral view . a , chrysotoxum montanum sp . nov . ; b , chrysotoxum orthostylum sp . nov . ; c , chrysotoxum vernale .
fig . 8 . thoracic scutum of males , lateral view . a , chrysotoxum vernale ; b , chrysotoxum montanum sp . nov .
head of male , anterior view . a , chrysotoxum festivum a ; b , chrysotoxum festivum b ; a , width of black vitta .
fig . 9 . thoracic pleura , lateral view . a , chrysotoxum orthostylum sp . nov . ; b , chrysotoxum vernale ; c , chrysotoxum montanum sp . nov . dotted areas indicate yellow colour ( kepst , katepisternum ) .
a small dumpy chrysotoxum looking like a diminutive c . cautum or chubby c . verralli . the first two antennal segments are very short ( like cautum ) but the third segment is very long . the male has dense , dark eye hairs . this is the only chrysotoxum with a northern - biased distribution and is frequent in sheltered , scrubby parts of heathland , moorland and open structured woodland , particularly in upland districts of britain . the larvae feed on ant - associated aphids . nbn map : urltoken bnsys0000006893
chrysotoxum festivum ( linnaeus , 1758 : 593 ) corresponds to c . festivum a
fig . 7 . scatter plot of factor loadings of two pc scores showing the positions of chrysotoxum montanum sp . nov . and chrysotoxum vernale in the environmental space .
right surstylus of male genitalia , lateral view . a , chrysotoxum festivum a ; b , chrysotoxum festivum b ; a , length of surstylus ; b , width of surstylus .
a review of the palaearctic species of the genus chrysotoxum mg . ( diptera , syrphidae )
ch . cautum should be correct . however , a good picture of the antennae is needed to compare the length of the segments ( this is the first question the key http : / / web . archiv . . . x _ key . html asks ) . in your fly the third segment seems to equal the first and second together , which is fine for ch . cautum . the colouring fits well , too . regards , sundew
the european chrysotoxum species can be grouped according to the proportional length of the basoflagellomere : species with basoflagellomere shorter than scape and pedicel together , chrysotoxum bicinctum ( linnaeus , 1758 ) , chrysotoxum elegans loew , 1841 , chrysotoxum festivum ( linnaeus , 1758 ) , chrysotoxum gracile becker , 1921 , chrysotoxum lineare ( zetterstedt , 1819 ) , c . montanum sp . nov . , chrysotoxum octomaculatum curtis , 1837 , c . orthostylum sp . nov . , chrysotoxum parmense rondani , 1845 , chrysotoxum tomentosum giglio - tos , 1890 , c . vernale and chrysotoxum verralli collin , 1940 ; species with basoflagellomere longer than scape and pedicel together , chrysotoxum arcuatum ( linnaeus , 1758 ) , chrysotoxum cisalpinum rondani , 1845 , chrysotoxum cautum ( harris , 1776 ) , chrysotoxum fasciolatum ( de geer , 1776 ) and chrysotoxum intermedium meigen , 1822 . within the group with a relatively short basoflagellomere , masseti et al . ( 2006 ) defined the morphological festivum group , c . bicinctum , c . festivum , c . elegans , c . octomaculatum , c . parmense and c . vernale , excluding characters such as the colour of femur and lateral margin of terga 3 and 4 . based on morphology , c . tomentosum , c . montanum sp . nov . , and c . orthostylum sp . nov . are likely to belong to the festivum group ( sensu masseti et al . , 2006 ) . these three species have the yellow fasciae on terga 3 and 4 not reaching the lateral margin of the tergum , but the festivum group ( sensu masseti et al . , 2006 ) includes some species with yellow fasciae reaching the lateral margin . further morphological and molecular analyses are required to access the concept of a festivum group .
fig . 1 . location of the 16 analysed landmarks on the right wing in the chrysotoxum vernale .
chrysotoxum tomentosum , male genitalia . a , epandrium , dorsal view ; b , hypandrium , lateral view .
chrysotoxum festivum , c . orthostylum sp . nov . ( femora wholly yellow ) , and c . tomentosum
sang - wook s , ho - yeon h . 2013 . clarification of previously confused chrysotoxum sapporense and chrysotoxum graciosum ( insecta : diptera : syrphidae ) in east asia based on morphological and molecular data . animal cells and systems 17 : 277 - 289 .
chrysotoxum montanum sp . nov . , c . orthostylum sp . nov . ( femora black basally ) , and c . vernale
boxplot of mean centroid size showing differences in wing size between chrysotoxum festivum a , c . festivum b , and c . elegans .
fig . 2 . location of the 30 analysed semi - landmarks on the right surstylus of the male genitalia in the chrysotoxum vernale .
the social wasp - mimicking hoverfly chrysotoxum cautum must be a strong contender for the most handsomely marked british fly species . the subterfuge ( batesian mimicry ) is enhanced by the adoption of a flight pattern and buzz very similar to that of the social wasps that are being mimicked and it is very likely that any predators already wise to the hazards of being stung will avoid the perceived risk and leave this fly well alone . there are several other members of the chrysotoxum genus with very similar markings so close examination of the relative lengths of the antennal segments will be required in order to unequivocally separate the species .
habitat and flowers visited . chrysotoxum orthostylum sp . nov . has been found flying with other chrysotoxum species in the grassy edges of conifer forests at high altitudes in serbia and montenegro . in fyr macedonia , c . orthostylum sp . nov . has been found resting on lapsana and chrysanthemum leaves in wet meadows .
giglio - tos e . 1890 . le specie europee del genere chrysotoxum meig . atti della r . accademia delle scienze di torino 26 : 134 - 165 .
refinements to couplets in the key to the palaearctic chrysotoxum in violovitsh ( 1974 ) are provided below , based on the results presented here , as well as those in nedeljković et al . ( 2013 ) . chrysotoxum montanum sp . nov . , c . vernale and specimens of c . orthostylum sp . nov . with femora black basally would key out to c . vernale ( sensu lato ) in violovitsh ( 1974 ) ; specimens of c . orthostylum sp . nov . with femora wholly yellow would key out to chrysotoxum festivum ( linnaeus , 1758 ) ( s . l . ) . chrysotoxum festivum and chrysotoxum tomentosum giglio - tos , 1890 , which also have yellow fasciae on terga 2 - 4 not reaching the lateral margins of tergum , would key out to c . festivum ( s . l . ) .
habitat and flowers visited . this species has been found with other chrysotoxum species in the grassy edges of conifer forests at high altitudes . flowers visited : no data .
additional material . holotype of chrysotoxum collinum : m , museo “la specola” coll . rondani holotypus ( red label ) / 357 / chrysotoxum vernale ( = collinum rnd . ) / rev . ( overwritten by hand on ‘det . ’ ) m . daccordi 1985 [ lsf ] . for further non - type additional material see table s1 .
two or three legs of each specimen were used for total genomic dna extraction from 25 chrysotoxum dry specimens ( table s2 ) , following chen et al . ( 2010 ) . material analysed included 22 specimens of the c . vernale a , two specimens of the c . vernale b and one of third chrysotoxum species . dna voucher specimens are deposited in fsuns .
fig . 12 . terga 3 - 5 , dorsal view , chrysotoxum orthostylum sp . nov . a , male ; b , female ( t , thorn - like process ) .
violovitsh na . 1974 . a review of the palaearctic species of the genus chrysotoxum mg . ( diptera , syrphidae ) . энтомологическое обозрение ( entomological review ) 53 : 196 - 217 .
chrysotoxum verralli - restricted to south - east england and the midlands . very similar to elegans but smaller and with black at front of tergite 2 mostly straight , not curved as in elegans .
sommaggio d . 2001 . the species of the genus chrysotoxum meigen , 1822 ( diptera , syrphidae ) described by giglio tos . bollettino del museo regionale di scienze naturali , torino 1 : 115 - 126 .
in the world approximately 150 chrysotoxum species has been recorded so far ( evenhuis et al . , 2008 ) . peck ( 1988 ) listed 69 species of this genus occurring in the palaearctic of which 23 occur in europe .
masetti a , luchetti a , sommaggio d , burgio g , mantovani b . 2006 . phylogeny of chrysotoxum species ( diptera : syrphidae ) inferred from morphological and molecular characters . european journal of entomology 103 : 459 - 467 .
amongst the most striking of all british hoverflies , chrysotoxum species are boldly marked with yellow and black and have distinctly elongated antennae . although the genus is distinctive , identification to species level can be tricky and may require close examination of the abdominal markings in combination with other characters .
nedeljković z , ačanski j , vujić a , obreht d , đan m , ståhls g , radenković s . 2013 . taxonomy of chrysotoxum festivum linnaeus , 1758 ( diptera : syrphidae ) - an integrative approach . zoological journal of the linnean society 169 : 84 - 102 .
a preliminary examination of named specimens of chrysotoxum vernale loew , 1841 resulted in the identification of two different morphotypes according to the colour of long pile on thoracic scutum of males : yellow ( morphotype a ) and black ( morphotype b ) ( nedeljković , 2011 ) . in addition , a new chrysotoxum taxon sharing with morphotypes a and b an antenna with basoflagellomere shorter than scape and pedicel together and abdominal terga with yellow fasciae not reaching the lateral margins was recognized . the main objective of the present study is to test using an integrated approach , whether these morphotypes correspond to different species .
musca arcuata and m . festiva linnaeus , 1758 ( currently chrysotoxum arcuatum and c . festivum ) and m . citrofasciata de geer , 1776 ( currently xanthogramma citrofasciatum ) ( insecta , diptera ) : proposed conservation of usage of the specific names by the designation of neotypes for m . arcuata and m . festiva
opinion 1982 ( case 3090 ) . musca arcuata linnaeus , 1758 and m . festiva linnaeus , 1758 ( currently chrysotoxum arcuatum and c . festivum ) and m . citrofasciata de geer , 1776 ( currently xanthogramma citrofasciatum ) ( insecta : diptera ) : specific names conserved by the designation of neotypes for m . arcuata and m . festiva
veličković n , obreht d , đan m , kočiš tubić n , vujić a . 2012 . cytochrome oxidase i gene variation in genus chrysotoxum ( diptera , syrphidae ) . pp . 63 - 68 in : sabin i , ed . , proceedings of the xivth international symposium ‘young people and multidisciplinary research’ , timis¸oara . timisoara , romania : editura politechnica .
chrysotoxum species are present in all bio - geographic regions except for australasia . in the palaearctic region 71 species are recorded , 24 of which are present in europe ( vockeroth , 1969 ; violovitsh , 1974 ; peck , 1988 ; nedeljković et al . , 2013 ) . these wasp - mimics can be separated from other syrphid genera by the yellow and black body colouration ( yellow maculae in thoracic pleura and yellow fasciae in abdominal terga ) , the very long antenna and the convex , margined abdomen ( thompson and rotheray , 1998 ) . chrysotoxum adults are found in many habitat types ( speight , 2013 ) and their poorly known larvae are associated with ants and / or root aphids ( rotheray , 1994 ) .
worldwide , syrphids ( diptera : syrphidae ) play important ecological roles in nature . adults are usually conspicuous flies that feed on pollen and nectar from a wide range of plants in numerous habitats ( rotheray and gilbert , 2011 ) . in europe , syrphids are one of the better studied diptera families . however , the taxonomy of certain groups is still provisional and uncertain , such as the genus chrysotoxum meigen , 1803 ( speight , 2013 ) .
diagnosis . medium to large species ( 15 - 17 mm , n = 10 ) ; r = 1 : 0 . 8 : 1 . 8 ; vertical and frontal triangles with black pile ; scutum with black pile ( fig . 8b ) ; katepisternum completely black ( fig . 9c ) or with a very small yellow macula ; scutellum with long black pile ; male genitalia very similar to that in c . vernale ; surstylus 1 . 3 times longer than hypandrium ( excluding superior lobes and aedeagus ) ; epandrium plus surstylus 1 . 2 - 1 . 3 times longer than hypandrium ( excluding superior lobes and aedeagus ) ; in female frons black , with two triangular pollinose maculae . chrysotoxum montanum has longer wings than c . vernale . chrysotoxum montanum occurs only at high altitude . ml tree based on mtdna coi gene revealed a clear separation of c . montanum from c . vernale .
three morphologically related taxa , c . elegans loew , 1841 , c . festivum , and c . gracile becker 1921 share the following morphological characters : first flagellomere short ( shorter than scape and pedicel together ) and yellow legs . chrysotoxum gracile , clearly separable by the longer scape than pedicel from c . elegans and c . festivum , is not included in the present study . based on mtdna and internal transcribed spacer 2 ( its2 ) , c . festivum and c . elegans appeared in the same clade ( masetti et al . , 2006 ) .
taxonomic notes . in this new species , the katepisternum colour ranges from completely black ( fig . 9c ) to black with a small yellow macula anteriorly . chrysotoxum montanum sp . nov . can be separated from c . orthostylum sp . nov . and c . vernale by the colour of the long pile on the scutum , which are black in c . montanum sp . nov . ( fig . 8b ) and yellow in the other species . for further morphological differences between c . montanum sp . nov . and c . vernale see table 2 .
paralectotypes ( designated here ) . 1f labelled as 16 / 547 / vernale f loew alte sammlung / austria alte sammlung / chrysotoxum vernale f loew * ( nhmw ) . 20 specimens labelled as ‘coll h . loew’ and seven specimens not labelled but supposed to be part of loew’s collection and have been collected in poznań ( j . ziegler , in lit . ) ; 10 of the 20 ‘coll h . loew’ specimens are labelled as ‘wien schiner’ ( 4 specimens ) , ‘florenz mann’ ( 3 specimens ) , ‘syrakus zeller’ ( 1 specimen ) , ‘spanien’ ( 1 specimen ) , ‘paris’ ( 1 specimen ) ( zhmb ) .
an integrative taxonomic approach revealed two taxa within chrysotoxum festivum ( linnaeus , 1758 ) ( diptera , syrphidae ) , c . festivum a and c . festivum b . in addition to morphological differences , results also showed significant distinction in geometric morphometrics of wings and surstyli , and in dna sequence data ( nuclear its2 sequences ) between c . festivum a , c . festivum b , and the closely related species c . elegans loew , 1841 . from examination of type material , the name c . tomentosum giglio - tos , 1890 is proposed for c . festivum b , and the taxon is redefined . © 2013 the linnean society of london
the taxon tomentosa was originally described as a variety of chrysotoxum festivum based on material from the alpine region of piemonte and surrounding localities alagna and valsesia , valdieri , val dell ' orco , santa redegunda in stiria ( giglio - tos , 1890 ) . z . nedeljković and a . vujić examined the lectotype designated by sommaggio ( 2001 ) labelled as follows : 20–68 / ch . festivum linn . var . tomentosa giglio - tos ( handwritten ) / ch . festivum var . tomentosa ( handwritten ) lectotype / mzut - mrsn torino , coll . bellardi box 12 . the redescription of c . tomentosum is based on the examination of lectotype and additional studied material .
chrysotoxum festivum linnaeus , 1758 . described based on a male specimen bm 1937 - 539 as musca festiva . type locality : ‘europa’ . the lectotype of musca festiva was designated by thompson et al . ( 1982 : 155 ) . neotype : male . type locality : germany , schneverdingen , lüneberg heath , n . germany , leg . t . h . rowsell and b . j . clifton in 1937 was designated by p . j . chandler ( iliff & chandler , 2000 ) and is deposited in bmnh . it was examined by n . wyatt in october 2012 , a . ricarte in january 2013 , and also via photos by z . nedeljković in october 2012 .
the pca reduces the bioclim variables ( table 1 ) to three pcs with eigenvalue ≥ 1 . 88 % of the variation . separation among species were statistically significant along pc1 and pc2 ( anova , pc1 : f 1 , 302 = 141 . 26 ; p < 0 . 00000 , fisher lsd post hoc test p < 0 . 00000 ; pc2 : f 1 , 302 = 155 . 27 ; p < 0 . 00000 , fisher lsd post hoc test p < 0 . 00000 ) , but not along pc3 ( f 1 , 302 = 1 . 90 ; p < 0 . 168397 ) . because of this , pc3 was excluded in the interpretation of results . the precipitation - related pc1 explained 61 % of the total variation , while the temperature - and altitude - related pc2 explained 17 % of the total variation ( table 1 ) . the scatter plot of pc1 against pc2 illustrated a clear niche separation between c . vernale and c . montanum sp . nov . chrysotoxum vernale occurs across a wide temperature range , precipitation and altitude ( fig . 7 ) , while c . montanum sp . nov . occurs in a much narrower range of temperatures . chrysotoxum montanum sp . nov . occurs only at high altitudes with high precipitation , and its distribution is strongly influenced by the precipitation in the coldest and wettest quarters of the year ( bio19 , bio12 , bio16 and bio13 ) .
chrysotoxum vernale was described from an unspecified but large ( “unter vielen exemplaren . . . ” ) number of specimens , both males and females . some males studied by hermann loew were collected in the place where he was working at the time of the description ( “ . . . hier gefangen . . . ” ) : poznań ( “posen” ) , poland . apart from this , no other data about the type locality of c . vernale are given in the original description . in the zmhb there are 27 specimens of c . vernale belonging to loew’s collection ( j . ziegler , in lit . ) ; of these specimens only 20 are labelled as ‘coll . h . loew’ and further information is not provided in the available data bases at the zmhb ( sven marotzke , in lit . ) . these specimens were examined here to confirm the colour of scutum pile , which was yellow . two named specimens ( male and female ) of c . vernale from the ‘old collection’ ( “alte sammlung” ) at the nhmw were also examined ; the female is labelled as ‘ chrysotoxum vernale f loew * ’ ( handwritten ) . no information about the locality where the nhmw specimens were collected is available , but loew’s type material is usually marked with an asterisk ( * ) ( a . vujić , pers . obs . ) . we consider that the specimens examined both in zmhb and nhmw belong to the type series of c . vernale and the nhmw male is designated as lectotype in order to stabilize this species concept .
for surstyli , outline shape was studied . surstyli of 51 chrysotoxum males were analyzed : 21 of morphotype a and 30 of morphotype b . the right surstylus was taken off using pins . surstyli were mounted in hoyer’s medium on a microscopic slide and immobilized with a cover slip . in the absence of clearly - identifiable , homologous , anatomical loci , 30 semi - landmarks were generated ( fig . 2 ) . for each specimen semi - landmarks were drawn twice to reduce statistical error . to superimpose semi - landmarks , coordgen7 . 14 and an integrated semiland module were used following a distance - minimising protocol to minimize the shape differences due to the arbitrary nature of semi - landmark positions along the curve ( bookstein , 1997 ; zelditch et al . , 2004 ) .
diagnosis . large to medium size species ( 11 - 15 mm , n = 10 ) ; r = 1 : 0 . 6 : 1 . 5 ; vertical and frontal triangle with yellow pile ; scutum with long yellow pile ( male ) ( fig . 8a ) ; katepisternum with a yellow macula ( fig . 9b ) ; scutellum mainly with long yellow pile ; basal third of pro - and mesofemora black ; surstylus 1 . 2 - 1 . 3 times longer than hypandrium ( excluding superior lobes and aedeagus ) ( fig . 13c ) ; epandrium plus surstylus 1 . 1 - 1 . 2 times longer than hypandrium ( excluding superior lobes and aedeagus ) ( fig . 10e ) ; in female , frons black , with two rectangular pollinose maculae . chrysotoxum vernale has larger wings than c . montanum . additionally , it has wide temperature , precipitation and altitude range .
significant differences in wing size and shape between c . montanum sp . nov . , c . vernale and c . orthostylum were detected ( fig . 5a ) . inconspicuous but statistically significant differences in surstylar shape between c . montanum sp . nov . and c . vernale were also revealed with a geometric morphometric analysis ; this is the second example after that of nedeljković et al . ( 2013 ) in which geometric morphometry of genitalia has successfully contributed to the separation of cryptic syrphid species . in addition , niche divergence analysis reveals that c . montanum sp . nov . is more specialised than c . vernale . chrysotoxum montanum sp . nov . occurs only at high altitudes with high precipitation , while c . vernale is able to thrive in a wider range of environmental conditions of temperature , precipitation and altitude . the sympatric ( e . g . in fyr macedonia , golema poljana ) and synchronic ( early may to mid - august ) distributions of c . montanum sp . nov . and c . vernale also support their taxonomic separation .
the anova of wing centroid size showed significant differences between c . vernale and c . montanum sp . nov . ( males : f 1 , 117 = 8 . 99 ; p < 0 . 00331 , tukey’s post hoc test p < 0 . 01227 ; females : f 1 , 117 = 81 . 12 ; p < 0 . 00000 , tukey’s post hoc test p < 0 . 00010 ) . anova test and tukey’s post hoc test showed significant differences in wing size between c . orthostylum sp . nov . and c . vernale ( f 1 , 182 = 6 . 69 ; p < 0 . 01045 , tukey’s post hoc test p < 0 . 00969 ) and c . orthostylum sp . nov . and c . montanum sp . nov . ( f 1 , 166 = 30 . 12 ; p < 0 . 00000 , tukey’s post hoc test p < 0 . 00000 ) . chrysotoxum vernale has larger wings than c . montanum sp . nov . and c . orthostylum sp . nov . ( fig . 4b ) .
thin - plate spline deformation grid allows recognition of wing regions that are contributing to the discrimination . the main differences in wing shape between c . festivum a and c . festivum b are in the basal and apical parts of the wing and in wing length . males of c . festivum a have a wider basal part ( landmarks 1 and 16 ) and narrow medial ( landmarks 7 and 8 ) and apical ( landmarks 3 and 6 ) parts of the wing ( fig . 10a , b ) . the major wing deformations affecting c . elegans and c . festivum a occur in the medial part of the wing and are associated with landmarks 7–13 , and in the apical part of wing associated with landmarks 3–5 ( fig . 10c , d ) . the deformation grid indicates that the shape difference between c . elegans and c . festivum b is due to wing width and associated with displacement of all landmarks . chrysotoxum elegans is characterized in having a generally narrower wing than c . festivum b , especially in the male ( fig . 10e , f ) .
taxonomic notes . chrysotoxum orthostylum sp . nov . can be distinguished from c . montanum sp . nov . and c . vernale by the abdomen shape , which is slender and nearly parallel - sided in c . orthostylum sp . nov . ( fig . 11b ) but more oval in other species ( fig . 11a , c ) . in addition , the posterior corners of tergites 3 and 4 in c . orthostylum sp . nov . are developed into thorn - like processes ( fig . 12a , b ) , while in similar species these corners are blunter . the katepisternum of c . orthostylum sp . nov . has a yellow macula ( fig . 9a ) , which is more reduced or even absent in c . vernale and c . montanum sp . nov . ( fig . 9b , c ) . males of c . orthostylum sp . nov . can be separated from other species by the long surstylus , which is 3 . 3 - 3 . 7× ( n = 5 ) longer than wide in c . orthostylum sp . nov . ( fig . 13b ) but 2 . 8× or less in the other species ( fig . 13a , c ) . in addition , the shape of the hypandrium apex is rectangular in c . orthostylum sp . nov . ( fig . 10a ) and more triangular in c . festivum ( fig . 10c ) and c . vernale ( fig . 10e ) .
da correctly classified the material studied in accordance with the three defined species . the overall classification success of the da was 92 . 95 % which indicates wing shape is a reliable predictor of interspecific discrimination . of the 249 digitalisations , only 21 were misclassified . canonical variate analysis conducted on shape variables gave two highly significant axes . the first canonical axis ( cv1 ) , with 65 . 4 % of total variation , separated c . vernale from c . orthostylum sp . nov . ( wilks’ lambda = 0 . 09515 ; χ 2 = 662 . 18 ; p < 0 . 00000 ) . the second canonical axis ( cv2 ) , with 34 . 6 % of total variation , separated c . vernale and c . orthostylum sp . nov . from c . montanum sp . nov . ( wilks’ lambda = 0 . 38388 ; χ 2 = 269 . 52 ; p < 0 . 00000 ) ( fig . 4a ) . the superimposed outline drawings showing major wing deformations between c . vernale and c . montanum sp . nov . occur in the central and distal part of wing and are associated with landmarks 2 , 4 , 5 and 9 - 11 ( fig . 5a ) . major differences in wing shape were found in the central part of the wing ( landmarks 2 , 7 - 11 ) of c . orthostylum sp . nov . and c . vernale ( fig . 5c ) , and in the central ( landmarks 7 , 8 , 12 and 13 ) and apical parts in c . orthostylum sp . nov . and c . montanum sp . nov . ( landmarks 4 and 5 ) ( fig . 5b ) . chrysotoxum vernale has wider wings , while c . orthostylum sp . nov . longer wings .
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a relatively large robust boldly marked grassland hoverfly species , widespread in southern britain and europe , usually seen flying near hedgerows , woodland edges and scrubby areas in small numbers where they visit a wide range of flowers . they are most often seen from june to october with a peak in mid july to early august . wing length 10 - 13mm ( 0 . 4 - 0 . 5in ) .
is similar to several hoverfly species and care is needed for identification . the abdominal patterning has two straight black bars thinly joined in the centre per segment , with the pattern towards the rear falling short of the segment edge and gradually becoming more like the shape of a whales fluke .
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hello , can this hoverfly be identified by foto ? i caught it on may 10th , 2011 , on the light . location : germany , rhineland - palatinum , 53572 unkel , river rhine valley , dry and warm habitat . i already posted this fly 1 1 / 2 years ago , with another photo , but never got an answer . perhaps this photo is better for an identification . thanks in advance , michael
why didn´t the upload work ? there were nop spaces , dots etc . in the name . one more try : michael stemmer attached the following image : [ 71 . 88kb ]
thank you , sundew ! this helps me a lot . greetings from the river rhine valley , michael
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the family syrphidae , also known as hover flies , flower flies , and syrphid flies , is one of the largest families of diptera with over 5 , 000 described species ( capinera 2004 ) . they are called hover flies because they can hover while in mid - flight , lingering in one place much like a hummingbird . while many diptera are detrimental to humans , most syrphid flies are beneficial . the larvae of some species ( for example , allograpta obliqua ) are natural predators of aphids , and many of the adults are important crop pollinators ( weems 1951 ) .
hover fly adults are often brightly colored and are commonly mistaken for the bees and wasps that they resemble ( mimic ) in appearance and behavior ( catts and mullen 2002 ) . many are black with either white , yellow , or orange bands across the abdomen and similar in shape to bees and wasps . they also have similar flight behavior and feed on pollen and nectar ( brower and brower 1965 , heal 1979 ) . this type of mimicry is called batesian mimicry because the mimic , although it is not dangerous to predators , benefits because the model is dangerous to predators ( bates 1961 ) . in other words , predators may avoid bees , wasps and yellowjackets because they can inject toxins by stinging , so predators also avoid the flower flies that are mistaken for bees and wasps ( gilbert 1986 ) .
figure 2 . adult female eastern yellowjacket , vespula maculifrons ( buysson ) . photograph by bruce marlin , http : / / cirrusimage . com .
the species eristalis tenax ( linnaeus ) , commonly known as the drone fly ( adult ) or rat - tailed maggot ( immature ) , is a mimic of the european honey bee apis mellifera linnaeus ( golding et al . 2001 ) , and was introduced from europe around 1875 ( gilbert 1986 ) . the rat - tailed maggot is probably the source of biblical writings that depict honey bees spontaneously developing from dead animals . this is because female drone flies can lay their eggs in carcasses ( osten - sacken 1893 ) .
figure 3 . adult female drone fly , eristalis tenax ( linnaeus ) . photograph by lloyd spitalnik , copyright 2006 . www . lloydspitalnikphotos . com , used with permission .
figure 4 . adult honey bee , apis mellifera linnaeus . photograph by zachary huang , urltoken .
adult e . tenax are important pollinators of many crops and wildflowers , while their larvae occasionally are pests around livestock ( day 2008 ) and cause accidental myiasis in humans ( catts and mullen 2002 ) . myiasis occurs when fly larvae infest humans and other vertebrate animals and feed on the host ' s living tissue ( lakshminarayana et al . 1975 ) , and is a common occurrence in certain other dipteran species , such as bot flies ( human bot fly , horse bot fly ) and screwworms .
the rat - tailed maggot is cosmopolitan , occurring on every continent except antarctica and ranges to the highest latitudes in the north ( metcalf 1913 ) . it is absent in the extreme southern latitudes and in arid areas of europe , asia , and africa ( thompson 1999 ) . in the united states , it is found as far north as alaska and south through california and florida ( milne and milne 1980 ) .
egg : the egg is white in color , has an elongate shape , and is covered in a sticky substance ( milne and milne 1980 ) .
larva : the following information is from metcalf ( 1913 ) . the aquatic larva has a cylindrical shape with patches of horizontal folds dividing the body into segments , between which the cuticle is smooth . at the division of each body segment , two rows of flexible hairs are visible . the larva has a highly specialized organ on the posterior end ( siphon ) that acts as a respiratory appendage and also looks like a tail , thus giving them their nickname
rat - tailed maggot .
the siphon can be several times the length of the body .
figure 5 . larva of the rat - tailed maggot , eristalis tenax ( linnaeus ) , about two and a half inches in length . photograph by walter reeves , the georgia gardener .
figure 6 . larvae of the rat - tailed maggot , eristalis tenax ( linnaeus ) , in dirty , polluted water within a tire . photograph by tami ansley , atlanta , ga .
figure 7 . larvae of the rat - tailed maggot , eristalis tenax ( linnaeus ) , in manure . photograph by j . keith waldron , cornell university .
pupa : the pupa looks very similar to the larva but is shorter and thicker ( gilbert 1986 ) . however , unlike the larva the pupa has two pairs of cornua , or horn - like bumps , located on the thorax ( metcalf 1913 ) . the siphon remains present in the pupa but generally locks in a curved position over the back ( metcalf 1913 ) .
figure 8 . rat - tailed maggot pupa , eristalis tenax ( linnaeus ) . photograph by unknown .
adult : the following information is from milne and milne ( 1980 ) . the adult drone fly can be over half an inch in length . they can be easily differentiated from honey bees because they lack a constricted waist between the thorax and the abdomen , and they only have two wings , while honey bees have four . short , brownish - yellow hairs are located on the thorax and the first segment of the abdomen . the body is dark brown to black in color , with yellow - orange marks on the side of the second abdominal segment while a narrow yellow - orange band crosses the third abdominal segment .
like many other fly species , males can easily be distinguished from females because they have larger eyes that almost touch , while females have smaller eyes that are spaced further apart . flies from the family syrphidae can be distinguished from all other fly species by the identification of a spurious vein , or
false vein .
this vein does not terminate at the end of the wing or at another vein but has a free end , and is not as sclerotized as the other wing veins ( metcalf 1913 )
figure 9 . adult male drone fly , eristalis tenax ( linnaeus ) . photograph by lloyd spitalnik , copyright 2006 . used with permission .
figure 10 . adult female drone fly , eristalis tenax ( linnaeus ) . photograph by lloyd spitalnik , copyright 2006 . www . lloydspitalnikphotos . com , used with permission .
figure 11 . spurious vein that is indicative of all syrphidae flies . illustration from maxwell lefroy , manual of entomology , 1923 .
the drone fly undergoes complete metamorphosis with three larval instars . usually two to three generations are produced each year ( gilbert 1986 ) . however , there are many gaps in our understanding of the drone fly life cycle and more research is needed to provide detailed information on the life cycle .
eggs : eggs are deposited near the surface of foul water or decaying organic material , and are laid in masses with the eggs side by side , perpendicular to the ground ( metcalf 1913 ) . it is not known how long it takes for eggs to hatch .
figure 12 . larvae of the drone fly , eristalis tenax ( linnaeus ) , sharing their habitat with mosquito larvae . the drone fly larvae are represented by the dimples where their siphons are attached to the surface of the water . photograph by phillip e . kaufman , university of florida .
pupae : pupation occurs in a drier environment than where the larvae develop . this is usually just below the soil surface , where they remain for eight to 10 days ( gilbert 1986 , milne and milne 1980 ) . cornua that appear on the pupa are thought to aid in respiration during the pupation period as the trachea within the siphon becomes unusable ( metcalf 1913 ) .
adults : females will feed on pollen once they emerge from the pupa in order to obtain the necessary nutrients to complete reproduction ( gilbert 1986 ) . subsequent meals will consist mainly of nectar to provide the energy necessary for activity ( gilbert 1986 ) . adult drone flies often feed on nectar from daisies , chrysanthemums , and asters ( gilbert 1986 ) . the adults prefer yellow flowers , leading to their importance in the pollination of yellow - flowered crops ( ilse 1949 ) .
male e . tenax tend to be territorial . observations suggest that males may live in the same territory their entire lives where they mate , feed , and groom , defending this area against other insects ( wellington and fitzpatrick 1981 ) . mating can occur while the pair is flying , with the male uppermost , or terrestrially while resting on foliage ( rogers and walker 1916 ) . after mating , adult rat - tailed maggot females lay clusters of about 20 eggs near dirty , contaminated water , sewage , or decomposing organic substances ( milne and milne 1980 ) .
adults can be found from late march to early december and most often in september and october ( gilbert 1986 ) . in the late autumn months , females from the latest generation will mate and find a secure place to overwinter . the sperm remain alive , nourished by fat reserves from the female , while her eggs remain undeveloped until the spring ( kendall and stradling 1972 ) . after overwintering , the female emerges and lays from 80 to 200 eggs , and the cycle begins again ( kendall and stradling 1972 ) .
eristalis tenax is usually not a serious pest , but occasionally the larvae can become a nuisance in livestock areas , where they are often abundant in manure lagoons and holding pits ( kaufman et al . 2000 ) . during the summer , larvae can migrate from these sites in massive numbers as they seek dry pupation sites ( day 2008 ) . the migrations can cause many problems , such as contamination of livestock feed , short circuits from accumulations in electrical boxes , and congregations in barn stalls , egg cartons , and other unwanted places ( kaufman et al . 2000 , day 2008 ) .
figure 13 . a dairy farm lagoon in upstate new york . photograph by j . keith waldron , cornell university .
figure 14 . rat - tailed maggots of the drone fly , eristalis tenax ( linnaeus ) , infesting a dairy farm lagoon in upstate new york . photograph by j . keith waldron , cornell university .
the drone fly is reported to have caused accidental myiasis , which occurs when fly larvae inhabit a living host by accident , usually because of the ingestion of contaminated food in humans ( lakshminarayana et al . 1975 ) . this can transpire in four ways : intestinal or gastric , nasal , auricular , or anal , with intestinal or gastric being the most common ( rogers and walker 1916 , herms 1969 ) . the larvae are able to survive gastric fluids , possibly due to the fact that they are adapted to living in polluted habitats ( aguilera et al . 1999 ) . myiasis from e . tenax becomes apparent when the host notices the larvae in their bowel movements ( lakshminarayana et al . 1975 , aguilera et al . 1999 ) . symptoms of this condition include diffuse abdominal pain and diarrhea , and can easily be treated with medication which expels the larvae from the body ( lakshminarayana et al . 1975 , aguilera et al . 1999 ) .
because e . tenax larvae live in highly polluted water , lagoons and manure pits need to be kept in the best condition possible . not allowing the manure to extend through the surface of the water can help prevent fly development ( lyon 1995 , day 2008 ) . physical blockades between lagoons and barns / coops can prevent larvae from migrating to the barns / coops when searching for pupation sites , and will keep the pupating larvae in a non - essential area ( lyon 1995 , day 2008 ) . agitating the lagoons frequently by pumping , especially during the warm summer months can disrupt larval development .
aguilera a , cid a , regueiro bj , prieto jm , noya m . 1999 . intestinal myiasis caused by eristalis tenax . journal of clinical microbiology 37 : 3082 .
bates hw . 1862 . contributions to an insect fauna of the amazon valley . lepidoptera : heliconidae . transactions of the linnean society 23 : 495 - 566 .
brower jz , brower lp . 1965 . experimental studies of mimicry . 8 . further investigations of honeybees ( apis mellifera ) and their dronefly mimics ( eristalis spp . ) . the american naturalist 99 : 173 - 187 .
capinera jl . 2004 . flies . pp . 875 - 883 . in encyclopedia of entomology , vol . 2 . capinera jl [ ed . ] . kluwer academic publishers , dordrecht , the netherlands .
catts ep , mullen gr . 2002 . myiasis ( muscoidea , oestroidea ) . pp . 319 - 348 . in mullen g , durden l [ eds . ] , medical and veterinary entomology . academic press an imprint of elsevier , san diego , ca .
day er . ( 2008 ) . livestock area fly control . virginia tech extension . urltoken ( 5 february 2009 ) .
gilbert fs . 1986 . hoverflies . cambridge university press , cambridge , england .
golding yc , ennos ar , edmunds m . 2001 . similarity in flight behaviour between the honeybee apis mellifera ( hymenoptera : apidae ) and its presumed mimic , the dronefly eristalis tenax ( diptera : syrphidae ) . the journal of experimental biology 204 : 139 - 145 .
heal j . 1979 . colour patterns of syrphidae ii . eristalis intricarius . heredity 43 : 229 - 238 .
herms wm . 1969 . herms ' s medical entomology 6th ed . james mt , harwood rf [ eds . ] , the macmillan company , london , england .
ibrahim ia , gad am . 1978 . the occurrence of paedogenesis in eristalis larvae ( diptera , syrphidae ) . journal of medical entomology 12 : 268 .
ilse d . 1949 . colour discrimination in the drone fly eristalis tenax . nature 163 : 255 - 256 .


brasilennea arethusae is a fossil species of air-breathing land snail , a terrestrial pulmonate gastropod mollusk in the family cerionidae , from the paleocene itaboraí basin , brazil .

brasilennea arethusae is the largest species in the genus brasilennea .

it was one of the first fossils found in itaboraí basin and its name makes reference to the fact that it is a terrestrial species : the name is in honor of arethusa , a sylvan nymph and one of the hesperides from greek mythology .

brasilennea arethusae

how can i put and write and define brasilennea arethusae in a sentence and how is the word brasilennea arethusae used in a sentence and examples ? 用brasilennea arethusae造句 , 用brasilennea arethusae造句 , 用brasilennea arethusae造句 , brasilennea arethusae meaning , definition , pronunciation , synonyms and example sentences are provided by ichacha . net .
clade panpulmonata clade eupulmonata clade stylommatophora informal group sigmurethra | superfamilia = urocoptoidea | familia = cerionidae | genus = { extinct } brasilennea | species = { extinct } b . arethusae | binomial = brasilennea arethusae | binomial _ authority = maury , 1935 | synonyms = } } brasilennea arethusae is a fossil species of air - breathing . . .
brasilennea arethusae var . minor trindade , 1956 : 18 ( pl . 3 , figs . 1e , 2e ) .
stylommatophora family cerionidae genus brasilennea maury , 1935 ( figs . 3 - 26 )

' brasilennea arethusae
' is a fossil species of air - breathing land snail , a arethusa , a sylvan nymph and one of the hesperides from greek mythology .
brasilennea arethusae maury , 1935 : 4 - 6 , figs . 1 - 5 ; salvador et al . 2011 : 445 - 447 , fig . 1 , a - g , l .
brasilennea guttula salvador & simone , 2012 : 2 ( figs . 1 - 4 ) .
included species : b . arethusae maury , 1935 ; b . guttula salvador & simone , 2012 ; b . minor trindade , 1956 .
- following salvador et al . ( 2011 ) , the genus brasilennea is considered a cerionidae , not a streptaxidae ;
b . arethusae usually has stronger ribs ( and fewer per whorl ) than other brasilennea species . quensen & woodruff ( 1997 ) attribute such strong ribs to protection against predators in cerion röding , 1798 ; the ribs strengthen the shell structure and make it harder for predators ( in this case , crabs ) to crush it . however , b . arethusae ' s possible predators are unknown : there is no record of crabs in itaboraí or other possible predators known for crushing shells , such as beetles ( symondson , 2004 ) ; still , some small mammals , also potential predators ( allen , 2004 ) , do occur in itaboraí , but possibly not in the same sequence as the brasilennea ( bergqvist et al . , 2006 ) . in any case , the shell of b . arethusae would be more resistant to predation due to its strong ribs .
etymology : maury ( 1935 ) considered the newfound species very similar to the african genus ennea and thus named the new genus brasilennea , meaning
brazilian ennea
.
shell shape , structure and sculpture : in respect to the general shell shape , brasilennea strongly resembles the typical cerionidae : a thick , pupiform and multispiral shell , with acuminated apex ( triphasic sensu gould , 1989 ) , hollow columella ( at least in first whorls ) , and the shape , placement and size of aperture ( schileyko , 1999b ) . also , b . arethusae has strong and regularlyspaced ribs that become less oblique towards the shell aperture , as commonly seen in cerionids ( schileyko , 1999b ) . the ribs in the other brasilennea species are weaker , but still raised and well - marked .
discussion : b . minor was originally described as a smaller sympatric variety of b . arethusae by trindade ( 1956 ) , but it was lately elevated to the category of species by brito ( 1967 ) , using the sympatry as one of the reasons for such . b . minor specimens are smaller than the smallest specimen of b . arethusae , showing no overlapping in size distribution . b . minor shows a smaller number of whorls than the other brasilennea species : usually 9 ( 8 or 10 whorls are not uncommon ) . the whorls of b . minor are more convex and its shell is weaker and thinner than the other species . it has a greater number of ribs and they are also weaker . the basal furrow in the body whorl is more weakly marked . b . minor also shows some variation in shell shape besides whorl number as , for example , thinner and more elongated specimens ( fig . 25 ) ; however , in a broader sense , this variation seems to be less than what is seen in b . arethusae .
os calcários da bacia de itaboraí ( paleoceno médio ) , rio de janeiro , brasil , abrigam uma rica fauna de gastrópodes pulmonados do paleoceno médio , tanto terrestres quanto dulciaquícolas . no presente trabalho realiza - se uma extensiva revisão taxonômica dessa paleofauna . duas novas espécies , eoborus fusiforme e gastrocopta itaboraiensis , são descritas , assim como um novo gênero , cortana . a classificação revisada encontra - se do seguinte modo : austrodiscus lopesi ( charopidae ) ; biomphalaria itaboraiensis ( planorbidae ) ;
brachypodella
britoi ( urocoptidae ) ; brasilennea arethusae , brasilennea guttula , brasilennea minor ( cerionidae ) ; bulimulus fazendicus , bulimulus trindadeae , cortana carvalhoi , cyclodontina coelhoi , itaborahia lamegoi , leiostracus ferreirai , plagiodontes aff . dentatus ( orthalicidae ) ; cecilioides sommeri ( ferussaciidae ) ; eoborus rotundus , eoborus sanctijosephi , eoborus fusiforme ( strophocheilidae ) ; gastrocopta mezzalirai , gastrocopta itaboraiensis ( gastrocoptidae ) ; temesa magalhaesi ( clausiliidae ) . a espécie strobilopsis mauryae foi considerada sinônimo de brasilennea arethusae ; bulimulus sommeri sinônimo de itaborahia lamegoi ; e vorticifex fluminensis sinônimo de eoborus sanctijosephi . a bacia conta com os registros fósseis mais antigos das famílias orthalicidae , gastrocoptidae , ferussaciidae e strophocheilidae . além disso , os registros de itaboraí das famílias charopidae , clausiliidae , cerionidae , e urocoptidae estão entre os mais antigos do mundo e , dentre esses , os de cerionidae , clausiliidae e urocoptidae merecem destaque por estarem bem afastados das distribuições atuais das famílias . ademais , os registros de itaboraí são os mais antigos para os gêneros austrodiscus , brachypodella , bulimulus , cecilioides , cyclodontina , eoborus , gastrocopta , leiostracus , plagiodontes e temesa . há três gêneros endêmicos na bacia : brasilennea , cortana e itaborahia . discussões adicionais sobre paleobiogeografia e evolução dessa paleofauna são oferecidas .
geographic and stratigraphic occurrence : known only from the type locality . the precise stratigraphic occurrence can ' t be assessed ; probably sequence s1 , the same sequence of occurrence of the other brasilennea species ( medeiros & bergqvist , 1999 ; bergqvist et al . 2006 ) .
diagnosis : shell bigger than other species . greatest width in central portion of shell . sculptured by stronger whorls , usually in lesser quantity . in some specimens , weaker ribs , in greater quantity ( similar to the other brasilennea species ) , may also occur in some specimens .
discussion : b . guttula is smaller than b . arethusae but larger than b . minor , and has more whorls than both ( b . guttula has 14 , while b . arethusae has 10 - 11 and b . minor has 8 - 9 ) . the most striking difference , however , is its
water drop
shape , i . e . , an acuminate spire ( the other species have wide , dome - shaped first whorls ) . regarding the other typical features of the genus , b . guttula shares all of them : the shell strength , the smooth protoconch , the sculpture pattern , the suture pattern and the most prominent feature of the genus , the two furrows on the body whorl . nevertheless , the holotype ( the only specimen known ) has the last portion of the body whorl broken , so for now there is no clue indicating if the peristome has a doubled aspect or if there was a parietal lamella and / or a columellar lamella as in the other two brasilennea species .
salvador , r . b . ; rowson , b . & simone , l . r . l . 2011 . rewriting the fossil history of cerionidae ( gastropoda , pulmonata ) : new family assignment of the brazilian paleocene genus brasilennea maury , 1935 . journal of molluscan studies , 77 : 445 - 447 . [ links ]
there are 20 species in the itaborahian fauna ( including four entirely new species ; marked in bold on the list below ) , which is a fair amount of diversity . almost all of them are land snails , with the exception of the freshwater biomphalaria species . the genera brasilennea , cortana and itaborahia are endemic to the basin .
salvador , r . b . ; rowson , b . ; simone , l . r . l . 2011 . re - writing the fossil history of cerionidae ( gastropoda , pulmonata ) : new family assignment of the brazilian paleocene genus brasilennea maury , 1935 . journal of molluscan studies 77 : 445 - 447 . [ pdf ]
therefore , here we propose that the specimens previously classified as s . mauryae are in fact juveniles of b . arethusae ( or fragments of it , namely the top of the spire ) . besides all the characters of the supposed specimens of s . mauryae being identical to what is found in the top region of the shell of b . arethusae ( like sculpture pattern , aperture shape , umbilicus shape , absence of teeth and lamellae etc . ) , they do not present a single character that could allow their classification as strobilopsids , such as long parietal lamellae , greatly extending themselves towards the shell ' s interior , and a thickened and reflected lip ( schileyko , 1998a ) . moreover , no deflection of the peristome was detected in the specimens identifiable as s . mauryae , another necessary character for confirming the generic attribution .
cerionidae : this family was not officially present in the basin until the recent revision of the genus brasilennea by salvador et al . ( 2011 ) , as it was previously classified in streptaxidae . there are three brasilennea species in itaboraí , which together consist on the second oldest record of cerionidae . the oldest is a probable cerion , named c . acherontis , from the upper cretaceous of montana , usa ( roth & hartman , 1998 ) . even if itaboraí ' s record is not the oldest , its location is very interesting , for the basin is greatly removed from the family ' s recent distribution : the islands of florida and the caribbean islands ( fig . 98 ) . despite the absence of recent cerionids in south america , it can be seen that the family ' s distribution included this area in the beginning of the cenozoic . it is even possible that , at that time , cerionidae had a more ample distribution , occurring from northwestern usa ( c . acherontis ) to rio de janeiro ( brasilennea ) .
salvador , r . b . ; rowson , b . ; simone , l . r . l . 2011 . rewriting the fossil history of cerionidae ( gastropoda : pulmonata ) : new family assignment of the brazilian palaeocene genus brasilennea maury , 1935 . journal of molluscan studies 77 : 445–447 . doi : 10 . 1093 / mollus / eyr021
salvador r . b . , rowson b . & simone l . r . l . ( 2011 ) . re - writing the fossil history of cerionidae ( gastropoda , pulmonata ) : new family assignment of the brazilian paleocene genus brasilennea maury , 1935 . journal of molluscan studies . 77 : 445 - 447 . , available online at urltoken [ details ]
important characters in the familiar allocation of b . britoi are : the conical - turriform shell shape , sculpture pattern and the body whorl detached from previous whorl , very common in urocoptidae ( schileyko , 1999a ) . the original illustration of ferreira & coelho ( 1971 : p . 471 , fig . 8 ) shows the body whorl greatly detached from the previous one . however , this degree of detachment is not seen in the specimens - the body whorl is only slightly detached ( fig . 88 ) . brachypodella britoi , as brasilennea arethusae , has few ribs per whorl , but they are raised and strong . quensen & woodruff ( 1997 ) relates these ribs to a greater protection against shell - breaking predators in cerion röding , 1798 . unfortunately , there is no known malacophagous predator in itaboraí in sequence s1 .
discussion : in the original description of e . sanctijosephi , the specific epithet had a hyphen , which has reappeared in some recent works , but must be omitted obeying the iczn ( 1999 ) rules ( articles 11 . 2 and 32 . 5 . 2 . 4 . 1 ) . e . sanctijosephi is certainly the largest pulmonate in itaboraí basin and the second species with the most numerous record ( the first is brasilennea arethusae ) . e . sanctijosephi differs from e . rotundus salvador & simone , 2012 by its larger size , conic shell , oval aperture , less reflected peristome and the way in which the upper palatal region of outer lip meets with the preceding whorl accompanying its outline . also , it differs from e . fusiforme sp . nov . by its much larger size , conic shell , less acuminated spire , slightly convex whorls and orthocline aperture .
discussion : b . arethusae is the type species of the genus by original designation and monotypy ( maury , 1935 ) . it is larger than the other species , presenting usually 11 whorls , though a few specimens have 10 whorls . a single specimen has 9 whorls ( fig . 13 ) , but it seems to be anomalous , since it also shows a slightly different shell shape , with the lip largely reflected and without the doubled aspect , and does not have the parietal lamella . the shell shape can vary slightly in the last whorls , which can be thinner and present the aperture more centrally located ( notably in the holotype , figs . 10 - 11 ) . in the same manner , the ribs in some specimens ( ~ 40 % ) can be weaker and more abundant ( figs . 10 - 12 ) , like those of other brasilennea species .
therefore , during the beginning of the cenozoic , it seems likely that the morphological variation in the shells of cerionidae would be greater , as shown by brasilennea . this would not be a complete surprise , since the family ' s ancestral stock may have also originated the urocoptidae as suggested by uit de weerd ( 2008 ) , a family with great morphological variation ( schileyko , 1999a ) .
spiral furrows on body whorl : the furrows in brasilennea ' s body whorl are well - marked and deeply set and comprise the most striking diagnostic feature of the genus . this feature is not known in cerionidae , but some urocoptids do show a single furrow in their body whorl ( schileyko , 1999a ) , such as in the genera apoma beck , 1837 , brachypodella beck , 1837 , mychostoma albers , 1850 and spirostemma pilsbry & vanatta , 1898 .
ferreira & coelho ( 1971 ) described strobilops mauryae ( figs . 14 - 15 ) , stating that it could be taken for fragments of b . arethusae and also that maury ( 1935 ) had committed such error when defining the paratype of b . arethusae ( amnh , 24239 ; fig . 16 ) as a juvenile . ferreira & coelho ( 1971 ) gave the key character to place their specimens in the genus strobilops pilsbry , 1893 : a tooth ( or
well - developed basal fold
) in the aperture ' s basal region . teeth and lamellae are typical of the family strobilopsidae , which , for many authors , contain only the genus strobilops , and are essential to the family ' s taxonomy ( schileyko , 1998a ) . in the illustration of s . mauryae presented by ferreira & coelho ( 1971 : 469 , fig . 6 ) , such tooth can be clearly seen . however , examining the type material , we saw that only the holotype ( figs . 14 - 15 ) presented such tooth . further examination revealed that the supposed tooth was in fact a grain of sediment attached to the shell .
brasilennea arethusae maury , 1935 : 4 ( figs . 1 - 5 ) ; oliveira , 1936 : 4 ; mezzalira , 1946 : 18 ; paula couto , 1949 : 11 ; magalhães & mezzalira , 1953 : 221 ( pl . 64 , fig . 262 , 262º ) ; trindade , 1956 : 15 ( pl . 3 , figs . 1c , 2c ) ; zilch in wenz , 1959 - 60 : 578 ( fig . 2025 ) ; brito , 1967 : 18 ( pl . 3 , fig . 6 ) ; jaeckel , 1969 : 822 ; parodiz , 1969 : 186 ( pl . 19 , figs . 3 , 12 ) ; palma & brito , 1974 : 396 ( pl . 1 , fig . 9 ) ; simone & mezzalira , 1994 : 51 ( pl . 15 , fig . 430 ) ; bergqvist et al . , 2006 : 59 ( fig . 74 ) ; salvador et al . , 2011 : 445 ( fig . 1a - g , l ) ; salvador & simone , 2012 : 2 ( figs . 5 - 6 ) .
aperture , peristome and lamellae : the basic shape of brasilennea ' s aperture is the same as in cerionidae , especially when considered together with the complete peristome , virtually straight parietally , and its duplicated aspect ( parallel lamella sensu maury , 1935 , projecting itself forwards , away from the lip , for a couple of millimeters ) . the apertural dentition is also very similar to the typical cerionidae , with a single , strong median parietal lamella reaching the peristome and also a spiral columellar lamella .
brasilennea minor : brito , 1967 : 19 ( pl . 3 , figs . 7 , 8 ) ; palma & brito , 1974 : 397 ( pl . 1 , fig . 10 ) ; simone & mezzalira , 1994 : 51 ( pl . 15 , fig . 431 ) ; bergqvist et al . , 2006 : 60 ( fig . 75 ) ; salvador et al . , 2011 : 445 ( fig . 1h - k ) ; salvador & simone , 2012 : 2 ( figs . 7 - 8 ) .
brasilennea maury , 1935 : 3 ; oliveira , 1936 : 4 ; mezzalira , 1946 : 18 ; magalhães & mezzalira , 1953 : 221 ; trindade , 1956 : 15 ; zilch in wenz , 1959 - 60 : 578 ; brito , 1967 : 18 ; jaeckel , 1969 : 822 ; parodiz , 1969 : 186 ; palma & brito , 1974 : 396 ; simone & mezzalira , 1994 : 51 ; bergqvist et al . , 2006 : 59 ; salvador et al . , 2011 : 445 ; salvador & simone , 2012 : 2 .
discussion : brasilennea was originally placed in the family streptaxidae . however , due to many morphological characters shared with cerionidae , and also to the fact that the similarities shared with streptaxidae were superficial , the genus was recently transferred to cerionidae by salvador et al . ( 2011 ) . since this topic was well explored elsewhere , it will not be discussed here ; instead we present only the formal diagnosis and description and limit ourselves to comparisons with other cerionidae ( and urocoptidae , when informative , since these two families were deemed sister taxa by uit de weerd , 2008 ) .
maury , c . j . ( 1935 ) . new genera and new species of fossil terrestrial mollusca from brazil . american museum novitates . 764 : 1 - 15 . , available online at urltoken ; = y page ( s ) : 4 - 6 , figs 1 - 5 [ details ]
( of strobilops mauryae ferreira & coelho , 1971 † ) ferreira , c . s . ; coelho , a . c . s . ( 1971 ) . novos gastrópodes pulmonados da bacia calcária de são josé de itaboraí , rj , brasil . geocronologia . anais da academia brasileira de ciências . 43 ( supl . ) : 463 - 472 . page ( s ) : 469 , fig . 6 [ details ]
( of strobilops mauryae ferreira & coelho , 1971 † ) salvador , r . b . ; simone , l . r . l . ( 2013 ) . taxonomic revision of the fossil pulmonate mollusks of itaboraí basin ( paleocene ) , brazil . papéis avulsos de zoologia . 53 ( 2 ) : 5 - 46 . page ( s ) : 9 - 11 [ details ] available for editors [ request ]
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maury ( 1935 : 6 ) considered the horizon from which this species was collected as
probably miocene , but possibly as young as pliocene .
salvador et al . ( 2011 : 445 ) report the itaborai limestones to be of paleocene age .
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urltoken
i museu de zoologia , universidade de são paulo . caixa postal 42 . 494 , 04218 - 970 , são paulo , sp , brasil ii e - mail : salvador . rodrigo . b @ urltoken iii e - mails : lrsimone @ urltoken , lrlsimone @ urltoken
key - words : itaboraí ; middle paleocene ; mollusca ; pulmonata ; rio de janeiro .
palavras - chave : itaboraí ; mollusca ; paleoceno médio ; pulmonata ; rio de janeiro .
itaboraí basin is located in the municipality of itaboraí ( fig . 1 ) , rio de janeiro state , brazil , and is one of the smallest basins in the country : it has an elliptical shape , with ~ 1 , 400 m in its biggest axis , ~ 500 m in the smallest and ~ 125 m of maximum depth ( rodrigues francisco & cunha , 1978 ; bergqvist et al . , 2006 ) . despite its small size , itaboraí basin houses an astounding fossil record in its limestones .
these limestones started to be explored in 1933 for the production of cement , and fossils were found there almost as soon as the excavations had begun . the first fossils discovered were the gastropods , but , as excavations proceeded , many other were found , including plants , mammals , reptiles , birds , palynomorphs , coprolites and , of course , more mollusks ( maury , 1929 , 1935 ; bergqvist et al . , 2006 ) . these fossils alongside the basin ' s geology were studied almost exclusively by researchers of two institutions from rio de janeiro : the museu nacional ( national museum ) and the divisão de geologia e mineralogia of the departamento nacional de produção mineral ( division of geology and mineralogy of the national department of mineral production ) .
the exploration lasted until 1984 , at which point the vast majority of itaboraí ' s outcrops were already destroyed by the quarrying , and little of its limestone remained . after the limestone extraction ceased , the quarry was abandoned and a lake formed in the basin , leaving the few remaining outcrops underwater ( bergqvist et al . , 2008 ) . therefore , it is currently hard to obtain new fossil specimens from itaboraí , as they can only be found in a few restricted sites above water level . as such , any posterior work depended heavily upon data from the literature and museum specimens ( e . g . , medeiros & bergqvist , 1999 ) . still , itaboraí is a very important fossiliferous site and on december 12 , 1985 a park was created for its preservation , called parque paleontológico de são josé de itaboraí ( beltrão et al . , 2001 ; bergqvist et al . , 2008 ) .
according to the last works that listed the fossil mollusks from itaboraí , there are 18 ( simone & mezzalira , 1994 ) or 17 species ( bergqvist et al . , 2006 ) in the basin . three more ( two new species and a new occurrence ) were added to this list by salvador & simone ( 2012 ) . in any case , up to this moment the studies dealing with itaboraí basin ' s molluscan fauna consist almost exclusively of the original descriptions ; no taxonomic revision under a larger scope has been so far produced . as such , many taxonomic problems and inconsistencies have been readily identified , most stemming from the fact that the original descriptions were overly based on comparisons with european genera and species . therefore , this work intends to review the taxonomy of the entire itaborahian molluscan fauna . as the original descriptions ( with the exception of maury , 1935 ) are too brief and incomplete , we expand and complement them , figuring all type material and other well - preserved specimens , and offering a proper diagnosis for each taxon . additionally , two new species , previously misidentified , were found in museum collections and are described herein .
itaboraí basin is a small tectonic depression dating from the mesozoic , probably related to the tectonic activities during the separation of gondwana ; it rests on a pre - cambrian crystalline basement , over the brazilian southeastern continental rift ( beurlen & sommer , 1954 ; rodrigues francisco & cunha , 1978 ; rodrigues francisco , 1989 ; medeiros & bergqvist , 1999 ; sant ' anna & riccomini , 2001 ; sant ' anna et al . , 2004 ) . the basin was then filled by carbonatic sediments during the paleocene ( rodrigues francisco & cunha , 1978 ; medeiros & bergqvist , 1999 ) . the resulting limestones were named
itaboraí formation
by oliveira ( 1956 ) , but this name was almost never used afterwards ; the majority of authors prefer simply
itaboraí basin
. we follow this practice here .
the first work dealing with the basin ' s geology ( leinz , 1938 ) defined three rock horizons : laminated limestone , gray limestone and eluvial sediment . this definition still remains basically valid , having received only posterior refinements . the most up - to - date geological profile ( medeiros & bergqvist , 1999 ) defines and names two stratigraphic sequences for the basin ' s paleocene limestones : sequence s1 lies at the bottom , directly above the pre - cambrian rocks ; and sequence s2 consists of sediments that filled fissures opened in s1 by water activity .
sequence s1 is composed of an intercalation of carbonatic rocks of chemical and detrital origins . the chemically originated rocks ( named facies a ) are mainly travertine , linked to hydrothermal activity inside the basin . the grey detrital limestones ( facies b ) are the product of gravitational and hydrodynamic fluxes towards the basin ' s interior and contain all known fossil mollusks as well as some plant , reptilian and mammalian remains . facies c is composed of oolitic - pisolitic limestone , being associated to facies a .
sequence s2 is also carbonatic and the result of torrent and gravitational flows . its clastic rocks contain the bulk of the basin ' s fossil vertebrate fauna as well as plants and palynomorphs . a third sequence , named s3 , is composed of conglomerates and was deposited on top of the others much later , in the eocene - oligocene . it harbors reptilian and mammalian fossils .
after much disagreement about itaboraí basin ' s age , the mammalian fossils allowed the correlation with the upper paleocene fauna of rio chico formation , argentina ( paula couto , 1952 ) . it is currently agreed that both sequences s1 and s2 belong to a time interval ranging from the end of the lower paleocene to the beginning of the upper paleocene ( from about 59 to 57 ma , according to bergqvist et al . 2006 ) , which has been informally named
middle paleocene
( marshall , 1985 ; medeiros & bergqvist , 1999 ) . sequence s1 is younger , of itaborahian age ( according to the nomenclature of the south american land mammalian age ) , while s2 is partly itaborahian and partly riochican ( bergqvist & ribeiro , 1998 ) .
institutional abbreviations : amnh : american museum of natural history , new york , usa . dgm : divisão de geologia e mineralogia ( recently transformed into the mct ) , rio de janeiro , rj , brazil . mct : museu de ciências da terra , rio de janeiro , rj , brazil . mnrj : museu nacional , rio de janeiro , rj , brazil . mzsp : museu de zoologia da universidade de são paulo , são paulo , sp , brazil .
examined material : the material from itaboraí analyzed here ( 79 lots ; 322 specimens ) includes all the types and is housed in the collections of the above mentioned institutions . unfortunately , the holotypes of two species ( austrodiscus lopesi and biomphalaria itaboraiensis ) are lost , i . e . they could not be located in the museums that housed them , respectively mnrj and dgm . only the reasonably well - preserved specimens , i . e . those useful for analysis , were included . the specimens in bad preservational state , including internal molds , were mostly excluded . since most of the basin ' s outcrops are gone and further explorations commonly do not result in good material , all the specimens presented here are from museum collections . in large part , this is the same material used in the species ' original descriptions , but it also contains specimens from later collecting .
shell terminology and measures : we follow here the terminology of moore et al . ( 1952 ) and cox ( 1955 ) for molluscan shells and their structures . additionally , we follow breure ( 1979 ; p . 10 , figs . 1 - 7 ) for the terminology of whorl convexity and the umbilicus and tillier ( 1989 ; p . 7 , fig . 1c ) for the terminology of the peristome regions ( fig . 2 ) . all shell measures presented here ( fig . 2 ) are in accordance with moore et al . ( 1952 ) : h = shell length ( or height ) ; d = greatest width ( or diameter ) of shell ; s = spire length ( without the aperture ) ; h = aperture height ; d = aperture width ; θ ' = spire angle ( the
mean spire angle
sensu moore , 1960 , approximated to the closest multiple of five ) . all measures were taken with a digital caliper or , in the case of microgastropods ( smaller than 5 mm ) , with the aid of coreldraw ® graphics suit x5 . unfortunately , not every shell is well - preserved or even completely preserved and thus some measures could not be precisely taken or taken at all ( such imprecise measures are indicated by italicized numbers ) . when specimens are abundant , the mean and standard deviation is presented . the whorl counting method follows janssen ( 2007 ) , i . e . , excluding the semicircular nucleus of the protoconch , and thus the total number of whorls can vary a little ( ¼ to ½ whorl ) when compared to other works .
nomenclatural update : here we follow the work of bouchet et al . ( 2005 ) for the nomenclature of families and sub - families , with the addition of the more recent work of uit de weerd ( 2008 ) with the urocoptoidea . therefore , a few nomenclatural changes had to be made in the classic literature of itaborahian molluscan fauna : ( 1 ) the family bulimulidae is now considered a subfamily of orthalicidae ; ( 2 ) likewise , odontostomidae is now considered a subfamily of orthalicidae ; ( 3 ) the family megalobulimidae is now considered a subfamily of strophocheilidae .
we also follow the works of schileyko ( 1998a , 1998b , 1999a , 1999b , 2000 , 2001 ) , which offers the description and diagnosis of families , subfamilies and genera based on morphological characters . other works , dealing more specifically with one or other taxon , were also followed ; among these , the works of breure ( 1978 , 1979 ) on the bulimulinae stand out .
geographic and stratigraphic occurrence : known only from itaboraí basin : limestone sequence s1 ( medeiros & bergqvist , 1999 ; bergqvist et al . , 2006 ) .
diagnosis : shell pupiform ( expect for b . guttula : spire sharply acuminated ) . ribs regularly spaced . greatest width in central portion of shell length . body whorl with two well - marked spiral furrows , one central and the other basal . peristome complete , well - marked and with duplicated aspect . single and strong median parietal lamella , reaching the peristome . columellar lamella present .
re - description : shell dextral , pupiform ( expect for b . guttula : spire sharply acuminated ) , with greatest width in central portion of shell length . columella hollow , at least in first whorls . whorls flat to slightly convex . suture well - marked , linear , practically perpendicular ( horizontal ) to columellar axis , becoming more oblique towards last whorls . shell sculptured by well - defined and raised ribs , regularly spaced , becoming less oblique towards last whorls . first two to three whorls ( protoconch ) smooth ; transition to teleoconch clear . body whorl with two well - marked spiral furrows ( one central and the other basal ) , that can be seen as two parallel folds in shell ' s inner surface . aperture large , approximately semicircular , with parietal and columellar lips straight ; other lip regions rounded . peristome complete , well - marked , virtually straight parietally , with duplicated aspect ( parallel lamella sensu maury , 1935 , projecting itself forward ) . single and strong median parietal lamella reaching peristome . columellar lamella present . umbilicus narrow .
strobilops mauryae ferreira & coelho , 1971 : 469 ( fig . 6 ) ; palma & brito , 1974 : 397 .
strobilopsis mauryae : simone & mezzalira , 1994 : 49 ( pl . 14 , fig . 413 ) ; bergqvist et al . , 2006 : 60 ( fig . 76 ) . ( error )
paratypes : amnh 24238 ( 1 specimen , examined ; fig . 12 ) , 24239 ( 1 specimen , examined ; fig . 16 ) .
type locality : limestones of parque paleontológico de são josé de itaboraí , rio de janeiro , brazil . park ' s center coordinates : 22º50 ' 20
s , 42º52 ' 30
w .
geographic and stratigraphic occurrence : known only from the type locality : sequence s1 ( medeiros & bergqvist , 1999 ; bergqvist et al . , 2006 ) .
etymology : due to the terrestrial habits of the species , maury ( 1935 ) dedicated it to arethusa ( sometimes also spelled arethousa ) , a nymph from greek mythology . there are mentions to two nymphs of such name in the myths : one is a nereid ( or even a goddess of springs ) while the other is one of the hesperides ( kerényi , 1951 ) . maury did not specify which one she was referring to , stating only that it was a
sylvan nymph
. thus , the name probably refers to the latter since the previous is a water - related being .
measures ( in mm ) : holotype : 11 whorls ; h = 23 . 6 ; d = 13 . 2 ; s = 15 . 9 ; h = 7 . 5 ; d = 6 . 1 . paratypes : amnh 24238 : 11 whorls ; d = 12 . 6 ; s = 15 . 8 . amnh 24239 ( juvenile ) : 6 or 7 whorls ( shell apex covered by sediment ) ; h = 4 . 1 ; d = 7 . 7 ; s = 1 . 8 ; d = 3 . 2 . mean ( n = 35 ) : 11 whorls ( eventually 10 ) ; h = 21 . 6 ± 2 . 2 ( max 24 . 9 ; min 17 . 3 ) ; d = 11 . 6 ± 1 . 2 ( max 14 . 4 ; min 9 . 4 ) ; s = 14 . 9 ± 1 . 7 ; h = 6 . 9 ± 0 . 9 ( max 7 . 8 ; min 5 . 2 ) ; d = 5 . 5 ± 1 . 0 ( max 6 . 9 ; min 3 . 2 ) .
examined material : types . dgm 4222 - i ( 1 specimen ) , 4998 - i ( 8 specimens ) , 5002 - i ( 25 specimens ) , unnumbered ( 7 specimens ) ; mnrj 3346 - i ( 2 specimens ) , 3348 - i ( 2 specimens ) , 4338 - i ( 5 specimens ) ; mzsp 86321 ( 20 specimens ) , 86322 ( 1 specimens ) , 86324 ( 4 specimens ) . type material of strobilops mauryae : mnrj 5020 - i ( holotype ) , 5021 - i ( paratype , 4 specimens ) .
holotype : mct 6940 - i ( examined ; figs . 17 - 21 ) .
etymology : the name refers to the species ' intriguing shell shaped as a water drop .
diagnosis : outline shaped like water drop ( spire sharply acuminate ) instead of pupiform . greatest width on body whorl . larger number of whorls ( about 14 ) .
measures ( in mm ) : holotype : 14 whorls ; h = 13 . 6 ( aperture broken ) ; d = 6 . 5 ; s = 11 . 3 .
diagnosis : shell small ( smallest species in genus ) . greatest width in central portion of shell . profile of whorls slightly convex . sculptured by large number fine ribs . basal furrow in body whorl more weakly marked .
re - description : shell small , multispiral , pupiform , with acuminated apex . greatest width in central portion of shell ; diameter ~ ½ shell length . spire angle ~ 45º . protoconch dome shaped , blunt , smooth ; transition to teleoconch clear . columella hollow ( at least on first whorls ) . profile of whorls slightly convex . suture well - marked , linear , practically perpendicular ( horizontal ) to columellar axis , becoming more oblique towards last whorls . sculptured by fine and raised ribs , regularly distributed , becoming less oblique towards last whorls , and in large numbers ( ~ 70 on penultimate whorl ) . body whorl with two spiral furrows , one central and the other basal , placed equidistantly from the upper furrow and the bottom of the shell ; basal furrow more weakly marked than the upper one . aperture large , orthocline , approximately semicircular ( parietal and columellar lips straight , others rounded ) ; ~ ⅓ shell length . peristome complete and well - marked , with duplicated aspect ( parallel lamella sensu maury , 1935 , projecting itself forwards away from the lip for a couple of millimeters ) . single and strong median parietal lamella , reaching the peristome and extending itself towards shell ' s interior up until ~ ¼ of body whorl . columellar spiral lamella extending itself towards interior . body whorl ~ ½ shell length . umbilicus narrow .
measures ( in mm ) : holotype : 9 whorls ; h = 9 . 0 ; d = 5 . 0 ; s = 6 . 3 ; d = 2 . 1 . mean ( n = 16 ) : 9 whorls ( eventually 8 or 10 ) ; h = 11 . 6 ± 1 . 9 ( max 14 . 9 ; min 8 . 4 ) ; d = 5 . 8 ± 0 . 6 ( max 6 . 7 ; min 4 . 8 ) ; s = 7 . 9 ± 1 . 2 ; h = 3 . 6 ± 0 . 2 ( max 3 . 9 ; min 3 . 2 ) ; d = 3 . 2 ± 0 . 4 ( max 4 . 0 ; min 2 . 5 ) .
examined material : holotype . dgm 4224 - i ( 1 specimen ) , 4999 - i ( 9 specimens ) , unnumbered ( 1 specimen ) ; mnrj 3346 - i ( 1 specimen ) , 4338 - i ( 2 specimens ) ; mzsp 86323 ( 2 specimens ) .
austrodiscus lopesi ferreira & coelho , 1989 : 193 ( figs . 1 - 2 ) ; simone & mezzalira , 1994 : 50 ( pl . 15 , fig . 417 ) ; bergqvist et al . , 2006 : 59 .
geographic and stratigraphic occurrence : known only from the type locality . there is no information in the literature about which sequence this species occurs .
etymology : species dedicated to the zoologist dr . hugo s . lopes ( instituto oswaldo cruz , rio de janeiro , brazil ) .
discussion : the genus austrodiscus parodiz , 1957 is normally allocated to the family endodontidae ( schileyko , 2001 ) , a family endemic to the pacific islands ( solem , 1976 , 1979 , 1981 ) . schileyko ( 2001 ) still considers that endodontidae contains some species in south america and on saint helena island , but other authors state that these species actually belong to charopidae , a family with a broader distribution : the americas , the pacific islands , oceania southern africa and saint helena island ( solem , 1981 ; fonseca & thomé , 1993 ) . it has been a common practice in revisionary work dealing with the supposed american endodontids to reallocate them in charopidae , as in fonseca & thomé ( 1993 ) , who dealt specifically with austrodiscus . such resolution is here adopted .
all type material of austrodiscus lopesi ( totaling 9 specimens , according to ferreira & coelho , 1989 ) has disappeared ; it could not be found in the museum ' s collection ( mnrj ) or in the records of lent material ( since it is common practice in this institution not to lend type material ) . unfortunately , no additional material exists .
ferreira & coelho ( 1989 ) , when describing the species , decided for its placement in the genus austrodiscus due to its smooth protoconch . however , this character was contested for the genus by fonseca & thomé ( 1993 ) , who stated that the protoconch is sculptured . however , this character cannot be confirmed only by the illustration in the species original description . without having the type material , it is impossible to conduct a proper taxonomic revision and thus the allocation of a . lopesi in the genus austrodiscus could not be confirmed or contested . as such , the only alteration proposed here is the change in a . lopesi ' s familiar allocation , transferring it from endodontidae to charopidae .
family clausiliidae subfamily neniinae genus temesa h . & a . adams , 1855 temesa magalhaesi ( trindade , 1953 ) comb . nov . ( figs . 27 - 33 )
clausilia magalhaesi trindade , 1953 : 40 ( fig . 1 ) ; brito , 1967 : 14 ( pl . 3 , figs . 4 , 5 ) ; palma & brito , 1974 : 397 ( pl . 1 , fig . 6 ) ; simone & mezzalira , 1994 : 50 ( pl . 14 , fig . 414 ) ; bergqvist et al . , 2006 : 59 ( fig . 71 ) .
etymology : species dedicated to prof . júlio de magalhães ( faculdade nacional de filosofia , rio de janeiro , brazil ) .
diagnosis : shell cylindrical - fusiform , broad and robust , with acuminated apex . ribs very thin and weak . parietal lamella high , vertical ; apparently the single lamella present .
re - description : shell medium - sized , sinistral , multispiral , thin ( but broad for genus ) , cylindricalfusiform , with acuminated apex . greatest width in central portion of shell ( antepenultimate whorl ) ; diameter ~ ⅓ shell length . spire angle ~ 20º . protoconch flattened , blunt , smooth . profile of whorls slightly convex , eventually flat . suture well - marked , oblique ( diagonal ) to columellar axis . sculptured by fine ribs ( ~ 85 on penultimate whorl ) , oblique to columellar axis . body whorl not thinned , adnate to the spire . peristome weakly reflected , supposedly complete . parietal lamella high , vertical , median , reaching the peristome and extending itself towards interior . apparently without other teeth and / or lamellae .
mean measures ( in mm ; n = 4 ) : 10 whorls ( eventually 11 ) ; d = 5 . 1 ± 0 . 2 ( min 4 . 8 ; max 5 . 3 ) .
discussion : the only clausiliid from itaboraí basin was originally placed in the genus clausilia dreparnaud , 1805 . however , clausilia is a recent genus from central europe , of the european subfamily clausiliinae . due to this biogeographical incongruence and additional morphological characters , here we opted for the reallocation of c . magalhaesi in a genus of neniinae , a strictly latin american subfamily ( schileyko , 2000 ) . it is also important to emphasize that all latin american species of clausilia have been reallocated in genera of neniinae ( schileyko , 2000 ) .
there is only a single recent clausiliid in brazil , nenia orbignyi ancey , 1892 , in the state of mato grosso ( simone , 2006 ) , despite nenia h . & a . adams , 1855 being a caribbean - endemic genus ( loosjes & loosjes - van bemmel , 1966 ) . moreover , schileyko ( 2000 ) restricted the genus to a single species , n . tridens ( chemnitz , 1786 ) , from puerto rico . as such , the most obvious choice for the generic reallocation of c . magalhaesi would perhaps be the genus nenia , which supposedly occurs in brazil . however , due to doubts regarding the classification of the brazilian species n . orbignyi and also to the astounding similarity of the itaborahian fossil to the recent south - american genus temesa h . & a . adams , 1855 , c . magalhaesi is here reallocated to this later genus . despite being absent in brazil , temesa occurs in various neighboring countries : peru , bolivia , colombia , and , with some doubt , in argentina ( loosjes & loosjes - van bemmel , 1966 ; schileyko , 2000 ; nordsieck , 2005 ) .
the decision for the reallocation in temesa is due to a vast array of shared morphological characters : cylindrical - fusiform and thin shell , spire with acuminated apex , protoconch present ( i . e . , non - decollated ) , number of whorls ( 10 or 11 ) , body whorl not thinned and adnate to the spire , peristome weakly deflected , absence of lamellae or teeth in the aperture ( besides the parietal lamella ) . usually , the lamellae of temesa species cannot be seen in the aperture ; they can be found in the inner portion of shell . unfortunately , the presence of additional lamellae could not be assessed in this fossil due to the specimens ' state of preservation : there is not even one specimen with an intact aperture , only one has part of the aperture preserved ( figs . 28 - 33 ) . however , to discover if the other lamellae are present or not , it would be necessary to break the aperture and part of the body whorl and , due to the small number of specimens , this course of action was discarded .
temesa magalhaesi differs from the other species in the genus mainly by its broader and more robust shell and the spire apex sharply acuminated . the genus does not possess the clausilial apparatus , a typical structure in the family ; however , such structure is not often preserved in the fossil record .
family ferussaciidae genus cecilioides férussac , 1814 cecilioides sommeri ( ferreira & coelho , 1971 ) comb . nov . ( figs . 34 - 35 )
carychium sommeri ferreira & coelho , 1971 : 467 ( fig . 4 ) ; palma & brito , 1974 : 391 ; simone & mezzalira , 1994 : 49 ( pl . 14 , fig . 405 ) ; bergqvist et al . , 2006 : 59 ( fig . 70 ) .
holotype : mnrj 5016 - i ( xamined ; figs . 34 - 35 ) .
etymology : species dedicated to prof . friedrich w . sommer ( divisão de geologia e mineralogia do departamento nacional de produção mineral , dnpm , rio de janeiro , brazil ) .
diagnosis : shell oval . aperture sub - oval . peristome reflected , slightly thickened .
re - description : shell diminutive , oval , smooth , with blunt spire apex ; 6 whorls ( eventually 5 ) . greatest width on body whorl ; width ~ ½ shell length . spire angle ~ 50º . protoconch smooth , blunt , broad , domeshaped . profile of whorls flattened . suture weakly marked , practically perpendicular ( horizontal ) to columellar axis . aperture small , orthocline , sub - oval ; ~ 2 / 5 shell length . teeth or lamellae absent . peristome reflected , slightly thickened . body whorl ~ ½ shell length . umbilicus imperforated .
measures ( in mm ) : holotype : 6 whorls ; h = 2 . 6 ; d = 1 . 3 ; s = 1 . 5 ; h = 1 . 0 ; d = 0 . 8 .
discussion : the species was originally described in the genus carychium , a holartic genus and one of the few exclusively terrestrial animals of the family ellobiidae ( morton , 1955 ; barker , 2001 ) . the apertural dentition is conspicuous in the family ( martins , 1996 ) and was the single character used by ferreira & coelho ( 1971 ) in their original classification . these authors stated that the species presented a
greatly evident
single columellar tooth . in the original illustration presented by them ( ferreira & coelho , 1971 : 468 , fig . 4 ) , such tooth can be clearly seen . however , after examining the type specimens , only the holotype seemed to possess this tooth . further examination under stereomicroscope revealed that the supposed tooth was in fact a grain of sediment placed in such a manner that a quick glance could take it for an actual shell structure . after this grain was removed , a toothless aperture was revealed . therefore , the original classification of this species in that genus , done exclusively due to the apertural dentition , is mistaken . in the absence of such dentition , we propose here the reallocation in the family ferussaciidae .
cecilioides consobrina ( orbigny , 1841 ) . the two species of geostilbia , g . gundlachi ( pfeiffer , 1850 ) and g . blandiana crosse , 1886 , were both previously classified in the genus cecilioides ( morretes , 1949 ; salgado & coelho , 2003 ) . we propose here that the itaborahian species is reallocated in the genus cecilioides .
the oval shell shape of cecilioides sommeri is very similar to the norm in this genus and is especially similar to c . consobrina due to the more cylindrical shell and to the aperture less elongated in the upper palatal region . meanwhile , geostilbia species show more acuminated spires , giving a more conical aspect to their shells . c . sommeri differs from the other species in the genus by the sub - oval aperture and by the reflected and lightly thickened peristome . such characters are not uncommon in the family , occurring in some species of the european genus ferussacia risso , 1826 ; however , the latter show more acuminated spires and much larger sizes .
family orthalicidae subfamily bulimulinae genus bulimulus leach , 1814 bulimulus fazendicus maury , 1935 ( figs . 36 - 37 )
bulimulus fazendicus maury , 1935 : 7 ( figs . 10 , 11 ) ; oliveira , 1936 : 5 ; mezzalira , 1946 : 18 ; magal hães & mezzalira , 1953 : 218 ( pl . 64 , fig . 257 ) ; trindade , 1956 : 14 ( pl . 3 , figs . 1d , 2d ) ; brito , 1967 : 16 ( pl . 2 , fig . 1 ) ; palma & brito , 1974 : 393 ( pl . 1 , fig . 3 ) ; breure , 1979 : 137 ; simone & mezzalira , 1994 : 50 ( pl . 15 , fig . 420 ) ; bergqvist et al . , 2006 : 57 ( fig . 64 ) .
bulimulus ( sensu lato ) fazendicus : parodiz , 1969 : 182 . itaborahia fazendicus : breure , 1978 : 237 .
paratype : amnh 24242 ( 1 specimen , examined ; fig . 37 ) .
geographic and stratigraphic occurrence : known only from the type locality : sequences s1 and s2 ( medeiros & bergqvist , 1999 ; bergqvist et al . , 2006 ) .
etymology : reference to place of discovery , the then called fazenda são josé , of which itaboraí basin was a part .
diagnosis : shell small , narrow . spire high . profile of whorls flat . aperture small and narrow , elliptical and slightly trapezoid .
re - description : shell small , conical - oval , narrow , with acuminated apex . spire high ; 8 whorls . shell smooth , except for growth lines . greatest width on body whorl ; width ~ ½ shell length . spire angle ~ 55º . protoconch apparently smooth ; transition to teleoconch not clear . suture well - marked , slightly oblique ( diagonal ) to columellar axis . profile of whorls flat . aperture small and narrow , orthocline , elliptical and slightly trapezoid ; ~ ⅓ shell length . lamellae or teeth absent . peristome thin , weakly reflected ( only in the columellar region ) . body whorl ~ ⅓ shell length . umbilicus narrow , partially covered by the lip .
measures ( in mm ) : holotype : 8 whorls ; h = 16 . 4 ; d = 9 . 1 ; s = 10 . 5 ; h = 6 . 1 ; d = 5 . 1 . paratype : 8 whorls ; h = 16 . 3 ; d = 9 . 1 ; s = 10 . 6 ; h = 6 . 1 ; d = 5 . 5 .
examined material : types . dgm unnumbered ( 8 specimens ) ; mnrj 4339 - i ( 4 specimens ) ; mzsp 86326 ( 1 specimen ) .
discussion : protoconch sculptural pattern is an important character in bulimulinae taxonomy ( breure , 1978 , 1979 ; schileyko , 1999a ) . the shell of bulimulus fazendicus is completely smooth , including the protoconch . this could be a diagnostic feature of this species but could also be a preservation artifact , since this kind of sculpture is very delicate and can be easily erased during fossil diagenesis . this same statement is valid for the others orthalicids from itaboraí . maury ( 1935 ) comments that one specimen showed vertical ribs on the second or third whorl and breure ( 1978 ) says that the protoconch of b . fazendicus shows vertical ribs ; however , this character could not be confirmed .
maury ( 1935 ) , citing a personal communication from henry a . pilsbry , stated that b . fazendicus does not seem to be closely related to any other bulimulus species . however , b . fazendicus is very similar to highspired species that also present a narrow aperture such as b . felipponei marshall , 1930 . b . fazendicus differs from the other species by its slightly flattened whorls , suture more weakly marked and trapezoid aperture ( broader than tall ) .
bulimulus trindadeae ferreira & coelho , 1971 : 470 ( fig . 7 ) ; palma & brito , 1974 : 394 ; breure , 1979 : 137 ; simone & mezzalira , 1994 : 50 ( pl . 15 , fig . 424 ) .
itaborahia trindadeae : bergqvist et al . , 2006 : 58 ( fig . 69 ) .
holotype : mnrj 5022 - i ( examined ; figs . 38 - 39 ) .
etymology : species dedicated to prof . nicéa m . trindade ( faculdade nacional de filosofia , rio de janeiro , brazil ) .
diagnosis : shell small , conical , narrow . spire high . aperture elliptical , tall and narrow .
re - description : shell small , conical , narrow . spire high . shell smooth , except for growth lines . greatest width ( supposedly ) on body whorl ; width ~ ⅓ shell length . spire angle ~ 35º . protoconch blunt , apparently smooth ; transition to teleoconch unclear . profile of whorls slightly convex . suture well - marked , oblique ( diagonal ) to columellar axis ; less oblique on first whorls . aperture supposed elliptical , tall and narrow , orthocline ; ~ 2 / 5 shell length .
measures ( in mm ) : holotype : probably 6 whorls ; h = 9 . 4 ; d = 3 . 3 ; s = 5 . 4 . paratype : probably 6 whorls ; h = 5 . 6 ; d = 2 . 6 .
discussion : the few specimens of b . trindadeae are in a bad state of preservation , including the type specimens : the apertures are broken or completely covered by sediment . it is impossible to even tell if a shell is an adult or not and , therefore , the precise number of whorls could not be counted . in the same manner , it is impossible to confirm the presence of teeth or lamellae . however , as stated below , there is a set of characters that guarantees its validity as a distinct species .