the apennine yellow-bellied toad ( bombina pachypus ) is a species of toad in the bombinatoridae family endemic to italy .

its natural habitats are temperate forests , temperate grassland , swamps , freshwater marshes , intermittent freshwater marshes , arable land , pastureland , ponds , open excavations , irrigated land , and seasonally flooded agricultural land .

it is threatened by habitat loss .

apennine yellow - bellied toad

fire - bellied toad & apennine yellow - bellied toad . george boulenger , tailless batrachians of europe , vol . 1 ( 1897 ) .
genetic diversity and phylogeography of the apennine yellow - bellied toad bombina pachypus , with implications for conservation .
minden pictures stock photos - apennine yellow - bellied toad ( bombina pachypus ) showing yellow underside , acqua cheta river , foreste casentinesi nation . . .
genetic diversity and phylogeography of the apennine yellow - bellied toad bombina pachypus , with implications for conservation . - pubmed - ncbi
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fire - bellied toad & apennine yellow - bellied toad . george boulenger , tailless batrachians of europe , vol . 1 ( 1897 ) . urltoken urltoken
reversing the decline of the endangered apennine yellow - bellied toad in liguria ( northern italy ) through site restoration and population reinforcement .
the yellow - bellied toad ( bombina variegata ) on the green moss . brown frog with yellow belly with green and brown backgroun .
information on the appenine yellow - bellied toad is currently being researched and written and will appear here shortly .
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yellow _ bellied _ toad
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cesbin srl , 2017 . reversing the decline of the endangered apennine yellow - bellied toad in liguria ( northern italy ) through site restoration and population reinforcement . final report .
the apennine yellow - bellied toad ( also called bombina pachypus ) is a species of toad in the bombinatoridae family endemic to italy . content licensed under creative commons attribution . source : urltoken pictures sources : urltoken urltoken
the european fire - bellied toad ( bombina bombina ) captured close up in moss .
canestrelli , d . , cimmaruta , r . , constantini , v . , and nascetti , g . ( 2006 ) . ' ' genetic diversity and phylogeography of the apennine yellow - bellied toad
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the toad’s odd coloring on its underbelly—black and white with large spots of yellow—serves a very distinct purpose . it is a warning to predators that the apennine yellow - bellied toad is poisonous and not to be messed with . it secretes a toxic substance when it is threatened . the substance is not all bad ; it is also antimicrobial and antifungal .
stagni , g . , dall ' olio , r . , fusini , u . , mazzotti , s . , scoccianti , c . , and serra , a . ( 2004 ) . ' ' declining populations of apennine yellow bellied toad
cartoon frog prince with golden crown and blue mantle . green toad with yellow belly . character of children s fairy tale . colorful flat vector design
barberio , c . , delfino , g . , and mastromei , g . ( 1987 ) . ' ' a low molecular weight protein with antimicrobial activity in the cutaneous ' venom ' of the yellow - bellied toad (
di cerbo , a . r . and ferri , v . 1996 . preliminary data on the ecological observations of the appennine yellow - bellied toad ( bombina pachypus ) in abruzzo , central italy . naturschutzreport 11 ( 1 ) : 91 - 99 .
guarino , f . m . , picariello , o . and pellegrini , m . 2006 . bombina pachypus ( bonaparte , 1838 ) . ululone appenninico / apennine yellow - bellied toad . in : sindaco , r . , doria , g . , razzetti , e . and bernini , f . ( eds ) , atlante degli anfibi e dei rettili d’italia . , pp . 272 - 277 . societas herpetologica italica , edizioni polistampa . , firenze .
( fire - bellied toad ) hybridise in overlapping areas in austria , hungary , bulgaria and western ukraine . the high proportion of hybrids , although often not abundant , demonstrates the ongoing speciation process in the basin .
) are found in the south with abundances decreasing from south to north . the natterjack toad occurs regularly along the river . the
stagni , g . , dall’olio , r . , fusini , u . , mazzotti , s . , scoccianti , c . and serra , a . 2005 . declining populations of apennines yellow - bellied toad bombina pachypus ( bonaparte , 1838 ) in northern appennines , italy : is batrachochytrium dendrobatidis the main cause ? . ital . j . zool . 71 ( suppl . 2 ) : 151 - 154 .
) are typical mediterranean species and occur from the delta marshes to montélimar . common frog populations have decreased since the 1950s . the midwife toad (
hofman , s . , spolsky , c . , uzzell , t . , cogălniceanu , d . , babik , w . , and szymura , j . m . ( 2007 ) . ' ' phylogeography of the fire - bellied toads
hofman , s . , spolsky , c . , uzzell , t . , cogălniceanu , d . , babik , w . and szymura , j . 2007 . phylogeography of the fire - bellied toads , bombina : independent pleistocene histories inferred from mitochondrial genomes . molecular ecology 16 : 2301 - 2316 .
is endemic to italy . it ranges from the southern tip of the italian mainland , north through the apennine region ( or in italian , appenine ) , where it stops just south of the po river valley ( dimartino and ferri 2002 ) . this species occurs in both terrestrial and freshwater habitats ( andreone and corti 2006 ) . it is commonly found in unshaded pools in forests and open areas ( andreone and corti 2006 ) , including pools formed in ditches , irrigation areas , farmland , or pasture land ( guarino et al . 1998 ) .
, is toxic . the secretions are discharged through the skin when the animal is threatened or stimulated ( barberio et al . 1987 ) . these secretions have also been found to be antimicrobial and antifungal ( barberio et al . 1987 , mastromei et al . 1991 ) . when threatened , the animal will arch its back in response to expose its colorful yellow and black underside as a warning of its toxicity to possible predators ( bajger 1980 ) . this arching of the body is known as the
unken reflex
and the behavior is found throughout the genus
the dorsal surface can range from a dark - tan to a dark - gray color , and the surface of the skin is tubercular . the eye has a triangular pupil . this species is regarded by some workers as a subspecies of
is diurnal , when it is active . hibernation takes place from november to late april . this species typically becomes active in may and begins breeding almost immediately , with the period of activity and breeding lasting until september . although
is considered a prolonged breeder due to the long breeding period , a single animal does not reproduce continuously within that period . multiple distinct periods of calling and oviposition alternate with non - reproductive activity . this reproductive plasticity presumably allows
populations to make maximum use of the temporary pools preferred for breeding ( guarino et al 1998 ) . in addition , sperm was found to be constantly present in male testes , but the testes do not always exhibit all stages of spermatogenesis . males apparently use sperm from the preceding fall for the earlier periods of breeding ( guarino et al . 1998 ) .
the clutch size ranges from a few eggs to a few tens of eggs . it is thought that each female lays a few eggs during each wave of reproduction , but in different temporary pools . however , it is not clear how often females reproduce in the wild . female frogs in captivity showed varying reproductive activity , with one female laying several eggs multiple times during the month of june , and a few eggs during a short period in september ; another female laid a few eggs during a single short period in july ; a third captive female did not lay any eggs ( guarino et al . 1998 ) .
this species is relatively long - lived , with an average lifespan of 8 years , and an apparent maximum lifespan of 16 years . sexual maturity is attained during the third year of life ( guarino et al . 1995 ) .
this species is thought to be common in suitable habitat ( iucn 2008 ) but has also been reported as becoming increasingly rare in some parts of its range in southern italy ( caputo et al . 1992 ) .
faces a threat from chytrid fungus . this species was the first italian amphibian to be confirmed with chytrid infection , in the summer of 2001 ( stagni et al . 2004 ) . neometamorph
collected from the hills in the province of bologna experienced high mortality , dying within one or two weeks from collection and a few days after experiencing symptoms ( skin desquamation , locomotor difficulties , dry skin , anorexia , hyperaemia of fingers ) . in captivity the infection was nearly always fatal for newly metamorphosed
froglets , but only sometimes for subadults and adults . in contrast , neometamorphs collected from tuscan - romagna areas during the same time period did not show infection or symptoms ( stagni et al . 2004 ) .
the species may be declining due to the loss of wetland habitat as a result of agricultural damage ( canestrelli et al . 2006 ) . it occurs in several protected areas in italy , including parco nazionale d ' abruzzo , parco nazionale pollino and parco nazionale del gran sasso e monti della laga ( iucn 2008 ) . parco nazionale del gran sasso e monti della laga national park began a long - term monitoring program for this species in 1999 ( ferri 2000 ; dimartino and ferri 2002 ) .
which were associated with man - made aquatic environments ( eg . troughs or washtubs ) underwent greater decline than those in natural habitats . maintenance and cleaning by agricultural workers had previously made these ideal for
reproductive success , by removing shading and vegetation conducive to tadpole predators such as dragonflies and newts . inland depopulation and agricultural abandonment since the 1960 ' s , however , has resulted in the overgrowth or complete loss of these cultivated habitats , resulting in decline of
man - made , regularly maintained water bodies in cultivated environments have become successful breeding sites for this species , as they provide a sustained , unshaded , vegetation free habitat for tadpole development ( stefano canessa , froglog 2012 ) .
by lanza and vanni ( 1991 ) due to genetic distance based on allozymes ( nascetti et al . 1982 ) and morphological differences ( vaccaneo 1931 ) . canestrelli et al . ( 2006 ) have argued for maintaining it as a separate species , based on analysis of mitochondrial and nuclear markers , and that calabrian populations deserve special conservation status as the reservoir of significant genetic diversity for this species however , phylogenetic analysis by hofman et al . ( 2007 ) , using mitochondrial cyt b sequences , found that italian populations were nested within
di martino , v . , and ferri , v . ( 2002 ) .
proceedings of third conference , save the amphibians , lugano , june 23 - 24 , 2000 , penne ( pescara ) .
nel parco nazionale del gran sasso e monti della laga . indicazioni per la conservazione .
ente parco nazionale gran sasso e monti della laga . relazione per l ' ente gestore , italia .
fromhage , l . , vences , m . , and veith , m . ( 2004 ) . ' ' testing alternative vicariance scenarios in western mediterranean discoglossid frogs . ' '
guarino , f . m . , angelini , f . , and cammarota , m . ( 1995 ) . ' ' a skeletochronological analysis of three syntopic amphibian species from southern italy . ' '
iucn ( 2008 ) . 2008 iucn red list of threatened species . www . iucnredlist . org . downloaded on 10 october 2008 .
iucn 2008 . 2008 iucn red list of threatened species . www . iucnredlist . org . downloaded on 27 november 2008 .
lanza , b . , and corti , c . ( 1993 ) . ' ' erpetofauna italiana : acquisizioni ed estinzioni nel corso del novecento . ' '
lanza , b . , and vanni , s . ( 1991 ) . ' ' notes on the biogeography of the mediterranean island amphibians . ' '
mastromei , g . , barberio , c . , pistolesi , s . , and delfino , g . ( 1991 ) . ' ' a bactericidal protein in
nascetti , g . , vanni , s . , bullini , l . , and lanza , b . ( 1982 ) . ' ' variabilità e divergenza genetica in popolazioni italiane del genere
simoncelli , f . , fagotti , a . , dall ' olio , r . , vagnetti , d . , pascolini , r . , and di rosa , i . ( 2005 ) . ' ' evidence of
bollettino dei musei di zoologia e di anatomia comparata della r . universitá di torino , series iii
dustin guericke , john cavagnaro ( dustingrey at gmail . com ) , black hills state university
> university of california , berkeley , ca , usa . accessed jul 9 , 2018 .
> university of california , berkeley , ca , usa . accessed 9 jul 2018 .
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frost , d . r . 2015 . amphibian species of the world : an online reference . version 6 . 0 . new york , usa . available at : urltoken .
a number of authors consider bombina pachypus to be a subspecies of bombina variegata ( bologna et al . 2000 ; ohler 1997 ; hofman et al . 2007 ) . the taxon was elevated to species level by lanza and vanni ( 1991 ) mainly through the use of preliminary electrophoretic data published in the form of a congress summary ( nascetti et al . 1982 ) and also on the basis of old morphological evidence ( vaccaneo 1931 ) .
justification : listed as endangered because of a population decline , estimated to be more than 50 % ( but presumably lower than 80 % ) inferred from a decline in the area of occupancy , and significant reduction in mature individuals , in the past 10 years , possibly because of susceptibility of this species to the chytridiomycosis pathogen .
this species is endemic to italy , where it occurs south of the po valley , through the appenine region , south to the southern tip of the italian mainland . it ranges from 20 m asl up to almost 1 , 700 m asl ( in latium ) .
it was formerly common in suitable habitat . however , the species has declined in almost all of its range ( with the exception of calabria , where populations remain stable ) over the last 10 years . a survey of representative sites across the species ' range showed that the species disappeared from > 50 % of surveyed sites between 1996 and 2004 ( 55 occupied sites in 1996 versus 23 occupied sites in 2004 ; barbieri et al . 2004 ) . significant recent declines in the populations of this species have been recorded from the province of siena ( piazzini et al . 2005 ) ; abruzzo ( ferri et al . 2007 ) ; ancona ( fiacchini 2003 ) ; lazio ( bologna et al . 2000 , bologna et al . 2007 ) and emilia - romagna ( stagni et al . 2005 ) .
the species occurs in shallow , unshaded pools in forests and open areas . spawning and larval development takes place in these pools . it also occurs in modified habitats , such as low - intensity farmland , pastureland , ditches , irrigations areas , drinking troughs , and ponds .
threats to this species are presumed to largely include loss and fragmentation of wetland habitat to drainage for intensive agricultural purposes . however many populations appear to have declined in areas of presumably intact habitat . in some places it may have very small populations ( 10 - 12 individuals ; mattoccia et al . 2005 ) ; these small populations are highly subject to stochastic extinctions . this species might also be threatened with chytridiomycosis , which has been associated with the deaths of several captive animals ( stagni et al . 2002 , stagni et al . 2005 ) , and further research into the possible impacts of this disease on populations of this species is needed . the cause of recent serious population declines remains incompletely known , but it has been speculated that chytridiomycosis is responsible .
this species is listed on appendix ii of the bern convention , and on annexes ii and iv of the eu habitats directive , in both cases under bombina variegata . it is known to occur in many protected areas including several national parks . the cause of recent severe declines in this species requires urgent investigation and action .
reformatted names of assessor ( s ) , reviewer ( s ) , contributor ( s ) , facilitator ( s ) and / or compiler ( s ) .
( errata version published in 2016 ) . the iucn red list of threatened species 2009 : e . t54450a86629977 .
to make use of this information , please check the < terms of use > .
1 . forest - > 1 . 4 . forest - temperate suitability : suitable 4 . grassland - > 4 . 4 . grassland - temperate suitability : suitable 5 . wetlands ( inland ) - > 5 . 4 . wetlands ( inland ) - bogs , marshes , swamps , fens , peatlands suitability : suitable 5 . wetlands ( inland ) - > 5 . 7 . wetlands ( inland ) - permanent freshwater marshes / pools ( under 8ha ) suitability : suitable 5 . wetlands ( inland ) - > 5 . 8 . wetlands ( inland ) - seasonal / intermittent freshwater marshes / pools ( under 8ha ) suitability : suitable 14 . artificial / terrestrial - > 14 . 1 . artificial / terrestrial - arable land suitability : marginal 14 . artificial / terrestrial - > 14 . 2 . artificial / terrestrial - pastureland suitability : suitable 15 . artificial / aquatic & marine - > 15 . 2 . artificial / aquatic - ponds ( below 8ha ) suitability : suitable 15 . artificial / aquatic & marine - > 15 . 5 . artificial / aquatic - excavations ( open ) suitability : suitable 15 . artificial / aquatic & marine - > 15 . 7 . artificial / aquatic - irrigated land ( includes irrigation channels ) suitability : suitable 15 . artificial / aquatic & marine - > 15 . 8 . artificial / aquatic - seasonally flooded agricultural land suitability : marginal
2 . agriculture & aquaculture - > 2 . 1 . annual & perennial non - timber crops - > 2 . 1 . 3 . agro - industry farming
2 . agriculture & aquaculture - > 2 . 3 . livestock farming & ranching - > 2 . 3 . 3 . agro - industry grazing , ranching or farming
8 . invasive and other problematic species , genes & diseases - > 8 . 1 . invasive non - native / alien species / diseases - > 8 . 1 . 1 . unspecified species
8 . invasive and other problematic species , genes & diseases - > 8 . 1 . invasive non - native / alien species / diseases - > 8 . 1 . 2 . named species
8 . invasive and other problematic species , genes & diseases - > 8 . 2 . problematic native species / diseases - > 8 . 2 . 1 . unspecified species
1 . research - > 1 . 1 . taxonomy 1 . research - > 1 . 2 . population size , distribution & trends 1 . research - > 1 . 3 . life history & ecology 1 . research - > 1 . 5 . threats 1 . research - > 1 . 6 . actions 3 . monitoring - > 3 . 1 . population trends
barbieri , f . , bernini , f . , guarino , f . m . and venchi , a . 2004 . distribution and status of bombina variegata in italy . italian journal of zoology suppl . 1 : 83 - 90 .
bologna , m . a . , capula , m . and carpaneto , g . m . 2000 . anfibi e rettili del lazio . fratelli palombi editori , roma .
bologna , m . a . , salvi , d . and pitzalis , m . 2007 . atlante degli anfibi e rettili della provincia di roma . gangemi editore , roma .
bonato , l . , fracasso , g . , pollo , r . , richard , j . and semenzato , m . 2007 . atlante degli anfibi e dei rettili del veneto . associazione faunisti veneti , nuovadimensione .
caputo , v . , guarino , f . m . , trecroci , t . and tripepi , s . 1992 . amphibian species of campania and calabria : their distribution , ecology and conservation . in : ferri , v . ( ed . ) , atti i convegno italiano sulla salvaguardia degli anfibi . 19 : 109 - 118 .
dupré , e . and la posta , a . 2008 . la direttiva habitat e lo stato di conservazione dell ' erpetofauna in italia . in : corti , c . ( ed . ) , herpetologia sardiniae . , pp . 235 - 240 . societas herpetologica italica / edizioni belvedere , latina , “le scienze” .
emanueli , l . 1994 . ululone dal ventre giallo bombina variegata ( linneaeus 1758 ) . in : doria , g . and salvidio , s . ( eds ) , atlante degli anfibi e dei rettili della liguria . , pp . 50 - 51 . regione liguria , genova .
ferri , v . , di tizio , l . and pellegrini , m . r . 2007 . atlante degli anfibi d’abruzzo . ianieri - talea edizioni , pescara .
fiacchini , d . 2003 . atlante degli anfibi e dei rettili della provincia di ancona . assessorato all ' ambiente della provincia di ancona . nuove ricerche editore .
fiacchini , d . 2008 . primo contributo per una “lista rossa” dell ' erpetofauna marchigiana ( italia centrale ) : amphibia . in : corti , c . ( ed . ) , herpetologia sardiniae . , pp . 258 - 261 . societas herpetologica italica / edizioni belvedere , latina , “le scienze” .
fromhage , l . , vences , m . and veith , m . 2004 . testing alternative vicariance scenarios in western mediterranean discoglossid frogs . molecular phylogenetics and evolution 31 ( 1 ) : 308 - 322 .
guarino , f . m . , bellini , l . , mazzarella , g . and angelini , f . 1998 . reproductive activity of bombina pachypus from southern italy . the italian journal of zoology 65 ( 4 ) : 335 - 342 .
iucn . 2009 . iucn red list of threatened species ( ver . 2009 . 1 ) . available at : urltoken . ( accessed : 22 june 2009 ) .
iucn . 2016 . the iucn red list of threatened species . version 2016 - 1 . available at : urltoken . ( accessed : 30 june 2016 ) .
lanza , b . , andreone , f . , bologna , m . a . , corti , c . and razzetti , e . 2007 . fauna d ' italia amphibia . vol . xlii . edizioni calderini de il sole 24 ore editoria specializzta s . r . l . , bologna .
lanza , b . and vanni , s . 1991 . notes on the biogeography of the mediterranean islands amphibians . atti conv . lincei , roma . 85 : 335 - 344 .
mattoccia , m . , cari , b . , romano , a . and angelini , c . 2006 . osservazioni sull’ecologia di alcune popolazioni di bombina pachypus ( amphibia : bombinatoridae ) nei monti lepini ( lazio ) . proceedings v congresso nazionale della societas herpetologica italica : 161 - 166 .
nascetti , g . , vanni , s . , bullini , l . and lanza , b . 1983 . variabilità e divergenza genetica in popolazioni italiane del genere bombina ( amphibia , discoglossidae ) . boll zool . 49 ( suppl . ) : 134 - 135 .
ohler , a . 1997 . supplement to the list of european amphibians . in : gasc , j . - p . et al . ( ed . ) , atlas of amphibians and reptiles in europe , pp . 404 - 40 . societas europea herpetologica & muséum national d’histoire naturelle ( iegp / spn ) .
piazzini , s . , favilli , l . and manganelli , g . 2005 . atlante degli anfibi della provincia di siena ( 1999 - 2004 ) . quaderni naturalistici 1 .
scillitani , g . , rizzi , v . and gioiosa , m . 1996 . atlante degli anfibi e dei rettili della provincia di foggia . gitto , foggia .
sindaco , r . , doria , g . , razzetti , e . and bernini , f . 2006 . atlas of italian amphibians and reptiles \ \ atlante degli anfibi e dei rettili d ' italia . societas herpetologica italica - edizioni polistampa , firenze .
stagni , g . , scoccianti , c . and fusini , r . 2002 . segnalazione di chytridiomicosi in popolazioni di bombina pachypus ( anura , bombinatoridae ) dell ' appennino tosco - emiliano . iv congresso della societas herpetologica italica , a ercolano .
vaccaneo , r . 1931 . ricerche sui caratteri morfologici dei bombinator italiani . bolletino dei musei di zoologia ed anatomia comparata della università di torino 41 ( 5 ) : 1 - 47 .
vanni , s . and nistri , a . 2006 . atlante degli anfibi e dei rettili della toscana . regione toscana & museo di storia naturale dell’università degli studi di firenze , sezione di zoologia “la specola” .
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kda ) , is the first amphibian member of the bv8 - prokineticin family . homologs of bv8 are present in the skin secretions of other amphibians such as
. the striking characteristics of these proteins are their identical n - terminal sequence , avitg , and the presence of 10 cysteines with identical spacing , which define a five - disulfide - bridged motif called a colipase fold .
the high degree of identity between amphibian bv8 peptides , fish peptides , and mamba mit - 1 ( 58 % ) suggested that similar peptides could also be present in other species , including mammals . in the mouse , rat , cattle , monkey , and human , cdna cloning identified orthologs of bv8 (
) . the two mammalian proteins similar to bv8 were named prokineticin 1 ( pk1 or eg - vegf ) and prokineticin 2 ( pk2 or mbv8 ) . a second form of pk2 has been identified and named pk2b ( because of an insert of 21 basic amino acids in its sequence ) . the name prokineticin refers to the ability of these peptides to contract guinea pig ileum , a property shared with amphibian bv8 .
expression patterns of rodent and human mrnas for pk1 and pk2 ( prok1 and prok2 ) have been reported in peripheral tissues ( dorsal root ganglia ( drg ) , gastrointestinal tract , endocrine glands , spleen , and human and murine leukocytes ) and the cns .
prokineticins are also expressed in the neurons of the medial preoptic area , nucleus of solitary tract , trigeminal and facial nuclei , and drg .
the two g - protein - coupled receptors for bv8 - pks , prokineticin receptor 1 ( pkr1 ) and prokineticin receptor 2 ( pkr2 ) have an overall identity in their amino acid sequences of 85 % , with most differences at the n - terminal , and are approximately 80 % identical to the previously described mouse orphan receptor gpr73 . in specific endothelial cells , neurons , and transfected cells expressing these receptors , bv8 - induced pkr activation stimulates ca 2 + mobilization and phosphoinositol turnover , indicating a g q / o - protein coupling . receptor binding studies showed that pkr2 is a mit - preferring receptor ; indeed , the affinity of mit for pkr2 ( in the picomolar range ) is approximately 10 times higher than that of pk2 and 50 times higher than that of pk1 . pkr1 is a mit - and pk2 - preferring receptor ; the affinity of mit for pkr1 ( ∼5–10 times lower than for pkr2 ) is comparable to that of pk2 and 60 times higher than that of pk1 . the affinity of bv8 for the receptors is comparable to that of pk2 and is approximately 40 times higher than that of pk1 .
human kallmann syndrome ( ks ) , which combines anosmia , related to defective olfactory bulb morphogenesis , with hypogonadism due to gonadotropin - releasing hormone deficiency , appears to be related to mutations in the genes coding for pk2 and pkr2 . in a cohort of 192 patients affected by ks , 10 different point mutations were identified in the genes encoding pkr2 and four were identified in the genes encoding pk2 . the mutations in pk2 were detected in the heterozygous state , whereas pkr2 mutations were found in the heterozygous , homozygous , or compound heterozygous state . in addition , one of the patients heterozygous for a pkr2 mutation also carried a missense mutation in kal1 , thus indicating a possible digenic inheritance of the disease in this individual . these findings reveal that insufficient prokineticin - signaling through pkr2 leads to the abnormal development of the olfactory system and reproductive axis also in humans .
pkr1 and pkr2 mrnas are expressed in the drg of neonatal and adult rats . pkr1 is mainly expressed in small and medium - size neurons and pkr2 in large neurons . pkr proteins are present in the drg , in the outer layers of the dorsal horns of the spinal cord , and in the peripheral terminals of nociceptor axons . activation of nociceptor pkrs by bv8 in rats and mice produces nociceptive sensitization to thermal and mechanical stimuli , without inducing any spontaneous , overt nocifensive behavior or local inflammation . a physiological role of bv8 - prokineticins as peripheral and central pain modulators is supported by the observation that mice lacking the pkrs or pk2 are less sensitive to noxious heat ( hot - plate test ) than are wild - type mice . pkr1 - null mice also exhibited impaired development of hyperalgesia after tissue injury . the inflammatory agents cfa and mustard oil produced comparable paw edema in wild - type and pkr1 - null mice , but induced inflammatory nociceptive sensitization to heat and pressure significantly lower in pkr1 - knockout than in wild - type mice , demonstrating a role of prokineticins and their receptors in inflammatory pain .
in situ mrna hybridization experiments demonstrated that pk2 is expressed by inflammatory cells , predominantly neutrophils , in the human inflamed tonsil and appendix and in the rat inflamed paw . pk2 released by inflammatory cells can bind and activate pkrs on the primary sensitive neurons contributing to inflammatory pain . neutrophil extracts , fractionated using ionic exchange chromatography , gel filtration , and reverse phase ( rp ) chromatography , displayed bv8 - like activity , displaced 125 i - mit binding from pkr1 - transfected chinese hamster ovary ( cho ) cell membranes and produced the bv8 - characteristic hyperalgesia when injected intrathecally in rats .
prokineticin receptors are potential targets for drugs which block the nociceptive information before it reaches the brain . identifying of the structural determinants required for receptor binding and hyperalgesic activity of bv8 - prokineticins is thus mandatory for the design of pkr antagonists . the highly conserved n - terminal sequence avitga and the triptophan residue in position 24 in all members of the bv8 / pk family are required for biological activity ; deletions and substitutions in these conserved residues produce antagonist molecules . preclinical studies are currently being carried out on these mutated pk peptides as pkr antagonists endowed with analgesic properties .
kda ) . the amino acid sequence was established by automated edman degradation and analysis of proteolytic fragments as well as by cdna cloning . homologs of bv8 have been predicted ( but not yet isolated ) in skin secretions of
( bm8a ) . these proteins are derived from simple precursors composed of a putative 19 - residue signal peptide and the mature protein . bo8 and bm8a show 96 % and 92 % sequence identity with bv8 . moreover , the putative signal peptides in all three proproteins are 100 % identical (
) . the primary sequence of bv8 is similar to mit ( mamba intestinal toxin ) , a constituent of the venom of the snake black mamba . the amphibian and snake proteins share the same pattern of cysteine distribution , and their overall identity is 58 % . cdna cloning showed the existence of mouse ( mbv8 ) and human ( hbv8 ) homologs of amphibian bv8 .
) . a similar cys - motif is also present in mammalian colipase and in the carboxy - terminal region of members of the dickkopf family of extracellular signaling proteins . however , the frog protein does not stimulate the activity of pancreatic lipase , and it is also inactive in an assay for dickkopf functions . starting from the 3d - structure of mammalian colipase , a model was built for frog bv8 in which the hydrophobic amino terminal sequence avitg forms a sort “beak” exposed at the surface of the tightly folded rest of the protein .
two forms of mbv8 and hbv8 have been characterized in the mouse and human testis . these forms differ in an exon coding for 21 amino acids , the majority of which are basic (
) . the predominant site of bv8 expression in both humans and mice is primary spermatocytes in the seminiferus tubules of the testis . the genomic structure of these murine and human bv8 genes has been determined , and the chromosomal localization was identified near a synteny breakpoint of mouse chromosome 6 and human 3p21 .
identified two human est sequences , one encoding the human protein already described and the other encoding a slightly different bv8 - like protein . the two proteins were named prokineticin 2 ( pk2 ) and prokineticin 1 ( pk1 ) (
) . the name prokineticin refers to the ability of these peptides to contract gpi , a property shared with amphibian bv8 .
identified a protein that induced proliferation , migration , and fenestration in the endothelial cells of steroid synthesizing glands ( ovary , testis , adrenals ) and named it endocrine - gland - derived vascular endothelial growth factor ( eg - vegf ) . eg - vegf and pk1 are the same protein and have an overall identity of 58 % and homology of 76 % with human pk2 and murine bv8 and 43 % identity with amphibian bv8 . pk1 / eg - vegf has been isolated and sequenced also from bovine milk . rat and human mrnas for pk1 and pk2 have been cloned and their expression patterns reported in peripheral tissues and the central nervous system .
the distribution of murine bv8 - like proteins , pk1 and pk2 , and their mrnas has been reported in the brain ( olfactory bulb , cerebral cortex , hippocampus , some thalamic and hypothalamic nuclei , purkinje cells of the cerebellum , many nuclei of the brain stem ) , spinal cord , gastrointestinal tract , endocrine glands , and other peripheral organs .
( being the lowest in the dark phase ) and is severely blunted in mutant mice deficient in clock or cryptochrome genes . pk2 / bv8 , constitutively expressed in cells and organs of myeloid origin , is dramatically upregulated by inflammatory processes .
receptors for pk1 and pk2 have been identified in humans , mice , and rats . these receptors , named prokineticin receptor 1 and 2 ( pkr1 and pkr2 ) , belong to the family of g protein - coupled receptors , share approximately 85 % amino acid identity , and are about 80 % identical to the previously described mouse orphan receptor gpr73 . in a study aimed to map the expression of pkr1 and pkr2 in rat and mouse nervous system , we
detected pkr2 mrna in the brain ( olfactory bulbs , scn , hippocampus , paraventricular thalamic and hypothalamic nuclei , nucleus arcuatus , subfornical organ ( sfo ) , amygdala , caudate putamen , and cortex ) and in dorsal root ganglia ( drg ) . pkr1 is mainly expressed outside the mammalian central nervous system , with the highest density in the neurons of drg . pkr1 is also expressed by murine macrophages . mice lacking the pkr2 show a dramatic reduction in olfactory bulbs ( ob ) size , and they do not survive ; mice lacking pkr1 are severely deficient in their responses to noxious thermal and chemical ( capsaicin , protons ) stimuli .
intensive investigation of the bv8 / prokineticin system over the past decade has revealed a dazzling array of physiological functions including the regulation of circadian rhythms , metabolism , angiogenesis , neurogenesis , muscle contractility , hematopoiesis , immune response , reproduction , and pain perception . in addition , the disruption of prokineticin system has been implicated in several pathological conditions , including cancer , immunological response , mood disorder ( anxiety / depression ) ,
cardiomyopathy , and persistent pain . point mutations in the genes encoding for prokineticins and / or prokineticin receptors are involved in human kallmann’s syndrome and hirschprung’s disease .
the isolation of an adequate amount of the amphibian bv8 protein has provided valuable research tools for characterizing the prokineticin pharmacology .
mm . in cell - based assays , bv8 and eg - vegf / pk1 induce proliferation , migration , and survival of primary adrenal cortical capillary endothelial ( ace ) cells . delivery of bv8 or eg - vegf to the mouse testis results in a potent angiogenic response resulting from activation of pkrs on vascular endothelial cells within the interstitial space .
pmol ) into the lateral ventricles of the rat brain suppress diurnal , nocturnal , deprivation - induced , and neuropeptide y - stimulated feeding . bv8 injected icv also stimulates drinking in a dose - dependent manner . we showed that bv8 stimulates drinking through its receptors in the sfo and inhibits feeding after injection into the arcuate nucleus . rat sensitivity to the bv8 - induced anorexogenic response increases significantly during the dark phase of the circadian cycle ,
in accordance with the physiological role of bv8 / pk2 in controlling circadian rhythm from the scn .
in rats , icv and sc injections of bv8 induce antidiuresis dependent on vasopressin release . bv8 injected sc also induces the release of oxytocin and corticosterone .
) , and intrabrain administration of amphibian bv8 induces hyperalgesia activating the prokineticin receptors that are present on peripheral nociceptors and in regions of the central nervous system associated with pain .
translocation , and sensitization of the vanilloid receptor trpv1 , hence decreasing the nociceptive threshold to mechanical and thermal stimuli .
amphibian bv8 injected into the periaqueductal gray ( pag ) exerts a pronociceptive action by increasing the intrinsic gabaergic tone which , in turn , is responsible for the inhibition of pag antinociceptive output neurons demonstrating that the bv8 / pkr system may also intervene in modulating central pain mechanisms .
we have shown that amphibian bv8 activates macrophages to migrate and to produce proinflammatory cytokines . it is also able to affect the t lymphocyte population , because the administration
of amphibian bv8 skews the th1 / th2 balance toward a th1 response . all these effects depend on the activation of pkr1 because they lack in the lymphocytes and macrophages from pkr1 - ko mice .
endogenous bv8 / pk2 is overexpressed in inflamed tissues and in animal and human neoplastic tissue , predominantly in infiltrating neutrophils . these cells can secrete it , since the sequence of the precursors contains the typical signal peptide for secretion .
ferrara’s group showed that bv8 regulates myeloid - cell - dependent tumor angiogenesis . we showed that bv8 has a crucial role in neutrophil - dependent inflammatory hypernociception .
to develop pkr antagonists as novel analgesic and antiinflammatory drugs , structural determinants required for the biological activity of bv8 must be identified .
studies from our group suggested that avitga family members could interact with pkr1 / pkr2 by orienting the protein region that comprises the avitga sequence and the conserved tryptophan residue in position 24 . the n - terminal deletion of the first two amino acids in amphibian bv8 molecule ( dav - bv8 ) yield an analog lacking any biological activity but still able to bind the receptors . dav - bv8 is a short acting antagonist
. substituting trp with ala in position 24 yields a molecule , a - 24 , that preferentially binds and activates pkr2 . it displays weak affinity and efficacy on pkr , acting as an antagonist . a - 24 has an extraordinary , long - lasting antihyperalgasic effect in animal models of postsurgical and inflammatory pain .
some nonpeptide molecules , triazine - guanidine derivatives , seem very promising . local or systemic injection of the lead compound , pc1 , abolishes the hyperalgesia in animal models of inflammatory
and postsurgical pain and speeds up the paw healing being , on a molar base , about a thousand times more potent than indomethacin ( unpublished ) . our results also show an important role for the bv8 / pkr in neuropathic pain at least in rodents . if this role is confirmed in humans , reducing bv8 levels in damaged tissues or antagonizing pkrs might be an innovative strategy to control persistent , invalidating pain .
) , well - distributed along the rhône river , has disappeared in the swiss rhône downstream of lake léman since 1980 , despite multiple re - introductions ( geneva herpetological society ) .
some amphibians are mainly found in the northern part of the rhône . smooth newt (
) are rare in the french upper rhône and lower rhône floodplains upstream of montélimar but are absent downstream . south great crested newts (
) are scarce because of the lack of reproduction areas . backwaters around arles have been identified as the last southern reproduction areas for this species . around geneva , south great crested newt populations have been declining since 1987 because of the non - native italian great crested newt (
) is found as far as montélimar . other amphibians are mainly present in the south part of the rhône river . the mediterranean tree frog (
is commonly found in camargue and in the lower rhône floodplain . until recently , the mediterranean tree frog and common tree frog co - occurred between lyon and valence but river regulation and the loss of wetlands led to their quasi - extinction . the western spadefoot is present from the delta to valence , and the parsley frog (
has disappeared in the mediterranean part of the rhône since the early 20th century . most amphibian species are protected .
, which is endangered also because of habitat reduction in many areas . among amphibians , the endemic sardinian brook newt (
) is present in the flumendosa basin , and the few populations are considered endangered .
there are four species of salamanders , the most widespread being the fire salamander ( salamandra salamandra ) , common in mountain ranges of continental and peninsular italy . the alpine salamander ( s . atra ) is still frequent in the upper ranges of the central - eastern alps , while the endemic lanza ' s salamander ( s . lanzai ) is restricted to a few locations in the headwaters of the po basin , in the western alps . lastly , the spectacled salamander ( salamandrina terdigitata ) is common along all the apennines , mostly between 300 and 900 m asl . the most common newts are : the alpine newt ( mesotriton alpestris ) , mostly in the western alps and northern apennines , the italian crested newt ( triturus carnifex ) widely distributed throughout italy , except the islands , the italian newt ( lissotriton italicus ) , endemic of the central and southern apennines , and the smooth newt ( l . vulgaris ) , common from the central apennines to the lower ranges of the alps .
) in the apennines . there also are some endemic frogs , including the tyrrhenian painted frog (
, which are common in other parts of northern italy , have reduced populations . the only priority species , according to the habitat directive , is the
) , still present along the whole adige valley but endangered because of a reduction in their specific habitats , represented by small , temporary pools in the river floodplains . reptiles associated with freshwater habitats include the common grass and dice snakes (
( stebbins & cohen 1995 ) . habitat loss and drought may have also contributed to the decline of these species . recent biomolecular research (
) . amphibian richness within a given habitat was significantly related to distance from islands , fish density and water temperature . vegetated islands and large woody debris played pivotal roles , directly and indirectly , in maintaining both habitat and amphibian diversity in this gravel - bed river .
) . two thirds of the amphibians and 1 / 3 of the reptiles prefer riverine landscape elements ; the remaining species occur in adjacent hillslope and upland areas . only three reptiles are truly aquatic :
despite the enormous interest in keeping exotic amphibians and reptilians as pets , which can result in abandoned or escaped animals , no introduced alien species have established reproducing populations in the basin until now . nevertheless , there are problems with the ongoing introduction of trachemys scripta elegans and other pond turtles , especially in germany and austria , because they most likely compete with the native turtle emys orbicularis .
) . many species are protected also by national laws ( e . g . ,
) . destruction of wetlands is the most serious threat to amphibian populations . even common species like
is locally threatened due to drainage , pollution , and destruction of breeding ponds and adjacent terrestrial habitats . in recent years ,
and called it ranatensin . in the same year , and shortly thereafter , erspamer
by 1978 , similar peptides had been isolated from other amphibians : the nonapeptide ( 9 amino acid peptide ) litorin , the undecapeptide ranatensin c , and the heptadecapeptide ( 17 amino acid peptide ) ranatensin r .
from porcine non - antral gastric tissue . the sequence of this peptide was identified by the same group in the next year ,
and because of the most prominent function of this peptide , which is stimulation of gastric acid secretion due to the release of gastrin , the peptide was given the name gastrin - releasing polypeptide . the next year , the same group identified grp in the chicken proventriculus as another 27 - amino acid peptide .
characterized three forms of grp ( 27 , 23 , and 10 ) from canine intestinal extracts .
) in the porcine spinal cord ; the next year , the same group identified grp - 10 in the porcine spinal cord , calling it nmc .
in 1984 , grp - 27 was isolated and characterized from human pulmonary carcinoid cells .
later on , similarity was found between the two classes of peptides , the amphibian peptides and grp , especially at the biologically active cooh - terminal . however , because the nh
the highest amphibian diversity is found in the floodplain areas and small waterbodies of the upper and middle rhine . in recent gravel ponds , rare species such as
in the northern middle rhine . in reed belts and willows of lake constance and floodplains of the upper rhine , tree frogs (
) can be heard during summer . early studies reported populations of the european water turtle
) . rare recent sightings of this species are due to the release of aquarium specimens . several non - native reptiles ( mostly the painted turtle
and other turtles , but even caimans and alligators ) are observed for the same reason ; however they rarely find suitable wintering conditions to maintian populations .
) . due to land - use changes and river regulation , many amphibian populations declined or gone extinct in floodplains along the rhine tributaries in the netherlands ( delta rhine ) . however , several common amphibian species still occur , such as
complex . the frequencies of occurrence and densities are still rather low in comparison with pristine areas .
hibernate in meadows , thickets and bushes on sand or clay in the higher as well as lower parts of floodplains .
clearly prefer sandy habitats in the higher parts of floodplains . the ringed snake snake is still frequently observed in floodplains along the nederrijn and ijssel (
the views of darwin , wallace , rensch , and mayr that geographic replacement forms , subspecies and semispecies , are in fact incipient species , has few critics today . most geographic replacement species ( i . e . , “semispecies , ” which intergrade only rarely when they meet ) must indeed have evolved from previously interbreeding subspecies . modern genetic data have done nothing to cast doubt on this idea . meanwhile , superspecies became a term reserved for groups of semispecies that intergraded little at their boundaries .
however , under the trinominal approach , taxonomists were now required to describe subspecies , which has never been seen as a particularly noble activity in comparison to the description of species , especially recently . a strong attack on the zoological subspecies was mounted by
. both were systematists working on ants , a group particularly riddled with poorly conceived trinominals and other named varieties at the time . wilson and brown argued that subspecies rarely , if ever , could be justified on the basis of multiple characters , and that therefore they were not “real taxa . ” the only “real taxa” were species , which in a sense were self - defining because interbreeding prevented divergent genes from flowing from one species to another . subspecies that interbred at their boundaries , on the other hand , were not so endowed , so that genes and morphological characters could flow between them . good examples of subspecies were put forward which undoubtedly would be hard to justify on multiple character grounds . this single paper was enormously influential on systematics in the united states , and generations of insect systematists trained at harvard and cornell , where wilson and brown worked , and their own many intellectual descendants , and their students’ students in turn , have eschewed the practice of naming subspecies .
across a broad front in europe . the two forms hybridize freely in the contact zone—although the hybrids can be shown to suffer some inviability—and so should really be classified as members of the same species under polytypic or biological species concepts . however , it has always seemed natural to place such well - defined forms in separate species in spite of the fact they have not truly “speciated . ” the two taxa differ strongly in call , morphology , skin thickness , the sizes of water bodies used for breeding , and egg size , as well as in mitochondrial deoxyribonucleic acid ( dna ) and protein sequence . the levels of differentiation suggest that the
taxa have evolved separately for many millions of years . this situation of multiple character changes has now been shown to be true across many examples of subspecies as well as species separated by hybrid zones . gene flow can be shown to be almost completely blocked by hybrid zones such as these , even if hybridization is frequent (


the wingspan is 17 – 20 mm .

the forewings are light purplish-grey .

the hindwings are deep purplish-brown in the apical third .

apsidophora

this is the place for apsidophora definition . you find here apsidophora meaning , synonyms of apsidophora and images for apsidophora copyright 2017 © urltoken
here you will find one or more explanations in english for the word apsidophora . also in the bottom left of the page several parts of wikipedia pages related to the word apsidophora and , of course , apsidophora synonyms and on the right images related to the word apsidophora .
apsidophora is a genus of moths belonging to the subfamily olethreutinae of the family tortricidae . it contains only one species , apsidophora purpurorbis , which is found in thailand , the malay peninsula , sumatra and new guinea .
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by t . m . gilligan 1 , j . baixeras 2 , j . w . brown 3 , and k . r . tuck 4
urltoken is pleased to offer the complete world catalogue of the tortricidae ( t @ rts ) ! this is a complete list of all world species , utilizing the world catalogue published in 2005 as the foundation for the database . version 3 . 0 of the online catalogue contains 15 , 099 records representing 10 , 883 species . more than 1 , 600 records have been updated from ver 2 . 0 ( jul , 2012 ) , and more than 3 , 000 records have been updated from the original catalogue . the database is completely searchable and contains photos of over 1 , 200 type specimens .
t @ rts will be updated regularly both with corrections from the original world catalogue and with additions since its publication . as such , these pages will serve as the most up to date information on current tortricid nomenclature . if you find any errors in the data presented here or have any questions / comments , please use the contact form to send the authors an email .
we are indebted to all of the original authors of the world catalogue ( j . w . brown , j . baixeras , r . brown , m . horak , f . komai , e . metzler , j . razowski , and k . tuck ) for providing the basis for this project . we would also like to thank the dozens of individuals who have provided corrections or updates to the database since it was first placed online in 2007 .
gilligan , t . m . , j . baixeras , j . w . brown & k . r . tuck . 2014 . t @ rts : online world catalogue of the tortricidae ( ver . 3 . 0 ) . urltoken
1 colorado state university , bioagricultural sciences and pest management , 1177 campus delivery , fort collins , co 80523 , usa 2 institut cavanilles de biodiversitat i biologia evolutiva , universitat de valencia , apartat oficial 2085 , 46071 valencia , spain 3 systematic entomology laboratory - usda [ retired ] , smithsonian institution , p . o . box 37012 , national museum of natural history , washington , dc 20013 , usa 4 curator - microlepidoptera [ retired ] , entomology department ( dc2 - 2n ) , natural history museum , cromwell road , london sw7 5bd , uk
unless noted , all images on these pages are copyright © 2003 - 14 by todd gilligan . please do not download , copy , print , or otherwise distribute any images from these pages without the permission of the author . contact form .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
the author ' s name is made up of a string of letters , with no blanks , and multiple authors ' names are separated by a comma . spelling of author ' s name is based on the original publication . if there are more than three authors , only the names of the first two authors are shown , followed by
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and the number of omitted authors .
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the wingspan is 17 - 20 mm . the forewings are light purplish - grey . the hindwings are deep purplish - brown in the apical third .
notable tortricids include the codling moth and the spruce budworm , which are among the most well - studied of all insects because of their economic impact . [ 3 ]
the head is where many sensing organs and the mouth parts are found . like the adult , the larva also has a toughened , or sclerotized head capsule . [ 22 ] here , two compound eyes , and chaetosema , raised spots or clusters of sensory bristles unique to lepidoptera , occur , though many taxa have lost one or both of these spots . the antennae have a wide variation in form among species and even between diffe
though the true dimensions of species diversity remain uncertain , estimates range from 2 . 6–7 . 8 million species with a mean of 5 . 5 million . [ 42 ]
the embryos of all arthropods are segmented , built from a series of repeated modules . the last common ancestor of living arthropods probably consisted of a series of undifferentiated segments , each with a pair of appendages that functioned as limbs . however , all known living and fossil arthropods have grouped segments into tagmata in which segments and their limbs are specialized in various ways . [
nearly all animals make use of some form of sexual reproduction . [ 20 ] they produce haploid gametes by meiosis ; the smaller , motile gametes are spermatozoa and the larger , non - motile gametes are ova . [ 21 ] these fuse to form zygotes , [ 22 ] which develop via mitosis into a hollow sphere , called a blastula . in sponges , blastula larvae swim to a new location , attach to the seabed , and develop into a new sp