woolly aphids ( subfamily : eriosomatinae ) are sucking insects that live on plant fluids and produce a filamentous waxy white covering which resembles cotton or wool .

the adults are winged and move to new locations where they lay egg masses .

the nymphs often form large cottony masses on twigs , for protection from predators .

they occur throughout the northern hemisphere .

many of the numerous species of woolly aphids have only one host plant species , or alternating generations on two specific hosts .

the woolly apple aphid , eriosoma lanigerum is a widespread pest of fruit trees , feeding principally on apple , but also , pears , hawthorn , ash , alders , elms and oaks .

in flight they have been described as looking like " flying mice " , and are given nicknames like " angel flies " , " fluff bugs " , " fairy flies " , and " ash bugs " " snow bugs " , " fluffer fairy " , " poodle fly " , " fluffy gnats " .

eriosomatinae

eriosomatinae <EOT> morphological phylogeny of gall - forming aphids of the tribe eriosomatinae ( aphididae : eriosomatinae ) .

another significant difference with other species of eriosomatinae is the shape and position of rectal bladder . in

les according to the 3 tribes of eriosomatinae and the out - groups . e genbank accession n

evolutionary relationships of pemphigus and allied genera ( hemiptera : aphididae : eriosomatinae ) and . . .

look at wing venation for further clues in distinguishing among these . below is a pemphigus ( eriosomatinae ) for comparison .

honeydew from root aphids ( aphidae : eriosomatinae ) is an important part of the diet of the subterranean ant lasius nearcticus . new york , usa

type genus : eriosoma leach , 1818 ; origin of priority for family - group names not recorded in database . first use as eriosomatinae by baker , a . c . , 1920 .

many woolly aphids are in subfamily eriosomatinae . however , there are several exceptions ( and according to andrew jensen , " there are examples of wax - covered aphids in almost all subfamilies . " ) :

. . . in addition , in all higher - level phylogenetic studies of aphididae that have been conducted , neither aphid data ( vondohlen & moran 2000 ; ortiz - rivas et al . 2004 ; ortizrivas & mart ınez - torres 2010 ) nor buchnera data ( nov akov a et al . 2013 ) supported the eriosomatinae as a monophyletic clade . its monophyly was also not corroborated in the morphological cladistic study ( zhang & chen 1999 ) and molecular phylogenetic analyses of eriosomatinae ( zhang & qiao 2008 ; li et al . 2014 ) . the subfamily eriosomatinae has been traditionally recognized as a monophylum based on the synapomorphy of sexuales apterous , dwarfish and lacking rostrum ( heie 1980 ; zhang et al . 1999a ) . . . .

. . . in addition , in all higher - level phylogenetic studies of aphididae that have been conducted , neither aphid data ( von dohlen & moran 2000 ; ortiz - rivas et al . 2004 ; ortizrivas & mart ınez - torres 2010 ) nor buchnera data ( nov akov a et al . 2013 ) supported the eriosomatinae as a monophyletic clade . its monophyly was also not corroborated in the morphological cladistic study ( zhang & chen 1999 ) and molecular phylogenetic analyses of eriosomatinae ( zhang & qiao 2008 ; li et al . 2014 ) . the subfamily eriosomatinae has been traditionally recognized as a monophylum based on the synapomorphy of sexuales apterous , dwarfish and lacking rostrum ( heie 1980 ; zhang et al . 1999a ) . . . .

) , except of eriosomatinae , which were not studied in this respect . the existing differences concern mainly the lengths of particular segments of the tract . due to the lack of filter chamber , the adhering of the loops of crenated intestine to the ventral wall of the stomach may play the role of osmoregulation , as hypothesized for the pea aphid ( rhodes et al .

eriosomatinae , the gall - forming aphid subfamily , traditionally consists of 3 tribes , eriosomatini , pemphigini , and fordini . however , the phylogenetic relationships among these tribes remain controversial , which has made it difficult to conduct further investigation regarding the evolution of galls and host alternations in this group . we analyzed the molecular phylogeny of the subfamily eriosomatinae , combining sequences from 2 mitochondrial genes ( coi and coii ) and 2 nuclear genes ( ef - 1 alpha and lwo ) . the reconstructions were implemented based on single - gene and multigene datasets through 3 different reconstructing algorithms , respectively ; analyses with 5 different out - groups were also conducted . results revealed a large paraphyletic clade , in which there were 4 out - groups clustering between eriosomatini and the other 2 tribes . however , the monophyly of the 3 tribes was well supported by the obtained trees , respectively .

. . . but the relationships among genera were not well supported with the position of the north american melaphis poorly resolved . using the mitochondrial coi and coii and the nuclear ef - 1a and lwo , li et al . ( 2014 ) constructed the relationships of subfamily eriosomatinae , sampling six genera of the melaphidina aphids . their results supported the monophyly of melaphidina , but the relationships among the melaphidina genera were only well supported by the bayesian posterior probabilities ( pp ) without strong bootstrap support ( bs ) . . . .

as a conclusion , it must be stated that the alimentary tract of apterous viviparous female of anholocyclic population of g . utricualria consists of all anatomical organs typical of aphid subfamily eriosomatinae and constitutes a continuous tube . it is unlikely that g . setulosa may have differently built alimentary tract , since existing data on its foregut and hindgut fully agree with that of presented study on g . utricularia . however , taking into account complicated life cycle and heteroecy of geoica spp , . there is essential need for further studies to confirm whether the situation of geoica spp . ( and also various morphs of species in this genus ) is similar to that of g . utricularia or to that described by ponsen .

the analysis of b . aphidicola genes that follow an evolutionary pattern that agrees with the molecular clock hypothesis [ 12 , 13 ] can be used to estimate the divergence time between pairs of aphids . this is possible because two aphid species , acyrthosiphon pisum and schizaphis graminum belonging to two tribes of the subfamily aphidinae , have an estimated divergence time calibrated from their fossil record of 50 to 70 my [ 14 ] . in addition , using molecular data from complete b . aphidicola genomes available , pérez - brocal and coworkers calculated the divergence time of aphids belonging to subfamilies eriosomatinae ( baizongia pistaciae ) and lachninae ( cinara cedri ) [ 15 ] . based on morphological traits , the subfamilies eriosomatinae and lachninae have traditionally been considered very divergent . in fact , most phylogenetic hypotheses based both on morphological and molecular data consider the lachninae as a sister group of the aphidinae [ 11 , 16 ] . however , the position of this subfamily remains controversial , as recent phylogenies based on molecular sequences located the subfamily in a basal position [ 17 – 19 ] . here , we follow a genomic approach to deepen the evolutionary analyses and propose a phylogeny of the three subfamilies of aphids based on the genome sequence of their primary endosymbionts b . aphidicola . in addition , in order to detect if there is any selective effect related to the specific role of the genes , we also gave a closer look to the acceleration pattern of each functional category .

this species is widely distributed and are quite often encountered in yards . yet details of its life history are , as far as my literature searches can tell me , mostly undescribed : when does the switch to sexual reproduction happen ? how does the foundress feed or is she fed by her kids ? how and where do they overwinter ? how many asexual generations are completed within a year ? what kinds of predators and parasites plague the colonies ? some the species’ close kin ( other wooly and gall - forming insects in the subfamily eriosomatinae of the aphididae ) have ant - like life histories , forming solider castes within the colony . grylloprociphilus seems less specialized , but without detailed life history studies , we can’t say more .

since the establishment of the symbiosis between the ancestor of modern aphids and their primary endosymbiont , buchnera aphidicola , insects and bacteria have coevolved . due to this parallel evolution , the analysis of bacterial genomic features constitutes a useful tool to understand their evolutionary history . here we report , based on data from b . aphidicola , the molecular evolutionary analysis , the phylogenetic relationships among lineages and a comparison of sequence evolutionary rates of symbionts of four aphid species from three subfamilies . our results support previous hypotheses of divergence of b . aphidicola and their host lineages during the early cretaceous and indicate a closer relationship between subfamilies eriosomatinae and lachninae than with the aphidinae . they also reveal a general evolutionary pattern among strains at the functional level . we also point out the effect of lifecycle and generation time as a possible explanation for the accelerated rate in b . aphidicola from the lachninae .

existing literature data report the lack of stomach and crenated intestine in the aphid species geoica setulosa ( passerini , 1860 ) , a representative of subfamily eriosomatinae . this odd anatomical feature seemed remarkable , due to the presence of fully developed intestine in closely related genera and mutualistic relationship with ants of this genus . the study aimed at repeated anatomical research of geoica utricularia ( passerini 1856 ) , in order to confirm what seemed to be a generic feature . standard histological methods were applied , with addition of oblique light microscopy , fluorescence microscopy and confocal laser scanning microscopy . the results indicated the existence of a fully developed intestine , with broad sac - shaped stomach and loops of the crenated intestine . the general anatomy of the alimentary tract of g . utricularia resembles that of other representatives of the tribe fordini . also well - developed rectal gland is present , most probably playing a role in modifying the carbohydrate composition of excreted honeydew .

our phylogenetic analysis supports the presence of one clade clustering b . aphidicola bbp and bcc , and another clade consisting of b . aphidicola bap and bsg . this result is consistent with a panorama of a rapid evolutionary radiation of the main subfamilies of aphids , during the early cretaceous ( 144 - 100 mya ) , which seems concordant with previous proposals [ 3 ] . in addition , our evolutionary molecular data from b . aphidicola point out that aphids belonging to subfamilies eriosomatinae and lachninae share a common ancestor more closely related than compared to the members of subfamily aphidinae . if true , our data refute the traditional phylogenetic reconstructions that placed aphidinae and lachninae as a monophyletic group [ 11 ] . however , we do not have evidence to conclude whether , within the subfamily lachninae , tribes feeding on conifers are ancestral or more recent than those living on herbaceous angiosperms , since our analysis does not resolve which strain ( and thus which host aphid ) is basal compared to the others . to solve this point , it would be necessary to sequence the genome of a greater number of b . aphidicola strains , including members of the different tribes from the subfamily lachninae ( work in progress ) . this would allow us to establish the date of divergence between those tribes and , thus , try to relate this fact to the change of vegetal host in either direction .

regarding synonymous substitutions , when pairs of strains of b . aphidicola were compared based on the average number of synonymous substitutions per site ( ) , a greater accumulation was observed in the b . aphidicola bbp strain compared to bacteria from aphids of the subfamily aphidinae ( b . aphidicola bap and bsg ) , while the smallest value is found between the b . aphidicola bap and bsg strains [ 15 ] . however , if the temporary factor is considered , the rates of synonymous nucleotide substitutions per site and million years are greater in the endosymbionts from the aphidinae ( b . aphidicola bap and bsg strains ) , registering the b . aphidicola bcc strain the smallest values . these results demonstrate that the synonymous substitution rate in b . aphidicola is a variable character , yet the explanation for these divergent patterns is not obvious . as stated elsewhere [ 7 , 14 , 44 ] , these differences can be attributed to differences in the host’s life cycle , as well as ecological factors such as host - alternation and variations in the effective population size showed by the two members of the aphidinae subfamily compared to the other two aphid lineages . additionally a differential mutation rate per generation cannot be ruled out . for example , endosymbionts from aphids with short generation times can accumulate more synonymous mutations per million years ( case of the aphidinae ) than those with longer generation times , such as the eriosomatinae and the lachninae . future studies are required to understand the evolutionary processes driving these patterns .

in our efforts to understand the biology of soldier - producing aphids , we attempted to maintain them in the laboratory using a chemically defined artificial diet . the ability of 16 species from the subfamilies eriosomatinae and hormaphidinae , most of which are soldier - producing species , to survive on the artificial diet was examined . some species neither fed nor grew on the diet , whereas other species accepted the diet , grew to some extent , and managed to produce a small number of short - lived offspring . although they performed poorly on the diet in general , aphid performance was correlated with the stage in the life cycle and the developmental stadium in that aphids of the gall generation tended to accept the diet and survive on it , whereas aphids of the non - gall generation did not . also , old insects tended to perform better on the diet than young nymphs . notably , only one species , tuberaphis styraci , a gall - forming aphid that produces 2nd instar sterile soldier , showed good performance on the diet . insects collected from galls ( generation g0 ) survived on the diet , grew well , and produced many progeny . three successive generations ( g1 , g2 and g3 ) were produced on the diet . developmental period , adult body size , and age of first reproduction were almost constant through g0 , g1 and g2 whereas fecundity , adult longevity and daily offspring production declined as the generations proceeded . these results are comparable to previous studies in which pest aphids have been maintained on similar artificial diets for several generations . therefore , it is suggested that the artificial - diet rearing system will provide a useful tool to investigate various biological aspects of the soldier - producing eusocial aphid , t . styraci .

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nomenclatural details : original name . with precedence over myzoxilinae amyot & audient - serville from which it takes the date of priority , 1843 ( iczn article 40 . 2 )

börner . 1944 . in brohmer . fauna von deutschland : ein bestimmungsbuch unserer heimischen tierwelt . fifth edition 218

eastop & van emden . 1972 . in van emden [ ed . ] . aphid technology 13 > > note : = pemphiginae

nieto nafría , mier durante & g . remaudière . 1997 . revue française d’entomologie nouvelle série 19 ( 3 - 4 ) : 82 , 88 > > note : kirkaldy 1905 : 418 ( 1843 )

zhang , guangxue , qiao & xiaolin chen . 1999 . in zhang , guangxue [ ed . ] . fauna of agricultural and forestry aphids of northwest , china : insecta homoptera aphidinea 121

nieto nafría , mier durante & pérez hidalgo . 2011 . in nieto nafría & favret [ ed . ] . registers of family - group and genus - group taxa of aphidoidea ( hemiptera sternorrhyncha ) 56 > > note : kirkaldy 1905 ( 1843 )

li , xingyi , jiang & qiao . 2014 . turkish journal of zoology 38 : 285 - 297

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formerly a separate family , eriosomatidae . tree of life places many ( all ? ) of the included genera in family pemphigidae .

many , but by no means all , root aphids belong to this subfamily . as of 4 / 12 / 10 , the only non - eriosomatine root aphids on bugguide are these aphis ( aphidinae ) :

subfamily hormaphidinae includes two species that cause galls on witch hazel . the manzanita leaf gall aphid is in subfamily tamaliinae .

( the following information is based on bugguide ' s images and may be modified based on future observations . )

the asian woolly hackberry aphid ( calaphidinae : shivaphis celti ) has red eyes , banded antennae , and distinctively patterned wings .

the woolly beech aphid ( phyllaphidinae : phyllaphis fagi ) forms woolly masses on the undersides of beech leaves ( primarily european varieties ) . we do not have adult images for this species yet .

the balsam twig aphid ( mindarinae : mindarus abietinus ) produces white fluff on fir twigs . adults have only a thin , light coating of this material .

alleyne , e . h . and f . o . morrison . 1977 . some canadian poplar aphid galls . the canadian entomologist 109 ( 3 ) : 321 - 328 . [ should help sort out some of the unidentified poplar galls ]

borror and delong ' s introduction to the study of insects norman f . johnson , charles a . triplehorn . 2004 . brooks cole .

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in all aphid species , the ectodermal part of the anterior region of the alimentary tract consists of the stylet bundle , pharynx , foregut and oesophageal valve . the pharynx consists of the pharyngeal duct , valve and pump ( ponsen

) . the midgut is the endodermal part of the alimentary tract consisting of the stomach , crenated intestine and descending intestine . the midgut is responsible for production and excretion of enzymes and the absorption of nutritional substances . in this part of the alimentary tract , often the so - called filter chamber develops ( ponsen

) . it is an adaptation to feeding on the phloem sap , and it takes part in regulation of the osmotic pressure due to the high concentration of sugars in the sap ( rhodes et al .

) . in various groups of aphids , where the filter chamber is present , it is developed variously , e . g . in the subfamily lachninae , the proximal part of the midgut is coiled inside the chamber , creating loops ( klimaszewski and wojciechowski

) . despite the high diversity of the structure of the filter chamber , it is known that foregut and hindgut are connected , so showing the continuity of the tract . in species without the filter chamber , the coiled segments of hindgut adhere the stomach , e . g .

) . the wall of the descending intestine is internally made up of epithelium , whose cells are adapted to resorption of water , amino - acids and minerals and ends with rectum .

) . there is no filter chamber . the very short ectodermal part of the posterior region of the alimentary tract consists of a rectum , epidermal invagination and an anal opening .

and secondary host being the roots of various grasses . their life cycle is two years , and on their primary host , they live in galls ( heie

) . in central europe , the anholocyclic population occurs , with parthenogenetic way of reproduction throughout the year . there are , however , data concerning

consists of a foregut , which is closed at its posterior end , and a blindly starting descending intestine ( fig .

) . it seemed to be odd , taking into account the necessity for constant resorption of vast quantity of protein - poor food ( cristofoletti et al .

) . these species have even the anal plate developed into a trophobiotic organ ( modified anal plate ) , enabling proper positioning of the honeydew droplet ( figs .

alimentary tract of g . utricularia : a extracted from the body ; b schematic reconstruction ; c location within the body ; f foregut , ov oesophageal valve , st stomach , al abdominal loop , di descending intestine

alimentary tract of g . utricularia : a , b longitudinal section through head and thorax ; c transverse section through salivary glands ; d location of stomach within the body ; f foregut , sg salivary glands , ov oesophageal valve , st stomach , di descending intestine

alimentary tract of g . utricularia : a cleared specimen in oblique light microscopy ; b view in clsm , rhodamine - b stained ; c , d cleared specimens in fluorescent microscopy , rhodamine - b stained ; sg salivary glands , ov oesophageal valve , st stomach , al abdominal loop

alimentary tract of g . utricularia : a frontal section through the body ; b longitudinal section through thorax and abdomen ; c , d longitudinal section through posterior part of the abdomen ; sg salivary gland , st stomach , al abdominal loop , di descending intestine , hg hindgut , rb rectal bladder , rg rectal gland , ei epidermal invagination , r rectum , ap anal plate

) , to verify data on the reported lack of the stomach and crenated intestine .

was used because it is more common and abundant , and both species differ only slightly in their morphology and biology . in

, where there are two rows of long setae . the purpose of these setae ( in both species ) is to keep the droplet of honeydew until it is collected by worker ant . their host plants , life mode and relations with ants are in their anholocyclic range very similar and significantly overlap .

the material was collected in october and november 2014 on moist meadows in the vicinity of piekary śląskie , in southern poland . in total , 32 individuals , apterous viviparous females of anholocyclic population on secondary host of

in order to analyse the anatomical structure of the digestive system of g . utricularia , the paraffin method was applied . the material was collected into eppendorf microtubes containing 70 % ethanol for keeping and preserving collected specimens . next , insects were dehydrated in increasing concentration of ethanol ( 90 , 96 , 100 % ) . in order to ensure transparency , specimens were kept in methyl benzoate for one night . then , material was consecutively transferred to benzene , benzene with paraffin ( in proportions 2 : 3 and 1 : 2 ) , paraffin i ( melting point : 56 °c ) and finally to paraffin ii ( melting point : 60 °c ) , where it was kept over night . after this process , material was immersed in paraffin ii . the bars obtained were sectioned into 5 µm strips , which were stuck on slides in a 0 . 5 % gelatine solution at temperature 50–52 °c . then , the slides were dried in 37 °c .

slides were next deparaffined in xylene and treated with a series of ethanol solutions ( 100–60 % ) . they were rinsed in distilled water , stained with ehrlich’s acid hematoxylin for about 20 min , rinsed again and differentiated with xylidine ponceau . after this process , preparations were treated with series of ethanol solutions ( 60–100 % ) , rinsed twice in xylene and embedded in canadian balm or dpx .

this process was applied to 15 individuals of g . urticularia . histological preparations were prepared : cross - sections from six individuals and longitudinal sections from nine individuals . in total , 49 microscopic slides were made , including 26 preparations of cross - section and 23 of longitudinal section . the cross - section series were also used to reconstruct the course of alimentary duct and in some cases the number and shape of cells .

the specimens of g . utricularia were put into a droplet of 30 % ethanol , on the microscopic glass , and with the mounting needles , the whole alimentary tract was extracted from the body . the extracted organs were preserved in glycerol and mounted , to make all the anatomical structures visible at the stereomicroscope . a total number of 13 preparations were made using this technique .

the documentation was prepared using nikon eclipse e6000 , with measurements made by lucia net program . the pictures of translucent specimens were taken under stereomicroscope equipped with monochromatic camera axio cam programmed with axio vision .

in order to investigate localization and structure of gastric tract in g . utricularia , specimens were examined with oblique light microscopy , fluorescence microscopy and confocal laser scanning microscopy ( clsm ) . before the examination , collected specimens were fixated overnight in cold 4 % pfa ( paraformaldehyde ) and subsequently underwent dehydration in graduated ethyl alcohol series , then optical clearing in methyl salicylate .

for fluorescent and clsm imaging , specimens were prior to dehydration treated with 30 % hydrogen peroxide for 24 h and thereafter stained with rhodamine - b before clearing in methyl salicylate . the staining with rhodamine - b was applied for visualizing elastic tissues in clsm ( shelley

) . all reagents used in procedures had analytical grade purity . fluorescent and oblique light imaging was performed with carl zeiss stereo lumar . v12 stereomicroscope and clsm with olympus fv1000 confocal system . acquired images were processed in fiji open source image processing package ( smolla et al .

begins with the external mouthparts and runs through the body cavity without any break or separately located organs . it ends at the anal orifice located above the dorsal edge of the anal plate ( figs .

a ) is a thin straight tube , about 27 μm long , with the diameter of 0 . 01–0 . 02 μm . in the prothorax , it runs between the two main salivary glands ( fig .

c ) , which are built of ca . 20 cells . they are in shape of fan , bottle or other and comprise one or two nuclei located in central or in basal position in the cell , which is typical of exocrine glands ( fig .

a–c ) . their diameter is 0 . 08–0 . 12 μm . the wall of the foregut is made up of one layer of small flat epithelial cells , without any folds into its lumen . in the thorax , the foregut joins the stomach , by invagination into the cavity of stomach ( figs .

the midgut is the longest part of the alimentary tract and consists of the stomach , crenated and descending intestine .

the stomach begins in the second segment of thorax ( mesothorax ) ( figs .

d ) and extends towards the abdomen , slightly dorsally within the body . in longitudinal section , it has a shape of ventro - dorsally flattened bag ( figs .

b ) , constricting gradually along its length . in its widest part , it has a diameter of 0 . 24–0 . 30 μm , and its length varies from 0 . 43 to 0 . 49 μm . the stomach is made up of two types of cells . the anterior part of stomach is built up of elongated , cone - shaped or finger - shaped cells , adhering to each other with lateral membranes at the cell bases ( figs .

a , b ) . their free tips , directed towards the cavity of stomach , broaden the internal surface of the stomach . the cytoplasm of these cells comprises visible granules , which altogether indicates their secretive function .

b ) are flat and hexagonal in shape and possess oval nuclei . the stomach narrows and opens into the crenated intestine , which runs distally ; then , it loops towards the head ( figs .

a ) , adhering the ventral surface of the stomach , and again loops towards the end of abdomen and opens into the descending intestine .

the descending intestine is a straight tube with thin walls , running through abdomen , and its wall is made up of flattened epithelial cells throughout its length ( b , c ) . at its distal end , the descending intestine is broadened , and a well - developed rectal bladder is formed ( fig .

c , d ) . it is positioned dorsally and has a roundish bubble shape , with a few secretive cells in its cavity . these cells are fan - shaped and secrete into the bladder ( fig .

d ) . the rectal bladder opens into the epidermal invagination . it is a straight tube opening with the anal orifice .

( von heyden 1837 ) , another representative of erisomatinae . the presence of doubled nuclei in some of the cells in salivary glands is not surprising , since it was proved to occur also in other aphid families ( ponsen

) , the rectal bladder consists of a ring of cells of polygonal shape , surrounding the hindgut near the rectum . it concerns only females of

spp . , because the presence of rectal bladder was not confirmed in males of this genus .

the role of rectal bladder is not sufficiently recognized . it is probably homologous with malphigian tubules ( ponsen

) , which have disappeared during the evolution of aphids . its possible role is that it takes part in regulation of the honeydew composition ( dixon

) , which contains carbohydrates , especially disaccharides : raffinose and melecitose , synthesized in aphid intestine , and their presence may play role in mutualistic relation of aphids and ants . in particular , the most attractive of ants is the honeydew rich in melecitose ( fisher and shingleton 2001 ) . aphids producing honeydew with 30–70 % of melecitose ( e . g .

stroyan 1950 ) are more often ant attended than those not having melecitose in their honeydew ( e . g .

( koch 1854 ) , with lower amount of melecitose in its honeydew . taking into account the high level of myrmecophily of both

spp . , the presence of well - developed rectal bladder may serve the purpose of regulating the composition of honeydew .

we wish to thank m . b . ponsen ( agricultural university wageningen ) for his kind response and all valuable suggestions and corrections made on the first version of the manuscript . we also thank two anonymous referees for their valuable comments and suggestions on the manuscript .

all applicable international , national and / or institutional guidelines for the care and use of animals were followed .

blackman rl , eastop vf ( 2006 ) aphids on world’s herbaceous plants and shrubs . wiley , chichester , p 1439

depa ł , wojciechowki w ( 2008 ) ant - root aphid relations in different plant associations . pol j entomol 77 : 151–163

dixon afg ( 1998 ) aphid ecology . chapman & hall , london , pp 8–23

fischer mk , shingleton aw ( 2001 ) host plant and ants influence the honeydew sugar composition of aphids . funct ecol 15 : 544–550

ponsen mb ( 1987 ) alimentary tract . in : minks ak , harrewijn p ( eds ) aphids . their biology , natural enemies and control . volume a . elsevier , amsterdam , pp 79–97

, and its bearing on the evolution of filter chambers in the aphidoidea . wagening agric univ pap 91 ( 5 ) : 1–61

ponsen mb ( 2015 ) a histological description of the salivary gland system of some aphid species of the adelgidae and aphididae . wagening agric univ pap 06 ( 1s ) : 64

shakesby aj , wallace is , isaacs hv , pritchard j , roberts dm , douglas ae ( 2009 ) a water - specific aquaporin involved in aphid osmoregulation . insect biochem mol 39 ( 1 ) : 1–10

shelley wb ( 1969 ) fluorescent staining of elastic tissue with rhodamine b and related xanthene dyes . histochemie 20 ( 3 ) : 244–249

smolla m , ruchty m , nagel m , kleineidam cj ( 2014 ) clearing pigmented insect cuticle to investigate small insects’ organs in situ using confocal laser - scanning microscopy ( clsm ) . arthropod struct dev 43 ( 2 ) : 175–181

völkl w , woodring j , fischer m , lorenz mw , hoffmann kh ( 1999 ) ant - aphid mutualisms : the impact of honeydew production and honeydew sugar composition on ant preferences . oecologia 118 ( 4 ) : 483–491

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three distantly related arthropod groups have converged on a most remarkable social lifestyle . these are gall thrips ( thysanoptera : phlaeothripidae ) , gall aphids ( sternorrhyncha : hormaphididae , pemphigidae , and a few aphididae ) , and snapping shrimp ( decapoda : alpheidae ) , groups in which specialized soldier or defender morphs have evolved in the defense of a static and critical resource , often a nest site that doubles as food source . all three taxa exhibit a range of social structures , the most striking of which involve a morphological division of labor into soldiers and nonsoldier forms . thrips and aphid soldiers have appendages that are enlarged and thickened relative to those of nonsoldiers in the same colony ; these are often armed with spines . similarly , snapping shrimp soldiers bear enlarged chelae . these social groups do not exhibit parental care or much general social interaction ; rather , sociality centers on defense of a valuable and largely irreplaceable resource : the colony domicile . for this reason , this form of sociality is termed

are evident . some social aphids , for example , engage in nest hygiene , expelling honeydew globules that accumulate as a result of the way these insects feed on sap and excrete excess water .

) is produced by a foundress that induces development of a hollow gall on young stem or leaf tissue of the host plant . her brood colony will develop within the safety of the hollow gall , feeding on the plant tissue from within . if the gall is breached , however , the defender morphs will emerge and patrol the surface , aggressively attacking any intruder they encounter . these species are beset by a host of predacious insects such as syrphid fly larvae ; gall thrips also face severe pressure by usurping kleptoparasitic thrips species that kill the foundress and make the gall their own . the domicile of snapping shrimp consists of crevices in sponges , a resource that is less ephemeral than galls but also highly susceptible to usurpation . all of these domiciles represent a valuable , if not critical , commodity that is not easily replaced if lost .

mortality stemming from predation and nest usurpation is likely the most important ecological pressure in the evolution of defender morphs , though it is important to point out that kin selection , in the case of thrips and aphids , may play a role as well . soldier - producing thrips species , which like all members of the order thysanoptera are haplodiploid , often exhibit high levels of inbreeding . aphid colony mates are clonally produced by the foundress , a mode of asexual reproduction called ‘parthenogenesis’ that is found in virtually all aphids and related groups . in social species of both thrips and aphids , there is , accordingly , a high degree of genetic relatedness between colony mates : an average of 0 . 75 in social thrips colonies , and 1 . 0 in colonies of social aphids . this may favor kin selection in these groups , in which indirect fitness – realized through the successful reproduction of relatives – more than compensates the loss of direct fitness that results from altruistic sacrifice ( soldier morphs reproduce less and have a greater chance of dying as a result of colony defense ) .

aphids with morphologically different life stages that exploit taxonomically different host plants are typically difficult to assign to the correct species .

( elm ) species , as the primary host , to identify species and associate the different life stages , thereby using them as a diagnostic tool . using sequence data already available from adult specimens , a phylogeny was produced and the unknown morph sequences clustered with

, confirming the species identity . the authors then described in detail the morphology of the secondary morph , allowing inclusion in future identification keys .

) communities in order to associate adult and larval life stages from tropical lowlands of nepal . although adult taxonomy is well developed and identification is possible ( albeit by taxonomic experts ) , new species are being discovered and the relation of the root - feeding larvae , which have limited morphologically characterising features , to the adults aboveground is unknown . these authors used more in - depth phylogenetic analyses to align sequences to species since intraspecific variation can make this difficult when using absolute methods of sequence divergence and especially for unknown , large assemblages of species . combined mitochondrial and nuclear gene trees ( using maximum likelihood ) and statistical parsimony , whereby populations are subdivided into subgroups , and population aggregation analysis , which combines populations that are uniform for a particular character state , percentage differences (

- distance ) and amova and species delimitation methods for estimating species boundaries from the tree were used to assign species membership . this resulted in 24 species of which 19 could be associated with adults and identified using linnaean names ( nearly 93 % of unknown larvae were identified to species ) . the

sequences , which has implications for gene marker choice in such studies . as for the aphid study , the use of these techniques for determining species and then searching for diagnostic morphological characters of both larvae and adults is advocated . in terms of relating this to species biology , the species assemblages found above - and belowground can depend upon larval mortality , which varies both spatially and temporally ; in particular , larvae and adults were often not found in the same season , which can confound attempts to associate larvae with adults . with a sampling scheme that takes these factors into account , it may be possible to use such dna - based taxonomy to further investigate spatiotemporal patterns and geographic variation .

host - plant resistance is not new to agriculture , but has been applied for at least 200 years . among the earliest successes were resistance to hessian fly ,

, and references therein ) . host - plant resistance is likely to become increasingly important given the continuing homogenization of agricultural landscapes . until the advent of the green revolution , crops were diverse and often hidden in heterogeneous landscapes that consisted of trees , woodland , or other natural and derived habitats . monocultures have eroded landscape diversity and resulted in a loss of habitat for natural enemies , which often increases the vulnerability of crops (

) . agricultural expansion , declining agricultural diversity , and declining crop genetic diversity have demanded a greater focus on intrinsic plant protection . this has coincided with an increasing emphasis on plant molecular biology since the 1990s , largely driven by advances in the molecular tools available to science , including molecular breeding and transgenic technologies .

indicates that for cpb , potato aphids , and tuber moths , the number and proportion of scientific publications related to hpr and insect - plant interactions have generally increased over the past 35 years . during the 2000s , research into conventional hpr and transgenic approaches to plant protection dominated all other management approaches ( cpb 31 % , tuber moths 38 % , and aphids / viruses 24 % of all papers published from 2000 to 2011 ) , including all research on pest natural enemies and biological control . for tuber moths , significant advances have been made in the use of pheromones and granulosis viruses for protection of stored potatoes (

f ) , but for field crops hpr remains the main focus of research . interestingly ,

suggests that research into conventional and transgenic resistance may be mutually competitive in terms of attention or funding opportunities . in the early 2000s , research into transgenics reached its peak ; however , more recently there has been a shift to research of natural induced responses to insect attack and plant biochemical defenses .

it is worth noting that aphids , the masters of phase polyphenism , exhibit yet another form of polyphenism . some aphid species facultatively produce a behaviorally and morphologically distinct soldier caste . since its original description (

abbot et al . , 2001 ; carlin et al . , 1994 ; pike and foster , 2008

foster and rhoden , 1998 ; fukatsu et al . , 2005 ; rhoden and foster , 2002 ; schutze and maschwitz , 1991

) . little is known about proximate mechanisms that induce soldier formation . positive correlations have been found between soldier proportion and colony size (

) , inferring proximate cues that already act during embryonic stages and gradually separating developmental pathways in first instar nymphs .

jhas can be broadly categorized into two groups : the terpenoid compounds , such as methoprene , hydroprene and kinoprene ; and phenoxy jhas , like fenoxycarb and pyriproxyfen . jhas have been found effective especially against adult stages of dipteran and homopteran insects but not against lepidopteran insect pests . as jhas are especially effective against adult stages of the insects , their poor efficacy against lepidopteran crop pests may be because most of these pests cause the damage as larvae (

terpenoid jhas , such as methoprene and kinoprene , have been used successfully against mosquito larvae , scales , and mealybugs . phenoxy jhas cause morphogenetic effects , ovicidal action , and sterility on treated insects . fenoxycarb offers effective control against fire ants , fleas , and sucking insect pests of crops . it is highly effective against sucking pests , such as apple wooly aphid ,

) . pyriproxyfen has been used for controlling aphids , whiteflies , and psyllids causing damage to crops . it has been used for control of locusts ,

most of the bpu compounds have been found very effective against lepidopteran insect pests . diflubenzuron has been found effective against a wide range of lepidopteran pests and locusts (

) . flucycloxuron is effectively used as an acaricide against spider mites . flufenoxuron and novaluron are particularly effective against apple codling moth ,

kanno et al . , 1981 ; yarom et al . , 1988 ; patel et al . , 2010

) . the basis of classical biocontrol is the “enemy release hypothesis . ” according to this hypothesis , the exotic species become pests in new environments by escaping the influence of those natural enemies that suppress their populations in their native range . thus , classical biocontrol reestablishes the top - down control by reintroducing the natural enemies of the pest into its new range (

van driesche and bellows , 1995 ; crawley , 1997 ; keane and crawley , 2002 ; hajek , 2004 ; naranjo et al . , 2015

thus , classical biocontrol primarily describes the releases of insect predators , parasitoids , and pathogens to control other insect pests and insect herbivores to control weeds . this form of biocontrol is appropriate when insects that spread or are introduced ( usually accidentally ) to areas outside of their natural range become pests mainly because of the absence of their natural enemies . the same strategy has also been called “importation” by

( homoptera : mgarodidae ) in california in the late 1800s . likewise , ash whitefly (

, which was introduced in june 1898 . although , the predator did not control soft green scale ,

, which was its specific target , in july 1951 , it controlled various mealybugs infesting fruit crops , coffee , ornamental plants , etc . , in south india . the predator is now effective in suppressing mealybug infestations on citrus , guava , grapes , mulberry , coffee , mango , pomegranate , custard apple , ber , etc . , and green shield scale on sapota , mango , guava , brinjal , and crotons in karnataka (

( native of eastern united states ) , was accidentally introduced to india from england . it soon spread to all of the apple - growing areas of the country and started causing severe damage . for the control of woolly aphid , exotic aphelinid parasitoid ,

( native of north america ) , was introduced from the united kingdom to saharanpur ( uttar pradesh , india ) . however , the parasitoid failed to establish itself because of the intense activity of a ladybird beetle ,

, a native of the caribbean region and central america , was introduced into india in 1995 . first reported from kerala , it soon spread to all of the southern states , causing serious damage to several plants . it started attacking more than 253 plant species belonging to 176 genera and 60 families . however , serious damage was caused to avocado , banana , cassava , guava , papaya , and mango in addition to several ornamental and avenue trees . as a result , exotic aphelinid parasitoids ,

causing a perceptible reduction in the pest population . although the parasitism levels due to both parasitoids vary from 29 % to 70 % and exceed 90 % during some parts of the year , the former is performing better than the latter and has established itself in kerala , karnataka , and several parts of andhra pradesh , where it was previously absent (

despite having many beneficial aspects , classical biocontrol is currently not being encouraged because negative environmental effects may arise through ill - considered introductions of exotic natural enemies . many introduced agents have failed to control pests ; for example , more than 60 predators and parasitoids have been introduced into the northeastern part of north america with little effect on the target gypsy moth ,

( lymantriidae ) . some introductions have strengthened the pest problems whereas others have become pests themselves . exotic introductions generally are irreversible , and nontarget species can suffer worse consequences from efficient natural enemies than from chemical insecticides , which are unlikely to cause total extinctions of native insect species . there are documented cases of introduced biocontrol agents wiping out native invertebrates . several endemic hawaiian insects ( target and nontarget ) have become extinct largely as a result of biocontrol introductions (

although initial classical biocontrol was predominantly focused on control of introduced pests with the goal of reestablishing host / natural enemy associations that keep pests in check in their areas of origin , the strategy has now also been applied against native pests ( e . g . , see

is used when an exotic natural enemy is introduced against a native pest . however , the introduction ( importation , augmentation , and release ) of exotic natural enemies against exotic pests with which they did not coevolve is termed

such new associations are supposed to be very effective at controlling pests because the pest has not coevolved with the introduced enemies . unfortunately , the exotic species that are most likely to be effective biocontrol agents because of their ability to utilize new hosts are also those most likely to be a threat to nontarget species . an example of the possible dangers of neoclassical control is provided by the work of jeffrey lockwood , who campaigned against the introduction of a parasitic wasp and an entomophagous fungus from australia as control agents of native rangeland grasshoppers in the western united states . the potential adverse environmental effects of such introductions include the suppression or extinction of many nontarget grasshopper species with probable concomitant losses of biodiversity and existing weed control and disruptions to food chains and plant community structure . the inability to predict the ecological outcomes of neoclassical introductions means that they are highly risky , especially in systems where the exotic agent is free to expand its range over large geographical areas .


tyler 's tree frog or the southern laughing tree frog ( litoria tyleri ) is an arboreal tree frog .

it is native to eastern australia .

it occurs from southeast queensland to the south coast of new south wales .

it is generally a coastal species and is not found inland .

tyler ' s tree frog

tyler ' s tree frog <EOT> tyler ' s tree frog ( litoria tyleri ) has large toe pads and webbed feet .

tyler ' s tree frog ( litoria tyleri ) is similar - looking but has a golden iris .

the revealed tree frog or whirring tree frog , ( litoria revelata ) is a species of tree frog native to coastal eastern australia .

the peron ' s tree frog is found in northern victoria , new south wales and southern queensland .

the peron ' s tree frog breeds in temporary pools , dams and ditches and sometimes tries to breed in suburban fish ponds .

genus . the male peron ' s tree frog is about 44 - 53 mm , while females are 46 - 65 mm .

the peron ' s tree frog lives in most forest habitats but will also forage for food in open grassland and other open areas .

the peron ' s tree frog has a drill - like call , which has also been described as a ' maniacal cackle ' .

southern laughing tree frog ( litoria tyleri ) by stephen zozaya d ' aguilar range , queensland .

australia has many species of tree frogs , here i film peron ' s tree frog in a breeding pond making a racket - the stuttered laugh call . also present but not seen were tyler ' s tree frog with the fast ' ark ark ark . . . . . ' call . there are a couple more frogs species in this small pond but i ' m not sure of them . filmed on the south coast of nsw on a warm october evening .

some frogs in the tropical rainforest are specially adapted for gliding from tree to tree or parachuting to the forest floor . typical of them is

close - up of frog ' s head showing eye , nostril , mouth and tympanum .

barker , j . , grigg , g . c . , and tyler , m . j . ( 1995 ) .

. some species are carnivorous at the tadpole stage , eating insects , smaller tadpoles , and fish . the cuban tree frog (

males make a high - pitched whirring noise , similar to the verreaux ' s tree frog ; calling occurs from spring to autumn , either from the ground or from vegetation bordering the breeding area .

tree frogs are frogs of the family hylidae . they are of small size and more elegant in form than the true frogs ( family ranidae ) . many tree frogs have bright coloration .

, a less toxic product , by most terrestrial adults . a few species of tree frog with little access to water excrete the even less toxic

another tactic used by some frogs is to " scream " , the sudden loud noise tending to startle the predator . the gray tree frog (

a barred frog provides surrogate eggs for cloning the southern gastric brooding frog . photo by bob beale

i always wondered what these pale grey frogs were called - now i know - a " peron ' s tree frog " . thank you ! i have seen these around but have not heard any ' maniacal cackle ' : )

please don’t let your boyfriend attempt to house a new baby bearded dragon with chameleons and tree frogs . it’s a bad combination because they all require different habitat temperature and humidity parameters .

. indonesia is the world ' s largest exporter of frog meat , exporting more than 5 , 000 tonnes of frog meat each year , mostly to france , belgium and luxembourg .

the southern gastric brooding frog ( rheobatrachus silus ) was discovered in 1972 in the mountains of queensland , australia . but the world only took notice of it in 1974 when mike tyler discovered how it reproduced .

archer has several reasons for trying . there’s the medical potential . there’s the unusual nature of the frog’s life cycle , which no other animal shares . but really , it’s a more “transcendent reason” that drives him . “if we were responsible for the extinction of the species , deliberately or inadvertently , we have a moral responsibility or imperative to undo that if we can , ” he says .

archer , however , is in the business of de - extinction . he’s going to clone the southern gastric brooding frog back into life .

unfortunately , this is not the case for a tree frog , box turtle , chameleon and bearded dragon . bearded dragons are from arid desert areas in australia . they require high heat and low humidity , and their daytime temperature range is 75 to 85 degrees fahrenheit with a focal hot spot of 110 degrees f . being amphibians , tree frogs require higher humidity and a cooler environment . if you tried to keep the temperature range high enough for the beardie , you would end up dehydrating your tree frog , resulting in its untimely demise . i don’t know what kind of tree frog you have , so i cannot offer more specific recommendations . it would be safest to house your chameleons and tree frogs in separate habitats .

" the frog prince " is a fairy tale about a frog that turns into a handsome prince after he has rescued a princess ' s golden ball and she has taken him into her palace .

still , archer is hopeful . whenever he has gone through the same technical motions with a living frog , and inserted the species’ nucleus into its own egg , the resulting embryo also paused at the same point . this suggests that there’s something wrong with the team’s techniques , rather than with tyler’s frozen gastric brooding frog tissues . busted tissues would be a deal breaker but technological problems can be fixed , and archer has brought in stem cell expert robert lanza to help him do so . “we retain our vibrant optimism , ” he says .

frog populations have declined dramatically since the 1950s . more than one - third of frog species are considered to be threatened with

. some frogs use their sticky tongues to catch fast - moving prey , while others push food into their mouths with their hands . a few species also eat plant matter ; the tree frog

the colour of a frog ' s skin is used for thermoregulation . in cool damp conditions , the colour will be darker than on a hot dry day . the

when news broke about this weird strategy , other scientists were incredulous . tyler provided vivid accounts of a young frog poking its head out of mum’s mouth like an amphibian russian doll , but even these were insufficient . “it just seemed to many zoologists absolutely impossible , ” he later wrote in a book . “there were frequently insinuations that somehow we were wrong . ”

tree frogs and some non - aquatic species have a rain call that they make on the basis of humidity cues prior to a shower .

frost , s . w . ( 1932 ) . " notes on feeding and molting in frogs " .

tree frogs are very acrobatic and can catch insects while hanging by one toe from a twig or clutching onto the blade of a windswept reed .

the peron ' s tree frog has the ability to quickly change colour . by day it is usually a pale green - grey colour that changes to a reddish brown with emerald green flecks at night . it also has bright black and yellow markings on its thighs . it has a cross - shaped pupil and a silver iris .

is a common defensive mechanism in frogs . most camouflaged frogs are nocturnal ; during the day , they seek out a position where they can blend into the background and remain undetected . some frogs have the ability to change colour , but this is usually restricted to a small range of colours . for example , white ' s tree frog (

stuart , s . n . ; chanson , j . s . ; cox , n . a . ; young , b . e . ; rodrigues , a . s . l . ; fischman , d . l . ; waller , r . w . ( 2004 ) . " status and trends of amphibian declines and extinctions worldwide " .

vancompernolle , s . e . ; taylor , r . j . ; oswald - richter , k . ; jiang , j . ; youree , b . e . ; bowie , j . h . ; tyler , m . j . ; conlon , m . ; wade , d . ; et al . ( 2005 ) .

hudson , n . j . ; lehnert , s . a . ; ingham , a . b . ; symonds , b . ; franklin , c . e . ; harper , g . s . ( 2005 ) . " lessons from an estivating frog : sparing muscle protein despite starvation and disuse " .

estrada , alberto r . ; hedges , s . blair ( 1996 ) . " at the lower size limit in tetrapods : a new diminutive frog from cuba ( leptodactylidae : eleutherodactylus ) " .

peron ' s tree frog is one of the most variably coloured frogs in australia , with the ability to change colour in less than one hour . it varies in shades of grey and brown , where its lightest is almost white . the frog has mottled yellow and black thighs , armpits , and groin . occasionally , emerald spots are found on the back , which increase in number with age . a characteristic uncommon in the genus

and a frog deprived of its lungs can maintain its body functions without them .

ford , l . s . ; cannatella , d . c . ( 1993 ) . " the major clades of frogs " .

anderson , jason s . ; reisz , robert r . ; scott , diane ; fröbisch , nadia b . ; sumida , stuart s . ( 2008 ) . " a stem batrachian from the early permian of texas and the origin of frogs and salamanders " .

, where they scramble around on the branches , twigs , and leaves , sometimes never coming down to earth . the " true " tree frogs belong to the family hylidae , but members of other frog families have independently adopted an arboreal habit , a case of

surface rendering of the head of the frog atelopus franciscus , with ear parts highlighted .

to clone the gastric brooding frog , the team first needed its dna . archer called up mike tyler , who rummaged through his freezers and found some old tissue samples . they were in shoddy condition—just bits of frog dropped in a container , without any antifreeze to protect them . the cells should have been useless , ruptured sacks but they had somehow stayed intact . “we thought it was worth a try , ” archer says .

federle , w . ; barnes , w . j . p . ; baumgartner , w . ; drechsler , p . ; smith , j . m . ( 2006 ) . " wet but not slippery : boundary friction in tree frog adhesive toe pads " .

marjanović , d . ; laurin , m . ( 2009 ) . " the origin ( s ) of modern amphibians : a commentary " .

emerson , s . b . ; diehl , d . ( 1980 ) . " toe pad morphology and mechanisms of sticking in frogs " .

some species of frog have adaptations that allow them to survive in oxygen deficient water . the

then , good news ! a second species—the northern gastric brooding frog ( rheobatrachus vitellinus ) —was discovered in 1984 in queensland’s eungella national park . but a year later , almost before anyone could uncork the celebratory champagne , it too went extinct .

it took many years , field surveys and photographs to persuade the naysayers . tyler eventually published a full description of the frog and its behaviour in 1981 . ( nature rejected the paper because they—wrongly—deemed it uninteresting . ) the medical community took notice . if this creature could deliberately stop making acid in its stomach , it might provide new ways of treating stomach ulcers or helping people who go through stomach surgeries to heal more quickly . several teams started studying the frog .

during the evolutionary history of the frog , several different groups have independently taken to the air .

is so powerful , one frog contains enough poison to kill an estimated 22 , 000 mice .

the atlas of living australia acknowledges australia ' s traditional owners and pays respect to the past and present elders of the nation ' s aboriginal and torres strait islander communities . we honour and celebrate the spiritual , cultural and customary connections of traditional owners to country and the biodiversity that forms part of that country .

the frog has a highly developed nervous system that consists of a brain , spinal cord and nerves . many parts of the frog ' s brain correspond with those of humans . it consists of two olfactory lobes , two cerebral hemispheres , a pineal body , two optic lobes , a cerebellum and a medulla oblongata . muscular coordination and posture are controlled by the

this might seem like an over - reaction . after all , millions of frog eggs divide into embryos every day , as they have done since before dinosaurs walked the earth . but this egg was special . archer’s team of scientists had loaded it with the dna of the southern gastric brooding frog —a bizarre creature that has been extinct for almost 30 years .

this article is about the group of amphibians . for other uses , see frog ( disambiguation ) .

youtube video of a frog eating a caterpillar gets close to 1 . 5 million views in three days

nauwelaerts , s . ; schollier , j . ; aerts , p . ( 2004 ) . " a functional analysis of how frogs jump out of water " .

every time the team has done this , the ball of cells starts to turn inwards on itself—a crucial moment called gastrulation—and stops . that’s where they are for now . they have the beginnings of a gastric brooding frog , but are a long way from even a simple tadpole .

, united states . in asia , frog legs are consumed in china , vietnam , thailand and indonesia .

are farmed and consumed on a large scale in some areas of china . frog legs are part of chinese

" national recovery plan for the southern corroboree frog ( pseudophryne corroboree ) : 5 . previous recovery actions "

tree frog with a small head and large eyes . mottled grey to brown with fine granular skin , sometimes with a few green spots on back . eye has golden coloured iris with cross shaped pupil . the underside is cream or yellowish . the hind surface of thighs is yellow and dark brown / black . large toe discs .

let’s assume archer succeeds . where would the new generation gastric brooding frogs live ? their habitat in the queensland mountains is being threatened by feral pigs , invasive weeds and polluted or diverted waters . and then there’s the chytrid fungus , which has spread to almost every part of the world . it would be like releasing lazarus into an ecological dystopia .

whether a frog sees in colour is debatable but it has been shown that it responds positively to blue light , perhaps because that colour is associated with bodies of water that can provide refuge when the frog feels threatened .

the team then needed something to put the dna into—the egg of another frog . he chose a barred frog —a reasonably close relative that produces large eggs of the right size . the downside—and it’s a big one—is that barred frogs only lay eggs once a year . “we had a few times when we went in and the frogs weren’t laying , and that was that for the year , ” says archer .

, as well as to water . there are blood vessels near the surface of the skin and when a frog is underwater , oxygen diffuses directly into the blood . when not submerged , a frog breathes by a process known as

the call or croak of a frog is unique to its species . frogs create this sound by passing air through the

after metamorphosis , young adults may disperse into terrestrial habitats or continue to live in water . almost all frog species are

so i just found this little guy in my backyard . i have no idea about frogs only snakes haha anyone know what it is ? or is it just a garden frog ? it ' s eyes where crazy looking . i ' ve never seen one like this in my yard before haha . thanks !

sandberger , l . ; hillers , a . ; doumbia , j . ; loua , n - s . ; brede c . ; rödel , m - o . ( 2010 ) .

warkentin , i . g . ; bickford , d . ; sodhi , n . s . ; corey , j . a . ( 2009 ) . " eating frogs to extinction " .

can you put chameleons and tree frogs in the same cage together ? i ' ve also heard that you’re not supposed to hold chameleons all that much . is that true ? also my dumb boyfriend is wanting to buy a bearded dragon , and he wants to keep it at my house . the only problem is he is trying to put it in the same cage as the chameleons and tree frogs . even though the cage is big enough , i want to know if it is ok to do .

the grip of the male frog during amplexus stimulates the female to release eggs , usually wrapped in jelly , as spawn .

named 2008 as the " year of the frog " in order to draw attention to the conservation issues faced by them .

because frog toxins are extraordinarily diverse , they have raised the interest of biochemists as a " natural pharmacy " . the alkaloid

) has short , slim hind limbs unsuited to jumping . it can move fast by using a running gait in which the two hind legs are used alternately . slow - motion photography shows , unlike a horse that can trot or gallop , the frog ' s gait remained similar at slow , medium , and fast speeds .

a canadian study conducted in 2006 suggested heavy traffic in their environment was a larger threat to frog populations than was habitat loss .

some of the arguments against de - extinction don’t apply to the gastric brooding frog . unlike the woolly mammoth or passenger pigeon , the frog isn’t a social creature that would need companions to learn from or travel among . unlike the mammoth , which would need to be born inside an elephant , the frog doesn’t need a complicated surrogate parent . and unlike many of the candidates for de - extinction , like the moa or saber - toothed cat , the frog is small and can be reared in a laboratory . archer has so much frozen tissue that once he successfully clones one frog , he could make a practically infinite supply of them .

archer’s goal is simple : to bring the extinct gastric brooding frog back from oblivion and , in doing so , provide hope for the hundreds of other frogs that are heading that way . getting the embryo was a milestone and archer is buoyantly optimistic that he’ll cross the finish line soon . lazarus , he says , will rise again .

. this allows the frog to climb on smooth surfaces , but the system does not function efficiently when the pads are excessively wet .

although few species are toxic enough to be used for this purpose . at least two non - poisonous frog species in tropical america (

shubin , n . h . ; jenkins , f . a . jr ( 1995 ) . " an early jurassic jumping frog " .

, shows how each frog family is related to other families , with each node representing a point of common ancestry . it is based on frost

) digs into the rear of a tadpole , causing a rearrangement of the limb bud cells and the frog develops an extra leg or two .

they may be the only part of an otherwise submerged frog to protrude from the water . each eye has closable upper and lower lids and a

different species of frog use a number of methods of moving around including jumping , running , walking , swimming , burrowing , climbing and gliding .

many frogs are able to absorb water and oxygen directly through the skin , especially around the pelvic area , but the permeability of a frog ' s skin can also result in water loss . glands located all over the body exude mucus which helps keep the skin moist and reduces evaporation . some glands on the hands and chest of males are specialized to produce sticky secretions to aid in

. similar glands in tree frogs produce a glue - like substance on the adhesive discs of the feet . some arboreal frogs reduce water loss by having a waterproof layer of skin , and several south american species coat their skin with a waxy secretion . others frogs have adopted behaviours to conserve water , including becoming

have become elongated to add to the leg length and allow the frog to push against the ground for a longer period on take - off . the

a distress call , emitted by some frogs when they are in danger , is produced with the mouth open resulting in a higher - pitched call . it is typically used when the frog has been grabbed by a predator and may serve to distract or disorientate the attacker so that it releases the frog .

is frog - like , being broad with large eye sockets , but the fossil has features diverging from modern frogs . these include a longer body with more

a frog may be startled by an unexpected noise but it will not usually take any action until it has located the source of the sound by sight .

features michigan j . frog , that will only dance and sing for the demolition worker who opens his time capsule , but will not perform in public .

a frog ' s skin is protective , has a respiratory function , can absorb water and helps control body temperature . it has many glands , particularly on the head and back , which often exude distasteful and toxic substances . the secretion is often sticky and helps keep the skin moist , protects against the entry of moulds and bacteria , and make the animal slippery and more able to escape from predators .

a frog of the family hylidae , of small size and more elegant in form than the true frogs ( family ranidae ) , often with a bright coloration .

many species also have a territorial call that is used to drive away other males . all of these calls are emitted with the mouth of the frog closed .

in small pools , predators are mostly absent and competition between tadpoles becomes the variable that constrains their survival . certain frog species avoid this competition by making use of smaller

) was placed on a treadmill which was turned at varying speeds . by measuring the toad ' s uptake of oxygen it was found that hopping was an inefficient use of resources during sustained locomotion but was a useful strategy during short bursts of high - intensity activity .

) on being approached by a snake is to crouch down and remain immobile . this is usually successful , with the snake passing by and the toad remaining undetected . if it is encountered by the snake ' s head , however , the toad hops away before crouching defensively .

lips has another concern : resurrection projects take up a lot of money . archer concedes that cloning research is initially expensive , but he says that costs will eventually fall . “i can’t think of what cloning dolly must have cost and now it’s a routine technique , ” he says .

a different call is emitted by a male frog or unreceptive female when mounted by another male . this is a distinct chirruping sound and is accompanied by a vibration of the body .

slow - motion photography shows that the muscles have passive flexibility . they are first stretched while the frog is still in the crouched position , then they are contracted before being stretched again to launch the frog into the air . the fore legs are folded against the chest and the hind legs remain in the extended , streamlined position for the duration of the jump .

) was first widely used in laboratories in pregnancy tests in the first half of the 20th century . a sample of urine from a pregnant woman injected into a female frog induces it to lay

) are traditionally used by native colombians to poison the darts they use for hunting . the tip of the projectile is rubbed over the back of the frog and the dart is launched from a

, environmental factors contributing to rates of disease , and vulnerability to attack by parasites . malformations impair mobility and the individuals may not survive to adulthood . an increase in the number of frogs eaten by birds may actually increase the likelihood of parasitism of other frogs , because the trematode ' s complex lifecycle includes the

. in some cases , the full leg still grows , but in others it does not , although the frog may still live out its normal lifespan with only three limbs . occasionally , a parasitic

frost , d . r . ; grant , t . ; faivovich , j . n . ; bain , r . h . ; haas , a . ; haddad , c . l . f . b . ; de sá , r . o . ; channing , a . ; wilkinson , m . ; donnellan , s . c . ; raxworthy , c . j . ; campbell , j . a . ; blotto , b . l . ; moler , p . ; drewes , r . c . ; nussbaum , r . a . ; lynch , j . d . ; green , d . m . ; wheeler , w . c . ( 2006 ) . " the amphibian tree of life " .

like other amphibians , the life cycle of a frog normally starts in water with an egg that hatches into a limbless larva with gills , commonly known as a tadpole . after further growth , during which it develops limbs and lungs , the tadpole undergoes metamorphosis in which its appearance and internal organs are rearranged . after this it is able to leave the water as a miniature , air - breathing frog .

frogs are valued as food by humans and also have many cultural roles in literature , symbolism and religion . frog populations have declined significantly since the 1950s . more than one third of species are considered to be

at the end of the tadpole stage , a frog undergoes metamorphosis in which its body makes a sudden transition into the adult form . this metamorphosis typically lasts only 24 hours , and is initiated by production of the

when a frog shoots out its tongue to catch an insect it is reacting to a small moving object that it cannot see well and must line it up precisely beforehand because it shuts its eyes as the tongue is extended .

the distant vision of a frog is better than its near vision . calling frogs will quickly become silent when they see an intruder or even a moving shadow but the closer an object is , the less well it is seen .

that amplifies the sound . species of frog that lack vocal sacs and that do not have a loud call tend to inhabit areas close to constantly noisy , flowing water . they need to use an alternative means to communicate . the

) lays her eggs on the forest floor . the male frog guards them from predation and carries water in his cloaca to keep them moist . when they hatch , the female moves the tadpoles on her back to a water - holding

and its population crashed in 1987 , along with about 20 other frog species in the area . this could not be linked directly to human activities , such as deforestation , and was outside the range of normal fluctuations in population size .

from early in its development , a gill pouch covers the tadpole ' s gills and front legs . the lungs soon start to develop and are used as an accessory breathing organ . some species go through metamorphosis while still inside the egg and hatch directly into small frogs . tadpoles lack true teeth , but the jaws in most species have two elongated , parallel rows of small ,

) are directly exposed or may be covered by a layer of skin and are visible as a circular area just behind the eye . the size and distance apart of the eardrums is related to the frequency and wavelength at which the frog calls . in some species such as the bullfrog , the size of the tympanum indicates the sex of the frog ; males have tympani that are larger than their eyes while in females , the eyes and tympani are much the same size .

the main reason for calling is to allow male frogs to attract a mate . males may call individually or there may be a chorus of sound where numerous males have converged on breeding sites . females of many frog species , such as the

and resting in a water - conserving position . some frogs may also rest in large groups with each frog pressed against its neighbours . this reduces the amount of skin exposed to the air or a dry surface , and thus reduces water loss .

because predation of eggs and larvae is high in large water bodies , some frog species started to lay their eggs on land . once this happened , the desiccating terrestrial environment demands that one or both parents keep them moist to ensure their survival .

the skin is shed every few weeks . it usually splits down the middle of the back and across the belly , and the frog pulls its arms and legs free . the sloughed skin is then worked towards the head where it is quickly eaten .

iskandar , d . t . ; evans , b . j . ; mcguire , j . a . ( 2014 ) . " a novel reproductive mode in frogs : a new species of fanged frog with internal fertilization and birth of tadpoles " .

) are also used for this purpose . these are less toxic and less abundant than the golden poison frog . they are impaled on pointed sticks and may be heated over a fire to maximise the quantity of poison that can be transferred to the dart .

while these smaller rearing sites are free from competition , they also lack sufficient nutrients to support a tadpole without parental assistance . frog species that changed from the use of larger to smaller phytotelmata have evolved a strategy of providing their offspring with nutritive but unfertilized eggs .

archer sees it slightly differently—the plight of living species gives him even more recourse to bring back extinct ones . “no matter how many resources we put into looking after the environment , wildlife is no longer safe in the wild , ” he says . “if we accept that maintaining biodiversity is important , we can’t assume that if you whack a fence up , everything’s going to be okay . you need to explore lots of parallel strategies . ”

two years ago , mike archer from the university of new south wales looked down a microscope and saw that a single fertilised frog egg had divided in two . then , it did it again . and again . eventually , the egg produced an embryo containing hundreds of cells .

frogs that live in or visit water have adaptations that improve their swimming abilities . the hind limbs are heavily muscled and strong . the webbing between the toes of the hind feet increases the area of the foot and helps propel the frog powerfully through the water . members of the family

are considered " true toads " . the use of the term " frog " in common names usually refers to species that are aquatic or semi - aquatic and have smooth , moist skins ; the term " toad " generally refers to species that are terrestrial with dry , warty skins .

in the throat . in most calling frogs , the sound is amplified by one or more vocal sacs , membranes of skin under the throat or on the corner of the mouth , that distend during the amplification of the call . some frog calls are so loud that they can be heard up to a mile away .

lays up to 40 eggs on the ground , where they are guarded by the male . when the tadpoles are about to hatch , they are engulfed by the male , which carries them around inside his much - enlarged vocal sac . here they are immersed in a frothy , viscous liquid that contains some nourishment to supplement what they obtain from the yolks of the eggs . they remain in the sac for seven to ten weeks before undergoing metamorphosis , after which they move into the male ' s mouth and emerge .

have pads located on the ends of their toes to help grip vertical surfaces . these are not suction pads , the surface consisting instead of columnar cells with flat tops with small gaps between them lubricated by mucous glands . when the frog applies pressure , the cells adhere to irregularities on the surface and the grip is maintained through

archer is unfazed . “we can ultimately fix the wild , ” he says . “even if we had to maintain most of the world’s wildlife in artificial environments , that would be a thousand times better than to let them slide off the brink . ” the frogs can wait until their homes are ready for them . in the meantime , scientists could perhaps engineer or breed them to be resistant to the chytrid fungus , or carry out experimental releases to see whether they would actually find a niche in this brave , new world .

is increased . this means that the limited amount of energy available to the comatose frog is used in a more efficient manner . this survival mechanism is only useful to animals that remain completely unconscious for an extended period of time and whose energy requirements are low because they are cold - blooded and have no need to generate heat .

among prolonged breeders , males usually arrive at the breeding site first and remain there for some time whereas females tend to arrive later and depart soon after they have spawned . this means that males outnumber females at the water ' s edge and defend territories from which they expel other males . they advertise their presence by calling , often alternating their croaks with neighbouring frogs . larger , stronger males tend to have deeper calls and maintain higher quality territories . females select their mates at least partly on the basis of the depth of their voice .

. it then travels on through the ureters , which are consequently known as urinogenital ducts . there is no penis , and sperm is ejected from the cloaca directly onto the eggs as the female lays them . the ovaries of the female frog are beside the kidneys and the eggs pass down a pair of oviducts and through the cloaca to the exterior .

faivovich , j . ; haddad , c . f . b . ; garcia , p . c . a . ; frost , d . r . ; campbell , j . a . ; wheeler , w . c . ( 2005 ) . " systematic review of the frog family hylidae , with special reference to hylinae : phylogenetic analysis and revision " .

once the team had their surrogate egg , they had to destroy the native nucleus so they could insert one from the frozen gastric brooding frog tissues . they either did the job manually with a very fine instrument , or bombarded the egg with ultraviolet ( uv ) radiation . they tried both techniques on hundreds of eggs and one of these eventually divided into an early embryo with hundreds of cells .

, located on the sides of their heads behind the eyes and other glands elsewhere on their bodies . these glands secrete mucus and a range of toxins that make frogs slippery to hold and distasteful or poisonous . if the noxious effect is immediate , the predator may cease its action and the frog may escape . if the effect develops more slowly , the predator may learn to avoid that species in future .

the structure of the feet and legs varies greatly among frog species , depending in part on whether they live primarily on the ground , in water , in trees or in burrows . frogs must be able to move quickly through their environment to catch prey and escape predators , and numerous adaptations help them to do so . most frogs are either proficient at jumping or are descended from ancestors that were , with much of the

being cold - blooded , frogs have to adopt suitable behaviour patterns to regulate their temperature . to warm up , they can move into the sun or onto a warm surface ; if they overheat , they can move into the shade or adopt a stance that exposes the minimum area of skin to the air . this posture is also used to prevent water loss and involves the frog squatting close to the substrate with its hands and feet tucked under its chin and body .

, the larvae of which damage and kill the canes . initial results in many of these countries were positive , but it later became apparent that the toads upset the ecological balance in their new environments . they bred freely , competed with native frog species , ate bees and other harmless native invertebrates , had few predators in their adopted habitats , and poisoned pets , carnivorous birds , and mammals . in many of these countries , they are now regarded both as pests and

at first sight , frogs seem rather defenceless because of their small size , slow movement , thin skin , and lack of defensive structures , such as spines , claws or teeth . many use camouflage to avoid detection , the skin often being spotted or streaked in neutral colours that allow a stationary frog to merge into its surroundings . some can make prodigious leaps , often into water , that help them to evade potential attackers , while many have other defensive adaptations and strategies .

frogs have maxillary teeth along their upper jaw which are used to hold food before it is swallowed . these teeth are very weak , and cannot be used to chew or catch and harm agile prey . instead , the frog uses its sticky , cleft tongue to catch flies and other small moving prey . the tongue normally lies coiled in the mouth , free at the back and attached to the mandible at the front . it can be shot out and retracted at great speed .

this frog reaches 40 mm in length . it is cream to red - brown on the back , with a darker band running down the middle . males turn bright yellow in colour during the breeding season . a dark strip runs from the nostril to the shoulder , across the tympanum . the back the legs are red and the thigh is yellow - orange . some large black dots occur on the thigh and the backs of legs . the belly is cream and the iris is golden .

still , funding is a zero - sum game . there’s only so much cash to go around and conservationists need it to monitor animals that are still alive , work out why they are disappearing , and develop ways of saving them . there are plenty of cases where we know how to save a species , but can’t afford to do so . “i can’t help but think that we can’t even take care of what we’ve got , and now we’re going to invest in very expensive techniques to recover a handful of special - interest species that may or may not be able to survive in the wild on their own , ” says lips . as a best - case scenario , she hopes that these high - profile projects will help to drum up interest in saving a broader swathe of imperilled wildlife .

) normally sink to the bottom of the pond where they lie , semi - immersed in mud but still able to access the oxygen dissolved in the water . their metabolism slows down and they live on their energy reserves . some frogs can even survive being frozen . ice crystals form under the skin and in the body cavity but the essential organs are protected from freezing by a high concentration of glucose . an apparently lifeless , frozen frog can resume respiration and the heart beat can restart when conditions warm up .

simply put , the mother frog converts her stomachs into a womb . she swallows her own eggs and stops making hydrochloric acid in her stomach to avoid digesting her own young . around 20 to 25 tadpoles hatch inside her and the mucus from their gills continues to keep the acid at bay . while the tadpoles grow over the next six weeks , mum never eats . her stomach bloats so much that her lungs collapse , forcing her to breathe through her skin . eventually , she gives birth to her brood through “propulsive vomiting” , spewing them into the world as fully - formed froglets .

the muscular system has been similarly modified . the hind limbs of ancestral frogs presumably contained pairs of muscles which would act in opposition ( one muscle to flex the knee , a different muscle to extend it ) , as is seen in most other limbed animals . however , in modern frogs , almost all muscles have been modified to contribute to the action of jumping , with only a few small muscles remaining to bring the limb back to the starting position and maintain posture . the muscles have also been greatly enlarged , with the main leg muscles accounting for over 17 % of the total mass of the frog .

eastern coast of australia from south - eastern queensland to southern new south wales . the extent of occurrence of the species is approximately 73000 km2

variety of habitats , but usually wet and dry forests . often found in trees and other vegetation overhanging permanent ponds . spring breeder . male call from vegetation surrounding a pond where spawn is deposited and tadpoles develop .

j - m hero et al . ( m . hero at mailbox . gu . edu . au ) , griffith university

> university of california , berkeley , ca , usa . accessed jul 9 , 2018 .

> university of california , berkeley , ca , usa . accessed 9 jul 2018 .

html public " - / / w3c / / dtd xhtml 1 . 0 strict / / en " " urltoken "

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justification : listed as least concern in view of its wide distribution , tolerance of a broad range of habitats , presumed large population , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .

this australian endemic occurs along the eastern coast of australia from southeastern queensland to southern new south wales . the extent of occurrence of the species is approximately 73 , 000 km 2 .

the species occurs in a wide range of habitats : melaleuca woodland , upland , and lowland wet sclerophyll forest . it is often found in trees and other vegetation overhanging permanent ponds . it is a spring breeder . males call from vegetation surrounding still water where spawn is deposited and tadpoles develop . it breeds in permanent and semi - permanent water . it also occurs in pasture lands .

this species was accidentally transferred to the genus nyctimystes due to confusion between litora michaeltyleri and l . tyleri . this error is corrected here .

( errata version published in 2015 ) . the iucn red list of threatened species 2004 : e . t41113a85820328 .

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urn : lsid : biodiversity . org . au : afd . taxon : 29ccf6db - 4cad - 4878 - b70c - 7b7e59d0dfde

urn : lsid : biodiversity . org . au : afd . taxon : 69f078cf - b0ff - 4842 - b5dc - 1a2fcd48c1e2

urn : lsid : biodiversity . org . au : afd . taxon : a591355f - 030a - 401d - 89ce - 96258d822782

urn : lsid : biodiversity . org . au : afd . taxon : c35b38e7 - d304 - 4b80 - 8a62 - 2b8ca054e27d

urn : lsid : biodiversity . org . au : afd . taxon : f5f7c98d - 045f - 42b7 - b88b - d3f65aecd905

urn : lsid : biodiversity . org . au : afd . taxon : 74d6d114 - 993d - 4e74 - 9786 - 245b68e4943d

urn : lsid : biodiversity . org . au : afd . name : 291914

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these frogs are also in south australia we go to the river murray quite often ( morgan ) & they are everywhere there . . .

this is another species calling heavily around our dam . during the breeding season the males adopt this brilliant yellow colour , in contrast to the grey / brown they are for the rest of the year .

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and over one hundred and twenty are believed to have become extinct since the 1980s .

" has no taxonomic justification . from a classification perspective , all members of the order anura are frogs , but only members of the family

species that fit within the anuran definition . the characteristics of anuran adults include : 9 or fewer presacral vertebrae , the presence of a urostyle formed of fused vertebrae , no tail , a long and forward - sloping ilium , shorter fore limbs than hind limbs ,

, by far the largest group , which contains the remaining 24 families of modern frogs , including most common species throughout the world . the neobatrachia suborder is further divided into the two superfamilies

, and the morphology of tadpoles . while this classification is largely accepted , relationships among families of frogs are still debated .

are similar in forming hybrids . these are less fertile than their parents , giving rise to a

the origins and evolutionary relationships between the three main groups of amphibians are hotly debated . a

. this would help account for the relative scarcity of amphibian fossils from the period before the groups split .

seemed to have originated in africa / india , the salamanders in east asia and the caecilians in tropical pangaea .

other researchers , while agreeing with the main thrust of this study , questioned the choice of calibration points used to synchronise the data . they proposed that the date of lissamphibian diversification should be placed in the

, rather less than 300 million years ago , a date in better agreement with the palaeontological data .

was estimated as taking place 292 million years ago , rather later than most molecular studies suggest , with the caecilians splitting off 239 million years ago .

) , " to jump " ) is the name of the total group that includes modern frogs in the order anura as well as their close fossil relatives , the " proto - frogs " or " stem - frogs " . the common features possessed by these proto - frogs include 14

. the tail has separate vertebrae unlike the fused urostyle or coccyx in modern frogs . the tibia and fibula bones are also separate , making it probable that

from the middle jurassic is slightly younger , about 155–170 million years old . the main evolutionary changes in this species involved the shortening of the body and the loss of the tail . the evolution of modern anura likely was complete by the jurassic period . since then , evolutionary changes in

evidence suggests they also inhabited antarctica in an earlier era when the climate was warmer .