phascolopsis gouldii is a species of unsegmented benthic marine worm , also known as a peanut worm or star worm .

it lives in burrows in muddy sand in shallow waters off north america .

phascolopsis gouldii

the mitochondrial genome of the sipunculid phascolopsis gouldii supports its association with annelida rather than mollusca .
the mitochondrial genome of the sipunculid phascolopsis gouldii supports its association with annelida rather than mollusca . - pubmed - ncbi
the crystal structures of phascolopsis gouldii wild type and l98y methemerythrins : structural and functional alterations of the o2 binding pocket .
the crystal structures of phascolopsis gouldii wild type and l98y methemerythrins : structural and functional alterations of the o2 binding pocket . - pubmed - ncbi
1450 phascolopsis gouldii ( ms ) virtual museum of natural history / marine biological laboratory send us an e - mail if you are interested in high resolution microct data of this organism . virtualmuseumofnaturalhistory @ urltoken
boore , j . l . and j . l . stanton . 2002 . the mitochondrial genome of the sipunculid phascolopsis gouldii supports its association with annelida rather than mollusca . molecular biology and evolution 19 : 127 - 137 .
( of phascolopsis gouldi pourtalès , 1851 ) integrated taxonomic information system ( itis ) . , available online at urltoken [ details ]
jeffrey l . boore , joseph l . staton ; the mitochondrial genome of the sipunculid phascolopsis gouldii supports its association with annelida rather than mollusca , molecular biology and evolution , volume 19 , issue 2 , 1 february 2002 , pages 127–137 , urltoken
fig . 2 . —abbreviated representation of the 7 , 470 nt determined of the mtdna of the sipunculid phascolopsis gouldii . center portions of each protein - or rrna - encoding gene have been replaced with a numeral indicating the number of omitted nucleotides . nucleotides forming stop codons , partial or complete ( see text ) , are marked with a caret
myohemerythrin from the sipunculid , phascolopsis gouldii : purification , properties and amino acid sequence .
long r . c . , zhang j . - h . , kurtz d . m . jr . , negri a . , tedeschi g . , bonomi f . biochim . biophys . acta 1122 : 136 - 142 ( 1992 ) [ pubmed ] [ europe pmc ] [ abstract ]
a leucine residue ' gates ' solvent but not o2 access to the binding pocket of phascolopsis gouldii hemerythrin .
farmer c . s . , kurtz d . m . jr . , phillips r . s . , ai j . , sanders - loehr j . j . biol . chem . 275 : 17043 - 17050 ( 2000 ) [ pubmed ] [ europe pmc ] [ abstract ]
the crystal structures of phascolopsis gouldii wild type and l98y methemerythrins : structural and functional alterations of the o2 binding pocket .
farmer c . s . , kurtz d . m . jr . , liu z . j . , wang b . c . , rose j . , ai j . , sanders - loehr j . j . biol . inorg . chem . 6 : 418 - 429 ( 2001 ) [ pubmed ] [ europe pmc ] [ abstract ]
( of phascolopsis gouldi pourtalès , 1851 ) trott , t . j . ( 2004 ) . cobscook bay inventory : a historical checklist of marine invertebrates spanning 162 years . northeastern naturalist . 11 , 261 - 324 . , available online at urltoken [ details ] available for editors [ request ]
moving p . gouldii to make it a sister taxon to an ( l . bleekeri , k . tunicata ) clade or to an ( ( l . bleekeri , k . tunicata ) , t . retusa ) clade increases tree lengths by seven steps in each case for this gene arrangement analysis .
( of phascolopsis gouldi pourtalès , 1851 ) brunel , p . ; bosse , l . ; lamarche , g . ( 1998 ) . catalogue of the marine invertebrates of the estuary and gulf of st . lawrence . canadian special publication of fisheries and aquatic sciences , 126 . 405 p . ( look up in imis ) [ details ] available for editors [ request ]
the primary structure of golfingia gouldii hemerythrin . oder of peptides in fragments produced by tryptic digestion of succinylated hemerythrin . complete amino acid sequence .
klippenstein g . l . , holleman j . w . , klotz i . m . biochemistry 7 : 3868 - 3878 ( 1968 ) [ pubmed ] [ europe pmc ] [ abstract ]
two other annelids , galathealinum brachiosum and helobdella robusta , were included in a separate analysis , since each lacks a complete sequence for cob ( boore and brown 2000 ) . their inclusion had no effect on the phylogenetic placement of p . gouldii , and these two taxa joined p . dumerilii and l . terrestris in a topology consistent with the results of the earlier study ( not shown ) .
mitochondrial genes often terminate with abbreviated stop codons , where a single t or ta is presumably completed by polyadenylation to a taa stop codon ( ojala , montoya , and attardi 1981 ) . this appears to be the case for three genes in p . gouldii mtdna . the least certain of these is for atp8 , since the next nucleotide ( a ) , otherwise part of trny , would complete the stop codon ( fig . 2 ) . if cob were to extend to the first complete stop codon , it would overlap trnp by 41 nt , and if nad6 were to do so , it would overlap cob by 20 nt . in each case , homologous genes of related animals are similar in predicted amino acid sequences to the abbreviated forms in p . gouldii and have no similarity to the overlapping extensions .
we report the sequence of about half ( 7 , 470 nt ) of the mtdna of the first representative of the phylum sipuncula , p . gouldii . we analyze genomic features in comparison with those of other animal mtdnas and compare both gene arrangement and inferred amino acid sequences to resolve the phylogenetic placement of this phylum . these comparisons strongly support the closer relationship of sipuncula with annelida to the exclusion of mollusca and other taxa .
overall , the 7 , 470 nt determined for p . gouldii mtdna are 63 . 1 % a + t , similar to the percentage found for the whole mtdna sequence of l . terrestris ( 61 . 6 % ; boore and brown 1995 ) . as is common among mtdnas , cg is the least frequent dinucleotide , occurring at only 53 % of expectation given the proportion of g and of c observed in this sequence .
live specimens of p . gouldii ( fisher 1950 ) were purchased from the marine biological laboratory at woods hole , mass . we isolated total dna from excised retractor muscles using a ctab method described in collins et al . ( 1996 ) . we also isolated total dna from the upper “nuclear” fraction of the cscl purification method for mitochondrial dna ( dowling et al . 1996 ) . the upper layer of that preparation contains both nuclear and nonsupercoiled ( i . e . , nicked ) mitochondrial dna .
most of the studied animal mitochondrial genomes contain one or more genes that initiate with some alternative to the standard atg . annelids appear as a notable exception : based on the few animals sampled , atg is used as the initiation codon for all , or nearly all , genes ( boore and brown 2000 ) . consistent with this and with the phylogenetic placement of sipuncula advanced here ( see below ) , each of the p . gouldii protein - encoding genes in this portion of the mtdna appears to start with atg ( the initiator of cox1 is not present in this sequence ) .
it is common for trna ( s ) in many mtdnas to lack a paired dhu arm and , in some cases , to have a six - member anticodon stem ( yokogawa et al . 1991 ) . there is potential for p . gouldii trna ( s2 ) to have such an extended anticodon stem and also to have a paired dhu arm with an unusual structure , with only one nucleotide separating the amino - acyl acceptor arm from the dhu arm . several other serine - specifying trnas have been identified in animal mtdnas that have this identical pairing potential ( boore and brown 2000 ) .
van der land , j . ( ed ) . ( 2008 ) . unesco - ioc register of marine organisms ( urmo ) . , available online at urltoken [ details ]
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j biol inorg chem . 2001 apr ; 6 ( 4 ) : 418 - 29 .
farmer cs 1 , kurtz dm jr , liu zj , wang bc , rose j , ai j , sanders - loehr j .
department of chemistry and center for metalloenzyme studies , university of georgia , athens , ga 30602 - 2556 , usa .
research support , u . s . gov ' t , p . h . s .
the adults are found on sand flats , just below low tide level . they are abundant at hadley harbor , near woods hole , mass . animals covered with a mass of sea - weed will remain in good condition , even after several hours of transportation . the sexes are separate , but similar in appearance .
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myohemerythrin is an oxygen - binding protein found in the retractor muscles of certain worms . the oxygen - binding site contains two iron atoms .
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hemerythrin is a respiratory protein in blood cells of certain marine worms . the oxygen - binding site in each chain contains two iron atoms .
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x - ray crystallography ( 1 . 80 angstroms ) in complex with iron , subunit .
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in addition to providing gene order characters for inferring phylogeny , this data set also allows comparison of sequences as an independent estimator of evolutionary relationships . furthermore , mitochondrial genome comparisons serve as a model system for genome evolution , where we can examine such factors as the following : what determines whether genes are all encoded by the same dna strand versus being distributed between the two strands ? how does mutation bias influence a + t - richness , dinucleotide frequency , codon usage , strand - skew between purines and pyrimidines , and amino acid composition of proteins ? how do the structures of trnas evolve ?
initially , primers designed to match generally conserved regions of the mtdna were used in the pcr to amplify four short ( 450–710 nt ) fragments from cox1 ( primers lco1490 and hco2198 ; folmer et al . 1994 ) , cox3 ( primers coiiif and coiiib ; boore and brown 2000 ) , cob ( primers cytbf and cytbr ; boore and brown 2000 ) , and rrnl ( primers 16sarl and 16sbrh ; palumbi 1996 ) . dna sequences obtained from these fragments were used to design oligonucleotides that were then employed in “long - pcr” ( barnes 1994 ) to amplify the portions of the mtdna spanning cox1 – cox3 , cox3 – cob , and cob – rrnl . reaction conditions and results , fragment purification , dna sequence determination and assembly , and gene identifications were as in boore and brown ( 2000 ) . all sequence was determined for both strands using synthetic oligonucleotides to “primer - walk” through the long - pcr - amplified fragments . this 7 , 470 - nt sequence was deposited in genbank under accession number af374337 .
leucine is inferred to be the most frequent amino acid , present 243 times in these six inferred amino acid sequences , followed by four amino acids in nearly equal numbers , each about half as common as leucine : alanine ( 124 ) , isoleucine ( 123 ) , phenylalanine ( 135 ) , and serine ( 125 ) . cysteine is the least frequently used amino acid , occurring only 14 times . these values are all similar to those for the amino acid usage of l . terrestris homologs ( see table 2 ) .
the dna strand shown in figure 2 is rich in pyrimidines ( c and t ) . one way to measure this is called skewness , an assessment of whether the g of gc pairs ( gc - skew , calculated as [ g − c ] / [ g + c ] ) and the a of at pairs ( at - skew , calculated as [ a − t ] / [ a + t ] ) are more commonly on the considered strand ( perna and kocher 1995 ) . gc - skew for the entire 7 , 470 nt is −0 . 239 , that for the protein - encoding portion only is −0 . 265 , and that for third codon positions ( see below ) , which , presumably , are more free to change and therefore better reflect mutational bias , is a remarkable −0 . 754 . at - skew values are much less biased : that for the entire region is −0 . 080 , that for the protein - encoding portion is −0 . 138 , and that for the third codon positions is −0 . 028 .
all genes are in a compact arrangement , with a total of only 13 noncoding nucleotides . these are distributed in six regions of 1–4 nt each . there is a cc between trny and trng , but otherwise , all are a or t . no genes are inferred to overlap except for the possibility of a single nucleotide shared between atp8 and trny ( see above ) .
there are 11 sequences identified with the potential for folding into trna - like structures ( fig . 3 ) . each has an anticodon matching exactly one trna in l . terrestris mtdna . each has a seven - member amino - acyl acceptor stem ( four with a single mismatch each ) and a five - member anticodon stem ( again , four with a single mismatch each ) . two have 5 nt in the extra arm , and all others have 4 nt . all but four have a immediately preceding the anticodon arm . for all except trna ( n ) and trna ( s2 ) , there are three to five nucleotide pairs in both the dhu and the tψc arms . all but four trnas have ta immediately preceding the dhu arm . the nucleotides preceding the anticodon are yt for all trnas , and the nucleotide following the anticodon is a for all but trna ( p ) .
discrepancies among the ml , mp , and nj analyses were in the placement of the brachiopod t . retusa and the relationship among the crustaceans d . pulex and artemia franciscana . in mp and nj trees , t . retusa was placed as sister to loligo bleekeri , albeit with less than 52 % bootstrap support in either case , whereas in the ml tree , t . retusa was placed as sister to a clade of the two mollusks ( 69 % of quartets sampled ) . the two crustaceans were paraphyletic in the mp analysis , albeit with poor ( < 50 % ) bootstrap support , but grouped together in both the ml and nj analyses . this association was found in 95 % of the quartets in ml and with 82 % of the bootstrap resamplings in nj .
our findings are in general agreement with several published studies ( winnepenninckx , backeljau , and de wachter 1995 [ parsimony - based analysis ] ; giribet et al . 2000 [ 18s - only tree ] ; regier and shultz 1998 ) in that sipunculans are closely associated with annelids to the exclusion of mollusks . other studies , or portions thereof , give results to the contrary . comparisons of 18s rdna sequences have placed sipuncula as basal and sister to an assemblage of worms and mollusks ( field et al . 1988 ; winnepenninckx , backeljau , and de wachter 1995 [ neighbor - joining analysis ] ; giribet et al . 2000 [ when combined with morphological characters ] ) or as sister to only one of two ectoprocts analyzed ( mackey et al . 1996 ) . the conflicts among these studies indicate that comparisons of short dna sequences lack the necessary resolving power at this level of relationship . furthermore , some of the associations between sipunculans and annelids found in these past studies were probably discounted based on the problematica associated with these data ( maley and marshall 1998 ) .
likewise , many characteristics that have been hypothesized to link sipunculans with mollusks , e . g . , developmental pattern , lack of segmentation , etc . , must be reevaluated . the presence of a “molluscan cross” may be ancestral in all eutrochozoans and subsequently lost in the annelids , or possibly is a convergent pattern that has independent origins in the two phyla . recent reports of segmental development in chitons ( jacobs et al . 2000 ) suggest that segmentation in some form may be present in all spiralians , so its loss in other mollusks and worm groups may not be surprising . certainly , the detailed examination of hypothesized synapomorphic larval and adult morphology in the sipunculans and molluscan groups ( scheltema 1993 , 1996 ) will need careful reconsideration to exclude convergence or oversight of annelid larval and adult homologs .
address for correspondence and reprints : jeffrey l . boore , doe joint genome institute and lawrence berkeley national laboratory , 2800 mitchell drive , walnut creek , california 94598 . jlboore @ urltoken .
fig . 5 . —phylogenetic tree based on gene arrangement comparisons using an adjacency matrix . this is the strict consensus of 30 equally parsimonious trees . the unusual placement of alligator mississippiensis is poorly supported ( see text )
thanks to inaki ruiz - trillo for assistance with the program tree - puzzle and to kevin helfenbein for helpful comments on the manuscript . this work was supported by nsf grants deb - 9807100 and deb - 9303301 . part of this work was performed under the auspices of the united states department of energy office of biological and environmental research by lawrence livermore national laboratory under contract de - ac03 - 76sf00098 .
the duplication / random loss model for gene rearrangement exemplified by mitochondrial genomes of deuterostome animals , pp . 133–147
d . sankoff and j . nadeau , eds . computational biology series , vol . 1 . comparative genomics . kluwer academic publishers , dordrecht , the netherlands
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études sur les types inférieurs de l ' embranchement des annelés . mémoire sur l’échiure de gaertner ( echiurus gaertnerii nob . )
memorie sulla storia e notomina degli animali senza vertebre del regno di napoli . vol . i fratelli fernandes , naples , italy
d . m . hillis , c . moritz , and b . k . mable , eds . molecular systematics . sinauer , sunderland , mass
m . e . rice and m . todorovic , eds . proceedings of the international symposium on the biology of the sipuncula and echiura . vol . 2 . naucno delo press , belgrade
p . harvey and l . partridge , eds . oxford surveys in evolutionary biology . oxford university press , oxford , england
histoire naturelle des animaux sans vertèbres , vol . 3 radiaires , vers , crustacés , insectes . verdière , paris
le t . h . , d . blair , t . agatsuma , et al . ( 14 co - authors )
mackey l . y . , b . winnepenninckx , r . de wachter , t . backeljau , p . emschermann , j . r . garey ,
a . giese and j . pearse , eds . reproduction of marine invertebrates , vol . 2 . entoprocts and lesser coelomates . academic press , new york
s . conway morris , j . d . george , r . gibson , and h . m . platt , eds . the origins and relationships of lower invertebrates . oxford university press , oxford , england
j . d . taylor , ed . origin and evolutionary radiation of the mollusca . oxford university press , oxford , england
paup * . phylogenetic analysis using parsimony ( * and other methods ) . version 4 sinauer , sunderland , mass
with special reference to the significance of the polar lobe . i , ii and iii . division chronology and development of the cell pattern in
n . h . verdonk , j . a . m . van den biggelar , and a . s . tompa , eds . the mollusca , vol . 3 . development . academic press , new york
yamazaki n . , r . ueshima , j . terrett , et al . ( 12 co - authors )
yokobori s . - i . , u . takuya , g . feldmaier - fuchs , s . pääbo , r . ueshima , a . kondow , k . nishikawa , k . watanabe ,
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according to cutler ( 1994 ) the unacknowledged type material appears to be in the museum of comparative zoology , harvard .
this species is only found on the atlantic coast of north america . this slender , smooth , light brown worm has been used as a source for teaching and research among the biological community . the deciduous hooks are present in juveniles but are absent in adults .
introvert about one - third of the trunk length , with deciduous hooks ( 26 - 40 µm tall ) , widely dispersed and arranged in ill - defined rings . present in juveniles ( in worms after 3 - 4 mm long ) but absent in adults . small papillae arranged in rings are present on the distal part of the introvert .
two pairs of retractor muscles , attached in the anterior half of the trunk . dorsal pair originating anteriorly to the ventral pair .
most common size is between 50 - 100 mm in trunk length , but it can be up to 150 mm ( cutler , 1994 ) .
atlantic coast of north america from florida to nova scotia . it is rare south of long island .
the breeding season in newport , road island begins in the middle of june and extends to middle of september ( gerould , 1907 ) .
for many years this species has been used as a teaching and research resource ( e . g . abercrombie and bagby , 1984 ; boore and staton , 2002 ; oglesby , 1982 ; vanin et al , 2006 ) .
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hemerythrin ( hr ) is a non - heme diiron oxygen transport protein found in four marine invertebrate phyla including priapulida , brachiopoda , sipunculida , and annelida , as well as in protozoa . myohemerythrin ( mhr ) , a hemerythrin homolog , is found in the muscle tissue of sipunculids as well as in polycheate and oligocheate annelids . in addition to oxygen transport , mhr proteins are involved in cadmium fixation and host anti - bacterial defense . hr and mhr proteins have the same
four alpha helix bundle
motif and active site structure . hr forms oligomers , the octameric form being most prevalent , while mhr is monomeric .
cdd / sparcle : functional classification of proteins via subfamily domain architectures .
department of biology , university of michigan . doe joint genome institute and lawrence berkeley national laboratory , walnut creek , california 94598 , usa . jlboore @ urltoken
research support , u . s . gov ' t , non - p . h . s .
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eernisse , d . j . 1998 . arthropod and annelid relationships re - examined . pages 43 - 56 in arthropod relationships ( r . a . fortey and r . h . thomas , eds . ) the systematics association and chapman and hall , london .
eernisse , d . j . , j . s . albert , f . e . anderson . 1992 . annelida and arthropoda are not sister taxa . a phylogenetic analysis of spiralian metazoan morphology . systematic biology 41 : 305 - 330 .
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garey , j . r . 2001 . ecdysozoa : the relationship between cycloneuralia and panarthropoda . zoologischer anzeiger 240 : 321 - 330 .
garey , j . r . and a . schmidt - rhaesa . 1998 . the essential role of
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phyla in molecular studies of animal evolution . american zoologist 38 : 907 - 917 .
garey , j . r . , m . krotec , d . r . nelson , and j . brooks . 1996 . molecular analysis supports a tardigrade - arthropod association . invertebrate biology 115 : 79 - 88 .
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giribet , g . and c . ribera . 1998 . the position of arthropods in the animal kingdom : a search for a reliable outgroup for internal arthropod phylogeny . molecular phylogenetics and evolution 9 : 481 - 488 .
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giribet , g . , d . l . distel , m . polz , w . sterrer and w . wheeler . 2000 . triploblastic relationships with emphasis on the acoelomates and the position of gnathostomulida , cycliophora , plathelminthes , and chaetognatha : a combined approach of 18s rdna sequences and morphology . systematic biology 49 : 539 - 562 .
giribet , g . , m . v . sørensen , p . funch , r . m . kristensen , and w . sterrer . 2004 . investigations into the phylogenetic position of micrognathozoa using four molecular loci . cladistics 20 : 1 - 13 .
giribet , g . , s . carranza , j . baguna , m . riutort and c . ribera . 1996 . first molecular evidence for the existence of a tardigrada plus arthropoda clade . molecular biology and evolution 13 : 76 - 84 .
halanych , k . 2004 . the new view of animal phylogeny . annual review of ecology , evolution , and systematics 35 : 229 - 256 .
halanych , k . m . , j . d . bacheller , a . m . a . aguinaldo , s . m . liva , d . m . hillis , and j . a . lake . 1995 . evidence from 18s ribosomal dna that the lophophorates are protostome animals . science 267 : 1641 - 1643 .
hanelt , b . , d . vanschyndel , c . m . adema , l . a . lewis , e . s . loker . 1996 . the phylogenetic position of rhopalura ophiocomae ( orthonectida ) based on 18s ribosomal dna sequence analysis . molecular biology and evolution 13 : 1187 - 1191 .
hausdorf , b . , m . helmkampf , a . meyer , a . witek , h . herlyn , i . bruchhaus , t . hankeln , t . h . struck , and b . lieb . 2007 . spiralian phylogenomics supports the resurrection of bryozoa comprising ectoprocta and entoprocta . molecular biology and evolution 24 ( 12 ) : 2723 - 2729 .
hejnol , a . and m . q . martindale . 2008 . acoel development supports a simple planula - like urbilaterian . philosophical transactions of the royal society series b 363 ( 1496 ) : 1493 - 1501 .
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helfenbein , k . g . , h . m . fourcade , r . g . vanjani , and j . l . boore . 2004 . the mitochondrial genome of paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protosotomes . proceedings of the national academy of sciences ( usa ) 101 : 10639 - 43 .
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irimia , m . , i . maeso , d . penny , j . garcia - fernandez , and s . w . roy . 2007 . rare coding sequence changes are consistent with ecdysozoa , not coelomata . molecular biology and evolution 24 : 1604 - 1607 .
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jenner , r . a . 2004 . towards a phylogeny of the metazoa : evaluating alternative phylogenetic positions of platyhelminthes , nemertea , and gnathostomulida , with a critical reappraisal of cladistic characters . contributions to zoology 73 ( 1 - 2 ) : 3 - 163 .
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mackey , l . y . , b . winnepenninckx , r . dewachter , t . backeljau , p . emschermann , and j . r . garey . 1996 . 18s rrna suggests that entoprocta are protostomes , unrelated to ectoprocta . journal of molecular evolution 42 : 552 - 559 .
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hypsopygia nostralis , the southern hayworm moth , is a species of snout moth in the genus hypsopygia .

it was described by guenée in 1854 .

it has a wide distribution and is found in most of south america , saint helena , puerto rico and in the southern united states , from texas to florida .

hypsopygia nostralis

species hypsopygia nostralis - southern hayworm moth - hodges # 5531 - bugguide . net
hypsopygia nostralis ( guenée , 1854 ) was a synonym of herculia sordidalis ( b . & mcd . , 1913 ) in the 1983 hodges checklist . this species was placed in the genus ocrasa in shaffer and solis ( 1995 ) and subsequently transferred to the genus hypsopygia in leraut ( 2006 ) .
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scholtens , b . g . & m . a . solis , 2015 . annotated check list of the pyraloidea ( lepidoptera ) of america north of mexico . zookeys , 535 : 1 - 136 .
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, missouri to texas ; west indies ; mexico to brazil ; asia ; st . helena .
dyar , h . g . 1917 . seven new pyralids from british guiana ( lepidoptera : pyralidae . insecutor inscitiae menstruus . p . 90 .
arthropods of florida and neighboring land areas : lepidoptera of florida j . b . heppner . 2003 . florida department of agriculture 17 ( 1 ) : 1 - 670 .
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guenée a . m . 1854 . histoire naturelle des insectes . viii . species général des lépidoptères . deltoites et pyralites . - — 8 : 1—448 .
ghesquière j . 1942a . lépidoptères microlépidoptères ( deuxième partie ) . - annales du musée du congo belge , zoologie [ 3 , arthropodes ] section 2 , catalogues raisonnés ( 3 ) 7 : 121—240 .
pinhey e . c . g . 1975 . moths of southern africa . descriptions and colour illustrations of 1183 species . - — : i—iv , 1—273 , pls . 1—63 .
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afrixalus fornasini is a species of frog in the hyperoliidae family and is native to africa .

its common name is fornasini 's spiny reed frog or the greater leaf-folding frog

afrixalus fornasini

encyclopedia of life . 2012 . afrixalus fornasini . encyclopedia of life . accessed 8 / 12 / 2012 at : urltoken
iucn ssc amphibian specialist group 2013 . afrixalus fornasini . the iucn red list of threatened species 2013 : e . t56063a3034378 . urltoken
fornasini ' s spiny reed frog ( channing , 2001 , amph . cent . s . afr . : 137 ) .
the skin of a . fornasini contains chemicals called tachykinins , which deter some mammalian predators ( channing and howell , 2006 ) .
fornasini ' s frog ( hewitt , 1937 , guide vert . fauna e . cape province , rept . amph . fishes : 113 ) .
unknown . it has been reported that a . fornasini preys on eggs and developing larvae of other frogs ( drewes and altig , 1996 ) .
pickersgill , m . , a . schiøtz , l . minter 2004 . afrixalus fornasini . in : iucn 2012 . iucn red list of threatened species . version 2012 . 2 . < www . iucnredlist . org > . accessed 3 / 5 / 2013 .
fuller , p . , 2018 , afrixalus fornasini ( bianconi , 1849 ) : u . s . geological survey , nonindigenous aquatic species database , gainesville , fl , urltoken revision date : 5 / 3 / 2018 , access date : 7 / 9 / 2018
table 1 . states with nonindigenous occurrences , the earliest and latest observations in each state , and the tally and names of hucs with observations† . names and dates are hyperlinked to their relevant specimen records . the list of references for all nonindigenous occurrences of afrixalus fornasini are found here .
afrixalus unicolor — pickersgill , 1996 , herptile , 21 : 81 ; pickersgill , 2007 , frog search : 469 .
schiøtz , a . 2004 . afrixalus equatorialis . 2006 iucn red list of threatened species . downloaded on 22 july 2007 .
this name is used with some uncertainty for one or several species of striped afrixalus from open vegetation in the forest belt and humid savanna in central africa . the name afrixalus brevipalmatus is used in cameroun , possibly for the same species ( for further discussion , see the account under a . fulvovittatus ) .
pickersgill , m . & , schiøtz , a . 2004 . afrixalus wittei . 2006 iucn red list of threatened species . downloaded on 22 july 2007 .
afrixalus fornasinii — guibé , 1948 , bull . mus . natl . hist . nat . paris , ser . 2 , 20 : 500 , by implication .
afrixalus unicolor — guibé , 1948 , bull . mus . natl . hist . nat . paris , ser . 2 , 20 : 500 , by implication .
schiøtz , a . & rödel , m . - o . 2004 . afrixalus weidholzi . 2006 iucn red list of threatened species . downloaded on 28 november 2010
fornasini ' s spiny reed frog ( lambiris , 1990
1989
, monogr . mus . reg . sci . nat . torino , 10 : 159 ; channing and howell , 2006 , amph . e . afr . : 134 ) .
afrixalus fornasinii — schiøtz , 1974 , vidensk . medd . dansk naturhist . foren . , 137 : 12 . rejected by schiøtz , 1999 , treefrogs afr . : 67 .
a large forest afrixalus ( males 25–34 mm , females 30–38 mm ) from r . d . congo . the pattern is rather constant with an almost square dark dorsal spot .
this form has previously been regarded as a subspecies of three other afrixalus species ( a . fornasini , a . dorsalis , a . congicus ) , but its status as a full species was established by laurent ( 1982 ) and schiøtz ( 1982 ) . in habitat preference and morphology it is quite similar to the west african a . nigeriensis but it lacks the dark lumbar band which is almost always present in a . nigeriensis .
pickersgill ( 1996 , 2007 ) recognized afrixalus unicolor ( boettger , 1913 ) as a distinct species ; thus afrixalus fornasinii from south africa to southern tanzania ( including the area around lake malawi ) and a . unicolor from southern tanzania ( including zanzibar and pemba islands ) to kenya . however , here we follow schiøtz ( 1999 ) in considering these to constitute a single species .
schiøtz , a . , amiet , j . - l . & fretey , t . 2004 . afrixalus laevis . 2006 iucn red list of threatened species . downloaded on 22 july 2007 .
a large forest afrixalus ( males 25 - 34 mm , females 30 - 38 mm ) from r . d . congo . the pattern is rather constant with an almost square dark dorsal spot .
fornasini ' s banana frog ( broadley , 1973 , j . herpetol . assoc . afr . , 10 : 23 ; passmore and carruthers , 1978 , j . herpetol . assoc . afr . , 19 : 7 ; passmore and carruthers , 1979 , s . afr . frogs : 238 ) .
a rather large ( females up to 33 mm ) afrixalus from lowland savanna in south - eastern r . d . congo and adjacent parts of angola , tanzania and zambia . dorsum with three regular light stripes , the dorsal one not meeting the two lateral ones on the head . voice : a typical afrixalus - voice , starting with a creaking sound and continuing with a number of clicks with an indistinct frequency - intensity maximum at 2500–3000 cps .
it is confined to forest interior , and does not occur in altered habitats . the eggs are deposited on a leaf ( which is not folded ) near flowing water ( unusually for afrixalus species which usually breed in still water ) .
laurent , r . f . ( 1982 ) . ' ' le genre afrixalus laurent ( hyperoliidae ) en afrique centrale . ' ' koninklijk museum voor midden - afrika tervuren belgie annalen zoologische wetenschappen , ( 235 ) , 1 - 58 .
afrixalus fornasinii unicolor — laurent , 1951 , ann . soc . r . zool . belg . , 82 : 24 . status rejected by loveridge , 1955 , j . e . afr . nat . hist . soc . , 22 : 195 .
largen , m . & schiøtz , a . ( 2004 ) . afrixalus enseticola . 2006 . iucn red list of threatened species . iucn 2006 . urltoken . retrieved on 5 mar 2007 . database entry includes a range map and justification for why this species is vulnerable
schiøtz , a . , pickersgill , m . , largen , m . , amiet , j . - l . , burger , m . & lötters , s . 2004 . afrixalus quadrivittatus . 2006 iucn red list of threatened species . downloaded on 22 july 2007 .
the ethiopian banana frog ( afrixalus enseticola ) is a small frog that is endemic to ethiopia . they live in altitudes of 1700 - 2750 meters on both sides of the great rift valley in the ethiopian highlands . it is classified as
vulnerable
on the iucn red list ( 2004 ) due to decline of forested habitat in the highlands .
afrixalus fornasinii fornasinii — guibé , 1948 , bull . mus . natl . hist . nat . paris , ser . 2 , 20 : 500 , by implication ; laurent , 1951 , ann . soc . r . zool . belg . , 82 : 24 ; loveridge , 1953 , bull . mus . comp . zool . , 110 : 345 .
euchnemis fornasini bianconi , 1849
1848
, nuovi ann . sci . nat . , bologna , ser . 2 , 10 : 107 . syntypes : mzub 100174 , 100174a , and 100064 , according to bonfitto , 1991 , boll . mus . reg . sci . nat . torino , 9 : 364 ; type locality :
mozambico
; restricted to
inhambane , mozambico
, by bonfitto , 1991 , boll . mus . reg . sci . nat . torino , 9 : 362 . ( only a single i in original orthography—drf . )
this species is known from eastern angola , southeastern democratic republic of congo , most of zambia , and southwestern tanzania adjacent to the zambian border . a record from mbega in the kilombero valley of southeastern tanzania probably refers to afrixalus quadrivittatus ( m . pickersgill pers comm . ) . a record from tulawaka in northwestern tanzania , and records from duque de braganca and cabiri in northwestern angola ( ruas 1991 ) require confirmation .
afrixalus fornasinii lives in the savanna , possibly preferring dense savanna with larger bushes and trees . it is found in the coastal lowlands , but can be found up to 1 , 300 m in elevation . to the south , it is confined to coastal tropical lowlands of south africa . moist savannah , bush land , grassland , and dry forest are inhabited . breeding occurs in permanent and semi - permanent pools , and marshes with reeds and sedges . ( pickersgill et al . 2004 ; amphibiaweb , 2012 ) .
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frost , d . r . 2013 . amphibian species of the world : an online reference . version 5 . 6 ( 9 january 2013 ) . electronic database . american museum of natural history , new york , usa . available at : urltoken .
justification : listed as least concern in view of its wide distribution , tolerance of a broad range of habitats and its presumed large population .
this species ranges from coastal kenya southward through eastern and southern tanzania , malawi , mozambique and eastern zimbabwe to coastal kwazulu - natal in south africa . though its presence is uncertain in swaziland , this area is still mapped on the species distribution . it is a lowland species generally occurring below 300 m , though it has been recorded up to 1 , 300 m asl .
it is a species of moist savannah , bush land , grassland , and dry forest , with a possible preference for dense savannah . it breeds in permanent and semi - permanent pools , dams , vleis and marshes with reeds and sedges , tending to favour permanent waterbodies . the eggs are laid on emergent vegetation , particularly on leaf blades of cyperus sedges .
in the south of its range it is threatened by sugar cane farming , urbanization , drainage of breeding sites , afforestation with pines , and the spread of eucalyptus ( drying up breeding sites ) . chemical spraying to control mosquitoes might impact some populations . however , overall it is not threatened . it is sometimes found in the international pet trade but at levels that do not currently constitute a major threat .
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gland , switzerland , 5 july 2018 ( iucn ) – australia’s unique reptiles – including lizards and snakes – face severe threats from invasive species and climate change , with 7 % of th . . .
the value of medicinal and aromatic plant trade has increased three - fold in the past 20 years , but traditional harvesting practices are being replaced by less sustainable alternatives . . . .
a recently released iucn technical brief recommends increasing investments in sustainable land management practices , as well as better cooperation between agriculturalists and conservationists to conse . . .
from the savannas of eastern and southern africa ; dark with broad light silverish dorsolateral bands from tip of snout to anus . the bands meet posteriorly , not anteriorly . about half the specimens in northern populations ( tanzania and kenya ) have the entire dorsum silverish white . upper side of tibia white .
the males have numerous large conspicuous black - tipped asperities on head , back , dorsal surfaces of limbs and around the anus . the females have smaller asperities .
( boettger 1913 ) , without arguing for its specific distinctness . drewes & altig ( 1996 ) have reported that
sound , but loud , slow and low - pitched as can be expected from such a large animal . the voice has been compared with the stuttering of a minute machine - gun by wager . there is a slow , creaking initial sound followed by a series of figures at a rate of 5–10 per second and with a frequency - intensity maximum at about 2500 cps .
is a characteristic element in the savanna , possibly preferring rather dense savanna with larger bushes and trees . it is found in the coastal lowlands , but also on localities up to 1300 metres in malawi ( stewart 1967 ) . to the south it is again confined to the coastal tropical lowlands of south africa .
development . – the eggs are glued in a leaf , the tadpoles are large , up to 65 mm ( 17 + 48 ) and streamlined with a terminal mouth . tooth formula 0 / 1 .
drewes , r . c . and altig , r . ( 1996 ) . ' ' anuran egg predation and heterocanibalism in a breeding community of east african frogs . ' '
steward , m . m . and wilson , v . j . ( 1966 ) . ' ' herpetofauna of the nyika plateau ( malawi and zambia ) . ' '
> 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 , darrel r . 2004 . amphibian species of the world : an online reference . version 3 . 0 ( 22 august , 2004 ) . electronic database accessible at urltoken american museum of natural history , new york , usa
amphibian species of the world : an online reference v5 . 3 , database ( version 5 . 3 )
frost , darrel r . 2009 . amphibian species of the world : an online reference . version 5 . 3 ( 12 february , 2009 ) . electronic database accessible at urltoken american museum of natural history , new york , usa
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
* will not find nomina inquirenda ; use basic search ( above ) for that purpose .
will find all uses of
hyl . . .
anywhere in a record : e . g . , hylarana , hyla , hylidae , hylinae , hylaedactyla .
will find all uses of
. . . hyla
anywhere in a record : e . g . , hyla , hylidae , plectrohyla , ptychadena hylaea , adenomera hylaedactyla
will find all records that contain stand - alone uses of hyla : e . g . , hyla , hyla arenicolor
interprets this as
lithobates or pipiens
so will find the union of all records that contain either
lithobates
or
pipiens
: e . g . , lithobates omiltemanus , hylorana pipiens
interprets this as
lithobates and pipiens
so will return all records that have the character string
lithobates pipiens
anywhere within a record : e . g . , all members of the lithobates pipiens complex .
hyperolius bivittatus peters , 1854 , ber . bekannt . verhandl . k . preuss . akad . wiss . berlin , 1854 : 627 . syntypes : zmb 4529 ( 7 specimens ) , according to bauer , günther , and klipfel , 1995 , in bauer et al . ( eds . ) , herpetol . contr . w . c . h . peters : 44 . type locality :
terra boror
, mozambique ; rendered as
tette
and
mozambique
by ahl , 1930 , sitzungsber . ges . naturforsch . freunde berlin , 1930 : 101 . synonymy by günther , 1859
1858
, cat . batr . sal . coll . brit . mus . : 87 ; peters , 1882 , naturwiss . reise mossambique , zool . 3 : 160 ; boulenger , 1882 , cat . batr . sal . coll . brit . mus . , ed . 2 : 130 .
hyperolius fornasinii — günther , 1859
1858
, cat . batr . sal . coll . brit . mus . : 87 . incorrect subsequent spelling .
rappia fornasinii — günther , 1869
1868
, proc . zool . soc . london , 1868 : 479 .
megalixalus fornasinii — peters , 1882 , naturwiss . reise mossambique , zool . 3 : 160 ; boulenger , 1882 , cat . batr . sal . coll . brit . mus . , ed . 2 : 130 .
megalixalus fornasinii var . unicolor boettger , 1913 , in voeltzkow ( ed . ) , reise ost - afr . , 3 ( 4 ) : 349 . syntypes : smf , mcz 15429 ( on exchange from smf , according to barbour and loveridge , 1946 , bull . mus . comp . zool . , 96 : 153 ) ; smf 7274 designated lectotype by mertens , 1967 , senckenb . biol . , 48 ( a ) : 48 . type locality : pemba island , tanzania .
megalixalus fornasinii — nieden , 1915 , mitt . zool . mus . berlin , 7 : 372 . loveridge , 1942 , bull . mus . comp . zool . , 91 : 395 .
megalixalus loveridgii procter , 1920 , proc . zool . soc . london , 1920 : 418 . holotype : bmnh 1947 . 2 . 9 . 83 , according to museum records . type locality :
morogoro
, kenya . synonymy by parker , 1931
1930
, proc . zool . soc . london , 1930 : 900 .
megalixalus unicolor — ahl , 1930 , sitzungsber . ges . naturforsch . freunde berlin , 1930 : 101 ; ahl , 1931 , das tierreich , 55 : 443 .
megalixalus fornasinii loveridgei — parker , 1931
1930
, proc . zool . soc . london , 1930 : 900 . suggested as a likely arrangement . status rejected by loveridge , 1955 , j . e . afr . nat . hist . soc . , 22 : 195 .
spiny reed frog ( wager , 1965 , frogs s . afr . : 192 ) .
greater leaf - folding frog ( passmore and carruthers , 1978 , j . herpetol . assoc . afr . , 19 : 7 ; passmore and carruthers , 1979 , s . afr . frogs : 238 ; pickersgill and bishop , 2004 , in minter et al . ( eds . ) , atlas frogs s . afr . lesotho and swaziland : 127 ; du preez and carruthers , 2009 , compl . guide frogs s . afr . : 230 ) .
brown and white spiny reed frog ( passmore and carruthers , 1978 , j . herpetol . assoc . afr . , 19 : 7 ; passmore and carruthers , 1979 , s . afr . frogs : 238 ) .
brown - striped spiny reed frog ( ananjeva , borkin , darevsky , and orlov , 1988 , dict . amph . rept . five languages : 70 ) .
zaire banana frog ( frank and ramus , 1995 , compl . guide scient . common names amph . rept . world : 65 ) .
mozambique banana frog ( channing and howell , 2006 , amph . e . afr . : 134 [ alternative name ] ) .
silver - banded banana frog ( channing and howell , 2006 , amph . e . afr . : 134 [ alternative name ] ) .
savannas of coastal southern kenya southward through eastern and southern tanzania , malawi , mozambique , and extreme eastern zimbabwe to extreme southern coastal kwazulu - natal , rep . south africa .
please note : these links will take you to external websites not affiliated with the american museum of natural history . we are not responsible for their content .
for access to available specimen data for this species , from over 350 scientific collections , go to vertnet .
copyright © 1998 - 2018 , darrel frost and the american museum of natural history . all rights reserved .
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if we can discover the meaning in the trilling of a frog , perhaps we may understand why it is for us not merely noise but a song of poetry and emotion .  - - -
thanks for that terry . i only got them a week ago - haven ' t heard any calling yet and they ' re quite secretive - i ' ll have to wrestle them out of their tank and see if i can spot any differences ! there ' s been an influx of all sorts of species from madagascar and africa in the uk recently ( boophis species for example ) , it ' s mighty tempting to get the old mastercard out !
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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 .
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it would not be a surprise if further studies show that a . equatorialis and a . nigeriensis are subspecies of one species but , strangely enough , a vicariant to these forms ( from dense forest habitats ) is not known from cameroun .
this account was taken from
treefrogs of africa
by arne schiøtz with kind permission from edition chimaira publishers , frankfurt am main .
schiøtz , a . ( 1999 ) . treefrogs of africa . edition chimaira , frankfurt am main .
schiã¸tz , a . ( 1999 ) . treefrogs of africa . edition chimaira , frankfurt am main .
this species is known from the central parts of the congo basin in the democratic republic of congo and southeastern cameroon ( amiet 2009 ) . it probably has a wider distribution than the few records suggest . it presumably occurs in congo and central african republic , and these areas are included in the species ' distribution on the map , however so far there are no records from these countries .
south - eastern r . d . congo , thus probably also r . congo . forest in central lowlands .
it is a species of lowland rainforest , and is unlikely to be tolerant of alteration to its habitat . its breeding is unknown , but it presumably lays its eggs on leaves above water , into which the tadpoles fall and develop .
listed as least concern in view of its wide distribution and its presumed large population .
the voice consists of an initial sound and a small number of identical figures with a frequency - intensity maximum at 2000 - 2500 cps . and a rate of 15 - 20 per second .
most of its habitat remains intact , but shifting agriculture might impact local population .
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this species ranges from southeastern nigeria to western ethiopia , through cameroon , chad , central african republic and south sudan . then south to southern tanzania and southern and western democratic republic of congo , including : equatorial guinea , gabon , congo , uganda , rwanda and burundi . it is presumed to occur in the cabinda enclave of angola , and this is included in the species ' distribution on the map , however there are no records from this area yet . it probably also occurs in northern angola , zambia and malawi .
this species is found in central africa . it inhabits open vegetation , in humid savanna and the forest belt . since both the taxonomy and nomenclature are unsettled , it is difficult to give an exact distribution with confidence .
it occurs widely in moist and dry savannah , and degraded former forest . the eggs are deposited on folded leaves above water , and the tadpoles fall into ponds where they develop .
listed as least concern in view of its wide distribution , tolerance of a broad range of habitats and its presumed large population .
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a typical savanna species , found infrequently both in the humid , rather dense guinea woodland and the open dry savanna . known from a few localities widely scattered over west africa as far east as garamba n . p . and ituri in eastern r . d . congo .
this species is known from isolated records from gambia and southern mali , east to northeastern democratic republic of congo ( on the sudan border ) in savannas . there do not appear to be records from guinea - bissau , guinea , liberia , burkina faso , chad or sudan , but it is likely to occur in these countries . the small number of records over much of its range probably reflects the lack of herpetological work in many parts of west and central africa .
it is a species of dense moist , and open dry , savannas . it is probably tolerant , at least to some extent , of habitat alteration . it breeds in temporary ponds , eggs can be found above water .
although this species is known mainly from isolated records from a large area , it is listed as least concern in view of its wide distribution and its presumed large population .
development : small batches of about 10 unpigmented eggs are placed in transversally folded grass leaves , glued together by the jelly . the tooth formula of the tadpole is 0 / 1 . newly metamorphosed frogs have a length of 10 , 5 mm .
overgrazing might impact local populations , but it is unlikely to be significantly threatened .
in its habitat , but it is quite common in localized areas , somewhat tolerant of habitat disturbance , and not considered very threatened .
this species is endemic to high - altitude areas of southern ethiopia occurring on both sides of the rift valley at altitudes between 1 , 700 and 2 , 750 m asl . recent surveys have confirmed its presence in previously known localities bore and bonga ; however , intervening areas in the southwest and east still need to be surveyed ( a . mengistu and s . loader pers . comms . june 2012 ) . it is estimated to occur in six threat - defined locations ( a . mengistu and s . loader pers . comm . june 2012 ) . taking its range as a proxy , its extent of occurrence ( eoo ) is estimated to be
; however , it should be noted that there is a level of uncertainty associated with the edges of its range .
open forest in south - western ethiopia at altitudes of about 1800 metres and above .
it is found in montane forest at high altitudes , and rarely in montane grassland after forest clearance , typically breeding amongst herbaceous vegetation surrounding pools in forest clearings . it can survive in plantations and rural gardens suggesting tolerance to moderate disturbance ( a . mengistu and s . loader pers . comms . june 2012 ) . for breeding it requires emergent vegetation in marshy pools , at least some of which are likely to become dry for part of the year .
, it is considered to occur in six threat - defined locations , and there is continuing decline in the extent and quality of its forest habitat in the ethiopian highlands due to a variety of threats .
it was locally common within its relatively restricted range at the time when the specimens were collected ( 1971 - 1975 ) ( largen 2001 ) . recent collections in 2009 have confirmed that this species is still locally abundant ( a . mengistu and s . loader pers . comms . june 2012 ) . its population is not considered to be severely fragmented .
the major threat is ongoing habitat loss due to selective logging , human settlement , and small - scale and large - scale agricultural encroachment ( a . mengistu and s . loader pers . comms . june 2012 ) . chytrid fungus occurs in high prevalence in amphibians in highland ethiopia and has been detected on this species , although its impact is not known ( gower et al . 2012 ) .
it is known to occur in the bale mountains national park , although area management is very limited and enforcement is required ( a . mengistu and s . loader pers . comm . june 2012 ) . additional actions needed include the protection of remaining montane forest habitats from subsistence exploitation . more information is needed on this species ' distribution , population status , natural history and the potential effects of chytrid fungus on its population .
an adult ethiopian banana frog only reaches a length of 20 - 28 mm .
a small species ( males 20–23 mm , females 23–25 mm ) ranging from cameroun to western uganda . head large with protruding eyes . posterior part of dorsum translucent , anterior part dense yellowish with a varying pattern in brown . the only constant pattern is a brown canthal stripe continuing behind the eye . posterior part of body and limbs translucent . gular flap very small . by day the ground colour is white or silverish , by night yellow .
this species occurs in southwestern and southern cameroon , south to northeastern gabon ( at m ' passa ) east to eastern democratic republic of congo and southwestern uganda . it is also recorded from the island bioko ( equatorial guinea ) ( r . c . drewes pers . comm . ) . there do not appear to be records from central african republic , congo , though it presumably occurs in these countries and they are included on the species distribution on the map . it is also likely to occur in rwanda and equatorial guinea ( mainland ) .
a forest form known from cameroun , eastern r . d . congo and south - western uganda , as well as burundi , liberia , and rwanda .
listed as least concern in view of its wide distribution and presumed large population .
it is reasonably common in suitable habitat in cameroon , but it is rare in kibale forest , uganda , where it is on the edge of its range .
the voice consists of single clacks with a frequency - intensity maximum at about 3000 cps . the eggs are deposited near flowing water . they are placed on a leaf which is not glued together . the tadpoles are flatter than the other members of the genus . tooth formula 0 / 0 .
local populations are no doubt affected by forest loss as a result of agricultural encroachment , expanding human settlements , and removal of wood .
it is found in cameroon , democratic republic of the congo , equatorial guinea , gabon , uganda , possibly central african republic , possibly republic of the congo , possibly nigeria , and possibly rwanda . its natural habitats are subtropical or tropical moist lowland forests and rivers .
lowland savanna in south - eastern r . d . congo and adjacent parts of angola , tanzania and zambia .
it is a species of moist savannah and shrubland that is probably tolerant of some alteration of its habitat . it is typically associated with flooded grassland , small marshes , and temporary pools , in which it breeds .
sujeevan ratnasingham , paul d . n . hebert , barcode of life data systems ( bold )
its natural habitats are moist savanna , subtropical or tropical moist shrubland , subtropical or tropical seasonally wet or flooded lowland grassland , swamps , intermittent freshwater marshes , arable land , pastureland , rural gardens , ponds , and canals and ditches .
southeastern africa , kenya , malawi , mozambique , south africa , tanzania , and the united republic of zimbabwe ( krysko et al . 2011 ) .
amphibiaweb : information on amphibian biology and conservation . 2012 . berkeley , california : amphibiaweb . available from : urltoken accessed 1 / 1 - 13 / 2012 .
channing , a . and k . howell . 2006 . amphibians of east africa . cornell university press , ithaca , ny . 418pp .
drewes , r . c . and r . altig . 1996 . anuran egg predation and heterocanibalism in a breeding community of east african frogs . tropical zoology 9 ( 2 ) : 333 - 347 .
harper , e . b . g . j . measey , d . a . patrick , m . menegon , and j . r . vonesh . 2010 . field guide to amphibians of the eastern arc mountains and coastal forests of tansania and kenya . camerapix publishers international , nairobi , kenya .
krysko , k . l . and 12 others . 2011 . verified non - indigenous amphibians and reptiles in florida from 1863 through 2010 : outlining the invasion process and identifying invasion pathways and stages . zootaxa 3028 : 1 - 64 .
this information is preliminary or provisional and is subject to revision . it is being provided to meet the need for timely best science . the information has not received final approval by the u . s . geological survey ( usgs ) and is provided on the condition that neither the usgs nor the u . s . government shall be held liable for any damages resulting from the authorized or unauthorized use of the information .
the data represented on this site vary in accuracy , scale , completeness , extent of coverage and origin . it is the user ' s responsibility to use these data consistent with their intended purpose and within stated limitations . we highly recommend reviewing metadata files prior to interpreting these data .
citation information : u . s . geological survey . [ 2018 ] . nonindigenous aquatic species database . gainesville , florida . accessed [ 7 / 9 / 2018 ] .
contact us if you are using data from this site for a publication to make sure the data are being used appropriately and for potential co - authorship if warranted . for queries involving fish , please contact pam fuller . for queries involving invertebrates , contact amy benson .
from southern tanzania ( including zanzibar and pemba islands ) to kenya . however , here we follow schiøtz ( 1999 ) in considering these to constitute a single species .
the iucn red list of threatened species . version 2018 - 1 . < urltoken > . downloaded on 09 july 2018 .
listed as data deficient in view of continuing doubts as to its taxonomic validity .
if it is a valid species , it is possibly extinct following the very extensive habitat destruction that has taken place around the city of douala . there have been no records since its discovery , despite extensive herpetological work in the area .