colomesus is a genus of pufferfishes confined to tropical south america .

apart from differences in size , the two species are superficially similar , being green above , white below , and patterned with black transverse bands across the dorsal surface .

c. asellus is commonly found in the aquarium trade while c. psittacus due to its size and more specialized requirements is not found as often .

colomesus

the analyses included the following taxa : tetraodon nigroviridis , tetraodon biocellatus , sphoeroides testudineus , lagocephalus laevigatus , colomesus asellus , colomesus psittacus , and the freshwater colomesus from the tocantins drainage .
a ) colomesus tocantinensis nov . sp . – tocantins ( holotype pnt . uerj . 405 highlighted in white ) ; b ) colomesus asellus – iquitos ; c ) colomesus asellus – belém .
colomesus psittacus is a component of the marine aquarium trade , however there no indications at present time of declines from harvesting .
there are no known species - specific conservation measures in place for colomesus psittacus however its distribution overlaps with several marine protected areas .
colomesus tocantinensis nov . sp . urn : lsid : zoobank . org : act : 9b8accb5 - ff55 - 4514 - 901b - 6366fb6ea307
the hydrodynamic trails of lepomis gibbosus ( centrarchidae ) , colomesus psittacus ( tetraodontidae ) and thysochromis ansorgii ( cichlidae ) investigated with scanning particle image velocimetry .
the hydrodynamic trails of lepomis gibbosus ( centrarchidae ) , colomesus psittacus ( tetraodontidae ) and thysochromis ansorgii ( cichlidae ) investigate . . . - pubmed - ncbi
here we used both morphological and molecular methodologies in an integrative taxonomical approach to investigate the diversity of the amazonian freshwater pufferfishes of the genus colomesus based on specimens collected from three distinct populations from both brazil and peru . additionally , we describe a new colomesus species from the upper tocantins drainage based on both morphological and molecular data .
the neighbor - joining ( nj ) and maximum - likelihood ( ml ) result trees are presented in figures 2 and 3 , respectively . the genus colomesus was recovered as monophyletic inside the group formed by the sampled sphoeroides species , in except for sphoeroides pachygaster . lagocephalus was recovered in a basal phylogenetic position in relation to sphoeroides and colomesus , therefore corroborating recent results such as those presented by [ 43 ] – [ 45 ] .
based on a comprehensive analysis including both morphological and molecular methodologies using the cytochrome c oxidase i gene , we were able to discuss aspects of the phylogeny and phylogeography of the south american freshwater pufferfishes of the genus colomesus .
the use of molecular systematic techniques together with morphological methodologies confirmed the identification of a new cryptic pufferfish species from the upper tocantins drainage , colomesus tocantinensis nov . sp . morphological features such as the color pattern , the absence of dermal flaps across the chin , the distinct ‘inverted v’ opercle shape , and the caudal peduncle morphology , all support the description of colomesus tocantinensis nov . sp . , as a new pufferfish species from the south american freshwater drainages .
tyler , j . c . , 1964 - proceedings of the academy of natural sciences of philadelphia 116 : 119 - 148 a diagnosis of the two species of south american puffer fishes ( tetraodontidae , plectognathi ) of the genus colomesus .
the color pattern of colomesus tocantinensis nov . sp . is essentially the same as that of colomesus asellus , with five transverse dark bars across the dorsal region of the body . a dark blotch on the underside of the caudal peduncle , which is a state used by [ 46 ] to diagnose colomesus asellus , is present or absent , being vestigial to unobservable or absent in several specimens . the interspaces between the dark bars are light yellow , with gradually decreasing pigmentation and becoming white in the ventral region ( figure 6 ) . however , the light yellow to pale pattern presented by c . tocantinensis nov . sp . clearly contrasts with the gold - yellow pattern present in specimens from iquitos and belém .
colomesus was recovered deep inside the group formed by the remaining sphoeroides species , therefore suggesting sphoeroides as paraphyletic , with s . pachygaster being recovered as basal in relation to all the remaining sphoeroides species in all the analyses . additionally , colomesus was also recovered as the sister - taxa of the group formed by the species sphoeroides nephelus , s . tyleri , and s . greeleyi in the nj result , although it was recovered as the sister - taxa of s . greeleyi in the ml results .
deep sequence divergence was observed regarding the freshwater colomesus from the tocantins drainage ( figure 5 ) . the mean sequence divergence of the specimens from both belém and iquitos was estimated at 1 . 079 % , while the tocantins distances ranged from 1 . 955 % to 3 . 063 % , with a mean distance of 2 . 166 % . the observed sequence divergence values together with the congruence observed from both molecular and morphological phylogenetic approaches used here suggest the existence of an overlooked species within the genus colomesus .
coi amplicons were obtained from all the specimens included in the analyses . the obtained sequences clearly identified both previous accepted colomesus species ( c . asellus and c . psittacus ) , therefore being in accordance with the previous morphological diagnose presented by [ 46 ] .
the neighbor - joining ( nj ) and maximum - likelihood ( ml ) trees that encompass the genera triodon , diodon , chilomycterus , lagocephalus , tetraodon , takifugu , sphoeroides , and colomesus , were constructed using the mega 5 . 06 software [ 40 ] .
our molecular results based on the coi marker agrees with the recent results such as [ 43 ] – [ 45 ] , and suggest that the genus sphoeroides should be revised , mainly regarding the phylogenetic position recovered for the genus colomesus , deeply nested within the sphoeroides tree , and the basal position recovered for s . pachygaster . we plan further investigations along these lines to reconcile any conflicts between these molecular hypotheses presented herein and morphologically based interpretations [ 47 ] of the phylogeny of the taxa of colomesus , sphoeroides , and lagocephalus .
citation : amaral crl , brito pm , silva da , carvalho ef ( 2013 ) a new cryptic species of south american freshwater pufferfish of the genus colomesus ( tetraodontidae ) , based on both morphology and dna data . plos one 8 ( 9 ) : e74397 . urltoken
araujo - lima , c . a . r . m . , d . savastano , and l . cardeliquio jordão , 1994 - revue d ' hydrobiologie tropicale 27 ( 1 ) : 33 - 38 drift of colomesus asellus ( teleostei : tetraodontidae ) larvae in the amazon river .
krumme , u . , h . keuthen , u . saint - paul , and w . villwock , 2007 - brazilian journal of biology 67 ( 3 ) : 383 - 392 contribution to the feeding ecology of the banded puffer fish colomesus psittacus ( tetraodontidae ) in north brazilian mangrove creeks .
specimens of colomesus asellus were collected from three distinct populations with about 2200 km of mean distance separating them . the collection localities were ilha do mosqueiro , belém , brazil ; upper tocantins river - porto nacional , tocantins , brazil ; and nanay river - iquitos , peru ( figure 1 ) .
amazon puffer fish aka colomesus asellus are awesome fish . they are generally considered as peaceful and get to about 4 inches and can live in planted tanks with other fish . if you ' ve enjoyed this video and haven ' t already subscribed to our channel , please click that subscribe button now .
amaral , c . r . l . , p . m . brito , d . a . silva and e . f . carvalho , 2013 - plos one 8 ( 9 ) : 1 - 15 a new cryptic species of south american freshwater pufferfish of the genus colomesus ( tetraodontidae ) , based on both morphology and dna data .
within the genus colomesus , it can be immediately identified from congeners by its larger adult size , possession of 17 - 19 ( vs . 13 - 16 ) pectoral - fin rays , presence of 6 ( vs . 5 ) transverse dark bands dorsally on the body , and predominantly brackish , coastal ( vs . freshwater , fluvial ) ecology .
colomesus : from the ancient greek χωλóς ( cholós ) , meaning ‘physically defective , crippled’ , and μέσος  ( mésos ) , meaning ‘middle’ , presumably in reference to the frontal bones being narrowed , not connected to the orbit , and with the elongated postfrontals connected to the prefrontals ( see gill 1884 , also note misspelling of χωλóς as κολoς ) .
oliveira , j . s . , s . c . rego fernandes , c . a . schwartz , c . bloch jr . , j . a . taquita melo , o . r . pires jr . , and j . c . de freitas , 2006 - toxicon 48 ( 1 ) : 55 - 63 toxicity and toxin identification in colomesus asellus , an amazonian ( brazil ) freshwater puffer fish .
in lateral view , the skull is characterized by the wide preopercle with about 110 degrees between both horizontal and vertical rami ( figure 7 ) , with the preopercular canal running along its anterior border , and by the opercle which is divided in two distinct regions , having ventral and posterior wings , the herein called “inverted v” shape , distinct from the condition found in all other examined specimens of colomesus ( figure 10 ) . the subopercle is sturdy , with two small dorsal processes .
puffer flesh is toxic and can cause clinical poisoning and human mortality , although it is regarded as a delicacy in certain countries . the predominant toxin , usually either tetrodotoxin or saxitoxin , is dependant on species , geographic area , and time of year . the toxins are not produced by the fishes themselves , but by bacteria living in symbiotic association , or they are acquired via the food chain . colomesus species accumulate saxitoxin , although it is unclear whether eating their flesh represents a danger to humans .
among the amazonian taxa exploited by the ornamental fish industry in south america are those of colomesus [ 2 ] , a genus confined to south america , with what is presently considered two species , c . asellus and c . psittacus . c . asellus [ 3 ] is spread in the entire amazon , tocantins - araguaia drainages , and coastal environments from the amazon mouth to venezuela , being the only freshwater puffers on that continent . c . psittacus [ 4 ] is found in coastal marine and brackish water environments from cuba and the northern coast of south america to sergipe in brazil .
within the genus colomesus , c . asellus can be immediately identified by possessing a unique transverse row of dermal flaps across the chin which is absent in its congeners c . psittacus and c . tocantinensis . it can be further told apart from the very similar c . tocantinensis by possession of 10 ( vs . 9 in c . tocantinensis ) anal - fin rays , 11 ( vs . 10 ) dorsal - fin rays , a triangular ( vs . notched ventrally , appearing as an inverted v ) opercle , base colour in dorsal portion of body golden yellow ( vs . light yellow to pale ) .
colomesus species diagnosed by six to seven basal pterygiophores and nine rays in the anal fin ( contra ten to eleven in both c . asellus and c . psittacus ) ; ten basal pterygiophores and rays in the dorsal fin ( contra eleven for both c . asellus and c . psittacus ) ; the absence of dermal flaps across the chin ( contra its presence uniquely in c . asellus ) ; a caudal peduncle with eight vertebrae ; and an opercle with a posterior ventral border subdivided in a ventral and a posterior region , the herein called “inverted v” shape ( contra the triangular opercle exhibited by both c . asellus and c . psittacus ) .
although the influence of marine incursions after the miocene is still under debate , the caribbean ( or miocene ) marine incursion , via the llanos basin ( colombia - venezuela ) , is well accepted based on both geological and paleontological evidence , suggesting that these incursions may have isolated marine taxa within the western south america freshwater environments [ 48 ] – [ 52 ] . this might be the case for the freshwater tetraodontids . as pointed by [ 53 ] , this scenario predicts that the distribution of the marine sister groups of marine lineages should be related with the caribbean or western atlantic , the age of freshwater taxa should be coincident with marine incursions , and the biogeographic congruence should be observed among multiple unrelated taxa , conditions only partially filled by the genus colomesus .
the skull is partially similar to those found in colomesus asellus described and figured by [ 46 ] , although the frontals exhibit a wide posterior border and prominently participate in the orbital margin ( figures 7 – 9 ) . the prefrontals are triangular and articulate medially with the ethmoid , which posteriorly articulates with the frontals and anteriorly with the palatines ( figure 8 ) . the supraoccipital is roughly triangular and well developed , with an elongate posterior process which covers the first vertebrae ( figure 8 ) . the sphenotics articulate postero - laterally with the frontals and , in the examined specimens , they neither contact nor closely approach the prefrontals . the lateral wing of the sphenotics is only partially developed ( figure 8 ) . posterior to the sphenotics , the pterotics ( figures 7 and 8 ) articulates posteriorly with the slender supracleithrum and medially with the epiotics , which articulates medially with the supraoccipital ( figure 9 ) .
the genus colomesus is restricted to south america ( amaral et al . 2013 ) . tetraodontids are characterized by a tough skin that is often covered with small spinulous scales , a beak - like dental plate divided by a median suture , a slit - like gill opening anterior to the base of the pectoral fin , no pelvic fins , no fin spines , a single usually short - based dorsal fin , a single usually short - based anal fin , and no ribs . they are capable of inflating their abdomens with water when frightened or disturbed and are capable of producing and accumulating toxins such as tetrodotoxin and saxitoxin in the skin , gonads , and liver . the degree of toxicity varies by species , and also according to geographic area and season ( allen and randall 1977 , allen and erdmann 2012 ) . fishes in the family tetraodontidae have the smallest vertebrate genomes known to date ( neafsey and palumbi 2003 )
greek , kolos = short , truncated + greek , mesos = a half ( ref . 45335 )
marine ; freshwater ; brackish ; demersal ; ph range : 7 . 0 - ? ; dh range : 10 - ? ; depth range 1 - 40 m ( ref . 5217 ) . tropical ; 23°c - 26°c ( ref . 2060 )
maturity : l m ? range ? - ? cm max length : 29 . 3 cm tl male / unsexed ; ( ref . 71685 ) ; common length : 25 . 0 cm tl male / unsexed ; ( ref . 5217 )
dorsal spines ( total ) : 0 ; dorsal soft rays ( total ) : 11 - 12 ; anal spines : 0 ; anal soft rays : 11 . body except for snout , pectoral base and caudal peduncle covered with prickles ; teeth fused into plates , two plates on each jaw ; nostril with two openings ; body dark green dorsally with six transverse black bars , white ventrally ; fins dusky green or dark brown ( ref . 13608 ) .
solitary or in groups of 2 or 3 individuals but never in schools ( ref . 35237 ) . inhabits shallow inshore waters usually on soft bottoms . frequently found in freshwater ( ref . 13608 ) . when threatened , it becomes inflated like a balloon , in order to ward off predators . carnivorous , feeding mainly on mollusks which crushes with its powerful teeth ( ref . 35237 ) . of negligible commercial importance and usually not marketed ( ref . 5217 ) .
cervigón , f . , r . cipriani , w . fischer , l . garibaldi , m . hendrickx , a . j . lemus , r . márquez , j . m . poutiers , g . robaina and b . rodriguez , 1992 . fichas fao de identificación de especies para los fines de la pesca . guía de campo de las especies comerciales marinas y de aquas salobres de la costa septentrional de sur américa . fao , rome . 513 p . preparado con el financiamento de la comisión de comunidades europeas y de norad . ( ref . 5217 )
) : 25 . 6 - 28 , mean 27 . 3 ( based on 210 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 7500 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 02570 ( 0 . 01637 - 0 . 04036 ) , b = 2 . 86 ( 2 . 73 - 2 . 99 ) , in cm total length , based on lwr estimates for this species & ( sub ) family - body ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 6 ±0 . 50 se ; based on food items .
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 25 of 100 ) .
freshwater ; demersal ; ph range : 5 . 5 - 7 . 2 ; dh range : 5 - 15 . tropical ; 22°c - 28°c ( ref . 13614 )
south america : amazon river basin from peru to marajó island , including tributaries araguaia and guaporé rivers ; orinoco river basin near the mouth ; essequibo river basin .
maturity : l m ? range ? - ? cm max length : 12 . 8 cm sl male / unsexed ; ( ref . 79673 )
found mostly in freshwater and coastal streams , but can tolerate brackish water . sometimes kept in aquariums ( ref . 26938 ) .
ortega , h . and r . p . vari , 1986 . annotated checklist of the freshwater fishes of peru . smithson . contrib . zool . ( 437 ) : 1 - 25 . ( ref . 6329 )
bayesian length - weight : a = 0 . 03236 ( 0 . 01891 - 0 . 05537 ) , b = 2 . 85 ( 2 . 71 - 2 . 99 ) , in cm total length , based on lwr estimates for this species & ( sub ) family - body ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 3 ±0 . 51 se ; based on food items .
vulnerability ( ref . 59153 ) : low vulnerability ( 14 of 100 ) .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
carpenter , k . e . , comeros - raynal , m . , harwell , h . & sanciangco , j .
is known from the gulf of paria to northern brazil where it is common and locally abundant in mangrove creeks and estuaries at depths ranging from 0 to 32 metres . it appears to be common and abundant in parts of its range .
is a component of the marine aquarium trade , however there are no indications of population declines from harvesting at present time . there have been no documented population declines in
, however due to its affinity with mangroves this species may be experiencing population declines due to habitat loss in parts of its range .
, however its distribution overlaps with several marine reserves in parts of its range . it is therefore listed as least concern .
2010 ) . its range extends eastward the gulf of paria and orinoco river region on the eastern coast of venezuela to the amazon river basin in northern brazil . it is found as far south as sergipe in brazil ( amaral 2013 ) . it is known from marine and freshwaters around trinidad and tobago ( tyler 1964 , philip
. 2013 ) . it is also found in cuba ( amaral 2013 ) . it can also be found in freshwater , and is known from several hundred kilometres upstream in the amazon and its tributary river system .
is a dominant component of northern mangrove estuaries , both numerically and by weight .
2007 ) accounting for between 19 and 52 % of total catch mass ( giarrizzo and krumme 2009 ) . this species is the second most abundant estuarine species in tidal mangrove creeks of the lower caeté estuary , with population density and biomass fluctuating seasonally ( barletta
2003 ) but with a continuous occurrence ( giarrizzo and krumme 2009 ) . in mangrove creeks in northern brazil , there were no major changes in the mean size of juvenile
2013 ) . this species exhibited a geographic consistence in abundance in northern brazilian estuaries ( between 7 . 7 % and 11 . 4 % ) ( vilar
are very common in museum collections ( accessed through the fishnet2 portal , www . fishnet2 . org , 2014 - 03 - 17 ) .
may be experiencing population declines due to habitat loss in parts of its range .
is thought to have relatively continuous reproductive activity in brazilian mangrove creeks ( giarrizzo and krumme 2009 ) . different estuarine habitats are inhabited by different size classes of resident
, and are likely connected by regular tidal migrations ( giarrizzo and krumme 2009 ) . the estimated consumption / biomass ratio of this species in a mangrove estuary in northeastern brazil was 8 . 8 , which was relatively low - q / b ratios varied from between 2 . 3 for
is a component of the aquarium trade , and is exported from brazil , colombia , and peru ( prang 2007 ) . in brazil , this species is of no commercial interest to fishermen and is rarely marketed due to its toxicity ( krumme
2007 ) . under the name baiacu , this species is valued for its medicinal properties . the bile and liver oil are used to treat breast cancer , backache , and warts ( alves and rosa 2007 ) . the japanese name for this species is kuro - obifugu .
may be experiencing population declines due to habitat loss in parts of its range . it is taken as bycatch in the laulao catfish (
globally , 16 % of mangrove species are at elevated risk of extinction . particular areas of geographical concern include the atlantic and pacific coasts of central america , where as many as 40 % of mangroves species present are threatened with extinction . ( polidoro et al . 2010 ) . in the caribbean , approximately 24 % of mangrove area has been lost over the past quarter - century ( fao 2007 ) .
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psittacus : from the ancient greek ψιττακός  ( psittakós ) , meaning ‘parrot’ , persumably in reference to this species’ beak - like mouthparts .
this species’ known range extends eastwards from the parque nacional natural tayrona in magdalena department , colombia , across northern venezuela , the gulf of paria and orinoco delta , then southwards via the guyanas and past the mouth of the amazon , with the southernmost records from sergipe state in northeastern brazil . it is also known from trinidad and tobago and caribbean islands including the greater antilles , lesser antilles , and bahamas .
records from the middle and upper amazon basins appear to represent misidentifications of the congener c . asellus .
although it does penetrate the lower basins of rivers , particularly the amazon where it has been collected from the rio xingu several hundred kilometres from its mouth , this species is predominantly an inhabitant of mangrove swamps , estuaries , and other such saline habitats .
it is particularly common in tidal channels , shallow inshore lagoons , and the lower reaches of rivers .
is suggested as a bare minimum , but even this may prove too small for long - term care .
choice of décor is not especially critical though it should be maintained in a well - decorated set - up , perhaps containing some driftwood roots or branches in order to mimic its natural mangrove habitats .
it is intolerant of organic waste and require spotless water in order to thrive . moderate levels of dissolved oxygen and water movement are also recommended , meaning additional powerheads , pumps , etc . , should be employed as necessary . a linear flow pump may prove a useful addition , while weekly water changes of 30 - 50 % should be considered mandatory .
wild examples can be delicate and sensitive to white spot / ich post - import , so a lengthy quarantine period may be required . maintenance in pure freshwater may also present problems , so the addition of marine salt to a standard gravity of ≥1 . 010 is recommended .
chiefly carnivorous and feeds almost exclusively on molluscs and crustaceans such as cirripedia ( barnacles ) and brachyuran crabs , taking increasingly mobile prey as it matures . there is also evidence to suggest seasonal changes in diet , with crabs favoured during drier periods .
in the aquarium offer unshelled crab legs , cockles , mussels , prawns , etc . tetraodontids lack true teeth , the jawbone itself being modified into four fused toothlike structures . these grow continuously at a surprising rate , so such a diet is essential in order to maintain them at a reasonable length .
juveniles apparently form loose aggregations in the wild , but in the confines of an aquarium this species is likely to prove aggressive in all but the largest systems .
this species is also referred to as ‘parrot puffer’ in the ornamental trade , although it is not a popular aquarium fish .
other defining characters of tetraodontids include a tough skin usually covered with small spines , a beak - like dental plate divided by a median suture , a reduced gill opening anterior to the pectoral - fin base , no pelvic fins or spinous fin rays , typically short - based dorsal and anal fins , and no ribs .
bloch , m . e . and j . g . schneider , 1801 - gottlob schneider , saxo . berolini . sumtibus auctoris impressum et bibliopolio sanderiano commissum : i - lx + 1 - 584 m . e . blochii , systema ichthyologiae iconibus cx ilustratum . post obitum auctoris opus inchoatum absolvit , correxit , interpolavit jo .
camargo , m . , t . giarrizzo , and v . isaac , 2004 - ecotropica 10 : 123 - 147 review of the geographic distribution of fish fauna of the xingu river basin , brazil .
gill , t . n . , 1884 - proceedings of the united states national museum 7 ( 26 - 27 ) : 411 - 427 synopsis of the plectognath fishes .
helfman , g . , b . b . collette , d . e . facey , and b . w . bowen , 2009 - wiley - blackwell : 1 - 736 the diversity of fishes : biology , evolution , and ecology , 2nd edition .
nelson , j . s . , 2006 - john wiley & sons , hoboken , n . j . : i - xix + 1 - 601 fishes of the world . 4th edition .
phillip , d . a . t . , d . c . taphorn , e . holm , j . f . gilliam , b . a . lamphere and h . lópez - fernández , 2013 - zootaxa 3711 ( 1 ) : 1 - 64 annotated list and key to the stream fishes of trinidad & tobago .
reis , r . e . , s . o . kullander and c . j . ferraris , jr . ( eds ) , 2003 - edipucrs , porto alegre : i - xi + 1 - 729 check list of the freshwater fishes of south and central america . cloffsca .
the best way to identify c . asellus from c . psittacus is the number of bands on their back . c . asellus having 5 when c . psittacus having 6 . the black band at the nose doesn’t count . even if the band is very faint , it counts .
asellus : derivation unclear ; possibly from the latin asellus , meaning ‘small donkey’ .
occurs throughout much of the amazon basin in brazil , colombia , peru , and ecuador , including the amazonas / solimões main channel plus the rios pará , tocantins , jari , xingu , tapajós , uatumã , madeira , trombetas , negro , purus , tefé , japurá / caquetá , juruá , jutaí , içá / putomayo , javary , ampiyacu , amacayacú , napo , nanay , marañón , and ucayali , with its range extending at least as far upstream as pucallpa in eastern peru .
there are also numerous records from drainages north of the amazon mouth including the essequibo and waini in guyana , and lower orinoco in venezuela . it appears to be absent from french guiana and suriname .
has been recorded in lower , middle , and upper river basins with habitats including sandbars , beaches , floodplain lakes , banks with overhanging vegetation , and fast - flowing rapids over bedrock , boulders , and stones . it is mostly collected from habitats with high oxygen levels , suggesting that it may be sensitive to low oxygen availability .
it is adaptable , penetrating into tributaries of the upper amazon , but also occuring in the amazon and orinoco delta regions , although it does not tend to be found in highly acidic black - waters .
the maximum recorded length in wild specimens is 128 mm , but aquarium reports suggest 70 – 80 mm to be typical .
choice of décor is not especially critical though it should be maintained in a well - decorated set - up . the addition of floating or overhanging vegetation and driftwood roots or branches also seems to be appreciated .
this species is intolerant of organic waste and require spotless water in order to thrive . moderate levels of dissolved oxygen and water movement are also recommended , meaning additional powerheads , pumps , etc . , should be employed as necessary . a linear flow pump may prove a useful addition , while weekly water changes of 30 - 50 % should be considered mandatory .
wild examples can be delicate and sensitive to white spot / ich post - import , so a lengthy quarantine period may be required .
tetraodontids lack true teeth , the jawbone itself being modified into four fused toothlike structures . these grow continuously at a surprising rate , so offer regular meals of shelled invertebrates such as snails , crab legs , cockles , etc . , in order to maintain them at a reasonable length . there is some evidence to suggest that aufwuchs form a significant proportion of the natural diet , therefore it may be worth permitting or even encouraging algal growth on hard items of décor .
additional foods can include chopped shellfish , small earthworms , and live or frozen chironomid larvae ( bloodworm ) , artemia , etc . dried products should not form the principal component of the diet , although pelleted formats with a very hard consistency may prove useful .
not aggressive as such but unsuitable for the general community aquarium , and best - maintained alone or in a larger set - up with other fluvial fishes .
this species naturally forms loose aggregations and can behave nervously in the absence of conspecifics . ideally a group of 6 or more should be purchased .
this species exhibits a spawning strategy comparable to that of marine puffers and in contrast to the majority of freshwater tetraodontids , with high fecundity , relatively small eggs , and no parental care . limited studies in the central amazon basin suggest that spawning occurs in main river channels or close to banks at the mouths of floodplain lakes and tributaries during periods of high water . the pelagic larvae are washed into nursery zones in floodplain lakes where they complete their development , returning to river channels when flood waters recede .
this species is also referred to as ‘south american puffer’ , ‘sap’ , ‘amazonian puffer’ , ‘peruvian puffer’ , or ‘brazilian puffer’ in the ornamental trade .
it is distinguished from c . psittacus by its smaller adult size ( max . 128 mm sl vs . 289 mm sl in c . psittacus ) , possession of 13 - 16 ( vs . 17 - 19 ) pectoral - fin rays , presence of 5 ( vs . 6 ) transverse dark bands dorsally on the body , and predominantly freshwater , fluvial ( vs . brackish , coastal ) ecology .
müller , j . and f . h . troschel , 1849 - im auftrag sr . mäjestat des königs von preussen ausgeführt von richard schomburgk v . 3 . berlin : 618 - 644 fische . in : reisen in britisch - guiana in den jahren 1840 - 44 .
ortega , h . and r . p . vari , 1986 - smithsonian contributions to zoology 437 : iii + 1 - 25 annotated checklist of the freshwater fishes of peru .
its really hard to identify c . asellus from c . psittacus , the best way i found out is to count the black bars on their body . c . asellus has 5 bars and c . psittacus has 6 .
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eschmeyer , w . n . ( ed ) . catalog of fishes . urltoken electronic version accessed 03 - nov - 2014
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 .
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froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of tetrodon psittacus bloch & schneider , 1801 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
my amazonian , a . k . a . patagonian puffer fish , i love him ! i breed snails for him in a separate 2 , 5 gallon tank . he also loves frozen bloodworms , but he really needs snail to keep his teeth at correct size .
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institut für zoologie der rheinischen friedrich - wilhelms - universität bonn , poppelsdorfer schloszligbeta ; , d - 53115 bonn , germany . hanke @ urltoken
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copyright : © 2013 amaral et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : this study was supported by the brazilian national counsel of technological and scientific development and fundação carlos chagas filho de amparo à pesquisa do estado do rio de janeiro . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
the tetraodontidae is an acantomorpha fish family with circumglobal distribution composed of 189 species in 19 genera , occurring in seas , estuaries , and rivers between the tropical and temperate regions [ 1 ] . they are mainly characterized by their typical four large dental plates ; the ability to inflate their body in stressful situations ; the presence of the neurotoxin tetrodotoxin / saxitoxin in its tissues , being responsible for numerous cases of fatal poisoning in many countries , including brazil ; and by having the smallest genome among vertebrates , therefore being considered as a model for the genome evolution of the group .
mainly located in tropical and subtropical regions all around the world , including the amazon region , the ornamental fish industry is one of the largest transporters of live animals and plants with an annual trade volume estimated at u $ 15–25 billion [ 5 ] – [ 7 ] , in a scenario where species identification problems , mainly related to border biosecurity are not rare .
the dna barcode is a widely accepted tool for species determination mainly due to its enhanced attention on standardization and data validation [ 8 ] , being a rapid and low cost method of identification [ 9 ] . the use of dna barcoding techniques has been utilized in many taxa , including bacteria , birds , bivalves , butterflies , fishes , flies , macroalgae , mammals , spiders , sprigtails , and also for plants [ 10 ] – [ 27 ] .
the dna barcode technique for metazoans uses a short ( ∼650 bp ) and standardized gene region from the mitochondrial 5′ region of the cytochrome c oxidase subunit i ( coi ) for a rapid and cost - effective animal identification . this has been demonstrated to be an effective fish identification tool in numerous situations , including consumer protection [ 28 ] – [ 30 ] , fisheries management / conservation [ 31 ] , border biosecurity in the ornamental fish trade [ 5 ] , and in the identification of overlooked or cryptic species [ 32 ] .
map of south america showing the northern hydrology and the localities where the specimens were collected ( grey and green marks ) .
no statement from an ethics committee was necessary , and the manuscript did not involve any endangered or protect species . all samples were extracted from dead specimens collected with appropriate permissions under authorization number 22512 issued by sisbio / instituto chico mendes de conservação da biodiversidade . we used the ice - slurry method for killing following [ 33 ] as they are tropical warm water species and the collected specimens are all smaller than 5 cm sl . all specimens were preserved in alcohol . the reported localities do not include protected areas .
the electronic version of this document does not represent a published work according to the international code of zoological nomenclature ( iczn ) , and hence the nomenclatural acts contained in the electronic version are not available under that code from the electronic edition . therefore , a separate edition of this document was produced by a method that assures numerous identical and durable copies , and those copies were simultaneously obtainable ( from the publication date noted on the first page of this article ) for the purpose of providing a public and permanent scientific record , in accordance with article 8 . 1 of the code . the separate print - only edition is available on request from plos by sending a request to plos one , 1160 battery street , suite 100 , san francisco , ca 94111 , usa along with a check for $ 10 ( to cover printing and postage ) payable to “public library of science” .
in addition , this published work and the nomenclatural acts it contains have been registered in zoobank , the proposed online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : 033a323a - 18f7 - 4788 - 8405 - 32d78bf65b13 .
specimens from all three localities were cleared and stained following the methodology of [ 34 ] .
the molecular systematic analyses used newly determined sequences obtained from the mitochondrial barcode marker coi as well as previously published sequences obtained from the ncbi and bold databases .
for the newly determined sequences , a fragment of epaxial musculature was submitted to the standard protocol for dna extraction and purification from the qiagen qiaamp dna ffpe tissue kit . the fragments were amplified and sequenced using the primers vf2 _ t1 and fishr2 _ t1 [ 35 ] – [ 37 ] . all primers were appended with m13 tails on sequencing reactions . the pcr profile consisted of 2 min at 95°c , 35 cycles of 30 sec at 94° , 40 sec at 52°c , and 1 min at 72°c , with a final extension step for 10 min at 72°c . sequencing reactions were performed with the use of the bigdye® terminator v . 3 . 1 cycle sequencing kit ( applied biosystems , inc . ) , with 25 cycles of 10 sec at 95°c , 5 sec at 50°c and 4 min at 60°c . sequencing products were processed in an abi 3500 capillary system ( applied biosystems , inc . ) .
the chromatograms were checked and aligned using the bioedit 7 . 053 [ 38 ] software with its built - in clustalw routine [ 39 ] . the alignment was visually inspected for accuracy and to minimize missing data . all the newly determined sequences are available at the bold database ( urltoken ) under the project acronym pufer . the genbank accession numbers for all newly determined and previously published sequences used in the present manuscript are summarized in table 1 . the dataset consisted of a 651 bp coi matrix , and we used the mega 5 . 06 software [ 40 ] to determinate the tn93 + g + i as the most appropriate model of sequence evolution based on the akaike criterion ( aic ) [ 41 ] .
the neighbor - joining sequence divergences were calculated based on the kimura two parameter ( k2p ) distance model [ 42 ] on bold workbench and mega 5 . 06 software [ 40 ] . the haplotype determination was carried with the use of the server fabox ( urltoken ) .
the k2p divergence distances between congeneric species ranged from 5 . 557 % to 12 . 394 % with a mean distance of 8 . 546 % , while the uncorrected k2p distance ranged from 0 to 4 . 472 % within species . the mean k2p distance within the analyzed populations was 0 . 657 % and the mean normalized distance within species is 1 . 079 % ( figure 4 ) .
distribution of k2p distances ( % ) for coi : a ) within species ; b ) within genera ; c , normalized distribution of k2p distance ( % ) within species .
the specific epithet tocantinensis refers to the type locality , porto nacional , state of tocantins , brazil .
the specimens are from the tocantins river near porto nacional , state of tocantins , brazil .
the holotype ( pnt . uerj . 405 ) is 29 , 62 mm sl ( figure 6 ) , with 10 , 37 mm hl ; the entire type - series ranges from 27 . 02 mm to 34 . 9 mm sl . the meristic and morphometric data of the type series is presented in table 2 . the extent of the dorsal and ventral lateral lines is similar to those found in c . asellus . the prickles extend along the dorsal , lateral , and ventral surfaces of the body , from the level of the eye to the origin of the dorsal fin .
the nasal sac is higher than that presented in the specimens of c . asellus . two large lateral and anteromedial nostrils are present . they are similar to those found on c . psittacus , rather than the two small nostrils exhibited by c . asellus . the anterior surface of the nasal sac is smooth while the posterior surface of it is folded as in c . psittacus , exhibiting a “t - shaped” ridge with a relatively small dorsal flap . this flap seems much smaller than the one found on c . asellus , although more flexible when compared to c . psittacus .
the presence of dermal flaps across the chin is another character used by [ 46 ] to distinguish c . asellus from c . psittacus . no dermal flaps could be seen in the examined specimens from the tocantins river , although they are always present in examined specimens from iquitos and belém .
a ) left photograph of the head ; b ) anatomical interpretations . abbreviations : ang , angular ; art , articular ; boc , basioccipital ; brstgs , branchiostegals ; cl , cleithrum ; den , dentary ; epi , epiotic ; ethm , ethmoid ; exo , exoccipital ; fr , frontal ; hyo , hyomandibula ; ecptg , ectopterygoid ; mept , mesopterygoid ; mtptg , metapterygoid ; mx , maxilla ; op , opercle ; pal , palatine ; pcl . l / r , ventral post - cleithrum left and right ; pfr , prefrontal ; pmx , premaxilla ; pop , preopercle ; pro , prootic ; psph , parasphenoid ; pto , pterotic ; qua , quadrate ; r , radials ; scl , supracleithrum ; soc , supraocciptal ; sop , subopercle ; sphe , sphenotic ; sym , sympletic ; vo , vomer . scale bar equals 5 mm .
a ) top photograph of the head ; b ) anatomical interpretations . abbreviations : epi , epiotic ; ethm , ethmoid ; exo , exoccipital ; fr , frontal ; pal , palatine ; pfr , prefrontal ; pmx , premaxilla ; pto , pterotic ; scl , supracleithrum ; soc , supraocciptal ; sphe , sphenotic . scale bar equals 5 mm .
the parasphenoid is elongate and does not exhibit any developed dorsal flange ( figures 7 and 9 ) . the hyomandibula is roughly triangular and has a slender ventral region ; its wide head articulates dorsally with the sphenotics , and its upper posterior edge with the anterior end of the opercle ( figure 7 ) .
the palatine is wide and somewhat triangular , with a robust anterior process for the maxilla ( figure 7 ) . the maxilla is robust , with an anterodorsal region articulating with the premaxilla and a posterior expanded region , medially concave for muscle insertion . the ectopterygoid articulates dorsally with the palatine and ventrally with the anterodorsal border of the quadrate . the metapterygoid is wide and composes almost the entire ventral orbital region ( figure 7 ) . it articulates anteriorly with the mesopterygoid ( figure 7 ) , and with the posterior end of the large and triangular quadrate ( figure 7 ) . the quadrate exhibits a well - developed posteroventral spine articulating posteriorly with the slender symplectic ( figure 7 ) , and anteriorly with the articular . the articular is “l” shaped and articulates anteriorly with the robust dentary and ventrally with the small angular ( figure 7 ) .
five branchiostegal rays ( figure 7 ) are present and the branchial apparatus is strikingly similar in all the examined specimens ( figure 11 ) .
the pectoral girdle is robust and formed by a wide cleithrum , somewhat triangular and posteriorly expanded , articulating dorsally with the slender supracleithrum . the supracleithrum articulates ventrally with the two postcleithra ; a slender dorsal postcleithrum , followed by the posteriorly expanded ventral postcleithrum ( figure 7 ) . there are four radials and sixteen pectoral fin rays ( figure 7 ) .
the axial skeleton has 19 vertebrae . the dorsal fin originates between vertebrae 7–8 and has ten basal pterygiophores and ten fin rays ( figures 12 – 14 ) . the anal fin is located beneath the 9 th vertebra and has six basal pterygiophores and nine fin rays .
a ) left photograph of the unpaired fins and caudal endoskeleton ; b ) anatomical interpretations . abbreviations : e , epural ; h , dorsal hypural plate ; h - h , ventral hypural plate fused with the ural centrum ; phy , parhypural ; pu , pre - ural vertebrae ; pbd , dorsal - fin basal pterigiophores ; pba , anal - fin basal pterigiophores . scale bar equals 5 mm .
the caudal skeleton ( figure 12 ) has a wide ural centrum formed by the preural centrum 1 , the ural centrum , the ventral hypural plate , and the postero - dorsal expansion which articulates anteriorly with the almost triangular epural , and posteriorly with the dorsal hypural plate ( figure 12 ) . eleven caudal fin rays , five dorsal and six ventral , are present in all of the specimens , both the uppermost and the two lowermost rays are unbranched .
it was recently proposed [ 55 ] , based on the distribution of characiforms , that recent marine incursions would have isolated fish populations in upland terrains or refuges , where lineage divergence is maximized , followed by dispersal episodes back to the lowlands . the “museum hypothesis” predicts that lowlands exhibit a higher number of species , but lower levels of endemism , than highlands , and that the upland refuges would represent areas of high endemism .
looking on the molecular phylogeny of the serrasalmids pygocentrus and serrasalmus , [ 56 ] proposed a phylogenetic test which predicts that basal lineages in a phylogeny of widespread fishes would occur in highland areas , and lowland lineages would have originated only during the last 5 ma . additionally , [ 57 ] studying the genetics of symphysodon cichlids , indicated the effects that marine incursions would have in population structure , stating that populations in upland terrains or refuges would exhibit reduced genetic variation , while populations in lowlands would represent multiple upland sources , therefore exhibiting a high level of genetic variation , and that populations in lowlands would show a demographic pattern of expansion .
our results recovered the upper tocantins lineages in a basal position in relation to all the remaining specimens , with the sequences being collapsed in uniquely two haplotypes ( figure 5 ) , the first one ( h1 ) , represented by eight sequences , and the second haplotype ( h2 ) , represented by a unique sequence . this suggests low genetic variation , at least among the studied sampling , and a history initially related with the eastern amazon , followed by a subsequently expansion to the western south america .
the tocantins - araguaia drainage is the fourth largest brazilian drainage , draining part of the northern end of the brazilian shield directly to the eastern end of the amazon basin . it exhibits a recent geomorphological history , within a still tectonically active sedimentary basin with recent subsidence episodes , which are related with the high load of sediments observed within the basin , leading the development of the bananal plain , in the lower part of the drainage , mainly during the quaternary [ 58 ] – [ 59 ] .
finally , our results reinforce the upper tocantins drainage as an area of high endemism within the tocantins - araguaia drainage , although the composite nature of the entire drainage is unquestionable .
we would like to thank dr . james c . tyler ( smithsonian institution , washington ) for his support and helpfull comments on the manuscript . we are also grateful to dr . francesco santini ( universitá degli studi di torino ) , dr . dorothée huchon ( tel aviv university ) , and an anonymous reviewer for the valuable suggestions during the review of the manuscript , dr . leonor gusmão and dr . antonio amorim ( universidade do porto ) for the comments during the initial discussion of the results , yuri modesto ( universidade do estado do rio de janeiro ) for the specimens from the tocantins drainage , lúcio paulo machado and diogo de mayrink ( universidade do estado do rio de janeiro ) for the specimens from iquitos ; ms . sandra raredon ( smithsonian institution , washington ) for the x - rays of tetraodontids , dr . richard pyle ( hawaii biological survey ) for the lsid numbers , and kleyton m . c . severiano and anna carolina chaves ( universidade do estado do rio de janeiro ) for the technical assistance .


the geelvink imperial pigeon ( ducula geelvinkiana ) is a bird in the family columbidae formerly considered conspecific with the spice imperial pigeon ( ducula myristicivora ) .

it is endemic to the islands of biak , numfoor and mios num in the indonesian province of papua .

geelvink imperial pigeon

islands of numfor , biak and mios num , in geelvink bay ( nw new guinea ) .
del hoyo , j . , collar , n . , kirwan , g . m . & garcia , e . f . j . ( 2018 ) . geelvink imperial - pigeon ( ducula geelvinkiana ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
not globally threatened . common on islands in geelvink bay . presence on nearby mainland ( see movements ) , whether as a wanderer or , presumably scarce , resident remains to be . . .
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del hoyo , j . , collar , n . j . , christie , d . a . , elliott , a . and fishpool , l . d . c . 2014 . hbw and birdlife international illustrated checklist of the birds of the world . volume 1 : non - passerines . lynx edicions birdlife international , barcelona , spain and cambridge , uk .
ducula myristicivora and d . geelvinkiana ( del hoyo and collar 2014 ) were previously lumped as d . myristicivora following sibley and monroe ( 1990 , 1993 ) .
justification : although this species may have a restricted range , it is not believed to approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . the population trend appears to be stable , and hence the species does not approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size has not been quantified , but it is not believed to approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
the global population size has not been quantified , but the species is reported to be common ( del hoyo et al . 1997 ) . trend justification : the population is suspected to be stable in the absence of evidence for any declines or substantial threats .
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hitherto considered conspecific with d . myristicivora , but differs on account of its lack of black knob on bill ( 3 ) ; lack of pale vinous wash on hindneck ( 2 ) ; yellow iris , based on photographic evidence ( at least 2 ) ; darker grey head , breast and mantle , with dirtier , darker lower underparts ( ns [ 1 ] ) ; greener - glossed vs bluer - glossed rump ( inconstant character ; entire upperparts of geelvinkiana tend to be more strongly green [ ns ] ) ; smaller size ( wing length effect size –1 . 636 ; score 1 ) . also closely related to d . rubricera ; a close relationship with d . concinna has also been suggested . monotypic .
( see taxonomy comments ) but smaller ( wing 227–232 mm , versus 257–260 mm ) . lacks pink on hindcrown , ear - coverts . . .
advertising call a medium - pitched , coarse “crwwooo” repeated incessantly at rate of one per second . . .
occurs in forest of all types , including secondary and logged areas , from sea - level to 500 m .
no information available , although presumably exclusively frugivorous , gathering at fruiting trees , usually alone or in pairs .
probably mainly resident , but there is at least one mainland record , from hill forests at c . 400 m . . .
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although this species may have a restricted range , it is not believed to approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . the population trend appears to be stable , and hence the species does not approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size has not been quantified , but it is not believed to approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
recommended citation birdlife international ( 2018 ) species factsheet : ducula geelvinkiana . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .
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