the seychelles frogs ( sooglossidae ) are a family of frogs found on the seychelles islands and india .

until recently , this family was believed to include the genera nesomantis and sooglossus , but following a major revision of amphibians in 2006 , the genus nesomantis was named a junior synonym of sooglossus ; conversely , the recently discovered purple frog , which was initially assigned to a distinct monotypic family ( nasikabatrachidae ) , is now included in the sooglossidae .

all of the species are relatively small terrestrial frogs , about 4 cm ( 1.6 in ) in length , hiding under fallen leaves or in rock crevices .

they are unusual for the neobatrachials , in that they undertake inguinal amplexus , a primitive version of amplexus .

except for n. sahyadrensis , they lay their eggs on moist ground , rather than in water .

several species lack tadpoles , with the eggs hatching directly into froglets .

the tadpoles of s. sechellensis are carried , abnormally , on the backs of the female frogs until metamorphosis .

there is no fossil record for the family .

ancestors diverged when the seychelles islands split from india about 100 million years ago .

sooglossidae

cannatella , d . 1996 . sooglossidae : tree of life . ( website . )
cannatella , david . 1995 . sooglossidae . version 01 january 1995 ( under construction ) .
the distribution of living members of the family sooglossidae is confined to the seychelles islands , as indicated in red .
pictures of seychellfrogs ( sooglossidae ) .
swiss herp . urltoken ( accessed on february 22 , 2005 ) .
seychelles frogs : sooglossidae .
grzimek ' s student animal life resource . . retrieved july 09 , 2018 from urltoken urltoken
seychelles frogs : sooglossidae .
grzimek ' s student animal life resource . . encyclopedia . com . ( july 9 , 2018 ) . urltoken
to cite this page : heying , h . 2003 .
sooglossidae
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
nussbaum , r . a . 1982 . heterotopic bones in the hindlimbs of frogs of the families pipidae , ranidae , and sooglossidae . herpetologica 38 : 312 - 320 .
sooglossidae — griffiths , 1963 , biol . rev . cambridge philos . soc . , 38 : 273 ; laurent , 1980
1979
, bull . soc . zool . france , 104 : 418 .
family sooglossidae no fossil record ; 8 presacral vertebrae ; vertebrae procoelous ; sacral diapophyses dilated ; intercalary cartilages absent ; larvae lacking spiracle ; seychelles ; 2 genera , 3 species ; length about 4 cm ( 1 . 5 inches ) . superfamily ranoidea pectoral girdle firmisternal ; ribs absent ; amplexus axillary ; larvae with single sinistral spiracle
ford and cannatella ( 1993 ) defined sooglossidae as a node - based name for the common ancestor of nesomantis and sooglossus , and all its descendants . the only published synapomorphy appears to be the presence of a sesamoid bone in the heel region ( os sesamoides tarsale ; nussbaum , 1982 ) , a feature shared with some petropedetine ranids . sooglossids have been allied to ranoids ( griffiths , 1959a , 1963 ; tyler , 1985 ) or to myobatrachids ( lynch , 1973 ) . duellman ( 1975 ) placed the sooglossids with ranoids on the basis of zoogeography . the two derived characters cited as evidence of a ranoid relationship ( griffiths , 1959a ) are the semitendinosus tendon passing dorsal to the m . gracilis and partial fusion of the epicoracoid cartilages and sternum of the pectoral girdle . nussbaum ( 1980 ) concluded that placement of sooglossidae in ranoidea was not supported , and preferred its placement near myobatrachids . hay et al . ( 1995 ) placed sooglossus in a trichotomy with 1 ) ranoids and 2 ) a clade composed of all other neobatrachians .
ford and cannatella ( 1993 ) described five shared derived characters allying sooglossids to myobatrachines . the first is a ventrally incomplete cricoid ring ( lynch , 1973 ) ; the ring is complete in rheobatrachus and limnodynastines ( heyer and liem , 1976 ) . the second is the semitendinosus tendon inserting dorsal to the m . gracilis . this is shared with all myobatrachines except for taudactylus , which has a ventral insertion ( plesiomorphic ) ; the insertion is also ventral in rheobatrachus ( heyer and liem , 1976 ) . a third shared derived character is the horizontal pupil , which sooglossids share with myobatrachines , except for uperoleia ( also vertical in rheobatrachus ) . limnodynastines primitively have a vertical pupil , but several genera have horizontal ones ( heyer and liem , 1976 ) . a fourth derived character shared with myobatrachines and rheobatrachus is the winglike alary processes of the hyoid ( griffiths , 1959a ) . the fifth is a divided sphenethmoid , which is present in sooglossids ( griffiths , 1959a ) and myobatrachines but not limnodynastines ( lynch , 1971 ) . ford and cannatella ( 1993 ) concluded that sooglossidae is the sister taxon of myobatrachinae ( table 1 ) , and they treated the two as an unnamed more inclusive taxon .
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sooglossids include two genera of small frogs that are restricted to the seychelles islands of the indian ocean ( silhouette and mahé ) . the frogs are relatively small and terrestrial . their phylogenetic placement is the subject of much controversy ; see below . they are unusual as neobatrachians in having inguinal amplexus . there are no fossils .
a character of uncertain status is the presence of notochordal vertebrae ( lynch , 1973 ; but noble [ 1926 ] reported procoelous vertebrae ) , reported in myobatrachines and sooglossids , but not limnodynastines . the polarity is uncertain because 1 ) other presumably closely related taxa have not been characterized adequately , and 2 ) observations on dried specimens differ from those made from serial sections ( cannatella , personal observation ) . sooglossids and myobatrachines share the plesiomorphic condition of inguinal amplexus . according to heyer and liem ( 1976 ) , mixophys is the only myobatrachid genus with axillary amplexus , but lynch ( 1973 ) incorrectly reported that myobatrachines have inguinal amplexus and limnodynastines have axillary amplexus .
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this tiny family of geographically restricted frogs has confounded all systematists who have attempted to discern its relationship to other anurans . with only three species in two genera , this family is restricted to two small islands ( silhouette and mahé ) in the seychelles , an island group roughly 250 miles north of madagascar in the indian ocean
seychelle frogs are small ( 40 mm snout - vent length ) and terrestrial , with digits that terminate in small , pointed discs . the only known synapomorphy of this group is the presence of a sesamoid bone in the heel region . several other characters have been used to unite the sooglossids with either the myobatrachids or the ranoid frogs . sooglossids , along with ranoids , have a semitendinosus tendon that passes dorsal to the m . gracilis , and partial fusion of the epicoracoid cartilages and sternum of the pectoral girdle . like myobatrachines , sooglossids have a ventrally incomplete cricoid ring , horizontal pupils , winglike alary processes on the hyoid , and a divided sphenethmoid . amplexus is inguinal . diploid number is 26 .
sooglossids live in mossy forests on the granitic islands of the seychelles . along with the caecilians , these comprise the oldest herpetofauna on the seychelles , an island group that has attracted much attention from biogeographers and systematists alike . two distinct reproductive modes are known from seychelle frogs . both
eggs , by contrast , hatch in two to three weeks into nonfeeding tadpoles that lack mouthparts and spiracles . these tadpoles crawl onto the mother ' s back , where they remain until they metamorphose . the reproductive mode of the third species ,
, is not known , although the large eggs are suggestive of direct development . males call not in choruses , but individually , though adults of both sexes appear to lack tympana ( eardrums ) .
sooglossids are unambiguously placed in the neobatrachia , but beyond that , nothing about their evolutionary relationships are resolved . most authors identify a superfamily , alternately called bufonoidea or hyloidea , which includes all the neobatrachians that are not ranoids or microhyloids . sooglossids have often been proposed to be related to the myobatrachids , in the bufonoidea , on the basis of several morphological characters , as well as karyotype and amplectic position . in particular , sooglossids may be sister to the subfamily myobatrachinae , the other subfamily of myobatrachids being more distantly related to both groups . alternately , some authors have hypothesized that the sooglossids are not bufonoids at all , but members of the ranoidea . this relationship is supported by soft tissue characters , and the biogeography of the families involved . finally , recent molecular work suggested an unresolved trichotomy between sooglossids , ranoids , and all other neobatrachians . it seems that the sooglossids will not be well understood phylogenetically any time soon .
duellman , w . e . , and l . trueb . 1986 . biology of amphibians . johns hopkins university press , baltimore , md .
ford , l . s . , and d . cannatella . 1993 . the major clades of frogs . herpetological monographs 7 : 94 - 117 .
hay , j . m . , i . ruvinsky , s . b . hedges , and l . r . maxson . 1995 . phylogenetic relationships of amphibian families inferred from dna sequences of mitochondrial 12s and 16s ribosomal rna genes . molecular biology and evolution 12 : 928 - 937 .
pough , f . h . , r . m . andrews , j . e . cadle , m . l . crump , a . h . savitzky , and k . d . wells . 1998 . herpetology . prentice - hall , inc . , upper saddle river , nj .
zug , g . r . 1993 . herpetology : an introductory biology of amphibians and reptiles . academic press , san diego .
having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .
a large change in the shape or structure of an animal that happens as the animal grows . in insects ,
incomplete metamorphosis
is when young animals are similar to adults and change gradually into the adult form , and
complete metamorphosis
is when there is a profound change between larval and adult forms . butterflies have complete metamorphosis , grasshoppers have incomplete metamorphosis .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
a small family , the seychelles frogs include only four species . they are gardiner ' s frog , which is one of the tiniest frogs in the world ; a species known simply as seychelles frog ; thomasset ' s frog ; and the family ' s newest member , seychelles palm frog , which scientists named in 2002 . the frogs have a typical frog appearance with hind legs that are longer than the front legs , long toes on the hind feet and shorter ones on the front feet , and large , bulging eyes on the head . they also have a somewhat pointy snout and horizontal pupils in their eyes .
the four species come in different colors . the upper body of the seychelles frog is usually yellowish brown with black spots and blotches . the gardiner ' s frog may be reddish brown or tan with or without spots or stripes and sometimes with noticeable , small warts . the newly named seychelles palm frog is light brown with a dark , diamond - shaped pattern in the middle of its back and faded dark patterns on its hind legs . finally , thomasset ' s frog is dark brown to reddish brown with a thin , light stripe running from its snout down the middle of its slightly warty back to its rump . the back also sometimes has small light - colored specks on either side of the line . whatever their color or pattern , however , the four species blend in quite well with their habitat . males and females look alike .
depending on the species , the adults may be very small or medium - sized . the gardiner ' s frog , which is tiny enough to completely fit on a u . s . dime , only reaches about 0 . 4 to 0 . 5 inch ( 1 . 0 to 1 . 3 centimeters ) long from the tip of its slightly pointy snout to the end of its rump . the largest member of the family is thomasset ' s frog . this species usually grows to 1 . 8 inches ( 4 . 5 centimeters ) in length . as in many other types of frogs , the females of each species are a bit larger than the males . for example , a female gardiner ' s frog usually grows to 0 . 47 inch ( 1 . 19 centimeters ) long and sometimes reaches 0 . 5 inch ( 1 . 3 centimeters ) long , while the male typically grows to 0 . 4 inch ( 1 centimeter ) long with a maximum length of 0 . 43 inch ( 1 . 1 centimeters ) .
in 2003 , scientists announced the discovery of a new species of frog , known only by its scientific name— nasikabatrachus sahyadrensis . they placed this species in its own family , but have since decided that its nearest relatives are the seychelles frogs . in other words , the new species and the seychelles frogs have the same ancestors . nasikabatrachus sahyadrensis is a very odd - looking purple frog that apparently stays underground for all but two weeks a year , when it comes out to mate . the world conservation union ( iucn ) considers it to be endangered , or facing a very high risk of extinction in the wild , because it lives in a small area of mountain forests , and its habitat is disappearing as the forest is turned into farmland . as of 2004 , scientists had only found 135 individuals , and only three of those were females .
seychelles frogs live only in the country called seychelles , which is a group of islands in the western indian ocean about 580 miles ( 930 kilometers ) northeast of madagascar . scientists believe that these islands may have been part of india far in the past , but about 55 to 65 million years ago , india began to slowly move away to its current location , about 1 , 800 miles ( 2 , 900 kilometers ) north and now part of asia . like many other islands , the seychelles islands are actually the tops of mountains that are mostly underwater . scientists believe that the common ancestor of the seychelles frogs and the new species of nasikabatrachus sahyadrensis , lived more than 130 million years ago when seychelles and india were still connected . as of yet , they have found no fossils of any of the four seychelles frogs or of the new purple species .
within seychelles , these frogs live only on mahé and silhouette islands , and usually more than 656 feet ( 200 meters ) above sea level , although a single thomasset ' s frog was found lower on the mountain , at about 312 feet ( 95 meters ) above sea level . rainforests are home for all four species . the seychelles palm frog only lives in those areas that have plenty of palms , and thomasset ' s frog likes to remain in forests near rocky streams .
they will eat mosquitoes , fruit flies , and other small insects , as well as mites and other invertebrates ( in - ver - teh - brehts ) , which are animals without backbones , that they find in the forest . thomasset ' s frog also hunts for insects alongside streams .
the four seychelles frogs usually stay out of sight under piles of leaves lying on the rainforest floor , inside cracks in rocks , and even within hollow plant stems or on the base of a leaf where it attaches to a stem . the seychelles palm frog , for example , hunkers down in the leaves of palm and sometimes banana trees . usually , only rains will bring the seychelles frogs out of their hiding places . during these wet periods , the frogs will hop about day or night looking for food . thomasset ' s frog often settles on a streamside rock after sunset and waits for flying insects to zip by closely enough for it to capture and eat them .
the mating season occurs during the rainy season . males may call during the day or at night from under leaves or from one of their other hiding spots . unlike the males of many other types of frogs , males in the seychelles frog family do their calling alone and from their own personal , on - land location . in other frogs , the males often group together in one place—usually in the water—and all call at the same time .
in 2003 , scientists announced the discovery of a new , red - eyed , purple frog that is so unusual , they even created a separate family for it . the frog , which is only known by its scientific name— nasikabatrachus sahyadrensis —was described in national geographic news as
a bloated doughnut with stubby legs and a pointy snout .
villagers in a small village in western india found the odd frog while digging a well and turned the purple creature over to scientists . after studying it , the scientists agreed that it was not only a new species , but was so different that it needed its own family , which is now known as nasikabatrachidae . of all the other frogs in the world , they think it is most closely related to the seychelles frogs , which live 1 , 900 miles ( 3 , 000 kilometers ) away on an island country in the indian ocean .
from the
wrracck toc toc toc toc
of the thomasset ' s frog to the high
peep
of the gardiner ' s frog , each species has its own call . to mate , the male climbs onto the female ' s back and uses his front legs to hang just in front of her hind legs . although scientists do not know how some of these frogs lay their eggs or how those eggs develop into frogs , they do have details about gardiner ' s frog and the seychelles frog species . the female gardiner ' s frog lays her eight to 15 eggs in a hiding place on the ground and stays with them . instead of hatching into tadpoles , these eggs hatch in three to four weeks right into tiny froglets , each one about 0 . 12 inch ( 3 millimeters ) long—no bigger than a grain of rice . as the froglets hop away , the female ' s job is done and she leaves . in the seychelles frog species , the female lays her eggs on land and stays with them just as the gardiner ' s frog does , but her eggs hatch in two to three weeks into tadpoles . without water to swim in , the tadpoles instead wiggle up and cling to the mother ' s back . they stay there until a short while after they turn into froglets , and finally hop off to live on their own .
as yet , scientists are not sure how thomasset ' s frogs or the seychelles palm frogs mate , where the females lay their eggs , whether their eggs develop into tadpoles or right into froglets , and if the adult female or adult male watch over their young . they do know , however , that the female thomasset ' s frog lays large eggs , and they think this may mean that her eggs skip the tadpole stage and hatch right into froglets . all frog species with eggs that hatch into tadpoles go through what is known as indirect development . those whose eggs skip the tadpole phase and develop directly into froglets go through direct development .
according to the world conservation union ( iucn ) , all four species are vulnerable , which means that they face a high risk of extinction in the wild . because all these frogs live in small areas , and some have small populations , changes to their habitat could possibly be dangerous to them . fortunately , many live in morne seychellois national park or inside the boundaries of a conservation project on silhouette island , and some seem to be able to survive in newly growing forests that were once cut down .
physical characteristics : the seychelles frog is yellowish - brown with black spots and blotches on its head , legs , and back . the blotches on its back legs may look like bands . it usually has a triangular - shaped , dark spot that runs from one of its rather large eyes across its forehead to the other eye . its snout is somewhat pointed . the toes on its long hind legs and shorter , thinner front legs have no webbing between them . the females usually grow to about 0 . 8 inch ( 2 cen - timeters ) long from the tip of the snout to the end of the rump , while the typical size of the males is about 0 . 6 inch ( 1 . 5 centimeters ) .
geographic range : seychelles frogs live on mahé and silhouette islands in seychelles , a country in the indian ocean .
habitat : they make their homes in the rainforests of the island mountains at least 660 feet ( 200 meters ) above sea level . they usually hide in leaf piles on the ground , often in areas where cinnamon grows .
diet : seychelles frogs ' diet includes small insects and other invertebrates that the frogs find on the forest floor .
behavior and reproduction : they usually remain hidden , except on wet days and nights . during these rainy periods , they hop out of their hiding spots to search for food . to mate , the males begin calling from under the leaves at any hour of the day or night . a male climbs onto the back of a female and mates with her while hanging onto her body just above her hind limbs . she lays her eggs on land , and her six to 15 eggs hatch into tadpoles , which scramble onto her back . the tadpoles stay on her back and soon turn into froglets . the froglets leave her back to grow up on their own . at one time , scientists thought that the tadpoles rode on the back of an adult male . a closer look , however , showed that it was the female who was the caretaker of her young .
conservation status : according to the world conservation union ( iucn ) , this species is vulnerable , which means that it faces a high risk of extinction in the wild . the frog lives in a small area on only two islands , but it is quite common there . some of the frogs live in morne seychellois national park and others within the boundaries of a conservation project on silhouette island . while it is doing quite well , a change to its habitat could possibly hurt some populations or the entire species . as people move closer to its habitat , conservationists are keeping a watchful eye on this species . ∎
halliday , tim , and kraig adler , eds . the encyclopedia of reptiles and amphibians ( smithsonian handbooks ) . new york : facts on file , 1991 .
noble , gladwyn k . the biology of the amphibia . new york : mcgraw - hill , 1931 .
nussbaum , r . a .
amphibians of the seychelles .
in biogeography and ecology of the seychelles islands , edited by d . r . stoddart . hague : dr . w . junk , 1984 .
' weird stuff ' found in india .
current science january 16 , 2004 ( 89 ) : 12 .
amphibians .
silhouette island . urltoken ( accessed on february 22 , 2005 ) .
the animal life .
virtual seychelles , republic of seychelles . urltoken ( accessed on february 22 , 2005 ) .
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sooglossinae noble , 1931 , biol . amph . : 494 . type genus : sooglossus boulenger , 1906 .
seychelles frogs ( ananjeva , borkin , darevsky , and orlov , 1988 , dict . amph . rept . five languages : 135 ; frank and ramus , 1995 , compl . guide scient . common names amph . rept . world : 114 ) .
seychelles island frogs ( ananjeva , borkin , darevsky , and orlov , 1988 , dict . amph . rept . five languages : 135 ) .
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there are only four species of tiny frogs in this family . they are only found in mountain forests on silhouette and mahé islands in the seychelles . the smallest frog in the world , the sooglossus gardeneri , is in this family . it is less than a half an inch in length .
least concern near threatened vulnerable endangered critically endangered extinct in wild extinct status and range is taken from icun redlist . if no status is listed , there is not enough data to establish status .
key : profile photos video audio gardiner ' s seychelles frog - sechellophryne gardineri gardiner ' s seychelles frog is found on mahé and silhouette islands in the seychelles . source : edge intended audience : general reading level : middle school teacher section : no
gardiner ' s seychelles frog - sechellophryne gardineri gardiner ' s seychelles frog is a little bit under a half an inch in length . source : amphibiaweb intended audience : general reading level : high school teacher section : no
seychelles palm frog - sechellophryne pipilodryas the seychelles palm frog is only found on silhouette island in the seychelles . source : edge intended audience : general reading level : middle school teacher section : no
seychelles palm frog - sechellophryne pipilodryas the seychelles palm frog has only been recognized as a species since 2002 . source : amphibiaweb intended audience : general reading level : high school teacher section : no
seychelles frog - sooglossus sechellensis the seychelles frog is found in leaf litter on the forest floor . source : edge intended audience : general reading level : middle school teacher section : no
seychelles frog - sooglossus sechellensis the seychelles frog is found in wet , small forests . source : amphibiaweb intended audience : general reading level : high school teacher section : no
seychelles frog - sooglossus sechellensis the seychelles frog lays its eggs on land and guards them until they hatch . source : arkive intended audience : general reading level : middle school teacher section : yes
thomasset ' s frog - sooglossus thomasseti thomasset ' s frog is found on mahé and silhouette islands in the seychelles . source : edge intended audience : general reading level : middle school teacher section : no
thomasset ' s frog - sooglossus thomasseti thomasset ' s frog is the largest species in this family of frogs . source : amphibiaweb intended audience : general reading level : high school teacher section : no
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 .
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animal , ( kingdom animalia ) , any of a group of multicellular eukaryotic organisms ( i . e . , as distinct from…
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frost , darrel r . , 1997 : null . amphibian species of the world , a taxonomic and geographic reference . second edition .
parker , s . p . ( ed ) . ( 1982 ) . synopsis and classification of living organisms . mcgraw - hill , new york . 2 volumes .


latibulocrinis is a genus of moths belonging to the subfamily tortricinae of the family tortricidae .

it contains only one species , latibulocrinis curiosa , which is found in brunei and indonesia ( buru ) .

latibulocrinis

tuck 1985 . latibulocrinis , a new genus of tortricini with a sexually dimorphic hindwing anal tuft ( lepidoptera : tortricidae ) . entomologica scandinavica , 16 ( 4 ) 1985 : 325 - 332
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by t . m . gilligan 1 , j . baixeras 2 , j . w . brown 3 , and k . r . tuck 4
urltoken is pleased to offer the complete world catalogue of the tortricidae ( t @ rts ) ! this is a complete list of all world species , utilizing the world catalogue published in 2005 as the foundation for the database . version 3 . 0 of the online catalogue contains 15 , 099 records representing 10 , 883 species . more than 1 , 600 records have been updated from ver 2 . 0 ( jul , 2012 ) , and more than 3 , 000 records have been updated from the original catalogue . the database is completely searchable and contains photos of over 1 , 200 type specimens .
t @ rts will be updated regularly both with corrections from the original world catalogue and with additions since its publication . as such , these pages will serve as the most up to date information on current tortricid nomenclature . if you find any errors in the data presented here or have any questions / comments , please use the contact form to send the authors an email .
we are indebted to all of the original authors of the world catalogue ( j . w . brown , j . baixeras , r . brown , m . horak , f . komai , e . metzler , j . razowski , and k . tuck ) for providing the basis for this project . we would also like to thank the dozens of individuals who have provided corrections or updates to the database since it was first placed online in 2007 .
gilligan , t . m . , j . baixeras , j . w . brown & k . r . tuck . 2014 . t @ rts : online world catalogue of the tortricidae ( ver . 3 . 0 ) . urltoken
1 colorado state university , bioagricultural sciences and pest management , 1177 campus delivery , fort collins , co 80523 , usa 2 institut cavanilles de biodiversitat i biologia evolutiva , universitat de valencia , apartat oficial 2085 , 46071 valencia , spain 3 systematic entomology laboratory - usda [ retired ] , smithsonian institution , p . o . box 37012 , national museum of natural history , washington , dc 20013 , usa 4 curator - microlepidoptera [ retired ] , entomology department ( dc2 - 2n ) , natural history museum , cromwell road , london sw7 5bd , uk
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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 .
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the eastern frogfish , batrachomoeus dubius is a bottom-dwelling fish endemic to coastal eastern australia , from fraser island , queensland to kiama , new south wales .

it is an ambush predator with a large expandable stomach , capable of swallowing crustaceans , molluscs and other fishes whole .

batrachomoeus dubius

dana campbell set
image of batrachomoeus trispinosus
as an exemplar on
batrachomoeus trispinosus ( günther , 1861 )
.
the face of an eastern frogfish , batrachomoeus dubius , in chowder bay , sydney harbour . source : graham short . license : cc by attribution - noncommercial - sharealike
lophius dubius white 1790 , v oyage to new south wales : 265 . type locality : near sydney , nsw .
citation :
eastern frogfishes , batrachomoeus dubius ~ marinebio . org .
marinebio conservation society . web . accessed monday , july 9 , 2018 . < urltoken > . last update : 1 / 14 / 2010 2 : 22 : 00 pm ~ contributor ( s ) : marinebio
ogilby , j . d . 1908 . revision of the batrachoididae of queensland . annals of the queensland museum 9 ( 2 ) : 43 - 57 ( as batrachomoeus minor )
( of lophius dubius white , 1790 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of batrachomoeus minor ogilby , 1908 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of pseudobatrachus dubius ( white , 1790 ) ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
macleay , w . j . 1881 . descriptive catalogue of the fishes of australia . part 2 . proceedings of the linnean society of new south wales 1 5 ( 4 ) : 510 - 629 pls 13 - 14 ( as batrachus dubius )
research batrachomoeus dubius » barcode of life ~ bioone ~ biodiversity heritage library ~ cites ~ cornell macaulay library [ audio / video ] ~ encyclopedia of life ( eol ) ~ esa online journals ~ fishbase ~ florida museum of natural history ichthyology department ~ gbif ~ google scholar ~ itis ~ iucn redlist ( threatened status ) ~ marine species identification portal ~ ncbi ( pubmed , genbank , etc . ) ~ ocean biogeographic information system ~ plos ~ siris ~ tree of life web project ~ unep - wcmc species database ~ worms
richardson , j . 1844 . ichthyology . pp . 1 - 16 pls 1 - 6 , 7 - 8 ( parts ) , 9 - 10 in richardson , j . & gray , j . e . ( eds ) . the zoology of the voyage of h . m . s . erebus and terror under the command of captain sir james clark ross , r . n . , f . r . s . , during the years 1839–43 . london : e . w . janson vol . 2 139 pp . ( as batrachus dubius )
representative hoot train of batrachomoeus trispinosus . ( a ) waveform ( amplitude is in dimensionless arbitrary units ) and ( b ) spectrogram ( hz ) of 13 boatwhistle calls in a train of 36 s in duration . box indicates the position of the individual call represented in c . ( c ) power spectrum of an individual , representative hoot on an expanded time base , shown in the inset ( scale bar represents 250 ms ) . ( d ) close - up of boatwhistle waveform , showing fine - scale call structure . sounds were recorded at 44 . 1 khz .
batrachomoeus trispinosus ( günther ) were acquired through the aquarium trade and maintained in freshwater aquaria ( 150 and 380 l ) at 26°c . the room was maintained on a 13 h : 11 h light : dark cycle along with incandescent moonlight timer replicating the lunar cycle ( solar 1000 dimmer , blueline aquatics , ambler , pa , usa ) . the overhead lights in the aquarium room were off from 20 : 00 h to 07 : 00 h . fish were fed live minnows and small goldfish weekly . all procedures were approved by the institutional animal care and use committee at cornell university .
the spectral properties of the acoustic beats observed here are reminiscent of the spectra for the two - voice / biphonation calls in mockingbirds ( zollinger et al . , 2008 ) . in batrachomoeus calls , the difference in frequency between the two major high frequency peaks , rather than between each of the three peaks , approximates the call ' s modulation rate ; hence our interpretation of the signals as beats rather than as amplitude - modulated signals . interestingly , there is also a significant amount of energy near 200 hz , close to the fundamental frequency of non - beat grunts . whether these non - linear phenomena ( see fitch et al . , 2002 ; zollinger et al . , 2008 ) arise from the activity of one or both swimbladders remains to be investigated .
the eastern frogfish has a large mouth , fleshy lips and a tasseled ' beard ' . the species is endemic to australia , occurring from southern queensland to central new south wales .
the eastern frogfish has a large mouth , fleshy lips and a tasseled ' beard ' . it has a large head with raised eyes and a wide , slightly concave interorbital region . the pectoral fins are large . the three short first dorsal fin spines are reported to be venomous . there is a pair of diverging spines on the upper edge of the operculum .
adult colour varies from a mottled brown to pale grey or a deep blue - grey . juveniles are pale with broad dark bands .
it is an endemic species that occurs from southern queensland to central new south wales . it is relatively common in sydney harbour .
the map below shows the australian distribution of the species based on public sightings and specimens in australian museums . source : atlas of living australia .
it is known from very shallow waters down to at least 150 m in depth . it has been observed sheltering in holes on rocky reefs , kelp beds and silty areas .
hutchins , b . & r . swainston . 1986 . sea fishes of southern australia . complete field guide for anglers and divers . swainston publishing . pp . 180 .
kuiter , r . h . 2000 . coastal fishes of south - eastern australia . gary allen . pp . 437 .
kuiter , r . h . 1996 . guide to sea fishes of australia . new holland . pp . 433 .
thanks for confirming that the fish was a juvenile . i ' ve only ever seen adults . agreed , you don ' t often see this fish in the open . i ' ll email you offline about the video . thank you for trying to post it .
yep , this one was a juvenile . probably about 4 - 5cm across the head . it ' s been in the same hole / spot for about 2 months now . on 26 july , 2009 , there was an adult around that was out in the open . quite strange to see them exposed rather than hiding under a ledge or rock . only got video of it as my camera was being repaired . couldn ' t get the video down to a size that this site would accept .
thank you for posting the image jonathan . you did very well spotting the fish in what appears to be a very silty habitat . is the fish a juvenile ? it certainly appears to be a young individual .
photo taken at chowder bay , 29 august , 2009 , at a depth of 9m .
a mottled brown to pale grey or a darker bluish - grey frogfish with two broad bands and small scattered spots and blotches on the body . juveniles are much paler with broad dark bands .
eastern frogfish have large eyes on top of the broad , flattened head , and an enormous wide mouth surrounded by fleshy flaps and barbels . they have expandable stomachs and can swallow very large prey .
video of an eastern frogfish with young in the shiprock aquatic reserve , port hacking , may 2013 .
endemic to eastern australia , from swains reefs , queensland , to kiama , new south wales , in depths of 1 - 150 m . the species is relatively common in central new south wales .
the eastern frogfish are found in estuaries and on inshore reefs , and have been trawled in 140 m . these bottom - dwellers usually shelter on rocky reefs amongst macroalgae , or under rocks and in caves . they are rarely seen out in the open and are possibly nocturnal .
dorsal fin iii , 19 - 20 ; anal fin 16 - 17 ; pectoral fin 22 - 23 ; caudal fin ( segmented ) 13 - 15 ; lateral line pores 25 - 32 ; vertebrae 9 + 17 - 18 .
colour variable , from a mottled brown to pale grey or a darker bluish - grey , with two broad bands across the body and small scattered spots and blotches . juveniles are much paler with broad dark bands .
ambush predators - feed on a range of crustaceans , molluscs , echinoderms and fishes . they have very expandable stomachs and swallow their prey whole .
females lay their eggs onto rocky surfaces in caves , and she protects the developing eggs and the young for a period of time after they hatch .
castelnau , f . l . de 1875 . researches on the fishes of australia . intercolonial exhibition essays . 2 . pp . 1–52 in , philadelphia centennial exhibition of 1876 : official record . melbourne . ( as pseudobatrachus striatus )
coleman , n . 1980 . australian sea fishes south of 30ºs . lane cove , nsw : doubleday australia pty ltd 309 pp .
de vis , c . w . 1884 . new fishes in the queensland museum . no . 3 . proceedings of the linnean society of new south wales 1 9 ( 3 ) : 537 - 547 ( as thalassophryne coeca )
grant , e . m . 1975 . guide to fishes . brisbane : queensland government , co - ordinator general’s department 640 pp .
grant , e . m . 2002 . guide to fishes . redcliffe : em grant pty ltd 880 pp .
greenfield , d . w . 1999 . family batrachoididae . pp . 1999 - 2003 in carpenter , k . e . & niem , v . h . ( eds ) . the living marine resources of the western central pacific . fao species identification guide for fisheries purposes . rome : fao vol . 3 1397 - 2068 pp .
greenfield , d . w . , winterbottom , r . & collette , b . b . 2008 . review of the toadfish genera ( teleostei : batrachoididae ) . proceedings of the california academy of sciences 4 59 ( 15 ) : 665 - 710
hutchins , j . b . 1976 . a revision of the australian frogfishes . records of the western australian museum 4 ( 1 ) : 3 - 43 figs 1 - 17
hutchins , b . & r . swainston . 1986 . sea fishes of southern australia . complete field guide for anglers and divers . swainston publishing . pp . 180 .
kuiter , r . h . 1993 . coastal fishes of south - eastern australia . bathurst : crawford house press 437 pp .
kuiter , r . h . 1996 . guide to sea fishes of australia . new holland . pp . 433 .
kuiter , r . h . 2000 . coastal fishes of south - eastern australia . gary allen . pp . 437 .
johnson , j . w . 1999 . annotated checklist of the fishes of moreton bay , queensland , australia . memoirs of the queensland museum 43 ( 2 ) : 709 - 762 .
ogilby , j . d . 1906 . honorary curator ' s report . amateur fishing club association of queensland . annual report 1905–6 : 3 - 13 ( as pelophiletor caloundrae )
white , j . 1790 . journal of a voyage to new south wales with sixty - five plates of non - descript animals , birds , lizards , serpents , curious cones of trees and other natural productions . london : debrett 299 pp . 65 pls
whitley , g . p . 1957 . a kennel of frogfishes . australian museum magazine 12 ( 5 ) : 139 - 142
greek , batrachos = frog + greek , momai , mao = to get excited , to be roused ( ref . 45335 )
maturity : l m ? range ? - ? cm max length : 35 . 0 cm tl male / unsexed ; ( ref . 33839 )
inhabits the continental shelf . also found in inshore waters ( ref . 7300 ) .
paxton , j . r . , d . f . hoese , g . r . allen and j . e . hanley , 1989 . pisces . petromyzontidae to carangidae . zoological catalogue of australia , vol . 7 . australian government publishing service , canberra , 665 p . ( ref . 7300 )
) : 16 - 24 . 3 , mean 20 . 4 ( based on 14 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5312 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00389 ( 0 . 00180 - 0 . 00842 ) , b = 3 . 12 ( 2 . 94 - 3 . 30 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 6 ±0 . 5 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : medium , minimum population doubling time 1 . 4 - 4 . 4 years ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : moderate to high vulnerability ( 45 of 100 ) .
urltoken an eastern frogfish and its young filmed at night at a depth of 6 m , shiprock aquatic reserve , port hacking , sydney , new south wales , 26 may 2013 . video © gary dunnett .
if you have images for this taxon that you would like to share with atlas of living australia , please upload using the upload tools .
hutchins , j . b . 1976 ,
a revision of the australian frogfishes
, records of the western australian museum , vol . 4 , no . 1 , pp . 3 - 43 figs 1 - 17
mcculloch , a . r . 1929 ,
a check - list of the fishes recorded from australia . part iii
, records of the australian museum , vol . 5 , pp . 329–436
de vis , c . w . 1884 ,
new fishes in the queensland museum . no . 3
, proceedings of the linnean society of new south wales , ser . 1 , vol . 9 , no . 3 , pp . 537 - 547
ogilby , j . d . 1906 ,
honorary curator ' s report
, amateur fishing club association of queensland . annual report , vol . 1905–6 , pp . 3 - 13
ogilby , j . d . 1907 ,
honorary curator ' s report and list of specimens
, amateur fishing club association of queensland . annual report , vol . 1906–7 , no . 3 , pp . 8 - 14
castelnau , f . l . de 1875 ,
researches on the fishes of australia . intercolonial exhibition essays . 2
richardson , j . 1844 ,
ichthyology
, ed . richardson , j . & gray , j . e . ( eds ) , the zoology of the voyage of h . m . s . erebus and terror under the command of captain sir james clark ross , r . n . , f . r . s . , during the years 1839–43 , vol . 2 , pp . pp . 1 - 16 pls 1 - 6 , 7 - 8 ( parts ) , 9 - 10 , e . w . janson , london
ogilby , j . d . 1908 ,
revision of the batrachoididae of queensland
, annals of the queensland museum , vol . 9 , no . 2 , pp . 43 - 57
urn : lsid : biodiversity . org . au : afd . taxon : 656e088a - 7a2d - 4f09 - ab41 - 0313ad3c4d06
urn : lsid : biodiversity . org . au : afd . taxon : a7bc17c4 - dbda - 4f8d - 8dc2 - 95d95d63784f
urn : lsid : biodiversity . org . au : afd . name : 374436
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in possessing a darker colour and in lacking pinkish bands between the eyes and on the sides of the head . both species have an expandable mouth that can accommodate very large prey . females guard their eggs , which are placed on the ceiling of rock crevices .
occurrence describes how often the species is found on surveys within its distribution . it is calculated as the % of reef sites surveyed by rls divers across all the ecoregions in which the species has been observed
abundance is calculated as the average number of individuals recorded per rls transect , where present .
edit by : gj edgar . 2008 . australian marine life . new holland , sydney
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special publication of the center for biodiversity research and information , no . 1 , vol 1 - 3
greenfield , d . w . / carpenter , kent e . , and volker h . niem , eds
fao species identification guide for fishery purposes : the living marine resources of the western central pacific , vol . 3 : batoid fishes , chimaeras and bony fishes , part 1 ( elopidae to linophrynidae )
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 .
the webpage text is licensed under a creative commons attribution - noncommercial 4 . 0 license
froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of lophius nigricans forster , 1794 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of thalassophryne coeca de vis , 1884 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of pelophiletor caloundrae ogilby , 1907 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
( of pseudobatrachus striatus castelnau , 1875 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
inhabits reefs , mangrove estuaries , and offshore trawling grounds ( ref . 3132 ) . found over muddy bottoms in shallow estuaries and along coasts . marketed fresh ( ref . 12693 ) .
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the batrachoidid vocal repertoire also includes sounds that function in aggression . aggressive vocalizations are usually emitted during nest defense and are distinct from courtship calls in two specific temporal properties : duration and call rate . in agonistic interactions , both male and female toadfishes produce ` grunts ' : short duration , typically non - harmonic calls . in some cases , grunts can be rapidly repeated to form a ` grunt train ' that may last for several seconds ( e . g . see amorim et al . , 2008 ; brantley and bass , 1994 ; gray and winn , 1961 ) . unlike courtship sounds , agonistic grunts are not seasonally dependent . midshipman also produce an amplitude - and frequency - modulated call known as a ` growl ' that has grunt and hoot - like portions like the boatwhistle of some toadfishes ( see above ) , can last for several seconds , and probably functions in agonistic contexts ( bass et al . , 1999 ) .
while broad comparisons have been made across species between the different toadfish sounds and divergent neural and peripheral vocal mechanisms ( amorim et al . , 2008 ; bass and marchaterre , 1989 ; bass and baker , 1991 ; bass and mckibben , 2003 ; dos santos et al . , 2000 ; mann et al . , 2002 ; tavolga , 1958 ; tavolga , 1965 ) , no robust quantitative methods have been used to rigorously investigate the species - level diversity of vocalizations in the family , as have been used in studying acoustics in other vertebrate taxa ( e . g . buck and tyack , 1993 ; clark et al . , 1987 ; mitani and marler , 1989 ; nowicki and nelson , 1990 ; rendell and whitehead , 2003 ; young et al . , 1999 ) . additionally , the majority of this work has focused only on three species within the batrachoididae ( o . beta , o . tau and p . notatus ) ( amorim , 2006 ; bass and mckibben , 2003 ) , despite the moderate diversity of the family , which contains 25 genera and 78 species ( greenfield et al . , 2008 ) ( see also nelson , 2006 ) .
this investigation of b . trispinosus has two main goals . the first is to characterize the vocal repertoire and quantitatively compare it with that of other toadfishes . the second goal is to describe the unique morphology of the sonic swimbladder and its functional implications . by generating a molecular phylogeny of the toadfishes , we demonstrate that b . trispinosus has evolved both a novel vocal signal ( acoustic beat ) and a novel swimbladder ( bilaterally divided ) . the diversity in vocal behaviors and supporting biomechanical mechanisms in this family broadens our understanding of the evolution of acoustic communication in toadfishes , and among closely related groups of fishes in general ( malavasi et al . , 2008 ) .
recordings were made in aquaria using a hydrophone ( aquarian aq - 6 , aquarian audio products , anacortes , wa , usa , frequency response 20 hz to 100 khz ; or 171 / 1 / 5 hydrophone , high tech , gulfport , ms , usa , sensitivity –168 . 2 db re : 1 v μpa –1 , frequency response 2 hz to 30 khz ) suspended 20–30 cm above fish nests . sounds were recorded either through the audacity 1 . 2 . 5 software package ( urltoken ) on a macbook pro or on an olympus ls - 10 digital recorder ( olympus imaging america , center valley , pa , usa ) , at a sampling rate of 44 . 1 khz as 16 - bit wav files . the hydrophone passively recorded in aquaria for 12–15 h periods to detect mainly undisturbed conspecific sounds ; as reported , a few sounds were recorded from individual fish while they were chased by an observer . sounds were then analyzed using raven 1 . 3 ( bioacoustics research program , cornell laboratory of ornithology , cornell university , ithaca , ny , usa ) .
sounds were cross - correlated in raven 1 . 3 using the batch correlator function . sounds were normalized and bandpass filtered between 0 and 5000 hz to reduce the effects of any incidental background noise ( cortopassi and bradbury , 2000 ) . both spectrograms ( hann window , 3 db bandwidth 49 . 6 hz , fft 1280 samples ) and waveforms were cross - correlated . a total of 166 sounds were included in the spcc analysis resulting in 27 , 556 sound comparisons . the resulting output is a similarity matrix , consisting of the similarity score between all possible pair - wise comparisons of sounds . this matrix was converted to a distance matrix ( distance = 1–similarity ) and analyzed with a pco analysis using the pcoord script in the r package ( casgrain and legendre , 2004 ) following the method used by cortopassi and bradbury ( cortopassi and bradbury , 2000 ) .
the relative temporal position of the maximum amplitude of each batrachoidid sound was calculated as a function of its occurrence within the sound ( time of maximum amplitude / duration = relative temporal position of maximum amplitude ) , and differences among species were analyzed with an anova .
specimens preserved in 70 % ethanol ( n = 28 ) were dissected to examine the swimbladder morphology . standard length , body mass and sex were recorded . two specimens were cleared and stained [ following the protocol of song and parenti ( song and parenti , 1995 ) ] and the swimbladder was left in place to allow its morphology to be visualized in relation to the rest of the body . the length and width of swimbladders were measured in situ with calipers and photographed under a dissecting microscope . sexual dimorphism of the swimbladder characteristics was tested with an ancova using jmp 5 . 0 . 1 . 2 ( sas institute , cary , nc , usa ) with standard length as the covariate .
swimbladders were dissected out from the peritoneal cavity ( n = 11 ) and the muscles attached to the walls were dissected free , blotted twice with filter paper , and then weighed . differences in mass between the left and right swimbladder muscles were tested with student ' s paired t - test using the residuals of the swimbladder muscle mass from a regression against swimbladder length ( to account for differences in swimbladder size ) . differences in muscle mass between the sexes were tested using an anova on the muscle residuals .
to visualize the light microscopic structure of the swimbladder and muscles , swimbladders were sectioned at 70 μm on a sledge microtome ( microm hm440e , neuss , germany ) . sections were mounted on glass slides , stained with methylene blue , dehydrated and coverslipped , and then photographed under a microscope at ×4 and ×10 magnification .
over a period of 6 months in captivity ( january to june 2008 ) , a total of 1094 sounds produced by b . trispinosus were recorded . the sounds were initially classified mainly on the basis of the similarity of two temporal characters , duration and repetition rate , that apply to other toadfish sounds ( see introduction ) : short duration grunts and longer duration hoots produced either alone or in succession as grunt trains . together with additional analyses of spectral ( e . g . either broad - band or multiharmonic ) and temporal ( e . g . sound pulse number and inter - pulse intervals ) characters , four major classes of calls were identified : hoot trains , single broad - band grunts , multiharmonic grunt trains , and single broad - band grunts with acoustic beats . of the total calls recorded ( n = 198 calls , comprising 1094 individual sounds ) , 34 . 8 % were hoot trains ( n = 69 trains , composed of 592 individual hoots ) , 48 . 0 % were single grunts ( n = 95 ) , 7 . 1 % were grunt trains ( n = 14 trains , composed of 392 grunts within the train ) and 10 . 1 % were grunts with acoustic beats ( n = 20 ) . there were only three occurrences of single hoots and hence these were not considered a separate class of calls .
) . individual hoot duration ranged from 0 . 285 to 6 . 077 s ( 1 . 15±0 . 04 s mean±s . e . m . ,
= 295 hoots analyzed ) . the mean fundamental frequency of hoots was 151 . 35±0 . 39 hz , while the mean dominant frequency ( either the second or third harmonics ) was 426 . 40±10 . 10 hz . hoots typically had around 10 pronounced harmonics , with the strongest power contained in the fundamental and the first three to four harmonics (
as mentioned above , hoots mainly occurred as trains . the mean number of hoots in a train was 8±1 , and the mean interval between hoots in the train was 1 . 95±0 . 08 s (
= 32 trains ) . over the course of trains , hoots in immediate succession within a sequence significantly decreased in duration ( one - way anova : d . f . = 292 ;
) . however , there was no relationship between the duration of the inter - hoot interval as a function of the temporal position in the train .
temporal patterns in different b . trispinosus vocalizations . ( a ) duration of individual boatwhistle hoots within a train versus the order in which they occur within the train . ( b ) duration of individual grunts within a grunt train versus the order in which they occur within the train . the duration of calls decreases with increasing order within the train ( i . e . later calls are shorter ) , while there is no correlation with interval duration within the train .
= 1 ) . we refer to the latter collectively as single grunts because they had an irregular periodicity when occurring in groups of two to five , unlike grunt trains ( see below ) . singly produced grunts were often coincident with gravel - like movement sounds suggestive of a rapid body movement and an agonistic role as observed in other toadfishes ( e . g .
) . on average , single grunts were much shorter in duration than single hoots within a train , with a mean of 0 . 276±0 . 035 s ( range : 0 . 029–0 . 775 s ,
( a ) waveform , ( b ) spectrogram and ( c ) power spectrum of a representative individual grunt of b . trispinosus . sounds were recorded at 44 . 1 khz .
= 330 grunts analyzed from 12 grunt trains ) . the mean fundamental frequency of grunts within the train was 182±1 hz , with a dominant frequency of 1002±53 hz . the mean interval between grunts was 0 . 353±0 . 014 s . different from single grunts , grunts within trains had a clear and well - defined harmonic structure (
( a ) waveform and ( b ) spectrogram of a representative grunt train ( composed of 45 individual grunts ) of b . trispinosus . boxes in a represent individual grunts within the train in c and e . ( c ) the waveform and ( d ) corresponding power spectrum of an individual grunt at the beginning of the grunt train . ( e ) waveform of an individual grunt in the middle of the grunt train . sounds were recorded at 44 . 1 khz .
the first grunt in a train was significantly longer in duration than subsequent ones ( 0 . 529±0 . 084
) , with no relationship between the duration of the inter - grunt interval and the temporal position in the train .
the fundamental frequency was significantly different between the component sounds within hoot and grunt trains ( one - way anova : d . f . = 520 , f = 1230 . 50 , p < 0 . 0001 ) , with the fundamental frequency of hoots about 30hz lower than that of grunts ( see above ) . the dominant frequency of hoots was also significantly lower than that of grunts either alone or in trains ( one - way anova : d . f . = 571 , f = 63 . 06 , p < 0 . 0001 , tukey ' s hsd post - hoc test : q = 2 . 35 , α = 0 . 05 , p < 0 . 05 ) , but grunts and grunt trains were not different from each other ( tukey ' s hsd post - hoc test : q = 2 . 35 , α = 0 . 05 , p > 0 . 05 ) .
) , to our knowledge previously undescribed in individual fishes . although beats are known for the plainfin midshipman fish , they are formed by the temporal overlap in the advertisement hums of neighboring males ( see
= 20 ) because of their overall duration ( mean 0 . 147±0 . 026 s , range : 0 . 024–0 . 371 s ) and lack of repetition as trains (
) . grunts with beats were recorded from fish in the different sized aquaria ( see materials and methods ) . the short duration of these calls , along with their stable temporal (
) for stability of call structure for individual toadfish ] . moreover , individual fish produced grunts with beats when physically chased around the aquarium with a small net by an observer (
= 4 grunts from two individuals , not shown ) . these grunts had a longer duration ( 0 . 39±0 . 04 s , range : 0 . 31–0 . 482 s ) and much lower dominant and beat frequencies ( 176±1 and 21±2 hz , respectively ) than the conspecifically elicited ones .
examples of two representative grunts exhibiting acoustic beats from b . trispinosus . ( a ) waveform , ( b ) spectrogram , ( c ) power spectrum , ( d ) and waveform with expanded time base of a individual grunt . box in a indicates enlarged section in d . arrows in c represent peaks at 194 , 2196 , 2326 and 2498 hz . ( d ) waveform , ( e ) spectrogram , ( f ) power spectrum and ( g ) waveform with expanded time base of a second grunt , showing beat structure and clear harmonics . box in e indicates enlarged section in h . arrows in g represent peaks at 194 , 2196 , 2348 and 2520 hz . for spectrograms ( b , f ) and power spectra ( d , h ) , fft size = 1028 samples .
) . the number of hoot trains declined in occurrence during the early morning hours until 09 : 00 h . single grunts were produced at higher levels during more hours of the day , and were the most common call type throughout the day , with an elevated number of calls between 19 : 00 and 07 : 00 h (
( a ) diel patterns of calling frequency of different call types in b . trispinosus . horizontal bar beneath x - axis labels indicates light cycle : open bar represents the period of ambient lights on , filled bar represents the period of ambient lights off ( with only moonlight on ) . ( b ) number of different call types over the lunar cycle in b . trispinosus .
the hoot portion of the calls of all species considered exhibited different patterns of amplitude modulation . the time point at which the maximum amplitude occurred within the harmonic , hoot portion of all calls was significantly different among the batrachoidid species sampled (
( a ) representative waveforms of different toadfish vocalizations . horizontal scale bars represent 250 ms ; amplitude of the different calls is not to scale . ( b ) relative temporal position of the occurrence of the maximum amplitude in toadfish harmonic calls , for b . trispinosus ( bt ) , halobatrachus didactylus ( hd ) , opsanus beta ( ob ) , o . phobetron ( op ) , o . tau ( ot ) , and the growl and hum of porichthys notatus ( pn ) .
boatwhistles ) showed that the first three principal coordinates accounted for 27 . 96 % of the total variation ( pco1 : 13 . 83 % , pco2 : 7 . 47 % , pco3 : 6 . 66 % ) . cross - correlation analysis of the waveform produced similar results to spectrographic cross - correlation . vocalizations from within species were clustered closely together in the pco (
formed a distinct cluster apart from all the other species with a positive distribution along pco1 , and some dispersion along pco2 ( 0 . 1 to –0 . 1 ) .
formed clusters extending to the lower left of the plot ( with negative pco1 and pco2 values ) . though there were fewer samples of
hums were tightly clustered near the center of the plot . when maximum amplitude position was regressed against pco1 , the result was a strong and highly significant relationship between these variables (
results from a principal coordinates analysis ( pco2 versus pco1 ) of spectrographic cross - correlation ( spcc–pco ) ( cortopassi and bradbury , 2000 ) of 166 vocalizations from six species of toadfishes . sounds analyzed included harmonic hoots from the boatwhistle vocalizations of b . trispinosus ( filled blue squares ) , h . didactylus ( filled red circles ) , o . beta ( open green circles ) , o . phobetron ( open purple squares ) , o . tau ( open blue triangles ) and the hum of p . notatus ( filled orange triangles ) . due to their temporal similarity to toadfish boatwhistles , ` growls ' of p . notatus ( filled yellow diamonds ) were also analyzed .
) . within the peritoneal cavity , the swimbladder extended anteriorly to the pelvic girdle and posteriorly to the caudal third of the cavity . the intrinsic swimbladder muscles spanned the entire lateral wall of each swimbladder and were whitish in coloration in unfixed material . the vocal nerve inserted on the dorsomedial edge of each bladder muscle . each swimbladder had its own rete mirabele , located on the posterior dorsal end of bladder , rather than a single rete as in other batrachoidids [ described by greene (
( a ) ventral view of a cleared and stained b . trispinosus specimen ( 6 . 6 cm standard length ) showing the bilateral swimbladder . the skin and viscera of the abdominal cavity were removed and the swimbladder was left in place . specimen was cleared and stained for cartilage ( with alcian blue ) and bone ( with alizarin red ) following the protocol of song and parenti ( song and parenti , 1995 ) . scale bar represents 1 cm . the right pelvic and pectoral fins are labeled for reference orientation . ( b ) ventral view of dissected swimbladders from a 9 . 9 cm standard length female , showing asymmetry between left and right bladders . arrows indicate points of attachment , where the bladders are connected to each other by connective tissue . scale bar represents 1 cm .
a single muscle was attached to the lateral wall of each swimbladder . when viewed in cross - section , the lateral wall was concave in smaller fish ( 7 . 1 cm standard length ;
) , becoming more convex in larger body size fish ( 15 . 1 cm standard length ;
) . the bladder muscle was thicker in the smaller fish ( standard length of 7 . 1 cm ) , while it was larger in the dorsoventral axis in larger fish ( standard length of 15 . 1 cm ;
cross - section ( 70 μm thick sections ) of swimbladder wall and intrinsic swimbladder muscle of b . trispinosus , stained with methylene blue from 7 . 1 cm standard length male ( a ) and 15 . 1 cm standard length male ( b ) . the medial side of the swimbladder is to the left . scale bar represents 1 mm for a and b . ( c ) high magnification view of junction of swimbladder wall and muscle from the area represented by the box in a . medial side of the swimbladder is to the left . scale bar represents 1 mm .
) , with the anterior poles of the two adjacent ; however , one side was not consistently longer . of the specimens measured (
= 25 ) , the left swimbladder was longer than the right in 15 fish ( nine males and six females ) , the right was longer in nine ( seven males and two females ) , and only one ( a female ) had equal - sized swimbladders . differences in the width of the left and right bladders did not necessarily correspond to differences in length . of the 15 with the longer left bladder , seven also had a wider left bladder ; of the nine with the longer right bladder , six also had a wider right bladder .
for results from statistical tests ) ; however , while males had wider swimbladders , females had longer ones . the degree of asymmetry of either length or width ( i . e . the difference for each measurement between the two bladders ) showed no strong relationship with body size (
results from analyses of covariance of swimbladders of males and females , with the fish ' s standard length as the covariate . differences in muscle mass were tested with an analysis of variance using the residuals of muscle mass regressed against body size
ontogenetic changes in swimbladder morphology of b . trispinosus males and females . ( a ) length of left swimbladder versus standard length ; ( b ) length of right swimbladder versus standard length ; ( c ) left swimbladder width versus standard length ; ( d ) right swimbladder width versus standard length . solid lines represent regression lines of males ( black ) and females ( gray ) . for all parameters measured , males were significantly different from females ( ancova , p < 0 . 0001 ) ; females had longer swimbladders and wider swimbladder muscles than males , while males had wider swimbladders than females .
there was no consistent difference in the proportional muscle mass ( i . e . corrected for body mass ) between the left and right bladders ( two - tailed paired
) . we attempted to get accurate volumetric measurements for each swimbladder , but were unable to successfully separate the vocal muscle from each bladder and keep the entire bladder intact .
the maximum likelihood analysis resulted in a topology with a negative log likelihood score of 17838 . 52 (
) . likelihood bootstrapping analysis showed high support ( > 75 ) in 9 of 11 nodes .
phylogenetic relationships of toadfishes generated from a maximum likelihood analysis of genes from available sequences ( 16s ; 28s ; cytochrome oxidase subunit i , coi ; cytochrome b , cytb ) in genbank . branch lengths are drawn proportional to the amount of character change . bootstrap values are shown next to nodes . the gadid , gadus morhua , was used as an outgroup for the batrachoididae . boxes around species names indicate taxa used in the comparative sound analysis , and a representative waveform of the species ' call and swimbladder is shown for each taxon analyzed .
like other toadfishes , b . trispinosus produces an array of different vocalizations . however , in contrast to other studied members of the family , b . trispinosus has evolved a novel swimbladder morphology – laterally divided into two separate bladders – and is capable of producing sounds with beats : both unique features among vocal fish . as discussed below , these unique morphological and behavioral adaptations of b . trispinosus may represent one of the most highly derived vocal systems in fishes , convergent with tetrapod taxa in their ability to produce acoustic beats .
) , in which the two halves of the swimbladder are connected by a short hollow tube in the posterior third of the bladders , and perpendicular to their long axis . the available phylogenetic evidence suggests that the swimbladder of
sexually dimorphic bladders have been reported in h . didactylus , o . tau , o . beta and p . notatus ( brantley and bass , 1994 ; brantley et al . , 1993 ; fine , 1975 ; fine et al . , 1990 ; modesto and canário , 2003 ; walsh et al . , 1987 ; walsh et al . , 1989 ) . the sexual dimorphism in the swimbladders of b . trispinosus is complex ; males had wider swimbladders , while females had longer ones . the potential functional significance of differences in either of these dimensions remains unclear . more detailed biomechanical studies of each swimbladder ( e . g . fine et al . , 2001 ) will be required to better understand the role of dimorphisms in sound production .
vocalizations produced by b . trispinosus resemble the major acoustically and behaviorally defined classes of vocalizations described in other batrachoidids , namely agonistic grunts that are brief in duration , and comparatively longer duration , multiharmonic advertisement boatwhistles and hums ( e . g . brantley and bass , 1994 ; dos santos et al . , 2000 ; gray and winn , 1961 ; ibara et al . , 1983 ; tavolga , 1958 ; thorson and fine , 2002b ) . because of the similarities of the acoustical properties of b . trispinosus calls to those of other toadfishes , we suggest that the different classes of calls have a homologous social context : the long duration , multiharmonic hoot serves as an advertisement or courtship call , while the short duration grunts and grunt trains function in aggression and territorial defense . a number of dissected females from the tanks were found to have ripe eggs in the ovaries ( n = 4 ) , which is suggestive of courtship activity and reproductive behavior , as observed in other toadfishes ( e . g . brantley and bass , 1994 ; gray and winn , 1961 ) . the presence of reproductively mature females within the population further supports the hypothesis that the hoots serve a role in courtship behavior .
with the exception of p . notatus hums , b . trispinosus hoots are substantially longer than those in other toadfishes : individual b . trispinosus hoots last up to 6 s ( compared with < 1 s for halobatrachus , opsanus and sanopus ) ( amorim and vasconcelos , 2008 ; amorim et al . , 2008 ; mann et al . , 2002 ) ; the hoot - like hums of p . notatus last upwards of an hour ( bass et al . , 1999 ; brantley and bass , 1994 ; ibara et al . , 1983 ) . the rich harmonic structure of b . trispinosus grunt trains also appears to be distinct from that of most other toadfish ( see above references ) . like other toadfishes , b . trispinosus showed increased levels of calling at night ( e . g . brantley and bass , 1994 ; breder , 1968 ; ibara et al . , 1983 ; thorson and fine , 2002b ) . however , it appears that other toadfish species do not display the same degree of lunar synchrony in calling patterns as b . trispinosus ( breder , 1968 ) .
the quantitative comparison of toadfish vocalizations offers an interesting insight into the diversity and perhaps the evolution of toadfish acoustic communication . taxonomically , the between - genus diversity of toadfish sounds is primarily distributed along pco1 , whereas within - genus diversity (
) . as such , it is impossible to determine which specific components of the sound ( e . g . fundamental frequency , call duration , etc . ) are specifically influencing their distribution in the pco analysis . however , the correlation between amplitude modulation pattern and pco1 suggests that amplitude envelope shape may be influencing the statistical discrimination of the different sounds (
shared the latest position of maximum amplitude in their calls among the species studied . the behavioral importance of amplitude modulation is also shown by playback studies in midshipman fish (
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