lyle 's flying fox ( pteropus lylei ) is a species of flying fox in the family pteropodidae .

it is found in cambodia , thailand , and vietnam .

it can do fairly well in some urban areas .

1 it is killed for bushmeat in parts of its range .

lyle ' s flying fox

lyle’s flying fox ( pteropus lylei ) is a medium - sized flying fox which forms large colonies high up in trees . lyle’s flying fox has a long dark muzzle , large eyes and a ring of orange fur around the neck , and bears some resemblance to a fox , hence the common name ‘flying fox’ ( 5 ) .
bbc two - lyle’s flying fox - thailand : earth ' s tropical paradise , the central heartland - in pictures . . .
farmers also pose a threat as they consider lyle’s flying fox to be a crop pest , resulting in persecution of this species ( 3 ) .
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lyle ' s flying fox is classified as vulnerable ( vu ) on the iucn red list ( 1 ) and is listed on appendix ii of cites ( 4 ) .
the diet of lyle’s flying fox consists mainly of ripe fruit . however , this species will also feed on nectar , pollen and blossoms to ensure it gets enough energy . fruit is very low in protein and sodium , so the salivary gland of lyle’s flying fox has become specially adapted to ensure this species can extract the required nutrients ( 7 ) .
the population of lyle’s flying fox is believed to be in decline , a trend which is expected to continue due to human pressures on its environment ( 1 ) . habitat loss is a major threat to the population ( 1 ) , with deforestation and construction projects irreversibly destroying the forests that lyle’s flying fox relies on for roosting and for food ( 3 ) .
unlike other bat families , fruit bats do not hibernate . instead , lyle’s flying fox produces heat by shivering , which keeps its body temperature between 33 and 37 degrees celsius ( 6 ) .
the wings and back of lyle ' s flying fox are dark brown or black , which strongly contrast against the bright fur around the head and neck . its lower body varies from a deep dark - brown to a brighter yellow - brown . its breast and belly are black - brown , which is similar to the large flying fox ( pteropus vampyrus ) ( 3 ) .
lyle’s flying foxes are the largest bats in the world , as some species can have a wingspan of up to two metres and weigh around 1 . 5 kg .
the lyle ' s flying fox is listed as least concern ( lr / lc ) , lowest risk . does not qualify for a more at risk category . widespread and abundant taxa are included in this category , on the iucn red list of threatened species
lanlua , p . , sricharoenvej , s . , niyomchan , a . and chico , d . e . ( 2007 ) unique cellular structures in the parotid gland of an old world fruit bat , pteropus lylei ( lyle ' s flying fox ) . italian journal of anatomy and embryology , 112 : 179 - 90 .
lyles’s flying fox can be found in some of southeast asia’s most secluded jungles , as well as in the middle of villages or even large urban metropolises such as bangkok . the species usually roosts in temples in the middle of urban areas ( 3 ) .
unlike the majority of animals in the chiroptera order , which are nocturnal , insectivores and orient themselves using ultrasounds , lyle’s flying foxes are crepuscular , feed on fruit , flowers and nectar and use their sight to guide them while flying and to find food .
the habitat of lyle’s flying foxes is limited to thailand , cambodia , viet nam , and the yunnan province of china . even though little is known about populations within china , lyle’s flying foxes have been listed by iucn as vulnerable or one step away from being considered endangered . until recent hunting restrictions were put into place , fruit bats were considered a delicacy and were hunted legally .
hondo , e . , inoue , n . , maeda , k . , rerkamnuaychoke , w . and duengkae , p . ( 2010 ) movement of lyle ' s flying fox ( pteropus lylei ) in central thailand . journal of wildlife in thailand , 17 : 55 - 63 .
lyle’s flying fox is native to southeast asia and is found in thailand , vietnam , cambodia , malaysia , and yunnan in china . this species has been mostly documented in thailand , where at least 11 colonies have been identified , the largest containing around 3 , 000 individuals ( 1 ) .
although it is a nocturnal species , lyle’s flying fox is very sociable and noisy during the day , as this is when females suckle their young ( 5 ) . large noisy colonies are very conspicuous , but they have few natural predators and so can hang safely up in the trees all day ( 6 ) .
lyle’s flying fox’s primary sense when foraging is vision , as it lacks the echolocation abilities of insectivorous bats . it has well developed teeth which are used to chew fruit while spitting out most of the seeds and pulp ( 5 ) . some seeds are ingested , which is important for the local ecosystem as it allows for the dispersal of seeds into other areas ( 2 ) .
lyle’s flying fox , and other species of bat , have been found to harbor large reservoirs of the nipah virus . although the virus does not harm the bat , it can be deadly to humans , and there is substantial data suggesting that recent outbreaks were caused by bat to person transmission of the virus ( 8 ) .
lyle’s flying fox is listed in appendix ii of the convention on international trade in endangered species ( cites ) , so trade in this species should be carefully monitored ( 4 ) . there are no known populations within protected areas in vietnam or cambodia . however , this species is protected by monks in thailand where populations roost in temple roofs ( 1 ) .
lyle’s flying foxes are the largest bats in the world , as some species can have a wingspan of up to two metres and weigh around 1 . 5 kg . they do not have tails , have a claw on the second digit of each wing and have two incisors on each maxilla . their faces are reminiscent of a small fox and , hence , its common name .
hunting still occurs in some places , but habitat degradation and destruction has also contributed to the 30 % population decline over the past 15 years ( three generations ) . in thailand and cambodia , lyle’s flying foxes are considered pests and are not protected except on temple grounds . for this reason , large , noisy colonies of lyle’s flying foxes can be found colonizing the trees over some buddhist temples in cambodia and thailand . up to 11 colonies are known to exist in thailand and the largest contains close to 3000 individuals .
lyle’s flying foxes will colonize trees in large groups , where they nest and suckle their young during the day . at night they depart to forage for food and have been known to travel 40 miles away to find food . rather than using echolocation as their insectivore cousins do , fruit bats rely on well - developed visual and olfactory senses .
in central thailand ' s forests , plains and city streets , nature lives alongside people .
fujita , m . ( 1988 ) flying foxes and economics . bats magazine , 6 : 1 . available at : urltoken
the size of fruit bats can be truly startling when first encountered at night by strangers to tropical climes and has resulted in reports of vampire bats or flying pterodactyls , but unless you’re a fruit or flower , or smell like one , you have little to fear . okay , a little fear is appropriate . over the past 10 years or so several research studies have found that fruit bats , including lyle’s flying foxes , are hosts for emerging viruses that can infect and kill livestock and
) belongs to the large family of mammalian old world fruit bats or flying foxes . the spanish name is zorro valador de lyle . as the common name suggests fruit bats live on nectar , pollen , blossoms , and fruit , and for this reason their habitats are limited to tropical and subtropical regions of the world . old world fruit bats are the largest bats in the world , with wing spans up to 6 feet ( 1 . 8 meters ) and weighing as much as 2 . 2 pounds ( 1 . 5 kg ) . by comparison , the size of lyle’s flying foxes is about mid - sized within the fruit bat family .
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luby , s . p . et al . ( 2009 ) recurrent zoonotic transmission of nipah virus into humans , bangladesh , 2001 - 2007 . emerging infectious diseases , 15 : 1229 - 1235 .
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this species is listed on appendix ii of cites . there are no known populations within protected areas in viet nam ( son nguyen truong pers . comm . ) or cambodia . the species is protected by monks in thailand ( s . bumrungsri pers . comm . ) .
nothing is known from china about the ecology of this species ( smith et al . 2008 ) . in other areas , it is known to form large colonies in trees that can become stripped of leaves by the bats ' activity . it feeds in orchards and is regarded as a serious pest in thailand ( s . bumrungsri pers . comm . ) . it occurs in mangrove forest in viet nam ( son nguyen truong pers . comm . ) . this species travels up to 50 km between colonies ( s . bumrugsri pers . comm . ) .
in thailand the species is threatened through loss of roosting habitat , as existing trees die and are not replaced , it is also subject to hunting ( s . bumrungsri pers . comm . ) . it is also threatened in cambodia by hunting ( p . bates pers . comm . ) .
echolocation detecting objects by reflected sound . used by bats and odontocete cetaceans ( toothed whales , dolphins and porpoises ) for orientation and to detect and locate prey . gland an organ that makes and secretes substances used by the body . hibernation a winter survival strategy in which an animal’s metabolic rate slows down and a state of deep sleep is attained . while hibernating , animals survive on stored reserves of fat that they have accumulated in summer . insectivorous insect - eating . nocturnal active at night .
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hutson , a . m . , racey , p . a . ( chiroptera red list authority ) , chanson , j . & chiozza , f . ( global mammal assessment team )
justification : listed as vulnerable as it is estimated that a decline of over 30 % has occurred in a fifteen year time period ( three generations ) including the past and the present , and this decline is expected to continue due to increasing trade and hunting pressure and ongoing decline in the extent and quality of its habitat .
this species is known from yunnan in china ( smith et al . 2008 ) , extending in to cambodia , thailand , and viet nam .
the largest known colony in thailand is about 3 , 000 individuals and up to 11 colonies have been identified . there are three known colonies in viet nam ( son nguyen truong pers . comm . ) .
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colonies inhabiting the forest roost in the tops of tall canopy trees , which they revisit year after year ( 5 ) ( 6 ) .
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wilson , e . d . ( 1997 ) bats in question : the smithsonian answer book ( second edition ) . smithsonian institution press , washington d . c .
allen , g . m . ( 1980 ) bats : biology , behavior and folklore . harvard university , dover publications inc .
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with an isolated population in yunnan ( china ) and extending through cambodia , thailand and vietnam .
gestation lasts from 140 to 190 days and only one baby is generally born in each birth , occasionally two . the mother takes the baby right when it is born and it remains clinging to her body for several days . after this period , the baby pup remains on tree branches and holes when the mother goes out to feed .
gregarious animals , they form large colonies that come together to rest in the immense trees in the jungle . the largest known colony is in thailand , comprised of some 3000 specimens .
their populations are shrinking considerably in recent years due to direct hunting , deforestation and the progressive deterioration of their habitat . according to the iucn ( international union for the conservation of nature ) , the species has diminished by some 30 % in the last 15 years .
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one specimen was at least 20 . 4 years old when it died in captivity [ 0671 ] .
[ 0730 ] smith et al . ( 2003 ) , body mass of late quaternary mammals
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heterobranchus boulengeri is a species of airbreathing catfish found in the democratic republic of the congo , zambia and zimbabwe .

it is found in lake mweru , the lukonzolwa river and the upper congo river .

heterobranchus boulengeri

heterobranchus boulengeri ( pellegrin 1922 , bu : 147 ) . . . heterobranchus longifilis . . . urltoken 6 eschmeyer reference - new / other species . . . boulengeri . to species summary . heterobranchus boulengeri . . . heterobranchus boulengeri . to species summary . microlepidogaster bourguyi . more
jennifer hammock chose to hide data on
heterobranchus longifilis valenciennes , 1840
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( 1 ) large chrysichthys sharpii 6 , clarias sp . ( t ) heterobranchus boulengeri serranochromis spp . ( b ) ( 2 ) c . sharpii , clarias gariepinus ; c ngamensis ( t ) auchenoglanis occidentalis ( b )
c . ngamensis ; c . gariepinus ; h . boulengeri ; serranochromis sp . 2 . o . mweruensis ( t ) ; tilapia rendalli and clariids ( b )
c . michael hogan selected
range description
to show in overview on
heterobranchus longifilis valenciennes , 1840
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c . michael hogan marked the classification from
species 2000 & itis catalogue of life : april 2013
as preferred for
heterobranchus longifilis valenciennes , 1840
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heterobranchus : from the greek heteros , meaning different and branchos , meaning fin ; in reference to the presence of the two dorsal fins ( rayed and adipose ) . named to honour the belgian - british zoologist , george albert boulenger .
heterobranchus : from the greek heteros , meaning different and branchos , meaning fin ; in reference to the presence of the two dorsal fins ( rayed and adipose ) . the specific epithet comes from the latin longus , meaning long and filum , meaning thread ; in reference to the long barbels of the fish .
teugels , g . g . , b . denayer and m . legendre ( 1990 ) a systematic revision of the african catfish genus < i > heterobranchus < / i > geoffroy - saint - hilaire , 1809 ( pisces : clariidae ) . : zool . j . linn . soc . 98 : 237 - 257 .
head generally longer and much more flattened than in other heterobranchus species . in dorsal outline , spatulate . snout rectangular and lower jaw extending beyond upper . dark brown dorsally , on the flanks and on the upper side of the paired fins , light brown to greyish on the belly and the lower side of the paired fins . underparts of the head can be bright yellow .
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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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confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
diet of adults consists of fishes . juveniles feed on insect larvae and other invertebrates ( ref . 78218 ) .
africa : lake mweru system below johnston falls , including mweru wantipa ( ref . 78218 ) , in democratic republic of the congo and zambia
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600mm or 23 . 6
sl . find near , nearer or same sized spp .
the genital papilla , which is right below the anus , is elongated and pointy in males while it is round and relatively larger ( and larger than the anus ) in females . in ventral view especially , females are much broader in the body than males of equal age and rearing practices .
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pellegrin , j . 1922 . poissons nouveaux ou rares du musée du congo . revue zoologique africaine 10 : 272 - 280 .
chinese academy of fishery sciences ( 2003 ) chinese aquatic germplasm resources database . : urltoken
froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication .
mwape , l . m . ( 2003 ) bemba local names of lake fishes in northern zambia . : p . 246 - 249 . in m . l . d . palomares , b . samb , t . diouf , j . m . vakily and d . pauly ( eds . ) fish biodiversity : local studies as basis for global inferences . acp - eu fish . res . rep . 14 : 281 p .
varjo , m . , l . koli and h . dahlström ( 2004 ) kalannimiluettelo ( versio 10 / 03 ) . : suomen biologian seura vanamo ry .
wu , h . l . , k . - t . shao and c . f . lai ( eds . ) ( 1999 ) latin - chinese dictionary of fishes names . : the sueichan press , taiwan . 1028 p .
hardly anything is known about the biology of this species ( ref . 78218 ) . maximum reported size for west africa : 800 mm tl ( ref . 57129 ) .
africa : nile ( ref . 3820 ) , senegal , gambia , volta , niger , benue rivers and lake chad ( ref . 57129 )
this database was established according to official pieces of work and with the help of famous scientists . however , there might be some errors .
the vernacular names were collected in the field and in the colonial literature from the first part of the 20th century . the monks who established the first dictionaries were not necessarily informed naturalists . therefore , errors must have been committed .
we invite everyone who could help us to improve this working tool to contact us in order to correct us and share her / his knowledge with us .
1500mm or 59 . 1
sl . find near , nearer or same sized spp .
dorsal - fin rays 26 - 35 ; anal - fin rays 42 - 52 ; vertebrae 55 - 61 . head long , broad and somewhat rectangular in dorsal outline ; snout broadly rounded ; eyes with supero - lateral position . frontal fontanelle long and narrow ; occipital fontanelle oval - shaped . toothplates wide . suprabranchial organ well developed . pectoral spine strongly serrated on anterior side . adipose fin with blackish posterior part , nearly as long as dorsal fin ; caudal fin barred . openings of secondary canals hardly visible but display a regular pattern .
africa : nile , niger , sénégal , congo system , upper and middle zambezi . also from lakes tanganyika and edward gambia and bénoué river , chad and volta basins , and the coastal basins of guinea to nigeria .
simply anything digestible it can fit into its mouth . will therefore take all normal aquarium foods .
not really an aquarium fish . although largely inactive and so doesn ' t require a great deal more room than simply to turn around do bear in mind though that this fish can easily reach the size of 1m in captivity .
will attack surprisingly large fish often with fatal results , a greedy , voracious predator . one for a species tank .
during spawning , the male first follows the female , swimming with the head against the side of the female , interspersed with periods of solitary swimming or inactivity ) . the pair then swim head against head with the male on top . this is followed by the male folding the body first around the head , and then the body of the female . during the latter stage , eggs and sperm are released . the female then pushes her head into the substrate and beats her tail vigorously , mixing the eggs and sperm .
hist . nat . poiss . v . 15 - pp394 - pl . 447 poncin , petitfrere , vandewalle & ruwet ( 2002 ) , the reproductive behaviour of the african catfish heterobanchus longifilis ( siluriformes , clariidae ) in an aquarium - preliminary results .
( 1 ) arapaima , ( 2 ) dazza3 , who also notes :
fast growing . can be aggressive . very active . wet pet
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max . size in west africa reported as 510 mm tl ( ref . 57129 ) .
africa : coastal basins from the konkouré ( guinea ) to the cross ( cameroon ) ; present in the upper senegal , but apparently absent from the gambia and the niger ( ref . 57129 )
jennifer hammock removed an association between
zambezi river demersal habitat
and
hippopotamyrus discorhynchus ( peters , 1852 )
.
jennifer hammock added an association between
zambezi river demersal habitat
and
zaireichthys rotundiceps ( hilgendorf , 1905 )
.
management , co - management or no management ? major dilemmas in southern african freshwater fisheries . part 2 : case studies .
the high diversity of methods used could be complementary to each other in exploiting different parts of the fish community . this diversity in fishing patterns can be considered as adaptations to the large diversity of species in the fishery . characteristic of the specialization is that all gears require relatively low - investments ( traps , hooks and lines , etc . ) or , if requiring more investment as in case of gillnets , the gears can be used in a large variety of fishing patterns , thus making them highly adaptable to changing circumstances . all these gears also do not last long , a few years at the most , before they need to be replaced , again contributing to the high adaptability to changing circumstances . the only fishery that requires larger investments , the pelagic light fishery , has converged after initial years of experimentation on a low cost method , both in terms of capital and organization of labour . in the following paragraph we will discuss the result of both this adaptability and the huge increase in effort in particular of the gillnet based fishery .
1 . large variety of species ( t ) 2 . oreochromis mweruensis ( t ) 3 . serranochromis spp . ( t ) 4 . tylochromis mylodon ( t ) ; all areas : clarias or the other cichlids mentioned ( b ) , many other species ( o )
hydrocynus vittatus ; alestes macrophthalmus ( t ) cichlids , clarias spp . ( b ) large barbus and a variety of other species ( o )
o . mweruensis ( t ) serr . spp . ( b ) many other species e . g . hydrocynus , barbus etc . ( o )
t . mylodon ( t ) o . mweruensis and many other species ( b )
m . moeruensis ( t ) ; serr . macrocephalus ( b ) ; a . macrophthalmus ( b )
small and juvenile cichlids ( e . g . pseudocrenilabus philander ) ; small barbus spp . two main types : 1 . ( dominant ) clarias theodorae ; c . buthupogon ( t ) ctenopoma sp . ( b )
tilapia rendalli ( t ) . t . sparmanni ( t ) ; serranochromis sp . ( b ) pseudocrenilabrus philander
1 most active methods are illegal . reported effort through framesurveys for these methods is probably an underestimate ; 2 mapira = gillnetting at mid - water depths ; 3 kutumpula = chasing fish into the net by beating on the water with a knobbed stick ; 4 seining = three types of seining are used . the number of fishermen refers to all methods ; 5 chisense gears = all are light fisheries except chasing . japan = a net hung in between two loose bamboo poles sticking from the side of the canoe , is lowered under the light and hauled inside by at least 2 - 4 people pulling ropes and poles . a boat seine is operated from two boats by 7 - 8 persons ; method comparable to purse seine but the net is closed at the bottom only at hauling onto the canoe ; mutobi = handheld scoopnet operated by one person ; chasing = two women walking at kneedepth in the water who haul a piece of netting or cloth in between ; 6 previously chrisichthys mabusi ( rich , 1986 ) .
figure 9 . development of catch rates in the fishery and in experimental surveys with gillnets of dominant mesh sizes in the fishery ( 76 - 102 mm in 1970s and 63 - 76 mm in 1980s and 1990s ) . catch rates from the fishery are standardized to 100 kg / m gillnet ; catch rates from experimental gear = kg / 100 m 2 . water level is shown as annual deviations of the long term mean . grey bars indicate years over which biomass - size distributions are calculated ( see text )


oenomaus ortygnus , the aquamarine hairstreak , is a species of butterfly of the lycaenidae family .

it is found from brazil through central america to tamaulipas , mexico .

rare strays may be found up to southern texas .

the habitat consists of low elevation wet and dry tropical forests .

the wingspan is 30 – 38 mm .

the upperside is blue with black borders and the underside is gray with a pink sheen .

there are no submarginal or postmedian lines .

there are three or four black spots on the hindwings .

adults are on wing from january to november in mexico .

the larvae feed on annona species .

they bore into and feed upon the buds , flowers and fruits of their host plant .

oenomaus ortygnus

new host plant records for oenomaus ortygnus ( cramer ) ( lepidoptera : lycaenidae ) in mexico .
new host plant records for oenomaus ortygnus ( cramer ) ( lepidoptera : lycaenidae ) in mexico . - pubmed - ncbi
this is the first record of oenomaus ortygnus ( cramer ) damaging fruits of ilama ( annona diversifolia ) and extends the butterfly distribution for three states in mexico .
this is the first record of oenomaus ortygnus ( cramer ) damaging fruits of ilama ( annona diversifolia ) and extends the butterfly distribution for three states in mexico .
oenomaus ortygnus ; [ nacl ] , # 4294 ; [ bow ] : pl . 67 , f . 20 ; [ opler ] ; [ nl4a ] , # 880
calvo r ( 1998 ) reproducción de oenomaus ortygnus ( lepidoptera : lyacaenidae ) en barva , heredia , costa rica . rev biol trop 46 : 101 - 104 . [ links ]
oenomaus isabellae faynel , 2006 ; bull . soc . ent . fr . 111 ( 2 ) : ( 137 - 156 ) ; tl : french guiana
faynel c ( 2006 ) le genre oenomaus hübner , 1819 , en guyane française ( lepidoptera , lycaenidae ) . bull soc entomol fr 111 : 137 - 156 . [ links ]
fennah rg ( 1937 ) lepidopterous pests of the sour - sop in trinidad . ( 2 ) thecla ortygnus cramer . trop agric 14 : 244 - 245 . [ links ]
faynel c ( 2008 ) le genre oenomaus hübner , 1819 , en guyane française . 2 e partie ( lepidoptera , lycaenidae ) . bull soc entomol fr 113 : 15 - 32 . [ links ]
peña & bennett ( 1995 ) reported 296 species of neotropical insects associated with annona ( annonaceae ) , among them , the borers , such as bephratelloides cubensis ( ashmead ) ( hymenoptera : eurytomidae ) , cerconota anonella ( sepp ) ( lepidoptera : elachistidae ) , and oenomaus ortygnus ( cramer ) ( lepidoptera : lycaenidae ) , are the main pests causing economically important damages to the crops .
oenomaus ortygnus has been found in 12 states of mexico ( fig 2 ) ( godman & salvin 1887 - 1901 , kendall 1975 , raguso & llorente - bousquets 1990 ) . the report of kendall ( 1975 ) of females ovipositing and larvae feeding on the fruits of annona globiflora schltdl . ( annonaceae ) in ciudad mante , tamaulipas , is the only information available so far for mexico . to update the status of this pest in mexico , we inspected the most important annona growing areas from march 2008 to february 2009 .
all fruits of a . reticulata and a . diversifolia attacked by o . ortygnus showed necrosis ( fig 1a ) . larvae are usually found under the dead tissue ( fig 1b ) , feeding on the fruit pulp . one to five larvae may be found per fruit . in the laboratory pupation occurred on the external surface of the fruit ( fig 1c ) as observed by calvo ( 1998 ) with a population of o . ortygnus at the estación experimental santa lúcia , barva , heredia , costa rica .
five fruits of annona reticulata l . that showed damage by o . ortygnus were collected in march 2008 in tepalcingo , morelos ( 18º35 ' n , 98º50 ' w , elevation 1169 m ) . six fruits of a . diversifolia safford with the same damages were found in september 2008 in cacahuamilapa , guerrero ( 18º40 ' n , 99º30 ' w , elevation 1163 m ) . three fruits of a . reticulata were attacked by o . ortygnus in february 2009 in zacapala , puebla ( 18º35 ' n , 98º03 ' w , elevation 1254 m ) . collected fruits were placed in plastic containers ( 40 x 21 x 12 cm ) and covered with cheesecloth so that adults could not escape . the samples were incubated at 26 ± 1ºc and relative humidity of 50 % . five adults of o . ortygnus , from larvae reared on these fruits , emerged in the lab .
the present report increases the knowledge about o . ortygnus in two ways : it adds a new record of annona as larval host plant and extends the known distribution of the butterfly for the morelos state in mexico . adults were deposited in the fruit pest collection at fundación salvador sánchez colín cictamex , s . c . , coatepec harinas , estado de méxico , méxico .
although the presence of o . ortygnus has been often noticed on bullock heart ( a . reticulata l . ) , soursop ( a . muricata l . ) and cherimola ( a . cherimola mill . ) ( domínguez - gil 1978 , calvo 1998 , coto & saunders 2001 , beccaloni et al 2008 ) , we found no evidence that this butterfly damaged soursop and cherimola in our surveys . we visited orchards in las varas , nayarit ( 21º11 ' n , 105º08 ' w , elevation 40 m ) and coatepec harinas , estado de méxico ( 18º45 ' n , 99º45 ' w , elevation 2119 m ) in july , august , and november 2008 , other insect pests were recorded attacking a . muricata as b . cubensis and talponia batesi heinrich ( lepidoptera : tortricidae ) on cherimola , respectively .
the authors acknowledge dr . robert k . robbins ( national museum of natural history , smithsonian institution ) for important comments , corrections , and for reviewing the manuscript . for sharing with us information on the geographical distribution of o . ortygnus in mexico , we thank dr . jorge llorente - bousquets , dr . moisés armando luis - martínez , dr . isabel vargas - fernández ( museo de zoología
alfonso l . herrera
, departamento de biología evolutiva , facultad de ciencias , universidad nacional autónoma de méxico ) , and dr . robert k . robbins . for financial support , md thanks fundação de amparo à pesquisa do estado de são paulo / fapesp ( as part of the project
systematics , bionomy , and evolution of neotropical lepidoptera  process number 02 / 13898 - 0 ) and pró - reitoria de pesquisa of the universidade de são paulo / usp / projeto 1 and m . sc . jorge váldez - carrasco for helping with the illustrations . thanks also to anonymous reviewers for valuable comments .
upperside blue with black borders . underside gray with a pink sheen ; no submarginal or postmedian lines . hindwing has 3 or 4 black spots .
eggs are laid singly on the host plant ; caterpillars bore into and feed upon the buds , flowers and fruits .
january - november in mexico , strayed to south texas in mid - december .
brazil north through central america to tamaulipas , mexico . rare stray to south texas .
g4 - apparently secure globally , though it might be quite rare in parts of its range , especially at the periphery .
s . texas , mexico - brazil , surinam , trinidad . see [ maps ]
panama - brazil ( pará , rio de janeiro ) . see [ maps ]
thecla geba hewitson , 1877 ; ill . diurn . lep . lycaenidae ( 7 ) : 198 , pl . 79 , f . 641 - 642 ; tl : ecuador
thecla atena hewitson , 1867 ; ill . diurn . lep . lycaenidae ( 3 ) : 92 , pl . 36 , f . 93 , pl . 37 , f . 101 ; tl : brazil , amazon
e - mail : jshuey at tnc . org ; director of conservation science ; indiana office of the nature conservancy ;
butterflies of north america . 2 . scientific names list for butterfly species of north america , north of mexico .
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .
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urltoken
upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
upperside iridescent greenish - blue with black borders in males ; pale blue in females . underside gray with an aquamarine band along the trailing edge of the hindwing . hindwing has 3 or 4 black spots .
very rare vagrant in the lower rio grande valley of southernmost texas ; normally found from tamaulipas , mexico south to brazil .
adults are nectar feeders ; us records have mostly been on eupatorium betonicifolium . caterpillars found on plants in the genus annona ; no known host plants or documented reproduction in the us .
most recent us record was november 5 2005 , at bentsen - rio grande valley state park . prior to that , november 22 2003 at the naba international butterfly park . two individuals were seen and one collected in brownsville in december 1962 .
wauer , roland h .
butterflies of the lower rio grande valley
boulder co : johnson books , 2004 .
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
contributors own the copyright to and are solely responsible for contributed content . click the contributor ' s name for licensing and usage information . everything else copyright © 2003 - 2018 iowa state university , unless otherwise noted .
naba checklist of north american butterflies occurring north of mexico - edition 2 . 3
[ cite : 1318989 this link is a list of just the names including typo corrections , additions and updates to the following checklist : cassie , b . , j . glassberg , a . swengel & g . tudor . 2001 . north american butterfly association ( naba ) checklist & english names of north american butterflies . second edition . north american butterfly association , morristown , nj .
the second edition of the checklist includes all 722 species of butterflies that have been recorded in north american , north of mexico and in hawaii , giving both english and scientific names .
a catalogue of the butterflies of the united states and canada - - by jonathan p . pelham
taxonomic checklist used on bugguide for butterflies - - does not seem to have a link , so here it is . link updated 16 october 2016 as per comments .
a downloadable pdf file with extensive data on habitats , hostplants , flight dates , and range in the carolinas . you must give an e - mail address to reach the download - - the authors want to inform you of updates . ( this is produced by the north carolina wildlife resources commission , a state agency . ) ( i thought i had posted this before , but cannot find it . )
sammlung europäischer schmetterlinge , errichtet von jacob hübner in augsburg ( 1793–1841 ) : updated plates legend .
seachable database with information on the general classification and the diet and feeding behaviour of the orders of insects and arachnids found in canada’s forest environments . cite : 1463600
an invaluable site for synonyms , links to original descriptions , literature , and other information . covers the entire world , but tends to have more complete information for european and north american species . a great place to begin literature searches . cite : 1329955
the best online resource for moths of great britain and ireland . cite : 1329952
publisher ' s page glassberg , j . 2018 . a swift guide to butterflies of mexico and central america , second edition . princeton university press , princeton , nj . 304 pp . 3 , 250 color photos and maps a groundbreaking photographic field guide to almost all of mexico ' s butterfly species and many of central america ' s this is a revised second edition of a groundbreaking photographic field guide to the butterflies of mexico and central america . written by jeffrey glassberg , the pioneering authority on the field identification of butterflies , the guide covers more than 2 , 000 species and features over 3 , 700 large , gorgeous color photographs , the very best images available , accompanied by authoritative facing - page text . this second edition includes more species , more than 1 , 500 new photos , and updated text , maps , and species names . and range maps , field marks , and host plants are included for all mexican butterflies . the result is an ideal field guide that will enable you to identify almost every butterfly you see .
a systematic revision of some of the american butterflies , with brief notes on those known to occur in essex co . , massachusetts .
full text scudder , s . h . 1872 . a systematic revision of some of the american butterflies , with brief notes on those known to occur in essex county , massachusetts . annual report . peabody academy of science 4 : 24 - 83 .
by kendall , r . o . , & w . w . mcguire .
full pdf kendall , r . o . , & w . w . mcguire . 1984 . some new and rare records of lepidoptera found in texas . bulletin of the allyn museum 86 : 1 - 50 . introduction some of the records treated here have appeared in the lepidopterists ' news ( season summary ) and others in the southern lepidopterists ' news . for historical and biological abstracting purposes they are formalized in this paper , together with appropriate credits and reference citations . it will be noted that these particular species , except for staphylus azteca ( scudder ) and atrytone mazai freeman ( oversightsl , were not included in the 1981 miller and brown catalogue / checklist because their original appearance in the literature was not in a formal publication .
an annotated checklist of the butterflies of bentsen - rio grande valley state park and vicinity .
texas parks & wildlife department , austin . mimeograph pp . 1 - 22 . , 1974
mcguire , w . w . & m . a . richard . 1974 . an annotated checklist of the butterflies of bentsen - rio grande valley state park and vicinity . texas parks & wildlife department , austin . mimeograph pp . 1 - 22 .
the species listed here in are primarily a result of the collecting by the authors during the period 1972 - 1973 . certain important records of the previous several years are also included . additionally , the checklist incorporates records of a number of other lepidopterists . the primary focus of the checklist , then , is upon recent collecting , rather than being an attempt to list all known records from the mid - valley area .
durden , c . j . 1990 . guide to butterflies of austin . texola 6 : 1 - 109 . invaluable resource for information on austin ' s butterflies . see : travis county butterfly checklist
the butterfly fauna of barton creek canyon on the balcones fault zone , austin , texas , and a regional list .
journal of the lepidopterists ' society 36 ( 1 ) : 1 - 17 . , 1982
full pdf durden , c . j . 1982 . the butterfly fauna of barton creek canyon on the balcones fault zone , austin , texas , and a regional list . journal of the lepidopterists ' society 36 ( 1 ) : 1 - 17 . abstract . diversity of substrate , topography and water supply , and climate and vegetation account for the occurrence of 74 % of the regional fauna along a 1 . 1 km stretch of barton creek . monthly mean weather records for 40 years are analyzed on a bioclimagram and the modes matched with habitat types . 172 species are listed for the ten counties around austin , and range , habitat , abundance , and residency are indicated for the 127 found at the study site .
butterflies of the upper frio - sabinal region , central texas , and distribution of fauna elements across the edwards plateau .
journal of the lepidopterists ' society , 52 ( 3 ) : 229 - 261 . , 1998
full pdf gaskin , d . e . 1998 . butterflies of the upper frio - sabinal region , central texas , and distribution of fauna elements across the edwards plateau . journal of the lepidopterists ' society , 52 ( 3 ) : 229 - 261 . abstract . a survey of the butterfly fauna ( 1988 - 96 ) of the upper frio - sabinal region of the southern edwards plateau , texas , is presented . butterflies were observed along transects at five study sites and during repeated opportunistic transects at 12 secondary localities . a total of 28 , 03 . 5 specimens , comprising 100 species was recorded .
geyata ajilvsgi . 1990 . butterfly gardening in the south : cultivating plants that attract butterflies . taylor publishing co . , dallas . xii + 342 pp . = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = select quotes from miller ' s review ( full pdf ) : novice and master gardeners in the southern u . s . , particularly in the rio grande valley , texas , have an extraordinary treat in store with this volume . there are special sections of the book devoted to creating a personal butterfly garden and to methods of attracting butterflies , highlighted by personal observations on such topics as the important characteristics of floral nectaries ( color , shape and fragrance ) and how to choose the appropriate plants , with one of my favorites - adopt a weed .
warning : the ncbi web site requires javascript to function . more . . .
castañeda - vildózola a 1 , nava - díaz c , duarte m , franco - mora o , hernández - fuentes lm .
centro de investigación y estudios avanzados en fitomejoramiento , facultad de ciencias agrícolas , campus el cerrillo , universidad autónoma del estado de méxico , toluca , estado de méxico , méxico . acastanedav @ urltoken
neotrop . entomol . vol . 40 no . 4 londrina july / aug . 2011
beccaloni gw , viloria al , hall sk , robinson gs ( 2008 ) catalogue of the hostplants of the neotropical butterflies . catálogo de las plantas huésped de las mariposas neotropicales . zaragoza , sociedad entomológica aragonesa vol . 8 , 536p . [ links ]
clench hk ( 1964 ) a new hairstreak for the united states . j lepid soc 18 : 189 - 190 . [ links ]
coto d , saunders jl ( 2001 ) insectos plaga de la guanabana ( annona muricata ) en costa rica . man integr plagas 61 : 60 - 68 . [ links ]
domínguez - gil oe ( 1978 ) insectos perjudiciales del guanábano ( annona muricata l . ) en el estado zulia , venezuela . rev fac agron ( luz ) 4 : 149 - 163 . [ links ]
godman fd , salvin o ( 1887 - 1901 ) biologia centrali - americana . insecta . lepidoptera - rhopalocera . fam . lycaenidae 2 : 1 - 112 . [ links ]
janick j , paull re ( 2008 ) the encyclopedia of fruit and nuts . cambridge , cabi publishing , 800p . [ links ]
kendall o ( 1975 ) larval foodplants for seven species of hairstreaks ( lycaenidae ) from mexico . bull allyn mus 24 : 1 - 4 . [ links ]
opler pa , lotts k , naberhaus j ( 2010 ) butterflies and moths of north america . ( urltoken ) , accessed on 11 . ix . 2010 . [ links ]
peña je , bennett fd ( 1995 ) arthropods associated with annona spp . in the neotropics . fla entomol 78 : 329 - 349 . [ links ]
raguso ra , llorente - bousquets je ( 1990 ) the butterflies ( lepidoptera ) of the tuxtlas mts . , veracruz , mexico , revisited : species - richness and habitat disturbance . j res lepid 29 : 105 - 133 . [ links ]
sociedade entomológica do brasil r . harry prochet , 55 86047 - 040 londrina pr brasil tel . : ( 55 43 ) 3342 3987 editor @ urltoken


rhinodipterus is an extinct genus of prehistoric dipnoan sarcopterygians or lobe-finned fish , that lived in the devonian period , between 416 and 359 million years ago .

it is believed to have inhabited shallow , salt-water reefs , and is one of the earliest known examples of marine lungfish .

research published in 2010 based on an exceptionally well-preserved specimen from the gogo formation of australia has shown that rhinodipterus has cranial ribs attached to its braincase and was probably adapted for air-breathing to some degree .

this could be the only case known for a marine lungfish with air-breathing adpatations .

rhinodipterus

( a ) rhinodipterus sp . skull in ventral view . ( b ) rhinodipterus sp . mandible in dorsal view . ( c ) rhinodipterus sp . ceratohyal in lateral view . ( d ) the extant protopterus showing arrangement of cranial ribs attached to neurocranium ( adapted from bemis 1986 ) .
rhinodipterus secans ( gross , 1956 ) : lukševičs , 2001 , lk . 503 , joon .
rhinodipterus secans ( gross , 1956 ) : krupina , 2004 , lk . 394 , joon .
rhinodipterus secans : gross , 1956 , lk . 20 - 32 , joon . text fig . 12 - 23 ; 5 - 7
in griphognathus whitei from the gogo formation , an exact contemporary of rhinodipterus with a similar long - snouted morphology , parts of the cranial cavity and labyrinth spaces ( fig . 4e ) have been described from acid - prepared specimens [ 52 ] . the information is more limited than that for rhinodipterus , but certain comparisons can be made . the olfactory canals , which are very long , merge posteriorly with a short and narrow telencephalic cavity ( [ 52 ] figs . 56 , 61 , 63 ) . proportionately , this cavity appears somewhat smaller than that of rhinodipterus , with a less pronounced ventral bulge ( [ 52 ] fig . 10 ) . there is a large anterodorsally directed pineal recess ( [ 52 ] fig . 10 ) ; this region of the cranial cavity is unfortunately damaged in rhinodipterus , so the two taxa cannot be compared in this respect . the labyrinth cavity and supraotic cavity of griphognathus have been figured in some detail ( [ 52 ] fig . 46 ) . the supraotic cavity is broadly similar to that of rhinodipterus but is not as bulbous and sac - like . however , the labyrinth cavity has an enlarged utricular recess ( fig . 4e ) , very similar to that of rhinodipterus .
palaeozoic global oxygen levels ( berner 2006 ) and the rise of air - gulping . shaded area represents global oxygen levels of 15 % or less . rhinodipterus has been added to the pruned tree of ahlberg et al . ( 2006 ) to show monophyly of air - gulping dipnoans .
as can be seen , these comparisons yield a remarkably consistent picture : on every point , the endocast of rhinodipterus is more similar to the brain of neoceratodus than to the lepidosirenids . given that rhinodipterus is a stem lungfish , this strongly suggests that the similarities reflect retained primitive characters in neoceratodus , relative to a more derived morphology in lepidosirenids . comparison with other members of the lungfish stem group , and with the non - dipnoan sarcopterygians eusthenopteron , tungsenia and spodichthys , support this conclusion and add further information about character evolution in the early part of the lungfish stem lineage .
in chirodipterus wildungensis ( figs . 4c , 5c ) , which comes from the latest frasnian [ 53 ] of bad wildungen , germany , and is thus the youngest of the fossil lungfish discussed here , the telencephalic cavity appears to be somewhat larger and deeper than that of rhinodipterus . the olfactory bulbs were pedunculate , though it is difficult to determine the exact length of the tracts . dorsally there is a prominent pineal recess . the labyrinth cavity of chirodipterus is similar to that of rhinodipterus except that the utricular cavity is slightly smaller ( figures 4 and 5 ) . supraotic cavities have not been described .
our work represents the first virtual endocast of any lungfish . despite being late devonian in age , rhinodipterus kimberleyensis is one of the most derived near - complete fossil lungfish braincases known . not surprisingly , of all the known stem lungfish endocasts , rhinodipterus resembles the similarly - aged chirodipterus wildungesis and griphognathus whitei most closely [ 21 ] , [ 52 ] . with respect to extant taxa , the endocast of rhinodipterus consistently resembles the brain of neoceratodus more than the lepidosirenids , suggesting neoceratodus has retained more primitive characters than the other extant lungfish taxa . the stark difference in morphology between the australian lungfish and the lepidosirenids is again noted . the early evolution of the lungfish stem group is characterized by expansion of the ventral part of the telencephalon , possibly related to evolution of an enhanced sense of smell , and slightly later , expansion of the utricular recess . undoubtedly these changes in labyrinth proportions relate to increased sensitivity for some sensory input , whether it be acceleration , gravitational or auditory , or a combination of all three .
the distinct ‘stepped’ shape of the ceratohyal ( indicated by the ventral notch ) in rhinodipterus ( figure 1 c ) is more similar to neoceratodus ( günther 1871 ) than other devonian genera . the strong posterior margin of this bone indicates a robust connection of the sternohyoideus muscle to the pectoral girdle . the large lateral crest on this bone provides a larger area for interhyoideus muscle attachment , and these features indicate highly controlled mobility of the hyoid arch . as neoceratodus uses its ceratohyals to push air from the branchial chambers into the lung when breathing ( den blaauwen et al . 2005 ) , we suggest that rhinodipterus also had increased capacity for this action when compared with other marine dipnoans .
the form of the sacculolagenar pouch in rhinodipterus is also worth mentioning . most extant fish and amphibians possess separate saccular and lagenar maculae , however modern lungfish are unusual in this respect in having just one [ 65 ] . primitive osteichthyans , including the earliest lungfish and tetrapods , also have a common sacculolagenar chamber as opposed to the large , separate lagena outgrowth seen in modern tetrapods [ 58 ] and actinopterygians [ 47 ] . although rhinodipterus maintains a common sacculolagenar pouch , its ventral margin shows a distinct ventral notch ( figs . 4f , 5d ) . this is particularly unusual as none of the other stem lungfish with labyrinth morphology known appear to do so , nor do the extant taxa .
the presence of the large articulations for cranial ribs , an elongate parasphenoid ( figure 1 a ) and a mobile ceratohyal and pectoral girdle in rhinodipterus indicate air - gulping behaviour . cranial ribs anchor the pectoral girdle during air - gulping in extant forms ( figure 1 d ) and are a key feature in all air - breathing forms ( long 1993 ) .
here we present an investigation by x - ray computed tomography ( µct ) of the braincase and cranial cavity of rhinodipterus kimberleyensis [ 30 ] , [ 31 ] , a recently described stem - group dipnoan from the late devonian gogo formation in northern western australia . the gogo formation is a lagerstätte known for its perfect 3d preservation and high diversity of taxa [ 32 ] , [ 33 ] . rhinodipterus is one of the most crownward stem dipnoans with a near - complete ossified braincase known . lungfish endoskeletons have undergone a radical reduction in ossification since the devonian , possibly as a result of paedomorphosis [ 34 ] , with the result that many post - devonian lungfish are known only from dermal bones and teeth , and none preserves a complete three - dimensional braincase .
holodipterus gogoensis , also from the gogo formation , shows the shape of the braincase particularly well in one weathered specimen ( anu 49102 ) [ 24 ] . the authors have identified numerous nerve canals and approximate endocranial proportions . holodipterus shows two short , robust , and broadly diverging olfactory canals ( [ 24 ] fig . 6b ) , contrasting with the narrower and more elongate canals in rhinodipterus . the space for the telencephalon is relatively short and does not allow for any considerable ventral expansion . like griphognathus , holodipterus also exhibits a large , anterodorsally directed pineal recess ( [ 24 ] fig . 6b ) . the labyrinth region more closely resembles chirodipterus wildungensis ( fig . 4c ) with space for a high superior sinus but lacking a distinctively enlarged utriculus , such as that in rhinodipterus .
when the cranial cavity of rhinodipterus is compared with the brain morphology of the extant lungfish ( figure 1 ) , substantial differences in telencephalic morphology immediately become apparent . all the extant genera have large telencephalic lobes : in protopterus and lepidosiren they are oblong [ 12 ] , [ 16 ] , [ 18 ] , whereas in neoceratodus they are rounder , and very deep ventrally [ 14 ] . the olfactory bulbs are sessile in the lepidosirenids but separated from the telencephalon by short olfactory tracts in neoceratodus ; in all three genera the olfactory bulbs ( or tracts ) attach anterodorsally on the telencephalon . in rhinodipterus the telencephalic lobes must have been much smaller , though the predominantly ventral expansion of the telencephalic cavity suggests a shape somewhat like the brain in neoceratodus , and the olfactory tracts must have been long .
the labyrinth region of modern lungfish are distinguished from those of other gnathostomes by possession of an enlarged utriculus [ 51 ] , [ 52 ] . in the lepidosirenids this is enormous , as big as or bigger than the ( undifferentiated ) sacculolagena , whereas neoceratodus shows a less extreme morphology with a somewhat smaller utriculus ( figure 4 ) . rhinodipterus has a labyrinth cavity resembling that of neoceratodus , except that its utricular cavity is slightly smaller and does not extend so far posteriorly ( figure 4f , g ) . the sinus superior is tall and narrow , as in all modern lungfish . the only surprising feature of the ear of rhinodipterus is the shallow notch , situated approximately between the saccular and lagenar portions , which is not matched either by modern lungfish or by other fossil stem lungfish ( see below ) .
figure 1 shows the characteristic elongated parasphenoid and articulations for cranial ribs of rhinodipterus . the lengthening of the parasphenoid extending posterior to the neurocranium significantly enlarges the buccal cavity , would have enabled the animal to hold a large air bubble . the slot between the tooth plates for a tongue pad ( figure 1 b ) acts as a stop - valve when air is being forced into the lungs ( campbell & barwick 1988 ) .
it is popularly believed that vertebrate air - gulping originated in a tropical lowland setting ( thomson 1969 ) . most fossil lungfish inferred to be air - gulpers have all been from freshwater deposits . the exceptions are the escuminac bay fauna , and partial remains of rhinodipterus from marginal marine environments from the late devonian of europe ( gross 1956 ; ørvig 1961 ; cloutier 1996 ) . usually , marine dipnoi lack the morphological features that would indicate that they were air - breathers ( den blaauwen et al . 2005 ) , and no evidence of cranial ribs in the common late devonian marine gogo fauna ( holodipterus , chirodipterus , griphognathus ) ( miles 1977 ) . however , cranial rib articulations and other features suggesting air - gulping behaviour have been found in a new species of rhinodipterus recently discovered from the shallow marine reef environment of the gogo formation , western australia .
a new species of rhinodipterus from the gogo formation of western australia is known from a single incomplete specimen ( wam 09 . 6 . 149 ) and will be described in detail in a separate paper ( clement in preparation ) . the specimen was prepared by one of us ( j . l . ) using weak acetic acid ( 7–10 % ) , fortified by mowital b30 consolidant , and coated in a sublimate of ammonium chloride for photography .
the pineal and parapineal organs primitively reached close to the skull roof , though there may not have been a pineal foramen ( a foramen is present in dipnorhynchus and powichthys , but absent in all other members of the lungfish total group ) . this condition is characteristic of all known stem group members ( though unknown in rhinodipterus because of specimen damage ) ; the reduced size and height of the pineal - parapineal complex in extant lungfish is derived ( as recognised by stensiö 1963 [ 50 ] ) . lepidosirenids are more derived than neoceratodus in this regard .
figures 2 and 3 show a virtual endocast of the neurocranial cavity of rhinodipterus kimberleyensis . it measures approximately 45 mm in length , and 20 mm at the widest point across the horizontal semicircular canals . the neurocranium is incomplete anteriorly ( snout tip missing ) , and damaged dorsally in the orbito - temporal region [ 31 ] , but otherwise it is complete and uncompressed with many smaller canals for cranial nerves or blood vessels identifiable . the posterior portion of the neural cavity corresponding to the myelencephalic region or hindbrain ( including the labyrinth region ) is particularly well preserved .
together , these patterns build up a picture of rather modest , directional change during the early evolution of lungfish , centered principally on ventral telencephalic expansion and enlargement of the utriculus . this was a prolonged process , as shown by the substantial differences between even the most derived devonian stem group members such as rhinodipterus , and the extant lungfish . stensiö ' s ( 1963 : p . 82 [ 50 ] ) assertion , based on chirodipterus wildungensis , that the extant ‘dipnoan brain had evolved prior to the beginning of the devonian and that since that time it has remained practically unchanged’ is incorrect .
a , youngolepis praecursor ( from chang [ 25 ] , fig . 19 ) ; b , dipnorhynchus sussmilchi ( from campbell and barwick [ 22 ] , fig . 25 ) ; c , ‘ chirodipterus’ australis ( from miles [ 52 ] , fig . 48 ) ; d , chirodipterus wildungensis ( from säve - söderbergh [ 21 ] , fig . 9 ) ; e , griphognathus whitei ( from miles [ 52 ] , fig . 46 ) ; f , rhinodipterus kimberleyensis ( wam 09 . 6 . 149 ) ; g , neoceratodus forsteri ; and h , protopterus annectens ( both from retzius [ 51 ] , plate xxiv ) . not to scale , anterior is to the right .
the middle part of the cranial cavity of rhinodipterus is less informative , partly because it is damaged dorsally , but it nevertheless allows some comparisons to be made with the extant genera . the hypophysis appears to have been oriented ventrally as in neoceratodus , not posteroventrally as in lepidosiren [ 14 ] , [ 18 ] and protopterus [ 12 ] . the shape of the mesencephalic - metencephalic cavity suggests the presence of quite large auricles extending forward on either side of a raised middle region of the brain , as in neoceratodus or latimeria . the raised middle region would have comprised the optic tectum and cerebellum , but unfortunately nothing can be said about their morphology and relative proportions beyond the fact that they were narrow in dorsal view .
the distribution of cranial ribs is known from the middle devonian to recent lungfish . recorded from howidipterus and barwickia from a middle devonian lacustrine site in australia ( long 1992 ) , they are also present in later taxa such as sagenodus ( schultze & chorn 1997 ) and gnathorhiza ( berman 1976 ) . like the extant taxa , these fishes inhabited freshwater systems exposed to seasonal drying or anoxia , possibly caused by rotting vegetation . fleurantia and scaumenacia from the marginal marine late devonian deposits of quebec also possess cranial ribs ( cloutier 1996 ) , their presence in dipterus from the fluvial old red sandstone of scotland is not confirmed ( ahlberg & trewin 1995 ) . schultze ( 1975 ) was first to record the presence of cranial ribs in rhinodipterus .
the evolution of labyrinth morphology over time begs an obvious question : how does it relate to function and behaviour ? the vestibular system in fish is involved in detecting body orientation in three dimensions , and respond to other cues such as gravity and acceleration . clement [ 31 ] previously discussed semicircular morphology and its bearing on locomotive behaviour . the similarity between the labyrinth region of neoceratodus and rhinodipterus ( fig . 4f , g ) drew her to conclude that some late devonian lungfish may have obtained similar functional abilities of the labyrinth as seen in extant taxa . narrower semicircular canals increase sensitivity and are associated with more rapid swimming or accurate maneuverability [ 66 ] . the loss of ampullae , narrowing and heightening ( effectively lengthening ) of the canals - a trend seen over time in lungfish - almost certainly enhances sensitivity .
the semicircular canals have already been described in r . kimberleyensis [ 31 ] , the sacculolagena and utricular recess could not be modeled at that time and were thus neglected in that description . these regions are in fact ossified , well preserved and visible from the scan data , measuring close to 20 mm across their widest point ( fig . 2 ) . the sinus superior is laterally compressed and stands vertically ( fig . 3c ) . both the utricular recess and sacculolagena are large , rounded structures . the sacculolagena of rhinodipterus protrudes ventrally below the notochordal chamber , and there is a distinct ventral notch in the sacculolagena ( fig . 3c ) . an expansion in the anterior semicircular canal exists for the anterior ampulla , but the absence of a significant expansion for the posterior ampulla on the posterior semicircular canal [ 31 ] is confirmed .
description : this is a fine example of the lungfish rhinodipterus . it has long been believed that air breathing developed in lungfish living in freshwater habitats , and then spread to marine taxa . specimens of this genus found in australia last year have raised doubts about this long - held hypothesis . the long mouth cavity of this fish coupled with the articulation of the cranial ribs are much like those of the extant freshwater lungfishes of today . while the invasion of freshwater habitats from the marine realm was previously thought to be the driving force in the development of air breathing , this new data has led some researchers to theorise that a drop in ambient oxygen levels was more likely . examples such as this are rarely available . the repaired crack detracts but little from the specimen offered here , the only one i have ever been able to secure .
rhinodipterus provides the earliest unequivocal evidence of air - gulping adaptations in a marine lungfish . extant lungfish shed little light on the selective pressures of the earliest air - breathing dipnoans . despite the ambiguity of the environment in which air - gulping evolved , this observation raises some interesting phylogenetic implications . do all the air - gulpers belong to a single lineage or did air - gulping evolve more than once in the dipnoi ? such a complex suite of characters as those involved in aerial respiration most probably evolved only once , suggesting that these fishes comprise a monophyletic group ( figure 2 ) , based on the recent phylogenetic analysis of lungfishes by ahlberg et al . ( 2006 ) . we suggest that it was a combination of increased metabolic activity combined with low global oxygen levels that drove the evolution of air - gulping in lungfishes during the devonian period .
the presence of rhinodipterus in marine conditions might indicate that these environments were oxygen stressed . many of the exceptionally preserved specimens from gogo show asymmetry ( e . g . variable shape and number of dermal skull bones ) with high levels of variability often present within the same individual ( long & trinajstic in press ) . increasing asymmetry has been shown to occur in animals in stressed habitats ( parsons 1992 ) . the superb state of preservation of the gogo fossils also indicates quick burial and low oxygen levels . while marine conditions were generally better oxygenated than many freshwater environments , there was still sufficient hypoxia in some marine environments to select for air - gulping ( playford & wallace 2001 ) . a large spiracular notch suggests accessory air intake by comparison with extant forms like polypterus . the tetrapodomorph fish gogonasus shows large spiracular openings on top of the skull ( long et al . 2006 ) , indicating that some other gogo fishes may also have been adapting to hypoxic conditions .
a , youngolepis praecursor ( from chang [ 25 ] , fig . 19 ) ; b , dipnorhynchus sussmilchi ( from campbell and barwick [ 22 ] , fig . 25 ) ; c , chirodipterus wildungensis ( from säve - söderbergh [ 21 ] , fig . 9 ) ; d , rhinodipterus kimberleyensis ( wam 09 . 6 . 149 ) ; and e , eusthenopteron foordi ( from jarvik [ 69 ] , figs . 57 , 59 ) . not to scale , anterior is to the right , grey arrows indicate telencephalic ventral expansion . abbreviations : n . i , olfactory nerve ; n . ii , optic nerve ; n . iii , oculomotor nerve ; n . iv , trochlear nerve ; n . v , trigeminal nerve ; n . vii , facial nerve ; n . viii , auditory nerve ; n . ix , glossopharyngeal nerve ; n . x , vagus nerve ; pin , pineal gland . colour key : green , telencephalic region ; red , diencephalic region ; blue , mesencephalic region ; yellow , metencephalic and myelencephalic regions ; and orange , labyrinth region .
the reconstruction of internal structures in vertebrate fossils has undergone a technical revolution in recent years with the shift from physical serial sectioning or grinding [ 35 ] , [ 36 ] to tomographic techniques based on x - rays or other types of radiation that penetrate the specimen [ 37 ] – [ 42 ] . while brains themselves are only very rarely preserved [ 43 ] , the study of cranial endocasts as a proxy for brain morphology [ 44 ] is a thriving field . of course , one must take care when interpreting endocasts , especially those of fishes where the brain can incompletely fill the neurocranial cavity [ 14 ] , [ 43 ] . nevertheless , endocasts of exquisitely preserved fossils can provide much information about the relative sizes of different regions of the brain , and thus provide a basis for tentative inferences about the sensory and motor capabilities of the animal [ 45 ] . endocasts are also highly character rich , highlighting an underutilized source of anatomical traits for comparative morphology and cladistic analysis . the cranial endocast of rhinodipterus presented here is the first near - complete cranial cavity of a stem lungfish to be imaged tomographically . it demonstrates both the utility of the technique and the high quality of the anatomical data obtainable from gogo formation material .
recent discoveries of tetrapod trackways in 395 myr old tidal zone deposits of poland ( niedźwiedzki et al . 2010 nature 463 , 43–48 ( doi : 10 . 1038 / nature . 08623 ) ) indicate that vertebrates had already ventured out of the water and might already have developed some air - breathing capacity by the middle devonian . air - breathing in lungfishes is not considered to be a shared specialization with tetrapods , but evolved independently . air - breathing in lungfishes has been postulated as starting in middle devonian times ( ca 385 ma ) in freshwater habitats , based on a set of skeletal characters involved in air - breathing in extant lungfishes . new discoveries described herein of the lungfish rhinodipterus from marine limestones of australia identifies the node in dipnoan phylogeny where air - breathing begins , and confirms that lungfishes living in marine habitats had also developed specializations to breathe air by the start of the late devonian ( ca 375 ma ) . while invasion of freshwater habitats from the marine realm was previously suggested to be the prime cause of aerial respiration developing in lungfishes , we believe that global decline in oxygen levels during the middle devonian combined with higher metabolic costs is a more likely driver of air - breathing ability , which developed in both marine and freshwater lungfishes and tetrapodomorph fishes such as gogonasus .
dipnoans , or lungfish , are an ancient lineage of osteichthyan fish that first appeared in the early devonian ( chang & yu 1984 ) . today they are represented by three genera , all of which respire bimodally to some extent ( graham 1997 ) . all exist in freshwater environments exposed to seasonal drying or anoxic conditions . lungfish take in air by gulping a bubble in the mouth , thus a large buccal cavity is an intrinsic feature of all modern air - breathing species .
the gogo formation is an early to mid - frasnian shallow marine inter - reef shale deposit within a large reef complex in the canning basin , western australia ( playford 1980 ) . gogo has yielded an incredibly diverse fish fauna with over 50 species described so far , more than 10 of which are dipnoans .
air - gulping specializations first appeared within the dipnoi during the middle devonian ( figure 2 ) , a time of maximum low global oxygen levels ( berner 2006 ) . air - gulping is thought to have evolved to supplement conventional aquatic respiration during this time of low oxygen ( thomson 1969 ) . their adaptations have also been interpreted as for buoyancy control ( graham 1997 ) , to allow the expulsion of excessive carbon dioxide ( thomson 1969 ) , or to supply the heart with an additional source of oxygen ( farmer 1999 ) . however , in many extant fishes including neoceratodus , activity is a stronger stimulus for air breathing than aquatic hypoxia ( johansen et al . 1967 ) . the question remains as to whether such adaptations were environmentally or metabolically induced .
marine waters are usually better oxygenated than most freshwater environments because of increased wind - driven mixing and tidal flushing ( graham 1997 ) . this trend contributed to the belief that air - gulping must have evolved in hypoxic or anoxic freshwater conditions . however , while hypoxia in marine environments is generally not as extreme , widespread or prolonged as in freshwater habitats , it does still exist . an expansion of hypoxic waters across continental shelves occurred during the warm climate of the palaeozoic ( berry et al . 1989 ) , with hypoxic conditions common in shallow marine and estuarine areas . despite this , few workers considered that aerial respiration could have evolved in marine waters .
ecological radiations of fishes in the devonian are likely to have been associated with increased metabolic costs for greater mobility , and more intensive competition and predation pressures ( graham 1997 ) . the ability to extract additional oxygen could have afforded air - gulping lungfishes the selective advantage of higher rates of metabolic activity , and does not confine their evolution to specific ecological conditions .
we thank k . campbell for useful discussion , g . young for comments on the manuscript , and a . warren who discovered the specimen . this work was funded by arc discovery grant dp0772138 .
structure and phylogenetic significance of diabolichthys speratus gen . et sp . nov . , a new dipnoan - like form from the lower devonian of eastern yunnan , china
the postcranial anatomy of two middle devonian lungfishes ( osteichthyes , dipnoi ) from mt . howitt , victoria , australia
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citation : clement am , ahlberg pe ( 2014 ) the first virtual cranial endocast of a lungfish ( sarcopterygii : dipnoi ) . plos one 9 ( 11 ) : e113898 . urltoken
copyright : © 2014 clement , ahlberg . 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 .
data availability : the authors confirm that all data underlying the findings are fully available without restriction . all relevant data are within the paper .
funding : this work was funded by a wallenberg scholarship from the knut and alice wallenberg foundation , awarded to pea . urltoken . the funder had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
of the extant dipnoans , the neoceratodontid lineage is thought to have diverged from the lepidosirenids ( protopterus and lepidosiren ) as long ago as the permian [ 10 ] , [ 11 ] . as would be expected , the lepidosirenid lungfish are more similar to each other than to neoceratodus , the sole remaining species of the neoceratodontidae . the two lungfish families differ strikingly in a number of morphological features , including their overall body shape , both body and paired fins , skull roof , dentition , and in a number of soft tissue elements including their nervous systems [ 12 ] – [ 18 ] .
a , latimeria ; b , neoceratodus ; and c , protopterus in dorsal view , modified from northcutt [ 16 ] ; and schematic representations of extant sarcopterygian brains showing brain divisions , latimeria in d , dorsal and e , lateral view ( adapted from niewenhuys [ 19 ] ) ; neoceratodus in f , dorsal ( adapted from northcutt [ 16 ] , [ 17 ] ) and g , lateral view ( adapted from holmgren and van der horst [ 14 ] ) ; and protopterus in h , dorsal view ( adapted from northcutt [ 16 ] ) and i , lepidosiren in lateral view ( adapted from collin [ 18 ] ) . scale bar equals 1 cm . abbreviations : a , auricle ; c , cerebellum ; n . ii , optic nerve ; n . v , trigeminal nerve ; n . x , vagus nerve ; t , telencephalon .
the first illustrated endocast of any devonian lungfish was that of chirodipterus wildungensis [ 21 ] . later , a number of uncrushed specimens were used to reconstruct the neurocranial cavity of dipnorhynchus illustrating the relationship between the braincase and the canals for nerves and blood vessels [ 22 ] , [ 23 ] , and again similarly for holodipterus from a single specimen [ 24 ] . partial endocasts of the extinct dipnomorphs youngolepis [ 25 ] and powichthys [ 26 ] , and the porolepiform glyptolepis [ 27 ] are also known , as are those of primitive tetrapodomorphs such as tungsenia [ 28 ] and eusthenopteron [ 29 ] .
clement 2012 ( wam 09 . 6 . 149 ) from the frasnian gogo formation , was acid - prepared by prof . john a . long ( mv ) , and is currently housed at the west australian museum , perth . the uncrushed specimen was scanned at the australian national university ( anu ) high resolution x - ray computed tomography facility [ 46 ] , with a scan resolution of 55 . 5 microns . three - dimensional modeling and segmentation was completed using the software vgstudio max , version 2 . 2 ( volume graphics inc . , germany ) . permits were not required for the described study , which complied with all relevant regulations .
we use the following brain terminology herein : telencephalon and diencephalon comprise the forebrain , the mesencephalon the midbrain , and the hindbrain is composed of the metencephalon and myelencephalon . anteriorly , housed within the olfactory canal , the olfactory tract is an extension of the brain itself ( not a nerve ) . the tract can be expanded into an olfactory bulb anteriorly , usually contained within a nasal sac or cavity [ 47 ] .
( wam 09 . 6 . 149 ) cranial endocast in a , dorsal ; and b , ventral views .
abbreviations : n . ii , optic nerve ; n . v , trigeminal nerve ; n . x , vagus nerve .
( wam 09 . 6 . 149 ) cranial endocast in a , anterior ; b , posterior ; and c , lateral views .
the telencephalic region is long ( at least longer than 14 mm ) and slightly broader than the midbrain cavity ( by approx . 0 . 5 mm ) . anteriorly , the endocast has long , slender olfactory canals that are circular in cross - section . they lie close to each other and diverge only narrowly ( fig . 2 ) . the anterior portion of the snout is missing , so the nasal capsules are unknown . the olfactory canals merge with the telencephalic cavity proper in the posterior third of the telencephalic region ; relative to the canals , the cavity bulges ventrally as well as ( to a lesser degree ) laterally and dorsally .
posterior to the telencephalon , the anterior margin of the diencephalic region is marked by the canals for the optic nerves ( n . ii ) that emerge into the orbits . a small , smoothly rounded ventral protuberance , separated from the telencephalic cavity by a distinct notch , represents the hypophysial fossa ( fig . 2b , 3a , c ) . two pairs of canals for blood vessels emerge from the hypophysial fossa ; anteriorly possibly for the ophthalmic artery and posteriorly possibly for the pituitary vein ( fig . 2b ) . unfortunately this region of the braincase has suffered some damage dorsally , so canals for nerves iii and iv that should emerge from the dorsal part of the cranial cavity are not preserved . the diencephalic , mesencephalic and metencephalic regions are strongly laterally compressed ( between 3 . 5–5 . 1 mm in width ) , all slightly narrower than the telencephalic region ( ≈5 . 5 mm ) .
wide canals for the trigeminal nerves ( n . v ) anterior to the labyrinth region mark the anterior boundary of the myelencephalic region ( fig . 2 ) . while the mesencephalic and metencephalic regions are short , the myelencephalon is longer ( almost the same length as all three preceding regions combined , ≈10 mm ) . dorsally , the supraotic cavities are well defined and carry broad , bulging anterior and posterior prominences ( figs . 2a , 3b ) . in anterior view two deep indentations where the supraotic cavities attach to the cranial cavity can be seen ( fig . 3a ) .
on either side of the specimen , behind the labyrinth region , a thick canal for the vagus nerve ( n . x ) extends posterolaterally . posterior to the vagus nerve , five narrow canals for spinal nerves can be seen extending laterally from the cranial cavity ( figs . 2 , 3b , c ) . moving from the posterior margin of the labyrinth region to the posterior end of the endocast , the cranial cavity gradually narrows by nearly half its width , and also becomes lower . at its most posterior point the cranial cavity has a narrow upright - oval cross - section ( fig . 3b ) .
the canal for the notochord is wide and circular in cross - section posteriorly . it gradually narrows and becomes shallower towards the anterior , while simultaneously flexing ventrally to accommodate the ventral expansion of the mesencephalic - metencephalic region of the cranial cavity ; its anterior end lies just posterior to the hypophyseal fossa ( fig . 2b ) .
obviously care must be taken when interpreting cranial cavity endocasts , especially as the shape of fish brains are generally not constrained as tightly as in other groups such as mammals and birds [ 48 ] , [ 49 ] . however , stensiö [ 50 ] noted that the telencephalic and diencephalic regions in neoceratodus very closely reflected the shape of the cranial cavity . moreover , no part of the brain can be larger than the cavity that houses it , and this simple fact places strong constraints on the inferred brain morphology .
chirodipterus australis from the gogo formation , which has been figured in the same manner as griphognathus whitei [ 52 ] shows some interesting differences from chirodipterus wildungensis . the overall shape of the cranial cavity is similar , again with a well - developed pineal recess , but the telencephalic cavity appears to be proportionately shorter and the olfactory canals are long ( [ 52 ] figs . 17 , 66 ) . in the labyrinth cavity , the utricular recess is much smaller than in the aforementioned taxa ( figure 4c , d ) and the supraotic cavity is small and simple in shape ( [ 52 ] fig . 47 ) . as previously recognized [ 54 ] , these and other features suggest that ‘chirodipterus’ australis is not closely related to c . wildungensis and should not be assigned to that genus , ‘c . ’ australis is probably a less crownward member of the stem group than c . wildungensis .
the stem - group members discussed above are all forms that from a traditional ‘key character’ perspective would be ( and have been ) deemed conventional ‘fossil lungfish’ : they possess major components of the modern lungfish character complex such as autostyly , absence of an intracranial joint , and a palatal bite . this contrasts with youngolepis , powichthys and porolepiforms - considered in the next section - which are assigned to the most basal part of the dipnoan stem group in the majority of recent analyses [ 28 ] , [ 55 ] but are not so obviously lungfish - like . the ‘fossil lungfish’ are all undisputed members of the stem group [ 22 ] , [ 55 ] , [ 56 ] and are always recovered crownward to youngolepis , powichthys and porolepiforms .
the cranial cavity of youngolepis ( fig . 5a ) has been described in its entirety [ 25 ] , whereas in powichthys and porolepiforms ( represented here by glyptolepis groenlandica ) the part posterior to the diencephalon is unknown [ 26 ] , [ 27 ] . the telencephalic regions of all three taxa resemble each other , and differ from those of the ‘fossil lungfish’ as well as extant lungfish , on several points . the telencephalic cavity proper is very short , and the olfactory canals correspondingly long . furthermore , there is no ventral expansion of the telencephalic cavity ; insofar as there is a vertical expansion at all ( in powichthys and glyptolepis , but not in youngolepis ) it is dorsal to the exits of the olfactory canals . in eusthenopteron ( figure 5e ) , tungsenia and spodichthys , which are members of the tetrapod stem group and can thus be used as an outgroup in this context , the telencephalic cavity is longer than in youngolepis , powichthys and glyptolepis , but there is again no obvious ventral expansion [ 28 ] , [ 29 ] , [ 57 ] . all six genera have large anterodorsally directed pineal recesses . the otic region of youngolepis closely resembles that of eusthenopteron ( fig . 5a , e ) : the utricular recess is not enlarged , the sinus superior is low , and the supraotic cavity resembles that of griphognathus .
primitively , the nasal cavities are separated from the telencephalic cavity by long olfactory canals . although the exact location of the olfactory bulb within the cranial cavities of the fossil forms is not strongly constrained ( see jarvik 1980 : fig . 89 , for one attempt at reconstruction [ 29 ] ) , it is clear that pedunculate olfactory bulbs are primitive for the lungfish total group . sessile bulbs are a derived feature of lepidosirenids .
the telencephalon proper is primitively short . all extant lungfish have a longer telencephalon than the stem group members . lepidosirenids are more derived than neoceratodus in this regard .
a ventral expansion of the telencephalon is primitively absent ( as shown by youngolepis , powichthys and glyptolepis ) , but begins to develop in conjunction with other aspects of lungfish morphology such as autostyly and a palatal bite . dipnorhynchus and all more crownward stem lungfish have such an expansion , but it is smaller than that of modern lungfish .
an expanded utriculus develops gradually within the lungfish stem group , crownward of dipnorhynchus . the utriculi of extant lungfish are larger than those of any stem group members , but once again the lepidosirenids are much more derived than neoceratodus .
the neuroanatomy of extant lungfish allow us to draw some tentative conclusions about the functional significance of the telencephalic expansion . in neoceratodus the part of the telencephalon that lies ventral to the plane of the olfactory tract consists entirely of the subpallium [ 14 ] . this region of the brain appears to deal substantially with olfaction , and we can probably conclude that the ventral expansion of the telencephalon during lungfish evolution related functionally to the development of an enhanced sense of smell [ 19 ] . conversely , the mesencephalon , which receives visual input , is proportionately small in both stem and crown lungfish , relative to most actinopterygians and chondrichthyans .
there are surprisingly few studies on lobe - finned fish labyrinths , despite the likely insights they could provide into the transition of the early tetrapods [ 58 ] – [ 60 ] . retzius [ 51 ] was the first to describe them in lungfish , millot and anthony [ 61 ] for the coelacanth . later authors have expanded this work [ 62 ] – [ 65 ] . latimeria is considered to have an labyrinth region “with tetrapod affinities” [ 62 ] , whereas the dipnoan labyrinth has been described as having a mix of chondrichthyan and lissamphibian features [ 64 ] .
the functional significance of the striking utricular expansion in the dipnoi is surprisingly difficult to determine . early literature considered the utricule most important for postural control ( particularly horizontal movement ) , whereas the sacculolagena was considered to have a more auditory role [ 67 ] . however , more recent literature suggests that all otolith organs may have both auditory and vestibular roles [ 67 ] , often varying between different fish species [ 68 ] . although conclusions can ' t be drawn without experimental evidence , the changing proportions of labyrinth organs likely indicate an altered sensitivity of the vestibular system with respect to posture control , and may or may not have had an additional role in hearing .
thanks are due to a number of people , firstly prof . john long who led the 2008 museum victoria / australian national university gogo expedition , funded by arc discovery grant dp 0772138 , during which dr . anne warren discovered the specimen , prof . john long also prepared the specimen . we are indebted to prof . t . senden of australian national university for scanning the specimen , and many thanks are also due to dr . sophie sanchez for assistance with three - dimensional modeling and rendering of the specimen . the manuscript was improved by three helpful reviews , and amc wishes to acknowledge dr . tom challands for valuable discussions concerning lungfish endocast morphology .
conceived and designed the experiments : amc pea . performed the experiments : amc . analyzed the data : amc pea . contributed reagents / materials / analysis tools : amc pea . wrote the paper : amc pea .
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