the african pitta ( pitta angolensis ) is an afrotropical bird of the family pittidae .

it is a locally common to uncommon species , resident and migratory in the west , and an intra-african migrant between equatorial and southeastern africa .

they are elusive and hard to observe despite their brightly coloured plumage , and their loud , explosive calls are infrequently heard .

the plump , somewhat thrush-like birds forage on leaf litter under the canopy of riparian or coastal forest and thickets , or in climax miombo forest .

they spend much time during mornings and at dusk scratching in leaf litter or around termitaria , or may stand motionless for long periods .

following rains breeding birds call and display from the mid-canopy .

african pitta

stamps showing african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis african pitta pitta angolensis 109 . 009 african pitta ioc v2 . 4 : 4217 links will open countrypage in new window - cambodia 01 . 12 . 1994 birds ms - gabon 15 . 10 . more
african pitta ( pitta angolensis ) is a species of bird in the pittidae family .
nsabagasani c . a migratory species , african pitta “pitta angolensis” passed buhanga relict forest .
the african pitta ( pitta angolensis ) also called angola pitta is certainly one of the top prizes for birdwatchers going for africa .
pittas of the world . a monograph on the pitta family | african bird club
museum specimens indicate historical distributions . the map below shows locations from which museum specimens of african pitta were collected . you can see more information on the individual museum specimens of african pitta here .
we will dedicate 3 mornings and afternoons to try and photograph the elusive african pitta in the zambezi valley .
the interest for the african pitta migration of 2012 started with a possible siting on the pugu hills nature centre trail during the eastern holidays . due to doubts about this african pitta a query among the
dar birders
revealed that actually an african pitta had landed in the
peninsula
, the part of dar es salaam with adequately invitive gardens .
the african pitta’s are intra - african migrants and they migrate to the zambezi valley each year between late nov and early april to breed . one can only really locate the pitta’s at the start of the rainy season when they begin their breeding cycle .
the untamed african wilderness – both south luangwa and kasanka national parks are unfenced with free roaming animals that offers the authentic african wilderness experience .
possible hybrid with green - breasted pitta as breast more green than buff and voice was as pitta reichenowi . however shows . . .
english : green - breasted pitta , black - headed pitta ; french : brève a capuchon ; german : kappenpitta ; spanish : pita encapuchada .
the african pitta is classified as least concern . does not qualify for a more at risk category . widespread and abundant taxa are included in this category .
english : black - and - scarlet pitta , black - crowned pitta ; french : brève gacieuse ; german : granatpitta ; spanish : pita de corona negra .
the african pitta is a very special bird ; it is one of 32 other pitta species . it is very aggressive and will do anything to be the best pitta around . it is known as “the holy grail of african birds” because it is very shy , even the best birdwatchers have a hard time catching a glimpse of this bright bird . it communicates in a repeating trail of two note whistles , trills , or grunts . they are often compared with thrushes because of the similarities . the african pitta is diurnal and migrates at night . this makes it hard for them to see buildings as they fly over african cities . this is the reason why in kenya , the number of african pittas have decreased because lighted buildings pose a threat at night . african pittas make themselves feel at home by using fallen trees and branches as perches because they are birds that stay on the ground rather than in the air . in the rare occasion that it is sighted , the african pitta will usually be seen in zimbabwe . when it walks , it flits its tail from side - to - side in a very arrogant way . in the zambezi valley , the breeding of the african pitta has been affected by the large number of elephants and also by increased farming .
the pitta did not allow anymore photographs in the
wild
after being released .
rheindt , f . e . , and j . a . eaton . 2010 . biological species limits in the banded pitta pitta guajana . forktail 26 , 86 - 91 .
malawi 1992 birds 75t african pitta used sg 892 watch this auction meet the seller seller not yet uploaded a picture . borucha feedback tier 5 ( 251 - > 750 rating ) address verified . more
unique opportunities – this photo safari offers the only time and place where these three events , the elephants through reception , african pitta , and biggest mammal migration in the world , can be photographed .
the african pitta , pitta angolensis , is a species of bird in the pittidae family . it is found in angola , burundi , cameroon , central african republic , republic of the congo , democratic republic of the congo , ivory coast , equatorial guinea , ethiopia , gabon , ghana , kenya , liberia , malawi , mozambique , nigeria , rwanda , sierra leone , south africa , tanzania , togo , uganda , zambia , and zimbabwe . references - * birdlife international 2004 . pitta angolensis . more
from 3000 m² in the african pitta to 10 , 000m² in the rainbow pitta . pittas will perform territory defence displays on the edges of their territories , although fights between rivals have only been recorded once . migratory species will defend non - breeding feeding territories in addition to their breeding ones . more
dutson , g . , and j . newman .
observations on the superb pitta ( pitta superba ) and other manus endemics .
bird conservation international 1 ( 1991 ) : 215–222 .
the coming of the rains heralds “change” . the african pitta arrives in the zambezi valley to breed . migrants abound in the bush and hundreds of migrants flock to feed on the new bounty that the rains bring in . come with us in search of one africa’s most elusive bird species the african pitta , we will journey to the zambezi valley to try to locate it at the start of its breeding season .
south african special forces ( elite ) and over 2 million other books are available for amazon kindle . learn more
but i also want to take images . to photograph the african pitta is still a very different thing . i worked hard for one hour and more . i kept changing my positon with my guides support on the edges of the bush . unfortunately the african pitta seemed a bit reluctant . not really shy , but at least it could not be approached easily . i tried to stay under the bush , which was located on a termite mound . after a while laying calm in the heat , i really could take fabulous looks , took pictures and even flash the african pitta after some time . finally the pitta came so close to me that even the 300 - lens with its close focus was to big . really i was very impressed . a thrill experience !
the african pitta , pitta angolensis , is a species of bird in the pittidae family . it is found in angola , burundi , cameroon , central african republic , republic of the congo , democratic republic of the congo , ivory coast , equatorial guinea , ethiopia , gabon , ghana , kenya , liberia , malawi , mozambique , nigeria , rwanda , sierra leone , south africa , tanzania , togo , uganda , zambia , and zimbabwe .
kari pihlaviita marked the finnish common name
afrikanpitta
from
pitta angolensis vieillot 1816
as trusted .
the southern african bird atlas project ( sabap1 ) in namibia gathered a huge amount of distribution data between 1970 and 1993 .
modern african wars vol . 2 : angola and mozambique , 1961 - 74 ( men - at - arms no 202 )
english : black - backed pitta ; french : brève superbe ; german : mohrenpitta ; spanish : pita soberbia .
english : angolan pitta ; french : brève d ' angola ; german : angolapitta ; spanish : pita africana .
jewel of the forest – it’s not difficult to understand why the spectacular , elusive and rare african pitta is the most sought after bird for photographers in africa , and now for the first time you’ll get a realistic chance to photograph it .
pittas are found alone or in pairs and are territorial . territories may vary widely in size depending on the species and habitat ; african pitta ( pitta angolensis ) territories may be as small as 0 . 75 acre ( 0 . 3 ha ) , rainbow pittas ( pitta iris ) defend areas larger than 2 . 5 acres ( 1 ha ) , and for some species only a single pair may be found in an area as large as 50 acres ( 20 ha ) .
english : bengal pitta ; french : brève du bengale ; german : neunfarbenpitta ; spanish : pita de alas azules .
english : black - breasted pitta ; french : brève iris ; german : rogenboenpitta ; spanish : pita arco iris .
modern african wars vol . 3 : south west africa ( men - at - arms series 242 ) : south west africa vol 3
english : black - breasted pitta ; french : brève de gurney ; german : goldkehlpitta ; spanish : pita de gurney .
african pitta pitta angolensis = described by : vieillot ( 1816 ) alternate common name ( s ) : angolan pitta old scientific name ( s ) : none known by website authors photographs no photographs are available for this species range w . , cw . and se . africa ; three populations ; ( 1 ) sierra leone , se . guinea and liberia e . to w . cameroon . ( 2 ) sw . cameroon to nw . angola . ( 3 ) e . zaire , c . and e . more
erritzoe , j . ( 2018 ) . african pitta ( pitta angolensis ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
the african pitta is an intra - african breeding migrant , arriving in southern and south - central africa to breed around november - december , then departing around march - april for its non - breeding grounds in equatorial africa . in southern africa it is scarce and localized , mainly occupying evergreen forest or dense thickets , often on the banks of rivers or streams , foraging for invertebrates in the leaf litter . more
rozendaal , f .
species limits within the garnet pitta - complex .
dutch birding 16 ( 1994 ) : 239–245 .
like humans . pitta chicks don’t beg for food , they just wait with their mouth open until food is given to them .
modern african wars ( 1 ) : rhodesia 1965 - 80 : rhodesia , 1965 - 80 no . 1 ( men - at - arms )
this book provides a highly detailed account of the history , organisation , uniforms and insignia of south african special forces from their origins up to the early 90s – units such as the 44 parachute brigade , the hunter group and the infamous swa police coin unit ' koevoet ' . these elite units of the south african defence force and the special anti - terrorist units of the south african police forces comprised the largest , best trained and best equipped of any country in southern africa . robert pitta and jeff fannell provide the text in a volume packed with photographs and illustrations .
this book provides a highly detailed account of the history , organisation , uniforms and insignia of south african special forces from their origins up to the early 90s - units such as the 44 parachute brigade , the hunter group and the infamous swa police coin unit ' koevoet ' . these elite units of the south african defence force and the special anti - terrorist units of the south african police forces comprised the largest , best trained and best equipped of any country in southern africa . robert pitta and jeff fannell provide the text in a volume packed with photographs and illustrations .
gretton , a . , m . kohler , r . lansdown , t . pankhurst , j . parr , and c . robson .
the status of gurney ' s pitta ( pitta gurneyi ) 1987–1989 .
bird conservation international 3 ( 1993 ) : 351–367 .
it african pitta exclusively eats invertebrates , usually foraging in leaf litter , searching for prey . once it spots a prey item it stabs it with its bill , killing it instantly . its diet has not been studied very well , however it is thought to eat the following prey items :
the domed nest typical of the pitta family is the size and shape of a
rugby football .
both the male and female participate
the asian mainland also has impressive pittas , including blue pitta ( above in an excellent shot by j . j . harrison ) . this pitta appears to be at the spot in khao yai where some elusive species become accustomed to being fed mealworms . we visited the spot in january 2013 — by then a rare orange - headed thrush geokichla citrina greeted us but the ' acclimatized ' blue pitta did not appear .
i was there in the green season from december to march . this time is one of the prime birding season in malawi . summer is the best season to experience avian diversity to the max with visitors like palearctic migrants and even african pitta . the best month is said to be december – if there are abundant rains . i was lucky , as there are years with no sighting of the angola pitta in malawi at all .
on sunday april 29th , 2012
an annual african pitta hunt in dar
was organised in the pugu forest and very successful with three sightings in a part of the forest which hardly deserves the name any longer . the picture shows one of this year ' s annual visitors to pugu .
os - c checklist of the birds of east africa . ornithological sub - committee of the east african natural history society , nairobi ( in press ) .
pittas have always been thought of as a cohesive group of 24 - 32 species ( e . g . , lambert & woodcock 1996 , erritzoe 2003 ) . irestedt et al . ( 2006 ) used molecular evidence to show that there were three major clades within the family , and divided them into three genera . the first - named genus pitta had the 14 species . the two african species are in this genus , as is indian pitta p . brachyura . pittas in this genus have green upperparts with a blue wing - patch and contains all the migratory species , like blue - winged pitta p . moluccensis and noisy pitta . noisy pitta ( left , in a photo be arthur grosset ) breeds in eastern australia in the southern summer and then ' winters ' in new guinea . many of the island endemics are presumably derived from migratory species .
african pitta is reliably found here in december when they migrate into riverine forests to breed . our zambia tour focuses on the zambezi endemics found primarily in the miombo woodlands cloaking large parts of the country . the tour also targets zambia ' s only two endemics , chaplin ' s barbet and black - cheeked lovebird . we have run a conservation tour for the african bird club . for more information on our zambia birding tour itineraries , please contact us . more
[ african tailorbirding cc ( ck2003 / 020710 / 23 ) trading as birding africa ] p . o . box 22727 , scarborough , 7975 , south africa .
distribution of african pitta in southern africa , based on statistical smoothing of the records from first sa bird atlas project ( © animal demography unit , university of cape town ; smoothing by birgit erni and francesca little ) . colours range from dark blue ( most common ) through to yellow ( least common ) .
a number of world checklists within the last 50 years listed as few as 24 species . this was after the great heydays of lumping in the mid - 20th century . this contrasts with the lists of of sclater and elliot at the end of the 19th century that contained 48 and 47 species , respectively ( erritzoe 2003 ) . now , in 2014 , we are back with the numbers envisioned by 19th century ornithologists . it began with the book on pittas and relatives ( lambert & woodcock 1996 ) that split sula pitta e . dohertyi from the widespread indo - pacific red - bellied pitta on morphological grounds . then rheindt and eaton ( 2010 ) split banded pitta hydrornis guajanus is split into three species : malayan banded - pitta h . irena , bornean banded - pitta h . schwaneri , and javan banded - pitta h . guajanus . these splits were widely accepted ( as was the split of black - headed e . ussheri from garnet pitta e . granatina in the sundaic region ) , bringing the list to 34 species .
the former
red - bellied pitta
was widespread in range from the philippines to the solomon island . the other indo - pacific pitta with numerous races is hooded pitta ( left ; a photo from papua new guinea by steve wilson ) . there are at least 12 races , some black - crowned and some chestnut - crowned . no one has yet done a molecular phylogeny of this complex . are there still more
new
pittas to come ?
mackworth - praed c . w . and grant , c . h . b . ( 1964 ) african handbook of birds . series 1 . vols 1 & 2 .
the pittidae is a fairly small family of mid - sized , short - tailed , long - legged ground - dwelling jewels in the old world tropics . i ' ve spent hours trying to track down calling birds inside bornean forests , and encounters there with blue - headed pitta hydrornis baudii , giant pitta h . caerulea , and black - headed pitta were absolute highlights ( see also the bottom of this page ) . here ( left ) is a very nice shot of black - headed pitta by gareth knass . if pressed , i ' d choose pittas as my favorite family of birds in the world .
our guides are at least 6th generation african . who better to show you the african continent than people who live there ? they have explored their home and neighbouring african countires as travellers even before digital photography and workshops became popular , and with a camera have spent more than 10 years chasing the best light in south africa , namibia , botswana , zimbabwe , zambia , tanzania and kenya . therefore they know the people , the culture and we have developed relationships with the local people that gains them access to special places .
some years ago i spent 3 days at the end of the so - called green ( i . e . rainy ) season in the liwonde national park . i decided to stay in the mvuu lodge . one of the highlights of the area of the lodge and the mvuu camp is the irregular appearance of the enigmatic african pitta (
irestedt et al ' s ( 2013 ) proposal of the 17 - way split was adopted by the clements and ebird world checklists in 2014 , bringing the world pitta total to 49 species . as there are several other endemic pittas in the philippines , e . erythrogaster was not called
philippine pitta
but , instead , the old name blue - breasted pitta was reasonably adopted for this new philippine endemic . personally , i had recorded 4 of the 17 taxa in the old
red - bellied pitta
complex , but one of those was
heard only ,
so my net was only two extra lifers , with some 14 still unseen .
almost all pittas breed seasonally , with breeding timed to coincide with the onset of the rainy season . an exception to this pattern is the superb pitta ( pitta superba ) , which apparently nests throughout the year on the island of manus . in most species there are relatively few unique displays prior to copulation , and most pittas probably are monogamous . however , the african pitta performs a unique display prior to the breeding season . during display bouts , this species repeatedly jumps about 10 in ( 25 cm ) into the air , parachuting back to the perch with several shallow wing - beats . during this display the red belly is prominently displayed and the birds often give a
prrt - wheet
vocalization .
deep in the gloom of an old world jungle a pitta stands motionless , giving a characteristic whistle . you , a potential observer , know that if you could see it , it will be a memorable experience , as pittas are wonderfully colorful . but they are also characteristically elusive , and more often than not , the pitta disappears unseen . photography would seem impossible . indeed , for all my visits to tropical rainforests , this shot of sulawesi pitta ( above ) is my best effort . [ sulawesi pitta is a recent split from red - bellied pitta ; more on that below ] . the collection of photos below are from other photographers — i stand in awe at their inspiring accomplishments . alas , much of my time in old world was pre - digital . digital photography has made pittas a bit more accessible — yet , still , each encounter is memorable .
most pittas are nonmigratory or make local movements outside the breeding season . however , indian pittas ( pitta brachyura ) , blue - winged pittas ( p . moluccensis ) , and fairy pittas ( p . nympha ) , as well as a subspecies of the african pitta ( p . a . longipennis ) and populations of hooded pittas ( p . sordida ) and red - bellied pittas ( p . erythrogaster ) are migrants . although most species migrate over land , it is believed that the fairy pitta may fly nonstop from vietnam across the ocean to borneo , a flight of approximately 620 mi ( 1 , 000 km ) ! given the short , rounded wings of pittas , it is somewhat surprising these birds make long migratory flights .
pittas are so wonderful that i list two pittas — gurney ' s pitta h . gurneyi and giant pitta h . caeruleais — among my
top 50 birds
in the world . gurney ' s has an exceptionally limited range along the s . thailand - burma border and is gorgeous . it is critically endangered in thailand but there is more hope for the population in remote burma . a further eight species ( including the elusive giant pitta ) are listed by the iuc as vulnerable . the main threat to pittas is habitat loss by deforestation .
reichenow , 1901 – se tanzania s to e zambia , n & e zimbabwe and s mozambique ; migrates n to region from s central african republic and n & ne drcongo e to s kenya and n tanzania .
erritzoe , j . ( 2018 ) . superb pitta ( pitta superba ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
a very short summary of south african special forces with lots of photographs and illustrations showing the aforementioned forces . when read with other books about this time and place of conflict throughout southern africa it definitely enhances the knowledge base .
it is an intra - african breeding migrant , arriving in southern and south - central africa to breed around november - december , then departing around march - april for its non - breeding grounds in the drc , uganda and kenya .
pitta joffe , a botanist by profession with a keen interest in horticulture , authored a number of popular books on indigenous plants , including the first edition of this book . pitta ' s plant photographs have appeared in numerous publications on the subject . tinus oberholzer holds a degree in horticulture and has had experience in propagation and production of nursery plants as well as plant marketing . he has contributed to the writing and editing of various gardening books , and also writes monthly articles for some south african gardening magazines . tinus is co - owner of plantae orchids , a nursery specialising in orchids and rare and unusual plants , with a special interest in indigenous flora .
our image database has been created to bring together photographs taken in the african region of as many bird species as possible . we hope this will provide both a useful tool for researchers and a splendid collection of photographs for anyone to browse .
experience three once - in - a - life opportunities in one dedicated photo safari . join c4 photo safaris and specialist wildlife photographic guide , isak pretorius , to help you take world class photos of elephants walking through mfuwe lodge reception , the spectacular and elusive african pitta , and a bat migration where 10 million fruit bats creates the largest mammal migration on earth . what makes this even better is that you’ll experience this in the wild and untamed wilderness of zambia’s south luangwa and kasanka national parks .
historically , this species was trapped for the cage bird trade . however , with increasing awareness of the dwindling population size , the economic value of gurney ' s pitta has begun to shift from illegal trade to conservation - based ecotourism .
the paoc12 scientific programme committee has nominated the following symposium topics for the 12th pan - african ornithological congress . to find out what each symposium incorporates click on the symposium title ; titles not hyperlinked do not have outlines at present but will be added in due course .
one of the ' new ' species accepted out of the split of the former
red - bellied pitta
is on sulawesi . my distant photo of a half - hidden adult is at the top of this page . here ( below , a photo by jason thompson from wiki - commons ) is a very fine photo of a sulawesi pitta in juvenal plumage , just started to molt into adult plumage with new green , blue , and red feathers appearing on the underparts .
the african pitta is an intra - african breeding migrant , arriving in southern and south - central africa to breed around november - december , then departing around march - april for its non - breeding grounds in equatorial africa . in southern africa it is scarce and localized , mainly occupying evergreen forest or dense thickets , often on the banks of rivers or streams , foraging for invertebrates in the leaf litter . its nest is a dome - shaped structure made of twigs , leaves and plant debris , usually placed in the uppermost branches of a tree sapling . it lays 2 - 4 eggs , which hatch into chicks with black skin and orange bills . strangely enough , the chicks do not beg for food , they just patiently wait with their mouths open until they are given something .
once we arrive at bilimungwe bushcamp , we’ll have lunch and then a chance to unpack and settle into our rooms . after high tea we’ll go on our first game drive in search of the african pitta and everything else that will make great photos . the afternoon game drives extend into the evenings for stunning night drives . here we’ll have the chance to photograph nocturnal animals , like leopard , genets , owls and elephant shrews , using a spotlight . after the night drive we’ll return to camp for dinner and bed .
handle seamlessly the logistical challenges that african touring presents . expertise , personalised service , attention to detail and best value tours are why , after 13 years , we ' re arranging more birding tours for top international companies and small groups than ever before . in sum , have a look at our
earthworms figure prominently in the diets of many pittas , especially during the nesting season . in australia , the diet of the rainbow pitta varies seasonally ; earthworms comprise most of the diet during the wet season , while other invertebrates are more important during the dry season .
recommended citation birdlife international ( 2018 ) species factsheet : pitta angolensis . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .
family of birds . it is an elusive bird with an unknown lifespan . it is unknown if they are good at camouflage since they are hard to find even by experienced birdwatchers . some think they may be very shy . . it is a special bird with very characteristic features , which include : very long legs , large feet , and sharp talons . they are 6 . 5 to 7 inches tall , they have a beak that is 1 inch long and looks like a pointy sunflower seed and they weigh 3 oz . with their scarlet belly , black stripe over their eye , green and blue feathers , orange - buff breast , and dark brown crown they are very colorful and most pittas will have up to 9 colors on it . even though they are so colorful , you rarely get to see an african pitta . interestingly , both the male and the female pitta look the same .
there are also two pittas in tropical africa . this one ( below ) is green - breasted pitta , in another fine shot by gareth knass . again , i have spent a day searching for it at a
known
location in uganda , and again the great bird has eluded me .
the final genus , hydrornis , includes some variable oriental species that are sexual dimorphic in plumage and have a cryptic juvenile plumage in the species that have been studied . this genus includes eared pitta which had often been placed into its own genus [
anthocincla
] because of its alleged primitive characters .
then , irestedt et al . ( 2013 ) did a complete review of the entire red - bellied pitta set . they used both nuclear and mtdna , combined with vocal and morphological evidence , and
based on all available evidence
they proposed that the e . erythrogaster complex is composed of 17 species .
at 6 : 00 all ( few ) guests of the lodge had booked a trip on foot . the guide was richard . we rattled the whole area with the scrub vegetation for the pitta and a broadbill ( smithornis sp . ) . unfortunately in vain . instead , for example we observed grey - headed kingfisher (
a late morning road transfer from mfuwe lodge to bilimungwe bushcamp means that we’ll have time for a short morning drive at mfuwe lodge before we change camps . bilimungwe bushcamp is a three hour drive south from mfuwe lodge in the south luangwa national park , on the banks of the luangwa river . this is where we’ll look for the african pitta . there are a number of breeding pairs in the thickets close to the camp , and although they are still elusive , their mating calls usually give their positions away . photographers in the past few years have had the best success their in photographing them than anywhere else in africa . the game viewing around bilimungwe bushcamp is great , so we will spend the next few days looking for these birds while on game drives , with a good chance of seeing and photographing more than just the bird . once we arrive at bilimungwe bushcamp , we’ll have lunch and then a chance to unpack and settle into our rooms . after high tea we’ll go on our first game drive in search of the african pitta and everything else that will make great photos . the afternoon game drives extend into the evenings for stunning night drives . here we’ll have the chance to photograph nocturnal animals , like leopard , genets , owls and elephant shrews , using a spotlight . after the night drive we’ll return to camp for dinner and bed .
this tour is tailored towards photographers who are keen on bird photography and passionate about birding in general . photographers can be from a beginner level to advanced dslr users . we will search for the pitta in its breeding location , and spend quite a bit of time specifically focussing on attempting to photograph this rare and special bird .
with its concentration and diversity of wildlife , south luangwa national park offers an unrivalled safari experience . during november and mid december , three “once - in - a - lifetime” opportunities present themselves to safari goers in zambia . go in search of the “jewel of the forest” – the rare , migratory african pitta , while also being able to witness the unique , annual phenomenon of elephants regularly walking through mfuwe lodge reception . another highlight will be when we are joined by renowned , dutch - born ornithologist , frank willems to watch the world’s greatest migration – the highest density of mammals on earth – as 10 million straw - coloured fruit bats descend upon zambia’s kasanka national park… just a short flight away .
experiences with species this much sought after bird is difficult to locate unless it is calling , which occurs in a small time window , which is usually shortly after the first rains in early december . it hops on the branch , making a loud sound with it ' s wings , followed by a frog - like squeak . once it lands , it looks around , quite proud of itself , as if to say
did you see what i just did ? !
. click here to watch a video of the pitta displaying - be warned you may get seasick , a handheld video on a big lens does not work ! other names : angola pitta
another clade was named erythropitta . these are mostly small species with short tails , lots of red on the underparts , and greenish or bluish backs . there was initially six species in this genus , but that has now tripled when red - bellied pitta e . erythrogaster was split into 17 species ( more on that below ) .
with its concentration and diversity of wildlife , south luangwa national park offers an unrivalled safari experience . during november and mid december , three “once - in - a - lifetime” opportunities present themselves to safari goers in zambia . go in search of the “jewel of the forest” – the rare , migratory african pitta , while also being able to witness the unique , annual phenomenon of elephants regularly walking through mfuwe lodge reception . go one step further and extend your safari to join renowned , dutch - born ornithologist , frank willems to watch the world’s greatest migration – the highest density of mammals on earth – as 10 million straw - coloured fruit bats descend upon zambia’s kasanka national park… just a short flight away .
serious climate change challenges for birds and other biodiversity in africa are much less well understood than those in europe , north america and australia . given the magnitude of these challenges in africa , especially as compounded by existing conservation threats from land use change , invasive species , pollution and harvesting , a focus on the vulnerability of african birds and bird habitats is critical . we propose drawing on the insights from detailed biodiversity / climate change work in europe and australia to kick - start a wave of accelerated , collaborative and more detailed demographic , behavioural , ecological , biogeographic and conservation research on the vulnerability of african birds and bird habitats to climate change . we also hope to supplement and follow the symposium with an in - depth round - table discussion on this theme .
robert pitta has been a writer and editor in the defence / aerospace industry for over sixteen years . he is a former professional photographer and holds an advanced university degree . he has given lectures to the military and written on the history , development , and use of camouflage clothing and concealment techniques . he lives and works in maryland , usa .
in a recent visit to thailand , guide nik upton located a pair of eared pitta ( right ; photo by n . upton ) deep in the understory . like many pittas , it was not particularly shy but just cryptic and very difficult to view after each hop across the littered ground . this photo is of the adult female of the pair .
creative gardening with indigenous plants , now in its second edition , is an invaluable handbook for south african gardeners . in this comprehensive guide the authors set out to popularise the use of indigenous south african plants in gardens , parks , on roadsides and anywhere where previously exotic plants were preferred . this book covers more than 300 plants , all illustrated by stunning photographs in full colour showing the whole plant as well as selected features such as flowers , fruit , leaves and bark . the text comprises a description of each plant , its usages - including medicinal uses and snippets of folklore - and advice on cultivation . introductory chapters cover topics such as propagating plants from seeds and cuttings , gardening in the different climate regions , and using indigenous plants to attract birds and insects to the garden . taking a very practical approach , using symbols , distribution maps , plant size and flower colour , the authors lead the reader to the correct plant to choose for the application , whether this be tree , shrub , annual , perennial , climber or water - loving plant . this all - inclusive book is a must for gardeners , horticulturists , landscape designers and anyone involved in the south african nursery industry .
bird interactions with industry e . g . power lines and aircraft is an important factor to consider both from a conservation as well as an industry safety and quality of supply perspective . little has been done on the african continent with respect to bird power line and bird aircraft interactions . in south africa the endangered wildlife trust has pioneered the establishment of two unique industry partnerships to forge relationships with industry and to establish programs to effectively deal with the issue . as more and more power lines are constructed across the african continent and airports adhere to international safety standards , knowledge needs to be shared and assistance provided on how to deal with bird industry interactions ! the aim of such a symposium will be to present papers from what has been achieved in africa and to plan what still needs to be done .
the ' fffffrrrt ' sounds on the recording , which are the displaying pitta ( rapid wing - flapping ? ) , were detected from xc419773 by michael mills of go - away - birding . recording made before dawn , but , as michael puts it , pittas quite often start displaying when it is still dark . many thanks to michael for this cute contribution !
my personal best memory was at danum valley , borneo , and i wrote of it in my daily notes that evening ( below , notes from 4 aug 2003 ) . i really really really wanted to see the endemic blue - headed pitta , so after a rainstorm one afternoon , i hike out a trail to a spot where we had heard one the day before . i sat down quietly near the end of a log , and briefly played a tape . to my shock ,
the male blue - headed pitta appeared at the end of the log where it looked around , shook itself , and puffed out breast / belly ( see sketch ) . absolutely crippling views . it hopped down log towards me a bit before flying off .
a late morning road transfer from mfuwe lodge to bilimungwe bushcamp means that we’ll have time for a short morning drive at mfuwe lodge before we change camps . bilimungwe bushcamp is a three hour drive south from mfuwe lodge in the south luangwa national park , on the banks of the luangwa river . this is where we’ll look for the african pitta . there are a number of breeding pairs in the thickets close to the camp , and although they are still elusive , their mating calls usually give their positions away . photographers in the past few years have had the best success their in photographing them than anywhere else in africa . the game viewing around bilimungwe bushcamp is great , so we will spend the next few days looking for these birds while on game drives , with a good chance of seeing and photographing more than just the bird .
chris gooddie accomplished his goal of seeing all the pittas at the time when 32 species were widely accepted as the full set . now there are 49 species and counting . i wondered how many he still has left to see if he used the newest taxonomy . so i wrote him and the answer was that , except for the
red - bellied
group , none . he was aware of pending splits in
banded pitta
and potential splits in elegant pitta , so he saw all of them except the vigorsii race of elegant [ tanimbar and kai islands in eastern indonesia ; this was once split by clements but is lumped again in recent editions ] . chris says that he has continued to travel and see additional races of several pittas , but he is still 12 short of the entire 17 - species - set of the now - split
red - bellied pitta .
he agreed that it is likely there are some
cryptic species
in the complex , but thought that there is
so much vocal variation
within the known taxa , and so much to be learned about the remote island birds , that it was
too early to split red - bellied complex .
if i expanded by
top 50
birds in the world to a hundred , i ' d surely add several more pittas . so many are just drop - dead gorgeous , including rainbow pitta of northern australia ( right , in an exceptional shot by david fisher ) . they are all shy and secretive , with ethereal voices that draw one far from trails into primeval tropical forests in the old world .
although classification at the family level is widely accepted , there are conflicting opinions regarding the appropriate number of genera and species . although as many as six genera have been proposed , and preliminary estimates of genetic divergence support these distinctions , most authors have chosen to recognize only the genus pitta . recent taxonomic treatments recognize 29–31 species . this number will undoubtedly change as molecular methods generate a better understanding of the evolutionary history of the pittidae .
ringing activities have been conducted in africa for 60 years , and ringing is an important tool in continuing studies on the biology , survival and movement of birds . ringing has often been used in survey work , particularly in forest habitats . this symposium endeavours to present overviews of some recent work , showing how ringing will still be relevant as a tool to study african birds for decades to come . 2008 marks the 60th anniversary of safring , thus the paoc is an appropriate time to review ringing activities .
returning home , i even tried to do a painting of the encounter with the blue - headed pitta ( below , now embedded in color among my field notes ) . still , i ' m far from chris gooddie in terms of success — i ' ve barely seen a quarter of the world ' s pittas . just another good reason to take yet another long trans - pacific flight to a forest in the lowlands of southeast asia . . . .
physical characteristics : african pittas have a black head with a yellow side stripe ; white throat with pink wash ; blackish brown bill with a reddish base ; deep buff breast and flanks ( sides ) ; whitish color under the bill and throat that turns yellow at breast ; bright olive green upperparts with blue and black banding on wings ; dark azure - blue rump ; blackish flight feathers with paler tips ; black tail with red underside and blue upperside ; and pinkish to grayish white feet . males and females look alike . more
breeds may to august . domed nest is located 3–10 ft ( 1–3 m ) above the ground , often in palm trees . nest constructed from dead leaves and twigs on a base of larger sticks ; lined with fine rootlets . clutch size usually three to four . eggs similar to those of the hooded pitta ; white with dark purple or brownish spots over gray markings , most numerous on widest end . female and male share incubation , brooding , and provisioning of young .
p . a . longipennis : migratory ; breeds in central tanzania , malawi , southeast democratic republic of congo , eastern zambia , zimbabwe , and possibly into northern south africa ; nonbreeding migrant in northern tanzania , rwanda , burundi , democratic republic of congo , central african republic , uganda , and coastal kenya . p . a . pulih : west africa ; resident in sierra leone lowlands , ghana , liberia , ivory coast , nigeria , and coastal cameroon . p . a . angolensis : west africa ; southern cameroon , guinea , congo , democratic republic of congo , and angola
pittas give short calls , usually one , two , or occasionally three syllables , which can be either whistled or buzzy . the role that these calls play in territorial defense is evidenced by the fact that many species can be drawn out of dense vegetation by playing a recording of their call . in a natural setting , such a response may lead to encounters between males on adjacent territories . for rainbow pittas and elegant pittas ( pitta elegans ) biologists have described displays in which males from adjacent territories face off and perform bowing displays sometimes in conjunction with
purring
vocalizations .
much of avian life history theory has been developed from studies of birds in temperate climates of the northern hemisphere , where food availability is regarded as a critical driver of life history variation . yet , nearly two - thirds of the word’s bird species are confined to the tropics and temperate regions of the southern hemisphere , where they exhibit life histories that differ from north - temperate birds . also , traditional life history theory regards age - specific survival as the key driver of life history variation . this symposium will highlight the insights that recent studies of ‘southern’ , particularly african birds , provide to our understanding of global drivers of avian life history variation , and the conservation implications thereof .
bliimungwe’s four raised thatched rooms have been carefully designed to ensure the height of comfort whilst still retaining that authentic bushcamp feel . the tangle of mature mahogany trees that surround camp are echoed in the rich , wood interiors and the beautiful wooden furniture , handmade by local artisans . bright african textiles bring splashes of colour to the elegant rooms . after an early morning game drive , head back to your room for a refreshing , open - air waterfall shower , or continue the wildlife spectacle from your private deck . two of the chalets have twin beds ( queen - sized beds ) , while the other two each have a king bed . bilimungwe sleeps up to eight guests and is open from may to december .
winter offers the only chance of malagasy pond - heron , mascarene martin and short - tailed pipit . it is also possible to see madagascar cuckoo in winter and all the resident species are available in winter . while there is no replacing the exciting rush of full activity in mid - summer , winter birding can be surprisingly good and we have had good views of great bittern , thick - billed cuckoo , african golden oriole , rufous - bellied heron , allen ' s gallinule and even black coucal in the winter period . carmine bee - eater , madagascar bee - eater , collared pratincole , black - winged lapwing , wattled crane , osprey , terek sandpiper , grey - rumped swallow , mangrove kingfisher are all regularly seen on winter trips .
african pittas are monogamous like humans . that means , that they stay with one mate their entire life . they tend to usually breed between september and february . when it is mating season , the male pittas will show off their bright colors in a dance - like manner , spreading their wings and hopping on branches . their gestation period in unknown and it breeds once a year . there are typically 1 to 4 eggs in a litter . their eggs are glossy and round and their nest is usually hidden in a bush so that nothing can find it . the nest looks like a messy , leafy dome , which makes it even better for camouflage . when born , baby pittas are brown with pink legs and an orange beak . the baby pittas are
pittas are particularly associated with the islands of the indo - pacific . borneo , the philippines , new guinea , and many indonesian and melanesian islands host endemic pittas . the absolutely gorgeous ivory - breasted pitta ( right , a wonderful shot by rob hutchinson ) is endemic to halmahera and nearby moluccan islands . i was with rob when he called in this individual with a tape , and the bird appeared only briefly . rob managed this great shot ; mine was distant and marginal . it is truly remarkable how difficult these colorful birds are to see . unless they decide to fly briefly up to a perch ( as this one has done ) they seemingly melt away in the understory .


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 .
gen . et sp . nov . , a new dipnoan - like form from the lower devonian of eastern yunnan , china . proceedings of the linnean society of new south wales 107 : 171–184 .
from the xishancun formation ( early lochkovian ) of qujing , china . geobios ms 19 : 293–299 .
clack ja ( 2011 ) the fossil record of lungfishes . in : jørgensen jm , joss jeditors . the biology of lungfishes . enfield , usa : science publishers .
cloutier r ( 1996 ) dipnoi ( akinetia : sarcopterygii ) . in : schultze hp , cloutier reditors . devonian fishes and plants of miguasha , quebec , canada munich : verlag dr friedrich pfeil . pp . 198–226 .
ahlberg pe ( 1991 ) a re - examination of sarcopterygian interrelationships , with special reference to porolepiformes . zoological journal of the linnean society 103 : 241–287 .
meyer a , zardoya r ( 2003 ) recent advances in the ( molecular ) phylogeny of vertebrates . annual review of ecological and evolutionary systems 34 : 311–338 .
long j , gordon ms ( 2004 ) the greatest step in vertebrate history : a paleobiological review of the fish - tetrapod transition . physiological and biochemical zoology 77 : 700–719 .
as the oldest coelacanth : implications for early osteichthyan interrelationships . journal of systematic palaeontology 5 : 289–343 .
betancur - r r , broughton re , wiley eo , carpenter k , lopez ja , et al . ( 2013 ) the tree of life and a new classification of bony fishes . plos currents . 1 .
marshall c , schultze h - p ( 1992 ) relative importance of molecular , neontological , and paleontological data in understanding the biology of the vertebrate invasion of land . journal of molecular evolution 35 : 93–101 .
heinicke mp , sander jm , blair hedges s ( 2009 ) lungfishes ( dipnoi ) . in : hedges sb , kumar seditors . the timetree of life : oxford university press . pp . 348–350 .
wilder b ( 1887 ) the dipnoan brain . the american naturalist 21 : 544–548 .
rudebeck b ( 1945 ) contributions to forebrain morphology in dipnoi . acta zoologica 26 : 9–156 .
northcutt rg ( 1986 ) lungfish neural characters and their bearing on sarcopterygian phylogeny . journal of morphology supplement 1 : 277–297 .
northcutt rg ( 2002 ) understanding vertebrate brain evolution . integrative and comparative biology 42 : 743–756 .
collin sp ( 2007 ) nervous and sensory systems . in : mckenzie dj , farrell ap , brauner cjeditors . primitive fishes , fish physiology . pp 121–179 .
nieuwenhuys r , donkelaar hj , nicholson c ( 1998 ) the central nervous system of vertebrates : springer . 2214 p .
joss jmp ( 2005 ) lungfish evolution and development . general and comparative endocrinology 148 : 285–289 .
gross , an upper devonian dipnoan from wildungen . kunglinga svenska vetenskapsakademiens handlingar 4 3 : 1–29 .
from wee jasper , new south wales . records of the australian museum 52 : 103–128 .
pridmore pa , campbell ksw , barwick re ( 1994 ) morphology and phylogenetic position of the holodipteran dipnoans of the upper devonian gogo formation of northwestern australia . philosophical transactions of the royal society of london ( biol ) 344 : 105–164 .
, a lower devonian crossopterygian from yunnan south - western china : university of stockholm and section of palaeozoology , swedish museum of natural history . 113 p .
from spitsbergen , based on ct scanning . in : elliott dk , maisey jg , yu x , miao deditors . morphology , phylogeny and paleobiogeography of fossil fishes : honoring meemann chang . munich : dr . friedrich pfeil . pp 363–377 .
n . sp . and a discussion on the structure of the head in the porolepiformes . meddelelser om gronland 187 : 1–307 .
lu j , zhu m , long ja , zhao w , senden tj , et al . ( 2012 ) the earliest known stem - tetrapod from the lower devonian of china . nature communications 3 : 1160 .
jarvik e ( 1980 ) basic structure and evolution of vertebrates . london : academic press .
clement am , long ja ( 2010 ) air - breathing adaptation in a marine devonian lungfish . biology letters 6 : 509–512 .
clement am ( 2012 ) a new species of long - snouted lungfish from the late devonian of australia , and its functional and biogeographical implications . palaeontology 55 : 51–71 .
playford pe ( 1980 ) devonian ‘great barrier reef’ of canning basin , western australia . bulletin of the american association of petroleum geologists 64 : 814–840 .
long ja , trinajstic k ( 2010 ) the late devonian gogo formation lägerstatten of western australia - exceptional vertebrate preservation and diversity . annual review of earth & planetary sciences 38 : 665–680 .
bemis we ( 1984 ) paedomorphosis and the evolution of the dipnoi . paleobiology 10 : 293–307 .
sollas wj ( 1903 ) a method for the investigation of fossils by serial sections . philosophical transactions of the royal society , biological sciences 196 : 259–265 .
stensiö ea ( 1927 ) the downtonian and devonian vertebrates of spitzbergen . skrifter om svalbard og nordishavet 12 : 1–31 .
young gc ( 2005 ) x - ray microtomography of 410 million - year - old optic capsules from placoderm fishes . micron 36 : 551–557 .
tafforeau p , boistel r , boller e , bravin a , brunet m , et al . ( 2006 ) applications of x - ray synchrotron microtomography for non - destructive 3d studies of paleontological specimens . applied physics a 83 : 195–202 .
long ja , trinajstic k , young gc , senden tj ( 2008 ) live birth in the devonian period . nature 453 : 650–652 .
sanchez s , ahlberg pe , trinajstic km , mirone a , tafforeau p ( 2012 ) three - dimensional synchrotron virtual paleohistology : a new insight into the world of fossil bone microstructures . microscopy and microanalysis 18 : 1095–1105 .
sanchez s , dupret v , tafforeau p , trinajstic km , ryll b , et al . ( 2013 ) 3d microstructural architecture of muscle attachments in extant and fossil vertebrates revealed by synchrotron microtomography . plos one . 8 .
dupret v , sanchez s , goujet d , tafforeau p , ahlberg pe ( 2014 ) a primitive placoderm sheds light on the origin of the jawed vertebrate face . nature .
pradel a , langer m , maisey jg , geffard - kuriyama d , cloetens p , et al . ( 2009 ) skull and brain of a 300 - million - year - old chimaeroid fish revealed by synchrotron holotomography . pnas 106 : 5224–5228 .
witmer lm , chatterjee s , franzosa j , rowe t ( 2003 ) neuroanatomy of flying reptiles and implications for flight , posture and behaviour . nature 425 : 950–953 .
sakellariou a , sawkins tj , limaye a , senden tj ( 2004 ) x - ray tomography for mesoscale physics . physica a - statistical mechanics and its applications 339 : 152–158 .
kardong kv ( 2006 ) vertebrates : comparative anatomy , function , evolution . new york : mcgraw hill . 782 p .
( merycoidodontidae ; oreodontoidea ) and its implications for apomorphy - based diagnosis of isolated , natural endocasts . journal of vertebrate paleontology 29 : 1199–1211 .
milner ac , walsh sa ( 2009 ) avian brain evolution : new data from palaeogene birds ( lower eocene ) from england . zoological journal of the linnean society 155 : 198–219 .
stensiö e ( 1963 ) the brain and the cranial nerves in fossil , lower craniate vertebrates . skrifter utgitt av det norske videnskaps - akademi : 1–120 .
retzius g ( 1881 ) das gehororgan der wirbeltieren : das gehororgan der fische und amphibien vol . 1 . stockholm .
miles rs ( 1977 ) dipnoan ( lungfish ) skulls and the relationships of the group : a study based on new species from the devonian of australia . zoological journal of the linnean society 61 : 1–328 .