rinconsaurus is a genus of dinosaur from the late cretaceous .

it was a titanosaurid sauropod which lived in what is now argentina .

the type species , rinconsaurus caudamirus , was described by calvo and riga in 2003 , and is based on three partial skeletons .

rinconsaurus

skeletal reconstruction showing known parts of rinconsaurus caudamirus . from calvo & gonzalez riga , 2003 .
without the ability to grind food ,
rinconsaurus
, like other sauropods , would have stripped foliage .
sin la capacidad de moler el alimento ,
rinconsaurus
, como otros saurópodos , habría pelado el follaje .
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rinconsaurus was a relatively small titanosaur that lived in south america during the late cretaceous period .  not too much is known about rinconsaurus other than it appears to have had a more gracile ( lightweight ) build than some titanosaurs .  teeth associated with the type specimen are also termed chisel - like which means that rinconsaurus probably snipped off fronds of vegetation rather than stripping it between peg - like teeth like some sauropods .  rinconsaurus is thought to be closely related to aeolosaurus which has sometimes seen it described as a member of the aeolosaurini . another titanosaur from the same formation as rinconsaurus is antarctosaurus .
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rinconsaurus
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rinconsaurus is a dinosaur from the late cretaceous period . it was a titanosaurian sauropod which lived in what is now argentina . unlike its relatives brachiosaurus and diplodocus rinconsaurus was very slender . it was a herbivorous animal that fed on plants like cycads and fern . its egg , possibly the rinch egg is featured in dino run
text - fig . 2 . skeletal reconstruction of rinconsaurus caudamirus gen . et sp . nov . , showing preserved bones ( estimated total length : 11 m ) .
revista geológica de chile - rinconsaurus caudamirus gen . et sp . nov . , a new titanosaurid ( dinosauria , sauropoda ) from the late cretaceous of patagonia , argentina
text - fig . 1 . map of neuquén basin ( patagonia , argentina ) showing the locality where the holotype of rinconsaurus caudamirus gen . sp . nov . was found .
like all sauropods , rinconsaurus was a large long - necked quadrupedal animal , with a long , whip - like tail and four pillar - like legs . although fossil discoveries are incomplete , and no complete necks or heads have been found , fully grown rinconsaurus are estimated to have been 11 meters ( 36 ft ) long and approximately 2 . 5 meters ( 8 ft ) high at the shoulder . though only a portion of one skull has been recovered , rinconsaurus may have had a long , narrow skull , based on fossil evidence of the skulls of related titanosaurs .
rodrigo m . santucci y antonio c . de arruda - campos ( 2011 ) en su análisis cladístico encontró que
aeolosaurus
,
gondwanatitan
,
maxakalisaurus
,
panamericansaurus
y
rinconsaurus
son aeolosáuridos .
rodrigo m . santucci and antonio c . de arruda - campos ( 2011 ) in their cladistic analysis found
aeolosaurus
,
gondwanatitan
,
maxakalisaurus
,
panamericansaurus
and
rinconsaurus
to be aeolosaurids .
rinconsaurus information tells us about its characteristics , diet , habitat etc . rinconsaurus diet tells us whether it is carnivorous , herbivorous or omnivorous . herbivores feed on plants and grass , whereas the carnivores feed on other animals or even other dinosaurs . omnivores eat both animal and plants . depending on its diet , we can say that it is herbivore . the meaning of the name will also tell you something about that dinosaur . the correct pronunciation of it is rin - con - sore - us .
text - fig . 4 . cladogram determined by cladistic analysis ( 68 steps ; ci 0 . 79 ; ri 0 . 78 ) showing the phylogenetic relationships of rinconsaurus caudamirus gen . et sp . nov . synapomorphies supporting each node are listed and discussed in the text .
j . o . calvo and b . j . gonzález riga . 2003 . rinconsaurus caudamirus gen . et sp . nov . , a new titanosaurid ( dinosauria , sauropoda ) from the late cretaceous of patagonia , argentina . revista geológica de chile 30 ( 2 ) : 333 - 353 .
full reference : j . o . calvo and b . j . gonzález riga . 2003 . rinconsaurus caudamirus gen . et sp . nov . , a new titanosaurid ( dinosauria , sauropoda ) from the late cretaceous of patagonia , argentina . revista geológica de chile 30 ( 2 ) : 333 - 353
los titanosaurios fueron un linaje diverso de dinosaurios herbívoros que incluye desde las especies más pequeñas , como rinconsaurus o saltasaurus de alrededor de 6 toneladas ( el equivalente aproximado en tamaño a un elefante adulto ) , hasta las más grandes conocidas , tales como argentinosaurus o puertasaurus , especies de más de 60 toneladas .
text - fig . 3 . rinconsaurus caudamirus gen . et sp . nov . a , right ischium in lateral view , mrs - pv 101 . b , left ilium in lateral view , mrs - pv 96 , c , right femur in posterior view , mrs - pv 49 . scale bars equal 10 cm .
further reading - rinconsaurus caudamirus gen . et sp . nov . , a new titanosaurid ( dinosauria , sauropoda ) from the late cretaceous of patagonia , argentina . - revista geológica de chile 30 ( 2 ) : 333 - 353 . - j . o . calvo & b . j . gonzález riga - 2003 .
rinconsaurus belongs to the family of titanosaurid sauropod that has been found existed in the turonian to coniacian age of late cretaceous period . the fossil remains of this species was first recovered in argentina , neuquén province - neuquén formation . this herbivore species comes under the classification of chordata , reptilia , dinosauria , saurischia , sauropodomorpha , sauropoda , titanosauria .
' aeolosaurus rionegrinus belongs to aeolosaurus ' according to a . w . a . kellner et al . 2005 ' alamosaurus sanjuanensis belongs to alamosaurus ' according to a . w . a . kellner et al . 2005 ' andesaurus delgadoi belongs to andesaurus ' according to a . w . a . kellner et al . 2005 ' antarctosaurus brasiliensis is a nomen dubium ' according to a . w . a . kellner et al . 2005 ' baurutitan britoi belongs to baurutitan ' according to a . w . a . kellner et al . 2005 ' baurutitan belongs to titanosauridae ' according to a . w . a . kellner et al . 2005 ' epachthosaurus sciuttoi belongs to epachthosaurus ' according to a . w . a . kellner et al . 2005 ' gondwanatitan faustoi belongs to gondwanatitan ' according to a . w . a . kellner et al . 2005 ' laplatasaurus araukanicus is recombined as titanosaurus araukanicus ' according to a . w . a . kellner et al . 2005 ' lirainosaurus astibiae belongs to lirainosaurus ' according to a . w . a . kellner et al . 2005 ' mendozasaurus neguyelap belongs to mendozasaurus ' according to a . w . a . kellner et al . 2005 ' opisthocoelicaudia skarzynskii belongs to opisthocoelicaudia ' according to a . w . a . kellner et al . 2005 ' pellegrinisaurus powelli belongs to pellegrinisaurus ' according to a . w . a . kellner et al . 2005 ' rinconsaurus caudamirus belongs to rinconsaurus ' according to a . w . a . kellner et al . 2005 ' saltasaurus loricatus belongs to saltasaurus ' according to a . w . a . kellner et al . 2005 ' titanosaurus australis is recombined as neuquensaurus australis ' according to a . w . a . kellner et al . 2005 ' titanosaurus colberti is recombined as isisaurus colberti ' according to a . w . a . kellner et al . 2005
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urltoken
revista geológica de chile , vol . 30 , no . 2 , p . 333 - 353 , 4 figs . , 3 plates , 1 appendix , 2 tables , december 2003 .
laboratorio de paleovertebrados , ianigla - cricyt , centro regional de investigaciones científicas y tecnológicas , avda . r . leal s / n , parque gral . san martín ( 5500 ) , mendoza , argentina ( bgonriga @ urltoken ) .
has preserved short articulated posterior caudal series with amphicoelous , opisthocoelous and biconvex centra . this unusual morphology represents the first such occurrence in sauropod dinosaurs . from the systematic point of view , this discovery is important because titanosauridae were traditionally defined , among other characters , by strongly developed procoelia in their caudal vertebrae .
key words : dinosauria , sauropoda , titanosauridae , late cretaceous , neuquén , argentina .
gen . et sp . nov . , un nuevo titanosáurido ( dinosauria , sauropoda ) del cretácico tardío de patagonia , argentina .
gen et sp . nov . ( dinosauria , saurpoda ) es un nuevo y esbelto titanosauridae proveniente de la formación río neuquén , cretácico superior de rincón de los sauces , provincia de neuquén , patagonia , argentina . sus restos están integrados por 13 vértebras caudales articuladas y materiales craneales , cervicales , dorsales y apendiculares desarticulados , que pertenecen a tres individuos , dos adultos y un juvenil . este nuevo taxón se caracteriza por la siguiente asociación de autapomorfías : ( 1 ) vértebras dorsales medioanteriores con espinas neurales inclinadas posteriormente más de 60 grados con respecto a la vertical , ( 2 ) vértebras caudales medias con procesos óseos que sostienen las carillas articulares de las poszigapófisis y ( 3 ) vértebras caudales procélicas con intercalaciones de series con centros anficélicos y bioconvexos , o con centros anficélicos , opistocélicos y biconvexos . un análisis filogenético cladista permite incluir a
presenta dos cortas secuencias caudales posteriores articuladas con centros anficélicos , opistocélicos y biconvexos . esta inusual morfología se registra por primera vez en dinosaurios saurópodos . es importante desde el punto de vista sistemático , ya que titanosauridae ha sido tradicionalmente definido , entre otros caracteres , por la presencia de fuerte procelia en sus vértebras caudales .
palabras claves : dinosauria , sauropoda , titanosauridae , cretácico tardío , neuquén , argentina .
) . in south america the record of titanosauridae is particularly abundant . they are known since the xix century (
) . during the late cretaceous , titanosaurids were important primary consumers in the ecosystems of patagonia , as the ornithischians were in the north america . this difference has been explained by the gradual isolation of south america during most of the late cretaceous (
in the last five years , extraordinary titanosaurid fossils were found in the region of rincón de los sauces , northern neuquén province . some of these discoveries include articulated specimens (
) . the excavations of these discoveries were coordinated by investigators of the national university of comahue and they have been partially supported by the local town hall .
) . they were extracted by the paleontologist j . calvo and his team of the paleontology museum of the national university of comahue . fossil remains are housed at the laboratory of rincón de los sauces museum under the abbreviation mrs - pv .
( pl . 1 , a - h ; pl . 2 , a - c ; pl . 3 , a - b ; text -
( latin ) , astonishing , amazing , in reference to the unusual morphology of posterior caudal vertebrae ; freely ' the dinosaur from rincón with amazing tail ' .
mrs - pv 26 , 13 articulated anterior - middle and middle - posterior caudal vertebrae and two ilia .
the following bones associated with the holotype are included : teeth ( mrs - pv 117 , 263 ) , prefrontal ( mrs - pv 102 ) , angular and surangular ( mrs - pv 112 ) , cervical vertebrae ( mrs - pv 2 , 3 , 8 , 4 and 21 ) , dorsal vertebrae ( mrs - pv 5 , 6 , 9 , 11 , 13 , 16 , 17 , 18 and 19 ) , anterior caudal vertebrae ( mrs - pv 22 , 23 , 24 , 25 and 27 ) , middle caudal vertebrae ( mrs - pv 27 , 28 and 31 ) , posterior caudal vertebrae ( mrs - pv 29 , 30 , 32 - 40 ) , haemal arches ( mrs - pv 20 , 42 , 93 ; 99 , 109 , 113 ) , scapula and coracoid ( mrs - pv 43 ) , sternal plates ( mrs - pv 46 , 103 , 104 ) , humerus ( mrs - pv 47 ) , metacarpals ( mrs - pv 98 ) , ischia ( mrs - pv 94 , 101 ) , ilia ( mrs - pv 96 ) , pubes ( mrs - pv 97 , 100 ) , femora ( mrs - pv 49 , 92 ) and metatarsal ( mrs - pv 111 ) .
correspond to three individuals , two adults and one juvenile . all fossil remains ( dorsal vertebrae , articulated caudal vertebrae and appendicular bones of different sizes ) were found associated , exhibit similar morphologies and constitute a monospecific assemblage . the best preserved bones , the articulated caudal series mrs - pv 26 , were associated ' in life position ' with two ilia . moreover , associated with them , several articulated dorsal vertebrae ( mrs - pv 05 ) were preserved that probably correspond to the same specimen , as well as several limb bones . in the same fossil assemblage were found several articulated vertebrae and limb bones that correspond to other two specimens . duplicate bones represented by caudal vertebrae , ilia , sternal plates , ischium and femur exhibit the same morphological characters . these paleontological evidences confirm the presence of a monospecific assemblage . for this reason , all bones recovered were included in the same type series ( holotype and paratypes ) such as the
) . a cranial fragment , one mandible and teeth have been recovered . the right prefrontal ( mrs - pv 102 ) is small and anteroposteriorly elongated . its posterior end is transversally wide and it is arched posteroventrally . the anterior end is acute in its exposed portion . the articular surface for the nasal is extensive . its posterior border is concave and of triangular shape . the lateral border of the prefrontal contributes to the skull roof and the posterodorsal border of the orbit . its rugose surface contains small grooves .
) . the surangular is a delicate and elongated bone lamina . posteriorly , the ventral border is slightly convex in lateral view and it becomes concave anteriorly . in medial view , the angular forms a weakly concave surface .
) . one of them ( mrs - pv 117 ) has a suboval section , and has two crests that allow us to differentiate slightly convex lingual and more convex labial surfaces . its apex is incomplete . the other tooth ( mrs - pv 263 ) , more complete and bowed lingually , presents the same features as the anterior one but in more marked form . the worn surface is parallel to the tooth axis . it is remarkable with respect to the development of sharp crests forming the anterior and posterior borders . in general , titanosaurid teeth are pencil - chisel like (
) with a sharp worn surface placed almost parallel to the tooth axis . some have oval section with rounded borders depending the position in the jaws . the suboval teeth with labial and lingual faces well differentiated by crests present in
) is an unusual character for derived titanosaurids . however , a similar morphology , with some differences in the cross - sectional shape , is also present in the basal titanosaurid
) . two anterior cervical vertebrae were recovered ( mrs - pv 08 , 21 ) . they possess relatively short , opisthocoelous centra , wider than high . in their lateral side is a deep , acute depression that defines the pleurocoels . the parapophyses are located at the base of the centra . laterally , the centroprezygapophyseal and centrodiapophyseal laminae are inclined strongly forward . this inclination differs considerably from that of the saltasaurinae (
) , but it is very similar to the inclination observed in the titanosauridae indet . ' series b ' from brazil (
posteriorly , the opisthocoelous centra increase in length . this character is observed in the middle cervical vertebra mrs - pv 02 and in the posterior cervical vertebrae mrs - pv 03 and mrs - pv 04 , the last one not complete (
. the ventral face is wide and smoothly concave anteroposteriorly . in the lateral face of the centum there is a long anteroposterior depression with small pleurocoels . this depression is divided by a posteroventrally directed lamina . the parapophyses are laminar and subtriangular . they are located in the anterior half of the vertebral body and extend lateromedially . in anterior view , the spinodiapophyseal and spinoprezygapophyseal laminae reach each other at the level of the postzygapophysis , defining a subtriangular depression . the diapophysis is located on the anterior half of the body , at the level of the prezygapophysis . it is reinforced by the anterior and posterior centrodiapophyseal laminae , the prezygadiapophyseal lamina and the postzygadiapophyseal lamina . in this way , these laminae form four deep depressions around the diapophysis . the prezygapophyses are relatively long , extending beyond the anterior end of the vertebral body ; the postzygapophyses do not extend beyond the center . the opposite is observed in
) . the articular surfaces of the prezygapophyses are elongated anteroposteriorly and inclined toward the medial plane . the prezygapophyses are reinforced by two centroprezygapophyseal laminae that in anterior view define a deep depression . the neural spine is low and distally expanded .
) . the authors have collected three articulated anteromedial dorsal neural arches ( mrs - pv 05 ) . the neural arches are wider than long and have suffered dorsoventral compression . the prezygapophyseal facets are reduced and have an oval outline . the centroprezygapophyseal lamina is absent . in contrast ,
) . in the most anterior dorsal , an accessory centrodiapophyseal lamina extends parallel to the centrodiapophyseal lamina . at this level , the anterior and posterior centrodiapophyseal laminae are not forked as they occur in the middle - posterior dorsal arches ( mrs - pv 06 ) . the prezygadiapophyseal lamina presents a straight border in dorsal view . the almost horizontal postygadiapophyseal lamina has a concave border in dorsal view .
the neural spine is reduced and transversely expanded distally . it is reinforced ventrally by a spinopostzygapophyseal lamina . the neural spines is strongly inclined posteriorly . in spite of that , in most of titanosaurs an inclination of the neural spine in anterior dorsal vertebrae is present (
the prespinal lamina extends to the base of the neural arch . the accessory spinodiapophyseal laminae are not present . in contrast , these laminae are present in
the authors have also recovered an isolated middle - posterior dorsal neural arch ( mrs - pv 06 ) . in lateral view , there is a bifurcation of the centrodiapophyseal lamina . the neural spine is reinforced by a prespinal and postspinal laminae . in dorsal view , the spinodiapophyseal and spinoprezygapophyseal laminae form a wide and deep cavity .
) . in their lateral faces , they have well developed , eye - shaped pleurocoels . the pleuro coels are elongated and deep , and they occupy 60 percent of the centrum length . the hyposphene - hypantrum articulation is absent . in only few posterior dorsal vertebrae the neural arch is partially preserved . the neural arch has a wide lateral base , which occupies more than 60 percent of the centrum . the transverse process is relatively thin . the diapophyses are directed laterally and upward . they are reinforced ventrally by centrodiapophyseal , postygadiapophyseal and paradiapophyseal laminae , and are supported dorsally by the spinodiapophyseal lamina . the distal end of the diapophysis present a horizontal and plane surface , similar to that in
) . the parapophyses are not well preserved but they are reinforced ventrally by robust anterior and slender posterior centroparapophyseal laminae . the last one unites to the inferior portion of the centrodiapophyseal lamina , similar to that of
. the articular surface of the postzygapophysis prolongs as the postzygadiapophyseal laminae . the postzygapophyses are well separated . the neural spine is undivided . it is composed by the spinodiapophyseal , spinopostzygapophyseal , prespinal and postspinal laminae ; all unite in the transversal broadening of the distal end . the prespinal lamina is well developed , extending to the base of the neural spine . besides the described materials we have collected 6 dorsal centra , some with incomplete neural arches ( mrs - pv 9 , 11 , 13 , 16 , 19 ) .
two fused centra have been preserved ( mrs - pv 41 ) without any diagnostic character .
. the authors have collected several caudal vertebrae ; some are isolated , but others are articulated . below , the authors describe their shape and morphology in different parts of the tail . all the caudals collected belong to two individuals of
. the first part of the tail is represented by two poorly preserved centra and neural arches . the ? first caudal ( mrs - pv 22 ) has thin , laminar and laterally directed transverse processes . at the base of the postzygapophyses , the transverse processes possess two foramina . the second caudal ( mrs - pv 23 ) is strongly procoelous and has smaller lateral expansion than the first one . in lateral view , the centrum presents a depression with the shape of a pleurocoel .
the series of anterior caudals is not complete . three articulated series have been collected . mrs - pv 23 is represented by five vertebrae , mrs - pv 24 by six vertebrae and mrs - pv 25 by three vertebrae . the holotype (
) includes an articulated series of caudals ( mrs - pv 26 ) represented by four anterior caudals , eight middle caudals and five posterior ones .
) . the holotype ( mrs - pv 26 ) preserves eight middle caudal vertebrae and five posterior ones . there is also an articulate series of three middle caudals ( mrs - pv 27 ) , two incomplete disarticulated middle caudals ( mrs - pv 28 ) and an isolated one ( mrs - pv 31 ) .
middle caudals are similar to the anterior caudals except for the neural spine , that is directed posteriorly , and for the absence of transverse processes . middle caudal centra are strongly procoelous , as high as wide . the lateral faces are flat , slightly concave anteroposteriorly . the ventral face of the centra is narrow and flat , with a strong compression in its middle part . the prezygapophyses are directed horizontally or slightly upward . both prezygapophyses fuse proximally , developing a small platform .
the articular faces of the postzygapophyses are inclined lateroventrally . they are placed at the level of the posterior border of the centra . as in anterior caudals , the articular faces of the postzygapophyses are separated from the neural spine by a bony process which can be seen clearly in dorsal view (
) . the postzygapophyses and the prezygapophyses are well separated from the axial plane due to the development of the postzygapophyseal processes .
) . there is an articulated section of five posterior vertebrae that are part of the holotype ( mrs - pv 26 ) . there are also two series of three articulated caudals each one ( mrs - pv 29 , 30 ) and several isolated posterior caudals ( mrs - pv 32 , 33 , 34 , 35 , 36 , 37 , 38 , 39 , 40 ) . all these posterior caudals are similar to the middle caudals in having the typical procoelous centrum . the bony process of the postzygapophysis is not present .
in contrast to the caudals described above , the authors have recovered two articulated series with an unusual centrum . one of the series ( mrs - pv 29 ) is composed of a procoelous vertebra , an amphicoelous vertebra and a biconvex vertebra (
a ) . the centrum , dorsoventrally compressed , is wider than high . the lateral faces are convex laterodorsally . the ventral border is slightly convex . the posterior border of the neural arch is placed in the posterior half of the centrum . the postzygapophyses are placed at the level of the posterior border of the centra . the distal ends of the neural spine reach the posterior border of the centra . the anterior border of the neural spine is at a lower level than the posterior one . the processes for the chevrons are not present . the prezygapophyses fuse in their proximal end , developing a small platform as in anterior and middle caudal vertebrae . measurements of the caudal mrs - pv 29 / 2 : length , width and height of the centrum : 104 , 82 and 38 mm , respectively .
) . they present characteristics similar to the anterior series , although their neural spines are more reduced due to their more posterior position in the sequence .
) . this unusual morphology can be interpreted as an autapomorphy of this new taxon . as indicated
, the finding of non - procoelous isolated caudal vertebrae in the late cretaceous must be interpreted carefully because they can belong to a titanosaurid .
measurements of the caudal mrs - pv 30 / 1 : length , width and height of the centrum : 114 , 57 and 33 mm , respectively .
. there are several haemal arches ; some are deformed and others are incomplete ( mrs - pv 20 , 42 , 93 ; 99 , 109 , 113 ) . in general , they are relatively long and thin , open proximally , and lack strong articular faces .
. the authors have collected a complete right scapula ( mrs - pv 43 ) . it is a relatively long and laminar bone with its external face convex . the scapular blade is thin and slender . the contact between the scapular blade and the proximal end is narrow and there is a crest that separates both of them . the ventral border is straight up to its union with the proximal end , forming an angle of approximately 140 degrees . the supraglenoid process is prominent . the supracoracoid depression is wide because the diagonal acromion coincides with the border of the acromial process . the proximal end of the scapula contacts the coracoid . the distal end , although incomplete , has a convex border . the coracoid has a square shape . its posteromedial border forms a 90 degree angle with the anteromedial border of the scapula . the coracoid foramen , in the external face , is surrounded by a depression directed toward the anteromedial border . measurements : length : 820 mm ; proximal width : 440 mm ; distal width : 215 mm ; minimum width of the blade : 130 mm .
) . three sternal plates have been preserved ( mrs - pv 46 , 103 and 104 ) . the right sternal plate mrs - pv 46 is relatively complete . it is laminar and it has a semilunar shape . the concave external border is very thin , while the internal one is convex . measurements : length : 560 mm ; width : 260 mm . the other sternal plates are not well preserved .
) . the incomplete left humerus is relatively slender ( mrs - pv 47 ) . its anterior face presents a prominent deltopectoral crest anteriorly projected . in its distal end , the radial condyle is well developed . measurements : estimated length : 790 mm ; preserved length : 740 mm ; proximal width : 260 mm ; distal width : 210 mm ; perimeter and diameter of the diaphysis : 300 and 120 mm , respectively .
. the authors have recovered five isolated metacarpals with their ends eroded . their lengths range between 260 and 240 mm .
mrs - pv 98 / 5 ) exhibits a robust proximal end of subtriangular shape . one of its lateral sides is slightly rounded . its distal end is relatively reduced and it has been crushed .
( mrs - pv 98 / 4 ) presents a subtriangular shape at the distal end . both extremities are well developed .
( mrs - pv 98 / 1 ) presents an incomplete proximal end of subtriangular shape . it is characterized by slender shape . the diaphysis has two flat faces , with the sharp angle directed toward the posterior side .
( mrs - pv 98 / 2 ) exhibits a very robust distal end of subtriangular shape . the proximal end is incomplete .
( mrs - pv 98 / 3 ) , strongly crushed , is characterized by having the distal end more developed than the proximal one .
) . the authors have recovered four incomplete ilia corresponding to two individuals . one pair ( mrs - pv 26 ) is associated with a series of caudal vertebrae . the other pair is incomplete ( mrs - pv 96 ) .
specimen mrs - pv 96 possesses a left ilium that preserves the acetabulum and great part of the preacetabular lamina . this lamina is wide , and it is directed upward and outward . the incomplete right ilium also preserves the acetabulum , and the lower part of the preacetabular and postacetabular laminae .
specimen mrs - pv 26 possesses a right ilium that preserves the preacetabular lamina , most of the postacetabular one , and also the complete acetabula and their peduncles . the preacetabular lamina exhibits a straight ventral border and a curved anterodorsal border . the left ilium lacks part of the iliac lamina , but it is complete in the lower part of the postacetabular and preacetabular laminae , the acetabulum and their peduncles . the length of the left ischium is 480 mm and the diameter of the acetabulum is 110 mm . the postacetabular lamina presents a rounded posterior border . the pubic peduncle is wide transversally .
) . the authors have collected three pubes . one specimen has both pubes preserved ( mrs - pv 97 ) , and the other specimen has just the right pubis ( mrs - pv 100 ) .
the right pubis of mrs - pv 97 is a relatively thin and laminar bone . the oval pubic foramen is closed . the pubic blade is wide and flattened , with its thick lateral border concave . the medial border is thin , and not well preserved . the distal end becomes wider anteroposteriorly . the acetabulum is reduced . measurements : length : 770 mm ; pubic foramen : 70 x 35 mm ; distal with : 255 mm .
the incomplete pubis mrs - pv 100 lacks its distal end , part of the proximal one in the area of the pubic foramen and part of the pubic lamina ; it corresponds to a smaller specimen than the one describe above .
) . two ischia have been collected ( mrs - pv 94 and 101 ) corresponding to two individuals . specimen mrs - pv 101 is an almost complete right ischium of small size . it lacks only a small central portion of the contact region with the pubis . this contact occupies 50 percent of the total length of the ischium . the acetabulum is complete and the thin iliac pedicel is well developed ; it is as large as that of
) . the distal lamina of the ischium is relatively wide with regard to the width of the ischiac articulation . measurements : length : 360 mm , width 130 mm .
specimen mrs - pv 94 , from larger animal , is not well preserved .
) . two femora have been collected ; the right one is complete and the left one is incomplete ( mrs - pv 49 and 92 ) . the most complete is relatively slender . the anteroposterior diameter of the traverse section at the level of the diaphysis is shorter than the posteromedial one . the fourth trochanter is well developed and it is 490 mm below the femoral head . the femoral head is placed at a right angle with respect to the axis . on the lateral margin , the lateral bulge is present as in
) . measurements : length : 990 mm ; greatest diameter and perimeter of the diaphysis : 140 and 340 mm , respectively .
. just one metatarsal has been recovered . it is probably metatarsal iii ? ( mrs - pv 111 ) . it is relatively slender and twisted . the proximal end , more developed than the distal one , has a subtriangular shape . the distal end presents a convex articular surface . measurements : length : 160 mm ; diameter of the diaphysis : 30 mm .
most of sauropod titanosaurs are represented by incomplete and fragmentary skeletal elements . in this context , the discovery of
gen . et sp . nov . , integrated by cranial , vertebral and appedicular remains , is relevant from a systematic viewpoint .
, in cladistic analyses already proposed by other authors . the authors have improved characters proposed by
, and have added new characters according to the evidence presented by this new taxon . this paper does not comprise a phylogenetic reevaluation of the titanosauridae family , because more taxa and characters should be included . the phylogenetic relationships of
gen . et sp . nov . ( this paper ) formed the ingroup .
) with a length of 68 steps and high consistency and retention indices ( c . i . = 0 . 79 ; r . i . = 0 . 78 ) . the multi - state characters were considered unordered .
, it presents differences due to the inclusion of new taxa and characters . they allow the authors to improve the knowledge of the relationships among
and titanosauria and all of its descendants ' . the clade is supported by nine synapomorphies defined by delayed optimization : teeth with sharply inclined wear facets ( 2 . 1 ) , single ( non - bifurcated ) neural spine in cervical vertebrae ( 6 . 1 ) , elongate cervical centra ( 7 . 1 ) , single ( non - bifurcated ) neural spine in anterior dorsal vertebrae ( 10 . 1 ) , prespinal lamina present in the distal end of neural spines in dorsal vertebrae ( 12 . 1 ) , neural arches placed anteriorly in middle and posterior caudal centra ( 26 . 1 ) , pubic peduncle of the ilium perpendicular to the sacral axis ( 40 . 1 ) , preacetabular lobe of ilium expanded and dorsally directed ( 41 . 1 ) and lateral bulge of femur below the greater trochanter ( 46 . 1 ) .
) . in the authors ' analysis it is united by five unambiguous synapomorphies : centroparapophyseal lamina in posterior dorsal vertebrae ( 13 . 1 ) , slightly forked centrodiapophyseal laminae in posterior dorsal vertebrae ( 14 . 1 ) , acuminate ( eye shaped ) pleurocoels in dorsal vertebrae ( 16 . 1 ) , bone internal structure of somphospondylous - camellate type ( 18 . 1 ) , and pubis longer than ischium ( 39 . 1 ) .
saltasaurinae and all of its descendants . in this analysis , it is supported by five synapomorphies defined by delayed optimization : presence of pencil - chisel like teeth ( 1 . 2 ) , absence of cervical pleurocoels divided by septa ( 5 . 1 ) , absence of hyposphene - hypantrum articulation in posterior dorsal vertebrae ( 15 . 1 ) , anterior caudal centra strongly procoelous with prominent posterior condyles ( 24 . 2 ) , and semilunar sternal plates ( 34 . 1 ) .
) have ambiguous distributions in the authors ' analysis and might be excluded from the diagnosis of this clade . they are : middle and posterior caudal centra strongly procoelous with prominent condyles ( 25 . 1 ) , absent in
, and six sacral vertebrae ( 19 . 1 ) , which is unknown in most of the analyzed taxa .
) , show that the caudal procoelia was not a permanent and uniform character . in these sauropods , the mid and posterior section of the tail exhibit complex morphological variation that must be analyzed carefully . for example ,
shows a typical strongly procoelous caudal sequence discontinued by amphicoelous , opisthocoelous , and biconvex centra . on the other hand , a new titanosaurid from mendoza province has slightly procoelous middle caudal centra with reduced posterior condyles , associated with typical strongly procoelous anterior caudal vertebrae (
from the lower cretaceous of africa . it has strongly procoelous anterior caudal centra apparently associated with gently amphicoelous or platycoelous middle and posterior caudals (
. it is united by three unambiguous characters : presence of prespinal lamina in posterior dorsal vertebrae up to the base of the neural spine ( 12 . 2 ) , middle caudal centra strongly procoelous with prominent condyles ( 25 . 1 ) , and slender and well - developed iliac pedicel of ischium ( 43 . 1 ) . the first two characters were cited by
. the third is proposed in this paper . this character has allowed recognition of morphological variation in the ischium of titanosauridae .
it is supported by two unambiguous characters : humerus with slightly curved proximal border ( 33 . 1 ) and coracoid of quadrangular shape ( 36 . 1 ) .
it is supported by three characters defined by delayed optimization : teeth with cylindrical cross - section ( 3 . 1 ) , reduced neural spines on posterior dorsal vertebrae ( 17 . 1 ) , and absence of phalangeal articular facets on the metacarpals ( 38 . 1 ) . node 7 includes node 11 (
. it is supported by three characters defined by delayed optimization : posteriorly inclined neural spines ( 20 to 50 degree from vertical ) on anterior and middle dorsal vertebrae ( 11 . 1 ) , 35 or fewer caudal vertebrae ( 21 . 1 ) , and metacarpal i longer than metacarpal iv ( 37 . 1 ) .
+ saltasaurinae . this clade is supported by three unambiguous characters : the presence of depressed middle and posterior caudal centra ( 23 . 1 ) , prominent lateral crest in the base of the neural arch in middle caudals ( 27 . 1 ) and wide and well developed iliac pedicel of ischium ( 43 . 2 ) .
) . it is diagnosed by two unambiguous synapomorphies : depressed anterior caudal centra ( 22 . 1 ) and the posterior orientation of the anterodorsal border of the neural spine in middle caudals ( 28 . 1 ) .
) . it is supported by one character : the presence of relatively long prezygapophyses ( 29 . 1 ) . this character is not exclusive of these taxa , since it is also present in
shows that the caudal morphology of titanosaurs is a complex subject to analyze . the inclusion of
among titanosauridae is well recorded according to the morphological evidence . however , unique among sauropods ,
has procoelous posterior caudal centra with intercalation of a series of amphicoelous - biconvex or amphicoelous - opisthocoelous - biconvex centra . this unusual morphology is important because titanosauridae were usually defined , among other characters , by having procoelous caudal vertebrae .
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( sauropoda , titanosauridae ) en la formación los alamitos , cretácico superior de la provincia de río negro , argentina .
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wilson , j . a . ; sereno , p . 1998 . early evolution and higher - level phylogeny of sauropod dinosaurs . society of vertebrate paleontology , memoir 5 , journal of vertebrate paleontology , vol . 18 , no . 2 , p . 1 - 68 . [ links ] manuscript received : january 22 , 2002 ; accepted : november 25 , 2003 .
tooth in transverse section ( c ) and lateral view ( d ) , mrs - pv 117 .
anteromedial dorsal neural arch in lateral ( g ) and anterior ( h ) views , mrs - pv 05 / 3 .
references : dp ; diapophysis , ne : neural spine , poz : postzygapophysis . scale bar equals 1 cm in figure d and 5 cm in the other figures .
articulated series of anterior - middle caudal vertebrae in lateral view , mrs - pv 26 .
articulated series of middle caudal vertebrae in lateral view , mrs - pv 27 .
references : cc : caudal centrum , ne : neural spine , poz : postzygapohysis , pozp : process of the postzygapophysis . scale bars equal 5 cm .
articulated posterior caudal vertebrae with procoelous , amphicoelous and biconvex centra , mrs - pv 29 .
articulated posterior caudal vertebrae with opisthocoelous , biconvex and procoelous centra , mrs - pv 30 .
left pubis in dorsal view , mrs - pv 97 . scale bars equal 5 cm .
) the distribution of 46 characters corresponding to 12 taxa of sauropods is shown . in this work , characters 32 and 43 are proposed , and characters 1 , 7 , 11 , 18 , 24 , 25 and 44 are redefined . the other characters have been proposed by the authors indicated on the list . the data matrix was analyzed with the computer programs paup ( swofford , 1989 ) and nona ( goloboff , 1993 ) . the application of heuristic method produced one most parsimonious tree with a length of 68 steps , c . i . = 0 . 79 and r . i . = 0 . 78 . the multi - state characters were considered unordered .
7 . middle cervical centra , anteroposterior length / height of the posterior face : between 2 . 5 - 3 ( 0 ) ; more than 3 ( 1 ) ; less than 2 . 5 ( modified from
32 . humerus , breadth of proximal end : less than 50 percent of total length ( 0 ) ; more than 50 percent of total length ( 1 ) .
43 . iliac pedicel of ischium : short and poor developed ( 0 ) ; slender and well developed ( 1 ) ; wide and well developed ( 2 ) .
45 . humerus / femoral ratio of 0 , 90 or more : absent ( 0 ) ; present ( 1 ) . (
46 . lateral bulge of femur below the greater trochanter : absent ( 0 ) ; present ( 1 ) ( mcintosh , 1990 ; salgado , 1993 ; calvo and salgado , 1995 ; salgado et al . , 1997a ; wilson and sereno , 1998 ) .
2 departments of palaeobotany and palaeozoology , swedish museum of natural history , se - 104 05 stockholm , sweden .
3 centre of advanced study in geology , panjab university , chandigarh - 160014 , india .
science 18 nov 2005 : vol . 310 , issue 5751 , pp . 1177 - 1180 doi : 10 . 1126 / science . 1118806
departments of palaeobotany and palaeozoology , swedish museum of natural history , se - 104 05 stockholm , sweden .
centre of advanced study in geology , panjab university , chandigarh - 160014 , india .
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silica particles from grass in fossil dung from cretaceous sauropods suggest that grasses evolved earlier than had been thought , providing food for dinosaurs and early mammals .
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is derived from
rincón de los sauces
( the area where its fossils were discovered ) and the greek
sauros
( lizard ) . the
, is derived from the latin
cauda
( tail ) and
mirus
( amazing , wonderful , marvellous ) , in reference to the unusual shape of its tail vertebrae .
( mrs - pv 26 ) is a series of 13 tail vertebrae and two ilia ( hip bones ) .
• jorge o . calvo and b . j . g . riga ( 2003 )
gen . et sp nov . , a new titanosaurid ( dinosauria , sauropoda ) from the late cretaceous of patagonia , argentina
revista geológica de chile , vol . 30 , no . 2 , p . 333 - 353 .
• vandana prasad , caroline a . e . strömberg , habib alimohammadian and ashok sahni ( 2005 )
science 18 nov 2005 : vol . 310 , issue 5751 , pp . 1177 - 1180 .
• casal , g . ; martínez , r . d . ; luna , m . ; sciutto , j . c ; and lamanna , m . c . ( 2007 )
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see also calvo and gonzález riga 2003 , calvo et al . 2007 , calvo and porfiri 2010 , campos et al . 2005 , casal et al . 2007 , filippi and garrido 2008 , franco - rosas et al . 2004 , gonzález riga et al . 2009 , kellner et al . 2005 , kellner et al . 2006 , lopes and buchmann 2008 , mannion and otero 2012 , salgado and carvalho 2008 and santucci and bertini 2006
type specimen : mrs - pv 26 . its type locality is cañadón río seco , which is in a santonian fluvial - lacustrine sandstone in the bajo de la carpa formation of argentina .
average measurements ( in mm ) : femur length 900 . 0 , femur diameter 140 . 0 , femur circumference 340 . 0 , humerus length 790 . 0 , humerus diameter 120 . 0 , humerus circumference 300 . 0 , scapula 820 . 0 x 130 . 0
estimated body mass : 3 . 59 tons based on femur circumference , femur length , and humerus circumference
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in reference to rincón de los sauces ( neuquén province , argentina ) where the dinosaur was found ; saurus ( greek ) , lizard .


wallace 's flying frog or the abah river flying frog ( rhacophorus nigropalmatus ) is a moss frog found at least from the malay peninsula into western indonesia , and is present in borneo and sumatra .

it is named for the biologist , alfred r. wallace , who collected the first specimen to be formally identified .

r. dennysii , r. maximus and polypedates feae were once contained within wallace 's flying frog as subspecies .

similar frogs also occur in laos , vietnam , minnesota , thailand and southern china ; these may be r. nigropalmatus or an undescribed , closely related species .

wallace ' s flying frog

wallace ' s flying frog is named after alfred russel wallace , naturalist who described this species in the 19th century .
wallace ' s flying frog can reach 4 inches in length . females are larger than males .
beautiful painting of wallace ' s flying frog ( rhacophorus nigropalmatus ) by the unique carel brest van kampen .
wallace ' s flying frog is carnivore ( meat - eater ) . its diet is based on various insects .
wallace ' s flying frog
( on - line ) . accessed ( date unknown ) at urltoken .
the flying frog has a diet that consists of insects , and other small invertebrates ( wallace ' s 1999 ) .
wallace ' s flying frog is it jumping or gliding ? habitat a habitat is where an animal lives . my frog , the wallace ' s flying frog lives in dark asian rainforests and in china because of the wet grounds and trees . adaptations adaptations are characteristics an animal has to survive and live . the wallace ' s flying frog has adaptations like webbed feet to glide , sticky feet to hop from branch to branch if threatened or in search of prey . prey prey is an insect or animal another animal may eat but some animals are plant eater ( omnivore ) . the wallace ' s flying frog eats insects and some meat from what it finds off the ground . when it ' s getting insects predators predators are animals that eat other animals . the wallace ' s flying frog ' s predator is a tree snake . the tree snake gets to the wallace ' s flying frog because the tree snake lives it ' s life in trees and so does the wallace ' s wallace ' s flying frog my frog is the wallace ' s flying frog and it actually glides not flies but it can jump up to 50 feet in the air and can glide down .
wallace ' s flying frog , also known as parachuting frog , is a type of asian frog that belongs to the family rhacophoridae . this frog can be found in the southeastern parts of asia . wallace ' s flying frog inhabits moist tropical rainforests of borneo and malaysia . habitat destruction ( as a result of increased deforestation ) and pet trade negatively affect number of wallace ' s flying frogs in the wild . despite that , population of wallace ' s flying frogs is still large and stable . these animals are not on the list of endangered species .
wallace ' s flying frog has broad head with rounded snout , large eyes and eardrums . it has slender body and long legs .
wallace ' s flying frog or the abah river flying frog ( rhacophorus nigropalmatus ) is a moss frog found at least from the malay peninsula into western indonesia . it is named for the biologist , alfred r . wallace , who collected the first specimen to be formally identified .
wallace ' s flying frogs mate during the rainy season . they gather in large groups called armies .
pakcenter , * . 1999 .
wallace ' s flying frog
( on - line ) . accessed november 12 , 1999 at urltoken .
1854 illustration of reinwardt ' s flying frog rhacophorus reinwardtii by jean gabriel prêtre . image in public domain .
tiny but colorful wallace’s flying frogs are bright green with yellow sides . they can grow up to four inches .
where do they live ? wallace’s flying frogs live in the tropical rain forests of malaysia and borneo . wallace’s flying frogs stay in the trees almost all the time . they only come to the ground to mate and lay eggs .
subscribe for more bbc highlights : urltoken more about this programme : urltoken bill watches wallace ' s flying frog - with feet as parachutes - falling with style .
wallace ' s flying frog spends its time in the trees where it can glide in the air due to the adaptations of its extremely webbed feet and skin folds along side of the body .
the webbing of the feet and skin folds helps catch the air like a small sail thus giving the ability to glide . this species has the ability to glide up to 50 ft . wallace ' s flying frog
wallace ' s flying frogs are arboreal animals ( adapted to the life in the treetops ) . they move to the ground only to reproduce and lay eggs .
wallace ' s flying frog does not fly - it glides through the air to escape from predators such as large snakes . this animal is able to travel distance of 50 feet and safely land on another branch or tree . wallace ' s flying frog glides diagonally ( at the angle of less than 45 degrees ) and gets closer to the ground with each gliding session . this type of movement is also known as
parachuting
.
wallace ' s flying frog has extremely large feet with fully webbed toes and fingers . toe pads are oversized and disc - shaped . flap of loose skin ( on the sides of the body ) stretches between the limbs .
wallace ' s flying frog is brightly green colored . lateral sides of the body , thighs , toes and snout are covered with yellow patches . throat and belly are white or pale yellow colored . skin is smooth or slightly granulated .
also known as parachute frogs , wallace ' s flying frogs inhabit the dense tropical jungles of malaysia and borneo . they live almost exclusively in the trees , descending only to mate and lay eggs .
the overachieving wallace ' s flying frog wasn ' t content to just hop and swim . thousands of years of watching birds navigate the rain forest and avoid predators by taking to the sky appears to have convinced this unique amphibian that air travel is the way to go .
unusual feet and flap of skin serve as a parachute which facilitates movement through the air . soft , disc - like pads ensure gentle landing . wallace flying frog also uses large pads to stick to the surface of trees .
wallace ' s flying frogs are not the only frogs who have developed this ability , but they are among the largest . the black color of their foot webbing helps distinguish them from their similarly aerial cousins .
a vanishing resource wallace’s flying frogs prefer breeding and laying eggs in the smelly wallowing holes of the asian rhinoceros . however , the asian rhinoceros is almost extinct . this may harm the population of these frogs .
wallace ' s flying frogs have external fertilization . female lays eggs into the foam and waits for male to cover them with sperm . foamy nests are then attached to the branches above the shallow pools of water .
the wallace ' s flying frog population is considered stable , and they have special status only in certain localities . however , they are partial to breeding and laying eggs in the fetid wallowing holes of the nearly extinct asian rhinoceros , and further decreases in rhino populations may negatively affect the species .
females usually lay eggs above the wallowing holes of asian rhinoceros . unfortunately , asian rhinoceros are on the brink of extinction which means that survival of remaining wallace ' s flying frogs in the wild may be questioned also .
episode 2 of david attenborough’s conquest of the skies was on tv the other day , and i watched it ( i livetweeted throughout , mostly because i wanted to talk about their portrayal of pterosaurs and mesozoic theropods ) . and hence i have rhacophorid frogs on my mind – the mostly tropical afro - asian group that includes the famous rhacophorus flying frogs , the best known member of which is wallace’s flying frog r . nigropalmatus from indonesia , thailand and adjacent countries . as usual , flying frog were used by sir david to help illustrate the diversity of animals that have evolved a gliding ability .
its back is bright shiny green and the underside is white to pale yellow . the upper sides of the inside toes , as well as the outer parts of the toe and finger webbing , are brilliant yellow . the base of the webs and one flank spot per side are jet black . overall , this frog looks much like the green flying frog ( r . reinwardtii ) and r . kio , which even if full grown do not reach the size of wallace ' s flying frog , though , and have more orange web fringes .
a moss - textured rhacophorid frog . specimens of vietnamese mossy frog or tonkin bug - eyed frog ( theloderma corticale ) . top photo by steven g . johnson , cc by - sa 3 . 0 . lower image by václav gvoždík , cc by - sa 2 . 5 .
the most famous illustration of a rhacophorid ever published : a gliding rhacophorus nigropalmatus from alfred russell wallace ' s 1869 the malay archipelago . wallace wrote about this species and illustrated it in his notes ( he didn ' t discover it though - that honour goes to charles hose ) .
tapley , b . 2009 . aspects of captive husbandry of taylor’s bug - eyed frog , theloderma stellatum ( taylor , 1962 ) . herpetological bulletin 108 , 31 - 33 .
this is the flying frog of borneo , mentioned by evolutionary biologist alfred russel wallace in his 19th century book on the malay archipelago . this emerald green frog is easy to recognize : the big eyes are pale yellow ; the hands and feet are huge and appear oversized . there is bright yellow color along arms and legs , contrasting with the black webbing . fingers and toes terminate in large adhesive pads .
a master falconer shows how his bird protects valuable u . s . crops ( hbo )
the views expressed are those of the author ( s ) and are not necessarily those of scientific american .
do they really fly ? when a predator threatens them or when they are looking for prey , wallace’s flying frogs will leap from a branch . the membranes ( thin layer of skin ) between their toes and the loose skin flaps on their sides help them glide , sometimes 50 feet or more . their toe pads help provide a soft landing , and also help them stick to tree trunks .
size of adults : up to 90 mm in males , 100 mm snout - vent length in females . the largest tree frog on borneo .
holotype specimen of philautus maia , showing eggs preserved in contact with ventral surface . did this frog really carry its eggs around like this ? illustration from günther ( 1876 ) .
hertwig , s . t . , lilje , k . e . , min , p . y . , haas , a . & das , i . 2012 . molecular evidence for direct development in the rhacophorid frog , philautus acutus ( rhacophoridae , anura ) from borneo . the raffles bulletin of zoology 60 , 559 - 567 .
byrne , p . g . & whiting , m . j . 2011 . effects of simultaneous polyandry on offspring fitness in an african tree frog . behavioral ecology 22 , 385 - 391 .
little is known about the habits of this frog because it lives high up in the canopy of primary lowland rainforest . it comes down from the trees to reproduce in forest ponds such as pig wallows .
wallace’s flying frog and the other gliding rhacophorus species are pretty remarkable . they’re very big compared to most other members of the group , svls being 90 - 100 mm in females and 80 - 90 mm in males . their fully webbed hands and feet are enormous , but they also have flaps of skin – winglets , if you like – on the arms and legs , and sometimes on the body . glides of more than 15 m have been recorded . exactly how many rhacophorus species are true gliders is uncertain : the ability is confirmed for just a handful of species but more may have it ( inger & stuebing 2005 ) . several smaller ones ( including r . angulirostris , r . cyanopunctatus and r . gauni ) have only partial digital webbing and either lack those skin flaps or only have small versions .
the outsides of these foam nests dry to form a hard crust , thereby protecting the eggs within . however , monkeys , snakes and other predators will break into the nests and eat the eggs if they can . most surprisingly , fornasini’s spiny reed frog afrixalus fornasini ( a member of hyperoliidae ) is a documented foam nest predator , though it can only eat from the nest before the foam has dried ( channing 2001 ) .
makes a long , low final approach to the ground in order to slow down and make a smooth landing . special bones help the frog snugly press the tiny suction pads on feet and toes against the surface of a tree , giving it a firm grip for landing .
episode 2 of david attenborough ' s conquest of the skies appeared on tv the other day , and i watched it ( in fact , i livetweeted throughout , mostly because i wanted to talk about their portrayal of pterosaurs and mesozoic theropods ) .
grosjean , s . , delorme , m . , dubois , a . & ohler , a . 2008 . evolution of reproduction in the rhacophoridae ( amphibia , anura ) . journal of zoological systematics and evolutionary research 462 , 169 - 176 .
- . , manamendra - arachchi , k . , schneider , c . j . & pethiyagoda , r . 2007 . new species amongst sri lanka’s extinct shrub frogs ( amphibia : rhacophoridae : philautus ) . zootaxa 1397 , 1 - 15 .
this flying frog is about 90 - 100 mm in total length , with distinct large eyes . the tympanum is prominent , at 1 / 2 to 2 / 3 of the eye diameter . snout is rounded and is not projected outward . head is as broad as it is long , sometimes slightly broader . finger tips are expanded into large , oval discs with third finger wider than tympanum ( berry 1975 ) . hands and feet are fully webbed . toes discs are smaller than finger discs . there is an oval inner metatarsal tubercle present while the outer metatarsal tubercle is absent . no supratympanic fold is present ( berry 1975 ) .
images of the juvenile and the perching adult © kurt - hock ping guek . metamorphs and juveniles look dramatically different from the adults . note that the webbing in the juvenile appears less extensive than the hand webbing of the adult . this might indicate a different microhabitat and different predation pressure in juveniles . flying adult from below by © ch ' ien lee .
records from northern thailand , lao people ' s democratic republic , viet nam and china are tentatively referred to rhacophorus reinwardtii for the purposes of this assessment , but further taxonomic work is needed . previously , r . maximus , r . dennysii and r . feae were considered to be subspecies of r . nigropalmatus .
a nice illustration of this is provided by meegaskumbura et al . ’s ( 2002 ) documentation of more than 100 new rhacophorid species on sri lanka alone ( just 18 sri lankan rhacophorid species were known prior to their work ) , a discovery that makes sri lanka on par with madagascar , new guinea and borneo in terms of anuran diversity .
rhacophorids are sometimes called flying frogs , shrub frogs , bush frogs , moss frogs , old world treefrogs , or afro - asian treefrogs , and occur across sub - saharan africa , china , much of tropical asia , japan , the philippines and sulawesi . about 380 species are recognised as of early 2015 . the last time i wrote about this group – december 2008 – this number was more like 290 , so the rate at which new species are discovered and named is pretty impressive .
haas , a . , hertwig , s . t . , krings , w . , braskamp , e . , dehling , j . m . , min , p . y . , jankowski , a . , schweizer , m . & das , i . 2012 . description of three rhacophorus tadpoles ( lissamphibia : anura : rhacophoridae ) from sarawak , malaysia ( borneo ) . zootaxa 3328 , 1 - 19 .
the latter is most developed in the grotesque rough treefrog theloderma horridum of thailand , peninsula malaysia and borneo . indeed , this is one of several species ( most of which belong to theloderma ) that resemble moss or bark in external texture and colour [ adjacent photos of t . corticale by steven g . johnson and václav gvoždík ] . t . asperum – patterned in brownish and pale blotches – superficially resembles a bird dropping and is sometimes called the bird poop frog . vocal sacs are absent in some taxa ( like nyctixalus ) but big and obvious in others .
external appearance is variable in rhacophorids . for all their fame as ‘flying frogs’ , it has to be said that the vast majority look – to those unenlightened in anuran diversity – like standard ‘treefrogs’ . they’re generally small ( svls of 40 mm or less ) , wide - headed , big - eyed frogs with expanded digit - tips and a ( normally ) prominent tympanum . many are smooth - skinned but spiny tubercles cover the skin in some taxa , and others are notably warty , with a rough , bumpy skin that aids camouflage .
then there are those rhacophorids that manufacture arboreal foam nests [ adjacent nest photos by alpsdake and brian gratwicke ] . the females exude a secretion that they ( and their male partners ) whip up with their legs to form a foam clump that’s attached to leaves , branches or aerial roots . it seems that the production of this secretion is quite costly and that a female needs to take a break and re - hydrate herself by soaking up standing water before she can complete a single nest .
this strategy is present in the afro - asian foam - nest frogs chiromantis , most rhacophorus species , and members of polypedates . in some species – most famously the grey foam nest treefrog c . xerampelina of south - eastern africa – large numbers of these frogs sometimes choose to nest in the same place , meaning that branches or aerial roots can be festooned with whole lines of dripping foam nests . actually , it isn’t just that the frogs ‘choose’ to nest in the same place – males will deliberately get in on the action if they see a pair working to make a nest , and the result is that single egg clutches are invariably fertilised by more than one male . byrne & whiting ( 2011 ) showed that this multiple paternity – technically , it’s simultaneous polyandry – assists in the survival of the resulting offspring , so it’s certainly in the interests of females to solicit as much male attention as possible during these breeding events . [ photos below by brian gratwicke , kapenta and chintan sheth . ]
the great paradox is that amphibians are in chronic global decline at the same time , and many species can no longer be located at all . despite meegaskumbura et al . ’s ( 2002 ) 100 + new rhacophorid species , they were unable to find many that had been described in the 19th century , a discovery which implies that the species concerned have gone extinct . as you should also know , amphibian species are currently being ‘lost’ on a regular and worrying basis – we don’t talk of a ‘global amphibian crisis’ for nothing .
this flying frog is about 90 - 100 mm in total length , with distinct large eyes . the tympanum is prominent , at 1 / 2 to 2 / 3 of the eye diameter . snout is rounded and is not projected outward . head is as broad as it is long , sometimes slightly broader . finger tips are expanded into large , oval discs with third finger wider than tympanum ( berry 1975 ) . hands and feet are fully webbed . toes discs are smaller than finger discs . there is an oval inner metatarsal tubercle present while the outer metatarsal tubercle is absent . no supratympanic fold is present ( berry 1975 ) . skin is smooth or finely granulated above , coarsely granulated below except smooth on the throat . a broad flap of skin on outer edge of forearm and tarsus , and another is at the heel and above the vent . a narrow flap on the inner edge of the arm is present ( berry 1975 ) . body color is shiny green with minute white markings and a couple large white patches on the thighs . the flanks , the inside of the thighs , and the undersurfaces of the body are yellow . the interdigital membranes are jet black at the base . these membranes are yellow and veined with black towards the border . the upper surfaces of toes on to four are yellow . ventral surface of the head and body is whitish ( berry 1975 ) .
a really interesting thing that’s been noted for rheophilous tadpoles is that their limb development seems to be offset , time - wise , relative to the condition in related , non - rheophilous species . this is presumably an evolutionary response to the fact that developing hindlimbs might affect their streamlining and ability to cling to rocks in fast - flowing water . they also keep sucker - like mouths and other features for longer than do other tadpoles ( nodzenski & inger 1990 ) . accordingly , it can be difficult to say reliable things about their age and estimated metamorphic stage .
- . , grant , t . , faivovich , j . , bain , r . h . , haas , a . , haddad , c . f . b . , de sá , r . o . , channing , a . , wilkinson , m . , donnellan , s . c . , raxworthy , c . j . , campbell , j . a . , blotto , b . l . , moler , p . , drewes , r . c . , nussbaum , r . a . , lynch , j . d . , green , d . m . & wheeler , w . c . 2006 . the amphibian tree of life . bulletin of the american museum of natural history 297 , 1 - 370 .
finally , there are yet other rhacophorids where egg clumps are laid in arboreal settings , but not in foam nests . in some theloderma species , egg clumps are laid in water - filled tree hollows , and the tadpoles complete their development here . in captivity , these frogs will lay their egg clumps attached to the bark , just above the hollow , the hatching tadpoles then dropping into the water ( tapley 2009 ) . oh , there are also a few foam - nesting rhacophorus species that lay their eggs in tree hollows , the most famous of which is r . vampyrus from vietnam ( after hatching inside a foam nest , the tadpoles drop into a water - filled tree hollow ) . this species saw international stardom a couple of years ago when it was revealed that the tadpoles have black , hooked fangs on the lower jaw that –it ' s presumed – are used when feeding on unfertilised eggs provided by their mother : these tadpoles , it seems , practise obligate oophagy , eating r . vampyrus eggs and nothing else ( vassilieva et al . 2013 ) .
when threatened or in search of prey , they will leap from a branch and splay their four webbed feet . the membranes between their toes and loose skin flaps on their sides catch the air as they fall , helping them to glide , sometimes 50 feet or more , to a neighboring tree branch or even all the way to the ground . they also have oversized toe pads to help them land softly and stick to tree trunks .
they are generally bright green with yellow sides and grow to about 4 inches . they survive mainly on insects .
we tried amazon key . the strangers it let in our door wasn ' t the worst part .
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mating takes place on the ground , during the night . females produce liquid substance and stir it with hind legs until it becomes foamy .
when embryonic development completes , foamy nests fall apart and tadpoles fall directly into the water where they can finish metamorphosis and turn into adult frogs .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
frost , d . r . 2014 . amphibian species of the world : an online reference . version 6 . 0 ( 7 july 2014 ) . electronic database . american museum of natural history , new york , usa . available at : urltoken .
justification : listed as least concern in view of its wide distribution , presumed large population , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .
this species is known from mainland southeast asia with certainty from the thai - malay peninsula from ranong south to endau - rompin ( taylor , 1962 and berry , 1975 ) , from borneo from eastern sabah south to southeastern kalimantan and west to central sarawak , and also from sumatra , in indonesia . it probably occurs more widely than current records suggest , especially in areas between known sites . it occurs up to 600m asl .
it is generally a rarely encountered species ; this is because it is only seen when it comes down from the trees to its breeding aggregations where it can be locally common ( grandison , 1972 and dring , 1979 ) .
it is found typically in primary evergreen rainforest , but it has also been found in old shifting cultivation , but not in open areas . breeding aggregations form in vegetation near forest pools , descending from higher strata in the forest to breed at rain pools , and also animal wallows , and usually egg masses are attached to low vegetation overhanging these pools .
deforestation and further degradation of rainforest habitat constitute the most significant threats . there is small - scale pet trade of this species , but this is not considered a threat .
it occurs in a number of protected areas . safeguarding the integrity of protected areas is a primary objective .
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berry , p . y . ( 1975 ) . the amphibian fauna of peninsular malaysia . tropical press , kuala lumpur .
jurgen , f . , richter , c . , and jacob , u . ( 1988 ) . atlas of reptiles and amphibians for the terrarium . tfh publications , neptune , n . j .
habitat ranges from trees , bushes , palms , etc . in primary rain forests and logged rain forests . can be found at elevations ranging from 700 to 1800 feet ( berry 1975 ) .
before mating , the female produces a fluid that she beats into a foam with her hind legs . she then lays her eggs in this bubble nest , at the same time the male fertilizes the eggs with his sperm . the egg nest is then hung over a source of water . when the embryos inside the eggs have developed into tadpoles , the nest falls apart . the young fall into the water and begin life as tadpoles . it is important that the tadpoles don ' t drop onto dry surface , if so the tadpoles will simply dry out and die ( pakcenter 1999 ) .
listed as least concern in view of its wide distribution , presumed large population , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .
reproduction : female produces fluid and converts it into a foam nest by the beating action of her hind legs . she then lays her eggs into the foam nest and at that time the male fertilizes these eggs . the nest is made on branches or leaves above the water . when the embryos inside the eggs have developed into tadpoles , the nest deteriorates and the larvae drop into the water . larvae live in the water until metamorphosis is completed ( richter & jacob 1988 ) .
are large , its limbs are very long , and its fingers and toes are webbed right to the tips . together with a fringe of skin stretching between the limbs , this
can parachute to the forest floor from high in the trees where it is normally found .
( amphibia , anura ) : identity of red - webbed forms and description of a new species from assam .
. version of 2005 - jun - 29 . retrieved 2007 - jun - 22 .
. version of 2003 - apr - 12 . retrieved 2007 - jun - 22 .
van dijk , p . p . ; iskandar , d . & inger , r .
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according to inger & stuebing ( 2005 ) the tadpole grows up to 50 mm in total length . it has a conspicuously spotted tail and a grayish color on head and trunk . tadpoles are most often found in turbid , muddy water , as it is characteristic for pig and rhino wallows . tadpoles hatch from a foam nest attached to overhanging vegetation or to the mud banks .
cladogram from meegaskumbura et al . ( 2002 ) , the sri lankan taxa being shown in blue . note the huge number of species that had not been named when this study was published .
and , yes , more than 100 new species announced in a single paper . if we look at the discovery record of various of the rhacophorid lineages , we see that – for example – 43 new species of raorchestes , 30 new species of rhacophorus , and 51 new species of pseudophilautus have been named since 2000 . . . 9 new raorchestes species were named in 2014 alone ( frost 2014 ) . as should be well known , the number of recognised amphibian species has sky - rocketed in recent years , and this really is because of newly discovered species , not just the result of splitting , taxonomic elevation of subspecies , or the recognition of cryptic species that can only be distinguished genetically .
it has to be said that some rhacophorids are not really all that remarkable when compared to , say , familiar ranid frogs . this montage shows tadpoles and an adult of rhacophorus arboreus . however , this species is a foam nester , on which see below . tadpole photo at lower left by σ64 , other photos by alpsdake . all cc by - sa 3 . 0 .
specialised reproductive strategies are widespread across these frogs , and some of the techniques they use mean that they don’t have to come down to the ground to breed . some ( like some philautus species ) stick their eggs to the undersides of leaves above the ground and some philautus species ( like p . mjobergi ) have been reported to be nepenthiphilous – that is , to lay their eggs inside pitcher plants . while some frogs definitely are nepenthiphilous , the only alleged rhacophorid eggs discovered inside a pitcher plant and subjected to molecular testing turned out to be from the microhylid species microhyla borneensis ( hertwig et al . 2012 ) . [ photo below by katja rembold . ]
frogs belonging to several lineages are documented as users of pitcher plants ( this photo shows a heterixalus inside a dead nepenthes madagascariensis . heterixalus is a hyperoliid , not a rhacophorid ) . it has been claimed that rhacophorids of several species use pitchers in this way , but the cases are either controversial or turned out to be erroneous . photo by katja rembold , cc by - sa 3 . 0 .
those philautus eggs , by the way , don’t produce free - swimming tadpoles : philautus species are direct developers , which means that the embryos change to froglets within the eggs , a free - living tadpole phase being absent ( the developing embryos are lecithotrophic or endotrophic , meaning that they depend on a yolk store ) . direct development is also the case in pseudophilautus and raorchestes .
while ( as just mentioned ) some of these direct developers stick their eggs to leaves that are alive and well above ground - level , others come down to the ground and lay their eggs beneath dead leaves . meegaskumbara et al . ( 2007 ) said that these ground - breeding species “deposit their eggs in nests excavated on the forest floor” ( p . 9 ) . waitaminute – frogs excavating nests ? really ? i have to look into this . . .
rhacophorid foam nests in asia and africa . at left : nests of rhacophorus arboreus in japan , photo by alpsdake , cc by 3 . 0 . at right : chiromantis rufescens foam nest in gabon , photo by brian gratwicke , cc by 2 . 0
montage of chiromantis and kin . at left : c rufescens ( photo by brian gratwicke , cc by 2 . 0 ) . top right : c . xerampelina ( image by kapenta , cc by - sa 4 . 0 ) . lower right : feihyla vittata ( photo by chintan sheth , in public domain ) . the feihyla species were once included within chiromantis but have since been recovered in several alternative placements in molecular phylogenies .
polypedates leucomystax pair in amplexus . i think i spot some subtle sexual dimorphism . image taken in java , indonesia , by w . a . djatmiko , cc by - sa 3 . 0
the eggs hatch inside the clump , the tadpoles dropping into the stream or pool ( sometimes originally formed by rhinos or pigs ) below after several days . polypedates leucomystax bucks the trend by sometimes making foam nests on the ground ( inger & stuebing 2005 ) . [ adjacent photo by w . a . djatmiko ] . it seems that foam - nesting evolved just once within rhacophorids , since all foam - nesters belong to a single clade ( frost et al . 2006 , grosjean et al . 2008 , pyron & wiens 2011 ) .
foam - nester tadpoles are ectotrophic : free - swimming and completing development outside the egg , and often with a schooling habit . some live in muddy pools and are of typical , non - specialised morphology . others ( like those of rhacophorus penanorum ) are specialised stream - dwellers with streamlined bodies , sucker - like mouths and elongate , muscular tails . these tadpoles are rheophilous ( associated with fast - flowing streams ) and inhabit rocky pools that are sometimes also home to megophrys / xenophrys spadefoot tadpoles and ansonia toad tadpoles ( haas et al . 2012 ) .
amazing ' vampire tadpoles ' of rhacophorus vampyrus . ( a ) schematic view of tadpole as seen from the front , ( b ) photo of the real thing . image from vassilieva et al . ( 2013 ) .
finally finally , the possibility exists that a completely unique reproductive strategy was present in a species that now seems to be extinct . the holotype female specimen of
, collected on sri lanka prior to 1876 , had a disc - shaped mass of eggs attached to its belly , raising the possibility that members of this species carried their eggs around with them ( günther 1876 ) . alas , meegaskumbura
. ( 2007 ) discussed how unlikely this proposal was , concluding that a more plausible possibility is that the individual concerned was collected while in the process of laying and positioning her egg clutch on a leaf . alas , only further observations can establish the ‘truth’ and . . . sadly ,
century discovery and the present . don’t forget : global . amphibian . crisis .
finally , where do rhacophorids fit within the anuran radiation ? molecular studies find them to be close to ranidae , the familiar neobatrachian clade that includes european water frogs , brown frogs , the american bullfrog , leopard frogs and so many others ( frost et al . 2006 , pyron & wiens 2011 ) . they’re clearly not at all close to hylid treefrogs ( hylids are part of the same clade as glassfrogs , toads and kin ) . i also need to say that a huge amount of work – scarcely any of which is cited in the article you’re reading now – has recently been devoted to the in - group relationships of rhacophoridae , several conventional ‘genera’ being the subject of substantial disagreement due to proposals that they might be paraphyletic or polyphyletic . at the risk of elaborating further , i must stop here . oh , i seem to have blogged about anurans again .
channing , a . 2001 . amphibians of central and southern africa . cornell university press , ithaca and london .
günther , a . 1876 . note on the mode of propagation of some ceylonese tree - frogs , with description of two new species . annals and magazine of natural history ( 4 ) 17 , 377 - 380 .
inger , r . f . & stuebing , r . b . 2005 . a field guide of the frogs of borneo . natural history publications ( borneo ) , kota kinabalu .
meegaskumbura , m . , bossuyt , f . , pethiyagoda , r . , manamendra - arachchi , k . , bahir , m . , milinkovitch , m . c . & schneider , c . j . 2002 . sri lanka : an amphibian hotspot . science 298 , 379 .
nodzenski , e . & inger , r . f . 1990 . uncoupling of related structural changes in metamorphosing torrent - dwelling tadpoles . copeia 1990 , 1047 - 1054 .
pyron , a . r . & wiens , j . j . 2011 . a large - scale phylogeny of amphibia including over 2800 species , and a revised classification of extant frogs , salamanders , and caecilians . molecular phylogenetics and evolution 61 , 543 - 583 .
vassilieva , a . , galoyan , e . & poyarkov , n . 2013 . rhacophorus vampyrus ( anura : rhacophoridae ) reproductive biology : a new type of oophagous tadpole in asian treefrogs . journal of herpetology 47 , 607 - 614 .
darren naish is a science writer , technical editor and palaeozoologist ( affiliated with the university of southampton , uk ) . he mostly works on cretaceous dinosaurs and pterosaurs but has an avid interest in all things tetrapod . his publications can be downloaded at darrennaish . wordpress . com . he has been blogging at tetrapod zoology since 2006 . check out the tet zoo podcast at tetzoo . com !
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would this diet “bug” you ? these frogs eat only insects , and at the zoo , they are fed crickets .
don abbey ( author ) , michigan state university , james harding ( editor ) , michigan state university .
having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .
forest biomes are dominated by trees , otherwise forest biomes can vary widely in amount of precipitation and seasonality .
a large change in the shape or structure of an animal that happens as the animal grows . in insects ,
incomplete metamorphosis
is when young animals are similar to adults and change gradually into the adult form , and
complete metamorphosis
is when there is a profound change between larval and adult forms . butterflies have complete metamorphosis , grasshoppers have incomplete metamorphosis .
the area in which the animal is naturally found , the region in which it is endemic .
found in the oriental region of the world . in other words , india and southeast asia .
rainforests , both temperate and tropical , are dominated by trees often forming a closed canopy with little light reaching the ground . epiphytes and climbing plants are also abundant . precipitation is typically not limiting , but may be somewhat seasonal .
to cite this page : abbey , d . 2000 .
rhacophorus nigropalmatus
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
skin is smooth or finely granulated above , coarsely granulated below except smooth on the throat . a broad flap of skin on outer edge of forearm and tarsus , and another is at the heel and above the vent . a narrow flap on the inner edge of the arm is present ( berry 1975 ) .
body color is shiny green with minute white markings and a couple large white patches on the thighs . the flanks , the inside of the thighs , and the undersurfaces of the body are yellow . the interdigital membranes are jet black at the base . these membranes are yellow and veined with black towards the border . the upper surfaces of toes on to four are yellow . ventral surface of the head and body is whitish ( berry 1975 ) .
jurgen , f . , richter , c . , and jacob , u . ( 1988 ) .
sunny shah and rachna tiwari ( sunnys at uclink . berkeley . edu ) , amphibiaweb
> university of california , berkeley , ca , usa . accessed jul 9 , 2018 .
> university of california , berkeley , ca , usa . accessed 9 jul 2018 .
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