BIOGEOGRAPHY GEOG 167
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A giant frog with South American affinities from the Late Cretaceous of Madagascar Susan E Evans Malt E H Jones and David W Krause nat m BulldlngUCL United Gower street London WClE dBT t ri Brook NY i WHOM Edited by Dayid 3 Wake university or California Berkeley CA and approved lanuary 7 2002 reeled ror reyiew August i i 2007 Madagascar has a diverse but mainly endemic frog fauna the biogeographic history of whi h h g e ed intense debate r ce av 3 a E in tion of mantellids and microhylids that now dom fauna in addition the presence of a ceratophryine provides ceous it also suggests that the initial radiation of hyioid anurans began earlier than proposed by some recent estimates Anura i ceratophryinae i Gondwaria i south America i Hyloidea The MadagascareSeychellesIndia Plate separated from Afr rica 160 M a and bega ragment into it y n to f s Uomponent s 1 hese ev ts the pla e was also contact with AntarcticaeAustralia to the south but the e and nat of th39 is deb 173 Ultimately the Indian subwnt39 t n d h t ntact Eurasia a d Madagasc was 1 ft 430 r lated by the deep Mozambiq e C el Th arkable diversity and en s micity ofthe Malagasy biota has prompted intense debate as to the role of vicariance nd more recently of dispersal in it analyses have clarified the relatiorships ofkey groups leg the monophny of Malagasy mantellids 1043 Of gt200 docus o the ranid sis o fe n t of the endemic Malagasy mantellids and microhylids is equivocal and both vicariantsbased 7 12 16e18 and dispersalsbased 5 6 19 hypotheses have been good fossil record would aid the debate but apart from the instead seems to be related to South American hyloids of the clade Ceratophryinae note t at we use a standard taxonomic nomenclature 23 rather than the oomprehensive but not yet widely adopted classification of Frost at al Sysienratit Paleontology Systematic paleontology is as follows Anura Rafinesque atrachia Reig 1958 Hyloidea Ford and Cannatella 199 Ceratophryinae Tschudi 1838 Beale zebufg umpmgu gen et sp nov Ilololype The holotype is as follows fused z ervical atlantal d presaoral z entra Universite d39Ant ananal39ivo oolleos K 8 l and tions Madagascar specimen no UA 9600 Fig 1 El mology The generic name is based on Beerzebuz Greek Devil and Bufa Latin toad in reference to t e size and probable life appearance of this anuran the specific epithet umpmgu Malagasy means shield in reference to cranial hyperossification Loalily and lloriron The holotype was collected from locality MAD e25 15 5439 17714quot 3 46 3439 55946 E Berivotra Study Area Anembalemba em er Maevarano m a e janga Basin adagascar The Anembalemba Member is Late Cretaceous Maastriohtian in age 122 Diagnosis The anuran resembles adults ofall extant Ceratophryis nae and differs from all other hyperossified anurans in o e in39 exostosed skull roofing bone u 39 lateral parieta nsion an r e palatine shelfont e adult premaxilla and anterior maxilla 25 r 39 inmu la 0 maturity it resem Cemmphrys an amp rys in tightly interlocking maxillaspremaxilla articulation it resembles tz7 in having cervical oot es oontinuous in Ventral 39 n in h possession of posttemporal fenestrae but differs in that rostral tips of nasals fail to un39 nde ly39 39 r lized nasal cart39 a es39 39 in 39 E s i ges tween pits an more slender session of strong r quadratojugal and in much larger body size Description and comparison The external cranial elements of the Malagasy fossil anuran show a distinctive coarse pitsandsridge Author ontrlbu on EE designed rexezrth EE M E Hi and DWK perrormed researchsEE anaiyzed data andsEE and DWK wrom the paper identified Late Cretaceous finds 21 this record has been lacking Here w d rib a very large hyperossified anuran from the Up er Creta Maevarano Formation Fm of Madagascar me 65 Mya 1 22 This anuran differs from extant Malagasy taxa all of which are ranoid neobatrachians and wwwpnas olggldollo l073pria 0707599l05 rhisartitieisarNAsoirettsuomission trowhomtorrespondenteshouid be addressed Ermali ultgame ultl 2uk rhis artlde ontaln supporting information oniine at wwprlaxorgglontentmii 0707599l05Dtl and oyrhe National Atademy or tientes or the UiA PNAS i rebruaryzszone i vol in i no 8 i 19514956 Evolllrloll Fig 1 Beelzebufo ampinga holotype fused cervical and second presacral centra UA 9600 stereophotographic pairs of anterodorsal A dorsal B and ventral C views cc confluent cotyles cv cervical vertebra v2 second pre sacral vertebra spn spinal nerve foramen between arch pedicels The small arrow indicates the line of central fusion Scale bar 5 mm sculpture that in conjunction With the size and robustness of the bones permits attribution of elements and is consistent With the presence of a single large hyperossified anuran species in the Maevarano Fm Accordingly the description and reconstruction are based on gt60 bones collected from 26 localities Within a 18km radius see supporting information SI Text for specimen list These bones include parts of all cranial roofing bones the palatoquadrate braincase jaws vertebrae pelvis and limb elements Figs 1 2A and B and 3 As reconstructed the skull of Beelzebufo is Wider than long Fig 2A With thick tightly sutured dermal roofing bones The premaxillae Fig 3 A and B are unsculptured but their attri bution to Beelzebufo is confirmed by their size and matching maxillary articulation They are distinctive in lacking projecting palatine shelves and closely match corresponding elements of extant Ceratophrys The maxilla Fig 3 C and D is exostosed except for the pars dentalis and had strong articulations With the premaxilla nasal squamosal quadratojugal and presumably neopalatine and pterygoid not identified Like the premaxilla the anterior half of the maxilla lacks a palatal shelf Teeth were present on the premaxilla 13 or 14 teeth and maxilla 50 60 teeth but are damaged Their implantation resembles that of the nonpedicellate Ceratophrys In one partial maxilla FMNH Field Museum of Natural History PR 2506 sharp monocus pid unankylosed tooth tips are preserved in Sim in the broken tooth bases The large nasals Fig 3 E G are L shaped With a tapering rostral process a long recurved maxillary process and a short dorsomedial plate that was coossified With the sphen ethmoid and covered it completely Medially the nasals and frontoparietals both met their counterparts in a strong hori 2952 wwwpnasorgcgidoi101073pnas0707599105 Fig 2 Beelzebufo ampinga Late Cretaceous of Madagascar A Skull reconstruction showing parts preserved white areas Left and distribution of pitandridge ornament stippling Right B Skeletal reconstruction and inferred body outline of averagesized skull width 200 mm SVL 425 mm adult female B ampinga based mainly on Lepidobatrachus asper 32 White areas indicate parts represented by fossil specimens For size comparison dorsal view silhouettes of Ceratophrys aurita the largest extant ceratophry ine C and Mantidactylus guttulatus the largest extant Malagasy frog D are shown cp crista parotica fm foramen magnum frp frontoparietal mx maxilla n nasal pmx premaxilla qj quadratojugal qu quadrate sq squa mosal Scale bars 50 mm zontally laminated suture Posteromedially each frontoparietal Fig 3 was fused to the underlying endocranium UA 9675 Fig 3P but posterolaterally the bone extended into a shelf that was separated from the crista parotica by a distinct posttemporal fenestra 25 26 As preserved UA 9640 the shelf is embayed posteriorly but less so than in Ceratophrys It probably met the squamosal behind the orbit to form a parietosquamosal bridge but the lateral tip of the bone has not been identified With certainty The squamosals were triradiate With a zygomatic ramus that was strongly sutured to the maxilla Fig 3 H and I a flangelike horizontal otic ramus Fig 310 and preserved only at the base a ventrolateral ramus Whether the zygomatic process of the squamosal met the nasal below the orbit is uncertain The only nasal With a complete ventrolateral tip UA 9680 Fig 3G suggests abutment rather than a suture but the bone is from a juvenile and the relationship may have changed With maturity A slender quadratojugal met the maxilla anteri orly and was coossified posteriorly With a robust mineralized quadrate pars quadrata palatoquadrati The cervical vertebra atlas is characterized by large confluent anterior cotyles Fig 1A Type 3 25 and by fusion of its centrum to that of the second presacral Fig 1 B and C as in some large Ceratophrys 25 Known presacrals are pro coelous The sacral vertebra Fig 3L is bicondylar posteriorly and has a dorsoventrally compressed diapophysis that is slightly flared distally unlike the cylindrical diapophyses of ranids The presence of an anterior cotyle rather than a condyle on the sacral centrum suggests that the last presacral was not biconcave as it is in many ranoid anurans 16 The stout urostyle Fig 3 M and Evans et al Fig 3 Representative elements of Beelzebufo ampinga Late Cretaceous of Madagascar A and B Left premaxilla UA 9622 labial and lingual views C and D Left maxilla anterior region FMNH PR 2510 labial and lingual views E Right nasal rostral process UA 9674 dorsal view reflected F Partial left nasal UA 9629 dorsal view within scaled nasal shape G Immature right nasal maxillary process UA 9625 reflected for comparison with F dorsolateral view H Right squamosal maxillary process FMNH PR 1959 lateral view I Left squamosal partial maxillary process UA 9639 lateral view J Left frontoparietal anterior region FMNH PR 2512 dorsal view K Right squamosal otic process FMNH PR 2536 dorsal view L Sacral vertebra right half with left side added by reflection FMNH PR 2003 dorsal view M and N Urostyle anterior part UA 9636 anterior and dorsal views 0 Left tibiofibula UA 9628 posterior view P Left frontoparietal and exoccipital in posterior view with right side added by reflection UA 9675 Small arrows indicate unbroken edges ams absence of medial shelf ap alary process aps absence of palatal shelf mxa maxillaryarticulation occ occipital condyle pa premaxillary articulation pp posterior process Scale bar 10 mm N has no transverse processes The pelvis is represented by a partial right ilium UA 9681 that lacks a dorsal crest and the short broad tibiofibula Fig 30 resembles that of living anurans that are predominantly ambulatory Unlike Ceratophrys and Lepidobatrachus in which the skull bones coossify early the sutures remained patent in the largest specimens of Beelzebufo suggesting an enhanced potential for prolonged growth Extant Ceratophryinae range in size from snoutvent length SVL 40 50 mm Chacophryspierottii 28 to 170 mm Ceratophrys aurita 29 Estimates based on extant taxa SI Tables 1 and 2 indicate that most specimens of B ampinga belonged to individuals with an estimated posterior skull width SkW of 80 120 mm SVL 160 270 mm but a few bones represent anurans of significantly larger size SkW 150 200 mm SVL gt400 mm Figs 28 and 3 E and In extant ceratophryines 29 the smaller but more widely represented size class consists of subadults and adult males adult females are significantly larger The size distribution in Beelzebufo suggests a similar pattern The largest individuals would have dwarfed the largest extant ceratophryine Ceratophrys aurita Fig 2C and the largest extant Malagasy anuran Mantidactylus guttulatus Fig 2D rivaling Miocene representatives of Calyptocephalella sp Caudiverbera 30 from Argentina 31 and the extant West African Conraua goliath Discussion and Conclusions Phylogenetic Position Taken together the skeletal characters of Beelzebufo are consistent with neobatrachian affinity 25 33 The characters of the vertebral column procoely sacro urostylar articulation bicondylar weakly dilated sacral diapoph yses no urostylar transverse processes render attribution to leiopelmatids bombinatorids discoglossids pipids or peloba toids 23 24 33 unlikely and also exclude Cretaceous Asian gobiatines 34 Some characters cranial exostosis skull wider Evans et al than long palatoquadrate mineralized fused to quadratojugal parietosquamosal bridge maxillary pars facialis large with strong nasal squamosal and quadratojugal articulations cervical cotyles approaching one another in ventral midline cervical and second presacral centra fused occur in different combinations in hyperossified taxa from a range of lineages including some pelobatoids the hyloids Calyptocephalella and Hemiphractus some hylids eg Triprion Osteopilus some ranids eg Pyxi cephalusAubria and Ceratobatrachus and Ceratophryinae 24 25 33 35 see also SI Text This hyperossification complicates comparison 35 but other characters constrain the possibilities Beelzebufo lacks the derived vertebral characters of most ranoid anurans and the basal hyloid Calyptocephalella lacks posttem poral fenestrae The presence of maxillary teeth and large orbits differentiate Beelzebufo from the hyperossified Indian Nasika batrachus 36 37 although the latter s osteology remains largely undescribed Conversely a unique combination of skull charac ters posterolaterally expanded frontoparietals premaxilla strongly articulated with maxilla premaxilla and anterior maxilla without palatal shelves link Beelzebufo to Ceratophryinae 25 a small clade with three extant genera Ceratophrys Lepidoba trachus and Chacophrys and two attributed fossils Baumbatra chus Late Cretaceous 67 65 Mya Argentina 27 Wawelia Miocene 14 12 Mya Argentina 38 Other characters of Beelzebufo teeth unicuspid nasal body short and broad with tapering rostral process strong internasal sutures nasal with long recurved maxillary process bearing rounded unorna mented tip presence of posttemporal fenestrae squamosal otic process prominent horizontal cervical cotyles confluent in ventral midline Type 3 25 ilium without prominent dorsal crest are consistent with this attribution 25 28 32 39 as are fragments of possible bony dorsal shield UA 9619 see SI T ext for further comparison This hypothesis of ceratophryine relationship is supported by phylogenetic analysis Fig 4 that nests Beelzebufo within Cera PNAS February262008 vol105 no8 2953 EVOLUTION Peurodema Physalaemus Dendropsophus Hypsiboas Flectonotus Isthmohyla Phyllomedusa Litoria Scinax Thaumastosaurus Triprion Osteopilus Calyptocephalella Hemiphractus Baurubatrachus Chacophrys Wawelia Lepidobatrachus Ceratophrys Beelzebufo Megophrys RANOIDEA Bombina PIPIDAE PELOBATOIDEA Rhinoderma Allophryne BUFONIDAE I Leptodactylus Pseudis Batrachophrynus Telmatobius Heleioporus Limnodynastes Telmatobufo Mixophyes Cyclorana Odontophrynus Fig 4 Seventy percent Majority Rule Consensus of 106 equally parsimonious trees length 633 consistency index 03 rescaled consistency index 0169 Monophyletic clades not directly relevant to the discussion have been col lapsed to single nodes Asterisks denote that the clade or part of it contains exostosed andor hyperossified taxa tophryinae as the sister taxon of Ceratophrys although this latter position may reflect the more generalized morphology of Cera tophrys in comparison with the derived Lepidobatrachus and paedomorphic Chacophrys 28 32 39 Baumbatrachus and Wawelia group together sometimes with Chacophrys Both South American fossil taxa show ceratophryine postcranial features strongly elongated anterior presacral transverse pro cesses short urostyle 25 32 39 but apart from a sculptured fragment Wawelia lacks a skull and this problem limits com parison As originally described 27 Baumbatrachus is more primitive than extant taxa and Beelzebufo in lacking a frontopa rietalsquamosal bridge and in having bicuspid teeth and a palatal shelf on the anterior maxilla However ongoing restudy suggests some of this information will change A Baez personal communication We also included Thaumastosaums Eocene Europe in our analyses as it has been discussed in relation to Ceratophryinae 40 but it was placed outside the group Ecology Ceratophryinae are found today throughout South America mainly in warm seasonally dry habitats with ephem eral pools 28 29 The environment of deposition of the Maevarano Fm was comparable 22 and Beelzebufo may have resembled Ceratophrys in being terrestrial with a tendency to 2954 wwwpnasorgcgidoi101073pnas0707599105 burrow exostosis thick strongly sutured nasals 33 Cerato phryines are ambush predators that include vertebrates in their diet 29 Their strong bite is correlated with hyperossification sharp teeth and stabilizing connections between the upper jaw and skull 33 Beelzebufo has the same morphology and large adults would have been formidable predators on small vertebrates Biogeography and the Evolution of Malagasy Anura The fossil record of Gondwanan anurans is patchy Pipoids dominate the South American record 31 34 but hyloid neobatrachians are represented in the Late Cretaceous by Baumbatrachus and Estesiella with bufonids and hylids reported from the Paleocene and Calyptocephalella from the Oligocene onwards 31 Disco glossoids are recorded from the earliest Cretaceous of Morocco 41 but otherwise the African Mesozoic and early Tertiary record is limited to pipoids 34 Nothing is known from Ant arctica but the earliest Australian anuran Eocene 546 Mya is attributed to the extant myobatrachid genus Lechriodus with pelodryadine hylids and microhylids reported from the late Oligocene onward 42 The Late Cretaceous fauna of India reportedly combines Laurasian gobiatine discoglossoid pelo batoid and Gondwanan hyloidranidrhacophorid elements 43 However with the exception of the putative myobatrachid Indobatrachus these records rely on incomplete ilia and are tentative Based on the distribution patterns of living and extinct taxa but allowing for hyperoliid dispersal the Cretaceous anuran fauna of the Madagascar Seychelles India Plate or of parts of this plate after its fragmentation would have included the ancestors of endemic Malagasy taxa mantellids dyscophine scaphiophrynine and cophyline microhylids ii endemic Sey chellian taxa sooglossids and iii ancient Indian lineages Nasikabatrachus ranixaline micrixaline and nyctibatrachine ranids rhacophorids with the possibility of pipids early Afri can ranids myobatrachids and basal hylids 7 12 16 19 31 35 44 45 A ceratophryine is unexpected In conventional paleo biogeographic models eg 2 the Madagascar Seychelles India plate lost contact with the AntarcticaAustralia landmass and thus also South America 120 Mya However an alter native hypothesis posits the existence of physical links between Madagascar the Indian subcontinent and South America that persisted late into the Late Cretaceous 80 Mya As early as 1927 paleontologists 46 noted similarities between the dinosaurs of these three regions and the fossil assemblage of the Maevarano Fm provides further support with the most striking links to the Indian subcontinent and South America involving theropod and sauropod dinosaurs crocodyliforms and mam mals The late persistence of a physical connection between Madagascar and southern Gondwana has also received support from molecular studies on ratite birds 47 and on iguanian lizards podocnemid turtles and boid snakes 48 The presence of a ceratophryine anuran with South American relatives in the Late Cretaceous of Madagascar provides strong and indepen dent support for this paleobiogeographic reconstruction Fig 5 Hyloid Diversification An early molecular analysis 49 provided minimum age estimates of 55 Mya early Eocene for the origin of extant Ceratophryinae and this result is reasonably consistent with the presence of Late Cretaceous ceratophryines in South America and Madagascar However more recent analyses have dated the divergence of Ceratophrys from Lepidobatrachus to the latest Oligocene or Miocene 127 261 45 50 results that are clearly incompatible with the attribution of either Baumbatra chus or Beelzebufo to the crown group Moreover based on the relatively low levels of genetic divergence among extant hyloids 45 some analyses also date the main hyloid radiation ie without myobatrachids or Calyptocephalella as occurring at or Evans et al Fig 5 Map showing positions and coastlines of the southern continents at 80 Mya 3 and indicating localities of Beelzebufo position 1 Baurubatrachus position 2 and Wawelia position 3 soon after the CretaceousPaleogene boundary 65 55 Mya confidence limits 52 84 Mya 45 50 51 This finding is difficult to reconcile With the presence of ceratophryines or even stemceratophryines in Madagascar at 70 65 Mya even allowing for confidence limits as the island was isolated from at least 80 Mya 3 Given that the estimated dates of hyloid origin 130 152 Mya confidence limits 108 208 Mya 7 19 36 45 51 substantially predate those for the beginning of the main hyloid nobleoba trachian 24 45 radiation long fuse 45 there is potential for some lineages to have arisen earlier It is of course possible that Baurubatrachus and Beelzebufo are hyperossified stem hyloids that are convergent on ceratophryines but the skeletal evidence for ceratophryine affinity at least for Beelzebufo is compelling The relationships of the component clades of Leptodactylidae including ceratophryines are still incom pletely resolved eg refs 24 35 36 45 50 and 52 Cerato phryines have been alternately placed as basal hyloids eg refs 25 53 and 54 or more deeply nested eg refs 18 24 35 and 36 and even their sister group is uncertain eg telmatobines 24 35 hylids 36 45 hemiphractines 50 This uncertainty makes it difficult to date their origin Moreover Ceratophryinae is a very small clade and thus resembles other such clades eg sooglossids Heleophryne ranixalids Calyptocephalella Rhino derma centrolenids posited to be remnants of older formerly more Widespread lineages 44 Under this hypothesis Bauru batrachus and Beelzebufo provide at least minimum constraints on the antiquity of Ceratophryinae Conclusions We suggest that extant ceratophryines are remnants of a Gondwanan hyloid clade that once ranged from at least South America to IndoMadagascar Whether this clade was more broadly distributed and on Which Gondwanan landmass it originated cannot be determined on current evidence However as the Late Cretaceous fauna of the Maevarano Fm 1 4 including its ceratophryine anuran bears little resemblance to that of modern Madagascar major biotic changes clearly oc curred on the island in the intervening period When and how the 1 Krause DW et al 2006 Late Cretaceous terrestrial vertebrates from Madagascar Implications for Latin American biogeography Ann Mo Bot Gard 93178 208 2 Smith AG Smith DG Funnel BM 1994 Atlas of Mesozoic and Cenozoic coastlines Cambridge University Cambridge UK 3 Hay WW etal 1999 Alternative global Cretaceous paleogeography Evolution ofthe Cretaceous OceanClimate System eds Barrera E Johnson CC Geological Society of America Boulder CO Special Paper 33 pp 1 47 4 Krause DW Hartman JH Wells NA 1997 Late Cretaceous vertebrates from Madagas car Implications for biotic changes in deep time Natural Change and Human Impact in Madagascar eds Goodman SM Patterson DB Smithsonian Institution Washington DC pp 3 43 5 Vences M 2004 Origin of Madagascar39s extant fauna A perspective from amphibians reptiles and other nonflying vertebrates ItalJZool Suppl 2210 228 6 Yoder AD Nowak MD 2006 Has vicariance or dispersal been the predominant biogeo graphic force in Madagascar Only time will tell Annu Rev Ecol Evol Syst 37405 431 7 Van Bocxlaer I Roelants K Biju SD Nagaraju J Bossuyt F 2006 Late Cretaceous vicariance in Gondwanan amphibians PLoS ONE 1e74 Evans et al ancestors of the endemic mantellid and microhylid anurans arrived on Madagascar remains controversial 5 6 8 12 18 19 but there is general agreement that these frogs did not diversify significantly until the Paleogene 5 6 12 18 19 Their radiation has been linked at least in part to the expansion of rainforests but may also have been facilitated by the extinction of archaic faunal elements 5 including ceratophryines Methods Beelzebufo Baurubatrachus and Wawelia were coded into an existing mor phological character matrix 39 with 81 characters and 62 taxa including basal anurans Bombina pipids pelobatoids and ranoid and hyloid neo batrachians Some genera were originally represented by several species but these taxa were run as single sometimes polymorphic units to make the analysis more manageable The matrix was then extended to broaden the sampling of microhylids myobatrachids and hyperossified taxa see SI Textfor character list and details see SI Table 3 for matrix Parsimony analysis was performed by using PAUPversion 401 b 55 in heuristic search mode because of matrix size with default settings but multistate characters coded as polymorphism Bombina Bombinatoridae Xenopus and Hymenochirus Pipidae and Megophrys Pelobates Spea and Scaphiopus Pelobatoidea were designated outgroups This analysis resulted in 106 equally parsimonious trees length L 633 consistency index CI 03 rescaled consistency index RC 0169 The full topology 70 majority rule tree is shown in SI Fig 9 In Fig 4 pipids pelobatoids bufonids and ranoids are collapsed to single nodes The overall tree topology shows some similarities with previous analyses and anomalies eg the positions of bufonids and Megophrys although as for other morphological trees eg refs 35 and 39 and in contrast to most molecular analyses eg refs 24 35 36 45 and 52 Hyloidea does not form a monophyletic sister taxon to Ranoidea Additionally as highlighted by others 35 characters relating to size and cranial hyperossification can have a disproportionate effect eg the placement of Calyptocephalella and the hylids Osteopilus and Triprion close to ceratophryines rather than with less ossified relatives We reran the analysis using a subset of 18 exostosed andor hyperossified taxa Branch and Bound search characters unordered and un weighted multistate characters treated as polymorphism This analysis yielded nine maximum parsimony trees L 168 CI 0565 RC 0307 in which ceratophryines including Beelzebufo Baurubatrachus and Wawelia consis tently grouped together SI Fig 10 In a bootstrap analysis 1000 replicates run on the same dataset support values for most clades were low Size was estimated by comparing Beelzebufo bones with equivalent ele ments from Ceratophrys and Calyptocephalella using a range of specimens from juvenile SkW 44 mm to full adult SkW 98 mm to allow for allometry see SI Tables 1 and 2 ACKNOWLEDGMENTS We thank field teams of the Mahajanga Basin Project for collecting specimens colleagues at the Universit d Antananarivo and the Madagascar Institute pour la Conservation des Environnements Tropicaux for logistical support R Symonds C J Bell B Clark A Resetar and H Chatterjee for comparative material F Glaw M Vences R Whately and V Hutchison for information and photographs A Baez for discussion of Baurubatrachus M Fabrezi for clarification of characters J Groenke V Heisey and J Sertich for specimen preparation andor curation A Pendjiky and A Smith for technical help C Forster for suggesting the generic name Beelzebufo V Ranaivo for help with the trivial name ampinga and A Baez L Trueb and M Vences who provided valuable comments on the manuscript This work was funded in part by grants from the National Science Foundation and the National Geographic Society to DWK 8 Vences M et al 2003 Multiple overseas dispersal in amphibians Proc R Soc London B 2702435 2442 9 Measey GJ etal 2007 Freshwater paths across the ocean Molecular phylogeny ofthe frog Ptychadena newtonigives insights into amphibian colonization of oceanic islands J Biogeogr 347 20 10 Glaw F Vences M Bohme W 1998 Systematic revision of the genus Aglyptodactylus Boulenger 1919 Amphibia Ranidae and analysis of its phylogenetic relationship to other Madagascan ranid genera Tompterna Boophis Mantidactylus and Mantela J Zool Syst Evol Res 3617 37 11 Glaw F Vences M Andreone F Vallan D 2001 Revision of the Boophis majori group Amphibia Mantellidae from Madagascar with descriptions
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