A giant frog with South American affinities from the Late Cretaceous of Madagascar

Abstract

Madagascar has a diverse but mainly endemic frog fauna, the biogeographic history of which has generated intense debate, fueled by recent molecular phylogenetic analyses and the near absence of a fossil record. Here, we describe a recently discovered Late Cretaceous anuran that differs strikingly in size and morphology from extant Malagasy taxa and is unrelated either to them or to the predicted occupants of the Madagascar-Seychelles-India landmass when it separated from Africa 160 million years ago (Mya). Instead, the previously undescribed anuran is attributed to the Ceratophryinae, a clade previously considered endemic to South America. The discovery offers a rare glimpse of the anuran assemblage that occupied Madagascar before the Tertiary radiation of mantellids and microhylids that now dominate the anuran fauna. In addition, the presence of a ceratophryine provides support for a controversial paleobiogeographical model that posits physical and biotic links among Madagascar, the Indian subcontinent, and South America that persisted well into the Late Cretaceous. It also suggests that the initial radiation of hyloid anurans began earlier than proposed by some recent estimates.

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 presacral vertebra; spn, spinal nerve foramen between arch pedicels. The small arrow indicates the line of central fusion. (Scale bar: 5 mm.)

Fig. 2.

Beelzebufo ampinga, Late Cretaceous of Madagascar. (A) Skull reconstruction showing parts preserved (white areas, Left) and distribution of pit-and-ridge ornament (stippling, Right). (B) Skeletal reconstruction and inferred body outline of average-sized (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 ceratophryine) (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, squamosal. (Scale bars: 50 mm.)

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. (O) 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, maxillary articulation; occ, occipital condyle; pa, premaxillary articulation; pp, posterior process. (Scale bar: 10 mm.)

Fig. 4.

Seventy percent Majority Rule Consensus of 106 equally parsimonious trees (length, 633; consistency index, 0.3; rescaled consistency index, 0.169). Monophyletic clades not directly relevant to the discussion have been collapsed to single nodes. Asterisks denote that the clade, or part of it, contains exostosed and/or hyperossified taxa.

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).