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Fig 10 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 10. Rhombophryne proportionalis sp. nov., holotype ZSM 1826/2010, in life. (a) dorsolateral, (b) dorsal, and (c) ventral view. https://doi.org/10.1371/journal.pone.0213314.g010

Fig 8 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 8. Osteology of Mini scule gen. et sp. nov. holotype (ZSM 5943/2005). (a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g008

Fig 6 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 6. Osteology of Mini mum gen. et sp. nov. holotype (ZSM 861/2014). (a-c) Whole skeleton in (a) dorsal, (b) lateral, and (c) ventral view. (d-g) Skull in (d) lateral, (e) ventral, (f) anterior, and (g) dorsal view. (h) Foot in ventral view. (i) Hand in ventral view. Abbreviations for all osteological figures: angspl, angulosplenial; col.pip, pars interna plectra of columella; col.pmp, pars media plectra of columella; cpl(s), carpal(s); den, dentary; exoc, exoccipital; exoc.occ, occipital condyle of exoccipital; fpar, frontoparietal; max, maxilla; max.fp, facial process of maxilla; mmk, mentomeckelian; n, nasal; n.mp, maxillary process of nasal; npl, neopalatine; oc, otic capsule; prsph. ap, parasphenoid alary process; prsph.cp, parasphenoid cultriform process; prsph.pmp, parasphenoid posteromedial process; pmx, premaxilla; pmx.ap, premaxilla ascending process; pmx.lp, premaxilla lateral process; pmx.pp, premaxilla palatine process; povom, postchoanal portion of vomer; pro, prootic; prvom, prechoanal portion of vomer; pt.ar, pterygoid anterior ramus; pt.mr, pterygoid medial ramus; pt.vr, pterygoid ventral ramus; qj, quadratojugal; smx, septomaxilla; spheth, sphenethmoid; sq. or, squamosal otic ramus; sq.vr, squamosal ventral ramus; sq.zr, squamosal zygomatic ramus; tsl(s), tarsals; vt, vomerine teeth. https://doi.org/10.1371/journal.pone.0213314.g006

Fig 4 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 4. Mini mum gen. et sp. nov. in life and its habitat in Manombo Special Reserve. (a-c) ZSM 861/2014, holotype, in (a) anterolateral view on a thumbnail, (b) dorsolateral view on a leaf, (c) ventral view. (d, e) ZSM 862/2014, paratype, in (d) ventral view, and (e) lateral view on a thumbnail. (f, g) ZMB 83194, paratype, in (f) dorsolateral view, and (g) ventral view. (h) ZMB 81993, paratype, in dorsolateral view. (i) ZMA 20172 in posterodorsolateral view. (j) Habitat of the new species in Manombo Special Reserve. https://doi.org/10.1371/journal.pone.0213314.g004

Fig 2 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 2. Holotypes of the new species described in this paper and their hands. Whole specimens in dorsal (left) and ventral (right) view. Hand images not to scale. https://doi.org/10.1371/journal.pone.0213314.g002

Fig 5 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 5. One-second spectrograms and oscillograms of the calls of the new species described here. Insets represent the respective species but not the calling specimens. (a) Mini mum gen. et sp. nov., paratype ZMB 81993 from Manombo, (b) Mini scule gen. et sp. nov., ZSM 265/2018 from Sainte Luce, (c) Rhombophryne proportionalis sp. nov., part of a call (note series) of a specimen from Bepia campsite, Tsaratanana (not collected), (d) Anodonthyla eximia sp. nov., specimen not collected, from Maharira (Ranomafana). https://doi.org/10.1371/journal.pone.0213314.g005

Fig 3 in Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Fig 3. Currently known localities of the new taxa described in this paper. The base map is USGS SRTM 1-Arc second digital elevation model. https://doi.org/10.1371/journal.pone.0213314.g003

Extended Data Fig. 10 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 10 | Reconstruction of Perucetus colossus in its coastal habitat. Because portions of the skeleton are unknown, several aspects of the reconstruction are tentative:the overall proportions of the axial postcranium are based on a close relative Cynthiacetus peruvianus, which was scaled-up and dilated according to the elements recovered for P.colossus (see Extended Data Fig. 8); the skull and limbs were only scaled-up; the tail fluke and forelimb use (bottom-walking) are based on the manatee (Trichechus), the extant marine mammal with the closest degree of pachyosteosclerosis in the postcranial skeleton; the hind limb of P.colossus was not recovered, but the anatomy of its innominate indicates the presence of a reduced, articulated leg. The associated sawfish (Pristis) was recovered from the same unit in the East Pisco Basin, the Yumaque Member of the Paracas Formation87. Reconstruction by A.Gennari.

Extended Data Fig. 9 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 9 | Estimates of the osteological range of motion. Extension and flexion of the preserved portion of vertebral column of Perucetus colossus holotype (MUSM 3248) is compared with an equivalent vertebral column portion of Cynthiacetus peruvianus holotype (MNHN.F.PRU10) using the respective 3D models.Intervertebral spaces were reconstructed based on the common dolphin (Delphinus delphis)81 (see Methods). Scale bar = 50 cm.

Extended Data Fig. 8 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 8 | Estimating the whole skeletal volume of Perucetus colossus based on Cynthiacetus peruvianus' holotype (MNHN.F.PRU10). a. Unmodified 3D model of Cynthiacetus peruvianus' holotype.b,c. vertebral column scaled-up (top) and dilated (bottom).c,d. Rib cage scaled-up (top) and dilated (bottom) identifying the scanned rib of P.colossus as R17 (c) or R20 (d). Scale bars = 2 m.

Extended Data Fig. 7 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 7 | Vertebral microanatomy illustrated with binarized sections (black = bone) obtained with physical core drills (a) or µCT data(b–f). a, Perucetus colossus (MUSM 3248, holotype, vertebra L-e for the centrum and transverse process and L-c for the neural spine).The global compactness (Cg) was measured in ten areas (orange outlines) to assess overall centrum compactness (see Supplementary Methods).b, common minke whale (Balaenoptera acutorostrata, LR M 523). c, Amazon river dolphin (Inia geoffrensis, ZMB_ Mam_41500).d, common dolphin (Delphinus sp. ZMB_Mam_697.59).e, dugong (Dugong dugon, ZMB_Mam_69340).f, manatee (Trichechus manatus, ZMB_ Mam_17377).(1) Centrum (anteroposterior mid-length, from its dorsal edge to its centre), external towards top; (2) Neural spine (dorsoventral mid-height), external towards right); (3) Transverse process (mediolateral mid-width), external towards top. Width of the virtual cross-sections were defined as representing 7.2% of the centrum dorsoventral height; this is the mean ratio between the centrum height of P.colossus and the width of the physical core drills.The core drills of the neural spine and transverse process do not reach the middle of the corresponding vertebral parts (but break surfaces indicate uniform structure throughout).See core drills' location in Supplementary Fig.S6c,d.Scalebars:a = 13.9 mm, b = 23.6 mm,c = 2.2 mm,d = 2.7 mm, e = 3.2 mm, f = 3.0 mm.

Extended Data Fig. 6 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 6 | Vertebral and costal morphology of Perucetus colossus (MUSM 3248, holotype) compared to that of other cetaceans. a, mean and maximumratioofvertebralcentrumanteroposteriorlength (CL) tomediolateral width (CW), computed with the thoracic and lumbar vertebrae of other cetaceans (Ba = Basilosaurinae; Dor = Dorudontinae; FM = fossil Mysticeti; FO = fossil Odontoceti;P = Pakicetidae+Ambulocetidae; Pa = Pachycetinae; Pr = Protocetidae;R = Remingtonocetidae).b, profile of vertebral centrum length in P.colossus (Pc) compared with the profiles of other basilosaurids (Aa = Antaecetus aithai; Bc = Basilosaurus cetoides; Bi = Basilosaurus isis; Cf = Chrysocetus fouadassii Cp = Cynthiacetus peruvianus; Da = Dorudon atrox; data of A. aithai and B. isis from Gingerich et al.15, fig.11).c, curvature (length/chord) of the best preserved rib, R-a (red horizontal line) compared to those of Cynthiacetus peruvianus (MNHN.F.PRU10).

Extended Data Fig. 5 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 5 | Location and stratigraphic position of Perucetus colossus MUSM 3248. a. Map showing the position of the Coastal Batholith and major trench-parallel structural highs along the coast of Peru (redrawn and modified after Travis et al.82. and Thornburg & Kulm83). b. Stratigraphic column of the Cenozoic succession exposed in the East Pisco Basin (redrawn and modified after Malinverno et al.84, Bianucci & Collareta85, and Bosio et al.86). c. Measured stratigraphic section of the type locality of P.colossus, indicating the stratigraphic height of the holotype MUSM 3248, the dated tephra layer, and the identified bioevents.

Extended Data Fig. 4 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 4 | Two vertebrae (a–l) and three posterior ribs (m; R-b,c,d) of Perucetus colossus (MUSM 3248, holotype). a,g. Anterior view.b,h. Posterior view.c,i. Dorsal view.d, j. Ventral view.e,k. Right lateral view.f,l Left lateral view. Tentative position along the vertebral column: a–f = L-j, eleventh lumbar;g–l = L-k,twelfth lumbar.a–l. Images generated from three-dimensional surface models. m. Photograph taken in the field. Scale bars = 50 cm.

Extended Data Fig. 3 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 3 | Three incomplete vertebrae of Perucetus colossus (MUSM 3248, holotype). a,g,m. Anterior view.b,h,n. Posterior view.c,i,o. Dorsal view.d,j,p. Ventral view.e,k,q. Right lateral view.f,l,r. Left lateral view.Tentative position along the vertebral column:a–f = L-e,fifth lumbar; g–l = L-f,sixth lumbar; m–r = L-g, seventh lumbar. All images were generated from three-dimensional surface models.Scale bar = 50 cm.

Extended Data Fig. 2 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 2 | Three vertebrae of Perucetus colossus (MUSM 3248, holotype). a,g,m. Anterior view. b,h,n., Posterior view.c,i,o. Dorsal view. d,j,p. Ventral view.e,k,q. Right lateral view.f,l,r. Left lateral view.Tentative position along the vertebral column:a–f = L-b,second lumbar; g–l = L-c,third lumbar; m–r = L-d, fourth lumbar. All images were generated from three-dimensional surface models.Scale bar = 50 cm.

Extended Data Fig. 1 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Extended Data Fig. 1 | Three vertebrae of Perucetus colossus (MUSM 3248 holotype). a,g,m. Anterior view. b,h,n. Posterior view. c,i,o. Dorsal view. d,j,p. Ventral view.e,k,q. Right lateral view.f,l,r. Left lateral view.Tentative position along the vertebral column:a–f = Th-a, penultimate thoracic; g–l = Th-b, last thoracic; m–r = L-a,first lumbar.All images were generated from three-dimensional surface models.Scale bar = 50 cm.

Fig. 3 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Fig. 3 | Bone histology of the P.colossus MUSM 3248 holotype. a, Half rib cross-section (in three parts), featuring very sparse porosity and multiple growth marks (arrows).b, Detail of the above, showing a highly convoluted, woven-parallel complex, devoid of remodelling (left, natural light; right, cross-polarized light). c, Broken peduncle of a lumbar vertebra in ventral view (note the fractures following growth lines), made of compact bone throughout. The small inset (vertebra parts in anterior view,not to scale) indicates the position of the illustrated break surface.Scale bars, 1 cm (a), 0.5 mm (b) and 5 cm (c).

Fig. 4 in A heavyweight early whale pushes the boundaries of vertebrate morphology

Fig. 4 | Scaling of the skeletal mass across amniotes' body mass range. Regressions based on extant terrestrial mammals (grey line) and extant cetaceans (blue line).The estimated values for extreme terrestrial amniotes, sirenians (either measured or estimated) as well as the results of our estimations for P.colossus are also shown. Each data point corresponds to a distinct species (specific mean for the largest extant cetaceans, B.musculus and Physeter macrocephalus), except for extant sirenians (represented by two species). The horizontal and vertical bars indicate the maximum and minimum estimates (some are too small to be visible).Scales are logarithmic. Human height, 1.8 m (scale for larger silhouettes); penny diameter, 2 cm.See the Supplementary Methods and Supplementary Data 2.