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Figure 5 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 5. Lateral and dorsal views of heads. A, B, Centrolene buckleyi, male, QCAZ 22388; C, D, Cochranella posadae, male, QCAZ 26023; E, Cochranella wileyi sp. nov., female, QCAZ 26028; F, Cochranella wileyi sp. nov., male, QCAZ 26029. Scale bar = 2 mm.

Figure 6 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 6. Ventral view of hands and feet. A, B, Centrolene buckleyi, male, QCAZ 22388; C, D, Cochranella posadae, males, QCAZ 25090 and 26023, respectively; E, F, Cochranella wileyi sp. nov., female, QCAZ 26028. Scale bar = 2 mm.

Figure 11 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 11. Dorsal (top) and ventral (bottom) views of the holotype of Cochranella wileyi sp. nov., adult female, SVL = 27.1 mm, QCAZ 26028.

Figure 1. Phylogenetic relationships among genera and species groups within Centrolenidae. A in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 1. Phylogenetic relationships among genera and species groups within Centrolenidae. A, tree topology suggested by Ruiz-Carranza & Lynch (1991a, b, c, 1996, 1998) and modified by Bolívar et al. (1999), Señaris (2001) and Duellman & Señaris (2003). B, single, most-parsimonious tree of the phylogenetic relationships of Centrolenidae (tree length = 13, CI = 0.923, RI = 0.9474, RC = 0.8745). Numbers refer to the following characters: (1) tibiale and fibulare, 0 = not fused, 1 = partially or completely fused; *(2) T-shaped terminal phalanges, 0 = absent, 1 = present; (3) dilated medial process on Metacarpal III, 0 = absent, 1 = present; (4) eggs deposition site, 0 = deposited in water, 1 = not deposited in water, 2 = deposited on underside of leaves; (5) shape of liver, 0 = liver lobed, 1 = liver bulbous; (6) humeral spine in males, 0 = absent, 1 = present; (7) relative size of disc of Finger III, 0 = disc small (<80% of eye diameter), 1 = disc large (> 80% of eye diameter); (8) coloration of hepatic peritoneum, 0 = clear, 1 = white; (9) coloration of peritoneum covering urinary blad- der, 0 = clear, 1 = white; (10) red heart visible in ventral view, 0 = heart not visible, 1 = red heart visible; (11) coloration of parietal peritoneum, 0 = white, 1 = clear. Character 12 (venter-to-venter fight behaviour) was hypothesized to be a synapomorphy shared by Centrolene and Cochranella (Bolívar et al., 1999); however, the distribution of this behaviour has been reported in only nine species (Guayasamin & Barrio-Amorós, 2005) and we did not include in B. Numbers next to tick marks represent bootstrap support values. Grey boxes denote characters that appear more than once in the tree. *T-shaped terminal phalanges are also present in Allophryne ruthveni.

Figure 4 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 4. Photographs of glass frogs. A, B, Centrolene buckleyi from Yanayacu Biological Station, male, dorsolateral and ventral views, QCAZ 26032 (WCF); C, Centrolene buckleyi from Carchi, male, dorsolateral view, MECN 1246 (MRB); D, E, Cochranella posadae, male, dorsolateral and ventral views, QCAZ 25090 (WCF); F, Centrolene bacatum, dorsolateral view, male, QCAZ 26056 (MRB); G, H, Cochranella wileyi sp. nov., males, dorsolateral and ventral views, QCAZ 26029 and 27441, respectively (MRB); I, Cochranella griffithsi, dorsolateral view, QCAZ 29525 (JMG).

Figure 2 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 2. Terminology (modified from Savage & Heyer, 1967) used for webbing formula in hands and feet. Roman numerals represent fingers or toes; Arabic numerals represent the number of phalanges completely or partially free of webbing. We use 0– to indicate that the web reaches the distal margin of the disc; 0 indicates that the web reaches the middle of the disc; 0+ indicates that the web reaches the proximal margin of the disc; 1– indicates that the web reaches the distal margin of the intercalary cartilage; 1 indicates that the web reaches the middle of the intercalary cartilage; 1+ indicates that the web reaches the proximal margin of the intercalary cartilage; 2– indicates that the web reaches the distal margin of the distal subarticular tubercle; 2 indicates that the web reaches the middle of the distal subarticular tubercle; 2+ indicates that the web reaches the proximal margin of the distal subarticular tubercle; the notation for 3 and 4 follow the same pattern described for 2. When, for example, the webbing reaches a midpoint between the intercalary cartilage and the distal subarticular tubercle in Finger IV, and the proximal margin of the disc in Finger V, the appropriate notation between these two fingers would be IV 11/2−0+ V. Scale bar = 2 mm.

Figure 14 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 14. Ventral view of two individuals of Hyalinobatrachium crurifasciatum showing intraspecific variation of the pericardium. A, white pericardium, MHNLS 16477; B, mostly clear pericardium, but see upper right corner of the heart, MHNLS 16475.

Figure 10 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 10. Ventral view of hand (left) and foot (right) of Cochranella posadae, male, QCAZ 25090. Note white pigments covering surface of arm, tarsus and heel. Additional white pigmentation is visible on external border of Finger IV and Toe V. Hand length = 11.0 mm; foot length = 15.6 mm.

Figure 7. A in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 7. A, power spectrum; B, sonagram; and C, oscillogram of the advertisement call of Centrolene buckleyi (QCAZ 26032). The call shown here consists of four notes, each note with two distinct pulses. The power spectrum was measured along the duration of the first note.

Figure 12 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 12. Kidneys of Cochranella wileyi sp. nov., male, QCAZ 26030, ventral view. Note that kidneys are covered with a white peritoneum and small, unpigmented spots. Length of kidneys = 4.7 mm.

Figure 15 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 15. Ventral view of hand of Cochranella wileyi sp. nov., QCAZ 26029. Arrow indicates medial process of Metacarpal III, a character present in all centrolenid frogs.

Figure 13 in Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

Figure 13. Egg clutches of Cochranella wileyi sp. nov. Note that egg clutches are hanging on the tip of leaves. Each egg mass contains between 19 and 28 eggs.

Figure 15 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 15. Comparison of the changes between the larval and adult body plans during larval development. The horizontal axis represents developmental stages (Gosner, 1960). The curves plotted depict structural modifications, and the grey area represents metamorphic events that take place during the metamorphic climax (stages 42–46) for most anurans. The early occurrence of metamorphic events (predisplacement) is observed in the ceratophryine frogs, especially in Lepidobatrachus spp., that have precocious metamorphosis. Delayed metamorphic events take place in the development of Pseudis platensis, a species in which some morphological changes that imply the end of metamorphosis for most anurans have yet to finish.

Figure 13 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 13. Variation in structural changes in some external features. A, dorsal view of Lepidobatrachus llanensis at the end of metamorphosis. A small tail stub is present. B, lateral view of the same specimen in (A) showing the angle of the mouth far beyond the posterior margin of the eye. C, dorsal view of Chacophrys pierottii during metamorphosis. The tail has started to reduce. D, lateral view of the same specimen in (C). The disappearance of the caudal fin is advanced, and the angle of the mouth reaches the posterior margin of the eye. E and F, ventral and lateral views of a Pseudis platensis tadpole at the beginning of metamorphosis. The oral disc and keratinized buccal structures are still present, and the forelimbs have emerged, but the anal tube remains well developed. G and H, ventral and lateral views of a P. platensis tadpole during metamorphosis. Larval mouthparts have disappeared; tail regression has started with the reduction of the fins, whereas the disappearance of the anal tube is delayed. I, dorsal view of P. platensis at an advanced metamorphic stage. The tail is conserved and has reduced caudal fins. J, ventral view of the same specimen showing the absence of the anal tube, and features of the mouth that are similar to those present in most anurans at the end of metamorphosis. K, detail in lateral view of the position of the angle of the mouth posterior to the eye.

Figure 12 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 12. Hematoxylin–eosin cross sections (6-Mm thick) at the diaphyseal level of metatarsal IV in postmetamorphic specimens. Black arrowheads indicate lines of arrested growth (LAGs). A, Lepidobatrachus llanensis, five LAGs male [data for two adult males; 5 ± 1 (SVL 74.5 mm) and 6 ± 1 (SVL 74.1 mm)]. B, Lepidobatrachus laevis, six LAGs female [data for two adult females; 6 ± 1 (SVL 144 mm) and 7 ± 1 (SVL 111 mm)]. C, Chacophrys pierottii, four LAGs male [data for two adult males; 6 ± 1 (SVL 49.9 mm) and 4 ± 1 (SVL 45.2 mm)]. D, Ceratophrys cranwelli, 11 LAGs male [data for three adult males; 14 ± 1 (SVL 84.6 mm), 11 ± 1 (SVL 81 mm), and 13 ± 1 (SVL 74.3 mm)]. E, Pseudis platensis, two LAGs female [data for two adults (female and male); 2 ± 1 (SVL 46 mm) and 3 ± 1 (SVL 44.6 mm)]. F, Telmatobius atacamensis, juvenile specimen, two LAGs [data for an adult male; 5 ± 1 (SVL 45.4 mm) and for the juvenile 2 ± 1 (SVL 45 mm)]. Abbreviations: mc; marrow cavity. Scale bars: 0.05 mm.

Figure 11 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 11. Mean, maximum, and minimum values of snout–vent lengths (SVLs) in advanced tadpoles (stages 38–41), during metamorphosis (stages 42–46), and in adults. Values of SVL are given in mm. Light-grey areas approximately represent the extension of larval development (LD) in months, and dark-grey areas refer to postmetamorphic growth (PG) in lines of arrested growth (LAGs). Lepidobatrachus llanensis: SVL in tadpoles (N = 15, SVL = 36.7 ± 3.4 mm); SVL of metamorphic specimens (N = 26, SVL = 36.3 ± 5.8 mm); SVL of adults (N = 16, SVL = 76.5 ± 7.5 mm). Larval development takes place over a period of 2 weeks. Postmetamorphic growth estimated in LAGs = 6 years. Lepidobatrachus laevis: SVL in tadpoles (N = 12, SVL = 46.6 ± 2.8 mm); SVL of metamorphic specimens (N = 26, SVL = 49.9 ± 7.8 mm); SVL of adults (N = 13, SVL = 99.2 ± 22.1 mm). Larval development takes over a period of 2 weeks. Postmetamorphic growth estimated in LAGs = 7 years. Chacophrys pierottii: SVL in tadpoles (N = 10, SVL = 46.3 ± 3.9 mm); SVL of metamorphic specimens (N = 28, SVL = 36.3 ± 2.6 mm); SVL of adults (N = 15, SVL = 47.8 ± 2.7 mm). Larval development takes place over a period of 2 weeks. Postmetamorphic growth estimated in LAGs = 6 years. Ceratophrys cranwelli: SVL in tadpoles (N = 8, SVL = 26.3 ± 3.8 mm); SVL of metamorphic specimens (N = 18, SVL = 27.5 ± 4.4 mm); SVL of adults (N = 10, SVL = 91.9 ± 12.7 mm). Larval development takes place over a period of 3 weeks. Postmetamorphic growth estimated in LAGs = 14 years. Pseudis platensis: SVL in tadpoles (N = 9, SVL = 46.3 ± 3.6 mm); SVL of metamorphic specimens (N = 8, SVL = 37.4 ± 2.7 mm); SVL of adults (N = 7, SVL = 45.3 ± 5.0 mm). Larval development takes place over a period of 6 months. Postmetamorphic growth estimated in LAGs = 3 years. Telmatobius atacamensis: SVL in tadpoles (N = 11, SVL = 32.9 ± 3.4 mm); SVL of metamorphic specimens (N = 30, SVL = 31.0 ± 1.8 mm); SVL of adults (N = 2, SVL = 45.2 ± 1.0 mm). Larval development takes place over a period of 8 months. Postmetamorphic growth estimated in LAGs = 5 years.

Figure 14 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 14. Presence and absence of m. suspensoriohyoideus among selected anuran tadpoles, following the criteria proposed by Haas (2003) at larval stage 37. A, Pseudis platensis. The muscle is absent. B–F, the muscle is present. B, Telmatobius atacamensis. C, Lepidobatrachus laevis. D, Lepidobatrachus llanensis. E, Ceratophrys cranwelli. F, Chacophrys pierottii. The absence of m. suspensoriohyoideus was reported for Ceratophrys ornata, L. laevis (Ruibal & Thomas, 1988; Haas, 2003), and C. cranwelli (Vera Candioti, 2005), but Palavecino (1999) described it in C. ornata and C. cranwelli. Haas (2003) proposed that the absence of m. suspensoriohyoideus is a synapomorphy for the Ceratophrys and Lepidobatrachus clade, and Frost et al. (2006) extended the synapomorphy for Ceratophryini. Our data demonstrate the presence of the m. suspensoriohyoideous in tadpoles of the three genera of Ceratophryinae. Abbreviations: oh, m. orbitohyoideus; sh, m. suspensoriohyoideus. Scale bars: 1 mm.

Figure 9 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 9. Palmar and plantar views of the autopodia in anurans at the end of metamorphosis, where subarticular turbercles are already well defined. A, Bombina variegata (foot): subarticular tubercles are absent and metatarsal tubercles are poorly developed. B, Bombina variegata (hand): subarticular tubercles are absent. C, Odontophrynus americanus (foot): subarticular tubercles are evident, and the inner metatarsal is not yet keratinized. D, Odontophrynus americanus (hand): subarticular tubercles are strongly developed. E, Pseudis platensis (foot): tubercles are absent and the inner metatarsal tubercle is defined. F, Pseudis platensis (hand): small subarticular tubercles are evident. G, Telmatobius atacamensis (foot): small subarticular tubercles and the inner metatarsal tubercle are present. H, Telmatobius atacamensis (hand): subarticular tubercles are better defined than in the foot. I, Lepidobatrachus llanensis (foot): subarticular tubercles are absent. The well-developed inner metatarsal tubercle and toe tips are keratinized. J, Lepidobatrachus llanensis (hand): subarticular tubercles are absent. K, Lepidobatrachus laevis (foot): subarticular tubercles are absent, and the inner metatarsal tubercle presents incipient keratinization. L, Lepidobatrachus laevis (hand): subarticular tubercles are absent. M, Ceratophrys cranwellii (foot): small subarticular tubercles are defined, and the inner metatarsal tubercles are poorly keratinized. N, Ceratophrys cranwelli (hand): small subarticular tubercles are present. O, Chacophrys pierottii (foot): subarticular tubercles are not prominent. Inner metatarsal tubercle has keratinization. P, Chacophrys pierottii (hand): subarticular tubercles are well developed. The pattern of distribution of subarticular tubercles is considered as diagnostic in most species, and development of these structures takes place at larval stages 38–40 (Gosner, 1960). Subarticular tubercles in manus and pes are also absent or poorly developed in Xenopus, Hymenochirus, Pipa, Ascaphus, Leiopelma, Alytes, Discoglossus, Madecassophryne, Stumpffia, Rhinoderma, Brachycephalus, Truebella, and Ansonia (Guibé, 1978; Graybeal & Cannatella, 1995; Matsui, Nabhitabhata & Panha, 1998; Pombal & Gasparini, 2006, among others). Scale bar: 0.2 mm.

Figure 10 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 10. Total length (TL) versus snout–vent length (SVL) plotted for larval specimens of six species between stages 39 and 41. Ceratophrys cranwelli tadpoles (N = 8, TL = 61.8 ± 8.2 mm, SVL = 26.3 ± 3.8 mm) are the smallest. Pseudis platensis larvae (N = 9, TL = 129.5 ± 14.4 mm, SVL = 46.3 ± 3.6 mm) are the largest. Lepidobatrachus laevis (N = 12, TL = 98.3 ± 7.2 mm, SVL = 46.6 ± 2.8 mm) and Chacophrys pierottii (N = 10, TL = 106.5 ± 8.0 mm, SVL = 46.3 ± 3.9 mm) share with P. platensis similar values of SVL. Lepidobatrachus llanensis (N = 15, TL = 83.7 ± 7.5 mm, SVL = 36.7 ± 3.4 mm) and Telmatobius atacamensis (N = 11, TL = 84.6 ± 4.8 mm, SVL = 32.9 ± 3.4 mm) are similar in total length, but L. llanensis tadpoles have greater body sizes.

Figure 1 in Morphological evolution in Ceratophryinae frogs (Anura, Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Figure 1. The hypothetic relationships among ten anuran taxa that resulted from the analyses of 102 morphological characters. A, the only tree that was obtained from the analysis of larval and adult characters. B, strict consensus of relationships obtained from the analysis of 61 larval characters. C, strict consensus of relationships obtained from the analysis of 41 adult characters.