possess moderately deflected alary processes on the premasillae, which produces a thick but not truncate snout; the deflection is much greater and the profile of the snout thinner in G, ruizi than in either any other member of the subgenus or Gastrotheca as a whole. Second, all taxa have a narrow otic plate, except G. bu ona, a taxon not included in the molecular analysis. All members of the Gastrotheca monticola clade (G. monticola sp. B, lojana, orophylax, and plumbea; Fig. 4.11) have neopalatines that are widely separated below the sphenethmoid. Likewise, they have parasphenoids with cultriform processes that narrow abruptly ante. rior to the optic fenestra; they share this trait with both Edaphotheca and most Gastrotheca. The clade (except G. lojana) has neopalatines with alow irregular ventral ridge. The group composed of G. monticola sp. A, sp. B, and lojana has a squamosal with a wide dorsal head, but the zygomatic ramus is narrow in its lateral aspect. Within this lineage, the postorbital process of the max- illa of G. orophylax and G. plumber is high and broad in its lateral aspect and has a horizontal articulation with the zygomatic ramus of the squamosal; in addi- tion, the articulation of the pars facialis of the maxilla and the ventrolateral edge of the nasal is only about 50% complete Although Gastrotheca ovifera is basal to the subge nera Bdaphotheca, Duellmania, and Gastrotheca in the molecular tree (Fig, 4.1), in the combined analy: of morphological and molecular data, G. ovifera is re- solved as the sister taxon to a clade composed of G. argenteovirens, aurcomaculata, bufona, danni, espeletia, nnicefor, riobambae, ruizt, and trachyceps. All members of, this group (except G. niceford) have parasphenoids with cultriform processes that narrow gradually anterior to the optic fenestra, and all (except G. espeletia) have a partially closed carotid canal; it is open in G. espeletia ‘The group comprising G. argenteovirens, aureomaculata, bufona, espeletia, ruizi, and trachyceps lacks an evident postorbital process on the maxilla, and all of these taxa (except G, trachyceps) have a long spinous medial pala tine process on the premaxilla. G. argenteovirens, dunt, espeletia, and trachyceps have a prominent postorbital process on the maxilla that has a broad horizontal articulation with the zygomatic ramus of the squamo. sal. G. dunni, espeletia, and trachyceps have a neopala. tine with a low irregular ridge ventrally, whereas in G. aureomaculata, buona, espeletia, and ruizi, the neopala. tine bears a distinct ventral flange. In G, argenteovirens, cspeletia, and trachyceps, the nasals are synostosed with the sphenethmoid, whereas in G. aureomaculata and G. bufona, the anterior terminus of the cultriform process of the parasphenoid is acuminate, rather than blunt, Osteology 49 and the pattern of dermal sculpturing is heterogeneous across the skull table, ‘The frontoparietals of both taxa extend over the anterior epiotic eminence and part of, the posterior eminence. Both have a pattern of vermi form ossification in the sphenethmoid; the skull table of G. bufona is smooth, whereas that of G. aureomacu- laia has a semicoarse pattern of shallow pits and low rounded ridges. Subgenus Gastrotheca Based on analyses of combined osteological and molecular data, this subgenus is represented by three major clades: (1) Gastrotheca zeugocystis, 2) G. phalarasa, and (3) the more advanced allies of the former taxa, Unfortunately, osteological specimens of G. zeugocystis and G. phalarosa were unavailable; thus ostcological characterization of these taxa and their ancestral nodes is not possible. The placement of G. zeugocystis is the same in both the combined and molecular trees (Fig. 4.1). G. phalarosa is basal to all remaining taxa in the subgenus (except G. zeugocystis) in the combined tree, whereas it is nested amid more advanced taxa in the molecular tree. The remaining species of Gastrotheca, all of which have a semicoarse pit-and-ridge pattern of dermal sculpturing on the skull table and a tendency toward reduced ossification of the skull, are repre- sented by two major lineages, as opposed to four in the molecular results. One is composed of G. antoniiochoai and its sister group, herein termed the G, excubitor clade, and the other, the G, marsupiata clade. Osteologically, the Gastrotheca excubitor clade (exclu sive of G, antoniiochoai, for which, in this discussion, a skeleton was unavailable) represents a montage of plesiomorphic characters. The G. excubitor group [G. excubitor + G. ochoai] and [G. pachackacae + G. re- beccae|—has neopalatines that have a moderate medial separation ventral to the sphenethmoid and beara low irregular ventral ridge. The dentigerous processes of the vomers are located at the posterior level of the choanae, ‘The sister taxa G. excubitor and G. ochoai lack a supraorbital flange on the frontoparietal. Three distantly related species in this lineage exhibit multiple signs of independent reversals to a less osseous and ro- bust skull: G. andaguiensis, nebulanastes, and ochoai. In all three, the pars facialis of the maxilla has a short articu- lation with the nasal. G. nebulanastes and G. ochoai have small nasals that do not articulate medially; likewise, the nasals do not articulate with the frontoparietals, and the frontoparietals have an incomplete medial articulation. The zygomatic ramus of the squamosal is slender in G. andaguiensis and G. nebulanastes, and the ramus is incomplete in G. ochoai, Both G, andaquiensis50 Marsupial Frogs and G. ochoai lack an otic flange on the frontoparietal ‘The sister species G. abdita and G. andaquiensis have an evident postorbital process on the maxilla. ‘The Gastrotheca marsupiata clade is somewhat more tractable than its sister group in terms of its osteolog. ical features. It is characterized by lack of supraorbital and otic flanges on the frontoparietal and an open carotid canal, all of which represent diminished ossifi cation of the cranium, Two sister-groups compose thi clade in the combined tree (Fig, 4.11). The more nar: rowly defined G. marsupiata~G. gracilis lineage is less well os the G. psychrophila G. piperata lineage. The G. marsupiata-G. gracilis clade has a high truncate snout in lateral profile, owing to the minimal posterior deflection of the alary processes of the premaxillae, and the postorbital process on the maxilla is either absent or scarcely evident. Addi tionally, these frogs lack a supraorbital flange on the frontoparietal; the carotid canal is open and, in most of, them, the frontoparietal does not cover the epiotic em. inences, In contrast, members of the G. psychrophila G. piperara clade have a conspicuous supraorbital flange, and most have a partially closed carotid canal and a frontoparietal that covers the anterior epiotic eminence. In addition, the alary processes of the pre maxillae are deflected posteriorly, such that the snout, isnot markedly truncate in lateral profile. One species-pair defined by osteological featu evident in the Gastrotheca marsupiata clade: G. griswoldi and G. pacchamama shate several features associated with reduction of the osseous cranium. ‘The skulls are smooth, not exostosed. The nasals do not articulate with the frontoparietals, and the latter lack a complete medial articulation. The head of the squamosal lacks a dorsal crest, and the anterior ramus of the pterygoid lacks a dorsal process. Within the Gastrotheca psychrophila-G. piperata clade, both G. angustifions and G. peruana possess a heterogeneous pattern of dermal sculpturing and a smooth non-exotosed sphenethmoid. The nasals do not articulate with the frontoparietals, and the latter Jack a complete medial articulation. The quadratojugal is short. G. lateonota and G. piperata are alike in having, hyperossified nasals; the bones are fused to one an. other medially, and the sphenethmoid posteriorly. ed than its sister clad ‘This analysis of evolutionary trends in cranial osteo! ogy in Gastrotheca has, on the one hand, been a journey of frustration, and, on the other, a promising exercise in the informative power of derstanding of a particular group of anurans, The frus mbined data on our un tration emanates from the many taxa for which little or no morphological information is available, from our meager knowledge of variation between sexes and within and between populations, and from our ex ceedingly limited knowledge of ontogenetic develop- ment of the skeleton among members of Gastrotheca. Further, we have no idea of the functional significance of the characters and character complexes that have been described here and elsewhere. All of these factors speak to the need for, and satisfaction of, producing, morphological studies to complement and provide evolutionary meaning to the relationships revealed by molecular analyses. Yet the relative ease with which genomic data can be retrieved and analyzed to produce trees provides unparalleled opportunities for morphologists to ply their trade. This has been one of the most rewarding, aspects of the Gastrotheca project. The concordance of the molecular and osteological data in the deep phylogeny of the marsupial frogs is both informative and significant, Cranial characters can be used to unite taxa, Among the more advanced taxa, the amount of homoplasy thwarts characterization, but it also doc ments the enormous evolutionary plasticity of crani development in anurans. LITERATURE CITED Alberch, B, and E. A. Gale, 1983, Size dependence during the evelopment of the amphibian foot; colehicine-induced Gigita loss and reduction. 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