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ACANTHOSOMATIDAE Signoret Historical: This predominantly Southern Hemisphere taxon includes three subfamilies: Acanthosomatinae, Blaudinae with two tribes Blaudini and Lanopini, and Ditomotarsinae, also with two tribes, Ditomotarsini and Laccophorellini (Kumar, 1974). Froeschner (1999) emendedthe spelling of Blaudusinae Kumar to Blaudinae and Blaudini because the higher-taxon name was based on Blaudus St��l; we follow Froeschner��s usage. In the cladistic analysis of Gapud (1991) the Acanthosomatidae is related to the Dinidoridae plus Tessaratomidae and Scutelleridae, coming out in a relatively basal position on the cladogram (Fig. 1f). Fischer (1994a,b), in a phylogenetic analysisof the family, stressed the monophyly of the Acanthosomatidae based on three non-homoplastic characters and one homoplastic character: (i) presence of Pendergrast��s organ (abdominal disc organ), (ii) segment VIII in males visible (not concealed by segment VII), (iii) females with a special organ for symbiont transmission, and (iv) openings of anterior abdominal scent glands shifted laterad, a feature shared with the Scutelleridae. Fischer (2006) described the biological context and evolution of Pendergrast��s organ in the Acanthosomatidae, presenting a surveyof these organsin more than 100 acanthosamatid species. Analytical result: Our morphological and total evidence analyses (Figs 42���44 and 51���55, respectively) always resolve the Acanthosomatidae as monophyletic, a theory concordant with most prior work. Our taxon sample for DNA sequences is biased toward the Australian fauna and the subfamilies Acanthosomatinae and Blaudinae, although we did sequence Elasmostethus Fieber from the Northern Hemisphere, a member of the Acanthosomatinae. The position of the group within the Pentatomoidea is variable, depending on the data set being analysed. The morphological analyses treat the group as relatively basal (Figs 42���44), whereas the molecular data always treat the group as closely associated with the Pentatomidae (Figs 45 and 46), although sometimes with a small number of other taxa involved (Fig. 45). The 52-taxon total evidence analyses (Figs 51 and 52) place the Acanthosomatidae + Lestoniidae as the sister group of the Pentatomidae, in the case of 1: 1 cost ratio also including Thaumastella. The result of the 92-taxon analysis under a 1: 1 cost ratio (Fig. 53) is similar to molecular and 52-taxon analyses, the 1: 2 cost ratio moves the Acanthosomatidae to a more basal position in the cladogram (Fig. 54), and the 2: 2 costratioincludesthe Dinidoridae + Tessaratomidae as part of the Acanthosomatidae + Pentatomidae complex. Morphological characters supporting the monophyly of the Acanthosomatidae in both the morphological and 92-taxon total evidence analyses are the membranous abdominal tergite VIII in males (411) and the triangulin absent with a smooth intergonocoxal membrane between gonapophyses 8 (531). The 92-taxon total evidence analyses offer additional support from the obsolete claval commissure (172) and the claws with bristles (311)., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 969-970, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Kumar, R., 1974. A revision of world Acanthosomatidae (Heteroptera: Pentatomoidea): Keys to and descriptions of subfamilies, tribes and genera with designation of types. Aust. J. Zool., Suppl. Ser. N. 34, 1 - 60.","Froeschner, R. C., 1999. Revision of the South American genus Hellica Stal (Heteroptera: Acanthosomatidae). J. NY Entomol. Soc. 107, 164 - 170.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Fischer, C., 1994 a. Phylogenetisch-systematische analyse der Acanthosomatidae (Heteroptera, Pentatomoidea). Verh. Deut. Zool. Ges. 87, 220.","Fischer, C., 2006. The biological context and evolution of Pendergrasts ̕ organs of Acanthosomatidae (Heteroptera, Pentatomoidea). In: Rabitsch, W. (Ed.), Hug the bug-For love of true bugs. Festschrift zum 70 Geburtstag von Ernst Heiss., Denisia 19, 1041 - 1054."]}

PENTATOMIDAE Leach sensu lato (including Aphylinae and Cyrtocorinae) Historical: Rolston and McDonald (1979) recognized five subfamilies in Pentatomidae from the Western Hemisphere (Asopinae, Discocephalinae, Edessinae, Pentatominae, and Podopinae). Schuh and Slater (1995) included two additional subfamilies from the Eastern Hemisphere (Phyllocephalinae, and Serbaninae) and included the Cyrtocoridae as a subfamily, giving a total of eight subfamilies. Rolston (1981) proposed Ochlerini as a new tribe in Discocephalinae. Ahmad and Kamaluddin (1988, 1990) and Kamaluddin and Ahmad (1988) established a tribal classification of the Phyllocephalinae recognizing four tribes: Cressonini, Megarrhamphini, Phyllocephalini, and Tetrodini. In the Pentatominae, Hassan and Kitching (1993) provided a cladistic analysis of some of the tribes but did not propose a revised formal classification. For the Podopinae, Davidov�� -Vilimov�� and ��tys (1994) recognized five tribes: Brachycerocorini, Deroploini, Graphosomatini, Podopini, and Tarisini; Schuh and Slater (1995) recognized eight tribes (Aeptini, Diemeniini, Halyini, Lestonocorini, Mecideini, Myrocheini, Pentatomini, and Sciocorini). Recently, Rider (2000) proposed a new subfamily, Stirotarsinae, for the monotypic genus Stirotarsus Bergroth, based on the unique antennal, rostral, and tarsal characters, along with the relatively rare ostiolar, tibial, and spiracular characters. Cassis and Gross (2002) summarized the suprageneric classifications of Pentatomidae of some earlier authors who had a broad concept of the family, incorporating the dinidorids, plataspids, tessaratomids, and scutellerids (e.g. Kirkaldy, 1909; Miller, 1956; China and Miller, 1959). Most recently, Rider (2006) recognized ten subfamilies within Pentatomidae [Aphylinae, Asopinae, Cyrtocorinae, Discocephalinae, Edessinae, Pentatominae, Phyllocephalinae, Podopinae, Serbaninae (see discussion under Phloeidae), and Stirotarsinae], the Pentatominae comprising 42 nominal tribes. Gapud (1991) considered the Pentatomidae to be probably the most ������advanced������ family in the Pentatomoidea, supported by six apomorphies: the dorsally membranous eighth abdominal segment in males, the rigid phallotheca, the vesica without a conjunctival sheath (an extremely variable character within the Pentatomoidea, as are the majority of male genitalic characters), the fixed position of the ejaculatory reservoir on the phallotheca, triangulin present, and the completely fused 2nd valvifers (shared with Scutelleridae). Aphylinae Bergroth: This exclusively Australian group, known from two genera and three species, was first reported in the description of Aphylum syntheticum Bergroth (1906). The author proposed a new subfamily, considering A. syntheticum to be an isolated taxon combining characters of the pentatomoid family-groups Scutellerinae, Graphosomatinae, Plataspinae, and Pent- atominae. Schouteden (1906b) described the new species A. bergrothi and was inclined to place Aphylinae in the neighborhood of the Scutellerinae. Aphylum was monographed by Schouteden (1906a) and subsequently raised to family rank by Reuter (1912). China (1955), in establishing his new subfamily Lestoniinae under the Plataspididae, compared it to Aphylum; China (1963) properly corrected his statement of 1955 concerning the absence of trichobothria in both subfamilies. McDonald (1970) discussed the morphology and relationships of Aphylum. Gross (1975), considering it to be closely related to the pentatomid genera Tarisa Amyot & Serville and Kumbutha Distant, returned the taxon to subfamily status. Schuh and Slater (1995) and Cassis and Gross (2002) accepted family rank for the group. ��tys and Davidov�� -Vilimov��(2001) described Neoaphylum to include the new species N. grossi. Rider (2006) treated the taxon as a pentatomid subfamily. Cyrtocorinae Distant: This exclusively Neotropical taxon was recently revised by Packauskas and Schaefer (1998); it includes four genera and 11 species. Besides the record of Cyrtocoris trigonus (Germar) from California (Banks, 1910; Horv��th, 1916; Brailovsky et al., 1988), Packauskas and Schaefer (1998) agreed with Henry and Froeschner (1988) that the lack of any subsequent discovery of Cyrtocoris White in the United States makes Bank��s record suspect. Packauskas and Schaefer (1998) stated that Kormilev (1955) appears to have been the first person to present evidence for raising Cyrtocorinae to family rank as distinct from the Pentatomidae (based on features of the fore- and hindwing venation, the position of the second abdominal spiracle in the membrane, and the placement of the abdominal trichobothria), allying Cyrtocoridae with Cydnidae; but, at the same time, they argued that placement of the anterior trichobothria lateral to the spiracles, also found in the pentatomid subfamily Discocephalinae, represents convergence. The differences in the venation of fore- and hindwings may be an issue of degree; the second abdominal spiracle, lying in the membranous part of the segment, needs a more complete survey, as this situation varies in different groups of pentatomids. Gapud (1991) separated Cyrtocorinae (as a pentatomid subfamily) from the rest of Pentatomidae by the absence of a triangulin, 2nd valvifers with a distinct median fusion line, and male phallotheca relatively flexible. Packauskas and Schaefer (1998) considered the presence of a triangulin, 2nd valvifers completely fused, and a rigid phallotheca as apomorphies of Pentatomidae minus Cyrtocoridae. Gapud (1991) placed the Cyrtocorinae + Pentatomidae sensu stricto as the most apical taxa, ������strongly separated from the rest of Pentatomoidea������ by the loss of first valvulae, the absence of the gonangulum, the invagination and dilation on the spermathecal duct, the retention of membranous flaps of the 2nd valvulae, and the presence of an antero-posterior pair of basal sclerites on the spermathecal base. Analytical result: This, the largest family-group within the Pentatomoidea, is resolved as monophyletic in every analysis���except the 16S, 18S, and CO1 partitions���testifying to the strength of character support for it. Morphological characters that consistently support the recognition of a broadly conceived Pentatomidae include: the loss of gonapophyses 8 and the first rami (452), gonapophyses 9 reduced and fused to gonocoxites 9 (492), gonangulum absent (502), and the ductus receptaculi dilated and invaginated, forming three distinct walls (511). The overall sample of taxa and characters for the Pentatomidae in this study is too small to provide a robust scheme at the subfamily and tribal level. We can comment, however, on the relationships of the Pentatomidae sensu stricto with the familygroup taxa Aphylinae and Cyrtocorinae. Discussions of the systematic position and rank of the Aphylinae and Crytocorinae have occupied considerable space in the literature. Many of those discussions have focused on differences instead of similarities. We cannot adduce information from sequence data for these two taxa. We can point out, however, that our morphological analysis offers strong character support for the grouping (Cyrtocorinae (Aphylinae + Pentatomidae sensu stricto)). Thus, it would seem that discussions concerning whether or not Cyrtocorinae and Aphylinae should be recognized at the family level, or as part of the Pentatomidae, simply amount to preference regarding degree of difference, rather than a substantive interpretation of relationships. We have chosen to treat both taxa at subfamily rank in recognition of their many shared similarities with the Pentatomidae sensu stricto. Sister-group relationships of the Pentatomidae sensu lato at the next higher level are not as clear-cut. Nonetheless, the totality of the evidence seems to point towards a sister-group relationship with the Acanthosomatidae + Lestoniidae, as suggested in Figs 45, 46, and 51���53., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 970-971, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Rolston, L. H., McDonald, F. J. D., 1979. Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). J. NY Entomol. Soc. 87, 189 - 207.","Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY.","Rolston, L. H., 1981. Ochlerini, a new tribe in Discocephalinae (Hemiptera: Pentatomidae). J. NY Entomol. Soc. 89, 40 - 42.","Kamaluddin, S., Ahmad, I., 1988. A revision of the tribe Phyllocephalini (Hemiptera: Pentatomidae: Phyllocephalinae) from Indo- Pakistan subcontinent with description for five new species. Orient. Insects 22, 185 - 240.","Ahmad, I., Kamaluddin, S., 1990. A new tribe for Phyllocephalinae genera Gellia Stal and Tetroda Amyot & Serville (Hemiptera: Pentatomidae) and their revision. Annot. Zool. Bot. Bratislava 195, 1 - 20.","Hassan, S. A., Kitching, I. J., 1993. A cladistic analysis of the tribes of Pentatomidae (Heteroptera). Jpn. J. Entomol. 61, 651 - 669.","Rider, D. A., 2000. Stirotarsinae, new subfamily for Stirotarsus abnormis Bergroth (Heteroptera: Pentatomidae). Ann. Entomol. Soc. Am. 93, 802 - 806.","Cassis, G., Gross, G. F., 2002. Hemiptera- Heteroptera (Pentatomomorpha). In: Houston, W. W. K., Wells, A. (Eds.), Zoological Catalog of Australia. CSIRO Publishing, B. Melbourne, Australia, Vol. 27.3 B, xiv + 737 pp.","Kirkaldy, G. W., 1909. Catalogue of Hemiptera (Heteroptera) with Biological and Anatomical References, List of Food Plants and Parasites, etc. vol. 1 Cimicidae. F. L. Dames, Berlin.","Miller, N. C. E., 1956. The Biology of Heteroptera. Leonard Hill Ltd., London.","China, W. E., Miller, N. C. E., 1959. Check-list and keys to the families and subfamilies of the Hemiptera-Heteroptera. Bull. Br. Mus. (Nat. Hist.), Entomol. 8, 1 - 45.","Rider, D. A., 2006. Pentatomoidea Home Page. North Dakota State University. http: // www. ndsu. nodak. edu / ndsu / rider / Pentatomoidea / [accessed on 21 July 2006].","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Bergroth, E., 1906. Aphylinae und Hyocephalinae, zwei neue Hemipteren-Subfamilien. Zool. Anz. 29, 644 - 649.","Schouteden, H., 1906 b. Une nouvelle espe`ce du genre Aphylum. Ann. Soc. Entomol. Belg. 50, 215 - 216.","Schouteden, H., 1906 a. Heteroptera, Fam. Pentatomidae, Subfam. Aphylinae. In Wytsman, M. P. (Ed.), Genera Insectorum, Fasc. 47, Brussels.","Reuter, O. M., 1912. Bemerkungen uber mein neues Heteropterensystem. Ofv. Finska Vet. - Soc. Forh. 54, 1 - 54.","China, W. E., 1955. A new genus and species representing a new subfamily of Plataspidae, with notes on the Aphylidae (Hemiptera, Heteroptera). Ann. Mag. Nat. Hist. (12) 8, 204 - 210.","China, W. E., 1963. Lestonia haustorifera China (Hemiptera: Lestoniidae) - a correction. J. Entomol. Soc. Queensland 2, 67 - 68.","McDonald, F. J. D., 1970. The morphology of Lestonia haustorifera China (Het. Lestoniidae). J. Nat. Hist. 4, 413 - 417.","Gross, G. F., 1975. Plant Feeding and Other Bugs (Hemiptera) of South Australia-Heteroptera pt. I. Handbook Flora Fauna South Australia. A. B. James, Adelaide.","Packauskas, R., Schaefer, C. W., 1998. Revision of the Crytocoridae (Hemiptera: Pentatomoidea). Ann. Entomol. Soc. Am. 91, 363 - 386.","Banks, N., 1910. Catalogue of the Nearctic Hemiptera Heteroptera. American Entomological Society, Philadelphia, PA.","Brailovsky, H., Cervantes, L., Mayorga, C., 1988. Hemiptera- Heteroptera de Me'xico XL: La Familia Cyrtocoridae Distant en la estacion de biologia tropical '' Los Tuxtlas' ' (Pentatomoidea). An. Inst. Biol. Univ. Auton. Mex., ser. zool., 58, 537 - 560.","Henry, T. J., Froeschner, R. C. (Eds.)., 1988. Catalog of Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, New York.","Kormilev, N. A., 1955. La subfamilia Cyrtocorinae Distant en la Argentina (Hemiptera Pentatomoidae) [sic]. Rev. Ecuat. Entomol. Parasitol. 2, 321 - 334."]}

LESTONIIDAE China Historical: China (1955) described the lestoniids as a new subfamily of Plataspididae based on characters of the hind wing venation and the two-segmented tarsi. China and Miller (1959) raised the group to family status. China (1963), McDonald (1970), and Schaefer (1993a) retained the familial ranking. Gapud (1991) considered Lestoniidae and Plataspididae as sister groups, with five homoplasious characters supporting the relationship: well-developed triangulin (an erroneous interpretation; see explanation of the characters), two-segmented tarsi, obsolete frena, enlarged scutellum that covers the abdomen, and contiguous ninth paratergites. The Plataspididae is unique in having almost the entire pleural region covered by the evaporative area, whereas the Lestoniidae has an extremely reduced evaporative area. Fischer (2000, 2006) considered the disc-like organs of the Lestoniidae to be homologous with the Pendergrast̕s organs of the Acanthosomatidae and associated the two taxa on that basis. Additional characters suggesting a common stem-species for Acanthosomatidae and Lestoniidae include the structure of the male genital segments, the arrangement of the abdominal scent-gland openings, and the number of tarsomeres. McDonald (1969, 1970), Schaefer (1993a), and other authors erroneously considered the spermathecal flanges to be absent in Lestoniidae (see explanation of characters); Fischer (2000) corrected this misinterpretation and illustrated the spermatheca of Lestonia haustorifera China as possessing a proximal flange. Analytical result: The placement of the Lestoniidae has been controversial, in large part because of its many novel morphological attributes. Because the most commonly collected species is small, and usually taken in very limited numbers, acquiring a decent sample of specimens takes considerable effort in the field. Our own fieldwork has allowed the present analysis to benefit from the recently published morphological analysis of Fischer (2000) and DNA sequence data for the group. Our morphological analyses (e.g. Figs 43 and 44) offer a less than convincing placement for the group, and only the PIWE result supports a sister-group relationship with Acanthosomatidae as proposed by Fischer (2000). However, all of our results that include DNA sequence data show the Lestoniidae + Acanthosomatidae forming a monophyletic group. The molecular data alone treat Lestonia as part of the Acanthosomatidae (Figs 45 and 46), whereas the combined analyses always place Lestonia as the sister group of the Acanthosomatidae sensu stricto (Figs 51–55). Morphological characters supporting the monophyly of the former grouping include: tarsi two-segmented (291), abdominal sternite VIII in males at most partially covered by segment VII (400), and Pendergrast̕s organ present in females (441).

DINIDORIDAE St��l Historical: Durai (1987), in a world revision of the Dinidoridae, recognized the subfamilies Dinidorinae, with the two tribes Dinidorini and Thalmini, and Megymeninae, also with two tribes, Megymenini and Eumenotini. Lis (1990) catalogued the Old World Dinidoridae. Rolston et al. (1996) summarized the current classification in their world catalogue of Dini- doridae. Gapud (1991) considered Dinidoridae and Tessaratomidae to be sister groups with two synapomorphies supporting the grouping: partially exposed spiracles on the second abdominal segment (these becoming completely exposed in Tessaratomidae), and greatly enlarged ninth paratergites. The Dinidoridae, on the other hand, is separated from the Tessaratomidae by the reduced gonangulum and the enlarged and often reticulate hemelytral membrane, both of which characters are not found exclusively in this group. Kocorek and Lis (2000), in a cladistic revision of the Megymeninae, proposed anew tribe, Byrsodepsini, and established Eumenotini as a junior synonym of Megymenini sensu stricto. Analytical result: After extensive searching, we were unable to secure, either through our own fieldwork or that of others, material adequate for sequencing other than for the genus Megymenum, leaving the Dinidorinae without molecular data. Our unweighted and successively weighted morphological analyses (Figs 42 and 43) treat the broadly conceived Dinidoridae as paraphyletic; analysis of the morphological data under implied weights using PIWE (Fig. 44) treats the Dinidoridae as a monophyletic subset of a paraphyletic Tessaratomidae. Megymenum is treated as the sister-group of Tessaratomidae in the 52-taxon molecular and combined analyses when using a 1: 1 cost ratio (Figs 45 and 51). In the 92-taxon combined analyses, the Dinidoridae is paraphyletic within a larger Tessaratomidae using a 1: 1 cost ratio (Fig. 53), is the monophyletic sister group of a monophyletic Tessaratomidae using a 1: 2 cost ratio (Fig. 54), and also includes Urochela (Urostylididae) when applying a 2: 2 cost ratio (Fig. 55). The close association of the Dinidoridae with the Tessaratomidae in our analyses is concordant with the conclusions of Gapud (1991), although there is no clearcut set of morphological characters that diagnoses that larger grouping. Future efforts should aim to produce a more broad-based taxon sample of sequence data for the group to test more rigorously its monophyly as well as its relationship with the Tessaratomidae as well as the remaining Pentatomoidea., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 967, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Durai, P. S. S., 1987. A revision of the Dinidoridae of the world (Heteroptera: Pentatomoidea). Orient. Insects 21, 163 - 360.","Lis, J. A., 1990. New genera, new species, new records and checklist of the Old World Dinidoridae (Heteroptera, Pentatomoidea). Ann. Upper Silesian Mus., Entomol. 1, 103 - 147.","Rolston, L. H., Rider, D. A., Murray, M. J., Aalbu, R. L., 1996. Catalog of the Dinidoridae of the World. Papua New Guinea J. Agric. For. Fish. 39, 22 - 101.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Kocorek, A., Lis, J. A., 2000. A cladistic revision of the Megymeninae of the world (Hemiptera: Heteroptera: Dinidoridae). Polskie Pismo Entomol. 69, 7 - 30."]}

PHLOEIDAE Amyot & Serville (+ SERBANINAE Leston), sensu Distant Historical: Amyot and Serville (1843) first recognized the neotropical ������Phleides������ as a suprageneric taxon. St��l (1872) treated the group as a subfamily of Pentatomidae. Leston (1953b) and Lent and Jurberg (1965) monographed the two genera and three species. Rolston and McDonald (1979) treated the group as a family. In Gapud��s (1991) analysis, the Phloeidae came out in a basal position following the Urostylididae. The interpretation of at least seven characters in Gapud��s matrix should be changed, based on more thorough morphological observations. Apparently, Gapud (1991) exam- ined specimens of Phloea corticata (Drury), and made the same erroneous interpretations of the female genitalia found in Lent and Jurberg (1965) and Rolston and McDonald (1979). Thegonocoxites 9 (= 2nd gonocoxae, 2nd valvifers) of these authorsare infactthe laterotergites 9, whereas the 9th paratergites (Rolston and McDonald, 1979) correspond to the laterotergites 8 (= 8th paratergites). Character 13 (2nd valvifers) of Gapud (1991), coded as distinctly separate, should be changed to 2nd valvifers with a distinct median fusion line. Also, the Phloeidae (including Serbaninae) have a membranous gonangulum rather than this structure being well sclerotized, as suggested by Gapud (character 17). The Serbaninae, originally described in the Phloeidae by Distant (1906), is monotypic and restricted to Borneo. Serbana borneensis Distant was removed and established as a higher taxon within the Pentatomidae by Leston (1953b), based mostly in the morphology of the male genitalia; this placement has more recently been accepted by Rider (2006). Analytical result: Phloeidae sensu lato receives support from our morphological analyses, on the basis of the invariant characters: body foliations present (31) and eyes divided on sagittal plane (81); and the homoplasious characters: base of corium expanded (201) and spiracles of segment VIII present, concealed by segment VII (391). This is despite the fact some prior authors have rejected the association of Phloeidae sensu stricto with Serbana. Nonetheless, the novel morphology of the Phloeidae complicates its placement within the Pentatomoidea on the basis of morphology alone. In the total evidence analysis of 92 taxa under an equal-cost regime, Phloeidae (including Serbana) becomes the sister group of a clade including the Plataspididae + Cyrtocoris + Megaris. The inclusion of Crytocoris in this grouping is found nowhere else in our analyses, and we consider it to be an artifact relating to the large amount of missing data in the 92-taxon combined analysis. Cyrtocoris is ������correctly������ placed as the sister group of the Pentatomidae under 1: 2, 2: 1, and 2: 2 cost regimes. At the same time, the Phloeidae move into the clade containing the Dinidoridae, Tessaratomidae, and some of the ������cydnoid������ taxa (Figs 54 and 55). That result is similartothecombinedmolecularanalysesundera 2: 2 weighting scheme (Fig. 46) and the 52-taxon total evidence analysis under a 1: 1 weighting scheme (Fig. 51). Further sequencing may or may not help to resolve this issue., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 964-965, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Amyot, C. J. B., Serville, A., 1843. Histoire naturelle des insectes He mipte`res. Librairie Encyclope'dique de Roret, Paris.","Leston, D., 1953 b. '' Phloeidae' ' Dallas: systematics and morphology, with remarks on the phylogeny of '' Pentatomoidea' ' Leach and upon the position of '' Serbana '' Distant (Hemiptera). Rev. Brasil. Biol. 13, 121 - 140.","Lent, H., Jurberg, J., 1965. Contribuic ¸ ao ao conhecimento dos Phloeidae Dallas, 1851, com um estudo sobre genitalia (Hemiptera, Pentatomoidea). Rev. Bras. Biol. 25, 123 - 144.","Rolston, L. H., McDonald, F. J. D., 1979. Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). J. NY Entomol. Soc. 87, 189 - 207.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Distant, W. L., 1906. Oriental Heteroptera. Ann. Soc. Entomol. Belg. 50, 405 - 408.","Rider, D. A., 2006. Pentatomoidea Home Page. North Dakota State University. http: // www. ndsu. nodak. edu / ndsu / rider / Pentatomoidea / [accessed on 21 July 2006]."]}

MEGARIDIDAE McAtee and Malloch Historical: This exclusively Neotropical and monogeneric taxon includes 16 species. Recognized as a subfamily of the Pentatomidae by McAtee and Malloch (1928), it was raised to family status by McDonald (1979), who studied the genitalia of both sexes of the Megarididae, Canopidae, and Plataspididae, and concluded that the well-developed scutellum in the three taxa represents only superficial resemblance. The simple nature of several structures of the genitalia, which might well be interpreted as derived loss conditions, caused Rolston and McDonald (1979) to believe that the megaridids probably represent an early offshoot from the pentatomoid line of evolution. Analytical result: The data adduced in the present study for relationships of Megaris are not as complete as those for Canopus, because we were unable to obtain specimens of Megaris suitable for DNA sequencing. Because the association of Megaris with the Plataspididae in our morphological (Figs 42���44) and some of the total evidence 92-taxon analyses (Figs 53���54) is largely based on body form; a sequence data set for this taxon would go some distance toward producing a more credible result regarding its placement within the Pentatomoidea. Certainly in the cases of Canopus and the Plataspididae, the DNA sequence data suggest that body shape alone is misleading with regard to the establishment of phylogenetic affinities. In lieu of the acquisition of additional data, we continue to maintain Megaris at the family level., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 968, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["McAtee, W. L., Malloch, J. R., 1928. Synopsis of pentatomid bugs of the subfamilies Megaridinae and Canopinae. Proc. U. S. Natl. Mus. 72, 1 - 21.","McDonald, F. J. D., 1979. A new species of Megaris and the status of the Megarididae McAtee and Malloch and Canopidae Amyot and Serville (Hemiptera: Pentatomoidea. J. NY Entomol. Soc. 87, 42 - 54.","Rolston, L. H., McDonald, F. J. D., 1979. Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). J. NY Entomol. Soc. 87, 189 - 207."]}

SAILERIOLIDAE China and Slater, revised status Historical: This taxon has been treated by many authors as a subfamily of the Urostylididae. Saileriola China and Slater resembles the Urostylididae sensu stricto mostly in the position and shape of the antennal insertion and the placement of the ocelli close to one another, the condition seen in Urostylis Westwood. China and Slater (1956) were not entirely comfortable with the placement of their new subfamily Saileriolinae under the Urostylididae, mentioning that this group must represent the Proto-Trichophora at the base of the Pentatomidae, Coreidae, and Lygaeidae. Gapud (1991) considered the group Saileriolinae + Urostylinae as the most ‘‘primitive’’ family of the Pentatomoidea, having retained many of the plesiomorphic characters which occur in Coreoidea, namely separate 2nd valvifers, dorsolateral antennophores, small bucculae, poorly developed preocular part of the head, and widely separated middle and hind coxae. Gapud (1991) studied Ruckesona vitrella Schaefer and Ashlock (1970); we conclude that he erroneously interpreted the 2nd valvifers in this species. Analytical result: Whereas the Saileriolidae has previously been treated as part of the Urostylididae, our morphological analysis indicates that the Urostylididae sensu lato is not a monophyletic group. We therefore recommend recognition of Saileriolidae at the family rank, with available morphological data supporting its position as the sister-group of all non-urostylidid Pentatomoidea (see Conclusions for list of characters). The Saileriolidae is diagnosed, in our analysis, by sternite VII in females with a longitudinal cleft (420; Fig. 24) and trichobothria present on urosternites V– VII, lateral, 2 + 2 or at least 1 + 1 (343; Fig. 18) (also found in Amnestinae). stricto mostly in the position and shape of the antennal insertion and the placement of the ocelli close to one another, the condition seen in Urostylis Westwood. China and Slater (1956) were not entirely comfortable with the placement of their new subfamily Saileriolinae under the Urostylididae, mentioning that this group must represent the Proto-Trichophora at the base of the Pentatomidae, Coreidae, and Lygaeidae. Gapud (1991) considered the group Saileriolinae + Urostylinae as the most ‘‘primitive’’ family of the Pentatomoidea, having retained many of the plesiomorphic characters which occur in Coreoidea, namely separate 2nd valvifers, dorsolateral antennophores, small bucculae, poorly developed preocular part of the head, and widely separated middle and hind coxae. Gapud (1991) studied Ruckesona vitrella Schaefer and Ashlock (1970); we conclude that he erroneously interpreted the 2nd valvifers in this species. Analytical result: Whereas the Saileriolidae has previously been treated as part of the Urostylididae, our morphological analysis indicates that the Urostylididae sensu lato is not a monophyletic group. We therefore recommend recognition of Saileriolidae at the family rank, with available morphological data supporting its position as the sister-group of all non-urostylidid Pentatomoidea (see Conclusions for list of characters). The Saileriolidae is diagnosed, in our analysis, by sternite VII in females with a longitudinal cleft (420; Fig. 24) and trichobothria present on urosternites V– VII, lateral, 2 + 2 or at least 1 + 1 (343; Fig. 18) (also found in Amnestinae).

TESSARATOMIDAE St��l Historical: Sinclair (1989), in a generic revision and cladistic analysis of the exclusively Southern Hemisphere taxon Tessaratomidae, concluded that this group is polyphyletic. He raised the Oncomerinae to family status. The Tessaratomidae under Sinclair��s (1989) vision included two subfamilies, Tessaratominae and Natalicolinae, the latter with two tribes, Natalicolini and Prionogastrini, and the latter of those comprising two subtribes, Prionogastraria and Sepinaria. More recently, Sinclair (2000) restored Oncomerinae as a subfamily of Tessaratomidae. Rolston et al. (1993) summarized the current classification in their world catalogue of the Tessaratomidae. Following Leston (1955), Leston (1956b) and Kumar (1969), they recognized the subfamilies Natalicolinae, Oncomerinae, and Tessaratominae, the last subdivided into three tribes: Prionogastrini, Sepinini, and Tessaratomini. Schuh and Slater (1995) recognized the three subfamilies mentioned above, the Tessaratominae subdivided into five tribes (Eusthenini, Platytatini, Prionogastrini, Sepinini, and Tessaratomini), and the Oncomerinae into two tribes (Oncomerini and Piezosternini). Sinclair (2000) revised the Oncomerinae, including in this subfamily 15 genera, but did not recognize Leston��s division of the subfamily into Oncomerini and Piezosternini. Monteith (2006) described maternal care of eggs and nymphs in five genera of Oncomerinae, three of them also showing nymphal phoresy where the nymphs are carried on the modified body of the female for a period after hatching. Analytical result: Our unweighted parsimony analysis of morphological character data supports the monophyly of this group (Figs 42), as does analysis under successive weighting (Fig. 43), whereas analysis with PIWE under implied weights treats it as paraphyletic (Fig. 44). In the combined molecular results and 52- taxon total evidence analyses under a 1: 1 cost ratio (Figs 45 and 51, respectively) the Tessaratomidae is always monophyletic and is the sister group of the Dinidoridae. The Tessaratomidae is paraphyletic via inclusion of the Dinidoridae in the 92-taxon combined analysis using a 1: 1 cost ratio (Fig. 53), but is monophyletic using 1: 2 and 2: 2 cost ratios (Figs 54 and 55). As with the Dinidoridae, a broader sample of sequence data would provide a more rigorous test of the monophyly and sister-group relationships of the Tessaratomidae and help to resolve the historical indecision as to the composition of the group., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 967-968, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Sinclair, D. P., 1989. A Cladistic, Generic Revision of the Oncomeridae Stal n. stat. and Tessaratomidae Schilling n. stat. (Hemiptera: Heteroptera: Pentatomoidea). PhD dissertation, University of Sydney, Sydney, Australia.","Sinclair, D. P., 2000. Generic revision of the Oncomerinae (Heteroptera: Pentatomoidea: Tessaratomidae). Mem. Queensl. Mus. 46, 307.","Rolston, L. H., Aalbu, R. L., Murra, M. J., Rider, D. A., 1993. Catalog of the Tessaratomidae of the World. Papua New Guinea J. Agric. For. Fish. 36, 36 - 108.","Leston, D., 1955. A key to the genera of Oncomerini Stal (Heteroptera: Pentatomidae: Tessaratominae) with the description of a new genus and species from Australia and new synonymy. Proc. R. Ent. Soc. London, series B, 24, 62 - 68.","Leston, D., 1956 b. Results from the Danish expedition to the French Cameroons 1949 - 50. IX. Hemiptera, Pentatomoidea. Bull. Inst. Fr. Afr. Noire (A) 18, 618 - 626.","Kumar, R., 1969. Morphology and relationships of the Pentatomoidea (Heteroptera). IV. Oncomerinae (Tessaratomidae). Aust. J. Zool. 17, 553 - 606.","Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY.","Monteith, G., 2006. Maternal care in Australian oncomerine shield bugs (Insecta, Heteroptera, Tessaratomidae). In: Rabitsch, W. (Ed.), Hug the Bug-For Love of True Bugs. Festschrift zum 70 Geburtstag von Ernst Heiss., Denisia 19, 1135 - 1152."]}

PLATASPIDIDAE Dallas Historical: Dallas (1851) was the first worker to recognize this Old World taxon at the family-group level. St��l (1864) treated it as a subfamily of Pentatomidae. Leston (1952) raised it to family status (= Brachyplatidae). Most modern workers do not recognize an infrafamilial classification, with the exception of Rider (2006), who recognized the subfamilies Brachyplatidinae and Plataspidinae. Gapud (1991) considered Plataspididae as the sister group of the Lestoniidae (Fig. 1f), in spite of the variability of the characters that grouped them (see comments under Lestoniidae). One character that was treated as synapomorphic for the Pentatomoidea by Gapud (1991) was the presence of a pair of ring sclerites (chitinellipsen; Dupuis, 1955). Gapud treated a single ring sclerite as plesiomorphic because that is the condition he found in the Coreidae that he examined. His observations indicated that these structures were absent in the Plataspididae, Acanthosomatidae, and Cyrtocorinae, a condition that he treated as a reversal. Our observations in part contradict those of Gapud (1991), as we found a pair of ring sclerites to be present in the species of Plataspididae and Acanthosomatidae that we examined, but in accordance with the observations of Gapud (1991) we did not observe ring sclerites in the Cyrtocorinae. Because the presence of ring sclerites varies greatly within families and subfamilies, we have not included this character in our morphological matrix. Analytical result: The Plataspididae resembles, at least superficially, the Canopidae and Megarididae, and indeed groups with them in our morphological analyses on the basis of one or more of the following characters (Figs 42���44): body sphaeroid (12), scutellum well developed (163), frena obsolete (182), and gonocoxites 9 joined by membrane (461). In the molecular and total evidence analyses, the position of the Plataspididae is more basal and always dissociated from the Canopidae, although not from the Megarididae; here it forms the sister group (sometimes in conjunction with other taxa) of nearly all Pentatomoidea except Urostylididae and Saileriolidae. These results suggest that the enlarged scutellum is the result of convergence, a conclusion that can be drawn from its observed occurrence in many groups of pentatomoids which show little relation to one another on the basis of other characters. Although the relatively basal placement of the Plataspididae contradicts the theories of all prior authors (Fig. 1), this position is not altered by changing the taxon composition or the cost regimes in the combined analyses (Figs 51���55). The monophyly of the Plataspididae is supported in our analyses by the condition of laterotergites 9 being contiguous and partially or totally covering segment X (470)., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 964, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Dallas, W. S., 1851. List of Specimens of Hemipterous Insects in the Collection of the Bristish Museum, Pt. 1. Trustees of the British Museum, London.","Leston, D., 1952. Notes on the Ethiopian Pentatomoidea (Hemiptera). V. On the specimens collected by the A. L. Capener, mainly in Natal. Ann. Mag. Nat. Hist., 5, 512 - 520.","Rider, D. A., 2006. Pentatomoidea Home Page. North Dakota State University. http: // www. ndsu. nodak. edu / ndsu / rider / Pentatomoidea / [accessed on 21 July 2006].","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Dupuis, C., 1955. Les genitalia des He mipte`res-He'teropte`res (genitalia externe des deux sexes; voies ectodermique femmelles). Revue de la morphologie, lexique de la nomenclature. Index bibliographique analytique. Me m. Mus. Hist. Nat. Paris (A) 6, 183 - 278."]}

CYDNIDAE Billberg Historical: Of all pentatomoid family-group taxa, the composition of the Cydnidae has probably been the most controversial over time. The taxon Cydnidae was first recognized by Billberg (1820). The modern concept of the group has been heavily influenced by the works of Froeschner (1960) and Dolling (1981). Both of these authors, and especially the latter, argued for an inclusive approach to conceiving the family, even though such a grouping is structurally somewhat heterogeneous. Froeschner (1960) recognized five subfamilies for the Western Hemisphere: Amnestinae, Cydninae, Garsauriinae, Scaptocorinae, and Sehirinae. Dolling (1981) recognized eight subfamilies within the Cydnidae: Amnestinae, Corimelaeninae, Cydninae, Garsauriinae, Scaptocorinae, Sehirinae, Thaumastellinae (formerly in Lygaeoidea), and Thyreocorinae. Lis (1994, 1999a,b) recognized seven subfamilies with the following classification: Amnestinae, Cephalocteinae (Cephalocteini, Scaptocorini), Corimelaeninae, Cydninae (Cydnini, Geotomini), Garsauriinae, Parastrachiinae, and Sehirinae (Amaurocorini, Sehirini). In their interpretation of the literature, Schuh and Slater (1995) proposed an amalgam of existing classifications supporting the inclusion of the Thyreocorinae and Parastrachiinae, both at subfamily rank, and the exclusion of the Thaumastellidae (following Jacobs, 1989). Cassis and Gross (2002) summarized the suprageneric classifications of some earlier authors; and Rider (2006) recognized five subfamilies (Amnestinae, Cydninae, Garsauriinae, Scaptocorinae, Sehirinae), without tribal subdivisions. Analytical result: Our morphological analyses for the Cydnidae produce the grouping proposed by Dolling (1981), on the basis of characters 160, 261, and 271, as mentioned above. All remaining analyses fail to recognize the Cydnidae sensu Dolling, but there is no strong signal as to how the constituent taxa should be grouped. We propose that there are probably two reasons for this inconsistency of grouping: first, our taxon sample for DNA sequences does not include several taxa that have been accorded subfamily rank by prior authors, and second, of all pentatomoid taxa, we had the greatest difficulty obtaining complete sequences for members of the subfamily Cydninae. We suggest that there may be good reason to question the monophyly of the Cydnidae sensu Dolling and recommend a more strongly analytical approach to determining its limits and composition. As part of this protocol, securing a sequence data set more representative of the recognized subgroups would seem to be the first priority (see also discussion under Corimelaenidae, Parastrachiinae, and Thaumastellidae)., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 966, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Billberg, G. J., 1820. Enumeratio Insectorum in Museo Gust. Joh. Billberg. Stockholm, 138 pp.","Froeschner, R. C., 1960. Cydnidae of the Western Hemisphere. Proc. U. S. Natl Mus. 111, 337 - 680.","Dolling, W. R., 1981. A rationalized classification of the burrower bugs (Cydnidae). Syst. Entomol. 6, 61 - 76.","Lis, J. A., 1994. A Revision of the Oriental Burrower Bugs (Heteroptera: Cydnidae). Upper Silesian Museum, Bytom, POL.","Lis, J. A., 1999 a. Taxonomy and phylogeny of the Cephalocteinae with reference to their historical biogeography (Hemiptera: Heteroptera: Cydnidae). Pol. J. Entomol. 68, 111 - 131.","Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY.","Jacobs, D. H., 1989. A new species of Thaumastella with notes on the morphology, biology and distribution of the two southern African species (Heteroptera: Thaumastellidae). J. Entomol. Soc. S. Afr. 52, 301 - 316.","Cassis, G., Gross, G. F., 2002. Hemiptera- Heteroptera (Pentatomomorpha). In: Houston, W. W. K., Wells, A. (Eds.), Zoological Catalog of Australia. CSIRO Publishing, B. Melbourne, Australia, Vol. 27.3 B, xiv + 737 pp.","Rider, D. A., 2006. Pentatomoidea Home Page. North Dakota State University. http: // www. ndsu. nodak. edu / ndsu / rider / Pentatomoidea / [accessed on 21 July 2006]."]}

Cimicomorpha Our sample of Cimicomorpha in this analysis is extremely limited and designed only to contribute to the range of outgroup morphology for the Pentatomomor- pha. As such, it appears that the morphological and molecular data are not sufficient to make judgments concerning the monophyly of the group. That issue was previously addressed by Schuh and Štys (1991) and is being revisited by R. T. Schuh et al. (in press) on the basis of a larger taxon sample and a robust sample of morphological and molecular characters.

THAUMASTELLIDAE Seidenst��cker Historical: The superfamilial position of this family group has been controversial since the time of its description. Having been originally placed in the Lyg- aeoidea, the Thaumastellidae was transferred to the Pentatomoidea at family rank by ��tys (1964a) and later reduced to a subfamily of the Cydnidae by Dolling (1981) and Jacobs (1989). Jacobs (1989) mentioned the presence of a microchromosome (m-chromosome) in the Thaumastellidae, which is also present in many Coreoidea, Lygaeoidea, and Pyrrhocoroidea (Largidae). Henry (1997) suggested that the m-chromosome is a synapomorphy defining this broad group within the Pentatomomorpha, albeit lost in a few taxa (Berytidae, Lygaeinae, and Piesmatidae), and that the Thaumastellidae, due to the presence of an m-chromosome, may therefore not belong in the Pentatomoidea. Nonetheless, many apomorphies of the Pentatomoidea are found in the Thaumastellidae, including the presence of the foretibial apparatus, expanded mandibular plates, the base of the head not forming a ������neck������, post-ocular tubercles absent, the structure of the female genitalia, and the paired lateral trichobothria. Analytical result: Because of the controversy over placement of the Thaumastellidae, and their rarity in collections, we felt fortunate to be able to secure sequences of two species of Thaumastella Horv��th, offering new evidence to test the relationships of the taxon. In our morphological analyses (Figs 42���44) the Thaumastellidae is placed within the Cydnidae, in conformity with the theory proposed by Dolling (1981), an unsurprising result. The strict consensus of molecular data with a 1: 1 cost ratio places the Thaumastellidae as the sister group of the Pentatomidae (Fig. 45); changing the cost ratio to 2: 2 places Thaumastella in a clade that includes the Urostylididae, Dinidoridae, Tessaratomidae, Canopus, and Phloea (Fig. 46). These results are more or less duplicated in the 52-taxon combined analyses (Figs 51 and 52), although there Thaumastella becomes the sister group of the Acanthosomatidae + Lestoniidae under a 1: 1 cost ratio (Fig. 51), rather than of the Pentatomidae. The 92-taxon combined analyses always place the Thaumastellidae in a clade containing most of the other ������cydnoid������ taxa (Figs 53���55). Although it seems abundantly clear that Thaumastella belongs within the Pentatomoidea, as opposed to the Lygaeoidea, because it never falls outside the ingroup in any of our analyses, we continue to maintain it at the family level, in the absence of a more clear-cut signal regarding the details of its phylogenetic placement within the group, and more particularly a more rigorous test of the monophyly of the Cydnidae sensu Dolling. Morphological features treated as diagnostic for the Thaumastellidae in our analyses are the short scutellum that at most barely surpasses the posterior margin of the metathorax (160) and the claval commissure reduced, being no more than half the length of the scutellum (171)., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 966-967, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Stys, P., 1964 a. Thaumastellidae - a new family of pentatomoid Hemiptera. Acta Soc. Entomol. Cechoslov. 61, 236 - 253.","Dolling, W. R., 1981. A rationalized classification of the burrower bugs (Cydnidae). Syst. Entomol. 6, 61 - 76.","Jacobs, D. H., 1989. A new species of Thaumastella with notes on the morphology, biology and distribution of the two southern African species (Heteroptera: Thaumastellidae). J. Entomol. Soc. S. Afr. 52, 301 - 316.","Henry, T. J., 1997. Phylogenetic analysis of family groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera), with emphasis on the Lygaeoidea. Ann. Entomol. Soc. Am. 90, 275 - 301."]}

Pentatomoidea Our study supports the monophyly of the Pentatomoidea on the basis of the following characters (Figs 42, 43, 53 and 54): (i) scutellum reaching or surpassing an imaginary transverse line crossing the connexivum at apical angles of 3rd abdominal segment (161); (ii) claval commissure obsolete, claval apices close together but not contiguous (172); (iii) abdominal trichobothria lateral on urosternites II���VII, usually 2 + 2 or at least 1 + 1 (342); and (iv) tergite VIII covering tergite IX in females (431). This character suite is in close agreement with the views of earlier authors (see Schuh and ��tys, 1991). In Figure 44, under implied weighting, the Pentatomoidea is not monophyletic, and neither is the group supported by the characters listed above., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 963, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Schuh, R. T., Stys, P., 1991. Phylogenetic analysis of cimicomorphan family relationships (Heteroptera). J. NY Entomol. Soc. 99, 298 - 350."]}

CANOPIDAE Amyot and Serville Historical: This exclusively Neotropical taxon, containing a single genus and eight species, was most recently revised by McAtee and Malloch (1928). Its association with the Scutelleridae, as proposed by Gapud (1991), was discussed in Schuh and Slater (1995), who continued to maintain the family status of the group. The two defining characters for the group, presence of a prosternal sulcus and strongly laminate prosternal carinae, are shared with two other families of Pentatomoidea, the Megarididae, and Corimelaenidae, a situation that has resulting in ambiguity concerning its relationships with other members of the Pentatomoidea. Analytical result: The position of the monogeneric Canopidae in our morphological analyses (Figs 42���44) shows little agreement with that derived from combined molecular analysis (Figs 45 and 46) and total evidence analysis of both 52 (Figs 51 and 52) and 92 taxa (Figs 53���55). Canopus is allied with Megaris and the Plataspididae in the first and with the phloeid���cydniddinidorid���tessaratomid clade in the last three. Although Canopus resembles Megaris St��l and members of the Plataspididae in body form and the expansion of the scutellum, our phylogenetic results indicate that these similarities are almost certainly superficial, as most past classifications have implied. The biology of the group is novel within the Pentatomoidea and uncommon in the Heteroptera more broadly, showing an apparently obligate association with fungi. We continue to maintain family status for Canopus, in light of the somewhat ambiguous nature of relationships in the present analyses., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on page 968, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["McAtee, W. L., Malloch, J. R., 1928. Synopsis of pentatomid bugs of the subfamilies Megaridinae and Canopinae. Proc. U. S. Natl. Mus. 72, 1 - 21.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY."]}

PARASTRACHIINAE Schaefer, Dolling and Tachikawa Historical: Schaefer et al. (1988) treated the Asian Parastrachia Distant as a subfamily within the Cydni- dae. More recently, Sweet and Schaefer (2002) elevated Parastrachia to family status, arguing that it did not share diagnostic features with either the Cydnidae, or the Pentatomidae, where some authors had placed the group. The characters supporting Parastrachia as distinct from the Cydnidae are: venation of the fore wings (base of membrane with large basal cells formed by cross veins), stridulatory apparatus differing from that of most Cydnidae and resembling that of the Amnestinae (Cydnidae), metathoracic scent gland area with a reduced spout above the ostiole, dorsum of the abdo- men relatively desclerotized, epipleurites absent, and several unique features in the female and male genitalia (Schaefer et al., 1988). Sweet and Schaefer (2002) placed the Parastrachiidae only within the ������cydnoids or lower pentatomoids������, arguing that inclusion of Parastrachia within the Cydnidae produces a grouping that cannot be diagnosed. Analytical result: Our analyses suggest that Parastrachia Distant and Dismegistus Amyot & Serville form a monophyletic group, an issue that has not been addressed in most of the recent���and rather extensive���literature on Parastrachia. For the characters included in our morphological matrix, these two taxa receive the same coding, so this result represents a foregone conclusion. Nonetheless, molecular data���adduced here for both Dismegistus and Parastrachia ���support this same conclusion, as do the total evidence analyses. We therefore broaden the concept of Sweet and Schaefer (2002) for the Parastrachiinae to include Dismegistus. Our morphological analyses (Figs 42���44) always place Parastrachia + Dismegistus within a broadly conceived Cydnidae, in a sense similar to that used by Dolling (1981), on the basis of characters 160, 261, and 271. As noted above, the molecular data alone and combined analyses of 52 taxa usually group Parastrachia and Dismegistus with Allocoris (Corimelaenidae) and with equal consistency with one of the sequenced taxa we have treated as Cydninae. The combined analyses of 92 taxa places some additional cydnoid taxa with the Parastrachia clade, but notably���and consistently���exclude two cydnines and Thaumastella. Thus, we conclude that Parastrachia + Dismegistus is a monophyletic group, and that its inclusion in a broadly conceived Cydnidae may well render that group paraphyletic, a conclusion in concordance with the findings of Sweet and Schaefer (2002). That particular conclusion does not preclude our recommendation that the Parastrachiinae be treated as part of a more broadly conceived Corimelaenidae. Whether the Sehirinae should also be included as part of the grouping may be clarified through the inclusion of additional sequence data, not only for the Sehirinae, but also for the Corimelaeninae., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 965-966, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Schaefer, C. W., Dolling, W. R., Tachikawa, S., 1988. The shieldbug genus Parastrachia and its position within the Pentatomoidea (Insecta: Hemiptera). Zool. J. Linnean Soc. 93, 283 - 311.","Sweet, M. H., Schaefer, C. W., 2002. Parastrachiinae (Hemiptera: Cydnidae) raised to family level. Ann. Entomol. Soc. Am. 95, 441 - 448.","Dolling, W. R., 1981. A rationalized classification of the burrower bugs (Cydnidae). Syst. Entomol. 6, 61 - 76."]}

SCUTELLERIDAE Leach Historical: This widely distributed taxon was first recognized as a family group by Leach (1815), Fieber (1861), and St��l (1867). Kirkaldy (1909) maintained the group as a subfamily of Pentatomidae with five tribes: Odontotarsini, Tetyrini, Scutellerini, Sphaerocorini, and Elvisurini. Van Duzee (1917) restored the group to family status. McDonald and Cassis (1984) erected a new subfamily, the Tectocorinae, andaccepted the Elvisurinae asavalidsubfamily. Intheirmorerecentsummaries of the literature, Schuh and Slater (1995) and Rider (2006), following Leston (1953a), recognized four subfamilies, Eurygastrinae, Odontotarsinae, Pachycorinae, and Scutellerinae, the latter author subdividing Scutellerinae into three tribes: Elvisurini, Scutellerini, and Sphaerocorini. Gapud (1991) noted thatthe Scutelleridaeischaracterized by the completely fused 2nd valvifers. Our examination reveals that the gonocoxites 9 (= 2nd valvifers) are not completely fused, asstatedby Gapud, but joinedmedially with a distinct fusion line (except in the Eurygastrinae) (see explanation of characters; corrections to Gapud��s Fig. 12). Fischer (2001) recognized the monophyly of Pachycorinae, Sphaerocorinae, and Elvisurinae, as well as a sister-group relationship between Tectocoris and Odontotarsinae; he further concluded that the Scutellerinae and Odontotarsinae are non-monophyletic groups. Cassis and Vanags (2006) monographed the Australian genera of Scutelleridae, updating McDonald and Cassis (1984) and Cassis and Gross (2002) in relation to the homologies and terminology of morphological characters. They also discussed the current literature on the monophyly and supra-familial position of the Scutelleridae within the Pentatomoidea, as well as for the infrafamilial classification of the scutellerids, recognizing five subfamilies. Analytical result: Although the status of the Scutelleridae has been debated (Lattin, 1964; Kumar, 1965; McDonald and Cassis, 1984; Fischer, 2001; Cassis and Gross, 2002; Cassis and Vanags, 2006), our analyses offer support for the concept of a monophyletic taxon, one that is reinforced by the morphological, molecular, and combined analyses. In addition to the morphological data, this conclusion is based on a reasonably good sample of previously unavailable DNA sequence data for one of the six recognized subfamilies. Morphological characters supporting scutellerid monophyly include one synapomorphic character: areas surrounding orificium receptaculi, in pars communis, with an elongate, grooved sclerite (541). The exact relationship of the Scutelleridae within the pentatomoid hierarchy is less clear, however. Gapud (1991) treated the Scutelleridae as the sister group of the Canopidae, the two groups having a central position in the cladogram (Fig. 1f). Our total evidence analyses are ambiguous as to the precise placement of the Scutelleridae, but always put the group distal to the Plataspididae and Parastrachiidae (Figs 51��� 55) and in the analyses under 1: 1 cost ratios always basal to the Acanthosomatidae + Pentatomidae (Figs 51 and 53). An improved sample of DNA sequence data across the range of scutellerid subfamilies might help to resolve this ambiguity., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 968-969, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Leach, W. E., 1815. Hemiptera. In: Brewster, D. (Ed.), Brewster ̕ s Edinburgh Encyclopedia. Edinburgh Vol. 9, pp. 57 - 192.","Fieber, F. X., 1861. Die europaischen Hemiptera Halbfluger (Rhynchota, Heteroptera). C. Gerold ̕ s Sohn. Wien., pp. 113 - 444.","Kirkaldy, G. W., 1909. Catalogue of Hemiptera (Heteroptera) with Biological and Anatomical References, List of Food Plants and Parasites, etc. vol. 1 Cimicidae. F. L. Dames, Berlin.","Van Duzee, E. P., 1917. Catalogue of the Hemiptera of America North of Mexico, Excepting the Aphididae, Coccidae and Aleurodidae. University of California Publications, Entomology 2.","McDonald, F. J. D., Cassis, G., 1984. Revision of the Australian Scutelleridae Leach (Hemiptera). Aust. J. Zool. 32, 537 - 572.","Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY.","Rider, D. A., 2006. Pentatomoidea Home Page. North Dakota State University. http: // www. ndsu. nodak. edu / ndsu / rider / Pentatomoidea / [accessed on 21 July 2006].","Leston, D., 1953 a. The suprageneric nomenclature of the British Pentatomoidea (Hemiptera). Entomologist ̕ s Gaz. 4, 13 - 25.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Fischer, C., 2001. Ein Beitrag zum Grundmuster, phylogenetischen System und zur Verwandtschaft der Scutelleridae (Heteroptera, Pentatomoidea). Inaugural-Dissertation zur Erlangung der Doktorwurde. Freie Universitat, Berlin, Germany.","Cassis, G., Vanags, L., 2006. Jewel bugs of Australia (Insecta, Heteroptera, Scutelleridae). In: Rabitsch, W. (Ed.): Hug the bug-For love of true bugs. Festschrift zum 70 Geburtstag von Ernst Heiss., Denisia 19, 275 - 398.","Cassis, G., Gross, G. F., 2002. Hemiptera- Heteroptera (Pentatomomorpha). In: Houston, W. W. K., Wells, A. (Eds.), Zoological Catalog of Australia. CSIRO Publishing, B. Melbourne, Australia, Vol. 27.3 B, xiv + 737 pp.","Lattin, J. D., 1964. The Scutellerinae of America north of Mexico (Hemiptera: Heteroptera: Pentatomidae). PhD dissertation, University of California, Berkeley, CA.","Kumar, R., 1965. Contributions in the morphology and relationships of the Pentatomoidea (Hemiptera: Heteroptera). Part I. Scutelleridae. J. Entomol. Soc. Queensland 4, 41 - 55."]}

UROSTYLIDIDAE Dallas, sensu stricto Historical: The systematic position of this family has been ambiguous over time, as was stressed by Schuh and Slater (1995). Singh-Pruthi (1925) related it to the Acanthosomatidae. Yang (1938a,b, 1939) and Pendergrast (1957) related it to the Pyrrhocoridae, and Miyamoto (1961) to the Pentatomidae. Kumar (1971) believed that the group represented an early divergence from the other pentatomomorphans, possibly together with the Pyrrhocoridae, with which they share uniquely the fused 2nd valvifers that form an M- or W-shaped sclerite. As Gapud (1991) did not provide an illustration of what he considered as separate 2nd valvifers, we conclude that he erroneously interpreted the 2nd valvifers in the species of Urostylididae that he studied. Berger et al. (2001) proposed the emended spelling Urostylididae, in order to remove the homonymy between the heteropteran family name and Urostylidae B��tschli (Ciliophora, Hypotrichia). Analytical result: The treatment of the Urostylididae as the basal grouping within the Pentatomoidea by Gapud (1991) is a position supported by our morphological analyses (Figs 42���44) and combined analyses under an unweighted equal costs regime (Figs 51 and 53). Under unequal costs, Urochela is dissociated from Urostylus and moved up to a more central position within the pentatomoid cladogram. Characters supporting the treatment of the Urostylididae sensu strico as the sister group of all remaining Pentatomoidea are listed below in the Conclusions. The monophyly of the Urostylididae sensu strico is recognized in our analysis by the presence of a stridulitrum on A 1 in the hind wing (221) and the presence of bristles on the claws (311). Additionally, gonocoxites 9 (in females) form an Mor W-shaped sclerite (465; Fig. 28), which we suggest argues for its treatment at the family level., Published as part of Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), pp. 932-976 in Cladistics 24 on pages 963-964, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["Schuh, R. T., Slater, J. A., 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, NY.","Singh-Pruthi, H., 1925. The morphology of the male genitalia in Rhynchota. Trans. R. Entomol. Soc. Lond. 1925, 127 - 267.","Yang, We. - Y., 1938 a. A new method for the classification of urostilyd insects. Bull. Fan. Inst. Biol. Peking 8, 35 - 48.","Yang, We. - Y., 1939. A revision of Chinese urostilyd insects (Heteroptera). Bull. Fan. Inst. Biol. Peking 9, 5 - 66.","Pendergrast, J. G., 1957. Studies on the reproductive organs of Heteroptera with a consideration of their bearing on classification. Trans. R. Entomol. Soc. Lond. 109, 1 - 63.","Miyamoto, S., 1961. Comparative morphology of alimentary organs of Heteroptera, with the phylogenetic consideration. Sieboldia 2, 197 - 259, pls. 20 - 49.","Kumar, R., 1971. Morphology and relationships of the Pentatomoidea (Heteroptera). 5 - Urostylidae. Am. Midl. Nat. 85, 63 - 73.","Gapud, V., 1991. A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philippine Entomol. 8, 865 - 961.","Berger, H., Heiss, E., Kerzhner, I. M., 2001. Removal of homonymy between Urostylidae Dallas, 1851 (Insecta, Heteroptera) and Urostylidae Butschli, 1889 (Ciliophora, Hypotrichia). Ann. Naturhist. Mus. Wien 103 (B), 301 - 302."]}

Osteopilus Fitzinger, 1843 TYPE SPECIES: Trachycephalus marmoratus Duméril and Bibron, 1841 (5 Hyla septentrionalis Duméril and Bibron, 1841). Calyptahyla Trueb and Tyler, 1974. Type species: Trachycephalus lichenatus Gosse, 1851 (5 Hyla crucialis Harlan, 1826), by original designation. DIAGNOSIS: This genus is diagnosed by 43 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. No morphological synapomorphies are known for this genus. CONTENTS: Eight species. Osteopilus brunneus (Gosse, 1851); Osteopilus crucialis (Harlan, 1826); Osteopilus dominicensis (Tschudi, 1838); Osteopilus marianae (Dunn, 1926); Osteopilus pulchrilineatus (Cope ‘‘1869’’ [1870]); Osteopilus septentrionalis (Duméril and Bibron, 1841); Osteopilus vastus (Cope, 1871); Osteopilus wilderi (Dunn, 1925)., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Itapotihyla, new genus TYPE SPECIES: Hyla langsdorffii Duméril and Bibron, 1841. DIAGNOSIS: Molecular autapomorphies include 122 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. A possible morphological autapomorphy is the presence of a prominent subcloacal flap. ETYMOLOGY: From Itapoti 1 ­ Hyla. The generic name is an allusion to the resemblance of the unique known species of this genus with lichens and mosses. Itapoti is a Tupi­Guarani term, a composition of ‘‘ita´’’ (5 rock) with ‘‘poti’’ (5 flower or to flourish), which means lichen or moss. CONTENTS: Monotypic. Itapotihyla langsdorffii (Duméril and Bibron, 1841), new comb.

Nyctimantis Boulenger, 1882 TYPE SPECIES: Nyctimantis rugiceps Boulenger, 1882, by monotypy. DIAGNOSIS: Molecular autapomorphies include 139 transformations in mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Possible morphological autapomorphies are the development of an irregular orbital flange in the frontoparietal, and the sphenethmoid almost completely concealed dorsally by the frontoparietals and nasals (Duellman and Trueb, 1976). CONTENTS: Monotypic. Nyctimantis rugiceps Boulenger, 1882., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Plectrohyla bistincta Group DIAGNOSIS: Exemplars of this species group in our analysis are diagnosed by 16 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. We are not aware of any morphological synapomorphy supporting this group. CONTENTS: Twenty­one species. Plectrohyla ameibothalame (Canseco­Márquez, Mendelson, and Gutiérrez­Mayén et al., 2002), new comb.; Plectrohyla arborescandens (Taylor, ‘‘1938’’[1939]), new comb.; Plectrohyla bistincta (Cope, 1877), new comb.; Plectrohyla calthula (Ustach, Mendelson, McDiarmid, and Campbell, 2000), new comb.; Plectrohyla calvicollina (Toal, 1994), new comb.; Plectrohyla celata (Toal and Mendelson, 1995), new comb.; Plectrohyla cembra (Caldwell, 1974), new comb.; Plectrohyla charadricola (Duellman, 1964), new comb.; Plectrohyla chryses (Adler, 1965), new comb.; Plectrohyla crassa (Brocchi, 1877), new comb.; Plectrohyla cyanomma (Caldwell, 1974), new comb.; Plectrohyla cyclada (Campbell and Duellman, 2000) new comb.; Plectrohyla hazelae (Taylor, 1940), new comb.; Plectrohyla labedactyla (Mendelson and Toal, 1996), new comb.; Plectrohyla mykter (Adler and Dennis, 1972), new comb.; Plectrohyla pachyderma, (Taylor, 1942), new comb.; Plectrohyla pentheter (Adler, 1965), new comb.; Plectrohyla psarosema (Campbell and Duellman, 2000), new comb.; Plectrohyla robertsorum (Taylor, 1940), new comb.; Plectrohyla sabrina (Caldwell, 1974), new comb.; Plectrohyla siopela, (Duellman, 1968), new comb.; Plectrohyla thorectes (Adler, 1965), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 104-105, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Osteocephalus Steindachner, 1862 TYPE SPECIES: Osteocephalus taurinus Steindachner, 1862, by subsequent designation of Kellogg (1932). DIAGNOSIS: This genus is diagnosed by 34 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this genus. CONTENTS: Seventeen species. Osteocephalus buckleyi (Boulenger, 1882); Osteocephalus cabrerai (Cochran and Goin, 1970); Osteocephalus deridens Jungfer, Ron, Seipp, and Almendáriz, 2000; Osteocephalus elkejungingerae (Henle, 1981); Osteocephalus exophthalmus Smith and Noonan, 2001; Osteocephalus fuscifacies Jungfer, Ron, Seipp, and Almendáriz, 2000; Osteocephalus heyeri Lynch, 2002; Osteocephalus leoniae Jungfer and Lehr, 2001; Osteocephalus leprieurii (Duméril and Bibron, 1841); Osteocephalus mutabor Jungfer and Hödl, 2002; Osteocephalus oophagus Jungfer and Schiesari, 1995; Osteocephalus pearsoni (Gaige, 1929); Osteocephalus planiceps Cope, 1874; Osteocephalus subtilis Martins and Cardoso, 1987; Osteocephalus taurinus Steindachner, 1862; Osteocephalus verruciger (Werner, 1901); Osteocephalus yasuni Ron and Pramuk, 1999., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Tlalocohyla, new genus TYPE SPECIES: Hyla smithii Boulenger, 1902. DIAGNOSIS: This genus is diagnosed by 92 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy. ETYMOLOGY: From Tlaloc, the Olmec God of the rain, 1 connecting ­ o 1 Hyla. The gender is feminine. COMMENTS: The inclusion of Hyla godmani and H. loquax is tentative and based on its association with H. picta and H. smithii in the former H. godmani group by Duellman (2001). The larvae of H. loquax and H. smithii share a reduction in the length of the third posterior tooth row (Caldwell, 1986; Lee, 1996). This feature is not present in the larvae of H. godmani as described by Duellman (1970). CONTENTS: Four species. Tlalocohyla godmani (Günther, 1901), new comb.; Tlalocohyla loquax (Gaige and Stuart, 1934), new comb.; Tlalocohyla picta (Günther, 1901), new comb.; Tlalocohyla smithii (Boulenger, 1902), new comb.

Bokermannohyla circumdata Group DIAGNOSIS: This species group is diagnosed by 52 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A putative morphological synapomorphy of this group is the presence of (usually thin) dark vertical stripes on the posterior surface of the thigh (Heyer, 1985). CONTENTS: Fifteen species. Bokermannohyla ahenea (Napoli and Caramaschi, 2004), new comb.; Bokermannohyla astartea (Bokermann, 1967), new comb.; Bokermannohyla caramaschii (Napoli, 2005) new comb.; Bokermannohyla carvalhoi (Peixoto, 1981), new comb.; Bokermannohyla circumdata (Cope, ‘‘1870’’ [1871]), new comb.; Bokermannohyla feioi (Napoli and Caramaschi, 2004), new comb.; Bokermannohyla gouveai (Peixoto and Cruz, 1992), new comb.; Bokermannohyla hylax (Heyer, 1985), new comb.; Bokermannohyla ibitipoca (Caramaschi and Feio, 1990), new comb.; Bokermannohyla izeckshoni (Jim and Caramaschi, 1979), new comb.; Bokermannohyla lucianae (Napoli and Pimenta, 2003), new comb.; Bokermannohyla luctuosa (Pombal and Haddad, 1993), new comb.; Bokermannohyla nanuzae (Bokermann and Sazima, 1973), new comb.; Bokermannohyla ravida (Caramaschi, Napoli and Bernardes, 2001), new comb.; Bokermannohyla sazimai (Cardoso and Andrade, ‘‘1982’’ [1983]), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 82-83, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Scinax ruber Clade DIAGNOSIS: This clade is supported by 53 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A morphological synapomorphy suggested for this clade by Faivovich (2002) is the proctodeal tube positioned above the margin of the lower fin. COMMENTS: Faivovich (2002) was skeptical about the monophyly of the S. ruber clade; however, the present analysis recovers it as monophyletic, with a considerable number of transformations supporting its monophyly. As in the case of several of the synapomorphies suggested by Faivovich’s (2002) analysis for Scinax, we are unsure as to whether the suggested morphological synapomorphies are optimized identically in our analysis. In particular, we do not know the taxonomic distribution within Dendropsophini for two other synapomorphies proposed for this clade (Faivovich, 2002): the arytenoids with a dorsal prominence developed over the pharyngeal margin, and absence of the lateral m. extensor brevis distalis digiti V (pes). Preliminary observation on the larvae of some species of Sphaenorhynchus (Sphaenorhynchus bromelicola, S. orophilus, S. pauloalvini, and S. prasinus; Faivovich, personal obs.) indicate that their proctodeal tubes are attached to the free margin of the lower fin, similar to the S. catharinae clade, instead of having the characteristic position seen in larvae of the S. ruber clade. Scinax megapodius and S. trachythorax are considered here to be junior synonyms of S. fuscovarius for reasons discussed in appendix 4. There are two species, Hyla dolloi and H. karenanneae, that upon examination of their type series we consider to be species of Scinax (see appendix 4 for further comments on them). CONTENTS: Fifty­six species. Eleven assigned to two groups, 43 unassigned to any group., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 96, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa perinesos Group DIAGNOSIS: A possible synapomorphy of this group is the purple coloration on the hands, feet, flanks, and concealed surfaces, as well as the purple venter with white granules (Cannatella, 1982). COMMENTS: We did not include any exemplar of this group in the analysis. Its monophyly is tentatively assumed following Cannatella (1982) and is based on the evidence mentioned above. CONTENTS: Four species. Phyllomedusa baltea Duellman and Toft, 1979; Phyllomedusa duellmani Cannatella, 1982; Phyllomedusa ecuatoriana Cannatella, 1982; Phyllomedusa perinesos Duellman, 1973., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 117, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyloscirtus armatus Group DIAGNOSIS: This species group is diagnosed by 103 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Duellman et al. (1997) suggested four synapomorphies of the H. armatus group: the presence of keratincovered bony spines on the proximal ventral surface of the humerus, on the expanded distal element of the prepollex, and on the first metacarpal; forearms hypertrophied; tadpole tail long with low fins and bluntly rounded tip; and the presence of a ‘‘shelf’’ on the larval upper jaw sheath. COMMENTS: Our observations of breeding males of the two species of this group indicate the presence of darkly pigmented, keratinized spicules in the dorsum, head (particularly lips), forelimbs, undersides of forelimbs, and pectoral and abdominal region. As the breeding biology of this and the other two species groups of the genus becomes better known, it will be possible to understand if the presence of these spicules are a putative synapomorphy of the H. armatus group. CONTENTS: Two species. Hyloscirtus armatus (Boulenger, 1902), new comb.; Hyloscirtus charazani (Vellard, 1970), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 84, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyla arborea Group DIAGNOSIS: This species group is diagnosed by 37 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this group. COMMENTS: The contents of the Hyla arborea group are restricted to avoid its paraphyly. The inclusion of the species that were not included in the present analysis and do not show the NOR in chromosome 6 is tentative, because no evidence, other than the molecular data presented here, is known to support its monophyly. CONTENTS: Fourteen species. Hyla annectans (Jerdon, 1870); Hyla arborea (Linnaeus, 1758); Hyla chinensis Günther, 1858; Hyla hallowellii Thomson, 1912; Hyla immaculata Boettger, 1888; Hyla intermedia Boulenger, 1882; Hyla meridionalis Boettger, 1874; Hyla sanchiangensis Pope, 1929; Hyla sarda (De Betta, 1853); Hyla savignyi Audouin, 1827; Hyla simplex Boettger, 1901; Hyla tsinlingensis Liu and Hu, 1966; Hyla ussuriensis Nikolsky, 1918; Hyla zhaopingensis Tang and Zhang, 1984., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 101-102, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Aplastodiscus albosignatus Group DIAGNOSIS: This species group is diagnosed by 42 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A possible morphological synapomorphy of this group is the presence of elaborate tubercles and ornamentation around the cloacal region (Cruz and Peixoto, ‘‘1985’’ [1987]). CONTENTS: Seven species. Aplastodiscus albosignatus (A. Lutz and B. Lutz, 1938), new comb.; Aplastodiscus callipygius (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus cavicola (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus flumineus (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus ibirapitanga (Cruz, Pimenta, and Silvano, 2003), new comb.; Aplastodiscus leucopygius (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus sibilatus (Cruz, Pimenta, and Silvano, 2003), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 82, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyla cinerea Group DIAGNOSIS: This species group is diagnosed by 35 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this group. COMMENTS: Hyla femoralis is excluded from the H. cinerea group to avoid the paraphyly of the group. CONTENTS: Three species. Hyla cinerea (Schneider, 1799); Hyla gratiosa LeConte, ‘‘1856’’ [1857]; Hyla squirella Bosc, 1800., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 102, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Ecnomiohyla, new genus TYPE SPECIES: Hypsiboas miliarius Cope, 1886. DIAGNOSIS: This genus is diagnosed by 37 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this genus. ETYMOLOGY: From the Greek, ecnomios, meaning marvelous, unusual; an obvious reference to the incredible frogs of the Hyla tuberculosa group. The gender is feminine. COMMENTS: This new genus contains the Hyla tuberculosa group, excluding H. dendrophasma, and including one species of the H. miotympanum group as well. Erecting a new genus for this clade is the only way of being consistent with the new monophyletic taxonomy that is proposed for hylids. Although naming the former H. tuberculosa group as a genus constitutes a testable claim of monophyly, we expect that it will ultimately be found to be two or three different clades, with one of these being the one named here. CONTENTS: Ten species. Ecnomiohyla echinata (Duellman, 1962), new comb.; Ecnomiohyla fimbrimembra (Taylor, 1948), new comb.; Ecnomiohyla miliaria (Cope, 1886), new comb.; Ecnomiohyla minera (Wilson, McCranie, and Williams, 1985), new comb.; Ecnomiohyla miotympanum (Cope, 1863), new comb.; Ecnomiohyla phantasmagoria (Dunn, 1943); Ecnomiohyla salvaje (Wilson, McCranie, and Williams, 1985), new comb.; Ecnomiohyla thysanota (Duellman, 1966), new comb.; Ecnomiohyla tuberculosa (Boulenger, 1882), new comb.; Ecnomiohyla valancifer (Firschein and Smith, 1956), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 100, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa burmeisteri Group DIAGNOSIS: We are not aware of any synapomorphy of this group. COMMENTS: We included only a single species of this group in our analysis, and as such we did not test its monophyly, but we recognize it following Pombal and Haddad (1992), pending a rigorous test of its monophyly. CONTENTS: Four species. Phyllomedusa burmeisteri Boulenger, 1882; Phyllomedusa distincta B. Lutz, 1950; Phyllomedusa iheringii Boulenger, 1885; Phyllomedusa tetraploidea Pombal and Haddad, 1992., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 117, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla pseudopseudis Group DIAGNOSIS: This group is diagnosed by 48 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy for this group. COMMENTS: Considering our results, which show the undescribed species of the Hyla pseudopseudis group (Hyla sp. 6) to be the sister taxon of an undescribed species similar with H. alvarengai (Hyla sp. 9), we are tentatively including the H. alvarengai in this species group. Eterovick and Brandão (2001) characterized this group on the basis of the presence of short, lateral irregular tooth rows and for having more tooth rows (between six and eight rows) in the oral discs of the larvae than do those of the Bokermannohyla circumdata group. However, the tadpole of H. ibitiguara, included in this group by Caramaschi et al. (2001), has a labial tooth formula of 2/4 (Cardoso, 1983) and seems to lack the short, lateral irregular tooth rows, as do tadpoles of H. alvarengai (Sazima and Bokermann, 1977). CONTENTS: Four species. Bokermannohyla alvarengai (Bokermann, 1956), new comb.; Bokermannohyla ibitiguara (Cardoso, 1983), new comb.; Bokermannohyla pseudopseudis (Miranda­Ribeiro, 1937), new comb.; Bokermannohyla saxicola (Bokermann, 1964), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 83-84, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla claresignata Group DIAGNOSIS: We are not aware of any synapomorphy supporting the monophyly of this group; see below. COMMENTS: We did not include any exemplar of this group, and as such we did not test its monophyly. See the earlier discussion regarding its position in the South American I clade. Considering the topology of Cophomantini, synapomorphies suggested for the Bokermannohyla claresignata group can only be maintained if assumed to be reversals (i.e., a enlarged larval oral disc, complete marginal papillae, and large number of labial tooth rows are also present in Myersiohyla new genus and the Hyloscirtus armatus group; complete marginal papillae and large number of labial tooth rows are also present in the H. bogotensis and H. larinopygion groups; the marginal papillae are also complete or with an extremely reduced gap in other species of Bokermannohyla). This would be unproblematic if it were a result of the analysis, but we prefer not to assume it a priori. While these character states cannot be considered at this stage to support the monophyly of the B. claresignata group, considering that its two species are barely distinguishable from each other, we think its nonmonophyly is unlikely and we continue to recognize the group as a hypothesis to be tested. CONTENTS: Two species. Bokermannohyla claresignata (A. Lutz and B. Lutz, 1939), new comb.; Bokermannohyla clepsydra (A. Lutz, 1925), new comb.

Aplastodiscus perviridis Group DIAGNOSIS: This species group is diagnosed by 58 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Apparent morphological synapomorphies of this group include the bicolored iris and the absence of webbing between toes I and II (known instances of homoplasy within hylids occur in some Scinax and in various groups of Lophiohylini) and reduction of webbing between the other toes (Garcia et al., 2001). CONTENTS: Two species. Aplastodiscus cochranae (Mertens, 1952); Aplastodiscus perviridis A. Lutz in B. Lutz, 1950. Bokermannohyla, new genus TYPE SPECIES: Hyla circumdata Cope, ‘‘1870’’ [1871]. DIAGNOSIS: This genus is diagnosed by 65 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy. ETYMOLOGY: This genus is dedicated to Werner Carlos Augusto Bokermann (1929– 1995), as homage to his contribution to the knowledge of Brazilian anurans. He also described several species now included in the new genus. The name derives from Bokermann 1 connecting ­ o 1 Hyla. We are adopting the ending ­ hyla for several of the new genera described here, most of which contain species groups formerly placed in Hyla. The gender is feminine. COMMENTS: Bokermannohyla includes all species previously allocated in the Hyla circumdata, H. martinsi, and H. pseudopseudis groups. We include tentatively the H. claresignata group pending the inclusion of its species in the analysis, because it was associated to the H. circumdata group by Bokermann (1972) and Jim and Caramaschi (1979). Hyla alvarengai is also included because our analysis shows that Hyla sp. 9 (aff. H. alvarengai) is nested within this new genus. These species groups should be maintained within Bokermannohyla until their monophyly is rigorously tested. CONTENTS: Twenty­three species, placed in four species groups., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 82, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa tarsius Group DIAGNOSIS: We are not aware of any synapomorphy supporting the monophyly of this group. COMMENTS: We included a single species of this group, and as such we did not test its monophyly, but we continue to recognize it following De la Riva (1999) until its monophyly is rigorously tested. CONTENTS: Four species. Phyllomedusa boliviana Boulenger, 1902; Phyllomedusa camba De la Riva, 2000; Phyllomedusa sauvagii Boulenger, 1882; Phyllomedusa tarsius (Cope, 1868). Species of Phyllomedusa Unassigned to Group There are several species that are currently not assigned to any group. These are: Phyllomedusa atelopoides Duellman, Cadle, and Cannatella, 1988; Phyllomedusa bicolor (Boddaert, 1772); Phyllomedusa coelestis (Cope, 1874); Phyllomedusa palliata Peters, ‘‘1872’’ [1873]; Phyllomedusa tomopterna (Cope, 1868); Phyllomedusa trinitatis Mertens, 1926; Phyllomedusa vaillanti Boulenger, 1882; and Phyllomedusa venusta Duellman and Trueb, 1967., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 117-118, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus columbianus Group DIAGNOSIS: The only morphological synapomorphy suggested for this group is the presence of two close, triangular lateral spaces between the cricoid and arytenoids at the posterior part of the larynx (Kaplan, 1999). COMMENTS: We included a single exemplar of this group, and as such we did not test its monophyly, but following Kaplan (1999) we recognize it on the basis of the evidence mentioned above. CONTENTS: Three species. Dendropsophus bogerti (Cochran and Goin, 1970), new comb.; Dendropsophus carnifex (Duellman, 1969), new comb.; Dendropsophus columbianus (Boettger, 1892), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 90, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus labialis Group DIAGNOSIS: We are not aware of any synapomorphy for this group. COMMENTS: We included a single exemplar of this group in the analysis, and as such we did not test its monophyly. Following Duellman and Trueb (1983) and Duellman (1989), we continue to recognize the group pending a rigorous test of its monophyly. CONTENTS: Three species. Dendropsophus labialis (Peters, 1863), new comb.; Dendropsophus meridensis (Rivero, 1961) new comb.; Dendropsophus pelidna (Duellman, 1989), new comb.

Scinax catharinae Clade DIAGNOSIS: This clade is diagnosed by 90 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Morphological synapomorphies suggested for this clade by Faivovich (2002) are absence of the anterior process of the suprascapula, internal vocal sac, distal division of the middle branch of the m. extensor digitorum comunis longus, and insertion of the medial side of this branch on the tendon of the m. extensor brevis medius digiti IV. COMMENTS: Regardless of problems imposed by the present results to interpretation of the possible synapomorphies of Scinax resulting from Faivovich’s (2002) analysis, the sparse available knowledge on the taxonomic distribution of the transformations supporting the monophyly of the very distinctive S. catharinae clade suggests that most of them still hold in the present analysis. An exception is the m. depressor mandibulae without an origin from the dorsal fascia at the level of the m. dorsalis scapulae, which also occurs in Hyla uruguaya, rendering its optimization ambiguous in our analysis., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 95, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218