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The status of Elachistocleis pearsei, E. surinamensis, E. sikuani and E. tinigua Elachistocleissurinamensis and E. pearsei aretheoldest available names in this group. The former was named by Daudin (1802) as Bufo surinamensis, and the latter was named by Ruthven (1914) as Hypopachus pearsei. Both were considered synonyms of Elachistocleis ovalis by Parker (1934). Dunn (1944; 1949) commented on H. pearsei under the new combination Elachistocleis pearsei. Carvalho (1954) placed it into the new genus Relictivomer Carvalho, 1954 based on the presence of the posterior part of the prevomer and, recently, de Sá et al. (2012) reallocated it into Elachistocleis based on DNA data. Kenny (1969) redescribed E. surinamensis from specimens from Trinidad and cited a personal communication from A. Grandison, who had found no differences between specimens of E. surinamensis from Trinidad and a paratype of E. pearsei. Duellman (1997) also compared specimens of E. pearsei from Panama and specimens of Elachistocleis sp. from Gran Sabana, south-eastern Venezuela (which he did not assign to any species, but which falls in the range of occurrence of E. surinamensis; see Fig. 3) and did not find any noticeable difference in the external morphology of adults or in their advertisement calls. Jowers et al. (2021) recently redescribed and designated a neotype for E. surinamensis from Trinidad but, unfortunately, provided no comparisons of the species with E. pearsei. While the present paper was in review, two new species from Colombia were described and named: Elachistocleis sikuani and E. tinigua (AcostaGalvis et al., 2022). Although we did not access the vouchers and did not include the new sequences generated therein in our analyses, we confirmed that two lineages already represented in our tree correspond to them (based on genetic similarity, 99.8%). The authors made a brief comparison of the new species with E. pearsei and E. surinamensis, mentioning some differences in body size, quantity and colour of the blotches in live specimens, degree of conspicuousness of the post-commissural glands and presence/absence of the mid-dorsal white line (Acosta-Galvis et al., 2022). The latter feature was reported as absent in E. sikuani and E. tinigua but present (‘evident’) in E. surinamensis. However, as mentioned above in discussion about the variation of E. surumu, the presence of a mid-dorsal white line in E. surinamensis was not reported in its original description (see: Daudin, 1802) or in its redescription and neotype designation (see: Jowers et al., 2021). On the contrary, Jowers et al. (2021) used the absence of the mid-dorsal line in E. surinamensis to distinguish it from E. nigrogularis. We also have not observed a mid-dorsal white line in any examined specimens of E. surinamensis (Fig. 1). Acosta-Galvis et al. (2022) did not provide a comparison between E. pearsei and E. surinamensis. Instead, they added more pieces to the puzzle by giving new names to two allopatric lineages. They followed the prevailing tradition in the taxonomy of Elachistocleis of creating more and more nomina rather than conducting a thorough review addressing the validity and application of available names. We reiterate that a comprehensive morphological revision of Elachistocleis is needed and should take priority over the naming of additional new Elachistocleis species. Unfortunately, our sampling of specimens of those species is limited, and a reassessment of the taxonomic status of those species is beyond the scope of the present work. Therefore, for time being, we recognize E. pearsei, E. surinamensis, E. sikuani and E. tinigua as valid species., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 560, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Daudin FM. 1802. Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris: Levrault.","Ruthven AG. 1914. Description of a new engystomatid frog of the genus Hypopachus. Proceedings of the Biological Society of Washington 27: 77 - 80.","Parker H. 1934. A monograph of the frogs of the family Microhylidae. London: Trustees of the British Museum.","Dunn ER. 1944. Los generos de anfibios y reptiles de Colombia, 1. Primera parte: anfibios. Caldasia 2: 497 - 529.","Dunn ER. 1949. Notes on South American frogs of the family Microhylidae. American Museum NoVitates 1419: 1 - 21.","de Carvalho AL. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 555: 1 - 22.","de Sa RO, Streicher JW, Sekonyela R, Forlani MC, Loader SP, Greenbaum E, Richards S, Haddad CFB. 2012. Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC EVolutionary Biology 12: 241.","Kenny JS. 1969. The amphibia of Trinidad. Studies on the fauna of Curacao and other Caribbean islands 108: 1 - 78.","Duellman WE. 1997. Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers Natural History Museum the UniVersity of Kansas 2: 1 - 52.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Acosta-Galvis AR, Tonini JFR, de Sa RO. 2022. Two new species of Elachistocleis Parker, 1927 (Anura: Microhylidae: Gastrophryninae) from Colombia. Zootaxa 5099: 527 - 548."]}

Elachistocleis surumu Caramaschi, 2010 as a junior subjective synonym of Elachistocleis surinamensis (Daudin, 1802) Although we do not have samples from the type locality of E. surumu (Vila Surumu, Municipality of Pacaraima, State of Roraima, Brazil), we sampled many specimens from nearby localities surrounding the type locality (Fig. 3) and all of them are recovered in a single lineage that is widespread in the eastern portion of the Guiana Shield (Figs 1, 3). The proposed morphological differences between E. surumu and E. surinamensis are subtle. Caramaschi (2010) differentiated E. surumu from E. surinamensis based on the alleged presence of a light vertebral stripe in E. surinamensis (absent in E. surumu) and minor details in the ventral colour patterns. However, neither the original description of E. surinamensis (Daudin, 1802) nor the recent species redescription and neotype designation (Jowers et al., 2021) stated that E. surinamensis has any kind of dorsal stripe or line. Caramaschi (2010) did not list any voucher specimens of E. surinamensis in his list of examined material. Ventral colour patterns of E. surumu are contained in the variation found herein for specimens of E. surinamensis (see also: Jowers et al., 2021). Therefore, we consider Elachistocleis surumu Caramaschi, 2010 as a junior subjective synonym of Elachistocleis surinamensis (Daudin, 1802)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 558, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Daudin FM. 1802. Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris: Levrault.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206."]}

THE ELACHISTOCLEIS BICOLOR GROUP We recognize four nominal species in the E. bicolor group: E. bicolor, E. haroi, E. helianneae and E. nigrogularis. Except for E. nigrogularis, species of this group predominantly have an immaculate belly, but see comments below., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 557, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481

Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill, Peloso, Pedro (2023): A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis). Zoological Journal of the Linnean Society 197: 545-568, DOI: 10.1093/zoolinnean/zlac057

The (unsettling) status of Elachistocleis ovalis There has been a long controversy regarding the validity and applicability of this nomen, especially regarding whether it should be applied to immaculate or to maculate-bellied specimens of Elachistocleis [see Lavilla et al. (2003) and Caramaschi (2010) for lengthy discussions on the issue]. While some authors (e.g. Parker, 1927; Dunn, 1949; Kenny, 1969; De la Riva et al., 2000) applied this name to maculate-bellied specimens – thus conveniently distinguishing them from the immaculate-bellied E. bicolor – others did not. In a meeting abstract of his unpublished dissertation, Carcerelli (1992) regarded E. bicolor a junior synonym of E. ovalis; a view that was followed by a few other authors (e.g. Klappenbach & Langone, 1992; Olmos & Achaval, 1997; Kwet & Di-Bernardo, 1998). That decision was mainly grounded on the fact that Schneider (1799) stated that Rana ovalis had an ‘ inferne flavidus ’ (yellow venter). Lavilla et al. (2003) also remarked that, considering Schneider’s description, E. ovalis should be applied to the specimens with ‘immaculate, yellow, ventral colouration’; but they did not adhere to the idea of synonymizing E. ovalis and E. bicolor. Instead, they regarded E. ovalis as restricted to the ‘northern portion of the generic range’, whereas E. bicolor would be restricted to the ‘southern portion of the generic range’. Nevertheless, ventral patterns have been recorded idiosyncratically in the literature and ‘uniform’, ‘yellow’ and ‘immaculate’ bellies have often been treated as synonyms, although this is not necessarily the case. Some individuals of typically maculatebellied species can present the venter almost entirely covered by yellow blotches (e.g. Fig. 5G, J, L) and thus could also fit into the ‘ inferne flavidus ’ description. Thus, assuming that the type of E. ovalis had an immaculate venter, because Schneider (1799) stated that it has ‘ inferne flavidus ’, is not solid reasoning in itself. Moreover, as described above for our samples from Cuiabá (AAG-UFU 5953 and AAG-UFU 5954), sympatric specimens may present different ventral patterns and still be genetically similar, increasing the complexity of this puzzle. The identity of E. ovalis is still a conundrum, especially difficult to solve due to the lack of a type locality and the unknown whereabouts of the type specimen. For these reasons, among others, Caramaschi (2010) proposed an operational solution: he considered Rana ovalis Schneider, 1799 and the combination Elachistocleis ovalis a nomen dubium associated with a species inquirenda; a decision that was followed by Jowers et al. (2021). Since we could not find any new evidence on the contrary, we also adhere to that decision by Caramaschi (2010). AmphibiaWeb (2022) and Frost (2022), the two largest amphibian taxonomic catalogues, have essentially also followed this suggestion and, although both still include this species in their list of 22 recognized Elachistocleis species, the following notes are added for E. ovalis: ‘Nominally, Elachistocleis ovalis is a nomen inquirenda (see comment), not applied to a biological population. But, pending revision, this name is applied to populations from Bolivia, although a great deal of confusion surrounds the identification of specimens mentioned from other countries’ (Frost, 2022); ‘This ancient name (1799) lacks type material and there is no type locality. Despite wide usage it should be considered an invalid name and not used [...] Pending revision, AmphibiaWeb continues to use the name E. ovalis ’ (AmphibiaWeb, 2022)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 561-562, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284.","Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6.","Dunn ER. 1949. Notes on South American frogs of the family Microhylidae. American Museum NoVitates 1419: 1 - 21.","Kenny JS. 1969. The amphibia of Trinidad. Studies on the fauna of Curacao and other Caribbean islands 108: 1 - 78.","De la Riva I, Kohler J, Lotters S, Reichle S. 2000. Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature and iconography. ReVista Espanola de Herpetologia 14: 19 - 164.","Klappenbach MA, Langone JA. 1992. Lista sistematica y sinonimica de los anfibios del Uruguay con comentarios y notas sobre su distribucion. Anales del Museo Nacional de Historia Natural de MonteVideo (2. a Serie) 8: 163 - 222.","Olmos A, Achaval F. 1997. Anfibios y reptiles del Uruguay. Montevideo: Barreiro y Ramos.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Schneider JG. 1799. Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens ranas, calamitas, bufones, salamandras et hydros in genera et species descriptos notisque suis distinctos. Jena: Friederici Frommanni.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Frost DR. 2022. Amphibian species of the aeorld: an online reference, V. 6.1. New York: American Museum of Natural History. Doi: 10.5531 / db. vz. 0001. Electronic database available at https: // amphibiansoftheworld. amnh. org / index. php (accessed 2 April 2022)."]}

The status of Elachistocleis cesarii, E. piauiensis and E. magna Miranda-Ribeiro (1920) described two subspecies of Engystoma ovale based on maculate-bellied specimens from the state of São Paulo, Brazil, which were later synonymized by Parker (1934) with Elachistocleis ovalis. Toledo et al. (2010) recently resurrected and redescribed them under the new combination Elachistocleis cesarii. Herein we recognize E. cesarii as a species distributed in south-eastern and central-western Brazil (Fig. 3) and closely related to E. piauiensis and E. magna (Fig. 1). Elachistocleis piauiensis was described and named by Caramaschi & Jim (1983) from Picos, Piauí. We found four lineages distributed in north-eastern Brazil, but we could not confidently assign any to E. piauiensis, because we did not have access to data from the type locality. Specimens assigned to each of these four lineages are morphologically similar to each other and to the original species description of E. piauiensis. The ventral colour pattern varies widely and similarly along the clade composed of E. cesarii, E. magna and the four candidate species E. cf. piauiensis 1–4, with no clear difference between them. The presence/absence of the mid-dorsal line also varies, except for E. cf. piauiensis 3, in which all examined vouchers lack the line. The holotype of E. piauiensis also lacks the line, and this could be evidence to assign the name to this specific clade. However, we cannot rule out that the consistent absence in E. cf. piauiensis 3 is not an effect of our sampling, given the high variability of this character state in several of our delimited species (Fig. 1). Elachistocleis cesarii was distinguished from E. piauiensis by differences in the advertisement call, size of the post-commissural gland and details of the inguinal region colouration (Toledo et al., 2010) – phenotypic traits that we have not explored for the present contribution. On the other hand, E. magna was distinguished from E. cesarii and E. piauiensis solely by its size (male SVL above 31 mm in E. magna, against SVL below 29 mm in E. cesarii and E. piauiensis; Toledo, 2010). Among examined specimens, we observed a consistent difference in size between E. magna and E. cf. piauiensis 1–4, but not between E. magna and E. cesarii. Males of E. cesarii can have SVL up to 31 mm, whereas some males of E. magna have SVL below 29 mm. Therefore, a thorough evaluation of phenotypic data allied with a denser molecular sampling is needed to verify if the differential diagnoses among E. cesarii, E. piauiensis and E. magna will hold true with additional data., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 560-561, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Miranda-Ribeiro A. 1920. Os engystomatideos do Museu Paulista (com um genero e tres especies novos). ReVista do Museu Paulista. Sao Paulo 12: 281 - 288.","Parker H. 1934. A monograph of the frogs of the family Microhylidae. London: Trustees of the British Museum.","Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60.","Caramaschi U, Jim J. 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura), from northeastern Brasil. Herpetologica 39: 390 - 394."]}

The (mysterious) status of Elachistocleis corumbaensis Unfortunately, we have no samples confidently assignable to E. corumbaensis. We have two samples from two localities near (around 20 km and 80 km) but not from the type locality of the species (Parque Municipal de Piraputangas, Corumbá municipality; Piva et al., 2017) (Fig. 3). For one of these tissue samples (CFBHT 00078), we were unable to find the voucher specimen. The other (MAPT 1363) was identified by the collectors as E. matogrosso (see: Koroiva et al., 2020). So we assume, albeit tentatively, that it has an immaculate venter. Both samples fall with E. bicolor in our species delimitation (Fig. 1). However, E. corumbaensis was described as having a maculate belly (Piva et al., 2017). The closest locality that we have a sample of a maculate-bellied species (E. cesarii) is Campo Grande, state of Mato Grosso do Sul, Brazil, 357 km from the type locality of E. corumbaensis. It is also worth remarking that E. corumbaensis has several similarities with E. haroi, particularly the dorsal pattern with a dark mark in the shape of a pine tree (Pereyra et al., 2013; Piva et al., 2017). Our closest sample of E. haroi is from Puerto Carmelo Peralta, Department of Alto Paraguay (Paraguay), 275 km south of the type locality of E. corumbaensis. For the moment, we consider E. corumbaensis a valid species, although it is unclear if this species belongs to the E. bicolor group or to the E. surinamensis group. The (unresolved) status of Elachistocleis erythrogaster This species is apparently restricted to the Centre of Research and Nature Conservation Pró-Mata, state of Rio Grande do Sul, above 900 m a.s.l. (Kwet & Di-Bernardo, 1998). It was distinguished from E. bicolor [treated as E. ovalis in Kwet & Di-Bernardo (1998)], which occurs in sympatry, by the unique red belly, larger size and other bioacoustic and reproductive characteristics (Kwet & Di-Bernardo, 1998). We have sampled some individuals from localities close to the type locality – although from the lowland – and they all fall with E. bicolor. This species has not been collected for nearly 20 years and is considered to be rare (Kwet et al., 2010)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 561, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154.","Koroiva R, Rodrigues LRR, Santana DJ. 2020. DNA barcoding for identification of anuran species in the central region of South America. PeerJ 8: e 10189.","Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Kwet A, Lingnau R, Di-Bernardo M. 2010. Pro-Mata: anfibios da Serra Gaucha, sul do Brasil - Amphibien der Serra Gaucha, Sudbrasilien - amphibians of the Serra Gaucha, south of Brazil. Tubingen and Porto Alegre: Brasilien-Zentrum de Universitat Tubingen and EDIPUCRS."]}

Elachistocleis bumbameuboi Caramaschi, 2010 and Elachistocleis carvalhoi Caramaschi, 2010 as junior subjective synonyms of Elachistocleis magna Toledo, 2010 Caramaschi (2010) distinguished E. bumbameuboi and E. carvalhoi from E. magna based on subtle differences in the dorsal and ventral colour patterns, as follows: regarding the dorsal pattern, E. magna has ‘dorsum uniform dark greyish with scarce minute brighter dots in the outer boundaries’, whereas E. bumbameuboi and E. carvalhoi have, respectively, ‘dorsum uniformly dark grey or black without marks’ and ‘dorsum uniformly brown or dark grey without marks’; regarding the ventral pattern, E. magna has ‘venter grey with minute scattered white spots, mainly on the belly and ventral surfaces of legs’, whereas E. bumbameuboi and E. carvalhoi have, respectively, ‘venter grey with minute anastomosed whitish spots, producing a salt-and-pepper pattern’ and ‘venter greyish with large anastomosed yellow or whitish yellow blotches, producing a coarse marbled pattern, mainly in the chest area. Photographs of the holotypes of E. carvalhoi (Caramaschi, 2010: fig. 3) and E. bumbameuboi (Caramaschi, 2010: fig. 4) and a photograph of a paratype of E. bumbameuboi (Nunes et al., 2010: fig. 1) depict specimens with dorsa scattered with minute white dots. Thus, our examination of specimens of E. bumbameuboi, E. carvalhoi and E. magna show a broad overlap among the various shades of dorsal colour and the size and distribution pattern of ventral markings (e.g. see Fig. 5). Based on our phylogenetic hypothesis, on the broad geographic sampling, which included a paratype of E. bumbameuboi, topotypes of E. carvalhoi and E. magna, and on the examination of several voucher specimens, we consider that E. bumbameuboi Caramaschi, 2010 and E. carvalhoi Caramaschi, 2010 are junior subjective synonyms of E. magna Toledo, 2010., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 558-560, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi & Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34."]}

TAXONOMY OF ELACHISTOCLEIS – A COMPLEX MATTER Parker (1927) erected Elachistocleis to allocate Rana ovalis Schneider, 1799 and Engystoma ovale bicolor (Guérin-Méneville, 1838). As usual at the time, the descriptions of both species were brief and did not designate type specimens. Lavilla et al. (2003) suggested that these names apply to complexes of species, because species of this genus are morphologically similar, with few obvious external characters that can be used as reliable diagnostic characters (i.e. non-overlapping and fixed). Ultimately, as mentioned throughout our contribution, diagnoses in this genus rely heavily on the ventral colour pattern, head proportions and the presence/absence and extension of body lines, such as a mid-dorsal white line and lines on the hidden surface of hindlimbs (Parker, 1927; Lavilla et al., 2003; Caramaschi, 2010; Pereyra et al., 2013). Caramaschi (2010) and Piva et al. (2017) suggested the ventral colour pattern as a diagnostic character for two internal species groups: a group with immaculate bellies (uniformly clear, free of markings) and another with maculate bellies. We found that the groups proposed by the above-cited authors are not monophyletic. For example, Elachistocleis muiraquitan (immaculate-bellied) is not related to the remaining immaculate-bellied species; instead, it was recovered in the E. surinamensis species group, a clade composed of species with predominantly maculate bellies. Furthermore, whereas E. panamensis has a maculate belly (Dunn et al., 1948; Nelson, 1972), E. araios has an immaculate belly (Sánchez-Nivicela et al., 2020). Given their phylogenetic position as successive sistertaxa to the remaining species of the genus (but see our comments on the position of E. araios), the description of E. araios already rendered the immaculate-bellied species group as paraphyletic. Also, it implies that the optimization of the ventral pattern of the Elachistocleis ancestor is ambiguous, given our topology and that of Sánchez-Nivicela et al. (2020). More importantly, we found intraspecific variation of the ventral pattern in some species, such as E. bicolor and E. nigrogularis (Figs 1, 4). Ontogenetic variation of the ventral pattern has also been recently reported for E. haroi (Bueno-Villafañe et al., 2020). Therefore, the available phylogenetic and morphological evidence emphatically rejects the existence of groups diagnosed solely by the colour of their bellies. It is noteworthy that this discussion seems far from settled. Ventral patterns have been recorded idiosyncratically in the literature. Sánchez-Nivicela et al. (2020) cite E. cesarii as ‘having uniform, immaculate, ventral colouration’, although Toledo et al. (2010) when resurrecting the species clearly state that it has ‘ventral colouration white or yellow with grey marks and reticulations’, and their figure 3d depicts a specimen with a maculate belly. Jowers et al. (2021) state that E. cesarii has a ‘uniform’ belly. This same was said of E. erythrogaster and E. bicolor. While E. cesarii and E. erythrogaster are maculate (the latter comparatively less maculate), E. bicolor bellies usually present no markings whatsoever (but see discussion on E. bicolor below). Given the historical use of the ventral pattern in the taxonomy of Elachistocleis, we attempted to verify as many specimens as possible for this trait. We have not extensively checked our vouchers for other morphological diagnostic traits (male throat colour; presence and shape of the femoral stripe; post-commissural gland). Nevertheless, our preliminary data suggest that none of the above-cited morphological traits is as reliable as suggested in the literature [see also comments in Nunes et al. (2010) and Marinho et al. (2018)]. Data from other sources are scarce. Tadpoles of few species are described (e.g. Rossa-Feres & Nomura, 2006; Pereyra et al., 2013; Schulze et al., 2015; Ferreira & Weber, 2021), but it is difficult to confidently assign these descriptions to lineages recovered because most lack an association with available DNA sequences. Moreover, there is no comprehensive understanding of variation, although Gómez & Kehr (2012) reported some variability in larval morphology related to chemical cues in the presence of predators. Advertisement calls of several Elachistocleis have been described (e.g. Nelson, 1972; Duellman, 1997; Kwet & Di-Bernardo, 1998; Lavilla et al., 2003; Nunes et al., 2010; Toledo, 2010; Marinho et al., 2018; Pansonato et al., 2018; Jowers et al., 2021). Some of these are associated with vouchers present in our phylogeny (see: Marinho et al., 2018; Jowers et al., 2021). These are not adequately distributed in our topology and most are described from a few specimens and localities. Published data have already shown that even slight increases in sample size may affect recorded variation (see: Marinho et al., 2018). Finally, some localities present at least two sympatric species of Elachistocleis. For instance, both E. nigrogularis and E. surinamensis are syntopically found at the type locality of E. nigrogularis (Jowers et al., 2021), and both E. bicolor and E. erythrogaster are found at the type locality of E. erythrogaster (Kwet & Di-Bernardo, 1998) (also see Fig. 3). This makes it even more difficult to associate larvae with adults and advertisement calls of unvouchered specimens to named species without molecular data. A fine line A white or light yellow mid-dorsal line is present in many individuals of Elachistocleis, and other genera of Gastrophryninae, such as Chiasmocleis Méhelÿ, 1904 (e.g. Peloso et al., 2014), Ctenophryne Mocquard, 1904 (e.g. Duellman, 1978), Hamptophryne A.L. Carvalho, 1954 (e.g. Parker, 1927; Duellman, 1978), Hypopachus [e.g. Cope (1889); see also figure 5 in Greenbaum et al. (2011)], Dasypops Miranda-Ribeiro, 1924 and Stereocyclops Cope, 1870 (PP, personal observation). Many authors have noticed important intraspecific or intrapopulational variation (i.e. being present in some individuals and absent in others) in this character in other genera (e.g. Chiasmocleis, Peloso et al., 2014; Ctenophryne, Zweifel & Myers, 1989; Stereocyclops, GNF, personal observation) and in Elachistocleis (e.g. Nelson, 1972; Toledo, 2010; Toledo et al., 2010; Marinho et al., 2018). Notwithstanding, the occurrence of this line and the variation of its extension when present (e.g. from snout to vent or from post-cephalic fold to vent) were used in diagnoses of several Elachistocleis species (e.g. Caramaschi & Jim, 1983; Caramaschi, 2010; Nunes-de-Almeida & Toledo, 2012; Piva et al., 2017). We found a remarkable variation in the occurrence of the mid-dorsal white line in and among species (Fig. 1), similar to the variation reported for Chiasmocleis (Peloso et al., 2014). This variation challenges the reliability of the trait as a diagnostic feature and weakens the known diagnoses of several currently recognized species (see below). It was impossible to confidently evaluate the extension of the mid-dorsal line, thus we only scored the presence or the absence of the line. Hence, we recommend caution when using the mid-dorsal line in the systematics of Elachistocleis, especially for diagnostic purposes., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 554-555, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6.","Schneider JG. 1799. Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens ranas, calamitas, bufones, salamandras et hydros in genera et species descriptos notisque suis distinctos. Jena: Friederici Frommanni.","Guerin-Meneville F-E. 1838. Iconographie du regne Animal de G. CuVier ou representation d'apres nature de l'une des especes les plus remarquables et souVent non enVore figurees, de chaque genre d'animaux, aVec un texte descriptiff mis au courant de la science, Vol. 3 (Part - Reptiles). Paris: J. B. Balliere.","Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284.","Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544.","Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154.","Dunn ER, Trapido H, Evans H. 1948. A new species of the microhylid frog genus Chiasmocleis from Panama. American Museum NoVitates 1376: 1 - 8.","Nelson CE. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia 1972: 895 - 898.","Sanchez-Nivicela JC, Peloso PLV, Urgiles VL, Yanez- Munoz MH, Sagredo Y, Paez N, Ron S. 2020. Description and phylogenetic relationships of a new trans-Andean species of Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae). Zootaxa 4779: 323 - 340.","Bueno-Villafane D, Caballlero-Gini A, Ferreira M, Netto F, Fernandez Rios D, Brusquetti F. 2020. Ontogenetic changes in the ventral colouration of post metamorphic Elachistocleis haroi Pereyra, Akmentins, Laufer, Vaira, 2013 (Anura: Microhylidae). Amphibia-Reptilia 41: 191 - 200.","Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi & Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34.","Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375.","Rossa-Feres D de C, Nomura F. 2006. Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of Sao Paulo State, Brazil. Biota Neotropica 6: 1 - 26.","Schulze A, Jansen M, Kohler G. 2015. Tadpole diversity of Bolivia's lowland anuran communities: molecular identification, morphological characterisation, and ecological assignment. Zootaxa 4016: 1 - 111.","Ferreira JS, Weber LN. 2021. A survey of the external morphology, internal oral morphology, chondrocranium and hyobranchial apparatus of Elachistocleis larvae Parker, 1927 (Anura, Microhylidae). Journal of Morphology 282: 472 - 484.","Gomez VI, Kehr AI. 2012. The effect of chemical signal of predatory fish and water bug on the morphology and development of Elachistocleis bicolor tadpoles (Anura: Microhylidae). Biologia 67: 1001 - 1006.","Duellman WE. 1997. Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers Natural History Museum the UniVersity of Kansas 2: 1 - 52.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Pansonato A, Mudrek JR, Nunes CA, Strussmann C. 2018. Advertisement calls of topotypes of Elachistocleis matogrosso (Anura: Microhylidae). Salamandra 54: 92 - 96.","Peloso PLV, Sturaro MJ, Forlani MC, Gaucher P, Motta AP, Wheeler WC. 2014. Phylogeny, taxonomic revision, and character evolution of the genera Chiasmocleis and Syncope (Anura, Microhylidae) in Amazonia, with descriptions of three new species. Bulletin of the American Museum of Natural History 386: 1 - 112.","Duellman WE. 1978. The Biology of an Equatorial Herpetofauna in Amazonian Ecuador. Lawrence: University of Kansas.","de Carvalho AL. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 555: 1 - 22.","Cope ED. 1889. Batrachia of North America. Bulletin of the United States National Museum 34: 5 - 525.","Greenbaum E, Smith EN, de Sa RO. 2011. Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae). Molecular Phylogenetics and EVolution 61: 265 - 277.","Zweifel RG, Myers CW. 1989. A New Frog of the Genus Ctenophryne (Microhylidae) from the Pacific Lowlands of Northwestern South America. American Museum NoVitates 2947: 1 - 16.","Caramaschi U, Jim J. 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura), from northeastern Brasil. Herpetologica 39: 390 - 394.","Nunes-de-Almeida CHL, Toledo LF. 2012. A new species of Elachistocleis Parker (Anura, Microhylidae) from the state of Acre, northern Brazil. Zootaxa 3424: 43 - 50."]}

Myrmecicultor Ram��rez, Grismado, and Ubick, new genus TYPE SPECIES. Myrmecicultor chihuahuensis, new species. ETYMOLOGY. The generic name is a combination of myrmex (ancient Greek for ���ant���) and cultor (Latin for worshiper, follower), and refers to the association between these spiders and their host ants. Gender masculine. DIAGNOSIS. By the characters of the family. DESCRIPTION. See under the species description., Published as part of Ram��rez, Mart��n J., Grismado, Cristian J., Ubick, Darrell, Ovtsharenko, Vladimir, Cushing, Paula E., Platnick, Norman I., Wheeler, Ward C., Prendini, Lorenzo, Crowley, Louise M. & Horner, Norman V., 2019, Myrmecicultoridae, a New Family of Myrmecophilic Spiders from the Chihuahuan Desert (Araneae: Entelegynae), pp. 1-24 in American Museum Novitates 2019 (3930) on page 6, DOI: 10.1206/3930.1, http://zenodo.org/record/4598671

Myrmecicultor chihuahuensis Ram��rez, Grismado, and Ubick, new species Figures 1���12, map 1 Cf. Gnaphosoidea TEX, Ram��rez, 2014 (provisional name in phylogenetic analysis). TYPES. Male holotype (CAS 9067289; temporary preparations CJG-1504, 1508), from the United States: Texas, Brewster Co., Dalquest Desert Research Station (DDRS), Big Sandy #4, N 29��33���05.7���, W 103��47���38.3���, 1232 m, Oct. 16���Nov. 1, 2014, N. V. Horner, propylene glycol pitfall traps, in 100 % ETOH. Male and female paratypes (CAS 9067288), DDRS, Big Sandy #3, N 29��33���04.2���, W 103��47���38.3���, elev. 1232 m, Sept. 24���Oct. 15, 2014, N. V. Horner, in 100 % ETOH. Female paratype (CAS 9067293; temporary preparation CJG-1505), DDRS, below Alamo #4, N 29��33���20.00���, W 103��47���04.7���, 1143 m, Oct. 16���Nov. 1, 2014, N. V. Horner, propylene glycol pitfall traps, in 100 % ETOH. Female paratype (AMNH [ARANP19], temporary preparations CJG-620, 632, MJR-1382���1383), DDRS, Sandy Canyon Sept. 22���Oct. 4, 2005, N. V. Horner and G.H. Broussard, pitfall covered, in propylene glycol. ETYMOLOGY. The specific epithet is an adjective referring to the Chihuahuan Desert, where most specimens were collected. DIAGNOSIS. By the characters of the family. MALE (holotype): Total length 2.89. Carapace length 1.47, width 1.20. Clypeus 0.26 (at AME), 0.19 (at ALE). Eye sizes and interdistances: AME 0.13, ALE 0.10, PME 0.13, PLE 0.09, AME-AME 0.06, AME-PME 0.08, AME-ALE 0.02, PME-PME 0.03, PME-PLE 0.07, ALE-PLE 0.02, AER 0.39, PLE-PLE 0.03, PER 0.43. Leg lengths: Leg I: total 4.70 (femur 1.55, patella 0.56, tibia 1.32, metatarsus 1.27, tarsus broken); Leg II: 4.98 (1.32 0.56 1.10 1.18 0.78); Leg III: 4.43 (1.19 0.49 0.97 1.13 0.65); Leg IV: 5.70 (1.47 0.56 1.29 1.55 0.83). Leg formula 4123. Color in alcohol: Carapace whitish, black in eye region; sternum whitish. Abdomen uniformly whitish both dorsally and ventrally. Legs also whitish, but distal podomeres slightly darker. Live specimens darker, with pale brown coloration (fig. 11F). Vestiture: Carapace almost glabrous, chelicerae with few needlelike setae at promargin. Sternum with marginal setae, some setae on posterior extension. Abdomen dorsum with two types of setae, mostly needlelike, with some thickened setae scattered irregularly; appendages densely clothed with thick setae, but with few spines (see below); metatarsi III and IV with distal preening brush of plumose setae. Carapace broadly oval in dorsal view, highest in front of fovea; thoracic fovea narrow, dark longitudinal line. Eyes mostly surrounded by black pigment, contrasting sharply with pale carapace (fig. 4D); AME largest, most convex, lateral eyes subequal, partly surrounded by black pigment, PME nearly rectangular, with oblique tapeta at 90�� angle. AER procurved in dorsal view, PER strongly procurved in dorsal view; both eye rows procurved in anterior view. Clypeus high, about twice ALE diameter (fig. 4C). Chelicerae as in female. Sternum heart shaped, anteriorly truncate, with posterior pointed extension between coxae IV, with marginal setae. Precoxal triangles absent. Labium trapezoidal, wider than long; endites quadrate, with serrula on anterior margin. Abdomen lacking dorsal scute. Epiandrum with two spigots (observed in KOH-digested specimen). Respiratory system consisting of two book lungs and wide tracheal spiracle slightly separated from spinnerets, giving origin to four simple, unbranched tracheae, all limited to abdomen, gradually attenuated. Spinnerets superficially as in female, but not examined with SEM; PMS apparently lacking spigots (as observed with stereomicroscope). Leg spines: I: femur d1-1-0, p1-0-0, metatarsus v2ap; II: femur 1-1-0, p1 (very small)-0-0, metatarsus v2ap; III: femur d1-1-0, tibia v2ap, metatarsus v2ap (below preening brush); IV: femur d1-1-0, tibia v2ap, metatarsus v2ap (below preening brush). Palp: femur with 1-1 dorsal macrosetae, lacking ventral process; tibia cup shaped, with RTA forming procurved concavity with retrolateral dorsal small, pointed protrusion; cymbium with dorsoapical patch of chemosensory setae (fig. 9G) situated in a cavity, dorsal surface with widened bipectinate setae (probably scales) (fig. 9F), and seven trichobothria in two longitudinal rows. Copulatory bulb: subtegulum simple, visible from retrolateral side; tegulum simple bearing spermophore without loops; median apophysis articulated, small hook in retrobasal position; two conductors (C1 and C2; figs. 9D, 10D), latter larger than former, more sclerotized, coiled, anteriorly directed. Embolus articulated, bent at right angle to retrolateral side; tip widened. FEMALE (paratype CAS 9067293): Total length 2.73. Carapace length 1.26, width 1.09. Clypeus 0.13 (at AME), 0.07 (at ALE). Eye sizes and interdistances: AME 0.14, ALE 0.09, PME 0.12, PLE 0.09, AME-AME 0.05, AME-ALE almost touching, AME-PME 0.06, PME-PME 0.02, PMEPLE 0.06, ALE-PLE 0.03, AER 0.37, PER 0.41. Palpus and leg lengths: Palpus: 1.35 (0.39 0.21 0.29 0.46); Leg I: 4.86 (1.33 0.51 1.14 1.09 0.79); Leg II: 4.27 (1.12 0.45 0.97 1.02 0.71); Leg III: 3.91 (1.03 0.40 0.84 1.03 0.61); Leg IV: 5.04 (1.28 0.49 1.16 1.35 0.75). Leg formula 4123. Leg spines: Palp: tarsus d1-1; I: femur d1-1-0, p1 (very small)-0-0; metatarsus v1ap; II: femur 1-1-0; metatarsus v1ap; III: femur d1-1-0, metatarsus v1ap (below preening brush); IV: femur d1-1-0, tibia v1ap, metatarsus v1ap (below preening brush, fig. 6B). Tarsal organ (observed on leg II) as simple pit, not domed (fig. 7C). Patellar indentation narrow, leading to retrolateral lyriform organs (fig. 7A). Tarsal step opposing metatarsal sensory organ with irregular, mainly longitudinal ridges (fig. 6G). Two tarsal claws, pectinate, without claw tuft or any other adhesive setae (fig. 6E, F); only two large setae with many barbs oriented distally, but without expanded tips. Trichobothrial shaft with expanded base covered by bumps, hood with several proximal transverse ridges (fig. 7B), on single row on leg tarsi. Palpal claw well developed, with teeth, tarsus apically with dorsal cavity containing a patch of chemosensory setae (fig. 5F, G). Color and general morphology (fig. 3) as in male. Vestiture as in male, widened bipectinate setae as that of male cymbium present on prolateral sides of palpal tibia and tarsus. Chelicerae slightly expanded anteriorly, anterior surface (fig. 5B, D) with row of rake setae and row of whisker setae parallel to fang, one promarginal whisker seta large (the promarginal escort seta), weak promarginal mound in front of fang base, promargin with one very small tooth (seen in male digested with KOH), retromargin without teeth, retromarginal escort seta present (socket observed, fig. 5E). Cheliceral fang very long, falcate, flat, with shaft serrula, venom duct opening present (fig. 5B���E). Genitalia: epigynum as single sclerotized plate with unpaired, shallow median concavity (fig. 3E). Posterior margin sinuous, with two rounded cavities (fig. 10F). Copulatory openings not visible, probably near central depression. Copulatory ducts fused anteriorly, with tortuous lumen, leading to simple spermathecae, copulatory duct and spermatheca together in uniformly sclerotized, solid piece (fig. 10F). Spinnerets small, clustered together, arising from slightly projecting base (fig. 8). ALS with basal articles conical, adjacent to each other, distal articles crescent-shaped, sclerotized, ectal area with setae (fig. 8C); seven piriform gland spigots tightly grouped, with shafts well defined from base, two major ampullate gland spigots with shafts larger than those of piriforms; no setae associated with piriform bases. PMS, PLS much shorter than ALS. PMS conical, with single spigot with thick shaft, probably cylindrical gland spigot. PLS with basal spigot with thick shaft, probably cylindrical gland spigot, and four smaller ones, probably aciniform gland spigots. ADDITIONAL MATERIAL EXAMINED. MEXICO: Coahuila: Municipio de Cuatroci��negas de Carranza: Cuatroci��negas Protected Area, ca. 9.7 km S Cuatroci��negas, N 26��54���31.896���, W 102��07���6.718���, Sep. 23, 2011, D.C. Lightfoot, 3♂ (MSBA 51822). Aguascalientes: Municipio de Tepazal��: 12 km (7.5 mi) N Tepezal��, Aug. 30, 1965, W. Gertsch, R. Hastings, 3♂ (AMNH; discovered by N.I.P. in May of 2019 while sorting unidentified specimens). U.S.A.: Texas: Hudspeth Co.: Indio Mountains Research Station (UTEP), N 30��46���34.7���, W 105��01���09.3���, Jul. 28���Aug. 30, 2017, N. V. Horner, 3♂, 1 juv. (CAS 9063001). Presidio Co.: Big Bend Ranch State Park: Ojito Adentro, N 29��29.5���, W 104��03.7���, Oct. 14, 2000, under rock, P.W. Hyder, 3♀ (AMNH 1256). DDRS Camp, N 29��33���25.91���, W 103��47���37.95���, 1266 m (4154 ft), propylene glycol pitfall trap [PGPT], G.H. Broussard: Sep. 5, 1999, 1♂ (AMNH); Sep. 15, 2000, 1♀ (AMNH). White Canyon Rd. #6, N 29��33���46.9���, W 103��48���03.2���, 1273 m (4177 ft), PGPT, N. V. Horner: Aug. 15���Sep. 9, 2015, 1♂ (MSU); Oct. 4���20, 2015, 1♀ (MSU). Brewster Co.: Below Alamo Springs, N 29��33���15.6���, W 103��47���08.4��� to N 29��33���17.8���, W 103��47���11��� to N 29��33���20���, W 103��47���04.7���, 1121���1158 m (3742���3798 ft), PGPT, N. V. Horner: May. 30���Jun. 30, 2014, 8 juv. (MSU), 7 juv. (CAS 9067292); Jul. 18���Aug. 17, 2012, 4♂ (CAS 9076507); Jul. 4���Aug. 12, 2013, 3 juv. (CAS 9067290, 9067291); Jul, 30��� Sep. 2, 2014, 7♂ (MSU); Aug. 15���Sep. 9, 2015, 3♂ (MSU); Aug. 24���Oct. 1, 2016, 3♂, 1♀ (MACN-Ar 38650), 1♂ (MACN-Ar 38649); Sep. 2���27, 2007, 1♂ (AMNH); Sep. 21���Oct. 6, 2007, 1♀ (CAS 9076536); Sep. 14���Oct. 9, 2010, 1♀ (CAS 9076511); Sep. 10���Oct. 3, 2015, 8♂, 3♀ (MSU); Oct. 7���Nov. 5, 2012, 1♂, 1♀ (CAS 9076528); 1♀ (CAS 9076530); Oct. 16���Nov. 1, 2014, 1♀ (CAS 9067293); Oct. 21��� Nov. 21, 2015, 2♀ (MACN-Ar 38651, 38663); Main Canyon, N 29.55512��, W 103.78561��, 1137 m, inside ant nest, P.E. Cushing, N. V. Horner, K. Sanko, C. Dowling, 10���11 Jun 2015, 1♂ (DMNS ZA.40328), 11 Jun 2015, 1 juvenile (DMNS ZA.40329); same, N 29.55506��, W 103.78536��, 12 Jun 2015 (DMNS ZA.40330). Brewster and Presidio Counties: Sandy Canyon, [N 29��33���07���, W 103��47���36���, 1215 m (3988 ft)], PGPT, G.H. Broussard: Oct. 7, 1999, 2♂ (AMNH); Sep. 15, 2000, 2♂, 1♀ (CAS 9076539, ♂ used for SEM); Big Sandy Canyon, N 29��33���07.38���, W 103��47���37.51��� to N 29��33���01.2���, W 103��47���38.1���, 1207���1260m (3960���4133 ft), PGPT, N. V. Horner: May. 23���Jun. 16, 2010, 1 juv. (CAS 9076513, 1♂ used for SEM); May. 30���Jun. 30, 2014, 1♂, 1♀, 4 juv. (MSU); Jun. 27���Jul. 13, 2009, 1 juv. (AMNH); Jun. 17���Jul. 22, 2010, 1 juv. (CAS 9076510); Jun. 14���Jul. 9, 2011, 1 juv. (CAS 9076519); Jun. 21���Jul. 17, 2012, 6 juv. (CAS 9076522, 9076526); Jun. 25���Jul. 24, 2016, 4 juv. (MSU); Jul. 14���Aug. 22, 2009, 4♂ (AMNH); Jul. 23���Aug. 10, 2010, 2 juv. (CAS 9076512); Jul. 10���Aug. 12, 2011, 1 juvenile (CAS 9076532); Jul. 18���Aug. 17, 2012, 3♂ (CAS 9076516), 1♂, 1♀, 3 juv. (CAS 9076518); Jul. 30���Sep. 3, 2014, 2 juv. (MSU); Aug. 15���Sep. 3, 2006, 1♀ (MSU); Aug. 29���Sep. 19, 2008, 1♀ (CAS 9076523); Aug. 10���28, 2010, 3♂ (MSU); Aug. 29���Sep. 13, 2010, 1♀ (MSU); Aug. 13��� Sep, 14, 2011, 2♂, 1♀ (CAS 9076533, 9076540, 9076537); Aug. 24���Oct. 1, 2016, 1♂ (MSU), 1♀ (MACN-Ar 38636); Sep. 22���Oct. 4, 2005, 1♀ (AMNH); Sep. 26���Oct. 10, 2006, 1♀ (MSU); Sep. 28���Oct. 9, 2008, 1♀ (CAS 9076535); Sep. 19���27, 2008, 2♂ (CAS 9076538); Sep. 14��� Oct. 9, 2010, 2♂, 2♀ (CAS 9076509, 9076514, 9076517); Sep. 15���Oct. 19, 2011, 1♂, 2♀ (CAS 9076520, 9076525, 9076527); 24 Sep���15 Oct 2014, 4♂, 1 juv. (MSU), 1♂, 1♀ (CAS 9067288); 9 Oct���2 Nov 2008, 1♂, 2♀ (AMNH); 4���14 Oct 2009, 1♀ (AMNH); 6 Oct���5 Nov 2012, 4♀ (CAS 9076521, 9076529, 9076534); 16 Oct���1 Nov 2014, 1♂, 1♀ (DMNS ZA.40327), 1♂ (CAS 9067289). DISTRIBUTION. Known only from the Big Bend region (Presidio, Brewster and Hudspeth counties) of Texas, and in Mexico from Cuatro Ci��negas, Coahuila, and Tepezal��, Aguascalientes (map 1)., Published as part of Ram��rez, Mart��n J., Grismado, Cristian J., Ubick, Darrell, Ovtsharenko, Vladimir, Cushing, Paula E., Platnick, Norman I., Wheeler, Ward C., Prendini, Lorenzo, Crowley, Louise M. & Horner, Norman V., 2019, Myrmecicultoridae, a New Family of Myrmecophilic Spiders from the Chihuahuan Desert (Araneae: Entelegynae), pp. 1-24 in American Museum Novitates 2019 (3930) on pages 6-16, DOI: 10.1206/3930.1, http://zenodo.org/record/4598671

Camponotus chloroticusEMERY, 1897 (Figs. 15 �� 20; Tabs. 2, 3) Camponotus maculatus ssp. chloroticus EMERY, 1897. Combination in Camponotus (Myrmoturba), as Camponotus (Myrmoturba) maculatus chlorotica var. chlorogaster: EMERY, 1914. Camponotus (Myrmoturba) maculatus pallidus var. samoensis SANTSCHI, 1919, unavailable name. Homonym of Camponotus irritans samoensis (SMITH, 1857). Camponotus (Myrmoturba) maculatus ssp. sanctae crucis MANN, 1919. Subspecies of Camponotus irritans: EMERY 1920. Combination in Camponotus (Tanaemyrmex): EMERY 1925. Subspecies of Camponotus irritans: KARAVAIEV 1933. Raised to species: WILSON& TAYLOR 1967. Comments: Camponotus chloroticus was originally de�� scribed by EMERY (1897) as a subspecies of C. macula�� Tab. 3: Caste and label trans�� tus, as follows:"I bought from Godeffroy Museum[Hamburg, 1861 �� 1885] specimens of this form from the Tonga Islands and New Britain, under the name C. pallidus. ��� For the shape of the various parts of the body, for the pubescence, the very weak sculpture and the hairs, it is very close to the C. Kubaryi, MAYR[specific epithet capitalized in original], particularly the oceanic specimens and those from New Guinea. ��� Maximum size is 8 mm; reddish��yellow, dirt��like color; head darker and more red, abdomen more or less blackish in its rear." We do not know which aspects of the pilosity EMERY noticed as being similar to that of Camponotus kubaryi stat. rev., but the presence of standing hairs on the proximal hind femur and on the propleuron in both species is one of the few readily discernable synapomorphies of an important clade of Camponotus in the Pacific and one of the key characters used to distinguish C. chloroticus from C. micronesicus. Using this pilosity character, overall similarity in size, shape, and coloration, as well as our finding of only one such yellow Camponotus species in the same islands, we confirm here that the Tongan syntypes of C. chloroticus (Figs. 15, 16) match the species in Clade IV, which extends from New Guinea to Polynesia (Fig. 2). The Camponotus chloroticus syntype from Irupara, New Guinea (Figs. 17 �� 20), is not as clearly aligned with the Tongan syntypes or modern specimens from the Polynesian, Fijian, and Melanesian clade, partially due to its mounting, which limits our view of the important pilosity characters. However, the New Guinean syntype has distinctly shorter scapes than almost all C. micronesicus sp.n. specimens measured, measuring just at the lower limit of the range, and producing a scape index for the New Guinea syntype that is smaller than all C. micronesicus specimens measured but within the range for C. chloroticus. The petiole length of the New Guinean syntype is also similar to that of C. chloroticus specimens, and altogether we have more support for it being C. chloroticus than C. micronesicus sp.n. Other options for the identity of the New Guinean syntype include an undescribed from, or, if it is truly missing the hind femur and propleuron standing hairs, an oddly concolorous C. humilior (which tends to be bicolorous); C. novaehollandiae is too large, also usually bicolorous, and, from our PCA analysis, slightly different in shape. To the original description we add a summary of our morphological observations of this species, combining syntypes and modern specimens, as follows (also see Tabs. 2, 3). Majors: EL 0.48 (range 0.40 �� 0.51), EW 0.36 (0.25 �� 0.40), FCL 1.14 (1.00 �� 1.31), HL 2.22 (1.8 5 �� 2.40), HW 2.00 (1.45 �� 2.25), ML 2.69 (2.38 �� 2.85), MTL 1.68 (1.44 �� 2.10), PH 0.74 (0.59 �� 0.81), PL 0.58 (0.43 �� 0.75), SL 1.65 (1.44 �� 1.85); CI 90 (78 �� 9 5), SI 83 (73 �� 124). Mesosoma light yellow, gaster same color as mesosoma or slightly darker, head color usually darker than mesosoma; head tapering, vertex usually slightly concave; hind femur and propleuron with standing hairs. Minors: EL 0.42 (0.38 �� 0.55), EW 0.33 (0.30 �� 0.40), FCL 1.0 4 (0.90 �� 1.40), HL 1.66 (1.55 �� 2.10), HW 1.28 (1.18 �� 1.60), ML 2.39 (2.20 �� 3.05), MTL 1.58 (1.31 �� 1.95), PH 0.62 (0.50 �� 0.75), PL 0.57 (0.50 �� 0.70), SL 1.80 (1.45 �� 2.45); CI 77 (74 �� 82), SI 142 (123 �� 154). Mesosoma usually light yellow, gaster and head usually same color as mesosoma or slightly darker; head tapering, vertex convex and occipital carina present; hind femur and propleuron with standing hairs., Published as part of Ronald M. Clouse, Benjamin D. Blanchard, Rebecca Gibson, Ward C. Wheeler & Milan Janda, 2016, Taxonomic updates for some confusing Micronesian species of Camponotus (Hymenoptera: Formicidae: Formicinae), pp. 139-152 in Myrmecological News 23 on pages 144-147, DOI: 10.5281/zenodo.164974, {"references":["EMERY, C. 1897: Viaggio di Lamberto Loria nella Papuasia orientale. XVIII. Formiche raccolte nella Nuova Guinea dal Dott. Lamberto Loria. - Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova) (2) 18 (38): 546 - 594.","WILSON, E. O. & TAYLOR, R. W. 1967: The ants of Polynesia (Hymenoptera: Formicidae). - Pacific Insects Monograph 14: 1 - 109."]}

Camponotus tol sp.n. GIBSON & CLOUSE (Figs. 30 �� 35) Type material: Holotype major worker. Federated States of Micronesia: Chuuk, Tol Island at 120 m (7o 19' 27.3" N, 151o 36' 50.6" E), leg. R. Clouse, P. Sharma, and Techuo family. Paratypes, (3 major workers, 9 minor workers), same collection data as holotype. Twelve additional minors stored in 95% EtOH, as well as two minors each with one leg removed and used for DNA extraction, also same collection data as holotype. All specimens are deposited in the Museum of Comparative Zoology, Harvard University, Massachusetts, USA. Description of holotype major worker (Figs. 30 �� 32): EL 0.48, EW 0.30, FCL 1.05, HL 2.05, HW 1.84, ML 2.51, MTL 1.54, PH 0.54, PL 0.60, SL 1.77; CI 90, SI 96. Mandible outer margin gently curved to an apex of about 75 degrees, the masticatory margin straight in front view. Mandibles tightly closed, masticatory margin with five visible teeth that gradually diminish in size from apex. Clypeus continuing anteriorly past mandibular insertions a distance slightly less than length of apical tooth, then straight across. Posterior clypeus curved anteriorly forming bilobed appearance, antennal insertions separated from clypeus by a distance almost equal to the distance from nearest clypeal margin to clypeal midpoint. Head slightly longer than wide. Vertex weakly concave. In frontal view eyes located halfway between posterior clypeal margin and vertex; inner margins halfway between frontal lobes and sides of head; eyes not extending past lateral edge. Antennae 12��segmented. Antennal scape length extending past the vertex by a distance of 2 �� 3 times the width of the scape at the apex. Mesosoma in profile gently sloping from anterior pronotum to dorsal propodeum, with moderate propodeal declivity. Color: Distinct bicoloration: head mostly glossy brown, mesosoma uniform light yellow��brown, gaster glossy brown. Gradual lightening from vertex down to mandibles. Vertex to posterior clypeus same glossy brown as gaster. Black outlining along frontal carina. Anterior clypeus to mandibular insertions light yellow��brown. Mandibles lighter brown than vertex down to posterior clypeus. Teeth of mandibles, scrobes, sutures, and joints on the head darker than surrounding cuticle. Each gastral tergite with hyaline margin along posterior fifth. Pilosity: Layer of short, recumbent, light hairs all over head. Long, yellow, standing hairs numerous on front, back, and sides of head. In frontal view, area between eyes and frontal carina with two rows of long, standing hairs extending from vertex to mid��clypeus. Row of long hairs extending across anterior clypeal edge. From dorsal view, mesosoma with two side��by��side groups of standing hairs on pronotum, four long hairs with some small hairs per group; one group of standing hairs on mesonotum, with three long hairs and some small hairs; two groups of standing hairs clustered on propodeum, with three long hairs per group; layer of short recumbent hairs all over each appendage, decreasing in length from trochanter to tarsus. Propleuron standing hairs indistinct but present. Each gastral tergite with 20 to 30 long, yellow, standing hairs encircling tergite immediately before hyaline margin along posterior edge; 10 to 20 longer standing hairs encircling tergite halfway between hyaline margin and posterior edge of previous tergite. Standing hairs on hind femur indistinct but present. Sculpturing: Head and gaster surface glossy; genae weakly punctured. Mesosoma surface smooth and shiny, although not glossy. Description of paratypes: Majors closely resembling holotype except vertex and mandibles more reddish��brown; overall slightly larger (ML 2.6 �� 2.75 and HW 1.88 �� 1.95); CI same (90) but SI smaller (87 �� 92), indicating consistency in head shape and scape absolute length in larger specimens. Minors closely resembling majors in coloration and pilosity. Slightly smaller than majors (ML 2.00 �� 2.25) and with significantly narrower heads and longer, more variable scapes (CI 76 �� 80, SI 157 �� 172). Scapes extending beyond vertex by a range of 25% �� 50% of total scape length. Posterior clypeus lighter brown than holotype. Eyes extending past lateral outline of head. One minor with open mandibles has six visible teeth on masticatory margin that gradually diminish in size from the apex. Measurements of minor collected with holotype and pictured in Figures 33 �� 35: EL 0.45, EW 0.33, FCL 0.82, HL 1.34, HW 1.05, ML 2.08, MTL 1.31, PH 0.45, PL 0.45, SL 1.73; CI 78, SI 165. Differential diagnosis: The only specimens from Chuuk that might be mistaken for Camponotus tol sp.n. are very dark, small C. micronesicus sp.n., but C. tol sp.n. has hairs on the propleuron and hind femur, and C. micronesicus sp.n. does not. Among other similar species in Micronesia, C. tol sp.n. is not as starkly bicolorous as C. eperiamorum, and it is approximately 75% the overall size of C. kubaryi stat. rev. Habitat: This species was collected from low��elevation, mixed agroforest (120 m a.s.l.) on Tol Island in Chuuk Lagoon. This island is the largest in the region, and reaches a maximum elevation of 439 m, but it shows evidence of agroforesty and agro��native mixed forest at all elevations. Specimens were collected from one colony and appeared to be less abundant on the island than Camponotus micronesicus sp.n. Etymology: The specific epithet is a noun in apposition to the genus that refers to the type locality, Tol Island (pronounced " tōl," as in the common word "toll"). Tol is the largest island inside the main atoll and island group of Chuuk State in the Federated States of Micronesia. The island is sometimes written as "Ton," as the phonemes "l" and "n" are not distinct in Chuukese. Comments: Camponotus tol is part of a larger clade in the molecular phylogeny (Clade IV; Fig. 2) that includes C. chloroticus, C. kubaryi stat. rev., an undescribed species collected in Papua New Guinea and Indonesia, and an endemic species found on the nearby Micronesian island of Pohnpei, C. eperiamorum (CLOUSE 2007b, CLOUSE & al. 2015). These species are all larger than C. tol, and they are also distinguished by their coloration: C. chloroticus and the unnamed species are lighter, C. kubaryi stat. rev. is darker, and C. eperiamorum contrasts more in shade among the mesosoma, head, and gaster. All of these species have the distinctive hairs on the propleuron and hind femur, but only C. tol sp.n. has such a distinctive double row of long hairs down the front of the head. Overall, C. tol sp.n. is distinct as a dusky, small, member of the Camponotus maculatus ��like species in the Pacific., Published as part of Ronald M. Clouse, Benjamin D. Blanchard, Rebecca Gibson, Ward C. Wheeler & Milan Janda, 2016, Taxonomic updates for some confusing Micronesian species of Camponotus (Hymenoptera: Formicidae: Formicinae), pp. 139-152 in Myrmecological News 23 on pages 149-151, DOI: 10.5281/zenodo.164974, {"references":["CLOUSE, R. M. 2007 b: New ants (Hymenoptera: Formicidae) from Micronesia. - Zootaxa 1475: 1 - 19.","CLOUSE, R. M., JANDA, M., BLANCHARD, B., SHARMA, P., HOFFMANN, B. D., ANDERSEN, A. N., CZEKANSKI- MOIR, J. E., KRUSHELNYCKY, P., RABELING, C., WILSON, E. O., ECONOMO, E. P., SARNAT, E. M., GENERAL, D. M., ALPERT, G. D. & WHEELER, W. C. 2015: Molecular phylogeny of Indo-Pacific carpenter ants (Hymenoptera: Formicidae, Camponotus) reveals waves of dispersal and colonization from diverse source areas. - Cladistics 31: 424 - 437."]}

Metasiro savannahensis sp. nov. (Figs. 3 D���F; 4 B, E, H, K; 6 B, C; 7; 8; 10 C, D, H; 11 C, D; 12 A; 13 B) Metasiro americanus (Davis, 1933), partim Material examined. Males (n= 62), females (n= 72), and juveniles (n= 51), with the following collection details: Jasper County, South Carolina, USA, at Kingfisher Pond, Savannah National Wildlife Refuge, 300 m south of parking lot (lat. 32.18923, long. - 81.08008, elev. 3 m), leg. P. Sharma and R. Clouse, 16 March 2010. Holotype, male, specimen IZ- 133799 (105645)- 7184. Paratypes, from collections IZ- 133799 ���807, 6 males (SPM 007215, 7230, 7240, 7266, 7295, 7315) and 7 females (SPM 007191, 7222, 7234, 7252, 7275, 7301, 7322); two males dissected for genitalia (SPM 007185 and 7335), SPM 007185 disarticulated for appendage measurements and SEMs, and SPM 7214 and 7221 ��� 3 mounted for dorsal and ventral SEMs. A total of 85 specimens were sequenced for molecular analysis. All specimens are deposited in MCZ. Description. Morphology as in the original description of M. americanus (Davis, 1933), with the following recap of characters and additional details. Body and appendage article measurements and proportions available in Tables 5���7. Tuberculate granulations, irregular in shape and spacing, covering entire body (Figs. 3 D���F; 7 B) and legs with the following exceptions: on dorsal prosoma just anterior to ozophores and above anterior tip of coxa I; on irregular medial strip of anal plate of males (being glossy) (Figs. 6 B, C; 7 B); on inner distal femur and patella of leg II (being glossy); on inner trochanter of leg IV (being glossy), and on all tarsi (being generally smooth). Anal plate in males with smooth, raised, medial strip, tapering posteriorly to width of anal gland pore on tergite IX, not reaching anterior edge of anal plate (Figs. 6 B, C; 7 B). Tergite IX distinctly bilobed in males and females (Figs. 6 B, C; 8 B). Gonostome in males oval, with straight posterior edge and a toothlike projection on either side of anterior edge, edges otherwise smooth (Fig. 7 A); gonostome more rounded in females, without large toothlike lateral projections, anterior edge with fringe of small, sharp projections (Fig. 8 A). Setation pattern even over most of body and appendages; especially concentrated on tarsi (Fig. 4 B, E, H, K), these bearing different kinds of setae, including weakly defined solae on tarsus I (Fig. 4 B), and solenidia (thick, curved setae; Willemart & Giribet 2010) on dorsal surface of tarsi I and II (Fig. 4 B, E); hairs nearly absent from chelicerae, especially from second article. Microtrichial formula of spermatopositor: 3 ���4, 6���8, 4+ 4 + 4 (Figs. 10 C, D, H; 11 C, D; 12 A; 13 B); ventral microtrichia at nearly same level (middle slightly more distal) across middle of ventral surface (Fig. 10 C, D, H); apical microtrichia in some cases separated by deep medial cleft (Fig. 11 C). Molecular diagnostic characters. In our COI amino acid fragment (described above) no. 43 is isoleucine (not methionine), no. 80 is threonine (not serine), no. 135 is methionine (not isoleucine), no. 145 is asparagine (not serine or glycine), no. 163 is valine (not isoleucine), and no. 231 is glutamine (not histadine). See Table 3. Distribution and etymology. Known only from the type locality, the Savannah River delta in South Carolina, for which the species is named., Published as part of Clouse, Ronald M. & Wheeler, Ward C., 2014, Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates, pp. 177-201 in Zootaxa 3814 (2) on pages 187-188, DOI: 10.11646/zootaxa.3814.2.2, http://zenodo.org/record/250165, {"references":["Davis, N. W. (1933) A new opilionid from Florida (Arachnida, Cyphophthalmi). Journal of the New York Entomological Society, 41, 49 - 53.","Willemart, R. H. & Giribet, G. (2010) A scanning electron microscopic survey of the cuticle in Cyphophthalmi (Arachnida, Opiliones) with the description of novel sensory and glandular structures. Invertebrate Systematics, 129, 175 - 183. http: // dx. doi. org / 10.1007 / s 00435 - 010 - 0110 - z"]}

Metasiro americanus (Davis, 1933) (Figs. 3 A���C; 4 A, D, G, J; 6 A; 10 E, F, I; 11 E, F; 12 B) Siro americanus Davis, 1933 Parasiro americanus Hinton, 1938 Metasiro americanus Juberthie, 1960 Floridogovea americana Hoffman, 1963 Taxonomic notes: Giribet (2000) considered the synonymization of Metasiro and Floridogovea to have been done by Shear (1980), who noted the latter genus name but did not adopt it. Giribet and Kury (2007) formalized this synonymization and recombined Metasiro out of Sironidae into Neogoveidae. Shear (1980) examined the Sassafras Mt. collections by S. Peck and A. Fiske, and determined those specimens to be M. americanus, but the same has not been done with the Savannah River specimens, other than informally. Discussions and illustrations of M. americanus morphology can be found in these references, as well as in the original description, and as far as we can determine, they apply equally to all three species. Body and appendage article measurements are in Tables 5���6, respectively, and appendage article relative sizes are compared to similar data from Davis (1933) in Table 7. Material examined: Males (n= 112), females (n= 131), and juveniles (n= 74) collected at the type locality of Torreya State Park in the Florida panhandle (IZ- 133791 ��� 2, IZ- 133796 ��� 8, IZ- 134557 ��� 8), nearby localities east of the Apalachicola River (IZ- 133793 ��� 5, IZ- 134492 ��� 3), and at Florida Caverns State Park (IZ- 133808 ��� 13). Two specimens from collection IZ- 133812 were dissected for genitalia (SPM 007418 and 7420), the latter of which was disarticulated for appendage measurements and SEMs. A total of 124 specimens covering all localities were sequenced for molecular analysis. Diagnostic molecular characters: In our COI amino acid fragment (described above) no. 53 is isoleucine (not valine), no. 115 is methionine (rarely threonine; not leucine), and no. 248 is valine or leucine (not isoleucine). See Table 3. M. americanus M. savannahensis M. sassafrasensis sp. nov. sp. nov. male male male male male male female male male female holotype holotype paratype paratype paratype holotype paratype paratype M. americanus M. savannahensis M. sassafrasensis sp. nov. sp. nov. male male male male - holotype paratype paratype 7420 (from Davis, 7185 7176 range size 1933) (lengths) Chelicer I (whole) 0.6 / 2.9 0.7 / 3.3 0.7 / 3.3 0.1 I (from crest) 0.4 / 1.9 0.4 / 1.9 0.4 / 2.0 0 II 0.8 / 5.0 0.8 / 5.0 0.9 / 5.6 0.1 III 0.2 / 5.7 0.2 / 4.9 0.2 / 4.1 0 Total length 1.6 1.4 1.7 1.8 0.4 Palp Trochanter 0.3 / 3.6 0.2 / 2.9 0.3 / 3.4 0.1 II 0.4 / 5.0 0.4 / 5.0 0.4 / 5.7 0 III 0.2 / 3.4 0.2 / 3.0 0.3 / 3.2 0.1 IV 0.3 / 4.0 0.3 / 4.0 0.3 / 3.9 0 ......continued on the next page, Published as part of Clouse, Ronald M. & Wheeler, Ward C., 2014, Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates, pp. 177-201 in Zootaxa 3814 (2) on pages 184-186, DOI: 10.11646/zootaxa.3814.2.2, http://zenodo.org/record/250165, {"references":["Davis, N. W. (1933) A new opilionid from Florida (Arachnida, Cyphophthalmi). Journal of the New York Entomological Society, 41, 49 - 53.","Hinton, H. E. (1938) A key to the genera of the suborder Cyphophthalmi, with a description and figures of Neogovea immsi, gen. et sp. n. (Arachnida, Opiliones). Annals and Magazine of Natural History, Series 11, 2, 331 - 338.","Juberthie, C. (1960) Contribution a l'etude des opilions cyphophthalmes: description de Metasiro gen. n. Bulletin du Museum national d'histoire naturelle, 32, 235 - 241.","Hoffman, R. L. (1963) A new phalangid of the genus Siro from eastern United States, and taxonomic notes on other American sironids (Arach., Opiliones). Senckenbergiana Biologica, 44, 129 - 139.","Giribet, G. (2000) Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). Revista Iberica de Aracnologia, 2, 49 - 76.","Shear, W. A. (1980) A review of the Cyphophthalmi of the United States and Mexico, with a proposed reclassification of the suborder (Arachnida, Opiliones). American Museum Novitates, 2705, 1 - 34.","Giribet, G. & Kury, A. B. (2007) Taxonomy. In: Pinto-da-Rocha, R., Machado, G. & Giribet, G. (Eds.), Harvestmen: the biology of Opiliones. Harvard University Press, Cambridge, MA, pp. 88 - 246."]}

Clouse, Ronald M., Wheeler, Ward C. (2014): Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates. Zootaxa 3814 (2): 177-201, DOI: http://dx.doi.org/10.11646/zootaxa.3814.2.2

Metasiro sassafrasensis sp. nov. (Figs. 3 G���I; 4 C, F, I, L; 5; 6 D; 9; 10 A, B, G; 11 A, B; 13 A) Metasiro americanus (Davis, 1933), partim Taxonomic notes. Original 1969 collections by Peck and Fiske examined by Shear (1980) and reported as M. americanus. Material examined. Males (n= 8), females (n= 7), and juveniles (n= 1), with the following collection details: Pickens County, South Carolina, USA, at Jocosse Gorge, Sassafras Mt., off F. van Clayton Highway (lat. 35.06228, long. -82.79500, elev. 760 m), over ridge in old forest, around rotting log, leg. P. Sharma and R. Clouse, 15 March 2010. Holotype, male, specimen IZ- 134535 (105644 - 7171). Paratypes, from collections IZ- 134534 ���5, 7 males (SPM 007168 and 7172 ��� 7), 7 females (SPM 007169��� 70 and 7178 ��� 82); male specimens SPM 007176 and 7177 dissected for genitalia, and SPM 007176 disarticulated for appendage measurements. Twelve specimens were sequenced for molecular analysis. All specimens are deposited in MCZ. Description. Morphology as in the original description of M. americanus and of M. savannahensis sp. nov. (see above; leg characters of M. sassafrasensis sp. nov. illustrated in Fig. 5; chelicer, palp, claw, and solenidia of M. sassafrasensis sp. nov. illustrated in Fig. 9). Microtrichial formula of spermatopositor: 3, 6, 4 + 4 + 4 (Figs. 10 A, B, G; 11 A, B; 13 A). Body and appendage article measurements and proportions available in Tables 5���7. Molecular diagnostic characters. In our COI amino acid fragment (described above) no. 87 is valine (not leucine). See Table 3. Distribution and etymology. Known only from the type locality, Sassafras Mountain, South Carolina, for which the species is named. M. americanus M. savannahensis M. sassafrasensis sp. nov. sp. nov. male male male male holotype paratype paratype 7420 (from Davis, 1933) 7185 7176 range size (all) Chelicer Article I 0.7 0.8 0.8 0.8 0.1 (whole) Article III 0.3 0.3 0.3 0.2 0.1 Palp Patella 0.7 0.7 0.6 0.7 0.1 Tibia 0.8 0.9 0.8 0.8 0.1 Tarsus 0.8 0.7 0.8 0.8 0.1 Leg I Patella 0.6 0.6 0.5 0.5 0.1 Tibia 0.7 0.6 0.6 0.7 0.1 Metatarsus 0.6 0.6 0.5 0.5 0.1 Tarsus 0.7 0.7 0.7 0.7 0.0 Leg II Patella 0.6 0.5 0.5 0.5 0.1 Tibia 0.6 0.6 0.6 0.7 0.1 Metatarsus 0.5 0.5 0.5 0.5 0 Tarsus 0.8 0.6 0.7 0.7 0.2 Leg III Patella 0.6 0.6 0.7 0.5 0.2 Tibia 0.7 0.8 0.9 0.7 0.2 Metatarsus 0.7 0.8 0.7 0.6 0.2 Tarsus 0.9 0.8 1.0 0.8 0.2 Leg IV Patella 0.6 0.6 0.6 0.5 0.1 Tibia 0.7 0.7 0.7 0.7 0 Metatarsus 0.6 0.7 0.6 0.5 0.2 Tarsus 0.7 0.7 0.8 0.7 0.1, Published as part of Clouse, Ronald M. & Wheeler, Ward C., 2014, Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates, pp. 177-201 in Zootaxa 3814 (2) on pages 189-190, DOI: 10.11646/zootaxa.3814.2.2, http://zenodo.org/record/250165, {"references":["Davis, N. W. (1933) A new opilionid from Florida (Arachnida, Cyphophthalmi). Journal of the New York Entomological Society, 41, 49 - 53.","Shear, W. A. (1980) A review of the Cyphophthalmi of the United States and Mexico, with a proposed reclassification of the suborder (Arachnida, Opiliones). American Museum Novitates, 2705, 1 - 34."]}

Chiasmocleis supercilialba Morales and McDiarmid, 2009 Figure 35, plate 8 Chiasmocleis supercilialbus (Morales and McDiarmid, 2009). Incorrect spelling in the original description (see Peloso et al., 2013). Syncope supercilialbus (de Sá et al., 2012). HOLOTYPE: MHNSM 16174; examined solely from a photograph, in life (not shown). TYPE LOCALITY: Pakitza, Reserve Zone, Manu National Park, ca. 57 km northwestern from the mouth of Rio Manu (11 ° 569470S / 71 ° 179000W), on Rio Manu, Departamento Madre de Dios, Peru. DIAGNOSIS: A medium-sized species for the genus; SVL in males 18.4–18.7 mm (Morales and McDiarmid, 2009). Females unknown. Body ovoid to elongate; head triangular, snout rounded in dorsal and lateral views. Four distinctive fingers; FI well developed, subarticular tubercle may be absent or present; subarticular tubercles present on all remaining fingers, sometimes hardly visible; adpressed FI never extends past the distal margin of subarticular tubercle of FII; adpressed FIV does not reach distal tubercle of FIII; palmar tubercles present, not divided; relative finger lengths I,II, IV,III. Five distinctive toes present, first may be much reduced; toes may be slightly fringed; toes not webbed; TI lacks tubercle; adpressed TI does not touch subarticular tubercle of TII; adpressed TV does not touch middle subarticular tubercle of TIV; TII–IV with terminal discs, usually more developed in females, but also present in males; relative toe lengths I,II,V,III,IV. An inguinal blotch of varied shape is always present. Venter (belly and under surfaces of thighs) usually white or cream with black or dark brown irregular spots. VARIATION: Some variation in color pattern was observed from photos of live specimens (pl. 8) and examination of a paratype (USNM 342862: fig. 35). The dorsum can be dark brown, to almost uniformly whitish (silver) and may show reddish blotches and white spots (when not entirely whitish). A wide white stripe is almost invariably present from the snout to past the eyelids, sometimes extending to the inguinal region (forming the split stripe). A very narrow dorsal white line commonly present, extending from the occipital region to the cloacal region. Dorsal surfaces of forearm yellowish, orange or red, while dorsal surfaces of hind limbs usually follow the general dorsal pattern. As reported by Morales and McDiarmid (2009) the shape and number of inguinal and lateral blotches (or spots) is variable, but commonly dark brown. Throat region with intense brown reticulation over a white (or cream-colored) background. Chest, belly, and undersurfaces of limbs with large, dark brown spots or irregular blotches (fig. 35, pl. 8D, F, H). CALL AND TADPOLE: The advertisement call of C. supercilialba was described by Morales and McDiarmid (2009). Call consists of a fast series of multipulsed notes (mean note duration 31.0 ms; mean interval between notes 37.0 ms) with dominant frequency between 2985.8–3205.0 Hz. Number of pulses per note not given. Tadpoles are unknown. REMARKS: A population from Rio Formoso, state of Rondônia, Brazil (OMNH 37204–37206, 37308–37309, 37319), present dermal spines on dorsum, upper lips, cloacal region, fingers, and toes. The specimens also show well-developed fringes on the fingers and toes, and the toes are basally webbed. These specimens, therefore, agree (in part) with the diagnosis of Chiasmocleis supercilialba (see Morales and McDiarmid 2009, and text above). We tentatively assign the Rio Formoso specimens to C. supercilialba. DISTRIBUTION (fig. 36): Southern Peru (Madre de Dios) and western Brazil (Acre and Rondônia)., Published as part of Peloso, Pedro L. V., Sturaro, Marcelo José, Forlani, Mauricio C., Gaucher, Philippe, Motta, Ana Paula & Wheeler, Ward C., 2014, Phylogeny, Taxonomic Revision, And Character Evolution Of The Genera Chiasmocleis And Syncope (Anura, Microhylidae) In Amazonia, With Descriptions Of Three New Species, pp. 1 in Bulletin of the American Museum of Natural History 2014 (386) on pages 55-57, DOI: 10.1206/834.1, http://zenodo.org/record/5378601, {"references":["Morales, V. R., and R. W. McDiarmid. 2009. A new species of Chiasmocleis (Anura: Microhylidae) from southern Amazonian Peru with comments on some other michrohylids. Biotempo 9: 71 - 76.","Peloso, P. L. V., R. W. McDiarmid, and U. Caramaschi. 2013. The gender of '' cleis '': correct spelling of Chiasmocleis supercilialbus Morales and McDiarmid, 2009, and Elachistocleis magnus Toledo, 2010 (Anura, Microhylidae). Zootaxa 3718: 297 - 298.","de Sa, R. O., et al. 2012. Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology 12: 1 - 21."]}

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."]}

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 Pendergrast��s 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 Pendergrast��s organ present in females (441)., 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 969, DOI: 10.1111/j.1096-0031.2008.00224.x, http://zenodo.org/record/3968591, {"references":["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., 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.","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.","Schaefer, C. W., 1993 a. Notes on the morphology and family relationships of Lestoniidae (Hemiptera: Heteroptera). Proc. Entomol. Soc. Wash. 95, 453 - 456.","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., 2000. The disc-like organ of the Lestoniidae (Heteroptera, Pentatomoidea), with remarks on the relationship of Lestoniidae. Insect Syst. Evol. 31, 20 - 208.","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.","McDonald, F. J. D., 1969. A new species of Lestoniidae (Hemiptera). Pac. Insects 11, 187 - 190."]}

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."]}

Aradoidea Sweet (1996, 2006) argued that the Aradidae should be treated as a separate infraorder, Aradimorpha (originally Aradomorpha), because members of the group ������have a more plesiomorphous abdominal structure, than the Leptopodomorpha + Pentatomomorpha, and possess unique apomorphies������ (Sweet, 1996). This viewpoint has not been widely adopted in the literature on heteropteran classification, possibly because only a few workers have devoted effort to dealing with this particular subject during the intervening period (e.g. Cassis and Gross, 2002). Our evidence offers no support for Sweet��s (1996) theory, but rather corroborates the theory that the Aradoidea belong to the Geocorisae. As is the case for the Cimicomorpha, our sample for the Aradidae is small and does not offer strong evidence for placement of the Aradoidea within the Geocorisae. It is nonetheless largely concordant with that of Wheeler et al. (1993) in placing the Aradoidea as the sister group of the Trichophora. Astronger test of this theory will require analysis of a much larger taxon sample for the Aradoidea in conjunction with a broader sample from the Geocorisae and Leptopodomorpha., 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":["Sweet, M. H., 1996. Comparative external anatomy of the pregenital abdomen of the Hemiptera. In: Schaefer, C. W. (Ed.), Studies on Hemipteran Phylogeny. Thomas Say Publications in Entomology, Entomological Society of America, Lanham, MD, USA.","Sweet, M. H., 2006. Justification for the Aradimorpha as an infraorder of the suborder Heteroptera (Hemiptera, Prosorrhyncha) with special reference to the pregenital abdominal structure. In: Rabitsch, W. (Ed.): Hug the Bug-For Love of True Bugs. Festschrift zum 70 Geburtstag von Ernst Heiss., Denisia 19, 225 - 248.","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.","Wheeler, W. C., Schuh, R. T., Bang, R., 1993. Cladistic relationships among higher groups of Heteroptera: congruence between morphological and molecular data sets. Entomol. Scand. 24, 121 - 137."]}

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 ��tys (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., 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."]}

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)., 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":["China, W. E., Slater, J. A., 1956. A new subfamily of Urostylidae from Borneo. (Hemiptera: Heteroptera). Pac. Sci. 10, 410 - 414.","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.","Schaefer, C. W., Ashlock, P. D., 1970. A new genus and a new species of Saileriolinae (Hemiptera: Urostylididae). Pac. Insects 12, 629 - 639."]}

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.

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.

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