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Zootaxa 4044 (2): 224 240 www.mapress.com/zootaxa/ Copyright 2015 Magnolia Press Article http://dx.doi.org/10.11646/zootaxa.4044.2.3 http://zoobank.org/urn:lsid:zoobank.org:pub:592629b0-8c14-44cc-b243-ac2ac56ea86f ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) New records of Anthalona acuta Van Damme, Sinev & Dumont 2011 and Anthalona brandorffi (Sinev & Hollwedel, 2002) in Brazil, with description of a new species of the simplex-branch (Crustacea: Cladocera: Chydoridae) FRANCISCO DIOGO R. SOUSA 1,2,6, LOURDES M. A. ELMOOR-LOUREIRO 2, JOSÉ ROBERTO DEBASTIANI-JÚNIOR 3, RICCARDO MUGNAI 4 & ANDRÉ SENNA 5 1 Núcleo de Estudos em Biodiversidade Aquática, Programa de Pós-graduação em Biodiversidade Animal, Universidade Federal de Santa Maria UFSM, Av. Roraima 1000, Camobi, CEP 97105-900, Santa Maria, RS, Brazil 2 Laboratório de Biodiversidade Aquática, Universidade Católica de Brasília UCB, QS7 lote 1, Bloco M, sala 204, CEP 71966-700, Taguatinga, DF, Brazil 3 Departamento de Zoologia, Universidade Estadual Paulista UNESP, Rubião Júnior s/n, CEP 18618-970, Botucatu, São Paulo, Brazil 4 Laboratório de Aracnologia, Departamento de Invertebrados, Museu Nacional/Universidade Federal do Rio de Janeiro, Quinta da Boa Vista - São Cristóvão, Rio de Janeiro - RJ, 20940-040 5 Universidade Estado do Rio de Janeiro, Faculdade de Formação de Professores, Departamento de Ciências, Rua Francisco Portela, 1470, São Gonçalo RJ, 24435-005 6 Corresponding author. E-mail: sousa_bio@yahoo.com.br Abstract The range of geographical distribution of Anthalona acuta Van Damme, Sinev & Dumont 2011 and Anthalona brandorffi (Sinev & Hollwedel, 2002) in Brazil has increased by almost 2000 km to the south. New records of Anthalona verrucosa verrucosa (Sars, 1901) were also added. Populations of Anthalona brandorffi from Central Brazil showed a peculiar morphological variation, with some individuals having only a single denticle on the labral keel. A new species of the simplexbranch, Anthalona neotropica sp. nov., was described based on Brazilian material, and this is the first taxon of this branch registered in the Neotropics. It differs from Anthalona simplex Van Damme, Sinev & Dumont 2011, a Central African species, in the morphology of underneath sack of the lateral head pores, length of IDL setae and armature of first flamingtorch seta of limb IV. It could be distinguished from Anthalona sanoamuangae Sinev & Kotov, 2012 (distributed through the South- East Asia) by the morphology of the main head pores, length of IDL setae and armature of the pecten of postabdominal claw. Anthalona neotropica sp. nov. seems to have a benthic/hyporheic habit. All studied species have a wide geographical distribution and could be confused with Anthalona verrucosa Sars, 1901, thus at least some if not all previous records of this species on the continent must be revised. Key words: Anthalona verrucosa, Anomopoda, geographical distribution, head pores, terminal claw, taxonomy Introduction The taxonomy of genus Alona Baird, 1843 has passed through many modifications during the last 25 years (Van Damme et al. 2010), especially because of the paraphyletic or even polyphyletic nature of this genus evidenced in molecular and morphological analysis (Sacherová & Hebert 2003; Elmoor-Loureiro 2004; Eliás-Gutiérrez et al. 2008; Van Damme 2010). A solid diagnosis of the true Alona was proposed as an initial step to improve the knowledge of the morphological boundaries and to be used as the reference point for drawing out some natural groups (Van Damme & Dumont 2008a), for example: Phreatalona, which corresponds to the protzi-complex (Van Damme et al. 2009); Coronatella, which corresponds to the rectangula-complex (Van Damme & Dumont 2008b) and Brancelia (Van Damme & Sinev 2011) which corresponds to the hercegovinae-complex. Recent revision of the verrucosa-group, previously belonging to the Alona sensu lato, resulted in the description of the new genus Anthalona Van Damme, Sinev & Dumont, 2011 which is supported by: two main 224 Accepted by M. Alonso: 27 Oct. 2015; published: 18 Nov. 2015

head pores connected and lateral head pores with sacks below them (cosmaria); postabdomen S-shaped with deep concave anal margin, unmerged marginal denticles, lateral fascicles with long distal setulae; IDL with two setae; seta on the exopodite of limb II reduced or absent; limb III with six setae in exopodite of which third is the longest; limb IV with six setae; limb V with four setae on the exopodite and reduced gnathobase (Van Damme et al. 2011). However, the taxonomic history of this species-group is confusing: initially Alona verrucosa was considered a cosmopolitan species with many records in different regions of the world. Due to two main head pores, Smirnov (1971) included this species in the genus Biapertura Smirnov, 1971. Currently, this genus is considered an artificial assemblage (Frey 1987; Sinev 1999) because it bears some species of different lineages, such as Alona affinis (Leydig, 1860), Alona intermedia Sars, 1862, Karualona karua King, 1853 and Alona setigera Brehm, 1931. Now, it is known that the distinct stenothermic Anthalona is very rich and contains members distributed throughout warm zones in Central and South America, Africa, Asia, Europe and Oceania (Alonso 1996; Sinev & Hollwedel 2002; Van Damme et al. 2011; Sinev & Kotov 2012; Chatterjee et al. 2013). There are three known species of the Anthalona in South America: Anthalona acuta Van Damme, Sinev & Dumont, 2011, Anthalona brandorffi Sinev & Hollwedel, 2002, and Anthalona verrucosa verrucosa (Sars, 1901), in which the latter has the highest number of records in Brazil (Elmoor-Loureiro 2010). However, it is presumed that A. acuta and A. brandorffi also have a wide distribution in South America (Van Damme et al. 2011). The aim of this study was to analyze "verrucosa-like" populations found in Brazil, improving the knowledge on the geographical distribution of the genus Anthalona in South America, especially in Brazil. Herein, we also describe a species related to the simplex-branch which was unknown to science until now. Material and methods In this study, we analyzed 62 samples from the collection of Lourdes M. A. Elmoor-Loureiro, which encompassed several Brazilian regions and the additional sampling conducted in Argentina (San Pedro) and Brazil (Rio de Janeiro and Goiás States and in the Distrito Federal). The majority of samples were taken by traditional methods for littoral microcrustaceans, using a plankton net of 60-100 µm mesh size, and posterior preservation in ethanol or formalin. Samples from the hyporheic zone in the Tijuca National Park (Rio de Janeiro) were obtained using PVC mini-piezometers with 2.5 cm in diameter modified in order to allow a stratified sampling at three different depths: 10, 25 and 45 cm. These samples were elutriated in the field, filtered through a net of 68 µm mesh size and preserved in 75% ethanol solution. The selected animals were transferred to slides containing glycerin and dissected under a stereomicroscope. The morphology of appendages and other structures was studied using a phase contrast microscope Olympus BX41. Enumeration of limb setae and other structures proceeding from the epipodite to the gnathobase, without relation to homology, following the recent revision of the verrucosa-group (Van Damme et al. 2011). Drawings were prepared using a camera lucida attached to Olympus BX41 microscope. Two specimens of Anthalona neotropica sp. nov. were dehydrated in an alcohol series (50%, 70%, 90%, 95% and 100%) and dried using HMDS (Hexamethyldisilazane), then mounted on aluminum stubs, coated with platinum and examined under a JEOL-JSM 7001F scanning electron microscope. Abbreviations. Collections. CLLA: Slides collection of the GEEA, at Universidade Católica de Brasília, Brazil. MZUSP: Museu de Zoologia da Universidade de São Paulo, Brazil. EL: Personal collection of Lourdes M. A. Elmoor-Loureiro. FDRS: Personal collection of Francisco Diogo R. Sousa. In the illustrations and text. en: endite; ep: epipodite; ex: exopodite; fc: filter comb; gfp: gnathobasic filter plate; IDL: inner distal lobe; il; inner lobe; ODL: outer distal lobe; pep: pre-epipodite; s: sensillum; PA: postabdomen; PP: postpore distance (distance between posterior main head pore and posterior border of head shield); P1: limb I; P2: limb II; P4: limb IV. GEEA Research Group on Aquatic Environments, Universidade Católica de Brasília, Brazil. Results Taxonomy NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 225

Class Branchiopoda Letreille, 1817 Order Anomopoda Sars, 1865 Family Chydoridae Dybowsky & Grochowski, 1894 emend. Frey, 1967 Subfamily Aloninae Dybowsky & Grochowski, 1894 emend. Frey, 1967 Genus Anthalona Van Damme, Sinev & Dumont, 2011 Anthalona acuta Van Damme, Sinev & Dumont, 2011 (Figs 1, 6A) Alona verrucosa in Elmoor-Loureiro (2007), p. 347 348, figs. 8 9. Anthalona verrucosa (partim); Sousa & Elmoor-Loureiro (2012), p. 356. FIGURE 1. Anthalona acuta Van Damme, Sinev & Dumont, 2011, parthenogenetic female from Bonita Pond, Distrito Federal, Brazil. A, lateral view. B,h pores. C, antenna II apical segment of the exopodite. D, postabdomen. Scale bars = 50 µm. 226 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

Type locality. Temporary Dune pool near Atins, Lençóis Maranheses, NE Brazil (Van Damme et al. 2011). Holotype. Royal Belgian Institute for Natural Sciences, Brussels (RBIN) under the accession number IG 31782, INV 96691-93. Material examined. Distrito Federal. Fifteen parthenogenetic females from Bonita Pond (15 35'22.1"S, 47 41'50.1 W), Ecological Station of Águas Emendadas, Planaltina, Distrito Federal, Brazil, collected by Lourdes M. A. Elmoor-Loureiro (LMAEL) (EL00743, EL02506) and GEEA. Goiás. A parthenogenetic female from a pond at Paranã River valley (15 09 15.8"S, 47 28 04.7"W), Formosa, Goiás, Brazil, collected by LMAEL on 19.iv.2003 (EL02507). Two parthenogenetic females and one ephippial female from Jataí swamp at Paranã River valley (15 09 15.8"S, 47 28 04.7"W), Formosa, Goiás, Brazil, collected by Ciro Joko on 20.viii.2003 (EL02508) and LMAEL on 19.09.2004. One parthenogenetic female from a swamp at Paranã River valley (14 32 03.0"S, 47 01 50.1"W), Flores de Goiás, Goiás, Brazil, collected by LMAEL on 07.02.2005 (EL02509). One parthenogenetic female from a pond at Chapada dos Veadeiros National Park (14 6'8.57" S, 47 42'7.35"W), Alto Paraíso, Goiás, Brazil, collected by GEEA on 17.06.2012. Mato Grosso. A parthenogenetic female from the vicinity of Poconé (16 21'36"S, 56 38'48"W), Mato Grosso, Brazil, collected by LMAEL on 02.02.1986 (EL02510). Comments. Morphology of the specimens in these populations completely agrees with the initial description of the species (Van Damme et al. 2011). The morphology of A. acuta is very peculiar, and this taxon is easily differentiated from other species of Anthalona due to the strong and chitinized apical setae on the exopodite of the antenna II (Figs 1A, C). Distribution and Ecology. So far, A. acuta has been found from Venezuela to Central Brazil (Fig 6A). Our results confirm the suggestion of Van Damme et al. (2011) that A. acuta is widely distributed in the Neotropics. Anthalona acuta may be confused with A. verrucosa; thus, records of the latter from the south of Brazil need to be checked (for morphology of A. verrucosa, see Van Damme et al. 2011). Anthalona acuta seems to have a preference for lentic water bodies colonized by macrophytes. It was observed in the vegetation formed by plants from the families Cyperaceae Juss. and Pontederiaceae Kunth in coexistence with A. verrucosa verrucosa and A. brandorffi. Anthalona brandorffi (Sinev & Hollwedel, 2002) (Figs 2, 6B) Alona verrucosa (partim) in Sousa & Elmoor-Loureiro (2008); Sousa et al. (2009), p. 103. Anthalona verrucosa (partim) in Sousa & Elmoor-Loureiro (2012), p. 356; Sousa et al. (2013), p. 226. Type locality. Small water body beside the road BR 174 Boa Vista Caracaraí, 43 km southwest of Boa Vista, Brazil (Sinev & Hollwedel 2002). Holotype. Instituto National de Pesquisas da Amazônia, Manaus, Brazil, under accession number INPA 808. Material examined. Distrito Federal. Two parthenogenetic females from Bonita pond (15 35'22.1" S, 47 41'50.1 W), Ecological Station of Águas Emendadas, Planaltina, Distrito Federal, Brazil, collected by GEEA on 27.05.2008. Fourteen parthenogenetic females and one ephippial female from Henrique pond (15 41'16.5"S, 47 56'22.2"W), Brasília National Park, Distrito Federal, Brazil, collected by LMAEL (EL00723, EL01716) and GEEA. Goiás. Two parthenogenetic females from Capivaras pond (18 16.245'S, 52 50.529'W), Emas National Park, Mineiros, Goiás, Brazil, collected by Guilherme Miranda on 09.12.2000, (EL00728). Two parthenogenetic females from Feia Pond (15 34'25.05"S, 47 18'9.00"W), Instruction Field of the Brazilian Army, Formosa, Goiás, Brazil, collected by LMAEL on 12.12.2009 (EL02511). Thirty parthenogenetic females from Cabloca II pond (15º48'22.6''S, 47º14'10.6''W), Instruction Field of the Brazilian Army, Formosa, Goiás, Brazil, collected by GEEA on December/2009. Two parthenogenetic females from Estiva stream (14 06'40,3"S, 47 44'02,2"W), Chapada dos Veadeiros National Park, Alto Paraíso, Goiás, Brazil, collected by GEEA on 13.03.2013, (EL02377). One parthenogenetic female from a pond at Chapada dos Veadeiros National Park (14 04'26,5"S, 47 39'40,7"W), Alto Paraíso, Goiás, Brazil, collected by GEEA on 14.03.2013 (EL2512). Ceará. One parthenogenetic female from a lagoon at Pecém (3 33 22 S, 38 49 26 W), São Gonçalo do Amarante, Ceará, Brazil, collected by Maria Beatriz Gomez e Souza on 18.12.2007. NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 227

FIGUR 2. Anthalona brandorffi (Sinev & Hollwedel, 2002), parthenogenetic female from Bonita Pond, Distrito Federal, Brazil. A, lateral view. B, lateral head pore. C, labral keel. D, postabdomen. E, limb I, IDL. F, limb II, scraper 1 and exopodite. G, limb II, scraper 7 8 and gnathobase. Abbreviations in Material and Methods. Scale bars = 50 µm. Comments. One short seta on the exopodite of limb II (Fig 2F). Morphology of the IDL setae (Fig 2E), number of setae on the filter comb of limb II (Fig 2G), and morphology of scraper 8 (Fig 2G), as observed by Van Damme et al. (2011). However, the anal margin of the postabdomen is concave, not straight (Fig 2D), and the seventh scraper on limb II has no denticles (Fig 2G). Anthalona brandorffi has also a variation on the labral keel and spinulae at the base of the postabdominal claw; some individuals of the populations from Central Brazil had a small denticle on the labral keel (Fig 2C) as observed in A. acuta and A. verrucosa (Van Damme et al. 2011). A spinule on the base of the postabdominal claw longer than the basal spine was observed in the studied populations (Fig 2D), which had not been previously observed in any other species of Anthalona (see Van Damme et al. 2011; Sinev & Kotov 2012). Anthalona brandorffi may be confused with A. verrucosa, but differs in carapace setae (in A. brandorffi the posterior setae are long) and postabdomen (A. brandorffi has long lateral spinules on the postanal fascicles). Besides that, scrapers 7 8 are strongly modified and different from those in other species of Anthalona (for morphology of A. verrucosa, see Van Damme et al. 2011). Distribution and ecology. Anthalona brandorffi has a wide geographical distribution in the Neotropics (Fig 6B). The range of the species goes from Central America to Central Brazil. Records of A. verrucosa from southern South America need to be checked, since the two species may have been confused. In Bonita pond we observed A. brandorffi, A. acuta and A. verrucosa coexisting in the macrophytes from the family Cyperaceae Juss. and Pontederiaceae Kunth beds. Thus, A. brandorffi may coexist with other South American species of the genus. 228 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

Anthalona verrucosa verrucosa (Sars, 1901) Type locality. Vicinity of São Paulo, Brazil (Sars 1901). Lectotype. Single parthenogenetic female, Zool. Mus. Oslo, Accession number F12338a, Collection G.O. Sars, São Paulo, Brazil (Van Damme et al. 2011). Material examined. Bahia. A parthenogenetic female from a pond in Roda Velha (12 47'18"S, 45 57'06"W), Bahia, Brazil, collected by LMAEL on 30.06.1985, (EL00719). Nine parthenogenetic females from a pond in Nova Viçosa (17 53'55"S, 39 22'7"W), Bahia, Brazil, collected by LMAEL on 22.01.1991 and 13.03.1995, (EL00720, EL02213). Distrito Federal. A parthenogenetic female from Paranoá Lake (15 46'59"S, 47 50'53"W), Distrito Federal, Brasília, Brazil, collected by LMAEL on 01.01.1984, (EL00742). Twenty-seven parthenogenetic females from Cedro Pond (15 53'50.2"S, 47 56'37.7"W), Distrito Federal, Park Way, Brazil, collected by LMAEL on 03.04.2002; 07.06.2002; 14.01.2006, (EL00721, EL00722, EL00759, EL00760). Two parthenogenetic females from Acampamento Dam (15 44'53.0"S, 47 57'49.1"W), Parque Nacional de Brasília, Distrito Federal, Brazil, collected by LMAEL on 27.05.2002, (EL00724). Four parthenogenetic females from Bonita pond (15 35'22.1"S, 47 41'50.1 W), Ecological Station of Águas Emendadas, Planaltina, Distrito Federal, Brazil, collected by LMAEL on 03.05.2005 (EL00725). A parthenogenetic female from Recanto das Águas Stream (15 55'21.25"S, 47 58'56.43"W), Park Way, Distrito Federal, Brazil, collected by Erivaldo Casado on 15.06.2005, (EL00726). Fourteen parthenogenetic females from a pool at Botanic Horto (15 51'39.48"S, 48 1'46.81"W), Universidade Católica de Brasília, Taguatinga, Distrito Federal, Brazil, collected by LMAEL on 24.04.2013, (EL02385). Goiás. Three parthenogenetic females from Meia-Lua Farm (15 50'10"S, 48 54'28" W), Pirenópolis, Goiás, Brazil, collected by Valéria Barros on 04.06.2000, (EL02231). Two parthenogenetic females from Capivaras pond (18 16.245'S, 52 50.529'W), Emas National Park, Mineiros, Goiás, Brazil, collected by Guilherme Miranda on 09.07.2000 (EL00727). A parthenogenetic female from Macacos River (14 27 42.9"S, 47 00 15.2"W), Paranã River Valley, Flores do Goiás, Goiás, Brazil, collected by LMAEL on 11.04.2003, (EL00729). A parthenogenetic female from a pond in Paranã River valley (15 09 15.8"S, 47 28 04.7"W), Formosa, Goiás, Brazil, collected by LMAEL on 19.04.2003, (EL00730). Thirteen parthenogenetic females from Jataí swamp in Paranã River valley (15 09 15.8"S, 47 28 04.7"W), Formosa, Goiás, Brazil, collected by Ciro Joko on 20.08.2003, (EL00731). Sixteen parthenogenetic females from Jataí swamp in Paranã River valley (15 09 15.8"S, 47 28 04.7"W), Formosa, Goiás, Brazil, collected by LMAEL on 19.09.2004, (EL00724). Ten parthenogenetic females from swamp in Paranã River valley (14 32 03.0"S, 47 01 50.1"W), Flores de Goiás, Goiás, Brazil, collected by LMAEL on 07.02.2005, (EL00732). Two parthenogenetic females from swamp in Paranã River valley (14 32 03.0"S, 47 01 50.1"W), Flores de Goiás, Goiás, Brazil, collected by LMAEL on 07.02.2005, (EL00733). Eleven parthenogenetic females from a pond in Paranã River valley (14 54 56.0"S, 46 57 02.9"W), Flores de Goiás, Goiás, Brazil, collected by LMAEL on 07.02.2005, (EL00733). Eigth parthenogenetic females from Feia Pond (15 34'25.05"S, 47 18'9.00"W), Instruction Field of the Brazilian Army, Formosa, Goiás, Brazil, collected by LMAEL on 12.12.2009, (EL01758). A parthenogenetic female from Grande Pond (15 49'59.29" S, 47 13'55.42"W), Instruction Field of the Brazilian Army, Formosa, Goiás, Brazil, collected by GEEA on 07.08.2009, (EL01751). Five parthenogenetic females, two males and one ephipial female from Pond at Parque Nacional da Chapada dos Veadeiros (14 6'5.84"S, 47 42'4.89"W), Alto Paraíso, Goiás, Brazil, collected by GEEA on 17.12.2012, (EL02238). A parthenogenetic female from Pond at Parque Nacional da Chapada dos Veadeiros (14 6'8.57"S, 47 42'7.35"W), Alto Paraíso, Goiás, Brazil, collected by GEEA on 17.12.2012, (EL02266). Eight parthenogenetic females from Pond at Parque Nacional da Chapada dos Veadeiros (14 6'8.37"S, 47 42'12.32"W), Alto Paraíso, Goiás, Brazil, collected by GEEA on 17.12.2012, (EL02268). Thirty six parthenogenetic females from Sete Lagoas Pond (14 04'26,5"S, 47 39'40,7"W), Parque Nacional da Chapada dos Veadeiros, Alto Paraíso, Goiás, Brazil, collected by GEEA on 18.06.2012 and 14.03.2013, (EL02299, EL02363). Maranhão. Three parthenogenetic females from Azalão Pond (2º39'34"S, 42º39'55"W), Lençóis Maranhenses, Maranhão, Brazil, collected by Odete Rocha on 28.11.1996, (EL01578). Ninety parthenogenetic females from Pond at Tutoia, Caju Island (2º43'38"S, 42º04'44"W), Maranhão, Brazil, collected by Maria do Socorro Ibañez on 01.03.1999, (EL02135). Minas Gerais. Two parthenogenetic females from Jatobá Pond (14 56'13"S, 44 37'35"W), Parque Estadual Veredas do Peruaçú, Januária, Minas Gerais, Brazil, collected by Maria Beatriz Gomez e Sousa on 01.09.2003, (EL01979). Two parthenogenetic females from Vereda do Peruaçú (14 55'58"S, 44 34'49"W), Parque Estadual NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 229

Veredas do Peruaçú, Januária, Minas Gerais, Brazil, collected by Maria Beatriz Gomez e Sousa on 01.09.2003, (EL01970). A parthenogenetic female from Formosa Pond (14 55'58"S, 44 34'49"W), Parque Estadual Veredas do Peruaçú, Januária, Minas Gerais, Brazil, collected by Maria Beatriz Gomez e Sousa on 25.03.2003, (EL01984). Two parthenogenetic females from Preto River (17 55'6.20"S, 43 48'50.36"W), Parque Nacional das Sempre Vivas, Bocaiúva, Minas Gerais, Brazil, collected by Adriana Fernandes on 02.05.2010, (EL02409). Two ephipial females and one male from Carrascão Pond (17 56'34.73"S, 43 49'49.76"W), Parque Nacional das Sempre Vivas, Bocaiúva, Minas Gerais, Brazil, collected by Adriana Fernandes on 05.05.2010, (EL01873). Seven parthenogenetic females from Redonda Pond (17º46'39"S, 43º36'59"W), Parque Nacional das Sempre Vivas, Bocaiúva, Minas Gerais, Brazil, collected by Adriana Fernandes on 06.05.2010, (EL02406). Four parthenogenetic females from Silipe River (17 29 S, 43 21 W), Parque Nacional das Sempre Vivas, Bocaiúva, Minas Gerais, Brazil, collected by GEEA on 07.09.2010, (EL02410). Mato Grosso do Sul. A parthenogenetic female from Garças Pond (22 43'33"S, 53 13'34"W), Boitaporã, Mato Grosso do Sul, Brazil, collected by LMAEL on 26.09.2009, (EL01763). Mato Grosso. Thirteen parthenogenetic females from the vicinity of Poconé (16 21'36"S, 56 38'48"W), Mato Grosso, Brazil, collected by LMAEL on 02.02.1986, (EL00735, EL00736). Nineteen parthenogenetic females from swamp at Santo Antônio do Levenger (15 50'26"S, 56 4'33"W), Mato Grosso, Brazil, collected by LMAEL on 26.07.1988, (EL00737). Pará. Fourteen parthenogenetic females and one male from Serra dos Carajás (6 24'30.3"S, 50 21'4.4"W), Carajás, Pará, Brazil, collected by Reinaldo Luiz Bozzeli on 08.05.2005, (EL00738). Pernambuco. Two parthenogenetic females from Dama at Escada (8 19'30"S, 35 14'21.5"W), Pernambuco, Brazil, collected by LMAEL on 23.12.1989, (EL01574). Paraná. A parthenogenetic female from Pond in Curitiba (25 31'46"S, 49 13'19"W), Paraná, Brazil, collected by LMAEL and Gilmar Perbiche-Neves on 18.02.2008, (EL02129). A parthenogenetic female from Ressaco do Pau Véio (2 44'54"S, 53 15'25"W), Porto Rico, Paraná, Brazil, collected by LMAEL on 26.03.2009, (EL02128). Thirty parthenogenetic females from Ressaco do Leopoldo (22 45'27"S, 53 16'16"W), Porto Rico, Paraná, Brazil, collected by LMAEL on 26.03.2009, (EL02047). Rio de Janeiro. One hundred and thirty parthenogenetic females from Cabiúnas pond (22 18 S, 41 42 W), Macaé, Rio de Janeiro, collected by Paloma Lopes on 01.06. 2004, (EL00739). Santa Catarina. Twenty-five parthenogenetic females from Mapiju farm (27 41'30"S, 48 46'55"W), Imperatriz, Santa Catarina, Brazil, collected by Gilberto Pereira Júnior on 27.09.2001, (EL02185). São Paulo. Seventeen parthenogenetic females from Lagoa do Coqueiral (23 29'22.64"S, 48 37'6.65"W), Angatuba, São Paulo, Brazil, collected by Silvia M. C. Casanova and LMAEL on 22.01.2002 and 30.01.2001, (EL00740, EL02348). Comments. See Van Damme et al. (2011) for descriptions and diagnosis. Distribution and ecology. Anthalona verrucosa verrucosa has a wide distribution range in Brazil (Fig. 6C). This species was found in several types of water bodies such as streams, rivers, dams, reservoirs, lakes and swamps. According to Van Damme et al. (2011), this species may be associated with a lower acidity water but prefers neutral environments. Anthalona neotropica sp. nov. Sousa, Elmoor-Loureiro & Debastiani-Júnior (Figs 3 5, 6C) Anthalona verrucosa (partim) in Sousa & Elmoor-Loureiro (2012), p. 356. Anthalona sp. in Debastiani-Júnior et al. (2015). Etymology. The species name derives from the zoogeographical region where the species is found. Anthalona neotropica sp. nov. is the first species of the simplex-branch described for the Neotropics. Type locality. Sobradinho Stream (15 38'27"S, 47 46'40 W), Distrito Federal, Brazil. Type material. Holotype: undissected adult parthenogenetic female in a tube, preserved in 92% ethanol solution, deposited at the Museum of Zoology of the University of São Paulo under the accession number MZUSP33195. The label of the holotype is: Anthalona neotropica, 1 parth. from to Sobradinho Stream, Distrito Federal, Brazil. Holotype. 230 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

Paratypes: Distrito Federal. Two parthenogenetic females from São Bartolomeu River (15 47'34.7''S, 47 41'28.2''W), Planaltina, Distrito Federal, Brazil, collected by GEEA on 26.08.2006, (CLLA194 195; FDRS0370). Three adult parthenogenetic females from Sobradinho Stream (15 38'27"S, 47 46'40 W), Distrito Federal, Brazil, collected by GEEA on 5.09.2008 and 06.03.2009 (CLLA177 186, 192; FDRS0371). An adult parthenogenetic female from Pipiripau Stream (15 33'45,5'' S, 47 30'39,7''W), Distrito Federal, Brazil, collected by GEEA on 17.03.2009 (FDRS0372). Goiás. A juvenile parthenogenetic female from Cachoeira do Palmito (15 51'8.5"S, 48 57'58.1"W), Pirenópolis, Goiás, Brazil, collected by LMAEL on 01.10.2000 (CLLA193). Rio de Janeiro. Six parthenogenetic females from hyporheic zone in Solidão River (22º57 36.89 S, 43º17 1870 W), Tijuca National Park, Rio de Janeiro, Brazil, collected on March/2012, leg. Ricchardo Mugnai and André Senna (FDRS0363). Ten parthenogenetic females from the hyporheic zone of the Tijuca River (22 57'13.08"S, 43 16'55.45"W), Tijuca National Park, Rio de Janeiro, Brazil, collected on June and November/ 2012 (CLLA187-189; FDRS0365; 0367-0369). Rio Grande do Sul. A parthenogenetic female from Machadinho reservoir (27 32'26.71"S, 51 37'52.31"W), Upper Uruguai River, Machadinho, Rio Grande do Sul, Brazil, collected by Danilo Augusto Naliato on 01.06.2010 (CLLA190-191). Argentina. Two juvenile females from a lateral lake of Lower Paraná River (33 40 49.00ʺS, 59 38 48.80ʺW), San Pedro, Argentina, collected by José Roberto Debastiani-Júnior on 14.04.2010 (FDRS0366). Diagnosis. Female. Of medium size for the genus, length 0.30 0.37 mm, elongated, about 1.6 times as long as high. Head presenting ocellus and eye of similar size; two connected main pores of similar size, underneath sack of lateral head pores lobed (cosmaria). Labral keel without setulae, notches or denticles. Carapace covered by longitudinal lines at posterior region; 31 37 naked setae on ventral valve margin. Antennule 2.5 times longer than wide; aesthetascs exceeding the length of the antennule. Antenna II with antennal formula: spines 001/101; setae 113/003; first segment of the exopod with one cluster of long setulae; a spine on first segment of endopod exceeding the middle-length of the second segment; apical spines of similar length. Postabdomen relatively short, postanal margin rounded with 5 6 denticles; 5 6 lateral fascicles present; first spinule of each fascicle longer and thicker than the others; all fascicles exceeding the postanal margin. Postabdominal claw longer than the anal margin, with a group of at least three long spinules at the base; claw pecten divided in two groups; basal spine armed with spinules along its dorsal margin. Limb I: ODL with a thin seta; IDL (en4) with setae 2 3 of similar length, bisegmented and armed with thin setulae. Limb II: exopodite without seta, scrapers 7 8 with bent apexes. Limb III: exopodite with six setae, setae 1 2 of different length; basal endite armed with a sensillum near the first soft seta. Limb IV: exopodite with six setae, sixth seta markedly shorter than fifth seta, first flaming-torch seta robust. Limb V: exopodite with setae 1 3 long and plumose, seta 4 short, gnathobase armed with a curved element. Description. Parthenogenetic female. Habitus (Figs 3A B, 5A). Animal of medium size for the genus, length 0.30 0.37 mm; elongated, the carapace is oval in lateral view, maximum height near to anterior portion, about 1.6 times as long as high. Dorsal margin convex. Dorsal keel absent. Head (Figs 3A C, 5C D). Ocellus and eye of similar size. Headshield not studied. Rostrum short, rounded, projected towards ventral margin of carapace. Head pores as figured (Figs 3E I, 5D). Two main pores of similar size connected; PP about 0.35 0.5 IP; underneath sacks of lateral head pores lobed (cosmaria). Labrum (Figs 3J, 5E). Morphology as in simplex-branch, keel without setules, notches or denticles; anterior margin convex. Maxilla I (Fig 4A). Well developed, with two setulated setae. Antennule (Fig 3N). Antennular body elongated, about 2.5 times longer than wide, three rows of short setules. Antennular sensory seta slender and short, about 0.4 times smaller than antennular body, inserted at 2/3 length from the base of antennule. Nine aesthetascs of different length; longer aesthetascs projecting beyond the tip of rostrum, none exceeds antennules length. Antenna II (Figs 3O P, 5F). Coxal setae of equal length. Basipod thick, armed with one short spine. First segment of exopod with a cluster of long setulae exceeding the length of segment; second and third segments armed with relatively short and slender setulae in their terminal portion. First segment of endopod naked; second and third segments with short spinulae at the terminal portion; spine on first segment of endopod exceeding the middle-length of the second segment. Antennal formula: spines 001/101, setae 113/003. Setae on first segment of exopod not studied; seta on second segment of exopod bisegmented, long. Apical spines of endopod and exopod NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 231

long, of similar size and longer than apical segments. Apical setae of exopod and endopod bisegmented and of similar size. Carapace (Figs 3D, 5B) covered by longitudinal lines at posterior region; ventral margin with a concavity in the middle; 31 37 not plumose setae per valve; short spines between marginal setae present; long setae in anterior group; long intermediate group contains relatively short setae; posterior group contains long setae. Posteroventral corner armed with spinulae not arranged in groups nor projected beyond the margin. Abdomen about 2.5 times shorter than thorax with at least two rows of abdominal setulae (Figs 3A C; L). Postabdomen (Figs 3K L, 5H I). Relatively short, about 2.5 times as long as wide; ventral margin slightly rounded, with two rows of spinulae. Pre-anal margin longer than anal and postanal margin. Anal margin concave, with evident angles, supplied with 3 4 groups of robust setulae. Postanal margin convex, armed with 5 6 lateral fascicles; first setule of each fascicle longer and thicker than the others; all fascicles exceeding postanal margin; also the margin bears 5 6 short marginal denticles. Postabdominal seta not studied. Postabdominal claw (Figs 3K M, 5G). Implanted at the short projection from the postabdomen. Claw longer than anal margin of postabdomen, uniformly curved, with a group of at least two long spinules on the base; pecten divided in two groups; proximal group armed with longer spines compared to distal group. Basal spine length more than width of claw. Spine armed with spinulae along its dorsal margin. Limb I (Figs 4B D). Epipodite thin, oval, with a relatively long projection. ODL with a thin, distally serrulated seta, about the same length as IDL setae; accessory seta not studied. IDL (en4) with a group of long spinulae on its face, two setae of similar length, bisegmented and armed with thin setulae. Third endite armed with short spinulae and four setae; two posterior setae densely setulated (a b), the third posterior seta (c) and anterior seta 1 shorter. Second endite with three posterior setae (d f), setae (f) and (e) armed with thick spinules on the lateral face; seta (d) armed with short setulae; seta (e) about 1.5 times longer than seta (f); seta (d) about 2.8 times shorter than seta (e). First endite with three marginal posterior setae (g i), two bisegmented and setulated in distal part (g h) and the third seta relatively shorter (i). No sensillae were found on the endites. Ejector hooks of different length armed with short denticles. Ventral face of limb armed with seven groups of setulae organized in clusters, decreasing towards the distal portion; distal clusters of short and strong setulae. Gnathobase elongated, apex as a projection with short setulae. Limb II (Figs 4E H). Exopodite elongated, with two rows of setulae; without seta. Inner limb portion with eight scrapers; scraper 6 8 markedly shorter than others and with bent apex; scraper 7 longer than 6 and 8; scrapers 1 4 armed with fine setulae; scrapers 5 8 armed with thin denticles. Proximal portion of the gnathobase short, wide and densely setulated; distal portion armed with four elements, first element as a small sensillum. Filter comb with seven setae; two posterior setae markedly shorter than other setae. Limb III (Figs 4I K). Epipodite oval, with a short projection. Exopodite relatively large, subquadrangular, with six marginal setae, subdivided into two groups: 2 lateral setae and 4 distal setae. Setae 1 2 of different length, both setulated. Third seta long, setulated, about 5.7 times longer than the sixth seta and 2.3 longer than fourth and fifth setae. Fourth and fifth setae of similar length, relatively long, setulated. Sixth seta short, about 2.4 times shorter than the fifth setae, naked. Distal endite (as initially proposed by Kotov 1999) with three anterior setae (1 3), two scraper-like similar in length (1 2); third seta slightly curved and armed with setulae bilaterally implanted (3); four plumose posterior setae increasing in length towards distal gnathobase (a d). Basal endite with a sensillum and four soft setae (Fig. 4J). Gnathobase armed with three elements, the first being a cylindrical sensillum, the second a geniculate seta, third element naked and with acute tip. Filter comb with seven setae (Fig. 4K). Limb IV (Figs 4L N). Pre-epipodite round, densely setulated. Epipodite oval, with long projection. Exopodite round, with six marginal setae; lateral setae 1 4 plumose; seta 1 slightly longer than seta 2; third seta about same length of the first seta; fourth seta about 1.6 times longer than fifth seta; two distal setae (5 6) short, fifth seta naked, markedly longer than sixth seta; sixth seta naked. Distal endite with four anterior setae (1 4), one scraperlike (1) and three flaming-torch-like (2 4); first flaming-torch seta (2) robust, with long setulae. Basal endite with three posterior soft setae increasing in size towards the base. Gnathobase armed with a sensillum and a setulated seta implanted on a robust base. Filter plate with five slender setae. Limb V (Fig 4O). Pre-epipodite subrectangular and densely setulated. Epipodite oval, with a long projection. Exopodite moderately wide, not divided into lobes and armed with four plumose setae; setae 1 3 similar in length; fourth seta about 3.4 times shorter than third seta. Internal lobe very wide, oval, bearing long setulae; two setulated setae on inner face of this lobe, one about two times longer than the other. Gnathobase setulated, armed with a curved element; a filter comb absent. 232 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

FIGURE 3. Anthalona neotropica sp. nov., parthenogenetic female. A, lateral view, Sobradinho Stream, Distrito Federal, Brazil. B, lateral view, Tijuca River, Rio de Janeiro, Brazil. C, lateral view, juvenile from San Pedro, Argentina. D, carapace, posteroventral corner. E I, head pores. J, labral keel. K, postabdomen, Sobradinho Stream, Distrito Federal, Brazil. L, postabdomen, Tijuca River, Rio de Janeiro, Brazil. M, terminal claw, Sobradinho Stream, Distrito Federal, Brazil. N, antennule, Sobradinho Stream, Distrito Federal, Brazil. O, antenna II, Sobradinho Stream, Distrito Federal, Brazil. P, antenna II, Cachoeira do Palmito, Pirenópolis, Goiás, Brazil. Scale bars = 50 µm. NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 233

FIGURE 4. Anthalona neotropica sp. nov., parthenogenetic female from Sobradinho stream, Distrito Federal, Brazil. A, maxilla. B, limb I. C, limb I, IDL and ODL. D, limb I, IDL and ODL of the juvenil (Argentina). E, limb II. F H, limb II, scapers with apex bent. I, limb III, exopodite. J, limb III, distal endite. K, limb III, basal endite and gnathobase. L, limb IV, exopodite. M, limb IV, distal and basal endites. N, limb IV, gnathobase. O, limb V. Abbreviations in Material and Methods. Scale bars = 50 µm. 234 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

FIGURE 5. Anthalona neotropica sp. nov., parthenogenetic female from Tijuca River, Rio de Janeiro, Brazil. A, lateral view. B, carapace ornamentation. C, head rostrum. D, head pores. E, labral keel. F, antenna. G, postabdomen terminal claw. H, postabdomen postanal margin. I, postabdomen anal margin. Scale bars: A = 50 µm; B I = 10 µm. NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 235

FIGURE 6. Geographical distribution of four species of Anthalona in the Neotropics. A, Anthalona acuta Van Damme, Sinev & Dumont, 2011. B, Anthalona brandorffi (Sinev & Hollwedel, 2002). C, Anthalona verrucosa verrucosa (Sars, 1901). D, Anthalona neotropica sp. nov. Black circles are previous records in Van Damme et al. (2011) and Sinev & Hollwedel (2002). White circles are records observed in this study. 236 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

Male, ephipial female and ephipium. Unknown. Variability. We observed variability in the morphology of lateral head pores (Figs 3E I). In smaller individuals, we observed variability in the spine length of the first segment of endopodite of antenna II (Fig 3P). In juveniles, a variation was observed in the proportion of the IDL setae (Figs 4C D). Differential diagnosis. Anthalona neotropica sp. nov. is a member of the simplex-branch and differs from the other more specialized species in IDL setae armed with fine setulae (Figs 4C D) (see Van Damme et al. 2011; Sinev & Kotov 2012). Anthalona neotropica sp. nov. resembles A. sanoamuangae Sinev & Kotov, 2012 and A. simplex Van Damme, Sinev & Dumont, 2011 but differs from A. sanoamuangae in: (1) carapace setae organized in three groups; (2) spine on the first segment of endopod of antenna II very long and almost reaching the end of the third segment; (3) main head pores of similar size; (4) IDL setae of similar length; (5) setae 5 6 of limb IV with markedly different length and (6) pecten of postabdominal claw organized in two groups. Anthalona neotropica sp. nov. can be differentiated from A. simplex by: (1) long spinulae on the base of terminal claw; (2) spine on the first segment of endopodite of antenna II very long and almost reaching the end of third segment; (3) IDL setae of similar length; (4) setae 5 6 of limb IV with markedly different length and (5) first flaming-torch of limb IV robust. In contrast to other species of the simplex-branch, Anthalona neotropica sp. nov. has the apex of scrapers 6 8 markedly curved (Figs 4E, G H). More similarities and differences are presented in Table 1. TABLE 1. Morphological differences and similarities among species from the simplex-branch of Anthalona. Abbreviations in Material and Methods. Anthalona neotropica sp. nov. Anthalona simplex Size (mm) 0.30 0.37 0.31 0.38 0.30 0.34 Main head pores size similar similar distinct Lateral head pores shape lobed circular lobed Carapace setae n of groups three three two A1 aesthetascs long short long P1 ODL/IDL setae ODL = IDL ODL > IDL ODL < IDL P1 IDL setae, length similar 2 < 3 2 < 3 P2 scrapers 6 8, apex bent sharp sharp P4 exopodite setae 5/6 length 5 > 6 similar similar Anthalona sanoamuangae P4 first flaming torch seta robust thin robust PA spinules on the base of the two, long three, short two, short terminal claw PA anal angles strongly marked slightly marked strongly marked PA pecten on the terminal claw two groups two groups continue Distribution and ecology. Anthalona neotropica sp. nov. was observed only in Brazil and Argentina (Fig 6D). It is possible that it has as wide distribution range, as A. verrucosa verrucosa, A. acuta and A. brandorffi. Anthalona neotropica sp. nov. seems to be present in low density in ponds and other lentic ecosystems. However, it was observed with relatively high abundance in samples collected in the hyporheic zone of lotic ecosystems, at depths between 10 and 25cm. In Central Brazil, A. neotropica sp. nov. was only observed in hyporheic habitats of lotic ecosystems, once few specimens could be removed from the sediments by a water flow. Such as A. simplex and A. sanoamungae, A. neotropica sp. nov. is less specialized than other species from the genus, and probably is a fine sediment dweller, adapted to collect fine materials, such as food. The morphology of IDL setae, presence of a cluster of long setulae on the first segment of exopod of antenna II and the long and thick spine on postabdominal lateral fascicles, confirms its preference for benthic habitat (see Kotov 2006). Notes on the biogeography and relationship between species of the simplex-branch. Anthalona neotropica sp. nov. is a new member of the simplex-branch, which includes also African A. simplex and Asian A. sanoamuangae. The following synapomorphies support the simplex-branch: (1) IDL setae armed with thin setulae; (2) exopodite of limb II without setae and (3) cluster of setulae on the first segment of exopod of the antenna II. In addition, species from the simplex-branch share long lateral spinules on postabdominal lateral fascicles and very NEW RECORDS OF ANTHALONA ACUTA VAN DAMME Zootaxa 4044 (2) 2015 Magnolia Press 237

plumose long setae on limbs IV and V (Van Damme et al. 2011; Sinev & Kotov 2012). The affinities of A. neotropica sp. nov. with other species of the simplex-branch are obvious, i.e.: Anthalona neotropica sp. nov. and A. sanoamuangae have similar morphology of the underneath sacks of lateral head pores; length of aesthetascs of antennule; marked pre- and postanal angles of postabdomen; similar limbs I, III and V and robust flaming-torch on limb IV. However, the main head pores of different sizes is a unique trait of A. sanoamuangae (Sinev & Kotov 2012). Anthalona neotropica sp. nov. and A. simplex have the same morphology of main head pores, armature of ventral setae of carapace, pecten on postabdominal claw organized in two groups and similar limbs I, III and V (Van Damme et al. 2011). In the simplex-branch, circular sacks of lateral head pores and short and thin first flaming-torch seta on limb IV are morphological traits exclusive of A. simplex. It is known that A. sanoamuangae only occur through South-East Asia (Thailand, Vietnam and Laos) and that A. simplex was only recorded from Congo (Africa) (Van Damme et al. 2011; Sinev & Kotov 2012). Thus, the distribution of the species from the simplex-branch point to a tropical range and could suggest a divergence from the disjunction of the zones connected in the past. Usually such pattern is associated with the Gondwana disruption (see Bayly 1995; Fernando et al. 1987; Dumont & Negrea 2002), although some investigators have serious doubts in "Gondwanian" interpretation of such distributional ranges (Korovchinsky 2006). Morphology of A. neotropica sp. nov., A. sanoamuangae and A. simplex is very conservative, without important reductions or specializations (possibly due to association with similar habitat) that can be used as an argument pro close in-group relationships. Identification key for South American species of Anthalona 1. IDL setae 2 3 armed with spines......................................................................... 2 - IDL setae 2 3 armed with thin setulae.................................................... A. neotropica sp. nov. 2. IDL seta 3 armed with one large proximal spine followed by short spines or spinulae................................ 3 - IDL seta 3 armed with 3 4 large spines of similar length....................... A. brandorffi (Sinev & Hollwedel, 2005) 3. Apical setae on the antenna II chitinized and strongly thickened............. A. acuta Van Damme, Sinev & Dumont, 2011 - Apical setae on the antenna II thin............................................ A. verrucosa verrucosa (Sars, 1901) Conclusion Our results confirm the suggestion of Van Damme et al. (2011), indicating that A. acuta and A. brandorffi are species widely distributed in the Neotropics. The range of distribution of A. acuta and A. brandorffi in Brazil was expanded by almost 2000 km to the south. It is probable that the aforementioned species occur in a subtropical portion of South America also. Due to the possibility of confusion with A. verrucosa, records of this species on the continent need to be revised. The South American Chydoridae fauna, especially in Brazil, has recently received a lot of attention from taxonomists, which aided an initial understanding of the true diversity of species (e.g. Sinev and Elmoor-Loureiro, 2010; Van Damme et al. 2011; Elmoor-Loureiro, 2014; Elmoor-Loureiro et al. 2013; Sousa et al. 2015). The description of A. neotropica sp. nov. demonstrates that more studies need to be performed. Although a significant advance in the taxonomic knowledge in Brazil was achieved recently, the fauna of Chydoridae is still inadequately known. Acknowledgements The authors thank Fundo Nacional do Meio Ambiente (FNMA) (Process 02000.005571/2005-89), FAPESP (Process 2011/23444-6), CAPES and FAPERJ for partial financial support. Also, Grupo de Estudos de Ecossistemas Aquáticos (GEEA) for field work assistance. Many thanks to Dr. Adriano Kury from the Museu Nacional/UFRJ for allowing the research activities of RM in the Arachnology Laboratory and Mario J. Gatti, M. Sc., Curator of the Trichocomaceae Mycological Collection IOC/FIOCRUZ for technical support for RM. We thank Dr. Danilo Naliato for donation of material from Rio Grande do Sul. We are deeply grateful to Dr. Alexey A. Kotov and an anonymous reviewer for their valuable criticism, comments and suggestions, which improved this manuscript. The first author of this paper received a scholarship from the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES). 238 Zootaxa 4044 (2) 2015 Magnolia Press SOUSA ET AL.

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