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C. d. vizottoi Taddei and Lim, 2010 Synonyms: Chiroderma nov. sp. Cruz, Martínez, and Fernandes, 2007: 615. Chiroderma sp. Gregorin, Carmignotto, and Percequillo, 2008: 372. Chiroderma vizottoi Taddei and Lim, 2010: 384; type locality “Teresina (ca. 05° 02 ′ S and 42° 45 ′ W), in the State of Piauí, Brazil.” Distribution and Habitat. This subspecies is in the Brazilian states of Ceará, Maranhão, and Piauí (Fig. 23). A specimen from the Caatinga of Piauí was captured in an area with low xerophytic trees (3 to 5 meters high), next to a rocky outcrop (Gregorin et al. 2008). Two specimens from Maranhão were captured in mist nets set on the beach, in a mosaic of sandbank and mangrove vegetation. Remarks. Lund (1842a: 134, 1842b: 200), in his studies on the extant and fossil fauna of Lagoa Santa, Minas Gerais, recorded Phyllostoma dorsale among the living species of the region. In both publications, Lund clearly uses Phyllostoma dorsale as a new name, as evidenced by the “m.” (= mihi) following the name. Because the name is present only in a faunal list, and no formal description is given, Phyllostoma dorsale Lund is considered a nomen nudum for nomenclatural purposes (see article 12 of the ICZN 1999). Gray (1866), in his revision of Phyllostomidae, listed two species in Chiroderma, C. villosum and C. pictum. The latter name is a nomen nudum, because it lacks a description. We include Chiroderma pictum in the synonymy of Chiroderma doriae because, in the Dobson’s (1878) catalogue of bats in the British Museum, the only specimen of Chiroderma in the collection at the time of Gray’s (1866) publication was the specimen that later became the type of Chiroderma doriae. The Latin word pictus means “painted”, or “decorated”, suggesting that Gray referred to the conspicuous facial stripes, observable that specimen, a characteristic differentiating this species from Peters’ C. villosum, which was the only other Chiroderma known at that time. Dobson (1878), in his description of Chiroderma salvini, compared the type with a specimen from Minas Gerais (Brazil) identified by him as Chiroderma villosum. However, Thomas (1891), based on an unpublished plate by Wilhelm Peters, available in the Genova Natural History Museum, recognized that the Minas Gerais specimen Dobson identified as Chiroderma villosum did not represent Chiroderma villosum Peters, 1860. Thomas (1891) described that specimen as Chiroderma doriae. Winge (1892), studying the material sent by Lund to the Copenhagen museum, identified as Chiroderma villosum the material referred to Lund as Phyllostoma dorsale. Winge (1892: 9) located, among the Lagoa Santa specimens representing extant taxa, five fluid-preserved specimens, two skeletons, and three skins. Among the fossil material, Winge mentioned only post-cranial elements from Lapa da Escrivaninha and a cave he did not identify. Thomas (1893), after receiving part of the Lagoa Santa material Winge identified as Chiroderma villosum, correctly identified the skull as belonging to C. doriae. This publication is the first to ascertain that Phyllostoma dorsale of Lund was the same taxon that Thomas (1891) described as Chiroderma doriae. The specimen collected by Lund at Lagoa Santa (BMNH 93.1.9.16) and examined by us is probably the same specimen donated by Winge to the British Museum and mentioned by Thomas (1893). Aside from the Lagoa Santa specimens and the holotype, all from Minas Gerais, no additional specimen of C. doriae were mentioned in the literature for 86 years. Probably for this reason, publications that mentioned the species cited only Minas Gerais as a locality of occurrence and repeated the morphological characteristics mentioned in the original description (Vieira 1942; Cabrera 1958). It is important to note, however, that the BMNH houses specimens of C. doriae collected in the beginning of the 20 th century, by Alphonse Robert in Ypanema, currently Floresta Nacional de Ipanema, São Paulo (BMNH 3.7.1.163), and by Wilhelm Ehrhardt in Joinville, Santa Catarina (BMNH 9.11.19.15). Taddei (1979) published the first study including new data on C. doriae since Thomas’s report in 1893. When he redescribed the taxon based on 39 specimens of C. d. doriae collected in northwestern São Paulo, Taddei (1979) confirmed the presence of a conspicuous dorsal stripe previous authors had considered to be absent. Natural History. C. d. doriae is secondarily granivorous, specialized in chewing and grinding seeds of the fruits of Ficus to extract their nutritional content (Taddei 1980; Nogueira & Peracchi 2001, 2003). Four species of Ficus have been documented in the diet of C. d. doriae: Ficus clusiaefolia, F. cyclophylla, F. organensis, and F. tomentella (Sipinski & Reis 1995; Esbérard et al. 1996; Nogueira & Peracchi 2001). Individuals of C. d. doriae have been captured near to or visiting fruiting trees of Cecropia glaziovii (Urticaceae), Ficus enormis, F. gomeleira, and Muntingia calabura (Muntingiaceae), suggesting that this bat also consumes the fruits of these species (Taddei 1980; Esbérard et al. 1996). In addition to figs, fruits and infructescences of Cecropia pachystachya, Chlorophora tinctoria (Moraceae), Piper sp., and Psychotria suterella (Rubiaceae) are also consumed (Taddei 1973; Esbérard et al. 1996; Nogueira & Peracchi 2001; Novaes et al. 2010; Laurindo et al. 2017). Individuals of C. d. doriae have been found covered in the pollen of Mabea fistulifera (Euphorbiaceae) and of an unidentified species, suggesting that the bat also feeds on flowers (Esbérard et al. 1996; Olmos & Boulhosa 2000). Day roosts used by C. d. doriae are not known (see review in Garbino & Tavares 2018b), but there are two records from caves, the use of which may be occasional: one in the state of São Paulo (Arnone 2008) and the other in Minas Gerais (information on specimen UFMG 4953). In the Cerrado of São Carlos, southeastern Brazil, the species was found in pellets of the Stygian Owl (Asio stygius) by Motta-Junior & Taddei (1992). At localities in southeastern Brazil where they occur in sympatry, C. d. doriae is usually captured in much higher numbers than C. v. villosum when mist nets are set at ground level. At the Rio de Janeiro Botanical Garden, eight nights of ground-level mist netting resulted in the capture of 49 C. d. doriae and 5 C. v. villosum (Nogueira & Peracchi 2003). In Mirassol, São Paulo, 24 C. d. doriae and 6 C. v. villosum were captured in ground-level nets (Taddei 1979). In contrast, Gregorin et al. (2017), using canopy mist nets placed between 11 and 19.5 m above ground in the Parque Estadual do Rio Doce, Minas Gerais, captured 1 C. d. doriae and 24 C. v. villosum. The anecdotical report of Taddei (1980) describes the approach by C. d. doriae to fruiting trees as a low flight. Although the literature on the ecology of C. d. doriae is scarce when compared to that for C. gorgasi, C. trinitatum, and C. villosum, it suggests that the species is not as much a canopy frugivore as the other species. Five species of ectoparasites are known from C. d. doriae: Aspidoptera falcata (Streblidae), Megistopoda proxima (Streblidae), Strebla guajiro (Streblidae), and Trichobius joblingi (Streblidae) in Sergipe; and the mite Periglischrus iheringi (Spinturnicidae) recorded on individuals from Mato Grosso do Sul, Rio de Janeiro, and in Paraguay (Presley 2004; Lourenço et al. 2016; Lima Silva et al. 2017; Soares et al. 2017). Lourenço et al. (2018) found a trypanosomatid parasite in the blood of C. d. doriae from Distrito Federal, Brazil. The reproductive pattern is seasonal polyestry (Taddei 1980). Pregnancies were recorded in June through October, with earlier stages reported in June and July, and females with advanced fetuses recorded from August through October (Taddei 1980; Esbérard et al. 1996). Births of single young occur between October and January, suggesting a gestation period of at least three and half months (Taddei 1980). Females simultaneously pregnant and lactating are common, indicating postpartum estrus and the production of two young per year, with a second birth period in February and March (Taddei 1976, 1980; Esbérard et al. 1996). The natural history of C. d. vizottoi is poorly known. In Ceará, seven specimens were mist netted close to an unidentified fruiting Moraceae; one was pregnant and another lactating when captured in January (Silva et al. 2015). In Maranhão, a male and female were caught in a mist net placed along the beach in a small fishing community next to a mangrove swamp near regenerating rain forest and a “babaçual” (Attalea speciosa [Arecaceae]) palm grove. The mist-net was set in front of a tree locally called “agarra” or “amapá” (Apocynaceae?), and one of the bats had its abdomen covered with a sticky material resembling a milky sap, or latex. Almeida et al. (2016) recorded the mite Periglischrus iheringi (Spinturnicidae) on C. d. vizottoi in Ceará. Specimens Examined (N = 136): Brazil: Alagoas, Mata da Cachoeira, São José da Lage (UFPB 4348); Bahia, Reserva Particular do Patrimônio Natural Serra Bonita ([UFMG] RSB 21, [UFMG] RSB 22); Ceará, Reserva Natural Serra das Almas (ALP 10196, 10418, 10421, 10423, 10440, 10451, 10464); Goiás, Itumbiara (MCN-MQ 145); Maranhão, São Luís (ALP 6633–6635, [UFMG] VCT 373, 374); Mato Grosso do Sul, Corumbá ([UFMG] VCT 6069, [UFMG] VCT 6081), Fazenda Barma (MZUSP 28591, 28688), Morro do Paxixi (ZUFMS 2300), Parque Estadual Matas do Segredo (ZUFMS 493), Urucum (ZUFMS 800, 1058–1062, 1069), Urucum, Morro São Domingos (ZUFMS 912); Minas Gerais, without specific locality (BMNH 44.9.2.6 [holotype of doriae]), Área de Proteção Ambiental Coqueiral (CMUFLA 163), Barão de Cocais ([UFMG] VCT 5831), Belo Horizonte (UFMG 3537), Estação Ecológica de Pirapitinga in Morada Nova de Minas (UFMG 3383), Estação Ecológica de Pirapitinga in Três Marias (ALP 9154), Gruta do Salitre in Diamantina (UFMG 4953), João Monlevade (CMUFLA 965), Lagoa Santa (BMNH 93.1.9.16); Mariana (CMUFLA 938, [UFMG] VCT 6058), Parque Estadual do Rio Doce (CMUFLA 1157), Pompéu (MCN-MQ 253), Usina Hidrelétrica Retiro Baixo (CMUFLA 1415), Valos (CMFULA 496); Pará (not mapped; BMNH 7.1.1.723 [probably an error]); Paraíba, Fazenda Pacatuba, 10 km NE Sapé (UFPB 3), Reserva Biológica Guaribas (UFPB 7327, 7341); Pernambuco, Reserva de Saltinho ([MZUSP] AD119); Piauí, Parque Nacional de Sete Cidades (DZSJRP 11460 [paratype of vizottoi]), Teresina (DZSJRP 18054 [holotype of vizottoi]); Rio de Janeiro, Ilha da Marambaia (ALP 6121), Jardim Botânico do Rio de Janeiro (7508–7511, 7513–7515, 7517, 7519, 7520, 8059, 8062, 8064, 8077, 8079–8081), Morro Azul (ALP 9142), Parque Estadual da Pedra Branca (ALP 5784), Parque Natural Municipal da Prainha (ALP 6650); Rio Grande do Norte, Mata da Estrela (MZUSP 35027); Santa Catarina, Joinville (BMNH 9.11.19.15); São Paulo, Barra do Ribeirão Onça Parda (MZUSP 10632), Barão Geraldo (ZUEC 783), Cachoeira dos Índios (DZSJRP 3140, 3141), Cananéia (MZUSP 26354), Estação Experimental de Pindorama (DZSJRP 16506), Fazenda João XXIII ([LMUSP] GTG 01), Fazenda Paraguassu (ZUEC 1002), Fazenda Santa Carlota (MZUSP 35028), Fazenda Silvio Fazoli in Irapuã (DZSJRP 2924, 2937, 2945, 2946, 3611, 3728), Grota de Mirassol (DZSJRP 2640, 3997, 3998, 3999, 4029), Iguape (MZUSP 21802), Ilha do Cardoso (MZUSP 28037, ZUFMS 395, 397), Instituto de Biologia Marinha in São Sebastião (DZSJRP 10050), Itapetininga (USNM 542616), Parque Estadual da Ilha Anchieta (MZUSP 29456, 31582), Parque Estadual de Ilhabela (MZUSP 35029), Parque Estadual Turístico do Alto Ribeira (MZUSP 34012), São José do Rio Preto (DZSJRP 2469), São Roque (MZUSP 15112), Sítio Progresso (DZSJRP 3810, 3825, 3921, 4271–4273, 4381–4385, 4445), Ypanema (BMNH 3.7.1.163), Sergipe, Estação Ecológica de Itabaiana (ALP 6545), Floresta Nacional do Ibura (CMUFS SSB1 - FNI, SSB2 - FNI, SSB376 - FNI, SSB377 - FNI, SSB20 - FNI, SSB33 - FNI), Mata do Junco (CMUFS 51, 71, 73, 94), Parque Nacional Serra de Itabaiana (CMUFS 65). Paraguay: La Cordillera, Estancia Sombrero (TTU 75275, 99569); San Pedro, Yaguarete Forests, 0.5 km W headquarters (TTU 95747). Locality unknown: BMNH 7.1.1.698., Published as part of Garbino, Guilherme S. T., Lim, Burton K. & Tavares, Valéria Da C., 2020, Systematics of big-eyed bats, genus Chiroderma Peters, 1860 (Chiroptera: Phyllostomidae), pp. 1-93 in Zootaxa 4846 (1) on pages 37-39, DOI: 10.11646/zootaxa.4846.1.1, http://zenodo.org/record/4017497, {"references":["Taddei, V. A. & Lim, B. K. (2010) A new species of Chiroderma (Chiroptera, Phyllostomidae) from Northeastern Brazil. Brazilian Journal of Biology, 70 (2), 381 - 386. https: // doi. org / 10.1590 / S 1519 - 69842010000200021","Cruz, L. D., Martinez, C. & Fernandes, F. R. (2007) Comunidades de morcegos em habitats de uma Mata Amazonica remanescente na Ilha de Sao Luis, Maranhao. Acta Amazonica, 37 (4), 613 - 619. https: // doi. org / 10.1590 / S 0044 - 59672007000400017","Gregorin, R., Carmignotto, A. P. & Percequillo, A. R. (2008) Quiropteros do Parque Nacional da Serra das Confusoes, Piaui, nordeste do Brasil. Chiroptera Neotropical, 14 (1), 366 - 383.","Lund, P. W. (1842 a) Fortsatte Bemaerkninger over Brasiliens uddode Dyrskabning. Det Kongelige Danske videnskabernes selskabs skrifter, 9, 123 - 136.","Lund, P. W. (1842 b) Blik paa Brasiliens Dyreverden for sidste Jordomvaeltning. Det Kongelige Danske videnskabernes selskabs skrifter, 9, 140 - 208, pls 28 - 38.","ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp.","Gray, J. E. (1866) Revision of the genera of Phyllostomidae, or leaf-nosed bats. Proceedings of the Zoological Society of London, 1866, 111 - 118.","Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. British Museum of Natural History, London, xxxvi + 567 pp. https: // doi. org / 10.5962 / bhl. title. 55341","Thomas, O. (1891) Note on Chiroderma villosum with the description of a new species for the genus. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 10, 881 - 883.","Peters, W. (1860) Eine neute Gattung von Flederthieren, Chiroderma villosum, aus Brasilien vor. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1861,747 - 755.","Winge, H. (1892) Jordfundne og nulevende Flagermus (Chiroptera) fra Lagoa Santa, Minas Geraes, Brasilien. E Museo Lundii, Kjobenhavn, 2, 1 - 65.","Thomas, O. (1893). Further notes on the genus Chiroderma. Annals and Magazine of Natural History, Series 6, 11, 186 - 187. https: // doi. org / 10.1080 / 00222939308677490","Vieira, C. O. da C. (1942) Ensaio monografico sobre os quiropteros do Brasil. Arquivos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo, 3, 219 - 471.","Cabrera, A. (1958) Catalogo de los mamiferos de America del Sur. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \", Zoologia, 4, i-xvi + i-iv + 308.","Taddei, V. A. (1979) Phyllostomidae (Chiroptera) do Norte-ocidental do estado de Sao Paulo. III-Stenodermatinae. Ciencia e Cultura, 31 (8), 900 - 914.","Taddei, V. A. (1980) Aspectos da biologia de Chiroderma doriae, Thomas, 1891 (Chiroptera, Phyllostomidae). Sessoes da Academia Brasileira de Ciencias, 52, 643 - 644.","Nogueira, M. R. & Peracchi, A. L. (2001) The feeding specialization in Chiroderma doriae (Phyllostomidae, Stenodermatinae) with comments on its conservation implications. Chiroptera Neotropical, 8 (1 / 2), 143 - 156.","Nogueira, M. R. & Peracchi, A. L. (2003) Fig-seed predation by 2 species of Chiroderma: discovery of a new feeding strategy in bats. Journal of Mammalogy, 84 (1), 225 - 233. https: // doi. org / 10.1644 / 1545 - 1542 (2003) 084 2.0. CO; 2","Sipinski, E. A. B. & Reis, N. R. (1995) Dados ecologicos dos quiropteros da Reserva Volta Velha, Itapoa, Santa Catarina, Brasil. Revista Brasileira de Zoologia, 12 (3), 519 - 528.","Esberard, C. E. L., Chagas, A. S., Baptista, M., Luz, E. M. & Pereira, C. S. (1996) Observacoes sobre Chiroderma doriae Thomas, 1891 no municipio do Rio de Janeiro, RJ (Mammalia, Chiroptera). Revista Brasileira de Biologia, 56 (4), 651 - 654.","Taddei, V. A. (1973) Phyllostomidae da Regiao Norte-Ocidental do Estado de Sao Paulo. PhD dissertation. Universidade Estadual Paulista, Sao Jose do Rio Preto, 233 pp.","Novaes, R. L. M., Menezes Jr., L. F., Duarte, A. C. & Facanha, A. C. S. (2010) Consumo de Psychotria suturella Muell. Arg. (Rubiaceae) por morcegos no sudeste do Brasil. Chiroptera Neotropical, 16 (1), 535 - 538.","Laurindo, R. S., Gregorin, R. & Tavares, D. C. (2017) Effects of biotic and abiotic factors on the temporal dynamic of bat-fruit interactions. Acta Oecologica, 83, 38 - 47. https: // doi. org / 10.1016 / j. actao. 2017.06.009","Olmos, F. & Boulhosa, R. L. P. (2000) A meeting of opportunists: birds and other visitors to Mabea fistulifera (Euphorbiaceae) inflorescences. Ararajuba, 8 (2), 93 - 98.","Garbino, G. S. T. & Tavares, V. da C. (2018 b) Roosting ecology of Stenodermatinae bats (Phyllostomidae): Evolution of foliage roosting and correlated phenotypes. Mammal Review, 48 (2), 75 - 89. https: // doi. org / 10.1111 / mam. 12114","Arnone, I. S. (2008) Estudo da comunidade de morcegos na area carstica do Alto Ribeira - SP: uma comparacao com 1980. MSc thesis, Universidade de Sao Paulo, Sao Paulo, 116 pp.","Motta-Junior, J. C. & Taddei, V. A. (1992) Bats as prey of stygian owls in southeastern Brazil. Journal of Raptor Research, 26 (4), 259 - 260.","Gregorin, R., Lobao, K. W., Oliveira, L. F., Machado, F. S., Gil, B. B. & Tavares, V. da C. (2017) Vertical stratification in bat assemblages of the Atlantic Forest of south-eastern Brazil. Journal of Tropical Ecology, 33 (5), 299 - 308. https: // doi. org / 10.1017 / S 026646741700027 X","Presley, S. J. (2004) Ectoparasitic assemblages of Paraguayan bats: ecological and evolutionary perspectives. PhD dissertation, Texas Tech University, Lubbock, Texas, 326 pp.","Lourenco, E. C., Patricio, P. M. P. & Famadas, K. (2016) Community components of spinturnicid mites (Acari: Mesostigmata) parasitizing bats (Chiroptera) in the Tingua Biological Reserve of Atlantic Forest of Brazil. International Journal of Acarology, 42 (2), 63 - 69. https: // doi. org / 10.1080 / 01647954.2015.1117525","Lima Silva, C., Valim, M. P. & Graciolli, G. (2017) Acaros ectoparasitos de morcegos no estado de Mato Grosso do Sul, Brasil. Iheringia, Serie Zoologia, 107, 1 - 6. https: // doi. org / 10.1590 / 1678 - 4766 e 2017111","Soares, F. A. M., Rocha, P. A., Mikalauskas, J. S., Graciolli, G. & Ferrari, S. F. (2017) Ectoparasitic bat flies (Diptera, Streblidae) of bats (Chiroptera, Mammalia) from Mata do Junco Wildlife Refuge, Sergipe, northeastern Brazil. Oecologia Australis, 21 (4), 385 - 395. https: // doi. org / 10.4257 / oeco. 2017.2104.03","Lourenco, J. L. M., Minuzzi-Sauza, T. T. C., Silva, L. R., Olivei, A. C., Mendonca, V. J., Nitz, N., Aquiar, L. M. S. & Gurgel-Goncalves, R. (2018) High frequency of trypanosomatids in gallery forest bats of a Neotropical savanna. Acta Tropica, 177, 200 - 206. https: // doi. org / 10.1016 / j. actatropica. 2017.10.012","Taddei, V. A. (1976) The reproduction of some Phyllostomidae (Chiroptera) from the northwestern region of the State of Sao Paulo. Boletim de Zoologia, Universidade de Sao Paulo, 1 (1), 313 - 330. https: // doi. org / 10.11606 / issn. 2526 - 3358. bolzoo. 1976.121587","Silva, S. S. P., Dias, D., Martins, M. A., Guedes, P. G., Almeida, J. C., Cruz, A. P., Serra-Freire, N. M., Damascena, J. S. & Peracchi, A. L. 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148. Vizotto’s Big-eyed Bat Chiroderma vizottoi French: Chiroderme de Vizotto / German: Vizotto-GroRaugenfledermaus / Spanish: Quirodermo de Vizotto Taxonomy. Chiroderma vizottor Taddei & B. K. Lim, 2010, “Teresina (ca. 05° 02’ S and 42° 45' W), in the State of Piaui, Brazil.” This species is monotypic. Distribution. NE Brazil (Piaui and Ceara states). Descriptive notes. Forearm 45-50 mm. No specific data are available for other measurements and body weight. Dorsal fur of Vizotto’s Big-eyed Bat is yellowish brown. Dorsal hairs are tricolored, with dark brown bases, pale buff middles, and yellowish brown tips. Underparts are grayish. Prominent white median dorsal stripe extends from interscapular region to rump. Head has prominent supraocular and subocular stripes of entirely white hairs. Ears have yellowish bases and margins, and the remainder is blackish. Tragus is yellowish, about ¢.33% the ear length. Noseleafis simple, unnotched at tip, and brown. Underparts are grayish. Wing membranes are blackish except for translucent area between second and third fingers. Uropatagium is hairy and well developed, with notch near level of knees. Proximal two-thirds of forearm is hairy. Habitat. Xeric Brazilian caatinga. One Vizotto’s Big-eyed Bat from Piaui was captured in caatinga with relatively low xerophytic trees (3-5 m) near a rocky outcrop. Food and Feeding. There is no specific information available for this species, but all species of Chiroderma for which there is information primarily eat seeds from fruits of Ficus (Moraceae). In Ceara, seven Vizotto’s Big-eyed Bats were captured in mist nets set adjacent to fruiting trees belonging to an unidentified species of Moraceae. Breeding. A pregnant and a lactating Vizotto’s Big-eyed Bat were captured in January. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Vizotto’s Big-eyed bat is known from only ten individuals and virtually nothing is known about its diet, distribution, and life history. Bibliography. Gregorin, Carmignotto & Percequillo (2008), Silva et al. (2015), Taddei & Lim (2010)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2019, Phyllostomidae, pp. 444-583 in Handbook of the Mammals of the World – Volume 9 Bats, Barcelona :Lynx Edicions on page 555, DOI: 10.5281/zenodo.6458594