23 research outputs found
Review of the family Rivulidae (Cyprinodontiformes, Aplocheiloidei) and a molecular and morphological phylogeny of the annual fish genus Austrolebias Costa 1998
Get PDFLa familia Rivulidae es el cuarto clado más diverso dentro de los peces Neotropicales. Junto con algunos géneros de la familia Nothobranchiidae, muchos rivulidos presentan un característico ciclo de vida anual, con huevos resistentes a la desecación y embriones con diapausas que les permiten sobrevivir en los ambientes estacionales donde habitan. Los Rivulidae presentan también dos especies consideradas como los únicos vertebrados hermafroditas suficientes y algunas especies con inseminación interna. El primer objetivo de este artículo es actualizar la sistemática de la familia considerando las relaciones filogenéticas y las sinapomorfías de los clados que la componen, reuniendo información que se encuentra dispersa en la literatura. De esta revisión surge que las relaciones filogenéticas dentro de Rivulidae están todavía sin resolver, especialmente en uno de los grandes clados que la componen, la subfamilia Rivulinae, donde relaciones conflictivas entre géneros anuales y no anuales son evidentes. El segundo objetivo de este trabajo es presentar una hipótesis filogenética, basada en datos morfológicos, mitocondriales y nucleares, de uno de los géneros más diversos de la familia, el género Austrolebias. Nuestros resultados confirman la monofilia del género y de algunos clados subgenéricos previamente definidos, y propone nuevas relaciones entre ellos, particularmente de las especies del subgénero Acrolebias.The family Rivulidae is the fourth most diverse clade of Neotropical fishes. Together with some genera of the related African family Nothobranchiidae, many rivulids exhibit a characteristic annual life cycle, with diapausing eggs and delayed embryonic development, which allows them to survive in the challenging seasonal ponds that they inhabit. Rivulidae also includes two species known as the only the self-fertilizing vertebrates and some species with internal fertilization. The first goal of this article is to review the systematics of the family considering phylogenetic relationships and synapomorphies of subfamilial clades, thus unifying information that is dispersed throughout the literature. From this revision, it is clear that phylogenetic relationships within Rivulidae are poorly resolved, especially in one of the large clades that compose it, the subfamily Rivulinae, where conflicting hypotheses of relationships of non-annual and annual genera are evident. The second goal of this work is to present an updated phylogenetic hypothesis (based on mitochondrial, nuclear, and morphological information) for one of the most speciose genus of Rivulidae, Austrolebias. Our results confirm the monophyly of the genus and of some subgeneric clades already diagnosed, but propose new relationships among them and their species composition, particularly in the subgenus Acrolebias.Fil: Loureiro, Marcelo. Universidad de la República; UruguayFil: Sá, Rafael de. University of Richmond; Estados UnidosFil: Serra, Sebastián W.. Universidad de la República; UruguayFil: Alonso, Felipe. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Bio y Geociencias del NOA. Universidad Nacional de Salta. Facultad de Ciencias Naturales. Museo de Ciencias Naturales. Instituto de Bio y Geociencias del NOA; ArgentinaFil: Krause Lanés, Luis Esteban. Instituto Pro-pampa; BrasilFil: Volcan, Matheus Vieira. Instituto Pro-pampa; BrasilFil: Calviño, Pablo Antonio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; ArgentinaFil: Nielsen, Dalton. Universidade de Taubaté; BrasilFil: Duarte, Alejandro. Universidad de la República; UruguayFil: Garcia, Graciela. Universidad de la República; Urugua
Discovery of endangered annual killifish Austrolebias cheradophilus (Aplocheiloidei: Rivulidae) in Brazil, with comments on habitat, population structure and conservation status
Full text linkCrescimento e fecundidade do peixe anual Austrolebias Nigrofasciatus(cyprinodontiformes:rivulidae) sob condições de laboratório
No full textDissertação(mestrado)- Universidade Federal do Rio Grande, Programa de Pós-Graduação em Aqüicultura, Instituto de Oceanografia, 2009.O peixe anual Austrolebias nigrofasciatus é uma espécie endêmica do sistema lagunar Patos-Mirim, no Sul do Brasil, onde se encontra ameaçado de extinção. Os efeitos da temperatura sobre o crescimento e reprodução de A. nigrofasciatus foram estudados em cativeiro. Experimento 1 – crescimento:
Ovos fertilizados naturalmente, estocados em laboratório em ambiente úmido, foram expostos a água e dentro de 12h eclodiram. O comprimento padrão médio dos peixes recém eclodidos foi 4,67±0,25mm. Os juvenis foram
mantidos a 16 e 22°C por oito semanas e o crescimento foi mais rápido na temperatura mais elevada durante as primeiras semanas, mas tão logo tenham iniciado as desovas (quatro semanas após a eclosão), a taxa de crescimento foi reduzida e ao final de oito semanas eles mediam 23,68±3,73 e 22,68±5,36
mm (p>0,05), respectivamente em 16 e 22°C. Fêmeas mantidas a 22°C alcançaram 23,00±2,83mm e foram significativamente maiores (p<0,05) do que aquelas mantidas em 16°C (17,91±2,47 mm). A primeira desova na
temperatura reduzida foi observada somente na oitava semana após a eclosão. Não foi observada diferença significativa no crescimento para os machos. O sexo de A. nigrofasciatus parece ser determinado pela temperatura, houve uma maior proporção de machos (1:0,6) a 16°C do que a 22°C (1:1,1). Experimento 2 – reprodução: Doze casais foram coletados no ambiente e distribuídos em 12
aquários, mantidos a 17, 21 e 25°C (quatro replicas para cada temperatura) e acompanhados por cinco semanas. A média da fecundidade semanal foi de 30±15; 32±10 e 38±19 para 17, 21 e 25°C, respectivamente, sem apresentar diferença significativa entre os tratamentos. Ao término do experimento, a
fecundidade semanal acumulada também não diferiu entre os tratamentos e apresentou média de 150±49, 159±3 e 190±56 para 17, 21 e 25°C, respectivamente. Ao término do estudo, o crescimento dos machos não foi influenciado pela temperatura (p>0,05), entretanto fêmeas mantidas a 17 e 21°C foram significativamente maiores que aquelas mantidas a 25°C. O fator
de condição também foi reduzido na temperatura mais alta sugerindo que os
reprodutores de A. nigrofasciatus devem ser mantidos em temperatura reduzida para o sucesso reprodutivo.The annual fish Austrolebias nigrofasciatus, an endemic species to the coastal lagoons of Southern Brazil is currently endangered. The effects of temperature on growth and reproduction of A. nigrofasciatus were studied in captivity. Experiment 1 – growth: Naturally fertilized eggs were kept in the laboratory in a humid environment, once they were exposed to water, larvae hatched within 12 hours. Total length of newly hatched larvae was 4.67±0.25mm. They were kept at 16 e 22°C for eight weeks and larvae grew faster at the higher temperature during the first weeks, but as soon as they started to spawn (four weeks after hatching), growth rate was reduced and at the end of the eight weeks they measured 23.68±3.73 e 22.68±5.36 mm (P>0.05), respectively for 16 e 22°C.
Females reared at 22°C reached 23.0±2.83 and were significantly larger (P<0.05) than those reared at 16°C (17.91±2.47 mm). The first spawns at the lower temperature were only observed eight weeks after hatching. There was no significant difference in growth for males. Sex of A. nigrofasciatus seems to
be thermolabile determined, there was a higher proportion of males (1:0.6) at
16°C than at 22°C (1:1.1). Experiment 2 – reproduction: Twelve couples were collected in the wild and distribute in 12 aquaria kept at 17, 21 e 25°C (four replicates for each temperature) where they were followed for five weeks. As a result, the weekly average of eggs per gram of female was 30±15, 32±10 and 38±19 to 17, 21 and 25°C, respectively. There were no significant differences between treatments. At the end of five weeks the average number of eggs was 150±49, 159±3 and 190±56 for the treatments of 17, 21 and 25°C, respectively.At the end of the study, growth of males was not influenced by temperature (P>0.05), however, females kept at 17 and 21°C were significantly larger than those reared at 25°C. Condition factor was also reduced at the highest temperature suggesting that brrodstock of A. nigrofasciatus should be kept at lower temperatures for successful breeding
Austrolebias ephemerus (Cyprinodontiformes: Rivulidae), a new annual fish from the upper Rio Paraguai basin, Brazilian Chaco
No full textVolcan, Matheus Vieira, Severo-Neto, Francisco (2019): Austrolebias ephemerus (Cyprinodontiformes: Rivulidae), a new annual fish from the upper Rio Paraguai basin, Brazilian Chaco. Zootaxa 4560 (3): 541-553, DOI: https://doi.org/10.11646/zootaxa.4560.3.
Austrolebias ephemerus Volcan & Severo-Neto 2019, new species
No full text<i>Austrolebias ephemerus,</i> new species <p>urn:lsid:zoobank.org:act: 1EBF25E9-1D77-4609-A204-A7DEE66EEC35</p> <p> <b>Holotype</b>. ZUFMS 5465, male, 28.6 mm SL, Brazil, Mato Grosso do Sul State, Porto Murtinho, temporary pool near "do Bala" Road, 21°41'34"S 57°42'57"W, F. Severo-Neto, E. Ragalzi, P. Carvalho & D.J. Santana, 23 November 2017.</p> <p> <b>Paratypes</b>. All from Brazil, Mato Grosso do Sul, Porto Murtinho: ZUFMS 5730, 1 female, 28.1 mm SL, same data as holotype; ZUFMS 5463, 12 males, 14.8–19.7 mm SL (4 C&S); 9 females, 15.9–19.3 mm SL (4 C&S); 5 juveniles, sex undetermined, 13.2–19.3 mm SL, temporary pool at Fazenda Retiro Conceição, 21°42’10”S, 57°45’48”W, F. Severo-Neto, E. Ragalzi, P. Carvalho & D.J. Santana, 22 November 2017; MCP 53927, 1 male, 19.9 mm SL, 1 female, 17.7 mm SL, temporary pool at Fazenda Retiro Conceição, 21°42’10”S, 57°45’48”W, F. Severo-Neto, E. Ragalzi, P. Carvalho & D.J. Santana, 22 November 2017; ZUFMS 5464, 6 males, 15.2–31.7 mm SL, 3 females, 26.7–28.0 mm SL, 6 juveniles, sex undetermined, 12.5–15.3 mm SL, temporary pool near road to Fazenda Campo Florido, 21°38’35”S, 57°42’34”W, F. Severo-Neto, E. Ragalzi, P. Carvalho & D.J. Santana, 23 November 2017; ZUFMS 4144, 2 males, 15.7–22.2 mm SL, 3 females, 18.9–25.3 mm SL, temporary pool about 5 km of Rio Amonguijá, 21°41’08”S, 57°44’00”W, F. Severo-Neto & T.R.F Sinani, 12 December 2015; ZUFMS 5725, 1 male, 29.1 mm SL, 1 female, 26.6 mm SL, temporary pool at Fazenda Retiro Conceição, 21°42'08"S 57°45'52"W, F. Severo-Neto & D.J. Santana, 1 March 2018; ZUFMS 5462, 4 males, 15.9–19.8 mm SL, 7 juveniles, sex undetermined, 13.8–15.4, temporary pool at Fazenda Retiro Conceição, 21°42'08"S 57°45'52"W, F. Severo- Neto, E. Ragalzi, P. Carvalho & D.J. Santana, 22 November 2017.</p> <p> <b>Diagnosis.</b> <i>Austrolebias ephemerus</i> can be distinguished from all species belonging to the <i>A. bellottii</i> species group by presenting pectoral fin posterior margin on vertical between base of 5th and 7th anal fin rays in females (<i>vs</i>. between pelvic fin origin and base of 2nd anal fin ray), and by presenting a greater number of gill rakers in the first branchial arch (4–5+14–15 <i>vs</i>. 3–4+10–12; except in <i>A. vandenbergi</i>). It is distinguished from all species of the <i>A. bellottii</i> species group, except <i>A. accorsii</i> and <i>A. queguay</i>, by having a lower head width in males (55.3–61.1 % SL <i>vs</i>. 61.9–72.6 % SL) and females (58.4–62.8 % SL <i>vs</i>. 63.4–73.2 % SL). <i>Austrolebias ephemerus</i> also differs from all species of the <i>A. bellottii</i> species group, except <i>A. melanoorus</i> and <i>A. queguay,</i> by having fewer neuromasts in the preopercular series (16–23 <i>vs</i>. 24–29). In addition, the following characters are useful for diagnosing <i>A. ephemerus</i> from the other species of the <i>A. bellottii</i> group: from <i>A. accorsii</i> by having urogenital papilla attached to anal fin (<i>vs</i>. not attached), more scale rows in transversal series (13–14 <i>vs</i>. 12), position of dorsal fin origin in relation to the neural spines in males (7th–9th <i>vs</i>. 12th–13th) and females (9th–12h <i>vs</i>. 14th–15th), lower number of dorsal fin rays in males (24–25 <i>vs</i>. 26–28) and females (18–20 <i>vs</i>. 25–28), lower number of anal fin rays in males (27–30 <i>vs</i>. 32–34) and females (23–28 <i>vs</i>. 31–32), and position of anal fin origin in relation to the neural spines in males (6th–9th <i>vs</i>. 13th–14th). It is distinguished from <i>A. bellottii</i> by presenting basihyal cartilage width of about 75% of basihyal length (<i>vs.</i> 50–55%), lower head depth in males (103.3–110.0 % SL <i>vs</i>. 110.3– 118.1 % SL), and fewer neuromasts in the post-otic series (3 <i>vs</i>. 4). <i>Austrolebias ephemerus</i> can be distinguished from <i>A. melanoorus</i> by presenting a distinct colour pattern in males, consisting of flanks dark bluish gray with vertical rows of bright greenish blue dots (<i>vs.</i> light bluish gray bars alternating with dark gray bars), a single anterior rostral neuromast (<i>vs</i>. 2), and contact organs on anal fin in males (<i>vs</i>. contact organs absent on anal fin), dorsal fin origin between 1st and 4th anal fin ray in males (<i>vs</i>. slightly anterior or on vertical through anal fin origin), medial pelvic fin membranes about 75–100% coalesced (<i>vs</i>. 10–25 % coalesced), and a higher number of teeth on second pharyngobranchial (5–8 <i>vs</i>. 3–4). It is distinguished from <i>A. queguay</i> by presenting vertical rows of bright greenish blue dots in the flanks in males (vs. well-defined light blue bands contrasting with the sides of the body), pectoral fins hyaline, without iridescent blue sub-marginal band (vs. pectoral fin hyaline with iridescent blue sub-marginal band present), one to three rows of scales at anal fin base (vs. scales absent at anal-fin base). The new species can be distinguished from <i>A. univentripinnis</i> by the presence of contact organs on anal fin in males (<i>vs</i>. contact organs absent on anal fin), pectoral fins posterior margin on vertical between base of 6th and 8th anal-fin rays in males (<i>vs</i>. between 3th and 5th anal-fin rays), and fewer neuromasts in the preopercular series (16–23 <i>vs</i>. 24–26). Finally, <i>Austrolebias ephemerus</i> can be distinguished from <i>A. vandenbergi</i> by having urogenital papilla attached to anal fin (<i>vs</i>. not attached), fewer neuromasts in the infraorbital (2–3+21–26 <i>vs.</i> 3+31), preopercular (16–23 <i>vs</i>. 31) and mandibular series (9–16 <i>vs</i>. 18), fewer scales in longitudinal series (27–30 <i>vs</i>. 31–33), basihyal cartilage with about 50–60% of basihyal length (<i>vs.</i> 75%), and fewer teeth on second pharyngobranchial (5–8 <i>vs</i>. 11–13).</p> <p> <b>Description.</b> Morphometric and meristic data of the holotype and paratypes are presented in Table 1. Males larger than females, largest male examined 31.7 mm SL, largest female 28.1 mm SL. Dorsal profile slightly concave on head, convex from nape to end of dorsal fin base, straight on caudal peduncle. Ventral profile convex from lower jaw to origin of anal fin base, nearly straight on caudal peduncle. Greatest body depth at dorsal fin origin. Body deep and compressed. Snout blunt and jaws short.</p> <p>Dorsal fin rays in males 24–25; in females 18–20. Tip of dorsal fin rounded. Dorsal fin origin at vertical through 1st and 4th anal-fin rays in males and through 2nd and 5th anal-fin rays in females. Origin of dorsal fin at vertical through neural spines of 7th and 9th vertebrae in males and through neural spines of 9th and 12th vertebrae in females. Anal-fin rays in males 27–30; in females 23–28. Tip of anal fin rounded in males. Anteromedian rays of anal fin of females lengthened, resulting in nearly triangular anal fin shape. Origin of anal fin at vertical through pleural ribs of 6th and 9th vertebrae in males and in females through pleural ribs of 8th–11th vertebrae. Caudal fin rounded, 22–24 rays in males and 22–25 in females. Pectoral fin elliptical, with 11–13 rays in both sexes. Pectoral fin posterior tip reaching vertical through 6th to 8th anal-fin rays in males, and at vertical through 5th to 7th analfin rays in females. Pelvic fin rays 4–5. Pelvic fin posterior tip reaching 2nd to 4th anal-fin rays in both sexes. Medial pelvic fin membranes about 75–100 % coalesced. Urogenital papilla attached 60–100% of its length to anal fin in males.</p> <p>Scales cycloid. Body and head entirely scaled, except anterior portion of ventral surface of head. No scales on dorsal fin base. One to three rows of scales on anal-fin base and three rows of scales on caudal-fin base. Supraorbital scales 1–3. No transverse row of scales anterior to H-scale. Frontal squamation H-patterned; E-scales not overlapping medially. Scales arranged in transverse pattern. Lateral line of trunk complete, with one neuromast per scale. Longitudinal series of scales 27–30; transverse series of scales 13–14; scale rows around caudal peduncle 16–18. Contact organs present in body scales and opercular region in males. Row of minute contact organs in the 2–5 uppermost pectoral fin rays, sometimes also on distal portion of anteriormost 2–5 anal fin rays in males. No contact organs on pelvic, dorsal and caudal fins.</p> <p>Cephalic neuromasts: supraorbital 16–18, parietal 2–3, anterior rostral 1, posterior rostral 1, infraorbital 2–3+21– 26, preorbital 2, otic 1–3, post-otic 2–3, supratemporal 1, median opercular 1, ventral opercular 1–2, preopercular 16– 23, mandibular 9–16, lateral mandibular 5–7, paramandibular 1. Two neuromasts on caudal fin base.</p> <p>Six branchiostegal rays. One or two dermosphenotic ossifications. Urohyal deep. Total number of vertebrae 29–30. Gill rakers in first branchial arch 4–5+14–15. Basihyal subtriangular, width about 75% of length; basihyal cartilage about 50–60% of total basihyal length. Five to eight teeth on second pharyngobranchial. Vomerine teeth absent.</p> <p> <b>Colouration in life.</b> Males (Figs. 1–2): Body bluish gray, darker on dorsum; abdominal region bluish white or yellowish; sides of the body with 6 to 11 iridescent narrow bluish vertical bars, which become less conspicuous and discontinuous towards posterior portion of body. Number of vertical bars can be unequal between sides of the body. Urogenital papilla bluish gray. Pectoral fins hyaline, with distal black margin. Dorsal and anal fin bluish grey, with subtle black margins and a few scattered white spots at the base. Opercular region bright light blue. Hyaline to bluish gray caudal fin. Bluish pelvic fins. Suborbital and supraorbital bands dark; supraorbital band wider near the eye, reaching neuromast parietal series. Iris yellow, with a black vertical bar through the centre of the eye.</p> <p>Females (Figs. 3–4): Ground colour gray or light brown, sometimes with vertically elongated, diffuse darkgray to black blotches, forming small bars. One or two black spots on anterocentral portion of flank. Opercular region bright light blue. Iris light yellow, with inconspicuous vertical gray bar through centre of the eye. Fins hyaline. Dorsal and anal fins with small gray spots, sometimes inconspicuous. Supra and suborbital band light gray.</p> <p> <b>Distribution</b> (Fig. 5). Known only from six small temporary pools on the left bank of the Rio Paraguai basin, between the left bank of the Rio Amonguijá and the right bank of the Córrego Progresso, municipality of Porto Murtinho, Mato Grosso do Sul state, Brazil.</p> <p> <b>Etymology</b>. From the Greek <i>ephemeros</i> (ephemeral, short-lived), referring to the short life cycle of the new species and the highly seasonal environment that it inhabits.</p> <p> <b>Habitat</b> (Fig. 6 and 7). <i>Austrolebias ephemerus</i> was recorded in relatively flat and forested areas at altitudes ranging 80 to 100m a.s.l. with arbustive Chacoan vegetation both in open sites and in shaded areas. The pools where the species was recorded had a small area (between 10 m ² and 400 m ²), with low density of macrophytes, depth less than 40 cm, muddy substratum and slightly turbid water. The species was typically the single fish species at its habitat, or, less commonly, coexisting with other annual fish species [i.e., <i>Neofundulus</i> cf. <i>paraguayensis</i> (Myers) and <i>Trigonectes balzanii</i> (Perugia)].</p> <p> <b>Conservation</b>. <i>Austrolebias ephemerus</i> was recorded in small pools within private properties that are generally used for cattle farming. Since the discovery of the species in 2015, one third of the six pools with occurrence of the species were lost owing to their conversion to pasture areas. The species presents a low population density, occurring in small and isolated pools that, combined, have an area less than 1 hectare and in a region under strong pressure from livestock husbandry. <i>Austrolebias ephemerus</i> presents a reduced (AOO <10,000 m ²) and severely fragmented (a) area of occurrence and is experiencing continued decline (b) in the area of occupancy (ii) and in the quality of habitat (iii). In accordance with the IUCN (2011) criteria, <i>A. ephemerus</i> should be considered a “Critically Endangered” species under the category CR B2ab (ii, iii), and as such, the species should be included in future lists of the endangered fauna from Brazil.</p>Published as part of <i>Volcan, Matheus Vieira & Severo-Neto, Francisco, 2019, Austrolebias ephemerus (Cyprinodontiformes: Rivulidae), a new annual fish from the upper Rio Paraguai basin, Brazilian Chaco, pp. 541-553 in Zootaxa 4560 (3)</i> on pages 543-545, DOI: 10.11646/zootaxa.4560.3.6, <a href="http://zenodo.org/record/2627866">http://zenodo.org/record/2627866</a>
A new Dendrocephalus (Crustacea, Anostraca, Thamnocephalidae) from Rio Grande do Sul State, Brazil
No full textAbstract Dendrocephalus riograndensis n. sp., a new species of fairy shrimp, is described from a single temporary pool in Santa Vitória do Palmar, Rio Grande do Sul, Brazil. Males are readily separated from all other DendrocephalusDaday, 1908species by the form of the frontal appendage and the form of the first thoracopods. This new species is most similar to Dendrocephalus goiasensis Rabet & Thiéry, 1996 and could be confused with this species. This new species appears to be endemic to Rio Grande do Sul
Austrolebias botocudo Lanés & Volcan & Maltchik 2021, new species
No full text<i>Austrolebias botocudo</i>, new species <p>(Figs. 1–2, Table 1)</p> <p> <b>Holotype.</b> MCP 54450, male, 47.8 mm SL, Brazil, Rio Grande do Sul, Vacaria, temporary pool near rio Socorro, rio Apuaê-Inhandava drainage, upper rio Uruguay basin, 28°22’43.6” S, 50°52’21.8” W, altitude 934 meters a.s.l., L.E.K. Lanés & M. V. Volcan, 26 July 2017.</p> <p> <b>Paratypes.</b> All from Brazil, Rio Grande do Sul, Vacaria, rio Socorro, rio Apuaê-Inhandava drainage, upper rio Uruguay basin: MCP 54450, male, 51.6 mm SL, same locality as holotype, L.E.K. Lanés, R.S. Godoy & V. Weber, 13 October 2016; MCP 54451, 2 males, 35.7–40.8 mm SL, collected with holotype; MCP 54452, 3 males, 44.7–55.6 mm SL, 3 females, 28.5–39.3 mm SL, same locality as holotype, L.E.K. Lanés & M. V. Volcan, 26 Jul 2017; MCP 54453, 4 males, 36.5–54.6 mm SL (1 C&S), 6 females, 31.2–35.3 mm SL (1 C&S), 28°22’47.9” S, 50°52’ 07.9” W, altitude 926 meters a.s.l., L.E.K. Lanés, R. S. Godoy & M. Fogaça, 23 November 2013; MCP 54454, 3 males, 37.4–43.5 mm SL, 1 female, 33.2 mm SL, 28°22’47.9” S; 50°52’07.9” W, altitude 926 meters a.s.l, L.E.K. Lanés & R. S. Godoy & M. Fogaça, 30 Aug 2013; MCP 54455, 2 males, 28.0– 34.7 mm SL (1 C&S), 1 female, 25.6 (C&S), 28°22’47.9” S, 50°52’07.9” W, altitude 926 meters a.s.l., L.E.K. Lanés & A.B. Moraes, 10 May 2013; MCP 54456, 1 male, 34.8 mm SL (C&S), 1 female, 31.8 (C&S) 28° 22’43.6” S, 50°52’21.8” W, altitude 934 meters a.s.l., L.E.K. Lanés & A.B. Moraes, 10 May 2013.</p> <p> <b>Diagnosis.</b> <i>Austrolebias botocudo</i> differs from its congeners allocated in the subgenus <i>Acrolebias</i> by a unique male colour pattern consisting of body flank lilac grey with bluish white vertical bands (<i>vs</i>. flank pale yellowish brown, with dark brown bars in <i>A. araucarianus</i>; flank golden with purplish grey bars in <i>A. carvalhoi</i>; and flank dark bluish brown with yellowish golden or light grey vertical bands in <i>A. nubium</i>). The new species additionally differs from <i>A. araucarianus</i> and <i>A. carvalhoi</i> by males presenting melanophores irregularly distributed across the body (<i>vs</i>. melanophores absent in <i>A. araucarianus</i> and <i>A. carvalhoi</i>). <i>Austrolebias botocudo</i> differs from the remaining species of the subgenus <i>Acrolebias</i>, except from <i>A. nubium</i>, by presenting contact organs on anal fin in males (<i>vs</i>. contact organs absent). <i>Austrolebias botocudo</i> differ from <i>A. nubium</i> by the presence of discrete contact organs in the flank (<i>vs</i>. conspicuous contact organs), presence of generally only one contact organ by scale of lateral line (<i>vs</i>. two prominent contact organs by scale of lateral line), by presenting jaws slightly prognathous (<i>vs</i>. jaws short); by dorsal profile of head slight concave (<i>vs</i>. nearly convex), more neuromasts in the preopercular + mandibular series (36–43 <i>vs</i>. 26–35), a lower body depth in females (28.2–32.3 % SL <i>vs</i>. 32.6–37.8 % SL), and by presenting basihyal cartilage about 40–50% of basihyal length (<i>vs</i>. 60–70% of basihyal length). The new species differ from <i>A. araucarianus</i> by presenting neuromasts of the supraorbital series united (<i>vs</i>. first three neuromasts of the supraorbital series separated from the remaining neuromasts), preopercular and mandibular series of neuromasts united (<i>vs</i>. separated), by presenting pelvic fin well-developed (<i>vs</i>. pelvic-fin girdle rudimentary or absent), fewer scales in the longitudinal series (27–30 <i>vs</i>. 31–33), and by presenting more neuromasts in the ventral opercular series (2–4 <i>vs</i>. 1). <i>Austrolebias botocudo</i> differ from <i>A. carvalhoi</i> by presenting more neuromasts in the supraorbital (2–3+22–27 <i>vs</i>. 3+20) and lateral mandibular series (5–9 <i>vs</i>. 4), preopercular and mandibular series of neuromasts united (<i>vs</i>. separated), and by presenting basihyal cartilage about 40–50% of basihyal length (vs. 65% of basihyal length).</p> <p> <b>Description.</b> Morphometric data are presented in Table 1. Largest male examined 55.6 mm SL; largest female examined 39.3 mm SL. Dorsal profile slightly concave on head, convex from nape to the end of dorsal-fin base, nearly straight on caudal peduncle. Ventral profile convex from lower jaw to end of anal-fin base, about straight on caudal peduncle. Body moderately deep and compressed. Greatest body depth at vertical of pelvic fin origin. Lower jaw relatively long and prognathous, snout blunt.</p> <p>Dorsal-fin rays in males 22–24; in females 18–20. Dorsal fin sub-rectangular in males, with posterior tip rounded. Dorsal fin semi-circular in females. Origin of dorsal fin in males at vertical through neural spine of 7–9th vertebrae; in females through 10–11th vertebrae. Anal-fin rays in males 21–23, in females 18–20. Anal-fin sub-rectangular, with tip rounded in males. Anal-fin sub-triangular in females. Anteromedian rays of anal fin lengthened in females. Anal-fin origin at vertical through base of 3–5th dorsal-fin ray in males and through base of 2–4th dorsal-fin ray in females, and at vertical through pleural ribs of 8–9th vertebrae in males; and through 10–12th vertebrae in females. Caudal fin distal margin rounded, 24–27 rays in both sexes. Caudal-fin rays supported by last 4 vertebrae. Pectoral fin rays 12–14 in both sexes; pectoral-fin margin rounded. Pectoral-fin posterior tip reaching from before pelvic fin origin to 2nd anal-fin ray in males, from origin of pelvic fin or before it in females. Pelvic fins usually well-developed, with 5–6 rays, its tip extending from urogenital papilla to 2nd anal-fin ray in males, and to urogenital papilla in females. Urogenital papilla not attached to anal fin. Pelvic-fin bases medially separated by short interspace.</p> <p>Scales large and cycloid. Head and trunk entirely scaled, except ventral surface of head. Frontal squamation F, G, or H, sometimes irregularly arranged. No scales on dorsal and anal fin bases. Longitudinal series of scales 27–30, scales regularly arranged; transverse series of scales 11–16; 18–22 scale rows around caudal peduncle. Few, scattered contact organs on scales throughout body in males, when present located mainly at mid-ventral region of the flank from immediately distal tip of opercle to caudal peduncle. Usually one contact organ per scale of longitudinal series; rarely two contact organs along some scales of longitudinal series (present in only three males). Contact organs on distal portion of anteriormost 1–4 anal fin rays, present in 25% of males. Contact organs on outer region of 1–4 uppermost pectoral fin rays in males.</p> <p>Cephalic neuromasts: supraorbital 14–23, parietal 1–3, anterior rostral 1–2, posterior rostral 1, infraorbital 2–3+22–27, preorbital 2–3, otic 2–4, post-otic 2–5, supratemporal 1–2, median opercular 1, ventral opercular 2–4, preopercular + mandibular 36–43, lateral mandibular 5–9, paramandibular 1. Two neuromasts on caudal fin base.</p> <p>Six branchiostegal rays. Dermosphenotic ossification absent. Urohyal deep. Total number of vertebrae 28–29. Gill rakers in first branchial arch 3+8–9. Basihyal subtriangular, width about 60–75% of length; basihyal cartilage about 40–50% of total basihyal length. One to three teeth on second pharyngobranchial. Vomerine teeth absent.</p> <p> <b>Colouration in life. Males</b> (Fig. 2A–D). Ground colour of body lilac grey, brown in dorsal region, with 7–12 bluish white vertical bands, usually with narrower, overlapping or poorly defined stripes between well-defined bars. Some specimens in a dominant situation shown a darker coloured pattern, composed of a blue-purple background with vertical grey-green stripes. Pectoral and ventral region greyish. Melanophores irregularly distributed over the body, forming black dots, spots or irregular marks. Some specimens exhibit a concentration of melanophores in the middle of the flank, forming irregular black vertical bands. This pattern is more frequent in senile individuals. Opercle and preopercle greenish blue. Supraorbital and suborbital bars black. Pectoral fin hyaline, margin black. Pelvic fins dark blue or dark grey. Iris orange; dark vertical band crossing the eye. Dorsal fin dark blue with light grey elongated dots. Anal fin bluish-green with light, elongated grey dots; distal margin of anal fin darker. Caudal fin bluish grey with light grey dots and spots, distal margin hyaline.</p> <p> <b>Females</b> (Fig. 2E–H). Ground colour of body pale yellowish brown, generally with brown, dark brown or black spots distributed over the flank. Some individuals presenting a spotted pattern with several circular marks, which extends from dorsum to the medium flank. Dorsum generally pale yellowish brown. Some specimens present black melanophores in the dorsal region. Venter light yellowish grey. Sides of head yellowish brown, opercular region pale golden or bluish golden. Suborbital bar grey. Iris light yellow with transverse grey bar through centre of eye. Unpaired fins yellowish hyaline, with sparse and little conspicuous grey dots. Pectoral fin hyaline. Pelvic fin hyaline.</p> <p> <b>Distribution.</b> The species was recorded only in two temporary pools located in a highland grassland landscape at the rio Socorro, a tributary of upper rio Apuaê-Inhandava (upper rio Uruguay basin), in altitudes between 926 and 934 meters a.s.l., Vacaria municipality, Rio Grande do Sul State, southern Brazil (Fig. 3).</p> <p> <b>Etymology.</b> The specific epithet “botocudo” is a reference to an indigenous people, which formerly inhabited the region and belonged to the Xokleng Laklãnõ ethnic group, known during the 19th century as “Botocudos”, because adult men used a <i>botoque</i> (a plug-like wooden disk) to enlarge the lower lip. An allusion to a feature presented by the species, where the lower jaw seems more prominent than other species belonging to the subgenus <i>Acrolebias</i>.</p> <p> <b>Ecological notes.</b> <i>Austrolebias botocudo</i> was found only in two isolated high-altitude temporary wetlands, located within a private farmland, with size areas of 2,500 and 13,000 m 2. These two biotopes present clear (sometimes turbid), still water, with a latosol soil type, and are located in a grassland landscape (Fig. 4). The wetlands are close to each other (less than one kilometer apart) but are hydrologically isolated. A portion of the larger temporary pool was excavated, probably to become a water source for cattle. There are small intermittent drainages associated with the pools. The vegetation is composed mainly by stands of Peruvian watergrass (<i>Luziola peruviana</i>), peatbog (<i>Sphagnum</i> sp.), and other emergent, floating and submerse macrophytes. During winter and early spring, when the temporary pools present water, the minimum air temperature reaches 0°C (often negative temperatures are recorded), and frost, dew, fog and even snow can occur. The area where the biotopes inhabited by <i>A. botucudo</i> lie within are used for extensive cattle ranching. In the surroundings, mainly soybean crops and orchards (apples and pears) have drastically altered the natural high-altitude grasslands landscape.</p> <p> <b>Conservation.</b> In accordance with IUCN criteria, <i>Austrolebias botocudo</i> should be considered as “Critically Endangered”, with the criteria CR B2ab (ii, iii). The species presents an area of occupancy of 15,500 m 2 (AOO less than 10 km ²), present severely fragmented populations (a) with continued decline (b) in area of occupancy (ii) and quality of habitat (iii).</p>Published as part of <i>Lanés, Luis Esteban Krause, Volcan, Matheus Vieira & Maltchik, Leonardo, 2021, Two new annual fishes (Cyprinodontiformes: Rivulidae) unexpectedly discovered in the highlands of southern Brazil, pp. 499-520 in Zootaxa 4949 (3)</i> on pages 501-506, DOI: 10.11646/zootaxa.4949.3.4, <a href="http://zenodo.org/record/4640565">http://zenodo.org/record/4640565</a>
Austrolebias nubium Lanés & Volcan & Maltchik 2021, new species
No full text<i>Austrolebias nubium</i>, new species <p>(Figs. 5–6; Table 2).</p> <p> <b>Holotype.</b> MCP 54457, male, 41.1 mm SL, Brazil, Rio Grande do Sul State, Bom Jesus municipality, rio Santana, a tributary of upper rio Taquari-Antas drainage, upper rio Jacuí basin, Laguna dos Patos hydrographic system, 28°37’00” S, 50°26’42” W, altitude 1061 meters a.s.l., L.E.K. Lanés & M. V. Volcan, 20 July 2017.</p> <p> <b>Paratypes.</b> All from Brazil, Rio Grande do Sul State, Bom Jesus municipality, rio Santana, upper rio Jacuí drainage. MCP 54458, 1 male, 40.8 mm SL, 3 females, 34.5–38.4 mm SL, collected with holotype. MCP 54459, 6 males 27.8–35.5 mm SL (1 C&S), 7 females, 23.6–30.04 mm SL (1 C&S), 28°36’58.6” S, 50°26”41.7” W, altitude 1055 meters a.s.l., L.E.K. Lanés, R. S. Godoy, M. Reichard & L. Maltchik, 12 October 2012. MCP 54460, 6 males, 29.2–52.7 mm SL (2 C&S), 3 females, 24.9–27.7 mm SL (2 C&S), same locality as holotype, L.E.K. Lanés, 30 August 2013. MCP 54461, 3 males, 30.9–36.0 mm SL; 3 females, 28.7–29.7 mm SL, same locality as holotype, L.E.K. Lanés, R. S. Godoy, M. Reichard & L. Maltchik, 11 October 2012. MCP 54462, 1 male, 41.9 mm SL, same locality as holotype, L.E.K. Lanés & R.S. Godoy, 13 October 2016.</p> <p> <b>Diagnosis.</b> <i>Austrolebias nubium</i> differs from its congeners allocated in the subgenus <i>Acrolebias</i> by a unique male colour pattern consisting of flanks predominantly dark bluish brown with yellowish golden or light grey narrow vertical bands (<i>vs</i>. flank pale yellowish brown, with dark brown bars in <i>A. araucarianus</i>; flanks lilac grey with bluish white vertical bands in <i>A. botocudo</i>; and flanks golden with purplish grey bars in <i>A. carvalhoi</i>); presence of melanophores irregularly distributed across the body (<i>vs</i>. absence of melanophores in <i>A. araucarianus</i> and <i>A. carvalhoi</i>) (Fig. 7) (shared with <i>A. botocudo</i>); contact organs on anal fin in males (<i>vs</i>. contact organs absent) (shared with <i>A. botocudo</i>) and by prominent contact organs in the flank in males (<i>vs</i>. contact organs absent in <i>A. araucarianus</i> or discrete in <i>A. botocudo</i> and <i>A. carvalhoi</i>), and opercle (<i>vs</i>. absent in <i>A. araucarianus</i> and <i>A. carvalhoi</i> or discrete and rarely present in <i>A. botocudo</i>). In addition, <i>A. nubium</i> differ from all species of the subgenus <i>Acrolebias</i> by the presence of two prominent contact organs in the distal margin of scales of longitudinal series in larger males (<i>vs</i>. never a similar pattern) (Fig. 8). The new species differ from <i>A. botocudo</i> by presenting jaws short (<i>vs</i>. jaws slightly prognathous) (Fig. 9), dorsal profile of head nearly convex (<i>vs</i>. slight concave) (Fig. 9), fewer neuromasts in the preopercular + mandibular series (26–35 <i>vs</i>. 36–43), by a higher body depth in females (32.6–37.8% SL <i>vs</i>. 28.2–32.3% SL), and by presenting basihyal cartilage about 60–70% of basihyal length (<i>vs</i>. 40–50% of basihyal length). <i>Austrolebias nubium</i> differ from <i>A. araucarianus</i> by presenting neuromasts of the supraorbital series united (<i>vs</i>. first three neuromasts of the supraorbital series separated from the remaining neuromasts), preopercular and mandibular series of neuromasts united (<i>vs</i>. separated), pelvic fin well-developed (<i>vs</i>. pelvic-fin girdle rudimentary or absent), fewer scales in the longitudinal series (27–30 <i>vs</i>. 31–33), more neuromasts in the ventral opercular series (2–4 <i>vs</i>. 1). The new species differ from <i>A. carvalhoi</i> by having fewer scales around caudal peduncle (18–20 <i>vs</i>. 22), pelvic-fin posterior tip reaching urogenital papilla to 1st anal-fin ray in males, and from anterior to pelvic fin origin to anus in females (<i>vs</i>. tip of pelvic fin reaching base of 2nd anal-fin ray in both males and females), and preopercular and mandibular series of neuromasts united (<i>vs</i>. separated).</p> <p> <b>Description.</b> Morphometric data are presented in Table 2. Males larger than females, largest male examined 52.7 mm SL; largest female examined 38.4 mm SL. Dorsal profile slightly convex from nape to the end of dorsal-fin base, nearly straight on caudal peduncle. Ventral profile nearly convex from lower jaw to end of anal-fin base, about straight at caudal peduncle. Body moderately deep and compressed. Greatest body depth at vertical of pelvic fin. Snout blunt, jaws short. Dorsal-fin rays in males 22–24; in females 18–20. Dorsal fin sub-rectangular in males, with posterior tip rounded. Dorsal fin semi-circular in females. Origin of dorsal fin in males at vertical through neural spines of 8–9th vertebrae in males; in females through neural spines of 10–12th vertebrae. Anal-fin rays in males 21–23; in females 18–20. Anal-fin sub-rectangular, with tip rounded in males. Anal-fin sub-triangular in females. Anteromedian rays of anal fin of females lengthened. Anal-fin origin on vertical through base of 2nd to 5th dorsalfin ray in males and females, and at vertical through pleural ribs of 8–9th vertebrae in males; and through 10–13th vertebrae in females. Caudal-fin distal margin rounded, with 24–27 rays in both sexes. Caudal-fin rays supported by last 4 or 5 vertebrae. Pectoral-fin rays 12–14; pectoral-fin rounded. Pectoral-fin posterior tip reaching from pelvicfin origin to urogenital papilla in males and females. Usually well-developed pelvic fins. Pelvic fin rudimentary in one female paratype (28.2 mm SL); pelvic-fin present on a single side in one male paratype (male 33.1 mm SL). Pelvic fin rays 5–6 in both sexes. Pelvic-fin tip extending from urogenital papilla to 1st anal-fin ray in males, and from anterior to pelvic fin origin to anus in females. Urogenital papilla not attached to anal fin, sometimes with a thin membrane at the base. Pelvic-fin bases medially separated by short interspace.</p> <p>Scales large and cycloid. Head and trunk entirely scaled, except ventral surface of head. Frontal squamation F, G and H, sometimes irregularly arranged. No scales on anal and dorsal fin bases. Three rows of scales on caudal fin basis. Longitudinal series of scales 27–30, scales regularly arranged; transverse series of scales 11–14; 18–20 scale rows around caudal peduncle. Longitudinal series of trunk scales complete with one neuromast per scale. Large and evident contact organs on scales throughout most of body in males in the mid-lower region of the flank from the medial region of the operculum to the caudal peduncle. Usually one prominent contact organ per scale of body and two prominent contact organs on margin of scales along lateral line (Fig. 8). Contact organs on distal portion of anteriormost 3–5 anal fin rays present in 50% of males. Contact organs on outer region of 1–4 uppermost pectoral fin rays in males.</p> <p>Cephalic neuromasts: supraorbital 14–18, parietal 0–2, anterior rostral 1, posterior rostral 1, infraorbital 2– 3+18–24, preorbital 2, otic 1–4, post-otic 2–4, supratemporal 1–2, median opercular 1, ventral opercular 2–4, preopercular + mandibular 26–35, lateral mandibular 2–5, paramandibular 1. Two neuromasts on caudal fin base.</p> <p>Six branchiostegal rays. Dermosphenotic ossification absent. Urohyal deep. Total number of vertebrae 28–30. Gill rakers in first branchial arch 3+9. Basihyal subtriangular, width about 75–95% of length; basihyal cartilage about 60–70% of total basihyal length. Two to four teeth on second pharyngobranchial. Vomerine teeth absent.</p> <p> <b>Colouration in life. Males</b> (Fig. 5A–C, 7). Ground colour of body dark bluish brown, darker at dorsal region, with 9–13 yellowish golden or light grey narrow vertical bands, usually with narrower, poor defined or overlapping stripes between the well-defined bars. Pectoral and ventral region greyish. Melanophores occur irregularly distributed over the body, forming black dots, spots or irregular marks, more visible in stressed or faded individuals. Opercle and preopercle greenish blue. Supraorbital and suborbital bars black. Pectoral fin hyaline, ventral margin black. Pelvic fins dark blue or dark grey. Iris orange; dark vertical band crossing the eye. Dorsal and anal fins dark blue, with light grey dots; distal region of anal fin metallic green with the margin darker. Caudal fin bluish grey with light grey dots and spots, distal margin hyaline.</p> <p> <b>Females</b> (Fig. 5D–F). Ground colour of body pale yellowish brown, generally with vertically elongated grey or brown spots. Some specimens present several faded, irregular marks forming oval or vertically elongated blotches across the flank. Dorsum generally pale yellowish brown. Venter light yellowish grey. Side of head yellowish brown, opercular region pale golden or pale greenish golden. Suborbital bar grey. Iris light yellow with transverse grey bar through middle of eye. Unpaired fins yellowish hyaline, with sparse and discreet grey dots. Pectoral fins hyaline. Pelvic fins hyaline.</p> <p> <b>Distribution.</b> The species is only known from temporary pools located in highland grasslands of the rio Santana drainage, upper rio Taquari-Antas, upper rio Jacuí basin, Laguna dos Patos hydrographic system, in altitudes of ~ 1000 meters a.s.l., Bom Jesus municipality, Rio Grande do Sul State, Southern Brazil (Fig. 3).</p> <p> <b>Etymology.</b> The specific name “ <i>nubium</i> ” is alluding to cloud in latin [nubis]. The genitive plural of this i-stem noun is ‘nubium’ ‘of the clouds’. The specific epithet is in allusion to the fact that the species occurs at the highest altitude recorded for a member of genus <i>Austrolebias</i>. In addition, annual fish are popularly known as “peixes das nuvens” [cloud fishes], because they appear after rains in areas that were previously completely dry.</p> <p> <b>Ecological notes.</b> <i>Austrolebias nubium</i> was found only in isolated, natural temporary pools (with areas ranging between 780 and 1,100 m 2), with clear, lentic water, situated on cambisoils, and located in grassland landscape of private farmlands (Fig. 10). There are small intermittent drainages associated with the pools. The vegetation is composed by peatbog (<i>Sphagnum</i> sp.), emergent, floating, and submerse macrophytes. During the austral winter, temperatures can drop to below 0˚ C, with frost and even snow can occur in the species biotope. The surroundings of the biotopes are used for extensive cattle ranching, soybean crops, and orchards (apples and pears). Both populations are found in close proximity (less than one kilometer apart) but are hydrologically isolated.</p> <p> <b>Conservation.</b> In accordance with IUCN criteria, <i>Austrolebias nubium</i> is considered as “Critically Endangered” species, under the criteria CR B2ab (ii, iii). The species presents an area of occupancy of 1880 m 2 (AOO less than 10 km ²), populations severely fragmented (a) and continued decline (b) in area of occupancy (ii) and quality of habitat (iii).</p>Published as part of <i>Lanés, Luis Esteban Krause, Volcan, Matheus Vieira & Maltchik, Leonardo, 2021, Two new annual fishes (Cyprinodontiformes: Rivulidae) unexpectedly discovered in the highlands of southern Brazil, pp. 499-520 in Zootaxa 4949 (3)</i> on pages 507-512, DOI: 10.11646/zootaxa.4949.3.4, <a href="http://zenodo.org/record/4640565">http://zenodo.org/record/4640565</a>
Acrolebias Costa 2008
No full textIdentification key for the species of the subgenus <i>Acrolebias</i> <p>1a. Contact organs present on anal fin and opercle of males, preopercular and mandibular series of neuromasts united........ 2</p> <p>1b. Contact organs absent on anal fin and opercle in males, preopercular and mandibular series of neuromasts separated....... 3</p> <p> 2a. Sides of body dark bluish brown with yellowish golden or light grey narrow vertical bands, prominent contact organs in the flank in males, jaws short, dorsal profile of head nearly convex............................. <i>Austrolebias nubium</i> <b>sp. n.</b> (rio Taquari-Antas, upper rio Jacuí, Lagoa dos Patos basin, Brazil)</p> <p> 2b. Sides of body lilac grey with bluish white vertical bands, discrete contact organs in the flank in males, jaws slightly prognathous, dorsal profile of head slightly concave......................................... <i>Austrolebias botocudo</i> <b>sp. n.</b> (Upper rio Apuaê-Inhandava drainage, rio Uruguay basin, Brazil)</p> <p> 3a. Pelvic fin present, supraorbital series continuous, 27–29 scales in the longitudinal series............ <i>Austrolebias carvalhoi</i> (rio Iguaçu drainage, rio Paraná basin, Brazil)</p> <p> 3b. Absence or extreme reduction of pelvic fin, first three neuromasts of the supraorbital series separated from the remaining neuromasts by an interspace, 31–33 scales in the longitudinal series............................ <i>Austrolebias araucarianus</i> (rio Claro drainage, rio Iguaçu, rio Paraná basin, Brazil)</p>Published as part of <i>Lanés, Luis Esteban Krause, Volcan, Matheus Vieira & Maltchik, Leonardo, 2021, Two new annual fishes (Cyprinodontiformes: Rivulidae) unexpectedly discovered in the highlands of southern Brazil, pp. 499-520 in Zootaxa 4949 (3)</i> on page 513, DOI: 10.11646/zootaxa.4949.3.4, <a href="http://zenodo.org/record/4640565">http://zenodo.org/record/4640565</a>
