Neotropical Biology and Conservation 10(3):177-181, september-december 2015 Unisinos - doi: 10.4013/nbc.2015.103.08

SHORT COMMUNICATION

Establishment of non-native predator (Pisces, Erythrinidae) in a tributary of the Upper Paraná River basin, south Brazil

Estabelecimento de predador não nativo (Pisces, Erythrinidae) em tributário da bacia do Alto rio Paraná, sul do Brasil

Diego Azevedo Zoccal Abstract Garcia1,2* [email protected] Erythrinus erythrinus is an erythrinid native from the basins of the Amazon, Orinoco, coastal rivers of the Guyana and Lower Paraná, but allochthonous of Tibagi River region, Paranapanema basin. In the present work, we report the occurrence of E. erythrinus in Marina Carmona Água dos Tigres and Taquari rivers, in the Paranapanema basin, and aim to verify the Hernandes2 occurrence of constancy and the condition of health of its specimens in sub-tributaries of [email protected] the Lower Paranapanema River. The catch-per-unit-effort of the Água dos Tigres River mouth was about three times higher than the site upstream of the same river, and the Taquari River presented the lowest value. The species presented constancy of catches Ângela Teresa Silva-Souza3 in tributaries of the basin, where isometric growth and good health condition of the spec- [email protected] imens were observed. Its introduction into the Tibagi River basin may have occurred due to an intentional release as remaining live bait discarded in commercial fishery operations Mário Luís Orsi2 or may have dispersed after the flood of the Sete Quedas Falls. The presence of juveniles [email protected] along with adults indicates that the species is undergoing an establishment stage of the invasion process in the region.

Keywords: Tibagi River, biological invasion, fishes, diversity loss.

Resumo

Erythrinus erythrinus é um erythrinídeo nativo das bacias do Amazonas, Orinoco, rios costeiros da Guiana e Baixo Paraná, mas alóctone da região do rio Tibagi, bacia do Pa- ranapanema. No presente trabalho, registra-se a ocorrência da espécie nos rios Água dos Tigres e Taquari, na bacia do Paranapanema, e objetiva-se verificar a constância de captura e condições de saúde dos espécimes em subafluentes do baixo rio Paranapa- nema. A captura por unidade de esforço da foz do rio Água dos Tigres foi cerca de três

1 Programa de Pós-graduação em Ciências Biológicas da vezes maior do que a montante do mesmo rio, e o rio Taquari apresentou o menor valor. Universidade Estadual de Londrina. Rod. Celso Garcia A espécie apresentou constância de captura nos tributários da bacia, onde constatou-se Cid, PR 445, km 380, 86057-970, Londrina, PR, Brasil. crescimento isométrico e boa condição de saúde dos espécimes. Sua introdução na bacia 2 Universidade Estadual de Londrina, Centro de Ciências do rio Tibagi pode ter ocorrido devido a solturas intencionais de remanescentes de iscas Biológicas, Departamento de Biologia Animal e Vegetal, Laboratório de Ecologia de Peixes e Invasões Biológicas. vivas em pescarias comerciais, ou pode ter se dispersado após a inundação dos Saltos de Rod. Celso Garcia Cid, PR 445, km 380, 86057-970, Sete Quedas. A presença de juvenis junto aos adultos indica que a espécie encontra-se Londrina, PR, Brasil. em estabelecimento na região. 3 Universidade Estadual de Londrina, Centro de Ciências Biológicas, Departamento de Biologia Animal e Vegetal, Laboratório de Ecologia de Parasitos de Organismos Palavras-chave: Rio Tibagi, invasão biológica, peixes, perda de diversidade. Aquáticos. Rod. Celso Garcia Cid, PR 445, km 380, 86057- 970, Londrina, PR, Brasil. * Author for correspondence.

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 3.0), which permits reproduction, adaptation, and distribution provided the original author and source are credited. Diego Azevedo Zoccal Garcia, Marina Carmona Hernandes, Ângela Teresa Silva-Souza, Mário Luís Orsi

The biological invasion by fishes is a of the Upper Paraná River (Langeani occurred at two locations: upstream process associated with dam construc- et al., 2007), this is the first record of (22º59’48.8”S, 50º58’43.8”W) and tions, water transpositions, trade and this species considered allochthonous at the river mouth (22º59’02.9”S, international traffic, promoting chang- in the Tibagi River, Paranapanema 50º58’40.8”W). In the Taquari River, es in biological diversity, a common River basin. In its original distribution captures were accomplished only event in countries with more intense area, this species is found in different at the river mouth (23º11’03.6”S, trade activities (Xu et al., 2012). The habitats, especially among the veg- 50º56’48.5”W) (Figure 1). translocation of fishes outside their etation of shallow stretches of lentic The specimens were captured with the original areas can promote biotic ho- environments, where it builds its nests use of seines and sieves with meshes mogenization and, thus, put the native in association with the vegetation sub- of 0.5cm between adjacent knots. The diversity at risk by changing ecosys- strate (Oyakawa and Mattox, 2009). specimens were anesthetized and eu- tem processes (Roques, 2012; Vitule It shows facultative air-breathing and thanized in water saturated with clove and Pozenato, 2012). high adaptability to environmental oil, measured (standard length, SL, in In Brazil, introductions of fishes are changes (Brosset, 1997; Jucá-Chagas, centimeters) and weighed (weight to- common practices and, as a result of 2004). tal, WT, in grams), fixed in 10% for- economic interest, population growth Given the processes of biological in- malin and deposited in the Museu de and leisure, such events have been oc- vasion in freshwater ecosystems, the Zoologia da Universidade Estadual de curring more frequently (Agostinho present work aimed at verifying the Londrina (MZUEL 5743). Individuals et al., 2007). For example, introduc- occurrence of constancy and the con- having one-third of the maximum to- tions of the African catfish, Clarias dition of health of its specimens in tal length reached by the species (20 gariepinus (Burchel, 1822), and of the sub-tributaries of the Lower Paran- cm) (Graça and Pavanelli, 2007; Orsi, channel catfish, Ictalurus punctatus apanema River, to test the hypothesis 2010) were considered to be juveniles. (Rafinesque, 1818), occurred to attend that this species is undergoing an es- The UEL Ethics Committee for Ani- economic and fishery interests. As tablishment stage of the introduction mal Use authorized the sampling col- these fish are predators and drought- process in that basin. lections (CEUA Nº. 21149.2012.53). resistant, they entail considerable Captures were done seasonally be- The capture constancy was obtained environmental impacts (Vitule et al., tween August 2011 and October only for Água dos Tigres River, us- 2006; Zanatta et al., 2010; Vitule and 2012, in the Água dos Tigres and ing the formula C = p.100/P, where Pozenato, 2012). Taquari rivers, located in the lower C = constancy of species; p = number Among the causes of translocations portion of the Tibagi River basin. In of samplings containing the species; of allochthonous fish in northern Par- the Água dos Tigres River, captures P = total number of collections. aná State is the flooding of the Sete Quedas Falls, geographical barrier af- ter the construction of the Itaipu Dam on the Paraná River (Júlio Jr. et al., 2009); escapes from fish farms (Orsi and Agostinho, 1999); releases for sport fishing (Orsi and Britton, 2012), and releases of aquarium fish (Garcia et al., 2014a). Furthermore, the re- lease of remaining live baits at the end of the amateur and sport fishing sea- sons (Orsi and Britton, 2014) contrib- uted to the translocations of species in the environment. The predator Erythrinus erythrinus (BLOCH & SCHNEIDER 1801) (com- mon name: Jejú) occurs naturally in the basins of the Amazon and Ori- noco, coastal rivers of the Guyana (Oyakawa, 2003), and in the Lower Paraná River basin (Britski et al., Figure 1. Location of the sampling points in the Lower Tibagi River region, Paranapanema 2007; Júlio Jr. et al., 2009). Despite River basin, south Brazil. 1 = Mouth of the Água dos Tigres River; 2 = Upstream from the already registered to another region Água dos Tigres River; 3 = Taquari River.

178 Volume 10 number 3  september - december 2015 Establishment of non-native predator (Pisces, Erythrinidae) in a tributary of the Upper Paraná River basin, south Brazil

A given species is considered constant compared to the central value of Kn = characteristic used by adults to build in the environment when it is present 1.0, by the Student’s t test. This analy- their nests (Suzuki et al., 2004) and in over 50% of the samplings (Dajoz, sis involved 25 specimens of E. eryth- increase the chances for the survival 1978). The total sampling effort for rinus (Figure 2). of their offspring. Thus, environment this river was 11 collections. We recorded the occurrence of E. disruptions may facilitate the recruit- The relative abundance of the spe- erythrinus in both Água dos Tigres ment and establishment of E. erythri- cies was estimated by the catch-per- and Taquari rivers. Seventeen spec- nus. Furthermore, disturbances in the unit-effort for the two rivers (CPUE = imens were caught at the mouth of local community may promote and N*10/Effort), expressed by the num- the Água dos Tigres River and five facilitate invasions of these and other ber of individuals captured (N) in one of them were juveniles, whereas six species (Roy et al., 2013). The current hour/4.0 m2 of collection equipment. specimens were captured upstream conditions can allow the dispersal of The weight-length relationship was (five adults and one juvenile). In the the species to other tributaries and calculated using WT = a.SLb (Le Taquari River, only two adult speci- sub-tributaries in the Tibagi River ba- Cren, 1951), where a is the linear re- mens were captured. The individuals sin, as evidenced by the presence of gression coefficient and b the angular of E. erythrinus were captured in eight specimens in the Taquari River. regression coefficient of log trans- of the 11 collections accomplished in The presence of juveniles together formed weight and length. The physi- the Água dos Tigres River. Thus, the with adults of E. erythrinus indicates ological state of a fish is conditioned capture constancy value was equal to that there has been a recruitment of by the interaction of biotic and abiotic 72.7%, i.e., E. erythrinus was consid- the species in the region (Suzuki et factors. Variations of this state can ered constant in the analyzed region. al., 2004; Leal et al., 2010). It also be expressed by the condition factor The CPUE of the Água dos Tigres constitutes evidence that the species (K), which indicates the recent food River mouth was about three times is in an establishment stage (Coulatti conditions and varies according to higher than the CPUE on the site up- and MacIsaac, 2004; Blackburn et al., the sexual maturity cycle. Its value stream of the same river (42.5 and 2011). Its presence also indicates that reflects the nutritional status or the 15.0, respectively). The Taquari Riv- this new environment has favorable energy expended in cyclical activi- er presented the lowest value (CPUE characteristics for the development ties, or both, and can be related to the = 5.0). The confluence of the Água and maintenance of a viable popula- environmental conditions and behav- dos Tigres and Tibagi Rivers is a tion. Thus, the species would be ful- ioral aspects of the species (Vazzoler, popular place for sports and commer- filling the establishment stage that fol- 1996). Its calculation is dependent on cial fishermen. The occurrence of E. lows the invasion process (Lockwood fish weight and length (Verani et al., erythrinus, most frequently near the et al., 2007). 1997). Conversely, the relative condi- convergence of these two rivers, sug- The analysis of the weight-length re- tion factor (Kn) compares the meas- gests that the species may have been lationship provided the values of the ured value (WT) to the theoretically introduced when used as live baits. linear coefficients (a) = 0.036 and the expected value (WE) for each length, This area shows lentic characteris- angular coefficient (b) = 2.928, indic- and to the central value of Kn = 1.0 tics, since it receives the influence ative of isometric growth (Figure 3A). (Anderson and Neumann, 1996), indi- of the Escola Engenharia Mackenzie The mean relative condition factor cating the healthiness of the popula- hydroelectric power plant reservoir obtained (with its respective standard tion. After calculation of the Kn val- (Capivara Dam) (Figure 1). The con- error) was equal to 1.00 ± 0.09 (min. ues, the estimation of the mean value stant presence of dense grass near the – max. = 0.83 – 1.24) and presented was accomplished, and statistically river mouth is another environmental no difference when compared to the central value of Kn = 1.0 (p = 0.87). Thus, the fish had an average body weight equal to the theoretically ex- pected (Figure 3B), indicating that the fish was in good health condition. The weight-length relationship and the relative condition factor (Kn) of intro- duced species suffer the influence of environmental characteristics, popula- tion size, and intra and inter-specific Figure 2. Erythrinus erythrinus (MZUEL 5743) captured in the Água dos Tigres River, interactions (Le Cren, 1951; Gomiero Lower Tibagi River region, Paranapanema River basin, south Brazil. Total length = 11 cm, and Braga, 2003). Thus, the survival Standard length = 7.2 cm. of the species in a new area depends

Neotropical Biology and Conservation 179 Diego Azevedo Zoccal Garcia, Marina Carmona Hernandes, Ângela Teresa Silva-Souza, Mário Luís Orsi

The success of the invasion depends on the ecological environment resist- ance and propagule pressure, such as the amount and type of life strate- gies of the introduced species (Holle and Simberloff, 2005). The influ- ence of the reservoir on the sampled area (environmental damage) and the possibility of releases of live baits by fishermen (propagule pressure) are real. Therefore, the control and supervision of non-native species marketed and used as live baits are imperative. Furthermore, the eradica- tion of introduced species in aquatic environments becomes impossible after their establishment (Gozlan et al., 2010), especially in large reser- voirs. Actions to raise awareness of the riverine populations, fish farmers and fishermen are necessary in order to prevent greater dispersals and the consequences of new introductions and translocations in natural aquatic ecosystems. New surveys should be conducted over a longer period in or- Figure 3. Weight (g)-length (cm) relationship curve for Erythrinus erythrinus (A) and varia- der to continue monitoring the situa- tion of the relative condition factor (Kn), compared to the central value (Kn = 1.0) in relation tion of this species in this new place to the total weight of the species (B) in the Lower Tibagi River region, Paranapanema River of occurrence. basin, south Brazil. Acknowledgments

on its adaptability, which influences native community (Agostinho and We wish to thank Aparecido de Souza the reproductive and food events Júlio Jr., 1996; Orsi, 2010). The con- and Edson Santana for their technical (Gomiero et al., 2010). The E. eryth- sequences of E. erythrinus invasion support, and Alexandro Costa for his rinus specimens showed isometric are unknown. Although being small- help with the field samplings; to Ana growth (i.e., their body length incre- sized, this species is considered a vo- Maria Leal-Zanchet and anonymous ments were similar to their increases racious predator that can compete for referees for critical review. in body weight) and good health con- food and physical resources with the dition (Kn = 1.0), suggesting their ad- native species, including Hoplias ma- References aptation to local circumstances (Gar- labaricus (Bloch, 1794), which also AGOSTINHO, A.A.; JÚLIO JR., H.F. 1996. cia et al., 2014b). This compliance belongs to the Erythrinidae. Ameaças ecológicas: peixes de outras águas. with new environmental demands There was a decrease diversity and Ciência Hoje, 21(124):36-44. demonstrates the type of adjustability abundance of native fish in the Capi- AGOSTINHO, A.A.; GOMES, L.C.; PELICI- to new environments and hence suc- vara Power Plant Reservoir after the CE, F.M. 2007. Ecologia e manejo de recursos cess in the establishment (Vitule and introduction of eleven species. Among pesqueiros em reservatórios do Brasil. Marin- gá, Eduem, 501 p. Prodocimo, 2012). those are the piscivorous species Plagi- ANDERSON, R.O.; NEUMANN, R.M. 1996. The competition for food resources oscion squamosissimus (Heckel, 1840), Length, weight, and associated structural in- or locations for spawning seems to Astronotus crassipinnis (Heckel, 1840) dices. In: L.A. NIELSEN; D.L. JOHNSON be one of the most conspicuous ways and Cichla monoculus Spix, 1831 (Orsi (eds.), Fisheries techniques. Maryland, Fisher- whereby invasive species can displace and Britton, 2014). Thus, the introduc- ies Techniques, p. 447-481. BLACKBURN, T.M.; PYSEK, P.; BACHER, native species, influence the decrease tion of E. erythrinus can increase the S.; CARLTON, J.T.; DUNCAN, R.P.; JARO- of the native recruitment and, over ecological imbalance of the region al- SÍK, V.; WILSON, J.R.U.; RICHARDSON, time, of cause the decimation of the ready affected by fish invaders. D.M. 2011. A proposed unified framework for

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biological invasions. Trends in Ecology and littorale during aquatic hypoxia. Comparative grative Zoology, 7(3):227. Evolution, 26(7):333-339. Biochemistry Physiology, 139(1):49-53. http://dx.doi.org/10.1111/j.1749-4877.2012.00311.x http://dx.doi.org/10.1016/j.tree.2011.03.023 http://dx.doi.org/10.1016/j.cbpb.2004.06.019 ROY, K.; MARZORATI, M.; NEGRONI, BRITSKI, H.A.; DE SILIMON, K.Z.S.; JÚLIO JR., H.F.; DEI TÓS, C.; AGOSTINHO, A.; THAS, O.; BALLOI, A.; FAVA, F.; VER- LOPES, B.S. 2007. Peixes do Pantanal: ma- A.A.; PAVANELLI, C.S. 2009. A massive inva- STRAETE, W.; DAFFONCHIO, D.; BOON, nual de identificação. Brasília, Embrapa Infor- sion of fish species after eliminating a natural N. 2013. Environmental conditions and com- mação Tecnológica, 227 p. barrier in the upper rio Paraná basin. Neotropi- munity evenness determine the outcome of BROSSET, A. 1997. Aggressive mimicry by the cal Ichthyology, 7(4):709-718. biological invasion. Nature Communications, characid fish Eythrinus erythrinus. Ethology, http://dx.doi.org/10.1590/S1679-62252009000400021 4(1383):1-5. 103(11):926-934. LANGEANI, F.; CASTRO, R.M.C.; OYA- http://dx.doi.org/10.1038/ncomms2392 http://dx.doi.org/10.1111/j.1439-0310.1997.tb00134.x KAWA, O.T.; SHIBATTA, O.A.; PAVANEL- SUZUKI, H.I.; VAZZOLER, A.E.A.M.; COULATTI, R.I.; MACISAAC, H.J. 2004. A LI, C.S.; CASATTI, L. 2007. Diversidade da MARQUES, E.E.; LIZAMA, M.A.P.; INADA, neutral terminology to define “invasive” species. ictiofauna do Alto Rio Paraná: composição P. 2004. Reproductive ecology of the fish as- Diversity and Distributions, 10(2):135-141. atual e perspectivas futuras. Biota Neotropica, semblage. In: S.M. THOMAZ, A.A. AGOST- http://dx.doi.org/10.1111/j.1366-9516.2004.00061.x 7(3):181-197. INHO, N.S. HAHN (eds.), The Upper Paraná DAJOZ, R. 1978. Ecologia Geral. Rio de Janei- LE CREN, E.D. 1951. The length-weight rela- River and its floodplain: physical aspects, ecol- ro, Editora Vozes, 472 p. tionship and seasonal cycle in gonad weight and ogy and conservation. Leiden, The Netherlands GARCIA, D.A.Z.; COSTA, A.D.A.; YABU, condition in perch Perca fluviatilis. Journal of Backhuys Publishers, p. 271-292. M.H.S.; BALCONI, A.P.R.; ORSI, M.L. 2014a. Animal Ecology, 20:201-219. VAZZOLER, A.E.A.M. 1996. Biologia da re- Sobre como peixes de outras bacias chegam às LEAL, M.E.; KLEIN, G.F.; SCHULZ, U.H.; produção de peixes teleósteos: teoria e prática. nossas águas: o caso do rio Paranapanema, ba- ALBORNOZ, P.L. 2010. First record and eco- Maringá, Eduem, 169 p. cia do alto Paraná. Boletim da Sociedade Bra- logical aspects of Hoplerythrinus unitaeniatus VERANI, J.R.; SATO, Y.; FENERICH-VE- sileira de Ictiologia, 110:8-15. (Agassiz, 1829) (Characiformes, Erythrinidae) RANI, N.; VIEIRA, L.J.S. 1997. Avaliação GARCIA, D.A.Z.; ALMEIDA, F.S.; SILVA E as introduced species in Rio dos Sinos basin, de fêmeas de espécies ícticas aptas à indução SOUZA, Â.T.; BRITTON, J.R.; ORSI, M.L. RS, Brazil. Biota Neotroprica, 10(3):33-37. reprodutiva: critério embasado no fator de con- 2014b. Invasion characteristics of Pterygopli- LOCKWOOD, J.L.; HOOPES, M.F.; MAR- dição relativo. In: VIII Seminário Regional de chthys ambrosettii (Holmberg, 1893) in the CHETTI, M.P. 2007. Invasion Ecology. Oxford, Ecologia, São Carlos, 1997. Anais... São Carlos, lower Paranapanema River, Brazil. Journal of Blackwell Publishing, 312 p. p. 323-329. Applied Ichthyology, 30(5):1041-1044. ORSI, M.L.; AGOSTINHO, A.A. 1999. Intro- VITULE, J.R.S.; UMBRIA, S.C.; ARANHA, http://dx.doi.org/10.1111/jai.12468 dução de espécies de peixes por escapes aci- J.M.R. 2006. Introduction of the African catfish GOMIERO, L.M.; BRAGA, F.M.S. 2003. Re- dentais de tanques de cultivo em rios da Bacia Clarias gariepinus (Burchell, 1822) into South- lação peso-comprimento e fator de condição do Rio Paraná. Revista Brasileira de Zoologia, ern Brazil. Biological Invasions, 8(4):677-681. para Cichla cf. ocellaris e Cichla monoculus 16(2):557-560. http://dx.doi.org/10.1007/s10530-005-2535-8 (Perciformes, Cichlidae) no reservatório de ORSI, M.L. 2010. Estratégias reprodutivas de VITULE, J.R.S.; POZENATO, L.P. 2012. Ho- Volta Grande, Rio Grande - MG/SP. Acta Scien- peixes da região média-baixa do rio Paranapa- mogeneização biótica: misturando organismos tiarum Biological Sciences, 25(1):79-86. nema, Reservatório de Capivara. São Paulo, em um mundo pequeno e globalizado. Estudos http://dx.doi.org/10.4025/actascibiolsci.v25i1.2119 Blucher Acadêmico, 115 p. de Biologia, 34(83):239-245. GOMIERO, L.M.; VILLARES JR., G.A.; ORSI, M.L.; BRITTON, J.R. 2012. Length- http://dx.doi.org/10.7213/estud.biol.7336 BRAGA, F.M.S. 2010. Relação peso-com- weight relationships of 15 fishes of the Capi- VITULE, J.R.S.; PRODOCIMO, V. 2012. In- primento e fator de condição de Oligosarcus vara Reservoir (Paranapanema basin, Brazil). trodução de espécies não nativas e invasões bi- hepsetus (Cuvier, 1829) no Parque Estadual Journal of Applied Ichthyology, 28(1):146-147. ológicas. Estudos de Biologia, 34(83):225-237. da Serra do Mar - Núcleo Santa Virgínia, Mata http://dx.doi.org/10.1111/j.1439-0426.2011.01891.x http://dx.doi.org/10.7213/estud.biol.7335 Atlântica, estado de São Paulo, Brasil. Biota ORSI, M.L.; BRITTON, J.R. 2014. Long-term XU, H.; CHEN, K.; OUYANG, Z.; PAN, X.; Neotropica, 10(1):101-105. changes in the fish assemblage of a neotropical ZHU, S. 2012. Threats of invasive species GOZLAN, R.E.; BRITTON, J.R.; COWX, hydroelectric reservoir. Journal of Fish Biology, for China caused by expanding international I.; COPP, G.H. 2010. Current knowledge on 84(6):1964-1970. trade. Environmental Science & Technology, non-native freshwater fish introductions. Jour- http://dx.doi.org/10.1111/jfb.12392 46(13):7063-7064. nal of Fish Biology, 76(4):751-786. OYAKAWA, O.T. 2003. Family Erythrinidae. http://dx.doi.org/10.1021/es301996x http://dx.doi.org/10.1111/j.1095-8649.2010.02566.x In: R. REIS; S.O. KULLANDER; C.J. FER- ZANATTA, A.S.; RAMOS, I.P.; SILVA, R.J.; GRAÇA, W.J.; PAVANELLI, C.S. 2007. Peixes RARIS JR. (eds.), Check list of the freshwater LANGEANI, F.; CARVALHO, E.D. 2010. da planície de inundação do alto rio Paraná e fishes of South and Central America. Porto Ale- Pisces, Siluriformes, Ictaluridae, Ictalurus áreas adjacentes. Maringá, Eduem, 241 p. gre, Editora da Pontifícia Universidade Católica punctatus (Rafinesque, 1818): First record in HOLLE, B.V.; SIMBERLOFF, D. 2005. Eco- do Rio Grande do Sul, p. 241-244. middle Paranapanema river reservoir, aquacul- logical resistance to biological invasion over- OYAKAWA, O.T.; MATTOX, G.M.T. 2009. Re- ture and exotic species dispersion. Check List, whelmed by propagule pressure. Ecology, vision of Neotropical trahiras of the Hoplias lacer- 6(4):589-591. 86(12):3212-3218. dae species-group (Ostariophysi: Characiformes: http://dx.doi.org/10.1890/05-0427 Erythrinidae) with descriptions of two new spe- JUCÁ-CHAGAS, R. 2004. Air breathing of cies. Neotropical Ichthyology, 7(2):117-140. the neotropical fishes Lepidosiren paradoxa, http://dx.doi.org/10.1590/S1679-62252009000200001 Submitted on January 15, 2015 Hoplerythrinus unitaeniatus and Hoplosternum ROQUES, A. 2012. Biological Invasion. Inte- Accepted on August 31, 2015

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