A heterogeneidade ambiental explica o turnover de espécies, mas não seu aninhamento em assembleias de peixes de uma bacia Neotropical

Peláez, Oscar Eduardo; Azevedo, Filipe Manoel; Pavanelli, Carla Simone

Oscar Eduardo Peláez ^**e-mail: oscar_pelaez@hotmail.com

Programa de Pós-graduação em Ecologia de Ambientes Aquáticos Continentais, Universidade Estadual de Maringá – UEM, Av. Colombo, 5790, CEP 87020-900, Maringá, PR, Brazil

Filipe Manoel Azevedo

Programa de Graduação em Ciências Biológicas, Universidade Estadual de Maringá – UEM, Av. Colombo, 5790, CEP 87020-900, Maringá, PR, Brazil

Carla Simone Pavanelli

Programa de Pós-graduação em Ecologia de Ambientes Aquáticos Continentais, Universidade Estadual de Maringá – UEM, Av. Colombo, 5790, CEP 87020-900, Maringá, PR, Brazil

Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura – Nupélia, Universidade Estadual de Maringá – UEM, Av. Colombo, 5790, CEP 87020-900, Maringá, PR, Brazil

*e-mail: oscar_pelaez@hotmail.com

Figuras (4)
Tabelas (2)

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Figure 1
Sampling sites in the Upper Paraná River floodplain. Paraná River in grey. (1) Paranapanema; (2) Baía; (3) Ivinhema; (4) Ivinheminha; (5) Ivaí; (6) Amambai; (7) Maracaí; (8) Paracaí; (9) Pirajuí; (10) São João lagoon; (11) Xambrê lagoon; (12) Iguatemi; (13) Saraiva lagoon. Sampling sites in the main channel of the Paraná River were located upstream from each point in the map (except for 10 and 11).

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Figure 2
Sample-size-based rarefaction (solid lines) and extrapolation (dashed lines) of fish diversity at the Upper Paraná River (Hill numbers of order 0, 1 and 2). q = 0: species richness; q = 1: Shannon diversity; q = 2: inverse of Simpson’s concentration. Blue: tributaries; red: Paraná River.

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Figure 3
Non-metric multidimensional ordination of species composition of fish assemblages at the Upper Paraná River. Stress = 0.1495. Squares: sites in the northern portion of the sampled region (Paranapanema, Baía and Ivinhema rivers); triangles: sites in the middle portion of the sampled region (Ivinheminha, Ivaí and Amambaí rivers); diamonds: sites on the left side of the sampled region (Paracaí, São João and Xambrê); circles: sites in the southern and right side of the sampled region (Maracaí, Pirajuí and Iguatemi rivers). See species names in .

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Figure 4
Distance-based Redundancy Analysis (dbRDA) of species assemblages and environmental variables in the Upper Paraná River. (a) Total Bray Curtis; (b) Turnover and (c) Nestedness. Circles: Paraná River sites; Triangles: Tributaries. TEMP: water temperature; DO: dissolved oxygen; COND: conductivity; DEPTH: mean depth.

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Table 1
Fish species from the upper Paraná River, collected between August 2013 and May 2015. Abbreviations as in NMDS ordination (). NA: Not analyzed (species with only one individual, generally a juvenile). Classification follows Eschmeyer & Fong (2017)ESCHMEYER, J.D. and FONG, R. Species by family/subfamily [online]. 2017 [viewed 25 Oct. 2017]. Available from: http:// researcharchive.calacademy.org/research/ichthyology/catalog/SpeciesByFamily.asp, excepting Characidae subfamilies, which follow Mirande (2010)MIRANDE, J.M. Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology, 2010, 8(3), 385-568. http://dx.doi.org/10.1590/S1679-62252010000300001.
http://dx.doi.org/10.1590/S1679-62252010... .

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Table 2
Partition of beta diversity (mean and standard deviation of Bray Curtis dissimilarity) for fish assemblages and its relationship with environmental variables (R² and P-values) in the Upper Paraná River. TEMP: water temperature; DO: dissolved oxygen; COND: conductivity; DEPTH: mean depth. Significant relationships in bold.

MYLIOBATIFORMES	CHARACIFORMES
Potamotrygonidae	Tetragonopterinae
Potamotrygon amandae Loboda & Carvalho, 2013 (P.AMA)	Hyphessobrycon eques (Steindachner, 1882) (H.EQU)
CLUPEIFORMES	Hemigrammus ora Zarske, Le Bail & Géry, 2006 (H.ORA)
Clupeidae	Moenkhausia bonita Benine, Castro & Sabino, 2004 (M.BON)
Platanichthys platana (Regan, 1917) (P.PLA)	Moenkhausia forestii Benine, Mariguela & Oliveira, 2009 (M.FOR)
CHARACIFORMES	Moenkhausia gracilima Eigenmann, 1908 (M.GRA)
Crenuchidae	Moenkhausia aff. intermedia Eigenmann, 1908 (M.INT)
Characidium aff. zebra Eigenmann, 1909 (C.ZEB)	Clado Astyanax
Erythrinidae	Astyanax aff. fasciatus (Cuvier, 1819) (A.FAS)
Hoplias mbigua Azpelicueta, Benítez, Aichino & Mendez, 2015 (H.MBI)	Astyanax lacustris (Lütken, 1875) (A.LAC)
Hoplias sp. (Hoplias sp.)	Psellogrammus kennedyi (Eigenmann, 1903) (P.KEN)
Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829) (NA)	Stevardiinae
Parodontidae	Diapoma guarani Mahnert & Géry, 1987 (D.GUA)
Apareiodon affinis (Steindachner, 1879) (A.AFI)	Bryconamericus stramineus Eigenmann, 1908 (B.STR)
Cynodontidae	Knodus moenkhausii (Eigenmann & Kennedy, 1903) (K.MOE)
Rhaphiodon vulpinus Spix & Agassiz, 1829 (NA)	SILURIFORMES
Serrasalmidae	Auchenipteridae
Mylossoma duriventre (Cuvier, 1818) (M.DUR)	Parauchenipterus galeatus (Linnaeus, 1766) (P.GAL)
Metynnis lippincottianus (Cope, 1870) (M.LIP)	Auchenipterus osteomystax (Miranda Ribeiro, 1918) (A.OST)
Serrasalmus maculatus Kner, 1858 (S.MACU)	Doradidae
Serrasalmus marginatus Valenciennes, 1837 (S.MAR)	Trachydoras paraguayensis (Eigenmann & Ward, 1907) (T.PAR)
Hemiodontidae	Heptapteridae
Hemiodus orthonops Eigenmann & Kennedy, 1903 (H.ORT)	Pimelodella avanhandavae Eigenmann, 1917 (P.AVA)
Anostomidae	Pimelodella gracilis (Valenciennes, 1835) (P.GRA)
Leporinus friderici (Bloch, 1794) (L.FRI)	Pimelodella taenioptera Miranda Ribeiro, 1914 (NA)
Megaleporinus piavussu (Britski, Birindelli & Garavello, 2012BRITSKI, H.A., BIRINDELLI, J.L.O. and GARAVELLO, J.C. A new species of Leporinus Agassiz, 1829 from the upper Rio Paraná basin (Characiformes, Anostomidae) with redescription of L. elongatus Valenciennes, 1850 and L. obtusidens (Valenciennes, 1837). Papéis Avulsos de Zoologia, 2012, 52(37), 441-475.) (L.PIA)	Pimelodidae
Schizodon borellii (Boulenger, 1900) (S.BOR)	Sorubim lima (Bloch & Schneider, 1801) (S.LIM)
Curimatidae	Hemisorubim platyrhynchos (Valenciennes, 1840) (H.PLA)
Steindachnerina brevipinna (Eigenmann & Eigenmann, 1889) (S.BRE)	Pimelodus maculatus Lacepède, 1803 (P.MAC)
Steindachnerina insculpta (Fernández-Yépez, 1948) (S.INS)	Pimelodus mysteriosus Azpelicueta, 1998 (P.MYS)
Prochilodontidae	Pimelodus ornatus Kner, 1858 (NA)
Prochilodus lineatus (Valenciennes, 1837) (P.LIN)	Iheringichthys labrosus (Lütken, 1874) (I.LAB)
Bryconidae	Loricariidae
Salminus hilarii Valenciennes, 1850 (NA)	Loricariichthys platymetopon Isbrücker & Nijssen, 1979 (L.PLA)
Characidae	Loricariichthys rostratus Reis & Pereira, 2000 (L.ROS)
Aphyocharacinae	Loricaria prolixa Isbrücker & Nijssen, 1978 (L.PRO)
Aphyocharax anisitsi Eigenmann & Kennedy, 1903 (A.ANI)	Pterygoplichthys ambrosettii (Holmberg, 1893) (P.AMB)
Aphyocharax dentatus Eigenmann & Kennedy, 1903 (A.DEN)	Hypostomus ancistroides (Ihering, 1911) (NA)
Cheirodontinae	Hypostomus commersoni Valenciennes, 1836 (NA)
Serrapinnus calliurus (Boulenger, 1900) (S.CAL)	GYMNOTIFORMES
Serrapinnus heterodon (Eigenmann, 1915) (S.HET)	Rhamphicthyidae
Serrapinnus notomelas (Eigenmann, 1915) (S.NOT)	Gymnorhamphichthys britskii Carvalho, Ramos & Albert, 2011 (G.HYP)
GYMNOTIFORMES	PERCIFORMES
Rhamphicthyidae	Sciaenidae
Ramphichthys hahni (Meinken, 1937) (R.HAH)	Plagioscion squamosissimus (Heckel, 1840) (P.SQU)
Sternopygidae	Cichlidae
Eigenmannia guairaca Peixoto, Dutra & Wosiacki, 2015 (E.TRIL)	Cichla kelberi Kullander & Ferreira, 2006 (C.KEL)
Eigenmannia virescens (Valenciennes, 1836) (E.VIR)	Geophagus sveni Lucinda, Lucena & Assis, 2010 (G.PRO)
Sternopygus macrurus (Bloch & Schneider, 1801) (S.MACR)	Satanoperca pappaterra (Heckel, 1840) (S.PAP)
Apteronotidae	Apistogramma cf. commbrae (Regan, 1906) (A.COM)
Apteronotus ellisi (Alonso de Arámburu, 1957) (A.ELL)	Aequidens plagiozonatus Kullander, 1984 (A.PLA)
CYPRINODONTIFORMES	Laetacara araguaiae Ottoni & Costa, 2009 (NA)
Poeciliidae	Crenicichla britskii Kullander, 1982 (C.BRI)
Pamphorichthys sp. (PAMPH)	PLEURONECTIFORMES
Phalloceros harpagos Lucinda, 2008 (NA)	Achiridae
	Catathyridium jenynsii (Günther, 1862) (C.JEN)

	Bray Curtis	TEMP	DO	COND	TURB	pH	DEPTH
	Total	R² (P values)
All sites	0.71 ± 0.18	0.06 (0.07)	0.04 (0.33)	0.07 (0.05)	0.10 (0.01)	0.08 (0.04)	0.05 (0.19)
Paraná River	0.66 ± 0.25	0.15 (0.27)	0.07 (0.58)	0.20 (0.18)	0.06 (0.75)	0.04 (0.85)	0.02 (0.95)
Tributaries	0.72 ± 0.14	0.11 (0.09)	0.09 (0.18)	0.08 (0.27)	0.14 (0.03)	0.13 (0.04)	0.11 (0.07)
	Turnover
All sites	0.41 ± 0.19	0.06 (0.16)	0.02 (0.59)	0.18 (<0.01)	0.15 (<0.01)	0.03 (0.46)	0.10 (0.03)
Paraná River	0.27 ± 0.20	0.07 (0.54)	0.04 (0.62)	0.19 (0.34)	0.03 (0.65)	0.08 (0.54)	0.07 (0.55)
Tributaries	0.47 ± 0.20	0.10 (0.11)	0.10 (0.12)	0.12 (0.07)	0.22 (0.01)	0.07 (0.21)	0.20 (0.02)
	Nestedness
All sites	0.30 ± 0.22	0.08 (022)	0.06 (0.34)	0.05 (0.98)	0.08 (0.22)	0.13 (0.11)	0.02 (0.94)
Only Paraná	0.38 ± 0.24	0.15 (0.32)	0.11 (0.41)	0.16 (0.32)	0.10 (0.48)	0.01 (0.92)	0.04 (0.68)
Tributaries	0.23 ± 0.18	0.15 (0.27)	0.09 (0.43)	0.01 (0.81)	0.11 (0.35)	0.23 (0.18)	0.01 (0.92)

[1] *e-mail: oscar_pelaez@hotmail.com

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A heterogeneidade ambiental explica o turnover de espécies, mas não seu aninhamento em assembleias de peixes de uma bacia Neotropical

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