Relações tróficas em assembleia de peixes em lagoas de planície de inundação Neotropical: seletividade e sobreposição alimentar mediada pela disponibilidade de recursos alimentares

As relações entre o grau de sobreposição alimentar e disponibilidade de alimentos, e as implicações para a seletividade alimentar de espécies de peixes foram avaliadas em lagoas da planície de inundação do Alto rio Paraná, Brasil. As hipóteses testadas foram: i) as espécies tornam-se menos seletivas em lagoas com alta disponibilidade de recursos alimentares; e ii) espécies (interespecífica) ou indivíduos (intraspecífica) apresentam maior sobreposição alimentar em condições de alta disponibilidade de recursos alimentares. Em geral, com os resultados observou-se que as espécies se tornam menos seletivas quando o ambiente proporciona maior disponibilidade de recursos alimentares. A sobreposição interespecífica não demostrou padrão quando avaliado a disponibilidade de recursos alimentares nas lagoas. No entanto, a sobreposição intraespecífica foi mais acentuada nas condições de elevada disponibilidade de recursos alimentares.

PALAVRAS-CHAVE
Alimentação de peixes; eletividade; rio Paraná

Autoria

Rosa Maria Dias

Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura (Nupélia), Departamento de Biologia, Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900, Maringá, PR, Brazil. (rmdias2003@yahoo.com.br)Universidade Estadual de MaringáBrazilMaringá, PR, BrazilNúcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura (Nupélia), Departamento de Biologia, Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900, Maringá, PR, Brazil. (rmdias2003@yahoo.com.br)

Jean Carlo Gonçalves Ortega

Luiz Carlos Gomes

Angelo Antonio Agostinho

SCIMAGO INSTITUTIONS RANKINGS

Figuras | Tabelas

Figuras (5)
Tabelas (4)

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Fig. 1
Locations of the lakes on the upper Paraná River floodplain, Brazil: 1, Canal do Meio; 2, Carioca; 3, Zé Marinho; 4, Cidão; 5, Genipapo; 6, Tião.

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Fig. 2
Ordination by principal coordinate analysis (PCoA) of the food resource availability for six floodplain lakes along the Upper Paraná River, Paraná-Mato Grosso do Sul. AQI = aquatic insects; OAI = other aquatic invertebrates; OTI = other terrestrial invertebrates; PLA = plants; TRI = terrestrial insects.

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Fig. 3
Relationships between feeding selectivity by fish and the availability of food resources for six floodplain lakes along the Upper Paraná River, Paraná-Mato Grosso do Sul. Shape of data distribution (envelope effect) was significant.

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Fig. 4
Relationship of the mean the Schoener's Index (O) between pairs of species and the scores of the first PCoA axis of resource availability in isolated floodplain lakes along the upper Paraná River. The mean O was calculated based on 10 (Zé = Zé Marinho), 28 (Carioca = Car), 6 (Tião = Tia), 21 (Genipapo = Gen), 3 (Cidão = Cid) and 10 (Canal = Can) pairs of species. AQI = aquatic insects; PLA = plants.

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Fig. 5
Relationship between the mean of the proportional overlap Index (IS) and the scores of the first PCoA axis of resource availability in isolated floodplain lakes along the upper Paraná River.Values of IS closer to 1 indicates greater diet overlap. The mean IS was calculated based on individuals of 3 (Zé = Zé Marinho), 7 (Carioca = Car), 4 (Tião = Tia), 5 (Genipapo = Gen), 2 (Cidão = Cid) and 5 species (Canal = Can). AQI = aquatic insects; PLA = plants.

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Tab. I
Range (Max = maximum; Min = minimum) of standard length (Ls; cm) and abundance (individuals.m^-2) of the species captured in the study lakes (CA, Carioca; GE, Genipapo; ZM, Zé Marinho; CI, Cidão; TI, Tião; CM, Canal do Meio). †, values below 0.005

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Tab. II
Food resources identified for each species at the six isolated lakes of the upper Paraná River floodplain. N = total number of species caught, n = number of stomachs analyzed (ALG, Algae; DET, Detritus/sediment; AQI, Aquatic insects; ITR, Terrestrial insects; OAI, Other aquatic invertebrates; OTI, Other terrestrial invertebrates; MIC, Microcrustaceans; FIS, Fish; PLA, Plants).

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Tab. III
Principal coordinate analysis (PCoA) that summarized the food resource matrix (food availability) of the sampled lakes. It is presented the Pearson correlations of the scores of each axis with the original variables, the eigenvalue and the percentage of explanation of each axis. Variables with significant Pearson correlations are in bold.

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Tab. IV
Food selection of the ichthyofauna and availability of resources in six floodplain lakes of the upper Paraná River (ALG, Algae; PLA, Higher plants; AQI, Aquatic insects; ITR, Terrestrial insects; OAI, Other aquatic invertebrates; OTI, Other terrestrial invertebrates; MIC, Microcrustaceans; FIS, Fish; DET, Detritus/sediment).

Fig. 1 Locations of the lakes on the upper Paraná River floodplain, Brazil: 1, Canal do Meio; 2, Carioca; 3, Zé Marinho; 4, Cidão; 5, Genipapo; 6, Tião.

Fig. 2 Ordination by principal coordinate analysis (PCoA) of the food resource availability for six floodplain lakes along the Upper Paraná River, Paraná-Mato Grosso do Sul. AQI = aquatic insects; OAI = other aquatic invertebrates; OTI = other terrestrial invertebrates; PLA = plants; TRI = terrestrial insects.

Fig. 3 Relationships between feeding selectivity by fish and the availability of food resources for six floodplain lakes along the Upper Paraná River, Paraná-Mato Grosso do Sul. Shape of data distribution (envelope effect) was significant.

Fig. 4 Relationship of the mean the Schoener's Index (O) between pairs of species and the scores of the first PCoA axis of resource availability in isolated floodplain lakes along the upper Paraná River. The mean O was calculated based on 10 (Zé = Zé Marinho), 28 (Carioca = Car), 6 (Tião = Tia), 21 (Genipapo = Gen), 3 (Cidão = Cid) and 10 (Canal = Can) pairs of species. AQI = aquatic insects; PLA = plants.

Fig. 5 Relationship between the mean of the proportional overlap Index (IS) and the scores of the first PCoA axis of resource availability in isolated floodplain lakes along the upper Paraná River.Values of IS closer to 1 indicates greater diet overlap. The mean IS was calculated based on individuals of 3 (Zé = Zé Marinho), 7 (Carioca = Car), 4 (Tião = Tia), 5 (Genipapo = Gen), 2 (Cidão = Cid) and 5 species (Canal = Can). AQI = aquatic insects; PLA = plants.

Tab. I Range (Max = maximum; Min = minimum) of standard length (Ls; cm) and abundance (individuals.m^-2) of the species captured in the study lakes (CA, Carioca; GE, Genipapo; ZM, Zé Marinho; CI, Cidão; TI, Tião; CM, Canal do Meio). †, values below 0.005

Species	Ls Max-Min	Abundance
Species	Ls Max-Min	CA	GE	ZM	CI	TI	CM	Total
Characiformes
Astyanax altiparanae Garutti & Britski, 2000	8.0 -1.7	1.08	0.62	0.23	4.21	1.14	†	7.28
Aphyocharax anisitsi Eigenmann & Kennedy, 1903	3.7 - 1.1		0.07	0.04	0.02	0.29	4.06	4.48
Cyphocharax modestus (Fernández-Yépez, 1948)	4.3 - 4.2	0.02						0.02
Erythrinus erythrinus (Bloch & Schneider, 1801)	14.0 - 8.2	0.01			0.01			0.02
Hemigrammus marginatus Ellis, 1911	3.5 - 2.7		0.11	0.06				0.17
Hyphessobrycon eques (Steindachner, 1882)	3.5 - 0.9			0.01			0.53	0.54
Hopliasaff.malabaricus (Bloch, 1794)	5.8 - 1.8		0.02			†	0.05	0.07
Hoplerythrinus unitaeniatus (Agassiz, 1829)	15.7 - 13.5	0.02						0.02
Moenkhausia intermedia Eigenmann, 1908	3.4 - 1.6	0.01	0.16					0.17
Moenkhausia sanctaefilomenae (Steindachner, 1907)	3.9 - 0.9	0.11	1.22	0.04	0.56	1.12		3.05
Pyrrhulina australis Eigenmann & Kennedy, 1903	3.0 - 1.6	0.22	0.04					0.26
Roeboides descalvadensis Fowler, 1932	5.5 - 1.7	0.09	0.21	0.03				0.33
Serrasalmus marginatus Valenciennes, 1837	3.8 - 2.2	0.01	0.07					0.08
Serrapinnus notomelas (Eigenmann, 1915)	3.6 - 1.2		0.08	0.01		0.08	0.97	1.14
Serrapinus sp.1	1.4					0.02		0.02
Steindachnerina brevipinna (Eigenmann & Eigenmann, 1889)	5.6 - 1.5	0.56	0.85		0.07	0.08		1.56
Steindachner insculpta (Fernández-Yépez, 1948)	3.4 - 1.5		0.12					0.12
Perciformes
Astronotus crassipinnis (Heckel, 1840)	9.0 - 7.8		0.04					0.04
Cichlasoma paranaense Kullander, 1983	8.3 - 1.3	0.21	0.11			0.04	0.05	0.41
Crenicichla britskii Kullander, 1982	8.6		0.02					0.02
Crenicichla niederleinii (Holmberg, 1891)	2.3						0.01	0.01
Cichla kelberi Kullander & Ferreira, 2006	15.6		0.02					0.02
Satanoperca pappaterra (Heckel, 1840)	6.8 - 5.4		0.02	0.01				0.03
Laetacara sp.	4.8- 1.0	0.12		0.36		0.01	2.12	2.61
Siluriformes
Hoplosternum littorale (Hancock, 1828)	10.8 - 0.8	0.04	0.03		0.03	0.06	0.08	0.24
Pterygoplichthys ambrosettii (Holmberg, 1893)	11.3 - 1.2		0.05	0.03				0.08
Pimelodella sp.1	2.1		0.01					0.01
Loricariichthys platymetopon Isbrücker & Nijssen, 1979	15.1 - 7.3		0.08				†	0.08
Gymnotiformes
Eigenmannia trilineata López & Castello, 1966	9.3 - 5.1	0.01	0.02					0.03
Gymnotus sp.	17.7	0.13					0.01	0.14
Cyprinodontiformes
Poecilia reticulata Peters, 1859	2.1 - 1.4	0.04						0.04
Melanorivulus apiamici (Costa, 1989)	2.4 - 1.3	0.17						0.17
Synbranchiformes
Synbranchus marmoratus Bloch, 1795	48.2 - 7.8	0.03				0.01	0.13	0.17
Number of species		18	22	10	6	11	12	33

Tab. II Food resources identified for each species at the six isolated lakes of the upper Paraná River floodplain. N = total number of species caught, n = number of stomachs analyzed (ALG, Algae; DET, Detritus/sediment; AQI, Aquatic insects; ITR, Terrestrial insects; OAI, Other aquatic invertebrates; OTI, Other terrestrial invertebrates; MIC, Microcrustaceans; FIS, Fish; PLA, Plants).

Lakes/species	N	n	Food resources (% Volume)
Lakes/species	N	n	ALG	DET	AQI	ITR	OAI	OTI	MIC	FIS	PLA
Carioca
Astyanax altiparanae	116	20	6.46	9.33	2.66	26.04	0.96		0.07		54.47
Moenkhausia intermedia	1	1	56.37	22.41	18.87		2.36
Moenkhausia sanctaefilomenae	12	3	14.16	24.99		22.72	2.53		1.51		34.08
Erythrinus erythrinus	2	1			57.92		3.47	38.61
Eigenmannia trilineata	1	1			99.59				0.41
Gymnotus spp.	19	8	0.07		90.19	1.76	0.46		7.52
Cichlasoma paranaense	30	2	7.69	30.77	23.08	38.46
Pyrrhulina australis	32	8	1.25		54.55	28.21	1.25	6.27	8.46
Roeboides descalvadensis	10	7	1.68	11.49	78.02	6.29	2.52
Melanorivulus apiamici	26	3			43.48	34.78			21.74
Hoplosternum littorale	4	4	5.61	50.36	5.54	11.40	3.91		23.17
Synbranchus marmoratus	4	1			38.27	1.33	26.55			22.12	11.73
Cyphocharax modestus	2	1	10.80	85.40			3.80
Steindachnerina brevipinna	61	11	10.91	84.50			4.59
Laetacara sp.	18	1			23.08		7.69		69.23
Serrasalmus marginatus	1	1								100
Genipapo
Astyanax altiparanae	33	12			31.85	61.93		4.66		0.44*	1.11
Moenkhausia sanctaefilomenae	73	19			42.66	23.26			0.19		33.90
Pyrrhulina australis	3	1			100
Roeboides descalvadensis	11	6			85.59			14.41
Eigenmannia trilineata	1	1			50.00				50.00
Aphyocharax anisitsi	5	2			60.00				40.00
Moenkhausia intermedia	8	2	9.62	30.73	41.42		2.67		15.56
Hoplias malabaricus	1	1					100
Loricariichthys platymetopon	4	4	19.10	76.29			4.62
Pterygoplichthys ambrosettii	3	1	9.83	90.04			0.13
Steindachnerina brevipinna	44	11	33.04	63.41			3.55
Steindachnerina insculpta	6	3	33.54	56.34			10.12
Hoplosternum littorale	2	1							100
Cichlasoma paranaense	7	2				1.01				98.99
Serrasalmus marginatus	4	3								100
Astronotus crassipinnis	2	1	7.80				0.46				91.74
Zé Marinho
Astyanax altiparanae	31	22	0.65		3.44	6.18	13.10		1.36	0.84*	74.43
Serrapinnus notomelas	2	1	3.33						13.33		83.33
Hemigrammus marginatus	8	2							91.80	8.20*
Aphyocharax anisitsi	7	3			38.24				61.76
Laetacara sp.	68	12	0.71		8.50	19.26	69.41		0.71	1.42*
Moenkhausia sanctaefilomenae	8	5	1.54		27.16	36.42		9.26			25.62
Roeboides paranensis	4	2			49.33	30.67				20.0*
Pterygoplichthys ambrosettii	6	3	22.96	76.80			0.25
Cidão
Astyanax altiparanae	187	48	0.02	0.15	60.17	6.87	0.13	2.87	3.89	1.25*	24.65
Erythrinus erythrinus	4	1			52.94	3.49	26.14	17.43
Moenkhausia sanctaefilomenae	109	58	0.01	0.26	77.73	3.20	0.12	0.54	7.80	0.05*	10.29
Hoplosternum littorale	3	1			26.67				73.33
Steindachnerina brevipinna	4	2	38.91	61.09
Tião
Astyanax altiparanae	215	68	0.22		29.07	12.04	0.34	1.60	12.11	0.42*	44.19
Aphyocharax anisitsi	19	7	0.02		29.47		0.18		66.69		3.64
Steindachnerina brevipinna	7	4	4.00	95.85			0.15
Hoplosternum littorale	3	1	4.36			20.56	0.48		48.23		26.37
Hoplias aff. malabaricus	2	2			100
Laetacara sp.	6	1			100
Moenkhausia sanctaefilomenae	206	75	0.04		53.61	18.18	0.64	0.49	6.80	0.04*	20.20
Synbranchus marmoratus	4	2			37.74	58.18	3.30				0.79
Canal do Meio
Astyanax altiparanae	1	1	99.90				0.10
Crenicichla niederleinii	1	1	64.29								35.71
Laetacara sp.	319	5	40.49		10.71		10.39		16.37		22.04
Gymnotus spp.	2	1			98.94				1.06
Hyphessobrycon eques	81	1			100
Synbranchus marmoratus	19	18			97.90	0.57	1.52
Hoplias aff. malabaricus	8	2			1.72					98.28
Loricariichthys platymetopon	1	1	8.91	65.32			0.36		25.42
Hoplosternum littorale	15	5	32.18	5.09			18.52		21.30	22.9*
Serrapinnus notomelas	177	30	49.21	0.76	2.78		41.35		5.90
Aphyocharax anisitsi	684	72	9.83	0.43	40.84		36.47		12.21		0.21

Tab. III Principal coordinate analysis (PCoA) that summarized the food resource matrix (food availability) of the sampled lakes. It is presented the Pearson correlations of the scores of each axis with the original variables, the eigenvalue and the percentage of explanation of each axis. Variables with significant Pearson correlations are in bold.

Variable	PCoA 1	PCoA 2
ALG	0.343	-0.458
PLA	-0.946	-0.031
AQI	0.857	0.015
ITR	-0.007	0.842
OAI	-0.073	-0.872
OTI	0.297	0.944
MIC	0.540	-0.590
FIS	-0.169	0.636
DET	-0.234	0.673
Eigenvalue	0.351	0.247
Percent of explanation	44.32	33.28

Tab. IV Food selection of the ichthyofauna and availability of resources in six floodplain lakes of the upper Paraná River (ALG, Algae; PLA, Higher plants; AQI, Aquatic insects; ITR, Terrestrial insects; OAI, Other aquatic invertebrates; OTI, Other terrestrial invertebrates; MIC, Microcrustaceans; FIS, Fish; DET, Detritus/sediment).

Lake/Food resources	ALG	PLA	AQI	ITR	OAI	OTI	MIC	FIS	DET
Carioca
Astyanax altiparanae	0.10	0.27	-0.75	0.23	-0.61	-1	-0.94	-1	-0.32
Gymnotus sp.	-0.97	-1	0.65	-0.81	-0.80	-1	0.55	-1	-1
Pyrrhulina australis	-0.62	-1	0.49	0.26	-0.52	0.60	0.59	-1	-1
Roeboides descalvadensis	-0.52	-1	0.61	-0.45	-0.23	-1	-1	-1	-0.22
Steindachnerina brevipinna	0.34	-1	-1	-1	0.07	-1	-1	-1	0.65
Hoplosternum littorale	0.03	-1	-0.54	-0.18	-0.01	-1	0.83	-1	0.47
Moenkhausia sanctaefilomenae	0.45	0.04	-1	0.16	-0.23	-1	-0.18	-1	0.16
Melanorivulus apiamici	-1	-1	0.40	0.36	-1	-1	0.82	-1	-1
Availability	5.34	36.41	18.8	16.46	4.01	1.58	2.16	2.29	12.93
Genipapo
Astyanax altiparanae	-1	-0.81	0.01	0.19	-1	0.26	-1	-0.71	-1
Moenkhausia sanctaefilomenae	-1	0.52	0.15	-0.29	-1	-1	0.19	-1	-1
Roeboides descalvadensis	-1	-1	0.46	-1	-1	0.68	-1	-1	-1
Steindachnerina brevipinna	0.82	-1	-1	-1	0.75	-1	-1	-1	0.81
Loricariichthys platymetopon	0.70	-1	-1	-1	0.80	-1	-1	-1	0.84
Steindachnerina insculpta	0.82	-1	-1	-1	0.91	-1	-1	-1	0.79
Serrasalmus marginatus	-1	-1	-1	-1	-1	-1	-1	0.95	-1
Availability	3.31	10.73	31.39	42.00	0.50	2.73	0.13	2.60	6.62
Zé Marinho
Astyanax altiparanae	-0.03	0.08	-0.14	-0.15	-0.23	-1	0	-0.11	-1
Laetacara sp.	0.02	-1	0.30	0.40	0.54	-1	-0.31	0.16	-1
Aphyocharax anisitsi	-1	-1	0.79	-1	-1	-1	0.96	-1	-1
Pterygoplichthys ambrosettii	0.94	-1	-1.00	-1	-0.98	-1	-1	-1	1
Moenkhausia sanctaefilomenae	0.39	-0.42	0.71	0.63	-1	0.99	-1	-1	-1
Availability	0.69	63.02	4.57	8.32	20.88	0.05	1.35	1.04	0.09
Cidão
Astyanax altiparanae	-0.09	0.21	-0.08	0.19	-0.39	0.28	-0.22	0.40	-0.23
Moenkhausia sanctaefilomenae	-0.45	-0.22	0.05	-0.19	-0.43	-0.50	0.12	-0.84	0.06
Steindachnerina brevipinna	1	-1	-1	-1	-1	-1	-1	-1	0.99
Availability	0.03	16.08	70.36	4.70	0.30	1.61	6.15	0.54	0.23
Tião
Astyanax altiparanae	0.22	0.14	-0.16	-0.10	-0.17	0.18	0.11	0.25	-1
Aphyocharax anisitsi	-0.73	-0.80	-0.15	-1	-0.46	-1	0.74	-1	-1
Moenkhausia sanctaefilomenae	-0.58	-0.25	0.15	0.10	0.14	-0.39	-0.18	-0.70	-1
Steindachnerina brevipinna	0.93	-1	-1	-1	-0.52	-1	-1	-1	1
Availability	0.14	33.49	39.9	14.85	0.48	1.11	9.77	0.25	0.01
Canal do Meio
Aphyocharax anisitsi	-0.17	-0.50	0.01	-1	0.04	-	0.06	-1	-0.01
Hoplosternum littorale	0.40	-1	-1	-1	-0.29	-	0.33	0.92	0.84
Laetacara sp.	0.49	0.94	-0.58	-1	-0.53	-	0.21	-1	-1
Synbranchus marmoratus	-1	-1	0.42	0.88	-0.91	-	-1	-1	-1
Serrapinnus notomelas	0.56	-1	-0.87	-1	0.10	-	-0.29	-1	0.27
Availability	13.80	0.73	39.79	0.04	33.6	0	10.78	0.92	0.34

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Relações tróficas em assembleia de peixes em lagoas de planície de inundação Neotropical: seletividade e sobreposição alimentar mediada pela disponibilidade de recursos alimentares

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