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RESEARCH ARTICLE • Zoologia (Curitiba) 36 • 2019 • https://doi.org/10.3897/zoologia.36.e23539 copy

Influences of the area, shape and connectivity of coastal lakes on the taxonomic and functional diversity of fish communities in Southern Brazil

Authorship SCIMAGO INSTITUTIONS RANKINGS

ABSTRACT

In this study we investigated the influence of landscape variables on the alpha taxonomic and functional diversity of fish communities in coastal lakes. We built an analytical framework that included possible causal connections among variables, which we analyzed using path analysis. We obtained landscape metrics for the area, shape and connectivity (estuary connectivity and primary connectivity to neighboring lakes) of 37 coastal lakes in the Tramandaí River Basin. We collected fish data from 49 species using standardized sampling with gillnets and obtained a set of traits related to dispersal abilities and food acquisition. The model that best explained the taxonomic diversity and functional richness took into account the shape of the lakes. Functional richness was also explained by estuary connectivity. Functional evenness and dispersion were not predicted by area or connectivity, but they were influenced by the abundant freshwater species. This indicates that all lakes support most of the regional functional diversity. The results highlight the importance of the dispersal process in this lake system and allow the conclusion that considering multiple diversity dimensions can aid the conservation of local and regional fish communities.

KEY WORDS:
Alpha diversity; environmental filters; limiting similarity; neutral paradigm; geographic information system

Morphological traits measured (mm) or categorized	Functional trait	Ecological meaning	References
Standard length (Ls) Body depth (Bd)	Ratio of Ls/Bd	Hydrodynamic ability	Sibbing and Nagelkerke 2001Sibbing F, Nagelkerke L (2001) Resource partitioning by Lake Tana barbs predicted from fish morphometrics and prey characteristics. Reviews in Fish Biology and Fisheries 10: 393-437. https://doi.org/10.1023/A:1012270422092 https://doi.org/10.1023/A:1012270422092...
Length (Pl) and depth (Pd) of the pectoral fin	Ratio of Pl/Pd	Swimming ability, maneuverability at slow speeds and locomotion efficiency	Bellwood et al. 2002Bellwood D, Wainwright R, Fulton C, Hoey A (2002) Assembly rules and functional groups at global biogeographical scales. Functional Ecology 16: 557-562. https://doi.org/10.1046/j.1365-2435.2002.00655.x https://doi.org/10.1046/j.1365-2435.2002... , Dumay et al. 2004Dumay O, Tari P, Tomasini J, Mouillot D (2004) Functional groups of lagoon fish species in Languedoc Roussillon, southern France. Journal of Fish Biology 64: 970-983. https://doi.org/10.1111/j.1095-8649.2004.00365.x https://doi.org/10.1111/j.1095-8649.2004...
Length (Cl) and depth (Cd) of the caudal fin	Ratio of Cl/Cd	Decreases as the swimming ability of the fish declines; benthic species tend to have higher ratios whereas rapid swimmers have lower ratios	Dumay et al. 2004Dumay O, Tari P, Tomasini J, Mouillot D (2004) Functional groups of lagoon fish species in Languedoc Roussillon, southern France. Journal of Fish Biology 64: 970-983. https://doi.org/10.1111/j.1095-8649.2004.00365.x https://doi.org/10.1111/j.1095-8649.2004... , Mouillot et al. 2007Mouillot D, Dumay O, Tomasini J (2007) Limiting similarity, niche filtering and functional diversity in coastal lagoon fish communities. Estuarine, Coastal and Shelf Science 71: 443-456. https://doi.org/10.1016/j.ecss.2006.08.022 https://doi.org/10.1016/j.ecss.2006.08.0...
Total length (Lt)	Lt maximum (Standardized by log (x+1) transformation)	Species with longer bodies are more successful over long distance dispersers than species of smaller sizes	Lucas and Baras 2001Lucas M, Baras E (2001) Migration of Freshwater Fishes. Oxford, Blackwell Science, 440 pp.
Caudal fin shape	Cshape (forked, emarginate, rounded, truncated, semilunar)	Influence the forces exerted on the water	Lauder 2000Lauder G (2000) Function of the caudal fin during locomotion in fishes: kinematics, flow visualization, and evolutionary patterns. American Zoologist 40: 101-122. https://doi.org/10.1093/icb/40.1.101 https://doi.org/10.1093/icb/40.1.101...
Eye diameter (D)	Ratio D/Lt	Detection of food and visual acuity	Piet 1998Piet G (1998) Ecomorphology of a size-structured tropical freshwater fish community. Environmental Biology of Fishes 51: 67-86. https://doi.org/10.1023/A:100733853 https://doi.org/10.1023/A:100733853... , Boyle and Horn 2006Boyle K, Horn M (2006) Comparison of feeding guild structure and ecomorphology of intertidal fish assemblages from central California and central Chile. Marine Ecology Progress Series 319: 65-84. https://doi.org/10.3354/meps319065 https://doi.org/10.3354/meps319065...
Mouth depth (M)	Ratio M/Lt	Maximum size of the prey ingested	Sibbing and Nagelkerke 2001Sibbing F, Nagelkerke L (2001) Resource partitioning by Lake Tana barbs predicted from fish morphometrics and prey characteristics. Reviews in Fish Biology and Fisheries 10: 393-437. https://doi.org/10.1023/A:1012270422092 https://doi.org/10.1023/A:1012270422092...
Trophic Category	CT (invertivorous, planktophagous, piscivorous, omnivorous, benthophagous, detritivorous, insectivorous)	Position in the food web and feeding behavior	Gerking 1994Gerking S (1994) Feeding Ecology of Fish. San Diego, Academic Press, 416 pp.

Variable	Equation	Description
Area (A)		Calculated in hectares.
Shape (S)	$S h a p e = (0.282 \times p e r i m e t e r) / \sqrt{a r e a}$	Corrected relationship between perimeter/area. Ranges from 1 (a perfect circle) to infinite (a long narrow shape, Farina 1998Farina A (1998) Principles and methods in landscape ecology. London, Chapman & Hall, 412 pp.).
Primary Connectivity (PC)	${P C}_{i} = \sum [(R P)] \times [l o g (s m a l l e s t C D i s o f t h e s y s t e m / l o g 10) ({C D i s}_{i j})]$	Related to recolonization potential (RP) and to the cost distance (Adriaensen et al. 2003Adriaensen F, Chardon J, De Blust G, Swinnen E, Villalba S, Gulinck H, Matthysen E (2003) The application of ‘least-cost’ modelling as a functional landscape model. Landscape and Urban Planning 64: 233-247. https://doi.org/10.1016/S0169-2046(02)00242-6. https://doi.org/10.1016/S0169-2046(02)00... , Guimarães et al. 2014Guimarães T, Hartz SM, Becker F (2014) Lake connectivity and fish species richness in southern Brazilian coastal lakes. Hydrobiologia 740: 207-217. https://doi.org/10.1007/s10750-014-1954-x https://doi.org/10.1007/s10750-014-1954-... ). It considers the type of connection, lake area and the number and length of each connection. This index shows the degree of connection of each lake to its neighboring lakes.
	$R P = l o g ({A r e a}_{j}) / l o g (l a r g e s t l a k e o f t h e s y s t e m)$	RP assumes that the largest lake is also the largest reservoir of the species pool in the system. Therefore, the larger the lake the greater its potential to contribute to the recolonization of nearby lakes (Tonn and Magnuson 1982Tonn W, Magnuson J (1982) Patterns in the species composition and richness of fish assemblages in Northern Wisconsin lakes. Ecology 63: 1149-1166. https://doi.org/10.2307/1937251 https://doi.org/10.2307/1937251... ).
Estuarine Connectivity (EC)	$E C = 1 / l o g (C D i s)$	Measure of functional connectivity based on the cost distance from each lake to the estuary, considering the friction value for each connection (Adriaensen et al. 2003Adriaensen F, Chardon J, De Blust G, Swinnen E, Villalba S, Gulinck H, Matthysen E (2003) The application of ‘least-cost’ modelling as a functional landscape model. Landscape and Urban Planning 64: 233-247. https://doi.org/10.1016/S0169-2046(02)00242-6. https://doi.org/10.1016/S0169-2046(02)00... , Guimarães et al. 2014Guimarães T, Hartz SM, Becker F (2014) Lake connectivity and fish species richness in southern Brazilian coastal lakes. Hydrobiologia 740: 207-217. https://doi.org/10.1007/s10750-014-1954-x https://doi.org/10.1007/s10750-014-1954-... ).

Preditor	Response	Direct	Indirect	Net
Area	Shape	-0.0400	0.0000	-0.0400
Area	TD	0.0000	-0.0064	-0.0064
Area	FRic	0.0000	-0.0052	-0.0052
Area	FEve	0.0000	0.0002	0.0002
Area	FDis	0.0000	0.0001	0.0001
Area	CP	0.1230	0.0000	0.1230
Shape	TD	0.1610	0.0000	0.1610
Shape	FRic	0.0000	0.1288	0.1288
Shape	FEve	0.0000	-0.0048	-0.0048
Shape	FDis	0.0000	-0.0032	-0.0032
CE	FRic	0.2370	0.0000	0.2370
TD	FRic	0.8000	0.0000	0.8000
TD	FEve	-0.0200	0.0000	-0.0200
TD	FDis	-0.0300	0.0000	-0.0300

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E-mail: sbz@sbzoologia.org.br

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