This study describes the richness and density of zooplankton across temporary pools in an intermittent river of semiarid Brazil and evaluates the partitioning of diversity across different spatial scales during the wet and dry periods. Given the highly patchy nature of these pools it is hypothesized that the diversity is not homogeneously distributed across different spatial scales but concentrated at lower levels. The plankton fauna was composed of 37 species. Of these 28 were Rotifera, 5 were Cladocera, and 4 were Copepoda (nauplii of Copepoda were also recorded). We showed that the zooplankton presents a spatially segregated pattern of species composition across river reaches and that at low spatial scales (among pools or different habitats within pools) the diversity of species is likely to be affected by temporal changes in physical and chemical characteristics. As a consequence of the drying of pool habitats, the spatial heterogeneity within the study river reaches has the potential to increase β diversity during the dry season by creating patchier assemblages. This spatial segregation in community composition and the patterns of partition of the diversity across the spatial scales leads to a higher total diversity in intermittent streams, compared to less variable environments. 1. Introduction The Brazilian semiarid region represents one of the major dry lands in South America, being characterized by a high hydric deficit and low thermal amplitude [1]. These factors interact with broad climatic patterns to create important variation between dry and wet periods [2]. This affects the hydrology of the region which is characterized by intermittent watercourses [3]. The natural variation associated with the alternation of dry and wet periods and the intermittency of flow in rivers and streams create a mosaic of temporary natural aquatic habitats, mostly strings of ephemeral pools in the dry river bed and more permanent larger ones [4]. Therefore, these systems are highly temporally variable and spatially heterogeneous [5, 6]. During the wet season, the mechanical force of the water flow changes the physical habitat and modifies water chemical characteristics and nutrient dynamics [5, 7], whereas during the dry season, discontinuation of flow and water volume reduction influence community structure and diversity by concentrating nutrients and affecting physical and chemical characteristics of the water [5, 8]. Furthermore, during this phase, the water retention in semipermanent and temporary pools represents refugia for the maintenance of aquatic
References
[1]
L. Maltchik and M. Florin, “Perspectives of hydrological disturbance as the driving force of Brazilian semiarid stream ecosystems,” Acta Limnologica Brasiliensia, vol. 14, no. 3, pp. 35–41, 2002.
[2]
W. T. Liu and R. I. Negrón Juárez, “ENSO drought onset prediction in northeast Brazil using NDVI,” International Journal of Remote Sensing, vol. 22, no. 17, pp. 3483–3501, 2001.
[3]
E. R. Steffan, “Hidrografia,” in Regi？o Nordeste. Geografia Do Brasil, IBGE, Ed., pp. 111–133, SERGRAF-IBGE, Rio de Janeiro, Brazil, 1977.
[4]
L. Maltchik and E. S. F. Medeiros, “Conservation importance of semi-arid streams in north-eastern Brazil: implications of hydrological disturbance and species diversity,” Aquatic Conservation: Marine and Freshwater Ecosystems, vol. 16, no. 7, pp. 665–677, 2006.
[5]
E. S. F. Medeiros, M. J. Silva, and R. T. C. Ramos, “Application of catchment- and local-scale variables for aquatic habitat characterization and assessment in the Brazilian semi-arid region,” Neotropical Biology and Conservation, vol. 3, no. 1, pp. 13–20, 2008.
[6]
E. H. Stanley, S. G. Fisher, and N. B. Grimm, “Ecosystem expansion and contraction in streams: desert streams vary in both space and time and fluctuate dramatically in size,” BioScience, vol. 47, no. 7, pp. 427–435, 1997.
[7]
S. G. Fisher, N. B. Grimm, E. Martí, and R. Gómez, “Hierarchy, spatial configuration, and nutrient cycling in a desert stream,” Austral Ecology, vol. 23, no. 1, pp. 41–52, 1998.
[8]
M. C. Crispim and G. T. P. Freitas, “Seasonal effects on zooplanktonic community in a temporary lagoon of northeast Brazil,” Acta Limnologica Brasiliensia, vol. 17, no. 4, pp. 385–393, 2005.
[9]
E. S. F. Medeiros and L. Maltchik, “Diversity and stability of fishes (Teleostei) in a temporary river of the Brazilian semiarid region,” Iheringia, no. 90, pp. 157–166, 2001.
[10]
L. De Meester, S. Declerck, R. Stoks et al., “Ponds and pools as model systems in conservation biology, ecology and evolutionary biology,” Aquatic Conservation, vol. 15, no. 6, pp. 715–725, 2005.
[11]
E. S. F. Medeiros, M. P. Noia, L. C. Antunes, and T. X. Melo, “Zooplankton composition in aquatic systems of semi-arid Brazil: spatial variation and implications of water management,” Pan-American Journal of Aquatic Sciences, vol. 6, pp. 290–302, 2011.
[12]
D. G. Jenkins and A. L. Buikema Jr., “Do similar communities develop in similar sites? A test with zooplankton structure and function,” Ecological Monographs, vol. 68, no. 3, pp. 421–443, 1998.
[13]
C. E. Cáceres and D. A. Soluk, “Blowing in the wind: a field test of overland dispersal and colonization by aquatic invertebrates,” Oecologia, vol. 131, no. 3, pp. 402–408, 2002.
[14]
J. L. Brooks and S. I. Dodson, “Predation, body size, and composition of plankton,” Science, vol. 150, no. 3692, pp. 28–35, 1965.
[15]
R. Pourriot, C. Rougier, and A. Miquelis, “Origin and development of river zooplankton: example of the Marne,” Hydrobiologia, vol. 345, no. 2-3, pp. 143–148, 1997.
[16]
J. H. Stansfield, M. R. Perrow, L. D. Tench, A. J. D. Jowitt, and A. A. L. Taylor, “Submerged macrophytes as refuges for grazing Cladocera against fish predation: observations on seasonal changes in relation to macrophyte cover and predation pressure,” Hydrobiologia, vol. 342-343, pp. 229–240, 1997.
[17]
M. Seminara, D. Vagaggini, and F. G. Margaritora, “Differential responses of zooplankton assemblages to environmental variation in temporary and permanent ponds: zooplankton of temporary and permanent ponds,” Aquatic Ecology, vol. 42, no. 1, pp. 129–140, 2008.
[18]
A. C. B. Vieira, L. L. Ribeiro, D. P. N. Santos, and M. C. Crispim, “Correlation between the zooplanktonic community and environmental variables in a reservoir from the Northeastern semi-arid,” Acta Limnologica Brasiliensia, vol. 21, no. 3, pp. 349–358, 2009.
[19]
W. Sousa, J. L. Attayde, E. D. S. Rocha, and E. M. Eskinazi-Sant'Anna, “The response of zooplankton assemblages to variations in the water quality of four man-made lakes in semi-arid northeastern Brazil,” Journal of Plankton Research, vol. 30, no. 6, pp. 699–708, 2008.
[20]
S. Tavernini, “Seasonal and inter-annual zooplankton dynamics in temporary pools with different hydroperiods,” Limnologica, vol. 38, no. 1, pp. 63–75, 2008.
[21]
K. M. Jenkins and A. J. Boulton, “Connectivity in a dryland river: short-term aquatic microinvertebrate recruitment following floodplain inundation,” Ecology, vol. 84, no. 10, pp. 2708–2723, 2003.
[22]
R. L. Farias, L. K. Carvalho, and E. S. F. Medeiros, “Distribution of Chironomidae in a semiarid intermittent river of Brazil,” Neotropical Entomology, vol. 41, pp. 450–460, 2012.
[23]
L. G. Rocha, E. S. F. Medeiros, and H. T. A. Andrade, “Influence of flow variability on macroinvertebrate assemblages in an intermittent stream of semi-arid Brazil,” Journal of Arid Environments, vol. 85, pp. 33–40, 2012.
[24]
C. A. Frissell, W. J. Liss, C. E. Warren, and M. D. Hurley, “A hierarchical framework for stream habitat classification: viewing streams in a watershed context,” Environmental Management, vol. 10, no. 2, pp. 199–214, 1986.
[25]
C. M. Pringle, C. M. Pringle, R. J. Naiman, et al., “Patch dynamics in lotic systems: the stream as a mosaic,” Journal of the North American Benthological Society, vol. 7, pp. 503–524, 1988.
[26]
M. C. Thoms, P. J. Beyer, and K. H. Rogers, “Variability, complexity and diversity—the geomorphology of river ecosystems in dryland regions,” in Changeable, Changed, Changing: The Ecology of Desert Rivers, R. T. Kingsford, Ed., p. 368, Cambridge University Press, Cambridge, UK, 2004.
[27]
N. R. Sim？es, S. L. Sonoda, and S. M. M. S. Ribeiro, “Spatial and seasonal variation of microcrustaceans (Cladocera and Copepoda) in intermittent rivers in the Jequiezinho River Hydrographic Basin, in the Neotropical semiarid,” Acta Limnologica Brasiliensia, vol. 20, no. 3, pp. 197–204, 2008.
[28]
E. S. F. Medeiros and A. H. Arthington, “Allochthonous and autochthonous carbon sources for fish in floodplain lagoons of an Australian dryland river,” Environmental Biology of Fishes, vol. 90, no. 1, pp. 1–17, 2011.
[29]
AESA, “Agência Executiva de Gest？o das águas do Estado da Paraíba. Disponível em,” Paraíba, Brasil, 2012, http://www.aesa.pb.gov.br/.
[30]
I. R. Leal, M. Tabarelli, and J. M. C. Silva, Eds., Ecologia e Conserva？？o da Caatinga, EDUFPE, Recife, Brazil, 2nd edition, 2005.
[31]
M. C. Peel, B. L. Finlayson, and T. A. McMahon, “Updated world map of the K？ppen-Geiger climate classification,” Hydrology and Earth System Sciences, vol. 11, no. 5, pp. 1633–1644, 2007.
[32]
SUDENE, Dados Pluviométricos mensais do nordeste. Superintendência do Desenvolvimento do Nordeste, Recife, Brazil, 1990.
[33]
J. M. C. Silva, M. Tabarelli, M. Fonseca, and L. Lins, Eds., Biodiversidade da Caatinga: áreas e a？？es prioritárias para a conserva？？o, Ministério do Meio Ambiente/Universidade Federal de Pernanbuco, Brasília, DF, Brazil, 2003.
[34]
J. F. Haney and D. J. Hall, “Sugar-coated Daphnia: a preservation technique for Cladocerans,” Limnology and Oceanography, vol. 18, no. 2, pp. 331–333, 1973.
[35]
L. M. A. Elmoor-Loureiro, Manual de identifica？？o de cladóceros límnicos do Brasil, Editora Universa, Brasilia, DF, Brazil, 1997.
[36]
E. Suárez, J. W. Reid, T. M. Ilige, et al., Catálogo de los copépodos (Crustácea) continentales de la península de Yucatán, México, ECOSUR/CONABIO, Mexico City, Mexico, 1996.
[37]
H. Segers, “Annotated checklist of the rotifers (Phylum Rotifera), with notes on nomenclature, taxonomy and distribution,” Zootaxa, no. 1564, pp. 1–104, 2007.
[38]
P. S. Maitland, Field Studies: Sampling in Freshwaters, in Biology of Fresh Waters, Blackie, Glasgow, UK, 1990.
[39]
R. R. Sokal and F. J. Rohlf, Biometry: The Principles and Practice of Statistics in Biological Research, W.H. Freeman, San Francisco, Calif, USA, 1969.
[40]
J. A. Veech and T. O. Crist, “PARTITION: software for hierarchical partitioning of species diversity, version 3.0,” 2009, http://www.users.muohio.edu/cristto/partition.htm.
[41]
T. O. Crist, J. A. Veech, J. C. Gering, and K. S. Summerville, “Partitioning species diversity across landscapes and regions: a hierarchical analysis of α, β, and γ diversity,” American Naturalist, vol. 162, no. 6, pp. 734–743, 2003.
[42]
J. A. Veech and T. O. Crist, “PARTITION 3. 0 user’s manual,” unpublished document, 2009.
[43]
B. McCune and J. B. Grace, Analysis of Ecological Communities, Gleneden Beach, Ore, USA, MjM Software Design, 2002.
[44]
M. E. Biondini, C. D. Bonham, and E. F. Redente, “Secondary successional patterns in a sagebrush (Artemisia tridentata) community as they relate to soil disturbance and soil biological activity,” Vegetatio, vol. 60, no. 1, pp. 25–36, 1985.
[45]
M. Dufrêne and P. Legendre, “Species assemblages and indicator species: the need for a flexible asymmetrical approach,” Ecological Monographs, vol. 67, no. 3, pp. 345–366, 1997.
[46]
B. McCune and M. J. Mefford, PC-ORD. Multivariate Analysis of Ecological Data. Version 4.27, MjM Software Design, Gleneden Beach, Ore, USA, 1999.
[47]
L. Maltchik, L. E. K. Lanés, C. Stenert, and E. S. F. Medeiros, “Species-area relationship and environmental predictors of fish communities in coastal freshwater wetlands of southern Brazil,” Environmental Biology of Fishes, vol. 88, no. 1, pp. 25–35, 2010.
[48]
M. C. Crispim and T. Watanabe, “Caracteriza？？o Limnológica das Bacias doadoras e receptoras de águas do Rio S？o Francisco:1—Zooplancton,” Acta Limnologica Brasiliensia, vol. 12, no. 2, pp. 93–103, 2000.
[49]
V. L. S. Almeida, M. E. L. de Larrazábal, A. D. N. Moura, and M. de Melo Júnior, “Rotifera das zonas limnética e litoranea do reservatório de Tapacurá, Pernambuco, Brasil,” Iheringia, vol. 96, no. 4, pp. 445–451, 2006.
[50]
K. Fahd, L. Serrano, and J. Toja, “Crustacean and rotifer composition of temporary ponds in the Do？ana National Park (SW Spain) during floods,” Hydrobiologia, vol. 436, pp. 41–49, 2000.
[51]
J. Green, “Diversity and dominance in planktonic rotifers,” Hydrobiologia, vol. 255-256, no. 1, pp. 345–352, 1993.
[52]
J. D. Allan, “Life history patterns in zooplankton,” American Naturalist, vol. 110, no. 971, pp. 165–176, 1976.
[53]
K. G. Porter, J. D. Orcutt Jr., and J. Gerritsen, “Functional response and fitness in a generalist filter feeder Daphnia magna ( Cladocera: Crustacea),” Ecology, vol. 64, no. 4, pp. 735–742, 1983.
[54]
N. Walz and M. Welker, “Plankton development in a rapidly flushed lake in the River Spree system (Neuendorfer See, Northeast Germany),” Journal of Plankton Research, vol. 20, no. 11, pp. 2071–2087, 1998.
[55]
M. C. Crispim and T. Watanabe, “What can dry reservoir sediments in a semi-arid region in Brazil tell us about cladocera?” Hydrobiologia, vol. 442, no. 1–3, pp. 101–105, 2001.
[56]
D. Frisch and A. J. Green, “Copepods come in first: rapid colonization of new temporary ponds,” Fundamental and Applied Limnology, vol. 168, no. 4, pp. 289–297, 2007.
[57]
G. A. Cole, “Contrasts among calanoid copepods from permanent and temporary ponds in Arizona,” American Midland Naturalist, vol. 76, no. 2, pp. 351–368, 1966.
[58]
F. Sheldon and K. F. Walker, “Spatial distribution of littoral invertebrates in the lower Murray-Darling River system, Australia,” Marine and Freshwater Research, vol. 49, no. 2, pp. 171–182, 1998.
[59]
K. E. Esteves and S. Sendacz, “Rela？？es entre a biomassa do zooplancton e o estado trófico de reservatórios do estado de S？o Paulo,” Acta Limnologica Brasiliensia, vol. 2, no. 1, pp. 587–604, 1988.
[60]
M. Serafim-Júnior, G. Perbiche-Neves, L. de Brito, A. R. Ghidini, and S. M. C. Casanova, “Varia？？o espa？o-temporal de Rotifera em um reservatório eutrofizado no sul do Brasil,” Iheringia, vol. 100, no. 3, pp. 233–241, 2010.
[61]
P. Usseglio-Polatera, “Theoretical habitat templets, species traits, and species richness: aquatic insects in the Upper Rhone River and its floodplain,” Freshwater Biology, vol. 31, pp. 417–437, 1994.
[62]
J. A. Veech, T. O. Crist, and K. S. Summerville, “Intraspecific aggregation decreases local species diversity of arthropods,” Ecology, vol. 84, no. 12, pp. 3376–3383, 2003.
[63]
D. J. Murrell, D. W. Purves, and R. Law, “Uniting pattern and process in plant ecology,” Trends in Ecology and Evolution, vol. 16, no. 10, pp. 529–530, 2001.
[64]
R. E. Ricklefs and I. J. Lovette, “The roles of island area per se and habitat diversity in the species-area relationships of four Lesser Antillean faunal groups,” Journal of Animal Ecology, vol. 68, no. 6, pp. 1142–1160, 1999.
[65]
D. O. Hessen, B. A. Faafeng, V. H. Smith, V. Bakkestuen, and B. Walseng, “Extrinsic and intrinsic controls of zooplankton diversity in lakes,” Ecology, vol. 87, no. 2, pp. 433–443, 2006.
[66]
J. C. Marshall, F. Sheldon, M. Thoms, and S. Choy, “The macroinvertebrate fauna of an Australian dryland river: spatial and temporal patterns and environmental relationships,” Marine and Freshwater Research, vol. 57, no. 1, pp. 61–74, 2006.
