Comparision of different kind of semi-arid aquatic environments based on zooplankton communities 1

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1 REVISTA DE BIOLOGIA E CIÊNCI DA TERRA ISSN Suplemento Especial - Número 1-2º Semestre 2006 Comparision of different kind of semi-arid aquatic environments based on zooplankton communities 1 Maria Cristina Crispim 2, Leonardo L. Ribeiro 2, Sofia E. M. Gomes 2, Glecia Trinta de P. Freitas 2,3,Fabiano R. Serpe 2 RESUMO Foi feita a comparação entre diferentes tipos de ambientes aquáticos do semi-árido, com base nas comunidades zooplanctônicas. Estes ambientes são muito importantes porque são a única fonte de abastecimento de água superficial ao longo do ano para a população. Foram analisados 2 lagoas, 2 barreiros e 3 açudes, ambos nas estações seca e chuvosa. Os resultados mostraram que não existe um padrão para ambientes específicos. Muitas mudanças foram observadas no mesmo ambiente ao longo do ano e entre os anos. Apesar de algumas das espécies zooplanctônicas estarem presentes em todos os tipos de ambientes estudados, as densidades registradas variaram bastante. As densidades do zooplâncton total foram maiores no Barreiro Santa Clara, Açude Soledade, Açude da Barra, Lagoa Panati, Lagoa do Serrote, Açude Taperoá e Barreiro Soledade, respectivamente. O índice de Shannon foi maior nas lagoas, seguido pelos barreiros e açudes. A qualidade da água e a disponibilidade de habitat foi mais decisiva nas comunidades zooplanctônicas do que o tipo do ambiente estudado. Palavras-Chaves: Zooplâncton, biodiversidade, semi-árido, análise de cluster, ambientes aquáticos temporários ATRACT Comparison of different kinds of semi-arid aquatic environments based on zooplankton communities were done. These environments are very important because they are the only surface water supply throughout the year. We analyzed 2 lagoons, 2 barreiros and 3 dams in both dry and rainy seasons. Some of these studied areas are neighboring. The results showed that there is not a pattern for specific environments. Great changes were observed in the same environment throughout the year and over the years. The densities of total zooplankton communities were higher in Barreiro Santa Clara, Açude Soledade, Açude da Barra, Panati Lagoon, Serrote Lagoon, Açude Taperoá and Barreiro Soledade, respectively. The Shannon biodiversity indices were higher in lagoons, followed by barreiros and reservoirs. The water quality and the habitat availability seemed to be more decisive in determining the zooplankton s community structure than the environment type. Key words: Zooplankton, biodiversity, semi-arid, temporary aquatic environments. 1 INTRODUCTION Temporary environments are very important in semi-arid regions in northeast Brazil because they are ordinarily the unique source of water to the local population and they are rich in biodiversity. The region where the environments are located are characterized by the irregularity of rain distribution and they are often subjected to extensive drought periods. Dry and rainy seasons carry changes in the organization of these ecosystems because of indirect consequences caused by fluctuation of water level (Maltchik, 1999). In the rainy period a great number of aquatic environments emerge, while in the dry season the water level of these environments drops to a fraction of its capacity or they dry completely. 98

2 Water trophic level changes quickly in dams of paraiban semi-arid (Crispim et al., 2000). The dry period induces an increase of trophic degree due mainly to reduction of water level (this reduction range is according to the drought degree). Therefore, an increase in the concentration of nutrients in the water occurs and, consequently development of aquatic vegetation and dead algal biomass arises. During rainy season, the water level increases and the eutrophication process tends to decline (Martinez et al., 1991; Crispim et al., 2000; Vieira et al.,. 2000). Several dams (açudes) and little manmade ponds (barreiros) have been constructed in this area to decrease the impact of drought on humans (Esteves, 1998). During normal years, the reservoirs, especially the bigger ones, keep water during dry season, although their water levels largely decrease. However, dams, lagoons and barreiros almost always dry out during the dry season. Zooplanktonic organisms living in these environments must have life strategies that allow them to stay in diapause while their habitats change (Crispim & Watanabe, 2001). Many animals, such as crustaceans (Fryer, 1996; Crispim et. al., 2003) and rotifers (King & Snell, 1980; Gilbert, 1995) have the ability to produce diapausing stages that allow them to survive in temporarily adverse environments. Resting eggs permit recolonization of the habitat after a dry period. According to Hairston and Cáceres (1996), crustaceans found in inland water bodies, especially those that occasionally become dry, are more likely to possess a diapausing stage than those living in open oceans. The inducing of diapause stages is influenced by environmental quality; a higher or lower number of ephippial females in Moina minuta populations can be induced according to the environment conditions (Crispim et al., 2003). Species composition and abundance in zooplankton communities can be influenced by several factors such as the size of the water bodies (Patalas, 1971). Natural temporary ponds and barreiros are small, shallow and frequently show an extensive littoral zone; while the reservoirs have a larger capacity to receive water, are deeper and show an extensive limnetic zone. Temporary environments are those in which the entire habitat for aquatic organisms shifts from being available to being unavailable for a duration period which substantially affects the entire biota, due to evaporation or freezing water for some undefined period (Schwartz & Jenkins, 2000). The purpose of this study was to compare lagoons, dams and barreiros located in a semi-arid region in northeast Brazil, based on the zooplankton communities, to find out if the size or kind of the environment influenced the zooplankton community present in each of those ecosystems. 2 METHODOLOGY Two lagoons, 2 small man-made ponds (barreiros) and 3 dams (açudes) were studied. All these three different kinds of temporary aquatic environments are located in the semiarid region of the state of Paraíba, Northeast Brazil. Samples were taken weekly in the beginning of the rain period and monthly afterwards, for three years (2002 to 2004) by filtering 40 liters of littoral water through a 45 µm mesh size and the material was preserved in a 4% sugar saturated formaldehyde solution. In a work developed by Crispim & Watanabe (2000) in a semi arid dam, the zooplankton communities in littoral and limnetic zones were compared and they presented the same species, although in different densities. So the comparison among these environments using littoral samples can be accepted. In each environment three replicates were collected. Densities (ind.l -1 ) represent the mean of these three replicates. Identification and counting were performed using a Sedgewick-Rafter counting cell, under optical microscope. At least 100 organisms of rotifers and cladocerans were counted from subsamples of 1 ml. Rotifers were identified by Ruttner-Kolisko (1974) and Koste (1978), and cladocerans by Elmoor- Loureiro (1997). Shannon Diversity Index (H ) (nat.ind.l - 1 ) and Pielou s Eveness (E) were computed with the software Statistical Ecology (Ludwig and Reynolds, 1988), and Cluster Analysis, using Relative Euclidian Distance as dissimilarity measure, with Statistica 6.0 (StatSoft, 1998). Sørensen index was used to compare the species inventorial amongst each water body. A 99

3 Constancy Index was applied to each taxon recorded in each study site dividing the number of samples holding the taxon by the total number of samples collected, and multiplying by 100 (Dajoz, 1973). Taxa were considered constant when appearing in more than 50% of samples; occasional, when found between 25% and 50% of samples; and accidental when the taxon was present in less than 25% of the samples. The rainfall rates were obtained from the Meteorological Post of the State Government in Taperoá borough. Study Areas In paraiban semi-arid the aquatic environments suffer changes, in the trophic states along the year, and are strongly influenced by hydrological cycles, being less eutrophic in rainy periods and more in dry ones. The location of environments are showed in Figure 1. Açude Taperoá II - located in Taperoá borough (07 o S and 36 o W) and in the raining period it has a maximum capacity of 15,148.9 Km 3 (SEMARH/LMRS-PB) and a maximum depth of about 5 m. This environment was considered mesotrophic by the Carlson s trophic state index (TSI), along the year (Andrade, 2005). Açude da Barra - located in Juazeirinho borough (07 o S and 36 o W). Its maximum capacity is 3, Km 3 (SEMARH/LMRS-PB). This dam was considered eutrophic and hipereutrophic in 2002/2003, drying completely in the end of 2003 (Trophic State Index (TSI) from 55.8 to 86.1). Açude Soledade - located in the borough with the same name (07 o S and 36 o W). Its maximum capacity is 27,058.0 Km 3 (SEMARH/LMRS-PB). This environment present a TSI mesotrophic in the rainy period and eutrophic in the dry one (Andrade, 2005). In these dams water generally remains in the dry periods, although in especially dry years they completely dry out. In these years, the population digs artesian wells, in the river bed, to obtain water for their supply. Panati Lagoon - located near Taperoá borough (07 o S and 36 o W). The lagoon is oval, with about 100 m of maximum length and its maximum depth is about 1.5 m. A large quantity of macrophytes covers the littoral zone and aquatic birds can frequently be seen. This environment is oligotrophic most of the year and mesotrophic in special dry months (Andrade, 2005). Serrote Lagoon - located in Boa Vista borough, it was formed when the road was constructed, is oval, with close to 80 m of higher length and has about 1.0 m of maximum depth. There are less macrophytes in quantity and diversity than in the previous one. The TSI ranged from to 53 to 54, being considered mesotrophic. Barreiro Soledade - about 2 Km far from the Açude Soledade. It is small, oval with almost 30 m of diameter. Its maximum depth is about 1.5 m. There is only one species of macrophyte in small quantity. The TSI ranged from 38.2 to 58.8 being oligotrophic in the dry period and eutrophic in the rainy months. Barreiro Santa Clara - located neighboring Serrote Lagoon. It is small and elongated, with about 10 m of width and 30 m of length. The main differences between barreiros and lagoons are that the former have one abrupt margin, whereas the barreiro is built, and thin sediments confer a brown color to the water. In the latter there are more macrophytes. The TSI ranged from 55.5 to 58.8, being considered eutrophic. 3 RESULTS The precipitation regime was different in all 3 years of study and this is a common fact. In 2003 the winter was dry; in 2004, the rainfalls were greater than usual, the strongest rain in the last 40 years, and in 2002 the rainfall regime registered an intermediate situation (Table I). As a consequence of this, in 2002 the study sites did not dry, but after October 2003 even the bigger dams completely dried (except the Açude Soledade), and in 2004 a large quantity of water remained, even in the dry period. Thus, considerable changes in the zooplankton communities were observed. Copepods were observed, but as not all of them were identified, and this study is comparative, they were not considered in this work. 100

4 Fig. 1 - Taperoá River Drainage Basin in Paraíba, Brazil with location of sample sites. Table I - Annual precipitation indices (mm) in the studied aquatic environment region REGION/YEAR Taperoá Soledade Juazeirinho Boa Vista (not informed) SPECIES COMPOSITION We have found 70 species of rotifers and 15 of cladocerans. Table I shows the list of species and the constancy index for rotifers and cladocerans. C. cornuta, D. spinulosum and M. minuta were constant species in a great number of environments. The rotifer B. angularis was constant for all dams, and Lecane luna was constant in 2 dams (Açude da Barra and Açude Soledade and 1 lagoon (Serrote). All other rotifer species were not constant for more than 2 environments (Table II). The Panati Lagoon was the environment with the highest number of species, both cladocerans and rotifers, followed by Serrote Lagoon for rotifers and Açude da Barra for cladocerans (Fig. 2). This latter environment only registered D. spinulosum and M. minuta in 2002, and Biapertura karua in 2003, but in 2004, after the highest precipitation index, 6 species were present, all but D. spinulosum (Table III). The environments with the lowest number of species were Barreiro and Açude Soledade, for both rotifers and cladocerans; and Barreiro Santa Clara for rotifers (Fig. 2). DENSITIES AND BIODIVERSITY Generally higher densities were recorded for rotifers than for cladocerans, except in Barreiro Santa Clara and Serrote Lagoon where in one sample cladocerans were more abundant than rotifers. Density of Rotifera and Cladocera varied along the almost three years of study. In all environments, densities were smaller in the beginning of the rainy period, increasing 101

5 gradually along this period, and reaching its highest values in the beginning of dry period (Fig. 3). When the rotifer Conochilus natans was present, high densities of rotifers were observed in the beginning of the rainy period. The total cladoceran and rotifer densities were highest in Barreiro Santa Clara, where C. natans exhibited higher peaks of density (1,916 ind.l - 1 ), together with Keratella valga (1,418ind.L -1 ), in The density peak of the first species was recorded in the beginning of the rainy period and those of the other species at the end of this period. In 2004 the dominant species changed to Brachionus angularis and C. natans was not observed. Following this environment, the Açude Soledade and Açude da Barra recorded the highest densities. The Açude Taperoá and Barreiro Soledade recorded the lowest densities. Although in lagoons higher peaks were recorded, in these environments the densities observed along the years of study remained lower than in the other types of environments analysed (Fig. 3). Table. II - Rotifer species presence and constancy index for all analysed environments: Açude da Barra (), Açude Taperoá (), Açude Soledade (), Barreiro Soledade (), Barreiro Santa Clara (C), Panati Lagoon () and Serrote Lagoon (SL), from 2002 to == Accidental; /// Occasional; ++ Constant Blank cells - Absent species 102

6 Values of biodiversity indices vary highly throughout the year and over the studied years. For example in Açude da Barra, the most eutrophic one in 2003 (personal data), the diversity indices values highly decreased, compared with the previous and later year. On the contrary, the Açude Taperoá, the least eutrophic (personal data), showed less variation over the years (Fig. 4). Panati Lagoon was the environment showing the highest diversity indices values and the lowest were not as low as in the other ecosystems. The lowest indices throughout a longer period were obtained in Açude Soledade e Açude da Barra (Fig 4). Cluster analysis Cluster analysis to binary data (presence/absence) for showed high distances between environments (Fig. 5). Minimum relative distance found was 50%, between Açude Soledade and Barreiro Soledade. The Panati Lagoon was the most distant environment of all. On the other hand, Serrote lagoon was closest to Açude Soledade, Barreiro Soledade and Barreiro Santa Clara cluster than to Panati lagoon. 103

7 Table III - Cladoceran species and constancy index in Açude da Barra (), Açude Taperoá (), Açude Soledade (), Barreiro Soledade (), Barreiro Santa Clara (C), Lagoa Panati (LP) and Lagoa do Serrote (LS), from 2002 to Cladocera C SL Alona cambouei == A. poppey == == == Biapertura karua == Chydorus sp. == == Ceriodaphnia cornuta == /// /// /// Diaphanosoma spinulosum ++ /// /// ++ Dunhevedia odontoplax == == Ephemeroporus hybridus /// Ilyocryptus spinifer == Leydigia sp. == Macrothrix sp. == == M. laticornis ++ M. spinosa == == == == Moina minuta == /// == ++ Simocephalus acutirostris /// == Accidental; /// Occasional; ++ Constant Blank cells - Absent species Fig.2 - Number of species in all studied environments between 2002 and Açude da Barra; - Açude Soledade; - Barreiro Soledade; C - Barreiro Santa Clara; SL - Serrote Lagoon; - Açude Taperoá and - Panati Lagoon. 39 Rotifera 27 Cladocera C SL The Açude da Barra and the Açude Taperoá stayed in an intermediate position between Panati lagoon and the other water bodies. The high number of species which inhabit only Panati lagoon seemed to be decisive for making it an isolated cluster. This analysis indicates that the kind of environment is not the proximal cause influencing the zooplanktonic community 104

8 Fig.3 - Densities of Rotifera and Cladocera in the aquatic environments studied, from 2002 to Arrows indicate when there was no water in the environments A ç ud e d a B arra A ç ud e Tap e ro á II A ç ud e S o le d ad e B arre iro S o le d ad e B arre iro S ant a C lara P ana t i La g o o n S e rro t e P o nd Rotifera Cladocera Copepoda time (mo nth/year) 105

9 Fig.4 - Shannon s Index of Diversity and Pielou s Eveness in all study sites from 2002 to Arrows indicate when there was no water in the environments. 3,5 3,0 2,5 2,0 A ç ude da B a rra 0,8 0,6 3,5 3,0 2,5 2,0 A ç ude Ta pe ro á II 1,0 0,8 0,6 1,5 0,5 0,4 0,2 1,5 0,5 0,4 0,2 0,0 3,5 A ç ude S o le da de 1,0 3,5 B a rre iro S o le da de 3,0 2,5 2,0 0,8 0,6 3,0 2,5 2,0 0,8 0,6 1,5 0,5 0,4 0,2 1,5 1,0 0,5 0,4 0,2 0,0 0,0 3,5 3,0 2,5 2,0 B a rre iro S a nta C la ra 0,8 0,6 3,5 3,0 2,5 2,0 P a na ti La g o o n 0,8 0,6 1,5 0,5 0,4 0,2 1,5 0,5 0,4 0,2 3,5 S e rro te P o nd 3,0 2,5 2,0 1,5 0,5 0,8 0,6 0,4 0,2 Rotifera (H') Cladocera (H') Rotifera (E) Cladocera (E) time structures. Analysing the different seasonal periods in the 3 years one can observe that the most similar pairs change from one year to another, showing that the environments highly change over the years and that the kind of environment does not seem to define the community organization (Fig. 6). In 2004, the maximum distances between the environments were lower than in earlier years. 4 DISCUSSION In this study, all the samples were collected in the littoral zone of the different environments analyzed. The littoral region usually supports a variety of macrophytes and a large number of animal species associated with them, being the most diverse part of the lake ecosystem (Winterbourn & Lewis, 1975). The lagoons showed a larger littoral zone than the other kinds of aquatic ecosystems, with a large quantity and diversity of macrophytes and, 106

10 therefore, also showed a greater diversity of habitats. This way, the greater quantity and complexity of microhabitats observed in the lagoons, due to the presence of aquatic vegetation, could explain the higher zooplanktonic species richness in this kind of environment, especially in Panati Lagoon which exhibits a large quantity of macrophytes of different species and was the environment with a larger number of zooplanktonic species, both cladocerans and rotifers. The presence of macrophytes seemed to confer more stability to the environment. Wright et al. (2002) found less changes in macroinvertebrate fauna associated with macrophytes (Berula erecta) than with gravel in River Lambourn, Berkshire, when submitted to drought events. Dodson (1991) found a significantly positive correlation between crustacean zooplankton and lake size, although also influenced by trophic state. In eutrophic and hypereutrophic systems species richness decreases. These authors did not compare species richness with the presence or absence of macrophytes. We observed that in the more eutrophic environments (Açude Soledade e Açude da Barra) the diversity indices were low most of the time, increasing in 2004 when a higher precipitation was recorded, inducing a decrease in the trophic state, because of dissolution of nutrients (Martinez et al., 1991; Crispim et al., 2000). Fig. 5 - Sørensen index between all studied sites, based on presence/absence of species. - Açude da Barra; - Açude Soledade; - Barreiro Soledade; C - Barreiro Santa Clara; SL - Serrote Lagoon; - Açude Taperoá and - Panati Lagoon C SL The fact that the zooplankton communities are not related with a specific kind of environment is probably because, depending of the quantity of rain, these environments can, in some periods, become very similar in water quality and this is an important regulator of the zooplankton communities (Dodson, 1991). Actually we found lower distances between environments in rainy periods, which shows how these systems can change and become more similar to each other, depending on the hydrological cycle. Thus, zooplankton communities can quickly alter according to water quality, which influences food availability (quality and quantity) (Campbell & Haase, 1981 apud Sampaio et al., 2002) Although tropical lakes are often small, they can have extremely high productivities and few adjacent lakes and these factors tend to reduce species richness (Dodson, 1992). In our work we observed 70 rotifer and 15 cladoceran species in 7 different environments, which is higher than the values recorded by Fahd et al. (2000) in 18 temporary ponds in Doñana National Park in Spain, where 37 rotifer and 17 cladoceran species were found. At least for the rotifers more species were found but this group was also the least constant in the studied environments, since the majority of rotifer species were considered as accidental in the ecosystems. This can be an effect of the eutrophication process, carried out during the hydrological cycle evolution (Crispim et al., 2001), changing quickly the water quality and inducing an ecological 107

11 Fig. 6 - Cluster analysis (Euclidian Distance) between the aquatic environments studied from 2002 to 2004 in rainy, rainy-dry transition, and dry periods. - Açude da Barra; - Açude Taperoá II; - Soledade; - Barreiro Soledade; C - Barreiro Santa Clara; - Panati Lagoon; and SL - Serrote Lagoon Rainy Season (26/01/02) Rainy season (16/01/03) Rainy Season (28/02/04) C Max. Distance = 2580 Rainy-dry season (27/04/02) C Max. Distance = 1400 Dry season (30/10/02) SL C Max. Distance = 620 C Max. Distance = 650 Rainy-dry Season (31/03/03) SL C Max. Distance = 950 Dry season (30/10/03) C Max. Distance = 590 C Max. Distance = 170 Rainy-dry season (07/04/04) Max. Distance = 74 Dry season (18/06/04) C Max. Distance = 330 succession amongst species in the ecosystems, not allowing rotifer species to remain over the years. When the water level decreases in these environments, large quantities of macrophytes were exposed and died, decreasing the number of viable habitats. In cascade system reservoirs in the Paranapanema River, southeast Brazil, Sampaio et al. (2002) found 76 species of Rotifera and 26 of Cladocera, showing that in tropical or subtropical aquatic systems the number of species can be high. Maybe, since temporary aquatic environments quickly alter their trophic state, it is necessary to have a longer sampling period in order to collect all species present, or collect environment sediments to induce the hatching of diapause eggs (Crispim &Watanabe, 2000, 2001) and in Fahd work only two periods were analysed. Our results show that the temporary lagoons are the aquatic ecosystems with higher species richness and more stable than açudes and barreiros. Comparing to other kind of temporary environments located in Northeast Brazil, like dams, we can see that the zooplankton densities of lagoons are lower whereas the richness is higher (Crispim et al., 2000; Vieira et al., 2000), as opposed to dams. The Lecane genus is characteristic of the littoral zone, amongst aquatic vegetation (Sládecek, 1983); Colurella uncinata and Lepadella patella are also common in aquatic vegetation (Sládecek, 1983). Thus, the large quantity of aquatic macrophytes in lagoons would explain the frequent occurrence of these species in the samples analyzed. The cladoceran M. laticornis shows bentonic habits and would be influenced by aquatic vegetation present all around the lagoons littoral zone. This species and Macrothrix sp. were recorded only in lagoons, but Macrothrix spinosa was found in other environments like barreiros and dams. Colurella species were also found only in lagoons. In addition, the majority of Lecane, Lepadella and Cephalodella species were found in that kind of environment. The presence of these bentonic or semi bentonic species increased the number of species in Panati Lagoon. Predation can also influence the structure of the zooplankton communities. In dams we have a larger number of fish, but in 108

12 some of them the biodiversity is very low (Torelli personal communication). In Açude Soledade, in some periods, only Oreochromis niloticus was found. Lagoons also have O. niloticus, but their fish diversity is lower than in dams. Probably the lower fish predation in lagoons can contribute to the higher biodiversity in these environments. Applying cluster analysis to different periods, we can observe that the most similar pair changes from one year to another, showing that the environments highly change over the years. This way the kind of environment doesn t seem to interfere with the zooplankton communities, but probably the quality of the water, together with the habitat availability, were the main environmental processes determinig their structure. Sollberger & Paulson (1992) observed heterogeneous conditions in the presence of macrophyte vegetation, while little or no variability was found in aquatic ecosystems without vegetation. Duggan et al. (1998), analyzing rotifers in relation to littoral ecotone structure of a lake in New Zealand, observed that the characteristics of different macrophyte species were more responsible for spatial variability than the physical or chemical features. This allows the presence of a larger number of species as observed in Panati Lagoon. Shannon biodiversity indices ranged from 0.00 to The work of Sampaio et al. (2002) exhibited biodiversity indices from 1.9 to 2.7. We found lower values for both the highest and the lowest indices, but our inferior value is much lower. Our Shannon indices were considered by groups (Cladocera and Rotifera) to detect changes between them. But this can also be a result of the kind of the environment; the analyzed reservoirs were larger and not submitted to semi-arid conditions, and this could confer more stability to the environment which probably does not change much. AKNOWLEDGEMENTS This work was supported by CNPq/PELD Caatinga. We thank CNPq for the Master Degree granted to Glecia Trinta Freitas. REFERENCES Andrade, R.S Produção orgânica e o estado trófico dos sistemas aquáticos integrantes da bacia do Rio Taperoá. Monografia. UEPB, Campina Grande, PB. Crispim, M.C.; Leite, R.L & Watanabe, T Evolução do estado trófico em açudes temporários, no nordeste semi-árido, durante um ciclo hidrológico, com ênfase na comunidade zooplanctônica. Anais do V Simpósio de Ecossistemas Brasileiros: Conservação. Vitória - ES. Vol. III: Crispim, M.C. & Watanabe, T., Ovos de resistência de rotíferos presentes em sedimentos secos de um açude no semi-árido paraibano. Acta Limnologica Brasiliensia, 12(1): Crispim, M. C. & Watanabe, T What can dry reservoir sediments in a semi arid region in Brazil tell us about Cladocera? Hydrobiologia, 442: Crispim, M.C.; Paz, R. J. and Watanabe, T Comparison of different Moina minuta population dynamics ecloded from resting eggs in a semi-arid region in Brazil. Brazilian Journal of Ecology, 1/2: Dajoz, R Ecologia Geral. 3ª ed. Petrópolis. Vozes. 471p. Dodson, S Species richness of crustacean zooplankton in European lakes of different sizes. Verh. Internat. Verein. Limnol. 24: Dodson, D Predicting crustacean zooplankton species richness. Limnol. Oceanogr., 37 (4): Duggan, I.C.; Green, J.D.; Thompson, K. & Shiel, R.J Rotifers in relation to littoral ecotone structure in Lake Rotomanuka, North Island, New Zealand. Hydrobiologia, 387/388: Esteves, F.A Fundamentos de Limnologia. 2 nd edition. Interciência, Rio de Janeiro, 602p. 109

13 El Moor-Loureiro, L. M. A Manual de identificação dos cladóceros límnicos do Brasil. Brasília, UCB, 100p. Fryer, G Diapause, a potent force in the evolution of freshwater crustaceans. Hydrobiologia, 320: In: Diapause in the Crustacea, Alekseev, V.R. & Fryer, G. (ed). Kluwer Academic Publishers, Belgium, 241p. Fahd, K.; Serrano L. & Toja. J Crustacean and rotifer composition of temporary ponds in the Doñana National Park (SW Spain) during floods. Hydrobiologia, 436: Gilbert, J.J Structure, development and induction of a new diapause stage in rotifers. Freshwater Biology, 34: Hairston, N.G. and Cáceres, C.E Distribution of crustacean diapause: micro and macroevolutionary pattern and process. Hydrobiologia, 320: In: Diapause in the Crustacea, Alekseev, V.R. & Fryer, G. (ed). Kluwer Academic Publishers, Belgium, 241p. King, C.E. and Snell, T.W Densitydependent sexual reproduction in natural populations of the rotifer Asplanchna girodi. Hydrobiologia, 73: Koste, W Rotatoria: die Rädertiere Mittleuropas. Berlin, Gebruder Borntregger, 1147p. Ludwig & Reynolds Statistical Ecology - a premiere on methods and computing. Wiley & Sons Ed. USA. 337 p. Maltchik, L Biodiversidade e estabilidade em lagos do semi-árido. Ciência Hoje, 25: Martinez, C.P.; Baquero, R.M. & Castillo, P.S The effect of the volume decreasing on the trophic status in four reservoirs from Southern Spain. Verh.Internat.Verein.Limnol., 24: Patalas, K Crustacean plankton communities in forty-five lakes in the Experimental Lakes Area, Northwestern Ontario. J. Fish Res. Bd.Can., 28: Ruttner-Kolisko, A Plankton rotifers: Biology and taxonomy. Die Binnengewässer 26: (Supplement). Sampaio, E.V.; Rocha, O.; Matsumura-Tundisi, T & Tundisi J.G Composition and abundance of zooplankton in the limnetic zone of seven reservoirs of the Paranapanema River, Brazil. Braz. J. Biology. 62 (3): Schwartz, S.S. & Jenkins, D.G Temporary aquatic habitats: Constraints and opportunities. Aquatic Ecology. 34: 3-8. SEMARH/LMRS-PB. Secretaria Extraordinária do Meio Ambiente, Recursos Hídricos e Minerais / Laboratório de Meteorologia, Recursos Hídricos e Sensoriamento Remoto da Paraíba. URL: consulted in 29/jan/04. Sládecek, V Rotifers as indicators of water quality. Hydrobiologia, 100: Sollberger, P.J.; Paulson, P.J Littoral and limnetic zooplankton communities in Lake Mead, Nevada-Arizona, USA. Hydrobiologia, 237: StatSoft, Inc STISTICA for Windows [Computer program manual]. StatSoft, Inc.: Tulsa (U.S.A). URL: Vieira, D.M. Crispim, M.C. Watanabe, T Impacto da cheia e da seca sobre a comunidade zooplanctônica do açude São José dos Cordeiros, semi-árido paraibano. Anais do V Simpósio de Ecossistema Brasileiros: Conservação. Vitória - ES. Vol. III: Winterbourn, M.J. & Lewis, M.H Littoral fauna. In: Jolly, V.H. & Brown, M.A. (eds), New Zealand Lakes. Auckland University Press, Auckland: Wright, J.F.; Gunn, R.J.M.; Winder, J.M.; Wiggers, R.; Kneebone, N.T. & Clarke, R.T The impact of drought events in 1976 and 1997 on the macroinvertebrate fauna of a chalk stream. Verh.Internat.Verein.Limnol 110

14 [1] Project supported by CNPq/PELD Caatinga [2] Universidade Federal da Paraíba, Centro de Ciências Exatas e da Natureza, Depto. De Sistemática e Ecologia. Cidade Universitária João Pessoa-PB, Brazil. [3] CNPq Master Degree grant. 111

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