- Citado por Google
- Similares en SciELO
- Similares en Google
versión impresa ISSN 0716-078X
Rev. chil. hist. nat. v.81 n.2 Santiago jun. 2008
Revista Chilena de Historia Natural 81: 185-203,2008
Finding patterns of distribution for freshwater phytoplankton, zooplankton and fish, by means of parsimony analysis of endemicity
Encontrando patrones de distribución para fitoplancton, zooplancton y peces dulceacuícolas por medio de análisis de parsimonia de endemismos
J. PABLO OYANEDEL1, CAREN VEGA-RETTER1, SERGIO SCOTT1, LUIS FELIPE HINOJOSA1,2 & RODRIGO RAMOS-JILIBERTO1*
1Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Casilla 653 Santiago, Chile.
During the last decades, limnological studies on Chilean systems have contributed to know the species composition and main environmental variables of many water bodies distributed over a wide latitudinal interval, from 18º to 53º S. However, we still lack of a comprehensive view about the structure and functioning of regional freshwaters. In this work we review the available information about pelagic biota from Chilean basins, in order to reveal patterns of species distribution and their possible association with environmental variables. We built presence-absence matrices for phytoplankton, zooplankton and fish over lakes and basins. From this database, we performed parsimony analysis of endemicity as a tool for determining fundamental distribution patterns of freshwater biota. Also, we assessed the relationship between species occurrences and some available site-related variables. Our results indicated that latitude exerted the strongest influence on species distribution, although altitude, longitude, and area also exerted significant effects for some groups. On the other hand, our results suggest a relationship between the degree of vagility of the groups and the degree of metacommunity structuring, related to the number of endemicity areas.
Key words: limnology, lakes, plankton, biogeography, Chile, CCA.
Durante las últimas décadas, los estudios limnológicos en sistemas chilenos han contribuido al conocimiento de la composición de especies y de las principales variables ambientales de muchos cuerpos de agua distribuidos sobre un amplio intervalo latitudinal, desde los 18º a los 53º S. Asimismo, aún carecemos de una visión comprensiva acerca de la estructura y funcionamiento de las aguas dulces regionales. En este trabajo revisamos la información sobre la biota pelágica de las cuencas de Chile, con el propósito de revelar patrones de distribución de especies y su posible asociación con variables ambientales. Construimos matrices de presencia-ausencia para fitoplancton, zooplancton y peces en lagos y cuencas. Desde esta base de datos, realizamos análisis de parsimonia de endemismo, como medio para determinar patrones fundamentales de distribución de la biota dulceacuícola. También, determinamos la relación entre la presencia de especies y algunas variables relevantes de los sitios. Nuestros resultados sugieren que la latitud ejerce una fuerte influencia en la distribución de especies, aunque también la altitud, longitud y el área ejercen efectos significativos en algunos grupos. Por otro lado, los resultados sugieren una relación entre el grado de vagilidad de los grupos y el grado de estructuración de la metacomunidad, relacionada con el número de áreas de endemismos.
Palabras clave: limnología, lagos, plancton, biogeografía, Chile, CCA.
Understanding patterns of biodiversity and community structure on a given region, as much as developing guidelines for conservation of biota and environmental care, requires knowledge about patterns of distribution and abundance of entire taxonomic groups. In southern latitudes, data about freshwater biota are often scarce, incomplete, and dispersed. Particularly in Chile, the field of limnology is still in an early stage from the viewpoint of scientific development, and accordingly, most ecologically oriented work on freshwaters habitats has been mainly descriptive and comparative, with modest progress in experimental or theoretical research. In spite of several decades of valuable descriptive work, regional information about freshwater biota is fragmented and biased, both geographically and taxonomically. As a consequence, we lack of basic knowledge about patterns of distribution and biodiversity, which contrasts sharply with the abundance and variety of freshwater bodies found in the country. Chilean freshwaters exhibit a wide spectrum of physical features associated to strong gradients of climate, lithology, topography and vegetation cover, in agreement with their broad latitudinal range and the strong influences from high Andes and Pacific Ocean.
Here we intend for the first time to review the available information about key components of pelagic biota from Chilean freshwaters, in order to reveal patterns of species distribution and their possible association with environmental variables.
Parsimony analysis of endemicity (PAE) is a powerful biogeographical tool, useful for determining distributional patterns of species and for identifying biodiversity hotspots (Garrafoni et al. 2006). For these reasons, PAE can help for building up a comprehensive view on some aspects of the structure freshwater biota, as well as for guiding conservation planning. Originally developed by Rosen (1988), PAE aims to 80 classify areas by most parsimonious solutions based on the shared presence of species (Nihei 2006). This allows for the identification of areas with non-random distributional congruence among different species (Morrone 1994).
In this work we review the available information about distribution of freshwater phytoplankton, zooplankton, and fish species from mainland Chile. We use the compiled data for finding out basic distribution patterns for the analyzed biota through (a) testing whether the distribution of species across lakes and basins can be statistically explained by the environmental variables at hand, and (b) identifying areas of endemism for each of the groups of interest.
MATERIAL AND METHODS
We attempted to review all published information (in both indexed and not-indexed journals) about species distribution of phytoplankton, zooplankton and fish from Chilean freshwaters. We were able to find 41 articles with reliable data, and three additional papers containing environmental information needed for our analyses. The published data spanned from year 1973 to 2006. We studied 55 lakes located between 18.25º and 53.46º S, although most of them concentrate between 32.54º and 46.5º S (Fig. 1).
From this dataset we constructed presence/absence 103 matrices for each taxon (available upon request), with species as columns and sites as rows. Lakes and basins were treated separately as sites. For phytoplankton (found in 22 lakes), we constructed matrices for the three major taxa Bacillariophyceae (diatoms, 18 lakes), Chlorophyceae (green algae, 18 lakes), and Cyanophyceae (blue-green algae, 16 lakes). For zooplankton (52 lakes), we analyzed separately the three major taxa Cladocera (41 lakes), Copepoda (47 lakes), and Rotifera (31 lakes). Fish were treated as a single taxon and only data by basins were available.
In order to avoid anecdotic records, we did not considered lakes below percentile 10 of species richness, within each taxonomic group. For species names, we maintained the nomenclature contained in the original references.
We also reviewed the available environmental information associated to each site. For lakes, we recorded area, latitude, longitude, and altitude. Most of these data were obtained from the published information. For lakes Pichilafquen, Quillehue, and Chiguay, geographic coordinates were taken from Google Earth (http://earth.google.com). Surface area for lakes Pichilafquen, Quillehue, Chiguay, Atravesado, Bonita, Huilipilun, La Posada, Patos Bravos, and Lynch were measured with the software Image Tool 3.0 (http://ddsdx.uthscsa.edu/dig/itdesc.html) from images obtained with Google Earth.
A total of 40 basins were defined according to the official website of Dirección General de Aguas (http://www.dga.cl), from which we also obtained their surface area and perimeter. For determining the characteristic geographic coordinates of each basin, we followed the following protocol. For exhorreic basins between 22.47º and 48.20º S, basin coordinates were defined as those of the middle point of their main river. For coastal basins between 31.78º and 38.25º S, basin latitude was defined as the mean of those of the two rivers that limit the basin; and longitude was defined as that of the shoreline 127 at the basin latitude. Basins between 42.54º and 53.83º S present a large region of flooded land, and their coordinates correspond to that of the middle point of the entire area. This same procedure was used for basins Altiplanica and Budi. Chilean basins were also grouped into seven hydrographic zones, following official terms (http://www.igm.cl). Matrices with phytoplankton and zooplankton by basins were constructed from the information of species occurrences in the lakes that belong to each basin.
In order to establish the possible relationship between the presence/absence of several taxa in both Chilean lakes and basins and their environmental variables described below, we performed a multivariate direct gradient analysis, specifically a canonical correspondence analysis (CCA, Ter Braak 1986). CCA extracts continuous axes of variation from species occurrence in the light of known environmental variables (EV), by imposing the constraint that axes are linear combinations of EV. Hence, the relationship between species occurrences and EV are assumed to be linear. In order to determine whether these relationships are statistically significant, we performed Monte Carlo permutation tests with 1,000 runs (Manly 1991). The results of statistically significant CCA are displayed using biplots diagrams of sites (symbols) and EV (vectors). Both CCA and permutation tests were done using the package CANOCO v. 4.5 (Ter Braak & Smilauer 1998).
Parsimony analysis of endemicity
We performed parsimony analysis of endemicity (151 PAE, Rosen 1988, Morrone 1994) in order to determine possible areas of endemism. The analysis was based on a data matrix with species (characters) as columns and basins as rows (OGU's). The character states were coded as presence/absence (1/0) of the species on sites. For these analyses we used basins instead lakes as sites, in order to minimize the effect of incomplete sampling of species across lakes. A hypothetical basin without any species presence was used for rooting the tree. Data were analyzed with the software PAUP* 4.0b10 (Swofford 2001), using the heuristic search algorithm, randomizing the OGU's entries with 100 replicates. All characters were treated as ordered (Wagner parsimony). The strict consensus tree was calculated, that conserves the most robust grouping of localities and minimizes the influence of widely distributed species (Morrone 1994). A clade defined by at least two species was considered to be an endemicity area (Morrone 1994). We also performed PAE for a matrix that included all species and basins, where a clade defined by at least two species of each taxon was considered to be an endemicity area.
Seven CCA were statistically significant (Monte Carlo permutation test, P < 0.05, Table 1), from a total of 17 studied groups. Total zooplankton, copepods, rotifers, and fish consistently exhibited significant associations with the studied EV. In Fig. 2-4 we show bi-plots summarizing the relationship between the freshwater biota (symbols, classified according hydrographic Chilean zones) and EV (vectors).
In general terms, latitude represented the main gradient for all groups considered here. This gradient followed the north-to-south succession through the hydrographic zones, crossing the three main climatic regions of Chile: arid, Mediterranean and temperate. Considering the analyses based on lakes, the second gradient was dominated by altitude, except for rotifers where surface area was the main factor. On the other hand, the second gradient in basins-based analyses was dominated by either longitude (total zooplankton, copepods) or surface area (rotifers, fish). Note that for rotifers, surface area covaried with perimeter of the basin.
Results of CCA accounting distribution of total zooplankton on lakes indicated that the two first axes represent 80.6 % of total variance (Fig. 2A). The first axis was mainly correlated with latitude (r = -0.91) whereas second axis was correlated with altitude (r = 0.75). In CCA by basins (Fig. 2B), the first two axes represente 62.5 % of total variance; the first axis was mainly correlated with latitude (r = -0.80) and second axis was correlated with longitude (r = 0.75).
In CCA of copepod distribution on lakes (Fig. 3A), the first axis was mainly correlated with latitude (r = -0.82) and the second axis was correlated with altitude (r = 0.80). In CCA by basins (Fig. 3C), the two first axes represented 70.5 % of total variance. The first axis was mainly correlated with latitude (r = -0.58) whereas second axis was correlated with longitude (r = 0.88).
Multivariate analyses testing association of rotifer distribution on lakes and EV (Fig. 3B) showed that the two first axes represent 62.4 % of total variance, where the first axis was mainly correlated with latitude (r = 0.90) while the second axis was correlated with surface area (r = 0.60). In CCA by basins (Fig. 3D), the first two axes represented 60.8 % of total variance; the first axis was mainly correlated with latitude (r = -0.75) and second axis with area (r = -0.44).
Finally, in CCA of fish (Fig. 4) the first axis was mainly correlated with latitude (r = 0.83) whereas second axis was correlated with surface area (r = 0.35).
The analysis of the total information (i.e. a single matrix with all taxa and basins included) did not provided additional information, showing that latitude and area constituted the main driving gradients for the biota analyzed here.
Parsimony analysis of endemicity
Based on the analyzed information, endemicity areas were identified for all taxonomic groups with the exception of blue-green algae. Green algae (Chlorophyceae, Fig. 5A and 8A), exhibited two partially overlapping endemicity areas. The first clade (defined by Closterium acutum and Sphaerocystis schroeteri) is located between -19.913º and -51.247º S, and was formed by three joint basins (AA, AB, AC, see Table A-1 in the appendix) and four disjoint basins (A, N, S, and AL). The second clade defined for green algae (Ankistrodesmus falcatus, Scenedesmus quadricauda and Staurastrum polymorphum) is located between -33.317º and -39.015º S, and comprised four basins, inside which there were two sub-areas of endemicity: L, Z and R, T. For diatoms (Bacillariophyceae, Fig. 5B and 8B) we observed two endemicity areas. The first clade (defined by Aulacoseira granulata, Cocconeis placentula, and Synedra rumpens) comprised three disjoint basins (-19.913º and -42.542º S) A, N, and AD. The second clade (Melosira granulata, and Surirella guatimalensis) extends between -33.317º and -51,247º S and was composed by 10 basins (L, R, M, S, T, Z, AB, AC, AL, and AA).
Rotifers exhibited two endemicity areas. The first clade (defined by species Keratella valga and Lepadella ovalis) was built by two adjacent basins (J-K, -32.664º to - 32.735º S), while the second (Kellatera cochlearis, Conochilus unicornis, Hexarthra fennica and Polyarthra vulgaris, -33.317º to -42.542º S) was represented by a group of nine basins that included two continuous sub-areas comprising basins L-M-N and the region from basin Z till AC plus the island AD, respectively (Fig. 6A and 9A). Copepods exhibited three endemicity areas. The first clade (Tumeodiaptomus diabolicus and Tropocyclops prasinus meridionalis ) is a large group of eight basins located between - 33.711º and -42.542º S, inside which there was a continuous sub-area composed by basins Z, AA, AB, and AC. A second endemicity area for copepods (Parabroteas sarsii and Eucyclops Serrulatus) locates between -45.313º and -47.468º S and it was defined by the union of AH and AI. The last area (Diaptomus diabolicus and Mesocyclops longisetus) extends from -32.427º to -34.114º and corresponded to the disjoint basins J-L-N (Fig. 6B and 9B). The last zooplankton group, Cladocera, presented a single large endemicity area (defined by species Diaphanosoma chilense and Daphnia ambigua) covering basins A, L, M, N, V, Z, AA, AB, and AD, inside which there was a continuous sub-area composed by basins Z, AA, AB, located between -39.015º and -40.334º S (Fig. 6C and 9C).
If we consider only the native fish species for the identification of endemicity areas, the first clade would loss basin D and it would be defined by Diplomystes chilense, Basilichthys australis, Percichthys melanops. In the same vein, the third and fourth endemicity areas would disappear.
The analysis of the total information (all taxa included) showed four endemicity areas. A first area appeared in all groups analyzed (basins Z-AD). The second area (L, N) was already defined for plankton groups, the area M, S appeared for green algae, and the fourth area AL was defined for green algae, diatoms, and copepods. The number of retained trees was 7717, with a length of 819. Consistency Index of this tree was 0.37, and Retention Index was 0.46.
Our results revealed that basin AB (Bueno), N (Rapel), Z (271 Toltén), AA (Valdivia), and L (Coasta Aconcagua-Maipo) form part of endemicity areas for all groups studied here, excluding blue-green algae. Basins AC (Basin/Island Bueno-Puelo), and M (Maipo) form part of endemicity areas for five out of six groups (excluding blue-green algae).
PAE performed for lakes (not shown here) exhibited more confuse results, although they show agreement with the basin-based analyses regarding the main endemicity areas.
Our CCA results revealed that neither phytoplankton nor cladocerans exhibit a distribution trend that could be explained by the environmental variables considered here. On the other hand, latitude was the main factor explaining species distribution of total zooplankton, copepods, rotifers, and fish. Altitude/longitude constituted also an important gradient for total zooplankton and copepods, while lake surface did so for rotifers. Since this work only assessed general distributional trends of a regional freshwater biota, mechanisms generating the observed patterns are still unknown. Nevertheless, previous works emphasize that phytoplankton distribution is often driven by dispersal, while distribution of fish is driven by vicariance. This relates to the relative vagility of the two groups, with zooplankton locating midway between phytoplankton and fish. This agrees with the lack of significance in CCA results for algae.
Phytoplankton dispersal mechanisms are largely passive ones. Passive dispersal of algae rest on airborne transport operating at distances, while foretic movements mediated mostly by beetles, dragonflies and waterbirds are important mechanisms over shorter distances. Furthermore, local dispersal also occurs by means of drift among rivers and streams (Kristiansen 1996). On the other hand, dispersal mechanisms of zooplankton are often associated to resting stages (ephipia in cladocerans, diapausing eggs in rotifers, and criptobiosis in rotifers and copepod nauplii). Similar to algae, passive dispersal of zooplankton resting stages occurs mediated by wind, drift, and foretic transport through invertebrates and waterbirds (Bilton et al. 2001, Bohonak & Jenkins 2003, Havel & Shurin 2004). Dispersal determines at a large extent the distributional patterns of algae, rotifers and cladocerans; but distribution of copepods -being less vagile- is often determined by both vicariance and dispersal mechanisms (Rayner & Heeg 1994). On the other hand, fish disperse by active means and therefore their movements depend on the connectivity of the riparian network, as well as their life-history strategies. This explains the importance of vicariance for the current distribution of freshwater fish species (Anderson 1996, McDowall 2002, Cunha Ribeiro 2006).
Our results from endemicity analyses show that highly vagile groups due to their passive dispersal mechanisms such as algae, rotifers and cladocerans, exhibit at most two endemicity areas over the studied region. Intermediate levels of vagility associated to copepods agree with the finding of intermediate number of endemicity areas for this group, which are distributed along a latitudinal gradient. Fish exhibit the minimal relative vagility, and consistently they show the largest number of endemicity areas along the north-south gradient.
As a cautionary note, consider that some endemicity areas may have been defined by species that, in global sense, are widely distributed. Nevertheless, species that define endemicity areas within a region have a restricted area therein.
The results of our analyses reveal that latitude strongly influence distribution of some groups inhabiting Chilean freshwaters, and relative vagility among groups seems to influence the degree of metacommunity structure. Previous studies also support latitudinal patterns of distribution. For example, Henio (2001) found latitudinal gradient in species distribution of macrophytes, beetles, dragonflies, and stoneflies in the north hemisphere; Chengalath & Koste 1989, see also Segers 1996) found latitudinal variation in distribution patterns of rotifers in the circumpolar region; and Weckstroèm & Korhola (2001) found similar trends for diatoms in Fennoscandia.
In spite of the fact that dispersal abilities of organisms, particularly of planktonic ones, are important determinants of their current distribution (Fenchel et al. 1997, Finlay 2002, Fenchel & Finlay 2004), establishment on a given site strongly depends on the success of species in the front of abiotic constraints and natural enemies, as well as on their ability for exploiting local resources. On the other hand, new species can also appear by speciation mechanisms other than vicariance (Maidana et al. 2005, Pedrós-Alió 2006).
In this way, local physical and chemical conditions of waters bodies such as light penetration, pH, nutrients availability, and lake morphometry among others, might exert significant effects on species richness and distribution (Gutiérrez-Aguirre & Suárez-Morales 2001, Duggan et al. 2002). Unfortunately, we lack of such information for the bulk of systems analyzed here, and consequently we were not able to test their influence. Nevertheless our results may serve as a start point for future research on metacommunity structure of regional freshwaters, as well as for focal hypothesis testing. Latitude is associated with climate and soil composition, which partially determine basin and lake morphometry, water chemistry, and temperature. Organisms' biology defines their life histories as well as dispersal abilities and hence colonization probabilities. At the same time, environmental features interacting with biological traits influence the structure and dynamics of populations and communities, which in turn affects local persistence. Therefore, from the rough relationships between freshwater biota and environmental variables shown here, future research on local systems could advance towards revealing the actual ecological mechanisms driving distributional patterns of species occurrences. Finally, our analysis also identified geographic areas of endemicity for each of the groups considered, and the species that define them. These results should be considered for decision making in the field of natural reserve design and conservation planning.
We thank J.L. Fajardo for collecting an important amount of basal information. We also thank I. Vila, V. Montecino and L. Zúñiga for providing literature difficult to find. This work was supported by grants CONICYT-PBCT ACT 34/2006 and MULT-05-04/2 from Dirección de Investigación of Universidad de Chile. Sergio Scott acknowledges financial support by MECESUP UCO-214 doctoral fellowship.
ACUÑA P, I VILA, PARDO R & COMTE S (2005) Caracterización espacio temporal del nicho trófico de la fauna íctica andina del río Maule, Chile. Gayana (Chile) 69: 175-179. [ Links ]
ANDERSON M E (1998) A late cretaceous (Maastrictian) galaxiid fish from south Africa. Special Publication of the J.L.B. Smith Institute of Ichthyology 60: 1-12. [ Links ]
ANDREW TE, S CABRERA & V MONTECINO (1989) Diurnal changes in zooplankton respiration rates and the phytoplankton activity in two Chilean lakes. Hydrobiologia 175: 121-135. [ Links ]
ARAYA J & L ZÚÑIGA (1985) Manual taxonómico del zooplancton lacustre de Chile. Boletín Informativo Limnológico (Chile) 8: 1-110. [ Links ]
ARRATIA G (1981) Géneros de los peces de las aguas continentales de Chile. Publicación Ocasional, Museo Nacional de Historia Natural (Chile) 34: 3-108. [ Links ]
ARRATIA G (1983) Trichomycterus chungaraensis n. sp. (Pisces, Siluriformes) Trichomycteridae from the high Andean range. Studies on Neotropical Fauna and Environment 18: 65-87. [ Links ]
BERRA TM, RM FELTES & VH RUIZ (1995) Brachygalaxias gothei from sourth-central Chile, a synonym B. bulloki (Osteichthyes: Galaxiidae). Ichthyological Explorations of Freshwaters 6: 227-234. [ Links ]
BILTON DT, R FREELAND & B OKAMURA (2001) Dispersal in freshwater invertebrates. Annual Review of Ecology and Systematics 32: 159-810. [ Links ]
BOHONAK AJ & DG JENKINS (2003) Ecological and evolutionary significance of dispersal by freshwater invertebrates. Ecology Letters 6: 783-796. [ Links ]
CAMPOS H (1984) Limnological study of Araucanian lakes 385 (Chile) Internationale Vereinigung für Theoretische und Angewandte Limnologie 22: 1319-1327. [ Links ]
CAMPOS H (1985) Distribution of the fishes in the Andean rivers in the south of Chile. Archiv für Hydrobiologie 104: 169-191. [ Links ]
CAMPOS H, D SOTO, W STEFFEN, G AGÜERO, O PARRA & L ZÚÑIGA (1994) Limnological studies of lake El Toro (Chile). Morphometry, physics, chemistry and plankton. Archiv für Hydrobiologie Supplement 99: 199-215. [ Links ]
CAMPOS H, F GAVILÁN, V MURRILLO & P ALARCÓN (1996) Presencia de Cheirodon australes en el lago Tarahuin (isla grande de Chiloé, 42º40' S, Chile) y su significado zoogeográfico. Medio ambiente (Chile) 13: 69-79. [ Links ]
CAMPOS H, G DAZAROLA, B DYER, L FUENTE, L HUAQUIN, G MARTÍNEZ, R MELÉNDEZ, G PEQUEÑO, F PONCE, V RUIZ, W SIELFELD, D SOTO, R VEGA & I VILA (1998) Categorías de conservación de peces nativos de aguas continentales de Chile. Boletín del Museo Nacional de Historia Natural (Chile) 47: 101-122. [ Links ]
CAMPOS H, J ARENAS, W STEFFEN, C ROMAN & G AGÜERO (1982) Limnological study of lake Ranco (Chile). Morphometry, physics, chemistry and plankton. Archiv für Hydrobiologie 94: 137-171. [ Links ]
CAMPOS H, W STEFFEN, C ROMAN, L ZUÑIGA & G AGÜERO (1983) Limnological study of lake Villarrica. Morphometric, physical, chemical, planktonical factors and primary productivity. Archiv für Hydrobiologie Supplement 4: 371-406. [ Links ]
CAMPOS H, W STEFFEN, G AGÜERO, O PARRA & L ZÚÑIGA (1992b) Limnological studies of lake Rupanco (Chile) morphometry, physics, chemistry, plankton and primary productivity. Archiv für Hydrobiologie Suplement 90: 85-113. [ Links ]
CAMPOS H, W STEFFEN, G AGÜERO, O PARRA & L ZÚÑIGA (1992a) Limnology of lake Ranco (Chile). Limnologica 22: 337-353. [ Links ]
CAMPOS H, W STEFFEN, O PARRA, G AGÜERO & L ZÚÑIGA (1987a) Limnology of lake Riñihue. Limnologica 18: 339-357. [ Links ]
CAMPOS H, W STEFFEN, O PARRA, G AGÜERO & L ZÚÑIGA (1988) Limnological study of Lake Llanquihue (Chile). Morphometry, physics, chemistry, plankton and Primary Productivity. Archiv für Hydrobiologie Supplement 81: 37-67. [ Links ]
CAMPOS H, W STEFFEN, O PARRA, G AGÜERO & L ZÚÑIGA (1989) Estudios limnológicos en el lago Puyehue (Chile): morfometría, factores físicos y químicos, plancton y productividad. Medio Ambiente (Chile) 10: 36-53. [ Links ]
CAMPOS H, W STEFFEN, O PARRA, P DOMÍNGUEZ & G AGÜERO (1987b) Estudios limnológicos en el lago Caburgua (Chile). Gayana Botanica (Chile) 44: 61-84. [ Links ]
CHANG A, G ARRATIA & G ALFARO (1978) Percichthys lonquimayiensis n. sp. from the upper Paleocene of Chile (Pisces, Perciformes, Serranidae). Journal of Paleontology 52: 727-736. [ Links ]
CHENGALATH R & W KOSTE (1989) Composition and distributional patterns in Arctic rotifers. Hydrobiologia 186/187: 191-200. [ Links ]
CRUCES F, R URRUTIA, A ARANEDA, L TORRES, M CISTERNAS & W VYVERMAN (2001) Evolución trófica de laguna Grande de San Pedro (VIII Región, Chile) durante el ultimo siglo, mediante el análisis de registro sedimentarios. Revista Chilena de Historia Natural 74: 407-418. [ Links ]
CUNHA RIBEIRO A (2006) Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotropical Ichthyology 4: 225-246. [ Links ]
DOMÍNGUEZ P & D CASANOVA (1989) Advección superficial 433 del fitoplancton en la laguna Grande de San Pedro (Concepción, Chile) Anales Museo de Historia Natural de Valparaíso, Chile 20: 5-18. [ Links ]
DOMÍNGUEZ P & L ZÚÑIGA (1979) Perspectiva temporal de la entomostracofauna limnética de lago Ranco (Valdivia, Chile). Noticiero Mensual del Museo de Historia Natural de Valparaíso (Chile) 12: 53-58. [ Links ]
DOMÍNGUEZ P (1973) Contribución al estudio de los cladóceros Chilenos. I. Cladóceros del lago Chungará. Noticiero Mensual del Museo de Historia Natural de Valparaíso (Chile) 201-202: 3-10. [ Links ]
DONOSO TG & HK PHINNEY (1988) The phytoplankton of Lago Rupanco, Osorno, Chile. Archiv für Hydrobiologie 113: 189-211. [ Links ]
DUGGAN IC, JD GREEN & JS RUSSELL (2002) Distribution of rotifer assemblages in north island, New Zealand, lakes: relationships to environmental and historical factors. Freshwater Biology 47: 195-206. [ Links ]
DYER BS (2000a) Systematic review and biogeography of the freshwater fishes of Chile. Estudios Oceanológicos (Chile) 19: 77-98. [ Links ]
DYER BS (2000b) Revisión sistemática de los pejerreyes de Chile (Teleostei, Ateriniformes). Estudios Oceanológicos (Chile) 19: 99-127. [ Links ]
FENCHEL T & B J FINLAY (2004) The ubiquity of small species: patterns of local and global diversity. BioScience 54: 777-784. [ Links ]
FENCHEL T, GF ESTEBAN & B J FINLAY (1997) Local versus global diversity of microorganisms: cryptic diversity of ciliated protozoa. Oikos 80: 220-225. [ Links ]
FINLAY BJ (2002) Global dispersal of free-living microbial eukaryote species. Science 296: 1061-1063. [ Links ]
GARRAFONI ARS, SS NIHEI & PDC LANA (2006) Distribution patterns 457 of Terebellidae (Annelida: Polychaeta): an application of parsimony analysis of endemicity (PAE). Scientia Marina 70S3: 269-276. [ Links ]
GUTIÉRREZ-AGUIRRE MA & E SUÁREZ-MORALES (2001) Diversity and distribution of freshwater copepods (Crustacea) in southeastern Mexico. Biodiversity and Conservation 10: 659-672. [ Links ]
HABIT E (2005) Aspectos de la biología y hábitat de un pez endémico de Chile en peligro de extinción (Diplomystes nahuelbutaensis Arratia 1987) Interciencia 30: 1-10. [ Links ]
HABIT E, B DYER & I VILA (2006) Estado del conocimiento de los peces dulceacuícolas de Chile. Gayana (Chile) 70: 100-113. [ Links ]
HABIT E, P VICTORIANO & O PARRA (2002) Translocación de peces nativos en la cuenca del río Laja. Gayana (Chile) 66: 181-190. [ Links ]
HABIT E, P VICTORIANO, A RODRÍGUEZ & V RUIZ (2003) Variaciones espacio temporales del ensamble de peces de un sistema fluvial de bajo orden del centro-sur de Chile. Revista Chilena de Historia Natural 76: 3-18. [ Links ]
HAVEL JE & JB SHURIN (2004) Mechanisms, effects, and scales of dispersal in freshwater zooplankton. Limnology and Oceanography 49: 1229-1238. [ Links ]
HENIO J (2001) Regional gradient analysis of freshwater biota: do similar biogeographic patterns exist among multiple taxonomic groups? Journal of Biogeography 28: 69-76. [ Links ]
KELLER B & D SOTO (1998) Hydrogeologic influences on the preservation of Orestias ascotanenesis (Teleostei: Cyprinodontidae), in Salar de Ascotán, Northerm Chile. Revista Chilena de Historia Natural 71: 147-156. [ Links ]
KRISTIANSEN J (1996) Dispersal of freshwater algae - a review. Hydrobiologia 336: 151-157. [ Links ]
MAIDANA NI, I IZAGUIRRE, A VINOCUR, G MATALONI & H PIZARRO (2005) Diatomeas en una transecta patagónico-antártica. Ecología Austral (Argentina) 15: 159-176. [ Links ]
MANLY BF (1991) Randomization and Monte Carlo methods in biology. Champman & Hall, New York,USA. 281 pp. [ Links ]
MCDOWELL R M (2002) Accumulating evidence for a dispersal biogeography of southern cool temperate freshwater fishes. Journal of Biogeography 29: 207-219. [ Links ]
MORRONE JJ (1994) On the identification of areas of endemism. Systematic Biology 43: 438-441. [ Links ]
NIHEI SS (2006) Misconceptions about parsimony analyses of endemicity. Journal of Biogeography 33: 2099-2106. [ Links ]
OLGUÍN HF & P DOMÍNGUEZ (1995) Corrientes, turbulencias y distribución de las algas dominantes en el sector poniente del embalse Peñuelas (Valparaíso, Chile). Anales del Museo de Historia Natural de Valparaíso (Chile) 23: 5-20. [ Links ]
PARRA O, C VALDOVINOS, M URRUTIA, M CISTERNAS, E HABIT & M MARDONES (2003) Caracterización y tendencias tróficas de cinco lagos costeros de Chile central. Limnetica 22: 51-83. [ Links ]
PARRA OO & M GONZÁLEZ (1975) Synechocystis sauvageau, nuevo género de Cyanophyta para Chile. Boletín de la Sociedad de Biología de Concepción (Chile) 49: 153-155. [ Links ]
PARRA OO, V DELLAROSSA & E UGARTE (1976) Estudio 502 limnológico de las lagunas "Chica de San Pedro", "La Posada" y "Lo Méndez" I. Análisis cuali y cuantitativo del plancton invernal. Boletín de la Sociedad de Biología de Concepción (Chile) 50: 73-86. [ Links ]
PEDRÓS-ALIÓ C (2006) Marine microbial diversity: can it be determined? Trends in Microbiology 14: 257-263. [ Links ]
PEQUEÑO G (1997) Peces de Chile. Lista sistemática revisada y comentada: addendum. Revista de Biología Marina y Oceanografía (Chile) 32: 77-94. [ Links ]
RAYNER NA & J HEEG (1994) Distribution patterns of the Diaptomidae (Calanoida: Copepoda) in southern Africa. Hydrobiologia 272: 47-75. [ Links ]
ROSEN BR (1988) From fossils to earth history: applied historical biogeography. In: Myers A & P Giller (eds) Analytical biogeography: an integrated approach to the study of animal and plant distribution: 437-481. Chapman and Hall, London, United Kingdom. [ Links ]
RUBILAR A & E ABAD (1990) Percichthys sylviae n. sp. del Terciano de los Andes sur centrales de Chile (Pisces, Perciformes, Percichthydae). Revista Geológica de Chile 17: 197-204. [ Links ]
RUBILAR A (1994) Diversidad ictiológica en depósitos continentales. Revista Geológica de Chile 21: 3-29. [ Links ]
RUIZ R & N BAHAMONDE (1989) Cladóceros y copépodos límnicos en Chile y su distribución geográfica. Publicación ocasional, Museo Nacional de Historia Natural (Chile) 45: 1-48. [ Links ]
RUIZ R & N BAHAMONDE (2003) Distribución estacional de cladóceros y copépodos en el lago Rapel, Chile central. Publicación ocasional, Museo Nacional de Historia Natural (Chile) 58: 5-58. [ Links ]
SCHMID-ARAYA JM & LR ZÚÑIGA (1992) Zooplankton community 525 structure in two Chilean reservoirs. Archiv für Hydrobiologie 123: 305-335. [ Links ]
SCHMID-ARAYA JM (1993) Rotifer communities from some araucarian lakes of southern Chile. Hydrobiologia 255/256: 397-409. [ Links ]
SEGERS H (1996) The biogeography of littoral Lecane Rotifera. Hydrobiologia 323: 169-197. [ Links ]
SOTO D & L ZÚÑIGA (1991) Zooplankton assemblages of Chilean temperate lakes: a comparison with North American counterparts. Revista Chilena de Historia Natural 64: 569-581. [ Links ]
SWOFFORD DL (2001) PAUP*: phylogenetic analysis. Using parsimony (* and other methods), version 4.0b10. Sinauer Associates, Sunderland, Massachusetts, USA. [ Links ]
TER BRAAK C (1986) Canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis. Ecology 67: 1167-1179. [ Links ]
TER BRAAK JF & P SMILAUER (2002) CANOCO reference manual and CanoDraw for Windows user's guide: software for canonical community ordination (version 4.5). Microcomputer Power, Ithaca, New York, USA. 500 pp. [ Links ]
URRUTIA R, K SABBE, F CRUCES, K POZO, J BECERRA, A ARANEDA, W VYVERMAN & O PARRA (2000) Paleolimnological studies of Laguna Chica of San Pedro (VIII Region): diatoms, hydrocarbons and fatty acid records. Revista Chilena de Historia Natural 73: 717-728. [ Links ]
VALERO-GARCÉS JBB, R URRUTIA, K KELTS, H VEIT, PG APPLEBY & M GEYH (2002) Moisture changes and fluctuations of the westerlies in mediterranean central Chile during the last 2000 years: the Laguna Aculeo record (33º50` S). Quaternary International 87: 3-18. [ Links ]
VILA I & R PARDO (2003) Respuesta de la estructura fitoplanctónica a las perturbaciones antrópicas de un lago templado. Limnetica 22: 93-102. [ Links ]
VILA I, L FUENTES & M CONTRERAS (1999a) Peces límnicos de Chile. Boletín del Museo Nacional de Historia Natural (Chile) 48: 61-75. [ Links ]
VILA I, L FUENTES & M SAAVEDRA (1999b) Ichthyofauna in limnic system of isla grande, tierra del fuego. Revista Chilena de Historia Natural 72: 273-284. [ Links ]
VILLALOBOS L (1994) Distribution of Daphnia in high mountain and temperete lakes of South America. Internationale Vereinigung für Theoretische und Angewandte Limnologie 25: 2004-2404. [ Links ]
VILLALOBOS L, O PARRA, M GRANDJEAN, E JAQUES, S WOELFL & H CAMPOS (2003) A study of the river basins and limnology of five humic lakes on Chiloé island. Revista Chilena de Historia Natural 76: 563-590. [ Links ]
VILLWOCK W & SIENKNECHT (1996) Contribución al conocimiento e historia de los peces chilenos: los Cyprinodóntidos del género Orestias Val. 1839 (Téleostei: Cyprinodontidae) del altiplano Chileno. Medio Ambiente (Chile) 13: 119-126. [ Links ]
WECKSTROÈM J & A KORHOLA (2001) Patterns in the distribution, composition and diversity of diatom assemblages in relation to ecoclimatic factors in Arctic Lapland. Journal of Biogeography 28: 31-45. [ Links ]
ZUNINO S, M BAEZA, S QUIROZ & R RIVERA (1999) Ampliación distribucional de la carmelita, Percilia gillissi. Anales del Museo de Historia Natural de Valparaíso (Chile) 24: 119-120. [ Links ]
ZÚÑIGA L & P DOMÍNGUEZ (1977) Observaciones sobre el zooplancton de los lagos Chilenos. Anales del Museo de Historia Natural de Valparaíso (Chile) 10: 107-120. [ Links ]
ZÚÑIGA L, P DOMÍNGUEZ & MA CARVAJAL (1985) Distribución 572 espacio temporal del fitoplancton de la laguna El Plateado (Valparaíso, Chile). Anales del Museo de Historia Natural de Valparaíso (Chile) 16: 41-54. [ Links ]
Associate Editor: Horacio ZagareseReceived September 5, 2007; accepted November 17, 2007.