Water availability and seasonality shape elemental stoichiometry across space and time.

The interaction of climate change and increasing anthropogenic water withdrawals is anticipated to alter surface water availability and the transport of carbon (C), nitrogen (N), and phosphorus (P) in river networks. But how changes to river flow will alter the balance, or stoichiometry, of these fluxes is unknown. The Lower Flint River Basin (LFRB) is part of an interstate watershed relied upon by several million people for diverse ecosystem services, including seasonal crop irrigation, municipal drinking water access, and public recreation. Recently, increased water demand compounded with intensified droughts have caused historically perennial streams in the LFRB to cease flowing, increasing ecosystem vulnerability. Our objectives were to quantify how riverine dissolved C:N:P varies spatially and seasonally and determine how monthly stoichiometric fluxes varied with overall water availability in a major tributary of LFRB. We used a long-term record (21-29 years) of solute water chemistry (dissolved organic carbon, nitrate/nitrite, ammonia, and soluble reactive phosphorus) paired with long-term stream discharge data across six sites within a single LFRB watershed. We found spatial and seasonal differences in soluble nutrient concentrations and stoichiometry attributable to groundwater connections, the presence of a major floodplain wetland, and flow conditions. Further, we showed that water availability, as indicated by the Palmer Drought Severity Index (PDSI), strongly predicted stoichiometry with generally lower C:N and C:P and higher N:P fluxes during periods of low water availability (PDSI < -4). These patterns suggest there may be long-term and significant changes to stream ecosystem function as water availability is being dramatically altered by human demand with consequential impacts on solute transport, in-stream processing, and stoichiometric ratios.

Preferential retention of algal carbon in benthic invertebrates: Stable isotope and fatty acid evidence from an outdoor flume experiment.

According to the River Continuum Concept, headwater streams are richer in allochthonous (e.g. terrestrial leaves) than autochthonous (e.g. algae) sources of organic matter for consumers. However, compared to algae, leaf litter is of lower food quality, particularly ω-3 polyunsaturated fatty acids (n-3 PUFA), and would constrain the somatic growth, maintenance, and reproduction of stream invertebrates. It may be thus assumed that shredders, such as Gammarus, receive lower quality diets than grazers, e.g. Ecdyonurus, that typically feed on algae.The objective of this study was to assess the provision of dietary PUFA from leaf litter and algae to the shredder Gammarus and the grazer Ecdyonurus. Three different diets (algae, terrestrial leaves, and an algae-leaf litter mix) were supplied to these macroinvertebrates in a flume experiment for 2 weeks. To differentiate how diet sources were retained in these consumers, algae were isotopically labelled with 13C.Both consumers became enriched with 13C in all treatments, demonstrating that both assimilated algae. For Gammarus, n-3 PUFA increased, whereas n-6 PUFA stayed constant. By contrast, the n-3 PUFA content of Ecdyonurus decreased as a consequence of declining algal supply.Results from compound-specific stable isotope analysis provided evidence that the long-chain n-3 PUFA eicosapentaenoic acid (EPA) in both consumers was more enriched in 13C than the short-chain n-3 PUFA α-linolenic acid, suggesting that EPA was taken up directly from algae and not from heterotrophic biofilms on leaf litter. Both consumers depended on algae as their carbon and EPA source and retained their EPA from high-quality algae.

A patchy continuum? Stream processes show varied responses to patch- and continuum-based analyses.

Many conceptual syntheses in ecology and evolution are undergirded by either a patch- or continuum-based model. Examples include gradualism and punctuated equilibrium in evolution, and edge effects and the theory of island biogeography in ecology. In this study, we sought to determine how patch- or continuum-based analyses could explain variation in concentrations of stream macronutrients and system metabolism, represented by measures of productivity and respiration rates, at the watershed scale across the Kanawha River Basin, USA. Using Strahler stream order (SSO; continuum) and functional process zone (FPZ; patch) as factors, we produced statistical models for each variable and compared model performance using likelihood ratio tests. Only one nutrient (i.e., PO43- ) responded better to patch-based analysis. Both models were significantly better than a null model for ecosystem respiration; however, neither outperformed the other. Importantly, in most cases, a combination model, including both SSO and FPZ, best described observed variation in the system. Our findings suggest that several patch- and continuum-based processes may simultaneously influence the concentration of macronutrients and system metabolism. Nutrient spiral- ing along a continuum and the patch mosaic of land cover may both alter macronutrients, for example. Similarly, increases in temperature and discharge associated with increasing SSO, as well as the differences in light availability and channel morphology associated with different FPZs, may influence system metabolism. For these reasons, we recommend a combination of patch- and continuum-based analyses when modeling, analyzing, and interpreting patterns in stream ecosystem parameters.

More than a corridor: use of a main stem stream as supplemental foraging habitat by a brook trout metapopulation.

Coldwater fishes in streams, such as brook trout (Salvelinus fontinalis), typically are headwater specialists that occasionally expand distributions downstream to larger water bodies. It is unclear, however, whether larger streams function simply as dispersal corridors connecting headwater subpopulations, or as critical foraging habitat needed to sustain large mobile brook trout. Stable isotopes (δ(13)C and δ(15)N) and a hierarchical Bayesian mixing model analysis was used to identify brook trout that foraged in main stem versus headwater streams of the Shavers Fork watershed, West Virginia. Headwater subpopulations were composed of headwater and to a lesser extent main stem foraging individuals. However, there was a strong relationship between brook trout size and main stem prey contributions. The average brook trout foraging on headwater prey were limited to 126 mm standard length. This size was identified by mixing models as a point where productivity support switched from headwater to main stem dependency. These results, similar to other studies conducted in this watershed, support the hypothesis that productive main stem habitat maintain large brook trout and potentially facilitates dispersal among headwater subpopulations. Consequently, loss of supplementary main stem foraging habitats may explain loss of large, mobile fish and subsequent isolation of headwater subpopulations in other central Appalachian watersheds.

Gorgona, Baudó y Darién (Chocó Biogeográfico, Colombia): ecorregiones modelo para los estudios ecológicos de comunidades de quebradas costeras / Gorgona, Baudó and Darién (Biogeographic Chocó, Colombia): model ecoregions for coastal stream ecology studies

This paper summarizes the available information on the geology, hydrology, and biota of coastal streams located in the Gorgona National Natural Park (GNNP), and the Baudó and Darién ecoregions within the Biogeographic Chocó (Colombia). Despite of the scant number of studies, we hypothesized that these streams are consistent with the typology observed in volcanic oceanic islands in the Caribbean and the Pacific that do not conform to the tenets of the River Continuum Concept. Coastal streams in the Biogeographic Chocó are short (10(0)-10¹km) and steep and waterfalls and cascades are frequent due to tectonic origin. Step-and-pool sequences are common in montane reaches, while cobble-bed glides and riffles dominate in coastal plain reaches. Flow regimes are flashy in Pacific drainages (annual rainfall: <7000mm), but seasonally dry in the Caribbean drainages (<3500mm). Therefore, floods and droughts are important drivers of communities and ecosystem processes. Canopies are closed in low-order streams discharging directly to the ocean, thus contributing large amounts of litter. Diadromous fish and shrimp dominate consumer assemblages and various species are shared with streams in the Central America and the Antilles. These species play ecological roles probably equivalent to those found in other regions. These coastal streams are unique in terms of the number of primary freshwater fishes found (some endemics), and the functionally absence of diadromous gastropods. Rev. Biol. Trop. 62 (Suppl. 1): 43-64. Epub 2014 February 01.

A pesar de la escasez de estudios sobre la geología, hidrología y biota de las quebradas costeras ubicadas en el Chocó Biogeográfico, se hipotetiza que éstas se ajustan a la tipología observada en las islas oceánicas volcánicas del Caribe y el Pacífico, las cuales no cumplen con las predicciones del Concepto del Río Continuo. Estas quebradas son cortas (<10¹km) y muy pendientes debido a su origen tectónico, y presentan lechos dominados por bloques y cantos rodados. El régimen de caudal es torrencial en la vertiente Pacífico, pero estacionalmente seco en la vertiente Caribe. Los doseles cerrados aportan grandes cantidades de hojarasca a las quebradas que desembocan al mar con bajo orden. Las especies diádromas, con afinidades centroamericanas y antillanas, dominan los ensamblajes y posiblemente cumplen papeles ecológicos equivalentes a los establecidos en otras regiones. Particularmente, las quebradas costeras de esta provincia presentan varias especies de peces dulceacuícolas primarios, algunas endémicas, y carecen de gasterópodos. Los procesos geomorfológicos e hidrológicos de las cabeceras influyen sobre la dinámica ecosistémica de estas quebradas. Se proponen cinco hipótesis sobre la estructura y composición comunitaria. Se advierte que el PNNG es un modelo útil para la ecología lótica costera, pero que tiene particularidades.

Longitudinal use of feeding resources and distribution of fish trophic guilds in a coastal Atlantic stream, southern Brazil

This study described the use of food resources and the distribution of fish trophic guilds along the longitudinal gradient of a coastal Atlantic stream, southern Brazil. Allochthonous resources (terrestrial insects) predominated in the headwaters, whereas autochthonous food items (algae, fish) and detritus predominated in the mouth section. Detritivorous, aquaticinvertivorous, and terrestrial- and aquatic-insectivorous species occurred throughout the gradient, while omnivorous and piscivorous in the headwaters and middle stretches, respectively, and herbivorous in the middle and mouth. Detritivores and aquatic-insectivores were significantly more specialized than the other guilds, however, there was no evidence of a longitudinal increase in trophic specialization. Density and biomass of aquatic-invertivores and aquatic-insectivores decreased, whereas that of detritivores increased longitudinally. The distribution of trophic guilds was significantly associated with the stream section, where aquatic and terrestrial insectivorous were more frequent in rocky and flowed stretches from the headwater and detritivores in deeper environments with finer particles of substrate from the mouth. This suggests that fish assemblages in coastal streams with a steep longitudinal gradient may follow patterns in the use of food resources according with the food availability along the river, as predicted by the River Continuum Concept.

Este estudo descreve o uso dos recursos alimentares e a distribuição das guildas tróficas de peixes ao longo do gradiente longitudinal de um riacho da encosta Atlântica, Sul do Brasil. Recursos alóctones (insetos terrestres) foram consumidos com maior frequência na cabeceira, entretanto, recursos de origem autóctone (algas, peixes) e detritos foram mais frequentemente registrados na foz. Espécies detritívoras, invertívoras aquáticas, insetívoras terrestres e aquáticas ocorreram ao longo de todo gradiente, enquanto que as onívoras e piscívoras na cabeceira e meio, respectivamente, e herbívoras no meio e foz. Detritívoras e insetívoras aquáticas foram significativamente mais especializadas em relação às demais guildas; entretanto, não houve nenhuma evidência de aumento longitudinal da especialização trófica. A densidade e a biomassa de invertívoras e insetívoras aquáticas diminuíram, enquanto que das detritívoras aumentaram no sentido cabeceira-foz. A distribuição das guildas tróficas foi significativamente associada com a secção do riacho, onde insetívoras aquáticas e terrestres foram mais frequentes em trechos rochosos e de elevado fluxo da cabeceira, enquanto que detritívoras em ambientes mais profundos com partículas finas de substrato da foz. Tais resultados sugerem que assembleias de peixes de riachos tropicais de encosta, os quais apresentam acentuado gradiente longitudinal, podem seguir padrões de uso dos recursos alimentares de acordo com a disponibilidade alimentar ao longo do rio, como previstos no Conceito de Continuidade de Rios.

Longitudinal variation in the composition of the benthic macroinvertebrate fauna of a typical North coast Jamaican river

Benthic macroinvertebrate fauna plays a major role in river ecosystems, especially those of tropical islands. Since there is no information on the distribution of benthic invertebrates along a Jamaican river, we report here on the composition of the benthic fauna of the Buff Bay river, on the Northern coast of Jamaica. A total of 14 samples were collected from five sites, using kick nets and a Surber sampler, between May 1997 and October 1998. We also examined the applicability of the rhithron/potamon model, and some of the premises of the River Continuum Concept (RCC) in relation to the distribution of invertebrate taxa. The results showed a total of 38 taxa of identified invertebrates. A group of dominant taxa, composed mainly of immature stages of insects, occurred at all sites. Two notable characteristics of the river were the absence of a true potamonic fauna and the low representation of the shredder functional feeding group in the community We conclude that, while there was minor variation in the composition of the benthic macroinvertebrate fauna among the sites, this was a response to local conditions within the river system. The characteristics of the community did not conform to either of the models.

La fauna bentónica de macroinvertebrados juega un papel importante en los ecosistemas fluviales, especialmente los de las islas tropicales. En vista de que hay poca información disponible para los ríos de Jamaica, presentamos la composición de la fauna bentónica de la bahía riverina Buff, en la costa norte de Jamaica. Para ello, recolectamos un total de 14 muestras en cinco sitios, mediante el uso de redes de golpe y trampa Surber, entre mayo 1997 y octubre 1998. También se examinó la aplicabilidad del modelo de subdivisión de ríos ritrón/potamón y algunas de las premisas del concepto de Río como un Continuo, en relación con la distribución de los táxones de invertebrados. Los resultados mostraron un total de 38 táxones de invertebrados identificados, hay un grupo de taxones dominantes, compuesto principalmente por los estadios inmaduros de insectos, que se presentaron en todos los sitios. Dos características notables del río fueron la ausencia de una fauna de potamón verdadera y la escasa representación del grupo funcional alimenticio triturador en la comunidad. Llegamos a la conclusión de que, si bien hay una variación de menor importancia en la composición de la fauna de macroinvertebrados bentónicos entre los sitios, esto fue una respuesta a las condiciones locales dentro del sistema fluvial y las características de la comunidad que no se ajusta a ninguno de los modelos. Este estudio representa la primera publicación de una descripción longitudinal de la fauna bentónica de macroinvertebrados de un sistema fluvial de Jamaica.

Influence of the riparian zone phytophysiognomies on the longitudinal distribution of fishes: evidence from a Brazilian savanna stream

The structure and composition of a fish community in a tributary stream of the Aquidauana River, located in the plateau region of the Paraguay River basin, central west Brazil are described, with special attention to the differential effects of the phytophysiognomies of the Cerrado (a tropical savanna), which are predominant in riparian zones all along their longitudinal gradient. A total of 1,166 individuals belonging to 44 species were captured. Similarity analyses of composition and abundance showed two groups: upstream reaches (R1 and R2) and downstream reaches (R3, R4 and R5), which presented 85.8 percent of the average dissimilarity. Although physical habitat and physicochemical descriptors varied along the longitudinal gradient of the Correntes stream, shoreline vegetation was the most important environmental feature predicting fish structure and composition. The Mantel test revealed a correlation between shoreline vegetation structure and fish composition and quantitative structure of the fish community (R > 0.65; p < 0.04). This relationship is driven by the prevalence of species occupying microhabitats associated with shoreline vegetation in contact with water in upstream reaches. Structural differences in shoreline vegetation along the longitudinal gradient correspond to the phytophysiognomic dichotomy observed in the Correntes stream, where riparian vegetation is made up of wet grassland upstream and of gallery forest downstream; this reinforces the importance of the phytophysiognomic heterogeneity of the Cerrado in maintaining ichthyofauna diversity.

São descritas a estrutura e composição da comunidade de peixes em um riacho afluente do rio Aquidauana, localizado na região planáltica da bacia do rio Paraguai, centro-oeste do Brasil, com atenção especial aos efeitos diferenciais das fitofisionomias de Cerrado predominantes nas zonas ripárias ao longo do seu gradiente longitudinal. Foram capturados 1.166 indivíduos pertencentes a 44 espécies. As análises de similaridade com a composição e abundância evidenciaram dois grupos: os trechos a montante (R1 e R2) e trechos a jusante (R3, R4 e R5), os quais apresentaram 85,8 por cento de dissimilaridade média. Embora os descritores fisico-químicos e do habitat físico tenham variado ao longo do gradiente longitudinal do córrego Correntes, a vegetação marginal foi o componente ambiental mais importante predizendo a estrutura e composição da ictiofauna. O teste de Mantel revelou correlação entre a estrutura da vegetação marginal com a composição e estrutura quantitativa da comunidade de peixes (R > 0.65; p < 0.04). Essa relação é representada pela prevalência de espécies que exploram micro-habitats associados à vegetação marginal em contato com a água nos trechos a montante. As diferenças na estrutura da vegetação marginal ao longo do gradiente longitudinal correspondem à dicotomia fitofisionômica observada no córrego Correntes, cuja vegetação marginal é constituída de campos úmidos à montante e por fisionomias florestais nos trechos à jusante, reforçando a importância da heterogeneidade fitofisionomica do Cerrado brasileiro para a manutenção da diversidade ictiofaunística.

Fish assemblage structure in a first order stream, southeastern Brazil: longitudinal distribution, seasonality, and microhabitat diversity

This study was carried out on a first order stream in the Morro do Diabo State Park, southeastern Brazil. Three equidistant stream stretches - upper, middle, and lower - were chosen for underwater observations and fish collections that were performed in the dry (June and September 2000) and wet seasons (December 2000 and March 2001). Of the 18 fish species documented, 11 were considered resident year round. Phalloceros caudimaculatus, Hypostomus nigromaculatus, Hisonotus francirochai, Trichomycterus diabolus, and Hypostomus ancistroides represented 90 percent of the collected specimens. The highest richness and abundance occurred during the wet season. In general, the longitudinal addition of species follows the addition of microhabitats. Sexually mature individuals were present throughout the year for the majority of the species, which suggests broad reproductive activity. However, given that most juveniles specimens were collected during the wet season, it is hypothesized that reproduction and survival rates must be higher during this season due to higher availability of critical resources, most probably food and shelter.

Este estudo foi conduzido em três trechos - superior, médio e inferior - de um riacho de primeira ordem no Parque Estadual Morro do Diabo, sudeste do Brasil. Observações subaquáticas a partir de mergulho livre e coletas foram realizadas na estação seca (junho e setembro de 2000) e chuvosa (dezembro de 2000 e março de 2001). Das 16 espécies documentadas, 11 foram residentes ao longo das observações. Phalloceros caudimaculatus, Hypostomus nigromaculatus, Hisonotus francirochai, Trichomycterus diabolus e Hypostomus ancistroides representaram 90 por cento do total de indivíduos amostrados. A maior riqueza e abundância ocorreram durante a estação chuvosa. Em geral, a adição longitudinal de espécies seguiu a adição de micro-hábitats. Indivíduos sexualmente maduros foram encontrados ao longo do ano para a maioria das espécies, o que sugere prolongada atividade reprodutiva. Contudo, visto que a maioria dos jovens foi coletada na estação chuvosa, é provável que neste período ocorra um incremento de atividade reprodutiva, em razão de maior disponibilidade de recursos críticos, principalmente abrigos e alimentos.

Isotopic analysis of three food web theories in constricted and floodplain regions of a large river.

Analyses of stable isotope (δ13C and δ15N) and C:N ratios of food webs within a floodplain and a constricted-channel region of the Ohio River during October 1993 and July 1994 indicate that the increasingly influential flood pulse concept (FPC) does not, for either location, adequately address food web structure for this very large river. Furthermore, results of this study suggest that the riverine productivity model (RPM) is more appropriate than the widely known river continuum concept (RCC) for the constricted region of this river. These␣conclusions are based on stable isotope analyses of potential sources of organic matter (riparian C3 trees, riparian C4 grasses and agricultural crops, submerged macrophytes, benthic filamentous algae, benthic particulate organic matter, and transported organic matter containing detritus and phytoplankton) and various functional feeding groups of invertebrate and fish consumers. The FPC, which stresses the key contribution of organic matter, particularly terrestrial organic matter, originating from the floodplain to riverine food webs, was judged inappropriate for the floodplain region of the Ohio River for hydrodynamic and biotic reasons. The rising limb and peak period of discharge typically occur in November through March when temperatures are low (generally much less than 10°C) and greater than bank-full conditions are relatively unpredictable and short-lived. The major food potentially available to riverine organisms migrating into the floodplain would be decaying vegetation because autotrophic production is temperature and light limited and terrestrial insect production is minimal at that time. It is clear from our data that terrestrial C4 plants contribute little, if anything, to the consumer food web (based on δ13C values), and δ15N values for C3 plants, coarse benthic organic matter, and fine benthic organic matter were too depleted (∼7-12‰ lower than most invertebrate consumer values) for this organic matter to be supporting the food web. The RPM, which emphasizes the primary role of autotrophic production in large rivers, is the most viable of the remaining two ecosystem models for the constricted-channel region of the Ohio based on stable isotope linkage between sources and consumers of organic matter in the food web. The most important form of food web organic matter is apparently transported (suspended) fine (FTOM) and ultra-fine particulate organic matter. We propose that phytoplankton and detritus of an autochthonous origin in the seston would represent a more usable energy source for benthic (bivalve molluscs, hydropsychid caddisflies) and planktonic (microcrustaceans) suspension feeders than the more refractory allochthonous materials derived from upstream processing of terrestrial organic matter. Benthic grazers depend heavily on nonfilamentous benthic algae (based on gut analysis from a separate study), but filamentous benthic algae have no apparent connection to invertebrate consumers (based on δ13C values). Amphipod and crayfish show a strong relationship to aquatic macrophytes (possibly through detrital organic matter rather than living plant tissue). These observations contrast with the prediction of the RCC that food webs in large rivers are based principally on refractory FTOM and dissolved organic matter from upstream inefficiencies in organic-matter processing and the bacteria growing upon these suspended or dissolved detrital compounds. The conclusions drawn here for the Ohio River cannot yet be extended to other floodplain and constricted-channel rivers in temperate and tropical latitudes until more comparable data are available on relatively pristine and moderately regulated rivers.

Multiple-site and-year analyses of stream insect emergence: a test of ecological theory.

The use of existing data sets to test applicability of existing ecological theory is an uncommon but potentially cost-effective approach for exploitation of previously accumulated knowledge. Studies on the emergence of insects from small streams have been a major research topic in aquatic ecology, particularly in Austria and Germany; the availability of emergence data from these two countries, covering over 1 million identified specimens, from 18 sites, and for 32 collection years is an example of such exploitable information. Concurrent estimates of annual emergence biomass and annual benthic secondary production for 18 aquatic insect populations showed a statistically significant relationship, contradicting the premise that emergence data do not provide any quantitative measure for a given stream area. Therefore, the emergence data were examined to test various predictions from ecological theory. Observed richness of emerging species of three orders of lotic insects - the Ephemeroptera, Plecoptera, and Trichoptera (EPT) - over 15 years at one site did not agree with predictions based on either flow predictability or change in flow and the "habitat templet concept". Trends in observed richness of emerging EPT species over 1 year at 18 sites agreed weakly with predictions using either pH values or the annual temperature amplitude and the "intermediate disturbance hypothesis", or using either annual temperature amplitude or total biomass of EPT emergence and the "disturbance-productivity-diversity model". A prediction of the "river continuum concept" that abundance of the shredder functional-feeding group should decrease and that of grazers should increase along a dense- to open-canopy gradient was not consistently supported by the emergence biomass data. For shredders and grazers of all insects (12 sites) and EPT (18 sites), this trend was apparent (but not significant) only if sites with intermediate canopy density were omitted. We identified three critical elements in our study that generally interfere with such tests of these theoretical constructs: (i) species richness is a poor measure of resource limitation and/or utilization theories; (ii) restrictions of the taxonomic operational window (in our example usually to EPT) causes problems in extrapolation to a larger system (in our example to all insects); and (iii) historical constraints may affect the local result of tests of resource limitation and/or utilization theories simply because species that potentially interact are lacking in the region under examination. Problems notwithstanding, the use of existing data sets to test applicability of currently held ecological theories is a cost-effective and amenable approach for use in a variety of research topics in stream and general ecology. In this context, future tests should focus on: (i) measures that are more robust than just species richness, e.g., on measures commonly used to assign species to strategies such as r, K, or A; (ii) a variety of taxonomic groups; and (iii) gradients in historical constraints on current regional species composition.