Reproductive cycle of native viviparous fish species (Actinopterygii: Cyprinodontiformes: Goodeidae) in a subtropical Mexican lake

Reproductive tactics and strategies contribute to the persistence and maintenance of long-term populations in fish species. Members of the subfamily Goodeinae are a group of small-bodied freshwater fish with specialized reproduction (viviparity-matrotrophy). They are found in the highlands of central Mexico, most of them endemic. The aim of this study was to conduct a comprehensive investigation to evaluate the annual reproductive cycle of seven species of goodeines (splitfins). We carried out our study in the subtropical Lake Zacapu, Mexico, with bi-monthly sampling from May 2019 to March 2020. We obtain the fertility, size at first maturity (L50), sex ratio, and gonadosomatic index. Our result shows that populations of goodeines have high fertility compared to other populations of the same species in other aquatic systems and also to other species of goodein. We found that males mature at smaller sizes than females, the observed proportion of females was greater than males in all the goodeines. Lake Zacapu goodeines have two reproductive peaks, one in spring (April to June) and another in fall (September to November). These tactics (fertility rates, sex ratio, reproductive period) and strategies (viviparity-matrotrophy) favor reproductive success in this environmentally stable subtropical lake in the highlands of Mexico.(AU)

Las tácticas y estrategias reproductivas contribuyen a la persistencia y el mantenimiento de las poblaciones a largo plazo en especies de peces. Miembros de la subfamilia Goodeinae son un grupo de peces de agua dulce con reproducción especializada (viviparidad-matrotrofia). Se encuentran en el centro de México, la mayoría de ellos endémicos. El objetivo de este estudio fue realizar una investigación integral para evaluar el ciclo reproductivo anual de siete especies de goodeines o mexclapiques. Realizamos nuestro estudio en el lago subtropical de Zacapu, México, con muestreo bimestral de mayo de 2019 a marzo de 2020. Nuestros resultados muestran que las poblaciones de goodeines tienen una alta fertilidad en comparación con otras poblaciones de la misma especie en otros sistemas acuáticos y también con otras especies de goodeines. Encontramos que los machos maduran en tamaños más pequeños que las hembras, la proporción observada de hembras fue mayor que los machos en todas las goodeines. Los goodeines del lago Zacapu tienen dos picos reproductivos, uno en primavera (abril a junio) y otro en otoño (septiembre a noviembre). Estas tácticas (fertilidad, proporción de sexos, período reproductivo) y estrategias (viviparidad-matrotrofia) favorecen el éxito reproductivo en este lago subtropical ambientalmente estable en el altiplano de México.(AU)

Reproductive cycle of native viviparous fish species (Actinopterygii: Cyprinodontiformes: Goodeidae) in a subtropical Mexican lake

Reproductive tactics and strategies contribute to the persistence and maintenance of long-term populations in fish species. Members of the subfamily Goodeinae are a group of small-bodied freshwater fish with specialized reproduction (viviparity-matrotrophy). They are found in the highlands of central Mexico, most of them endemic. The aim of this study was to conduct a comprehensive investigation to evaluate the annual reproductive cycle of seven species of goodeines (splitfins). We carried out our study in the subtropical Lake Zacapu, Mexico, with bi-monthly sampling from May 2019 to March 2020. We obtain the fertility, size at first maturity (L50), sex ratio, and gonadosomatic index. Our result shows that populations of goodeines have high fertility compared to other populations of the same species in other aquatic systems and also to other species of goodein. We found that males mature at smaller sizes than females, the observed proportion of females was greater than males in all the goodeines. Lake Zacapu goodeines have two reproductive peaks, one in spring (April to June) and another in fall (September to November). These tactics (fertility rates, sex ratio, reproductive period) and strategies (viviparity-matrotrophy) favor reproductive success in this environmentally stable subtropical lake in the highlands of Mexico.(AU)

Las tácticas y estrategias reproductivas contribuyen a la persistencia y el mantenimiento de las poblaciones a largo plazo en especies de peces. Miembros de la subfamilia Goodeinae son un grupo de peces de agua dulce con reproducción especializada (viviparidad-matrotrofia). Se encuentran en el centro de México, la mayoría de ellos endémicos. El objetivo de este estudio fue realizar una investigación integral para evaluar el ciclo reproductivo anual de siete especies de goodeines o mexclapiques. Realizamos nuestro estudio en el lago subtropical de Zacapu, México, con muestreo bimestral de mayo de 2019 a marzo de 2020. Nuestros resultados muestran que las poblaciones de goodeines tienen una alta fertilidad en comparación con otras poblaciones de la misma especie en otros sistemas acuáticos y también con otras especies de goodeines. Encontramos que los machos maduran en tamaños más pequeños que las hembras, la proporción observada de hembras fue mayor que los machos en todas las goodeines. Los goodeines del lago Zacapu tienen dos picos reproductivos, uno en primavera (abril a junio) y otro en otoño (septiembre a noviembre). Estas tácticas (fertilidad, proporción de sexos, período reproductivo) y estrategias (viviparidad-matrotrofia) favorecen el éxito reproductivo en este lago subtropical ambientalmente estable en el altiplano de México.(AU)

Ectoparasitic copepod infestation on a wild population of Neotropical catfish Sciades herzbergii Bloch, 1794: Histological evidences of lesions on host.

We analyzed the infestation of the attached copepod species Lepeophtheirus sp on a wild population of Sciades herzbergii. The infestation and occurrence of attached copepods were related to body size, maturity and sex of host and the presence of lesions on fish skin were described. In 61 fish specimens (37 males and 24 females), total of 218 ectoparasitic copepods, including 204 mature and 14 immature stages were found. Copepods were attached to different regions of fish body without any regular pattern. The prevalence of infestation was 80.3% and intensity between 1 and 15 copepods/fish. No significant differences were found between sex or maturity and the presence of attached Lepeophtheirus sp. However, a contingency table including both sex and maturity status, and the total number of attached copepod per combined category showed a significant association. A positive correlation was found between body length of fish and intensity of infestation. Similarly, when comparing the infested fish according to group size, we found more copepods on larger individual. Attached copepods were associated with the presence of lesions visible to the naked eye. Histological analyses showed changes in cell architecture when sections of copepod-free tissues and attached copepods were compared.

Stable isotope evidence for trophic overlap of sympatric Mexican Lake Chapala silversides (Teleostei: Atherinopsidae: Chirostoma spp)

We explore the trophic role that a diverse sympatric group of fishes in the genus Chirostoma play in a large, shallow lake in central Mexico, Lake Chapala. We use δ13C and δ15N stable isotope - based food web analyses to explore how they relate to other components of the Lake Chapala ecosystem. We find five Chirostoma species in top trophic levels of the Chapala food web compared to other fishes, relying on a combination of zooplankton, fish and benthic resources as energy sources. Food web metric analyses showed generally overlapping trophic niches for members of Chirostoma, especially in terms of δ13C. However, C. jordani had lower mean δ15N isotopic values than C. promelas. As a group, "pescados blancos" (C. sphyraena and C. promelas) also had higher δ15N signatures than "charales" (C. consocium, C. jordani and C. labarcae) reflecting greater piscivory, but these differences were not strong for all food web metrics used. Trophic overlap among species of Chirostoma in Lake Chapala raises questions about the forces that might have led to a morphologically diverse but functionally similar and monophyletic group of species.

Exploramos el papel trófico de un diverso grupo de peces (género Chirostoma) que habita en simpatría en el Lago Chapala, México central. Utilizamos isótopos estables de δ13C y δ15N para explorar la relación que guardan éstos peces con otros componentes del ecosistema. Encontramos a Chirostoma en niveles altos de la red trófica de Chapala, dependiendo energéticamente de zooplankton, peces y recursos del bentos. Análisis de parámetros de la red trófica demostraron traslape de nicho trófico para cinco miembros de Chirostoma, especialmente con relación a δ13C. Sin embargo, C. jordani tuvo un menor valor promedio de δ15N que C. promelas. Al ser analizados como grupo, los "pescados blancos" (C. sphyraena, C. promelas) tuvieron niveles promedio de δ15N mayores que los "charales" (C. consocium, C. jordani, C. labarcae), indicando mayor ictiofagia, pero las diferencias no fueron significativas para todos los parámetros de red trófica utilizados. El traslape trófico entre las especies de Chirostoma en Chapala da origen a cuestionamientos acerca de las fuerzas que pueden haber intervenido en el surgimiento de un morfológicamente diverso pero funcionalmente similar grupo monofilético de peces.

Evaluación de la sustentabilidad en espacios rurales relacionados con el ecoturismo. estudio de caso en la selva lacandona, chiapas,México / Assesment of thee sutainability in rural spaces related to ecotouriism. case study in lacandon jungle, chiapas, Mexico

Esta investigación realiza una evaluación ex-post de la sustentabilidad en espacios rurales relacionados con el ecoturismo, centrada en un estudio comparativo de tres empresas enclavadas en la Reserva de la Biosfera Montes Azules (REBIMA) y una empresa más ubicada fuera de ella, todas en el municipio de Ocosingo, Chiapas, México. 75% de los entrevistados son hombres con edad promedio de 36 años, siete años de escolaridad y 1,9 hectáreas. La principal fuente de ingresos es el ecoturismo, que presenta una relación beneficio-costo venida a menos de 2,7 a 1,7, entre 2007 y 2011, respectivamente. La evidencia indica un desequilibrio en el acceso a subsidios, donde la pertenencia étnica desempeña un rol trascendental.

This research performs an ex-post evaluation of the sustainability in rural spaces related to ecotourism, focused on a comparative study of three companies located in the Montes Azules Biosphere Reserve (MABR) and another company located outside, all of them in the town of Ocosingo, Chiapas, México. 75% of respondents are male with an average age of 36 years, seven years of schooling and 1,9 hectares. The main source of income is ecotourism, which presents a cost-benefit relationship that has ran down from 2,7 to 1,7 between 2007 and 2011, respectively. The evidence indicates an imbalance in the access to subsidies, where the ethnical ownership plays a transcendental role.

Stable isotope evidence for trophic overlap of sympatric Mexican Lake Chapala silversides (Teleostei: Atherinopsidae: Chirostoma spp)

We explore the trophic role that a diverse sympatric group of fishes in the genus Chirostoma play in a large, shallow lake in central Mexico, Lake Chapala. We use δ13C and δ15N stable isotope - based food web analyses to explore how they relate to other components of the Lake Chapala ecosystem. We find five Chirostoma species in top trophic levels of the Chapala food web compared to other fishes, relying on a combination of zooplankton, fish and benthic resources as energy sources. Food web metric analyses showed generally overlapping trophic niches for members of Chirostoma, especially in terms of δ13C. However, C. jordani had lower mean δ15N isotopic values than C. promelas. As a group, "pescados blancos" (C. sphyraena and C. promelas) also had higher δ15N signatures than "charales" (C. consocium, C. jordani and C. labarcae) reflecting greater piscivory, but these differences were not strong for all food web metrics used. Trophic overlap among species of Chirostoma in Lake Chapala raises questions about the forces that might have led to a morphologically diverse but functionally similar and monophyletic group of species.(AU)

Exploramos el papel trófico de un diverso grupo de peces (género Chirostoma) que habita en simpatría en el Lago Chapala, México central. Utilizamos isótopos estables de δ13C y δ15N para explorar la relación que guardan éstos peces con otros componentes del ecosistema. Encontramos a Chirostoma en niveles altos de la red trófica de Chapala, dependiendo energéticamente de zooplankton, peces y recursos del bentos. Análisis de parámetros de la red trófica demostraron traslape de nicho trófico para cinco miembros de Chirostoma, especialmente con relación a δ13C. Sin embargo, C. jordani tuvo un menor valor promedio de δ15N que C. promelas. Al ser analizados como grupo, los "pescados blancos" (C. sphyraena, C. promelas) tuvieron niveles promedio de δ15N mayores que los "charales" (C. consocium, C. jordani, C. labarcae), indicando mayor ictiofagia, pero las diferencias no fueron significativas para todos los parámetros de red trófica utilizados. El traslape trófico entre las especies de Chirostoma en Chapala da origen a cuestionamientos acerca de las fuerzas que pueden haber intervenido en el surgimiento de un morfológicamente diverso pero funcionalmente similar grupo monofilético de peces.(AU)

Environmental Degradation in a Eutrophic Shallow Lake is not Simply Due to Abundance of Non-native Cyprinus carpio.

Non-native species are often major drivers of the deterioration of natural ecosystems. The common carp Cyprinus carpio are known to cause major changes in lentic systems, but may not be solely responsible for large scale changes in these ecosystems. We used data from extensive collection efforts to gain insight into the importance of carp as drivers of ecosystem change in Lake Patzcuaro, Mexico. We compared the structure (fish density, biomass, diversity, and evenness) of fish assemblages from six Lake Patzcuaro sites with different habitat characteristics. Intersite comparisons were carried out for both wet and dry seasons. We explored the relationships between non-carp species and carp; and studied multivariate interactions between fish abundance and habitat characteristics. From a biomass perspective, carp was dominant in only four of six sites. In terms of density, carp was not a dominant species in all sites. Further, carp density and biomass were not negatively related to native species density and biomass, even when carp density and biomass were positively correlated to water turbidity levels. Carp dominated fish assemblages in the shallowest sites with the highest water turbidity, plant detritus at the bottom, and floating macrophytes covering the lake surface. These results suggest that the effect of carp on fish assemblages may be highly dependent on habitat characteristics in Lake Patzcuaro. Watershed degradation, pollution, water level loss, and other sources of anthropogenic influence may be more important drivers of Lake Patzcuaro degradation than the abundance of carp.

Trophic ecology of the exotic Lerma livebearer Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) in the Lago de Pátzcuaro, Central Mexico.

Exotic fish species has caused several impacts on aquatic biodiversity. The Lago de Pátzcuaro has some well-studied exotic species, except the Lerma livebearer Poeciliopsis infans. This fish species was introduced into the Lago de Pátzcuaro before 1997 and the aspects of its biology are still unknown. In this study we assessed aspects of the trophic ecology of this exotic fish, P infans, using gut content and stable isotope analysis to understand its capacity to tolerate anthropogenic environmental degradation in the Lago de Pátzcuaro. We also determined its trophic guild position (TP) using the TrophLab Program and stable isotope. Niche breadth was calculated by standardized Levins' Index (Bi). Fish was captured with a seine during wet and dry seasons at six environmentally different sites and gut contents were obtained. We analyzed a total of 239 gut contents of P. infans. The contribution of each food item in the diet was quantified using frequency of occurrence and area percentage. The importance of each prey item was determined according to the index of relative importance (IRI), and the omnivory index (OI) was used to assess the feeding behavior. Fish were categorized by size and the diet was compared between fish sizes and sites. Dorsal muscle tissue and water hyacinth tissue was obtained for nitrogen isotope signature he fish can behave as a specialist (Bi = 0.39) or generalist (Bi = 0.68) and as a primary consumer (TROPH = 2.2; TP = 2.3) with a feeding strategy that was the same at dif- f erent sizes, seasons and sites. None of the evaluated sites showed good environmental quality. We argue that P infans can tolerate changes in water quality and feeding items availability, because it can exploit resources in multiple trophic webs. However, this species could be dependent on habitat complexity, especially in the aquatic vegetation cover.

Trophic ecology of the exotic Lerma livebearer Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) in the Lago de Pátzcuaro, Central Mexico / Ecología trófica del pez exótico Guatapote del Lerma Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) en el Lago de Pátzcuaro, Región Central de México

Exotic fish species has caused several impacts on aquatic biodiversity. The Lago de Pátzcuaro has some well-studied exotic species, except the Lerma livebearer Poeciliopsis infans. This fish species was introduced into the Lago de Pátzcuaro before 1997 and the aspects of its biology are still unknown. In this study we assessed aspects of the trophic ecology of this exotic fish, P. infans, using gut content and stable isotope analysis to understand its capacity to tolerate anthropogenic environmental degradation in the Lago de Pátzcuaro. We also determined its trophic guild position (TP) using the TrophLab Program and stable isotope. Niche breadth was calculated by standardized Levins’ Index (Bi). Fish was captured with a seine during wet and dry seasons at six environmentally different sites and gut contents were obtained. We analyzed a total of 239 gut contents of P. infans. The contribution of each food item in the diet was quantified using frequency of occurrence and area percentage. The importance of each prey item was determined according to the index of relative importance (IRI), and the omnivory index (OI) was used to assess the feeding behavior. Fish were categorized by size and the diet was compared between fish sizes and sites. Dorsal muscle tissue and water hyacinth tissue was obtained for nitrogen isotope signature (δ15N) analysis. Additionally, we measured water and habitat quality to evaluate environmental conditions at each site. We concluded that P. infans is an omnivore (OI=0.28) that consumes mainly detritus (44%), epiphytic diatoms (37%), and secondary on terrestrial insects (6%) and zooplankton (10%). The fish can behave as a specialist (Bi=0.39) or generalist (Bi=0.68) and as a primary consumer (TROPH=2.2; TP=2.3) with a feeding strategy that was the same at different sizes, seasons and sites. None of the evaluated sites showed good environmental quality. We argue that P. infans can tolerate changes in water quality and feeding items availability, because it can exploit resources in multiple trophic webs. However, this species could be dependent on habitat complexity, especially in the aquatic vegetation cover.

Las especies de peces exóticas han causado numerosos impactos sobre la biodiversidad acuática. El Lago de Pátzcuaro tiene especies exóticas bien estudiadas, excepto el Guatapote del Lerma, Poeciliopsis infans. Esta especie fue introducida en el Lago de Pátzcuaro antes de 1997 y los aspectos de su biología son aún desconocidos. Se estudiaron aspectos de la ecología trófica de este pez exótico, P. infans, mediante el uso de análisis de contenidos del tracto digestivo para entender su capacidad para tolerar la degradación ambiental antrópica en el Lago de Pátzcuaro. Lo que involucró la determinación del gremio y posición trófica (TP) con el programa TrophLab e isotopos estables. La amplitud de dieta fue calculada con el índice estandarizado de Levin (Bi). Un total de 239 tractos digestivos de P.infans fueron analizados. Los peces fueron capturados con una red tipo chinchorro durante la temporada de lluvias y estiaje en seis sitios ambientalmente distintos. La contribución de cada componente de la dieta fue cuantificada con la frecuencia de ocurrencia y el porcentaje de área. La importancia relativa de cada artículo alimentario fue determinada de acuerdo al índice de importancia relativa (IIR) y el índice de omnivoría (OI) fue usado para estimar la conducta en la forma de alimentarse. La dieta fue comparada entre tallas y entre sitios de estudio. El tejido de músculo dorsal y el tejido de lirio fueron obtenidos para determinar valores de δ15N. De manera adicional, la calidad del agua y del hábitat fue medida para evaluar la condición ambiental en cada sitio. P. infans es un omnívoro (OI=0.28) que consume principalmente detritus (44%), diatomeas epifíticas (37%) y de forma secundaria insectos terrestres (6%) y zooplancton (10%); puede comportarse como especialista (Bi=0.39) o generalista (Bi=0.68) y consumidor primario (TROPH=2.2; TP=2.3). La estrategia alimenticia fue similar entre tallas, temporadas y sitios. Ningún sitio mostró buena calidad ambiental. Los resultados del presente estudio permiten argumentar que P. infans puede tolerar cambios en la calidad del agua y en la disponibilidad de artículos alimenticios, porque puede usar recursos provenientes de múltiples redes tróficas. Sin embargo, esta especie podría ser dependiente de la complejidad del hábitat, especialmente de la cobertura de la vegetación acuática.

Spatial patterns of zooplanktivore Chirostoma species (Atherinopsidae) during water-level fluctuation in the shallow tropical Lake Chapala, Mexico: seasonal and interannual analysis

This study addresses the influence of water-level fluctuations on fish distribution at two temporal scales: seasonal (dry and rainy) and interannual (low and high volume conditions). The analysis of abundance relationships among three zooplanktivore Chirostoma species at fifteen sites in Lake Chapala, Mexico, revealed the significant influence of contrasting conditions (P=0.0002). Seasonally, segregation was more related to species dominance in the dry season and exclusively related to environmental characteristics in the rainy season. Interanually, biotic influence occurred in the shallowest and the deepest episodes of the lake. Environmental characteristics influenced species distribution when the lake reached 25 percent of its volume. Site, depth, temperature, and salinity were the leading factors influencing fish distribution. These results emphasize the necessity to implement different management strategies according to lake volume, particularly when a critical threshold is reached.

El presente estudio describe la influencia que tiene la fluctuación del nivel del agua en la distribución de los peces a dos diferentes escalas: estacional (estiaje y lluvias) e interanual (condiciones de bajo y alto volumen). El análisis de las relaciones de abundancia entre tres especies zooplanctófagas de Chirostoma en quince sitios del Lago de Chapala, México, reveló una influencia estadísticamente significativa en las condiciones contrastantes (P=0.0002). Estacionalmente, la segregación estuvo más relacionada a la dominancia de las especies en la época de estiaje y exclusivamente relacionada a las características ambientales en la época de lluvias. Interanualmente, la influencia biótica se presenta en los episodios más someros y más profundos del lago. Las características ambientales influenciaron la distribución de las especies cuando el lago alcanzó el 25 por ciento de su volumen. El sitio geográfico, profundidad, temperatura y salinidad fueron los principales factores que influenciaron la distribución de los peces. Estos resultados enfatizan la necesidad de implementar diferentes estrategias de manejo de acuerdo al volumen del lago, particularmente cuando se alcanza un umbral crítico.

Trophic interactions among sympatric zooplanktivorous fish species in volume change conditions in a large, shallow, tropical lake

Significant reductions in the water volume of shallow lakes impose a restriction on species segregation promoting more interactions in the trophic relationships. The diets of three closely related zooplanktivorous silversides belonging to the Atherinopsidae species flock of lake Chapala, Mexico, were analyzed at two sites (Chirostoma jordani, C. labarcae, andC. consocium). Diets were described in critical shallow (August 2000) and volume recovery conditions (August 2005). Diets included mainly cladocerans (Bosmina, Ceriodaphnia, and Daphnia) and copepods (Cyclops). A significant difference in diets was detected when comparing years (MRPP analysis, A = 0.22, p < 0.0001) and sites at different years (MRPP analysis, A = 0.17, p = 0.004). According to niche breadth mean values, species were classified as specialized and intermediate feeders. In shallow conditions, the small range of niche breadth (1.72 to 3.64) and high diet overlap values (D = 0.64, L = 8.62) indicated a high potential for interspecific exploitative interaction. When the lake volume recovered, an increase in the niche breadth range (1.04 to 4.96) and low niche overlap values (D = 0.53, L = 2.32) indicated a reduction of the species interaction. The Mann- Whitney U-test supported this pattern by showing a significant difference between years for niche overlap (p = 0.006). The increased interaction during the low volume suggests alternative segregation in life-history variations and other niche dimensions such as spatial or temporal distribution.(AU)

Spatial patterns of zooplanktivore Chirostoma species (Atherinopsidae) during water-level fluctuation in the shallow tropical Lake Chapala, Mexico: seasonal and interannual analysis

This study addresses the influence of water-level fluctuations on fish distribution at two temporal scales: seasonal (dry and rainy) and interannual (low and high volume conditions). The analysis of abundance relationships among three zooplanktivore Chirostoma species at fifteen sites in Lake Chapala, Mexico, revealed the significant influence of contrasting conditions (P=0.0002). Seasonally, segregation was more related to species dominance in the dry season and exclusively related to environmental characteristics in the rainy season. Interanually, biotic influence occurred in the shallowest and the deepest episodes of the lake. Environmental characteristics influenced species distribution when the lake reached 25 percent of its volume. Site, depth, temperature, and salinity were the leading factors influencing fish distribution. These results emphasize the necessity to implement different management strategies according to lake volume, particularly when a critical threshold is reached.(AU)

El presente estudio describe la influencia que tiene la fluctuación del nivel del agua en la distribución de los peces a dos diferentes escalas: estacional (estiaje y lluvias) e interanual (condiciones de bajo y alto volumen). El análisis de las relaciones de abundancia entre tres especies zooplanctófagas de Chirostoma en quince sitios del Lago de Chapala, México, reveló una influencia estadísticamente significativa en las condiciones contrastantes (P=0.0002). Estacionalmente, la segregación estuvo más relacionada a la dominancia de las especies en la época de estiaje y exclusivamente relacionada a las características ambientales en la época de lluvias. Interanualmente, la influencia biótica se presenta en los episodios más someros y más profundos del lago. Las características ambientales influenciaron la distribución de las especies cuando el lago alcanzó el 25 por ciento de su volumen. El sitio geográfico, profundidad, temperatura y salinidad fueron los principales factores que influenciaron la distribución de los peces. Estos resultados enfatizan la necesidad de implementar diferentes estrategias de manejo de acuerdo al volumen del lago, particularmente cuando se alcanza un umbral crítico.(AU)