Crater Lake Apoyo revisited--population genetics of an emerging species flock.

The polytypic Nicaraguan Midas cichlids (Amphilophus cf. citrinellus) have been established as a model system for studying the mechanisms of speciation and patterns of diversification in allopatry and sympatry. The species assemblage in Crater Lake Apoyo has been accepted as a textbook example for sympatric speciation. Here, we present a first comprehensive data set of population genetic (mtDNA & AFLPs) proxies of species level differentiation for a representative set of individuals of all six endemic Amphilophus species occurring in Crater Lake Apoyo. AFLP genetic differentiation was partitioned into a neutral and non-neutral component based on outlier-loci detection approaches, and patterns of species divergence were explored with Bayesian clustering methods. Substantial levels of admixture between species were detected, indicating different levels of reproductive isolation between the six species. Analysis of neutral genetic variation revealed several A. zaliosus as being introgressed by an unknown contributor, hereby rendering the sympatrically evolving L. Apoyo flock polyphyletic. This is contrasted by the mtDNA analysis delivering a clear monophyly signal with Crater Lake Apoyo private haplotypes characterising all six described species, but also demonstrating different demographic histories as inferred from pairwise mismatch distributions.

Introduced predator elicits deficient brood defence behaviour in a crater lake fish.

Introduced species represent one of the most serious global threats to biodiversity. In this field-based study, we assessed behavioural responses of brood tending cichlid fish to an invasive predator of their offspring. This was achieved by comparing parental defence responses of the endangered arrow cichlid (Amphilophus zaliosus), a fish species endemic to the crater lake Apoyo in Nicaragua, towards the bigmouth sleeper (Gobiomorus dormitor), a formidable predator of cichlid fry, and all other potential fish predators of offspring. The bigmouth sleeper was recently introduced into Apoyo but naturally co-exists with cichlids in a few other Nicaraguan lakes. Arrow cichlid parents allowed bigmouth sleepers to advance much closer to their fry than other predators before initiating aggressive brood defence behaviours. Interestingly, parents of a very closely related species, A. sagittae, which has coevolved with bigmouth sleepers in crater lake Xiloá, reacted to approaching bigmouth sleepers at comparable distances as to other predators of cichlid fry. These results provide a novel demonstration of the specific mechanism (i.e. naive parental behaviour) by which invasive predators may negatively affect species that lack the adequate behavioural repertoire.

Not a simple case - A first comprehensive phylogenetic hypothesis for the Midas cichlid complex in Nicaragua (Teleostei: Cichlidae: Amphilophus).

Nicaraguan Midas cichlids from crater lakes have recently attracted attention as potential model systems for speciation research, but no attempt has been made to comprehensively reconstruct phylogenetic relationships of this highly diverse and recently evolved species complex. We present a first AFLP (2793 loci) and mtDNA based phylogenetic hypothesis including all described and several undescribed species from six crater lakes (Apoyeque, Apoyo, Asososca Leon, Masaya, Tiscapa and Xiloá), the two great Lakes Managua and Nicaragua and the San Juan River. Our analyses demonstrate that the relationships between the Midas cichlid members are complex, and that phylogenetic information from different markers and methods do not always yield congruent results. Nevertheless, monophyly support for crater lake assemblages from Lakes Apoyeque, Apoyo, A. Leon is high as compared to those from L. Xiloá indicating occurrence of sympatric speciation. Further, we demonstrate that a 'three species' concept for the Midas cichlid complex is inapplicable and consequently that an individualized and voucher based approach in speciation research of the Midas cichlid complex is necessary at least as long as there is no comprehensive revision of the species complex available.

Territorial aggression can be sensitive to the status of heterospecific intruders.

Territorial animals are known to be able to differentiate between intruding individuals posing a low or high threat and adjust their aggressive response accordingly. However, plastic territorial aggression based on recognising individuals with different attributes is typically assumed to be relevant only in the context of conspecific interactions. In this study, we investigated territorial aggression of neotropical cichlid fish in their natural habitat to assess whether responses to different types of individuals of another species can also be plastic. We show that arrow cichlids (Amphilophus zaliosus) adjusted their territorial aggression regarding the status of heterospecific intruders: breeding individuals of Amphilophus astorquii received a lower level of aggression than non-breeders. The same pattern was also found for the two different types of A. astorquii individuals intruding into conspecific territories. These results suggest that heterospecific individuals should not be ignored when considering selection pressures shaping plasticity of aggressive behaviour in territorial animals.

Comparison of gastropod mollusc (Apogastropoda: Hydrobiidae) habitats in two crater lakes in Nicaragua.

The aquatic gastropod mollusc, Pyrgophorus coronatus, may perform an important role in the transmission of an emergent ocular pathology among fishes in Lake Apoyo, Nicaragua. This disease emerged after an introduction of tilapia (Oreochromis niloticus) and the subsequent loss of Chara sp. beds in the lake. We compared the mollusc population densities in three habitats (sandy/muddy substrates, rocks, and Chara vegetation) at varying depths (1.5, 10, 20, and 30 m) in two volcanic crater lakes in Nicaragua: Lake Apoyo and Lake Xiloa. where lower numbers of affected fishes were found and tilapia has not been introduced. Duplicate samples at 1.5 m depth were taken in each habitat monthly for a year, and triplicate samples for bathymetric analysis of snail populations were performed during August, 2005. Samples of fixed surface area were filtered in a 0.4 cm size screen and live snails were counted from each sample. The preferred snail habitat in both lakes, Chara beds, was vastly reduced in Lake Apoyo via consumption by introduced Nile tilapia (Oreochromis niloticus). Structureless sandy substrates (mean +/- standard error 3.1+/-1.3 ind/m2) had lower population densities than other habitats in Lake Xiloa (rocks 590.9+/-185.3 ind/m2; vegetation 3 686.5+/-698.2 ind/m2; ANOVA 1, p<0.01 in both cases) but this difference was attenuated in Lake Apoyo (sand 384.4+/-111.1 ind/m2; rocks 1 480.4+/-384.8 ind/m2: 0.0 1

Comparison of gastropod mollusc (Apogastropoda: Hydrobiidae) habitats in two crater lakes in Nicaragua

The aquatic gastropod mollusc, Pyrgophorus coronatus, may perform an important role in the transmission of an emergent ocular pathology among fishes in Lake Apoyo, Nicaragua. This disease emerged after an introduction of tilapia (Oreochromis niloticus) and the subsequent loss of Chara sp. beds in the lake. We compared the mollusc population densities in three habitats (sandy/muddy substrates, rocks, and Chara vegetation) at varying depths (1.5, 10, 20, and 30 m) in two volcanic crater lakes in Nicaragua: Lake Apoyo and Lake Xiloa. where lower numbers of affected fishes were found and tilapia has not been introduced. Duplicate samples at 1.5 m depth were taken in each habitat monthly for a year, and triplicate samples for bathymetric analysis of snail populations were performed during August, 2005. Samples of fixed surface area were filtered in a 0.4 cm size screen and live snails were counted from each sample. The preferred snail habitat in both lakes, Chara beds, was vastly reduced in Lake Apoyo via consumption by introduced Nile tilapia (Oreochromis niloticus). Structureless sandy substrates (mean +/- standard error 3.1+/-1.3 ind/m2) had lower population densities than other habitats in Lake Xiloa (rocks 590.9+/-185.3 ind/m2; vegetation 3 686.5+/-698.2 ind/m2; ANOVA 1, p<0.01 in both cases) but this difference was attenuated in Lake Apoyo (sand 384.4+/-111.1 ind/m2; rocks 1 480.4+/-384.8 ind/m2: 0.0 1

El gasterópodo acuático, Pyrgophorus coronatus, podría jugar un papel importante en la transmisión de una patología ocular emergente entre los peces de la laguna de Apoyo, Nicaragua. Esta enfermedad surgió después de una introducción de tilapia (Oreochromis niloticus) y la subsecuente pérdida de lechos de Chara sp. en la laguna. Comparamos las densidades poblacionales del caracol en tres hábitats (substratos arenosos/lodosos, rocas y vegetación de Chara) en dos lagunas cratéricas volcánicas en Nicaragua: La laguna de Apoyo y la laguna de Xiloá, donde no se encuentraron grandes cantidades de peces afectados y donde no se han introducido tilapias. Mensualmente, por un año tomamos muestras duplicadas a 1.5 m profundidad mensualmente por un año, y durante agosto de 2005, muestreos triplicados de las poblaciones del molusco, para análisis batimétrico (1.5, 10, 20 y 30 m). Para determinar el número de individuos por unidad de área superficial, las muestras fueron filtradas en un colador de 0.4 cm de apertura de malla y contamos los moluscos vivos en cada muestra. El hábitat preferido de los caracoles en ambas lagunas fue la vegetación de Chara, que en la laguna de Apoyo fue vastamente reducida al ser consumida por las tilapias nilóticas introducidas (Oreochromis niloticus). Los substratos arenosos, menos estructurados que los otros sustratos estudiados, tuvieron densidades poblacionales más bajas (media ± error estándar 3.1±1.3 ind/m2) que en los otros hábitats en la laguna de Xiloá (vegetation 590.9±185.3 ind/m2; rocas 3 686.5±698.2 ind/m2; ANOVA I, p<0.01 en ambos casos) pero esta diferencia fue atenuada en la laguna de Apoyo (arena 384.4±111.1 ind/m2; rocas 1 480.4±384.8 ind/ m2; 0.01

Mercury in fish from two Nicaraguan lakes: a recommendation for increased monitoring of fish for international commerce.

We measured total mercury concentrations in water and fish of Lake Managua and Lake Apoyo. Water mercury concentrations were 10-fold higher in Lake Managua than in Lake Apoyo, although differences in mercury concentration in the most common native fish were not significant. One-fourth of the commercially fished tilapia in Lake Managua exceeded maximum recommended mercury levels for consumption among pregnant women and other at-risk groups, although bioavailability to fishes was lower than in previously studied sites in Brazil and Western Maryland. The lower bioavailiability may present important information for management options to reduce mercury exposure to fishes and humans. We recommend closer mercury monitoring among freshwater fish destined for international commerce.