"Unraveling the Gut Microbiome of the Genus Herichthys (Pisces: Cichlidae): What Can We Learn from Museum Specimens?"

The use of museum preserved specimens to know microbiome in extinct and threatened species has been explored recently. The fishes of the genus Herichthys are distributed mainly in the Pánuco-Tamesí system in Northeastern Mexico, one of the most polluted basins in the country leading to near half of the species be considering as threatened. In this paper we used the hypervariable V4 region of the 16S rRNA gene from the 11 species of the genus Herichthys obtained from museum collections to evaluate the potential use of fixed preserved vouchers in the knowledge of gut microbiota diversity and the potential role of sympatric and allopatric speciation of the hosts in the gut microbiome evolution. The 100% of the samples were successfully amplified where the number of amplicons ranged from 4500 from a formaldehyde fixed specimen up to 55,000 in ethanol preserved specimens. Differences in gut microbiota were found between sympatric species and among the comparison of some trophic guilds. A non-random association between the gut host and their microbiome was found allow to suggest a potential phylosymbiosis relationship. In conclusion, the most abundant phyla recovered from the gut microbiota in this study were similar to those previously reported in other cichlids supporting the idea that a gut microbial core is conserved in this group of fishes despite millions of years of evolution and leading to support the potential use of museum specimens in microbiome studies.

The role of head shape and trophic variation in the diversification of the genus Herichthys in sympatry and allopatry.

In the present study we evaluated the putative cases of sympatric speciation in the genus Herichthys by studying the variation in head shape using principal component analysis, phylomorphospace and reconstructions of the ancestral states of feeding preferences. Herichthys includes both allopatric and sympatric sister species, as well as sympatric unrelated species and thus offers great potential for evolutionary studies of putatively sympatric speciation. Herichthys is the northernmost group of cichlids in America and one of the most ecologically disparate genera within Middle American cichlids. Fifteen anatomical points were recorded on the heads of 293 specimens of the 11 species recognized within the genus. The results show that in spite of having wide variation in consumed diets, most species of Herichthys are close in morphospace. However, morphological variation was great among the two pairs of sympatric sister species in agreement with the suggested sympatric model of speciation.

Molecular clocks, biogeography and species diversity in Herichthys with evaluation of the role of Punta del Morro as a vicariant brake along the Mexican Transition Zone in the context of local and global time frame of cichlid diversification.

Using molecular dated phylogenies and biogeographic reconstructions, the species diversity, biogeography and time frame of evolution of the genus Herichthys were evaluated. In particular, we test the role of Punta del Morro (PdM) as a vicariant brake along the Mexican Transition Zone in the context of local and global time frame of cichlid diversification using several sets of calibrations. Species diversity in Herichthys is complex and the here employed dating methods suggest young age and rapid divergence for many species while species delimitation methods did not resolve these young species including both sympatric species pairs. Based on our molecular clock dating analyses, Herichthys has colonized its present distribution area significantly prior to the suggested vicariance by PdM (10-17.1 Ma vs. 5 to 7.5 Ma). The PdM constraint is in conflict with all other paleogeographic and fossil constraints including novel ones introduced in this study that are, however, congruent among each other. Our study demonstrates that any cichlid datings significantly older or younger than the bounds presented by our analyses and discussion have to be taken as highly questionable from the point of view of Middle American paleogeography and cichlid biogeography unless we allow the option that cichlid biogeography is completely independent from ecological and geological constraints.

Feeding ecomorphologies in the fish genus Herichthys (Perciformes: Cichlidae) based on stomach content and lower pharyngeal jaw shape

Speciation is a multifactorial process with factors acting at different scales of space and time. Trophic niche segregation has promoted the diversification of cichlids fishes in lentic (lacustrine) environments, whether this is also the case in lotic (riverine) systems remains unknown. Herichthys is the genus of cichlids with the most boreal distribution in the Americas comprising 12 currently recognized species, most micro-endemic and only two with a wide distribution. In the present work, we analyzed the stomach content and lower pharyngeal jaw morphologies of the species of the genus to evaluate the possible role of feeding ecology in the diversification of the group. Trophic strategies varied widely, including omnivores, piscivores, invertivores, molluskivores, detritivores, herbivores and algivores. Low values of Pianka's index of niche overlap were found in the sympatric micro-endemic species, while in the widely distributed species the indices ranged from low to very high. The analysis of lower pharyngeal jaw morphologies allowed discriminating a shape associated with piscivorous species from other foraging groups. The results of this study suggest that trophic niche segregation is a factor that could promotes diversification within the genus Herichthys although additional studies need to be performed to fully understand the speciation process in this group of Neotropical cichlid fishes.

La especiación es un proceso con múltiples factores que actúan a diferentes escalas de espacio y tiempo. La segregación de nichos tróficos es un proceso que ha promovido la diversificación en cíclidos en entornos lacustres, pero en el caso de los ríos no está claro. Herichthys es un género de cíclidos cuya distribución es la más boreal en América, el cual comprende 12 especies actualmente reconocidas, la mayoría microendémicas y solo dos con una amplia distribución. En el presente trabajo, se analizó el contenido estomacal y las morfologías de la mandíbula faríngea inferior de las especies del género para compararlas y evaluar su posible papel en la diversificación del grupo. La dieta en dichas especies es muy variada e incluyó tanto especies que pueden ser consideradas omnívoras como especialistas. Se encontraron valores bajos del índice de solapamiento alimentario (índice de Pianka) en las especies simpátricas microendémicas, mientras que en las especies ampliamente distribuidas el índice fue muy variable. El análisis de morfometría geométrica de la mandíbula faríngea inferior permite discriminar dos formas principales, una que incluye la especie piscívora y otra que incluye a los otros grupos alimentarios. Los resultados encontrados en este estudio sugieren que la segregación de nicho trófico es un factor que promueve claramente la diversificación dentro del género Herichthys, aunque se deben realizar estudios adicionales para comprender completamente el proceso de especiación en este grupo de peces neotropicales.

Morphometric variation of the Herichthys bartoni (Bean, 1892) species group (Teleostei: Cichlidae): How many species comprise H. labridens (Pellegrin, 1903)?

Cichlids of the tribe Heroini have long been a source of taxonomical conflict. In particular, the species included in the Herichthys bartoni group have failed to be recovered as monophyletic in different molecular studies. In this paper we use traditional and geometric morphometrics to evaluate morphological variation in the species included in the H. bartoni complex in order to evaluate the number of species it contains. An update of a previously published DNA barcoding study suggests the existence of three genetic clusters that included the six recognized species analyzed in this study, none of them recovered as monophyletic. On the other hand, geometric morphometrics arise as a useful tool to discriminate species due that traditional morphometrics showed a high overlap in the characters analyzed that prevents the proposal of diagnostic characters.

Los cíclidos de la tribu Heroini han experimentado un largo conflicto taxonómico. En particular, las especies incluidas en el grupo Herichthys bartoni no han sido recuperadas como monofiléticas en diversos estudios moleculares. En este artículo nosotros usamos morfometría tradicional y geométrica para evaluar la variación morfológica de las especies incluidas en el complejo H. bartoni para evaluar el numero de especies que contiene. Una actualización de un estudio previo de código de barras de ADN sugiere la existencia de tres grupos genéticos que incluyen las seis especies reconocidas analizadas en este estudio, ninguna de las cuales fue recuperada como monofilética. Por otro lado, la morfometría geométrica surge como una herramienta de utilidad para discriminar especies debido a que la variación en caracteres morfométricos tradicionales muestra altos niveles de solapamiento que previene la propuesta de caracteres diagnósticos.