Phylogeny and species diversity of the genusHerichthys(Teleostei: Cichlidae)

Abstract

We provide a review of the systematics of Herichthys by evaluating the usefulness of several mitochondrial and nuclear genetic markers together with morphological data. The nDNA next‐generation sequencing ddRAD analysis together with the mtDNA cytochrome b gene provided well‐resolved and well‐supported phylogenies of Herichthys. On the other hand, the nDNA S7 introns have limited resolution and support and the COI barcoding analysis completely failed to recover all but one species of Herichthys as monophyletic. The COI barcoding as currently implemented is thus insufficient to distinguish clearly distinct species in the genus Herichthys that are supported by other molecular markers and by morphological characters. Based on our results, Herichthys is composed of 11 species and includes two main clades (the H. labridens and H. cyanoguttatus species groups). Herichthys bartoni is in many respects the most plesiomorphic species in the genus and has a conflicting phylogenetic position between mtDNA and nDNA markers, where the robust nDNA ddRAD data place it as a rather distant basal member of the H. labridens species group. The mtDNA of H. bartoni is on the other hand only slightly divergent from the sympatric and syntopic H. labridens, and the species thus probably have hybridized in the relatively recent past. The sympatric and syntopic Herichthys steindachneri and H. pame are supported as sister species. The Herichthys cyanoguttatus species group shows two well‐separated basal species (the northernmost H. minckleyi and the southernmost H. deppii) followed by the closely related and centrally distributed species H. cyanoguttatus, H. tepehua, H. carpintis, and H. tamasopoensis whose relationships differ between analyses and show likely hybridizations between themselves and the two basal species as suggested by conflicts between DNA analyses. Several instances of introgressions/hybridizations have also been found between the two main clades of Herichthys.

Molecular clocks, biogeography and species diversity inHerichthyswith 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

“…Our sampling for molecular analyses includes all described species of Herichthys including H. molango that has not been previously evaluated. The dataset is based on that of Říčan et al (2016) and Pérez-Miranda et al (2018) with the inclusion of many newly generated sequences (see supplemental Table S1). The dataset includes the complete cytb marker (1,137 bp) with 164 specimens (i.e., 141 haplotypes) from 62 localities ( Fig.…”

“…The dataset includes the complete cytb marker (1,137 bp) with 164 specimens (i.e., 141 haplotypes) from 62 localities ( Fig. 1; compared to 99 specimens from 32 localities in Říčan et al, 2016;Pérez-Miranda et al, 2018). The dataset is anchored by 13 additional species, i.e., Hypselecara coryphaenoides, Symphysodon aequifasciata, Mesonauta festivus, Heros sp., Uaru amphiacanthoides, Nandopsis tetracanthus, Nandopsis haitiensis, Caquetaia sp.…”

“…Unlike the two previous methods Starbeast analysis requires that the haplotypes are assigned to a priori species. We have used the traditionally recognized species (sensu Pérez-Miranda et al, 2018) for this analysis since they are not conflicted by the trees generated by RaxML and Beast (which all gave similar topologies). We have run all three delimitation analyses with the inclusion of only one, the most closely related, outgroup (the genus Vieja).…”

“…As a consequence of its isolation, Herichthys is the most diverse genus in terms of its ecomorphology among the Middle American cichlids (Říčan et al, 2016). In Herichthys, there are eco-morphologically polymorphic species, probably in early stages of speciation, and also sympatric and syntopic sister species that together with the nature of their isolation and ecomorphological divergence suggest sympatric speciation (Říčan et al, 2016;Pérez-Miranda et al, 2018). A large proportion of the diversity in Herichthys is therefore potentially the result of ecological opportunity afforded to the genus by the absence of other sympatric cichlid genera (Seehausen, 2015;Říčan et al, 2016;Burress et al, 2018;Piálek et al, 2019;Piálek et al, 2018).…”

“…While the phylogenetic position of Herichthys within Middle American cichlids is wellestablished (Říčan et al, 2016), its species diversity is complicated and remains contested (reviewed in Pérez-Miranda et al, 2018). Pérez-Miranda et al (2018) have evaluated the usefulness of several nDNA and mtDNA markers and concluded that the best resolution of both phylogenetic relationships and species-diversity issues was achieved by the nDNA ddRAD and mtDNA cytb datasets among those currently available. Based on the available data that also included morphological diagnosability of the species, Pérez-Miranda et al (2018) have concluded that Herichthys includes 11 species (one species has not been evaluated, H. molango, and several recently described species were placed in synonymy).…”