Heroine Cichlids Research Articles

Comparative analysis of cranial morphology in Middle-American heroine cichlids (Actinopterygii: Cichliformes).

Heroine cichlids are characterized by high morphological diversity, mainly in structures related to the capture and processing of food. The existence of ecomorphological groups has been proposed based on feeding behavior, where it is common for some phylogenetically unrelated species to show evolutionary convergence. Using geometric morphometrics and comparative phylogenetic methods, the variation in cranial morphology was evaluated for 17 species of heroine cichlids representing 5 ecomorphs. Cranial ecomorphs were recovered and significant differences were determined. Morphological variation of the ecomorphs was mainly explained by two axes: (1) the position of the mouth determined by the shape of the bones of the oral jaw and (2) the height of the head, defined by the size and position of the supraoccipital crest and the distance to the interopercle-subopercle junction. Cranial variation among species was related to phylogeny. To better understand the evolution of cranial morphology, it is necessary to evaluate the morphofunctional relationship of other anatomical structures related to feeding, as well as to increase the number of study species in each ecomorph by including other lineages.

Comparing Ultraconserved Elements and Exons for Phylogenomic Analyses of Middle American Cichlids: When Data Agree to Disagree.

Choosing among types of genomic markers to be used in a phylogenomic study can have a major influence on the cost, design, and results of a study. Yet few attempts have been made to compare categories of next-generation sequence markers limiting our ability to compare the suitability of these different genomic fragment types. Here, we explore properties of different genomic markers to find if they vary in the accuracy of component phylogenetic trees and to clarify the causes of conflict obtained from different data sets or inference methods. As a test case, we explore the causes of discordance between phylogenetic hypotheses obtained using a novel data set of ultraconserved elements (UCEs) and a recently published exon data set of the cichlid tribe Heroini. Resolving relationships among heroine cichlids has historically been difficult, and the processes of colonization and diversification in Middle America and the Greater Antilles are not yet well understood. Despite differences in informativeness and levels of gene tree discordance between UCEs and exons, the resulting phylogenomic hypotheses generally agree on most relationships. The independent data sets disagreed in areas with low phylogenetic signal that were overwhelmed by incomplete lineage sorting and nonphylogenetic signals. For UCEs, high levels of incomplete lineage sorting were found to be the major cause of gene tree discordance, whereas, for exons, nonphylogenetic signal is most likely caused by a reduced number of highly informative loci. This paucity of informative loci in exons might be due to heterogeneous substitution rates that are problematic to model (i.e., computationally restrictive) resulting in systematic errors that UCEs (being less informative individually but more uniform) are less prone to. These results generally demonstrate the robustness of phylogenomic methods to accommodate genomic markers with different biological and phylogenetic properties. However, we identify common and unique pitfalls of different categories of genomic fragments when inferring enigmatic phylogenetic relationships.

Diversity and evolution of the Middle American cichlid fishes (Teleostei: Cichlidae) with revised classification

We present a taxonomically complete and topologically robust molecular phylogeny of the Middle American heroine cichlids based on which we review their diversity and genus-level systematics. In order to ascertain the diversity of the group and its phylogeny we have used three nested taxon sampling analyses of the concatenated nDNA/mtDNA datasets and additionally to these analyses we present a summary of the results of a new Next Generation Sequencing-generated nuclear phylogeny based on a data set of ~ 140,000 informative characters. The NGS ddRAD phylogeny has a species-level sampling covering virtually all species (including the enigmatic Cichlasoma microlepis Dahl, 1960) with multiple sequenced specimens per species. Based on our results the Middle American heroine cichlids are made up of three main clades. The three clades (the herichthyines, the amphilophines, and the astatheroines) are however not each other sister groups since they are interspersed with South American (Australoheros, Caquetaia, Chocoheros, Heroina, Mesoheros) and Antillean (Nandopsis) genera and they represent probably two separate colonization events of Middle America from South America, probably via the Antilles. Our study reveals many cases of cytonuclear discordance and/or introgressive hybridization both at the genus and deeper levels stressing the importance to study the nuclear and mitochondrial phylogenetic signals independently and not solely in concatenated analyses. We have found that a great majority of morphological characters are ecologically correlated and that they form only a limited number of functionally-determined combinations – i.e. ecomorphs. We have found five main cranial ecomorphs but only two postcranial ecomorphs (the lotic and lentic ecomorphs, plus the undifferentiated ancestral character combination). The cranial and postcranial ecomorphs are not combined completely randomly having produced thirteen modular whole-body ecomorphs. Both the cranial and postcranial ecomorphs, and even their combinations, have evolved repeatedly in the Middle American cichlids in the same habitats both in sympatry as well as in allopatry. Our analyses of the diversity of Middle American cichlid clade support the existence of 31 genera in Middle America (plus six in South America and one in the Greater Antilles) as separate evolutionary lineages occupying separate adaptive zones. Nine new genera are described here for species and species groups that have lacked a genus level name to this day or were associated with other unrelated genera. We also review the species level diversity based on the mtDNA cytb gene population-level analysis. Furthermore, we provide a new biogeographical analysis of the group which explains their evolutionary history and demonstrates that biogeography is a much better indicator of evolutionary relationships in this fish group than are most morphological characters due to their ecological correlation.

Coordinated Dispersal and Pre-Isthmian Assembly of the Central American Ichthyofauna.

We document patterns of coordinated dispersal over evolutionary time frames in heroine cichlids and poeciliine live-bearers, the two most species-rich clades of freshwater fishes in the Caribbean basin. Observed dispersal rate ( values were estimated from time-calibrated molecular phylogenies in Lagrange+, a modified version of the ML-based parametric biogeographic program Lagrange. is measured in units of “wallaces” (wa) as the number of biogeographic range-expansion events per million years. estimates were generated on a dynamic paleogeographic landscape of five areas over three time intervals from Upper Cretaceous to Recent. Expected dispersal rate ( values were generated from alternative paleogeographic models, with dispersal rates proportional to target area and source-river discharge volume, and inversely proportional to paleogeographic distance. Correlations between and were used to assess the relative contributions of these three biogeographic parameters. estimates imply a persistent dispersal corridor across the Eastern (Antillean) margin of the Caribbean plate, under the influence of prevailing and perennial riverine discharge vectors such as the Proto–Orinoco–Amazon river. Ancestral area estimation places the earliest colonizations of the Greater Antilles and Central America during the Paleocene–Eocene (ca. 58–45 Ma), potentially during the existence of an incomplete Paleogene Arc (∼59 Ma) or Lesser Antilles Arc (∼45 Ma), but predating the GAARlandia land bridge (∼34–33 Ma). Paleogeographic distance is the single best predictor of . The Western (Central American) plate margin did not serve as a dispersal corridor until the Late Neogene (12–0 Ma), and contributed relatively little to the formation of modern distributions.

Species tree estimation and the historical biogeography of heroine cichlids

Heroine cichlids are major components of the fish faunas in both Central America and the Caribbean. To examine the evolutionary patterns of how cichlids colonized both of these regions, we reconstructed the phylogenetic relationships among 23 cichlid lineages. We used three phylogenetically novel nuclear markers (Dystropin b, Myomesin1, and Wnt7b) in combination with sequence data from seven other gene regions (Nd2, Rag1, Enc1, Sreb2, Ptr, Plagl2, and Zic1) to elucidate the species tree of these cichlids. The species examined represent major heroine lineages in South America, Central America, and the Greater Antilles. The individual gene trees of these groups were topologically quite discordant. Therefore, we combined the genetic partitions and inferred the species tree using both concatenation and a coalescent-based Bayesian method. The two resulting phylogenetic topologies were largely concordant but differed in two fundamental ways. First, more nodes in the concatenated tree were supported with substantial or 100% Bayesian posterior support than in the coalescent-based tree. Second, there was a minor, but biogeographically critical, topological difference between the concatenated and coalescent-based trees. Nevertheless, both analyses recovered topologies consistent with the Greater Antillean heroines being phylogenetically nested within the largely Central American heroine radiation. This study suggests that reconstructions of cichlid phylogeny and historical biogeography should account for the vagaries of individual gene histories.

Molecular systematics of the enigmatic Middle American genus Vieja (Teleostei: Cichlidae)

The genus Vieja represents a group of heroine cichlids (Teleostei: Cichlidae) distributed on the Atlantic and Pacific slopes of North and Central America from southern Mexico to Panama. Sixteen species of Vieja are presently recognized; however, based on long-standing taxonomic problems, the genus itself appears to be weakly defined. A number of different generic designations have been proposed for members of Vieja, and recent systematic studies of heroine cichlids have not specifically addressed the validity of the grouping and have not included all species in the genus. Therefore, the purpose of this study was to assess the monophyly of the genus Vieja by including all nominal species in the genus using the mitochondrial encoded cytochrome b gene and nuclear S7-1 intron. Results of Maximum Parsimony, Bayesian inference, and topology tests (constraint tree searches and post-burn-in Bayesian filtering) indicate that the genus is not monophyletic as it is currently recognized. The genus Herichthys was recovered as sister to a clade consisting of a number of Vieja species ( V. fenestrata, V. guttulata, V. zonata, V. hartwegi, V. bifasciata, V. breidohri, V. argentea, V. regani, V. melanura, V. synspila, and V. maculicauda, as well as Paraneetroplus bulleri). A clade consisting of V. intermedia, V. godmanni, and V. microphthalma was recovered sister to Theraps. Additionally, V. heterospila and V. tuyrensis were recovered outside of Vieja and Herichthys clades. Based on the results of this comprehensive study, we suggest a revised classification of Vieja species.

Temporal diversification of Central American cichlids

BackgroundCichlid fishes are classic examples of adaptive radiation because of their putative tendency to explosively diversify after invading novel environments. To examine whether ecological opportunity increased diversification (speciation minus extinction) early in a species-rich cichlid radiation, we determined if Heroine cichlids experienced a burst of diversification following their invasion of Central America.ResultsWe first reconstructed the Heroine phylogeny and determined the basal node to use as the root of Central American Heroine diversification. We then examined the influence of incomplete taxon sampling on this group's diversification patterns. First, we added missing species randomly to the phylogeny and assessed deviations from a constant rate of lineage accumulation. Using a range of species numbers, we failed to recover significant deviations from a pure-birth process and found little support for an early burst of diversification. Then, we examined patterns of lineage accumulation as nodes were increasingly truncated. We assumed that as we removed more recently diverged lineages that sampling would become more complete thereby increasing the power to detect deviations from a pure-birth model. However, truncation of nodes provided even less support for an early burst of diversification.ConclusionsContrary to expectations, our analyses suggest Heroine cichlids did not undergo a burst of diversification when they invaded from South America. Throughout their history in Central America, Heroine cichlids appear to have diversified at a constant rate.

Co-evolution of the premaxilla and jaw protrusion in cichlid fishes (Heroine: Cichlidae)

The ability of Perciform fishes to protrude their jaw has likely been critical to the trophic diversification of this group, which includes approximately 20% of all vertebrates. The length of the ascending process of the premaxilla is thought to influence the maximum extent that cichlids and other Perciforms protrude their oral jaw. Using a combination of morphometrics, kinematics, and new phylogenetic hypotheses for 20 Heroine cichlid species, we tested the evolutionary relationship between the length of the premaxillary ascending process and maximum jaw protrusion. In this clade, the length of the ascending process of the premaxilla ranged from 11.6–32.7% with respect to standard length whereas maximum jaw protrusion ranged from 3.5–23.4% with respect to standard length. The evolutionary relationships among the Heroine cichlids obtained from the genetic partitions cytochrome b, S7, and RAG1 showed limited concordance. However, correlations between the length of the ascending process and maximum jaw protrusion were highly significant when examined as independent contrasts using all three topologies. Evolutionary change in the length of the ascending process of the premaxilla is likely critical for determining the amount of jaw protrusion in Perciform groups such as cichlid fishes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 619–629.

Phylogenetic relationships of Middle American cichlids (Cichlidae, Heroini) based on combined evidence from nuclear genes, mtDNA, and morphology

Heroine cichlids are the second largest and very diverse tribe of Neotropical cichlids, and the only cichlid group that inhabits Mesoamerica. The taxonomy of heroines is complex because monophyly of most genera has never been demonstrated, and many species groups are without applicable generic names after their removal from the catch-all genus Cichlasoma (sensu Regan, 1905). Hence, a robust phylogeny for the group is largely wanting. A rather complete heroine phylogeny based on cytb sequence data is available [Concheiro Pérez, G.A., Říčan O., Ortí G., Bermingham, E., Doadrio, I., Zardoya, R. 2007. Phylogeny and biogeography of 91 species of heroine cichlids (Teleostei: Cichlidae) based on sequences of the cytochrome b gene. Mol. Phylogenet. Evol. 43, 91–110], and in the present study, we have added and analyzed independent data sets (nuclear and morphological) to further confirm and strengthen the cytb-phylogenetic hypothesis. We have analyzed a combined cytb-nuclear (RAG1 and two S7 introns) data set of 48 species representing main heroine lineages to achieve further resolution of heroine higher taxonomic levels and a combined cytb-morphological data set of 92 species to stabilize generic taxonomy. The recovered phylogenies supported the circumamazonian—CAM—Heroini (sensu Concheiro Peréz et al., 2007) as a monophyletic group, that could be divided into six main clades: (1) australoheroines (the southernmost heroine genus Australoheros), (2) nandopsines (the Antillean genus Nandopsis), (3) caquetaines (including the north western Amazonian genera Caquetaia and Heroina), (4) astatheroines (including Astatheros, Herotilapia and Rocio), (5) amphilophines (including Amphilophus and related genera), and (6) herichthyines (including Herichthyis and related genera). Nuclear and mitochondrial data partitions arrived at highly congruent topologies. Suprageneric relationships were influenced mainly by the nuclear signal, as well as the most basal phylogenetic position of Australoheros within CAM heroines. The new phylogeny of the tribe Heroini provides robust framework to stabilize the taxonomy of the group and for future comparative studies on these morphologically and ecologically diverse freshwater fishes. Morphology was mostly informative at the genus level and aid in determining the monophyly and composition of heroine genera. Upon acceptance of all putative genera, as recovered in this study, the Heroini would be with 35 genera the most genus-rich clade of Neotropical cichlids.

CONVERGENCE IN A MECHANICALLY COMPLEX PHENOTYPE: DETECTING STRUCTURAL ADAPTATIONS FOR CRUSHING IN CICHLID FISH

Morphological convergence provides strong evidence that evolution is adaptive. However, putatively convergent morphology is often examined in two dimensions with no explicit model of function. In this study, we investigated structural and mechanical similarities of the lower pharyngeal jaw (LPJ) in cichlid fish that have evolved the ability to crush hard-shelled molluscs. Using a novel phylogeny, we demonstrated molluscivory has been gained and/or been lost numerous times in Heroine cichlids. Within this comparative framework, we produced three-dimensional computed tomography (CT) scans for LPJs of both morphotypes in the trophically polymorphic Herichthys minckleyi and six evolutionarily independent pairs of closely related species. Like H. minckleyi, these species exhibit divergence between a molluscivore and a nonmolluscivore. Using the CT scans, we generated finite element models of papilliform H. minckleyi LPJs to determine where stress would concentrate in a jaw not modified to crush molluscs. Then, we examined whether stress in the papilliform LPJ predicted structural modifications in molariform H. minckleyi and other molluscivorous species. Despite potential constraints, stresses imposed during prey processing explain 40% of LPJ variation in mollusc crushing species. The structural and mechanical analyses also suggest divergence found in polymorphic species could provide the substrate for trophic differences found in reproductively isolated cichlids.

MICRO- AND MACROEVOLUTIONARY DECOUPLING OF CICHLID JAWS: A TEST OF LIEM'S KEY INNOVATION HYPOTHESIS

The extent to which elements of functional systems can change independently (modularity) likely influences the diversification of lineages. Major innovations in organismal design, like the pharyngeal jaw in cichlid fishes, may be key to a group's success when they relax constraints on diversification by increasing phenotypic modularity. In cichlid fishes, pharyngeal jaw modifications that enhanced the ability to breakdown prey may have freed their oral jaws from serving their ancestral dual role as a site of both prey capture and prey processing. This functional decoupling that allowed the oral jaws to become devoted solely to prey capture has been hypothesized to have permitted the two sets of cichlid jaws to evolve independently. We tested the hypothesis that oral and pharyngeal jaw mechanics are evolutionarily decoupled both within and among Neotropical Heroine cichlids. In the trophically polymorphic species Herichthys minckleyi, molariforms that exhibit enlarged molarlike pharyngeal jaw teeth were found to have approximately 400% greater lower jaw mass compared to H. minckleyi with the alternative papilliform pharyngeal morphology. However, oral jaw gape, lower jaw velocity ratios, anterior jaw linkage mechanics, and jaw protrusion did not differ between the morphotypes. In 40 other Heroine species, there was a weak correlation between oral jaw mechanics and pharyngeal jaw mass when phylogenetic history was ignored. Yet, after expansion of the cytochrome b phylogeny for Heroines, change in oral jaw mechanics was found to be independent of evolutionary change in pharyngeal jaw mass based on independent contrasts. Evolutionary decoupling of oral and pharyngeal jaw mechanics has likely played a critical role in the unparalleled trophic diversification of cichlid fishes.

Phylogeny and biogeography of 91 species of heroine cichlids (Teleostei: Cichlidae) based on sequences of the cytochrome b gene

Heroini constitute the second largest tribe of Neotropical cichlids and show their greatest diversity in Mesoamerica. Although heroine species are morphologically and ecologically very diverse, they were all historically assigned to one single genus, Cichlasoma that was never formally revised from a phylogenetic point of view. Here, we present the most comprehensive molecular phylogeny of the tribe Heroini to date, based on the complete DNA sequence of the mitochondrial gene cytochrome b, and the analysis of 204 individuals representing 91 species. Phylogenetic analyses did not support the monophyly of heroines because the genus Pterophyllum was placed as the sister group of all remaining heroines plus cichlasomatines. However, the recovered relative position of Pterophyllum was without strong statistical support. Within the remaining heroines, Hyspelecara and Hoplarchus are recovered with low support in a basal position with respect to a clade that includes Heros, Uaru, Mesonauta, and Symphysodon, and the circumamazonian (CAM) heroines. The first clade is restricted to South America. The largest clade of heroines, the CAM heroines, include more than 85% of the species within the tribe. This clade is mostly Mesoamerican, but also contains four species found in the Greater Antilles ( Nandopsis), and three genera found in South America (the ‘Heros’ festae group, Australoheros, and Caquetaia). Up to eight major lineages can be recovered within the CAM heroines, but the phylogenetic relationships among them remain unresolved. Two large suprageneric groups can be distinguished, the amphilophines and the herichthyines. The amphilophines include Amphilophus, Archocentrus, Hypsophrys, Neetroplus, Parachromis, Petenia, and five additional unnamed genera (the ‘Heros’ istlanus group, the ‘Amphilophus’ calobrensis group, the ‘Heros’ urophthalmus group, the ‘Heros’ wesseli group, and the ‘Heros’ sieboldii group). The herichthyines include the crown-group herichthyines ( Herichthys, Theraps, Vieja, and Paratheraps) and the genera Tomocichla, Herotilapia, and Thorichthys, together with three unnamed genera (the ‘Heros’ umbriferus group, the ‘Heros’ grammodes group, and the ‘Heros’ salvini group). Amphilophines are prevalent in southern Mesomerica south of the Motagua fault. Herichthyines have basal linages in Central America, whereas crown-group herichthyines and three related genera are found north from the Motagua fault. At least two independent origins are required to explain current Mesoamerican heroine distribution. Dispersal of heroines from South America into Mesoamerica was dated between 24 and 16 million years ago (MYA) based on geological calibrations and on standard fish mitochondrial cytochrome b rates, respectively. These datings cannot be reconciled with currently known geological evidence, and the existence of a connection between Central America and South America in the Miocene needs to be postulated in order to explain the origins of Mesoamerican heroine lineages. However, our datings agree with those estimated for the dispersal of other secondary freshwater fishes (Rivulidae, Synbranchus) into Mesoamerica, and predate the invasion of primary freshwater fishes by at least 10 myr.

Function of a key morphological innovation: fusion of the cichlid pharyngeal jaw.

The pharyngeal jaw of cichlids may represent a key innovation that facilitated their unparalleled trophic divergence. In cichlids, 'fusion' of the lower pharyngeal jaw (LPJ) results from suturing between the two lower ceratobranchials. To examine, what novel abilities a more extensively fused pharyngeal jaw may confer, the function of LPJ suturing was examined in Heroine cichlids. Greater LPJ suturing, pharyngeal jaw splitting under compression and the forces used to crush molluscs in the wild suggest increased LPJ fusion in the trophically polymorphic Herichthys minckleyi operates to strengthen the pharyngeal jaw. Among Heroine cichlid species, the presence of an external LPJ suture and feeding specialization on molluscs was evolutionarily quite variable, but greater LPJ fusion estimated from the amount of external suturing was highly correlated with molluscivory. Throughout cichlid diversification, increased pharyngeal jaw fusion via suturing has likely helped to reinforce the LPJ during pharyngeal processing thereby facilitating the ability of cichlids to exploit durable prey.

Cichlid jaw mechanics: linking morphology to feeding specialization

Summary The utility of the anterior jaw four‐bar linkage model and the trophic consequences of jaw protrusion were investigated in Heroine cichlids by examining the evolutionary relationships among maxillary kinematic transmission (KT), maximum jaw protrusion and dietary specialization on evasive prey. In 31 species of Heroine cichlids, a four‐bar linkage model was used to generate kinematic predictions of maxillary KT, the angular amount of maxilla rotation per unit lower jaw rotation, expected during mouth opening. Maxillary KT averaged 0·79 and ranged from 0·58 in Herichthys tamasopoensis to 1·06 in Petenia splendida. Because the maxilla pushes the toothed premaxilla out during jaw protrusion, we predicted higher maxillary KT should characterize species with greater maximum jaw protrusion. Maximum jaw protrusion ranged from 1·5 to 14·2% of cichlid standard length and was highly correlated with greater maxillary KT. The proportion of fish and crustaceans in the diet of these cichlids was correlated with maximum protrusion, suggesting jaw protrusion may aid in the capture of evasive prey. Phylogenetic independent contrasts indicate changes in anterior jaw mechanics may be necessary for diversification of cichlid jaw protrusion abilities, and the labile evolution of jaw protrusion in Heroine cichlids likely facilitated the repeated specialization on evasive prey during their diversification.

Temporal diversification of Mesoamerican cichlid fishes across a major biogeographic boundary

The Mexican Neovolcanic Plateau sharply divides the vertebrate fauna of Mesoamerica where the climate of both the neotropics and temperate North America gradually blend. Only a few vertebrate groups such as the Heroine cichlids, distributed from South America to the Rio Grande in North America, are found both north and south of the Neovolcanic Plateau. To better understand the geography and temporal diversification of cichlids at this geologic boundary, we used mitochondrial DNA sequences of the cytochrome b (cyt b) gene to reconstruct the relationships of 52 of the approximately 80 species of Heroine cichlids in Mesoamerica. Our analysis suggests several cichlids in South America should be considered as part of the Mesoamerican Heroine clade because they and the cichlids north of the Isthmus of Panama are clearly supported as monophyletic with respect to all other Neotropical cichlids. We also recovered a group containing species in Paratheraps + Paraneetroplus + Vieja as the sister clade to Herichthys. Herichthys is the only cichlid clade north of the Mexican Plateau and it is monophyletic. Non-parametric rate smoothing of cichlid cyt b sequence resulted in an estimated divergence time of approximately 6 million years for Herichthys. This temporal diversification is concordant with divergence times estimated for anurans in the genus Bufo, a group that exhibits a similar geographic distribution. Our results indicate the 5-million-year-old extension of the Mexican Neovolcanic Plateau to the Gulf Coast of Mexico has strongly influenced the current transition between the vertebrate faunas of the Neotropics and Nearctic.

Systematics and Evolution of Lower Central American Cichlids Inferred from Analysis of Cytochrome b Gene Sequences

Central American cichlids allied to the genus “Cichlasoma” are thought to be of South American ancestry. The group has apparently undergone extensive morphological, ecological, and behavioral differentiation in Central America following colonization. Uncertainties regarding the systematics of the group and the timing of colonization complicate interpretation of the biological history of cichlids in Central America. We determined complete cytochrome b gene sequences for 54 individual cichlids representing 21 species to test hypotheses regarding the time of origin and pattern of diversification of lower Central American cichlids. The data also bear on issues relating to the systematics and taxonomy of heroine cichlids. Our results suggest that cichlids have been in Central America since the middle to late Miocene. Moreover, the data provide evidence of a rapid radiation early in the history of the group. Similar ecomorphological types have evolved multiple times. Inferences of convergent morphological evolution may, in part, explain a lack of concordance between the mitochondrial gene tree and previous inferences of phylogenetic relationships based on observable characteristics. Phylogenetic inferences based on the molecular data provide support for the recognition of “sections” [erected by Regan (1905) and others] as distinct genera and suggest that thorough revision of Central American cichlids is necessary before the extent of biological diversity within the group can be fully appreciated.