Scale morphology and squamation patterns in cichlids (Teleostei, Perciformes): A comparative study

Problems in cichlid systematics call for new characteristics to be exploited. Scale surface morphology and squamation patterns could provide novel and useful information. A comparative study comprising 105 African, 10 American, and eight outgroup species was conducted to identify the most useful scale and squamation characters within the family and to clarify their phylogenetic significance. A great number of obviously genetically fixed characters were established. At least IS scale granulation types could be identified, proving to be ofparticular value for systematic purposes. Squamation patterns on the head, breast and fins were also found to be of interest in this context. Different character states are described and their distribution is reviewed, together with a discussion of their plesiomorphic status. From the results it can be concluded that for the Cichlidae scale and squamation studies can be valuable tools in investigating phylogenetic relationships.     

Molecular phylogeny and biogeography of the Malagasy and South Asian cichlids (Teleostei: Perciformes: Cichlidae)

Phylogenetic relationships of the Malagasy and South Asian cichlids are investigated using nucleotide characters from two mitochondrial genes, a 544 bp region of the large ribosomal subunit (16S) and a 649 bp region of cytochrome c oxidase subunit I (COI). This is the first molecular analysis to include a thorough taxonomic sampling of all Malagasy-South Asian genera, and a near complete taxonomic inventory of all valid species. Parsimony analysis of the combined data set (1193 aligned nucleotide positions) results in a single, completely resolved phylogenetic hypothesis. Results of this analysis, and that based on more comprehensive taxonomic sampling across Cichlidae for 16S alone (554 bp for 73 taxa), indicate that the Malagasy cichlids are paraphyletic, whereas the Malagasy and South Asian cichlids comprise a monophyletic group. In both analyses, the African and Neotropical assemblages are monophyletic. The Malagasy-South Asian cichlids are not recovered as plesiomorphic members of the family in either analysis. Two major clades are recovered within the Malagasy-South Asian assemblage and given subfamilial rank, Etroplinae, comprising Paretroplus (Madagascar) and Etroplus (southern India and Sri Lanka), and Ptychochrominae, comprising Ptychochromis, Ptychochromoides, and Oxylapia, all endemic Malagasy genera. Placement of the endemic Malagasy genus Paratilapia is equivocal depending on the gene fragment(s) analyzed. Inter- and intrageneric relationships within Ptychochrominae and Etroplinae are presented and discussed. The hypothesis of relationships for Cichlidae based on nucleotide characters from 16S alone, arguably the most comprehensive and broadly sampled data set across all major geographic assemblages to date, is congruent with prevailing hypotheses regarding the sequence of Gondwanan fragmentation and a vicariance scenario to explain the current distribution of cichlid fishes.     

A new species of Platygobiopsis from Vietnam (Teleostei: Perciformes: Gobiidae)

Platygobiopsis dispar sp. nov. is described on the basis of four specimens from the South China Sea off central Vietnam. The species differs from the only previously described species, P. akihito, in the morphology of the cephalic laterosensory system; in the distribution of scales on top of the head; in the absence of scales at the base of the pectoral-fin; in having 11 branched rays in the second dorsal and anal fins; in the lower predorsal, transverse, and lateral scale counts; and in other minor details. The genus Platygobiopsis is recorded for the South China Sea for the first time. The text was submitted by the author in English.     

Mating preferences in a species flock of Mexican pupfishes (Cyprinodon, Teleostei)

Female mating preferences were examined in three members of an evolutionarily young Cyprinodon lineage from Laguna Chichancanab, Yucatan, Mexico. In separate experiments, females of C. beltrani and C. labiosus , and C. beltrani and C. maya were simultaneously presented with conspecific and heterospecific males and given the opportunity to spawn with either or both. These three species showed different degrees of conspecific mate recognition: complete for C. maya , intermediate for C. labiosus , and indiscriminate for C. beltrani. We conclude that only C. maya is reproductively isolated and that C. beltrani and C. labiosus still hybridize in nature. Whereas between C. beltrani and C. maya mate choice is symmetric, it is asymmetric between C. beltrani and C labiosus. The results of the mating trials are consistent with mitochondrial DNA data which indicate that C. maya is genetically the most distinct species in the flock. They are also supported by experiments on visually and chemically mediated mate preferences.     

Phylogeny and species boundaries in the gobiid genus Gnatholepis (Teleostei: Perciformes)

Species of the goby genus Gnatholepis Bleeker, 1874, are common inhabitants of shallow tropical seas worldwide. In this study, mitochondrial DNA sequence (ND2 gene), from 349 Gnatholepis individuals sampled from across the South and Central Pacific and Caribbean, is used to infer phylogeny and determine species boundaries. Seven species of Gnatholepis are recognized: the Indo-Pacific G. anjerensis (Bleeker, 1851) [ G. cauerensis (Bleeker, 1853) is a synonym]; G. scapulostigma Herre, 1953; G. davaoensis Seale, 1910; G. knighti Jordan & Evermann, 1903; G. gymnocara Randall & Greenfield, 2001; G.  sp. Randall & Greenfield, 2001; and the Atlantic/Caribbean G. thompsoni Jordan, 1904. Results from the molecular phylogeny are compared with a previous morphology-based revision of the genus in order to establish which morphological characters diagnose species in correspondence with the molecular phylogeny.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 573–582.     

Phylogenomics of pike cichlids (Cichlidae: Crenicichla): the rapid ecological speciation of an incipient species flock

The rapid rise of phenotypic and ecological diversity in independent lake‐dwelling groups of cichlids is emblematic of the East African Great Lakes. In this study, we show that similar ecologically based diversification has occurred in pike cichlids (Crenicichla) throughout the Uruguay River drainage of South America. We collected genomic data from nearly 500 ultraconserved element (UCEs) loci and >260 000 base pairs across 33 species, to obtain a phylogenetic hypothesis for the major species groups and to evaluate the relationships and genetic structure among five closely related, endemic, co‐occurring species (the Uruguay River species flock; URSF). Additionally, we evaluated ecological divergence of the URSF based on body and lower pharyngeal jaw (LPJ) shape and gut contents. Across the genus, we recovered novel relationships among the species groups. We found strong support for the monophyly of the URSF; however, relationships among these species remain problematic, likely because of the rapid and recent evolution of this clade. Clustered co‐ancestry analysis recovered most species as well delimited genetic groups. The URSF species exhibit species‐specific body and LPJ shapes associated with specialized trophic roles. Collectively, our results suggest that the URSF consists of incipient species that arose via ecological speciation associated with the exploration of novel trophic roles.     

Systematics of Tanganyikan cichlid fishes (Teleostei: Perciformes)

The relationships among 53 genera of Tanganyikan cichlid fishes were analyzed based on internal and external morphological features. Comparison of the morphological cladistic tree with a previously proposed classification showed 5 of 12 tribes to be nonmonophyletic. Sixteen tribes were recognized, the changes in classification being that Trematocarini was treated as a junior synonym of Bathybatini; 5 new tribes were established for each of the following genera, Benthochromis, Boulengerochromis, Ctenochromis benthicola, Cyphotilapia, and Greenwoodochromis; Ctenochromis horei was transferred from Haplochromini to Tropheini; and Gnathochromis pfefferi was transferred from Limnochromini to Tropheini. The revised classification was supported by previously proposed molecular trees.     

AFLPs resolve phylogeny and reveal mitochondrial introgression within a species flock of African electric fish (Mormyroidea: Teleostei)

Estimating species phylogeny from a single gene tree can be especially problematic for studies of species flocks in which diversification has been rapid. Here we compare a phylogenetic hypothesis derived from cytochrome b (cyt b) sequences with another based on amplified fragment length polymorphisms (AFLP) for 60 specimens of a monophyletic riverine species flock of mormyrid electric fishes collected in Gabon, west-central Africa. We analyze the aligned cyt b sequences by Wagner parsimony and AFLP data generated from 10 primer combinations using neighbor-joining from a Nei-Li distance matrix, Wagner parsimony, and Dollo parsimony. The different analysis methods yield AFLP tree topologies with few conflicting nodes. Recovered basal relationships in the group are similar between cyt b and AFLP analyses, but differ substantially at many of the more derived nodes. More of the clades recovered with the AFLP characters are consistent with the morphological characters used to designate operational taxonomic units in this group. These results support our hypothesis that the mitochondrial gene tree differs from the overall species phylogeny due at least in part to mitochondrial introgession among lineages. Mapping the two forms of electric organ found in this group onto the AFLP tree suggests that posteriorly innervated electrocytes with nonpenetrating stalks have independently evolved from anteriorly innervated, penetrating-stalk electrocytes at least three times.     

The early development of substrate-brooding cichlids (Teleostei: Cichlidae) with a discussion of a new system of staging

The early development of five species of substrate-brooding cichlid (Aequidens pulcher; Cichlasoma bimaculatum; C. biocellatum; C. nigrofasciatum and Hemichromis bimaculatus) is described. Heterochrony in these species renders the standard system of staging inadequate for comparative study. A new system is proposed in which the period of development is resolved into five overlapping phases, each phase corresponding to a dissociable ontogenetic process. This form of staging permits comparison in a way which neither obscures the basic uniformity of cichlid development nor masks the differences between the species. Two special features are discussed: the universal presence in substrate-brooding cichlids of three pairs of embryonic adhesive organs; and the apparent presence of non-neural crest pigment on the embryos and alevins.     

Phylogeny and biogeography of cichlid fishes (Teleostei: Perciformes: Cichlidae)

Family level molecular phylogenetic analyses of cichlid fishes have generally suffered from a limited number of characters and/or poor taxonomic sampling across one or more major geographic assemblage, and therefore have not provided a robust test of early intrafamilial diversification. Herein we use both nuclear and mitochondrial nucleotide characters and direct optimization to reconstruct a phylogeny for cichlid fishes. Representatives of major cichlid lineages across all geographic assemblages are included, as well as nearly twice the number of characters as any prior family‐level study. In a strict consensus of 81 equally most‐parsimonious hypotheses, based on the simultaneous analysis of 2222 aligned nucleotide characters from two mitochondrial and two nuclear genes, four major subfamilial lineages are recovered with strong support. Etroplinae, endemic to Madagascar (Paretroplus) and southern Asia (Etroplus), is recovered as the sister taxon to the remainder of Cichlidae. Although the South Asian cichlids are monophyletic, the Malagasy plus South Asian lineages are not. The remaining Malagasy lineage, Ptychochrominae, is monophyletic and is recovered as the sister group to a clade comprising the African and Neotropical cichlids. The African (Pseudocrenilabrinae) and Neotropical (Cichlinae) lineages are each monophyletic in this reconstruction. The use of multiple molecular markers, from both mitochondrial and nuclear genes, results in a phylogeny that in general exhibits strong support, notably for early diversification events within Cichlidae. Results further indicate that Labroidei is not monophyletic, and that the sister group to Cichlidae may comprise a large and diverse assemblage of percomorph lineages. This hypothesis may at least partly explain why morphological studies that have attempted to place Cichlidae within Percomorpha, or that have tested cichlid monophyly using only “labroid” lineages, have met with only limited success. © The Willi Hennig Society 2004.     

Molecular phylogeny of the gobioid fishes (Teleostei: Perciformes: Gobioidei)

The phylogeny of groups within Gobioidei is examined with molecular sequence data. Gobioidei is a speciose, morphologically diverse group of teleost fishes, most of which are small, benthic, and marine. Efforts to hypothesize relationships among the gobioid groups have been hampered by the prevalence of reductive evolution among goby species; such reduction can make identification of informative morphological characters particularly difficult. Gobies have been variously grouped into two to nine families, several with included subfamilies, but most existing taxonomies are not phylogenetic and few cladistic hypotheses of relationships among goby groups have been advanced. In this study, representatives of eight of the nine gobioid familes (Eleotridae, Odontobutidae, Xenisthmidae, Gobiidae, Kraemeriidae, Schindleriidae, Microdesmidae, and Ptereleotridae), selected to sample broadly from the range of goby diversity, were examined. Complete sequence from the mitochondrial ND1, ND2, and COI genes (3573 bp) was used in a cladistic parsimony analysis to hypothesize relationships among the gobioid groups. A single most parsimonious topology was obtained, with decay indices indicating strong support for most nodes. Major phylogenetic conclusions include that Xenisthmidae is part of Eleotridae, and Eleotridae is paraphyletic with respect to a clade composed of Gobiidae, Microdesmidae, Ptereleotridae, Kraemeriidae, and Schindleriidae. Within this five-family clade, two clades are recovered. One includes Gobionellinae, which is paraphyletic with respect to Kraemeriidae, Sicydiinae, Oxudercinae, and Amblyopinae. The other contains Gobiinae, also paraphyletic, and including Microdesmidae, Ptereleotridae, and Schindleriidae. Previous morphological evidence for goby groupings is discussed; the phylogenetic hypothesis indicates that the morphological reduction observed in many goby species has been derived several times independently.     

The eastern Pacific species of Bathygobius (Perciformes: Gobiidae)

The circumtropical gobiid genus Bathygobius Bleeker is defined and three Eastern Pacific species are redescribed, with first dorsal fin pattern and postorbital blotches being shown to be additional characters of diagnostic value. Two mainland species are recognised, the Mexican-Panamanian B. ramosus Ginsburg 1947 and the Panamanian B. andrei (Sauvage 1880). B. ramosus is now reported from Clarión Island, Revillagigedos, and also from Cocos Island. Meristic variation of ramosus is tabulated for local populations and PCA analysis of their morphometry suggests regional differentiation in this species, with Tres Marias and Revillagigedos populations clustering away from mainland and Montuosa material. An insular species, B. lineatus (Jenyns 1842) from the Galapagos is defined, with B. arundelii (Garman 1899) from Clipperton Island and B. l. lupinus Ginsburg 1947 from Lobos de Afuera, off Peru, placed as nominal subspecies of lineatus. This species resembles the Indo-west Pacific B. fuscus and Atlantic basin B. soporator more closely than it does ramosus and andrei and may be the product of transpacific dispersal. A similar origin for B. ramosus is discussed but it seems more likely that both B. ramosus and B. andrei have Caribbean sister species.     

The oldest human fossil in Europe,from Orce (Spain)

The Orce region has one of the best late Pliocene and early Pleistocene continental paleobiological records of Europe. It is situated in the northeastern sector of the intramontane Guadix-Baza Basin (Granada, Andalusia, southern Spain). Here we describe a new fossil hominin tooth from the site of Barranco León, dated between 1.02 and 1.73 Ma (millions of years ago) by Electron Spin Resonance (ESR), which, in combination with paleomagnetic and biochronologic data, is estimated to be close to 1.4 Ma. While the range of dates obtained from these various methods overlaps with those published for the Sima del Elefante hominin locality (1.2 Ma), the overwhelming majority of evidence points to an older age. Thus, at the moment, the Barranco León hominin is the oldest from Western Europe.