Miniature Species Research Articles

Cryptic species diversity in the Hypsolebiasmagnificus complex, a clade of endangered seasonal killifishes from the São Francisco River basin, Brazilian Caatinga (Cyprinodontiformes, Aplocheilidae).

A great diversity of animal species adapted to life in the semi-arid Caatinga of northeastern Brazil, including seasonal killifishes, has been reported in the last three decades. More recently, field and molecular data have shown a high occurrence of cryptic species. The killifish group herein analysed, the Hypsolebiasmagnificus species complex, is endemic to the middle and southern portion of the Caatinga, occupying about 120 km along the floodplains of the middle São Francisco River and some adjacent tributaries. Species of this complex are rare and presently considered threatened with extinction, being uniquely found in pools protected by trees and bushes. Single-locus delimitation methods were used to test species limits of populations displaying different colour patterns along the whole distribution of the complex. All analyses consistently supported the three nominal species and two new, herein described: H.gardneri Costa, sp. n., from the floodplains of the middle São Francisco River and H.hamadryades Costa, sp. n., from the Gorotuba River floodplains. The phylogenetic analysis highly supports H.hamadryades as sister to a clade comprising H.gardneri and H.harmonicus. Our field observations suggest that H.hamadryades is a miniature species. This study indicates that the H.magnificus complex comprises cryptic species apparently endemic to small areas and extremely vulnerable to environmental changes, deserving high concern.

New remarkable sexually dimorphic miniature species of Hyphessobrycon (Characiformes: Characidae) from the upper Rio Tapajós basin.

A new miniature species of Hyphessobrycon is described from an affluent of the Rio Papagaio, tributary of the Rio Juruena, upper Rio Tapajós basin, Mato Grosso State, Brazil. The new species can be distinguished from its congeners by having a remarkable secondary sexual dimorphism in its live colouration (males are red and females yellow), well-defined and relatively narrow dark midlateral stripe extending from tip of snout to tip of middle caudal-fin rays, absence of humeral blotch, 15-18 branched anal-fin rays and five or six branched pelvic-fin rays. The sexually dimorphic colouration of the new species is briefly discussed regarding its temporal variation.

A new miniature species of geophilomorph centipede from the Ecuadorian Amazon rainforest (Chilopoda: Geophilomorpha: Ballophilidae)

ABSTRACTA new miniature species of geophilomorph centipede, Ityphilus grismadoi sp. nov. (Chilopoda: Geophilomorpha: Ballophilidae) is fully described and illustrated on the basis of specimens collected in Yasuní Scientific Station and Jatun Sacha Biological Station, both located in the Ecuadorian Amazon. The new species is characterized by having the internal edge of the forcipular tarsungulum completely smooth, this trait being shared by six other Neotropical members of the genus Ityphilus Cook, 1899; and another species (I. cifuentesi Cupul Magaña, 2014) from the transition area between the Neotropical and Nearctic regions. Ityphilus grismadoi sp. nov. is distinguished from these seven species by the low number of leg-bearing segments (37, 39), and the very small body size (7–9 mm in length) and for a confident identification it is compared in detail with those having a roughly similar range of leg-bearing segments. This is the first record of the genus Ityphilus from Ecuador and the second report of a ballophilid centipede from the country (the other represented by Zygethmus pantenus Chamberlin, 1957). A key to the Neotropical members of Ityphilus with forcipular tarsungulum smooth (including I. cifuentesi) is also given.http://zoobank.org/urn:lsid:zoobank.org:pub:32F77890-9DC5-47AC-A794-29C4437C3B93http://zoobank.org/urn:lsid:zoobank.org:act:3EB64970-F159-4C6E-B9FF-AF9D09128553

The complete mitochondrial genome of Juema pig (Suina:Suidae)

Juema pig (Suina:Suidae), a rare, miniature and primitive species, has been living and razing on the Qinghai-Tibet plateau completely wild for thousands of years. Owing to the cold plateau climate, low energy, and low carbohydrate diet, Juema pig has some unique genetic characteristics that differentiate it from other pig breeds. If the species should not be protected in a timely manner, it would become extinct in the future. In June 2006, No. 662 announcement of the People’s Republic ofChina has included the Tibetan pigs in the national protection list of poultry genetic resources. Developing reasonable conservation and utilization measures for Juema pig requires deeper understanding of molecular genetics. The complete mitochondrial genome of Juema pig was determined in this study. Similar to other vertebrate, the mitogenome is 16,690 bp in length and contains 13 protein-coding genes, 22 transfer RNA genes, 2 ribosomal RNA genes and a D-loop region. The overall base composition of the H-strand is 34.76% for A, 26.17% for C, 13.24% for G and 25.83% for T. To ensure the sequence is reliable, phylogenetic tree was constructed using MEGA 6 with Neighbor-Joiningmethod (NJ) methods, demonstrating that Juema pig was clustered in subfamily Suina.

Evolutionary Relationships and Cytotaxonomy Considerations in the Genus Pyrrhulina (Characiformes, Lebiasinidae).

Although fishes exhibit the greatest biodiversity among the vertebrates, a large percentage of this fauna is still underexplored on evolutionary cytogenetic questions, particularly the miniature species. The Lebiasinidae family is a particular example for such case. This study is the first one presenting differential cytogenetic methods, such as C-banding, repetitive DNAs mapping, comparative genomic hybridization (CGH), and whole chromosome painting in lebiasinid species. Pyrrhulina australis and Pyrrhulina aff. australis were deeply investigated concerning their chromosomal patterns and evolutionary relationships. These species have a very similar morphology, but they can be distinguished by a longitudinal midlateral faintly dark stripe exclusive for Pyrrhulina aff. australis. Both species presented 2n = 40 chromosomes (4st +36a), without heteromorphic sex chromosomes. However, despite their morphological and karyotype resemblance, it was evidenced that both species have already gone through a significant genomic divergence, thus corresponding to distinct evolutionary units. Furthermore, to give additional support to some proposals on evolutionary relationship among Lebiasinidae with other fish families, a chromosomal comparative approach with Erythrinus erythrinus, a representative species of the Erythrinidae family, was also performed. In addition to have similar karyotype structure, mainly composed by acrocentric chromosomes, both species share uncommon genomic similarities, such as (i) syntenic location of 5S and 18S rDNA sequences; (ii) huge dispersion of multiple 5S rDNA sites in the karyotypes; and (iii) complex association between 5S rDNA and Rex3 elements. CGH experiments, despite reinforcing some shared genomic homologies, also highlighted that both Pyrrhulina and Erythrinus have a range of nonoverlapping species-specific signals. The overall chromosomal data proved to be effective markers for the cytotaxonomy and evolutionary process among Lebiasinidae fishes.

A new species of Hoplomyzon (Siluriformes: Aspredinidae) from Maracaibo Basin, Venezuela: osteological description using high-resolution computed microtomography of a miniature species

ABSTRACT A new miniature species of banjo catfish of the genus Hoplomyzon is described from the Lake Maracaibo Basin in Venezuela. The new species is distinguished from all its congeners by the straight anterior margin of the mesethmoid (vs. a medial notch); a smooth and straight ventral surface of the premaxilla (vs. presence of bony knobs on the ventral surface of premaxilla); absence of teeth on dentary (vs. teeth present on dentary); configuration of ventral vertebral processes anterior to anal fin, which are composed of single processes anterior to anal-fin pterygiophore (vs. paired process); presence of several filamentous barbel-like structures on the ventral surface of head of adults (vs. small papillous structures in the ventral surface of head of adults); and 8 anal-fin rays (vs. 6 or 7). An extensive osteological description is made of the holotype using high-resolution x-ray computed microtomography (HRXCT).

Inferring Evolution of Habitat Usage and Body Size in Endangered, Seasonal Cynopoeciline Killifishes from the South American Atlantic Forest through an Integrative Approach (Cyprinodontiformes: Rivulidae).

Cynopoecilines comprise a diversified clade of small killifishes occurring in the Atlantic Forest, one of the most endangered biodiversity hotspots in the world. They are found in temporary pools of savannah-like and dense forest habitats, and most of them are highly threatened with extinction if not already extinct. The greatest gap in our knowledge of cynopoecilines stems from the absence of an integrative approach incorporating molecular phylogenetic data of species still found in their habitats with phylogenetic data taken from the rare and possibly extinct species without accessible molecular information. An integrative analysis combining 115 morphological characters with a multigene dataset of 2,108 bp comprising three nuclear loci (GLYT1, ENC1, Rho), provided a robust phylogeny of cynopoeciline killifishes, which was herein used to attain an accurate phylogenetic placement of nearly extinct species. The analysis indicates that the most recent common ancestor of the Cynopoecilini lived in open vegetation habitats of the Atlantic Forest of eastern Brazil and was a miniature species, reaching between 25 and 28 mm of standard length. The rare cases of cynopoecilines specialized in inhabiting pools within dense forests are interpreted as derived from four independent evolutionary events. Shifts in habitat usage and biogeographic patterns are tentatively associated to Cenozoic paleogeographic events, but the evolutionary history of cynopoecilines may be partially lost by a combination of poor past sampling and recent habitat decline. A sharp evolutionary shift directed to increased body size in a clade encompassing the genera Campellolebias and Cynopoecilus may be related to a parallel acquisition of an internally-fertilizing reproductive strategy, unique among aplocheiloid killifishes. This study reinforces the importance of adding morphological information to molecular databases as a tool to understand the biological complexity of organisms under intense pressure from loss of habitat.

Laimosemion leticia (Teleostei: Cyprinodontiformes: Rivulidae), a new species from the upper Rio Amazonas, Southern Colombia

Laimosemion leticia is described from the Rio Tacana drainage, upper Rio Amazonas basin, Southern Colombia. The new species was found in a shallow swamp within a tropical rainforest in the vicinity of Leticia, capital of the department of Amazonas and southernmost municipality of Colombia. This miniature species is considered to be a member of the Owiyeye subgenus, which is diagnosed by a unique frontal squamation. Laimosemion leticia can be distinguished from all other species of the genus by the unique colour pattern on the unpaired fins. The new species appears related to L. jauaperi, L. uatuman and L. ubim due to the similar flank colour pattern in males, but is easy distinguished by coloration of the unpaired fins and morphological characters.

First record of the endemic Neotropical genus Hyphydrophilus (Chilopoda: Geophilomorpha, Geophilidae) from Argentina, with description of H. minellii sp. nov., a new centipede with small tentorial process, inhabiting the Yungas

ABSTRACTA new miniature species of the endemic Neotropical genus Hyphydrophilus (Chilopoda: Geophilomorpha, Geophilidae) is described and illustrated, after the holotype female and paratype female, from the Yungas rainforest of Salta province, northwestern Argentina. H. minellii sp. nov. is compared in detail with the other two species currently assigned to the taxon, i.e. H. adisi Pereira, Minelli & Barbieri, 1994 and H. projectus Pereira, Foddai & Minelli, 2000, both from Brazil, from which it differs by the presence of a small slightly sclerotized and inwards directed process of internal limbs of tentorium. Other unique traits of the new taxon are the relatively high number of leg-bearing segments (59), antennal articles I–IV with numerous large setae on the latero-external side, apical medial edge of forcipular trochanteroprefemur with a well-developed slightly pigmented tooth, dorsal and ventral medial edges of the forcipular tarsungulum slightly serrate, sternal pore-fields present on anterior region of the trunk only. This is the first report of the genus Hyphydrophilus from Argentina and the Yungas forests; previously it was only known to occur in Central Brazilian Amazonia, thus the present record represents a remarkable extension of the known range of this taxon.

Osteology of Priocharax and remarkable developmental truncation in a miniature Amazonian fish (Teleostei: Characiformes: Characidae).

Establishing phylogenetic relationships of miniature fishes is challenging in taxa with developmental truncation. Within the Characiformes, developmental truncation appears to be relatively rare, with the Neotropical genus Priocharax being an example. Priocharax includes three miniature species among the smallest of the order and has been hypothesized to belong to the Heterocharacinae. The pronounced reduction in its skeleton, however, prevented a clearer evaluation of its relationships. The present detailed osteological study was designed to address this question and revealed that 21 bones are absent and nine other skeletal structures are simplified in Priocharax when compared to other characids. Comparison of the skeleton of adult Priocharax with early developmental stages of other characids demonstrated that most of the absences and simplifications can be interpreted as developmental truncations. The most striking developmental truncations are in the pectoral girdle, in which the endoskeleton remains entirely cartilaginous. Other interesting truncations are in the ethmoid region of the skull, infraorbital series, and Weberian apparatus, in which the claustrum is absent. Our study also revealed some unusual sexual dimorphisms in the pelvic girdle. Two cladistic analyses were performed to assess the relationships of Priocharax within the Heterocharacinae. The first consisted of a traditional analysis in which all absences and reductions of Priocharax were coded in the same way as in the remaining taxa. This resulted in three equally most parsimonious topologies, all of which have Priocharax as the most basal taxon of the Heterocharacinae. The second analysis incorporated ontogenetic information, and most absences and reductions of Priocharax were reinterpreted as apomorphic conditions and thus, coded differently from similar conditions in outgroups. This resulted in a single phylogenetic hypothesis with Priocharax and Gnathocharax as sister groups based on seven synapomorphies. Our approach demonstrates the importance of developmental studies to better understand morphological evolution of miniaturized, truncated taxa, and to generate hypotheses of their relationships.

Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores)

The first cladistic analysis of the Tricommatinae is presented here based on a matrix with 88 terminals and 117 characters. The subfamily, hitherto with 28 genera and 53 species, is not recovered as a clade, instead it is composed of two nonsister clades, corresponding to the two main physiognomies of Tricommatinae sensu lato: (1) most of the diversity of the Tricommatinae, including the emblematic genera Cryptogeobius Mello-Leitão, 1935, and Pseudopachylus Roewer, 1912, and (2) the less familiar species, including the type genus Tricommatus Roewer, 1912, plus a few relatives, totalling three genera with five species. The Tricommatinae sensu stricto are here reduced to their bare core, nested within Gonyleptidae, whereas Cryptogeobiidae fam. nov. is more basally placed within the Gonyleptoidea. The holotype and sole available material of Tricommatus brasiliensis Roewer, 1912, type species of Tricommatus, is restudied and is here redescribed and illustrated. Pherania Strand, 1942, is herein considered a junior subjective synonym of Tricommatus and a key is given to the species of Tricommatus. Voriax gen. nov. of Tricommatinae is described along with the type species Voriax popeye sp. nov., an unusual species from Bahia, showing strange sexual dimorphism. Some changes have been introduced in the taxonomy of Cryptogeobiidae: (1) 13 genera are synonymized (two of them, Berlesecaptus Mello-Leitão, 1940, and Simonoleptes, originally in Phalangodidae); (2) 18 new combinations are made; (3) a neotype is designated for the type species Berlesecaptus convexus Mello-Leitão, 1940; (4) a new miniature species, Paratricommatus lockei sp. nov., is described from an area of conservation, Reserva Ecológica de Guapi-Açu, in Rio de Janeiro state, south-eastern Brazil. © 2014 The Linnean Society of London

Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores)

The first cladistic analysis of the Tricommatinae is presented here based on a matrix with 88 terminals and 117 characters. The subfamily, hitherto with 28 genera and 53 species, is not recovered as a clade, instead it is composed of two nonsister clades, corresponding to the two main physiognomies of Tricommatinae sensu lato: (1) most of the diversity of the Tricommatinae, including the emblematic genera Cryptogeobius Mello-Leitão, 1935, and Pseudopachylus Roewer, 1912, and (2) the less familiar species, including the type genus Tricommatus Roewer, 1912, plus a few relatives, totalling three genera with five species. The Tricommatinae sensu stricto are here reduced to their bare core, nested within Gonyleptidae, whereas Cryptogeobiidae fam. nov. is more basally placed within the Gonyleptoidea. The holotype and sole available material of Tricommatus brasiliensis Roewer, 1912, type species of Tricommatus, is restudied and is here redescribed and illustrated. Pherania Strand, 1942, is herein considered a junior subjective synonym of Tricommatus and a key is given to the species of Tricommatus. Voriax gen. nov. of Tricommatinae is described along with the type species Voriax popeye sp. nov., an unusual species from Bahia, showing strange sexual dimorphism. Some changes have been introduced in the taxonomy of Cryptogeobiidae: (1) 13 genera are synonymized (two of them, Berlesecaptus Mello-Leitão, 1940, and Simonoleptes, originally in Phalangodidae); (2) 18 new combinations are made; (3) a neotype is designated for the type species Berlesecaptus convexus Mello-Leitão, 1940; (4) a new miniature species, Paratricommatus lockei sp. nov., is described from an area of conservation, Reserva Ecológica de Guapi-Açu, in Rio de Janeiro state, south-eastern Brazil. © 2014 The Linnean Society of London

Priocharax nanus, a new miniature characid from the rio Negro, Amazon basin (Ostariophysi: Characiformes), with an updated list of miniature Neotropical freshwater fishes

Priocharax nanus, new species, is described from the rio Negro, Brazil. It is a miniature fish that retains as an adult the larval rayless pectoral fin, a diagnostic character of the genus. Priocharax nanus possesses fewer reductive features compared to congeners, P. ariel and P. pygmaeus, from which it can be distinguished by the presence of i,6 pelvic-fin rays (vs. i,5), the presence of the claustrum (vs. claustrum absent) and the presence of two postcleithra (vs. postcleithra absent). An updated list of 213 species of miniature Neotropical freshwater fishes is presented. The greatest diversity among them is represented by the Characiformes with 87 miniature species.

A new genus of miniature cynolebiasine from the Atlantic Forest and alternative biogeographical explanations for seasonal killifish distribution patterns in South America (Cyprinodontiformes: Rivulidae)

The analysis of 78 morphological characters for 16 species representing all the lineages of the tribe Cynopoecilini and three out-groups, indicates that the incertae sedis miniature species ‘Leptolebias’ leitaoi Cruz & Peixoto is the sister group of a clade comprising the genera Leptolebias, Campellolebias, and Cynopoecilus, consequently recognised as the only member of a new genus. Mucurilebias gen. nov. is diagnosed by seven autapomorphies: eye occupying great part of head side, low number of caudal-fin rays (21), distal portion of epural much broader than distal portion of parhypural, an oblique red bar through opercle in both sexes, isthmus bright red in males, a white stripe on the distal margin of the dorsal fin in males, and a red stripe on the distal margin of the anal fin in males. Mucurilebias leitaoi is an endangered seasonal species endemic to the Mucuri river basin. The biogeographical analysis of genera of the subfamily Cynolebiasinae using a dispersal-vicariance, event-based parsimony approach indicates that distribution of South American killifishes may be broadly shaped by dispersal events. The analysis supports south-eastern Brazilian coastal plains as the centre of dispersal for cynolebiasine killifishes. Alternative biogeographical explanations are compared and possible dispersal routes and means of dispersal are discussed.

First report of geophilid centipedes of the genus Ribautia (Myriapoda: Chilopoda: Geophilomorpha) from the Atlantic Forest biome, with description of a new miniature species from Misiones Province, Northeastern Argentina.

Ribautia paranaensis sp. nov. a new miniature species of geophilid centipede from the Upper Paraná Atlantic Forest (the westernmost of the fifteen ecoregions comprising the Atlantic Forest biome sensu Di Bitetti et al. 2003), is herein described and illustrated. The new species is characterized by having the coxal organs grouped in clusters (one of these in each coxopleuron of the ultimate leg-bearing segment) and a claw-like pretarsus in the ultimate legs; these traits being shared by three other Neotropical members of the genus, i.e., R. combinata Pereira, Uliana & Minelli, 2006 (from the Amazonian rainforest of Peru), R. jakulicai Pereira, 2007 (from the Yungas rainforest of Northwestern Argentina), and R. lewisi Pereira, 2013 (collected in a gallery forest in the Mesopotamian region, Northeastern Argentina). R. paranaensis sp. nov. represents the first report of Ribautia Brölemann, 1909 in the entire Atlantic Forest biome, and the third confirmed record of the taxon from Argentina.

Miniaturization in Otothyris Myers, 1927 (Loricariidae: Hypoptopomatinae)

Many species of freshwater fishes are considered miniatures, nevertheless, their identification as miniatures is frequently based only on an arbitrary criterion of small body size. Although some species of Otothyris had been suggested to be miniature, informations about the process and consequences of miniaturization in the genus are lacking. Here, we detail developmental events of loss, fusion, and modifications in several bones, the laterosensory system, and even the brain, that in sum demonstrate miniaturization in all species of the genus. Our results may be useful in the recognition of other miniature species of loricariids.

Study on Activated Sludge Microorganisms of Northern Winter Sewage Treatment Plant

At the sewage temperatures of 7-15°C, it used biological video microscopy and scanning electron microscopy to detect activated sludge morphology, miniature animals and flora species in northern winter sewage treatment plant. The results showed that activated sludge had dense floc with sharp edge, and was bigger at the temperatures of 13 and 15°C than at the temperatures of 7, 9 and 11°C. When the autumn and winter seasons were alternate and the sewage temperature was instability, the edge of the floc was slightly fuzzy and there was a small amount of free bacteria in activated sludge floc. The study had also shown that the temperature was higher, the type of sludge miniature animal and flora species were richer.

Are diminutive turtles miniaturized? The ontogeny of plastron shape in emydine turtles

Miniaturization, or the evolution of a dramatically reduced body size compared to related lineages, is an extraordinarily widespread phenomenon among metazoans. Evolutionary biologists have been fascinated by miniaturization because this transition has occurred numerous times, often among close relatives, providing a model system for studying convergent evolution and its underlying mechanisms. Much of the developmental work describing the ontogeny of miniature species suggests that paedomorphosis is the predominant avenue of miniaturization. Nevertheless, specific alterations to ontogeny appear highly variable, so that even related lineages with similar miniaturized traits produce those similarities via distinct ontogenetic paths. One major vertebrate group that has been overlooked in research on miniaturization is turtles. In the present study, we examined patterns of shape change in the plastron (the ventral part of the shell) over the course of ontogeny in a small clade of turtles (Emydinae) aiming to investigate whether two independently evolved diminutive members of the clade (Glyptemys muhlenbergii and Clemmys guttata) should be considered as miniaturized. We employ geometric morphometric methods to quantify the patterns of shape change these potentially miniaturized species and their relatives undergo during ontogeny, and use molecular phylogenetic trees to reconstruct ancestral conditions and provide information on the polarity of shape changes. We find that differing changes in ontogenetic parameters relative to ancestral conditions accompany the evolution of small size in emydines: G. muhlenbergii changes the duration of ontogeny and rate of shape change, whereas C. guttata changes growth rate. The observed ontogenetic repatterning of these species is reminiscent of changes in ontogeny and life history often found in miniaturized taxa. However, we conclude that C. guttata and G. muhlenbergii are not truly miniaturized because they still produce typical adult shell morphologies, and larger emydines display comparable ontogenetic flexibility. Because no emydines carry juvenile shell features forward into adulthood, we speculate that few, if any turtles, will show paedomorphic shell traits without corresponding changes in defensive strategy because such shells may offer insufficient protection.

New species of miniature fish from Marajó Island, Pará, Brazil, with comments on its relationships (Characiformes: Characidae)

A new miniature species of the family Characidae from Marajó Island, Pará State, Brazil is described and assigned to the genus Tyttobrycon. The new species can be distinguished from its congeners primarily by having multicuspid teeth on jaws, and additionally by having the combination of 5-7 premaxillary teeth, dorsal fin only with scattered melanophores and the presence of an adipose fin.

Repeated trans-watershed hybridization among haplochromine cichlids (Cichlidae) was triggered by Neogene landscape evolution

The megadiverse haplochromine cichlid radiations of the East African lakes, famous examples of explosive speciation and adaptive radiation, are according to recent studies, introgressed by different riverine lineages. This study is based on the first comprehensive mitochondrial and nuclear DNA dataset from extensive sampling of riverine haplochromine cichlids. It includes species from the lower River Congo and Angolan (River Kwanza) drainages. Reconstruction of phylogenetic hypotheses revealed the paradox of clearly discordant phylogenetic signals. Closely related mtDNA haplotypes are distributed thousands of kilometres apart and across major African watersheds, whereas some neighbouring species carry drastically divergent mtDNA haplotypes. At shallow and deep phylogenetic layers, strong signals of hybridization are attributed to the complex Late Miocene/Early Pliocene palaeohistory of African rivers. Hybridization of multiple lineages across changing watersheds shaped each of the major haplochromine radiations in lakes Tanganyika, Victoria, Malawi and the Kalahari Palaeolakes, as well as a miniature species flock in the Congo basin (River Fwa). On the basis of our results, introgression occurred not only on a spatially restricted scale, but massively over almost the whole range of the haplochromine distribution. This provides an alternative view on the origin and exceptional high diversity of this enigmatic vertebrate group.