Search

In the welcome circumstance that species believed extinct are rediscovered, it is often the case that biological knowledge acquired before the presumed extinction is limited. Efforts to address these knowledge gaps, in particular to assess the taxonomic integrity and conservation status of such species, can be hampered by a lack of genetic data and scarcity of samples in museum collections. Here, we present a proof-of-concept case study based on a multidisciplinary data evaluation approach to tackle such problems. The approach was developed after the rediscovery, 40 years after its presumed extinction, of the enigmatic Lake Constance deep-water charr Salvelinus profundus. Targeted surveys led to the capture of further species and additional sympatric normal charr, Salvelinus cf. umbla. Since the lake had been subject to massive stocking in the past, an evaluation of the genetic integrity of both extant forms was called for in order to assess possible introgression. A two-step genomic approach was developed based on restriction site associated DNA (RAD). Diagnostic population genomic (single nucleotide polymorphism [SNP]) data were harvested from contemporary samples and used for RNA bait design to perform target capture in DNA libraries of archival scale material, enabling a comparison between extant and historic samples. Furthermore, life history traits and morphological data for both extant forms were gathered and compared with historical data from the past 60-120 years. While extant deep-water charr matched historical deep-water specimens in body shape, gill raker count, and growth rates, significant differences were discovered between historical and extant normal charr. These resulted were supported by genomic analyses of contemporary samples, revealing the two extant forms to be highly divergent. The results of population assignment tests suggest that the endemic deep-water charr persisted in Lake Constance during the eutrophic phase, but not one of the historical genomic samples could be assigned to the extant normal charr taxon. Stocking with non-endemic charr seems to be the most likely reason for these changes. This proof-of-concept study presents a multidisciplinary data evaluation approach that simultaneously tests population genomic integrity and addresses some of the conservation issues arising from rediscovery of a species characterized by limited data availability., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

A new species, Parakneria alytogrammus, is described from the main stream of the Upper Lufira River. This species is easily distinguished from its congeners from the Congo Basin by its unique colouration, consisting of a low number of transversal bands on each of the caudal-fin lobes, 2 (vs. 3-5) and the presence of an uninterrupted lateral mid-longitudinal black band in fresh and preserved specimens (vs. absent). In addition, the new species differs from its Upper Lualaba congeners by the narrow width of its pectoral-fin base, 4.8-5.6% L S [vs. wider, 8.2-10.1% for P. lufirae, 8.6% L S for P. damasi (holotype), and 7.6-7.9% L S for P. thysi]. Finally, it differs from the only species currently known from the Luapula-Mweru system, P. malaissei, by having a short post-dorsal distance, 36.4-36.6% L S (vs. longer, 38.6-41.1% L S ) and a short post-pelvic distance of 40.0-40.6% L S (vs. longer, 41.4-44.1% L S ). Mitochondrial DNA-haplotypes of P. alytogrammus sp. nov. form a clade, which is sister to the P. thysi clade, and from which it diverges by a genetic (Kimura 2-parameter and uncorrected p) distance of 0.7% in the COI-barcoding locus. The Upper Lufira, one of the sub-basins of the Upper Congo Basin, remains poorly explored relative to its fish fauna. In contrast, the region is well explored with regard to its mineral wealth. Unfortunately, mining exploitation is carried out in the region without proper concern for the environment. Thus, the discovery of this new species for science calls for increased protection and aquatic biodiversity exploration in this mining region., (© 2022 Fisheries Society of the British Isles.)

The small cyprinid genus Prolabeops Schultz, 1941 is restricted to the Nyong and Sanaga River systems in Cameroon. In the past, the genus had been suggested to be either a member of the Labeoninae, Torinae or the Smiliogastrinae mainly on the basis of morphological similarities, and it is nowadays considered as incertae sedis within the Cypriniformes. This study provides the first attempt to reveal the phylogenetic position of Prolabeops using molecular data. For this purpose, the authors sequenced a large fraction of the mitochondrial genome (c. 13,600 bp), including all mitochondrial protein coding genes, of two Prolabeops melanhypopterus specimens and an additional four Enteromius specimens. The large-scale phylogenetic analysis was based on an alignment including all mitochondrial protein coding genes of 902 specimens representing c. 899 cypriniform species. Prolabeops was clearly recovered within the African Smiliogastrinae, forming a weakly supported clade together with Enteromius jae, Enteromius hulstaerti and Barboides gracilis. The study data underline the urgent need of a thorough taxonomic revision of the small African barbs collectively placed in the genus Enteromius., (© 2022 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

An evidence-based annotated checklist of gobiid species (Teleostei: Gobiidae) inhabiting the South Caspian Sea and its catchment area (i.e., the South Caspian Sea sub-basin) is compiled. The South Caspian Sea sub-basin gobiofauna currently comprises 38 confirmed species in 11 genera (i.e., 88.4% of the Caspian gobiofauna); the most diverse genus is Benthophilus (16 species, 42.1%), followed by Ponticola (seven species, 18.4%), and Neogobius (four species, 10.5%). Ten species (26.3%) are endemic to the South Caspian Sea sub-basin, another 21 species (55.3%) are endemic in the Caspian Sea basin as a whole, six (15.8%) are native to the Ponto-Caspian region, and one species (2.6%) is exotic. According to the current IUCN Red List, 24 species (64.9%) are listed as being of "Least Concern", eight species (21.6%) are "Data Deficient", and five species (13.5%) as "Not Evaluated". Similar numbers of species are confirmed to inhabit the South Caspian Sea sub-basin waters of the three countries that border it: Iran harbors 25 species (nine genera), Azerbaijan has 28 species (10 genera), and Turkmenistan has 26 species (10 genera). The greatest known diversity of Benthophilus in South Caspian waters occurs in Azerbaijan and Turkmenistan (11 species each), whereas Iranian waters harbor seven species. In comparison, Iran, with six out of eight species (75%), has the greatest diversity of Ponticola known from the Caspian Sea basin. Species richness and endemism of the Caspian Sea gobiid-fauna varies considerably with latitude: the North, Middle and South sub-basins respectively harbor 21, 31, and 37 native species, of which 0, 3, and 10 species are endemic in that sub-basin alone. The high species diversity and endemism of Gobiidae in the South Caspian Sea sub-basin may have resulted from: (i) greater ecological diversity compared to the northern Caspian Sea marine areas (e.g., water depths) that may have led to differential niche adaptation and adaptive radiation in the Benthophilus-Anatirostrum species flock, (ii) lower historical extinction rate compared to Caspian higher latitudes, which had greater exposure to the Pleistocene's extreme climatic changes, (iii) geological history of freshwater habitats in the South Caspian Sea sub-basin that set the speciation and evolutionary stage for the genus Ponticola during these Pleistocene climatic oscillations, (iv) presently less limiting conditions compared to the North Caspian Sea, i.e., higher present winter minimum of water temperature and higher salinity, and (v) Iranian freshwater abundance, variability and habitat diversity. Contemporary gobiid diversity and endemism in the Caspian Sea basin suggests two higher-priority conservation areas: (i) freshwater habitats of the South Caspian Sea region in Iran and Azerbaijan, and (ii) shallow coastal and deep waters of the South and Middle Caspian Sea sub-basins. An identification key is provided for the updated gobiid species from the South Caspian Sea sub-basin.

Freshwater habitats of the Caucasus biodiversity hotspot represent a center of endemism for the gobiid genus Ponticola Iljin, 1927. Hitherto, large-scale molecular studies, owing to restricted taxon and geographical sampling, have failed to give an elaborate picture of diversity and evolutionary history of these species. Here, to contribute to filling this gap, we assessed taxonomic diversity, phylogeography and evolutionary history for the south Caspian populations of Ponticola presently classified as P. iranicus and P. patimari , using an integrative taxonomic approach comprising an entire geographic range sampling, and analyses of mitochondrial DNA haplotypes, the head lateral line system, otolith shape, and meristic and morphometric variation. All freshwater samples of the P. syrman group belong to a monophyletic clade with two main subclades: a small subclade confined to the upper Sefidroud sub-basin including the type locality of P. iranicus and a large subclade with three geographically constrained haplogroups (Hg1, Hg2, and Hg3), comprising the rest of the distribution. Hg1 showed an eastern distribution including the type locality of P. patimari , while Hg2 and Hg3 are sister groups with central and western-central distributions, respectively. The freshwater clade diverged from P. syrman during the Tyurkyanian low stand (~150 m b.s.l. lasting ~0.1 Myr), while the divergence of P. iranicus and P. patimari and radiations within P. patimari took place during the Bakunian high stand (up to 50 m a.s.l. lasting ~378-480 kya). Species delimitation analyses indicated two distinct species, corresponding to each main subclade. Although the otolith shape and lateral line analyses did not reflect with phylogeographic pattern, PCA and DFA plots of meristic and morphometric data showed a clear separation of the two major subclades corresponding to P. iranicus and P. patimari , suggesting the presence of significant morphological variation meriting formal taxonomic recognition. Overall, our findings (i) reveal the presence of two freshwater endemic species in the P. syrman group , and pending further investigation, hypothesize the presence of a third cryptic species; (ii) revise and document a narrow distributional range and low diversity for P. iranicus , in contrast to a wider distributional range and high diversity for P. patimari ; (iii) suggest that the climatic oscillations of the Pleistocene were associated with the cladogenesis within the P. syrman group ; and (iv) allowed for the recognition of conservation units and proposition of management measures., Competing Interests: We declare no conflict of interest., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Ponticola hircaniaensis sp. nov. is described as a new gobiid species from the Kaboudval Stream, southern Caspian Sea basin. The new species is diagnosed among Caspian Sea basin Ponticola species by the following combination of characters: second dorsal-fin branched rays 1416, anal-fin branched rays 1012, scales in lateral series 5259; lower jaw slightly, if at all, prognathous; head and body yellowish brown showing a reticulate brown pattern on a yellow background, first dorsal fin with a marginal bright orangish-yellow band and a dark anterior spot, upper part of pectoral-fin base with a distinct dark brown stripe; length of third spine in first dorsal fin 13.418.3 % of standard length (SL), second dorsal-fin spine length 11.113.8 % SL, caudal peduncle length and depth 16.420.1 % and 11.112.8 % SL, respectively, head depth at nape 70.981.0 % of head length (HL), and at eye 52.566.0 % HL; sagittal otolith dorsal rim with a broad concavity in the middle, dorsal depression absent or indistinct, sulcus length/sulcus height and sulcus height/otolith height ratios 1.471.82 and 0.340.40, respectively. It is also characterised by a K2P nearest neighbour distance of 5% to P. kessleri in the mtDNA COI barcode region. Mitochondrial and nuclear DNA analyses suggested extensive hybridization between P. hircaniaensis sp. nov. and P. gorlap at Kaboudval, providing evidence for the first record of hybridization in the Ponto-Caspian gobiids. Based on narrow geographic range isolated above the Zarrin Gol Dam (< 2 km2), extensive hybridization with P. gorlap, and other threats, the new species should be considered Critically Endangered.

Several hundred catfish species (order: Siluriformes) belonging to 11 families inhabit Africa, of which at least six families are endemic to the continent. Although four of those families are well-known to belong to the 'Big-Africa clade', no previous study has addressed the phylogenetic placement of the endemic African catfish family Austroglanididae in a comprehensive framework with molecular data. Furthermore, interrelationships within the 'Big-Africa clade', including the most diverse family Mochokidae, remain unclear. This study was therefore designed to help reconstruct inter- and intrarelationships of all currently valid mochokid genera, to infer their position within the 'Big Africa clade' and to establish a first molecular phylogenetic hypothesis of the relationships of the enigmatic Austroglanididae within the Siluriformes. We assembled a comprehensive mitogenomic dataset comprising all protein coding genes and representing almost all recognized catfish families (N = 33 of 39) with carefully selected species (N = 239). We recovered the monophyly of the previously identified multifamily clades 'Big Asia' and 'Big Africa' and determined Austroglanididae to be closely related to Pangasiidae, Ictaluroidea and Ariidae. Mochokidae was recovered as the sister group to a clade encompassing Auchenoglanididae, Claroteidae, Malapteruridae and the African Schilbeidae, albeit with low statistical support. The two mochokid subfamilies Mochokinae and Chiloglanidinae as well as the chiloglanid tribe Atopochilini were recovered as reciprocally monophyletic. The genus Acanthocleithron forms the sister group of all remaining Mochokinae, although with low support. The genus Atopodontus is the sister group of all remaining Atopochilini. In contrast to morphological reconstructions, the monophyly of the genus Chiloglanis was strongly supported in our analysis, with Chiloglanis macropterus nested within a Chiloglanis sublineage encompassing only other taxa from the Congo drainage. This is an important result because the phylogenetic relationships of C. macropterus have been controversial in the past, and because we and other researchers assumed that this species would be resolved as sister to most or all other members of Chiloglanis. The apparent paraphyly of Synodontis with respect to Microsynodontis provided an additional surprise, with Synodontis punu turning out to be the sister group of the latter genus., (© 2022 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

A new gobiid species, Benthophilus persicus sp. nov., is described from the southern Caspian Sea, Iran. The new species is diagnosed by the following character states: dermal fold on cheek well-developed, large, rectangular; chin barbel 1/32/3 of eye diameter; maximum body width 15.122.9% of standard length; mouth width, 36.355.8% of head length; second dorsal fin I+78; origin of anal fin in front of vertical through origin of second dorsal fin; dermal tubercles present on body, clearly larger than granules, with two posterior rows of spinules forming an acute angle, always less than right angle; dorsal row of tubercles complete, 2229; ventral row of tubercles 2225; ventrolateral row of tubercles absent; tubercles not present on temporal and occipital head regions; granules not present on flanks; transversal suborbital row 6i below posterior end of row b; anterior interorbital transversal row pa with one or two papillae and anterior interorbital transversal papilla row pp with two or three papillae; body with 2022 transversal ltm rows starting anteriorly behind pectoral axilla and alternating anteriorly with three longitudinal llm rows.

Corcyrogobius pulcher sp. nov. is described from off Île de Ngor, Dakar, Senegal. Corcyrogobius pulcher is distinguished from its two congeners by having the rear edge of the jaws ending posteriorly below mideye, second dorsal fin I/9, pectoral fin rays 17, pelvic fins oval or truncated posteriorly, scales in lateral series 26-27, anterior oculoscapular head canal with pore β, suborbital row b of sensory papillae anteriorly beginning below vertical of posterior edge of eye, dark vertical caudal bar, branchiostegal membrane without intense dark spot, cheek with two oblique whitish stripes, the first going from the eye downwards and forward to the posterior jaws, the second on the preopercular, alternating with brown oblique stripe going from behind the eye downwards and forward across the cheek. Furthermore, mitochondrial COI-barcoding data unambiguously support the species-level distinctiveness of the three Corcyrogobius species. A key to the species of Corcyrogobius is provided.

Two monotypic haplochromine cichlid genera (Teleostei: Cichlidae) of the Pseudocrenilabrus group are described from northern Zambia. One new genus is Palaeoplex gen. nov., with Pa. palimpsest sp. nov. as the type species, from the Luongo and Kalungwishi Rivers (Upper Congo drainage, Luapula subdrainage). It is diagnosed by a unique combination of morphological characters: (1) a fully developed infraorbital series without a distinct gap between the lachrymal and second infraorbital bone, (2) fused hypuralia 1+2 and hypuralia 3+4, (3) molariform teeth on the sagittal series of the lower pharyngeal jaw, and (4) a large maximum size. The second new genus, Lufubuchromis gen. nov., with L. relictus sp. nov. as the type species, is restricted to the upper Lufubu River catchment (Upper Congo drainage, Lake Tanganyika subdrainage). It is diagnosed by a unique combination of morphological characters: (1) a fully developed infraorbital series without a distinct gap between the lachrymal and second infraorbital bone, (2) fused hypuralia 1+2 (rarely with a visible suture) and fused hypuralia 3+4, (3) a unique male coloration pattern, i.e. deep crimson red colored areas on the anterior ventral flank parts, chest and belly and on the lower head; remaining parts of flanks and caudal peduncle bluish), and (4) a Pseudocrenilabrus blotch present in both sexes. Both new genera are compared with all remaining taxa of the Pseudocrenilabrus group and with all representatives of all other major haplotilapiine lineages.

A new goby species, Lebetus patzneri sp. nov. (Teleostei: Gobiidae) is described based on two preserved and two photographed specimens collected from two circalittoral bottom locations off the Balearic Islands, western Mediterranean. It differs from the one or both other Lebetus species by each of following characters: P 15-16; D2 segmented fin rays 7-8; snout slightly larger than eye diameter; eyes small, 23.3-25.5% of head length; body height low, body depth at pelvic fin 14.9-16.4%; lips thick, giving mouth a broad, almost rectangular shape in dorsal and lateral view; minute black "pepper" spots on predorsal area, dorsum, flanks and caudal peduncle, visible in fresh and preserved specimens; the most of the caudal peduncle similarly colored as the lateral area below D2, reddish or brown. We further report first records of Lebetus guilleti (Le Danois, 1913) from Norway and the Balearic Islands, and observations of its conspicuous courtship behavior ("D1 waving"), and coloration including the first description of its red fluorescence pattern.

The process of adaptive radiation was classically hypothesized to require isolation of a lineage from its source (no gene flow) and from related species (no competition). Alternatively, hybridization between species may generate genetic variation that facilitates adaptive radiation. Here we study haplochromine cichlid assemblages in two African Great Lakes to test these hypotheses. Greater biotic isolation (fewer lineages) predicts fewer constraints by competition and hence more ecological opportunity in Lake Bangweulu, whereas opportunity for hybridization predicts increased genetic potential in Lake Mweru. In Lake Bangweulu, we find no evidence for hybridization but also no adaptive radiation. We show that the Bangweulu lineages also colonized Lake Mweru, where they hybridized with Congolese lineages and then underwent multiple adaptive radiations that are strikingly complementary in ecology and morphology. Our data suggest that the presence of several related lineages does not necessarily prevent adaptive radiation, although it constrains the trajectories of morphological diversification. It might instead facilitate adaptive radiation when hybridization generates genetic variation, without which radiation may start much later, progress more slowly or never occur.

A new genus and species of goby, Gymnesigobius medits sp. nov., is described from the western Mediterranean slope bottoms at the Balearic Islands. The new goby belongs to the Gobius-lineage (Gobiinae). Examination of the single known specimen exhibits a unique combination of morphological characters which could not be fitted to any known genus. Gymnesigobius gen. nov. is morphologically distinguished from all other genera in the Gobius-lineage by the following combination of characters: chin without fold or barbels; mouth terminal with anterior tip above horizontal level of lower eye edge; predorsal area and first dorsal fin base naked; pelvic fin anterior membrane well developed; head with anterior oculoscapular and preopercular canals, posterior oculoscapular canal absent; pores of head canals enlarged, e.g. pores α and ρ larger or of about the same size as interspaces to pore β; six transverse suborbital rows of sensory papillae, four continuous suborbital rows in front of row b, fifth row divided in three parts but in front of row b, sixth row just as superior part above row b and below pore α; longitudinal suborbital row b barely reaching forward to the vertical from posterior edge of eye. A full description of the new genus and species is provided. The new species, collected at a depth of between 344 and 364 m (mean depth of 354 m), is one of very few gobiid species found at bathyal depths.

A new genus and species of fossil cichlid fishes of middle Miocene age (12.5 Ma) is described from the Ngorora fish Lagerstätte (Tugen Hills, Kenya) in the East African Rift Valley. Parsimony analysis of morphological characters using published phylogenetic frameworks for extant cichlids combined with the application of a comprehensive best-fit approach based on morphology was employed to place the new fossil taxon in the phylogenetic context of the African cichlids. The data reveal that the fossil specimens can be assigned to the tribe Oreochromini within the haplotilapiines. †Oreochromimos kabchorensis gen. et sp. nov. shows a mosaic set of characters bearing many similarities to the almost pan-African Oreochromis and the East African lake-endemic Alcolapia. As the striking diversity of present-day African cichlids, with 1100 recognised species, has remained largely invisible in the fossil record, the material described here adds significantly to our knowledge of the Miocene diversity of the group. It effectively doubles the age of a fossil calibration point, which has hitherto been used to calibrate divergence times of the East African cichlids in molecular phylogenetic investigations. Furthermore, the comparative dataset derived from extant cichlids presented here will greatly facilitate the classification of fossil cichlids in future studies.

Although the majority of cichlid diversity occurs in the African Great Lakes, these fish have also diversified across the African continent. Such continental radiations, occurring in both rivers and lakes have received far less attention than lacustrine radiations despite some members, such as the oreochromine cichlids (commonly referred to as 'tilapia'), having significant scientific and socio-economic importance both within and beyond their native range. Unique among cichlids, several species of the genus Oreochromis exhibit adaptation to soda conditions (including tolerance to elevated temperatures and salinity), which are of interest from evolutionary biology research and aquaculture perspectives. Questions remain regarding the factors facilitating the diversification of this group, which to date have not been addressed within a phylogenetic framework. Here we present the first comprehensive (32/37 described species) multi-marker molecular phylogeny of Oreochromis and closely related Alcolapia, based on mitochondrial (1583 bp) and nuclear (3092 bp) sequence data. We show widespread discordance between nuclear DNA and mitochondrial DNA trees. This could be the result of incomplete lineage sorting and/or introgression in mitochondrial loci, although we did not find a strong signal for the latter. Based on our nuclear phylogeny we demonstrate that adaptation to adverse conditions (elevated salinity, temperature, or alkalinity) has occurred multiple times within Oreochromis, but that adaptation to extreme (soda) conditions (high salinity, temperature, and alkalinity) has likely arisen once in the lineage leading to O. amphimelas and Alcolapia. We also show Alcolapia is nested within Oreochromis, which is in agreement with previous studies, and here revise the taxonomy to synonymise the genus in Oreochromis, retaining the designation as subgenus Oreochromis (Alcolapia)., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Phylogenetic studies focusing on Salmonidae have revealed significant obstacles in trying to clarify some interspecific relationships within the Salmoninae subfamily, due to a limited number of markers typed, conflicting phylogenetic signals and ancient hybridization events. To infer reliable phylogenetic relationships, evaluate several putative scenarios of ancient hybridization, and estimate divergence times within Salmoninae, we applied restriction-site associated DNA sequencing (RAD-seq) to 43 samples, including 26 genetic lineages across 21 species, largely representing the subfamily, with an emphasis on the genus Salvelinus. We identified 28,402 loci and 28,363 putatively unlinked SNPs, which were used in downstream analyses. Using an iterative k-means partitioned dataset and a Maximum Likelihood approach; we generated a well-supported phylogeny, providing clear answers to several previous phylogenetic uncertainties. We detected several significant introgression signals, presumably ancient, in the genus Salvelinus. The most recent common ancestor of Salmonidae dates back to approximately 58.9MY ago (50.8-64 MY) and the crown age of Salmoninae was estimated to be 37.7 MY (35.2-40.8 MY) using a Bayesian molecular dating analysis with a relaxed molecular clock. The divergence among genera of the subfamily occurred between the late Eocene and middle of the Miocene (≈38-11 MY) such as the divergence between the genus Oncorhynchus and Salvelinus, which we estimated to 21.2 MY ago (95% HPD: 19.8-23.0 MY), while species diversification took place mainly during the Neogene (≈22-1.5 MY), with more than half of these events occurring in the last 10 MY., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Didogobius janetarum sp. nov. is described from five specimens collected from small caves and rock crevices between 12 and 20 m depth off two locations of Santiago Island, Cape Verde Islands. The species differs from all currently described congeners by the combination of the following characters: (1) 27 vertebrae, (2) second dorsal fin I + 10, (3) posterior quarter of predorsal region in front of first dorsal fin origin scaled, with several rows of cycloid scales, (4) body squamation cycloid anteriorly and ctenoid posteriorly, (5) scales in the lateral series 30-32, (6) anterior oculoscapular canal present, (7) preopercular head canal absent, (8) suborbital row 7 each a single papilla, (9) suborbital rows 2 and 4 close to orbit, and by (10) branchiostegal membranes uniformely black below preopercle, forming a V-shaped mark. Definitions for all used meristic counts are presented to serve as a reference for gobioid meristic studies. The genus is rediagnosed to accommodate recently described Didogobius species.

A new gobiid species, Buenia lombartei sp. nov. (Teleostei: Gobiidae) is described from the continental slope off the Balearic Islands, western Mediterranean. The Atlantic species Buenia jeffreysii (Günther, 1867) is redescribed and the first record of Buenia affinis Iljin, 1930 is reported for the islands. A key for the species of genus Buenia is provided. The new species can be distinguished from congeneric species by morphological characters: anterior oculoscapular canal semi-closed with pores σ, λ, κ, α, ρ and additional pores and open furrows; suborbital row c of 6 papillae; anal fin I/7-I/8; scales in lateral series 25-27; scales in transverse series 6; pectoral fin rays 18; the second spine of the first dorsal fin longest, backwards reaching to middle second dorsal fin in males when folded down; pelvic fin anterior membrane reduced to less than one sixth of spinous ray in midline depth; tongue well developed and bilobed and several morphometric characters. The new species is also characterized by its reduced colouration and unique depth range and occurrence on upper slope muddy bottoms. Contrary to this, only a small number of gobiid species in the Mediterranean reach beyond the circalittoral to the deep shelf, and just a few have records below the shelf break. Phylogenetic analysis of their mitochondrial COI sequence grouped the two specimens within the same clade as B. jeffreysii, but one apomorphic transversion diagnoses the B. lombartei subclade. A hypothesis about rapid speciation of B. lombartei as well as alternative hypotheses are discussed. During the Last Glacial Maximum, boreal fish species such as cold-water adapted gadids, now restricted to northwestern Europe, entered the Mediterranean. This may have also been the case of the boreal population of B. jeffreysii. At the end of the glacial period, increasingly warming Mediterranean waters might restricted B. jeffreysii to deeper coldwater mud habitats of the upper slope, where it finally adapted to bathyal conditions.

Few studies have systematically investigated mid- or long-term temporal changes of biological characteristics in invasive alien species considering the different phases of an invasion. We studied the invasion performance of one of the most invasive species worldwide, the round goby Neogobius melanostomus, from total absence over first occurrence until establishment from 2010 to 2015 in the upper Danube River. After an upstream movement of the invasion front of about 30 river km within four years, the pattern that round goby pioneering populations significantly differ from longer established ones has been confirmed: Pioneering populations at the invasion front comprised more females than males, and adult specimens with a larger body size compared to those at longer inhabited areas. On the population-level, the proportion of juveniles increased with time since invasion. The results of this study provide support for the previously postulated ´bigger is better´ and ´individual trait utility´ hypotheses explaining invasion success in round goby. Pioneering invaders with their greater exploratory behavior, highly adaptive phenotypic plasticity and increased competitive ability seem to act as prime emperors of new habitats, strongly following and benefiting from man-made river-bank structures.

The relative importance of species-specific biological trait characteristics and environmental factors in invasions of nonindigenous species remains controversial because both have mostly been studied independently. Thus, the main objective of this study was to examine the correlation of biological traits with environmental variation in the globally invasive round goby Neogobius melanostomus from the upper Danube River. Based on a sample of 653 specimens along a continuous 200 km river pathway, links between nine environmental factors (substrate-type, six water measurements, and the communities of fishes and macroinvertebrates) and seven biological traits (nutritional and energetic status, trade-offs of parasite resistance and resource allocation, and three growth proxies) were analyzed. Biological trait values of N. melanostomus hardly correlated with the environment, could not explain invasion progress and imply a general low overall importance for invasion success. Instead, alternative individual life-history trajectories appear to determine invasion success. This is in line with up to 15% of all specimens having outlying biological trait values of potential adaptive value, suggesting a considerable importance of adaptive trait variation among single individuals for the whole invasion progress. This "individual trait utility hypothesis" gives an alternative explanation for success of invasive species by single individuals carrying particular traits, and it should be specifically targeted and analyzed at currently invaded sites.

A new miniature gobiid species, Buenia massutii sp. nov. (Teleostei: Gobiidae) is described from the circalittoral bottom off the Balearic Islands, western Mediterranean. Phylogenetic analysis of the mitochondrial COI-barcoding fragment supports its species-level distinctiveness and the monophyly of the genus Buenia. A description and diagnosis of the genus Buenia is for the first time provided. Material of the new species was collected in 57-67 m depth from beam trawl samples carried out on red algae beds. The traits that differentiate the new species from two congeneric species are: anterior oculoscapular canal only semiclosed with pores σ, λ, κ, ω, α, ρ and additional pores and open furrows from interorbital part to pore ρ; posterior oculoscapular canal absent; suborbital row c with 5 papillae; scales in lateral series 26-28; pectoral fin rays 16; the second spine of the first dorsal fin the longest, reaching to or behind posterior end of the second dorsal fin in males when folded down; pelvic fin anterior membrane one sixth or less of length of spinous ray in midline depth; head length 31.2-32.5% of standard length; eye 32.8-35.7% of head length; caudal fin length 21.5-24.0% of standard length.

Two new species of Bathyaethiops (Teleostei: Characiformes: Alestidae) are described. Bathyaethiops baka n. sp. is a dwarf species with the largest known specimen being only 24.4 mm SL. The species is characterized by an incomplete squamation and a large humeral spot. Bathyaethiops baka n. sp. is known so far only from the Ngoko River of Southeastern Cameroon, a tributary of the Sangha River in the northern Congo basin. The second species, Bathyaethiops flammeus n. sp., shows a diagnostic spot in front of the dorsal-fin base, which is devoid of melanophores and bright red in life. The species is described from the Bakéré River at Yambula-Bakéré, a locality north-west of Kisangani in the Central Congo basin. Other records of Bathyaethiops flammeus n. sp. from the Tshuapa respectively Ruki River at Boende and Eala, Central Congo basin, suggests a wider geographic distribution. A key to all species of Bathyaethiops is provided.

A new goby species, Speleogobius llorisi sp. nov. (Teleostei: Gobiidae) is described from the circalittoral sea bed at 46-69 m depth off the Balearic Islands, western Mediterranean. Six specimens, four females and two males, of the new species were collected from beam trawl samples carried out on the red algae beds off the south west coast of Mallorca Island. Among others, the main traits that differentiate the new species from the only congeneric species, S. trigloides, are the presence of preopercular head canal with pores γ, δ, ε vs. preopercular head canal absent; a longer snout, equal or longer than eye vs. shorter than eye; lower lip ending anteriorly slightly in front of upper lip vs. upper lip slightly protruding lower lip; scales in lateral series 28 or 29 vs. 26; scales in transverse series 6 vs.7-8. It also differs from S. trigloides in some non-overlapping morphometrics and in coloration. All individuals of the new species were collected from Peyssonnelia beds, beds of red algae dominated by species of the family Peyssonneliaceae. The generic diagnosis of Speleogobius is revised.

Strong disruptive ecological selection can initiate speciation, even in the absence of physical isolation of diverging populations. Species evolving under disruptive ecological selection are expected to be ecologically distinct but, at least initially, genetically weakly differentiated. Strong selection and the associated fitness advantages of narrowly adapted individuals, coupled with assortative mating, are predicted to overcome the homogenizing effects of gene flow. Theoretical plausibility is, however, contrasted by limited evidence for the existence of rugged adaptive landscapes in nature. We found evidence for multiple, disruptive ecological selection regimes that have promoted divergence in the sympatric, incipient radiation of 'sharpfin' sailfin silverside fishes in ancient Lake Matano (Sulawesi, Indonesia). Various modes of ecological specialization have led to adaptive morphological differences between the species, and differently adapted morphs display significant but incomplete reproductive isolation. Individual fitness and variation in morphological key characters show that disruptive selection shapes a rugged adaptive landscape in this small but complex incipient lake fish radiation., (© 2016 The Author(s).)

A long history of research focused on the East Africa cichlid radiations (EAR) revealed discrepancies between mtDNA and nuclear phylogenies, suggesting that interspecific hybridisation may have been significant during the radiation of these fishes. The approximately 250 cichlid species of Lake Tanganyika have their roots in a monophyletic African cichlid assemblage, but controversies remain about the precise phylogenetic origin and placement of different lineages and consequently about L. Tanganyika colonization scenarios. 3312 AFLP loci and the mitochondrial ND2 gene were genotyped for 91 species representing almost all major lacustrine and riverine haplotilapiine east African cichlid lineages with a focus on L. Tanganyika endemics. Explicitly testing for the possibility of ancient hybridisation events, a comprehensive phylogenetic network hypothesis is proposed for the origin and diversification of L. Tanganyika cichlids. Inference of discordant phylogenetic signal strongly suggests that the genomes of two endemic L. Tanganyika tribes, Eretmodini and Tropheini, are composed of an ancient mixture of riverine and lacustrine lineages. For the first time a strong monophyly signal of all non-haplochromine mouthbrooding species endemic to L. Tanganyika ("ancient mouthbrooders") was detected. Further, in the genomes of early diverging L. Tanganyika endemics Trematocarini, Bathybatini, Hemibatini and Boulengerochromis genetic components of other lineages belonging to the East African Radiation appear to be present. In combination with recent palaeo-geological results showing that tectonic activity in the L. Tanganyika region resulted in highly dynamic and heterogeneous landscape evolution over the Neogene and Pleistocene, the novel phylogenetic data render a single lacustrine basin as the geographical cradle of the endemic L. Tanganyika cichlid lineages unlikely. Instead a scenario of a pre-rift origin of several independent L. Tanganyika precursor lineages which diversified in ancient rivers and precursor lakes and then amalgamated in the extant L. Tanganyika basin is put forward as an alternative: the 'melting pot Tanganyika' hypothesis.

Background: The Malili Lakes system in central Sulawesi (Indonesia) is a hotspot of freshwater biodiversity in the Wallacea, characterized by endemic species flocks like the sailfin silversides (Teleostei: Atherinomorpha: Telmatherinidae) radiation. Phylogenetic reconstructions of these freshwater fishes have previously revealed two Lake Matano Telmatherina lineages (sharpfins and roundfins) forming an ancient monophyletic group, which is however masked by introgressive hybridization of sharpfins with riverine populations. The present study uses mitochondrial data, newly included taxa, and different external calibration points, to estimate the age of speciation and hybridization processes, and to test for phylogeographic relationships between Kalyptatherina from ancient islands off New Guinea, Marosatherina from SW Sulawesi, and the Malili Lakes flock., Results: Contrary to previous expectations, Kalyptatherina is the closest relative to the Malili Lakes Telmatherinidae, and Marosatherina is the sister to this clade. Palaeogeographic reconstructions of Sulawesi suggest that the closer relationship of the Malili Lakes radiation to Kalyptatherina might be explained by a 'terrane-rafting' scenario, while proto-Marosatherina might have colonized Sulawesi by marine dispersal. The most plausible analysis conducted here implies an age of c. 1.9 My for the onset of divergence between the two major clades endemic to Lake Matano. Diversification within both lineages is apparently considerably more recent (c. 1.0 My); stream haplotypes present in the sharpfins are of even more recent origin (c. 0.4 My)., Conclusions: Sulawesi's Telmatherinidae have most likely originated in the Sahul Shelf area, have possibly reached the island by both, marine dispersal and island/terrane-rafting, and have colonized the Malili Lakes system from rivers. Estimates for the split between the epibenthic sharpfins and the predominantly pelagic to benthopelagic roundfins in Lake Matano widely coincide with geological age estimates of this rift lake. Diversification within both clades clearly predates hybridization events with stream populations. For Lake Matano, these results support a scenario of initial benthic-pelagic divergence after colonization of the lake by riverine populations, followed by rapid radiation within both clades within the last 1 My. Secondary hybridization of stream populations with the sharpfins occurred more recently, and has thus most likely not contributed to the initial divergence of this benthic species flock.

The polytypic Nicaraguan Midas cichlids (Amphilophus cf. citrinellus) have been established as a model system for studying the mechanisms of speciation and patterns of diversification in allopatry and sympatry. The species assemblage in Crater Lake Apoyo has been accepted as a textbook example for sympatric speciation. Here, we present a first comprehensive data set of population genetic (mtDNA & AFLPs) proxies of species level differentiation for a representative set of individuals of all six endemic Amphilophus species occurring in Crater Lake Apoyo. AFLP genetic differentiation was partitioned into a neutral and non-neutral component based on outlier-loci detection approaches, and patterns of species divergence were explored with Bayesian clustering methods. Substantial levels of admixture between species were detected, indicating different levels of reproductive isolation between the six species. Analysis of neutral genetic variation revealed several A. zaliosus as being introgressed by an unknown contributor, hereby rendering the sympatrically evolving L. Apoyo flock polyphyletic. This is contrasted by the mtDNA analysis delivering a clear monophyly signal with Crater Lake Apoyo private haplotypes characterising all six described species, but also demonstrating different demographic histories as inferred from pairwise mismatch distributions.

Few studies have systematically investigated differences in performance, morphology and parasitic load of invaders at different stages of an invasion. This study analyzed phenotype-environment correlations in a fish invasion from initial absence until establishment in the headwater reach of the second largest European river, the Danube. Here, the round goby (Neogobius melanostomus) formed 73% of the fish abundance and 58% of the fish biomass in rip-rap bank habitats after establishment. The time from invasion until establishment was only about two years, indicating rapid expansion. Founder populations from the invasion front were different from longer established round goby populations in demography, morphology, feeding behaviour, sex ratio and parasitic load, indicating that plasticity in these traits determines invasion success. Competitive ability was mostly dependent on growth/size-related traits rather than on fecundity. As revealed by stable isotope analyses, specimens at the invasion front had a higher trophic position in the food web and seem to benefit from lower food competition. Somatic performance seems to be more important than investment in reproduction during the early stages of the invasion process and upstream-directed range expansion is not caused by out-migrating weak or juvenile individuals that were forced to leave high density areas due to high competition. This mechanism might be true for downstream introductions via drift. Greater abundance and densities of acanthocephalan endoparasites were observed at the invasion front, which contradicts the expectation that invasion success is determined by lower parasitic pressure in newly invaded areas. Overall, the pronounced changes in fish and invertebrate communities with a dominance of alien species suggest invasional meltdown and a shift of the upper Danube River towards a novel ecosystem with species that have greater resistance to goby predation. This seems to contribute to overcoming biological resistance and improve rapidity of dispersal.

African cichlids formerly referred to as "Tilapia" represent a paraphyletic species assemblage belonging to the so called haplotilapiine lineage which gave rise to the spectacular East African cichlid radiations (EARs) as well as to globally important aquaculture species. We present a comprehensive molecular phylogeny of representative haplotilapiine cichlids, combining in one data set four mitochondrial and five nuclear loci for 76 species, and compare it with phylogenetic information of a second data set of 378 mitochondrial ND2 haplotypes representing almost all important "Tilapia" or Tilapia-related lineages as well as most EAR lineages. The monophyly of haplotilapiines is supported, as is the nested sister group relationship of Etia and mouthbrooding tilapiines with the remaining haplotilapiines. The latter are consistently placed in eight monophyletic clades over all datasets and analyses, but several dichotomous phylogenetic relationships appear compromised by cytonuclear discordant phylogenetic signal. Based on these results as well as on extensive morphological evidence we propose a novel generic and suprageneric classification including a (re-)diagnosis of 20 haplotilapiine cichlid genera and nine tribes. New tribes are provided for the former subgenera Coptodon Gervais, 1853, HeterotilapiaRegan, 1920 and PelmatolapiaThys van den Audenaerde, 1969, in addition for "Tilapia" joka, Tilapia sensu stricto and Chilochromis, Etia, Steatocranus sensu stricto, the mouthbrooding tilapiines and for a clade of West African tilapiines., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Hybridization between invasive and native species accounts among the major and pernicious threats to biodiversity. The Mozambique tilapia Oreochromis mossambicus, a widely used freshwater aquaculture species, is especially imperiled by this phenomenon since it is recognized by the IUCN as an endangered taxon due to genetic admixture with O. niloticus an invasive congeneric species. The Lower Limpopo and the intermittent Changane River (Mozambique) drain large wetlands of potentially great importance for conservation of O. mossambicus, but their populations have remained unstudied until today. Therefore we aimed (1) to estimate the autochthonous diversity and population structure among genetically pure O. mossambicus populations to provide a baseline for the conservation genetics of this endangered species, (2) to quantify and describe genetic variation of the invasive populations and investigate the most likely factors influencing their spread, (3) to identify O. mossambicus populations unaffected by hybridization. Bayesian assignment tests based on 423 AFLP loci and the distribution of 36 species-specific mitochondrial haplotypes both indicate a low frequency of invasive and hybrid genotypes throughout the system, but nevertheless reveal evidence for limited expansion of two alien species (O. niloticus and O. andersonii) and their hybrids in the Lower Limpopo. O. mossambicus populations with no traces of hybridization are identified. They exhibit a significant genetic structure. This contrasts with previously published estimates and provides rather promising auspices for the conservation of O. mossambicus. Especially, parts of the Upper Changane drainage and surrounding wetlands are identified as refugial zones for O. mossambicus populations. They should therefore receive high conservation priority and could represent valuable candidates for the development of aquaculture strains based on local genetic resources.

Three new species of the lampeye genus Hylopanchax are described from the central Congo basin: H. leki, new species, H. ndeko, new species, and H. moke, new species. These differ considerably in body shape from the two previously de- scribed species, H. stictopleuron and H. silvestris, with two deep bodied and one small and slender species. A redefinition of the diagnostic characters of the genus Hylopanchax is presented, including pronounced sexual dimorphism.

Most freshwater diversity is arguably located in networks of rivers and streams, but, in contrast to lacustrine systems riverine radiations, are largely understudied. The extensive rapids of the lower Congo River is one of the few river stretches inhabited by a locally endemic cichlid species flock as well as several species pairs, for which we provide evidence that they have radiated in situ. We use more that 2,000 AFLP markers as well as multilocus sequence datasets to reconstruct their origin, phylogenetic history, as well as the timing of colonization and speciation of two Lower Congo cichlid genera, Steatocranus and Nanochromis. Based on a representative taxon sampling and well resolved phylogenetic hypotheses we demonstrate that a high level of riverine diversity originated in the lower Congo within about 5 mya, which is concordant with age estimates for the hydrological origin of the modern lower Congo River. A spatial genetic structure is present in all widely distributed lineages corresponding to a trisection of the lower Congo River into major biogeographic areas, each with locally endemic species assemblages. With the present study, we provide a phylogenetic framework for a complex system that may serve as a link between African riverine cichlid diversity and the megadiverse cichlid radiations of the East African lakes. Beyond this we give for the first time a biologically estimated age for the origin of the lower Congo River rapids, one of the most extreme freshwater habitats on earth.

Deep-sea Lantern Sharks (Etmopteridae) represent the most speciose family within Dogfish Sharks (Squaliformes). We compiled an extensive DNA dataset to estimate the first molecular phylogeny of the family and to provide node age estimates for the origin and diversification for this enigmatic group. Phylogenetic inferences yielded consistent and well supported hypotheses based on 4685bp of both nuclear (RAG1) and mitochondrial genes (COI, 12S-partial 16S, tRNAVal and tRNAPhe). The monophyletic family Etmopteridae originated in the early Paleocene around the C/T boundary, and split further into four morphologically distinct lineages supporting three of the four extant genera. The exception is Etmopterus which is paraphyletic with respect to Miroscyllium. Subsequent rapid radiation within Etmopterus in the Oligocene/early Miocene was accompanied by divergent evolution of bioluminescent flank markings which morphologically characterize the four lineages. Higher squaliform interrelationships could not be satisfactorily identified, but convergent evolution of bioluminescence in Dalatiidae and Etmopteridae is supported., (Copyright 2010 Elsevier Inc. All rights reserved.)

Nicaraguan Midas cichlids from crater lakes have recently attracted attention as potential model systems for speciation research, but no attempt has been made to comprehensively reconstruct phylogenetic relationships of this highly diverse and recently evolved species complex. We present a first AFLP (2793 loci) and mtDNA based phylogenetic hypothesis including all described and several undescribed species from six crater lakes (Apoyeque, Apoyo, Asososca Leon, Masaya, Tiscapa and Xiloá), the two great Lakes Managua and Nicaragua and the San Juan River. Our analyses demonstrate that the relationships between the Midas cichlid members are complex, and that phylogenetic information from different markers and methods do not always yield congruent results. Nevertheless, monophyly support for crater lake assemblages from Lakes Apoyeque, Apoyo, A. Leon is high as compared to those from L. Xiloá indicating occurrence of sympatric speciation. Further, we demonstrate that a 'three species' concept for the Midas cichlid complex is inapplicable and consequently that an individualized and voucher based approach in speciation research of the Midas cichlid complex is necessary at least as long as there is no comprehensive revision of the species complex available., (Copyright 2010 Elsevier Inc. All rights reserved.)

Background: For decades cichlid fishes (Perciformes: Cichlidae) of the East African cichlid radiations (Teleostei: Cichlidae) have served as natural experimental subjects for the study of speciation processes and the search for potential speciation key factors. Despite numerous phylogenetic studies dealing with their intragroup relationships, surprisingly little is known about the phylogenetic placement and time of origin of this enigmatic group. We used multilocus DNA-sequence data from five nuclear and four mitochondrial genes and refined divergence time estimates to fill this knowledge gap., Results: In concordance with previous studies, the root of the East African cichlid radiations is nested within the so called "Tilapias", which is a paraphyletic assemblage. For the first time, we clarified tilapiine intragroup relationships and established three new monophyletic groups:"Oreochromini", "Boreotilapiini" and a group with a distribution center in East/Central Africa, the "Austrotilapiini". The latter is the founder lineage of the East African radiations and emerged at the Miocene/Oligocene boundary at about 14 to 26 mya., Conclusion: Our results provide the first resolved hypothesis for the phylogenetic placement of the megadiverse East African cichlid radiations as well as for the world's second most important aquaculture species, the Nile Tilapia, Oreochromis niloticus. Our analyses constitute not only a robust basis for African cichlid phylogenetics and systematics, but provide a valid and necessary framework for upcoming comparative phylogenomic studies in evolutionary biology and aquaculture.

The Haplochromini are by far the most species-rich cichlid fish tribe that originated along with the so-called primary radiation of the Lake Tanganyika cichlid species flock, i.e. at the same time during which the majority of the endemic Lake Tanganyika cichlid tribes emerged. Unlike the other tribes, the haplochromines are not restricted to Lake Tanganyika but distributed throughout Africa, except for the northwestern part of the continent. Haplochromine cichlids seeded the adaptive radiation of cichlid fishes in Lakes Malawi, Kivu, Victoria, Turkana, as well as in the now extinct paleo-Lake Makgadikgadi. Here we present a comprehensive phylogenetic and phylogeographic analysis of haplochromine cichlids that is based upon DNA sequences of two mitochondrial gene segments of riverine taxa covering all major African biogeographic regions where haplochromines are found. Our analysis revealed that six lineages of haplochromines originated within a short period of time, about 5.3-4.4 MYA. These haplochromine lineages show a highly complex phylogeographic pattern, probably severely influenced by climate- and/or geology-induced changes of the environment, with river capture events most likely playing an important role for species dispersal.

The significance of sympatric speciation is one of the most controversial topics in evolutionary biology. Theory suggests that different factors can lead to speciation in full geographical contact, including selection and nonrandom mating. Strict criteria have been established for assessing sympatric speciation, which have been met in only a very few cases. Here, we investigate differentiation among sympatric morphospecies and color morphs of "roundfin" sailfin silversides (Telmatherinidae), small freshwater fish endemic to ancient Lake Matano in Central Sulawesi (Indonesia). Morphospecies are distinct according to body shape (geometric morphometrics), population structure (population-level amplified fragment length polymorphism [AFLP] markers), ecology, and mating behavior (habitat transects, stomach contents). Explorative genome scans based on AFLPs indicate that divergent selection affects only 1.3-4.2% of the analyzed loci, suggesting an early stage of speciation. Transect data demonstrate strong assortative mating and adaptive niche differentiation. However, we find no restrictions in gene flow among the conspicuous male color morphs. In summary, our data are consistent with a sympatric mode of divergence among three morphospecies under conditions effectively ruling out allopatric scenarios. Substantial, but incomplete, reproductive isolation suggests an early stage of speciation, most likely due to ecological selection pressure.

Adaptive radiations are extremely useful to understand factors driving speciation. A challenge in speciation research is to distinguish forces creating novelties and those relevant to divergence and adaptation. Recently, hybridization has regained major interest as a potential force leading to functional novelty and to the genesis of new species. Here, we show that introgressive hybridization is a prominent phenomenon in the radiation of sailfin silversides (Teleostei: Atheriniformes: Telmatherinidae) inhabiting the ancient Malili Lakes of Sulawesi, correlating conspicuously with patterns of increased diversity. We found the most diverse lacustrine species-group of the radiation to be heavily introgressed by genotypes originating from streams of the lake system, an effect that has masked the primary phylogenetic pattern of the flock. We conclude that hybridization could have acted as a key factor in the generation of the flock's spectacular diversity. To our knowledge, this is the first empirical evidence for massive reticulate evolution within a complex animal radiation.

BACKGROUND: Traditionally the rapid origin of megadiverse species flocks of extremely closely related species is explained by the combinatory action of three factors: Disruptive natural selection, disruptive sexual selection and partial isolation by distance. However, recent empirical data and theoretical advances suggest that the diversity of complex species assemblages is based at least partially on the hybridization of numerous ancestral allopatric lineages that formed hybrids upon invasion of new environments. That reticulate speciation within species flocks may occur under sympatric conditions after the primary formation of species has been proposed but not been tested critically. RESULTS: We reconstructed the phylogeny of a complex cichlid species flock confined to the tiny Cameroonian crater lake Barombi Mbo using both mitochondrial and nuclear (AFLP) data. The nuclear phylogeny confirms previous findings which suggested the monophyly and sympatric origin of the flock. However, discordant intra-flock phylogenies reconstructed from mitochondrial and nuclear data suggest strongly that secondary hybridization among lineages that primarily diverged under sympatric conditions had occurred. Using canonical phylogenetic ordination and tree-based tests we infer that hybridization of two ancient lineages resulted in the formation of a new and ecologically highly distinct species, Pungu maclareni. CONCLUSIONS: Our findings show that sympatric hybrid speciation is able to contribute significantly to the evolution of complex species assemblages even without the prior formation of hybrids derived from allopatrically differentiated lineages.