Phylogenetic inference in Odontesthes and Atherina (Teleostei: Atheriniformes) with insights into ecological adaptation

This contribution provides an insight into Atheriniformes systematics based on four mitochondrial regions: 12S rRNA, cytb, COI and control region (2794bp in total). In the Atherinopsoidei (New World silversides), comparisons among five species of Odontesthes, O. argentinensis, O. bonariensis, O. smitti, O. hatcheri and O. incisa revealed a putative marine-freshwater pairing pattern of Odontesthes species, possibly driven by sea level fluctuations of South American waters. This study represents the first data on molecular phylogeny of Odontesthes species that can be of usefulness to biodiversity conservation policies. In the Atherinoidei (Old World silversides), Atherina boyeri was corroborated as a species complex constituted by a marine form, a marine with dark spots form and a brackish form. Concretely, Odontesthes and Atherina may represent geographically replicated models to study genetic adaptation and speciation of marine species to brackish and freshwater habitats. In addition, phylogenetic analyses supported Odontesthes and Atherina as monophyletic taxa and their separation into two differentiated suborders Atherinopsoidei and Atherinoidei, respectively.     

Phylogenetic relationships and demographic histories of the Atherinidae in the Eastern Atlantic and Mediterranean Sea re-examined by Bayesian inference

The aim of our study is to examine the phylogenetic relationship, divergence times and demographic history of the five close-related Mediterranean and North-eastern Atlantic species/forms of Atherina using the full Bayesian framework for species tree estimation recently implemented in ?BEAST. The inference is made possible by multilocus data using three mitochondrial genes (12S rRNA, 16S rRNA, control region) and one nuclear gene (rhodopsin) from multiple individuals per species available in GenBank. Bayesian phylogenetic analysis of the complete gene dataset produced a tree with strong support for the monophyly of each species, as well as high support for higher level nodes. An old origin of the Atherina group was suggested (19.2 MY), with deep split events within the Atherinidae predating the Messinian Salinity Crisis. Regional genetic substructuring was observed among populations of A. boyeri, with AMOVA and MultiDimensional Scaling suggesting the existence of five groupings (Atlantic/West Mediterranean, Adriatic, Greece, Black Sea and Tunis). The level of subdivision found might be consequence of the hydrographic isolation within the Mediterranean Sea. Bayesian inference of past demographic histories showed a clear signature of demographic expansion for the European coast populations of A. presbyter, possibly linked to post-glacial colonizations, but not for the Azores/Canary Islands, which is expected in isolated populations because of the impossibility of finding new habitats. Within the Mediterranean, signatures of recent demographic expansion were only found for the Adriatic population of A. boyeri, which could be associated with the relatively recent emergence of the Adriatic Sea.     

On the reproductive biology of the sand smelt Atherina boyeri Risso (Pisces: Atherinidae) and its evolutionary potential

The small, typically estuarine fish Atherina boyeri Risso forms local populations which can differ greatly in their population biology and morphology; this species is viewed as being on the brink of speciation. This analysis of the reproductive strategy shows that A. boyeri can rapidly adapt its life history and morphology to environments ranging from freshwater to polyhaline coastal lagoons and oceanic waters. The optimum strategy is shown to be tied to temperate seasonality: the cold winters act to lengthen the life cycle, and result in cyclic switching of energy between reproduction, somatic growth and fat storage. The adaptability of the fish to a range of environmental characters is viewed as being due to natural selection acting to produce a genotype capable of a flexible phenotypic response. This flexibility is essential for survival in the highly variable inshore/brackish habitat. It is argued that such preadaptive plasticity, coupled with the potential for populations to become isolated, produces ideal conditions for speciation. The importance of the coastal, estuarine and lagoonal environment as a springboard for teleost evolution is discussed.     

Marine mites (Halacaroidea: Acari): a geographical and ecological survey

Halacarid mites (Acari), with almost 700 species described, inhabit marine and freshwater habitats. The majority of genera are recorded from at least two ocean basins or continents. There is no evidence of endemic genera, in either littoral faunal provinces or in deep-sea regions. Copidognathus, a genus comprising 1/4 of all species described, is found in almost all geographic regions, depths and habitats. Other genera dominate or are restricted to cold waters, to warm waters or to distinct habitats.Corresponding habitats on either side of the boreal Atlantic Ocean harbour congeneric, identical, sibling or morphologically similar species. The fauna in the western Atlantic is less diverse than that in the eastern. Amphiatlantics are restricted to certain genera. Transatlantic distribution is independent of the niche inhabited.Of the marine halacarid species recorded from the boreal western Atlantic, 41% are amphiatlantics, while only one species is recorded from both the Caribbean and the Mediterranean. The Caribbean and the Mediterranean faunas are dominated by genera in which amphiatlantics are unknown.Most of the Black Sea species of the genus Halacarellus also occur in the Baltic, North Sea or North Atlantic, whereas the Copidognathus fauna of the Black Sea is similar to that of the Mediterranean.Halacarids are thought to be an ancient taxon, with most genera probably having been present since the Mesozoic and with several species having an age of at least 50 million years. Evidence for their antiquity is found in the distributional pattern of marine and limnic genera and species, in the lack of endemic genera despite low fecundity and lack of dispersal stages, and in the fact that amphiatlantics are restricted to certain genera without relationships to the niches inhabited.     

Phylogenetic relationships of Atherina hepsetus and Atherina boyeri (Pisces: Atherinidae) populations from Greece, based on mtDNA sequences

The genetic divergence and the phylogenetic relationships of six Atherina boyeri (freshwater and marine origin) and five Atherina hepsetus populations from Greece were investigated using partial sequence analysis of 12s rRNA, 16s rRNA and control region mtDNA segments. Three different well divergent groups were revealed; the first one includes A. boyeri populations living in the sea, the second includes A. boyeri populations living in the lakes and lagoons whereas the third one includes all A. hepsetus populations. Fifty-seven different haplotypes were detected among the populations studied. In all three mtDNA segments examined, sequence analysis revealed the existence of fixed haplotypic differences discriminating A. boyeri populations inhabiting the lagoon and the lakes from both the coastal A. boyeri and the A. hepsetus populations. The genetic divergence values estimated between coastal (marine) A. boyeri populations and those living in the lagoon and the lakes are of the same order of magnitude as those observed among coastal A. boyeri and A. hepsetus populations. The results obtained by different phylogenetic methods were identical. The deep sequence divergence with the fixed different haplotypes observed suggests the occurrence of a cryptic or sibling species within A. boyeri complex.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 151–161.     

Deep phylogeographic structure may indicate cryptic species within the Sparid genus Spondyliosoma

Two geographically nonoverlapping species are currently described within the sparid genus Spondyliosoma: Spondyliosoma cantharus (Black Seabream) occurring across Mediterranean and eastern Atlantic waters from NW Europe to Angola and S. emarginatum (Steentjie) considered endemic to southern Africa. To address prominent knowledge gaps this study investigated range-wide phylogeographic structure across both species. Mitochondrial DNA sequences revealed deep phylogeographic structuring with four regionally partitioned reciprocally monophyletic clades, a Mediterranean clade and three more closely related Atlantic clades [NE Atlantic, Angola and South Africa (corresponding to S. emarginatum)]. Divergence and distribution of the lineages reflects survival in, and expansion from, disjunct glacial refuge areas. Cytonuclear differentiation of S. emarginatum supports its validity as a distinct species endemic to South African waters. However, the results also indicate that S. cantharus may be a cryptic species complex wherein the various regional lineages represent established/incipient species. A robust multilocus genetic assessment combining morphological data and detailing interactions among lineages is needed to determine the full diversity within Spondyliosoma and the most adequate biological and taxonomic status.     

Enzymatic survey of four populations of Atherina boyeri based on electrophoresis and the occurrence of a microsporidiosis

Biochemical genetics of a Atherinid species, Atherina boyeri were undertaken to determine variability. Among the 17 loci studied, 7 were polymorphic, of which three were retained for a more detailed comparison of populations inhabiting the French Mediterranean coastal lagoons. A microsporidiosis is present in these fishes and is limited to certain lagoons. Comparison of allelic frequencies at the polymorphic loci between fish populations of infected and healthy lagoons yielded no significant differences. Several hypotheses are discussed and attempt to agree parasite distribution with genetical results.     

Phylogeny of the cave shrimp Troglocaris: Evidence of a young connection between Balkans and Caucasus

The remarkably discontinuous distribution of the cave shrimp genus Troglocaris in South France, West Balkans, and West Caucasus has long been considered a biogeographic enigma. To solve it, its phylogeny was reconstructed by analyzing sequences from two mitochondrial (cytochrome oxidase I and 16S rRNA) and one nuclear gene (28S rRNA) using maximum likelihood, parsimony and Bayesian inference. The genus was found to be polyphyletic because the French taxon T. inermis had no direct common ancestry with other Troglocaris taxa but was sister to the epigean freshwater atyid Dugastella valentina. All other Troglocaris species constituted a well-supported monophylum, the second cave shrimp genus Spelaeocaris nested within. The monophylum had a well-defined structure: (1) a clade restricted to the Dinaric area of the Western Balkans containing the type species T. anophthalmus along with some unnamed species, and (2) a geographically mixed clade split between the Caucasian T. kutaissiana species complex on one, and T. hercegovinensis, S. pretneri, plus an unnamed taxon on the other side. It was surprising to find the dichotomy between the Caucasian and one of the West-Balkan lineages so low in the phylogenetic hierarchy of the genus. Taking into account molecular rates of other decapods, we tentatively dated this split at 6-11 Myr. This time is in agreement with the brackish and freshwater phase of the Paratethys thus allowing for a freshwater common ancestor of Caucasian and Dinaric cave shrimps. This would weaken the marine relicts hypothesis that has often been invoked to explain the distribution of freshwater cave species with close marine relatives.     

Life history strategies of four small-size fishes in the Suez Canal, Egypt

Of 35 species collected from the shores of the Suez Canal and its lakes, five were abundant year round. Sihouettea aegyptia and Aphanius dispar are of Red Sea (warm water) origin, whereas Pomatoschistus marmoratus, Aphanius fasciatus and Atherina boyeri are Mediterranean species. S. aegyptia and A. dispar are distributed throughout the Suez system whereas A. fasciatus is restricted to the northern canal. P. marmoratus and A. boyeri have not spread southward beyond the Bitter Lakes. Salinity is the main limiting factor for the distribution of Mediterranean species. S. aegyptia and A. dispar are dominant in the Bitter Lakes (salinity 44‰), whereas P. marmoratus and A. boyeri are abundant in Timsah Lake (salinity 7·8–41·6‰). There was considerable interannual and monthly variation in the relative abundance of each species. The populations were dominated by a single age group, and life spans were no more than 2 years. Rapid growth was evident during the first spring in P. marmoratus and A. boyeri and during the first summer, early autumn in S. aegyptia and A. dispar . The relative abundance of each food item in the diet varied with fish size and season. S. aegyptia, P. marmoratus and A. dispar fed mostly on harpacticoid copepods, polychaetes, demersal eggs, diatoms and blue-green algae whereas A. boyeri fed mostly on planktonic copepods. The warm water species are summer spawners, whereas the temperate species are autumn-winter spawners.     

Diverse migration strategy between freshwater and seawater habitats in the freshwater eel genus Anguilla

The freshwater eels of the genus Anguilla, which are catadromous, migrate between freshwater growth habitats and offshore spawning areas. A number of recent studies, however, found examples of the temperate species Anguilla anguilla, Anguilla rostrata, Anguilla japonica, Anguilla australis and Anguilla dieffenbachii that have never migrated into fresh water, spending their entire life history in the ocean. Furthermore, those studies found an intermediate type between marine and freshwater residents, which appear to frequently move between different environments during their growth phase. The discovery of marine and brackish-water residents Anguilla spp. suggests that they do not all have to be catadromous, and it calls into question the generalized classification of diadromous fishes. There has been little available information, however, concerning migration in tropical Anguilla spp. Anguilla marmorata, shows three fluctuation patterns: (1) continuous residence in fresh water, (2) continuous residence in brackish water and (3) residence in fresh water after recruitment, while returning to brackish water. Such migratory patterns were found in other tropical species, Anguilla bicolor bicolor and Anguilla bicolor pacifica. In A. b. bicolor collected in a coastal lagoon of Indonesia, two further patterns of habitat use were found: (1) constantly living in either brackish water or sea water with no freshwater life and (2) habitat shift from fresh water to brackish water or sea water. The wide range of environmental habitat use indicates that migratory behaviour of tropical Anguilla spp. is facultative among fresh, brackish and marine waters during their growth phases after recruitment to the coastal areas. Further, the migratory behaviours of tropical Anguilla spp. appear to differ in each habitat in response to inter and intra-specific competition. The results suggest that tropical Anguilla spp. have a flexible pattern of migration, with an ability to adapt to various habitats and salinities. The ability of anguillids to reside in environments of various salinities would be a common feature between tropical and temperate species without a latitudinal cline. Thus, the migration of Anguilla spp. into fresh water is clearly not an obligatory behaviour. This evidence of geographical variability among Anguilla spp. suggests that habitat use is determined by environmental conditions in each site.     

Atherina punctata and Atherina lagunae (Pisces,Atherinidae), new species found in the Mediterranean sea. 2. Molecular investigations of three Atherinid species

On the basis of morphoanatomical parameters, the sand smelt species (Atherina boyeri Risso, 1810) is viewed as a highly polymorphic complex. In this study, intraspecific sequence variation in a portion of the cytochrome b gene was examined in 88 individuals from Tunisia and France. The correlation between the results of statistical analysis of the sequence data using a variety of tree-building algorithms and morphoanatomical analyses demonstrated the subdivision into three putative species: A. boyeri, which only includes non-punctuated fishes, A. punctata, which corresponds to punctuated fishes and A. lagunae, which corresponds to atherines living in lagoons.     

MOLECULAR PHYLOGENY AND TAXONOMY OF THE AEGAGROPILA CLADE (CLADOPHORALES,ULVOPHYCEAE), INCLUDING THE DESCRIPTION OF AEGAGROPILOPSIS GEN. NOV. AND PSEUDOCLADOPHORA GEN. NOV.1

The Aegagropila clade represents a unique group of cladophoralean green algae occurring mainly in brackish and freshwater environments. The clade is sister to the species‐rich and primarily marine Cladophora and Siphonocladus lineages. Phylogenetic analyses of partial LSU and SSU nrDNA sequences reveal four main lineages within the Aegagropila clade, and allow a taxonomic reassessment. One lineage consists of two marine ‘Cladophora’ species, for which the new genus Pseudocladophora and the new family Pseudocladophoraceae are proposed. For the other lineages, the family name Pithophoraceae is reinstated. Within the Pithophoraceae, the earliest diverging lineage includes Wittrockiella and Cladophorella calcicola, occurring mainly in brackish and subaerial habitats. The two other lineages are restricted to freshwater. One of them shows a strong tendency for epizoism, and consists of Basicladia species and Arnoldiella conchophila. The other lineage includes Aegagropila, Pithophora and a small number of tropical ‘Cladophora’ species. The latter are transferred to the new genus Aegagropilopsis. Previously, polypyramidal pyrenoids had been suggested to be apomorphous for this clade, but we report the finding of both polypyramidal and bilenticular pyrenoids in members of the Pithophoraceae, and thus show that this character has no diagnostic value.     

Environmental factors controlling the spring immigration of two estuarine fishes Atherina boyeri and Pomatoschistus spp. into a Mediterranean lagoon

Atherina boyeri , a pelagic species, mainly entered a Mediterranean lagoon in the Rhone River Delta, during the day, whatever the flow direction. Flow velocity did not appear to be a significant factor and its swimming abilities seemed to be sufficient to cope with the observed velocity. The combined responses of Pomatoschistus spp., benthic species, to the different environmental factors clearly demonstrated a change in behavioural response during their ontogenic development, i.e. a shift from passive to active or active-passive migration. Young Pomatoschistus spp. (<20 mm L _T) exhibited a behaviour that correspond to the classical selective tidal stream transport of young fishes towards estuarine areas (passive drifting). Large Pomatoschistus spp. were also able to enter the lagoon with the flow, but they remained negatively affected by the flow velocity. The diel cycle appeared to be the main factor controlling the immigration of A. boyeri into the lagoon whereas Pomatoschistus spp. seemed more sensitive to hydrographic conditions prevailing at the connection.     

The biology of Boyer's sand smelt, Atherina boyeri Risso in the Bardawil Lagoon on the Mediterranean coast of Sinai

The present study describes some aspects of the life history of Boyer's sand smelt Atherina boyeri in the hypersaline Bardawil Lagoon on the Mediterranean coast of the Sinai peninsula, Egypt. Monthly samples were collected with a small-mesh experimental beach-seine during the years 1973–1974. Age determinations were based on otoliths readings and length frequency distributions which indicate that the Bardawil Lagoon population consists mostly of 0-age group. The largest recorded fish was 63 mm in standard length (S.L.). Length-weight relationship has been calculated as: W = 13.7 × 10⁻⁶× L ^2.93 where W = weight (g) and L =S.L. (mm). Atherina boyeri mature at the length of 34 mm. Spawning takes place from March to September. The average number of eggs in the ovary of a ripe female at the beginning of the spawning season was found to be 522. Females of all sizes were more numerous than males. In immature fish, less than 34 mm long, females constituted 53% of the population. In larger fish the number of females was higher; in fish over 34 mm long females constituted 64% and in fish over 51 mm—96%. Atherina boyeri in the Bardawil Lagoon feeds on both zoobenthos and zooplankton, mainly amphipods and copepods. Polychaetes, mysidaceans, insects and fish were also represented. Atherina boyeri is the host of endoparasitic Trematoda, Acanthocephala and Nematoda. The examined fish were found to have metacercaria in their mesenteries and liver.     

Biogeographical distribution of Hartaetosiga strains (Choanoflagellatea,Craspedida, Salpingoecidae) including morphological and transcriptomic data from a transect across the Atlantic Ocean

The genus Hartaetosiga Carr, Richter and Nitsche, 2017 comprised up to now only three species, H. gracilis (Kent) Carr, Richter, Nitsche, 2017, H. balthica (Wylezich and Karpov) Carr, Richter and Nitsche, 2017 and H. minima (Wylezich and Karpov) Carr, Richter and Nitsche, 2017. Based on distinct molecular data these species were relocated from the strictly freshwater genus Codosiga (Ehrenberg) Bütschli, 1878 to a new genus comprising brackish and marine species. During the cruise MSM82/2 across the Atlantic Ocean in 2019, surface water samples were taken from 15 stations along a transect ranging from 35°S to 23°N. We were able to isolate and cultivate 14 strains of the genus Hartaetosiga. Morphometric data showed no distinct morphological traits allowing for a species delineation, indicating a cryptic species complex within the genus. Based on cultivation, morphological data, and molecular analyses, we recorded H. gracilis for the first time from off-shelf waters of the Atlantic Ocean and could describe a new species, H. australis n. sp. This new species was recorded from sampling stations in the Southern Hemisphere only, which may indicate a potential biogeographic restriction likely caused by the Equatorial Counter Current (ECC), dividing the northern and southern surface waters.     

Phylogenetics and biogeography of the Balkan ‘sand gobies’ (Teleostei: Gobiidae): vulnerable species in need of taxonomic revision

Within the Atlantic–Mediterranean region, the ‘sand gobies’ are abundant and widespread, and play an important role in marine, brackish, and freshwater ecosystems. They include the smallest European freshwater fish, Economidichthys trichonis, which is threatened by habitat loss and pollution, as are several other sand gobies. Key to good conservation management is an accurate account of the number of evolutionary significant units. Nevertheless, many taxonomic and evolutionary questions remain unresolved within the clade, and molecular studies are lacking, especially in the Balkans. Using partial 12S and 16S mitochondrial ribosomal DNA sequences of 96 specimens of at least eight nominal species (both freshwater and marine populations), we assess species relationships and compare molecular and morphological data. The results obtained do not support the monophyly of Economidichthys, suggesting the perianal organ to be a shared adaptation to hole‐brooding rather than a synapomorphy, and urge for a taxonomic revision of Knipowitschia. The recently described Knipowitschia montenegrina seems to belong to a separate South‐East Adriatic lineage. Knipowitschia milleri, an alleged endemic of the Acheron River, and Knipowitschia cf. panizzae, are shown to be very closely related to other western Greek Knipowitschia populations, and appear conspecific. A distinct Macedonian–Thessalian lineage is formed by Knipowitschia thessala, whereas Knipowitschia caucasica appears as an eastern lineage, with populations in Thrace and the Aegean. The present study combines the phylogeny of a goby radiation with insights on the historical biogeography of the eastern Mediterranean, and identifies evolutionary units meriting conservation attention. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 73–91.     

A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast

A new species of the thermophylic Tethyan relict prawn Typhlatya is described from two anchialine caves near Perpignan (southern France). The new species is closely related to a congener known only from a freshwater cave at Castellón (eastern Spain), about 400 km to the south-west, differing apparently only in the size and shape of the rostrum and the armature of the dactylus of the fifth pereiopod. Based on palaeogeographical evidence and assuming a sister-group relationship between both species, we suggest that their common ancestor could not be older than early Pliocene in age, and that it was already a stygobiont taxon adapted to live in shallow-water marine crevicular habitats. This ancestor would have vanished from the western Mediterranean after the cooling associated with the onset of northern Hemisphere glaciation, about 3 Mya, as documented for other Mediterranean marine taxa. Indeed, the genus is completely stygobiont and does not occur in fluvial environments. The Pyrenees represent a watershed boundary that eliminates the possibility of the derivation of one species from the other by active dispersal after establishment in continental waters.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 387–414.     

Morphological variation in British Atherinids, and the status of Atherina presbyter Guvier (Pisces: Atherinidae)

A series of morphometric and meristic analyses conducted on specimens of sand smelt, Atherina , from populations around the British Isles, and including classic A. boyeri and A. presbyter forms, has shown that the characteristics variously used in the past to distinguish these two species are invalid. Multivariate analyses showed no significant splitting of the material into two groups which might correlate to these two species; classic A. boyeri and the very large Atlantic coast A. presbyter individuals represent the tails of a continuum of form. From these and published data, it is concluded that the A. boyeri morphology varies under the influence of conditions of temperature and salinity during embryo development, and the isolation of populations maintains comparative morphological distinctions resulting from local selection and random genetic drift.     

Relatedness of novel species of Myxidium Bütschli, 1882, Zschokkella Auerbach, 1910 and Ellipsomyxa Køie, 2003 (Myxosporea: Bivalvulida) from the gall bladders of marine fishes (Teleostei) from Australian waters

During a parasitological survey, Myxidium-like spores were identified in the gall bladders of marine fishes from Australian waters. This paper describes four novel species of Ellipsomyxa Køie, 2003, three novel species of Myxidium Bütschli, 1882 and six novel species of Zschokkella Auerbach, 1910 from teleosts from Australian waters using a combination of morphological, biological and molecular characters. Phylogenetic analyses showed a monophyletic relationship of all Ellipsomyxa spp. sequences with Sigmomyxa sphaerica (Thélohan, 1895) and Myxidium queenslandicus Gunter & Adlard, 2008 as sister species to the clade. The validity of genus Sigmomyxa Karlsbakk & Køie, 2012 is discussed. In phylogenetic analyses, the novel species of Myxidium fell within the ‘marine’ clade of Fiala (2006). However, the novel species of Zschokkella fell within the ‘freshwater’ clade of Fiala (2006) and formed a distinct clade with all other sequences of Zschokkella spp. from the gall bladder of marine fish and a sequence of a species of Myxobolus Bütschli 1882, also from the gall bladder of a marine fish. This is the second distinct marine lineage to emerge within the freshwater clade.     

Why are there so few fish in the sea?

The most dramatic gradient in global biodiversity is between marine and terrestrial environments. Terrestrial environments contain approximately 75-85% of all estimated species, but occupy only 30 per cent of the Earth's surface (and only approx. 1-10% by volume), whereas marine environments occupy a larger area and volume, but have a smaller fraction of Earth's estimated diversity. Many hypotheses have been proposed to explain this disparity, but there have been few large-scale quantitative tests. Here, we analyse patterns of diversity in actinopterygian (ray-finned) fishes, the most species-rich clade of marine vertebrates, containing 96 per cent of fish species. Despite the much greater area and productivity of marine environments, actinopterygian richness is similar in freshwater and marine habitats (15 150 versus 14 740 species). Net diversification rates (speciation-extinction) are similar in predominantly freshwater and saltwater clades. Both habitats are dominated by two hyperdiverse but relatively recent clades (Ostariophysi and Percomorpha). Remarkably, trait reconstructions (for both living and fossil taxa) suggest that all extant marine actinopterygians were derived from a freshwater ancestor, indicating a role for ancient extinction in explaining low marine richness. Finally, by analysing an entirely aquatic group, we are able to better sort among potential hypotheses for explaining the paradoxically low diversity of marine environments.