The evolutionary origin of the durophagous pelagic stingray ecomorph

Studies of the origin of evolutionary novelties (novel traits, feeding modes, behaviours, ecological niches, etc.) have considered a number of taxa experimenting with new body plans, allowing them to occupy new habitats and exploit new trophic resources. In the marine realm, colonization of pelagic environments by marine fishes occurred recurrently through time. Stingrays (Myliobatiformes) are a diverse clade of batoid fishes commonly known to possess venomous tail stings. Current hypotheses suggest that stingrays experimented with a transition from a benthic to a pelagic/benthopelagic habitat coupled with a transition from a non-durophagous diet to extreme durophagy. However, there is no study detailing macroevolutionary patterns to understand how and when habitat shift and feeding specialization arose along their evolutionary history. A new exquisitely preserved fossil stingray from the Eocene Konservat-Lagerstätte of Bolca (Italy) exhibits a unique mosaic of plesiomorphic features of the rajobenthic ecomorph, and derived traits of aquilopelagic taxa, that helps to clarify the evolutionary origin of durophagy and pelagic lifestyle in stingrays. A scenario of early evolution of the aquilopelagic ecomorph is proposed based on new data, and the possible adaptive meaning of the observed evolutionary changes is discussed. The body plan of †Dasyomyliobatis thomyorkei gen. et sp. nov. is intermediate between the rajobenthic and more derived aquilopelagic stingrays, supporting its stem phylogenetic position and the hypothesis that the aquilopelagic body plan arose in association with the evolution of durophagy and pelagic lifestyle from a benthic, soft-prey feeder ancestor.     

Cyrtocrinids (Cyrtocrinida,Crinoidea) and other associated crinoids from the Jurassic (Kimmeridgian–Tithonian)–Cretaceous (Berriasian–Barremian) of the Carpathian Foredeep basement (western Ukraine)

Upper Jurassic (Kimmeridgian–Tithonian) and Cretaceous (Berriasian–Barremian) strata of the Ukrainian part of the Carpathian Foredeep basement are rich, at least locally, in crinoid remains. Crinoids belonging to cyrtocrinids (Cyrtocrinida) are represented by whole cups, isolated remains of disarticulated cups, brachial plates and columnals. They are assigned to the following taxa: Cyrtocrinida indet., Eugeniacrinites cf. cariophilites (von Schlotheim), Lonchocrinus sp., Phyllocrinus stellaris Zaręczny, Ascidicrinus pentagonus (Jaekel), Gammarocrinites sp., Psalidocrinus armatus (Zittel), Psalidocrinus sp., and Hemibrachiocrinidae gen. indet. Cyrtocrinids are associated with other stalked (isocrinids, Isocrinida and millericrinids, Millericrinida) and stemless (saccocomids, Roveacrinida) crinoids. Columnals, pluricolumnals, brachial plates, and cirrals of isocrinids are assigned to Balanocrinus sp., Isocrinina fam. et subfam. indet., and columnals of millericrinids to Millericrinida indet. Free-living roveacrinids are assigned to Saccocoma sp. and Crassicoma sp. Knowledge on Jurassic and Cretaceous crinoids formerly described from Ukraine is discussed. Although majority of crinoids described herein seems to be allochthonous, autochthonous forms were also found with certainty in some intervals. These include some cyrtocrinids, which dominate in shallow-water environments of the Ukrainian Carpathian Foredeep basement. Isocrinids are also common in this shallow marine environment, whereas sessile saccocomids are assigned to low-energy, mud-supported bottom, open marine, outer-platform/upper slope, and relatively deep environments.     

IS SPECIALIZATION A DEAD END? THE PHYLOGENY OF HOST USE IN DENDROCTONUS BARK BEETLES (SCOLYTIDAE)

Ecological explanations for the prevalence of resource specialists are abundant, whereas phylogenetic evidence on their origins is scarce. In this paper, we provide a molecular phylogenetic study of the 19 specialist or generalist species in the bark beetle genus Dendroctonus, which collectively attack species in four different genera in the conifer family Pinaceae. Given substantial variation in diet breadth, we asked two general questions concerning the evolution of resource use in this group. How conservative is the evolution of host use in these insects? Does specialization tend to be derived (i.e., a “dead end”)? To answer these questions, we estimated the phylogeny of Dendroctonus using mitochondrial DNA sequences and mapped transitions in resource use on the resulting phylogeny estimate. The evolution of affiliations with Pinus and Picea hosts in Dendroctonus was conservative among beetle species (PTP test; P < 0.012), but there was no significant correspondence between the phylogeny of these beetles and the phylogeny among their Pinaceae hosts (among genera, P = 0.28; among Pinus species, P = 0.82). Degree of specialization, as measured in the proportion of hosts used, was bimodally distributed with “generalist” species utilizing < 60% of the congeneric hosts within their range and six specialist species utilizing < 40% of the available hosts. Among the generalists, we found a strong correlation between the number of hosts encountered and the number of hosts utilized (R = 0.97, P < 0.0001), whereas there was no significant correlation among the specialists (R = 0.27, P = 0.59). The evolution of specialization in Dendroctonus proved highly labile—specialists arose from generalists at least six separate times (without reversal) all in derived positions, and closer examination of some specialists revealed instances where they appear to have lost particular host species from their diet. However, evidence from the ecological literature also suggests that several Dendroctonus generalists may have increased their range of host genera within the Pinaceae.     

Sympatric speciation and radiative evolution of socially parasitic ants - Heretic hypotheses and their factual background

According to current hypotheses the main types of social parasitism among ants, namely slavery, temporary parasitism, and inquilinism, arose from such features as predation on other ants, or territorial behavior, both presumed precursors of slavemaking, and polygyny, a presumed precursor of temporary parasitism and inquilinism. The latter is believed also to represent a final instar in several evolutionary pathways leading from slavery, temporary parasitism, and xenobiosis to this permanently parasitic, workerless condition. Speciation, the origin of parasitic species from their usually closely related host species, is suggested to occur due to temporary geographic isolation and subsequent transition of one of the newly formed daughter species to parasitism in the nests of the other. Evidence is presented suggesting that the main types of social parasitism originated independently of each other. 15 ant genera are parasitized exclusively by inquilines, Eve other genera exclusively by temporary parasites. Only four groups of non-parasitic ant species (Formica, Tet-ramorium, Leptothorax subgenera Leptothorax and Myrafant) have parasites of several types each. Within these roups, however, there is little evidence of evolutionary transitions from one type to another. The few exceptions, mainly workerless species of the genera Epimyrma and Chalepoxenus, represent parasites which clearly derive from slave-making congeners, but differ from ordinary inquilines in that they eliminate the host colony queens like their actively dulotic ancestors. The new hypothesis suggests that all forms of interspecific true social parasitism (excluding xenobiosis) orginated from a common “preparasitic” stage, a subpopulation of reproductives in polygynous colonies and species, with diverging sexual behavior (near-nest mating vs. swarming) and caste ratios (production of more sexuals vs. workers). Arguments for sympatric speciation are compiled. Various features of the ancestral, and then host species (colony sizes, population density and structure, transition from polygyny to monoyny, etc.), and of the “preparasite” (production of few, or no workers, etc.) may shape the developing parasite to become a slave-maker, inquiline, or temporary parasite. These features usually leave open only one, or in a few genera, several options. The different types of parasitism within one host species group thus may have developed in a radiative manner from the common, preparasitic stage, which explains that independent colony foundation is a common feature of all true social parasites among ants.     

Evolutionary patterns in the terebratulid genus Caryona Cooper (Brachiopoda,Jurassic of France)

The study of ontogenetic and morphological changes in different species of the brachiopod genus Caryona COOPER (Terebratulidae) from the Lower Callovian‐Lower Oxfordian of northern France shows a succession (and an interference) of processes which vary according to the characters studied. This is an example of “mosaic evolution”;. The ontogenetic changes result from heterochrony, especially acceleration and hypermorphism. The main phylogenetic tendency is peramorphosis in most characters. This evolutionary pattern agrees with phyletic gradualism.

Moreover, analysis of the internal characters demonstrates the limited value of these in discrimination of the various genera.     

Reproductive mode evolution in nematodes: insights from molecular phylogenies and recently discovered species

The Phylum Nematoda has long been known to contain a great diversity of species that vary in reproductive mode, though our understanding of the evolutionary origins, causes and consequences of nematode reproductive mode change have only recently started to mature. Here we bring together and analyze recent progress on reproductive mode evolution throughout the phylum, resulting from the application of molecular phylogenetic approaches and newly discovered nematode species. Reproductive mode variation is reviewed in multiple free-living, animal-parasitic and plant-parasitic nematode groups. Discussion ranges from the model nematode Caenorhabditis elegans and its close relatives, to the plant-parasitic nematodes of the Meloidogyne genus where there is extreme variation in reproductive mode between and even within species, to the vertebrate-parasitic genus Strongyloides and related genera where reproductive mode varies across generations (heterogony). Multiple evolutionary transitions from dioecous (obligately outcrossing) to hermaphroditism and parthenogenesis in the phylum are discussed, along with one case of an evolutionary transition from hermaphroditism to doioecy in the Oscheius genus. We consider the roles of underlying genetic mechanisms in promoting reproductive plasticity in this phylum, as well as the potential evolutionary forces promoting transitions in reproductive mode.     

Phylogeny of the family Sialidae (Insecta: Megaloptera) inferred from morphological data,with implications for generic classification and historical biogeography

Sialidae (alderflies) is a family of the holometabolous insect order Megaloptera, with ca. 75 extant species in eight genera distributed worldwide. Alderflies are a group of “living fossils” with a long evolutionary history. The oldest fossil attributed to Sialidae dates back to the Early Jurassic period. Further, the global distribution of modern‐day species shows a remarkably disjunctive pattern. However, due to the rareness of most species and scarcity of comprehensive taxonomic revisions, the phylogeny of Sialidae remains largely unexplored, and the present classification system is in great need of renewal. Here we reconstruct the first phylogeny for Sialidae worldwide based on the most comprehensive sampling and broadest morphological data ever presented for this group of insects. All Cenozoic alderflies belong to a monophyletic clade, which may also include the Early Jurassic genus ?Dobbertinia, and the Late Jurassic genus ?Sharasialis is their putative sister taxon. Two subfamilies of Sialidae are proposed, namely ?Sharasialinae subfam. nov. and Sialidinae. Austrosialis is the sister of all other extant genera, an assemblage which comprises three monophyletic lineages: the Stenosialis lineage, the Ilyobius lineage, and the Sialis lineage. The revised classification of Sialidae is composed of 12 valid genera and 87 valid species. Ilyobius and Protosialis are recognized as valid generic names, while Nipponosialis is treated as a synonym of Sialis. Reconstruction of the ancestral area proposes a global distribution of alderflies in Pangaea before their diversification. The generic diversification of alderflies might have occurred before the breakup of Pangaea, but the divergence of some lineages or genera was probably promoted by the splitting of this supercontinent.     

Morphology and Molecular Phylogeny of a New Marine,Sand-dwelling Dinoflagellate Genus,Pachena (Dinophyceae), with Descriptions of Three New Species

Marine benthic dinoflagellates are interesting not only because some epiphytic genera can cause harmful algal blooms but also for understanding dinoflagellate evolution and diversification. Our understanding of their biodiversity is far from complete, and many thecate genera have unusual tabulation patterns that are difficult to relate to the diverse known phytoplankton taxa. A new sand-dwelling genus, Pachena gen. nov., is described based on morphological and DNA sequence data. Three species were discovered in distant locations and are circumscribed, namely, P. leibnizii sp. nov. from Canada, P. abriliae sp. nov. from Spain, and P. meriddae sp. nov. from Italy. All species are tiny (about 9–23 μm long) and heterotrophic. Species are characterized by their tabulation (APC 4′ 3a 6′′ 5c 5s 5′′′ 2′′′′), an apical hook covering the apical pore, an ascending cingulum, and a sulcus with central list. The first anterior intercalary plate is uniquely “sandwiched” between two plates. The species share these features and differ in the relative sizes and arrangements of their plates, especially on the epitheca. The ornamentation of thecal plates is species-specific. The new molecular phylogenies based on SSU and LSU rDNA sequences contribute to understanding the evolution of the planktonic relatives of Pachena, the Thoracosphaeraceae.     

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The cichlid fishes of Lake Malawi represent a prime example of a rapid evolutionary radiation via sexual selection. We show that in this group, acoustic communication significantly coincides with visual courtship displays. Specifically, individuals tend to use both multimodal and unimodal displays in successive courtship bouts. Behavioral analyses on six species of Malawian cichlids from two divergent genera revealed that five of the species use displays containing both an auditory and a visual component. Metriaclima zebra “katale” was the notable exception, along with a single individual of Metriaclima lombardoi, which utilized unimodal and multimodal displays with equal frequency. Taken together, these results highlight the importance of behavioral lability in multiple sensory signals at both the intra- and interspecific levels.     

Paraconvexitermes acangapua (Isoptera: Termitidae,Nasutitermitinae), a new genus and new species of the so-called “small Neotropical soil-feeding nasutes” from South America

A new genus Paraconvexitermes is described, comprising Paraconvexitermes acangapua, new species, P. nigricornis (Holmgren) and P. junceus (Emerson), both previously in Convexitermes Holmgren. Now, Convexitermes comprises C. manni Emerson and C. convexifrons (Holmgren). The worker’s digestive tube of both genera along with those of Atlantitermes species allowed to get a better definition of these three genera. Illustrations on the digestive tube and mandibles of workers, soldier’s head, alate’s head and secondary reproductive female of the new genus and species are provided. The phylogenetic relationships with the other nasute genera are discussed along with the “soil feeding” concept.     

Pleurotomaria DeFrance, 1826 (Gastropoda,Mollusca) from the Lower Bajocian (Middle Jurassic) sediments of Luxembourg,with considerations on its systematics,evolution and palaeobiogeographical history

Abstract: Pleurotomaria species from lower Bajocian (Middle Jurassic) sediments of south‐western Luxembourg housed in the National Natural History Museum of Luxembourg are described. Seven species are recognized, one of which is new, Pleurotomaria faberi sp. nov. A more detailed definition of the diagnostic characters of the genus is proposed and the morphological continuity between Talantodiscus and Pleurotomaria is demonstrated, suggesting that the former cannot be considered as a distinct taxon. The palaeoecology, evolution and palaeobiogeographical history of Pleurotomaria are outlined. Pleurotomaria presumably first appeared in late Middle Triassic of New Zealand where it underwent a relative diversification up to the Hettangian (Early Jurassic). From early Hettangian, most of its evolutionary history took place in Europe and western Tethys. In the European epicontinental seas, Pleurotomaria experienced two important radiations. The first occurred in the Early Jurassic, with a peak in the late Pliensbachian, and was marked by an expansion of the distribution to the central part of western Tethys. After a collapse in species diversity, probably related to the early Toarcian anoxic event, a second radiation occurred. This culminated in the early Bajocian and was mainly confined in a region encompassing southern England, Paris Basin and southern Germany. Low‐spired species, formerly attributed to Talantodiscus, probably originated independently and iteratively during the history of Pleurotomaria. The facies and associated benthic faunas suggest that Pleurotomaria probably lived on shallow soft bottoms composed of mixed calcareous–siliciclastic sediments. The two main Early Jurassic and early Middle Jurassic radiations of the genus took place in these environments. Records of the genus in Jurassic carbonate platform deposits are very few and concern mainly post‐Bajocian species.     

A new decapod crustacean faunule from the Middle Jurassic of north‐west France

Abstract: Decapod crustacean material collected recently from the lower Callovian (Middle Jurassic) in Maine‐et‐Loire (north‐west France) comprises two new species of prosopid and one new species of tanidromitid crabs, of the genera Nodoprosopon and Tanidromites, respectively. Also represented in this faunule is a probable paguroid anomuran, in the form of isolated chelae here assigned to the genus Orhomalus, as well as appendicular remains of unknown affinity; some of the latter might belong to prosopid crabs. These anomurans and brachyurans co‐occur with a diverse benthic fauna in limestones with abundant iron ooids; their main interest lies in the fact that they add valuable data to the rather poor record of Middle Jurassic decapod crustaceans.     

Speciation and Bursts of Evolution

A longstanding debate in evolutionary biology concerns whether species diverge gradually through time or by rapid punctuational bursts at the time of speciation. The theory of punctuated equilibrium states that evolutionary change is characterised by short periods of rapid evolution followed by longer periods of stasis in which no change occurs. Despite years of work seeking evidence for punctuational change in the fossil record, the theory remains contentious. Further there is little consensus as to the size of the contribution of punctuational changes to overall evolutionary divergence. Here we review recent developments which show that punctuational evolution is common and widespread in gene sequence data.     

Patterns of ontogenetic evolution in perianth diversification of Besseya (Scrophulariaceae)

Species of Besseya have perianth diversity centered largely in meristic variation, extreme corolla diminution, and corolla tube loss. Flowers of Besseya differ from those of their closest relative Synthyris in having a bilabiate corolla. The avenues of ontogenetic evolution that were important in creating these aspects of perianth diversity were explored by optimizing developmental transformations on cladograms. Three different evolutionary transformations were important in the diversification of calyx ontogenies in Besseya, including the derived reciprocal substitution of ontogenetic states in B. bullii, labile abaxial lobe expression in B. wyomingensis, and a heterotopic novel substitution in B. oblongifolia. The bilabiate corolla of Besseya arose via fractionation and localization of the zonal growth expressed in the more plesiomorphic Synthyris. Besseya plantaginea, B. ritteriana, and B. oblongifolia form a group characterized by diminished expression of the anterior corolla lobe. Four alternative scenarios equally explain the ontogenetic transformations underlying extreme corolla diminution and corolla tube loss. Corolla tube loss has been considered rare or impossible among angiosperms, but some alternative scenarios in Besseya demonstrate how this may have occurred via fractionation of zonal growth. Corolla tube loss may have arisen only indirectly from the sympetalous ancestors of Besseya following extreme corolla diminution and the reestablishment of a “full-size” corolla which had a novel developmental pattern that did not include the distribution of zonal growth present in ancestors.     

Out of the testis,into the ovary: biased outcomes of gene duplication and deletion in Drosophila

Gene turnover is a key source of adaptive variation. Yet most evolutionary studies have focused on gene duplication, dismissing gene deletion as a mechanism that simply eradicates redundancy. Here, I use genome‐scale sequence and multi‐tissue expression data from Drosophila melanogaster and Drosophila pseudoobscura to simultaneously assess the evolutionary outcomes of gene duplication and deletion in Drosophila. I find that gene duplication is more frequent than gene deletion in both species, indicating that it may play a more important role in Drosophila evolution. However, examination of several genic properties reveals that genes likely possess distinct functions after duplication that diverge further before deletion, suggesting that loss of redundancy cannot explain a majority of gene deletion events in Drosophila. Moreover, in addition to providing support for the well‐known “out of the testis” origin of young duplicate genes, analyses of gene expression profiles uncover a preferential bias against deletion of old ovary‐expressed genes. Therefore, I propose a novel “into the ovary” hypothesis for gene deletion in Drosophila, in which gene deletion may promote adaptation by salvaging genes that contribute to the evolution of female reproductive phenotypes. Under this combined “out of the testis, into the ovary” evolutionary model, gene duplication and deletion work in concert to generate and maintain a balanced repertoire of genes that promote sex‐specific adaptation in Drosophila.     

Crater lake cichlids individually specialize along the benthic–limnetic axis

A common pattern of adaptive diversification in freshwater fishes is the repeated evolution of elongated open water (limnetic) species and high‐bodied shore (benthic) species from generalist ancestors. Studies on phenotype‐diet correlations have suggested that population‐wide individual specialization occurs at an early evolutionary and ecological stage of divergence and niche partitioning. This variable restricted niche use across individuals can provide the raw material for earliest stages of sympatric divergence. We investigated variation in morphology and diet as well as their correlations along the benthic‐limnetic axis in an extremely young Midas cichlid species, Amphilophus tolteca, endemic to the Nicaraguan crater lake Asososca Managua. We found that A. tolteca varied continuously in ecologically relevant traits such as body shape and lower pharyngeal jaw morphology. The correlation of these phenotypes with niche suggested that individuals are specialized along the benthic‐limnetic axis. No genetic differentiation within the crater lake was detected based on genotypes from 13 microsatellite loci. Overall, we found that individual specialization in this young crater lake species encompasses the limnetic‐ as well as the benthic macro‐habitat. Yet there is no evidence for any diversification within the species, making this a candidate system for studying what might be the early stages preceding sympatric divergence.     

CHROMOSOME CYTOLOGY OF EUCHARIS,CALIPHRURIA, AND URCEOLINA (AMARYLLIDACEAE)

Karyotypes are presented for 19 taxa of Eucharis, Caliphruria, and Urceolina, a monophyletic group of Andean Amaryllidaceae within “infrafamily” Pancratioidinae Traub. All three genera are characterized by 2n = 46, the most common somatic number occurring in the “infrafamily.” Incidences of polyploidy are low. Only two tetraploid (2n = 92) species of Eucharis are so far known, E. bouchei and E. bonplandii, the northernmost species of E. subg. Eucharis. The 2n = 68 karyotype of E. amazonica is interpreted as triploid-derived (3x – 1). Chromosomal heteromorphism is reported for C. subedentata. Karyotype data is analyzed with principal component analysis and unweighted pairgroup cluster analysis. In a number of cases, phenetic relationships among the karyotypes correlate with phenetic and cladistic relationships based on morphological data. Karyotype evolution among the three genera is discussed in the context of classical theories of karyotypic symmetry. Stability of chromosome number in Eucharis and related genera suggests that chromosomal evolution has proceeded via nonreciprocal interchanges between chromosomes and infrachromosomal structural change. In at least one case (E. bakeriana), rapid sympatric speciation may have been vectored by chromosomal change.     

Evolutionary trajectory of fish Piscine novirhabdovirus (=Viral Hemorrhagic Septicemia Virus) across its Laurentian Great Lakes history: Spatial and temporal diversification

Piscine novirhabdovirus = Viral Hemorrhagic Septicemia Virus (VHSV) first appeared in the Laurentian Great Lakes with large outbreaks from 2005 to 2006, as a new and novel RNA rhabdovirus subgenogroup (IVb) that killed >30 fish species. Interlude periods punctuated smaller more localized outbreaks in 2007, 2010, and 2017, although some fishes tested positive in the intervals. There have not been reports of outbreaks or positives from 2018, 2019, or 2020. Here, we employ a combined population genetics and phylogenetic approach to evaluate spatial and temporal evolutionary trajectory on its G‐gene sequence variation, in comparison with whole‐genome sequences (11,083 bp) from a subset of 44 individual isolates (including 40 newly sequenced ones). Our results show that IVb (N = 184 individual fish isolates) diversified into 36 G‐gene haplotypes from 2003 to 2017, stemming from two originals (“a” and “b”). G‐gene haplotypes “a” and “b” differed by just one synonymous single‐nucleotide polymorphism (SNP) substitution, remained the most abundant until 2011, then disappeared. Group “a” descendants (14 haplotypes) remained most prevalent in the Upper and Central Great Lakes, with eight (51%) having nonsynonymous substitutions. Group “b” descendants primarily have occurred in the Lower Great Lakes, including 22 haplotypes, of which 15 (68%) contained nonsynonymous changes. Evolutionary patterns of the whole‐genome sequences (which had 34 haplotypes among 44 isolates) appear congruent with those from the G‐gene. Virus populations significantly diverged among the Upper, Central, and Lower Great Lakes, diversifying over time. Spatial divergence was apparent in the overall patterns of nucleotide substitutions, while amino acid changes increased temporally. VHSV‐IVb thus significantly differentiated across its less than two decades in the Great Lakes, accompanied by declining outbreaks and virulence. Continuing diversification likely allowed the virus to persist at low levels in resident fish populations, and may facilitate its potential for further and future spread to new habitats and nonacclimated hosts.     

Phylogeographic patterns and species delimitation in the endangered silverside “humboldtianum” clade (Pisces: Atherinopsidae) in central Mexico: understanding their evolutionary history

The Atherinopsidae is the second largest group of freshwater fishes occupying central Mexico and is one of biological, cultural, and economic importance. The “humboldtianum” clade (Genus Chirostoma) is a “species flock” of nine described species that inhabit lacustrine ecosystems in central Mexico. The high morphological polymorphism within the group makes species identification difficult and thereby limits the development of research and management projects focusing on this group. In this study, we used phylogeographic and coalescent-based methods to understand the evolution of genetic variation among these species. The results revealed taxonomic inaccuracies and genetic admixture among species. Genetic variation was structured geographically, rather than taxonomically, and five closely related genetic groups were recovered. Two evolutionary pathways were found. First, a novel geographical arrangement of haplotypes was recovered that gave rise to the five recently (Pleistocene, <?1 Myr) derived genetic groups. The second pathway showed a recent intra-lacustrine genetic differentiation that could be associated with sympatric or ecological speciation. The current classification of the group is revised and includes a reduction in the number of valid species in the “humboldtianum” clade. Moreover, this study provides new insight into the biogeography and evolutionary history of this important group of fishes.

     

Life history diversity and evolution in the Asterinidae

Asterinid sea stars have the greatest range of life historiesknown for the Asteroidea. Larval form in these sea stars hasbeen modified in association with selection for planktonic,benthic, or intergonadal developmental habitats. Life historydata are available for 31 species and molecular data for 28of these. These data were used to assess life history evolutionand relationships among asterinid clades. Lecithotrophy is prevalentin Asterinidae, with at least 6 independent origins of thisdevelopmental mode. Morphological differences in the attachmentcomplex of brachiolaria larvae were evident among species withplanktonic lecithotrophy. Some features are clade specific whileothers are variable within clades. Benthic brachiolariae aresimilar in Aquilonastra and Parvulastra with tripod-shaped larvae,while the bilobed sole-shaped larvae of Asterina species appearunique to this genus. Multiple transitions and pathways havebeen involved in the evolution of lecithotropy in the Asterinidae.Although several genera have a species with a planktonic feedinglarva in a basal phylogenetic position, relative to specieswith planktonic or benthic lecithotrophy, there is little evidencefor the expected life history transformation series from planktonicfeeding, to planktonic non-feeding, to benthic non-feeding development.Intragonadal development, a life history pattern unique to theAsterinidae, arose three times through ancestors with benthicor pelagic lecithotrophy. Evolution of lecithotrophy appearsmore prevalent in the Asterinidae than other asteroid families.As diverse modes of development are discerned in cryptic speciescomplexes, new insights into life history evolution in the Asterinidaeare being generated.