A review and synthesis on the systematics and evolution of jellyfish blooms: advantageous aggregations and adaptive assemblages

Pelagic gelatinous invertebrates in many diverse phyla aggregate, bloom, or swarm. Although typically portrayed as annoying to humans, such accumulations probably are evolutionary adaptations to the environments of pelagic gelatinous zooplankton. We explore this proposition by systematic analysis completed in three steps. First, using the current morphological taxonomic framework for Scyphozoa, we summarize relevant information on species that aggregate, bloom, and swarm and on those species that do not. Second, we establish a molecular phylogenetic framework for assessing evolutionary relationships among classes and many orders of Medusozoa and among most families of Scyphozoa (particularly Discomedusae). Third, we interpret the phylogenetic distribution of taxa and of characteristics of jellyfish that aggregate, bloom, or swarm, in terms of species diversity—a proxy for evolutionary success. We found that: (1) Medusae that occur en masse are not randomly distributed within the Phylum Cnidaria but instead they are found primarily within the Scyphozoa which have a metagenic life history. (2) Midwater and deep-sea medusae rarely bloom or swarm. (3) Epibenthic medusae do not swarm. (4) Large carnivores that feed on large prey do not bloom strongly. (5) Large medusae that feed exclusively on small prey both bloom and swarm. (6) Pelagia, the only holoplanktonic, epipelagic scyphomedusan, both blooms and swarms, demonstrating that a metagenic life cycle is not required for blooming or swarming at sea. (7) Environmental change (overfishing, species introductions, and eutrophication) may induce or inhibit blooms. (8) Taxa that bloom or swarm are often more diverse than taxa that do not. (9) Speciation in scyphozoans can occur rapidly. (10) Morphological stasis in holozooplankton masks genetic variability. (11) Selection for convergent evolution in the sea is strong because mass occurrence has evolved multiple times in independent evolutionary lineages under similar circumstances. Thus, attributes possessed by many taxa that occur en masse appear to be evolutionarily advantageous, i.e., adaptations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: K. A. Pitt & J. E. Purcell Jellyfish Blooms: Causes, Consequences, and Recent Advances     

Cladistic analysis of Medusozoa and cnidarian evolution

Abstract. A cladistic analysis of 87 morphological and life history characters of medusozoan cnidarians, rooted with Anthozoa, results in the phylogenetic hypothesis (Anthozoa (Hydrozoa (Scyphozoa (Staurozoa, Cubozoa)))). Staurozoa is a new class of Cnidaria consisting of Stauromedusae and the fossil group Conulatae. Scyphozoa is redefined as including those medusozoans characterized by strobilation and ephyrae (Coronatae, Semaeostomeae, and Rhizostomeae). Within Hydrozoa, Limnomedusae is identified as either the earliest diverging hydrozoan lineage or as the basal group of either Trachylina (Actinulida (Trachymedusae (Narcomedusae, Laingiomedusae))) or Hydroidolina (Leptothecata (Siphonophorae, Anthoathecata)). Cladistic results are highly congruent with recently published phylogenetic analyses based on 18S molecular characters. We propose a phylogenetic classification of Medusozoa that is consistent with phylogenetic hypotheses based on our cladistic results, as well as those derived from 18S analyses. Optimization of the characters presented in this analysis are used to discuss evolutionary scenarios. The ancestral cnidarian probably had a sessile biradial polyp as an adult form. The medusa is inferred to be a synapomorphy of Medusozoa. However, the ancestral process (metamorphosis of the apical region of the polyp or lateral budding involving an entocodon) could not be inferred unequivocally. Similarly, character states for sense organs and nervous systems could not be inferred for the ancestral medusoid of Medusozoa.     

Morphology and the evolution of cycadeoidales

OneCycadeoidea stem one cycadeoidalean gynoecium and a bisporangiate cone attached to a slender cycadeoidalean trunkCycadeoidella japonica Ogura from the Cretaceous of Japan shows well-preserved internal structure that provides evidence for a better understanding of the morphological architecture of the cycadeoidalean plant. Structural details of the cone were confirmed. The ovule has an intergument enclosing a free nucellus and a thin outer envelope. Both reproductive and vegetative structures support the medullosan affinity of Cycadeoidales. The cone is interpreted as a compressed fertile shoot. Axillary cones characterizing some Cretaceous genera such asCycadeoidea andMonanthesia consist of a lateral shoot subtended by a frond that is the first leaf of the cone shoot itself. The origin of axillary buds in the Cycadeoidales is discussed. Heterochrony may have mediated the morphological changes that resulted in the establishment of the Cycadeoidales.     

Morphological and phylogenetic analysis of Raillietina spp. in indigenous chickens (Gallus gallus domesticus) in Bangladesh

Raillietina spp. (Cestoda: Davaineidae), the most common cestodes in indigenous chickens, cause a substantial production loss in poultry industry in Bangladesh. Here, we estimated the prevalence, confirmed the species and determined the genetic pattern of species of Raillietina using molecular tools. We collected and examined 375 chickens randomly from household of different villages of Mymensingh sadar and Gouripur upazila, Mymensingh district and adult parasites were isolated and identified. Genomic DNA was extracted from collected parasites, amplified ITS-2 and ND-1 genes, sequenced and analyzed. Out of 375 samples, 270 (72.0%) were found positive with Raillietina species and mean worm burden was 10.46 ± 0.56. Microscopically, three species of Raillietina, such as R. cesticillus (37.9%), R. echinobothrida (41.1%) and R. tetragona (52.8%) were detected on the basis of their morphological features. The total length, length and width of scolex, sucker and rostellum were also measured. Among different factors, age, farming nature and flock size of chickens were significantly (p < 0.05) influenced Raillietina infections. For further validation, the sequences of ITS-2 gene generated in this study were matched with reference sequences of R. cesticillus, R. echinobothrida and R. tetragona and found 99.63% − 100% similarity. The phylogenetic analyses of ITS-2 and ND-1 sequences were clustered together with the reference sequences of R. cesticillus, R. echinobothrida and R. tetragona confirming microscopic identification. This is the first confirmation of species of Raillietina along with the prevalence of the species, which will be helpful for the formulation of a control strategy and provide basic information for further molecular study.     

Serotonin immunoreactivity in the nervous system of the Pandora larva, the Prometheus larva, and the dwarf male of Symbion americanus (Cycliophora)

Cycliophora is a recently described phylum of enigmatic metazoans with a very complex life cycle that includes several sexual and asexual stages. Symbion pandora and Symbion americanus are the only two cycliophoran species hitherto described, of which morphological and genetic knowledge is still deficient to clarify the phylogenetic position of the phylum. Aiming to increase the database on the cycliophoran neural architecture, we investigated serotonin immunoreactivity in the free swimming Pandora larva, the Prometheus larva, and the adult dwarf male of S. americanus. In the larval forms, serotonin is mainly expressed in a ring-shaped pattern at the periphery of the antero-dorsal cerebral ganglion. Additionally, several serotonergic perikarya emerge from both sides of the cerebral ganglion. Thin neurites project anteriorly from the cerebral ganglion, while a pair of ventral longitudinal neurites emerges laterally and runs along the anterior-posterior body axis. Posteriorly, the ventral neurites fuse and extend as a posterior projection. In the dwarf male, serotonin is found mainly in the commissural neuropil of the large anterior cerebral ganglion. In addition, serotonin immunoreactivity is present in the most anterior region of the ventral neurites. Comparative analysis of spiralian nervous systems demonstrates that the neuroanatomy of the cycliophoran larval stages resembles much more the situation of adult rather than larval spiralians, which may be explained by secondary loss of larval structures and heterochronic shift of adult components into the nervous system of the Pandora and the Prometheus larva, respectively.     

Distribution and immunochemical character of neurotensin-like material in representative vertebrates and invertebrates: Apparent conservation of the COOH-terminal region during evolution

Using a radioimmunoassay for bovine neurotensin (NT) and various region specific antisera which react selectively with different portions of the molecule, the presence of immunoreactive NT (iNT) in a wide range of vertebrate and invertebrate species has been demonstrated. While antisera directed towards the N-terminal region of NT recognized only mammalian forms of NT, antisera directed towards the C-terminal region of NT recognized materials from all species examined, including representatives of all vertebrates and invertebrate classes. When extracts of the brain and gut of the vertebrates examined were chromatographed on Sephadex G-25 multiple NT-like substances were observed, and the patterns of iNT obtained seemed to fall into three groups: (a) mammals, (b) birds and reptiles and (c) amphibians and fish. Extracts of invertebrates also exhibited multiple peaks of iNT on Sephadex G-25 and the profiles observed resembled those for lower vertebrates. Partially purified iNT obtained from chicken, turtle, dogfish and lobster was shown to increase hematocrit and induce cyanosis in anesthetized rats. These findings indicate (a) that NT-like substances appear to be present throughout the animal kingdom, (b) that the C-terminal region of NT is highly conserved while the N-terminal region varies, and (c) that in any one animal multiple substances sharing C-terminal homologies with NT exist. These findings are consistent with the notion that NT and related peptides participate in important processes basic to animal life and that their functioning depends highly upon elements located in their C-terminal regions. They further suggest the existence of an entire family of NT-related peptides in each animal form, possibly distributed differently and functioning differently in the various organs of the animal.     

果实类型多样性的形成机制和进化

果实是被子植物特有的繁殖器官,果实的类型影响种子传播的模式、有效性和距离。果实类型的多样性在一定程度上造成了不同植物类群在生态上的差异,并对被子植物的进化速度和分化模式产生重要影响。本文对果实类型多样性形成中起关键作用的通路及其进化机制进行了综述,讨论了影响果实类型进化的各种因素,介绍了有关果实类型进化方向和模式的研究进展,为研究被子植物果实多样性的形成和演化提供新的思路。     

Evolutionary convergences and parallelisms: &;their theoretical differences and the difficulty &;of discriminating them in a practical &;phylogenetic context

The importance of evolutionary parallelisms and their differences from evolutionary convergences have been historically underappreciated, as recently noticed in Gould's last book `The structure of evolutionary history'. In that book, Gould make an effort to distinguish and to reinterpret these concepts in the light of the new discoveries of the last decades on developmental biology and genetics, presenting the elegant metaphor of `Pharaonic bricks versus Corinthian columns'. In this paper I will briefly discuss these concepts, and will argue that, despite the advances that have been made to define them in theory, it is rather hard to differentiate them in a practical phylogenetic context. In order to do so, I will provide some few examples from my own empirical studies on the last years of one of the most morphologically and taxonomically diverse groups of Vertebrates, the catfishes.     

Phylogenetic signatures of adaptation: The Arabis hirsuta species aggregate (Brassicaceae) revisited

The mustard family, Brassicaceae, is well-known for its homoplasy in almost any morphological character at practically all taxonomic levels. The genus Arabis, within the largest tribe of the Brassicaceae, is such an example comprising numerous para- and polyphyletic groups of taxa. Past research during the last 15 years has unraveled many phylogenetic relationships among the ∼550 (or more) species within the notoriously difficult tribe Arabideae. The European Arabis hirsuta species aggregate has remained unexplored, however. Herein we analyze phylogenetic relationships using nuclear ITS and plastid DNA sequences of Eurasian Arabis to characterize Hairy rock cress (A. hirsuta) and its relatives. Representative geographic sampling is used to study character and trait evolution, and bioclimatic data are used to differentiate between species. Our overview puts European Arabis into a reliable evolutionary framework, and we provide some striking insights into evolutionary trends and correlating morphological characters from seeds and flowers with environmental data such as climate variables and elevation. We demonstrate independent parallel evolution of sets of traits, and, therefore, we could further elaborate our previous findings that within tribe Arabideae high speciation rates are correlated with perennial growth form and occurrence at higher elevation. Finally some taxonomical remarks are provided to give added context.     

Origin,evolution, phylogeny and taxonomy of Pulex irritans

The human flea Pulex irritans Linnaeus, 1758 (Siphonaptera: Pulicidae) is one of the most studied species together with the cat flea Ctenocephalides felis Bouché, 1835, because they have a cosmopolitan distribution and are closely related to humans. The present study aimed to carry out a comparative morphometric and molecular study of two different populations of P. irritans (Spain and Argentina). Accordingly, internal transcribed spacer (ITS)1 and ITS2 of rDNA and the partial cytochrome c oxidase subunit 1 (cox1) and cytochrome b (cytb) mtDNA genes of these taxa were sequenced. Furthermore, the taxonomy, origin, evolution and phylogeny of P. irritans was assessed. The morphometric data obtained did not show significant differences between P. irritans specimens from Spain and Argentina, even when these two populations were collected from different hosts; however, there was a considerable degree of molecular divergence between both populations based on nuclear and mitochondrial markers. Thus, it is proposed that P. irritans, in contrast with other generalist fleas, maintains a certain degree of morphological similarity, at least between Western Palearctic and Neotropical areas. Furthermore, two well defined geographical genetic lineages within the P. irritans species are indicated, suggesting the existence of two cryptic species that could be discriminated by a polymerase chain reaction‐linked restriction fragment length polymorphism.     

Comparative morphology and evolution of the lungless caecilian Atretochoana eiselti (Taylor) (Amphibia: Gymnophiona: Typhlonectidae)

Atretochoana eiselti is a radically divergent aquatic caecilian until recendy known from only a single specimen from South America. In addition to its status as the largest lungless tetrapod known, and the only known lungless caecilian, this species has a suite of highly unusual morphological features that sets it apart from all other tetrapods, including sealed choanae (internal nostrils), complete loss of pulmonary arteries and veins, novel cranial architecture, and a novel stapedial muscle. The external, buccal, skeletal, muscular and cardiovascular anatomies of Atretochoana eiselti are described and compared to these features in other caecilians, particularly representatives of all typhlonectid genera which are its closest relatives. The comparative morphological data are used as a basis for interpretations of the ecology and evolution of Atretochoana eiselti. It is argued that lunglessness and the transition to cutaneous gas exchange is correlated with life in cold, montane, fast-flowing streams. Here, high oxygen concentrations and reduced metabolic rate serve to relax the physiological limitations on body size imposed by a reliance upon cutaneous gas-exchange, and lungs can produce disadvantageous buoyancy. Cranial evolution has increased the gape of Atretochoana eiselti relative to other caecilians, and seems likely to be associated with a shift in prey size and possibly type. Several modifications of the cranium appear to be associated with enhanced cranial kinesis in which a distinct cheek unit is highly mobile. The novel stapedial musculature is interpreted as contributing to this cranial kinesis. Respiratory and cranial evolution are argued to be correlated, with the ventilatory function of the buccopharyngeal pump constraining the evolution of the skull. The evolution of lunglessness removed this constraint facilitating repatterning of the skull.     

Evolutionary relationships among the Scarabaeini (Coleoptera: Scarabaeidae) based on combined molecular and morphological data

The Scarabaeini is an old world tribe of ball-rolling dung beetles that have origins dating back to at least the mid-upper Miocene (19-8 million years ago). The tribe has received little to no attention in morphological or molecular phylogenetics. We obtained sequence data from the mitochondrial cytochrome oxidase subunit I (1,197 bp) and 16S ribosomal RNA (461 bp) genes for 25 species of the Scarabaeini in an attempt to further resolve broad phylogenetic relationships within this tribe. Sequence data from both markers along with 216 morphological and 3 biological characters were analysed separately and combined. Independent analyses showed poorly resolved trees with many of the intermediate and basal nodes collapsed by low bootstrap values. Many sites in both genes exhibited strong A+T nucleotide bias and high interlineage divergences. The combined analysis revealed a number of well supported relationships such as the monophyly of the nocturnal species Scarabaeus satyrus, S. [Neateuchus] proboscideus, and S. zambesianus. Furthermore, the total evidence tree suggested to elevate S. (Pachysoma) to the status of an independent genus, Pachysoma, as a sister taxon to a clade containing Pachylomerus femoralis and Scarabaeus sensu lato. Within the latter, the following subgenera were maintained by the combination of data sets: S. (Scarabaeolus), S. (Sceliages), and S. (Kheper). Both, feeding specialisation and food relocation behaviour, were inferred to be polyphyletic in the Scarabaeini. Total evidence analysis found no support for common ancestry of Scarabaeini and Eucraniini.     

A functional analysis of compound eye evolution

New data on the phylogenetic relationships of various arthropod groups have spurred interesting attempts to reconstruct the evolution of arthropod nervous and visual systems. Some of the relevant new data are cell identities and developmental processes in the nervous and sensory systems, which is particularly useful for reconstructing the evolution of these systems. Here, we focus on the structure of compound eye ommatidia, and make an evolutionary analysis with functional arguments. We investigate possible routes of evolution that can be understood in terms of selection for improved visual function, and arrive at a number of conclusions that are discussed in the light of recent phylogenetic hypotheses. On the basis of ommatidial focusing structures and the arrangement of receptor cells we show that the evolution of compound eyes proceeded largely independently along at least two lineages from very primitive ancestors. A common ancestor of insects and crustaceans is likely to have had ommatidia with focusing crystalline cones, and colour and/or polarization vision. In contrast, the compound eyes in myriapods and chelicerates are likely to date back to ancestors with corneal lenses and probably without the ability to discriminate colour and polarization.     

Phylogeny of the major tetrapod groups: Morphological data and divergence dates

Summary The phylogeny of the major groups of tetrapods (amphibians, reptiles, birds, and mammals) has until recently been poorly understood. Cladistic analyses of morphological data are producing new hypotheses concerning the relationships of the major groups, with a focus on the identification of monophyletic groups. Molecular phylogenies support some of these views and dispute others. Geological dates of the major evolutionary branching points are recalculated on the basis of the cladograms and new fossil finds.     

The road to the Janiroidea: Comparative morphology and evolution of the asellote isopod crustaceans

This paper presents a new phylogenetic estimate of isopod crustaceans of the suborder Asellota with the aim of clarifying the evolution of the superfamily Janiroidea, a large and diverse group inhabiting all aqueous habitats. The phylogenetic analysis is based on a morphological evaluation of characters used in past classifications, as well as several new characters. The evolutionary polarity of the characters was determined by outgroup analysis. The characters employed were from the pleopods, the copulatory organs, the first walking legs, and the cephalon. The resulting character data set was analyzed with numerical phylogenetic computer programs to find one most parsimonious clado-gram, which is translated into a classification using the sequencing convention. The new phylogenetic estimate is significantly more parsimonious than previous trees from the literature, and several of its monophyletic groups have robust confidence limits. The superfamily Stenetrioidea belongs to the clade including the Janiroidea, not with the Aselloidea as previously suggested. The sister group of the Janiroidea is the family Pseudojaniridae, which is elevated to superfamily rank. The clade including the families Gnathostenetroididae and Protojaniridae is not the sister group of the Janiroidea, and is derived earlier in janiroidean evolution than the Stenetrioidea. Within the Janiroidea, the family Janiridae is not the most primitive taxon as previously believed. The clade including the families Munnidae and Pleurocopidae contains the earliest derived janiroideans. The data also indicate that the unusual sexual morphology of the Janiroidea did not appear suddenly but developed as a series of independent steps within the Asellota.     

Phylogenetic relationships, character evolution, and taxonomic implications within the slipper lobsters (Crustacea: Decapoda: Scyllaridae)

The slipper lobsters belong to the family Scyllaridae which contains a total of 20 genera and 89 species distributed across four subfamilies (Arctidinae, Ibacinae, Scyllarinae, and Theninae). We have collected nucleotide sequence data from regions of five different genes (16S, 18S, COI, 28S, H3) to estimate phylogenetic relationships among 54 species from the Scyllaridae with a focus on the species rich subfamily Scyllarinae. We have included in our analyses at least one representative from all 20 genera in the Scyllaridae and 35 of the 52 species within the Scyllarinae. Our resulting phylogenetic estimate shows the subfamilies are monophyletic, except for Ibacinae, which has paraphyletic relationships among genera. Many of the genera within the Scyllarinae form non-monophyletic groups, while the genera from all other subfamilies form well supported clades. We discuss the implications of this history on the evolution of morphological characters and ecological transitions (nearshore vs. offshore) within the slipper lobsters. Finally, we identify, through ancestral state character reconstructions, key morphological features diagnostic of the major clades of diversity within the Scyllaridae and relate this character evolution to current taxonomy and classification.