A new species of Euclymene (Maldanidae, Annelida) from Brazil, with new combinations, and phylogenetic implications for Euclymeninae.

Maldanids are tube-building polychaetes, known as bamboo-worms; inhabit diverse marine regions throughout the world. The subfamily Euclymeninae was proposed to include forms with anal and cephalic plates, a funnel-shaped pygidium, and a terminal anus. Euclymene, the type genus of Euclymeninae, has about 18 valid species. Euclymene vidali sp. nov. is defined and members of the species described from Northeastern Brazil. Members of this species have 23 chaetigers, and one pre-pygidial achaetous segment; nuchal grooves extend through three quarters of the cephalic plate, and there is one acicular spine with a denticulate tip. Euclymene africana, and E. watsoni, are here recognized, respectively, as Isocirrus africana comb. nov., and I. watsoni comb. nov. Three monotypic genera are invalid: Macroclymenella, Eupraxillella, and Pseudoclyemene; their species should be recognized as Clymenella stewartensis com. nov., Praxillella antarctica com. nov., and Praxillela quadrilobata com. nov., respectively. An identification key and a comparative table for all species of Euclymene are provided. A comparative table for all genera of Euclymeninae is also furnished. The paraphyletic status of Euclymene and Euclymeninae is discussed. The taxon Maldanoplaca is not code compliant and should only be regarded as an informal name.

Phylogenetic Inference and the Misplaced Premise of Substitution Rates.

Three competing 'methods' have been endorsed for inferring phylogenetic hypotheses: parsimony, likelihood, and Bayesianism. The latter two have been claimed superior because they take into account rates of sequence substitution. Can rates of substitution be justified on its own accord in inferences of explanatory hypotheses? Answering this question requires addressing four issues: (1) the aim of scientific inquiry, (2) the nature of why-questions, (3) explanatory hypotheses as answers to why-questions, and (4) acknowledging that neither parsimony, likelihood, nor Bayesianism are inferential actions leading to explanatory hypotheses. The aim of scientific inquiry is to acquire causal understanding of effects. Observation statements of organismal characters lead to implicit or explicit why-questions. Those questions, conveyed in data matrices, assume the truth of observation statements, which is contrary to subsequently invoking substitution rates within inferences to phylogenetic hypotheses. Inferences of explanatory hypotheses are abductive in form, such that some version of an evolutionary theory(ies) is/are included or implied. If rates of sequence evolution are to be considered, it must be done prior to, rather than within abduction, which requires renaming those putatively-shared nucleotides subject to substitution rates. There are, however, no epistemic grounds for renaming characters to accommodate rates, calling into question the legitimacy of causally accounting for sequence data.

Who's who in Magelona: phylogenetic hypotheses under Magelonidae Cunningham & Ramage, 1888 (Annelida: Polychaeta).

Known as shovel head worms, members of Magelonidae comprise a group of polychaetes readily recognised by the uniquely shaped, dorso-ventrally flattened prostomium and paired ventro-laterally inserted papillated palps. The present study is the first published account of inferences of phylogenetic hypotheses within Magelonidae. Members of 72 species of Magelona and two species of Octomagelona were included, with outgroups including members of one species of Chaetopteridae and four of Spionidae. The phylogenetic inferences were performed to causally account for 176 characters distributed among 79 subjects, and produced 2,417,600 cladograms, each with 404 steps. A formal definition of Magelonidae is provided, represented by a composite phylogenetic hypothesis explaining seven synapomorphies: shovel-shaped prostomium, prostomial ridges, absence of nuchal organs, ventral insertion of palps and their papillation, presence of a burrowing organ, and unique body regionation. Octomagelona is synonymised with Magelona due to the latter being paraphyletic relative to the former. The consequence is that Magelonidae is monotypic, such that Magelona cannot be formally defined as associated with any phylogenetic hypotheses. As such, the latter name is an empirically empty placeholder, but because of the binomial name requirement mandated by the International Code of Zoological Nomenclature, the definition is identical to that of Magelonidae. Several key features for future descriptions are suggested: prostomial dimensions, presence/absence of prostomial horns, morphology of anterior lamellae, presence/absence of specialised chaetae, and lateral abdominal pouches. Additionally, great care must be taken to fully describe and illustrate all thoracic chaetigers in descriptions.

Taking a closer look: an SEM review of Levinsenia species (Polychaeta: Paraonidae) reported from California.

Scanning electron (SEM) and light microscope examinations of members of Levinsenia Mesnil, 1897, species from California yielded a new species, new characters, emended name and range extension for L. kirbyorum Lovell, 2002. Specimens of L. gracilis (Tauber, 1879) from Sweden, Iceland, and California were compared and could not be distinguished on the basis of morphology. Two other Californian species, L. multibranchiata (Hartman, 1957) and L. oculata (Hartman, 1957), were also examined. SEM revealed features previously undescribed for the genus. Additional prostomial ciliary bundles, dorsal transverse ciliary branchial connections, notopodial sensory pores, and neurochaetal fascicle configurations. Levinsenia barwicki n.sp. possessing a terminal sensory organ, 4-8 leaf-like ciliate branchiae, and recurved neurochaete with distal hood is described More SEM work is necessary to confirm if these features are present among other members of Levinsenia and other Paraonidae genera. The status of Levinsenia according to the phylogenetic analysis performed by Langeneck et al. (2019, Molecular Phylogenetics and Evolution, 136, 1-13) is discussed.

Sequence Data, Phylogenetic Inference, and Implications of Downward Causation.

Framing systematics as a field consistent with scientific inquiry entails that inferences of phylogenetic hypotheses have the goal of producing accounts of past causal events that explain differentially shared characters among organisms. Linking observations of characters to inferences occurs by way of why-questions implied by data matrices. Because of their form, why-questions require the use of common-cause theories. Such theories in phylogenetic inferences include natural selection and genetic drift. Selection or drift can explain 'morphological' characters but selection cannot be causally applied to sequences since fitness differences cannot be directly associated with individual nucleotides or amino acids. The relation of selection to sequence data is by way of downward or top-down causation from those phenotypes upon which selection occurs. The application of phylogenetic inference to explain sequence data is thus restricted to instances where drift is the relevant theory; those nucleotides or amino acids that can be explained via downward causation are precluded from inclusion in the data matrix. The restrictions on the inclusion of sequence data in phylogenetic inferences equally apply to species hypotheses, precluding the more restrictive approach known as DNA barcoding. Not being able to discern drift and selection as relevant causal mechanisms can severely constrain the inclusion and explanations of sequence data. Implications of such exclusion are discussed in relation to the requirement of total evidence.

Inference of phylogenetic relationships within Fabriciidae (Sabellida, Annelida) using molecular and morphological data.

Cladistic relationships among fabriciids have to date been explored in the context of adult morphology, but resolution has been declining as more species are described. In this study, we incorporated data on the reproductive system, including features related to the male sperm and sperm storage by females, to supplement existing data on adult morphology (for a total of 50 characters). Three nuclear DNA markers (18S rDNA approximately 1800 bp, the D1 region of 28S rDNA approximately 320 bp, and histone H3 approximately 330 bp) were sequenced from 21 species of fabriciids. We assessed the phylogeny of Fabriciidae based on an integrative analysis of these morphological and molecular characters. Our results show that, in addition to three previously recovered apomorphies for Fabriciidae (absence of ventral lips, modification of abdominal uncini to an elongate manubrium, and presence of branchial hearts), six more apomorphies associated with the reproductive system can be used to support this clade-spermiogenesis only in the thorax, spermiogenesis in large clusters with a central cytophore, single dorsal sperm duct, sperm nuclear projection, thickening of the sperm nuclear membrane and the sperm extra-axonemal sheath. The results require the erection of two new genera and two new species, which are described. © The Willi Hennig Society 2010.

Species as explanatory hypotheses: refinements and implications.

The formal definition of species as explanatory hypotheses presented by Fitzhugh (Marine Biol 26:155-165, 2005a, b) is emended. A species is an explanatory account of the occurrences of the same character(s) among gonochoristic or cross-fertilizing hermaphroditic individuals by way of character origin and subsequent fixation during tokogeny. In addition to species, biological systematics also employs hypotheses that are ontogenetic, tokogenetic, intraspecific, and phylogenetic, each of which provides explanatory hypotheses for distinctly different classes of causal questions. It is suggested that species hypotheses can not be applied to organisms with obligate asexual, parthenogenetic, and self-fertilizing modes of reproduction. Hypotheses explaining shared characters among such organisms are, instead, strictly phylogenetic. Several implications of this emended definition are examined, especially the relations between species, intraspecific, and phylogenetic hypotheses, as well as the limitations of species names to be applied to temporally different characters within populations.

Problems Determining the Phylogenetic Position of Echiurans and Pogonophorans with Limited Data.

The phylogenetic position of Echiura and Pogonophora was recently reconsidered using 346 bp of elongation factor 1-alpha (EF-1α). On the basis of these results alone it was suggested that the relationships of Anne lida be reconsidered and that the systematics of the group changed accordingly. An examination of these data, however, reveals difficulties in the original analysis that relate to weighting and the relative inclusion or exclusion of available characters and of taxa. Wholesale revision of the systematics of Annelida or of groups therein awaits the availability of more comprehensive work.