Rhabdocoels comprise a large group of flatworms that currently consists of two major subgroups: (1) Kalyptorhynchia with an anterior proboscis and (2) Dalytyphloplanida without a proboscis. Most ...genera of rhabdocoels can easily be classified into one of these two subgroups, except for the three monotypic representatives of the Mariplanellinae. Comparative morphological data and previous molecular phylogenetic analyses have so far been unable to determine the positions of these rarely encountered microturbellarians within the context of rhabdocoels. Here we describe a new species of Mariplanellinae,
Mariplanella piscadera
sp. nov. from Curaçao (Dutch Caribbean), and present an updated phylogeny inferred from 18S and 28S rDNA sequences, including data from
M. piscadera
sp. nov.,
M. frisia
, and a selection of rhabdocoels, proseriates and other trepaxonematan flatworms. Our molecular phylogenetic trees reveal that
Mariplanella
belongs to a distinct higher-level group of rhabdocoels that forms the sister lineage to a clade uniting Kalyptorhynchia and Dalytyphloplanida. Therefore, we update the status of Mariplanellinae to Mariplanellida
status novus
, containing the family Mariplanellidae
status novus
, in order to better reflect our current understanding of rhabdocoel phylogeny and the establishment of now three major subgroups.
Modern microbialites are complex microbial communities that interface with abiotic factors to form carbonate-rich organosedimentary structures whose ancestors provide the earliest evidence of life. ...Past studies primarily on marine microbialites have inventoried diverse taxa and metabolic pathways, but it is unclear which of these are members of the microbialite community and which are introduced from adjacent environments. Here we control for these factors by sampling the surrounding water and nearby sediment, in addition to the microbialites and use a metagenomics approach to interrogate the microbial community. Our findings suggest that the Pavilion Lake microbialite community profile, metabolic potential and pathway distributions are distinct from those in the neighboring sediments and water. Based on RefSeq classification, members of the Proteobacteria (e.g., alpha and delta classes) were the dominant taxa in the microbialites, and possessed novel functional guilds associated with the metabolism of heavy metals, antibiotic resistance, primary alcohol biosynthesis and urea metabolism; the latter may help drive biomineralization. Urea metabolism within Pavilion Lake microbialites is a feature not previously associated in other microbialites. The microbialite communities were also significantly enriched for cyanobacteria and acidobacteria, which likely play an important role in biomineralization. Additional findings suggest that Pavilion Lake microbialites are under viral selection as genes associated with viral infection (e.g CRISPR-Cas, phage shock and phage excision) are abundant within the microbialite metagenomes. The morphology of Pavilion Lake microbialites changes dramatically with depth; yet, metagenomic data did not vary significantly by morphology or depth, indicating that microbialite morphology is altered by other factors, perhaps transcriptional differences or abiotic conditions. This work provides a comprehensive metagenomic perspective of the interactions and differences between microbialites and their surrounding environment, and reveals the distinct nature of these complex communities.
Dinoflagellates represent a major lineage of unicellular eukaryotes with unparalleled diversity and complexity in morphological features. The monophyly of dinoflagellates has been convincingly ...demonstrated, but the interrelationships among dinoflagellate lineages still remain largely unresolved. Warnowiid dinoflagellates are among the most remarkable eukaryotes known because of their possession of highly elaborate ultrastructural systems: pistons, nematocysts, and ocelloids. Complex organelles like these are evolutionary innovations found only in a few athecate dinoflagellates. Moreover, the taxonomy of warnowiids is extremely confusing and inferences about the evolutionary history of this lineage are mired by the absence of molecular phylogenetic data from any member of the group. In this study, we provide the first molecular phylogenetic data for warnowiids and couple them with a review of warnowiid morphological features in order to formulate a hypothetical framework for understanding character evolution within the group. These data also enabled us to evaluate the evolutionary relationship(s) between warnowiids and the other group of dinoflagellates with complex organelles: polykrikoids.
Molecular phylogenetic analyses of SSU and LSU rDNA sequences demonstrated that warnowiids form a well-supported clade that falls within the more inclusive Gymnodinium sensu stricto clade. These data also confirmed that polykrikoids are members of the Gymnodinium sensu stricto clade as well; however, a specific sister relationship between the warnowiid clade and the polykrikoid clade was unresolved in all of our analyses. Nonetheless, the new DNA sequences from different isolates of warnowiids provided organismal anchors for several previously unidentified sequences derived from environmental DNA surveys of marine biodiversity.
Comparative morphological data and molecular phylogenetic data demonstrate that the polykrikoid and the warnowiid clade are closely related to each other, but the precise branching order within the Gymnodinium sensu stricto clade remains unresolved. We regard the ocelloid as the best synapomorphy for warnowiids and infer that the most recent common ancestor of polykrikoids and warnowiids possessed both nematocysts and photosynthetic plastids that were subsequently lost during the early evolution of warnowiids. Our summary of species and genus concepts in warnowiids demonstrate that the systematics of this poorly understood group is highly problematic and a comprehensive revision is needed.
Environmental DNA surveys have revealed a great deal of hidden diversity within the Cercozoa. An investigation into the biodiversity of heterotrophic flagellates in marine benthic habitats of British ...Columbia, Canada, demonstrated the presence of several undescribed taxa with morphological features that resemble the cercozoan genera Cryothecomonas and Protaspis. Nine novel species of marine interstitial cercozoans are described that are distributed into five genera, four of which are new. Phylogenetic analyses of small subunit rDNA sequences derived from two uncultured isolates of Protaspis obliqua and nine novel cercozoan species (within four novel genera) provided organismal anchors that helped establish the cellular identities of several different environmental sequence clades. These data, however, also showed that the rarity of distinctive morphological features in cryomonads, and other groups of cercozoans, makes the identification and systematics of the group very difficult. Therefore, a DNA barcoding approach was applied as a diagnostic tool for species delimitation that used a 618 bp region at the 5' end of the SSU rDNA sequence. Nucleotide sequence analysis of this region showed high intergeneric sequence divergences of about 7% and very low intraspecific sequence divergences of 0-0.5%; phylogenetic analyses inferred from this barcoding region showed very similar tree topologies to those inferred from the full-length sequence of the gene. Overall, this study indicated that the 618 bp barcoding region of SSU rDNA sequences is a useful molecular signature for understanding the biodiversity and interrelationships of marine benthic cercozoans.
Marine microturbellarians are an assemblage of meiofaunal flatworms abundant in sediments and on seaweeds around the world. The diversity and distribution of these animals in Japan are poorly ...understood. Here, we provide an overview of all recorded species in Japan and characterize two new species of the rhabdocoel genus Reinhardorhynchus based on morphological features and a molecular phylogeny inferred from 18S and 28S rDNA sequences. Reinhardorhynchus ryukyuensis sp. nov. can be distinguished from other species in the genus by the lack of an armed cirrus and by the presence of two larger opposing hooks and five smaller interconnected hooks in its male copulatory organ. Reinhardorhynchus sagamianus sp. nov. differs from its congeners because its male copulatory organ combines a bipartite cirrus armed with a belt of overlapping scale-like spines, an unarmed accessory cirrus, and two large distal accessory hooks. Our molecular phylogenetic analyses show that R. ryukyuensis sp. nov. and R. sagamianus sp. nov. form a clade with all the other species of Reinhardorhynchus for which DNA sequence data are available. Within this clade, R. sagamianus sp. nov. is in a clade that also includes R. riegeri and R. anamariae . The discovery of these new species highlights the importance of uncovering and documenting the hidden biodiversity along Japan’s coastal margin.
Surveys of kinorhynch diversity in the northeastern Pacific Ocean are rare, and only eight species have been described from the region so far. We explored the diversity of kinorhynchs from a pristine ...coastal habitat of British Columbia (Calvert Island) and discovered a new subtidal species,
Echinoderes hakaiensis
sp. nov. We also redescribed one of the oldest described echinoderid species,
E. pennaki
. Both species were characterized with high-resolution light microscopy (LM), scanning electron microscopy (SEM), and DNA sequences from the mitochondrial cytochrome c oxidase subunit 1 (COI) gene.
Echinoderes hakaiensis
sp. nov. can be differentiated from other species by a unique pattern of spines and tubes (i.e., four pairs of tubes on segment 2 combined with three middorsal spines).
Echinoderes pennaki
shows a configuration of traits on segment 2 that questions the reliability of conventional morphological characters used to distinguish different genera within the Echinoderidae. Reevaluation of these characters will require a resolved molecular phylogenetic context including a more comprehensive sampling of species within the group.
Kinorhynch segmentation differs from the patterns found in Chordata, Arthropoda and Annelida which have coeloms and circulatory systems. Due to these differences and their obsolete status as ...'Aschelminthes', the microscopic kinorhynchs are often not acknowledged as segmented bilaterians. Yet, morphological studies have shown a conserved segmental arrangement of ectodermal and mesodermal organ systems with spatial correspondence along the anterior-posterior axis. However, a few aberrant kinorhynch lineages present a worm-like body plan with thin cuticle and less distinct segmentation, and thus their study may aid to shed new light on the evolution of segmental patterns within Kinorhyncha.
Here we found the nervous system in the aberrant Cateria styx and Franciscideres kalenesos to be clearly segmental, and similar to those of non-aberrant kinorhynchs; hereby not mirroring their otherwise aberrant and posteriorly shifted myoanatomy. In Zelinkaderes yong, however, the segmental arrangement of the nervous system is also shifted posteriorly and misaligned with respect to the cuticular segmentation.
The morphological disparity together with the distant phylogenetic positions of F. kalenesos, C. styx and Z. yong support a convergent origin of aberrant appearances and segmental mismatches within Kinorhyncha.
The eugregarines are a group of apicomplexan parasites that mostly infect the intestines of invertebrates. The high level of morphological variation found within and among species of eugregarines ...makes it difficult to find consistent and reliable traits that unite even closely related lineages. Based mostly on traits observed with light microscopy, the majority of described eugregarines from marine invertebrates has been classified into a single group, the Lecudinidae. Our understanding of the overall diversity and phylogenetic relationships of lecudinids is very poor, mainly because only a modest amount of exploratory research has been done on the group and very few species of lecudinids have been characterized at the molecular phylogenetic level. In an attempt to understand the diversity of marine gregarines better, we surveyed lecudinids that infect the intestines of Pacific ascidians (i.e. sea squirts) using ultrastructural and molecular phylogenetic approaches; currently, these species fall within one genus, Lankesteria. We collected lecudinid gregarines from six ascidian host species, and our data demonstrated that each host was infected by a different species of Lankesteria: (i) Lankesteria hesperidiiformis sp. nov., isolated from Distaplia occidentalis, (ii) Lankesteria metandrocarpae sp. nov., isolated from Metandrocarpa taylori, (iii) Lankesteria halocynthiae sp. nov., isolated from Halocynthia aurantium, (iv) Lankesteria herdmaniae sp. nov., isolated from Herdmania momus, (v) Lankesteria cf. ritterellae, isolated from Ritterella rubra, and (vi) Lankesteria didemni sp. nov., isolated from Didemnum vexillum. Visualization of the trophozoites with scanning electron microscopy showed that four of these species were covered with epicytic folds, whereas two of the species were covered with a dense pattern of epicytic knobs. The molecular phylogenetic data suggested that species of Lankesteria with surface knobs form a clade that is nested within a paraphyletic assemblage species of Lankesteria with epicytic folds.
Several lineages of euglenozoans are enveloped with epibiotic bacteria and live in low oxygen and anoxic marine sediments, such as Bihospites bacati and Calkinsia aureus. A combination of shared ...ultrastructural traits and molecular phylogenetic inferences demonstrate that these lineages belong to a clade called the “Symbiontida.” Bihospites and Calkinsia possess all of the synapomorphies for the Euglenozoa plus several novel traits. Bihospites has a distinctive cell surface organization reminiscent of the pellicle strips in euglenids, a robust C-shaped feeding apparatus that encircles the nucleus, and a diverse community of epibiotic bacteria. Calkinsia has a novel “extrusomal pocket” and a thick (orange) extracellular matrix beneath a uniform layer of epibiotic bacteria. Despite the absence of molecular phylogenetic data, similar ultrastructural traits in Postgaardi mariagerensis and its epibiotic bacteria strongly suggest that this species is also a member of the Symbiontida. Molecular phylogenetic trees inferred from small subunit (SSU) ribosomal DNA sequences have shown that Bihospites and Calkinsia group strongly with a diverse set of environmental DNA sequences (eDNA) generated from low-oxygen marine samples collected at different depths from different locations around the world. These data demonstrate a diverse array of symbiontids that have yet to be characterized at the genomic, cellular, and behavior levels, which underscores how poorly we currently understand the biology and ecology of the group. Moreover, current data suggest that the communities of epibiotic bacteria associated with Bihospites, Calkinsia, and Postgaardi co-evolved with their hosts and are metabolically integrated with modified mitochondria positioned immediately beneath the host's plasma membrane. No symbiontid species has ever been cultivated, so improved knowledge about these eukaryotic organisms and their intimate relationships with bacteria in low oxygen environments will likely be achieved using culture-independent approaches, such as isolated-cell metagenomics.
Gregarine apicomplexans are unicellular parasites commonly found in the intestines and coeloms of invertebrate hosts. Traits associated with the conspicuous feeding stage of gregarines, known as the ...trophozoite, have been used in combination with molecular phylogenetic data for species delimitation and the reconstruction of evolutionary history. Trophozoite morphology alone is often inadequate for inferring phylogenetic relationships and delimiting species due to frequent cases of high intraspecific variation combined with relatively low interspecific variation. The current study combined morphological data with small subunit (SSU) rDNA sequences to describe and establish two novel marine gregarine species isolated from the intestine of a polychaete host Lumbrineris inflata collected in British Columbia (Canada): Paralecudina anankea n. sp. and Lecudina caspera n. sp. The sister species to the host is Lumbrineris japonica, which can be found on the opposite side of the Pacific Ocean (Japan) and contains two different species of gregarine parasites: Paralecudina polymorpha and Lecudina longissima. Molecular phylogenetic analyses placed P. anankea n. sp. as the sister species to P. polymorpha and L. caspera n. sp. as the sister species to L. longissima. This phylogenetic pattern demonstrates a co-evolutionary history whereby speciation of the host (Lumbrineris) corresponds with simultaneous speciation of the two different lineages of intestinal gregarines (Paralecudina and Lecudina).