A giant stem-group chaetognath.

Chaetognaths, with their characteristic grasping spines, are the oldest known pelagic predators, found in the lowest Cambrian (Terreneuvian). Here, we describe a large stem chaetognath, Timorebestia koprii gen. et sp. nov., from the lower Cambrian Sirius Passet Lagerstätte, which exhibits lateral and caudal fins, a distinct head region with long antennae and a jaw apparatus similar to Amiskwia sagittiformis. Amiskwia has previously been interpreted as a total-group chaetognathiferan, as either a stem-chaetognath or gnathostomulid. We show that T. koprii shares a ventral ganglion with chaetognaths to the exclusion of other animal groups, firmly placing these fossils on the chaetognath stem. The large size (up to 30 cm) and gut contents in T. koprii suggest that early chaetognaths occupied a higher trophic position in pelagic food chains than today.

Homology of the head sensory structures between Heterotardigrada and Eutardigrada supported in a new species of water bear (Ramazzottiidae: Ramazzottius).

Phylum Tardigrada is represented by microscopic eight-legged panarthropods that inhabit terrestrial and marine environments. Although tardigrades are emerging model animals for areas of research including physiology, evolutionary biology, and astrobiology, knowledge of their external morphology remains insufficient. For instance, homologies between marine and terrestrial relatives largely remain unexplored. In the present study we provide detailed pictures of the head sensory organs in a new tardigrade, Ramazzottius groenlandensis sp. nov. Specimens were collected from a mixed moss and lichen sample on Ella Island, East Greenland. The new species differs from congeneric species in the presence of polygonal sculpturing on the dorsal cuticle, which is accentuated in the posterior region of the body, a lateral papilla on leg IV, and distinctive egg morphology. A Bayesian phylogenetic analysis (18S rRNA + 28S rRNA + COI) places the new species within the genus Ramazzottius with high confidence. Interestingly, the new species shows a full set of well-developed cephalic organs, which correspond to all sensory fields found in eutardigrades. Details on the full set of head organs were present only for heterotardigrades. The surface of these organs is covered with small pores, which presumably play a sensory role. This discovery suggests the homology of head sensory structures between heterotardigrades and eutardigrades, implying that the distinctive arrangement and positioning of sensory organs on the head is a plesiomorphic feature of tardigrades. Moreover, we find that the Ramazzottius oberhaeuseri morphotype forms a morphogroup, not a monophyletic species complex.

Cambrian lobopodians shed light on the origin of the tardigrade body plan.

Phylum Tardigrada (water bears), well known for their cryptobiosis, includes small invertebrates with four paired limbs and is divided into two classes: Eutardigrada and Heterotardigrada. The evolutionary origin of Tardigrada is known to lie within the lobopodians, which are extinct soft-bodied worms with lobopodous limbs mostly discovered at sites of exceptionally well-preserved fossils. Contrary to their closest relatives, onychophorans and euarthropods, the origin of morphological characters of tardigrades remains unclear, and detailed comparison with the lobopodians has not been well explored. Here, we present detailed morphological comparison between tardigrades and Cambrian lobopodians, with a phylogenetic analysis encompassing most of the lobopodians and three panarthropod phyla. The results indicate that the ancestral tardigrades likely had a Cambrian lobopodian-like morphology and shared most recent ancestry with the luolishaniids. Internal relationships within Tardigrada indicate that the ancestral tardigrade had a vermiform body shape without segmental plates, but possessed cuticular structures surrounding the mouth opening, and lobopodous legs terminating with claws, but without digits. This finding is in contrast to the long-standing stygarctid-like ancestor hypothesis. The highly compact and miniaturized body plan of tardigrades evolved after the tardigrade lineage diverged from an ancient shared ancestor with the luolishaniids.

The complete mitochondrial genome of the Arctic fairy shrimp Branchinectapaludosa (Müller, 1788) (Anostraca, Branchinectidae) from Sirius Passet, North Greenland.

Here we report the complete mitochondrial genome of the Arctic fairy shrimp, Branchinectapaludosa (Müller, 1788) (Anostraca, Branchinectidae), which was collected in the High Arctic of North Greenland. A complete 16,059 bp mitochondrion of B.paludosa was sequenced and assembled with the Illumina next generation sequencing platform. The B.paludosa mitogenome contains 13 PCGs, 22 tRNAs and 2 rRNA genes that are commonly observed in most metazoans and shows the conserved gene arrangement pattern of Anostraca. Our results of the phylogenomic analysis are consistent with the previous phylogenetic relationship, based on nuclear 18S ribosomal DNA. The B.paludosa mitogenome will be useful for understanding the geographical distribution and phylogenetic relationship of anostracans.

Characterization of the complete mitochondrial genome of the scale worm, Eunoe nodosa (Phyllodocida; Polynoidae) from the Beaufort Sea.

To increase the mitogenome data available for robust phylogeny, we sequenced the complete mitochondrial DNA of the scale worm Eunoe nodosa (Sars, 1861) in the family Polynoidae of the order Phyllodocida. The complete mitogenome has 15,366 bp and has 28.9% A, 13.2% C, 19.0% G, and 38.8% T. Using MITOS and tRNAscan-SE, we identified the 13 typical protein-coding genes (PCGs), 2 ribosomal RNA (rRNA) genes, 22 transfer RNA (tRNA) genes, and a non-coding region. Phylogenomic analysis based on 27 in-group taxa belonging to five families of the subclass Errantia show congruence with the published phylogenetic relationship within the Polynoidae, in which E. nodosa lies in the clade of shallow water species.

Integrative description of a new Dactylobiotus (Eutardigrada: Parachela) from Antarctica that reveals an intraspecific variation in tardigrade egg morphology.

Tardigrades constitute one of the most important group in the challenging Antarctic terrestrial ecosystem. Living in various habitats, tardigrades play major roles as consumers and decomposers in the trophic networks of Antarctic terrestrial and freshwater environments; yet we still know little about their biodiversity. The Eutardigrada is a species rich class, for which the eggshell morphology is one of the key morphological characters. Tardigrade egg morphology shows a diverse appearance, and it is known that, despite rare, intraspecific variation is caused by seasonality, epigenetics, and external environmental conditions. Here we report Dactylobiotus ovimutans sp. nov. from King George Island, Antarctica. Interestingly, we observed a range of eggshell morphologies from the new species, although the population was cultured under controlled laboratory condition. Thus, seasonality, environmental conditions, and food source are eliminated, leaving an epigenetic factor as a main cause for variability in this case.

Brain and eyes of Kerygmachela reveal protocerebral ancestry of the panarthropod head.

Recent discoveries of fossil nervous tissue in Cambrian fossils have allowed researchers to trace the origin and evolution of the complex arthropod head and brain based on stem groups close to the origin of the clade, rather than on extant, highly derived members. Here we show that Kerygmachela from Sirius Passet, North Greenland, a primitive stem-group euarthropod, exhibits a diminutive (protocerebral) brain that innervates both the eyes and frontal appendages. It has been surmised, based on developmental evidence, that the ancestor of vertebrates and arthropods had a tripartite brain, which is refuted by the fossil evidence presented here. Furthermore, based on the discovery of eyes in Kerygmachela, we suggest that the complex compound eyes in arthropods evolved from simple ocelli, present in onychophorans and tardigrades, rather than through the incorporation of a set of modified limbs.

Post-embryonic development of the Early Ordovician (ca. 480 Ma) trilobite Apatokephalus latilimbatus Peng, 1990 and the evolution of metamorphosis.

In many marine invertebrates metamorphosis entails a shift from a free-swimming larva to a benthic juvenile or adult. However, how the metamorphosis-entailing "indirect development" in arthropods arose from direct-developing ancestor is poorly understood. Trilobites left a rich fossil record, and some trilobite lineages had a metamorphosis-undergoing early developmental stage, termed the "asaphoid protaspis"-stage, providing a good opportunity to elucidate the rise of indirect development. Among others, the Ordovician representatives of Remopleuridioidea are known to possess a highly bulbous "asaphoid protaspis," while the Furongian (Late Cambrian) remopleuridioidean genus Haniwa did not possess it. Here we show the post-embryonic development of the remopleuridioidean trilobite, Apatokephalus latilimbatus, from the Tremadocian (485.4 Ma-477.7 Ma) Dongjeom Formation, Korea. The post-embryonic development of A. latilimbatus contains a free-swimming "commutavi protaspis" (a term replacing "asaphoid protaspis"). Interestingly, the earlier protaspid stage shows more similar morphology and size to the meraspis than the commutavi protaspid stage does. This indicates that the commutavi protaspid stage was intercalated into the ancestral direct development as a specialized stage for a better dispersal, and thus the "commutavi protaspis" of A. latilimbatus represents the initial phase of the evolution of indirect development. The duration of the free-swimming phase became longer in more derived remoplueridioidean trilobites, implying that the intercalated free-swimming strategy became emphasized during subsequent evolution. The morphological gap between the commutavi protaspis and the subsequent earliest meraspis provides a convincing case for the "selective independence" of developmental stages, explaining the various morphologies of commutavi protaspides in many trilobite lineages.