Tergipedidae represents a diverse and successful group of aeolid nudibranchs, with approximately 200 species distributed throughout most marine ecosystems and spanning all biogeographical regions of the oceans. However, the systematics of this family remains poorly understood since no modern phylogenetic study has been undertaken to support any of the proposed classifications. The present study is the first molecular phylogeny of Tergipedidae based on partial sequences of two mitochondrial (COI and 16S) genes and one nuclear gene (H3). Maximum likelihood, maximum parsimony and Bayesian analysis were conducted in order to elucidate the systematics of this family. Our results do not recover the traditional Tergipedidae as monophyletic, since it belongs to a larger clade that includes the families Eubranchidae, Fionidae and Calmidae. This newly recovered clade is here referred to as Fionidae, the oldest name for this taxon. In addition, the present molecular phylogeny does not recover the traditional systematic relationships at a generic level, and therefore, systematic changes are required. We recognize the following clades within Fionidae: Calma, Cuthona, Cuthonella, Eubranchus, Fiona, Murmania, Tenellia, Tergipes, Tergiposacca gen. nov., Rubramoena gen. nov. and Abronica gen. nov. The type species of Tergiposacca, T. longicerata nov. sp. is described. The other two new genera have a previously described species as their type species. Most of these taxa, with the exceptions of Eubranchus, Tergipes and Fiona are composed of radically different constituent species from their traditional membership, but appear to be supported by morphological synapomorphies as well as molecular data. Aenigmastyletus, Catriona, Phestilla, Tenellia and Trinchesia are nested within other clades and, thus are here considered as synonyms of the larger clades. The phylogenetic position and validity of Myja, Guyvalvoria, Leostyletus and Subcuthona still need to be tested in future studies when material becomes available.
The taxonomy of common northern nudibranch molluscs of the genus Dendronotus in the vast cold regions of Eurasia remains largely unknown. Abundant material collected in many localities from the Barents Sea, via the Arctic region, to the north-west Pacific was analysed for the first time. An integrated approach combining morphological and ontogenetic data with molecular four-gene (COI, 16S, H3, and 28S) analysis reveals seven species, including three previously undescribed. Dendronotus frondosus (Ascanius, 1774) and Dendronotus dalli Bergh, 1879 were commonly considered as amphiboreal species; however, according to this study they are restricted to the North Atlantic and the North Pacific, respectively. In the north-west Pacific two new species were discovered, Dendronotus kamchaticus sp. nov. and Dendronotus kalikal sp. nov., that are externally similar to D. frondosus, but that show significant distance according to molecular analysis and are considerably different in radular morphology. In the North Atlantic a new species Dendronotus niveus sp. nov., sibling to North Pacific D. dalli, is revealed. The separate status of North Atlantic Dendronotus lacteus (Thompson, 1840) is confirmed, including considerable range extension. The essential similarity of early ontogenetic stages of radular development common for species with disparate adult radular morphology (such as D. frondosus and D. dalli) is shown, and its importance for taxonomy is discussed.
Species of the genus Dendronotus are among the most common nudibranchs in the northern Hemisphere. However, their distribution and composition in the Northwest Pacific remain poorly explored. In the present study, we observed Dendronotus composition in northwestern part of the Sea of Japan, using an integrative approach, included morphological and molecular phylogenetic analyses and molecular species delimitation methods. These multiple methods revealed high cryptic diversity within the genus. Two specimens of Dendronotus frondosus were found in Amursky Bay and therefore its amphiboreal status was confirmed. In three locations of the Sea of Japan we found specimens, which are very close externally to D. frondosus, but show significant distance according to molecular analysis. We show that these specimens belong to a new species Dendronotus dudkai sp.n. This species is sister to D. frondosus according to morphological and molecular data, therefore the question of sympatric coexistence is discussed. For the first time Dendronotus kamchaticus was registered in the Sea of Japan and updated information of some intraspecific variation of this species is provided. How to cite this article:
Revealing the mechanisms of life cycle changes is critical for understanding the processes driving hydrozoan evolution. Our analysis of mitochondrial (COI, 16S) and nuclear (ITS1 and ITS2) gene fragments resulted in the discovery of unique polymorphism in the life cycle of Sarsia lovenii from the White Sea. This polymorphic species exhibits two types of gonophores: hydroids produce both free-swimming medusae and attached medusoids (phenotypic polymorphism). Our phylogenetic analysis revealed the intrinsic genetic structure of S. lovenii (genetic polymorphism). Two haplogroups inhabiting the White Sea differ in their reproductive modes. Haplogroup 1 produces attached medusoids, and haplogroup 2 produces free-swimming medusae. Our experiments indicated the possibility of free interbreeding between haplogroups that likely is a rare event in the sea. We propose that inter-haplogroup crossing of S. lovenii in the White Sea may be limited by discordance in periods of spawning or by spatial differences in habitat of spawning specimens. Our finding can be interpreted as a case of nascent speciation that illustrates the patterns of repeated medusa loss in hydrozoan evolution. Life cycle traits of S. lovenii may be useful for elucidating the molecular mechanisms of medusa reduction in hydrozoans.
In this paper we perform an integrative investigation of the Eubranchus odhneri -Eubranchus sanjuanensis species complex. Several specimens morphologically similar to E. sanjuanensis were collected in the Sea of Japan. However, preliminary molecular data indicated that they were closer to the Arctic species E. odhneri. For our analysis we obtained new data on mitochondrial COI and 16S rRNA, and nuclear histone H3 and 18S rRNA molecular markers to reconstruct their phylogeny (Bayesian Inference and Maximum Likelihood approaches), delimit species with ASAP, bPTP and GMYC analyses and estimate divergence times of putative species. We also studied morphological and ecological features including external morphology, colouration, jaw and radular morphology, configuration of the reproductive system, ultrastructure of cnidosacs, and determination of possible diet preferences. Our results show that specimens from the Sea of Japan represent a new species described herein, Eubranchus malakhovi sp.n. It differs both morphologically and genetically from its closely related species, the Arctic E. odhneri and the North-East Pacific E. sanjuanensis. The new species is closely related to E. odhneri with low mtDNA divergence, and our divergence time estimations suggest these species split ~1.5 Mya.
Nudibranch molluscs of the genus Dendronotus Alder and Hancock, 1845 are widely distributed in the Northern Hemisphere. Taxonomic studies on the genus Dendronotus have been problematic due to high variability in the colour pattern of many species, as well as in the external morphology and anatomy. In the present paper, we studied specimens of Dendronotus from northern Pacific presumably belonging to the species Dendronotus albus MacFarland, 1966 (white frond-aeolis). Molecular and morphological data revealed the existence of two distinct species among the material examined: D. albus, which has a wide range from Kamchatka and the Kurile Islands (from where we report this species for the first time) to California in North America, and the pseudocryptic species Dendronotus diversicolor Robilliard, 1970 (multicolor frond-aeolis), which has been previously considered a junior synonym of D. albus. Dendronotus diversicolor occurs from California, USA, to British Columbia, Canada, in sympatry with D. albus. Dendronotus albus and D. diversicolor can be clearly distinguished by colour pattern, internal and external morphology, and molecular sequence data. Despite some similarities in radular and external morphology between D. albus and D. diversicolor, these two species are phylogenetically distant and belong to different clades within the genus Dendronotus, which suggests convergent evolution.
Nudibranch mollusks (Gastropoda: Heterobranchia) are widely known for their ability to incorporate some active biochemical compounds of their prey, or even organelles and symbionts of the prey, which assured biological success of this group. At the same time, the process of nematocysts obtaining and incorporation into specific structures called cnidosacs by cladobranch mollusks remain poorly studied. This highlights a necessity of additional ultrastructural studies of cnidosac and adjacent organs in various aeolid mollusks using modern microscopic methods as they may provide new insight into the cnidosac diversity and fine‐scale dynamics of nematocysts sequestration process. The present study is focused on the general and fine structure of the cnidosac area in cladobranch Aeolidia papillosa (Aeolidiidae). Specific goals of our study were to provide a detailed description of histological and ultrafine structure of epidermis, upper parts of the digestive glands and the cnidosac, its innervation and proliferation using standard histological techniques, confocal laser scanning microscopy (CLSM) and transmission electron microscopy. Our results clearly demonstrated that A. papillosa cnidosac is a much more complex structure, than it was thought, especially compared with simple cnidosacs found in flabellinids and facelinids. Using CLSM for functional morphological analysis provides a better resolution in visualization of structural elements within a cnidosac compared with traditional histological techniques. We revealed the presence of two cell types in the cnidophage zone: cnidophages and interstitial cells, which differ in ultrastructure and function. Our results also document the presence of a specific cnidopore zone, lined with differentiated cuboid epithelium bearing long microvilli, which likely provides a unidirectional flow of nematocysts during kleptocnides extrusion. For the first time, occurrence of vacuoles containing protective chitinous spindles in the cnidosac epithelium was shown.
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