Evolutionary History of Chemosensory-Related Gene Families across the Arthropoda.

Chemosensory-related gene (CRG) families have been studied extensively in insects, but their evolutionary history across the Arthropoda had remained relatively unexplored. Here, we address current hypotheses and prior conclusions on CRG family evolution using a more comprehensive data set. In particular, odorant receptors were hypothesized to have proliferated during terrestrial colonization by insects (hexapods), but their association with other pancrustacean clades and with independent terrestrial colonizations in other arthropod subphyla have been unclear. We also examine hypotheses on which arthropod CRG family is most ancient. Thus, we reconstructed phylogenies of CRGs, including those from new arthropod genomes and transcriptomes, and mapped CRG gains and losses across arthropod lineages. Our analysis was strengthened by including crustaceans, especially copepods, which reside outside the hexapod/branchiopod clade within the subphylum Pancrustacea. We generated the first high-resolution genome sequence of the copepod Eurytemora affinis and annotated its CRGs. We found odorant receptors and odorant binding proteins present only in hexapods (insects) and absent from all other arthropod lineages, indicating that they are not universal adaptations to land. Gustatory receptors likely represent the oldest chemosensory receptors among CRGs, dating back to the Placozoa. We also clarified and confirmed the evolutionary history of antennal ionotropic receptors across the Arthropoda. All antennal ionotropic receptors in E. affinis were expressed more highly in males than in females, suggestive of an association with male mate-recognition behavior. This study is the most comprehensive comparative analysis to date of CRG family evolution across the largest and most speciose metazoan phylum Arthropoda.

Morphological and genetic differentiation of heteromorphy in Labidocera rotunda (Copepoda, Calanoida, Pontellidae).

The pontellid calanoid Labidocera rotunda Mori, 1929 is relatively widespread in the inshore surface waters of East Asia. In this study, some heteromorphic specimens have been observed with extreme morphological discrepancies in the female, including the genital double-somite, the second urosomite, the caudal rami, and the fifth leg. To evaluate the validity of species assignment of the morphological variants, we analyzed the DNA sequences of two mitochondrial genes, 16S rRNA and cytochrome oxidase subunit I (mtCOI). The specimens were collected in the Yellow Sea, the East China Sea and the East Sea (Sea of Japan) around Korea. They differed by <2.9% for mt16S rRNA and by 3.7%-4.7% for mtCOI, suggesting that they are conspecific despite their considerable morphological differences. In contrast, Labidocera japonica Mori, 1935, a morphologically similar species to L. rotunda, displayed interspecific variation of 6.1%-7.2% for mt16S rRNA and 15.1%-16.7% for mtCOI.

Dynamics of Noctiluca scintillans blooms: A 20-year study in Jangmok Bay, Korea.

This 20-year study (2001-2020) conducted in Jangmok Bay, Korea, assessed the intricate relationships between environmental factors and Noctiluca scintillans blooms. Granger causality tests and PCA analysis were used to assess the impact of sea surface temperature (SST), salinity, dissolved oxygen (DO) concentration, wind patterns, rainfall, and chlorophyll-a (Chl-a) concentration on bloom dynamics. The results revealed significant, albeit delayed, influences of these variables on bloom occurrence, with SST exhibiting a notable 2-month lag and salinity a 1-month lag in their impact. Additionally, the analysis highlighted the significant roles of phosphate, ammonium, and silicate, which influenced N. scintillans blooms with lags of 1 to 3 months. The PCA demonstrates how SST and wind speed during spring and summer, along with wind direction and salinity in winter, significantly impact N. scintillans blooms. We noted not only an increase in large-scale N. scintillans blooms but also a cyclical pattern of occurrence every 3 years. These findings underscore the synergistic effects of environmental factors, highlighting the complex interplay between SST, salinity, DO concentration, and weather conditions to influence bloom patterns. This research enhances our understanding of harmful algal blooms (HABs), emphasizing the importance of a comprehensive approach that considers multiple interconnected environmental variables for predicting and managing N. scintillans blooms.

Reconstruction of arietellid copepod phylogenetic relationship, with description of a new species of Sarsarietellus (Copepoda, Calanoida, Arietellidae) from Korean waters.

A new species of the genus Sarsarietellus from Korean waters, S. orientalis n. sp. is described based on both sexes. This is very closely related to S. suluensis Ohtsuka, Nishida and Machida, 2005 , but is readily distinguished by the following characteristics: in the female (1) narrow head; (2) nearly symmetrical posterior corners of last pedigerous somite, not extending beyond the genital double-somite; (3) antennary endopod slightly longer than the exopod; (4) nearly equal length of three outer spines and the terminal spine of the exopod of the fifth leg, except for the proximal third outer spine; and in the male the left fifth leg with two whip-like setae on the tip of the second exopodal segment not being rotated outside. Sarsarietellus orientalis is the fourth species of the genus Sarsarietellus, while the male of Sarsarietellus species is the first reported. The habitat colonization of arietellids is reconsidered.

A new species of Heterochondria (Copepoda: Chondracanthidae) parasitic on many-banded sole, Zebrias fasciatus (Pleuronectiformes: Soleidae) from Korea, with a key to the species of the genus.

A new species of Chondracanthidae (Copepoda: Cyclopoida), Heterochondria orientalis sp. n., is described based on specimens of both sexes collected from the gill rakers and the inner side of the operculum of the many-banded sole, Zebrias fasciatus (Basilewsky), from the Yellow Sea, Korea. The new species resembles most closely H. zebriae (Ho, Kim et Kuman, 2000), but can be distinguished from this species and other congeners by the shape of the trunk and length of the antenna, the number of teeth on the mandible and the terminal process of the maxilla, and the structure of the male antennule and maxilliped. Heterochondria orientalis is the first copepod species reported from Z. fasciatus and the first heterochondrid species reported from sole fishes in the Northwest Pacific. A key to distinguish all 10 nominal species of the genus is provided.

Taxonomic reassessment of chaetognaths (Chaetognatha, Sagittoidea, Aphragmophora) from Korean waters.

Since the first record of chaetognaths (arrow worms) reported from Korean waters by Molchanov in 1907, three families, 12 genera and 21 species have been additionally described. Eighteen of the 21 recorded species have been reported under scientific names different from the latest taxonomic system. This study aimed to address this issue by conducting a taxonomic re-evaluation of chaetognaths collected from Korean waters. Furthermore, the taxonomic usefulness of morphological differences in corona ciliata and distribution of ciliary sense receptors were re-examined using specimens stained with 1% Chlorazol black E (CBE) solution. This study includes taxonomically-validated voucher specimens of 18 species from Korean waters. Based on the specimens, re-description including image data and CBE staining pattern, distribution, ecological information and improved key were provided for each species. However, Decipisagittadecipiens, Serratosagittaserratodentata and Sagittapseudoserratodentata from Korean waters is still questioned because of the paucity of the voucher specimen and scientific literature.

Short-term variation in abundance of four Acartia species (Copepoda, Calanoida) in a eutrophic bay.

The short-term variation in the abundance of Acartia copepods in the eutrophic Gamak Bay of South Korea was investigated with weekly measurements from October 2007 to September 2008. During this period, four Acartia species (A. erythraea, A. ohtsukai, A. omorii, and A. sinjiensis) were recorded as showing seasonally different peak abundance. The abundance of A. erythraea and A. sinjiensis was high in autumn, whereas that of A. omorii was high from winter to spring. In summer, A. erythraea, A. ohtsukai, and A. sinjiensis coexisted at peak abundance significantly related to water temperature and salinity. Results from the response curves of the four Acartia species to water temperature and salinity suggest that A. erythraea and A. sinjiensis increased in abundance at water temperatures >18 °C, whereas A. ohtsukai increased in abundance at water temperatures >27 °C. The occurrence of A. erythraea, A. ohtsukai, and A. sinjiensis decreased with increasing salinity, but chlorophyll-a concentration showed no effect on occurrence. Despite these findings, the coexistence of the three ecologically similar species may be due to prey abundance in summer and autumn (chlorophyll-a concentration >10 µg L-1). Notably, the wide range of the response curve of A. omorii indicates its occurrence at higher salinity levels than other species.

Three new species of the genus Stephos Scott, 1892 (Crustacea, Copepoda, Calanoida, Stephidae) from Jeju Island, Korea.

During general field surveys carried out recently to collect benthopelagic copepods from near the substrate of the shallow waters off Jeju Island, Korea, a few specimens of three new species of Stephos Scott, 1892, were collected. The new species are placed in the genus Stephos because of the following combination of features: absence of seta on the basal exite of maxillule, and male right leg 5 ending in an unarmed claw-like and/or mitten-like segment. Stephos jejuensis sp. nov. can be distinguished from its congeners by body length 0.92 mm, left side of the female genital double-somite with protruding lobes, antennule that extends beyond the distal area of the genital double-somite, and the male leg 5 terminal complex. Stephos concavus sp. nov. can be distinguished from its congeners by the genital double-somite with protruding lobes on both sides, and the presence of larger spinules on the distomedial margin of leg 5. Stephos fortipes sp. nov. can be distinguished from its congeners by its longer body length, 1.12 mm long in the female, antennules that extend to the end of the genital double-somite, and the presence of a covered row of minute spinules on the ventral surface of the genital operculum in the female. Until now, 35 species of stephids were known worldwide.

A new species in the genus Acartia Dana, 1846 (Crustacea, Copepoda, Calanoida, Acartiidae) from the South Pacific coastal waters of Nadi Bay, Fiji.

A new species in the genus Acartia, Acartia nadiensis sp. nov., is described from Fijian coastal waters. This species belongs to the subgenus Odontacartia based on the following morphological features: presence of a rostral filaments, a pointed process on the last prosomite, a serrated terminal spine on female P5, and the absence of a protrusion on the basis of the male right P5. This new species can be differentiated from its congeners by the combination of the absence of a spine on the first segment of the antennules, the short outer seta of female P5, and a medial spine on the exp-2 of the left male P5. Phylogenetic analyses using mitochondrial COI partial sequences show that the new species is distinct from its congeners.

New species of Caromiobenella Jeon, Lee & Soh, 2018 (Crustacea, Copepoda, Monstrilloida) from Chuja Island, Korea.

Male monstrilloid copepods belonging to the genus Caromiobenella Jeon, Lee & Soh, 2018 were collected from Chuja Island, Jeju, Korea, using a light trap. This paper describes a new species, Caromiobenellaohtsukai sp. n., based on the display of reduced, knob-like fifth legs on the ventral side of the first urosomal somite. A unique combination of male genitalia features and number of caudal setae further confirms its specificity. Molecular analysis based on two partial gene sequences of mitochondrial cytochrome c oxidase subunit I (mtCOI) and 28S ribosomal RNA (28S rRNA) also supports the designation of this species by showing a relevant divergence from known congeners. Caromiobenellaohtsukai sp. n. is the ninth member of this genus and also the ninth monstrilloid reported from Korea.

Three new species of Heteromastus (Annelida, Capitellidae) from Korean waters, with genetic evidence based on two gene markers.

Three undescribed species of Heteromastus Eisig, 1887 were collected from intertidal to sublittoral habitats in western and southern waters of Korea. Heteromastus namhaensis sp. nov. is distinguishable from other congeners by the presence of hemispheric notopodial lobes in the posterior abdomen. Heteromastus gusipoensis sp. nov. closely resembles H. tohbaiensis Yabe & Mawatari, 1998 in the absence of posteriorly extended abdominal notopodial lobes, but differs in the absence of eyespots on the prostomium and distinct node on the shaft of thoracic hooks in H. gusipoensis. Heteromastus koreanus sp. nov. is similar to H. filiformis sensu Hutchings & Rainer, 1982 in the shape of abdominal notopodia, but clearly differs in dentition of the abdominal hooks and methylene green staining pattern (MGSP). DNA sequences (mtCOI and histone H3) of these new Korean species were compared with all sequences of Heteromastus species available in the public database. Molecular results showed distinct genetic differences among these three new Korean species at species level. Comparison of mtCOI gene revealed significant genetic difference between H. filiformis and these Korean species. A comprehensive comparison between three Heteromastus species of present study and their closely related congeners is conducted based on morphological and genetic results.

Prorocentrum shikokuense Hada and P. donghaiense Lu are junior synonyms of P. obtusidens Schiller, but not of P. dentatum Stein (Prorocentrales, Dinophyceae).

In Japanese, Chinese and Korean coastal waters, recurrent blooms of a small, elongate Prorocentrum species have been observed in recent years. In previous studies, this species has been respectively identified as P. shikokuense, P. donghaiense and P. dentatum, despite morphological similarity and identical rDNA sequences. To resolve the confusion, morphological features, including the architectural details of the periflagellar area, were examined and ribosomal DNA (rDNA) sequences were obtained from specimens collected from the East China Sea and Korean coast, and a strain established in the waters off the Canary Islands of Spain. In addition, the descriptions of the three species and allied species were reviewed. Morphological observations and a phylogeny based on the SSU, ITS region and LSU rDNA sequences revealed that the previously confused species and our studied strains are conspecific and that the morphology of strains identified as P. dentatum in the phylogenetic trees does not coincide with P. dentatum sensu stricto. The confusion can be traced back to Dodge (1975, p. 116), who considered P. veloi, P. monacense and P. obtusidens as junior heterotypic synonyms of P. dentatum. However, Dodge's P. dentatum are closer to P. obtusidens sensu stricto, rather than P. dentatum sensu stricto. P. obtusidens sensu stricto can be distinguished from P. dentatum sensu stricto by its relatively small size, parallel sides towards the anterior and a blunt anterior extension on one side. This indicates that P. obtusidens should not be considered a synonym of P. dentatum sensu stricto. In addition, a comparison of the original descriptions of P. obtusidens and allied species allowed to conclude that small, elongate Prorocentrum from Japanese, Chinese and Korean coastal waters previously identified as P. shikokuense, P. donghaiense, P. dentatum sensu Yoo and Lee (1986) and the specimens studied herein, which share identical rDNA sequences, morphologically coincide with P. obtusidens. Therefore, it is proposed that P. shikokuense and P. donghaiense should be regarded as junior synonyms of P. obtusidens.

First use of molecular evidence to match sexes in the Monstrilloida (Crustacea: Copepoda), and taxonomic implications of the newly recognized and described, partly Maemonstrilla-like females of Monstrillopsis longilobata Lee, Kim & Chang, 2016.

Two forms of the monstrilloid copepod, one represented by males, the other by females, were dominant among the monstrilloids collected while sampling zooplankton with a light trap in southern coastal waters of Korea. Morphological examination revealed that the males are conspecific with the previously reported and rather specialized Korean species Monstrillopsis longilobata Lee, Kim & Chang, 2016, hitherto known only from males that have extremely long genital lappets. The females also show several diagnostic features of Monstrillopsis, such as prominent eyes, bilobed fifth legs with the inner lobe unarmed and reduced, and four urosomal somites. In addition, though, these females are extraordinary among all Monstrilloida in that their ovigerous spines are directed ventrally, not posteriorly as in most species or anteriorly as in those assigned to Maemonstrilla. Genetic divergence analyses based on partial mitochondrial cytochrome c oxidase subunit I, complete Internal Transcribed Spacer 1 (ITS1)-5.8S-ITS2, and partial 28S rRNA revealed little or no genetic divergence between the present males and females, thus demonstrating that they are mutually conspecific. The present report (1) provides the first morphological description of female M. longilobata with the proposal of a revised nomenclature for antennular setal armament; (2) presents the molecular evidence for conspecificity of the males and females; and (3) lists several morphological characteristics that are sexually dimorphic in this species, and thus likely also in other monstrilloids. Matters bearing on the validity of the genera Haemocera, Monstrillopsis, and Maemonstrilla are discussed.

A new Notomastus (Annelida, Capitellidae) species from Korean waters, with genetic comparison based on three gene markers.

Notomastus koreanussp. n., collected from the sublittoral muddy bottom of Korean waters, is described as a new species. The Korean new species closely resembles N. torquatus Hutchings & Rainer, 1979 in the chaetal arrangement and the details of abdominal segments, but differs in the position of genital pores and the absence of eyes. DNA sequences (mtCOI, 16S rRNA, and histone H3) of the new species were compared with all the available sequences of Notomastus species in the GenBank database. Three genes showed significant genetic differences between the new species and its congeners (COI: 51.2%, 16S: 38.1-47.3%, H3: 3.7-9.3%). This study also includes a comprehensive comparison of the new Korean Notomastus species with its most closely similar species, based on the morphological and genetic results.

Genetic identification and molecular phylogeny of Pseudodiaptomus species (Calanoida, Pseudodiaptomidae) in Korean waters.

Five species of the genus Pseudodiaptomus, P. inopinus, P. poplesia, P. marinus, P. nihonkaiensis, and P. sp. occur in Korea nearshore waters. Although the four species except for P. sp. have been classified into Lobus and Ramosus groups, two for each group, based on morphological characters, this classification had yet to be confirmed by molecular characters. Here, we determined molecular characters and phylogenetic relationships of the five species in order to evaluate the morphology-based groupings and the species identifications. For this, a 625-bp DNA region of the mitochondrial gene cytochrome oxidase subunit I (mtCOI) was sequenced and compared among the species. Intraspecific variation of the sequences is less than 0.6%, while interspecific variation ranges from 17.6-26.7%, indicating every species, including P. sp., is a genetically distinct, valid species. Phylogenetic trees of the mtCOI DNA reveal that the Lobus-group species including P. inopinus and P. poplesia form a well-supported clade and that P. sp. belongs to this group. On the other hand, the Ramosus group clade consisting of P. marinus and P. nihonkaiensis is not well supported by bootstrap analyses, suggesting that further evaluation of the validity of this group assignment is needed.

Two new species in the genus Phoxokalliapseudes Drumm & Heard, 2011 (Crustacea: Tanaidacea: Kalliapseudidae) from the southwestern and southern coasts of Korea.

Two new species of Phoxokalliapseudes from the southwestern and southern coasts of Korea are described. Phoxokalliapseudes aculeatus n. sp. is distinguished by the two robust spiniform setae on the ventral margin of the cheliped basis, pereonites 2-6 each with a hyposphaenium and small spinules, and the setiferous lobe on the mandibles with different numbers of serrated spiniform setae in the female and male (five and six, respectively). Phoxokalliapseudes cinctus n. sp. can be distinguished by the propodus of the male cheliped with the largest length to width ratio (2.5: 1) of the genus, and the fixed finger with small denticles on the cutting edge and near the insertion of the dactylus. The female has the largest dactylus to propodus length ratio of pereopod 6 in the genus. Only four species of Phoxokalliapseudes have been described or redescribed previously, of which some exhibit morphological differences with the type specimens, possibly indicating additional new species. To minimize identification difficulties caused by morphometric variations in developmental stages, the morphological characteristics of the new species are described in detail for a fixed size range and compared to different-sized individuals. Additionally, a comprehensive comparison of the new Phoxokalliapseudes species with the other known species is provided and discussed.

A new species of Eusirus from Jeju Island, Korea (Crustacea, Amphipoda, Eusiridae).

A new eusirid amphipod, Eusirus bulbodigitussp. n., from Jeju Island, Korea is described with a detailed description and illustrations. Eusirus bulbodigitussp. n. shows common features with the five known eusirid amphipods Eusirus abyssi Stephensen, 1944, Eusirus columbianus Bousfield & Hendrycks, 1995, Eusirus hirayamae Bousfield & Hendrycks, 1995, Eusirus laticarpus Chevreux, 1906, and Eusirus parvus Pirlot, 1934, such as the mandibular palp article 3 bearing a group of setae laterally. However, this new species is differentiated by the combination of the following characteristics: the eyes are poorly developed, the propodus on pereopod 4 is slightly shorter, the inner margin of dactylus on pereopod 4 is swollen, the length of pereopods 5-7 is moderate, the urosomite 1 has a dorsal protrusion distally, and the telson is shallowly cleft. This is the first record of the genus Eusirus Krøyer, 1845 from Korean waters.

On the occurrence of caligids (Copepoda: Siphonostomatoida) in the marine plankton: a review and checklist.

Members of the copepod genus Caligus Müller, 1785 (Siphonostomatoida: Caligidae) are commonly referred to as sea lice. Virtually all of the more than 450 species utilize marine fishes as hosts, however, an increasing number of records from marine plankton samples shows that at least some species reside in the water column during their adult phase. Members of three different genera, Caligus, Lepeophtheirus von Nordmann, 1832, and Metacaligus Thomsen, 1949 have been reported from plankton samples off eastern Asia and in the north-western Atlantic. Thirteen species have so far been consistently reported from the plankton only with no information on their hosts. Here we review the various hypotheses previously proposed to explain the presence of caligids in the water column, i.e. accidental occurrence, behavioral detachment from the host during mate location, and host-switching. The discovery of adults of two species of Caligus with no other developmental stages on their teleost hosts, suggests an ontogenetic host-switching after the final molt since both species also occurred in plankton samples. A checklist of all caligid records from the marine plankton, including known host data, is presented.

A new species of the genus Attheyella (Copepoda: Harpacticoida: Canthocamptidae) from Gosu cave in Korea.

A new species, Attheyella namkungi sp. nov., is described from Gosu cave at Danyang, Chungcheongbuk-do, South Korea. This species bears a superficial resemblance to A. coreana Miura, 1969 in having a 2-segmented mandibular palp with only four smooth apical setae, and in the armature formulae of the antennary exopod and thoracic legs. However the new species is clearly distinguishable by the non-sexually dimorphic caudal rami, two long plumose setae and a short seta on leg 6, and the absence of the setular row at the inner lateral margin of each caudal ramus. The new species is characterized from other congeners by having two adjacent inner spines on female leg 3 endopod-2, the caudal ramus twice as long as its width, absence of a barbed apophysis on male leg 3 endopod-2, and the seta number of legs 5 and 6. A key to the Korean species of the genus Attheyella is provided.

Five Sarsiellidae ostracods (Crustacea: Myodocopida) from the South Coast of Korea (East China Sea).

The East China Sea is part of the Warm Temperate Northwest Pacific zoogeographic province and, as such, has a high biodiversity and many tropical and subtropical biotic elements. Nevertheless, many invertebrate groups from this area remain poorly studied. Ostracods are one of them, especially those belonging to the subclass Myodocopa. In this paper we provide the first data on a diverse myodocopid family, Sarsiellidae, not only for the East China Sea, but also for Korea. Five species are reported in this paper from three Korean islands (Jeju, Chuja, and Maemul), and they are only a part of the ostracods collected during this study, indicating a high diversity of the group in this region. Three new species, Eurypylus koreanus sp. nov., Eusarsiella hanguk sp. nov., and Sarsiella nereis sp. nov., clearly stand apart from their respective congeners, mostly by prominent shell characters but also by details of the soft part morphology. Their affinity though clearly indicates a close connection of the region with the more southern zoogeographical realms, especially Central Indo Pacific and partly Temperate Australasia. Two species previously known from Japan (north part of the Sea of Japan and southeastern part of the Pacific Coast of Japan), Sarsiella japonica Hiruta, 1977 and S. misakiensis Kajiyama, 1912, are redescribed. Based on 11 newly obtained COI sequences we construct a preliminary phylogenetic tree, which supports previous hypotheses based on the morphological data, that Eusarsiella Cohen & Kornicker, 1975 is a polyphyletic taxon. With the maps of species distribution provided for each of the three genera, we give an overview of their current zoogeography, and clearly indicate areas that have no data, mostly due to the lack of investigation.