Rhizocephalan Species Research Articles

The host range and distribution pattern of rhizocephalan parasitic barnacles in Korean coasts and their relationship with geographical factors

IntroductionThis study undertakes a detailed examination of the host range and distribution patterns of Rhizocephala, a group of specialized parasitic barnacles, within Korean marine environments. It aims to expand the understanding of their biodiversity and ecological impact.MethodsEmploying both morphological observation and mitochondrial DNA sequencing, the study identifies the variety of rhizocephalan species infecting decapod hosts in Korean waters, aiming to document their diversity and distribution.ResultsThe investigation identifies 26 rhizocephalan species parasitizing 28 decapod hosts, including the discovery of three new parasite-host relationships not previously reported. It was observed that while a few rhizocephalan species infect multiple hosts, most have a narrow host range. Additionally, the study maps out regional differences in Rhizocephala diversity across three Korean marine ecoregions, noting the highest diversity in the warmer East China Sea ecoregion and lower diversity in the colder Yellow Sea and East Sea ecoregions.DiscussionThe findings underscore the necessity for continued taxonomic research on Rhizocephala and their host species to better understand and manage these parasites, which hold significant ecological and economic importance. The study suggests that geographical variations in host-parasite relationships are influenced by regional climatic conditions, particularly winter water temperatures.

Functional role of lacunar and muscular systems in the externa of Peltogasterella gracilis (Cirripedia: Rhizocephala).

One of the most conspicuous traits of parasitic organisms is a well-developed reproductive system. In Rhizocephala ("Crustacea": Cirripedia) it is believed to be nested in the externa-a "reproductive part" located outside of the host. However, it is not clear how nutrients are transported to the externa. Several authors described a system of lacunae in the externa, and muscular contractions probably enable transport through these cavities. The aim of our study was to visualize (using microcomputed tomography and confocal laser scanning microscopy) and describe lacunar and muscular systems in the externa of Peltogasterella gracilis (fam. Peltogasterellidae). The lacunar system consists of "ventral" lacuna and several protrusions. The "ventral" lacuna is probably responsible for visceral mass nutrition, and mantle protrusions are associated with the mantle nutrition. The gross organization of the muscular system mostly corresponds to previous descriptions in other rhizocephalan species. Nonetheless, we observed several features of the externa morphology that had not been described before such as a muscular thickening in the proximal externa's part and a stalk plug disk. The muscular thickening might play a role of a propulsatory organ, helping to transport liquid through the lacunar system. The plug disk might fill the hole in the host's cuticle after the old externa drop off. The results allow us to make first assumptions on transport mechanisms in Rhizocephala.

Morphological and molecular analyses of parasitic barnacles (Crustacea: Cirripedia: Rhizocephala) in Korea: preliminary data for the taxonomy and host ranges of Korean species.

Morphological and molecular analyses of Korean rhizocephalan barnacle species were performed to examine their host ranges and taxonomy. Morphological examination and molecular analysis of mtDNA cox1, 16S, and nuclear 18S rRNA sequences revealed nine rhizocephalan species from three genera of the two families, Sacculinidae and Polyascidae. Phylogenetic analysis of molecular sequences revealed two new species candidates in the genus Parasacculina, and three Sacculina species (S. pilosella, S. pinnotherae, and S. imberbis) were transferred to the genus Parasacculina. Examination of host ranges revealed higher host specificity and lower infestation rates in Korean rhizocephalan species than rhizocephalans from other geographic regions. This is the first report of the taxonomy, species diversity, and host ranges of Korean parasitic rhizocephalan barnacles based on their morphological and molecular analyses. More information from extensive sampling of parasitic barnacles from a wide range of crustacean host species is necessary to fully understand their taxonomy, prevalence on decapod hosts, and phylogenetic relationships among major rhizocephalan taxa.

Differences in the parasitic effects of a bopyrid isopod and rhizocephalan barnacle on the portunid crab, Charybdis bimaculata.

Bopyrid isopods and rhizocephalan barnacles are obligate parasite crustaceans which harm their decapod hosts. However, to the best of our knowledge, studies have not compared which of these parasites has a greater parasitic effect on its hosts. Here, the parasitic effect of the bopyrid isopod, Allokepon hendersoni, and an unidentified sacculinid rhizocephalan species, infesting the same population of portunid crabs, Charybdis bimaculata, was investigated and compared for the first time. Samples were collected from the bycatch of a trawl fishery in Tosa Bay, Japan. A total of 2601 crabs were collected, of which 14 (0.55%) were parasitized by the bopyrid and 21 (0.82%) by the rhizocephalan. One of the two female crabs parasitized by the bopyrid was ovigerous (with much fewer eggs than unparasitized females). No ovigerous crab was found from the eight females parasitized by the rhizocephalan. Because only two female crabs were parasitized by the bopyrid, the following analyses were made using the male crabs. Both parasites reduced the wet weight (crab condition) and the cheliped size (secondary growth) of C. bimaculata, but the impact of the parasitism did not differ between the parasite species. The size of the abdominal flap of male hosts was reduced by the bopyrid infestation; however, rhizocephalan infestation caused enlargement of the abdominal flap, which is an indication of feminization. The present study provides information on how the effect of these two parasitic castrators on the same host crab varies. A moderate decrease in crab condition and cheliped development was common among the parasites, suggesting that the degree of impact might be favorable for the survival of the two parasites species.

Sympatric Two-species Infestation by Rhizocephalan Barnacle Parasites in the Spider Crab Pugettia aff. ferox Ohtsuchi & Kawamura, 2019 from Peter the Great Bay (Northwestern Sea of Japan).

Sympatric two-species infestation by rhizocephalan parasites in the spider crab Pugettia aff. ferox Ohtsuchi & Kawamura, 2019 (Brachyura: Epialtidae) was investigated in the Vostok Bay (Peter the Great Bay, northwestern Sea of Japan). Morphological and molecular analyses showed that this crab was infested simultaneously by Sacculina pugettiae Shiino, 1943 and Parasacculina pilosella (Van Kampen et Boschma, 1925) (Cirripedia: Rhizocephala). Sacculina pugettiae was found in the northwestern Sea of Japan for the first time. The two rhizocephalan species are clearly distinguishable by the morphology of their external cuticles, the shape and position of their receptacles, and the structure of their colleteric glands. Retinacula are present in the mature externae of both species. Molecular analysis showed that these rhizocephalans are unrelated, although both species parasitize Pugettia aff. ferox and are sympatric. Sacculina pilosella should be placed in the genus Parasacculina Høeg & Glenner, 2019, belonging to the family Polyascidae Høeg & Glenner, 2019. The intensity of infestation reached two externae in P. pilosella and three externa in S. pugettiae per host. A simultaneous settlement of two rhizocephalans on the same crab specimen was shown for the first time. The intensity of the two-species multiple infestations reached four externae per host. Externae with developing embryos occurred from June to September in P. pilosella and July to September in S. pugettiae, at water temperatures of 15-24°C, indicating that the reproductive periods of these species are confined to the summer months in the investigated locality.

Phylogenetic position, complete larval development and larval sexual dimorphism in a rhizocephalan barnacle, Lernaeodiscus rybakovi sp. nov. (Cirripedia: Rhizocephala: Peltogastridae), parasitizing the crab Pachycheles stevensii Stimpson, 1858 (Decapoda: Anomura: Porcellanidae)

Molecular and morphological methods are used to describe Lernaeodiscus rybakovi, a new rhizocephalan species parasitizing the porcellanid crab Pachycheles stevensii Stimpson, 1858, collected in Russian waters of the Sea of Japan. Molecular analysis of three species, including the new one, confirms the monophyly of the genus Lernaeodicus Müller, 1862 and its recent transfer to the family Peltogastridae. The main morphological features of the new species are also common with characters of the other species of the genus Lernaeodiscus Müller, 1862. Externa of L. rybakovi differs from the well studied Lernaeodiscus porcellanae from the E. Pacific by molecular markers, color and the absence of pronounced marginal lobes. Retinacula on the internal cuticle of L. rybakovi are found in the genus Lernaeodiscus for the first time. Some of the female hosts with adult externae were unusual in also carrying their own eggs on the pleopods, a rare situation among rhizocephalans. The complete larval development in the genus Lernaeodiscus is described here for the first time and includes five naupliar and one cypris instar. The main morphological features of Lernaeodiscus nauplii (the presence of flotation collar, morphology of the frontolateral horns and furcal spines, and the arrangement of dorsal setae on the shield head) are common with those of other known peltogastrid larvae, but also resemble nauplii of Peltogasterella in the presence of naupliar eyes and thin structure of the flotation collar. The sizes of male and female cyprids of L. rybakovi overlap slightly. In summer months, larval sex ratio is male-biased. We briefly review important larval characters in the Rhizocephala.

Specialized structures on the border between rhizocephalan parasites and their host\u2019s nervous system reveal potential sites for host-parasite interactions

Rhizocephalan barnacles are a unique group of endoparasitic crustaceans. In their extreme adaptation to endoparasitism, rhizocephalan adults have lost almost all features of their free-living relatives but acquired an outstanding degree of control over the body of their hosts (mostly decapods). The subtle influence exercised by rhizocephalans on the physiology, morphology and behaviour of their hosts is a vivid example of the most intimate host-parasite interactions but their mechanisms are very poorly known. In this study we examined the morphology and the adaptive ultrastructure of the organs invading the nervous system of the host in two rhizocephalan species from the families Peltogastridae, (Peltogaster paguri) and Peltogasterellidae (Peltogasterella gracilis). We found two essentially different types of structures involved in interactions of these two rhizocephalans with the nervous system of their hosts: modified rhizocephalan rootlets lying inside the ganglia and the neural fibres of the host enlacing the trophic rootlets of the parasites. We suggest that both these structures may be highly specialized tools allowing the parasite to interact with the host on the humoral level via neuromediators, hormones, attractants and trophic factors.

Evolution of Feeding Shapes Swimming Kinematics of Barnacle Naupliar Larvae: A Comparison between Trophic Modes.

SynopsisA central goal in evolutionary biology is connecting morphological features with ecological functions. For marine invertebrate larvae, appendage movement determines locomotion, feeding, and predator avoidance ability. Barnacle larvae are morphologically diverse, and the morphology of non-feeding lecithotrophic nauplii are distinct from those that are planktotrophic. Lecithotrophic larvae have a more globular body shape and simplified appendages when compared with planktotrophs. However, little is known about whether and how such morphological changes affect kinematics, hydrodynamics, and ecological functions. Here, we compared the nauplii kinematics and hydrodynamics of a lecithotrophic Rhizocephalan species, Polyascus planus, against that of the planktotrophic nauplii of an intertidal barnacle, Tetraclita japonica. High-speed, micro-particle image velocimetry analysis showed that the Polyascus nauplii swam faster and had higher amplitude and more synchronous appendage beating than the Tetraclita nauplii. This fast swimming was accompanied by a faster attenuation of induced flow with distance, suggesting reduced predation risk. Tetraclita nauplii had more efficient per beat cycles with less backward displacement during the recovery stroke. This “anchoring effect” resulted from the anti-phase beating of appendages. This movement, together with a high-drag body form, likely helps direct the suction flow toward the ventral food capturing area. In sum, the tradeoff between swimming speed and predation risks may have been an important factor in the evolution of the observed larval forms.

A new species of the genus Peltogaster Rathke, 1842 (Cirripedia: Rhizocephala: Peltogastridae) parasitizing the hermit crab Pagurixus boninensis (Melin, 1939) from the Bonin Islands, Japan.

A new rhizocephalan species, Peltogaster unigibba n. sp., is described from the host hermit crab, Pagurixus boninensis (Melin, 1939), from the Bonin Islands, Japan. Of the16 known species of Peltogaster now currently recognised, P. unigibba n. sp., and P. contorta Boschma, 1958 share a left lobe that projects beyond the mantle aperture. The two species can be distinguished from one another by the position of the opening of the mantle aperture. The new species most closely resembles P. naushonensis Reinhard, 1946 in its internal structure, but clearly differs in the relative length of the colleteric glands. Peltogaster unigibba n. sp. represents the first record of a rhizocephalan from the oceanic Bonin Islands, and the second record of a rhizocephalan from an oceanic island in the northern hemisphere.

Hermit crab host prevalence by species of Peltogastridae (Cirripedia: Rhizocephala): hosts vary with locations on the Pacific coast in mainland Japan

Of 2282 individuals in 23 hermit crab species collected from four coasts of the southern part of mainland Japan along the Pacific coast, 87 individuals in seven species were infested by six rhizocephalan species. Among the ten combinations between hosts and rhizocephalans, seven combinations were new records. Morphological identification of the rhizocephalan species from different host species and/or sites was consistent with molecular identification using DNA sequences (COI and 16S rRNA genes). Pagurus filholi and P. nigrivittatus were infested by both Peltogaster postica and P. lineata. The host preference of the two rhizocephalan species reciprocally shifted from one species to the other among the four collection sites. Rhizocephalans may choose the host inhabiting the preferred environment for the parasite within the range of their host species.

A new peltogastrid rhizocephalan parasitising a hermit crab from the Japanese coast: a second species of Dipterosaccus Van Kampen & Boschma, 1925 (Crustacea: Cirripedia)

A new rhizocephalan species of the genus Dipterosaccus Van Kampen & Boschma, 1925, is described from the host hermit crab Calcinus vachoni Forest on the coasts of mainland Japan and the Ryukyu Archipelago. Dipterosaccus shiinoi n. sp. differs from the previously described species, D. indicus Van Kampen & Boschma, 1925, which is redescribed, principally in the morphology of the mantle aperture and the disposition of the receptacle ducts. The distribution ranges of the two species partly overlap and they are sympatric at the same reef site near Okinawa Island. Their differentiation as two distinct species was confirmed by partial sequences of the COI gene. The monophyly of Dipterosaccus is supported by high Bayes posterior possibility and bootstrap values.

Anatomy of virgin and mature externae of Loxothylacus texanus, parasitic on the dark blue crab Callinectes rathbunae (Crustacea: Cirripedia: Rhizocephala: Sacculinidae)

Rhizocephalan parasites are dioecious organisms, in that one or several dwarf males are implanted into the external part of the female parasite soon after it emerges from the interior of the host animal. The structure of the female externa and its resident males is crucial for understanding both the reproductive biology and the taxonomy of these specialized parasites. We use scanning electron microscopy and histological methods to study the anatomy of juvenile and the mature externae of the rhizocephalan barnacle Loxothylacus texanus parasitizing the blue crab Callinectes rathbunae. We put emphasis on the implantation of males and the histology of the female reproductive organs. In the virgin externae, male cyprids attach around a cuticular hood covering the mantle aperture, which is partially blocked by a plug of cuticle so only trichogon larvae, not cyprids, can access the mantle cavity. This resembles the situation known from Sacculina carcini. The mature externa is characterized by a visceral mass that contains the ovary, paired colleteric glands, a single male receptacle, but paired receptacle ducts. The proximal attachment of the visceral mass is located at some distance from the basal stalk, as is characteristic for the genus Loxothylacus. The internal anatomy of the mature externa of L. texanus is in most features similar to that seen in other species of the Sacculinidae, which comprises the majority of rhizocephalan species. However, the single receptacle creates a situation where the two implanted males cannot be kept separate as in most other rhizocephalans, but pass through spermatogenesis in a common chamber. This may have unknown effects on the reproductive biology such as male-male competition.

Cirripede cleavage patterns and the origin of the Rhizocephala (Crustacea: Thecostraca)

Several aspects of phylogenetic relationships among barnacles (Cirripedia) are still unresolved. One of the contentious issues is the position of the parasitic Rhizocephala. In most molecular phylogenies Rhizocephala are resolved as sister group to a monophyletic Thoracica. However, since Rhizocephala are morphologically highly derived there is not a single morphological character supporting this view. Here we present data on the early cleavage patterns and the gastrulation of two rhizocephalan species. Based on our results and data from the literature we suggest that early cleavage and gastrulation indicate a monophyletic group comprising the thoracican Iblomorpha and the Rhizocephala. This renders thoracicans paraphyletic with respect to Rhizocephala. Based on this, we develop a new hypothesis for the origin of parasitism in the rhizocephalan stem lineage starting with parasite-like dwarf males of an iblomorph-like ancestor which already had the ability to penetrate the surface of a host animal – originally conspecifi c females, but later the decapod, probably a pagurid, host.

Parasitic barnacles (Cirripedia: Rhizocephala) from New Zealand off‐shore waters

Nine genera and species of rhizocephalans were recorded from the off‐shore waters around New Zealand. Mitochondrial and nuclear gene sequences were used to examine base differences between the European and New Zealand species of Parthenopea. Serial sections to study the internal structures of the reproductive organs were made for the genera Thylacoplethus and Thompsonia. Two species, Parthenopea australis n. sp. and Thylacoplethus novaezealandiae n. sp. are new to science and described in detail. Parthenopea australis n. sp. is the first rhizocephalan species recorded from the vicinity of active cold seeps. Three rhizocephalans could not be identified as they were parasitised by hyperparasitic cryptoniscine isopods. The decapodhost species comprised the taxa Paguridae, Lithodidae, Galatheidae, Chirostylidae, and Callianassidae.

Effects of temperature and salinity on the larvae of two species of rhizocephalan (Crustacea: Cirripedia)

Summary Responses of larvae of two rhizocephalan species to changes in seawater temperature and salinity were studied under laboratory conditions. Peltogasterella gracilis parasitizes the hermit crab Pagurus pectinatus, which occurs at stable salinity and gradually changing temperature in summer. Sacculina polygenea is a parasite of the crab Hemigrapsus sanguineus, which lives in the intertidal zone in summer where salinity and temperature can fluctuate during the day. The development of both species is comprised of five naupliar stages and the cyprid stage, and it was considered successful if more than 50% of the nauplii attained the cyprid stage. P. gracilis nauplii successfully developed at 12–20°C and 30–34‰, but at 22°C successful development occurred in a narrower salinity range (32–34‰). All nauplii died both at 25°C and in 26‰. S. polygenea nauplii successfully reached the cyprid stage at higher temperatures (18–25°C) and a wider salinity range (18–34‰) than P. gracilis nauplii, but at 12°C and 16‰ larval development of S. polygenea was suppressed. Under favorable conditions, naupliar development lasted 3.5 days in P. gracilis and 2–3 days in S. polygenea. The cyprids of both rhizocephalan species demonstrated a greater resistance to temperature and salinity changes than nauplii. However, P. gracilis cyprids were active in a narrower salinity range (16–34‰), as compared to S. polygenea cyprids (8–34‰). Under favorable conditions the cyprids of both species survived for 6 to 10 days.

Rhizocephala (Crustacea, Cirripedia) of the Faroe Islands

Abstract The rhizocephalan fauna of the Faroe lsiands is evaluated in the light of the BIOFAR program. The BIOFAR material includes six species of Rhizocephala of which the five are new to Faroese waters: Lemaeodiscus ingolfi, Peltogaster paguri, Sylon hippolytes, Tortugaster boschmai and Triangulus galatheae. The number of rhizocephalan species in the area is raised from two to seven and a gap in the knowledge of their North Atlantic distribution is filled. The vertical distribution of Clistosaccus paguri and P. paguri is revised. An abnormally positioned S. hippolytes is documented for the first time. Also new to the Faroe islands is the hyperparasite of P. paguri: Liriopsis (Crustacea, Isopoda).

Rhizocephalans (Crustacea: Cirripedia) from Taiwan

One species of shrimp and sixteen species of brachyuran crabs are parasitized by rhizocephalans in Taiwanese waters. Ten of the crabs have never before been recorded as hosts of rhizocephalans. Eight species of rhizocephalans occur in Taiwan. Four species, Diplothylacus taiwanensis, D. calappae, Sacculina scutigera and S. docleae are new to science. The paper records the first example of simultaneous infection of the same host individual by two rhizocephalan species, and provides further evidence of successive moulting of the external mantle cuticle in several species of sacculinids.

Ecological correlates of body size and egg size in parasitic Ascothoracida and Rhizocephala (Crustacea)

In order to identify key factors in the evolution of life history traits in Ascothoracida and Rhizocephala (two groups of crustacean parastes of invertebrates), comparative analyses were performed using phylogenetically independent contrasts. Among 59 ascothoracidan species, latitude correlated positively with body size, whereas there was no relationship between water depth and body size. Body size also correlated strongly with egg size; however, once corrected for body size, egg size was not related to either latitude or water depth. Among 91 rhizocephalan species, neither latitude nor water depth correlated with body size. However, host species of larger sizes harboured larger species of rhizocephalan parasites. Egg size of rhizocephalans did not correlate with body size, and was not influenced by either latitude or water depth. The patterns observed in this study show both differences from an similarities to those reported for other groups of crustacean parasites, and suggest that adaptations to similar selective pressures are not always identical among distantly-related taxa.

Scanning electron microscopy of lattice organs in cyprids of the Rhizocephala Akentrogonida (Crustacea: Cirripedia)

The morphology and disposition of lattice organs, putative sensory structures in the carapace of cyprids, are described with scanning electron microscopy from six rhizocephalan species representing four of the five families in the suborder Akentrogonida: Clistosaccus paguri and Sylon hippolytes (Clistosaccidae), Arcturosaccus kussakini (Duplorbidae), Mycetomorpha vancouverensis (Mycetomorphidae), and Diplothylacus sinensis and Thylacoplethus reinhardi (Thompsoniidae). The disposition of pores and setae on the cyprid carapace is also surveyed. In T. reinhardi, poor quality of the specimens allowed only confirmation that lattice organs are present. Cyprids of all other species except M. vancouverensis have two anterior and three posterior pairs of lattice organs on the carapace, just as in almost all other Cirripedia. Cyprids of C. paguri and S. hippolytes are nearly identical in both general shape, carapace setation, and morphology of the lattice organs. In both species the lattice organs are oval–circular pore fields, except the second pair which is very elongate and match-shaped, a morphology unique to the Clistosaccidae. The similarities in cyprid morphology are interpreted as apomorphies supporting the monophyly of the recently rediagnosed Clistosaccidae. Both thompsoniid species have oval–circular lattice organs, indicating a possible sister-group relationship between the Clistosaccidae and the Thompsoniidae. None of the lattice organs of A. kussakini are oval or match-shaped. In M. vancouverensis there is only a single anterior pair of lattice organs, a probable autapomorphy. But the morphology of the M. vancouverensis cyprid is otherwise very reminiscent of those in the Rhizocephala suborder Kentrogonida, such as in being ornamented with numerous (> 100) carapace setae in no fixed pattern compared with the few (< 80) setae in a fixed disposition seen in the other akentrogonids. In cyprids of all Akentrogonida examined, the lattice organs lack a large terminal pore. By out-group comparison with all other cirripedes and with the Ascothoracida, this is regarded as an apomorphy that supports the monophyly of the Akentrogonida.

Phase relations of KC10 4 and KBF 4 to high pressures

Bopyrid isopods and rhizocephalan barnacles are obligate parasite crustaceans which harm their decapod hosts. However, to the best of our knowledge, studies have not compared which of these parasites has a greater parasitic effect on its hosts. Here, the parasitic effect of the bopyrid isopod, Allokepon hendersoni, and an unidentified sacculinid rhizocephalan species, infesting the same population of portunid crabs, Charybdis bimaculata, was investigated and compared for the first time. Samples were collected from the bycatch of a trawl fishery in Tosa Bay, Japan. A total of 2601 crabs were collected, of which 14 (0.55%) were parasitized by the bopyrid and 21 (0.82%) by the rhizocephalan. One of the two female crabs parasitized by the bopyrid was ovigerous (with much fewer eggs than unparasitized females). No ovigerous crab was found from the eight females parasitized by the rhizocephalan. Because only two female crabs were parasitized by the bopyrid, the following analyses were made using the male crabs. Both parasites reduced the wet weight (crab condition) and the cheliped size (secondary growth) of C. bimaculata, but the impact of the parasitism did not differ between the parasite species. The size of the abdominal flap of male hosts was reduced by the bopyrid infestation; however, rhizocephalan infestation caused enlargement of the abdominal flap, which is an indication of feminization. The present study provides information on how the effect of these two parasitic castrators on the same host crab varies. A moderate decrease in crab condition and cheliped development was common among the parasites, suggesting that the degree of impact might be favorable for the survival of the two parasites species.