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Abstract Fish parasites are an important part of aquatic biodiversity and knowing these species and their interactions with their hosts helps in monitoring the aquatic biota. The present study investigated the ectoparasite crustacean fauna of ten fish species from the upper Araguari River, in the state of Amapá, northern Brazil. A total of 508 fish were collected and analyzed from July to November 2014, of which 82.6% (109) were parasitized by one or more crustacean ectoparasite species. In the ten host fish species, a total of 308 ectoparasite specimens were collected, from 12 taxa, such as Argulus multicolor Stekhoven, 1937, Argulus spinulosus Silva, 1980, Argulus sp.1, Argulus sp.2, Argulus sp.3, Dipteropeltis sp., Dipteropeltis hirundo Calman, 1912, Dolops bidentata Bouvier, 1899, Dolops striata Bouvier, 1899 (Argulidae), Braga fluviatilis Richardson, 1911, Braga amapaensis Thatcher, 1996 (Cymothoidae) and Excorallana berbicensis Boone, 1918 (Corallanidae). Higher levels of prevalence and abundance were recorded for Hoplias aimara (Valenciennes, 1847) and Tometes trilobatus Valenciennes, 1850, respectively. These ectoparasites were found in the fins, integument, mouth, and anus of the host fish. Argulus sp.2 and D. bidentata were the most abundant parasites (65.1%), and had the highest species richness. This study registered 36 novel host-parasite interactions, and thus represents a new record for all host species here examined.

Chonopeltis lisikili Van As and Van As, 1996 was original described from the Eastern Caprivi (Namibia) and the Okavango System (Botswana), collected from five Synodontis Cuvier, 1816 (Mochokidae) species. This fish genus is endemic to Africa, with 130 valid species, making it one of the most species-rich and widely distributed mochokid catfish family. During parasitological surveys conducted in the Phongolo River (South Africa), a Chonopeltis Thiele, 1900 species was collected from Synodontis zambezensis Peters, 1852. In total, 21 adult females, four adult males, as well as representatives of the larval developmental stages were found. After morphological comparison, this branchiuran was identified as C. lisikili. This paper provides additional information on body measurements and morphological information of all the life stages of the Phongolo material. Information on Chonopeltis material collected from the Okavango River, Botswana, not yet published is also included as well as the first molecular characterisation of a Chonopeltis species using COI and 18 S rRNA partial sequencing. The endemicity of the host and fish lice necessitates a discussion on the host origin and distribution in more than one river system, particularly referring to host specificity for Chonopeltis species.

Dolops intermediaSilva, 1978 (Crustacea: Argulidae) was described from Rio Grande do Sul state in the southernmost state of Brazil. Here, we report a new occurrence of this species from Maranhão state, 1 of the northernmost states of Brazil. The infection of new hosts Colossoma macropomum (Cuvier, 1816), Oreochromis niloticus (Linnaeus, 1758), Hoplias malabaricus (Bloch, 1794), Leporinus friderici (Bloch, 1794), and Pygocentrus nattereri (Kner, 1858) is reported for the first time. Additional morphological information is provided for some structures, and the ratios for body measurements are provided for the first time. [ABSTRACT FROM AUTHOR]

Abstract Background It is hypothesised that being a blood-feeding ectoparasite, Argulus foliaceus (Linnaeus, 1758), uses similar mechanisms for digestion and host immune evasion to those used by other haematophagous ecdysozoa, including caligid copepods (e.g. sea louse). We recently described and characterised glands associated with the feeding appendages of A. foliaceus using histological techniques. The work described in the present study is the first undertaken with the objective of identifying and partially characterising the components secreted from these glands using a proteomic approach. Methods Argulus foliaceus parasites were sampled from the skin of rainbow trout (Oncorhynchus mykiss), from Loch Fad on the Isle of Bute, Scotland, UK. The proteins from A. foliaceus secretory/excretory products (SEPs) were collected from the supernatant of artificial freshwater conditioned with active adult parasites (n = 5–9 per ml; n = 560 total). Proteins within the SEPs were identified and characterised using LC-ESI-MS/MS analysis. Data are available via ProteomeXchange with identifier PXD016226. Results Data mining of a protein database translated from an A. foliaceus dataset using ProteinScape allowed identification of 27 predicted protein sequences from the A. foliaceus SEPs, each protein matching the criteria of 2 peptides with at least 4 contiguous amino acids. Nine proteins had no matching sequence through OmicsBox (Blast2GO) analysis searches suggesting that Argulus spp. may additionally have unique proteins present in their SEPs. SignalP 5.0 software, identified 13 proteins with a signal sequence suggestive of signal peptides and supportive of secreted proteins being identified. Notably, the functional characteristics of identified A. foliaceus proteins/domains have also been described from the salivary glands and saliva of other blood-feeding arthropods such as ticks. Identified proteins included: transporters, peroxidases, metalloproteases, proteases and serine protease inhibitors which are known to play roles in parasite immune evasion/induction (e.g. astacin), immunomodulation (e.g. serpin) and digestion (e.g. trypsin). Conclusions To our knowledge, the present study represents the first proteomic analysis undertaken for SEPs from any branchiuran fish louse. Here we reveal possible functional roles of A. foliaceus SEPs in digestion and immunomodulation, with a number of protein families shared with other haematophagous ectoparasites. A number of apparently unique secreted proteins were identified compared to other haematophagous ecdysozoa.

Koi carp are globally known for their colors and cultural significance. The introduction of these fish to new environments poses a threat to local biodiversity, in addition to releasing parasites, such as argulid ectoparasites. This study presents a record of Argulus japonicus infecting carp in an artificial lake in Southern Brazil using morphological and molecular methods, with a 100% prevalence (n = 3) and a mean intensity of 21.6 parasites per host, distributed over the body surface. The invasion history of hosts in the study locality indicates that the introduction of A. japonicus occurred decades before its first formal record in Brazil., Competing Interests: Declaration of competing interest None., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Abstract The aim of this study was to evaluate the impacts of metazoan parasites on hematological and biochemical parameters and relative condition factor of tambaqui (Colossoma macropomum ) farmed in northern Brazil. A total of 32 juvenile fish were captured from a commercial fish farm located in the municipality of Rio Preto da Eva, Amazonas state, Brazil. Parasite prevalence was 100% for Anacanthorus spathulatus, Mymarothecium boegeri and Notozothecium janauachensis, 100% for Neoechinorhynchus buttnerae and 53.13% for Dolops geayi. The greatest mean parasite intensity was found in acantocephalans followed by monogeneans and branchiuran crustaceans. A negative correlation was observed between abundance of N. buttnerae and hematocrit percentage, hemoglobin concentration, total thrombocyte count and glucose and between abundance of the monogenean and glucose concentration. Parasitic infections caused damage in tambaqui in terms of the observed hematological parameters that were characterized by hypochromic anemia and thrombocytopenia, which are important parameters to be used in parasitic diagnosis. This study is the first record of the occurrence of Dolops geayi in farmed tambaqui in the Amazon.

Chonopeltis lisikili Van As and Van As, 1996 was original described from the Eastern Caprivi (Namibia) and the Okavango System (Botswana), collected from five Synodontis Cuvier, 1816 (Mochokidae) species. This fish genus is endemic to Africa, with 130 valid species, making it one of the most species-rich and widely distributed mochokid catfish family. During parasitological surveys conducted in the Phongolo River (South Africa), a Chonopeltis Thiele, 1900 species was collected from Synodontis zambezensis Peters, 1852. In total, 21 adult females, four adult males, as well as representatives of the larval developmental stages were found. After morphological comparison, this branchiuran was identified as C. lisikili. This paper provides additional information on body measurements and morphological information of all the life stages of the Phongolo material. Information on Chonopeltis material collected from the Okavango River, Botswana, not yet published is also included as well as the first molecular characterisation of a Chonopeltis species using COI and 18 S rRNA partial sequencing. The endemicity of the host and fish lice necessitates a discussion on the host origin and distribution in more than one river system, particularly referring to host specificity for Chonopeltis species. [Display omitted] • First COI sequence for genus Chonopeltis. • First phylogenetic analysis of Branchiurans using 18 S rRNA. • New host and locality record for Chonopeltis lisikili. • First illustrations of the majority of C. lisikili life stages, in fact for any of the known Chonopeltis species. [ABSTRACT FROM AUTHOR]

Se registra por primera vez para Colombia (cuenca del Orinoco), cuatro especies de crustáceos ectoparásitos en rayas de agua dulce (Potamotrygonidae), género Potamotrygon. Dos que pertenecen a la subclase Branchiura, familia Argulidae: Dolops reperta (Bouvier, 1899) y D. striata (Bouvier, 1899) y dos al orden Isopoda, correspondientes a las familias Cymothoidae y Corallanidae, representadas con las especies Braga cigarra (Szidat & Schubart, 1960) y Excorallana delaneyi (Stone & Heard, 1989), respectivamente.

Background: It is hypothesised that being a blood-feeding ectoparasite, Argulus foliaceus (Linnaeus, 1758), uses similar mechanisms for digestion and host immune evasion to those used by other haematophagous ecdysozoa, including caligid copepods (e.g. sea louse). We recently described and characterised glands associated with the feeding appendages of A. foliaceus using histological techniques. The work described in the present study is the first undertaken with the objective of identifying and partially characterising the components secreted from these glands using a proteomic approach. Methods: Argulus foliaceus parasites were sampled from the skin of rainbow trout (Oncorhynchus mykiss), from Loch Fad on the Isle of Bute, Scotland, UK. The proteins from A. foliaceus secretory/excretory products (SEPs) were collected from the supernatant of artificial freshwater conditioned with active adult parasites (n = 5–9 per ml; n = 560 total). Proteins within the SEPs were identified and characterised using LC-ESI-MS/MS analysis. Data are available via ProteomeXchange with identifier PXD016226. Results: Data mining of a protein database translated from an A. foliaceus dataset using ProteinScape allowed identification of 27 predicted protein sequences from the A. foliaceus SEPs, each protein matching the criteria of 2 peptides with at least 4 contiguous amino acids. Nine proteins had no matching sequence through OmicsBox (Blast2GO) analysis searches suggesting that Argulus spp. may additionally have unique proteins present in their SEPs. SignalP 5.0 software, identified 13 proteins with a signal sequence suggestive of signal peptides and supportive of secreted proteins being identified. Notably, the functional characteristics of identified A. foliaceus proteins/domains have also been described from the salivary glands and saliva of other blood-feeding arthropods such as ticks. Identified proteins included: transporters, peroxidases, metalloproteases, proteases and serine protease inhibitors which are known to play roles in parasite immune evasion/induction (e.g. astacin), immunomodulation (e.g. serpin) and digestion (e.g. trypsin). Conclusions: To our knowledge, the present study represents the first proteomic analysis undertaken for SEPs from any branchiuran fish louse. Here we reveal possible functional roles of A. foliaceus SEPs in digestion and immunomodulation, with a number of protein families shared with other haematophagous ectoparasites. A number of apparently unique secreted proteins were identified compared to other haematophagous ecdysozoa. [ABSTRACT FROM AUTHOR]

Four species of ectoparasite crustaceans are registered for the first time from the Orinoco River Basin (Colombia) in freshwater stingrays of the family Potamotrygonidae, genus Potamotrygon. Two belong to the Branchiura subclass, Argulidae family: Dolops reperta (Bouvier, 1899) and D. striata (Bouvier, 1899), and two to the Isopoda Order, corresponding to the families Cymothoidae and Corallanidae, represented by the species Braga cigarra (Szidat & Schubart, 1960) and Excorallana delaneyi (Stone & Heard, 1989), respectively. [ABSTRACT FROM AUTHOR]

The aim of this study was to investigate parasites crustacean fauna in Arapaima gigas, Cichla monoculus, Cichla ocellaris, Cichla jariina, Satanoperca jurupari, Leporinus friderici, Leporinus fasciatus, Hoplias malabaricus, Phractocephalus hemioliopterus, Serrasalmus altispinis, Pseudoplatystoma tigrinum and Potamotrygon motoro of the State Amapá and Pará, in northern Brazil. A total of 242 parasites, including Argulus elongatus, Argulus multicolor, Argulus juparanaensis, Argulus nattereri, Dolops discoidalis, Dolops longicauda, Braga patagonica, Braga fluviatilis, Livoneca guianensis and undetermined Lernaeidae, were collected from these hosts. The Argulus species had the greatest richness among the community of parasitic crustaceans. There was a low abundance of parasites among the hosts, other than D. discoidalis, was most abundant in the integument of A. gigas and P. tigrinum. Finally, the present study reported nine new hosts for the crustacean parasite species and expanded knowledge of the occurrence of some parasite species in the Jari River basin, in eastern Amazon. [ABSTRACT FROM AUTHOR]

An argulid branchiuran, Argulus quadristriatus Devaraj and Ameer Hamsa, 1977, is reported on the basis of specimens of both sexes isolated from two species of goatfish [Upeneus tragula Richardson, 1846 and Parupeneus ciliatus (Lacepède, 1802) (Perciformes: Mullidae)] and the goby Favonigobius reichei (Bleeker, 1854) (Perciformes: Gobiidae) collected in Kagoshima Bay and off both the east and west coasts of Okinawa-jima island, southern Japan. The branchiuran is easily distinguished from its nine previously recorded congeners in Japanese waters by the presence of one posterior spine on the antennule and scales ornamenting the basal half of the mouthtube. These are the first records of this branchiuran from the North Pacific Ocean including Japanese waters, and the three fish species mentioned above are new hosts for it. [ABSTRACT FROM AUTHOR]

An adult male of Argulus coregoni Thorell, 1864, was collected from the body surface of an ayu, Plecogiossus altivelis altivelis (Temminck & Schlegel, 1846), in the middle reaches of the Ani River, a tributary of the Yoneshiro River, Akita Prefecture, northern Honshu, Japan. The previous northernmost record of A. coregoni in Japan is from Fukushima Prefecture, and the present collection extends its geographical distribution from Fukushima Prefecture northward to Akita Prefecture in Japan. [ABSTRACT FROM AUTHOR]

Hemocyanin transports oxygen in the hemolymph of many arthropod species. Within the crustaceans, this copper-containing protein was thought to be restricted to Malacostraca, while other crustacean classes were assumed to employ hemoglobin or lack any respiratory protein. Only recently it has become evident that hemocyanins also occur in Remipedia and Ostracoda. Here we report for the first time the identification and characterisation of hemocyanin in the fish louse Argulus, which belongs to the class of Branchiura. This finding indicates that hemocyanin was the principal oxygen carrier in the stem lineage of the pancrustaceans, but has been lost independently multiple times in crustacean taxa. We obtained the full-length cDNA sequences of two hemocyanin subunits of Argulus foliaceus by a combination of RT-PCR, RACE and Illumina sequencing of the transcriptome. In addition, one full-length and one partial cDNA sequence were derived from the transcriptome data of Argulus siamensis. Western blot analysis confirmed the presence of at least two hemocyanin subunits in A. foliaceus, which are expressed at the mRNA level at a 1:3.5 ratio. The addition to the branchiuran hemocyanin subunits to a multiple sequence alignment of arthropod, hemocyanins improved the phylogenetic resolution within the pancrustacean hemocyanins. Malacostracan, ostracod and branchiuran hemocyanins are distinct from the hexapod and remipede hemocyanins, reinforcing the hypothesis of a close relationship of Remipedia and Hexapoda. Notably, the ostracod hemocyanins are paraphyletic with respect to the branchiuran hemocyanins, indicating ancient divergence and differential loss of distinct subunit types. [ABSTRACT FROM AUTHOR]

Several cases of epibiosis by macro-invertebrates on Orconectes limosus were recorded during our research on the biology of populations of this invasive crayfish species in the Czech Republic. (1) In 2001, we observed a high infestation of O. limosus by native species of branchiobdellidan worms (Annelida: Branchiobdellidae) in the river Elbe at Obríství (Central Bohemia). Four European Branchiobdella species were collected and identified from three crayfish specimens: B. pentodonta (52%), B. balcanica (24%), B. parasita (18%) and B. hexodonta (6%). In 2003, only a single crayfish was found bearing branchiobdellidans (5 specimens of B. parasita only) in the same locality. No branchiobdellidans on O. limosus have been confirmed since. (2) A flooded sandpit Lhota near Brandýs nad Labem is the only Czech locality where the settlement of Dreissena polymorpha (Mollusca, Bivalvia) on an O. limosus body was recorded. The crayfish lost the bivalves by moulting in summer; new mussels had settled by late summer and early autumn. (3) Females of the fish louse Argulus cf. foliaceus (Crustacea: Branchiura) lay egg-strings on any hard substrate, including the crayfish exoskeleton. Such egg-masses were found on up to 65% of O. limosus specimens in the above-mentioned sandpit. (4) Bryozoan colonies of Plumatella repens were found twice on crayfish in the rivers Elbe (Labe) and Cidlina.

Argulus coregoni Thorell, 1864 was collected from the body surface of ayu, Plecoglossus altivelis altivelis (Temminck & Schlegel, 1846), from six rivers in Gifu, Achi, Mie, and Shiga prefectures, central Honshu, Japan. The collection of A. coregoni from Mie Prefecture represents its new prefectural record. Salmonids have been regarded as the major hosts of A. coregoni in Japanese rivers, but the present collections suggest that ayu is also important as a riverine host of the parasite. [ABSTRACT FROM AUTHOR]

The lakeweed chub, Ischikauia steenackeri (Sauvage, 1883), was originally distributed in the Yodo River system including Lake Biwa, central Japan, and has recently established its populations as a domestic alien species in various inland waters in western and central Japan, including Kyushu. An adult female of Argulus japonicus Thiele, 1900 was collected from the body surface of a lakeweed chub in an irrigation canal connected to the Hase River, a tributary of the Nishi River within the Onga River system in Fukuoka Prefecture, northern Kyushu. This repersents a new prefectural and host records for A. japonicus. This parasite is considered not to have been introduced to the sampling site by I. steenackeri but to naturally occur in the river system. [ABSTRACT FROM AUTHOR]

The paper presents an overview of the history of aquatic parasitology of freshwater fish in southern Africa. Aquatic parasitology has become a very popular field of specialisation in South Africa and is currently practised by a number of individual research groups at different universities. So far most research deals with parasites of wild fish rather than pathogens of aquaculture. It is written as a narrative and summarises the aquatic parasitology research mostly on fish parasites carried out at the universities of Cape Town, Free State, Johannesburg, Limpopo, Pretoria, North West and Sefako Makgatho Health Sciences. It is not intended to be a review in the classical way because it deals with parasites ranging from Protozoa to Arthropoda. [ABSTRACT FROM PUBLISHER]

Abstract: The present study has been carried out to describe the cell morphology of the developing male gametes in a fish ectoparasite, Argulus bengalensis Ramakrishna, 1951. With respect to cell volume and nucleoplasmic index, spermatogonia are the smallest and primary spermatocytes are the largest in this lineage. The spermatogonia and the differentiating spermatogenic cells are in separate niches and confined to different enclaves within each testicular lobe. Spermiogenesis occurs within the inner enclave of each testicular lobe. During this process the nucleus becomes streamlined; an acrosome is formed, axoneme is originated, and residual cytoplasm is discarded through the flagellum. The sperm cell morphology displays a general pattern comprising head, mid-piece, and a full length flagellum. In the axoneme 9+2 arrangement of the microtubule is conserved. In addition to the axoneme, some more singlet microtubules are found surrounding a fiber sheath and around one of the mitochondria adjacent to the axoneme. This arrangement indicates a close phylogenetic relationship with pentastomida. In the present study, structure and formation of spermatophore are described in this branchiuran parasite. [Copyright &y& Elsevier]

Efficacy of in-feed administration of emamectin benzoate (EMB) was evaluated in commercial culture farms of Asian seabass (Lates calcarifer), grey mullet (Mugil cephalus), pearlspot (Etroplus suratensis), Indian major carp (Labeo rohita), goldfish (Carassius auratus) and koi carp (Cyprinus rubrofuscus) in different parts of India. Fish naturally infested with Caligus minimus , Argulus quadristriatus , Argulus siamensis , Lernanthropsis mugilii and Lernaea cyprinacea were administered medicated feed for 7–10 consecutive days (@ 50 μg kg−1 body weight day−1). Efficacy of the treatment was monitored by recording the parasitic intensity (PI). The study revealed PI was zero on head and gills within four days while parasites remained on trunk and fins till six days both in seabass and pearlspot. However, zero PI was achieved on 6th day on head and gills, and on 8th day on trunk and fins in seabass and Indian major carp. Treatment with EMB for seven days was effective against C. minimus , A. quadristriatus , A. siamensis and L. mugilii in E. suratensis , L. calcarifer, L. rohita and M. cephalus , respectively while L. cyprinacea infestation in seabass, goldfish and koi carp needed an extended treatment up to 10 days. In all the cases no recurrence was observed for 60 days post medication. Study revealed that there is a need for integrated pest management (IPM) program for effective prevention and control of parasitic infestation in commercially important food and ornamental fish cultures. • Efficacy of emamectin benzoate was determined in parasite infected fish species reared in different culture systems. • The efficacy was 100% against Argulus spp., Caligus spp. and Lernanthropsis spp. on 7th day, and against Lernaea spp. following 7 days treatment. • Administration for 10 days was required for the control of Lernaea spp. • No re-infestation of parasites was observed for a period of 60 days post medication. [ABSTRACT FROM AUTHOR]

The genus Dolops is one of only four genera in Branchiura, and as most other members of the group it is also ectoparasite on freshwater fish. The large majority of investigations of the genus concern the single species oí Dolops found in southern Africa, D. ranarum, although most of the remaining species are found in South America. In this paper, we present the first morphological description of the larva of D. carvalhoi using light- and scanning electron microscopy. This is the first detailed account of the larvae of any non-African species of Dolops, and our comparison of the larvae of D. carvalhoi and D. ranarum based on literature data show them to be surprisingly alike. The general implications of these findings are discussed in a phylogenetic context, with special emphasis on the different larval types found in the other genera of Branchiura. [ABSTRACT FROM AUTHOR]

Abstract: The process of sperm transfer is somewhat enigmatic in Argulus, even though copulation has been witnessed. A breeding colony of Argulus japonicus was kept under laboratory conditions in order to study reproduction in the species. Pairs in copula were removed and studied with histology and scanning electron microscopy to describe the mechanism of sperm transfer. Sections of copulating pairs revealed sperm on the accessory copulatory structures of the male’s swimming legs; and scanning electron microscopy showed that sperm transfer occurs in three phases which can be differentiated to 10 different stages. Sperm transfer occurs via a spermatophore which is extruded from the genital aperture of the male and is then transferred to the socket on the third pair of legs of the male, before being transferred into the spermathecae of the female via the spermathecal spines. This is the first observation of a spermatophore in Argulus. [ABSTRACT FROM AUTHOR]

Abstract: The Branchiura are fish ectoparasites which include the well-known and very commonly found genus Argulus. But the Branchiura also contains the monotypic genus Dipteropeltis, about which only very little is known. In this investigation of two female paratype specimens we describe for the first time the external morphology of Dipteropeltis based on Scanning Electron Microscopy. We show that Dipteropeltis has a pre-oral spine, unique suction disc support structures, an Argulus-like mouth cone and second maxillae with a combination of features seen in both Argulus and Chonopeltis. These data are compared with data from the three remaining branchiuran genera Dolops, Chonopeltis and Argulus and discussed in a phylogenetic context based on a recently proposed phylogeny of the group. [Copyright &y& Elsevier]

Pentastomida are parasites of tetrapods (especially reptiles) and have had long contentious relationships. Traditionally seen as a separate phylum with arthropod affinities, recent ultrastructural and molecular evidence unites them to branchiuran crustaceans (fish lice). However, the discovery of Cambrian pentastomids has been interpreted to refute the view that pentastomids are closely related to advanced crustaceans (of presumably recent origins). Bayesian phylogenetic and relaxed-clock analyses of molecular data (18S and 28S ribosomal RNA, elongation factor 1α and 2, RNA polymerase II subunit) help reconcile these apparently contradictory views: a pentastomid-branchiuran grouping is recovered, but this clade is the sister group to other Tetraconata, whereas all other crustaceans form a monophyletic group that is sister to hexapods. Crustacean monophyly is widely supported by morphology, but has rarely been retrieved in other molecular studies. Estimated molecular dates for pentastomid origins range from ∼490 to ∼520 million years ago, consistent with Cambrian fossils. The basal position of branchiurans within tetraconatans means pentastomids can be simultaneously related to branchiurans, and also have a deep fossil record. All Cambrian pentastomids are tiny, resembling the larvae of living pentastomids that infect intermediate hosts including fish; as definitive hosts (tetrapods) of modern pentastomids were absent in the Cambrian, it is possible that the small Cambrian forms were adults rather than larvae, and their entire life cycle was confined to small fish-like vertebrates that were then present. [ABSTRACT FROM AUTHOR]