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Example images of ectoparasites infecting various host species. A) Antimora rostrata with a lernaeopodid copepod (1059 m, Alvin Canyon); B) Synaphobranchus sp. with a sphyriid copepod (820 m, open slope); C) Nezumia bairdii with a sphyriid copepod parasitized by eight leeches (1035 m, Phoenix Canyon; D), Black fish, Diaphus sp. with an unknown siphonostomatid copepod (1130 m, cold seep) attached behind dorsal fin; E) N. bairdii with aegid isopod (780 m, open slope); F) Hoplostethus mediterraneus with a cymothoid isopod (744 m, Nygren Canyon); G) Amblyraja radiata with aegid isopod (1010 m, Alvin Canyon); and H) Cottunculus thomsonii with gnathiids (1210 m, Oceanographer Canyon).
Source publication
A complete understanding of how parasites influence marine ecosystem functioning requires characterizing a broad range of parasite-host interactions while determining the effects of parasitism in a variety of habitats. In deep-sea fishes, the prevalence of parasitism remains poorly understood. Knowledge of ectoparasitism, in particular, is limited...
Contexts in source publication
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... families of siphonostomatoid copepods were identified on three species of fishes (Fig. 2AeC). Copepods from the family Ler- naeopodidae, sized at approximately 1e3 cm total length (TL), were observed on A. rostrata ( Fig. 2A). Copepods were attached to anal fins, below second dorsal fins, and behind the eyes. At least two species (likely Lophoura spp.) from the family Sphyriidae infected Synaphobranchus spp. (Fig. 2B) and N. ...
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... families of siphonostomatoid copepods were identified on three species of fishes (Fig. 2AeC). Copepods from the family Ler- naeopodidae, sized at approximately 1e3 cm total length (TL), were observed on A. rostrata ( Fig. 2A). Copepods were attached to anal fins, below second dorsal fins, and behind the eyes. At least two species (likely Lophoura spp.) from the family Sphyriidae infected Synaphobranchus spp. (Fig. 2B) and N. bairdii (Fig. 2C). The copepods (~2e3 cm TL) infecting N. bairdii were attached directly behind the dorsal fin, whereas larger ...
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... species of fishes (Fig. 2AeC). Copepods from the family Ler- naeopodidae, sized at approximately 1e3 cm total length (TL), were observed on A. rostrata ( Fig. 2A). Copepods were attached to anal fins, below second dorsal fins, and behind the eyes. At least two species (likely Lophoura spp.) from the family Sphyriidae infected Synaphobranchus spp. (Fig. 2B) and N. bairdii (Fig. 2C). The copepods (~2e3 cm TL) infecting N. bairdii were attached directly behind the dorsal fin, whereas larger copepods (~4e5 cm TL) infecting Synaphobranchus spp. were attached mid body; either laterally or ventrally. Unidentified siphonostomatoid copepods were observed (~2 cm TL) on mesopelagic fishes, ...
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... Copepods from the family Ler- naeopodidae, sized at approximately 1e3 cm total length (TL), were observed on A. rostrata ( Fig. 2A). Copepods were attached to anal fins, below second dorsal fins, and behind the eyes. At least two species (likely Lophoura spp.) from the family Sphyriidae infected Synaphobranchus spp. (Fig. 2B) and N. bairdii (Fig. 2C). The copepods (~2e3 cm TL) infecting N. bairdii were attached directly behind the dorsal fin, whereas larger copepods (~4e5 cm TL) infecting Synaphobranchus spp. were attached mid body; either laterally or ventrally. Unidentified siphonostomatoid copepods were observed (~2 cm TL) on mesopelagic fishes, including two Diaphus sp. in- ...
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... The copepods (~2e3 cm TL) infecting N. bairdii were attached directly behind the dorsal fin, whereas larger copepods (~4e5 cm TL) infecting Synaphobranchus spp. were attached mid body; either laterally or ventrally. Unidentified siphonostomatoid copepods were observed (~2 cm TL) on mesopelagic fishes, including two Diaphus sp. in- dividuals ( Fig. 2D) and four unidentified myctophids. Copepods were attached behind the dorsal fins of myctophids. In all cases, only one copepod was observed on a single fish. Hyperparasitism was observed on N. bairdii, with each sphyriid copepod infected by at least three to eight leeches (Fig. ...
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... on mesopelagic fishes, including two Diaphus sp. in- dividuals ( Fig. 2D) and four unidentified myctophids. Copepods were attached behind the dorsal fins of myctophids. In all cases, only one copepod was observed on a single fish. Hyperparasitism was observed on N. bairdii, with each sphyriid copepod infected by at least three to eight leeches (Fig. ...
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... the three families of isopods that infected demersal fishes, gnathiids were the most common, with 1 to 45 individuals infect- ing at least one side of each individual fish. Gnathiids were trans- lucent, attached to all fins, heads, and sides of bodies ( Fig. 2H), and ranged in size from 1 to 3 mm TL. Distinct species of aegids infected A. radiata and N. bairdii. One large (~2.5 cm TL) aegid was attached mid-way on the body of A. radiata, at the juncture of the left pec- toral wing and the central disk (Fig. 2G). The other fish individual had at least 15 smaller (1e2 mm TL) aegids attached to ...
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... fish. Gnathiids were trans- lucent, attached to all fins, heads, and sides of bodies ( Fig. 2H), and ranged in size from 1 to 3 mm TL. Distinct species of aegids infected A. radiata and N. bairdii. One large (~2.5 cm TL) aegid was attached mid-way on the body of A. radiata, at the juncture of the left pec- toral wing and the central disk (Fig. 2G). The other fish individual had at least 15 smaller (1e2 mm TL) aegids attached to both the wings and the central disk. Five N. bairdii were observed each with an aegid isopod. Each aegid (~2e3 cm TL) was attached behind the dorsal fin (Fig. 2E). One cymothoid isopod (~4 cm TL) was observed attached on the side of the body below the ...
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... mid-way on the body of A. radiata, at the juncture of the left pec- toral wing and the central disk (Fig. 2G). The other fish individual had at least 15 smaller (1e2 mm TL) aegids attached to both the wings and the central disk. Five N. bairdii were observed each with an aegid isopod. Each aegid (~2e3 cm TL) was attached behind the dorsal fin (Fig. 2E). One cymothoid isopod (~4 cm TL) was observed attached on the side of the body below the dorsal fin of H. mediterraneus (Fig. ...
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... individual had at least 15 smaller (1e2 mm TL) aegids attached to both the wings and the central disk. Five N. bairdii were observed each with an aegid isopod. Each aegid (~2e3 cm TL) was attached behind the dorsal fin (Fig. 2E). One cymothoid isopod (~4 cm TL) was observed attached on the side of the body below the dorsal fin of H. mediterraneus (Fig. ...
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... thomsonii] or swimming normally (e.g., sharks, chi- maeras, ophidiids, morids, macrourids, synaphobranchids) either close to or just a few meters above the seafloor. Only a few of the fishes that had large ectoparasites appeared to be behaving abnormally. One N. bairdii individual with a sphyriid copepod hyperparasitized by eight leeches (Fig. 2C) appeared to be under- weight than other individuals of similar total lengths (~15 cm TL). This individual was observed swimming in circles and appeared to be leaning towards one side (Suppl. Video). One Hoplostethus mediterraneus with a large cymothoid isopod on its left side was observed making short, erratic movements using its ...
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Citations
... They are among the most common ectoparasites of teleost and elasmobranch fishes and are ubiquitous in marine environments (Grutter 1994;Ferreira et al. 2009;Coile and Sikkel 2013). As such, their biomass likely exceeds that of terrestrial blood-feeding arthropods (Smit and Davies 2004;Tanaka 2007;Kirkim et al. 2011;Quattrini and Demopoulos 2016). Considering the profound effects blood-feeding arthropods have on terrestrial communities, investigating the dynamics of gnathiid-host interactions is essential to our understanding of ecological processes in marine environments. ...
Appreciation for the role of cryptofauna in ecological systems has increased dramatically over the past decade. The impacts blood-feeding arthropods, such as ticks and mosquitos, have on terrestrial communities are the subject of hundreds of papers annually. However, blood-feeding arthropods have been largely ignored in marine environments. Gnathiid isopods, often referred to as “ticks of the sea”, are temporary external parasites of fishes. They are found in all marine environments and have many consequential impacts on host fitness. Because they are highly mobile and only associated with their hosts while obtaining a blood meal, their broader trophic connections are difficult to discern. Conventional methods rely heavily on detecting gnathiids on wild-caught fishes. However, this approach typically yields few gnathiids and does not account for hosts that avoid capture. To overcome this limitation, we sequenced blood meals of free-living gnathiids collected in light traps to assess the host range and community-dependent exploitation of Caribbean gnathiid isopods. Using fish-specific COI (cox1) primers, sequencing individual blood meals from 1060 gnathiids resulted in the identification of 70 host fish species from 27 families. Comparisons of fish assemblages to blood meal identification frequencies at four collection sites indicated that fishes within the families Haemulidae (grunts) and Lutjanidae (snappers) were exploited more frequently than expected based on their biomass, and Labrid parrotfishes were exploited less frequently than expected. The broad host range along with the biased exploitation of diel-migratory species has important implications for the role gnathiid isopods play in Caribbean coral reef communities.
... In recent years, with the exploration of offshore oil and gas resources continuously expanding into deep-sea areas [1,2], remotely operated underwater vehicles (ROVs) have become increasingly favored by companies and organizations engaged in offshore oil and gas industries due to their advantages of low economic cost, high work efficiency, and the ability to perform long-duration operations in deep, complex, and hazardous underwater environments [3][4][5][6]. ROVs are now intensively applied in various fields such as the sea oil industry for the purpose of infrastructure installation, device maintenance and repair, and pipeline inspection, etc. [7][8][9]. In science research activities, many researchers [10][11][12][13][14][15] have also used ROVs to accomplish inspection and sampling tasks like scanning seabed geomorphy and submarine biosampling in the deep-sea environment. ...
Remotely operated underwater vehicles (ROVs) have been widely used in deep-sea resource exploitation and industrial engineering operations. To perform these tasks accurately in the deep-sea environment, stable motion control has become a key area of research on ROV systems, which has led to the importance of analyzing the hydrodynamic characteristic of ROVs. But a systematic methodology for analyzing the hydrodynamic characteristics of ROVs is still lacking nowadays. In this paper, systematic numerical simulation methods for analyzing hydrodynamic characteristics and shape optimization of a work-class ROV are conducted, and details of simulation procedures based on computational fluid dynamics are studied, which can be a foundation for robust controller design.
... With improvements in photographic technology, especially the ability to photograph small organisms at night, photography of free-living fishes is now a viable tool for characterizing host-parasite associations. Indeed, high-resolution cameras mounted on submarines have been used to detect gnathiids and other external parasites on fishes in the deep ocean (Quattrini and Demopoulos 2016). A necessary next step is for divers to learn to recognize these parasites. ...
We present observations of fish-parasitic gnathiid isopods infesting bothid flounders at night in shallow reef habitat in the central Philippines.
... They are among the most common ectoparasites of teleost and elasmobranch shes and are ubiquitous in marine environments (Grutter 1994;Ferreira et al. 2009;Coile and Sikkel 2013). As such, their biomass likely exceeds that of terrestrial blood-feeding arthropods (Smit and Davies 2004;Tanaka 2007;Kirkim et al. 2011;Quattrini and Demopoulos 2016). Considering the profound effects blood-feeding arthropods have on terrestrial communities, investigating the dynamics of gnathiid-host interactions is essential to our understanding of ecological processes in marine environments. ...
Appreciation for the role of cryptofauna in ecological systems has increased dramatically over the past decade. The impacts blood-feeding arthropods, such as ticks and mosquitos, have on terrestrial communities are the subject of hundreds of papers annually. However, blood-feeding arthropods have been largely ignored in marine environments. Gnathiid isopods, often referred to as “ticks of the sea”, are temporary external parasites of fishes. They are found in all marine environments and have many consequential impacts on host fitness. Because they are highly mobile and only associated with their hosts while obtaining a blood meal, their broader trophic connections are difficult to discern. Conventional methods rely heavily on detecting gnathiids on wild-caught fishes. However, this approach typically yields few gnathiids and does not account for hosts that avoid capture. To overcome this limitation, we sequenced blood meals of free-living gnathiids collected in light traps to assess the host range and community-dependent exploitation of Caribbean gnathiid isopods. Using fish-specific COI ( cox 1) primers, sequencing individual blood meals from 1,060 gnathiids resulted in the identification of 70 host fish species from 27 families. Comparisons of fish assemblages to blood meal identification frequencies at four collection sites indicated that fishes within the families Haemulidae (grunts) and Lutjanidae (snappers) were exploited more frequently than expected based on their biomass, and Labrid parrotfishes were exploited less frequently than expected. The broad host range along with the biased exploitation of diel-migratory species has important implications for the role gnathiid isopods play in Caribbean coral reef communities.
... Viruses have been reported from various fish helminths [48,49] and many more may exist [32]; all these helminth viruses are thus hyperparasites. Hyperparasites were frequently reported from parasitic copepods, including microsporidians [41] and a leech [79]. However, Ohtsuka et al. (2018) recently reviewed all organisms living on parasitic copepods but considered that most of them were epibionts, not parasites [70]. ...
Cyclocotyla bellones Otto, 1823 (Monogenea, Diclidophoridae) is one of the few monogenean species reported as hyperparasitic: the worms dwell on cymothoid isopods, themselves parasites of the buccal cavity of fishes. We present here observations based on newly collected monogenean specimens from Ceratothoa parallela (Otto, 1828), an isopod parasite of Boops boops off Algeria and also investigated its diet to address whether Cy. bellones is indeed a hyperparasite, i.e., whether it feeds on the isopod. We also compared the body shape of various monogeneans belonging to the same family as Cy. bellones , the Diclidophoridae, including Choricotyle cf. chrysophryi Van Beneden & Hesse, 1863, collected from Pagellus acarne off Algeria. No morphological character of the anterior organs suggested any special adaptation in Cy. bellones to the perforation of the crustacean cuticle. The wall of the oesophagus and of the intestine of Cy. bellones was lined with a dark pigment similar to what is usually observed in haematophagous polyopisthocotyleans, and which is derived from ingested fish blood. We noticed that an anterior elongate stem exists only in diclidophorids dwelling on parasitic isopods and never in those attached to the gills. We hypothesize that the anterior stem of the body of Cy. bellones is an anatomical adaptation for the monogenean to feed on the fish while dwelling on the isopod. We thus consider that Cy. bellones is an epibiont of the parasitic crustacean, as it uses it merely as an attachment substrate, and is not a true hyperparasite.
... In some extent, the prevalence is varying between the infested sites of the same fish (Figure 4) [26]. Ectoparasitism is prevalent in a variety of aquatic habitats and depths; however, the diversity of both the hosts and the ectoparasites decline in the deep-sea communities; in addition, ectoparasites are often detached during the collection of their hosts, which hinders their proper investigation [46]. In addition, illumination and the slow swimming speed in the dark are factors influencing the fish's susceptibility to the parasitic infestations [47]. ...
Crustaceans during a parasitism relationship with fish cause biological disruption and diseases to the hosting fish that threaten their life. Several crustacean groups were reported to parasitize different fish species as definitive hosts. Copepods, Isopods, and Branchiura are the major fish parasitic groups under the crustacea. They are mainly ectoparasites and occasionally, the small and microscopic members can infest the internal organs of their fish hosts. The infected fish are suffering from serious clinical signs and the infested organs are usually suffering from severe pathological disruptions and lesions. Treatment is taking place by several chemical and biological facilities. Here in the current article, the major crustaceans that parasitize on fish are reviewed by describing their corresponding harms to the hosting fish. Factors affecting the prevalence of the crustacean parasites and some of their hazardous effects on the infected fish were considered. In addition, trials for the chemical and biological control of the crustacean infestations are summarized.
... Among the parasitic isopods, gnathiids are potentially the most important ecologically, having even been included among priority "species" to support the functional integrity of coral reefs (Wolfe et al., 2020). Gnathiids are found in temperate, polar, and tropical oceans, from tide pools to the deep ocean (Smit and Davies, 2004;Tanaka, 2007;Quattrini and Demopoulos, 2016). They differ from other marine ectoparasites in two fundamental ways. ...
The distribution and abundance of organisms is typically shaped by multiple biotic and abiotic processes. Micropredators are parasite-like organisms that are smaller than their hosts and/or prey and feed on multiple hosts during a given life stage. Unlike typical parasites, however, they spend much or most of their time free-living, associating only temporarily with hosts. In the ocean, micropredators can impact multiple fish species, and in particular can have significant lethal and sub-lethal effects on newly settled fish. Although gnathiid isopods are abundant and primary micropredators in coral reef ecosystems, their impacts are relatively unexplored within sub-tidal temperate rocky reefs. We investigated the distribution of juvenile gnathiid isopods along sub-tidal temperate rocky reefs and tested trap methodology. We also quantified both the sub-lethal and lethal impacts of feeding-stage juvenile gnathiid isopods on juvenile, post settlement reef fish, Heterostichus rostratus (giant kelpfish). We were most interested in determining the relationship between gnathiid infestation level and fish swimming performance, in particular swimming metrics relevant to predator avoidance maneuvers. We found that Gnathia tridens was present in rocky reefs rather than embayments along the Southern California coastline and that within rocky reefs, gnathiids occurred in the highest densities in lighted traps. Surprisingly, we observed almost no influence of fish size or gnathiid sub-lethal infestation level on ambient or burst swimming performance metrics. However, burst duration was reduced by gnathiid infestation, which is important in predator avoidance. There were significant differences in survivorship among small fish compared to large fish as a result of gnathiid infestation. Larger fish survived higher numbers of gnathiids than smaller fish, indicating that parasite-induced mortality is greater for smaller fish. Investigations of the effects of micropredators on subsequent predator-mediated mortality, including the susceptibility of fishes and their individual responses to micropredators, can further contribute to our understanding of processes affecting recruitment in resident reef fish populations. Further research, especially within temperate sub-tidal ecosystems, is needed to understand and highlight the overlooked importance of micropredation in shaping fish populations within a reefscape.
... In fact, no Lophoura-parasitized C. caelorhincus were recorded from the entire eastern Mediterranean until, beginning in 2014, parasitized fish were noted, contemporaneously with higher abundance and higher incidence of infection in the Strait of Sicily. As this ectoparasite may adversely influence the population dynamics (Quattrini & Demopoulos, 2016) of a key deep-sea fish species in the Mediterranean, recording its prevalence is but a first step to understanding its possible effects on its host population. ...
Lophoura edwardsi, a rarely recorded deep-water parasitic copepod, is reported for the first time from the Eastern Mediterranean Sea. We document its incidence of infection in hauls made between 2014 and 2020 on the Israeli upper slope, southeastern Mediter-ranean Sea. All 15 post-metamorphic females parasitized the deep-water macrourid Coelorinchus caelorhincus. Molecular analysis of 5 specimens revealed four 18S rRNA and four COI haplotypes. Comparisons of 18S rRNA to the Siphonostomatoida sequences available in NCBI confirm L. edwardsi is closely related to the cofamilial Tripaphylus elongatus. The recent emergence of a small but stable population may augur changes in the Levantine slope ecosystem.
... Gnathiid isopods (Crustacea: Isopoda: Gnathiidae) are one of the most common crustacean parasites infesting marine fishes (Sikkel and Welicky 2019) and are found in all oceans at depths ranging from less than one meter to over 3000 m (Tanaka 2006;Quattrini and Demopoulos 2016). Gnathiids are parasitic only during the three larval instars of their life cycle, where they feed on blood and lymph from a host fish (Smit and Davies 2004). ...
Gnathiid isopods are common crustacean parasites that inhabit all oceans from shorelines to depths of over 3000 m and use chemical cues to find their marine fish hosts. While gnathiids are host-generalists, hosts vary in their susceptibility to infestation. However, the mechanisms that mediate differential susceptibility are unknown. Here we used a combination of field and laboratory experiments to investigate if the chemical attractiveness of hosts explains differences in susceptibility of Caribbean reef fishes to infestation by a common Caribbean gnathiid isopod, Gnathia marleyi. We showed that while G. marleyi can detect and locate hosts using only chemical cues, they do not exhibit a preference for chemical cues produced by more susceptible fish species. We conclude that species-specific chemical cues are not the main mechanism driving differences in host susceptibility to gnathiid isopod infestation and that visual or post-attachment factors such as ease of obtaining a blood meal are likely mediators.
... More than 369 species of fish isopods exist in 43 genera (see Chap. 3). They occur in and on fishes around the world, but mostly in the tropics and subtropics, in coastal waters, with some in freshwaters largely in South America with a few species in Africa and Asia (Bunkley- Williams and Williams 1998a;Smit et al. 2014), and even the deep ocean (Quattrini and Demopoulos 2016;Williams and Bunkley-Williams 2003). Some almost complete life cycles are known (e.g. ...
Different parasitic life strategies are described including four new life cycles: complex rebrooding, micro-male, mesoparasite and prey-predator transfer. Four new life cycle behaviours are named: nursery hiding, mid-moult stage, positive precursor (intraspecific antagonism) and negative precursor (ambush strategy). Further strategies discussed are opossum attack, double parasitism (doubling of the normal reproductive set), duplex arrangement (separated male-female pairs), simple rebrooding, and describing how displaced parasites and superinfections may partly elucidate life cycles. Proportional stunting masks life history effects of parasitism; cuckoo copepods are true parasites and not just associates; burrowing barnacles (acrothoracicans) are not parasites. Further findings based on life cycle information: branchiurans and pentastomes are possibly not related; firefly seed shrimp are not parasites; copepod pre-adult life cycle stages are common in the western pacific but rare in Caribbean; harpacticoids on vertebrates are not parasites; cuckoo copepods are true parasites; explained the importance of pennellid intermediate hosts. Crustacean parasite life cycles are largely unknown (1% of species). Most crustacean life cycles represent minor modifications from the ancestral free-living mode. Crustacean parasites have less complex and less modified life cycles than other major parasite groups. This limits their exploitation of, and effectiveness, in parasitism. However, these life cycles will be an advantage in Global Change. Most metazoan parasites will be eliminated while crustaceans (and nematodes) will inherit the new world of parasites.
