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Systematics of the Pearlfishes (Pisces: Carapidae)

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A cladistic classification based on swimbladder morphology, developmental and osteological characters of the vertebral column, fins, pectoral and pelvic girdles, gill arches, ethmoid, jaws, and habitat is presented. A total of 31 species in 7 genera are recognized in 2 subfamilies. The family is distributed over broad depth (0-2000 m) and latitudinal (65°N to 60°S) ranges. In general taxa with more plesiomorphic characters are found in deeper water and higher latitudes while taxa with more apomorphic characters are found in tropical, shallow water. Highly specialized inquiline behavior is found in both tribes and the phylogenetic hyopthesis specifies independent acquisition of molluscan inquilinism, at least. Within the Carapini the genus with the most apomorphic characters, Encheliophis, is the most successful holothurian inquiline and the only holothurian inquiline in the Indo-Pacific. Species in its sister genus, Carapus, "switch' to non-holothurian primary hosts (asteroids or ascidians) when sympatric with Encheliophis in the Indo-Pacific. Atlantic species of Carapus retain holothurians as primary hosts. Vicariance of sister species is frequently along depth or latitudinal gradients. -from Authors
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... 3a, 4). The genus Pyramodon includes four species (Markle and Olney, 1990): P. lindas, P. parini, P. punctatus, and P. ventralis. Vexillifer larvae of P. ventralis and P. punctatus have been described in the literature and their pigmentation is significantly different from that in our specimen (the midline of the body is unpigmented in the known larvae of the other species, in contrast with P. lindas that has spots along the midline of the body). ...
... Another character for the adults of Pyramodon is the position of the anal-fin origin relative to the dorsal-fin origin (DRAO). According to the key to the adults of the Carapidae (Markle and Olney, 1990), the anal fin begins at the level of the 11 th -18 th ray of the dorsal fin for P. lindas. In our juvenile, the number of DRAO is 6-7. ...
... This discrepancy is because the juvenile has not yet acquired the adult body proportions. Markle and Olney (1990) pointed out that the relative position of the fins can change ontogenetically. Therefore, the relative position of the fins cannot be used to identify larvae and juveniles in this case. ...
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An illustrated description of the juvenile (9.7 mm head length) of Pyramodon lindas from the Indian Ocean is given for the first time. The specimen is compared with the previously described vexillifer larvae of P. ventralis and P. punctatus. The juvenile of P. lindas is clearly distinguished from the other species of Pyramodon by the number of pectoral-fin rays and its pigmentation pattern. This is the first record of this species in the western Indian Ocean north of Madagascar.
... Even though there are some free-living species, most carapid fishes are found as commensal inquilines (i.e., with no metabolic dependency) or parasites of bivalve mollusks, sea slugs, sea hares, sea stars, sea cucumbers, and ascidians (Glynn, Enochs, McCosker, & Graefe, 2008;Parmentier, Lanterbecq, & Eeckhaut, 2016;Parmentier, Mercier, & Hamel, 2006). These remarkable associations, as well as a unique early life history (i.e, planktonic vexillifer larvae, whit a latter tenuis stage that usually requires a host for metamorphosis), account for the notoriety of the group (Markle & Olney, 1990;Parmentier & Vandewalle, 2005;Trott, 1981). ...
... Echinoderms were identified based on specialized taxonomic literature for the region (Borrero-Pérez & Vanegas- Massin, Zulfigar, Tan, & Rizzal, 2002;Solís-Marín, Arriaga-Ochoa, Laguarda-Figueras, Fontana-Uribe, & Durán-González, 2009;Solís-Marín et al., 2014;Martín-Cao-Romero, Solís-Marín, Laguarda-Figueras, & Buitrón-Sánchez, 2017;Woo, 2013;Woo, 2018;Woo, Zulfigar, Tan, Kajihara, & Fujita, 2015); the morphological data and terminology employed follow Solís-Marín et al. (2014), Woo (2013), and Woo et al. (2015). Fishes were identified following Nielsen, Cohen, Markle, and Robins (1999), and ; the morphological data and terminology employed follow Olney (1990), andNielsen et al. (1999). Morphometric measurements ( Table 2) from all fishes were taken with a digital caliper, after fixation in formalin 10 %, and preserved in ethanol 70 %. ...
... Morphometric measurements ( Table 2) from all fishes were taken with a digital caliper, after fixation in formalin 10 %, and preserved in ethanol 70 %. Counts were taken following Markle and Olney (1990), on three specimens that were cleared and stained following Taylor and Dyke (1985). Species valid S223 Revista de Biología Tropical, ISSN: 2215-2075: S219-S233, October 2021(Published Oct. 30, 2021 names, authorities, and year of description follows the WoRMS Editorial Board (2021), for invertebrates, and Fricke, Eschmeyer, and Fong (2021), for fishes. ...
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The North Pacific and the South Pacific of Nicaragua is a region of great biological, geological, economic and social wealth. There the dry forest mixes with the rain forest, the sea with the islands and nature with the people. It is a region influenced by the Papagayo upwelling, the emergence of cold marine waters during the dry season, which generates an abundance of life in the sea. As a sample of this marine wealth, in this Special Issue, more than half of the contributions are dedicated to advances in the knowledge of the marine biodiversity of the region. Contributions from the social sciences, geology and physics of the region are also included. Within these areas, the publications provide information on maritime border management, archaeology and sustainable tourism, coastal geology, projected climate changes, as well as various oceanographic aspects of the area. We hope that this Special Issue on the North Pacific of Costa Rica and the South Pacific of Nicaragua will promote more research in the region and help inform decision-making processes and educational activities. We thank the authors for their manuscripts, as well as the more than sixty reviewers who with their comments and suggestions helped to improve the quality of the manuscripts.
... Even though there are some free-living species, most carapid fishes are found as commensal inquilines (i.e., with no metabolic dependency) or parasites of bivalve mollusks, sea slugs, sea hares, sea stars, sea cucumbers, and ascidians (Glynn, Enochs, McCosker, & Graefe, 2008;Parmentier, Lanterbecq, & Eeckhaut, 2016;Parmentier, Mercier, & Hamel, 2006). These remarkable associations, as well as a unique early life history (i.e, planktonic vexillifer larvae, whit a latter tenuis stage that usually requires a host for metamorphosis), account for the notoriety of the group (Markle & Olney, 1990;Parmentier & Vandewalle, 2005;Trott, 1981). ...
... Echinoderms were identified based on specialized taxonomic literature for the region ( Woo et al. (2015). Fishes were identified following Nielsen, Cohen, Markle, and Robins (1999), and Robertson and Allen (2015); the morphological data and terminology employed follow Markle and Olney (1990), and Nielsen et al. (1999). Morphometric measurements ( Table 2) from all fishes were taken with a digital caliper, after fixation in formalin 10 %, and preserved in ethanol 70 %. ...
... Morphometric measurements ( Table 2) from all fishes were taken with a digital caliper, after fixation in formalin 10 %, and preserved in ethanol 70 %. Counts were taken following Markle and Olney (1990), on three specimens that were cleared and stained following Taylor and Dyke (1985). Species valid names, authorities, and year of description follows the WoRMS Editorial Board (2021), for invertebrates, and Fricke, Eschmeyer, and Fong (2021), for fishes. ...
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Introduction: The family Carapidae includes about 40 species of marine fishes distributed in coastal habitats worldwide. The family includes some free-living species, however, most of them are found as commensal inqui-lines or parasites of marine invertebrates, including several echinoderm species. In the Eastern Tropical Pacific, the biology and host use of the representatives of the Carapidae is relatively poorly known. Objective: The present study reports the occurrence of the Star pearlfish Carapus mourlani within three previously unknown hosts in the region: the sea stars Nidorellia armata, Phataria unifascialis, and the sea cucumber Stichopus horrens. Some ecological implications and considerations regarding such symbiotic relationships are raised and discussed. Additional morphometric and meristic data for the fish and the echinoderms are also provided and discussed. Methods: Echinoderms were collected, from 25 localities along the North Pacific coast of Costa Rica, and were carefully examined searching for commensal/parasitic fishes. Echinoderms and fishes were identified and characterized in accordance with the specialized literature. Results: A total of 497 echinoderms, including about 60 species, were collected and examined. Commensal/ parasitic fish (a single species represented by 13 specimens) were found in three echinoderm specimens/species. Conclusions: The list of echinoderm hosts for this carapid fish, through its whole distribution range, rises to 12 species (six sea stars and six sea cucumbers) and that could be a consequence of its wide geographic distribution, its generalist feeding habits and opportunistic commensal behavior.
... The adults of most species live symbiotically or hide inside various invertebrates. The genus Onuxodon Smith 1955 comprises four nominal species, three of them are recognized as valid: Onuxodon fowleri (Smith 1955), Onuxodon margaritiferae (Rendahl 1921) and Onuxodon parvibrachium (Fowler 1927); all distributed in the Indo-Pacific region (Markle and Olney 1990). Adults of this genus are characterized by presence of a rocker bone, predorsal bone, numerous small distal radials supporting the pectoral-fin rays, and one to several large symphysial fangs on the jaws. ...
... Counts and measurements followed Markle & Olney (1990). All measurements were made on the left side using digital calipers and rounded to the nearest 0.1 mm. ...
... nov. shares the following characters with other congeners which distinguish the genus Onuxodon from other carapids: lack of pelvic fins and ventral patch of posterior tunic ridges on the swim bladder and presence of a rocker bone, predorsal bone, and single large, symphysial fangs on each jaw. The new species resembles O. fowleri, which is widely distributed in the Indo-Pacific; both species have a slender body with higher vertebral counts than those two other congeners (Markle & Olney 1990). However, a comparison of O. albometeori with the data of lectotype of O. fowleri (BPBM 5001) given by Markle & Olney (1990) indicated clear differences in the following characters: precaudal vertebrae 23 in O. albometeori (vs. ...
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Onuxodon albometeori sp. nov. (Ophidiiformes: Carapidae) is described from a single specimen collected by commercial trawl off southwestern Taiwan. The new species is most similar to the Indo-Pacific species Onuxodon fowleri (Smith 1955), both process a remarkably slender body, and higher precaudal vertebral counts and a longer pectoral fin, although the two latter features are even more extreme in the former. Onuxodon albometeori sp. nov. is further distinguished from O. fowleri by its lesser body depth, greater head width, higher counts of precaudal vertebrae, and uniformly whitish coloration only on the posterior part of the body.
... commonly known as "pearlfish" (Markle and Olney 1990), is among the species that infest the sea cucumbers. They can live in association with several species of Holothuriida (Holothuria tubulosa, H. poli, H. helleri, H. sanctori and H. stellati); however, it has a preference for the species of the order Synallactida (Parastichopus regalis) (Meyer-Rochow 1977;Mezali 1998;Eeckhaut et al. 2004;Parmentier et al. 2006). ...
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Commensalism between the pearlfish Carapus acus and the holothurian Parastichopus regalis is one of the most common associations between vertebrate and an invertebrate in the Mediterranean Sea. Seven individuals of Carapus acus were found inside the Parastichopus regalis coelomic cavity at three stations off the western Algerian coast with a depth of between 53 and 117 m. The wet weight of these individuals varied from 1.24 to 4.11 g and the total length from 12.30 to 16.60 cm. These values and other morphometric characteristics of C. acus individuals were compared with those reported on the Tunisian and Moroccan coasts.
... Remarks: A separate set corresponding to the anteroventral section of the epaxialis that connects the swim bladder to the first three vertebrae and ribs has been described and illustrated in Ophidiiformes (e.g. Courtenay & McKittrick, 1970;Tyler, 1970;Markle & Olney, 1990;Howes, 1992: figs 17, 18, 20-26;Parmentier & Diogo, 2006) and shown to be involved in a sexually dimorphic sound-production complex. Nevertheless, the distribution of this specialization has never been surveyed comprehensively across the order. ...
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More than half the ray-finned fishes and about one-quarter of all living vertebrates belong to Percomorphacea. Among its 30 orders, Stromateiformes encompass 77 species in 16 genera and six families. Stromateiform monophyly has never been tested using morphology, and it has been rejected by molecular analyses. This comprehensive revision of Stromateiformes includes all its valid genera of all percomorph families previously aligned with the order. We sampled 207 phenotypic characters in 66 terminal taxa representing 14 orders and 46 acanthopterygian families. This dataset significantly surpasses all previous phenotype-based phylogenies of Stromateiformes, which analysed only a fraction of these characters. Stromateiformes is recovered as monophyletic, supported by eight unequivocal synapomorphies. Amarsipidae is the sister group of all other Stromateiformes (= Stromateoidei). Centrolophidae is paraphyletic, with three of its genera allocated into an early-diverging clade and the other four appearing as successive sister groups to a lineage containing the remaining stromateiforms. All other stromateoid families are monophyletic, with the following cladistic arrangement: (Nomeidae (Stromateidae (Tetragonuridae, Ariommatidae))). Our analysis convincingly refutes recent molecular phylogenetic interpretations that fail to recover a monophyletic Stromateiformes. These findings call into question large-scale conclusions of percomorph relationships and trait evolution based solely on molecular data.
... are known to feed off their host's gonad (Murdy and Cowan 1980;Parmentier et al. 2003;Pamentier and Vandewalle 2005;Parmentier et al. 2016 (Smith and Tyler 1969;Trott 1970;Tyler et al. 1992;Hasbun and Lawrence 2002)distribution, size, structure, nutritional quality and feasibility of exploitation of a sea cucumber population, an alternative fishery resource for Banco Chinchorro (Mexican Caribbean, Actinopyga agassizi (Arnold 1956;Van Meter and Ache 1974;Hasbun and Lawrence 2002), Isostichopus badionotus (Smith and Tyler 1969;Vergara et al. 2016); Eostichopus arnesoni, Holothuria lentiginosa (Miller and Pawson 1979;Valentine and Goeke, 1983) distribution, size, structure, nutritional quality and feasibility of exploitation of a sea cucumber population, an alternative fishery resource for Banco Chinchorro (Mexican Caribbean, Selenkothuria glaberrima, Theelothuria princeps and Thyone sp. (Phyllophoridae, Dendrochirotida) (Smith et al. 1981;Trott 1970;Markle and Olney 1990;Tyler et al. 1992), Astichopus multifidus (Trott 1970) distribution, size, structure, nutritional quality and feasibility of exploitation of a sea cucumber population, an alternative fishery resource for Banco Chinchorro (Mexican Caribbean; Holothuria glaberrima (Trott 1970) distribution, size, structure, nutritional quality and feasibility of exploitation of a sea cucumber population, an alternative fishery resource for Banco Chinchorro (Mexican Caribbean and Holothuria princeps (Dawson 1971) distribution, size, structure, nutritional quality and feasibility of exploitation of a sea cucumber population, an alternative fishery resource for Banco Chinchorro (Mexican Caribbean. ...
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This study investigates the sounds and the anatomy of the sound‐producing organ in the male and female sand‐dwelling cusk‐eel Parophidion vassali. Although both sexes have similar external phenotype, they can be distinguished by their sonic apparatus and sounds. As in many Ophioidei, Parophidion vassali presents a panel of highly derived characters. Fish possess three pairs of sonic muscles, and males have mineralized swimbladder caps on which inserts the ventral sonic muscle, a neural arch that pivots, a stretchable swimbladder fenestra, an osseous swimbladder plate and a rounded pressure‐release membrane in the caudal swimbladder. Females, however, do not possess anterior swimbladder caps, a swimbladder fenestra and the caudal rounded membrane. Males possess the unusual ability to produce sounds starting with a set of low amplitude pulses followed by a second set with higher amplitudes clearly dividing each sound unit into two parts. Females do not vary their sound amplitude in this way: they produce shorter sounds and pulse periods but with a higher peak frequency. Morphology and sound features support the sound‐producing mechanism is based on a rebound system (i.e. quick backward snap of the anterior swimbladder). Based on features of the sounds from tank recordings, we have putatively identified the sound of male Parophidion vassali at sea. As these species are ecologically cryptic, we hope this work will allow assessment and clarify the distribution of their populations. This study investigates the sounds and the anatomy of the sound‐producing organ in the dwelling cusk‐eel. Males produce sounds starting with a set of low amplitude pulses followed by a second set with higher amplitudes clearly dividing each sound unit into two parts. Females produce shorter sounds and pulse periods but with a higher peak frequency. Differences in sounds correspond to differences in sound‐producing mechanisms.
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