image/svg+xml101New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023Neotropical HelminthologyNeotropical Helminthology, 2023, vol. 17 (2), 101-116ORIGINAL ARTICLE / ARTÍCULO ORIGINALANEW SPECIES OF UROCLEIDOIDES(MONOGENOIDEA: DACTYLOGYRIDAE) PARASITE OF PSECTROGASTER AMAZONICA(CHARACIFORMES: CURIMATIDAE) AND CAENOTROPUS LABYRINTHICUS(CHARACIFORMES: CHILODONTIDAE) FROM TOCANTINS RIVER, MARANHÃO STATE, BRAZIL WITH CHECKLIST OF THE GENUSUNA NUEVA ESPECIE DE PARÁSITO UROCLEIDOIDES (MONOGENOIDEA: DACTYLOGYRIDAE) DE PSECTROGASTER AMAZONICA(CHARACIFORMES: CURIMATIDAE) Y CAENOTROPUS LABYRINTHICUS(CHARACIFORMES: CHILODONTIDAE) DEL RÍO TOCANTINS, ESTADO DE MARANHÃO, BRASIL Y UNA LISTA DE VERIFICACIÓN DEL GÉNEROCarine Almeida Miranda Bezerra1, Álvaro José Bittencourt de Freitas2, Diego Carvalho Viana3,Simone Chinicz Cohen2*& Marcia Cristina Nascimento Justo2ISSN Versión Impresa 2218-6425 ISSN Versión Electrónica 1995-1403DOI: https://dx.doi.org/10.24039/rnh20231721591Volume 17, Number 2 (jul - dec) 2023Este artículo es publicado por la revista Neotropical Helminthology de la Facultad de Ciencias Naturales y Matemática, Universidad Nacional Federico Villarreal, Lima, Perú auspiciado por la Asociación Peruana de Helmintología e Invertebrados Af nes (APHIA). Este es un artículo de acceso abierto, distribuido bajo los términos de la licencia Creative Commons Atribución 4.0 Internacional (CC BY 4.0) [https:// creativecommons.org/licenses/by/4.0/deed.es] que permite el uso, distribución y reproducción en cualquier medio, siempre que la obra original sea debidamente citada de su fuente original.ABSTRACTParasites are important components of global biodiversity and among them, the Monogenoidea stands out as a parasites of f shes, commonly af ecting the gills of their hosts. T e present study aimed to analyze the presence of species of Urocleidoidesparasitizing characiform f shes from the Tocantins River. During the study, specimens of Psectrogaster amazonicaEigenmann & Eigenmann and Caenotropus labyrinthicus(Kner) were collected from the urban zone of the municipality of Imperatriz, state of Maranhão. T e gills of the f sh were removed and placed in vials containing hot 1 Programa de Pós-Graduação em Ciência Animal, Universidade Estadual do Maranhão (UEMA). Cidade Universitária Paulo VI, Avenida Lourenço Vieira da Silva 1000, 65055-310 São Luís, MA, Brazil.2 Laboratório de Helmintos Parasitos de Peixes, Instituto Oswaldo Cruz (IOC), FIOCRUZ. Avenida Brasil 4365, 21040-900 Rio de Janeiro, RJ, Brazil.3 Núcleo de Estudos Morfof siológicos Avançados (NEMO), Universidade Estadual da Região Tocantina do Maranhão (UEMASUL). Campus Imperatriz, Rua Godofredo Viana 1300, 65901-480 Imperatriz, MA, Brazil. * Corresponding author: scohen@ioc.f ocruz.brCarine A. M. Bezerra:https://orcid.org/0000-0003-4842-1595 Álvaro José B. de Freitas:https://orcid.org/0000-0003-4338-7893Diego C. Viana:https://orcid.org/0000-0002-3302-9892 Simone C. Cohen:https://orcid.org/0000-0001-8204-336X Marcia C. N. Justo:https://orcid.org/0000-0002-0684-3389
image/svg+xml102Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.water (65°C), and posteriorly, absolute ethanol was added to reach a concentration of 70%. A new species of Urocleidoideswas described parasitizing both hosts. Urocleidoides psectrogasteri n. sp. can be distinguished by its congeners species principally by the morphology of the MCO. Besides, the known species U. tocantinensis also was found parasitizing C. labyrinthicus, which was reported as a new host for this species. In addition, a checklist containing all species of Urocleidoidesis presented to compile the known data on this monogenoid genus.Keywords: Fish parasites –Monogenoidean – Neotropical Region – Tocantins-Araguaia Basin – Urocleidoidespsectrogasterin. sp.RESUMENLos parásitos son componentes importantes de la biodiversidad global y entre ellos, se destacan los Monogenoidea como parásitos de los peces, afectando comúnmente las branquias de sus hospederos. El objetivo del presente estudio fue analizar la presencia de especies de Urocleidoidesque parasitan peces Characiformes del río Tocantins. Durante el estudio, se recolectaron ejemplares de Psectrogaster amazonica Eigenmann & Eigenmann y Caenotropus labyrinthicus(Kner) de la zona urbana del municipio de Imperatriz, estado de Maranhão, Brasil. Se extrajeron las branquias de los peces y se colocaron en viales con agua caliente (65°C), y posteriormente se añadió etanol absoluto hasta alcanzar una concentración del 70%. Se describió una nueva especie de Urocleidoidesque parasita a ambos hospedadores. Urocleidoidespsectrogasteri n. sp. se puede distinguir por sus especies congéneres principalmente por la morfología de la MCO. Además, la especie conocida U. tocantinensis, también fue encontrada parasitando a C. labyrinthicus, la cual fue reportada como un nuevo hospedero para esta especie. Además, se presenta una lista de verifcación que contiene todas las especies de Urocleidoides para recopilar los datos conocidos sobre este género monogenoideo.Palabras clave:Bacia del Tocantins-Araguaia –Monogenoideos – Parásitos de peces – Region Neotropical – Urocleidoidespsectrogasterin. sp.INTRODUCTIONUrocleidoides Mizelle & Price, 1964 was proposed to accommodate the species U. reticulatusMizelle & Price, 1964 from the gills of Poecilia reticulataPeters, 1859 (Poeciliidae), which was collected from Capitol Aquarium, Sacramento, California, USA (Mizelle & Price, 1964). Kritsky & Tatcher (1983) listed 30 species of the genus described from four diferent orders of hosts: Characiformes, Cyprinodontiformes, Gymnotiformes e Siluriformes (Mizelle et al., 1968; Mizelle & Kritsky, 1969; Molnar et al.,1974; Kritsky & Tatcher 1974, 1976, 1983). Kritsky et al. (1986) revised the genus, described two new species, U. eremitusKritsky, Tatcher & Boeger, 1986 and U. paradoxus Kritsky, Tatcher & Boeger, 1986 and added new morphological details in species described previously: U. reticulatus, U. anops Kritsky & Tatcher 1974andU. curimatae Molnar, Hanek & Fernando, 1974. Te authors emended the diagnosis of the genus, characterizing the species by the presence of vaginal sclerite, male copulatory organ with counterclockwise ring, tandem gonads, no modifcated anchors, hook with dilated shank and pairs 1 and 5 of hooks usually reduced in size. Kritsky et al.(1986) considered 22 species as incertae sedis, mainly by the absence of the vaginal sclerite.Posteriorly, Ferreira et al. (2017) added U. advenaiMendoza-Franco & Reina, 2008 to this status of incertae sedis.Further studies reallocate some of these species on newly described genera as Ameloblastella Kritsky, Mendoza-Franco & Scholz, 2000,Aphanoblastella Kritsky, Mendoza-Franco & Scholz, 2000,CharacitheciumMendoza-Franco, Reina & Torchin, 2009,Diaphorocleidus Jogunoori, Kritsky & Venkatanarasaiah, 2004, Nanayella Acosta, Mendoza-Palmero, Silva & Scholz, 2019,Philocorydoras Suriano, 1986 and Sciadicleithrum Kritsky, Boeger & Tatcher, 1989 (Price & Bussing, 1968; Kritskyet al., 1989, 2000; Jogunoori et al., 2004; Mendoza-Franco et al., 2009; Yamada et al., 2015; Acosta et al., 2019).Recently, Santos-Neto & Domingues (2023) proposed four new species of Urocleidoidesfrom Characiformes and Gymnotiformes and revalidate the incertae sedisspecies, Urocleidoides gymnotus Mizelle, Kritsky & Crane, 1968 and Urocleidoides carapus Mizelle, Kritsky & Crane, 1968, based in molecular sequences. One of the new species, Urocleidoides vaniniSantos-Neto & Domingues, 2023 did not present vaginal sclerite as the incertae sedisrevalidated by these authors. From the species previously
image/svg+xml103New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023assigned as incertae sedis by Kritsky et al.(1986), six have not yet been relocated to other genera.Species of Urocleidoides have been described from Psectrogaster amazonica Eigenmann & Eigenmann and Caenotropus labyrinthicus (Kner), as U. tocantinensisFreitas, Bezerra, Meneses, Justo, Viana & Cohen, 2021and U. paratriangulusFreitas, Bezerra, Meneses, Viana, Justo & Cohen 2021 and the latter also from C. labyrinthicus (Freitas et al., 2021).In this present study, a new species of Urocleidoides was found parasitizing P. amazonicaand C. labyrinthicus and is described herein. In addition, C. labyrinthicus is reported as new host for U. tocantinensisand a checklist of Urocleidoides species with all current information about their hosts, localities and validity is presented. MATERIAL AND METHODSFishes were collected in Tocantins River at the urban zone of Imperatriz municipality, state of Maranhão,129 specimens of P. amazonica(standard length 11.5–16.1 cm; total weight 42–100 g)and 20 specimens C. labyrinthicus(standard length 12.1–14.5 cm; total weight 38–70 g). Te fshes were caught with the aid of local fshermen and were carried to the “Núcleo de Estudos Morfológicos Avançados (NEMO)” at the Universidade Estadual da Região Tocantina do Maranhão (UEMASUL), Maranhão State, Brazil, where they were identifed. Te gills of fsh were removed and placed in vials containing hot water (65°C), and vigorously shaken to detach the parasites from the gill flaments. Posteriorly, absolute ethanol was added to reach a concentration of 70%. Tose vials containing the gills were then sent to the “Laboratório de Helmintos Parasitos de Peixes, Instituto Oswaldo Cruz, FIOCRUZ”, Rio de Janeiro state, Brazil. Te gills and sediments were analyzed in Petri dishes. Monogenoids were collected under a stereoscopic microscope, and the specimens mounted in Hoyer’s medium to study the sclerotized structures (Humason, 1979). Parasitological indexes were calculated as proposed by Bush et al. (1997), followed by standard deviation. Measurements are presented in micrometers; range values are followed by mean and number of structures measured in parentheses. Morphometric studies were performed through ImageJ software (Wayne, 2010), distributed by the National Institutes of Health (NIH), and available at https://imagej.nih.gov/ij/download.html. Te authorship of the taxa followed the recommendation of Article 50.1 of the International Code of Zoological Nomenclature (ICZN), which deals with the identity of the authors. A checklist of valid Urocleidoides species is presented herein, providing information about hosts and geographical distribution. For construction of this checklist, published papers with original descriptions and/or reports of species, taxonomic revisions, and checklists were used and online biodiversity databases as Taxonomic Catalog of Brazilian Fauna, PubMed, Scopus, Google Scholar and CAB Abstracts. Posgraduate theses and abstract scientifc meetings do not constitute formal publications and were thus not considered. For Brazilian states, abbreviations are presented as follows: AC: Acre; AL: Alagoas; AM: Amazonas; AP: Amapá; BA: Bahia; CE: Ceará; DF: Distrito Federal;ES: Espírito Santo; GO: Goiás; MA: Maranhão;MG: Minas Gerais; MS: Mato Grosso do Sul; MT: Mato Grosso; PA: Pará; PB: Paraíba; PE: Pernambuco; PI: Piauí; PR: Paraná; RO: Rondônia; RJ: Rio de Janeiro; RN: Rio Grande do Norte; RR: Roraima; RS: Rio Grande do Sul; SC: Santa Catarina; SE: Sergipe; SP: São Paulo; TO: Tocantins.Ethic aspects: Te fsh sample collection protocol and laboratory procedures were approved by the Research Ethics Committee of “Universidade Estadual do Maranhão” (UFMA), Maranhão State, Brazil under protocol number 43/2021, and the environmental collection license was obtained from the System for Authorization and Information on Biodiversity (SISBIO), under protocol number 79538-1. RESULTSTAXONOMY Class Monogenoidea Bychowsky, 1937 Subclass Polyonchoinea Bychowsky, 1937 Order Dactylogyridea Bychowsky, 1937 Dactylogyridae Bychowsky, 1933UrocleidoidesMizelle & Price, 1964Urocleidoides psectrogasterisp. nov Bezerra, Freitas, Cohen & Justo(Fig. 1) Type host.Psectrogaster amazonicaEigenmann & Eigenmann (Curimatidae).
image/svg+xml104Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.Type locality.Tocantins River (5°27’50’S; 47°33’48’W), at the urban zone of Imperatriz municipality.Other hosts: Caenotropus labyrinthicus(Kner) (Chilodontidae)Infestation parameters.P. amazonica: Total number of hosts: 129; prevalence: 17%; total number of parasites: 25; mean intensity: 1.14 (±0.34); mean abundance: 0.19 (±0.45); range of intensity: 1–2. C. labyrinthicus: Total number of hosts: 20; prevalence: 5%; intensity: 3 (only one host was parasitized); mean abundance: 0.15 (±0.67).Deposited material.Holotype CHIOC 40210. Paratypes CHIOC 40211, 40212, 40213, 40214, 40215, 40216, 40217 a-c. Etymology: Te specifc name refers to the scientifc name of the host Psectrogaster amazonicaFigure 1. Urocleidoides psectrogasterisp. nov from Psectrogaster amazonica(Curimatidae) from Tocantins River, at the urban zone of Imperatriz municipality. A. Total, ventral view (composite). B. Copulatory complex; C. Vaginal sclerite; D. Dorsal bar; E. Ventral Bar; F. Ventral anchor; G. Dorsal anchor; H. Hook. Description based on 23 specimens mounted in Hoyer’s medium: Body fusiform, 250–564 (384, n= 23) long by 97–201 (139, n= 23) wide. Tree pairs of cephalic glands. Two pairs of eyespots, equidistant, anterior pair smaller than posterior. Pharynx 14–22 (18, n=5) ovate and muscular. Peduncle short. Haptor sub-hexagonal 75–184 (117, n= 22) wide. Anchors robust, dissimilar in shape. Ventral anchor with well-developed roots, protruding superfcial root, semi-circular shape, rounded deep root, evenly curved shaft and short point, 24–53 (39, n=46) long and base 15–22 (18, n=46); Dorsal anchor with rectangular-shaped superfcial root and deep root distally round, evenly curved shaft and acute point 23–34 (29, n= 44) long and base 12–19 (16, n= 44). Ventral bar robust, slightly curved, with enlarged extremities, concave posteromedial 27–77 (46, n=9) and drop-like shaped posteromedian projection, short and thin 5–14 (8, n= 21) long. Dorsal bar slender V-shaped, slightly curved, 37–70 (54, n= 23) long. Seven pairs of marginal hooks, with distribution of ancyrocephaline, fve pairs ventral and two dorsal. Hooks similar in shape, divided in two subunits, protruded thumb, curved point, shank straight with proximal dilation, comprising ⅔ of shank length; FH loop ¼ of shank length; pairs 1 and 5 slightly smaller than others, pair 1 12–27 (19, n = 34); pair 2 19–30 (25, n= 35); pair 3 19–34 (26, n= 33); pair 4 19–39 (28, n= 33); pair 5 14–28 (21, n= 28); pair 6 19–35 (27, n= 32); pair 7 19–34 (27, n= 34) long. Copulatory complex composed of male copulatory organ (MCO) with auriculate base, sickle shape, 28–44
image/svg+xml105New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023(36, n=18) and accessory piece bifurcated, with two elongated parts, articulated to MCO base, 27–41 (33, n=18) long. Testis dorsal to germarium; seminal vesicle a distal dilation of vas deferens; single prostatic reservoir present. Germarium elongated. Vaginal pore sinistral; vagina comprising a vaginal vestibule cup-shaped, sclerotized and a vaginal canal long; seminal receptacle round. Vaginal sclerite present in the middle of the body, with a longitudinal groove in the middle, erected thumb, 18–44 (29, n= 18) long. Eggs not observed. Vitellaria present, heavily distributed throughout the body, except in reproductive organs. Remarks: Te new species is allocated in Urocleidoides by the presence of vaginal sclerite, vagina in sinistral position, and hook pairs 1 and 5 reduced in size. Te MCO of the new species is similar to those of some species by the absence of coils on the MCO, as observed in U. surianoae Rossin & Timi, 2016, U. falxus Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva, 2020and U. tocantinensis. Te new species is evidently distinguished from these species by the morphology of the male copulatory organ, which is pincer-shaped (reverse J-shaped, with expanded proximal end in U. surianoae, male copulatory organ appearing to twist into itself and resembling the shape of a sickle and base enlarged in U. falxusand straight tube with enlarged base in U. tocantinensis). Te new species also resembles U. tocantinensisconsidering the bifurcation of the accessory piece and morphology of vagina. Considering the projection of the ventral bar, the new species is similar to Urocleidoides boullengerelae Freitas, Bezerra, Meneses, Justo, Viana & Cohen, 2021, Urocleidoides piriatiu Mendoza-Franco & Reina, 2008 and U. tocantinensis, difering that in Urocleidoides psectrogasteri sp. nov. the posteromediam projection is reduced in size. Despite the projection in Urocleidoides surianoae Rossim & Timi, 2016is similar in shape and size, it difers from U. psectrogasteri sp. nov. by the anteromedial position.Checklist of Urocleidoides spp. from Neotropical RegionUrocleidoides spp. have been recorded from 3 diferent orders (34 in Characiformes, eight in Gymnotiformes and four in Cyprinodontiformes), associated with 14 families and 48 species of fshes (Fig. 2). Most of them were reported from fshes belonging to Characiformes, from eight diferent families with predominance of Erythrinidae, mainly from Hoplias malabaricus (Bloch). Regarding the geographical distribution, most species are reported from Brazil, followed by Mexico and Panama, and records in Argentina, Colômbia, El Salvador, Honduras and Trindad and Tobago, thus presenting a broad host and locality spectrum. Until now, thirty-one species of Urocleidoides are referred from Brazil in diferent localities, rivers and basins within the country.Figure 2. Distribution of Urocleidoidesspp. within host families of three orders of freshwater fshes. Blue: Characiformes, Orange: Cyprinodontiformes, Grey: Gymnotiformes.
image/svg+xml106Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.Urocleidoides aimarai Moreira, Scholz & Luque, 2015Type-host and locality: Hoplias aimara (Valenciennes) (Characiformes: Erythrinidae) [Xingu River, around Altamira, PA,Brazil].Other hosts and locality: Hoplias malabaricus(Characiformes: Erythrinidae);Leporinus friderici (Bloch) (Characiformes, Anostomidae) [Batalha River, SP, Brazil].Records: Moreira et al.(2015); Dias et al.(2017); Gião et al.(2020).Urocleidoides anops Kritsky & Tatcher, 1974Type-host and locality: Characidium caucanumEigenmann (Characiformes: Crenuchidae) [Pance River, Cali, Valle, Colombia].Other host and localiy: Psalidodon fasciatus (Cuvier) (Characiformes: Characidae) [Noc-choncunchey and Dzonot Cervera cenotes, Mexico].Record: Kritsky & Tatcher (1974); Mendoza-Franco et al. (1999).Urocleidoides atilaiamarinoiSantos-Neto & Domingues, 2023Type host and locality: Hoplerythrinus unitaeniatus(Agassiz) (Characiformes: Erythinidae) [Igarapé Maratininga (Moju River Basin), municipality of Tailândia, PA, Brazil].Other locality: Balneário Aracu (Guamá River Basin), municipality of Ourém, Pará, Brazil. Record: Santos-Neto & Domingues (2023).Urocleidoides boulengerellaeFreitas, Bezerra, Meneses, Justo, Viana & Cohen, 2021Type-host and locality: Boulengerella cuvieri(Spix & Agassiz) (Characiformes: Ctenoluciidae) [Tocantins River, near the municipality of Tupiratins, TO, Brazil].Other localities: Arraias River, close to the municipality of Babaçulândia; Tocantins River, close to the municipalities of Aguiarnópolis and Estreito; at the mouth of the Itaueiras River, in the municipality of Estreito, MA, Brazil.Record: Freitas et al.(2021).Urocleidoides brasiliensis Rosim, Mendoza-Franco & Luque, 2011Type-host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Cuiabá River, MT,Brazil].Otherlocalities: Cristalino River, MT; Caeté River, Bragança; Maracanã River, Nova Timboteua; Piriá and Guripi Rivers,Viseu; Maparanim River, Municipality of Terra Alta; Itabocal River, Irituia, PA; Upper Paraná River Floodplain, PR, MS; Paraná River, PR; Guandú River, RJ, Brazil.Records: Rosim et al.(2011); Graça et al. (2013); Ferreira et al.(2017).Urocleidoides bulbophallus Ferreira, Rodrigues, Cunha & Domingues, 2017Type-host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Caeté River, Bragança, PA, Brazil].Other localities: Itabocal River, Irituia; Piriá River, Viseu; Maparanim River, Terra Alta; Gurupi River, Viseu, PA, Brazil.Records: Ferreira et al.(2017).Urocleidoides carapusMizelle, Kritsky & Crane, 1968Type-host and locality: Gymnotus carapo(Linnaeus) (Gymnotiformes, Gymnotidae) [Amazon Basin, Brazil].Other host and locality: Sternopygus macrurus(Bloch & Schneider) (Gymnotiformes, Gymnotidae) [Guamá River, Ourém, PA, Brazil].Records: Mizelle et al.(1968); Santos-Neto & Domingues (2023).Urocleidoides cuiabai Rosim, Mendoza-Franco & Luque, 2011Type host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Cuiabá River, MT, Brazil].Other localities: Maracanã River, Nova Timboteua; Gurupi and Piriá Rivers, Viseu, PA; Araguaia River, MT; Jaguari-Mirim River, MG, SP; Upper Paraná River Floodplain, PR, MS; Paraná River, PR; Guandu River, RJ; Batalha River, SP, Brazil. Records: Rosim et al.(2011); Graça et al.(2013); Gasques et al.(2016); Ferreira et al.(2017); Dias et al.(2017); Gião et al.(2020).Urocleidoides cultellus Mendoza-Franco & Reina, 2008Type-host and locality:Brachyhypopomus occidentalis(Regan) (Gymnotiformes: Hypopomidae) [Águas Claras River at its confuence with the Bayano Lake, Republic of Panama].
image/svg+xml107New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023Record: Mendoza-Franco & Reina (2008).Urocleidoides curimatae Molnár, Hanek & Fernando, 1974Type-host and locality:Steindachnerina argentea(Gill) (= Curimata argentea) (Characiformes: Curimatidae) [Arouca River near D’Abadie] after Trinidad.Record: Molnar et al.(1974).Urocleidoides digitabulum Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva, 2020 Type-host and locality: Leporinus friderici (Characiformes: Anostomidae) [Sapucaí-Mirim River, Upper Parana River Basin, SP, Brazil].Other hosts and localities: Leporinus octofasciatusSteindachner; Megaleporinus elongatus (Valenciennes) (Characiformes: Anostomidae)[Jurumirim Reservoir, Paranapanema River, Upper Paraná River Basin, SP, Brazil].Record: Zago et al.(2020).Urocleidoides eremitus Kritsky, Tatcher & Boeger, 1986Type-host and locality: Hoplias malabaricus(Characiformes: Erythrinidae) [Janauaca Lake near Manaus, AM, Brazil].Other hosts and localities: Leporinus friderici; Megaleporinus macrocephalus(Garavello & Britski) (Characiformes: Anostomidae) [Chascomus Lake, Buenos Aires province,Argentina; fsh farms inCruzeiro do Sul and Rio Branco, AC; Igarapé Fortaleza basin, in the municipality of Macapá, AP; Jaguari Mirim River, MG and SP; Machado River, MG; Cuiabá River, MT; Maracanã River, Nova Timboteua; Piriá River, Viseu; Caeté River, Bragança, PA; Upper Paraná River Floodplain in the PR, MS; Guandu River, RJ; Batalha River, Peixe River; Pirassununga lagoons, SP, Brazil; Tambopata River, Madre de Dios, Perú].Records: Kritsky et al. (1986); Iannacone & Luque (1993); Suriano (1997a); Rosim et al.(2011); Corrêa et al. (2013), Alcântara & Tavares-Dias (2015); Dias et al.(2017); Ferreira et al.(2017); Martins et al.(2017 a, b); Gião et al.(2020); Negreiros et al.(2021).Urocleidoides falxus Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva, 2020Type-host and locality:Megaleporinus elongatus(Characiformes: Anostomidae) [Sapucaí-Mirim River, Upper Paraná River basin, SP, Brazil].Other locality: Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin, SP, Brazil.Record: Zago et al.(2020).Urocleidoides fegomai Mendoza-Franco, Aguirre-Macedo & Vidal-Martínez, 2007Type-host and locality: Piabucina panamensisGill (Characiformes: Lebiasinidae) [Frijolito River, Panama].Record: Mendoza-Francoet al. (2007).Urocleidoides gymnotusMizelle, Kritsky & Crane, 1968Type-host and locality: Gymnotus carapo(Gymnoti-formes, Gymnotidae) [Amazon Basin, Brazil].Other host and locality: Sternopygus macrurus(Gymnotiformes, Gymnotidae) [Guamá River, Ourém, PA, Brazil].Records: Mizelle et al.(1968); Santos-Neto & Domingues (2023).Urocleidoides hypopomi Suriano, 1997Type-host and locality: Brachyhypopomus brevirostris (Steindachner) (Gymnotiformes: Hypopomidae) [Paraná River, Puerto Italia, Corrientes Province, Argentina].Record: Suriano (1997a).Urocleidoides indianensis Oliveira, Silva, Vieira & Acosta, 2021Type-host and locality: Parodon nasusKner (Characiformes: Parodontidae) [Indiana stream (Capivara River, Tietê River, Upper Paraná River basin), Botucatu, SP, Brazil].Record: Oliveira et al.(2021).Urocleidoides jariensis Oliveira, Santos-Neto, Tavares-Dias & Domingues, 2020Type-host and locality:Schizodon fasciatusSpix & Agassiz (Characiformes: Anostomidae) [Lower Jari River, near Jarilândia district, in municipality of Vitória do Jari, AP, Brazil].Record: Oliveira et al. (2020).Urocleidoides macrosomaSantos-Neto & Domingues, 2023Type host and locality: Hoplias malabaricus(Characiformes: Erythrinidae) [Vila Segredo – Segredo
image/svg+xml108Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.River (Quatipuru River Basin), Tauari, municipality of Capanema, PA, Brazil].Record: Santos-Neto & Domingues (2023).Urocleidoides malabaricusi Rosim, Mendoza-Franco & Luque, 2011Type-host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Cuiabá River, MT, Brazil].Other localities: Caeté River, Bragança; Gurupi River, Viseu; Maracanã River, Nova Timboteua; Maparanim River, Terra Alta; Piriá River, Viseu; Itabocal River, Irituia, PA; Upper Paraná River Floodplain PR, MS; Batalha River, SP; Paraná River, PR, Brazil.Records: Rosim et al.(2011); Graça et al.(2013); Gasques et al. (2016); Ferreira et al.(2017); Dias et al.(2017); Gião et al.(2020).Urocleidoides naris Rosim, Mendoza-Franco & Luque, 2011Type-host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Cuiabá River, MT, Brazil].Other localities: Guandu River, RJ; Jaguari-Mirim River, MG and SP; Machado River, MG, Brazil.Record: Rosim et al. (2011).Urocleidoides nataliapasternakaeSantos-Neto & Domingues, 2023Type host and locality: Brachyhypopomus brevirostris(Steindachner) (Gymnotiformes: Hypopomidae) [Balneário Aracu (Guamá River Basin), municipality of Ourém, PA, Brazil]. Record: Santos-Neto & Domingues (2023).Urocleidoides neotropicalis Mendoza -Franco & Reina, 2008Type-host and locality:Saccodon dariensis Meek & Hildebrand (Characiformes: Parodontidae) [Piriati River at its confuence with the Chagres River Basin, Republic of Panama].Record: Mendoza-Franco & Reina (2008).Urocleidoides paradoxus Kritsky, Tatcher & Boeger, 1986Type-host and locaÕlity:Rhytiodus microlepisKner (Characiformes, Anostomidae) [SolimÕes River near Ilha Marchantaria, Manaus, AM, Brazil].Other hosts and localities: Leporinus friderici; Leporinus lacustrisAmaral Campos; Megaleporinus elongatus; Megaleporinus macrocephalus, Megaleporinus obtusidens(Valenciennes) (Characiformes, Anostomidae) [Parana River, Puerto Italia, Corrientes province, Argentina; fsh farms in Cruzeiro do Sul and Rio Branco, AC; Furo do Catalão, near Encontro das Aguas, Manaus; Coari Manaus section, Lake Baixio; Lake Ananá, Lake Preto and Lake Catalão, Iranduba, AM; lower Jari River, near Jarilândia district, in municipality of Vitória do Jari, AP; Upper Paraná River foodplain, PR; Lake Guaiba, RS; Batalha River, SP; Brazil].Records: Kritsky et al.(1986); Suriano (1997a); Guidelli et al.(2006, 2011); Takemoto et al.(2009); Souza et al.(2017); Martins et al.(2017a,b); Wendt et al.(2018); Oliveira et al.(2020); Negreiros et al.(2021). Urocleidoides paranae Ferreira, Rodrigues, Cunha & Domingues, 2017Type-host and locality:Hoplias malabaricus(Characiformes: Erythrinidae) [Upper Paraná River foodplain (Paraná River Basin; Paraná, Paranapanema Sub-basin), PR and MS, Brazil].Records: Graça et al.(2013); Ferreira et al.(2017).Remarks: Te authors examined vouchers of U. eremituscollected from H. malabaricus from the Upper Paraná River foodplain of Graça et al.(2013) and indicated that these specimens are members of U. paranae. Urocleidoides paratriangulusFreitas, Bezerra, Meneses, Justo, Viana & Cohen, 2021Type hostand locality: Psectrogaster amazonica (Characiformes: Curimatidae) [Tocantins River, close to the municipality of Embiral, MA, Brazil].Other hosts and localities:Cyphocharax gouldingi Vari (Characiformes: Curimatidae); Caenotropus labyrinthicus (Chraciformes: Chilodontidae);Mylesinus pauscisquamatus Jégu & Santos (Characiformes: Serrasalmidae) [Farinha River, close to the municipality of Estreito, MA; Tocantins River, close to the urban perimeter of Imperatriz, MA; Tocantins River, close to the municipalities of Aguiarnópolis and Estreito, MA; at the mouth of the Itaueiras River, in the municipality of Estreito, MA; Tocantins River, close to the municipalities of Aguiarnópolis and Estreito, MA; at the mouth of the Itaueiras River, municipality of Estreito, MA; João Aires River, close to the municipality of Palmeirantes, TO; Arraias River, close to the municipality of Babaçulândia, TO, Brazil].Record: Freitas et al.(2021).
image/svg+xml109New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023Urocleidoides parodoni Oliveira, Silva, Vieira & Acosta, 2021Type-host and locality:Parodon nasus(Characiformes: Parodontidae) [Indiana stream (Capivara River, Tietê River, Upper Paraná River basin), Botucatu, SP, Brazil].Record: Oliveira et al.(2021).Urocleidoides piriatiu Mendoza-Franco & Reina, 2008Type-host and locality:Ctenolucius beani(Fowler) (Characiformes: Ctenoluciidae) [Piriati River at its confuence with the Chagres River Basin, Republic of Panama].Record: Mendoza-Franco & Reina (2008).Urocleidoides psectrogasteri Bezerra, Freitas, Cohen & Justo 2023Type-host and locality: Psectrogaster amazonica (Characiformes: Curimatidae); [Tocantins River, at the urban zone of Imperatriz municipality, MA; Brazil].Other hosts: Caenotropus labyrinthicus (Chraciformes: Chilodontidae) Record: Present studyUrocleidoides ramentacuminatus Oliveira, Santos-Neto, Tavares-Dias & Domingues, 2020Type-host and locality:Schizodon fasciatus(Characiformes: Anostomidae) [Lower Jari River, near Jarilândia district, in municipality of Vitória do Jari, AP, Brazil].Other host:Laemolyta proximaGarman (Characiformes: Anostomidae).Record:Oliveira et al.(2020).Urocleidoides reticulatus Mizelle & Price, 1964Type-Host and locality:Poecilia reticulata(Peters) [=Lebistes reticulatus(Peters)] (Cyprinodontiformes: Poeciliidae) [Host obtained from Capitol Aquarium, Sacramento, California, United States].Other hosts and localities:Belonesox belizanus Kner; Poecilia mexicana Steindachner; Poecilia petenensis Gunther; Xiphophorus helleriHeckel; Xiphophorus maculatus (Günther) (Cyprinodontiformes: Poeciliidae) [Private Aquaria, Prague and Ceske Budejovice, Czech Republic, imported freshwater ornamental fsh,Australia;Imported freshwater ornamental fsh, Sri Lanka].Records: Mizelle & Price (1964); Evans & Lester (2001); Pineda-López et al.(2005); Trujillo-González et al.(2018).Urocleidoides sapucaiensis Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva 2020Type-host and locality:Megaleporinus elongatus(Characiformes: Anostomidae) [Sapucaí-Mirim River, Upper Paraná River basin, SP, Brazil].Other locality: Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin, SP, Brazil.Record: Zago et al.(2020).Urocleidoides similuncus Mendoza-Franco, Aguirre-Macedo, & Vidal-Martínez, 2007Type-host and locality:Poecilia gillii(Cyprinodon-tiformes, Poeciliidae) [Frijolito River, Panama]. Record:Mendoza-Franco et al.(2007).Urocleidoides simonae Mendoza-Franco, Caspeta-Mandujano, Salgado-Maldonado & Matamoros, 2015Type-host and locality:Profundulus punctatus(Günther) (Cyprinodontiformes: Profundulidae) [Nandalumi River, Chiapa de Corso, Chiapas, Mexico].Other hosts and localities: Profundulus punctatus (=Profundulus balsanus); Profundulus guatemalensis(Günther), Profundulus kreiseriMatamoros, Shaefer, Hernández & Chakrabarty,Tlaloc labialis (=Profundulus labialis)(Günther), Profundulus oaxacae (Meek); Profundulussp. 1, Profundulussp. 2. (Cyprinodontiformes: Profundulidae) [Cauta River (Lempa River Basin); Cauca River, Nonoalpa River, Quebrada Los Tecomates (Lempa River Basin), El Salvador; Nil River (Guacalate River Basin); El Cantil River (Guacalate Rivr Basin); Creek El Platanar (Guacalate River basin), Guatemala; River at rancho San Antonio, Chicoasén; Arroyo Ojo de Agua, El Canelar, La Frailesca, Chiapas, Creek Tres Picos, Copainalá and River Nandalumi, Chiapa de Corso, Chiapas State; Ahl from La Laca River (Papagayo River Basin); Cahoapan River, Tamarindo River (Nexpa River basin), Guerrero State; Creek Los Sabinos (Atoyac-Verde River Basin), Chicaguaxtla River, Concepción del Progreso (Atoyac-Verde River Basin), Chico River, San Lorenzo Albarradas (Atoyac-Verde basin), Aguacate River, Juquilá, Manialtépec River Basin, Oaxaca State; Mexico].Record: Mendoza-Franco et al.(2015).
image/svg+xml110Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.Urocleidoides sinus Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva, 2020Type-host and locality:Schizodon nasutusKner (Characiformes: Anostomidae) [Sapucaí-Mirim River, Upper Paraná River basin, SP,Brazil].Other hosts and localities: Leporinus striatus Kner, Schizodon intermedius Garavello & Britski (Characiformes: Anostomidae) [Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin, SP,Brazil].Record: Zago et al.(2020).Urocleidoides solarivaginatus Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva, 2020Type-host and locality: Leporinus friderici(Characiformes: Anostomidae) [Sapucaí-Mirim River, Upper Paraná River basin, SP,Brazil].Other hosts and locality: Leporinus octofasciatus, Leporinus striatus (Characiformes: Anostomidae) [Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin, SP,Brazil].Record: Zago et al.(2020).Urocleidoides surianoae Rossin & Timi, 2016Type-host and locality:Cyphocharax voga(Hensel) (Characiformes: Curimatidae) [Chascomus Lake, Buenos Aires province, Argentina].Record: Rossin & Timi (2016).Urocleidoides tenuis Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva 2020Type-host and localityApareiodon piracicabae(Eigenmann) (Characiformes: Parodontidae) [Streams of the Middle Paranapanema River, Upper Paraná River basin, SP, Brazil].Other hosts and localities: Apareiodon afnis(Steindachner), Parodon nasus(Characiformes: Parodontidae) [Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin; Indiana stream (Capivara River, Tietê River, Upper Paraná River basin), Botucatu, SP, Brazil].Records: Zago et al.(2020); Oliveira et al.(2021).Urocleidoides tocantinensisFreitas, Bezerra, Meneses, Justo, Viana & Cohen, 2021Type host and locality:Psectrogaster amazonica (Characiformes, Curimatidae) [Tocantins River, close to the municipality of Embiral, MA, Brazil].Other host and localities:Caenotropus labyrinthicus (Characiformes, Chilodontidae); Mylesinus pauscisquamatus (Characiformes, Serrasalmidae) [Farinha River, close to the municipality of municipality of Estreito, MA; Tocantins River, close to the urban perimeter of Imperatriz; Tocantins River, at the urban zone of Imperatriz municipality, MA; Arraias River, close to the municipality of Babaçulândia, TO; Brazil].Records: Freitas et al.(2021); Present study.Urocleidoides triangulus (Suriano, 1981) Rossin & Timi, 2016Type host and locality: Cyphocharax gilbert(Quoy & Gaimard) (Characiformes: Curimatidae) [Chascomus Lake, Buenos Aires province, Argentina].Other hosts and localities: Cyphocharax modestus(Fernández-Yépez); Cyphocharax nagelii(Steindachner); Cyphocharax voga(Characiformes: Curimatidae) [Guandu River, RJ; Batalha River and Peixe River, Anhembi, SP, Brazil].Records: Suriano (1981); Suriano (1997b); Vieira et al.(2013); Abdallah et al.(2015); Rossin & Timi (2016); Dias et al.(2017); Freitas et al.(2021).Urocleidoides uncinus Zago, Yamada, Yamada, Franceschini, Bongiovani & Silva2020Type-host and locality:Gymnotus sylviusAlbert & Fernandes-Matioli (Gymnotiformes: Gymnotidae) [Sapucaí-Mirim River, Upper Paraná River basin, SP,Brazil].Other locality: Jurumirim Reservoir, Paranapanema River, Upper Paraná River basin, SP,Brazil.Record: Zago et al.(2020).Urocleidoides vaginoclaustroides Mendoza-Franco, Caspeta-Mandujano, Salgado-Maldonado & Matamoros, 2015Type-host and locality: Pseudoxiphophorus bimaculatus(Heckel) (Cyprinodontiformes: Poeciliidae) [Danta River, a tributary of the Río Lacantún basin in the Biosphere Reserve of Montes Azules (BRMA), Chiapas, Mexico].Other host and locality: Poeciliopsis retropinna(Regan) (Cyprinodontiformes: Poeciliidae) [Arroyo Jose, Chiriquicito River, Panama].Record: Mendoza-Franco et al.(2015).
image/svg+xml111New species of UrocleidoidesNeotropical Helminthology (Lima). Vol. 17, Nº2, jul - dic 2023Urocleidoides vaginoclaustrum Jogunoori, Kritsky & Venkatanarasaiah, 2004Type-host and locality:Xiphophorus hellerii (Cyprinodontiformes: Poeciliidae) [Aquarium fshes in the twin cities of Hyderabad and Secunderabad, India].Other hosts and localities: Pseudoxiphophorus bimaculatus(=Heterandria bimaculata), Xiphophorus birchmanni; Xiphophorus malinche(Cyprinodontiformes, Poeciliidae), Tlaloc labialis(=Profundulus labialis); Tlaloc portillorum (=Profundulus portillorum) (Cyprinodontiformes, Profundulidae) [Creek at Ojojona, Francisco Morazán (Nacaome River basin); Lepaterique, Francisco Morazán (Nacaome River basin),Honduras. Danta River, a tributary of the Lacantún River, Chiapas; River at rancho San Antonio, Chicoasén, Chiapas; Los Berros Spring, San Pedro River Mezquital basin, Durango; Arroyo Xontla and Conzintla River, Pánuco drainage, Hidalgo; Benito Juárez, part of Zontecomatlán River; La Antigua River basin, Veracruz, Mexico]. Records: Jogunoori et al.(2004); Salgado-Maldonado et al.(2014, 2016) Mendoza-Palmero & Aguilar-Aguilar (2008); Mendoza-Franco et al.(2015); Bautista-Hernández et al.(2014; 2019); Mendoza-Garfas et al.(2017). Remarks:Tis species was referred by Rodríguez-Santiago et al.(2016) parasitizing Pterygoplichthys pardalis(Siluriformes, Loricariidae). Considering that Urocleidoidesspp. are not reported from Siluriformes and these authors stated that only three specimens were collected, we did not consider this report. Urocleidoides vaniniSantos-Neto & Domingues, 2023Type host and locality: Erythrinus erythrinus(Bloch & Schneider) (Characiformes, Erythrinidae) [Vila Perseverança, Palheta River (Guamá River Basin), municipality of São Domingos do Capim, PA, Brazil].Record: Santos-Neto & Domingues (2023).Urocleidoides visiofortatus Mendoza-Franco & Reina, 2008Type-host and locality:Brachyhypopomus occidentalis(Regan) (Gymnotiformes: Hypopomidae) [Águas Claras River at its confuence with the Bayano Lake, Panama].Record: Mendoza-Franco & Reina (2008).Urocleidoides xinguensis Moreira, Scholz & Luque, 2015Type-host and locality:Hoplias aimara(Characiformes, Erythrinidae) [Xingu River, near Altamira, PA, Brazil].Record: Moreira et al.(2015).DISCUSSION Urocleidoideswas originally described by Mizelle & Price (1964) to accommodate the type species, U. reticulatus. Te authors proposed that species of the genus are restricted to those that present, besides vaginal sclerite, overlapping (tandem) gonads, counterclockwise cirrus rings, unmodifed anchors and hooks (pairs 1, 5 usually reduced) with enlarged shanks. Afterwards, some authors identifed diferent characteristics to the genus, Rosim et al. (2011) stated that the ventral bar with enlarged extremities may be considered a generic character for this genus. Zago et al. (2020) added an amended diagnosis, considering the recent discovery of new species of Urocleidoides, including “vagina lateral (dextral or sinistral) sclerotized; receptacle seminal present”. Besides these characteristics, Rossim & Timi (2016), Zago et al.(2020) and Freitas et al.(2021) described species with absence of coils in the male copulatory organ, and this character may be included as diagnostic for the genus, according to the amendment made by Zago et al. (2020). Santos-Neto & Domingues (2023) postulated that, although the presence of vaginal sclerite is a diferential character for Urocleidoides, it cannot be considered a main characteristic for the species of the genus. According to these authors, the species described from gymnotiform hosts, even without vaginal sclerite, share some morphological characteristics.Te type species of Urocleidoideshave been described from aquarium fshes, found in native habitats in the Neotropical Region, as U. reticulatusfrom Poecilia reticulatafrom Capitol Aquarium, Sacramento, California, USA, and another species of this genus was also reported from aquarium fshes, U. vaginoclaustrum, described from Xiphophorus hellerii. Te hosts from both species were collected from Neotropical Region and are available in aquarium in diferent countries: Israel, Czech Republic, India, California (USA) and in Central Mexico (Kritsky et al., 1986; Jogunoori et al., 2004; Mendoza-Palmero & Aguilar-Aguilar, 2008).From the total number of species, 27 were reported parasitizing a single host species, while 19 species are
image/svg+xml112Neotropical Helminthology (Lima). Vol. 18, Nº2, jul - dic 2023Bezerra et al.found in more than one host species, ranging from 2 to 6 species, belonging or not to the same family. All species of Urocleidoidesfound in more than one host species are specifc to the fsh order that they parasitize, and within the order, 12 species are restricted to one family, while 7 were recorded in more than one family, mainly in 2 diferent families from the same order, with emphasis in U. paratriangulus and U. tocantinensis, the only species recorded in 3 diferent families. Tis demonstrated that, despite the generalist behavior of Urocleidoides, in which all known species are found in 3 diferent orders, each species shows a specifcity to the host. Hoplias malabaricusis the host species which present the highest diversity in species: seven species from Urocleidoideswere described from diferent river basins in Brazil (Rosim et al. 2011; Ferreira et al., 2017; Santos-Neto & Domingues, 2023). Variations in size and shape of haptoral structures were observed by Rosim et al. (2011) from diferent river basins and were considered to be intraspecifc variation. According to Ferreira et al. (2017), this is probably due to geographic distance, phenotypic plasticity of parasites or host induced morphological change. Rosim et al.(2011) found their new species in H. malabaricusfrom diferent basins. Nevertheless, H. malabaricusappears to be a species complex when comparing cytogenetic and molecular studies. Tus, the diversity of species presented in this host may be artifcial, considering that they represent diferent Hopliasspecies that may harbour diferent Urocleidoidesspp. Considering Hoplias, nine species were reported in 10 associations from H. malabaricus and H. aimara. Anostomidae is the most representative family housing Urocleidoidesspp., represented by 12 diferent species in 23 associations host-parasite, followed by Poeciliidae, with 11 species in 13 associations and Curimatidae, with 8 species and 10 associations. Previous studies reported Urocleidoidesspp. as parasites of P. amazonica and C. labyrinthicus. In the present study,besides the new species, Urocleidoides tocantinensis were found parasitizing C. labyrinthicus, amplifying the host spectrum of this monogenoidean.In this paper, the number of valid species in genus Urocleidoides sensu strictowas increased to 46, as well as that of monogenoids in the Tocantins River basin, contributing to knowledge of the diversity of these parasites in this important region, which includes an endemic diversity of fsh and consequently, monogenoideans parasites. Further studies on other host species from this basin, whose fauna of parasites is underestimated, will provide interesting and valuable information about biogeographical and evolutionary research on dactylogyrids in the Neotropics.Author contributions: CRediT (Contributor Roles Taxonomy)CAMB= Carine A. M. Bezerra AJBF= Álvaro José B. de Freitas DCV= Diego C. Viana SCC= Simone C. Cohen MCNJ= Marcia C. N. Justo Conceptualization:CAMB, AJBF, SCC, MCNJData curation: CAMB, AJBFFormal Analysis: SCC, MCNJFunding acquisition: SCC, MCNJ, DCVInvestigation: CAMB, AJBF, SCC, MCNJMethodology: CAMB, AJBFProject administration: SCC, MCNJResources: SCC, MCNJ, DCVSupervision: SCC, MCNJValidation: SCC, MCNJWriting – original draft: CAMB, AJBFWriting – review & editing: SCC, MCNJ
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