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Parasite in Geoff roy’s cat
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Neotropical Helminthology
Neotropical Helminthology, 2026, vol. 20 (2), 227-234
ORIGINAL ARTICLE / ARTÍCULO ORIGINAL
EXPANDING MORPHOLOGICAL KNOWLEDGE OF TWO NEMATODE
SPECIES PARASITIZING SOUTHEASTERN2 BRAZILIAN MARINE FISHES
AMPLIACIÓN DEL CONOCIMIENTO MORFOLÓGICO DE DOS ESPECIES
DE NEMATODOS PARÁSITOS DE PECES MARINOS DEL SURESTE
DE BRASIL
Patrícia Silva de Oliveira1*, Arthur Bessi Machado2 & José Luis Luque3
ISSN Versión Impresa 2218-6425 ISSN Versión Electrónica 1995-1403
DOI: https://doi.org/10.62429/rnh20262022183
Este 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 Afi 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.
ABSTRACT
Histological approaches have increasingly been incorporated into integrative taxonomy as complementary sources of
morphological information. However, their application to parasitic nematodes remains limited. e present study provides
a histological characterization of two nematode taxa recovered from marine fi shes collected along the coast of the State
of Rio de Janeiro, southeastern Brazil: larval specimens of Anisakis sp. from Pogonias cromis (Linnaeus, 1766) and adult
specimens of Dichelyne (Cucullanellus) sciaenidicola Timi, Lanfranchi, Tavares & Luque, 2009 from Micropogonias furnieri
(Desmarest, 1823). Recovered nematodes were fi xed in neutral buff ered formalin, processed using routine histological
techniques, sectioned at 5 µm, and stained with hematoxylin and eosin and Gomori’s trichrome. Histological examination
of Anisakis sp. revealed a thick multilayered cuticle, well-developed somatic musculature, ventriculus, intestine, and boring
tooth, corroborating the diagnostic morphology of anisakid larvae. In D. (C.) sciaenidicola, the esophagus exhibited a
characteristic division into an anterior isthmus-like region and a posterior bulbous portion, while the intestine displayed
small villus-like mucosal projections and a cuticular lining in the anterior digestive tract. e observations obtained were
consistent with previous morphological descriptions and provided additional information on the internal organization
of both species. e results demonstrate that histological analyses can contribute useful complementary characters for
nematode taxonomy, while expanding knowledge of the morphofunctional organization of marine fi sh parasites from
the Brazilian coast.
Keywords: Anisakidae – Cucullanidae – fi sh parasites – histology – integrative taxonomy – morphology
1 Programa de Pós-Graduação em Ciências Veterinárias, Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil.
2 Programa de Pós-Graduação em Biologia Animal, Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil.
3 Departamento de Parasitologia Animal, Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil.
* Corresponding author: patricia.oliveira@faa.edu.br
Patrícia Silva de Oliveira: https://orcid.org/0009-0004-7232-0692
Arthur Bessi Machado: https://orcid.org/0009-0003-8793-7380
José Luis Luque: https://orcid.org/0000-0003-3515-1127
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Silva de Oliveira et al.
RESUMEN
Los enfoques histológicos se han incorporado cada vez más a la taxonomía integrativa como fuentes complementarias
de información morfológica. Sin embargo, su aplicación al estudio de nematodos parásitos sigue siendo limitada. El
presente estudio proporciona una caracterización histológica de dos taxones de nematodos recuperados de peces marinos
recolectados a lo largo de la costa del estado de Río de Janeiro, en el sureste de Brasil: ejemplares larvarios de Anisakis sp.,
procedentes de Pogonias cromis (Linnaeus, 1766), y ejemplares adultos de Dichelyne (Cucullanellus) sciaenidicola Timi,
Lanfranchi, Tavares & Luque, 2009, procedentes de Micropogonias furnieri (Desmarest, 1823). Los nematodos recuperados
fueron fijados en formalina tamponada neutra, procesados mediante técnicas histológicas de rutina, seccionados a 5 µm
y teñidos con hematoxilina-eosina y tricrómico de Gomori. El examen histológico de Anisakis sp. reveló una cutícula
gruesa y multilaminada, musculatura somática bien desarrollada, ventrículo, intestino y diente perforador, corroborando
la morfología diagnóstica de las larvas anisákidas. En D. (C.) sciaenidicola, el esófago presentó una división característica
en una región anterior semejante a un istmo y una porción posterior bulbosa, mientras que el intestino mostró pequeñas
proyecciones mucosas similares a vellosidades y un revestimiento cuticular en la porción anterior del tracto digestivo.
Las observaciones obtenidas fueron consistentes con las descripciones morfológicas previas y aportaron información
adicional sobre la organización interna de ambas especies. Los resultados demuestran que los análisis histológicos pueden
aportar caracteres complementarios útiles para la taxonomía de nematodos, además de ampliar el conocimiento sobre la
organización morfofuncional de los parásitos de peces marinos de la costa brasileña.
Palabras clave: Anisakidae – Cucullanidae – histología – morfología – parásitos de peces –taxonomía integrativa
INTRODUCTION
With the advent of new techniques and technologies,
an increasing number of tools have been incorporated
into species descriptions and redescriptions. Within the
framework of integrative taxonomy, multiple complementary
approaches, including diverse morphological analyses,
molecular methods, ecological data, histology, and scanning
electron microscopy, have been applied concurrently in
the study of parasitic species (Dayrat, 2005; De León &
Poulin, 2018; Machado et al., 2024ab). Recent studies have
demonstrated that the integration of multiple sources of
morphological and molecular evidence provides a more
comprehensive understanding of parasite morphology
and contributes to more robust species diagnoses and
redescriptions (Machado et al., 2024ab), while histological
analyses have emerged as an additional complementary
source of taxonomic information. is methodological
integration allows a more comprehensive assessment of
taxonomic characters at both species and higher taxonomic
levels, while promoting the exchange of knowledge among
distinct research fields. Furthermore, it may facilitate the
recognition of novel taxonomically informative characters
through the application of methodologies that have been
rarely or never previously employed in the study of a given
group, as well as support evolutionary interpretations
(Schlick-Steiner et al., 2010; Rojas et al., 2025).
In this context, histological analyses may expand the range
of descriptive characters available for parasitic species
by incorporating a cellular and tissue-level perspective
into morphological studies, thereby enabling functional
interpretations of organs and structures. Although
histological approaches have occasionally been employed
in taxonomic studies of parasites, their use has only recently
received renewed attention (Machado et al., 2024ab, 2026;
Silva De Oliveira et al., 2026). However, most of these
investigations have focused on platyhelminths, leaving
substantial gaps in knowledge regarding the histological
organization of other parasitic groups. In addition, a
large proportion of parasitological studies employing
histological techniques have been directed toward the
assessment of host-parasite interactions and associated
pathological alterations rather than the intrinsic structural
characterization of the parasites themselves (Dezfuli et al.,
2005; Magdálek et al., 2021; Ramdani et al., 2022).
Accordingly, the present study provides a histological
characterization of two nematode taxa, Anisakis sp. and
Dichelyne (Cucullanellus) sciaenidicola Timi, Lanfranchi,
Tavares & Luque, 2009, recovered from the marine
fishes Pogonias cromis (Linnaeus, 1766) (Black drum) and
Micropogonias furnieri (Desmarest, 1823) (Whitemouth
croaker), respectively, collected along the coast of the State
of Rio de Janeiro, southeastern Brazil. Anisakis sp. was
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selected because of its worldwide distribution, zoonotic
importance, and the limited histological information
available for its larval stages, whereas D. (C.) sciaenidicola
represents a common parasite of sciaenid fishes and provides
an opportunity for comparative histological analysis between
distinct nematode taxa. By complementing the existing
morphological knowledge of these taxa, this study seeks
to encourage the incorporation of histological approaches
into nematode taxonomy, contributing additional sources
of descriptive information and broadening the scope of
future taxonomic and comparative investigations.
MATERIAL AND METHODS
Host Necropsy and Parasite Collection
A total of eight specimens each of Micropogonias furnieri
(Desmarest, 1823) and Pogonias cromis (Linnaeus, 1766)
were obtained from local fishermen at the Central de
Abastecimento do Estado do Rio de Janeiro (CEASA),
Rio de Janeiro State, Brazil. e fish were maintained
under refrigeration and transported to the Laboratório
de Parasitologia de Peixes e Mamíferos Silvestres e
Sinantrópicos, Universidade Federal Rural do Rio de Janeiro,
Seropédica, Brasil, where necropsies were performed. Host
identification followed Menezes & Figueiredo (1985).
e digestive tract of each fish was removed, opened
longitudinally, and washed under running water through
a 150 µm mesh sieve. e retained material was transferred
to Petri dishes and examined under a stereomicroscope
for the presence of parasites. Recovered specimens were
collected and fixed in 10% neutral buffered formalin for
subsequent histological processing. All procedures involving
formalin were carried out in a chemical fume hood using
appropriate personal protective equipment, in accordance
with institutional biosafety guidelines for the handling and
disposal of hazardous chemical substances. Preliminary
identification was based on the available taxonomic
literature. Larvae of Anisakis sp. were identified following
Berland (1961), Smith (1983), and Mattiucci & Nascetti
(2008), whereas specimens of Dichelyne (Cucullanellus)
sciaenidicola were identified according to the original
description by Timi et al. (2009) and subsequent taxonomic
revisions.
Histological Processing
Histological processing of the nematodes followed the
protocol described by Abrahamsohn (2016). Specimens
were dehydrated through a graded ethanol series, cleared
in xylene, and embedded in paraffin wax. Longitudinal
and transverse sections, 5 µm thick, were obtained using
an automatic microtome. Sections were stained with
hematoxylin-eosin (HE) and Gomori’s trichrome (GT).
Permanent slides were mounted using Entellan® mounting
medium. Histological preparations were examined and
photographed using an Olympus BX51 light microscope
equipped with the Capture 2.4 image analysis software.
Voucher specimens (histological slides) of both specimens
were deposited in the Helminthological Collection of the
Oswaldo Cruz Institute (CHIOC), Rio de Janeiro (Brazil),
under the following deposit numbers: Anisakis sp. - 41120,
Dichelyne (Cucullanellus) sciaenidicola - 41121.
Ethical Aspects
All applicable institutional, national, and international
guidelines for the care and use of animals were followed.
As no live animals were used and all fish specimens were
obtained from a local fish market, approval by an animal
ethics committee was not required, in accordance with
Article 3, Item III, of Brazilian Federal Law No. 11.794.
RESULTS
Anisakis sp. (L3, third-stage larvae)
Based on 06 specimens.
Host: Pogonias cromis (Linnaeus, 1766), black drum.
Site of infection: Gut.
Prevalence: 25%.
Deposited material: FIOC number 41120.
Acellular tegument covered by a thick multilaminar
cuticle composed of regular transverse striations (Fig.
1a–e). Subcuticular region formed by extensive bundles
of transversely striated muscle fibers arranged longitudinally
(Fig. 1a–e). e interior of the pseudocoelomic cavity is
composed of stroma consisting of loose connective tissue
and free acidophilic cells with prominent nuclei (Fig.
1a–c). In transverse section, two large lateral muscle groups
are observed, whose dorsal and ventral junctions appear
associated with attachment sites for internal structures
and the subcuticular region (fig. 1a). Esophagus lined
by columnar epithelium (fig. 1d). Ventriculus lined by
epithelium of variable thickness (fig. 1a, b). Intestine lined
by columnar epithelium (fig. 1c, e). e digestive tract
contains luminal content throughout its extension (fig.
1a, c, e). Boring tooth consisting of a cuticular projection
(fig. 1b). Latero-terminal anus represented by a region of
cuticular invagination (fig. 1e).
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Silva de Oliveira et al.
Figure 1. Histological transversal (a) and longitudinal sections (b–e) of Anisakis sp., stained with Gomori’s trichrome
(GT). a middle body, b entire body, c middle body, d esophagus in detail, and e terminal portion in detail. Abbreviations:
c cuticle, m musculature, e stroma, v ventriculus, bt boring tooth, i intestine, and * indicates anus.
Dichelyne (Cucullanellus) sciaenidicola Timi, Lanfranchi,
Tavares & Luque 2009
Based on 27 adult specimens
Host: Micropogonias furnieri (Desmarest, 1823),
whitemouth croaker.
Site of infection: Gut.
Prevalence: 37.5%.
Deposited material: FIOC number 41121.
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in syncytial tegument covered by thin acidophilic
cuticle (Fig. 2a–h). Subepidermal region composed of
loose connective tissue (Fig. 2a–h). Supporting musculature
not evident. Interior of pseudocoelomic cavity composed
of stroma consisting predominantly of loose connective
tissue and small free cells (Fig. 2a–h). Large acidophilic
coelomocytes with prominent, apparently lobulated nuclei
and cytoplasmic granules observed in the region lateral to
the intestine (fig. 2a). In transverse section, two discrete
lateral muscle groups observed, whose dorsal and ventral
junctions appear associated with attachment points of
filling stroma and internal structures (fig. 2a). Cephalic
structures consisting of small projections of thin lining
epithelium and underlying stroma (fig. 2b, c). Mouth
terminally located in the anterior region, with the presence
of small cells embedded in surrounding connective tissue
(fig. 2b, c). Pseudobuccal cavity composed of two lateral
groups of small longitudinal muscle bundles, basophilic
cells, and mucous glands, with a hollow cavity present
in the most anterior portion of the structure (fig. 2b,
c). Esophagus elongated, divided into two portions; the
anterior portion isthmus-shaped and posterior portion
bulb-shaped; esophagus lined by acidophilic cuticle, with
directly underlying region formed by acidophilic glandular
cells arranged as tubular structures (fig. 2b–e). Intestine
lined by cuboidal epithelium composed of small cells of
variable thickness; in transverse section, small mucosal
projections similar to villi are observed (fig. 2a, e–h). e
digestive tract contains luminal content throughout its
extension (Fig. 2e–h).
DISCUSSION
is study provides a histological characterization of two
nematode species recovered from marine fishes: larval
specimens of Anisakis sp. and adult specimens of D.
(Cucullanellus) sciaenidicola. Histological techniques have
traditionally been employed in helminthological studies
primarily to assess the pathological effects of parasites
on their hosts (Dezfuli et al., 2005; Magdálek et al.,
2021; Ramdani et al., 2022). More recently, however,
these approaches have also been incorporated into
integrative taxonomic studies to provide morphofunctional
information that complements conventional morphological
descriptions (Machado et al., 2024ab, 2026; Silva De
Oliveira et al., 2026).
e histological characterization of the Anisakis sp. larvae
recovered from P. cromis revealed an internal organization
consistent with the diagnostic morphology traditionally
recognized for the genus, despite the absence of species-
level identification (Bicudo et al., 2005). e presence of
a thick, acellular, multilayered cuticle associated with well-
developed somatic musculature represents a characteristic
feature of anisakid nematodes, as described by Moravec
(1998). In addition, the visualization of the boring tooth
and ventriculus in longitudinal sections provided anatomical
evidence supporting generic identification, highlighting the
value of histological preparations for the examination of
structures that may be obscured in whole-mounted larval
specimens. For D. (Cucullanellus) sciaenidicola, histological
examination allowed a detailed assessment of the esophageal
morphology, including an anterior isthmus-like region
and a posterior bulbous portion, in agreement with the
diagnostic features reported by Timi et al. (2009). An
additional feature documented in the present study was
the occurrence of villus-like projections of the intestinal
mucosa, a character rarely emphasized in descriptions based
exclusively on external morphology or cleared specimens.
e agreement between the present observations and the
descriptions provided by Moravec (1998), Bicudo et al.
(2005), and Timi et al. (2009) demonstrates that histological
data can provide a valuable source of complementary
taxonomic information. Beyond corroborating species
identity, the examination of internal tissue organization may
reveal additional characters that contribute to comparative
morphological studies and broaden the range of traits
available for integrative taxonomic analyses.
e muscular organization of nematodes has been
investigated since the early development of nematode
systematics, including attempts to classify taxa according
to the arrangement of muscle cells and fibers (Chitwood,
1931). In the present study, the somatic musculature
exhibited morphological similarities to striated muscle
tissue described in vertebrates. Comparable observations
have also been reported in other parasitic invertebrates
(Landman et al., 2023; Machado et al., 2024a; Machado
et al., 2024b; Machado et al., 2026; Silva De Oliveira et
al., 2026). Although these similarities are based solely on
histological appearance, they may indicate the conservation
of certain structural features among metazoans. Further
studies integrating histological, molecular, and physiological
approaches are necessary to determine whether these
similarities extend beyond morphology.
Regarding muscular development, larval specimens of
Anisakis sp. exhibited proportionally more developed somatic
musculature than adult specimens of D. (C.) sciaenidicola.
is difference may reflect distinct functional demands
associated with different life stages and ecological strategies.
However, previous studies on Ascaris Linnaeus, 1758 have
demonstrated substantial changes in musculature throughout
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Silva de Oliveira et al.
Figure 2. Histological transversal (a) and longitudinal sections (b–h) of Dichelyne (Cucullanellus) sciaenidicola, stained
with hematoxylin and eosin (HE). a middle body, b anterior portion, c anterior portion in detail, d forebody, e digestive
structures in detail, f intestine in detail, g posterior body section, and h terminal portion in detail. Abbreviations: c
cuticle, e stroma, i intestine, pc pseudobuccal cavity, es esophagus, ce cephalic structures, and * indicates the mouth.
ontogenetic development (Stretton, 1976). Consequently,
comparative investigations involving multiple developmental
stages of the same species are required to determine whether
the differences observed here are primarily related to
ontogeny or represent species-specific traits.
In Dichelyne (Cucullanellus) sciaenidicola, small projections
of the intestinal mucosa extending toward the lumen were
observed. Similar structures have previously been reported
in a digenean trematode (Machado et al., 2024a), where
they were interpreted as increasing the absorptive surface
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of the digestive tract. Although functional studies are
lacking, the structures observed in the present material
may perform a comparable role. Furthermore, the cuticular
lining observed in the anterior portion of the digestive tract
agrees with previous descriptions for other nematode taxa
(Basyoni & Rizk, 2016).
e histological analyses performed on Anisakis sp. larvae
and D. (Cucullanellus) sciaenidicola adults demonstrated
that the internal organization of these nematodes is
consistent with diagnostic features previously established
through classical morphological studies. e visualization
of structures such as the cuticle, somatic musculature,
and digestive tract provided complementary evidence for
taxonomic characterization while revealing anatomical
details that are often difficult to observe in conventional
whole-mounted specimens. Although restricted to two
species, the present study highlights the usefulness
of histological approaches as a complementary tool
in nematode taxonomy and morphology, expanding
the morphofunctional knowledge available for marine
fish parasites from the Brazilian coast and providing a
comparative basis for future integrative studies involving
related taxa.
ACKNOWLEDGMENTS
Arthur B. Machado was supported by a student fellowship
from the Coordenação de Aperfeiçoamento de Pessoal de
Nível Superior (CAPES), Brazil (Finance Code 001). José
L. Luque received support from a research grant provided
by the Conselho Nacional de Desenvolvimento Científico
e Tecnológico (CNPq), Brazil. No funding was received
for conducting this study. e authors declare no conflicts
of interest.
Author contributions: CRediT (Contributor Roles
Taxonomy)
PSO = Patrícia Silva de Oliveira
ABM = Arthur Bessi Machado
JLL = José Luis Luque
Conceptualization: PSO, ABM, JLL
Data curation: PSO
Formal Analysis: PSO, ABM, JLL
Funding acquisition: PSO, ABM, JLL
Investigation: PSO, ABM, JLL
Methodology: PSO, ABM, JLL
Project administration: JLL
Resources: PSO
Software: PSO, ABM, JLL
Supervision: JLL
Validation: PSO, ABM, JLL
Visualization: PSO, ABM, JLL
Writing – original draft: PSO, ABM, JLL
Writing – review & editing: PSO, ABM, JLL
BIBLIOGRAPHIC REFERENCES
Abrahamsohn, P. (2016). Histologia. 1
st
ed. Guanabara
Koogan.
Basyoni, M. M., & Rizk, E. M. (2016). Nematodes
ultrastructure: complex systems and processes.
Journal of Parasitic Diseases, 40, 1130–1140.
Berland, B. (1961). Nematodes from some Norwegian
marine fishes. Sarsia, 2, 1–50.
Bicudo, A. J. A., Tavares, L. E. R., & Luque, J. L. (2005).
Larvas de Anisakidae (Nematoda: Ascaridoidea)
parasitas da cabrinha Prionotus punctatus (Bloch,
1793) (Osteichthyes: Triglidae) do litoral do
Estado do Rio de Janeiro, Brasil. Revista Brasileira
de Parasitologia Veterinária, 14, 109–118.
Chitwood, B. G. (1931). A comparative histological study
of certain nematodes. Zeitschrift für Morphologie
und Ökologie der Tiere, 23, 237–284.
Dayrat, B. (2005). Towards integrative taxonomy. Biological
Journal of the Linnean Society, 85, 407–417.
De León, G. P. P., & Poulin, R. (2018). An updated look at
the uneven distribution of cryptic diversity among
parasitic helminths. Journal of Helminthology, 92,
197–202.
Dezfuli, B. S., Giari, L., Simoni, E., Shinn, A. P.,
Manera, M., & Bosi, G. (2005). Histopathology,
ultrastructure and immunohistochemistry of
Coregonus lavaretus hearts naturally infected with
Ichthyocotylurus erraticus (Trematoda). Diseases of
Aquatic Organisms, 66, 245–254.
Landman, W., Verneau, O., Vences, M., & du Preez,
L. (2023). Metapolystoma ohlerianum n. sp.
(Monogenea: Polystomatidae) from Aglyptodactylus
madagascariensis (Anura: Mantellidae). Acta
Parasitologica, 68, 344–358.
Machado, A. B., Simões, R. O., Maldonado, A., Jr.,
Santos, M. A. J., & Luque, J. L. (2024a). Integrative
234
Neotropical Helminthology, Vol. 20, Nº2, jul - dec 2026
Silva de Oliveira et al.
Taxonomy of Prosogonotrema bilabiatum Vigueras,
1940 (Digenea: Sclerodistomidae): A Parasite in
Atlantic Spadefish Chaetodipterus faber (Broussonet,
1782) (Acanthuriformes: Ephippidae) from Brazil.
Acta Parasitologica, 69, 898–909.
Machado, A. B., Simões, R. O., Maldonado, A., Jr., Santos,
M. A. J., & Luque, J. L. (2024b). Morphological and
Molecular Analyses of Aponurus laguncula Looss,
1907 (Digenea: Lecithasteridae) Parasitic in Atlantic
Spadefish Chaetodipterus faber (Broussonet, 1782)
(Acanthuriformes: Ephippidae) from Brazilian
Coastal Zone. Acta Parasitologica, 69, 1600–1612.
Machado, A. B., Simões, R. O., Chero, J. D., Maldonado,
A., Jr., Santos, M. A. J., & Luque, J. L. (2026).
Integrative redescription of Sprostoniella micrancyra
Cezar, Luque and Amato, 1999 (Monopisthocotylea:
Capsalidae) and the phylogenetic position of
Sprostoniella Bychowsky and Nagibina, 1967. Acta
Parasitologica, 71, 124.
Magdálek, J., Makovický, P., & Vadlejch, J. (2021).
Nematode-induced pathological lesions and
alterations of mucin pattern identified in abomasa
of wild ruminants. International Journal for
Parasitology: Parasites and Wildlife, 14, 62–67.
Mattiucci, S., & Nascetti, G. (2008). Advances and trends
in the molecular systematics of anisakid nematodes,
with implications for their evolutionary ecology and
host–parasite co-evolutionary processes. Advances
in Parasitology, 66, 47–148.
Menezes, N. A., & Figueiredo, J. L. (1985). Manual de
peixes marinhos do sudeste do Brasil. V. Teleostei (4).
Museu de Zoologia, Universidade de São Paulo.
Moravec, F. (1998). Nematodes of freshwater fishes of
the Neotropical region. Praha: Academia. Kluwer
Academic Publishers.
Ramdani, S., Trilles, J. P., & Ramdane, Z. (2022).
Histopathological changes from parasitic Nematoda
infestation in the musculature of some marine
teleost fishes from the Algerian coast. Fisheries &
Aquatic Life, 30, 209–216.
Rojas, A., Bass, L. G., Campos-Camacho, J., Dittel-Meza,
F. A., Fonseca, C., Huang-Qiu, Y. Y., Olivares, R.
W. I., Romero-Vega, L. M., Villegas-Rojas, F., &
Solano-Barquero, A. (2025). Integrative taxonomy
in helminth analysis: protocols and limitations in
the twenty-first century. Parasites & Vectors, 18, 87.
Schlick-Steiner, B. C., Steiner, F. M., Seifert, B., Stauffer,
C., Christian, E., & Crozier, R. H. (2010).
Integrative taxonomy: a multisource approach to
exploring biodiversity. Annual Review of Entomology,
55, 421–438.
Silva de Oliveira, P., Bessi Machado, A., & Luque, J. L.
(2026). Histological diagnosis of Lecithochirium
species parasitizing Trichiurus lepturus Linnaeus,
1758 from southeastern Brazil. Neotropical
Helminthology, 20, 89-96.
Smith, J. W. (1983). Anisakis simplex (Rudolphi, 1809,
det. Krabbe, 1878) (Nematoda: Ascaridoidea):
Morphology and morphometry of larvae from
euphausiids and fish, and a review of the life-history
and ecology. Journal of Helminthology, 57, 205–224.
Stretton, A. O. W. (1976). Anatomy and development of
the somatic musculature of the nematode Ascaris.
Journal of Experimental Biology, 64, 773–788.
Timi, J. T., Lanfranchi, A. L., Tavares, L. E. R., & Luque, J.
L. (2009). A new species of Dichelyne (Cucullanellus)
(Nematoda: Cucullanidae) parasitizing the
whitemouth croaker Micropogonias furnieri
(Desmarest) (Pisces: Sciaenidae) from the Southwest
Atlantic. Journal of Parasitology, 95, 892–898.
Received June 25, 2026.
Accepted July 10, 2026.
Published online July 13, 2026.
