ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043

ORIGINAL ARTICLE / ARTÍCULO ORIGINAL

OLIVACOTYLE HEMANTHIASI N. GEN. N. SP. (MONOGENEA: DICLIDOPHORIDAE) FROM THE

GILLS OF DAMSEL BASS HEMANTHIAS SIGNIFER (GARMAN, 1899) (TELEOSTEI: SERRANIDAE)

IN THE SOUTH AMERICAN PACIFIC OCEAN

OLIVACOTYLE HEMANTHIASI N. GEN. N. SP. (MONOGENEA: DICLIDOPHORIDAE) DE LAS

BRANQUIAS DE LA CABRILLA DONCELLA HEMANTHIAS SIGNIFER (GARMAN, 1899)

(TELEOSTEI: SERRANIDAE) EN EL OCÉANO PACÍFICO DE AMÉRICA DEL SUR

1Laboratorio de Parasitología General y Especializada, Facultad de Ciencias Naturales y Matemática, Universidad

Nacional Federico Villarreal (UNFV), Código postal 15007, El Agustino, Lima, Perú.

2Laboratorio de Ecología y Biodiversidad Animal (LEBA). Facultad de Ciencias Naturales y Matemática (FCNNM).

Universidad Nacional Federico Villarreal (UNFV). El Agustino, Lima, Perú.

4Laboratorio de Parasitología. Facultad de Ciencias Biológicas. Universidad Ricardo Palma. Santiago de Surco, Lima,

Perú.

3Departamento de Parasitologia Animal, Universidade Federal Rural do Rio de Janeiro, Caixa Postal 74.540, Seropédica,

RJ, Brazil, CEP 23851-970.

*Corresponding author: E-mail: luqueufrrj@gmail.com

1 1 1 2,3 4,

Celso L. Cruces ; Jhon D. Chero ; Gloria Sáez ; José Iannacone & José L. Luque *

ABSTRACT

Keywords: damsel bass – Diclidophoridae – ectoparasites – Hemanthias signifer – Olivacotyle hemanthiasi – Pacific Ocean –

Peru – Serranidae

A new genus and species of Monogenea, Olivacotyle hemanthiasi n. gen., n. sp., are proposed to

accommodate diclidophorids collected from the gills of damsel bass Hemanthias signifer (Garman, 1899)

(Serranidae) in Peru, South American Pacific Ocean (45°54'S, 81°05'W). While clearly a member of the

Diclidophorinae Cerfontaine, 1895 that includes parasites of marine teleosts, O. hemanthiasi n. sp. differs

from the other 15 members of this subfamily by the combination of the following six characteristics: (1)

haptoral clamps with the anteromedial sclerite and one anterolateral sclerite fused forming a ring, with

other anterolateral sclerite not fused with any sclerites of inner anterior quadrant, lamellate extension

partially reduced and distal posterolateral sclerites not fused distally; (2) clamps arranged in a semicircle

around the posterior edge of the haptor; (3) testes extend into the haptor; (4) vagina absent; (5) seminal

receptacle well developed; and (6) prostatic vesicle absent.

Neotropical Helminthology

387

Neotropical Helminthology, 2017, 11(2), jul-dic: 387-394

INTRODUCTION

388

RESUMEN

Palabras claves: cabrilla doncella – Diclidophoridae – ectoparásitos – Hemanthias signifer – Océano Pacífico –

Olivacotyle hemanthiasi – Perú – Serranidae

Un nuevo género y especie de Monogenea, Olivacotyle hemanthiasi n. gen., n. sp., se proponen para

acomodar especímenes colectados de las branquias de la cabrilla doncella Hemanthias signifer (Garman,

1899) (Serranidae) en el Perú, Océano Pacífico Sudamericano (45°54'S, 81°05'W). Si bien es un miembro

de Diclidophorinae Cerfontaine, 1895 que incluye parásitos de teleósteos marinos, O. hemanthiasi n. sp.

difiere de los otros 15 miembros de esta subfamilia por la combinación de las siguientes seis

características: (1) pinzas haptorales con la esclerita anteromedial y una esclerita anterolateral fusionadas

formando un anillo, la otra esclerita anterolateral no fusionada con ninguna esclerita del cuadrante

anterior interno, extensión lamelar parcialmente reducida y escleritas posterolateral distal no fusionadas

distalmente; (2) pinzas dispuestas en semicírculo alrededor del borde posterior del haptor; (3) testículos

extendiéndose hacia el haptor; (4) vagina ausente; (5) receptáculo seminal bien desarrollado; y (6)

vesícula prostática ausente.

these specimens was compared to related species of

the Diclidophoridae herein.

Fish were collected between July 2014 to January

2015 off the coast of Puerto Pizarro, Tumbes, Peru

(45°54'S, 81°05'W), using gillnets and were

dissected immediately after capture. The gills were

excised and placed in Petri dishes with sea water

and examined for monogeneans with the aid of a

stereomicroscope. Monogeneans found were fixed

in hot 4% formalin, stained with Semichon's

carmine or Gomori's trichrome, clarified in

Eugenol and mounted in Canada balsam. Other

specimens were mounted in Gray and Wess

medium (Humason, 1979) for the study of

sclerotized structures. Specimens were examined

T M

using a compound O l y m p u s B X 5 1

photomicroscope equipped with normal light and

differential interference contrast microscopy

(DIC) optics and drawings were made with the aid

of a drawing tube. Measurements are in

micrometers, unless otherwise indicated, using

straight-line distances between extreme points of

the structures measured and are expressed as the

range followed by the mean and number (n) of

structures measured in parentheses.

Nomenclature of the clamp sclerites considered its

relative position follows Kritsky & Klimpel

Currently, 45 species belonging to 18 genera of the

Diclidophoridae Fuhrmann, 1928 have been

described from several marine South American

teleost fishes. From them, only Hemitagia

galapagensis (Meserve, 1938), has been reported

on fishes of the Serranidae: the Peruvian rock

seabass Paralabrax humeralis (Valenciennes,

1828) and the creole-fish Paranthias furcifer

(Valenciennes, 1828) (Cohen et al., 2013; Eiras et

al., 2016; Luque et al., 2016; Gómez del Prado-

Rosas et al., 2017).

The damsel bass Hemanthias signifer (Garman,

1899) (Serranidae) is a demersal fish distributed in

the Eastern Pacific waters, from southern

California and the Gulf of California to northern

Peru, including Malpelo Island (Chirichigno &

Cornejo, 2001); like other serranids, the damsel

bass inhabits rocky reef areas from 25 to 305m in

depth (Heemstra, 1995; Anderson & Heemstra,

2012). This species is listed as presenting least

concern by International Union for Conservation

of Nature and Natural Resources (IUCN Red List)

(Smith-Vaniz et al. 2010).

During a parasitological survey on monogeneans

parasites of demersal fish species off Tumbes,

northern Peru, specimens were collected from the

gills of H. signifier, representing a new species and

genus of the Diclidophoridae. The morphology of

Neotropical Helminthology, 2017, 11(2), jul-dic Cruces et al.

MATERIALS AND METHODS

Neotropical Helminthology, 2017, 11(2), jul-dic

(2007). Fishes were identified according to

Chirichigno & Vélez (1998) and Anderson &

Heemstra (2012). Prevalence and mean intensity of

infection were calculated according to Bush et al.

(1997).

For comparative purposes, the following

specimens deposited in the Helminthological

Collection of the National Museum of Natural

History, Smithsonian Institution (USNM), USA

and in the Colección Nacional de Helmintos,

Instituto de Biologia, Universidad Nacional

Autónoma de México (CNHE), Mexico were

examined: Lampanyctophilus wisneri Payne, 1986

(USNM 79178, holotype) and Mamaevicotyle

villalobosi Lamothe, 1984 (CNHE 0205,

holotype). Type material was deposited in the

Helminthological Collection of the Museum of

Natural History, San Marcos University (MUSM),

Lima, Peru, and in the Helminthological Collection

of Instituto Oswaldo Cruz (CHIOC), Rio de

Janeiro, Brazil.

Diclidophoridae Fuhrmann, 1928

Diclidophorinae Cerfontaine, 1895

Olivacotyle n. gen.

Diagnosis. Body symmetrical, not divided. Haptor

not separated from body, bearing 4 pairs of slightly

pedunculated clamps, arranged in semicircle

around posterior edge of haptor. Clamps

asymmetrical with opposable valves. Anterior

valve formed by anterolateral sclerite and anterior

ring-shape sclerite (fusion of anteromedial sclerite

and one anterolateral sclerite); other anterolateral

sclerite not fused to distal and proximal ends of

anterior ring-shape sclerite, lamellate extension

partially reduced; slightly muscular circular pad

present in inner anterior quadrant of clamp.

Posterior valve formed by posteromedial sclerite,

two distal posterolateral sclerites not fused distally

and two proximal posterolateral sclerites. Terminal

lappet absent. Three pairs of anchors, two outer

pairs simple and an inner pair sickle-shaped with

long shank, curved shaft and point; arranged in

posterior edge of haptor. Buccal suckers paired, not

septate. Pharynx present. Ceca diverticulated,

extending into haptor, not confluent posteriorly.

Testes numerous, postovarian, extend into the

haptor. Copulatory organ armed with 6 curved

hooks with bifid base. Prostatic vesicle absent.

Ovary tubular, elongate, double inverted

U–shaped. Seminal receptacle well developed.

Vagina absent. Elongated uterus. Eggs with single

long, strongly coiled filament at abopercular pole.

Parasites on gills of serranids.

Type species: Olivacotyle hemanthiasi n. gen. n.

sp.

Etymology: The generic epithet is named in honor

of Marcelo Enrique Oliva Moreno (Universidad de

Antofagasta, Chile) for his contribution to the

marine ichthyoparasitology.

Remarks

Currently, the Diclidophoridae includes 52

recognized genera, 15 of them belonging to the

Diclidophorinae, i.e., Diclidophora Kroyer, 1838;

Diclidophoroides Price, 1943; Osphyobothrus

Yamaguti, 1958; Flexophora Prost & Euzet, 1962;

Allotagia Dillon & Hargis, 1965; Upenicola

Unnithan, 1966; Lebboia Mamev & Paruchin,

1975; Polyipnicola Mamaëv & Paruchin, 1975;

Tribuliphorus Mamaev & Parukhin, 1977;

In bj umia Mamaev & Parukhin, 1984;

Mamaevicotyle Lamothe-Argumedo, 1984;

Camp ec hi a Zhukov & Mamaev, 1985;

L a m p a n y c t o p h i l u s P a y n e , 1 9 8 6 ;

Mamaevodiclidophora Rubec, 1991 &

Macrouridophora Rubec & Dronen, 1994

(Mamaev, 1976; Payne, 1986; Rubec & Dronen

1994).

The new genus differs from Allotagia because the

species of this genus has the posterior valve of the

clamp with the distal posterolateral sclerites and

the proximal posterolateral esclerites fused to form

a single pair of long sclerites (not fused in members

of Olivacotyle n. gen.).

The species of Campechia, Flexophora, Inbjumia

and Polyipnicola have the anterior valve of the

clamp with the anterolateral sclerites fused distally

(these are not fused in members of the new genus).

The new genus differs from Tribuliphorus because

the species of this genus has the anterolateral

sclerites and the distal posterolateral sclerites

equipped with large curved teeth (absent in

members of Olivacotyle n. gen).

389

RESULTS

A new genus of the Diclidophorinae (Monogenea).

Neotropical Helminthology, 2017, 11(2), jul-dic

390

Figures. 1–8. Olivacotyle hemanthiasi n. gen. n. sp. (Monogenea: Diclidophoridae) from gill

laments of the damsel bass Hemanthias signifer (Garman, 1899) (Serranidae), holotype. 1.

Body, ventral view. 2. Clamp posterolateral view. 3. Clamp dorsal view. 4. Anterior ring-shape

sclerite (Ar). 5. Anterolateral sclerite (Al). 6. Posteromedial sclerite (Ps). 7. Proximal

posterolateral sclerite (Ppl). 8. Distal posterolateral sclerite (Dpl).

Cruces et al.

Olivacotyle n. gen. differs from Diclidophora

because the species of this genus have the anterior

valve of the clamp with complete lamellate

extension (lamellate extension partially reduced in

member of the new genus). In addition,

Diclidophora species have a prostatic vesicle

(absent in member of Olivacotyle n. gen.) and the

clamps are arranged in the lateral margins of the

haptor (the new genus has the clamps arranged in a

semicircle around posterior edge of haptor).

The members of Mamaevicotyle have the anterior

valve of the clamp with a plate-shape sclerite (ring-

shape sclerite in species of the new genus)

(confirmed by the study of the holotype of M.

villalobosi, CNHE 0205). In addition,

Mamaevicotyle species have two vaginal pores

(absent in member of Olivacotyle n. gen.).

In the speci e s o f D i c l i d o p h o ro i d e s ,

Mamaevodiclidophora and Macrouridophora the

axial anterolateral sclerite and anteromedial

sclerite are not fused distally (fused in Olivacotyle

n. gen.). Moreover, in Diclidophoroides species the

posterior pair of clamps is larger than the other

clamps (the clamps are similar in size in members

of the new genus). In Mamaevodiclidophora and

Macrouridophora species have a prostatic vesicle

(absent in member of Olivacotyle n. gen.).

According to Mamaev (1976), Lebboia is

considered the most primitive genus of the

Diclidophorinae, and includes species with the

morphology of the clamps different from that of

Olivacotyle n. gen. In the species of Lebboia the

anteromedial sclerite is not fused to lamellate

extension (fused in members of the new genus).

Olivacotyle n. gen. differs from species of

Upenicola, Osphyobothrus and Lampanyctophilus

(confirmed by the study of the holotype of L.

wisneri, USNM 79178) by having distal

posterolateral esclerites fused distally (not fused in

the new genus).

Olivacotyle hemanthiasi n. sp. (Figs. 1-7)

Description (based on 4 specimens mounted in

Gray and Wess medium and 22 fixed in 4% hot

formalin and stained with Semichon's carmine or

Gomori's trichrome): Body piriform, symmetrical,

not divided (Fig. 1), 0.832–1.098 (1.008; n = 22)

mm long, 226–402 (338; n = 22) maximum width

at level of ovary. Tegument smooth. Buccal suckers

oval, muscular, aseptate, lying in posterolateral

wall of buccal cavity, 35–51 (45; n = 16) long,

39–59 (51; n =16) wide. Haptor 106–196 (148; n =

12) long, 344–469 (423; n = 12) wide, with 4 pairs

of subequal slightly pedunculate clamps. Clamps

slightly asymmetrical, similar in shape (Figs. 2–3),

40–67 (56; n = 15) long, 49–69 (62; n = 15) wide;

anterior ring-shape sclerite (Ar) with proximal end

subtriangular (Fig. 4); anterolateral sclerite (Al)

strongly curved (Fig. 5); posteromedial sclerite

(Ps) fluted, rod-shaped (Fig. 6); proximal

posterolateral sclerite (Ppl) curved and widened

(Fig. 7); distal posterolateral sclerite (Dpl) arced

(Fig. 8). Anchors between first pair of clamps, two

outer pairs (Fig. 9) 7–8 (8; n = 3) long; inner pair

(Fig. 9) 19–21 (20; n = 3) long.

Mouth subterminal. Pharynx ovoid, muscular,

immediately posterior to the prohaptoral suckers

36–49 (43; n = 10) long, 37–45 (41; n = 10) wide.

Esophagus short. Intestinal bifurcation anterior to

male copulatory organ; ceca laterally diverticulate,

extending to level of third pair of clamps (Fig. 1).

Testes subspherical, 54–76 in number, intercecal

and extracecal, extending into haptor. Vas deferens

extending sinuously along median line, dorsal to

uterus. Male copulatory organ (Fig. 10) 36–49 (43;

n = 15) long, 42–52 (46; n = 12) wide, muscular,

142–221 (161; n = 10) from anterior end; armed

with 6 curved hooks (Fig. 10), with bifid base,

1318 (16; n = 10) long.

Ovary tubular, 102–153 (129; n = 7) long, 136–189

(162; n = 7) wide. Ootype surrounded by Mehlis's

gland cells. Seminal receptacle pyriform, 173–222

(195; n = 17) long, 68–101 (83; n = 17) wide,

immediately anterior to proximal portion of ovary

(Fig. 11). Genito-intestinal canal not observed.

Vitelline follicles laterally distributed, extending

from level of cecal bifurcation to haptor (Fig. 1).

Vitelline reservoir Y-shaped (Fig. 11), 167–298

(224; n =10) long. Eggs oval (Fig. 12), 156–174

(166; n = 11) long, 50–98 (79; n = 11) wide.

Taxonomic summary

Type host: Hemanthias signifer (Garman, 1899)

(Serranidae), damsel bass.

Site of infection: Gill filaments.

Neotropical Helminthology, 2017, 11(2), jul-dic

391

A new genus of the Diclidophorinae (Monogenea).

Neotropical Helminthology, 2017, 11(2), jul-dic

392

Figures. 9–12. Olivacotyle hemanthiasi n. gen. n. sp. (Monogenea: Diclidophoridae) from gill

laments of damsel bass Hemanthias signifer (Garman, 1899) (Serranidae), holotype. 9.

Anchors. 10. Male copulatory organ. 11. Detail, female reproductive system. Abbreviations: oot,

ootype; mg, Mehlis' gland; ov, ovary; ovi, oviduct; sr, seminal receptacle; ut, uterus; vr, vitelline

reservoir. 12. Egg.

Cruces et al.

Neotropical Helminthology, 2017, 11(2), jul-dic

Type locality: Puerto Pizarro, Tumbes, Peru

(45°54'S, 81°05'W), South America.

Prevalence: 4 infected fish of 7 (57%) with a total

of 52 worms.

Mean intensity of infection: 13 monogeneans per

infected fish (range 9–22).

Specimens deposited: Holotype, MUSM 3300; 4

paratypes, MUSM 3301–3304; 2 paratypes

CHIOC 38882a, b.

Etymology: The specific epithet refers to the genus

of the fish host.

The Diclidophoridae contains several species that

are still in a controversial position within the family

and their placement into subfamilies. Many of the

original descriptions of the genera are absolutely

inadequate for comparative purposes and some

type specimens are not available or are in poor

conditions. Consequently, some species initially

classified within of a genus in the Diclidophorinae

have been redescribed, or transferred to other

genera, or both (Rubec, 1989, 1991; Rubec &

Dronen, 1994). For example, species originally

accommodated in Diclidophora from macrourid

and morid fishes have been transferred to

Macrouridophora on the basis of clamp

morphology (Rubec & Dronen, 1994). Therefore,

further revision of all species referenced into the

Diclidophoridae based on morphological and

molecular data (integrative taxonomy) may be

required (Oliva et al., 2014; Tambireddy et al.,

2016).

Olivacotyle n. gen. is added to the 15 genera

currently recognized in the Diclidophorinae.

Species of these genera have been described

parasitizing marine fish of the orders Gadiformes,

Myctophiformes, Perciformes, Pleuronectiformes

and Salmoniformes from different geographical

regions, mainly in the Indian and Atlantic Ocean

(Mamaev, 1976; Payne, 1986; Rubec & Dronen,

1994). According to Rubec & Dronen (1994) the

clamp morphology is considered as a characteristic

of generic importance in diclidophorids and

warrant the separation of genera. The clamps

morphology in Olivacotyle n. gen. is readily

distinguishable from all other presently know

clamp morphology in t h e sub f a m i l y

Diclidophorinae by including species with the

anteromedial sclerite and one anterolateral sclerite

fused forming a ring, the other anterolateral sclerite

not fused with any sclerites of inner anterior

quadrant, lamellate extension partially reduced and

the distal posterolateral sclerites not fused distally.

To date, 4 species belonging to 3 genera of the

Diclidophorinae (Diclidophora, Diclidophoroides

and Tribuliphorus) have been reported from South

American, i.e, Diclidophora micromesisti Suriano

& Martorelli, 1984 from the Southern blue whiting

Micromesistius australis Norman, 1937 (Gadidae)

in Argentina; D. minor (Olsson, 1876) from an

unidentified host in Falkland-Patagonian Region;

Diclidophoroides maccallumi Price, 1943 from the

Brazilian codling Urophycis brasiliensis (Kaup,

1858) (Phycidae) in Argentina and Brazil;

Tribuliphorus salilotae Mamaev & Parukhin, 1977

from the tadpole codling Salilota australis

(Günther, 1878) (Moridae) and Salilota sp.

(Moridae) in Argentina and Falkland-Patagonian

Region, respectively (Cohen et al., 2013).

José L. Luque was supported by a research

fellowship from CNPq (Conselho Nacional de

Desenvolvimento Científico e Tecnológico,

Brazil).

393

DISCUSSION

ACKNOWLEDGEMENTS

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Publicación Especial del Instituto del Mar.

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Received September 6, 2017.

Accepted October 31, 2017.

Cruces et al.