New species of Protomicrocotyle (Monogenea: Protomicrocotylidae), and new information on P. mirabilis, parasites of Caranx spp. from Veracruz, México

Ramírez-Cruz, Edgar Salvador; Monks, Scott; Manríquez-Morán, Norma Leticia; Violante-González, Juan; Pulido-Flores, Griselda

doi:10.1590/S1984-29612023066

Edgar Salvador Ramírez-Cruz

Laboratorio de Morfología Animal, Centro de Investigaciones Biológicas, Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, México

Scott Monks

Laboratorio de Morfología Animal, Centro de Investigaciones Biológicas, Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, México

Norma Leticia Manríquez-Morán

Laboratorio de Sistemática Molecular, Centro de Investigaciones Biológicas, Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, México

Juan Violante-González

Centro de Ciencias de Desarrollo Regional, Facultad de Ecología Marina, Universidad Autónoma de Guerrero, Acapulco, Guerrero, Mexico

Griselda Pulido-Flores ^**Corresponding author: Griselda Pulido-Flores. E-mail: gpulido@uaeh.edu.mx

Laboratorio de Morfología Animal, Centro de Investigaciones Biológicas, Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, México

http://orcid.org/0000-0002-0454-8466

*Corresponding author: Griselda Pulido-Flores. E-mail: gpulido@uaeh.edu.mx

Ethics declaration

The authors declare to follow ethical standards of the Universidad Autónoma del Estado de Hidalgo for scientific publication.

Conflict of interest

The authors declare that they have no conflict of interest.

Locality (number)	Species	Number of fish collected	Infected with P. mirabilis	*Infected with P. veracruzensis* sp. nov.**	Collection date	Coordinates
Tuxpan (1)	Caranx hippos	4	4	‒	11 Mar 2010	20° 58' 09.2” N; 97° 19' 43.4 “W
		6	5	‒	12 Nov 2013
		12	8	‒	19 Aug 2019
Tecolutla (2)	C. hippos	1	‒	‒	27 May 2015	20° 29' 00.8” N; 97° 00' 07.2” W
		14	14	‒	12 Nov 2021
Casitas (3)	C. hippos	22	18	‒	14 Nov 2021	20° 15' 31.5” N; 96° 47' 49.5” W
	C. latus	2	‒	1	4 May 2005
		3	‒	‒	19 Aug 2019
Puerto de Veracruz (4)	C. hippos	8	7	‒	24 May 2022	19° 13' 11.2” N; 96° 09' 24.4” W
	C. latus	1	‒	1	24 May 2022

Species of monogeneans	Host	GenBank ID		Reference for sequences
		CO1	28S
Protomicrocotylidae
Protomicrocotyle mirabilis	Caranx hippos	OR282821– OR282832	OR282885– OR282894	Present study
Protomicrocotyle veracruzensis sp. nov.	C. latus	OR282833– OR282836	OR282895– OR282898	Present study
Neomicrocotyle pacifica (Meserve, 1938Meserve FG. Some Monogenetic trematodes from the Galapagos Islands and the Neighboring Pacific. Allan Hancock Found Publ 1938; 2: 31-72.) Yamaguti, 1968	C. hippos	–	AF382043	Olson & Littlewood (2002)Olson PD, Littlewood DTJ. Phylogenetics of the Monogenea–evidence from a medley of molecules. Int J Parasitol 2002; 32(3): 233-244. http://dx.doi.org/10.1016/S0020-7519(01)00328-9. PMid:11835967. http://dx.doi.org/10.1016/S0020-7519(01)...
Neomicrocotyle sp.	C. sexfasciatus	–	KF378589	Justine et al. (2013)Justine J-L, Rahmouni C, Gey D, Schoelinck C, Hoberg EP. The monogenean which lost its clamps. PLoS One 2013; 8(11): e79155. http://dx.doi.org/10.1371/journal.pone.0079155. PMid:24278118. http://dx.doi.org/10.1371/journal.pone.0...
Lethacotyle vera Justine, Rahmouni, Gey, Schoelinck and Hoberg, 2013^* * Reported as Lethacotyle sp. in GenBank.	C. papuensis	–	KF378588	Justine et al. (2013)Justine J-L, Rahmouni C, Gey D, Schoelinck C, Hoberg EP. The monogenean which lost its clamps. PLoS One 2013; 8(11): e79155. http://dx.doi.org/10.1371/journal.pone.0079155. PMid:24278118. http://dx.doi.org/10.1371/journal.pone.0...
Bilaterocotyloides carangis Ramalingam, 1961Ramalingam K. A new genus of trematode Bilateracotyloides (Monogenea) from the gills of Caranx rottleri (Bloch). Zool Anz 1961; 166(7-8): 286-290.	Megalaspis cordyla	KF804043	–	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...
		–	KJ201022	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...
Bilaterocotyloides madrasensis Radha, 1966Radha E. On a new species of Bilaterocotyloides (Monogenea) from gills of Caranx rottleri. Zool Anz 1966; 177(S): 311-316.	M. cordyla	KF804041	KF804029	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...
		–	KF804037	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...
Outgroup
Allodiscocotylidae
Allodiscocotyla diacanthi Unnithan, 1962Unnithan RV. On the functional morphology of a new fauna of Monogenoidea on fishes from Trivandrum and environs Part II. Opisthogynidae fam.nov. (Gastrocotyloidea) and Abortipedinae subfam.nov. (Protomicrocotyloidea). Parasitology 1962; 52(3-4): 315-351. http://dx.doi.org/10.1017/S0031182000027190. http://dx.doi.org/10.1017/S0031182000027...	Scomberoides commersonnianus	KF804045	KF804033	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...
		KF804046	KF804038	Tambireddy et al. (2016)Tambireddy N, Gayatri T, Gireesh-Babu P, Pavan-Kumar A. Molecular characterization and phylogeny of some mazocraeidean monogeneans from carangid fish. Acta Parasitol 2016; 61(2): 360-368. http://dx.doi.org/10.1515/ap-2016-0047. PMid:27078660. http://dx.doi.org/10.1515/ap-2016-0047...

Species	*Protomicrocotyle mirabilis*	*P. mirabilis*	*P. mirabilis*
Locality	Veracruz (this study)	Florida Bay, Everglades National Park, Florida	New York Aquarium, USA
Host	*C. hippos*	*Caranx hippos*	*C. hippos*
Site	Gill filaments	Gill lamellae	Gill lamellae
	µm	µm	µm
Total length	2861 (805–4673, n =79)	2240 (1470–3080)	3128
Total width	295 (134–537, n =80)	322 (214–449)	200
Mouth length	26 (11–52, n =70)	16	‒
Mouth width	31 (12–67, n =73)	25	‒
Cecal bifurcation to anterior end	606 (380–775, n =66)	346	‒
Pharynx length	40 (14–64, n =77)	32	‒
Pharynx width	35 (14–52, n =77)	47 (39–57)	‒
Prohaptoral sucker length (right)	44 (13–68, n =77)	23	31
Prohaptoral sucker width (right)	31 (10–48, n =77)	32	22
Prohaptoral sucker length (left)	44 (14–67, n =77)	24	30
Prohaptoral sucker width (left)	31 (11–48, n =77)	33	17
Clamp length	45.7 (14.2‒65, n =71 )^* * Average of all anatomical structures measured.	57 (49–67)	55
Haptor length	477 (165–815, n =76)	338
Haptor width	255 (70–425, n =76)	161	110
Right side of haptoral lappet length	117 (60–217, n =72)	64	73
Left side of haptoral lappet length	120 (40–202, n =73)	72	61
Central haptoral lappet length	125 (40–214, n =74)	88	134
Haptoral lappet width	437 (105–785, n =75)	241	330
Lateral anchor length	41.5 (22.5‒65, n = 153)*	33 (29–37)	‒
Lateral anchor width	5 (2‒7.5, n = 153)*	3	‒
Medial anchor length	27.5 (9–37, n = 151)*	21 (19–22)	‒
Medial anchor width	4 (2–6, n = 151)*	2	‒
Hook length	17 (7–23, n = 144)*	18 (16–19)	‒
Hook width	3 (1–3, n = 138)*	2	‒
Number of testes	31 (21–38, n =77)	27 (23–33)	14–35
Testes length	45 (17–79, n =81)	43 (28–54)	34.6
Testes width	50 (14–86, n =80)	43 (28–54)	49.3
Testes-Ratio length: wide	1:1.1	1:1	1:1.4
Male copulatory organ length	54 (22–82, n =80)	77	‒
Male copulatory organ width	41 (11–66, n =80)	50 (42–56)	‒
Genital pore to anterior end distance	370 (140–660, n =81)	250	452
Vaginal vestibule length	69 (28–94, n =80)	82 (66–99)	52.5
vaginal vestibule width	51 (19–77, n =80)	59 (41–75)	48
Vaginal opening to anterior end	464 (175–740, n =80)	330	567
Vaginal opening to genital pore	112 (40–180, n =80)	88	134
Egg length	157 (88–206, n =41)	180 (167–193)	183
Egg width	49 (24–68, n =41)	68 (66–70)	85.5
Number of spines in male copulatory organ (MCO)	22 (16–29, n =71)	19 (16–21)	20
Number of spines in vaginal vestibule	45 (31–65, n =63)	49	‒
Reference	Present study	Kritsky et al. (2011)Kritsky DC, McAleese WJ, Bakenhaster MD. Heteronchoineans (Monogenoidea) from the gills of crevalle jack, Caranx hippos (Perciformes, Carangidae), from Everglades National Park, Florida, with a redescription of Protomicrocotyle mirabilis (Gastrocotylinea, Protomicrocotylidae). Comp Parasitol 2011; 78(2): 265-274. http://dx.doi.org/10.1654/4505.1. http://dx.doi.org/10.1654/4505.1...	MacCallum (1918)MacCallum GA. Notes on the genus Telorchis and other trematodes. Zoopathologica 1918; 1(3): 81-97.

Species	*P. mirabilis*	P. cf. mirabilis	P. cf. mirabilis
Locality	Tuxpan, Veracruz	Sontecomapan, Veracruz, Mexico	Las Cabanas, Veracruz, Mexico
Host	*C. latus*	*C. hippos*	*C. hippos*
Site	Gill lamellae	Gill lamellae	Gill lamellae
	µm	µm	µm
Total length	1254‒2451	1482‒3706	1881‒2052
Total width	149‒387	171‒399	285‒342
Mouth length	19	‒	‒
Mouth width	44	‒	‒
Cecal bifurcation to anterior end	375	‒	‒
Pharynx length	43	‒	‒
Pharynx width	44	‒	‒
Prohaptoral sucker length (right)	44	‒	‒
Prohaptoral sucker width (right)	29	‒	‒
Prohaptoral sucker length (left)	44	‒	‒
Prohaptoral sucker width (left)	27	‒	‒
Clamp length	48‒52	‒	‒
Haptor length	342	‒	‒
Haptor width	149‒298	‒	‒
Right side of haptoral lappet length	49	‒	‒
Left side of haptoral lappet length	44	‒	‒
Central haptoral lappet length	71	‒	‒
Haptoral lappet width	283	‒	‒
Lateral anchor length	‒	‒	‒
Lateral anchor width	‒	‒	‒
Medial anchor length	‒	‒	‒
Medial anchor width	‒	‒	‒
Hook length	‒	‒	‒
Hook width	‒	‒	‒
Number of testes	20‒30	18‒29	17‒26
Testes length	37‒41	‒	‒
Testes width	33‒59	‒	‒
Testes-Ratio length: wide	1:0.8	‒	‒
Male copulatory organ length	49	‒	‒
Male copulatory organ width	38	‒	‒
Genital pore to anterior end distance	231	‒	‒
Vaginal vestibule length	88	‒	‒
vaginal vestibule width	60	‒	‒
Vaginal opening to anterior end	268‒402	‒	‒
Vaginal opening to genital pore	77	‒	‒
Egg length	‒	‒	‒
Egg width	‒	‒	‒
Number of spines in male copulatory organ (MCO)	18‒20	17‒19	16‒17
Number of spines in vaginal vestibule	‒	‒	‒
Reference	Caballero y Caballero & Bravo-Hollis (1965)Caballero y Caballero E, Bravo-Hollis M. Monogenea (van Beneden, 1858) Carus, 1863 de peces marinos del litoral mexicano del Golfo de México y del Mar Caribe. II. Rev Biol Trop 1965; 13(1): 101-121.	Bravo-Hollis (1989)Bravo-Hollis M. Monogenea (Van Beneden, 1858) Carus 1863, de peces del Golfo de México y del mar Caribe XII. Nuevas localidades de colecta de especies conocidas de gastrocotilidos. An Inst Biol Univ Nac 1989; 59(1): 1-14.	Bravo-Hollis (1989)Bravo-Hollis M. Monogenea (Van Beneden, 1858) Carus 1863, de peces del Golfo de México y del mar Caribe XII. Nuevas localidades de colecta de especies conocidas de gastrocotilidos. An Inst Biol Univ Nac 1989; 59(1): 1-14.

Species	P. cf. mirabilis	P. cf. mirabilis	P. cf. mirabilis	P. cf. mirabilis	P. cf. mirabilis
Locality	Chetumal, Quintana Roo, Mexico	Isla Cozumel, Quintana Roo, Mexico	Isla Mujeres, Quintana Roo, Mexico	Port Aransas, Texas, USA	Ebrié Lagon, near Abidjan
Host	*C. latus*	Caranx sp.	*C. crysos*	*C. hippos*	*C. hippos*
Site	Gill lamellae	Gill lamellae	Gill lamellae	Gill lamellae	Gill lamellae
				µm	µm
Total length	1938‒3192	2565‒3078	2223‒2280	5400	2000‒2900
Total width	171‒285	228‒399	470‒620	275	250‒550
Mouth length	‒	‒	‒	‒	‒
Mouth width	‒	‒	‒	‒	‒
Cecal bifurcation to anterior end	‒	‒	‒	‒	345
Pharynx length	‒	‒	‒	‒	‒
Pharynx width	‒	‒	‒	‒	‒
Prohaptoral sucker length (right)	‒	‒	‒	‒	‒
Prohaptoral sucker width (right)	‒	‒	‒	‒	‒
Prohaptoral sucker length (left)	‒	‒	‒	‒	‒
Prohaptoral sucker width (left)	‒	‒	‒	‒	‒
Clamp length	‒	‒	‒	60	60
Haptor length	‒	‒	‒	‒	138
Haptor width	‒	‒	‒	‒	‒
Right side of haptoral lappet length	‒	‒	‒	‒	166
Left side of haptoral lappet length	‒	‒	‒	‒	178
Central haptoral lappet length	‒	‒	‒	‒	154
Haptoral lappet width	‒	‒	‒	200	416
Lateral anchor length	‒	‒	‒	‒	‒
Lateral anchor width	‒	‒	‒	‒	‒
Medial anchor length	‒	‒	‒	‒	‒
Medial anchor width	‒	‒	‒	‒	‒
Hook length	‒	‒	‒	‒	‒
Hook width	‒	‒	‒	‒	‒
Number of testes	20‒25	20‒21	24‒25	‒	28‒32
Testes length	‒	‒	‒	‒	75‒85
Testes width	‒	‒	‒	‒	45‒55
Testes-Ratio length: wide	‒	‒	‒	‒	1:0.6
Male copulatory organ length	‒	‒	‒	‒	107
Male copulatory organ width	‒	‒	‒	‒	71
Genital pore to anterior end distance	‒	‒	‒	‒	309
Vaginal vestibule length	‒	‒	‒	‒	71
vaginal vestibule width	‒	‒	‒	‒	95
Vaginal opening to anterior end	‒	‒	‒	‒	300
Vaginal opening to genital pore	‒	‒	‒	‒	‒
Egg length	‒	‒	‒	‒	‒
Egg width	‒	‒	‒	‒	‒
Number of spines in male copulatory organ (MCO)	18‒19	16‒20	18‒19	‒	15‒20
Number of spines in vaginal vestibule	‒	‒	‒	‒	‒
Reference	Bravo-Hollis (1989)Bravo-Hollis M. Monogenea (Van Beneden, 1858) Carus 1863, de peces del Golfo de México y del mar Caribe XII. Nuevas localidades de colecta de especies conocidas de gastrocotilidos. An Inst Biol Univ Nac 1989; 59(1): 1-14.	Bravo-Hollis (1989)Bravo-Hollis M. Monogenea (Van Beneden, 1858) Carus 1863, de peces del Golfo de México y del mar Caribe XII. Nuevas localidades de colecta de especies conocidas de gastrocotilidos. An Inst Biol Univ Nac 1989; 59(1): 1-14.	Bravo-Hollis (1989)Bravo-Hollis M. Monogenea (Van Beneden, 1858) Carus 1863, de peces del Golfo de México y del mar Caribe XII. Nuevas localidades de colecta de especies conocidas de gastrocotilidos. An Inst Biol Univ Nac 1989; 59(1): 1-14.	Koratha (1955)Koratha KJ. Studies on the monogenetic trematodes of the Texas Coast. II. Descriptions of species from marine fishes of Port Aransas. Int Mar Sci 1955; 4(1): 251-278.	Wahl (1972)Wahl EJ. Protomicrocotyle mirabilis (Mac Callum 1918) Johnston et Tiegs 1922 et P. ivoriensis n. spec., Monogènes parasites de Caranx hippos dans la lagune Ebrié (Côte d’Ivoire). Z Parasitenkd 1972; 38(4): 319-332. http://dx.doi.org/10.1007/BF00329279. http://dx.doi.org/10.1007/BF00329279...

Species	Protomicrocotyle veracruzensis sp. nov.	*P. mirabilis* (redescription)
Hosts	Caranx latus	C. hippos and C. latus
Location	Casitas and Puerto de Veracruz, Veracruz, México	Florida, EUA and Veracruz, Mexico
Total length	3858 (3209‒4709)	805‒4673
Total Width	804 (586‒1098)	134‒537
Haptor width	575 (415‒980)	70‒425
Right side of haptoral lappet length	173 (136‒240)	60‒217
Left side of haptoral lappet length	194 (124‒259)	40‒202
Central haptoral lappet length	206 (164‒259)	40‒214
Haptoral lappet width	769 (580‒920)	105‒785
Clamps length	64 (47‒74)^* * Data obtained from the average length of the four clamps.	14.2‒67
Number of testes	47 (36‒69)	14‒38
Testes length	47 (31‒59)	17‒79
Testes width	142 (96‒176)	14‒86
Testes-Ratio length: width	01:03	1:0.8‒1:1.1
Vaginal opening to anterior end	610 (440‒730)	175‒740
Vaginal opening to genital pore	506 (340‒615)	140‒660
Number of spines in MCO	23 (16‒28)	16‒29
Number of spines in vestibule vaginal	49 (33‒59)	31‒65
Reference	Present study	Present study

Species	*P. manteri*	*P. nayaritensis*	*P. ivoriensis*
Hosts	Trachinotus paitensis	C. caninus	C. hippos
Location	La Paz, BCS, Mexico	Nayarit, Mexico	Ivory Coast
Total length	1360‒3680	6820‒8580	3200‒4600
Total Width	400‒800	770‒1990	600‒1200
Haptor width	590	1050	442
Right side of haptoral lappet length	136	250	173
Left side of haptoral lappet length	121	150	178
Central haptoral lappet length	181	300	134
Haptoral lappet width	348	643‒801	500
Clamps length	50‒55	45‒48	44
Number of testes	22‒33	46‒48	50
Testes length	60.2	88	58.8
Testes width	116.5	129	99.5
Testes-Ratio length: width	01:01.8	01:01.4	01:01.6
Vaginal opening to anterior end	84‒155	1072‒1258	442
Vaginal opening to genital pore	289‒471	858‒1001	365
Number of spines in MCO	33‒38	48‒54	25‒26
Number of spines in vestibule vaginal	15	22‒46	‒
Reference	Bravo-Hollis (1966)Bravo-Hollis M. Helmintos de peces del Pacífico mexicano XXV. Descripción de tres monogéneos del Golfo de California. An Inst Biol Univ Nac 1966; 37(1-2): 107-123.	Bravo-Hollis (1979)Bravo-Hollis M. Monogéneos de peces de la colección Winter II. Sobre un protomicrocotylido nuevo de Caranx hippos caninus Gunther del Pacífico mexicano. An Inst Cienc Mar Limnol [internet]. 1979 [cited 2023 Sep 27]; 6(1): 189-196. Available from: http://biblioweb.tic.unam.mx/cienciasdelmar/centro/1979-1/articulo64.html http://biblioweb.tic.unam.mx/cienciasdel...	Wahl (1972)Wahl EJ. Protomicrocotyle mirabilis (Mac Callum 1918) Johnston et Tiegs 1922 et P. ivoriensis n. spec., Monogènes parasites de Caranx hippos dans la lagune Ebrié (Côte d’Ivoire). Z Parasitenkd 1972; 38(4): 319-332. http://dx.doi.org/10.1007/BF00329279. http://dx.doi.org/10.1007/BF00329279...

Species	*P. celebensis*	*P. madrasensis*	*P. mannarensis*
Hosts	Caranx sp.	C. affinis	C. sexfasciatus
Location	Indonesia	India	India
Total length	4000‒5100	2390	2880‒3640
Total Width	600‒830	410	480‒1140
Haptor width	‒	300	434
Right side of haptoral lappet length	111	250	130
Left side of haptoral lappet length	111	250	152
Central haptoral lappet length	166	112	213
Happtoral lappet width	‒	240	320‒440
Clamps length	‒	32‒48	30‒46
Number of testes	40‒55	64	42‒52
Testes length	‒	33‒38	21‒33
Testes width	‒	58‒81	67‒181
Testes-Ratio length: width	‒	1:1.7-1:2.1	1:3-1:5.4
Vaginal opening to anterior end	500‒830	462	526
Vaginal opening to genital pore	375‒630	362	447
Number of spines in MCO	16‒22	24	24
Number of spines in vestibule vaginal	‒	31	37
Reference	Yamaguti (1953)Yamaguti S. Parasitic worms mainly from Celebes. Part 2. Monogenetic trematodes of fishes. Acta Med Okayama 1953; 8(3): 203-265. http://dx.doi.org/10.18926/AMO/31878. http://dx.doi.org/10.18926/AMO/31878...	Ramalingam (1960)Ramalingam K. Morphological descriptions of a new genus Neomicrocotyle and three new species of the genus Protomicrocotyle (Monogenea) with a discussion on their taxonomic position. Proc Nat Inst Sci India B Biol Sci 1960; 26(6): 367-378.	Ramalingam (1960)Ramalingam K. Morphological descriptions of a new genus Neomicrocotyle and three new species of the genus Protomicrocotyle (Monogenea) with a discussion on their taxonomic position. Proc Nat Inst Sci India B Biol Sci 1960; 26(6): 367-378.

Species	*P. minutum*	*P. carangis*
Hosts	C. sexfasciatus	C. sansun
Location	India	India
Total length	1500	2540‒3100
Total Width	300	460‒540
Haptor width	239	‒
Right side of haptoral lappet length	65	‒
Left side of haptoral lappet length	65	‒
Central haptoral lappet length	119	‒
Haptoral lappet width	360	320‒410
Clamps length	52	48
Number of testes	34	20‒24
Testes length	14	‒
Testes width	34‒44	‒
Testes-Ratio length: width	1:2.4-1:3.1	‒
Vaginal opening to anterior end	304	‒
Vaginal opening to genital pore	260	‒
Number of spines in MCO	12	22
Number of spines in vestibule vaginal	19	14‒16
Reference	Ramalingam (1960)Ramalingam K. Morphological descriptions of a new genus Neomicrocotyle and three new species of the genus Protomicrocotyle (Monogenea) with a discussion on their taxonomic position. Proc Nat Inst Sci India B Biol Sci 1960; 26(6): 367-378.	Ramalingam (1960)Ramalingam K. Morphological descriptions of a new genus Neomicrocotyle and three new species of the genus Protomicrocotyle (Monogenea) with a discussion on their taxonomic position. Proc Nat Inst Sci India B Biol Sci 1960; 26(6): 367-378.

	Right lateral anchor (Ac1)	Left lateral anchor (Ac1’)	Right medial anchor (Ac2)	Left medial anchor (Ac2’)	Right central hook (Ho)	Left central hook (Ho’)
Total length (a)	43 (38‒50, n = 24)	42 (40‒47, n = 24)	27 (25‒29, n = 20)	27 (24‒29, n = 20)	18 (17‒19, n = 22)	18 (17‒19, n = 22)
Total width (b)	5 (5‒7, n = 24)	6 (5‒7, n = 24)	4 (2‒5, n = 22)	3 (2‒4, n = 20)	2 (2‒2, n = 22)	2 (2‒2, n = 22)
Opening length (c)	10 (7‒13, n = 24)	10 (7‒13, n = 23)	5 (4‒8, n = 22)	6 (4‒11, n = 20)	-	-
Internal root length (d)	16 (11‒19, n = 24)	15 (11‒17, n = 24)	7 (5‒11, n = 22)	9 (7‒12, n = 20)	-	-
Internal root width (e)	7 (5‒10, n = 24)	7 (5‒8, n = 24)	3 (2‒4, n = 22)	3 (2‒5, n = 20)	-	-
External root length (f)	16 (13‒20, n = 24)	15 (12‒19, n = 24)	9 (7‒13, n = 22)	9 (7‒11, n = 20)	-	-
External blade length (g)	23 (18‒29, n = 24)	24 (17‒34, n = 24)	16 (13‒19, n = 22)	18 (14‒20, n = 20)	-	-
Internal blade length (h)	20 (16‒25, n = 24)	20 (13‒31, n = 24)	13 (11‒17, n = 22)	15 (12‒20, n = 20)	-	-
Point length (i)	13 (5‒19, n = 24)	15 (8‒24, n = 24)	9 (5‒12, n = 22)	11 (8‒13, n = 20)	-	-

A
Principal Component	Eigenvalue	% variance	% Accumulate variance
1	0.81	44.10	44.10
2	0.16	8.651	52.75

B
Factor	Eigenvalue	% variance	% Accumulate variance
1	55.73	46.88	46.88
2	49.49	41.64	88.52

A
	KF804045	KF804046	KF804041	KF804043	OR282821^* * Sequences of Protomicrocotyle mirabilis;	OR282822*	OR282823*	OR282824*
KF804045
KF804046	3.55%
KF804041	19.86%	22.34%
KF804043	18.09%	20.21%	12.41%
OR282821*	20.21%	23.05%	16.67%	15.25%
OR282822*	20.21%	23.05%	16.31%	14.89%	1.00%
OR282823*	20.21%	23.05%	16.31%	14.89%	1.24%	0.74%
OR282824*	20.92%	23.76%	16.31%	14.89%	0.77%	1.28%	1.28%
OR282825*	20.21%	23.05%	16.31%	14.89%	0.50%	0.50%	0.74%	0.77%
OR282826*	20.21%	23.05%	16.31%	15.25%	0.75%	0.75%	1.00%	1.02%
OR282827*	20.21%	23.05%	16.31%	14.89%	0.50%	0.50%	0.75%	0.77%
OR282828*	20.21%	23.05%	16.31%	15.25%	1.00%	0.99%	1.24%	1.28%
OR282829*	20.57%	23.40%	16.67%	15.25%	1.00%	0.99%	1.24%	0.77%
OR282830*	20.21%	23.05%	16.31%	14.89%	0.75%	0.74%	0.99%	1.02%
OR282831*	19.86%	22.70%	16.67%	14.54%	1.00%	1.00%	1.25%	1.29%
OR282832*	20.92%	23.76%	16.31%	15.60%	1.26%	1.75%	2.01%	1.03%
OR282833^ Sequences of Protomicrocotyle veracruzensis sp. nov.	20.57%	23.40%	15.60%	16.31%	9.20%	8.71%	9.45%	9.21%
OR282834**	20.92%	23.76%	15.25%	16.67%	9.45%	8.68%	9.43%	9.46%
OR282835**	20.92%	23.76%	15.25%	16.67%	8.96%	8.68%	9.43%	8.95%
OR282836**	20.92%	23.76%	15.25%	16.67%	9.20%	8.93%	9.68%	9.21%

	OR282825*	OR282826*	OR282827*	OR282828*	OR282829*	OR282830*	OR282831*	OR282832*
OR282826*	0.75%
OR282827*	0.00%	0.75%
OR282828*	0.99%	0.25%	1.00%
OR282829*	0.99%	0.75%	1.00%	0.74%
OR282830*	0.74%	0.50%	0.75%	0.50%	0.50%
OR282831*	0.50%	1.26%	0.50%	1.00%	1.25%	1.00%
OR282832*	1.25%	1.52%	1.26%	1.50%	0.75%	1.25%	1.50%
OR282833**	8.71%	9.52%	8.73%	9.70%	9.70%	9.45%	9.27%	10.05%
OR282834**	9.18%	9.27%	8.98%	9.68%	9.68%	9.43%	9.75%	10.53%
OR282835**	8.68%	9.27%	8.48%	9.68%	9.68%	9.43%	9.25%	10.03%
OR282836**	8.93%	9.52%	8.73%	9.93%	9.93%	9.68%	9.50%	10.28%

	OR282833**	OR282834**
OR282834**	0.75%
OR282835**	0.75%	0.50%
OR282836**	0.75%	0.74%	0.25%

B
	KF804033	KF804038	KF804029	KF804037	KJ201022	AF382043	KF378589	KF378588
KF804033
KF804038	0.00%
KF804029	5.25%	5.25%
KF804037	5.25%	5.25%	0.00%
KJ201022	5.29%	5.29%	0.44%	0.44%
AF382043	6.06%	6.06%	2.42%	2.42%	2.94%
KF378589	5.93%	5.93%	2.20%	2.20%	2.79%	0.14%
KF378588	6.21%	6.21%	3.03%	3.03%	3.38%	1.79%	1.65%
OR282885*	5.25%	5.25%	2.55%	2.55%	3.23%	2.69%	2.48%	2.20%
OR282886*	5.14%	5.14%	2.43%	2.43%	3.10%	2.83%	2.63%	2.35%
OR282887*	5.26%	5.26%	2.56%	2.56%	3.24%	2.69%	2.48%	2.21%
OR282888*	5.26%	5.26%	2.56%	2.56%	3.24%	2.70%	2.48%	2.21%
OR282889*	5.15%	5.15%	2.44%	2.44%	3.08%	2.84%	2.62%	2.34%
OR282890*	5.17%	5.17%	2.45%	2.45%	3.09%	2.85%	2.62%	2.34%
OR282891*	5.31%	5.31%	2.59%	2.59%	3.23%	2.72%	2.48%	2.20%
OR282892*	5.27%	5.27%	2.56%	2.56%	3.23%	2.70%	2.48%	2.20%
OR282893*	5.20%	5.20%	2.46%	2.46%	3.10%	2.87%	2.63%	2.35%
OR282894*	5.16%	5.16%	2.44%	2.44%	3.09%	2.85%	2.62%	2.34%
OR282895**	5.38%	5.38%	2.96%	2.96%	3.67%	2.82%	2.62%	2.48%
OR282896**	5.38%	5.38%	2.96%	2.96%	3.67%	2.82%	2.62%	2.48%
OR282897**	5.38%	5.38%	2.96%	2.96%	3.67%	2.82%	2.62%	2.48%
OR282898**	5.41%	5.41%	2.97%	2.97%	3.68%	2.83%	2.62%	2.48%


OR282886*	0.13%
OR282887*	0.00%	0.13%
OR282888*	0.00%	0.14%	0.00%
OR282889*	0.14%	0.00%	0.14%	0.14%
OR282890*	0.14%	0.00%	0.14%	0.14%	0.00%
OR282891*	0.00%	0.14%	0.00%	0.00%	0.14%	0.14%
OR282892*	0.00%	0.14%	0.00%	0.00%	0.14%	0.14%	0.00%
OR282893*	0.14%	0.00%	0.14%	0.14%	0.00%	0.00%	0.14%	0.14%
OR282894*	0.14%	0.00%	0.14%	0.14%	0.14%	0.14%	0.14%	0.14%
OR282895**	0.81%	0.94%	0.81%	0.81%	0.95%	0.95%	0.82%	0.81%
OR282896**	0.81%	0.94%	0.81%	0.81%	0.95%	0.95%	0.82%	0.81%
OR282897**	0.81%	0.94%	0.81%	0.81%	0.95%	0.95%	0.82%	0.81%
OR282898**	0.81%	0.95%	0.81%	0.81%	0.95%	0.95%	0.82%	0.81%

	OR282893*	OR282894*	OR282895**	OR282896**	OR282897**
OR282894*	0.14%
OR282895**	0.96%	0.95%
OR282896**	0.96%	0.95%	0.00%
OR282897**	0.96%	0.95%	0.00%	0.00%
OR282898**	0.96%	0.95%	0.00%	0.00%	0.00%

[1] *Corresponding author: Griselda Pulido-Flores. E-mail: gpulido@uaeh.edu.mx

Brasil

Brasil

New species of Protomicrocotyle (Monogenea: Protomicrocotylidae), and new information on P. mirabilis, parasites of Caranx spp. from Veracruz, México

Uma nova espécie de Protomicrocotyle (Monogenea: Protomicrocotylidae), e novas informações sobre P. mirabilis, parasitos de Caranx spp. de Veracruz, México

Abstract