taxonID	type	description	language	source
0399878EFF90E6691A96FC67FAC9FB18.taxon	discussion	Family Anthocephaliidae, order Rhinebothriidea	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	description	(Figs. 3 A, 4 A – C, 5 A – E, 6 A – B)	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	biology_ecology	Type host: Hypanus guttatus (Bloch & Schneider) – Longnose stingray.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	materials_examined	Type locality: Coast of Maceió, Alagoas, Brazil (09 ° 40 ’ 25.32 ” S, 035 ° 44 ’ 07.08 ” W). Additional locality: Colares, Pará, Brazil (01 ◦ 00 ’ 33.88 ” S, 048 ◦ 16 ’ 47.28 ” W).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	biology_ecology	Site of infection: Spiral intestine. Prevalence of infection: 44.9 % (31 of 69 valves).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	materials_examined	Specimens deposited: Holotype (MZUSP 8076) and 11 paratypes (MZUSP 8054 a – e, 8055, 8056 a – b, 8057, 8058, 8059); 12 paratypes (HWML 217938 – 217949); and, 12 paratypes (LRP 11270 – 11281).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	etymology	Etymology: This species honors Dr. Denis Jacob Machado for his contribution to the systematics of cestodes through his expertise in bioinformatics.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	description	Description. [Based on 37 specimens: 29 whole mounts of mature worms, 2 scolexes prepared for SEM and cross sections of 6 mature proglottids]: Worms apolytic, 5.8 – 13.6 mm (n = 29) long with 20 – 45 proglottids (n = 29) in number, maximum width 439.5 – 761 (n = 10) at level of scolex (Fig. 3 A). Scolex (Figs. 4 B, 5 A) with four stalked bothridia, each one with 76 – 96 (n = 10) marginal loculi and one oval apical sucker; apical sucker 33.6 – 45.1 (n = 4) long by 29.5 – 32.7 (n = 4) wide. Short cephalic peduncle present. Proximal surfaces of marginal loculi next to the bothridial rims covered with acicular filitriches and scolopate spinithriches (Fig. 5 B, C); bothridial rims and proximal surface of marginal loculi covered with acicular filitriches (Fig. 5 D); distal surfaces of bothridia covered with capiliform filitriches (Fig. 5 E). Proglottids slightly craspedote. Immature proglottids wider than long, becoming longer than wide with maturity, 16 – 42 in number (n = 29) (Fig. 3 A). Mature proglottids (Fig. 4 A) 1,066 – 1,749.6 (n = 15) long by 125.2 – 225.3 (n = 15) wide, 1 – 4 in number (n = 29). Testes arranged in two columns from near anterior margin of proglottid to level of genital pore, and 1 row-deep in cross-section (Figs. 4 A, 6 A). Testes 35.5 – 72.5 (n = 17) long by 28.1 – 55.8 (n = 17) wide, 35 – 46 (n = 17) in number. Cirrus-sac pyriform, bent posteriorly, 75.8 – 121.3 (n = 8) long by 43.8 – 112.5 (n = 8) wide, containing coiled cirrus (Fig. 4 C). Cirrus armed with spinitriches. Genital pores lateral, irregularly alternating, 27.6 – 35.5 % (n = 15) of proglottid length from posterior end (Fig. 3 A, 4 A). Ovary near posterior end of proglottid, H-shaped in frontal view (Figs. 4 A), tetralobed in cross-section (Fig. 6 B), slightly asymmetrical, 229.6 – 432 (n = 16) long by 68 – 165.8 (n = 16) wide; ovarian bridge near middle of ovary. Mehli’s gland immediately posterior to ovarian bridge. Vagina thick-walled, sinuous, extending medially in proglottid from ootype to anterior margin of cirrus-sac, then laterally to open into genital atrium anterior to cirrus (Fig. 4 C). Vitellarium follicular; vitelline follicles 13.6 – 33.1 (n = 17) long by 6.5 – 17.3 (n = 17) wide, arranged in two lateral bands, each band consisting of one dorsal and one ventral column of vitelline follicles, extending from near anterior margin of the anterior-most row of testes to near posterior margin of proglottid, interrupted by genital pore and ovary both dorsally and ventrally (Figs. 4 A, C). Uterus saccate, ventral, extending along median line of proglottid from near the ovarian bridge to posterior margin of first or second anterior-most row of testes (Fig. 4 A).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF90E6691A96FC67FAC9FB18.taxon	discussion	Remarks: Anthocephalum machadoi n. sp. is distinguishable among the 24 other species of the genus Anthocephalum by its distinct combination of morphological features which include: 35 – 46 testes, 76 – 96 loculi, a total length of 5.8 – 13.6 mm, and vitelline follicles organized into two lateral bands with one dorsal and one ventral column each. In comparison with Anthocephalum blairi Herzog & Jensen, 2018, Anthocephalum gravisi Herzog & Jensen, 2018, Anthocephalum haroldsoni Herzog & Jensen, 2018, and Anthocephalum papefayei, the new species possesses more marginal loculi (76 – 96 vs. 65 – 73, 43 – 52, 41 – 57, and 45 – 60; respectively). On the other hand, Anthocephalum machadoi n. sp. posses fewer marginal loculi compared to Anthocephalum gracile (Wedl, 1855) Ruhnke, 1994, Anthocephalum healyae Ruhnke, Caira & Cox, 2015, Anthocephalum lukei Ruhnke and Seaman, 2009, Anthocephalum meadowsi Ruhnke, Caira & Cox, 2015, Anthocephalum odonnellae Ruhnke, Caira & Cox, 2015, Anthocephalum philruschi Ruhnke, Caira & Cox, 2015, Anthocephalum cairae, and Anthocephalum hobergi (Zamparo, Brooks & Barriga, 1999) Marques & Caira, 2016 (76 – 96 vs. 116 – 121, 150 – 171, 107 – 138, 98 – 134, 135 – 159, 200 – 219, 197 – 198, and 100; respectively). Anthocephalum machadoi n. sp. differs from Anthocephalum duszynskii Ruhnke, 1994 and Anthocephalum mounseyi Herzog & Jensen, 2018, in its total length (5.8 – 13.6 mm vs. 18 – 31 and 2.6 – 3.4 mm, respectively), and from Anthocephalum decrisantisorum Ruhnke, Caira & Cox, 2015, Anthocephalum jensenae Ruhnke, Caira & Cox, 2015, Anthocephalum ruhnkei Herzog & Jensen, 2018, and Anthocephalum centrurum, in number of testes (35 – 46 vs. 17 – 24, 14 – 20, 22 – 34, and 48 – 78, respectively). In comparison with Anthocephalum alicae, A. machadoi n. sp. possesses more proglottids in number (20 – 45 vs. 9 – 15, respectively). There are only five species — Anthocephalum currani Ruhnke & Seaman, 2009, Anthocephalum michaeli Ruhnke & Seaman, 2009, Anthocephalum jeancadenati Boudaya, Neifar & Euzet, 2018, Anthocephalum kingae (Schmidt, 1978) Ruhnke & Seaman, 2009, and Anthocephalum mattisi — that closely resembles A. machadoi n. sp. in number of marginal loculi, number of testes, total length, and proglottid number. However, they can be distinguished based on the position and number of the vitelline follicles. Anthocephalum machadoi n. sp. possesses vitelline follicles organized into two lateral bands with single dorsal and ventral column. In contrast, A. michaeli, A. jeancadenati, A. kingae, and A. mattisi also possess vitelline follicles in two lateral bands but with 2 – 3 dorsal and ventral columns. Finally A. currani possesses vitelline follicles in two lateral bands with 3 – 5 dorsal and ventral columns.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	discussion	Family Anthocephaliidae, order Rhinebothriidea	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	description	(Figs. 3 B, 4 D – F, 5 F – J, 6 C – D)	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	biology_ecology	Type host: Hypanus longus (Garman) — Longtail stingray.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	materials_examined	Type locality: Coast of Cebaco Island, Veraguas, Panama (7 ◦ 29 ’ 37.9 ” N, 81 ◦ 13 ’ 21.9 ” W). Additional locality: Coast of Mariato, Veraguas, Panama (7 ◦ 36 ’ 06.3 ” N, 80 ◦ 59 ’ 33.7 ” W).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	biology_ecology	Site of infection: Spiral intestine. Prevalence of infection: 50 % (four of eight valves).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	materials_examined	Specimens deposited: Holotype and one paratype (MIUP H 0086); three paratypes (HWML 217950 – 217952); three paratypes (LRP 11282 – 11284); and, two paratypes (MZUSP 8060 a – b).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	etymology	Etymology: This species honors Miriam Fukumitsu Sakai, mother of Lilian Sakai (first author), for her support and encouragement during graduation activities.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	description	Description. [Based on 13 specimens: 9 whole mounts of mature worms, 1 scolex prepared for SEM and cross sections of 3 mature proglottids]: Worms euapolityc, 12.5 – 26.8 mm (n = 9) long with 40 – 67 proglottids (n = 9) in number, maximum width 721.5 - 1985 (n = 4) at level of scolex (Fig. 3 B). Scolex (Figs. 4 E, 5 F) with four stalked bothridia, each one with 90 – 95 (n = 2) marginal loculi and one oval apical sucker; apical sucker 57 – 66.5 (n = 2) long by 48.5 – 76.5 (n = 2) wide. Short cephalic peduncle present. Proximal surfaces of marginal loculi next to the bothridial rims covered with acicular filitriches and scolopate spinithriches (Fig. 5 G, H); bothridial rims and proximal surface of marginal loculi covered with acicular filitriches (Fig. 5 I); distal surface of bothridium covered with capiliform filitriches (Fig. 5 J). Proglottids slightly craspedote. Immature proglottids wider than long, becoming longer than wide with maturity, 38 – 64 in number (n = 9) (Fig. 3 B). Mature proglottids (Fig. 4 F) 1402.5 – 2043.5 (n = 6) long by 172 – 448 (n = 6) wide, 1 – 4 in number (n = 9). Testes arranged in two columns from near anterior margin of proglottid to anterior margin of genital pore, and 1 row-deep in cross section (Figs. 4 F, 6 C). Testes 45.5 – 72 (n = 4) long by 35.5 – 58.5 (n = 4) wide, 28 – 38 (n = 4) in number. Cirrus-sac pyriform bent posteriorly, 114 – 161 (n = 4) long by 60 – 89 (n = 4) wide, containing coiled cirrus (Fig. 4 D). Cirrus armed with spinitriches. Genital pores lateral, irregularly alternating, 32.1 – 45.4 % (n = 6) of proglottid length from posterior end (Fig. 4 D, F). Ovary near posterior end of proglottid, H-shaped in frontal view (Figs. 3 B, 4 F), tetralobed in cross-section (Fig. 6 D), slightly asymmetrical, 388 – 620.5 (n = 6) long by 111.5 – 270.5 (n = 6) wide; ovarian bridge near middle of ovary. Mehli’s gland immediately posterior to ovarian bridge. Vagina thick-walled, sinuous, extending medially in proglottid from ootype to anterior margin of cirrus-sac, then laterally to open into genital atrium anterior to cirrus (Fig. 4 D). Vitellarium follicular; vitelline follicles 22 – 75.5 (n = 6) long by 8.5 – 18.5 (n = 6) wide, arranged in two lateral bands, each band consisting of one dorsal and one ventral column of vitelline follicles, extending from near posterior margin of the first or second anterior-most row of testes to near posterior margin of proglottid, interrupted by genital pore and by ovary both dorsally and ventrally (Figs. 4 D, F). Uterus saccate, ventral, extending along median line of proglottid from near the Mehli’s gland to posterior margin of first or second anterior-most row of testes (Fig. 4 F).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF93E66C1A96FAF4FA9BFE0C.taxon	discussion	Remarks: Anthocephalum miriamae n. sp. possesses a unique combination of morphological features. This new species, can be distinguishable from 15 of its congeners in number of proglottids [40 – 67 vs. 13 – 21 (A. blairi), 20 – 33 (A. decrisantisorum), 120 – 160 (A. duszynskii), 500 – 600 (A. gracile), 9 – 16 (A. gravisi), 17 – 29 (A. haroldsoni), 105 – 138 (A. healyae), 12 – 28 (A. jensenae), 7 – 10 (A. mounseyi), 86 – 120 (A. odonnellae), 11 - 30 (A. ruhnkei), 16 – 28 (A. jeancadenati), 9 – 15 (A. alicae), 80 – 110 (A. cairae), and 106 – 177 (A. papefayei)]. In number of testes, Anthocephalum miriamae n. sp. possesses more testes than A. meadowsi and A. philruschi (28 – 38 vs. 15 – 25, 17 – 25, respectively) but fewer than A. centruturum (28 – 38 vs. 48 – 78). Anthocephalum miriamae n. sp. differs from A. hobergi based on the position of the genital pore on the proglottid (32 – 45 % vs. 66 – 79 %). The new species also differs in the arrangement of vitelline follicles by possessing a single dorsal and a single ventral column versus the 3 – 5 dorsal and 3 – 5 ventral columns of A. currani, 4 – 6 dorsal and 4 – 6 ventral columns of A. lukei, and 2 – 3 dorsal and 2 – 3 ventral columns of A. michaeli, A. kingae, and A. mattisi. Finally, Anthocephalum miriamae n. sp. differs from Anthocephalum machadoi as the former is proportionally larger, with different scolex shape (ellipsoid vs. deltoid, sensu Clopton (2004 )), and associated with a distinct biogeographical region (Tropical Eastern Pacific vs. Tropical Western Atlantic), evidences that are corroborated by their phylogenetic positions based on molecular data.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	description	(Figs. 3 C, 6 E – F, 7 A – C, 8 A – E)	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	biology_ecology	Type host: Styracura schmardae (Werner) – Chupare stingray.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	materials_examined	Type locality: Coast of Tobacco Caye, Stann Creek, Belize (16 ◦ 54 ’ 15.2 ” N, 88 ◦ 03 ’ 38.2 ” W).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	biology_ecology	Site of infection: Spiral intestine. Prevalence of infection: 20 % (one of five valves).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	materials_examined	Specimens deposited: Holotype (HWML 217953) and five paratypes (HWML 217954 – 217958); six paratypes (LRP 11285 – 11290); and, six paratypes (MZUSP 8061 a – f).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	etymology	Etymology: This species honors Dr. Natalia Luchetti for her contributions to cestodes taxonomy.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	description	Description. [Based on 28 specimens: 17 whole mounts of mature worms, 9 scolexes prepared for SEM and cross sections of 2 mature proglottids]: Worms euapolityc, 4,6 – 8.1 (n = 17) long with 13 – 21 proglottids (n = 17) in number, maximum width 661.5 – 902 (n = 3) at level of scolex (Fig. 3 C). Scolex (Figs. 7 B, 8 A) with four stalked bothridia, each one with 58 – 66 (n = 3) marginal loculi and one oval apical sucker; apical sucker 23.5 – 57 (n = 12) long by 39 – 60.5 (n = 12) wide. Short cephalic peduncle present. Proximal surfaces of marginal loculi next to the bothridial rims covered with acicular filitriches and scolopate spinithriches (Fig. 8 B, C); bothridial rims covered with acicular filitriches (Fig. 8 D); distal surface of bothridium covered with capiliform filitriches (Fig. 8 E). Proglottids slightly craspedote. Immature proglottids wider than long, becoming longer than wide with maturity, 12 – 20 in number (n = 17) (Fig. 3 C). Mature proglottids (Fig. 7 A) 1195.3 – 1751 (n = 11) long by 153 – 204.8 (n = 11) wide, 1 in number (n = 17). Testes arranged in two columns from near anterior margin of proglottid to anterior margin of genital pore, and 1 row-deep in cross section (Figs. 7 A, 6 E). Testes 40 – 51.3 (n = 9) by 32.8 – 50 (n = 9) wide, 36 – 50 (n = 9) in number. Cirrus-sac pyriform slightly bent posteriorly, 100.5 – 187.3 (n = 5) long by 71 – 95 (n = 5) wide, containing coiled cirrus (Fig. 7 C). Cirrus armed with spinitriches. Genital pores lateral, irregularly alternating, 29.4 – 42.1 % (n = 10) of proglottid length from posterior end (Fig. 3 C, 7 A). Ovary near posterior end of proglottid, H-shaped in frontal view (Figs. 3 C, 7 A), tetralobed in cross-section (Fig. 6 F), slightly asymmetrical, 238.5 – 528 (n = 9) long by 82.3 – 137.5 (n = 9) wide; ovarian bridge near middle of ovary. Mehli’s gland immediately posterior to ovarian bridge. Vagina thick-walled, sinuous, extending medially in proglottid from ootype to anterior margin of cirrus-sac, then laterally to open into genital atrium anterior to cirrus, expanded proximally (Fig. 7 C). Vitellarium follicular; vitelline follicles 21.5 – 48.3 (n = 6) by 5.3 – 18 (n = 6), arranged in 2 lateral bands, each band consisting of one dorsal and one ventral column of vitelline follicles, extending from posterior margin of first or second anterior-most row of testes to anterior margin of the ovary to near posterior margin of proglottid, interrupted by genital pore and partially by ovary both dorsally and ventrally (Figs. 7 A, C). Uterus saccate, ventral, extending along median line of proglottid from near the ovarian bridge to posterior margin of first or second anterior-most row of testes (Fig. 7 A).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF96E6721A96FDC0FD77FE54.taxon	discussion	Remarks: Anthocephalum nataliae n. sp. can be distinguished from 16 of its 26 congeners in number of proglottids (13 – 21 vs. 35 – 70 A. currani, 120 – 160 A. duszynskii, 500 - 600 A. gracile, 105 – 133 A. healyae, 28 – 56 A. lukei, 30 – 40 A. meadowsi, 23 – 41 A. michaeli, 7 – 10 A. mounseyi, 86 – 120 A. odonnellae, 27 – 40 A. philruschi, 80 – 110 A. cairae, 33 – 50 A. kingae, 35 – 40 A. mattisi, 106 – 177 A. papefayei, 53 – 98 A. hobergi and, 40 – 67 A. miriamae). Furthermore, A. nataliae n. sp. differs from A. blairi, A. decrisantisorum, A. haroldsoni, A. jensenae, and A. ruhnkei by possessing more testes (36 – 50 vs. 10 – 15, 17 – 24, 25 – 32, 14 – 20, and 22 – 34, respectively). Anthocephalum nataliae n. sp. closely resembles A. jeancadenati, A. alicae, A. gravisi, and A. machadoi in total length, number of proglottids, and number of testes. However, A. nataliae n. sp. possesses a unique vitelline follicle arrangement: two lateral bands each with one dorsal and one ventral column. This contrasts with A. jeancadenati, which possesses two lateral bands with 2 – 3 dorsal and 2 – 3 ventral columns each, and A. alicae, with two lateral bands each with two dorsal and two ventral columns. Lastly, Anthocephalum nataliae n. sp. differs from A. machadoi by possessing fewer marginal loculi (58 – 66 vs. 76 – 96, respectively) and from A. gravisi by possessing a uterus that extends to the anterior margin of the proglottid.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	description	(Figs. 3 D, 6 G – H, 7 D – F, 8 F – J)	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	biology_ecology	Type host: Styracura schmardae (Werner, 1904) — Chupare stingray.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	materials_examined	Type locality: Coast of Tobacco Caye, Stann Creek, Belize (16 ° 54 ’ 15.2 ” N, 88 ° 03 ’ 38.2 ” W). Additional localities: Head Caye, Toledo, Belize (16 ° 49 ’ 43.1 ” N, 88 ° 04 ’ 48.1 ” W).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	biology_ecology	Site of infection: Spiral intestine. Prevalence of infection: 40 % (two of five valves).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	materials_examined	Specimens deposited: Holotype (HWML 217959) and three paratypes (HWML 217960 – 217962); four paratypes (LRP 11291 – 11294); and three paratypes (MZUSP 8062, 8063 a – b).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	etymology	Etymology: This species honors Dr. Veronica Mantovani Bueno for her contributions to cestodes taxonomy and systematics.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	description	Description. [Based on 18 specimens: 9 whole mounts of mature worms, 8 scolexes prepared for SEM and cross sections of 1 mature proglottid]: Worms euapolityc, 22.6 – 43.5 mm (n = 9) long with 116 – 180 proglottids (n = 9) in number, maximum width 2472 – 2828 (n = 3) at level of scolex (Fig. 3 D). Scolex (Figs. 7 D, 8 F) with four stalked bothridia, each one with 180 – 190 (n = 3) marginal loculi and one oval apical sucker; apical sucker 75 – 107 (n = 2) long by 56.5 – 78 (n = 2) wide. Short cephalic peduncle present. Proximal surfaces of marginal loculi next to the bothridial rims covered with acicular filitriches and scolopate spinithriches (Fig. 8 G, H); bothridial rims covered with acicular filitriches (Fig. 8 I); distal surface of bothridium covered with capiliform filitriches (Fig. 8 J). Proglottids craspedote. Immature proglottids wider than long, becoming longer than wide with maturity, 104 – 178 in number (n = 9) (Fig. 3 D). Mature proglottids (Fig. 7 E) 1667 – 2230 (n = 6) long by 405 – 577 (n = 6) wide, 1 – 3 in number (n = 9). Testes arranged in four columns from near anterior margin of proglottid to anterior margin of genital pore, and 1 row-deep in cross section (Figs. 7 E, 6 G). Testes 44 – 79 (n = 5) by 31 – 66 (n = 5) wide, 64 – 90 (n = 4) in number. Cirrus-sac pyriform bent posteriorly, 80.5 – 272 (n = 6) long by 76.5 – 161 (n = 6) wide, containing coiled cirrus (Fig. 7 F). Cirrus armed with spinitriches. Genital pores lateral, irregularly alternating, 29.5 – 38.6 % (n = 6) of proglottid length from posterior end (Fig. 3 D, 7 F). Ovary near posterior end of proglottid, H-shaped in frontal view (Figs. 3 D, 7 E), tetralobed in cross-section (Fig. 6 H), slightly asymmetrical, 393.5 – 649.5 (n = 6) long by 209.5 – 340.5 (n = 6) wide; ovarian bridge near middle of ovary. Mehli’s gland posterior to ovarian bridge. Vagina thick-walled, sinuous, extending medially in proglottid from ootype to anterior margin of cirrus-sac, then laterally to open into genital atrium anterior to cirrus (Fig. 7 F). Vitellarium follicular; vitelline follicles 46 – 74.5 (n = 6) by 10.5 – 22.5 (n = 5) wide arranged in two lateral bands, each band consisting of one dorsal and one ventral column of vitelline follicles, extending from posterior margin of the second or third anterior-most row of testes to near posterior margin of the proglottid, interrupted by genital pore both dorsally and ventrally (Figs. 7 E, F). Uterus saccate, ventral, extending along median line of proglottid from near the ovarian bridge to posterior margin of first or second anterior-most row of testes (Fig. 7 E).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF88E6721A96FE38FDA1F848.taxon	discussion	Remarks: Anthocephalum veronicae n. sp. is morphologically distinct from all except two species within its genus — A. haroldsoni and A. gravisi — by possessing uninterrupted vitelline follicles by the ovary. However, the new species is clearly different from A. haroldsoni and A. gravisi in total length (22.6 – 43.5 mm vs. 1.8 – 3.7 mm and 3.6 – 7.9 mm, respectively), by possessing a larger number of proglottids (116 – 180 vs. 9 – 16 and 17 – 29, respectively), a higher number of loculi (180 – 190 vs. 43 – 52 and 41 – 57, respectively), and more testes in number (64 – 90 vs. 23 – 38 and 25 – 32, respectively).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF89E6701A96F9DBFECEFBA0.taxon	description	Worms weakly craspedote, apolytic or euapolytic. Scolex with 4 bothridia and short or absent cephalic peduncle. Bothridia stalked, with numerous marginal loculi and single apical sucker; facial loculi present or absent. Proximal surfaces of marginal loculi with acicular filitriches and with or without scolopate spinitriches, or with capilliform filitriches and narrow gladiate spinitriches; proximal non-locular surfaces of bothridia with acicular filitriches, and with or without narrow gladiate spinitriches; distal bothridial surfaces with acicular or capilliform filitriches and small gladiate spinitriches. Mature proglottids longer than wide. Testes entirely pre-ovarian, medullary; postporal field of testes lacking. Genital pores lateral, usually in posterior half of proglottid; genital atrium present. Internal seminal vesicle present or absent. Vagina sinuous, opening into genital atrium anterior to cirrus sac. Ovary posterior in position, H-shaped in frontal view, tetralobed in cross-section. Vitellaria follicular; vitelline follicles in 2 lateral bands; each band consisting of 1 or more columns of vitelline follicles, uninterrupted by ovary, or partially or completely interrupted by ovary, and interrupted or uninterrupted by genital pore. Uterus median, ventral, sacciform, extending from level of cirrus sac to stopping short of anterior limit of field of testes or to anterior margin of proglottid. Parasites of stingrays (Myliobatiformes) and electric rays (Torpediniformes).	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
0399878EFF89E6701A96F9DBFECEFBA0.taxon	discussion	Remarks: The revision of the Anthocephalum diagnosis was necessary to accommodate the unique scolex morphology of Anthocephalum grabatum n. comb., and Anthocephalum zarzisense n. comb., which possess facially loculated bothridia. The incorporation of these two species raises the total number of species within the genus to 30 (Boudaya et al., 2018; Herzog and Jensen, 2018). Alveobothrium was established by Boudaya et al. (2018) to include species morphologically akin to Anthocephalum, but characterized by a scolex bearing staggered facial loculi arranged in multiple rows, a trait absent in Anthocephalum sensu stricto (Ruhnke et al., 2015; Herzog and Jensen, 2018). Despite this distinguishing feature, Alveobothrium shares all other morphological characteristics with Anthocephalum, leading the authors to recognize Alveobothrium on the basis of its unique scolex morphology alone. Phylogenetic analyses have previously supported the inclusion of tapeworms with varying scolex morphologies within a single genus of chondrichthyan parasites (e. g., Reyda et al., 2016; Trevisan et al., 2017; Caira et al., 2020). For example, Anindobothrium lisae Marques, Brooks & Lasso, 2001, a parasite of freshwater stingrays, lacks facial loculi and longitudinal septa, which contrast with marine species of the same genus that exhibit these features. Despite these differences, Anindobothrium lisae shares key morphological traits with other members of the genus and is confirmed by molecular evidence to belong to Anindobothrium (Trevisan et al., 2017). Additionally, in a recent redefinition of Scyphophyllidium Woodland, 1927 (Phyllobothriidea), Caira et al. (2020) examined six genera with similar proglottid anatomy and unique ultrastructural characteristics but divergent bothridial morphologies, such as the presence or absence of apical suckers and loculi. They concluded that bothridial morphology represented intra-genus diversity and proposed synonymizing the six genera under Scyphophyllidium, organizing the species into eight groups based on bothridial morphology. Similarly, despite differences in bothridial morphology, species of Anthocephalum and Alveobothrium display comparable proglottid anatomies. In our phylogenetic analysis, the positioning of Alveobothrium grabatum alongside other Anthocephalum species is strongly supported. This result, when considered in conjunction with the aforementioned evidence, substantiates the proposed taxonomic synonymy.	en	Sakai, Lilian F., Marques, Fernando P. L., Trevisan, Bruna (2025): Diversity and phylogenetic position of the amphi-American lineages of the tapeworms of the genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae). Zootaxa 5584 (2): 151-178, DOI: 10.11646/zootaxa.5584.2.1, URL: https://doi.org/10.11646/zootaxa.5584.2.1
