taxonID	type	description	language	source
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	description	ZooBank number for genus: urn: lsid: zoobank. org: act: B 2 B 660 B 1 - E 30 D- 4208 - 935 E-F 89 C 06 A 2 D 16 D	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	diagnosis	Diagnosis: Rhinebothriidea. Worms euapolytic, small. Scolex consisting of scolex proper and 4 bothridia; cephalic peduncle absent; short germinative zone present; apical organ absent. Bothridia stalked, consisting of anterior and posterior regions with distinctly different arrangement of loculi and septa; bothridial margins with thin rim. Anterior region with horizontally oriented loculi (i. e. loculi wider than long) with two (Figs. 2 B, 4 B) or more (Figs. 6 B, 8 B, 10 B, 12 B, 15) complete transverse septa, with (Figs. 2 B, 4 B) or without (Figs. 6 B, 8 B, 10 B, 12 B, 15) single partial medial longitudinal septum. Posterior region lacking medial longitudinal septum, divided into odd number of vertically oriented loculi (i. e. loculi longer than wide) by even number of nonmedial longitudinal septa; nonmedial longitudinal septa all incomplete (Figs. 6 B, 8 B, 10 B, 12 B, 15) or a combination of incomplete and complete (Figs. 2 B, 4 B); incomplete nonmedial longitudinal septa either abut posteriormost transverse septum of anterior region of bothridia (Figs. 2 B, 4 B, 10 B,), or overlap one or more posteriormost transverse septa (Figs. 6 B, 8 B, 12 B, 15). Lateral margins of posterior region of bothridium divided into additional loculi by marginally (Figs. 2 B, 4 B) or diagonally (Figs. 6 B, 15) oriented septa in some species. Longitudinal septa of posterior region appear as ridges in section (Fig. 13) with proximal and distal portions different; proximal portion of septa formed by underlying bothridial wall, consisting of radial muscles oriented with proximal ends of fibres adjacent to each other; distal portion of septa formed by separate muscle bundle; proximal and distal portions of septa separated by triangular gap. Testes numerous, arranged in two columns, one layer deep in cross section, restricted to pre-poral region of proglottid. Cirrus sac extending medially to or past midline of proglottid. Cirrus spinitriches present. Vas deferens extending posteriorly to ovarian isthmus, entering cirrus sac at anterior margin. Vagina opening anterior to cirrus sac; vaginal sphincter absent. Ovary H-shaped in dorsoventral view, tetralobed in cross section. Vitellarium follicular; follicles in 2 lateral bands; bands interrupted by terminal genitalia and usually also by ovary. Uterus saccate, medial, extending from posterior margin of proglottid or ovarian isthmus, anteriorly to near anterior margin of proglottid. Parasites of batoid elasmobranchs (Rhinobatidae, Zanobatidae and Dasyatidae); Indo-Pacific and coastal Afro-tropics.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	type_taxon	T y p e s p e c i e s: Stillabothrium ashleyae sp. n.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	materials_examined	A d d i t i o n a l s p e c i e s: Stillabothrium amuletum (Butler, 1987) comb. n.; Stillabothrium cadenati (Euzet, 1954) comb. n.; Stillabothrium campbelli sp. n.; Stillabothrium davidcynthiaorum sp. n.; Stillabothrium hyphantoseptum sp. n.; Stillabothrium jeanfortiae sp. n.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	etymology	E t y m o l o g y: From the Latin ‘ stilla’, meaning drop, for the teardrop shape of the bothridia of species the genus.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF4FFA2DF958485FD6A2B58.taxon	discussion	Remarks. Stillabothrium gen. n. is generally consistent with the diagnosis of the order Rhinebothriidea as given by Healy et al. (2009): Species of Stillabothrium possess facially loculated bothridia borne on stalks and possess a vas deferens that enters the cirrus sac at the anterior, rather than the medial, margin. Stillabothrium can be distinguished from all rhinebothriidean genera except Escherbothrium Berman and Brooks, 1994, Phormobothrium Alexander, 1963 and Tritaphros Lönnberg, 1889 in its possession of bothridia that are fully facially loculate, with posterior loculi that are longer than wide. Stillabothrium differs from Phormobothrium and Tritaphros in lacking an apical organ on the scolex. Stillabothrium is most similar to Escherbothrium, but Stillabothrium can be distinguished from the latter genus in lacking a medial longitudinal septum in the posterior region of the bothridium, thereby possessing an odd number of loculi. In Escherbothrium, the posterior region of the bothridium includes a short medial longitudinal septum (see fig. 6 in Berman and Brooks 1994) and an even number of loculi. In addition, Escherbothrium was described as possessing an apical sucker and rounded protrusions on its distal bothridial surfaces (see both in fig. 8 in Berman and Brooks 1994). Conversely, in Stillabothrium the feature on the anteriormost portion of the bothridium is considered to be a loculus, rather than a sucker, and no rounded protrusions were observed on the scolex of any surfaces of any of the six species of Stillabothrium examined with SEM in this study. Based on the recent designation of families within Rhinebothriidea by Ruhnke et al. (2015), Stillabothrium belongs to family Escherbothriidae Ruhnke, Caira et Cox, 2015. Species of Stillabothrium have appeared in previous works under different temporary names. The genus was first recognised by Healy (2006) in her dissertation, under a preliminary name which, as recommended by Article 8 of the ICZN (1999), she disclaimed. In addition to providing preliminary morphological characterisation of species, Healy (2006) included partial 28 S rDNA sequence data for eight species of Stillabothrium. Subsequently, four molecular phylogenetic studies (Healy et al. 2009, Caira et al. 2014, Ruhnke et al. 2015, Marques and Caira 2016) have included the sequence data originally generated by Healy (2006). Each study, which refers to Stillabothrium as ‘ Rhinebothriinae New genus 3 ’, supported recognition of those eight species as an independent, novel, genus.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF2FFA0DF958145FD9D2ED7.taxon	description	Figs. 1 – 3, 16 A ZooBank number for species: urn: lsid: zoobank. org: act: B 316 EE 69 - A 04 B- 4 A 09 - 9 AB 5 - 658752 FDC 431 Description (based on whole mounts of 25 complete mature worms and 14 free proglottids, cross sections of 2 strobila, longitudinal sections of 1 scolex, and 3 scoleces prepared for SEM): Worms (Fig. 2 A) euapolytic, acraspedote, 1.11 – 2.78 mm (1.66 ± 0.39; n = 23) long, greatest width 353 – 590 (455 ± 70; n = 24) at level of scolex; 6 – 10 (7.7. ± 1; n = 25) proglottids per worm. Cephalic peduncle lacking; darkly staining germinative zone 17 – 217 (54 ± 37; n = 24) long. Scolex (Fig. 2 B) consisting of scolex proper bearing 4 stalked bothridia. Stalks 46 – 164 (76 ± 31; n = 13) long by 53 – 100 (75 ± 17; n = 13) wide, attached slightly posterior to middle of bothridia. Bothridia (Fig. 2 B) varying in shape with degree of contraction, from shallowly-deltoid (Fig. 3 A) to deeply-deltoid (Fig. 3 B), facially loculated, 140 – 227 (175 ± 26; n = 20) long by 190 – 334 (270 ± 44; n = 24) wide; bothridial margins with thin rim. Anterior region of bothridia (Fig. 2 B) with 3 (n = 18) horizontally oriented loculi (i. e. loculi wider than long) with 2 complete transverse septa and one partial medial longitudinal septum. Anteriormost loculus 30 – 46 (39 ± 5; n = 15) long by 41 – 72 (59 ± 11; n = 18) wide. Posterior region of bothridia with 8 (n = 21) nonmedial longitudinal septa dividing bothridia into 9 primary loculi longer than wide; outermost primary loculi on each side subdivided by 3, or occasionally 2 (2.94 ± 0.2; n = 16) relatively short marginal septa into 3 – 4 small subloculi; longitudinal septa of posterior region not overlapping transverse septa of anterior region. Loculi (Fig. 3 D) and septa of distal bothridial surfaces bearing capilliform filitriches and coniform spinitriches. Proximal bothridial rim (Fig. 3 E) bearing capilliform filitriches greater in length than those on distal bothridial surfaces (Fig. 3 D). Proximal bothridial surfaces (Fig. 3 E, F) away from rim bearing acicular filitriches and coniform spinitriches. Isolated cilia observed on proximal bothridial surfaces. Bothridial stalks (Fig. 3 G) and strobila (Fig. 3 H) bearing capilliform filitriches only. Strobila with 2 – 4 (3.2 ± 0.8; n = 25) proglottids wider than long followed by 3 – 7 (4.5 ± 1; n = 25) proglottids longer than wide. Strobila widest at terminal proglottid; terminal proglottid 320 – 880 (591 ± 125; n = 25) long by 68 – 150 (101 ± 20; n = 24) wide; genital pore located 35 – 46 % (40 ± 3; n = 25) of proglottid length from proglottid posterior margin. Immature proglottids 4 – 9 (6.4 ± 1.2; n = 25) in number. Mature proglottids 1 – 2 (1.2 ± 0.4; n = 25) in number, including 0 – 1 (0.3 ± 0.5; n = 25) vas deferens-mature proglottids. Testes in mature proglottids 18 – 27 (22 ± 2; n = 25) in total number, 1 layer deep in cross section (Fig. 2 D), arranged in 2 columns (Fig. 2 C); columns extending from anterior margin of proglottid to anterior margin of cirrus sac, 19 – 37 (25 ± 4; n = 24) long by 20 – 45 (31 ± 6; n = 22) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending from level of ovarian isthmus to overlap posteriormost testes (Fig. 2 C). Cirrus sac thin-walled, oval, extending medially to near midline of proglottid; cirrus sac in terminal mature proglottid 26 – 42 (33 ± 5; n = 17) long by 26 – 47 (32 ± 6; n = 17) wide; cirrus sac in vas deferens-mature proglottids 28 – 38 (34 ± 4; n = 8) long by 36 – 50 (43 ± 5; n = 8) wide. Cirrus spinitriches present. Vagina (Fig. 2 C) thick-walled, weakly sinuous, somewhat overlapping anterior margin of cirrus sac (Fig. 16 A), extending along midline of proglottid from ootype region to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, H-shaped in frontal view, tetralobed in cross section (Fig. 2 E); ovarian lobes asymmetrical; poral and aporal ovarian lobes in terminal mature proglottids 80 – 215 (161 ± 33; n = 16) and 85 – 220 (163 ± 34; n = 16) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 146 – 265 (194 ± 47; n = 7) and 141 – 282 (190 ± 51; n = 7) long, respectively. Maximum width of ovary 35 – 96 (56 ± 17; n = 17). Ovarian isthmus at or anterior to midpoint of ovary; poral lobe of ovary stopping 26 – 63 (41 ± 12; n = 23) short of genital pore. Mehlis’ gland well posterior to ovarian isthmus, 19 – 35 (26 ± 5; n = 21) long by 14 – 30 (18 ± 4; n = 21) wide. Vitellarium follicular; vitelline follicles arranged in 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by terminal genitalia, and mostly interrupted by ovary (Fig. 2 C). Uterus ventral, sacciform, extending from near isthmus of ovary to near anterior margin of proglottid. Free proglottids with greatly expanded vas deferens and atrophied testes, 521 – 919 (724 ± 136; n = 12) long by 134 – 202 (165 ± 21; n = 12) wide. Free gravid proglottids 750 (n = 1) long by 210 – 280 (245 ± 50; n = 2) wide; unembryonated eggs oval, without filaments, 14 – 17 (15 ± 1; n = 6; from same proglottid) long. T y p e h o s t: Dasyatis biasa (Last, White et Naylor) (Myliobatiformes: Dasyatidae). A d d i t i o n a l h o s t: None. Ty p e l o c a l i t y: South China Sea off Mukah (02 ° 53 ' 52 '' N; 112 ° 05 ' 44 '' E), Sarawak, Malaysian Borneo (BO- 477). A d d i t i o n a l l o c a l i t i e s: Java Sea off Selakau (01 ° 03 ' 31 '' N; 108 ° 58 ' 26 '' E), West Kalimantan, Indonesian Borneo (KA- 182, KA- 184). Java Sea off Sukadana (01 ° 14 ' 33 '' S; 109 ° 57 ' 00 '' E), West Kalimantan, Indonesian Borneo (KA- 378). S i t e o f i n f e c t i o n: Spiral intestine. Ty p e m a t e r i a l: Holotype MZUM (P) No. 2016.6 (H). Paratypes: IPCAS No. C- 737; LRP Nos. 8992 – 8994; 9005 – 9037 (including molecular vouchers, cross sections and SEM specimens); MZB Nos. Ca 197 – Ca 199; MZUM (P) Nos. 2016.7 (P) – 2016.8 (P); SBC No. P- 00069; USNM Nos. 1420453 – 1420460. E t y m o l o g y: This species is named in honour of Ashley Willsey Attoma, sister of D. D. Willsey, for her support of her sister’s studies.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF2FFA0DF958145FD9D2ED7.taxon	discussion	Remarks. Stillabothrium ashleyae sp. n. is the type and first described species of the genus. The distribution of S. ashleyae includes both the Malaysian and Indonesian portions of the island of Borneo. Stillabothrium ashleyae appears to be highly host specific, as it was only encountered in specimens of D. biasa. It was not found parasitising any of the multiple other species of dasyatids (e. g. Dasyatis Rafinesque and Himantura Müller et Henle) examined during our survey work. We note that specimens with a morphology similar to S. ashleyae were found in D. zugei (VN- 23, VN- 34) from Vietnam. However, preliminary molecular and morphological data suggest that the specimens from Vietnam represent a distinct species of Stillabothrium that awaits formal study and description. Three specimens of S. ashleyae were included in the phylogenetic analysis (Fig. 1, Table 1).	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF0FFADDF9584C5FE632C37.taxon	description	Figs. 1, 4, 5, 16 B ZooBank number for species: urn: lsid: zoobank. org: act: D 36 AE 2 D 9 - C 4 A 6 - 4 C 4 D- 8 F 82 - E 9 C 0 FAF 1 F 92 C Description (based on whole mounts of 32 complete mature worms, cross sections of 1 strobila and longitudinal sections of 1 scolex and 3 scoleces prepared for SEM): Worms (Fig. 4 A) euapolytic, acraspedote, 0.62 – 2.59 mm (1.29 ± 0.57; n = 30) long, greatest width 195 – 561 (332 ± 84.7; n = 32) at level of scolex; 4 – 10 (7 ± 2.1; n = 32) proglottids per worm. Cephalic peduncle lacking; darkly staining germinative zone 12 – 40 (22 ± 7; n = 29) long. Scolex (Fig. 4 B) consisting of scolex proper bearing 4 stalked bothridia. Stalks 25 – 100 (52 ± 20; n = 23) long by 27 – 83 (55 ± 17; n = 23) wide, attached slightly posterior to middle of bothridia. Bothridia (Fig. 4 B) varying in shape with degree of contraction, from shallowly-deltoid (Fig. 5 A) to deeply-deltoid (Figs. 4 B, 5 B), facially loculated, 72 – 175 (108 ± 26; n = 20) long by 140 – 325 (202 ± 42; n = 32) wide; bothridial margins with thin rim. Anterior region of bothridia (Fig. 4 B) with 3 (n = 28) horizontally oriented loculi (i. e. loculi wider than long) with 2 complete transverse septa and one partial medial longitudinal septum. Anteriormost loculus 22 – 49 (31 ± 8; n = 18) long and 37 – 78 (47 ± 9; n = 25) wide. Posterior region of bothridia with 10 (n = 30) nonmedial longitudinal septa dividing bothridia into 11 primary loculi longer than wide, outermost primary loculi on each side subdivided by 2, or occasionally 3 (2.04 ± 0.2; n = 24) relatively short marginal septa into 3 – 4 small subloculi; longitudinal septa of posterior region not overlapping transverse septa of anterior region. Loculi and septa of distal bothridial surfaces (Fig. 5 C) bearing capilliform filitriches and coniform spinitriches. Proximal bothridial rim (Fig. 5 D) bearing capilliform filitriches greater in length than those on distal bothridial surfaces (Fig. 5 C). Proximal bothridial surfaces (Fig. 5 D, E) away from rim bearing acicular filitriches and coniform spinitriches. Isolated cilia observed on proximal bothridial surfaces. Bothridial stalks (Fig. 5 F) bearing capilliform filitriches and coniform spinitriches; strobila (Fig. 5 G) bearing capilliform filitriches only. Strobila with 1 – 5 (2.6 ± 0.9; n = 32) proglottids wider than long followed by 2 – 8 (4.1 ± 1.8; n = 32) proglottids longer than wide. Strobila widest at terminal proglottid; terminal proglottid 255 – 790 (438 ± 124; n = 31) long by 60 – 144 (108 ± 18; n = 32) wide; genital pore located 42 – 54 % (48 ± 3.5; n = 21) of proglottid length from proglottid posterior margin. Immature proglottids 3 – 9 (5.5 ± 1.9; n = 32) in number. Mature proglottids 1 – 3 (1.2 ± 0.5; n = 32) in number, including 0 – 2 (0.2 ± 0.5; n = 32) vas deferens-mature proglottids. Testes in mature proglottids 11 – 21 (16 ± 3; n = 32) in total number, 1 layer deep in cross section, arranged in 2 columns (Fig. 4 C); columns extending from anterior margin of proglottid to anterior margin of cirrus sac, 13 – 49 (25 ± 8; n = 32) long by 24 – 50 (35 ± 6; n = 32) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending from level of ovarian isthmus to overlap posteriormost testes. Cirrus sac thin-walled, oval, extending medially to near midline of proglottid; cirrus sac in terminal mature proglottid 18 – 35 (26 ± 5; n = 26) long by 24 – 38 (31 ± 4; n = 24) wide; cirrus sac in vas deferens-mature proglottids 32 – 45 (39 ± 5; n = 5) long by 33 – 46 (40 ± 5; n = 5) wide. Cirrus spinitriches present. Vagina (Fig. 4 C) thick-walled, weakly sinuous, somewhat overlapping anterior margin of cirrus sac (Fig. 16 B), extending along midline of proglottid from ootype region to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, H-shaped in frontal view, tetralobed in cross section; ovarian lobes asymmetrical; poral and aporal ovarian lobes in terminal mature proglottids 75 – 254 (144 ± 40; n = 24) and 75 – 272 (152 ± 42; n = 24) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 145 – 269 (198 ± 48; n = 5) and 152 – 270 (206 ± 46; n = 5) long, respectively. Maximum width of ovary 39 – 139 (65 ± 21; n = 32). Ovarian isthmus near midpoint of ovary; poral lobe of ovary stopping 19 – 85 (31 ± 16; n = 28) short of genital pore. Mehlis’ gland well posterior to ovarian isthmus, 12 – 43 (27 ± 8; n = 29) long by 10 – 35 (20 ±; n = 29) wide. Vitellarium follicular, vitelline follicles arranged in 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by genital genitalia, and mostly interrupted by ovary (Fig. 4 C). Uterus ventral, sacciform, extending from near isthmus of ovary to near anterior margin of proglottid. I n f o r m a l s y n o n y m s: Rhinebothriinae New genus 3 sp. n. 6 of Healy et al. (2009), Caira et al. (2014), Ruhnke et al. (2015), Marques and Caira (2016). T y p e h o s t: Himantura heterura (Bleeker), dwarf whipray (Myliobatiformes: Dasyatidae). A d d i t i o n a l h o s t s: Himantura macrura, Himantura gerrardi, Himantura uarnak 3. Ty p e l o c a l i t y: South China Sea off Sematan (01 ° 48 ’ 15 '' N; 109 ° 46 ' 47 '' E), Sarawak, Malaysian Borneo (hosts BO- 19, BO- 141, BO- 170). A d d i t i o n a l l o c a l i t i e s: South China Sea off Mukah (02 ° 53 ' 52 '' N; 112 ° 05 ' 44 '' E), Sarawak, Malaysian Borneo (BO- 47, BO- 66, BO- 67, BO- 237, BO- 238). Java Sea off Kalapseban (03 ° 14 ' 30 '' S; 112 ° 54 ' 52 '' E), Central Kalimantan, Indonesian Borneo (KA- 99, KA- 111). Java Sea off Singkawang (00 ° 55 ' 06 '' N; 108 ° 58 ' 60 '' E), West Kalimantan, Indonesian Borneo (KA- 145). S i t e o f i n f e c t i o n: Spiral intestine. Ty p e m a t e r i a l: Holotype MZUM (P) No. 2016.9 (H). Paratypes: IPCAS No. C- 739; LRP Nos. 8986 – 8991; 9038 – 9062 (including molecular vouchers, cross sections and SEM specimens); MZB Nos. Ca 200 – Ca 201; MZUM (P) No. 2016.10 (P); SBC No. P- 00070; USNM Nos. 1420461 – 1420466. E t y m o l o g y: This species is named in honour of David and Cynthia Daigler, parents of A. L. Daigler, for their support of his education and his interests.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF0FFADDF9584C5FE632C37.taxon	discussion	Remarks. Stillabothrium davidcynthiaorum sp. n. can be distinguished from S. ashleyae in its possession of 10, rather than eight, longitudinal septa on the central posterior region of the bothridium. While S. davidcynthiaorum and S. ashleyae are similar in possessing a range of 2 – 3 marginal septa on both of the lateral sides on the posterior region of the bothridium, the former differs from the latter in mean number (2.04 [n = 24] vs 2.94 [n = 16]). In the present study we report specimens of S. davidcynthiaorum from potentially four species of Himantura, H. heterura, H. macrura, H. gerrardi and H. uarnak 3. Although S. davidcynthiaorum was encountered in four potential species of Himantura, we point out having encountered specimens morphologically similar to S. davidcynthiaorum in four additional species of Himantura that were examined during the survey work in Borneo, including Himantura cf. pastinacoides (BO- 61, BO- 168), H. undulata (BO- 24), U. lobistomus (BO- 247) and H. oxyrhyncha (KA- 252). Further examination of the Stillabothrium specimens from those additional four species of Himantura is needed, in combination with DNA sequence data, to establish whether those cestodes are conspecific with S. davidcynthiaorum or represent an undescribed species of the genus. Seven specimens of S. davidcynthiaorum were included in the phylogenetic analysis (Fig. 1, Table 1). One of the seven specimens (LRP 3926) was previously included in the analysis provided by Healy et al. (2009) as ‘ Rhinebothriinae New genus 3 sp. n. 6 ’.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFDFFAADF958665FD9D2ED7.taxon	description	Figs. 1, 6, 7, 16 C ZooBank number for species: urn: lsid: zoobank. org: act: 8 EA 1 CC 15 - 6 A 59 - 4 FDC- 96 DE- 7 B 8389 B 40 F 7 C Description (based on whole mounts of 14 complete mature worms, cross sections of 2 strobila and longitudinal sections of 3 scoleces and 3 scoleces prepared for SEM): Worms (Fig. 6 A) euapolytic, acraspedote, 1.06 – 2.60 mm (1.68 ± 0.4; n = 14) long, greatest width 170 – 360 (261 ± 55; n = 14) at level of scolex; 5 – 8 (6 ± 1.0; n = 14) proglottids per worm. Cephalic peduncle lacking; small darkly staining germinative zone present. Scolex (Fig. 6 B) consisting of scolex proper bearing 4 stalked bothridia. Stalks 20 – 90 (49 ± 2; n = 13) long by 25 – 110 (68 ± 25; n = 13) wide, attached slightly posterior to middle of bothridia. Bothridia varying in shape with degree of contraction; ovoid, broadly ovoid, finely ovoid (Fig. 6 B), deeply ovoid (Fig. 7 B), very deeply ovoid, to finely deltoid (Fig. 7 A), facially loculated, 166 – 240 (205 ± 22; n = 15) long by 110 – 235 (150 ± 32; n = 14) wide; bothridial margins with thin rim. Anterior region of bothridia (Fig. 6 B) with 10 – 12 (10.6 ± 0.7; n = 11) horizontally oriented loculi (i. e. loculi wider than long) with 10 – 12 complete transverse septa. Anteriormost loculus 15 – 25 (20 ± 4; n = 13) long and 24 – 35 (31 ± 3; n = 12) wide. Posterior region of bothridia with 4 (n = 11) nonmedial longitudinal septa dividing bothridia into 5 primary loculi longer than wide; outermost primary loculi on each side subdivided by 3 – 4 (3.7 ± 0.5; n = 9) diagonal septa into 4 – 5 small subloculi; central 2 longitudinal septa of posterior region overlapping 3 – 4 (3.3 ± 0.5; n = 6) posteriormost transverse septa of anterior region. Loculi of distal bothridial surfaces (Fig. 7 C) bearing acicular and capilliform filitriches; septa on distal bothridial surfaces (Fig. 7 C) bearing acicular and capilliform filitriches and coniform spinitriches. Proximal bothridial rim (Fig. 7 D) bearing capilliform filitriches greater in length than those on distal bothridial surfaces (Fig. 7 C). Proximal bothridial surfaces away from rim bearing acicular filitriches throughout (Fig. 7 D). Posterior margin of proximal bothridial surfaces bearing patch of coniform spinitriches (Fig. 7 E) near, but not extending to, bothridial rim. Isolated cilia observed on distal and proximal bothridial surfaces. Bothridial stalks and strobila (Fig. 7 F) bearing capilliform filitriches only. Strobila with 1 – 4 (2.3 ± 1.0; n = 14) proglottids wider than long followed by 3 – 5 (3.8 ± 0.6; n = 14) proglottids that are longer than wide. Strobila widest at terminal proglottid; terminal proglottid 490 – 1 250 (798 ± 209; n = 14) long by 78 – 180 (125 ± 31; n = 14) wide, genital pore located 41 – 55 % (49 ± 4.7; n = 14) of proglottid length from proglottid posterior margin. Immature proglottids 3 – 6 (4.5 ± 0.9; n = 14) in number. Mature proglottids 1 – 2 (1.6 ± 0.5; n = 14) in number, including 0 – 1 (0.7 ± 0.5; n = 14) vas deferens-mature proglottid. Testes in mature proglottids 12 – 19 (15 ± 2.1; n = 14) in total number, 1 layer deep in cross section, arranged in 2 columns; columns extending from anterior margin of proglottid to anterior margin of cirrus sac, 23 – 42 (33 ± 6; n = 13) long by 28 – 50 (37 ± 8; n = 13) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending from level of ovarian isthmus to overlap posteriormost testes. Cirrus sac thin-walled, oval, extending medially well past midline of proglottid; cirrus sac in terminal mature proglottid 42 – 80 (66 ± 17; n = 4) long by 50 – 70 (62 ± 9; n = 4) wide; cirrus sac in vas deferens-mature proglottids 45 – 90 (71 ± 13; n = 9) long by 50 – 100 (75 ± 18; n = 9) wide. Cirrus spinitriches present. Vagina (Fig. 6 C) thick-walled, sinuous, somewhat overlapping anterior margin of cirrus sac (Fig. 16 C), extending from ootype past midline to aporal side of proglottid to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, H-shaped in frontal view, tetralobed in cross section, overlapping cirrus sac; ovarian lobes symmetrical; poral ovarian lobe somewhat overlapping cirrus sac; poral and aporal ovarian lobes in terminal mature proglottids 135 – 270 (219 ± 61; n = 5) and 140 – 275 (219 ± 60; n = 5) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 135 – 425 (258 ± 103; n = 8) and 155 – 450 (264 ± 108; n = 8) long, respectively. Maximum width of ovary 52 – 110 (78 ± 22; n = 13). Ovarian isthmus at or anterior to midpoint of ovary; poral lobe of ovary stopping 45 – 150 (81 ± 21; n = 13) short of genital pore. Mehlis’ gland posterior to ovarian isthmus, 30 – 45 (39 ± 7; n = 7) long by 25 – 37 (29 ± 4; n = 7) wide. Vitellarium follicular; vitelline follicles arranged in 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by terminal genitalia, and mostly interrupted by ovary (Fig. 6 A, C). Uterus ventral, sacciform, extending from near isthmus of ovary to near anterior margin of proglottid. T y p e a n d o n l y k n o w n h o s t: Himantura cf. pastinacoides (Myliobatiformes: Dasyatidae). Ty p e l o c a l i t y: Sulu Sea off Kampung Tetabuan (06 ° 01 ' 10 '' N; 117 ° 42 ' 15 '' E), Sabah, Malaysian Borneo (BO- 98, BO- 100, BO- 119). A d d i t i o n a l l o c a l i t i e s: Sulu Sea off Sandakan (05 ° 50 ' 20 '' N; 118 ° 07 ' 16 '' E), Sabah, Malaysian Borneo (BO- 79). South China Sea off Sematan (01 ° 48 ' 15 '' N; 109 ° 46 ' 47 '' E), Sarawak, Malaysian Borneo (BO- 168). S i t e o f i n f e c t i o n: Spiral intestine.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFDFFAADF958665FD9D2ED7.taxon	materials_examined	Ty p e m a t e r i a l: Holotype MZUM (P) No. 2016.11 (H). Paratypes: IPCAS No. C- 738; LRP Nos. 8995 – 8997; 9063 – 9082; 9149 (including molecular vouchers, sections and SEM specimens); MZUM (P) No. 2016.12 (P); SBC No. P- 00071; USNM Nos. 1420467 – 1420470.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFDFFAADF958665FD9D2ED7.taxon	etymology	E t y m o l o g y: This species is named in honour of Dr. William Campbell, Nobel laureate, for his role in the development of drugs to fight parasitic infections.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFDFFAADF958665FD9D2ED7.taxon	discussion	Remarks. Stillabothrium campbelli sp. n. is the only Stillabothrium species treated in this study that possesses a greater number of anterior loculi than posterior loculi on the bothridia. Stillabothrium campbelli can be distinguished from both S. ashleyae and S. davidcynthiaorum in the unique configuration of the septa and loculi on its bothridia. In terms of the anterior region of the bothridia, S. campbelli possesses a total of 10 – 12 loculi (Fig. 6 B) whereas both S. ashleyae and S. davidcynthiaorum each possess only 3 loculi (Figs. 2 B, 4 B). In the posterior region of the bothridia, S. campbelli possesses 5 loculi that are longer than wide whereas both S. ashleyae and S. davidcynthiaorum respectively possess 7 or 9 loculi that are longer than wide (Figs. 2 B, 4 B). In addition, the longitudinal septa in the posterior bothridia of S. campbelli extend anteriorly such that they overlap with the 3 – 4 posteriormost transverse septa of the anterior region of the bothridia. In S. ashleyae and S. davidcynthiaorum, the septa do not overlap. Scolex microtriches can also be used to distinguish S. campbelli from S. ashleyae and S. davidcynthiaorum. The posterior proximal surface of bothridia in S. campbelli possess a patch of coniform spinitriches (in addition to acicular filitriches throughout) near the rim, that is restricted in distribution, whereas the proximal bothridial surface of S. ashleyae and S. davidcynthiaorum is evenly covered with coniform spinitriches and acicular filitriches. Finally, proglottid morphology can also be used to distinguish S. campbelli from S. ashleyae and S. davidcynthiaorum. The cirrus sac of S. campbelli sp. n. is larger (45 – 90 µm long by 50 – 100 µm wide) than those of S. ashleyae (26 – 42 µm long by 26 – 47 µm wide) and of S. davidcynthiaorum (18 – 35 µm long by 24 – 38 µm wide), and it extends further across the midline (Figs. 6 C, 16 C), than in the latter two species (Figs. 2 C, 4 C, 16 A, B). Stillabothrium campbelli is reported from H. cf. pastinacoides from three different localities in Malaysian Borneo. It should be noted that the stingray host specimens of S. campbelli in this study, BO- 61, BO- 79, BO- 98, BO- 100, BO- 119 and BO- 168, are identified as H. cf. pastinacoides (Caira et al. 2012), but that identification awaits expert verification. Given that Naylor et al. (2012 b) showed that two species of stingray similar to H. pastinacoides (i. e. H. cf. pastinacoides 1 and H. cf. pastinacoides 2) co-occur in the region sampled, it is possible that S. campbelli sp. n. parasitises either or both. Additional specimens morphologically similar to S. campbelli were encountered in H. uarnacoides (BO- 118). Further studies are needed to determine whether Stillabothrium specimens from H. uarnacoides are conspecific with S. campbelli or represent an undescribed species of Stillabothrium. Four specimens of S. campbelli were included in the phylogenetic analysis (Fig. 1, Table 1).	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFAFFA8DF9584C5FB592D37.taxon	description	Figs. 1, 8, 9, 16 D ZooBank number for species: urn: lsid: zoobank. org: act: 97362 AA 0 - 762 D- 4 C 87 - B 5 ED- 0 A 3664 D 69 E 98 Description (based on whole mounts of 29 complete mature worms, cross sections of 2 strobila and longitudinal sections of 2 scoleces and 2 scoleces prepared for SEM): Worms (Fig. 8 A) euapolytic, acraspedote, 1.10 – 2.28 mm (1.64 ± 0.3; n = 29) long, greatest width 368 – 933 (601 ± 140; n = 29) at level of scolex; 5 – 9 (7.3 ± 1.1; n = 29) proglottids per worm. Cephalic peduncle lacking; small darkly staining germinative zone present. Scolex (Fig. 8 B) consisting of scolex proper bearing 4 stalked bothridia. Stalks 49 – 270 (119 ± 54; n = 22) long by 48 – 101 (73 ± 15; n = 27) wide, attached slightly posterior to middle of bothridia. Bothridia varying in shape with degree of contraction, from finely ovoid (Fig. 8 A), broadly ovoid (Fig. 8 B), to broadly deltoid (Fig. 9 A), facially loculated, 247 – 390 (319 ± 35; n = 24) long by 213 – 360 (288 ± 37; n = 24) wide; bothridial margins with thin rim. Anterior region of bothridia (Figs. 8 A, B, 9 B) with 6 – 8 horizontally oriented loculi (i. e. loculi wider than long) with 6 – 8 complete transverse septa (7.2 ± 0.5; n = 26). Anteriormost loculus 30 – 51 (40 ± 6; n = 27) long by 54 – 90 (71 ± 10; n = 29) wide. Posterior region of bothridia with 8 (n = 21) nonmedial longitudinal septa dividing bothridia into 9 loculi longer than wide; majority of longitudinal septa overlapping posteriormost 3 – 6 (4.3 ± 0.9; n = 13) transverse septa of anterior region of bothridia, resulting in a grid-like pattern of septa and loculi in the centre of bothridium (Fig. 8 A, B). Loculi and septa of distal bothridial surfaces (Fig. 9 B) bearing capilliform filitriches and coniform spinitriches. Proximal bothridial rim (Fig. 9 C, D) bearing capilliform filitriches (Fig. 9 B). Proximal bothridial surfaces away from rim bearing acicular filitriches throughout bothridium (Fig. 9 D, E, F). Posterior margin of proximal bothridial surfaces bearing a patch of coniform spinitriches (Fig. 9 C, E) near, but not extending to, bothridial rim. Bothridial stalks and strobila (Fig. 9 G) bearing capilliform filitriches only. Strobila with 2 – 5 (3.5 ± 0.9; n = 29) proglottids wider than long followed by 2 – 6 (3.9 ± 0.8; n = 29) proglottids longer than wide. Strobila widest at terminal proglottid; terminal proglottid 381 – 840 (601 ± 127; n = 29) long by 78 – 155 (12 ± 18; n = 29) wide; genital pore located 50 – 61 % (55 ± 3.6; n = 29) of proglottid length from proglottid posterior margin. Immature proglottids 4 – 8 (6.0 ± 1.1; n = 29) in number. Mature proglottids 1 – 2 (1.3 ± 0.5; n = 29) in number, including 0 – 1 (0.6 ± 0.5; n = 29) vas deferens-mature proglottid. Testes in mature proglottids 9 – 16 (13 ± 1.9; n = 28) in total number, 1 layer deep in section, arranged in 2 columns; columns extending from anterior margin of proglottid to anterior margin of cirrus sac, 23 – 57 (33 ± 7; n = 25) long by 27 – 49 (39 ± 7; n = 25) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending from level of ovarian isthmus to overlap posteriormost testes. Cirrus sac thin-walled, oval, extending medially past midline of proglottid; cirrus sac in terminal mature proglottid 45 – 69 (57 ± 6; n = 12) long by 38 – 60 (49 ± 7; n = 12) wide; cirrus sac in vas deferens-mature proglottids 52 – 79 (63 ± 8; n = 16) long by 45 – 63 (54 ± 5; n = 16) wide. Cirrus spinitriches present. Vagina (Figs. 8 C, 16 D) thick-walled, weakly sinuous, somewhat overlapping medial portion of cirrus sac (Fig. 8 A), extending along midline of proglottid from ootype region to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, H-shaped in frontal view, tetralobed in cross section. Ovarian lobes asymmetrical; poral ovarian lobe somewhat overlapping cirrus sac; poral and aporal ovarian lobes in terminal mature proglottids 125 – 201 (155 ± 28; n = 9) and 140 – 213 (165 ± 27; n = 9) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 175 – 290 (216 ± 27; n = 15) and 175 – 290 (228 ± 34; n = 15) long, respectively. Maximum width of ovary 68 – 114 (92 ± 12; n = 24). Ovarian isthmus at or anterior to midpoint of ovary; poral lobe of ovary stopping 30 – 80 (53 ± 14; n = 24) short of genital pore. Mehlis’ gland posterior to ovarian isthmus, 24 – 70 (40 ± 11; n = 20) long by 14 – 30 (21 ± 5; n = 18) wide. Vitellarium follicular, 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by terminal genitalia, and interrupted by ovary (Fig. 8 A, C). Uterus ventral, sacciform, extending from posterior margin of proglottid to near anterior margin of proglottid. T y p e a n d o n l y k n o w n h o s t: Pastinachus solocirostris Last, Manjaji et Yearsley, Roughnose stingray (Myliobatiformes: Dasyatidae). Ty p e l o c a l i t y: South China Sea off Mukah (02 ° 53 ' 52 '' N; 112 ° 05 ' 44 '' E), Sarawak, Malaysian Borneo (BO- 267). A d d i t i o n a l l o c a l i t i e s: Java Sea off Singkawang (Pasar Bringin) (00 ° 55 ' 06 '' N; 108 ° 58 ' 58 '' E), West Kalimantan, Indonesian Borneo (KA- 148). S i t e o f i n f e c t i o n: Spiral intestine.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFAFFA8DF9584C5FB592D37.taxon	materials_examined	Ty p e m a t e r i a l: Holotype MZUM (P) No. 2016.13 (H). Paratypes: IPCAS No. C- 741; LRP Nos. 9003 – 9004; 9083 – 9117 (including molecular vouchers, sec-tions, and SEM specimens); MZUM (P) No. 2016.14 (P) – 2016.15 (P); SBC No. P- 00072; USNM Nos. 1420471 – 1420477.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFAFFA8DF9584C5FB592D37.taxon	etymology	E t y m o l o g y: This species is named for the grid-like pattern of septa in the centre of the bothridium. The name is a combination of the Latin ‘ hyphantos’, meaning woven, and the Latin ‘ septum’.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFFAFFA8DF9584C5FB592D37.taxon	discussion	Remarks. The most striking characteristic of S. hyphantoseptum sp. n. is the extensive grid-like appearance of the septa on the centre of the bothridia (Fig. 8 A, B). This configuration results from the extensive overlap of transverse (anterior) septa and longitudinal (posterior) septa. This feature readily distinguishes S. hyphantoseptum from S. ashleyae and S. davidcynthiaorum, both of which lack overlapping septa. The only other described species of Stillabothrium with overlapping transverse and longitudinal septa is S. campbelli (Fig. 6 A, B). Although the centre of bothridia of S. campbelli is grid-like, the grid-like area is less extensive than in S. hyphantoseptum (compare Figs. 6 B and 8 B). In S. campbelli, the posteriormost 3 – 4 transverse septa are overlapped by its central 2 longitudinal septa, whereas in S. hyphantoseptum the posteriormost 3 – 6 transverse septa are overlapped by most of its eight longitudinal septa. The number of loculi that are wider than long in the anterior region of bothridia is also different between the two species (6 – 8 in S. hyphantoseptum vs 10 – 12 in S. campbelli). In terms of proglottid morphology, S. hyphantoseptum sp. n. can be distinguished from S. ashleyae, S. davidcynthiaorum and S. campbelli in its possession of a uterus that extends to the posterior margin of the proglottid (Fig. 8 C), whereas in the latter three species the uterus only extends posteriorly to the ovarian isthmus (Figs. 4 C, 6 C; not illustrated for S. ashleyae, but observed). Stillabothrium hyphantoseptum is the only species of Stillabothrium reported from a species of Pastinachus Rüppell. The host species, P. solocirostris (roughnose stingray) was described by Last et al. (2005) as part of the same survey work that made the current study possible. Two specimens of S. hyphantoseptum were included in the phylogenetic analysis (Fig. 1, Table 1).	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF8FFB5DC2F8765FD6C2E37.taxon	description	Figs. 1, 10, 11, 16 E ZooBank number for species: urn: lsid: zoobank. org: act: D 2 B 25027 - 058 D- 4 E 3 D-B 880 - F 1 FA 5 AB 763 DB Description (based on whole mounts of 20 complete mature worms, cross sections of 1 strobila and longitudinal sections of 1 scolex and 2 scoleces prepared for SEM): Worms (Fig. 10 A) euapolytic, acraspedote, 1.7 – 3.9 mm (2.5 ± 0.6; n = 20) long, greatest width 428 – 822 (588 ± 121; n = 20) at level of scolex; 5 – 10 (7.1 ± 1.2; n = 20) proglottids per worm. Cephalic peduncle lacking, small darkly staining germinative zone present. Scolex (Fig. 10 B) consisting of scolex proper bearing four stalked bothridia. Stalks 30 – 125 (77 ± 25; n = 16) long by 65 – 140 (94 ± 18; n = 16) wide, attached slightly posterior to middle of bothridia. Bothridia (Figs. 10 A, B, 11 A) always constricted but varying in shape with degree of contraction, broadly (Fig. 10 B), shallowly (Fig. 10 A), very shallowly (Fig. 11 A), or depressed deltoid, facially loculated, 218 – 305 (259 ± 21; n = 20) long by 255 – 460 (339 ± 58; n = 20) wide. Anterior region of bothridia (Figs. 10 A, B, 11 A) with 3 horizontally oriented loculi (i. e. loculi wider than long) with 3 complete complete transverse septa (n = 20). Anteriormost loculus 35 – 60 (46 ± 7; n = 18) long by 64 – 115 (81 ± 16; n = 18) wide. Posterior region of bothridia (Figs. 10 A, B, 11 A) with 6 (n = 20) nonmedial longitudinal septa dividing bothridia into 7 loculi longer than wide; longitudinal septa not overlapping transverse septa in anterior region of bothridia. Loculi and septa of distal bothridial surfaces (Fig. 11 B) bearing coniform spinitriches and, less conspicuously, capilliform filitriches. Proximal bothridial rim (Fig. 11 C) bearing capilliform filitriches greater in length than those on distal bothridial surfaces (Fig. 11 B). Proximal bothridial surfaces away from rim bearing densely arranged capilliform filitriches throughout bothridium (Fig. 11 E). Proximal bothridial surfaces near bothridial rim bearing coniform spinitriches (Fig. 11 D) that do not extend to stalks. Bothridial stalks and strobila (Fig. 11 F) bearing capilliform filitriches only. Strobila with 2 – 5 (3.2 ± 0.8; n = 20) proglottids wider than long followed by 3 – 7 (3.9 ± 1.0; n = 20) proglottids longer than wide. Strobila widest at terminal proglottid; terminal proglottid 724 – 1 260 (1 013 ± 185; n = 20) long by 92 – 192 (132 ± 24; n = 20) wide; genital pore located 42 – 56 % (48 ± 4; n = 20) of proglottid length from proglottid posterior margin. Immature proglottids 4 – 8 (5.8 ± 1.0; n = 20) in number. Mature proglottids 1 – 2 (1.4 ± 0.5; n = 20) in number, including 0 – 1 (0.3 ± 0.5; n = 20) vas deferens-mature proglottid. Testes in mature proglottids 20 – 29 (23 ± 2; n = 20) in total number, in 1 layer deep in section, arranged in 2 columns (Fig. 10 C), columns extending from anterior margin of proglottid to anterior margin of cirrus sac, 26 – 52 (38 ± 8; n = 16) long by 29 – 55 (42 ± 7; n = 16) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending from level of ovarian isthmus anteriorly to overlap posteriormost testes. Cirrus sac thin-walled, oval, extending medially past midline of proglottid. Cirrus sac in terminal mature proglottid 55 – 81 (72 ± 8; n = 15) long by 47 – 73 (59 ± 8; n = 15) wide; cirrus sac in vas deferens-mature proglottids 80 – 87 (84 ± 3; n = 5) long by 62 – 80 (72 ± 7; n = 5) wide. Cirrus spinitriches present. Vagina (Fig. 10 C) thick-walled, non-sinuous, somewhat overlapping medial portion of cirrus sac (Fig. 10 C), extending along midline of proglottid from ootype region to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, H-shaped in frontal view, tetralobed in cross section; ovarian lobes somewhat asymmetrical; poral and aporal ovarian lobes in terminal mature proglottids 238 – 396 (304 ± 47; n = 12) and 265 – 394 (318 ± 45; n = 12) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 355 – 486 (412 ± 58; n = 6) and 370 – 488 (434 ± 48; n = 6) long, respectively. Maximum width of ovary 60 – 122 (91 ± 19; n = 17). Ovarian isthmus at or posterior to midpoint of ovary; poral lobe of ovary stopping 18 – 70 (51 ± 14; n = 17) short of genital pore. Mehlis’ gland well posterior to ovarian isthmus, 32 – 72 (47 ± 13; n = 16) long by 22 – 35 (28 ± 4; n = 16) wide. Vitellarium follicular; vitelline follicles arranged in 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by terminal genitalia and mostly interrupted by ovary (Fig. 10 A, C). Uterus ventral, sacciform extending from near ovarian isthmus to near anterior margin of proglottid. T y p e a n d o n l y k n o w n h o s t: Himantura australis Last, White et Naylor, Reticulate whipray (Dasyatidae: Myliobatiformes). T y p e a n d o n l y k n o w n l o c a l i t y: Gulf of Carpentaria (Indian Ocean) off Weipa (12 ° 35 ' 11 '' S; 141 ° 42 ' 34 '' E), Queensland, Australia (CM 03 - 3, CM 03 - 25, CM 03 - 65). S i t e o f i n f e c t i o n: Spiral intestine.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF8FFB5DC2F8765FD6C2E37.taxon	materials_examined	Ty p e m a t e r i a l: Holotype QM No. G 235198. Paratypes: LRP Nos. 8999; 9118 – 9133 (including molecular vouchers, sections, and SEM specimens); QM Nos. G 235199 – G 235200; USNM Nos. 1420478 – 1420483; IPCAS No. C- 740.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF8FFB5DC2F8765FD6C2E37.taxon	etymology	E t y m o l o g y: This species is named in loving memory of Jean Forti, mother of K. S. Forti, for her support of her daughter’s education.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFF8FFB5DC2F8765FD6C2E37.taxon	discussion	Remarks. Stillabothrium jeanfortiae sp. n. is the fifth new Stillabothrium species described in this study. Its configuration of bothridial septa is unique relative to its four described congeners. Stillabothrium jeanfortiae is distinguished from S. ashleyae and S. davicynthiae in that it lacks, rather than possesses, marginal septa in the posterior region of bothridia. It also differs from S. ashleyae and S. davicynthiae in that the three loculi in its anterior region of bothridia are oriented in tandem, instead of occurring as a row of one and then two loculi. Stillabothrium jeanfortiae can be distinguished from S. campbelli and S. hyphantoseptum in that it lacks, rather than posseses, septa that overlap one another. Stillabothrium jeanfortiae is described from the stingray Himantura australis from the Gulf of Carpentaria in northern Australia. Specimens similar in morphology to S. jeanfortiae were encountered in H. leoparda (NT- 117), also from the Gulf of Carpentaria, but confirmation of the identity of this material awaits further study. Specimens with a similar bothridial morphology to that of S. jeanfortiae were also encountered in a diversity of species of Himantura during survey work in Borneo. Those specimens can be readily distinguished from S. jeanfortiae based on proglottid morphology, but their taxonomic treatment also awaits further study. One specimen of S. jeanfortiae was included in the phylogenetic analysis (Fig. 1, Table 1). It grouped with two undescribed species of Stillabothrium from Senegal.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE5FFB2DF958465FC322D97.taxon	description	I n f o r m a l s y n o n y m s: Rhinebothriinae New genus 3 cadenati of Healy et al. (2009), Caira et al. (2014), Ruhnke et al. (2015), Marques and Caira (2016). ZooBank number for species: urn: lsid: zoobank. org: act: 9274 C 612 - 40 F 1 - 4 F 66 - 839 C-DE 145 BC 0 C 01 C Redescription (based on specimens collected from Z. schoenleinii consisting of whole mounts of 6 complete mature worms, 5 strobilae and 2 scoleces; cross sections of 1 strobila and longitudinal sections of 2 scoleces [including 1 in situ scolex], and 1 specimen prepared for SEM): Worms euapolytic, slightly craspedote (Fig. 12 A), 1.04 – 1.77 mm (1.35 ± 0.3; n = 6) long, greatest width 439 – 694 (568 ± 96; n = 7) at level of scolex; 5 – 7 (5.7 ± 0.8; n = 6) proglottids per worm. Cephalic peduncle lacking; small darkly staining germinative zone present. Scolex (Figs. 12 A, 14 A) consisting of scolex proper bearing four stalked bothridia. Stalks 39 – 128 (86 ± 33; n = 8) long by 40 – 77 (60 ± 13; n = 8) wide, attached slightly posterior to middle of bothridia. Bothridia (Figs. 12 A, B, 14 A, B) varying in shape with degree of contraction, deeply (Fig. 12 A), finely deltoid or broadly deltoid (Figs. 12 B, 14 A, B), facially loculated, 285 – 378 (328 ± 38; n = 6) long by 201 – 355 (274 ± 64; n = 7) wide. Anterior region of bothridia (Figs. 12 A, B) with 3 horizontally oriented loculi (i. e. loculi wider than long) with 3 – 4 (3.8 ± 0.4; n = 6) complete transverse septa; fourth complete but reduced transverse septum (Fig. 12 B) observed in 5 of 6 specimens. Anteriormost loculus 58 – 105 (77 ± 14; n = 8) long and 74 – 118 (108 ± 14; n = 8) wide. Posterior region of bothridia (Fig. 12 A, B) with 4 (n = 8) nonmedial longitudinal septa dividing bothridia into 5 loculi longer than wide; longitudinal septa in posterior region of bothridia overlapping posteriormost (fourth) transverse septum in anterior region of bothridia (Fig. 12 B). Muscle fibres other than septa but parallel to bothridial surface also observed on bothridia (Fig. 12 A). All septa appear as ridges in section with proximal and distal portions different (Fig. 13); proximal portion of septa formed by underlying bothridial wall, consisting of radial muscles oriented with proximal ends of fibres adjacent to each other; distal portion of septa formed by separate muscle bundle; proximal and distal portions of septa separated by a triangular gap. Loculi (Fig. 14 D) and septa (Fig. 14 E) of distal bothridial surfaces bearing coniform spinitriches and capilliform filitriches; spinitriches lacking on distal bothridial margin and rim (Fig. 14 C). Proximal bothridial rim (Fig. 14 C) bearing capilliform filitriches greater in length than those on distal bothridial surfaces (Fig. 14 D, E). Proximal bothridial surfaces away from rim and stalks bearing coniform spinitriches and capilliform filitriches (Fig. 14 F, G). Strobila bearing capilliform filitriches only. Strobila with 3 – 4 (3.2 ± 0.4; n = 6) proglottids wider than long followed by 2 – 4 (2.5 ± 0.8; n = 6) proglottids longer than wide. Strobila widest at terminal proglottid; terminal proglottid 467 – 1 005 (n = 11) long by 103 – 139 (n = 11) wide; genital pore located 64 – 73 % (n = 11) of proglottid length from proglottid posterior margin. Genital atrium expansive, 38 – 68 (51 ± 10; n = 11) long by 31 – 49 (40 ± 7; n = 11) wide, with convoluted, muscular walls. Immature proglottids 4 – 5 (4.5 ± 0.5; n = 6) in number. Mature proglottids 1 – 2 (1.4 ± 0.5; n = 11) in number, including 0 – 1 (0.7 ± 0.5; n = 11) vas deferens-mature proglottids. Testes in mature proglottids 7 – 13 (10.8 ± 2; n = 12) in total number, 1 layer deep in section, arranged in 1 – 3 (2 ± 0.4; n = 11) though usually 2, columns (Fig. 12 C); columns extending from near anterior margin of proglottid to anterior margin of vagina, 22 – 36 (30 ± 4; n = 11) long by 29 – 50 (38 ± 6; n = 11) wide. Vas deferens coiled, entering anterior margin of cirrus sac, extending anterior to ovarian isthmus to near anterior vagina. Cirrus sac thick-walled and relatively large, oval and bent posteriorly, extending medially past midline of proglottid; cirrus sac in terminal mature proglottid 85 – 152 (119 ± 47; n = 2) long by 58 – 62 (60 ± 3; n = 2) wide; cirrus sac in vas deferens-mature proglottids 103 – 171 (136 ± 23; n = 8) long by 50 – 75 (67 ± 8; n = 8) wide. Cirrus spinitriches present, 3.5 – 6.1 (4.3 ± 0.9; n = 10) long by 2.0 – 3.7 (2.8 ± 0.5; n = 10) wide at base. Vagina (Fig. 12 A, C) thick-walled, sinuous, not overlapping cirrus sac, recurved (Figs. 12 C, 16 F); poral portion expanded; medial and posterior portion abruptly narrowed; extending from ootype past midline to aporal side of proglottid to anterior margin of cirrus sac, then laterally to open into genital atrium anterior to cirrus sac; vaginal sphincter absent. Seminal receptacle present. Ovary near posterior end of proglottid, lobulated, H-shaped in frontal view, tetralobed in cross section, not overlapping cirrus sac; ovarian lobes asymmetrical; poral and aporal ovarian lobes in terminal mature proglottids 125 – 215 (170 ± 64; n = 2) and 125 – 225 (175 ± 71; n = 2) long, respectively. Poral and aporal ovarian lobes in vas deferens-mature proglottids 120 – 350 (222 ± 71; n = 9) and 125 – 350 (218 ± 69; n = 9) long, respectively. Maximum width of ovary 48 – 87 (62 ± 13; n = 10). Ovarian isthmus at or anterior to midpoint of ovary; poral lobe of ovary stopping 90 – 190 (145 ± 35; n = 10) short of genital pore. Mehlis’ gland posterior to ovarian isthmus, 25 – 40 (33 ± 5; n = 10) long by 16 – 25 (20 ± 3; n = 10) wide. Vitellarium follicular; vitelline follicles arranged in 1 dorsal and 1 ventral column on each side of proglottid; columns extending from anterior to posterior margin of proglottid, interrupted by terminal genitalia, not interrupted by ovary (Fig. 12 A, C). Uterus ventral, sacciform, extending from posterior margin of proglottid to near anterior margin of proglottid. T y p e h o s t: Zanobatus schoenleini [sic!] (= schoenleinii) (Müller et Henle), striped panray (Rhinopristiformes: Zanobatidae). A d d i t i o n a l h o s t: Rhinobatos rhinobatos (Linnaeus), Common guitarfish (Rhinopristiformes: Rhinobatidae). Ty p e l o c a l i t y: Gorée Island, Senegal. A d d i t i o n a l l o c a l i t i e s: Atlantic Ocean off Ouakam (14 ° 42 ' 54 '' N; 17 ° 29 ' 28 '' W) (SE – 28) and Soumbédioune (14 ° 40 ' 42 '' N; 17 ° 27 ' 42 '' W) (SE – 201) in Dakar, and Joal (14 ° 10 ' 30 '' N; 16 ° 51 ' 12 '' W) (SE – 299) and Kafountine (12 ° 55 ' 41 '' N; 16 ° 45 ' 10 '' W) (SE – 289), Senegal. S i t e o f i n f e c t i o n: Spiral intestine.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE5FFB2DF958465FC322D97.taxon	type_taxon	Ty p e m a t e r i a l: Unknown.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE5FFB2DF958465FC322D97.taxon	materials_examined	M a t e r i a l e x a m i n e d (v o u c h e r s): LRP Nos. 9000 – 9002; 9134 – 9148 (including molecular vouchers, sections and SEM specimens); MNHN Nos. HEL 580 – HEL 581; USNM No. 1420484.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE5FFB2DF958465FC322D97.taxon	discussion	Remarks. Type specimens of S. cadenati comb. n. were not available for this study. In the original description of S. cadenati, Euzet (1954) did not provide specimen deposition information. It is possible that the specimens were in the personal collection of the late Louis Euzet, a collection now transferred to the Museum National d’Histoire Naturelle (MNHN) in Paris, France. Examination of that collection by the curator, however, did not reveal the presence of any specimens of S. cadenati. The material on which this redescription was based were newly collected, topotypic specimens of S. cadenati The specimens were collected from the type host at Ouakam and Soumbédioune on the west coast of Dakar, Senegal, approximately 12 and seven km, respectively away from Gorée island, the type locality of this species. Gorée lies 2.5 km off the eastern coast of Dakar. Specimens of S. cadenati were also obtained from the type host in Joal, Senegal. Two specimens of S. cadenati were included in the phylogenetic analysis (Fig. 1, Table 1). One was from the type host, Z. schoenleinii, and one was from R. rhinobatos (the common guitarfish). Given that R. rhinobatos and the type host, Z. schoenleinii, belong to different orders of elasmobranchs, this host record requires verification, ideally by the examination of additional specimens. The bothridial morphology of S. cadenati was not comprehensively described by Euzet (1954). While he noted the presence of 3 transverse septa, Euzet (1954) stated that the bothridia of the specimens he examined were tightly folded and clamped in four scallops. Subsequently, when Euzet and co-authors (Ball et al. 2003) erected the genus Scalithrium Ball, Neifar et Euzet, 2003, for species of Rhinebothrium that lack a median longitudinal septum, they concluded that the scolex morphology of S. cadenati was too poorly known to allow them to confidently place it in their new genus at that time. The work conducted here enabled comprehensive characterisation of the bothridial morphology of S. cadenati for the first time. A fourth, less muscular transverse septum (Figs. 12 B, 13) lies posterior to the three transverse septa noted by Euzet (1954), and this reduced transverse septum is crossed by some, but not all, non-medial longitudinal septa (Fig. 12 B). We believe the scalloped appearance in the posterior part of the bothridium of specimens with folded bothridia noted by Euzet (1954) is due to the presence of these non-medial longitudinal septa. This redescription also provides additional data on the reproductive morphology and microthrix distribution patterns of this species. Some measurements reported by Euzet (1954) suggest that his specimens of S. cadenati were somewhat larger than those examined here. For example, Euzet’s (1954) specimens were 4 – 6 mm long in total length and possessed 13 – 16 proglottids, whereas the specimens examined here were 1.04 – 1.77 mm long and possessed 5 – 7 proglottids. In spite of these differences, we believe that the specimens we obtained for the current study are conspecific with those examined by Euzet (1954), based on the illustrations he provided, and considering that that the specimens we used for this study are from the same host, and within 11 km of the type locality. The bothridia of S. cadenati are unlike any previously described Stillabothrium species in that they possess septa that differ in thickness. In S. ashleyae, S. davidcynthiaorum, S. campbelli, S. hyphantoseptum and S. jeanfortiae, all transverse and longitudinal septa appear uniform in thickness. In S. cadenati the 3 anteriormost transverse septa are thicker than the fourth transverse septa and the longitudinal septa (Fig. 12 B). Stillabothrium cadenati can also be distinguished from each of its five congeners in its conspicuously different proglottid morphology. Unlike those of its congeners, the genital atrium in S. cadenati has convoluted walls that are more muscular in appearance (compare Fig. 12 C with Figs. 2 C, 4 C, 6 C, 8 C, 10 C), and the genital pore is located more anteriorly in S. cadenati (64 – 73 % of proglottid length from proglottid posterior margin) than it is in S. ashleyae, S. davidcynthiaorum, S. campbelli, S. hyphantoseptum and S. jeanfortiae (35 – 46 %, 42 – 54 %, 41 – 55 %, 50 – 61 % and 42 – 56 %, respectively). In S. cadenati, the vitellarium (Fig. 12 C) occurs along the length of the ovary, whereas in S. ashleyae, S. davidcynthiaorum, S. campbelli, S. hyphantoseptum and S. jeanfortiae, the vitelline columns are interrupted by the ovary (Figs. 2 C, 4 C, 6 C, 8 C, 10 C). The cirrus sac is relatively larger in S. cadenati when compared to its 5 congeners (compare Fig. 12 C to Figs. 2 C, 4 C, 6 C, 8 C, 10 C), and reaches a greater length (103 – 171 µm vs 28 – 38 µm, 32 – 45 µm, 45 – 90 µm, 52 – 79 µm and 80 – 87 µm, respectively). Stillabothrium cadenati is the only species of the genus reported from the Atlantic Ocean. Specimens that appear to represent at least two additional species of Stillabothrium were encountered in stingrays of the genus Fontitrygon Last, Naylor et Manjaji-Matsumoto over the course of survey work in Senegal. These remain to be examined in detail. Four specimens of S. cadenati were included in the phylogenetic analysis (Fig. 1, Table 1). One of the four specimens (LRP 3924) was previously included in the analysis of Healy et al. (2009) as ‘ Rhinebothriinae New genus 3 cadenati ’.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE2FFB1DC2F8405FC482AB7.taxon	description	Figs. 1, 15 ZooBank number for species: urn: lsid: zoobank. org: act: DFCD 1070 - 9190 - 4890 - ADEE- 5474390 DB 2 BF	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE2FFB1DC2F8405FC482AB7.taxon	description	T y p e a n d o n l y k n o w n h o s t: Glaucostegus typus [Anonymous (Bennett)], Giant shovelnose ray (Rhinopristiformes: Rhinobatidae) (= Rhinobatos armatus). Ty p e l o c a l i t y: Moreton Bay, Northern Territory, Australia. A d d i t i o n a l l o c a l i t y: Fog Bay, Timor Sea (Indian Ocean) off Dundee Beach (12 ° 45 ' 33 '' S; 130 ° 21 ' 7 '' E), Queensland, Australia (AU- 56). S i t e o f I n f e c t i o n: Spiral intestine.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE2FFB1DC2F8405FC482AB7.taxon	materials_examined	S p e c i m e n s e x a m i n e d: AU- 56, QM GL 4621 (holotype), GL 4622 (paratype).	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
03EB87AEFFE2FFB1DC2F8405FC482AB7.taxon	discussion	Remarks. Butler (1987) provided a comprehensive description of this species, which included scanning electron micrographs and drawings, and represents the only treatment of this species, to date. Although examination of the holotype for the present study confirmed many of Butler’s (1987) observations, it refuted some of her observations, and revealed additional details missing from the original description. According to Butler (1987), the anterior region of the bothridia lack transverse septa. However, at least 4 transverse septa are visible in the anterior region of the scolex of this holotype (Fig. 15). These septa, and other septa not described by Butler (1987), are best visible using differential interference contrast microscopy (DIC). Examination of additional material of this species and, ideally, histological sections of the scolex, would be very useful in determining the exact number and extent of the septa (and the loculi) in this species. Not stated in the description, but visible in the holotype, are the following features: the vas deferens joins the cirrus sac at the middle of its anterior margin; the genital atrium has internally convoluted margins; when at least partially expanded with spermatozoa, the vas deferens extends from between the anterior lobes of the ovary to the level of the recurvature of the vagina. The morphology of the ovary, posterior vitelline follicles and posterior part of the uterus as drawn by Butler (1987) differ from that observed in the terminal proglottid of the holotype: the ovary is tetralobed and H-shaped; vitellarium is interrupted at the level of the ovary and some vitelline follicles are present in the lateral margins of the proglottid, posterior to the ovary; uterus extends posterior to the ovary. This species possesses transverse septa and non-medial longitudinal septa, as well as other characteristics consistent with its placement in Stillabothrium, such as a posterior row of loculi that are longer than wide (Fig. 15). Thus, this species is herein transferred into this genus as Stillabothrium amuletum comb. n. Stillabothrium amuletum possesses diagonal septa on the sides of the posterior bothridia, which overlap several longitudinal septa (Fig. 15). This feature distinguishes it from S. ashleyae (Fig. 2 B) and S. davidcynthiaorum (Fig. 4 B), and from S. campbelli (Fig. 6 B), which possess marginal (S. ashleyae and S. davidcynthiaorum) or diagonal (S. campbelli) septa that abut but do not overlap longitudinal septa, and from S. hyphantoseptum (Fig. 8 B), S. jeanfortiae (Fig. 10 B) and S. cadenati (Fig. 12 B) which each lack marginal or diagonal septa. One sequence of Stillabothrium from G. typus, the type host of S. amuletum, was included in the phylogenetic analysis in this study (Table 1, Fig. 1). It was from Healy et al. (2009), who referred to it as ‘ Rhinebothriinae New genus 3 sp. n. 7 ’.	en	Reyda, Florian B., Healy, Claire J., Haslach, Andrew R., Ruhnke, Timothy R., Aprill, Tara L., Bergman, Michael P., Daigler, Andrew L., Dedrick, Elsie A., Delgado, Illari, Forti, Kathryn S., Herzog, Kaylee S., Russell, Rebecca S., Willsey, Danielle D. (2016): A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. Folia Parasitologica (038) 63 (38): 1-28, DOI: 10.14411/fp.2016.038, URL: http://dx.doi.org/10.14411/fp.2016.038
