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FIGURE 3 in Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

FIGURE 3. Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of Benedeniella species. A. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. macrocolpa (Lühe, 1906) Yamaguti, 1963 showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1). B. The left anterior attachment organ of B. macrocolpa showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1). C. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. posterocolpa (Hargis, 1955) Yamaguti, 1963 showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13). D. The left anterior attachment organ of B. posterocolpa showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm.

FIGURE 4 in Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

FIGURE 4. Scanning electron micrographs of dorsal region of juvenile specimen of Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated. B. Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv). C. Enlargement of Figure 4B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm.

FIGURE 2 in Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3A). Scale bars: A–B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm.

FIGURE 5 in Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

FIGURE 5. Diagrammatic outlines of adult Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963 (A) and B. posterocolpa (Hargis, 1955) Yamaguti, 1963 (B) in dorsal view showing relative positions of paired anterior tapering horns (dh), common genital papilla (cgp; absent in B. posterocolpa), paired excretory papillae (ep) and paired conical structures (co). Drawn from several wholemounted specimens viewed dorsally. Scale bar: 4 mm.

FIGURE 1 in Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition

FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm.

FIGURE 4 in Review of the Capsalinae (Monogenea: Capsalidae)

FIGURE 4. Light photomicrograph of posterior margin of anterior attachment organ of Capsala laevis (CNHE 2738) showing finger-like projections. Scale bar: 100µm.

FIGURE 2 in Review of the Capsalinae (Monogenea: Capsalidae)

FIGURE 2. Light photomicrographs of dorsomarginal body sclerites. A. Capsala notosinense (BMNH 1993.5.18.2). B. Capsala martinierei (USNPC 8153). C. Capsala gregalis (USNPC 61642). D. Capsala biparasitica (USNPC 38134). E. Capsala poeyi (USNPC 38131). F. Capsala pricei (USNPC 37709); inner row of larger multicuspid sclerites indicated by arrowheads. G. Capsala magronum (USNPC 63595). H. Tristoma coccineum (USNPC 35645). I. Tristoma integrum (HWML 37935). J. Capsaloides cristatus (USNPC 63597). Scale bars: A, E, G, I, J = 50µm; B, F, H = 100µm; C = 10µm; D = 20µm.

FIGURE 3 in Review of the Capsalinae (Monogenea: Capsalidae)

FIGURE 3. Light photomicrograph of position of common genital pore in 2 closely related capsaline species. A. Capsala gregalis (USNPC 61642); common genital pore (arrowhead) opens on body margin a short distance from left anterior attachment organ. B. Capsala manteri (USNPC 37229); common genital pore (arrowhead) opens directly posterior to left anterior attachment organ. Scale bars: 200µm.

FIGURE 1 in Review of the Capsalinae (Monogenea: Capsalidae)

FIGURE 1. Light photomicrographs of capsaline haptors. A. Capsala magronum (USNPC 63595) showing large accessory sclerites and large papillae distributed on ventral surface of haptor and septa. Note that papillae are absent on the posterior half of the anterior 4 loculi. B. Finger-like fringe (arrowhead) on haptoral marginal valve of Capsala magronum. C. Capsala onchidiocotyle (USNPC 80214) with small papillae distributed across entire ventral surface of haptor and septa. D. Capsaloides sinuatus (CNHE 2739) showing bifid posterior pair of septa (arrow). Scale bars: A, C, D, = 500µm; B = 100µm.

FIG. 16 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 16. Haptors of Yamaguti's Hawaiian Benedenia species and B. epinepheli drawn to same scale from various specimens. (A) Benedenia bodiani; (B) B. epinepheli; (C) B. hawaiiensis; (D) B. lolo; (E) B. scari. Scale bar 5 250 Mm. Note tendon (te).

FIG. 14 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 14. Haptor and sclerites of Benedenia bodiani: (A) haptor; (B) accessory sclerite; (C) anterior hamulus; (D) posterior hamulus, all drawn from holotype; (E) haptor of one specimen of several from MPM: No. 15392; (F), (G), (H), anterior hamuli from various paratypes on USNPC: No. 63586 showing considerable morphological variation between individuals. Scale bar for haptors (A, E) 5 150 Mm. Scale bar for sclerites (B, C, D, F, G, H) 5 35 Mm.

FIG. 13 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 13. Body outline of entire Benedenia bodiani in ventral view showing widely spaced testes (t), large glands of Goto (G) and wavy muscles (wm) in the haptor. Scale bar 5 500 Mm. Composite drawn from several specimens.

FIG. 15 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 15. Male copulatory complex and terminal portion of vagina (v) of Benedenia bodiani. Composite drawn from several specimens. Scale bar 5 100 Mm. agr, accessory gland reservoir; da, duct of accessory gland reservoir; pe, penis; vd, vas deferens.

FIG. 11 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 11. Orientation of co-copulants during sperm exchange as observed for (A) Entobdella soleae (after Kearn, 1970); scale bar 5 1 mm. (B) Likely orientation of co-copulants for a species of Benedenia with a vaginal pore anterior to the common genital pore (based on B. lutjani); scale bar 5 500 Mm. Note that the probable orientation by which benedeniines with a vaginal pore opening anterior to the common genital pore may align themselves is likely to be similar to that of species which possess a vaginal pore that opens posterior to the common genital pore, such as E. soleae.

FIG. 12 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 12. Comparative ®gure showing positions of the vaginal pore (vp) of six Benedenia species demonstrating the continuum of variation from (A) B. ovata in which the vaginal pore is located in the mid-body between the germarium and the common genital pore to (F) where the vaginal pore in B. lutjani is located well anterior to the common genital pore. The other drawings depict gradations between these two extremes for: (B) Benedenia sp. of Grutter (1998); (C) B. epinepheli; (D) B. monticellii and (E) B. acanthopagr i. Scale bar 5 500 Mm. All drawings made from material we examined.

FIG. 10 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 10. Anterior ends of Benedenia species with a vaginal pore (vp) anterior to the common genital pore. These ®gures also show lobes (l) associated with the genital openings. (A) Benedenia acanthopagr i; (B) B. anticavaginat a; (C) B. lutjani. Scale bar 5 500 Mm.

FIG. 9 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 9. Haptors and haptoral sclerites of Benedenia species possessing a vagina that opens anterior to the common genital pore. Benedenia acanthopagr i: (A) haptor; (B) accessory sclerite; (C) anterior hamulus; (D) posterior hamulus. Benedenia anticavaginata: (E) haptor; (F) accessory sclerite; (G) anterior hamulus; (H) posterior hamulus with associated musculature (m). Benedenia lutjani: (I) haptor; (J) accessory sclerite; (K) anterior hamulus; (L) posterior hamulus. Scale bar for haptors (A, E, I) 5 250 Mm. Scale bar for sclerites (B, C, D, F, G, H, J, K, L) 5 100 Mm.

FIG. 7 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 7. Scanning electron micrographs of Benedenia sciaenae. (A). Ventral view of left anterior attachment organ (aa) with the male pore (mp) and the uterine pore (up) on the left ventral margin of the parasite. Scale bar 5 300 Mm. (B). Enlargement of left ventral margin showing the male pore (mp) and the uterine pore (up). Scale bar 5 50 Mm. (C). Dorsal view of left margin of parasite showing the vaginal pore (vp). Scale bar 5 100 Mm. (D). Enlargement of the dorsal vaginal pore (vp). Scale bar 5 10 Mm.

FIG. 5 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 5. Haptoral sclerites of Benedenia sciaenae drawn from various specimens. (A) Accessory sclerite; (B) anterior hamulus; (C) posterior hamulus. Scale bar 5 200 Mm. (D) Hooklet. Scale bar 5 10 Mm.

FIG. 4 in A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae)

FIG. 4. Benedenia sciaenae (van Beneden, 1856) Odhner, 1905, whole adult animal, ventral view. Gut not shown. Scale bar 5 1 mm. aa, anterior attachment organ; agr, accessory gland reservoir; ah, anterior hamulus; as, accessory sclerite; e, pigmented eye; g, germarium; G?, possible glands of Goto?; h, haptor; ho, hooklet; m, marginal valve; mp, male genital pore (i.e. opening of penis canal); oo, ootype; p, pharynx; ph, posterior hamulus; rt, ridge of tissue; t, testis; te, tendon; up, uterine pore; vd, vas deferens; vf, vitelline follicle; vp, vaginal pore; vr, vitelline reservoir. Drawn from voucher specimen QM: No. G 217921.