Benedenia

Deveney, Marty R. & Whittington, Ian D., 2010, Three new species of Benedenia Diesing, 1858 from the Great Barrier Reef, Australia with a key to species of the genus, Zootaxa 2348, pp. 1-22 : 16-17

Key to the described species of Benedenia

This key is accurate for the 25 currently described species of Benedenia . Numerous undescribed species of the genus exist and it is possible that many are likely to be incorrectly identified as described species if this key is used to identify them. Continual growth throughout their life, temperature dependent growth and host induced morphological variation in capsalids (see Discussion) are complicating factors. We strongly encourage the deposition of voucher specimens in recognised, curated museum collections of any capsalid taxa encountered ( Whittington 2004).

1a. Terminal part of penis adorned with sclerite or papillae ............................................................................................. 2

1b. Penis unadorned .......................................................................................................................................................... 4

2a. Penis with terminal sclerite ........................................................................................................................................ 3

2b. Penis with three terminal, conical papillae (see Fig. 24A of Whittington et al. 2001) ............................. B. jaliscana

3a. Vaginal pore posterior to testes ( Figs. 1 A, 2 in present study)....................................................................... B. ernsti

3b. Vaginal pore anterior to testes, posterior to common genital pore................................................................ B. rohdei

4a. Vaginal pore anterior to common genital pore (Fig. 10 of Whittington et al. 2001) ................................................. 5

4b. Vaginal pore adjacent or posterior to common genital pore but anterior to germarium (e.g. Fig. 12A-D of Whitting- ton et al. 2001) ............................................................................................................................................................ 7

5a. Anterior hamuli span almost half total length of haptor (Fig. 9A of Whittington et al. 2001); vaginal pore ventral .. ............................................................................................................................................................ B. acanthopagri

5b. Anterior hamuli less than half total length of haptor (Fig. 9E, I of Whittington et al. 2001); vaginal pore dorsal .. 6

6a. Posterior hamuli invested in muscle fibres (Fig. 9H of Whittington et al. 2001); single lobe associated with com- mon genital pore and vaginal pore (Fig. 10B of Whittington et al. 2001) ...................................... B. anticavaginata

6b. Posterior hamuli not invested in muscle fibres (Fig. 9L of Whittington et al. 2001); pair of lobes associated with common genital pore and single lobe associated with vaginal pore (Fig. 10C of Whittington et al. 2001). B. lutjani

7a. Separate uterine, male and vaginal pores .................................................................................................................... 8

7b. Uterine and male copulatory organ pores common forming common genital pore and separate vaginal pore ......... 9

8a. Large (> 5 mm total length) species with accessory sclerites placed posteriorly on haptor (Fig. 4A of Whittington et al. 2001); long tapering penis .................................................................................................................... B. sciaenae

8b. Small (<2.5 mm total length) species with broad anterior hamuli and accessory sclerites (Fig. 31A, B of Whitting- ton et al. 2001) .......................................................................................................................................... B. synagris

9a. Vaginal pore in mid-body between common genital pore and germarium (Fig. 12A, B of Whittington et al. 2001). .......................................................................................................................................................................... B. ovata

9b. Vaginal pore at a level between pharynx and common genital pore (e.g. Fig. 12B-D of Whittington et al. 2001). 10

10a. Penis with distinctive teat-shaped tip (Fig. 25A of Whittington et al. 2001) ........................................ B. monticellii

10b. Penis without teat-shaped tip, but with blunt or tapering tip ................................................................................... 11

11a. Accessory sclerites with distal hooks or with three points ...................................................................................... 12

11b. Accessory sclerites with tapering, pointed distal tips ............................................................................................. 13

12a. Distal tips of accessory sclerites a recurved hook (Fig. 17A, B, E of Whittington et al. 2001) ................ B. elongata

12b. Distal tips of accessory sclerites trident-like, with three points ( Fig. 7 A, B of present study)............... B. ishikawae

13a. Anterior hamuli with very broad roots; distal tips strongly recurved (Fig. 23B of Whittington et al. 2001) ............... ................................................................................................................................................................ B. innobilitata

13b. Anterior hamuli without very broad roots................................................................................................................ 14

14a. Small lobe near vaginal pore (Fig. 18 of Whittington et al. 2001) ......................................................... B. epinepheli

14b. Lobe near vaginal pore absent................................................................................................................................... 15

15a. Body broad, almost circular; anterior and posterior hamuli tiny relative to haptor (Fig. 29A, C, D of Whittington et al. 2001) .......................................................................................................................................................... B. sekii

15b. Body elongate, comparatively narrow; anterior and posterior hamuli not tiny relative to haptor ............................ 16

16a. Anterior and posterior hamuli not overlapping ........................................................................................................ 17

16b. Anterior and posterior hamuli overlapping .............................................................................................................. 18

17a. Anterior hamuli recurved distally, with slender shafts (Fig. 21 of Whittington et al. 2001) ................... B. hendorffii

17b. Anterior hamuli recurved distally with comparatively stout shafts (Fig. 30A, C, E, G of Whittington et al. 2001).... ..................................................................................................................................................................... B. seriolae

18a. Anterior attachment organs strongly hooded (Fig. 22B of Whittington et al. 2001); wide vitellarium-free margin to body; medium-sized (2.5–7.5 mm total length) species ............................................................................ B. hoshinai

18b. Anterior attachment organs weakly hooded or not hooded; vitellarium extends to near margin of body ................ 19

19a. Medium-large species (>2.5 mm total length, e.g. Fig. 3 A of Whittington et al. 2001) with relatively large anterior attachment organs.................................................................................................................................... B. pompatica

19b. Small-medium (<7.5 mm) species (e.g. Fig. 3 C of Whittington et al. 2001) with relatively small anterior attachment organs ........................................................................................................................................................................ 20

20a. Distal tips of accessory sclerites located in posterior half of haptor (Fig. 16E of Whittington et al. 2001) ... B. scari

20b. Distal tips of accessory sclerites located near centre of haptor ................................................................................ 21

21a. Penis broad proximally, with pointed or tapering tip ............................................................................................... 22

21b. Penis not broad proximally, with blunt tip (Fig. 15 of Whittington et al. 2001) ...................................................... 23

22a. Pair of prominent lobes near common genital pore; accessory sclerites with curved distal region; tip of penis pointed (Figs. 5A and 6A of present study) ............................................................................................. B. haywardi

22b. No lobes near common genital pore; accessory sclerites straight (Figs. 16C and 18G of Whittington et al. 2001); tip of penis tapering but not pointed (Fig. 20 of Whittington et al. 2001) ................................................ B. hawaiiensis

23a. Prominent wavy musculature near haptor margin; anterior and posterior hamuli of similar length (Figs. 13, 14, 16A, 19B, C of Whittington et al. 2001) .............................................................................................................. B. bodiani

23b. Prominent wavy musculature absent from haptor margin ........................................................................................ 24

24a. Marginal valve comprising a single large lobe between positions of each hooklet (Fig. 16D of Whittington et al. 2001) ................................................................................................................................................................. B. lolo

24b. Marginal valve comprising multiple lobes between positions of hooklets ( Fig. 3 A of present study; adults very small (<1.5 mm in total length) (e.g. Fig. 3 C of Whittington et al. 2001)..................................................... B. fieldsi

Figures

FIGURE 1. Benedenia ernsti n. sp. A. Entire adult animal, ventral view. Abbreviations: aao, anterior attachment organ; agr, accessory gland reservoir; ah, anterior hamulus; as, accessory sclerite; cgp, common genital pore; e, eye; g, germarium; gl, glandular tissue; ho, hooklet; ifc, internal fertilisation chamber; lo, lobe; m, marginal valve; mb, muscular bulb; Mg, Mehlis’ gland; ö, ootype; p, pharynx; pe, penis; ph, posterior hamulus; scl, sclerite; sr, seminal receptacle; t, tendon; te, testis; u, uterus; v, vagina; vp, vaginal pore; vr, vitelline reservoir. B. Penis sclerite. C. Accessory sclerite. D. Anterior hamulus. E. Posterior hamulus. Scale bars: A, 500 Μm; B, 40 Μm, C – E, 60 Μm.

FIGURE 3. Benedenia fieldsi n. sp. A. Adult specimen, ventral view. Composite drawing from type material and living specimens. Abbreviations as in previous Figures. B. Accessory sclerite. C. Anterior hamulus. D. Posterior hamulus. Drawn from paratypes. Scale bars: A, 300 Μm; B – D, 25 Μm.

FIGURE 7. Benedenia ishikawae. A. Ventral view of entire adult specimen. Re-drawn from Goto (1894). Abbreviations: hd, hood. B. Accessory sclerite. C. Anterior hamulus. D. Posterior hamulus. Other abbreviations as in previous Figures. Scale bars: A, 1 mm; B – D, 200 Μm.