Dipylidiidae, Stiles, 1896
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publication ID |
https://doi.org/ 10.1016/j.ijppaw.2024.100929 |
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persistent identifier |
https://treatment.plazi.org/id/03CB9E47-B953-2D4F-E717-7A9E67AD62AD |
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treatment provided by |
Felipe |
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scientific name |
Dipylidiidae |
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3.1.4. Dipylidiidae View in CoL - Joyeuxiella spp .
Four different species of Joyeuxiella were identified. Joyeuxiella sp. I was the most common and found in six different hosts from South Africa (one African wildcat, one African civet, one small-spotted genet and three rusty-spotted genets). Fragments of Joyeuxiella sp. II, obtained from a single domestic cat from South Africa, did not provide sufficient material suitable for morphological examination. The descriptions of Joyeuxiella sp. III are based on two worms isolated from an African wildcat from South Africa. Morphological information on Joyeuxiella sp.
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IV is based on four incomplete specimens found in a domestic cat and two African wolves from Ethiopia.
The complete cox1 gene was obtained from Joyeuxiella spp . I and III (1629 bp), whereas only a 964 bp long cox1 fragment of Joyeuxiella sp. II, a 1600 bp sequence of Joyeuxiella sp. IV and a 955 bp fragment of J. sp. aff. pasqualei could be sequenced. The complete nad1 gene was received from J. sp. aff. pasqualei (891 bp), as well as partial sequences from Joyeuxiella spp . I (741 bp) and III (833 bp). An additional 468 bp fragment of cob was obtained from Joyeuxiella spp . I, III and J. sp. aff. pasqualei . This allowed the construction of a concatenated sequence tree with Joyeuxiella spp . I and III, combining the sequences of these three mitochondrial genes for a more precise phylogenetic positioning of Joyeuxiella among other genera ( Fig. 6 View Fig ). For alignment, the length of the cox1 and nad1 sequences of Joyeuxiella spp . was reduced to 1554 bp and 737 bp, respectively. The final dataset of Fig. 6 View Fig uses 2768 positions, including gaps from insertions or deletions. While Joyeuxiella spp . had been positioned separately from Dipylidium spp. based on a short cox1 fragment in Fig. 1 View Fig , they are placed as a sister clade to Dipylidium spp. when the combined sequences are used ( Fig. 6 View Fig ). As seen in Fig. 1 View Fig , J. sp. aff. pasqualei clusters with sequences of J. pasqualei published by Bezerra-Santos et al. (2022) and Schuster et al. (2023), but only correspond to a maximum of 93.3% (with OR081744 - Schuster et al., 2023). The sequence of J. gervaisi (OR081751), synonymised with J. fuhrmanni , that was published by Schuster et al. (2023) takes the most basal position in the Joyeuxiella clade. This can also be seen when using untrimmed sequences. However, the placement of Joyeuxiella IV , which moves to the second most basal position of the Joyeuxiella clade, changed (Supplementary Fig. S1 View Fig ).
In addition to protein coding genes, partial sequences of the 18S rRNA gene of the species I (534 bp), III (422 bp) and J. sp. aff. pasqualei (534 bp) and the 12S rRNA (289, 287 and 298 bp respectively) could be sequenced. Only the 12S rRNA sequence of Joyeuxiella sp. aff. pasqualei correlated to an entry in GenBank. It was 100% identical to an undescribed Mesocestoides species from Turkey (MH992704.1).
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To investigate the phylogenetic position of Joyeuxiella , especially with respect to Dipylidium , based also on ribosomal DNA, a phylogenetic tree was constructed using 402 bp long sequences, including alignment gaps, of the 18S rRNA gene from 23 species of different cyclophyllidean families. In this phylogram, Joyeuxiella spp . and Anonchotaenia spp. (of the family Paruterinidae ) form sister clades, neighbouring the Dipylidium and Raillietina ( Davaineidae ) clades and Catenotaenia pusilla ( Catenotaeniidae ) (Supplementary Fig. S5 View Fig ).
The comparative numbers and measurements of various morphological features observed in adult Joyeuxiella spp . are listed in Table 4.
Most Joyeuxiella specimens were incomplete or fragmented, thus the full strobila length can only be estimated. Joyeuxiella sp. I attains a length of at least 5 cm. Two specimens of Joyeuxiella sp. III were 7.5 cm and approximately 12 cm long. Merely short fragments were available of both Joyeuxiella spp . II and IV. Based on the size of the proglottids of Joyeuxiella sp. IV, the full strobila length was estimated to be> 15 cm. The proglottids of Joyeuxiella sp. IV are considerably larger in both length and width than those of the other Joyeuxiella spp . I-III and measured up to 2 mm in width in mature segments. Gravid and pregravid proglottids of Joyeuxiella spp . grow more slender and longer than the posterior mature segments, which are wider than long ( Fig. 7B–F View Fig ). The width of the strobilae of Joyeuxiella sp. I specimens range between 0.1 and 0.3 mm at the neck and 0.5–1 mm at the widest part of the posterior mature proglottids. For Joyeuxiella sp. III, similar measurements ranging from 0.17 mm (anterior) to 1 mm (posterior) were made. Rostellar hooks of Joyeuxiella spp . I, III and IV are thorn-shaped and decrease in size towards the base of the rostellum ( Fig. 7D and E View Fig ). No scolex was available of Joyeuxiella sp. II and the small fragments could not be stained due to their poor condition. The stained specimens of Joyeuxiella spp . I and III reveal two vasa deferentia that are located close to the anterior margin of the mature proglottid and each coil toward the cirrus sacks, ending in the genital pores at the upper third of the proglottids. Some mature proglottids of one specimen of Joyeuxiella sp. III have long cirri protruding from the genital openings ( Fig. 7B View Fig ). The testes are found in the space posterior to the vasa deferentia and between the longitudinal excretory vessels ( Fig. 7A and B View Fig ). In contrast to Joyeuxiella spp . I and III, the vasa deferentia of Joyeuxiella sp. IV do not reach the anterior margins of the segments and the genital pores are located closer to mid proglottid ( Fig. 7C View Fig ). Exact numbers of testes could not be determined for Joyeuxiella sp. IV due to the condition of the specimens,
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l.e.v. = longitudinal excretory vessels.
though testes could be spotted anterior to the vasa deferentia sporadically. No gravid proglottids were available of Joyeuxiella sp. IV. Gravid proglottids of both Joyeuxiella spp . I and III are similar in appearance, with eggs that are confined to the space between the longitudinal excretory ducts ( Fig. 7F View Fig ). The egg capsules of both species are round or slightly oval in shape with average diameters ranging between 30 and 40 μm ( Fig. 5 View Fig , Table 4).
Fig. 7 View Fig depicts the general structure of the scolex, hooks and mature proglottid of Joyeuxiella sp. I (A, D and E), a younger mature segment and a gravid proglottid of one specimen of Joyeuxiella sp. III (B and F) and a mature proglottid of Joyeuxiella sp. IV found in this study (C).
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