Figure 1:
Facsimile reproduction of the original drawings of Cephalobus oxyuroides (A, H from de Man (1881) and B-G from de Man (1984)). (A) Neck; (B) Stoma; (C) Entire female; (D) Excretory pore; (E) Female posterior end; (F-H) Male posterior end.
Figure 2:
Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936a (line drawing). (A) Neck; (B) Lip region; (C) Female posterior end; (D) Female reproductive system; (E) Stoma; (F) Entire female; (G) Entire male; (H) Male posterior end; (I) Lateral field.
Figure 3:
Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936a (light microscopy). (A) Neck (black arrow pointing to the excretory pore, white arrow pointing to the deirid); (B) Lip region; (C) Stoma; (D) Lateral field; (E) Female reproductive system; (F) Uterine egg; (G) Female posterior end (arrow pointing the phasmid); (H) Male posterior end (arrows pointing genital papillae).
Figure 4:
Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936a (scanning electron microscopy). (A, B) Lip region in lateral and frontal view, respectively; (C) Female phasmid (arrow); (D) Neck (arrow pointing the excretory pore); (E) Excretory pore (arrow); (F, G) Female posterior end in lateral and ventral view, respectively (arrow pointing the phasmid); (H) Lateral field; (I) Anus.
Table 1.
Morphometrics of Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936a from Spain.
| Province | Navarra | Jaén | |
|---|---|---|---|
| Habitat | Oak forest | Whine forest | |
| Sex | Female | Male | Female |
| n | 11 | 9 | 2 |
| Body length | 610 ± 44.7 (554-695) | 567 ± 35.7 (513-629) | 493, 602 |
| a | 28.7 ± 1.9 (26.6-32.4) | 29.1 ± 2.9 (25.1-32.9) | 21.4, 26.2 |
| b | 4.2 ± 0.3 (3.9-4.7) | 4.0 ± 0.2 (3.8-4.3) | 3.5, 4.3 |
| c | 8.8 ± 0.6 (8.0-10.0) | 12.7 ± 1.0 (10.9-14.3) | 7.1, 8.4 |
| c' | 5.3 ± 0.3 (5.0-5.8) | 2.9 ± 0.3 (2.4-3.4) | 4.9, 5.5 |
| V | 60.3 ± 1.1 (58-64) | – | 58, 60 |
| Lip region width | 6.0 ± 0.0 (6) | 6.0 ± 0.0 (6) | 6, 7 |
| Stoma length | 13.5 ± 0.5 (13-14) | 13.2 ± 0.8 (12-14) | 14, 14 |
| Pharyngeal corpus length | 86.2 ± 3.4 (82–91) | 84.9 ± 5.6 (77-97) | 88, 86 |
| Isthmus length | 28.2 ± 3.0 (23-31) | 27.2 ± 2.9 (22-32) | 21, 25 |
| Bulbus length | 17.0 ± 1.2 (14-18) | 16.4 ± 0.9 (15-18) | 17, 15 |
| Pharynx length | 131 ± 5.4 (123–140) | 128.6 ± 8.0 (119-145) | 126, 126 |
| Nerve ring – anterior end | 95.1 ± 4.0 (89–103) | 93.0 ± 5.3 (84-101) | 100, 102 |
| Excretory pore – anterior end | 98.3 ± 43.8 (94-107) | 95.4 ± 5.2 (86-102) | 103, 101 |
| Deirid – anterior end | 114 ± 3.5 (106-117) | 110.4 ± 6.5 (97-119) | 119, 117 |
| Neck length (stoma + pharynx) | 145 ± 5.4 (137-153) | 141.8 ± 8.3 (132-158) | 140, 140 |
| Body diameter at midbody | 21.4 ± 2.1 (19-25) | 19.7 ± 2.2 (17-23) | 23, 23 |
| Lateral field width | 3.5 ± 0.6 (3-4) | 3.4 ± 0.5 (3-4) | 4, 4 |
| Anterior ovary/testis length | 135 ± 17.4 (116-160) | 115.6 ± 14.9 (99-144) | 118, 110 |
| Spermatheca length | 33.4 ± 7.9 (23-47) | – | 44, 55 |
| Post-vulval uterine sac length | 47.1 ± 4.0 (40-54) | – | 48, 57 |
| Vulva – anterior end | 368 ± 25.3 (328-405) | – | 285, 363 |
| Rectum length | 18.9 ± 1.4 (16-20) | 5.0 ± 0.7 (4-6) | 20, 20 |
| Anal body diameter | 13.1 ± 0.8 (12-14) | 15.7 ± 1.1 (14-18) | 14, 13 |
| Tail length | 69.5 ± 5.1 (60-79) | 44.9 ± 4.2 (36-49) | 69, 72 |
| Spicules length | – | 21.4 ± 0.5 (21-22) | – |
| Gubernaculum length | – | 13.7 ± 1.9 (12-16) | – |
1 Notes: Measurements in μm and in the form: mean ± standard deviation (range) where appropriate. Demanian indices (de Man, 1881): a, body length/body diameter; b, body length/pharynx length; c, body length/tail length; c′, tail length/anal body diameter; V = (distance from anterior region to vulva/body length) ×100.
Table 2.
Compendium of Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936a populations.
| Reference | Country | n | Body length | a | b | c | c ' | V | Stoma | Pharynx length | Neck length | Nerve ring − ant. end (%) | Excretory pore − ant. end (%) | Body width at midbody | Sper-matheca length | Post-vulval uterine sac / body diameter | Vulva-ant. end | Rectum length (female) | Anal body width | Tail length | Tail mucro | Spicules length | Gubernaculum length |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 1 | Spain | 13 females | 493-695 | 21.4-32.4 | 3.5-4.7 | 7.1-10.0 | 4.9-5.8 | 58-64 | 13-14 | 123-140 | 137-153 | 61-69 | 65-72 | 19-25 | 23-47 | 2.0-2.5 | 285-405 | 16-20 | 12-14 | 60-79 | Conoid elongate | – | – |
| 9 males | 513-629 | 21.1-32.9 | 3.8-4.3 | 10.9-14.3 | 2.4-3.4 | – | 12-14 | 119-145 | 132-158 | 64-68 | 64-71 | 17-23 | – | – | – | – | 14-18 | 36-49 | Conoid elongate | 21-22 | 12-16 | ||
| 2 | The Netherlands | 5 males | 447-514** | 20.0-21.8** | 3.1-3.2** | 10.4-11.9** | 2.5-2.7** | – | 12** | 125-149** | 145-161** | ? | 59** | 22-24** | – | – | – | – | 16-17** | 33-45** | Conoid elongate | 22** | 12* |
| 3 | The Netherlands | ? females | 700 | 20.0-25.0 | 4.0 | 8.0-9.0 | 5.1* | 61* | ? | ? | ? | 66* | 65* | ? | ? | 2.5* | ? | ? | ? | ? | Conoid elongate | – | – |
| ? males | 640 | 20.0-25.0 | 4.0 | 12.0-13.0 | 2.4* | – | ? | ? | ? | ? | ? | ? | – | – | – | – | ? | ? | Conoid elongate | ? | ? | ||
| 4 | Hungary | 1 female | 620 | 23.8** | 3.9** | 8.9** | ? | 56** | ? | ? | 160 | ? | ? | 26 | ? | ? | 350** | ? | ? | 70 | ? | ? | ? |
| 2 males | 450-500 | 20.8-21.4** | 3.6-3.8** | 11.1-11.3** | ? | – | ? | ? | 125-132 | ? | ? | 21-24 | – | – | – | – | ? | 40-45 | Conoid elongate | – | – | ||
| 5 | Australia | 1 female | 640 | 17.8** | 4.5* | 7.1** | 5.3** | 51 | 10** | ? | 143** | ? | 69** | 36** | ? | ? | ? | ? | 17** | 90 | ? | – | – |
| 6 | Switzerland | 1 male | 342 | 19.0 | 2.8 | 6.0 | ? | – | 11 | 112** | 122** | ? | ? | 16** | – | – | – | – | ? | 57** | ? | ? | ? |
| 7 | Austria | 1 female | 459 | 22.0 | 3.1 | 8.1 | ? | 61 | 8** | 141** | 149** | 57** | ? | 21** | ? | ? | ? | ? | 10** | 57** | ? | – | – |
| 8 | Rumania | 5 females | 500-565 | 19.4-21.8 | 3.7-4.1 | 6.8-7.8 | ? | 56-59 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | – | – |
| 1 male | 527 | 19.0 | 3.0 | 17.8 | ? | – | ? | ? | 176** | ? | ? | 28** | ? | ? | ? | ? | ? | 30** | ? | ? | ? | ||
| 9 | Austria | 19 females | 450-610 | 18.6-29.0 | 3.0-4.9 | 6.8-18.0 | 5.0* | 58-72 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | Short conoid | – | – |
| 11 males | 410-520 | 17.7-24.0 | 3.5-5.9 | 9.6-12.8 | ? | – | ? | ? | ? | ? | ? | ? | – | – | – | – | ? | ? | ? | ? | ? | ||
| 10 | Brazil | 1 female | 1926 | 21.4 | 4.2-5.0 | 12.4 | ? | 60 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | – | – |
| 1 male | 1485 | 29.0 | 4.9 | 16.5 | ? | – | ? | ? | ? | ? | ? | ? | – | – | – | – | ? | ? | ? | 45 | ? | ||
| 11 | UK | 1 female | 650 | 19.0 | 3.9 | 8.3 | 5.2* | 60 | ? | ? | 167** | ? | ? | 34** | ? | ? | 390** | ? | ? | 78** | Conoid elongate | – | – |
| 1 male | 550 | 21.0 | 3.5 | 11.6 | 2.7* | - | ? | ? | ? | ? | ? | 29** | – | – | – | – | ? | 47** | Conoid elongate | ? | ? | ||
| 12 | D. R. Congo | 1 female | 356 | 22.2 | 3.4 | 8.0 | ? | ? | ? | ? | 104** | ? | ? | 16** | ? | ? | ? | ? | ? | 45** | ? | – | – |
| 13 | Sweden | 1 female | 787 | 28.4 | 4.8 | 16.0 | ? | 65 | ? | ? | 164** | ? | ? | 28** | ? | ? | 512** | ? | ? | 49** | ? | – | – |
| 14 | Norway | 1 female | 800 | 38.1 | 3.8 | 12.3 | ? | ? | ? | ? | 213 | ? | ? | 21 | ? | ? | ? | ? | ? | 65 | ? | – | – |
| 1 male | 787 | 35.7 | 4.2 | 15.7 | ? | ? | ? | ? | 213 | ? | ? | 22 | ? | ? | ? | ? | ? | 50 | ? | ? | ? | ||
| 15 | Italy | ? females | 467-683 | 16.8-22.4 | 3.3-4.3 | 7.7-9.0 | ? | 59-65 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | – | – |
| 16 | South Africa | ? females | 410-460 | 20.0-22.0 | 3.8-3.9 | 8.7-13.1 | ? | 63 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | – | – |
| 17 | Mongolia | 1 female | 540 | 20.0 | 3.4 | 10.7 | 3.6 | 60 | ? | ? | 158** | ? | ? | 27** | ? | 1.4 | 326** | ? | ? | 50** | Short conoid | ? | ? |
| 1 male | 490 | 20.0 | 2.6 | 13.5 | 2.2 | – | ? | ? | 188** | ? | ? | 25** | – | – | – | – | ? | 36** | Short conoid | – | – | ||
| 18 | Hungary | ? females | 450-500 | 20.0-23.0 | 4.0-4.2 | 8.5-9.5 | 5.0-6.0 | 60-62 | 12 | ? | ? | ? | 74-77 | ? | ? | 0.8* | ? | ? | ? | ? | Conoid elongate | – | – |
| 19 | Tadjikistan | ? females | 460 | 20.0-22.0 | 3.4-3.8 | 8.0 | 4.3* | 60 | ? | ? | ? | ? | ? | ? | ? | ? | 276** | ? | ? | 58** | ? | – | – |
| 20 | Uzbekistan | 1 female | 450 | 22.2 | 39.41 | 9.1 | ? | 62 | ? | ? | ? | ? | ? | 20** | ? | ? | 279** | ? | ? | 49** | ? | – | – |
| 21 | Russia, Turkmenistan, Moldavia | ? females | 750-900 | 22.0-24.0 | 3.8-4.4 | 7.0-9.6 | ? | 58-62 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | – | – |
| ? males | 700-800 | 24.0-25.2 | 5.0-5.6 | 7.0-10.7 | ? | – | ? | ? | ? | ? | ? | ? | – | – | – | ? | ? | ? | ? | ? | ? | ||
| 22 | Russia | 1 female | 353 | 17.6 | 4.0 | 5.7 | ? | 50 | ? | ? | 88** | ? | ? | 20** | ? | ? | ? | ? | ? | 62** | ? | – | – |
| 2 males | 480-488 | 16.0-20.0 | 3.0-3.5 | 10.0 | 2.9* | - | 13 | 131* | 144 | 65* | 66* | 30- | – | – | – | – | 18* | 48-49** | Conoid elongate | 20 | 12 | ||
| 23 | Spain | 42 females | 450-700 | 17.0-23.0 | 3.0-5.0 | 6.5-9.0 | 6.0* | 56-72 | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | Conoid elongate | – | – |
| 3 males | 450-650 | 19.0-28.0 | 3.0-5.0 | 7.0-11.0 | 2.3* | – | ? | ? | ? | ? | ? | ? | – | – | – | – | ? | ? | Absent | ? | ? | ||
| 24 | Sweden | ? females | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | ? | 1.0-3.0 | ? | ? | ? | 59-73 | Conoid elongate | – | – |
| 25 | Brazil | 74 females | 370-650 | 18.0-32.0 | 3.3-5.0 | 5.0-8.5 | 7.3* | 55-63 | 10-14 | 140* | 108-170 | 61* | 65* | 19-23* | 16-32* | 1.1-1.3* | ? | 20-24 | 13* | 57-105 | Short conoid | – | – |
| 28 males | 390-600 | 21.0-35.0 | 3.2-4.2 | 14.6-19.8 | 2.5* | – | 10-14 | ? | 117-144 | ? | ? | 21* | – | – | – | – | 14* | 24-37 | Seta-like or wart-like | 14-20 | 9-12 | ||
| 26 | Austria | 6 females | 510-560 | 19.0-22.0 | 3.4-3.7 | 7.1-8.0 | 4.0-5.0 | 60–62 | ? | – | – | ||||||||||||
| 1 male | 520 | 20.0 | 3.4 | 9.5 | ? | – | 1.0-1.5 | ? | 23 | 12 | |||||||||||||
| 27 | Turkey | 1 female | 609 | 23.0 | 4.0 | 8.7 | 4.4 | 63 | 12 | 154 | 166** | 61 | 63 | 27 | 34 | 1.2 | ? | 19 | 16** | 70 | Short conoid | – | – |
| 1 male | 525 | 28.0 | 3.9 | 16.4 | 1.9 | – | 11 | 133 | 144** | 63 | 65 | 19 | – | – | – | – | ? | 32 | Short conoid | 17 | 10 | ||
| 28 | Spain | 4 females | 524-535 | 21.4-27.6 | 3.5-3.7 | 8.4-8.9 | 4.2-5.3 | 60-64 | 9-14 | 143-150 | 156-161** | 54-64 | 54-67 | 19-25 | 26-36 | 0.7-1.7 | 317-338 | 18-20 | 12-15 | 60-64 | Short conoid | – | – |
| 29 | Bulgaria | 5 females | 588-729 | 19.0-26.0 | 3.8-4.3 | 5.9-7.4 | 5.0-6.8 | 57-61 | ? | 127-136 | ? | ? | 66-74 | 17-22 | ? | 0.5 | ? | 13-20 | ? | 32-36 | ? | – | – |
| 3 males | 436-556 | 25.0-30.0 | 3.2-3.8 | 10.9-15.7 | 1.9-2.3 | – | ? | 124-139 | ? | ? | 64-69 | 16-17 | ? | ? | ? | – | ? | 30-36 | ? | 14-21 | 8-10 | ||
| 30 | Hungary | ? females | 380-740 | 18.0-28.0 | 3.3-5.0 | 6.0-10.0 | 4.2-5.8 | 56-66 | 10-14 | 120-150 | ? | ? | 74-78 | ? | ? | < 1.0 | ? | ? | ? | 70-90 | ? | – | – |
| ? males | 300-620 | 20.0-28.0 | 3.2-4.2 | 14.0-18.0 | ? | – | 10-14 | 120-150 | ? | ? | 74-78 | – | – | – | – | ? | ? | ? | 14-20 | 9-11 |
1 Notes: References: 1. Present paper, 2. de Man (1875), 3. de Man (1881, 1984), 4. Örley (1880), 5. Cobb (1893a), 6. Stefan´ski (1914), 7. Micoletzky (1914), 8. Micoletzky (1917), 9. Micoletzky (1921), 10. Rahm (1928, 1929), 11 Thorne (1937), 12. Schuurmans-Stekhoven and Teunissen (1938), 13. Allgén (1950), 14. Allgén (1953), 15. Meyl (1953a, 1953b), 16. Heyns (1961), 17. Andrássy (1964), 18. Andrássy (1967), 19. Ivanova (1968), 20. Mavljanov (1976), 21. Nesterov (1979), 22. Mukhina (1981), 23. Monreal and Campoy (1982), 24. Boström (1984), 25. Rashid et al. (1984), 26. Gerber (1991), 27. Boström (1993), 28. Abolafia and Peña-Santiago (2002), 29. Iliev et al. (2003), 30. Andrássy (2005), 31. Kim et al. (2017). *Measurements from drawings; **Measurements from other measurements; ?Unknown measurement; –Character absent; 1Probably a mistake.
Figure 5:
Lip region, post-vulval uterine sac, male and female tails of the species of the genus Eucephalobus Steiner, 1936a including species inquirendae. Based on original descriptions except E (male tail) cf. Amirzadi et al. (2013); G (lip region), M (lip region, tails) cf. Andrássy (1967); H (post-vulval sac), M (post-vulval sac) cf. Boström (1984); H (female tail) cf. de Man (1984). Not to scale.
Figure 6:
Bayesian inference tree showing the phylogenetic position of Eucephalobus and related taxa based on sequences of the 18S rDNA region. Bayesian posterior probabilities (%) are given for each clade. Scale bar shows the number of substitutions per site.
Figure 7:
Bayesian Inference tree showing the phylogenetic position of Eucephalobus and related taxa based on sequences of the 28S rDNA region. Bayesian posterior probabilities (%) are given for each clade. Scale bar shows the number of substitutions per site.