Systematic revision of dactylogyridean parasites (Monogenea) from cichlid fishes in Africa, the Levant and Madagascar Author Pariselle, Antoine Institut de Recherche pour le Développement, Laboratoire d’écologie et de systématique, Centre de Belair, BP 1386, 18524 Dakar (Sénégal) antoine. pariselle @ ird. fr pariselle@ird.fr Author Euzet, Louis Station méditerranéenne de l’Environnement littoral, Université Montpellier II, 1 Quai de la Daurade, F- 34200 Sète (France) euzet @ univ-montp 2. fr text Zoosystema 2009 2009-12-31 31 4 849 898 http://www.bioone.org/doi/abs/10.5252/z2009n4a6 journal article 8061 10.5252/z2009n4a6 d4ef2df1-7e15-49c7-9c1b-a2f1bf23af38 1638-9387 4520413 IDENTIFICATION KEY FOR STUDIED GENERA AND SPECIES Following Dossou (1985) , Pariselle & Euzet (1995c) and Pariselle (1996) the 71 species forming Cichlidogyrus , recorded from 66 different host species (64 Cichlidae , one Cyprinodontidae , one Nandidae ), may be clustered in different groups according to the relative size of their haptoral sclerites. This has been confirmed by Pouyaud et al . (2006) using both genetic and morphometrical data: “This division into four groups based on morphological criterions is also confirmed by the genetic distances”. These groups can be easily characterised by morphometrical characters: length of the transverse dorsal bar auricles, of the uncinulus pairs III-VII, and of uncinuli pairs I. The following may introduced morphometrical differences into the size range for these characters: developmental stage of the examined parasite; influence of temperature on the size of the sclerites ( Ergens & Gelnar 1985 ; Appelby 1996); equipment used; method by which the measurements are taken, or other constraints (see Caltran et al . 1995 ). To avoid these problems, the length of each examined sclerite, according to Pariselle & Euzet (2003) , will be “standardised” by dividing its total length by the total length of uncinuli pair II, which retain its size from larval to adult stages of development in ancyrocephalid Monogenea . – Length of transverse dorsal bar auricles: the distribution of this character values ( Fig. 1A ) shows a clear gap between species belonging to Scutogyrus (> 2.9) ( Fig. 2E ) and Cichlidogyrus (<1.9) ( Fig. 2F ). This unequivocal character, therefore, is relevant to diagnose Scutogyrus . – Size of uncinuli pairs III-VII: the gap between large and small standardized sizes of uncinuli pairs III-VII is located near 2 ( Fig. 1B ). – Size of uncinuli pair I: the gap between large and small standardized sizes of uncinuli pairs I is located near 1.7 ( Fig. 1C ). 1. One transverse bar (ventral) ....................................................................................... 2 — More than one transverse bar (ventral and dorsal) ...................................................... 3 2. One asymmetrical pair of anchors (ventral) ( Fig. 2A ) ..................................... Urogyrus — Two pairs of symmetrical anchors ( Fig. 2B ) ............................................. 6 Enterogyrus 3. Two transverse bars (one dorsal and one ventral) ........................................................ 4 — Three transverse bars (one dorsal, two ventral) ( Fig. 2C ) ...................... 15 Onchobdella 4. Dorsal bar with wings (= auricles) ( Fig. 2D, a ) ........................................................... 5 — Dorsal bar lacking wings ( Fig. 2D, b ) ................................................... 13 Insulacleidus 5. Dorsal bar winged with very long auricles, ventral bar associated with a plate ( Fig. 2E ) ................................................................................................................ 22 Scutogyrus — Dorsal bar with short auricles ventral bar not associated with a plate ( Fig. 2F ) .............. ........................................................................................................... 27 Cichlidogyrus 6. Dorsal anchor total length more than 50 µm , ventral bar straight, more than 40 µm ( Fig. 3A, a ) ................................................................................................................. 7 — Dorsal anchor total length less than 50 µm , ventral bar V-shaped, less than 40 µm ( Fig. 3A, b ) ............................................................................................................................... 8 7. Ventral anchor total length less than 27 µm .............................. Enterogyrus malmbergi — Ventral anchor total length more than 34 µm ................................... Enterogyrus crassus 8. Dorsal anchor with shaft longer than blade ( Fig. 3B, a ) .............................................. 9 — Dorsal anchor with shaft shorter than blade ( Fig. 3B, b ) .......................................... 11 9. Haptor cup-shaped .................................................................. Enterogyrus cichlidarum — Haptor in two parts: one bulbous and one peduncular shaped ( Fig. 3C ) .................. 10 10. Penis with distal turns well separated ( Fig. 3D, a ) ........................ Enterogyrus coronatus — Penis with distal turns adjoining ( Fig. 3D, b ) .................................. Enterogyrus foratus 11. Ventral bar total length less than 14 µm ........................................... Enterogyrus amieti — Ventral bar total length more than 16 µm ................................................................ 12 12. Penis with 3-2-3 turns ( Fig. 3E, a ) .............................................. Enterogyrus melenensis — Penis with 6-2-4 turns ( Fig. 3E, b ) ......................................... Enterogyrus barombiensis 13. Looped penis ( Fig. 4B, a ) ....................................................... Insulacleidus paratilapiae — Straight penis ( Fig. 4B, b ) ........................................................................................ 14 14. Simple accessory piece ( Fig. 4C, a ) ............................................. Insulacleidus paretropli — Accessory piece complex ( Fig. 4C, b ) ............................... Insulacleidus ptychochromidis 15. Dorsal bar clearly arched (M-shaped) ( Fig. 5A, a ) ..................................................... 16 — Dorsal bar straight or slightly V-shaped ( Fig. 5A, b ) ................................................. 19 16. Ventral bars longer than dorsal one ( Fig. 5B, a ) ......................... Onchobdella pterigyalis — Ventral bars shorter than dorsal one ( Fig. 5B, b ) ....................................................... 17 17. Penis J-shaped ( Fig. 5C, a ) ........................................................... Onchobdella voltensis — Penis long and sinuous or spirally coiled ( Fig. 5C, b ) ............................................... 18 18. Penis sinuous, no visible vagina ( Fig. 5D, a ) ............................... Onchobdella spirocirra — Penis spirally coiled (4 to 5 turns), long and sinuous vagina ( Fig. 5D, b ) ....................... ...................................................................................................... Onchobdella bopeleti 19. Dorsal bar slightly V-shaped, length more than 43 µm ( Fig. 5E, a ) ... Onchobdella silverai — Dorsal bar straight, length less than 40 µm ( Fig. 5E, b ) ............................................ 20 20. Ventral anchor total length less than 11 µm ( Fig. 5F, a ) .................. Onchobdella melissa — Ventral anchor total length more than 20 µm ( Fig. 5F, b ) ......................................... 21 21. Penis thin, G-shaped, total length 80 µm ( Fig. 5G, a ) .................... Onchobdella krachii — Penis large, J-shaped, total length less than 60 µm ( Fig. 5G, b ) .... Onchobdella aframae 22. Penis and vagina very long and thin ( Fig. 6A, a ) .............................. Scutogyrus ecoutini — Penis and vagina shorter ( Fig. 6A, b ) ........................................................................ 23 23. Penis length more than 70 µm ( Fig. 6B, a ) ............................................................... 24 — Penis length less than 70 µm ( Fig. 6B, b ) .................................................................. 25 24. Vagina tubular, thin-walled ( Fig. 6C, a ) ............................................ Scutogyrus bailloni — Vagina large, thick-walled ( Fig. 6C, b ) ...................................... Scutogyrus gravivaginus 25. Accessory piece ending in two curved pincer-like hooks ( Fig. 6D, a ) ... Scutogyrus chikhii — Accessory piece ending in pincers with one component hooked, the other blunt ( Fig. 6D, b ) ............................................................................................................................. 26 26. Accessory piece basal extension well marked ( Fig. 6E, a ) .................... Scutogyrus minus — No accessory piece extension ( Fig. 6E, b ) .................................... Scutogyrus longicornis 27. Uncinuli pairs III-VII long (“standardised length” more than 2) ( Fig. 7A, a ) ............ 28 — Uncinuli pairs III-VII short (“standardised length” less than 2) ( Fig. 7A, b ) .............. 53 28. Uncinuli pairs I large (“standardised length” more than 1.7) ( Fig. 7B, a ) .................. 29 — Uncinuli pairs I small (“standardised length” less than 1.7) ( Fig. 7B, b ) .................... 30 FIG. 2. — Illustration of the identification key for studied genera: A , haptoral morphology of Urogyrus Bilong Bilong, Birgi & Euzet, 1994 species; B , haptoral morphology of Enterogyrus Paperna, 1963 species; C , haptoral morphology of Onchobdella Paperna, 1968 species; D , shape of dorsal bars (not winged [ a ], winged [ b ]); E , haptoral morphology of Scutogyrus Pariselle & Euzet, 1995 species; F , haptoral morphology of Cichlidogyrus Paperna, 1960 species. Abbreviations: DA , dorsal anchor; DB , dorsal transverse bar; VA , ventral anchor; VB , ventral transverse bar. FIG. 3. — Illustration of identification key for Enterogyrus Paperna, 1963 species: A , size of dorsal anchor (large [ a ], small [ b ]) and shape of dorsal transverse bar (straight [ a ], V-shaped [ b ]); B , relative length of dorsal anchor blade and shaft (long shaft [ a ], short shaft [ b ]); C , morphology of haptor with bulbous and peduncular parts; D , penis distal turns (separated [ a ] or adjoining [ b ]); E , number of turns in the different parts of the penis. Abbreviations: bl , blade; s , shaft; 2 , 3 , 4 and 6 , number of turns in the parts of the penis. FIG. 4. — Illustration of identification key for Insulacleidus Rakotofiringa & Euzet, 1983 species: A , haptor morphology; B , shape of the penis (looped [ a ], straight [ b ]); C , shape of the accessory piece (simple [ a ], complex [ b ]). FIG. 5. — Illustration of identification key for Onchobdella Paperna,1968 species: A , shape of dorsal transverse bar (arched [ a ], straight or V-shaped [ b ]); B , relative length of dorsal and ventral bars (ventral> dorsal [ a ], ventral <dorsal [ b ]); C , shape of penis (J-shaped [ a ], sinuous or spirally coiled [ b ]); D , shape of penis (sinuous [ a ], spirally coiled [ b ]); E , shape and size of dorsal bar (long and V-shaped [ a ], short and straight [ b ]); F , size of ventral anchor (small [ a ], large [ b ]); G , shape of the penis (G-shaped [ a ], J-shaped [ b ]). FIG. 6. — Illustration of identification key for Scutogyrus Pariselle & Euzet,1995 species: A , size of the penis (long [ a ], short [ b ]); B , size of the penis (long [ a ], short [ b ]); C , shape of the vagina (tubular and thin-walled [ a ], large and thick-walled [ b ]); D , shape of the accessory piece ending (two curved pincer-like hooks [ a ], one component hooked, the other blunt [ b ]); E , shape of the accessory piece (basal extension well marked [ a ], no extension [ b ]). Scale bar: 30 μm. 29. Vagina not visible, single-looped penis ( Fig. 7C, a ) ............. Cichlidogyrus arthracanthus — Sclerotised vagina, curved penis ( Fig. 7C, b ) ........................ Cichlidogyrus inconsultans 30. Very large body and sclerite size, large and S-shaped penis ( Fig. 7D, a ) ... Cichlidogyrus halli — Other penis shapes ( Fig. 7D, b ) ................................................................................ 31 31. Male apparatus auxiliary plate(s) present ( Fig. 8A, a ) ................................................ 32 — Male apparatus without auxiliary plate ( Fig. 8A, b ) .................................................. 39 32. Presence of two small auxiliary plates ( Fig. 8B, a ) .......................... Cichlidogyrus guirali — Presence of one auxiliary plate ( Fig. 8B, b ) ............................................................... 33 33. Reduced auxiliary plate (smallest axis < 10 µm ) ( Fig. 8C, a ) ... Cichlidogyrus microscutus — Large auxiliary plate (smallest axis> 10µm ) ( Fig. 8C, b ) .......................................... 34 34. Accessory piece slightly S-shaped, ending bottle-opener-like ( Fig. 8D, a ) ...................... ................................................................................................ Cichlidogyrus thurstonae — Accessory piece bent at right angle ( Fig. 8D, a ) ........................................................ 35 35. Penis length less than 80 µm .................................................................................... 36 — Penis length more than 95 µm .................................................................................. 38 36. Extremity of accessory piece with large perpendicular diverticles, vagina thin walled and amphora-shaped ( Fig. 8E, a ) .................................................... Cichlidogyrus aegypticus — Extremity of accessory piece without large perpendicular diverticles ( Fig. 8E, b ) ...... 37 37. Accessory piece convex face with large semi-circular extension ( Fig. 8F, a ) ..................... ....................................................................................................... Cichlidogyrus gallus — Accessory piece convex face without large semi-circular extension ( Fig. 8F, b ) ............... .................................................................................................... Cichlidogyrus paganoi 38. Penis length less than 105 µm , vagina length less than 50 µm ....................... C. bilongi — Penis length more than 130 µm , vagina length more than 60 µm ................... C. agnesi 39. Long and narrow vagina ( Fig. 9A, a ) ........................................................................ 40 — Large and/or short vagina ( Fig. 9A, b ) ...................................................................... 42 40. Penis length more than 85 µm , ................................................. Cichlidogyrus flexicolpos — Penis length less than 60 µm ................................................................................... 41 41. Accessory piece T-shaped ( Fig. 9B, a ) ...................................... Cichlidogyrus testificatus — Accessory piece ending in a hook ( Fig. 9B, b ) ............................ Cichlidogyrus lemoallei 42. Short conical and right angle folded vagina ( Fig. 9C, a ) ........................................... 43 — Tubular vagina (almost constant diameter) ( Fig. 9C, b ) ............................................ 46 43. Penis length less than 61 µm .................................................................................... 44 — Penis length more than 65 µm .................................................................................. 45 44. Accessory piece ending with smooth convex face ( Fig. 9D, a ) ........ Cichlidogyrus bouvii — Accessory piece ending in a bifurcated hook with crenulated convex face ( Fig. 9D, b ) ... ..................................................................................................... Cichlidogyrus dossoui 45. Accessory piece ending folded back ( Fig. 9E, a ) ............................... Cichlidogyrus vexus — Accessory piece ending in a hook ( Fig. 9E, b ) .......................... Cichlidogyrus douellouae 46. Short annulated vagina ( Fig. 9F, a ) .................................................. Cichlidogyrus hemi FIG. 7. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , length of uncinuli III to VII (long [ a ], short [ b ], pair II is given as a reference); B , size of uncinuli I (large [ a ], small [ b ], pair II is given as a reference); C , C. arthracanthus Paperna, 1960 penis ( a ); C. inconsultans ( Birgi & Lambert, 1986 ) sclerotised genitalia ( b ); D , shape of penis ( C. halli ( Price & Kirk, 1967 ) [ a ], other species [ b ]). Abbreviations: I , II and III -VII , uncinuli numbering. — Vagina with smooth wall ( Fig. 9F, b ) ........................................................................ 47 47. Straight or slightly curved vagina ( Fig. 10A, a ) .......................... Cichlidogyrus legendrei — Well curved or looped vagina ( Fig. 10A, b ) .............................................................. 48 48. Looped vagina ( Fig. 10B, a ) ..................................................................................... 49 — Curved vagina (not looped) ( Fig. 10B, b ) ................................................................. 51 49. Vagina forming a single loop ( Fig. 10C, a ) ............................... Cichlidogyrus tiberianus — Vagina forming two semi-loops ( Fig. 10C, b ) ........................................................... 50 50. Vagina semi-loops in the same plane, crenulated distal accessory piece extremity ( Fig. 10D, a ) ................................................................................................. Cichlidogyrus kouassii — Vagina semi-loops in perpendicular planes, accessory piece distal extremity with a single hook ( Fig. 10D, b ) ................................................................ Cichlidogyrus bonhommei 51. Curved accessory piece ending in two points, vagina with characteristic enlargement ( Fig. 10E, a ) ...................................................................... Cichlidogyrus anthemocolpos — Accessory piece bent at right angle, ending in a single point, vagina without enlargement ( Fig. 10E, b ) ............................................................................................................. 52 52. Accessory piece with triangular extension at proximal third ( Fig. 10F, a ) ....................... ...................................................................................................... Cichlidogyrus ergensi — Accessory piece without triangular extension at proximal third ( Fig. 10F, b ) .................. ................................................................................................ Cichlidogyrus ouedraogoi 53. Auricles reduced, in continuity with dorsal bar anterior face ( Fig. 11A, a ) ................ 54 — Auricles developed ( Fig. 11A, b ) ............................................................................... 56 54. Uncinulus I “standardised length” more than 2 ( Fig. 11B, a ) ....... Cichlidogyrus kothiasi — Uncinulus I “standardised length” less than 2 ( Fig. 11B, b ) ...................................... 55 55. Accessory piece with narrow base, penis with large heel ( Fig. 11C, a ) ... Cichlidogyrus berrebii — Accessory piece with wide base, penis with thin heel ( Fig. 11C, b ) ... Cichlidogyrus pouyaudi 56. Uncinulus I “standardised length” more than 2 ( Fig. 11D, a ) ................................... 57 — Uncinulus I “standardised length” less than 2 ( Fig. 11D, b ) ...................................... 76 57. Penis with swollen portion ( Fig. 11E, a ) ................................................................... 58 — Penis without swollen portion (thin and tubular all along) ( Fig. 11E, b ) .................. 59 58. Penis length less than 35 µm ............................................. Cichlidogyrus papernastrema — Penis length more than 45 µm .................................................. Cichlidogyrus philander 59. Penis length less than 50 µm .................................................................................... 60 — Penis length more than 60 µm .................................................................................. 69 60. Sclerotised vagina ..................................................................................................... 61 — Vagina not visible ..................................................................................................... 62 61. Penis with developed heel, accessory piece S-shaped ( Fig. 11F, a ) ... Cichlidogyrus dageti — Penis with poorly developed heel, accessory piece C-shaped ( Fig. 11F, b ) ...................... ..................................................................................................... Cichlidogyrus falcifer FIG. 8. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , presence ( a ) or absence ( b ) of an auxiliary plate associated with the male apparatus; B , presence of two ( a ) or one ( b ) auxiliary plate associated with the male apparatus; C , size of the auxiliary plate associated with the male apparatus (small [ a ], large [ b ]); D , shape of the accessory piece (S-shaped [ a ], bent at right angle [ b ]); E , shape of the extremity of the accessory piece (presence [ a ] or absence [ b ] of large perpendicular diverticles); F , shape of the accessory piece convex face (presence [ a ] or absence [ b ] of a large semi-circular extension). FIG. 9. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , shape of the sclerotised vagina (long and narrow [ a ], large and/or short [ b ]); B , shape of the accessory piece (T-shaped [ a ], ending in a hook [ b ]); C , shape of the vagina (short conical and right angle folded [ a ], tubular [ b ]); D , shape of the accessory piece ending convex face (smooth [ a ], crenulated [ b ]); E , shape of the accessory piece ending (folded back [ a ], hook shaped [ b ]); F , shape of vagina wall (annulated [ a ], smooth [ b ]). FIG. 10. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , shape of the vagina (straight or slightly curved [ a ], well curved or looped [ b ]); B , shape of the vagina (looped [ a ], curved [ b ]); C , shape of vagina (one [ a ] or two semi [ b ] loops); D , shape of vagina and accessory piece extremity (semi-loops in the same plane and crenulated [ a ], semi-loops in perpendicular planes and single-hooked [ b ]); E , shape of accessory piece extremity and vagina (ending in two points and characteristic enlargement [ a ], ending in a single point and no enlargement [ b ]); F , presence ( a ) or absence ( b ) of a triangular extension at proximal third of the accessory piece. 62. Accessory piece forming a large gutter-like portion at its base ( Fig. 12A, a ) ................... .................................................................................................... Cichlidogyrus nuniezi — Narrow accessory piece ( Fig. 12A, b ) ........................................................................ 63 63. Accessory piece with a rounded bulge at distal third ( Fig. 12B, a ) .................................. ................................................................................................... Cichlidogyrus dionchus — Accessory piece without a rounded bulge at distal third ( Fig. 12B, b ) ....................... 64 64. Accessory piece with double ending ( Fig. 12C, a ) ....................... Cichlidogyrus digitatus — Accessory piece with single ending ( Fig. 12C, b ) ...................................................... 65 65. Accessory piece ending in a thin and long hook ( Fig. 12D, a ) .... Cichlidogyrus berradae — Accessory piece ending in a wide and short hook ( Fig. 12D, b ) ................................ 66 66. Penis with large and rounded heel ( Fig. 12E, a ) ........................................................ 67 — Penis with narrow heel ( Fig. 12E, b ) ......................................................................... 68 67. Penis extremity folded back, accessory piece closed C-shaped ( Fig. 12F, a ) .................... ................................................................................................... Cichlidogyrus reversati — Penis and accessory piece open C-shaped ( Fig. 12F, b ) ................. Cichlidogyrus halinus 68. Accessory piece> 27 µm ( Fig. 12G, a ) ........................................... Cichlidogyrus yanni — Accessory piece < 26 µm ( Fig. 12G, b ) ........................................ Cichlidogyrus quaestio 69. Spirally coiled penis, length more than 350 µm ( Fig. 13A, a ) ................................... 70 — Not spirally coiled penis, length less than 150 µm ( Fig. 13A, b ) ............................... 72 70. Penis with 4 or 5 turns, not spirally coiled vagina ( Fig. 13B, a ) ...... Cichlidogyrus euzeti — Penis with more than 4 or 5 turns, spirally coiled vagina ( Fig. 13B, b ) ..................... 71 71. Penis with 8 or 9 turns, length about 450 µm ( Fig. 13C, a ) ..... Cichlidogyrus longicirrus — Penis with 14 or 15 turns, length about 1500 µm ( Fig. 13C, b ) ... Cichlidogyrus sanseoi 72. Penis with large diameter, slightly sinuous, without heel ( Fig. 13D, a ) .......................... ......................................................................................................... Cichlidogyrus arfii — Filiform penis ( Fig. 13D, b ) ..................................................................................... 73 73. Nearly straight penis ( Fig. 13E, a ) ............................................... Cichlidogyrus albareti — Curved penis ( Fig. 13E, b ) ....................................................................................... 74 74. G-shaped penis ( Fig. 13F, a ) ......................................................... Cichlidogyrus teugelsi — Single-looped penis ( Fig. 13F, b ) .............................................................................. 75 75. Long vagina, accessory piece round ended ( Fig. 14A, a ) ............ Cichlidogyrus nandidae — Short vagina, accessory piece ending in a hook ( Fig. 14A, b ) ... Cichlidogyrus bychowskii 76. Penis with swollen portion ( Fig. 14B, a ) ................................................................... 77 — Tubular penis without swollen portion ( Fig. 14B, b ) ................................................ 84 77. Well marked swollen portion ( Fig. 14C, a ) ............................................................... 78 — “Diffuse” swollen portion ( Fig. 14C, b ) .................................................................... 82 78. Penis length less than 60 µm .................................................................................... 79 — Penis length more than 80 µm .................................................................................. 81 FIG. 11. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species; A , size of auricle on the dorsal bar (reduced [ a ], developed [ b ]); B , relative size of uncinulus pair I (large [ a ], small [ b ]); C , shape of the accessory piece base and penis heel (narrow and large [ a ], wide and thin [ b ]); D , relative size of uncinulus pair I (large [ a ], small [ b ]); E , presence ( a ) or absence ( b ) of a swollen portion in the penis; F , size of the penis heel and of the accessory piece (well developed and S-shaped [ a ], poorly developed and c-shaped [ b ]). FIG. 12. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , shape of the accessory piece base (gutter-like [ a ], narrow [ b ]); B , presence ( a ) or absence ( b ) of a round bulge at distal third of the accessory piece; C , shape of the accessory piece ending (double [ a ], single [ b ]); D , shape of the accessory piece ending hook (thin and long [ a ], wide and short [ b ]); E , shape of penis heel (wide [ a ], narrow [ b ]); F , shape of penis extremity and accessory piece (folded back and close C [ a ], straight and open C [ b ]); G , size of accessory piece (long [ a ], short [ b ]). FIG. 13.— Illustration of identification key for Cichlidogyrus Paperna,1960 species: A , shape of the penis (spirally coiled [ a ], not spirally coiled [ b ]); B , number of turns in spirally coiled penis and shape of the sclerotised vagina (4-5 and sinuous [ a ], more than 4-5 and spirally coiled [ b ]); C , number of turns in spirally coiled penis (8-9 [ a ], 14-15 [ b ]); D , penis diameter (large [ a ], filiform [ b ]); E , shape of the penis (straight [ a ], curved [ b ]); F , shape of the penis (G [ a ], single looped [ b ]). 79. No visible vagina, developed swollen portion (more than half penis total length) ( Fig. 14D, a ) ................................................................................................ Cichlidogyrus sanjeani — Visible S-shaped vagina, reduced swollen portion (less than one fifth of penis total length) ( Fig. 14D, b ) ............................................................................................................ 80 80. Accessory piece ending in a hook ( Fig. 14E, a ) ...................... Cichlidogyrus amphoratus — Accessory piece bifurcated, lined along convex face with tubercles ( Fig. 14E, b ) ............ .................................................................................................... Cichlidogyrus ornatus 81. Accessory piece with heavily sclerotised portion L-shaped, uncinuli pairs II and III to VII (11 and 13 µm ) ( Fig. 14F, a ) ........................................................ Cichlidogyrus giostrai — Accessory piece with heavily sclerotised portion C-shaped, uncinuli pairs II and III to VII (16 and 20 µm ) ( Fig. 14F, b ) .......................................................... Cichlidogyrus njinei 82. Penis length less than 40 µm .................................................... Cichlidogyrus lagoonaris — Penis length more than 55 µm .................................................................................. 83 83. Short, large and conical vagina ( Fig. 14G, a ) ......................... Cichlidogyrus zambezensis — Short, thin and S-shaped vagina ( Fig. 14G, b ) ............................. Cichlidogyrus karibae 84. Penis length less than 60 µm .................................................................................... 85 — Penis length more than 60 µm .................................................................................. 92 85. Sclerotised vagina ..................................................................................................... 86 — Vagina not visible ..................................................................................................... 88 86. Single C-shaped accessory piece with a thin extension at its base ( Fig. 15A, a ) .............. .................................................................................................... Cichlidogyrus levequei — Complex accessory piece with well-sclerotised C-shaped portion ( Fig. 15A, b ) ......... 87 87. Accessory piece with a C-shaped portion sharply arched, with thin plates at each extremity ( Fig. 15B, a ) .......................................................................... Cichlidogyrus slembroucki — Accessory piece with C-shaped portion slightly arched, without plates at its extremities, and a digitations at its middle ( Fig. 15B, b ) .................................. Cichlidogyrus acerbus 88. Penis with heel ( Fig. 15C, a ) .................................................................................... 89 — Penis without heel ( Fig. 15C, b ) ............................................................................... 91 89. Very thin heel, accessory piece ending in claws ( Fig. 15D, a ) ...... Cichlidogyrus fontanai — Developed heel, accessory piece with forked ending ( Fig. 15D, b ) ............................ 90 90. Penis length less than 40 µm , large dorsal bar auricles ( Fig. 15E, a ) ............................... .................................................................................................... Cichlidogyrus rognoni — Penis length more than 40 µm , dorsal bar auricles smaller ( Fig. 15E, b ) ........................ ................................................................................................. Cichlidogyrus bifurcatus 91. Penis length more than 40 µm , accessory piece thin at level of insertion on basal bulb ( Fig. 15F, a ) ......................................................................... Cichlidogyrus haplochromii — Penis length less than 37 µm , accessory piece large at level of insertion ( Fig. 15F, b ) ...... .................................................................................................... Cichlidogyrus tilapiae 92. Accessory piece straight or slightly curved ( Fig. 16A, a ) ............................................ 93 — Accessory piece sharply curved ( Fig. 16A, b ) ............................................................ 95 FIG. 14. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , size of the sclerotised vagina and shape of accessory piece ending (long and rounded [ a ], short and hooked [ b ]); B , presence ( a ) or absence ( b ) of a swollen portion in the penis; C , well marked ( a ) or diffuse ( b ) swollen portion in the penis; D , relative size of the swollen portion in the penis (developed [ a ], reduced [ b ]); E , shape of accessory piece ending (hooked [ a ], bifurcated [ b ]) and presence of tubercles along the convex face; F , shape of accessory piece heavily sclerotised portion (L [ a ], C [ b ]); G , shape of the sclerotised vagina (large and conical [ a ], thin and S-shaped [ b ]). FIG. 15. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , shape of the accessory piece (simple with extension [ a ], complex [ b ]); B , presence ( a ) or absence ( b ) of thin plates at the extremity of the accessory piece; C , presence ( a ) or absence ( b ) of a penis heel; D , size of the penis heel and shape of the accessory piece ending (thin and presence of claws [ a ], developed and forked [ b ]); E , size dorsal bar auricles (large [ a ], small [ b ]); F , size of the penis and width of accessory piece insertion at the level of the penis (more than 40 μm and thin [ a ], less than 37 μm and large [ b ]). FIG. 16. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A , shape of accessory piece (straight [ a ], sharply curved [ b ]); B , absence ( a ) or presence ( b ) of a penis heel; C , shape of the sclerotised vagina (straight and large [ a ], sinuous and thin [ b ]); D , shape of the accessory piece ending and of the penis (rounded and looped [ a ], forked and C [ b ]); E , accessory piece extremities shape and length (smooth and equal [ a ], rough and unequal [ b ]). 93. Penis with no visible heel ( Fig. 16B, a ) ..................................... Cichlidogyrus longipenis — Penis with developed heel ( Fig. 16B, b ) .................................................................... 94 94. Straight and large vagina, penis length 70 µm ( Fig. 16C, a ) ......... Cichlidogyrus sclerosus — Long, thin and sinuous vagina, penis length 145 µm ( Fig. 16C, b ) ... Cichlidogyrus cirratus 95. Accessory piece with rounded ending, looped penis ( Fig. 16D, a ) ... Cichlidogyrus amieti — Accessory piece forked ending, C-shaped penis ( Fig. 16D, b ) ................................... 96 96. Extremities of accessory piece smooth and about equal in length ( Fig. 16E, a ) .............. ..................................................................................................... Cichlidogyrus cubitus — Extremities of accessory piece highly unequal in length, the longer one with rough patch on its convex face ( Fig. 16E, b ) .............................................. Cichlidogyrus louipaysani