Tyrophagus perniciosus Zakhvatkin, 1941: 104

Masoudian, Farshad, Khanjani, Mohammad, Saboori, Alireza & Nejad, Karim Haddad Irani-, 2017, Ontogeny of Tyrophagus perniciosus Zakhvatkin (Acari: Acaridae) from Western Iran, Persian Journal of Acarology 6 (4), pp. 225-243 : 226-240

Tyrophagus perniciosus Zakhvatkin, 1941: 104 .

Diagnosis

Dorsal seta d1 considerably longer than seta c1, 2.4–3.2 times; eyespots absent; supracoxal seta scx tapering from base to tip or slightly widened in basal 2/3; spermathecal duct very wide along its entire length; solenidion ωΙoftarsi I-II cylindrical and obviously widened apically. Coxal plates II broadly triangular and with a well-developed apodeme near base of trochanter seta; aedeagus obviously curves, sickle-shaped.

Female ( Figs. 1–12 , 53–54, 63; n = 6) – Idiosoma oval. Length of body including gnathosoma 549– 687, excluding gnathosoma 465–575; width 263–375.

Dorsum ( Figs. 1–3 ). Prodorsal shield punctate, with 2 pairs of setae (vi and ve) almost pentagonal in shape with lateral margins a slightly concave; 78–102 long, 90–125 wide between setae ve -ve. Eyespots absent ( Figs. 1, 3 ); Basal lobe of Grandjean's organ with one large tooth and four small teeth, 17–20, 5–7, 4–5, 2–3 and 2 long, respectively ( Fig. 2 ). Supracoxal seta scx pectinate with 8 rays on each side and broadly expanding from the base but gradually tapers to a fine point distally ( Figs. 1, 3 ). All dorsal setae finely serrated. All opisthosomal setae whip-like except c1, d1 and d2. Opisthosoma with 3 pairs of lyrifissures (ia, im and ip) and 1 pair of opisthosomal gland (gla) at level of seta e1; Seta c1 is the shortest whereas seta h1 is the longest dorsal setae. Length of dorsal setae: vi 80–112; ve 41–57; sci 178–237; sce 122–150; scx 37–50; c1 27–45; c2 180–225; cp 125–153; d1 95–133; d2 28–45; e1 250–330; e2 195–270; f2 310–350; h1 365–400; h2 325–375. Distances: vi -vi 8–12; vi -ve 45–60; ve -ve 96–110; sce -sce 95–110; sci -sce 25–27; sci -sci 35–45; sce -ve 67–82; c1-c1 110–155; c1-c2 38–50; c2-c2 185–255; c2-cp 28–40; cp -cp 252–330; c1-d1 48–57; d1-d1 65–90; d1- d2 78–112; d2-d2 195–265; d2-gla 50–60; gla -gla 210–305; gla - e1 47–65; e1-e1 115–170; e1-e2 87– 135; e2-e2 210–310; e2-f2 45–55; f2-f2 200–255; e1-h1 118–155; h1-h1 88–112; h1-h2 40–65; h2- h2 85–100. Ratio: d1 / c1 2.95–3.51; d1 / d2 2.95–3.40; d2 / c1 1.00–1.04.

Gnathosoma ( Figs. 4–5 ) – Punctate; palp two-segmented, palp tarsus with a simple seta (pt) and one solenidion (ω) 15–20 and 4–7 long, respectively; palp tibia with two simple setae (d 18–25 and l 17–20 long); infracapitulum with one simple seta (m) 44–50 long, distance: m -m 18–25. Rutellum distinct and developed ( Fig. 4 ); chelicerae 95-110 long, cheliceral seta cha spine-like 5–7 long, movable and fixed digits with 4–5 teeth ( Fig. 5 ). Palp coxa with one elcp setae 17–23 ( Fig. 4 ).

Venter ( Figs. 6–8 , 63 ) – Coxal plates I divided with apodemes I anteriorly on each side with 3 nodules; coxal plates II broadly triangular and with an well-developed apodeme near base of trochanter seta ( Fig. 63 ); between coxae II and III with a pair of thin sclerotized sejugal apodemes, 45–65 long; coxal plates III-IV each with apodemes ( Fig. 6 ); genital region (posterior to sejugal apodemes to coxae IV) with two pairs of genital papillae 18-25 long and 13-15 wide, a pair of setae (g) and genital folds ( Figs. 6–7 ). Anal region with three pairs of adanal setae (ad1-3) and three pairs of pseudoanal setae (ps1-3), seta ps1 the longest anal setae ( Fig. 6 ); a pair of lyrifissures (ih) at level of seta ad2 base. Copulatory opening (8–11 in diameter) located posterior to anal opening, spermathecal duct (38–50 long) very wide along its entire length and join base of spermathecal sac and with a long neck, 80 long ( Figs. 6, 8 ); Length of ventral setae: 1a 37–50, c3 37–50, 3a 20–25, 3b 31–50, 4a 38–47, g 15-20, ad3 14–23, ad2 22–37, ad1 23–37, ps3 25–35, ps2 125–170, ps1 200– 260, h3 270–300.

Legs ( Figs. 9–12 , 53–54) – Setal formulae of leg segments I-IV as follows (solenidia and special setae in parentheses): coxae 1-0-2-1; trochanters 1-1-1-0; femora 1-1-0-1, genua 2(2 σ)-2(1 σ)-1(1 σ)- 0; tibiae 2(1 ϕ)- 2 (1 ϕ)-1 (1 ϕ)-1 (1 ϕ); tarsi 13 (3ω,18)-12 (1ω)-10-10. Solenidion ωΙand ωof legs I- II cylindrical with obviously widened apex ( Figs. 9–10 , 53–54). Tarsi I-II with solendidion ωΙcylindrical and distinctly widened apically ( Figs. 9–10 , 53–54). Measurements of leg segments and setae as follows: Leg I: Tr 45–55, pR 23–37, Fe 52–70, vF 37–50, Ge 35–44, cG 30–41, mG 44– 60, 01 55–65, 02 27–34, Ti 30–45, gT 25–35, hT 30–37, ϕ110–137, Ta (L.) 60–72, Ta (W.) 21–27, ωΙ 17–21, ω25–8, ω329–35, 83, aa 16–24, ba 15–23, wa 37–42, ra 30–37, la 19–22, d 29–40, e 6– 10, f 16–20, p 4–6, q 4–6, s 6–7, u 5–6, v 5–6, empodial claws 11–14; Leg II: Tr 37–45, pR 28–35, Fe 50–60, vF 50–63, Ge 30–44, cG 26–35, mG 40–50, 022–26, Ti 30–40, gT 25–33, hT 27–40, ϕ 125–135, Ta (L.) 55–65, Ta (W.) 18–22, ω16–21, ba 17–21, wa 35–42, ra 30–33, la 19–24, d 23– 27, e 5–7, f 14–19, p 4–5, q 4–5, s 6–8, u 5–6, v 5–6, empodial claws 10–12; Leg III: Tr 35–40, sR 33–40, Fe 35–50, Ge 30–40, nG 42–50, 0 20–25, Ti 32–40, kT 27–41, ϕ 125–141, Ta (L.) 60–75, Ta (W.) 16–20, w 27–35, r 23–30, d 18–25, e 6–9, f 15–21, p 3, q 3, s 5–7, u 4–6, v 4–6, empodial claws 9–12; Leg IV: Tr 35–50, Fe 50–60, Ge 40–50, Ti 35–45, kT 28–35, ϕ 121–150, Ta (L.) 75–90, Ta (W.) 15–18), w 27–36, r 17–23, d 24–35, e 7–8, f 17–20, p 3–4, q 3–4, s 5–7, u 5–6, v 5–6, empodial claws 9–12; Fe, Ge and Ti IV with minute preapical process ( Fig. 12 ).

Male ( Figs. 13–24 , 55–56, 64–66; n = 5) – Idiosoma oval, length of body including gnathosoma 433–502, excluding gnathosoma 360–437; width 210–225.

Dorsum ( Figs. 13–15 ). Prodorsal shield punctate, with two pairs of seta (vi and ve) nearly pentagonal in shape with lateral margins slightly concave; 57–70 long, 73–90 wide between setae ve– ve. Eyespots absent; Basal lobe of Grandjean’s organ with one large tooth and four small teeth, 13– 16, 6–7, 6–7, 3 and 3 long, respectively ( Fig. 14 ). Supracoxal seta scx pectinated with 9–10 branches on each side ( Figs. 14–15 ). All dorsal setae finely serrated. All opisthosomal setae whip like except c1 and d2. Opisthosoma with three pairs of lyrifissures (ia, im and ip) and one pair of opisthosomal glands (gla) at level of seta e1; setae c1 the shortest and seta h1-2 the longest dorsal setae. Length of dorsal setae: vi 70–80; ve 38–43; sci 150–170; sce 82–100; scx 25–35; c1 25–26; c2 180–228; cp 100–120; d1 95–120; d2 24–25; e1 270–302; e2 212–218; f2 312–330; h1 363–385; h2 337–370. Distances: vi -vi 8–10; vi -ve 30–38; ve -ve 68–85; sce -sce 70–72; sci -sce 15–18; sci -sci 28–30; c1-c1 87–105; c1-c2 24–32; c2-c2 150–165; c2-cp 18–30; cp -cp 170–215; c1-d1 37–45; d1-d1 47–65; d1- d2 58–80; d2-d2 147–165; d2-e2 75–90; d2-gla 35–50; gla -gla 150–162; d2- e1 33–38; e1-e1 85– 102; e1- e2 57–62; e2-e2 150–175; e2-f2 31–40; f2-f2 140–155; e1-h1 83–112; h1-h1 65–75; h1-h2 30–42; h2-h2 40–45.

Gnathosoma ( Figs. 16–17 ) – Punctate; palp two-segmented, palp tarsus with a simple seta (pt) and one solenidion (ω) 15–16 and 4–5 long, respectively; palp tibia with 2 simple setae (d 18–23 and l 15–18 long); infracapitulum with 1 simple seta (m) 32–42 long, distance: m -m 16–20. Rutellum distinct and developed ( Fig. 16 ); chelicerae 75–83 long, cheliceral seta cha spine like 5 long, movable and fixed digits with 3–4 teeth ( Fig. 17 ). Palp coxa with one elcp setae 14–15 ( Fig. 18 ).

Venter ( Figs. 18–20 , 64–66 ) – Coxal plates I divided with apodemes I anteriorly with 4 nodes on each side; coxal plates II with an well-developed apodeme near base of trochanter seta ( Fig. 64 ); between coxae II and III with a pair of thin sclerotized sejugal apodemes, 50–55 long; coxal plates III-IV each one with apodemes ( Fig. 18 ); genital region (between coxae IV) with two pairs of genital papillae 15 long and 10 wide, and a pair of setae (g); aedeagus obviously curves, sickle-shaped, aedeagus shaft 24–30 long, lateral arms supporting aedeagus turning inwards ( Figs. 19–20 , 65–66 ). Anal region with three pairs of pseudoanal setae (ps1-3), seta ps1 and ps3 the longest and the shortest anal setae respectively, a pair of anal suckers 20 and a pair of anal discs 4 in diameter ( Figs. 18–19 ). Length of ventral setae: 1a 27–35, c3 22–30, 3a 18–23, 3b 30–36, 4a 35–40, g 13–15, ps3 13–15, ps2 35–43, ps1 172–180, h3 250–280.

Legs ( Figs. 21–24 , 55–56) – Setal formulae of leg segments I-IV as follows (solenidia and special setae in parentheses): coxae 1-0-2-1; trochanters 1-1-1-0; femora 1-1-0-1, genua 2(2σ)-2(1σ)-1(1σ)- 0; tibiae 2(1ϕ)- 2(1ϕ)-1(1ϕ)-1(1ϕ); tarsi 13(3ω,18)-12(1ω)-10-10; solenidion ωΙand ωof legs I-II cylindrical with obviously widened apex ( Figs. 21–22 , 55–56). Measurements of leg segments and setae as follows: Leg I: Tr 35–41, pR 25–33, Fe 49–50, vF 38–41, Ge 35–37, cG 27–30, mG 36–41, 01 38–50, 02 17–20, Ti 28–30, gT 20–23, hT 23–27, ϕ105–117, Ta (L.) 70–82, Ta (W.) 18–19; ωΙ 15–17, ω25–7, ω325–28, 83–4, aa 16–17, ba 20–24, wa 29–31, ra 24–30, la 16–17, d 28–30, e 4– 6, f 13–16, p 3–4, q 3–4, s 5–7, u 4–5, v 4–5 empodial claws 9–11; Leg II: Tr 29–40, pR 25–28, Fe 40–45, vF 43–50, Ge 30–35, cG 23–27, mG 30–37, 018–19, Ti 25, gT 17–19, hT 24–25, ϕ117–125, Ta (L.) 49–50, Ta (W.) 15–17, ω 16–18, ba 21–22, wa 27–33, ra 26–30, la 15–18, d 26–28, e 4–6, f 13–16, p 3–4, q 3–4, s 5–7, u 4–5, v 4–5, empodial claws 8–12; Leg III: Tr 28–35, sR 30–37, Fe 33– 38, Ge 28–30, nG 37–42, 0 15–20, Ti 25, kT 27–33, ϕ135–140, Ta (L.) 56–60, Ta (W.) 13, w 26–30, r 24–28, d 23–24, e 5–6, f 13–14, p 3–4, q 3, s 5, u 4–5, v 4–5, empodial claws 8–10; Leg IV: Tr 30– 35, Fe 40–45, wF 25–35, Ge 35–37, Ti 25–30, kT 23–29, ϕ118–130, Ta (L.) 67–70, Ta (W.) 10–12, w 21–30, r 16–20, d 2–3, e 2–3, f 17–20, s 5, p 3–5, q 3, u 4–5, v 4–5, empodial claws 7–9. Tarsus IV with two suckers (seta d on proximal and seta e on distal sucker), distance between base of seta d and proximal segment of tarsus 15–21, e -d 15–18, e -f 25–30 ( Fig. 24 ).

Tritonymph ( Figs. 25-35 , 57-58; n= 6) – Idiosoma oval. Length of body including gnathosoma 330– 385, excluding gnathosoma 265–320; width 154–190.

Dorsum ( Figs. 25–27 ) – Prodorsal shield punctate, 55–60 long, 60–80 wide between setae ve -ve, nearly pentagonal in shape with lateral margins slightly concave and with two pairs of seta (vi and ve); eyespots absent; basal lobe of Grandjean's organ with one large tooth and three small teeth, 12– 15, 4–5 long, 3 and 1–2 respectively ( Fig. 27 ). Supracoxal seta scx pectinated with 7–8 rays on each side ( Figs. 25, 27 ). All dorsal setae finely serrated. All opisthosomal setae whip like except c1, d1 and d2. Opisthosoma with three pairs of lyrifissures (ia, im and ip) and one pair of opisthosomal glands (gla) at level of seta e1; setae c1 the shortest and setae f2, h1 and h2 the longest dorsal setae. Length of dorsal setae: vi 50–61; ve 28–39; sci 100–124; sce 58–73; scx 24–27; c1 16–17; c2 115– 135; cp 70–85; d1 36–44; d2 16–20; e1 175–215; e2 135–165; f2 237–275; h1 260–295; h2 250–295. Distances: vi -vi 6–8; vi -ve 28–30; ve -ve 60–75; sce -sce 56–67; sci -sce 15; sci -sci 23–30; sci -c1 55– 68; c1-c1 70–90; c1-c2 16–25; c2-c2 110–140; c2-cp 15–20; cp -cp 145–180; c1-d1 33–41; d1-d1 33– 45; d1-d2 44–55; d2-d2 100–131; d2- e2 58–67; d2-gla 18–30; gla -gla 97–138; d2- e1 22–28; e1- e1 53–80; e1- e2 44–60; e2-e2 120–150; e1-f2 60–80, e2-f2 24–35; f2-f2 91–130; e1-h1 56–70; h1-h1 44–60; h1-h2 17–30; h2-h2 24–40.

Gnathosoma ( Figs. 28–29 ) – Palp two-segmented, palp tarsus with a simple seta (pt) and one solenidion (ω) 8–11 and 4–5 long, respectively; palp tibia with two simple setae (d 14–18 and l 11– 14 long); infracapitulum with one simple seta (m) 28–30 long, distance: m -m 15–18. Rutellum distinct and developed ( Fig. 28 ); chelicerae 65–70 long, cheliceral seta cha spine like 5–6 long, movable and fixed digits with 3–4 teeth ( Fig. 28 ). Palp coxa with one elcp setae 9–12 ( Fig. 29 ).

Venter ( Figs. 30–31 ). Coxal plates I divided with apodemes I anteriorly with 3 nodes on each side; coxal plates II with a well-developed apodeme; a pair of thin sclerotized and narrow sejugal apodemes between coxae II and III 36–40 long; genital region with two pairs of genital papillae 10– 13 long and 6–9 wide and a pair of setae (g) ( Figs. 30–31 ). Anal region with three pairs of pseudoanal setae (ps1-3), setae ps1 and ps3 the longest and shortest anal setae, respectively ( Fig. 30 ); a pair of lyrifissures (ih) at level of setae ps2. Length of ventral setae: 1a 20–26, c3 25–30, 3a 12–17, 3b 23– 25, 4a 25–27, g 11–13, ps3 14–17, ps2 18–24, ps1 68–100, h3 160–205.

Legs ( Figs. 32–35 , 57–58) – Setal formulae of leg segments I-IV as follows (solenidia and special setae in parentheses): coxae 1-0-2-1; trochanters 1-1-1-0; femora 1-1-0-1, genua 2(2σ)-2(1σ)-1(1σ)- 0; tibiae 2(1ϕ)- 2(1ϕ)-1(1ϕ)-1(1ϕ); tarsi 13(3ω,18)-12(1ω)-10-10 ( Figs. 32–35 ). Leg I-II with cylindrical and apex obviously widened solenidion (ωΙand ω, respectively) ( Figs. 32–33 , 57–58). Measurements of leg segments and setae as follows: Leg I: Tr 25–30, pR 16–19, Fe 30–37, vF 27– 30, Ge 24–27, cG 17–21, mG 25–28, 01 30–38, 02 15–17, Ti 20–23, gT 14–19, hT 15–20, ϕ 80–95, Ta (L.) 35–45, Ta (W.) 13–15, ωΙ 12–14, ω2 3–4, ω315–20, 83, aa 9–11, ba 11–15, wa 22–30, ra 17–22, la 13–17, d 18–23, f 10–13, e 4, p 3, q 3, s 4–5, u 4–5, v 4–5, empodial claws 8–10; Leg II: Tr 20–25, pR 16–18, Fe 30–35, vF 30–40, Ge 20–25, cG 10–15, mG 16–22, 012–15, Ti 20–23, gT 13–14, hT 13–18, ϕ76–90, Ta (L.) 33–40, Ta (W.) 11–13, ω13–14, ba 11–14, wa 19–26, ra 18–21, la 12–16, d 15–19, e 3–4, f 9–10, p 2–3, q 2–3, s 4, u 3–4, v 3–4 empodial claws 7–9; Leg III: Tr 20–27, sR 17–25, Fe 24–25, Ge 20–24, nG 20–25, 0 10–12, Ti 19–24, kT 18–21, ϕ95–105, Ta (L.) 38–43, Ta (W.) 9–10, w 17–21, r 13–18, d 14–16, e 3–4, f 8–10, p 2, q 2, s 3–4, u 3, v 3, empodial claws 7; Leg IV: Tr 25–30, Fe 27–33, wF 17–21, Ge 20–25, Ti 21–24, kT 17–20, ϕ74–90, Ta (L.) 50–55, Ta (W.) 9–10, w 17–23, r 10–14, d 15–17, e 3, f 8–11, p 2, q 2, s 3–4, u 3, v 3, empodial claws 7–9; Fe, Ge and Ti IV with minute preapical process ( Fig. 35 ).

Deutonymph (Hypopus) – Absent.

Protonymph ( Figs. 36–44 , 59–60; n = 4) – Idiosoma oval. Length of body including gnathosoma 270–295, excluding gnathosoma 210–250; width 110–156.

Dorsum ( Figs. 36–37 ) – Prodorsal shield present and eyespots absent. Basal lobe of Grandjean's organ with one large and three small teeth, 9–10, 4–5, 3 and 3 long, respectively ( Fig. 37 ). Supracoxal seta scx pectinate with 6–7 rays seven branches on each side ( Figs. 36, 37 ). All dorsal setae finely serrated. All opisthosomal setae whip like except c1, d1 and d2. Opisthosoma with three pairs of lyrifissures (ia, im and ip) and one pair of opisthosomal gland (gla) at level of seta e1; setae c1 and d2 the shortest and setae h1 the longest dorsal setae. Length of dorsal setae: vi 39–42; ve 26–29; sci 80–91; sce 39–43; scx 22–24; c1 12–13; c2 80–100; Cp 48–60; d1 16–20; d2 10–13; e1 145–170; e2 80–100; f2 180–190; h1 210–232; h2 180–195. Distances: vi -vi 5–6; vi -ve 19–20; ve -ve 42–45; sce - sce 50–63; sci -sce 14–16; sci -sci 20–22; sci -c1 48–55; c1-c1 60–75; c1-c2 15–21; c2-c2 90–115; c2- Cp 12–17; Cp -Cp 115–150; c1-d1 25–35; d1-d1 25–35; d1-d2 33–46; d2-d2 75–100; d2- e2 55–60; d2-gla 20–24; gla -gla 90–105; d2- e1 23–28; e1- e1 48–60; e1- e2 30–48; e2-e2 75–100; e2-f2 17–23; f2-f2 65–80; e1-h1 55–62; h1-h1 37–40; h1-h2 12–15; h2-h2 20–30.

Gnathosoma ( Figs. 38–39 ) – Palp two-segmented, palp tarsus with a simple seta (pt) and one solenidion (ω) 6–7 and 3–4 long, respectively; palp tibia with two simple setae (d 11–12 and l 8–9 long); infracapitulum with one simple seta (m) 18–22 long, m -m 13–16. Rutellum visible ( Fig. 39 ); chelicerae 50–53 long, cheliceral seta cha spine like 4 long ( Fig. 38 ). Palp coxa with one elcp setae 8–9.

Venter ( Fig. 40 ) – Coxal plates I divided with apodemes I anteriorly without nodes; coxal plates II with a well-developed apodeme; a pair of thin sclerotized and narrow sejugal apodemes between coxae II and III 28–34 long; genital region with one pair of genital papillae 10–11 long and 7 wide and a pair of setae (g) ( Fig. 40 ). Anal region with three pairs of pseudoanal setae (ps1-3) sub equal in length ( Fig. 40 ); a pair of lyrifissures (ih) between setae ps1-2. Length of ventral setae: 1a 17, 3b 18– 19, c3 13–16, g 10–12, ps3 8–11, ps2 10–12, ps1 12–15, h3 100–103.

Legs ( Figs. 41–44 , 59–60) – Setal formulae of leg segments I-IV as follows (solenidia and special setae in parentheses): coxae 1-0-1-0; trochanters 0-0-0-0; femora 1-1-0-0, genua 2(2σ)-2(1σ)-1(1σ)- 0; tibiae 2(1ϕ)-2(1ϕ)-1(1ϕ)-0; tarsi 13(2ω,18)-12(1ω)-10-9; solenidion ωΙand ω of legs I-II cylindrical with obviously widened apex ( Figs. 41–44 , 59–60). Measurements of leg segments and setae as follows: Leg I: Tr 21–23, Fe 28–30, vF 16–22, Ge 17–22, cG 14–15, mG 15–16, 01 20, 02 8, Ti 16– 18, gT 12–15, hT 11, ϕ 65–69, Ta (L.) 30–33, Ta (W.) 11–12, ωΙ 10–12, ω2 4, 8 3, aa 6–8, ba 8– 11, wa 18–19, ra 13–17, la 9–10, d 17, e 3–5, f 9, p and q 3, s 4, v and u 3–4, empodial claws 7–10; Leg II: Tr 20, Fe 25–29, vF 23–24, Ge 17–18, cG 10–11, mG 14, 08–9, Ti 15–17, gT 11–13, hT 9, ϕ59–70, Ta (L.) 25–30, Ta (W.) 9–10, ω10, ba 8–10, wa 13, ra 12–13, la 8–9, d 14–15, e 2–3, f 9, p and q 2–3, s 3–4, v and u 3–4, empodial claws 7–10; Leg III: Tr 15–18, Fe 17–20, Ge 15–17, nG 16–18, 0 7, Ti 15, kT 12–16, ϕ70–73, Ta (L.) 29–31, Ta (W.) 8–9, w 15–18, r 10, d 11–13, e 3, f 8, p and q 2, s 3, v and u 2–3, empodial claws 6–10; Leg IV: Tr 23–24, Fe 20–21, Ge 16–17, Ti 15–17, Ta (L.) 34–35, Ta (W.) 8, w 12–15, r 8–11, d 11–13, e 2–3, p and q 2, s 2–3, v and u 2–3, empodial claws 6–10; Ge IV with minute preapical process ( Fig. 44 ).

Larva ( Figs. 45–52 , 61–62; n = 5) – Idiosoma oval. Length of body including gnathosoma 180–198, excluding gnathosoma 145–160; width 79–85.

Dorsum ( Figs. 45–46 ) – Prodorsal shield present and eyespots absent. Basal lobe of Grandjean’s organ finger like and with one large and two small teeth, 5, 1 and 1 long, respectively ( Fig. 45 ). Supracoxal seta scx pectinate with 4–5 rays branches on each side ( Fig. 45–46 ). All dorsal setae finely serrated. Opisthosomal setae e1, h1 whip like. Setae f2 and h3 absent. Opisthosoma with one pair of opisthosomal glands (gla) at level of seta e1; setae c1, d1 and d2 the shortest and setae h1 the longest dorsal setae. Length of dorsal setae: vi 24–25; ve 17–20; sci 50; sce 21–22; scx 12–15; c1 10–12; c2 47–50; Cp 30–35; d1 10–11; d2 7–10; e1 95; e2 18 –20; h1 185–188; h2 50. Distances: vi -vi 4; vi -ve 15–18; ve -ve 32–38; sce -sce 43–45; sci -sce 11–14; sci -sci 18–19; sci -c1 40–45; c1-c1 44–46; c1-c2 10–12; c2-c2 68–70; c2-Cp 5–12; Cp-Cp 80–85; c1-d1 25–27; d1-d1 20; d1-d2 20–25; d2-d2 60–65; d2- e2 35–40; d2-gla 12–14; gla -gla 58–62; d2- e1 22–25; e1- e1 31–35; e1- e2 20–25; e2- e2 55; e1– h1 35; h1-h1 30; h2-h2 11–15.

Gnathosoma ( Figs. 47–48 ) – Palp two-segmented, palp tarsus with a simple seta (pt) and one solenidion (ω) 5–6 and 2–3 long, respectively; palp tibia with two simple setae (d 9–10 and l 6–7 long); infracapitulum with one simple seta (m) 16–17 long, m -m 9–10. Rutellum visible ( Fig. 48 ); chelicerae 35–37 long, cheliceral seta cha spine like 2–3 long, movable and fixed digits with 4 teeth ( Fig. 47 ). Palp coxa with one elcp setae 5–6.

Venter ( Fig. 49 ) – Coxal plates I-III with narrow apodemes; Claparède organ between coxae I- II, with base tube like and 15–16 in length terminating in a spherical knob distally, 4 in diameter), setae 3a, 4a, genital papillae and seta, adanal and pseudoanal setae absent; a pair of lyrifissures (ih) anterolateral of setae h2. Length of ventral setae: 1a 12–13, 3b 15–16, c3 10–14, h2 50.

Legs ( Figs. 50–52 , 61–62) – Setal formulae of leg segments I-IV as follows (solenidia and special setae in parentheses): coxae 1-0-1; trochanters 0-0-0; femora 1-1-0, genua 2(2σ)-2(1σ)-1(1σ); tibiae 2(1ϕ)- 2(1ϕ)-1(1ϕ); tarsi 13(1ω,18)-12(1ω)-10; solenidion ωΙand ω of legs I-II cylindrical with obviously widened apex ( Figs. 50–52 , 61–62). Measurements of leg segments and setae as follows: Leg I: Tr 16–17, Fe 20, vF 12–15, Ge 12–154, cG 12–13, mG 8–10, 01 12–14, 02 8, Ti 11–12, gT 5–7, hT 7–10, ϕ 52–55, Ta (L.) 25–26, Ta (W.) 8, ω 8, 83, aa 5–6, ba 7–8, wa 10–12, ra 8–10, la 5– 7, d 10, e 3, f 7–8, q and p 2, s 3–4, v and u 2–3, empodial claws 5–6; Leg II: Tr 15, Fe 18–20, vF 16–17, Ge 13–15, cG 7–12, mG 8–9, 04–6, Ti 10–11, gT 7, hT 6–8, ϕ48–49, Ta (L.) 20, Ta (W.) 7– 8, ω6–8, ba 7–8, wa 9–12, ra 8, la 6–7, d 10–12, e 2, f 6–9, q and p 2, s 3, v and u 2–3, empodial claws 5–6; Leg III: Tr 15–16, Fe 15, Ge 13–14, nG 15–17, 0 5, Ti 13–15, kT 11–14, ϕ53–54, Ta (L.) 24–28, Ta (W.) 6, w 13, r 6–7, d 9, e 2, f 6, q and p 1, s 2–3, v and u 1–2, empodial claws 5 long.

Figures 53–62. Tyrophagus perniciosus Zakhvatkin, 1941 – Solenidion ωΙon tarsus I-II: 53–54. Female; 55–56. Male; 57–58. Tritonymph; 59–60. Protonymph; 61–62. Larva.

DISCUSSION and CONCLUSION

This is the first re-description of all stages of this species from Iran. This re-description shows minor differences between Iranian specimens and re-description from New Zealand ( Fan and Zhang 2007), however differs from the later in: The ratio of hysterosomal setae d1 about 3.1–3.9 × length of d2 whereas 2.7–3.8; 2. Leg I-IV with empodial claws 10–12 and 9–12 long respectively in Iranian specimens vs. 15–23 and empodial claws 17–24 long in New Zealand specimens. Male: The ratio of hysterosomal setae d1 about 3.7–5.0 × length of c1 and 3.7–5.4 × length of d2; d2 about 0.92–1.0 × length of c 1 in Iranian species whereas 3.2–4.0, 2.9–3.4 and 1.1–1.2 in New Zealand specimens, respectively. This study has been clearly shown the differential chaetotaxy especially hysterosomal region, organotaxy develops during ontogenetic (larva, protonymph, tritonymph and adults [♀ & ♂]) and also agree with the hypothesis of Griffiths et al. (1990). The ontogenetic characters of this species are presented in Table 1.

Figures

Figures 1-8. Tyrophagus perniciosus Zakhvatkin, 1941 (female) – 1. Dorsal view of idiosoma; 2. Grandjean's organ; 3. Prodorsal shield; 4. Gnathosoma; 5. Chelicera; 6. Ventral view of idiosoma; 7. Genital region; 8. Anal region, copulatory opening and spermatheca.

Figures 9–12. Tyrophagus perniciosus Zakhvatkin, 1941 (female) – 9. Leg I; 10. Leg II; 11. Leg III; 12. Leg IV.

Figures 63-66. Tyrophagus perniciosus Zakhvatkin, 1941: 63. Coxa II (female); 64. Coxa II (male); 65. Aedeagus; 66. Lateral view of aedeagus.

Figures 13–20. Tyrophagus perniciosus Zakhvatkin, 1941 (male) – 13. Dorsal view of idiosoma; 14. Grandjean's organ; 15. Prodorsal shield; 16. Gnathosoma; 17. Chelicera; 18. Ventral view of idiosoma; 19. Genital and anal regions; 20. Lateral view of aedeagus.

Figures 21–24. Tyrophagus perniciosus Zakhvatkin, 1941 (male) – 21. Leg I; 22. Leg II; 23. Leg III; 24. Leg IV.

Figures 25-31. Tyrophagus perniciosus Zakhvatkin, 1941 (tritonymph) – 25. Dorsal view of idiosoma; 26. Supracoxal seta (scx); 27. Grandjean's organ; 28. Chelicera; 29. Gnathosoma; 30. Ventral view of idiosoma; 31. Genital region.

Figures 32-35. Tyrophagus perniciosus Zakhvatkin, 1941 (tritonymph) – 32. Leg I; 33. Leg II; 34. Leg III; 35. Leg IV.

Figures 36-40. Tyrophagus perniciosus Zakhvatkin, 1941 (protonymph) – 36. Dorsal view of idiosoma; 37. Supracoxal seta (scx) and Grandjean's organ; 38. Chelicera; 39. Gnathosoma; 40. Ventral view of idiosoma.

Figures 41-44. Tyrophagus perniciosus Zakhvatkin, 1941 (protonymph) – 41. Leg I; 42. Leg II; 43. Leg III; 44. Leg IV.

Figures 45-52. Tyrophagus perniciosus Zakhvatkin, 1941 (larva) – 45. Dorsal view of idiosoma; 46. Supracoxal seta (scx) and Grandjean's organ; 47. Chelicera; 48. Gnathosoma; 49. Ventral view of idiosoma; 50. Leg I; 51. Leg II; 52. Leg III.