Morphological ontogeny of Oribatella similesuperbula (Acari: Oribatida: Oribatellidae) Author Seniczak, Stanisław Author Ivan, Otilia Author Marquardt, Tomasz Author Seniczak, Anna text Zootaxa 2021 2021-12-30 5086 1 90 110 journal article 2822 10.11646/zootaxa.5086.1.8 4d87b377-73cb-41b2-9cb6-613b62b5ae26 1175-5326 5814523 09DEF5BF-650B-4E58-9F2A-ACAE882AA72B Oribatella similesuperbula Weigmann, 2001 ( Figs. 1–15 ) Morphology of adult Adults studied here ( Figs. 1 , 2 , 3a , 4 , 5a ) similar to those investigated by Weigmann (2001) , but see Remarks . Mean length (and range) of females—374.2±8.5 (361–391, N= 30) and males—350.3±6.6 (range 343–361, N= 10); mean width of females—245.9±7.7 (235–259) and males—231.7±3.2 (229–235). Subcapitular setae h , m and a short and smooth ( Fig. 2 ). Chelicera chelate, seta cha longer than chb , both barbed ( Fig. 3b ). Palp relatively short, palpal setae smooth, except for slightly barbed l” on tibia ( Fig. 3c ). Formula of palp setae [trochanter to tarsus (+ solenidion ω)]: 0-2-1-3-9(1). Most epimeral setae short and smooth, except for longer and barbed 3b , 3c , 4a and 4b and clearly longer, thicker and barbed 4c ( Fig. 2 ). Most leg setae barbed, l” on genua and tibiae I and II, and l’ on genu and tibia IV longer and thicker than other setae ( Fig. 6 ). Formulae of leg setae [trochanter to tarsus (+ solenidia)]: I—1-5-3(1)-4(2)-20(2); II—1-5-3(1)-4(1)-15(2); III—2-2-1(1)-3(1)-15; IV—1-2-2-3(1)-12. Legs heterotridactylous. FIGURE 1. Oribatella similesuperbula , female. Dorsal aspect, legs partially drawn, scale bar 50 μm. FIGURE 2. Oribatella similesuperbula , female, ventral aspect, legs partially drawn, scale bar 50 μm. FIGURE 3. Oribatella similesuperbula , female. (a) Lateral aspect, legs partially drawn, scale bar 50 μm; mouthparts, right side, antiaxial aspect, scale bars 20 μm; (b) chelicera, (c) palp. Remarks : Females and males of O . similesuperbula investigated here are slightly longer than those investigated by Weigmann (2001) —length of females 340–380, and males 320–340. In our adults, the lamellar complex is located more anterior than depicted by Weigmann (2001) , and rostrum is not visible in dorsal aspect (versus visible). Other characters of adults are similar as in Weigmann (2001) . Diagnosis of juveniles Juveniles light-brown and nymphs apopheredermous, i.e. they carry exuvial scalps of previous instars away from dorsal integument, using modified setal pair da . Prodorsal and most gastronotal setae of juveniles long and barbed, except for medium sized h 2 and minute h 3 in larva, and medium sized c 1 , p 2 and p 3 in nymphs. Larva with 12 pairs of gastronotal setae, and central setae thicker than other setae, nymphs with 15 pairs, and long d -series. In all juveniles, humeral organ present. In nymphs, seta d on femora I and II, l” on genu and tibia I and genu II, and l’ on genu III clearly thicker than other leg setae. Description of juvenile stages Larva oval in dorsal aspect ( Figs. 5b–d , 7 ) and light brown. Prodorsum subtriangular, all prodorsal setae long and barbed, in longest, ro shortest ( Table 1 ). Mutual distance between setal pairs le slightly longer than between pair ro , and between pair in about two times longer than between pair ro . Seta le inserted midway between pairs ro and in . Bothridium rounded, bothridial seta narrowly clavate and barbed. TABLE 1. Measurements of some morphological characters of juvenile stages and adult of Oribatella similesuperbula (mean values of 10 individuals in μm, except for single tritonymph whose several setae are broken); Nd—not developed.
Morphological characters Larva Protonymph Deutonymph Tritonymph Adult
Body length 191 244 267 364 371
Body width 106 162 178 234 244
Length of prodorsum 64 57 109 116 112
Length of: seta ro 45 34 41 ? 75
seta le 50 75 72 64 80
seta in 69 86 96 ? 154
seta ex 44 54 72 ? 10
seta bs 66 71 115 98 93
seta c 1 109 32 38 56 Lost
seta c 2 106 91 128 152 25
seta c 3 93 78 109 ? Lost
seta da 110 93 95 ? Lost
seta dp 82 77 85 ? Lost
seta la 91 91 109 ? 19
seta lp 62 83 88 99 20
seta h 1 50 55 69 ? 11
seta h 2 11 46 72 ? 15
seta h 3 1 69 86 117 19
seta p 1 Nd 38 39 38 11
seta p 2 Nd 16 35 36 12
seta p 3 Nd 11 32 22 9
Genital opening Anal opening Nd 41 23 55 33 70 40 82 39 55
FIGURE 4. Oribatella similesuperbula , transmitted-light photomicrographs, adult, dorsal aspect. (a) Whole body, (b) anterior part, (c) medial part, (d) posterior part. Scale bars 50 μm. FIGURE 5. Oribatella similesuperbula , transmitted-light photomicrographs, dorsal aspect. (a) Leg II of adult; larva, (b) whole body, (c) anterior and medial part, (d) medial and posterior part. Scale bars 50 μm. FIGURE 6. Oribatella similesuperbula , segments of adult (part of femur to tarsus), right side, antiaxial aspect, seta on the opposite side not illustrated is indicated in the legend, scale bar 20 μm. (a) Leg I, tarsus ( pl’ ); (b) leg II; (c) leg III; (d) leg IV. FIGURE 7. Oribatella similesuperbula , larva, dorsal aspect, legs partially drawn, scale bar 20 μm. FIGURE 8. Oribatella similesuperbula , ventral part of hysterosoma, legs partially drawn, scale bars 20 μm. (a) Larva, (b) protonymph. FIGURE 9. Oribatella similesuperbula , lateral aspect, legs partially drawn, scale bars 50 μm. (a) Larva, (b) deutonymph. FIGURE 10. Oribatella similesuperbula , leg segments of larva (part of femur to tarsus), right side, antiaxial aspect, seta on the opposite side not illustrated is indicated in the legend, scale bar 10 μm. (a) Leg I, tarsus ( pl’ ); (b) leg II; (c) leg III. Gastronotum of larva with 12 pairs of setae, including minute h 3 inserted laterally to medial part of anal valves ( Figs. 7 , 8a , 9a ). Most gastronotal setae long and thick, with clear barbs, except for medium sized h 2 and minute h 3 ( Table 1 ), central setae thicker than lateral and posterior setae ( Figs. 5b–d , 7 , 9a ). Anal valves (segment P) glabrous. Cupule ih lateral to anterior part of anal valves, cupules ia , im , and ip not observed, gland opening lateroventral to seta lp , humeral organ anterior to seta c 3 ( Figs. 8a , 9a ). Most leg setae barbed, l” on genu and tibia I and genu II, and l’ on genu III thicker than other leg setae ( Fig. 10 ). Seta ft’ on tarsus I and ft on tarsi II and III slightly thicker than other tarsal setae. Nymphs more stocky than larva, with setiform bothridial seta. Gastronotum of protonymph with 15 pairs of setae because setae of p -series appearing ( Fig. 8b ), and remaining in deutonymph and tritonymph ( Figs. 9b , 11–13a, 13d , 14 ). In all nymphs, most gastronotal setae long and barbed, except for medium sized and slightly barbed c 1 , p 2 and p 3 , p 1 shorter than other posterior setae. Setal pair da long, barbed, inserted close to each other, with apical end adapted for bearing exuvial scalps of previous instars. Mutual distance between setal pair c 1 about two times longer than between setal pair da , mutual distance between setal pair dm and dp about 5.5, and 5 times longer than between setal pair da , respectively. Setae c 3 , lm , lp , h -series and p 1 in marginal position on gastronotum. Protonymph, deutonymph and tritonymph carry exuvial scalps of previous instars ( Fig. 9b ), using long setal pair da . In protonymph, one pair of setae appearing on genital valves, and two pairs added in deutonymph and tritonymph each ( Figs. 8b , 11 ); all short and smooth. In deutonymph, one pair of aggenital setae and three pairs of adanal setae appearing, and present in tritonymph; all short and smooth. Anal valves of protonymph (segment AD) and deutonymph (segment AN) glabrous, but in tritonymph two pairs of setae present, all short and smooth. In all nymphs, humeral organ placed as in larva. In deutonymph and tritonymph, cupule iad lateral to anterior part of anal valves, cupules ia , im , ip , ih and ips not observed, opisthonotal gland opening as in larva ( Figs. 9b , 11 ). In deutonymph and tritonymph, leg setae d on femora I and II, l” on genua I and II and tibia I, and l’ on genu III clearly thicker than other setae ( Figs. 13a, 13b , 14 , 15 ). FIGURE 11. Oribatella similesuperbula , ventral part of hysterosoma, legs partially drawn, scale bar 50 μm. (a) Deutonymph, (b) tritonymph. FIGURE 12. Oribatella similesuperbula , deutonymph, dorsal aspect, legs partially drawn, scale bar 50 μm. FIGURE 13. Oribatella similesuperbula , transmitted-light photomicrographs, deutonymph, dorsal aspect. (a) Whole body, (b) anterior part, dorsal aspect, (c) medial part, (d) posterior part. Scale bars 50 μm. Summary of ontogenetic transformations In all instars of O . similesuperbula , the prodorsal setae are long, and le is as thick as ro and in , but in the adult le is clearly thicker than other setae. In all juveniles, seta ex is long, whereas in the adult it is short. The opening of bothridium is rounded in all instars, but in the adult it gains small scales. In the larva, the bothridial seta is narrowly clavate, in the nymphs it is setiform, whereas in the adult it is fusiform. The larva has 12 pairs of gastronotal setae, including minute h 3 , the nymphs have 15 pairs ( p -series appear), whereas the notogaster of adult loses setae c 2 , c 3 and of d -series, such that 10 pairs of setae remain. The formula of gastronotal setae is 12-15-15-15-10 (from larva to adult), and formulae of epimeral setae are 3-1-2 (larva, including scaliform 1c ), 3-1-3-1 (protonymph), 3-1-3- 2 (deutonymph) and 3-1-3-3 (tritonymph and adult). Formula of genital setae is 1-3-5-6 (protonymph to adult), aggenital setae is 1-1-1 (deutonymph to adult), and formula of segments PS−AN is 03333-0333-022. The ontogeny of leg setae and solenidia of O . similesuperbula is given in Table 2 . Distribution, ecology, and biology
Oribatella similesuperbula has Central European distribution ( Subías (2004 , update 2021). It was recorded from Germany (Weigmann 2006) and Romania ( Călugăr & Ivan 2013 ). Weigmann (2001 , 2006) noted the preference of O . similesuperbula to moss and decaying wood. In Romania this species was first recorded from a forest plantation, with dominant Robinia pseudoacacia L. ( Călugăr & Ivan 2013 ) , but was more abundant in bryophyte covering calcareous cliffs, which suggests its preference to such a habitat. In this study, O . similesuperbula was distinctly more abundant in Horia (42 individuals per 500 cm 3 ) than in Dâmbovicioara Gorges (2 individuals per 500 cm 3 ). In both habitats, the adults dominated the juveniles, which constituted 49% and 20% of sample populations, respectively. In the samples in Horia, the stage structure of this species was the following: 25 larvae , 45 protonymphs, 33 deutonymphs, one tritonymph and 107 adults . Among 40 investigated adults, the sex ratio (females: males) of O . similesuperbula was 1:0.25, and 16% of females were gravid. The gravid females carried 1–2 relatively large eggs, each 152 x 104, which constitutes about 41% of the total body length of females. In all habitats, females of O. similesuperbula were significantly longer and wider than males. TABLE 2. Ontogeny of leg setation (Roman letters) and solenidia (Greek letters) of Oribatella similesuperbula .
Leg Trochanter Femur Genu Tibia Tarsus
Leg I
Larva d , bv” ( l ), σ ( l ), v’ , φ1 ( ft ), ( tc ), ( p ), ( u ), ( a ), s , ( pv ), ( pl ) ε, ω 1
Protonymph ω2
Deutonymph φ2
Tritonymph v’ ( l ) v’ v” ( it )
Adult v” l” , v’
Leg II
Larva d , bv” ( l ), σ l’, v’ , φ ( ft ), ( tc ), ( p ), ( u ), ( a ), s , ( pv ), ω 1
Protonymph ω2
Deutonymph l”
Tritonymph v ( l ) v’ v” ( it )
Adult v”
Leg III
Larva d , ev’ l’, σ v’ , φ ( ft ), ( tc ), ( p ), ( u ), ( a ), s , ( pv )
Protonymph
Deutonymph l’ l’
Tritonymph v’ v’’ ( it )
Adult
Leg IV
Protonymph ft’’ , ( p ), ( u ), ( pv )
Deutonymph d, ev’ d , l’ v’ , φ ( tc ), ( a ), s
Tritonymph v’ l’ , v”
Adult
Note: structures are indicated where they are first added and are present through the rest of ontogeny; pairs of setae in parentheses, dash indicates no additions.