Acari: Oribatida: Achipteriidae - BioOne

Systematic & Applied Acarology 22(2): 224–240 (2017) http://doi.org/10.11158/saa.22.2.7 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:38ED74F1-5B7A-4F61-841D-9C04B69F4BD6

Morphological ontogeny of Cerachipteria iturrondobeitiai sp. nov. (Acari: Oribatida: Achipteriidae) from northern Spain, with comments on Cerachipteria Grandjean ANNA SENICZAKa & STANISŁAW SENICZAKb a

Department of Ecology, UTP University of Science and Technology, Bydgoszcz, Poland Department of Evolutionary Biology, Faculty of Natural Sciences, Kazimierz Wielki University, Bydgoszcz, Poland Corresponding author; E-mail: [email protected]

b

Abstract The morphological ontogeny of Cerachipteria iturrondobeitiai sp. nov. from northern Spain is described and illustrated. The adult of this species has wide, long prodorsal lamellae, protruding beyond the rostrum, as in other genera of Achipteriidae. However, the lamellae are separated as in other species of Cerachipteria Grandjean, 1935. In C. iturrondobeitiai, the translamella can be present, incomplete or absent, whereas most congeners lacks the translamella, except for C. jugata Mihelčič, 1956, in which it is well-formed. The ovipositor of C. iturrondobeitiai is unusual because it bears seven pairs of coronal setae, instead of three pairs in other species of Achipteriidae.The adult of C. iturrondobeitiai is similar to C. digita Grandjean, 1935, but differs from it mainly by the location of porose areas A2 and A3 and seta lm on the notogaster and the shape of the bothridial seta. The juveniles of C. iturrondobeitiai are plicate, with most prodorsal setae minute, as in other species of Achipteriidae, and the larva lacks seta h3. In C. iturrondobeitiai, the gastronotal setae of c-series are also minute, whereas in other species these setae are longer and thicker, or at least c1. The leg claws of this species are smooth. The ontogeny of C. iturrondobeitiai is compared with some other species of Achipteriidae, and the adult is compared with congeners. Keywords: oribatid mites, soil mites, juveniles, leg setation, stage structure

Introduction Cerachipteria Grandjean, 1935 is a small and well defined genus of Achipteriidae, in which Subías (2016) included four species and one subspecies. He listed six additional genera in this family: Achipteria Berlese, 1885; Anachipteria Grandjean, 1932; Campachipteria Aoki, 1995; Dentachipteria Nevin, 1974; Cubachipteria Balogh & Mahunka, 1979 and Parachipteria Hammen, 1952. However, Seniczak & Seniczak (2016) proposed Cubachipteria and Parachipteria as synonyms of Achipteria, so that five genera remain. Cerachipteria was proposed by Grandjean (1935) on the base of C. digita Grandjean, 1935, with the main diagnostic characters monodactyle legs and separated prodorsal lamellae, with the translamella present or absent (Grandjean 1935; Mihelčič 1956). Some species of Achipteria and Campachipteria are also monodactyle, but these genera have the lamellae fused in the middle part (Seniczak & Seniczak 2016; Aoki 1995), as in most genera of Achipteriidae. Separated lamellae occur in Anachipteria geminus Lindo, Clayton & Behan-Pelletier, 2008 and Dentachipteria, but both taxa are heterotridactylous. Moreover, A. geminus lacks an anterior pteromorphic projection, and Dentachipteria has the prodorsum strongly curved ventrally.

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Cerachipteria has 10 pairs of notogastral setae, including c2, as in other genera of Achipteriidae, and four pairs of porose areas, but the latter character is considered species specific by Seniczak & Seniczak (2016). The juveniles of Cerachipteria are unknown (Norton & Ermilov 2014), but according to Weigmann (2006), Norton & Behan-Pelletier (2009) and Seniczak & Seniczak (2016) all juveniles of Achipteriidae are plicate. The aim of this paper is to describe C. iturrondobeitiai sp. nov. and its morphological ontogeny, compare it with that of some species of Achipteria and compare the morphology of the adult with congeners.

Material, methods and terminology The juveniles and adults of C. iturrondobeitiai sp. nov. originate from an abundant population of this species (1973 individuals), which was collected by St. S. on 25 June 2011 in spruce litter in spruce forest (Picea abies (L.) H. Karst), near Saldropo (Gorbea Natural Park, Basque country, northern Spain, 43°03'02"N, 02°42'81"E, 1986 m a. s. l.). The morphological ontogeny of C. iturrondobeitiai is similar to that of Achipteria gigantea Seniczak & Seniczak, 2016, except for the absence of seta h3 in the larva, which is present in A. gigantea. Therefore, the illustrations of instars are limited to the body regions of mites that show substantial differences, including the dorsal and lateral aspects of the larva, tritonymph and adult, and ventral regions of all instars. Some parts of the body and legs of the larva, tritonymph and adult are also illustrated. Illustrations were prepared from individuals mounted temporarily on slides in lactic acid. We measured the total length of the body (excluding chelicerae if extended) and maximum body width. All measurements are given in μm. In the text and figures we used the following abbreviations: rostral (ro), lamellar (le), interlamellar (in) and exobothridial (ex) setae, lamella (La), translamella (Tr), bothridium (bo), bothridial seta (bs), pteromorph (Ptm), pedotectum (Pd), tutorium (Tut), notogastral or gastronotal setae (c-, d-, l-, h-, pseries), lyrifissures or cupules (ia, im, ip, ih, ips, iad), porose areas (Aa, A1, A2, A3), opisthonotal gland opening (gla), subcapitular setae (a, m, h), palp setae (sup, inf, l, d, cm, acm, it, vt, ul, su) and solenidion ω, epimeral setae (1a–c, 2a, 3a–c, 4a–c), genital setae (g), setae k, τ and ψ on the ovipositor, adanal and anal setae (ad-, an-series), aggenital seta (ag), leg solenidia (σ, φ, ω), famulus (ε) and setae (bv, ev, d, l, ft, tc, it, p, u, a, s, pv, pl, v). Terminology used follows that of Grandjean (1932, 1935, 1936, 1953) and Norton & Behan-Pelletier (2009). The species nomenclature partly follows Subías (2004, 2016), Weigmann (2006) and Norton & Ermilov (2014).

Description of Cerachipteria iturrondobeitiai sp. nov. and its morphological ontogeny Diagnosis Adult medium sized (248–429), brown, with long and pointed anterior pteromorphic projection. Lamella long, wide, separated, protruding beyond rostrum in dorsal and ventral views, and distally pointed, translamella present, incomplete or absent. Seta in long, smooth and inserted on inner part of base of lamella. Notogastral setae (10 pairs, including c2) of medium size and smooth, c2 and la longer than other dorsal setae. Genua I and II without anteroventral projection, seta v'' on femur II distinctly longer than bv'', seta l' on femur III absent, seta ev' on femur IV short. Genu IV clearly shorter than tibia IV. Tarsi monodactyle, claws smooth. Juveniles stocky, light brown, with plicate cuticle and granular cerotegument. Most prodorsal and gastronotal setae of c-series of juveniles minute, whereas other setae on dorsal side of

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gastronotum short or medium sized and barbed, seta h1 in larva absent. Leg segments stocky, most setae smooth or with short barbs, seta d at solenidion φ on tibia IV present, leg claws smooth. Description of adult Body of medium size, with characters of Cerachipteria (Grandjean 1935), females usually slightly larger than males. Measurements. Length of holotype (female 449), mean length of 50 females—443.3 (range 436–462) and 50 males—413.4 (range 390–429); notogaster width of holotype (307), mean width of 50 females—298.2 (range 251–314) and 50 males—261.4 (range 248–287). Gravid females (52% females) thicker (notogaster width of 300–314) than engraved (notogaster width of 251–307). Integument. Body color brown, main body microporose with light dots (Fig. 1c).

FIGURES 1–2. Cerachipteria iturrondobeitiai n. sp. female. 1. (a) Dorsal aspect, legs partially drawn, scale bar 50 μm, (b) shape of distal and medial parts of seta in, (c) fragment of gastronotum with seta h3 (b, c enlarged). 2. (a), (b) Prodorsum and anterior part of notogaster of different individuals, dorsal aspect, legs partially drawn, scale bar 50 μm.

Prodorsum. Rostrum rounded. Lamella (La) relatively long (91–95) and wide (37–41), with longitudinal striae, protruding beyond rostrum in dorsal and ventral views (Fig. 1a), curved ventrally in distal part and pointed; translamella (length 18–20) present (well-formed or thin as line), incomplete, or absent (Figs 1a, 2a, b). Seta ro of medium size (55–58), curved inwards, barbed unilaterally and positioned laterally. Seta le (31–35), straight and smooth, inserted on inner and ventral side of lamellae, in 2/3 of lamella from its base. Seta in long, straight, with short setulae in medial and distal parts (Fig. 1b), basally covered or uncovered by anterior part of notogaster (Figs 226

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1a, 2a, b). Bothridium (bo) with two small lateral scales, inner scale longer than outer one; bothridial seta (bs) short (Table 1), clavate, with smooth stalk, and narrow head with short barbs. Seta ex short (6) and covered by pteromorph in lateral aspect.

FIGURE 3. Cerachipteria iturrondobeitiai sp. nov. (a) Female with extended ovipositor, lateral aspect, right side, legs partially drawn, scale bar 50 μm, (b) tutorium, (c) anterior part other tutorium (b, c enlarged).

Notogaster. Slightly longer (339–401) than wide (248–314), and convex dorsomedially, pteromorph with long and pointed anterior projection, strongly curved ventrally (Figs 1a, 2a, b, 3a), anteroventral edge of pteromorph pointed (Fig. 4a). Notogastral setae (10 pairs, including c2) smooth (Figs 1a, 3a), c2 and la longer (34–36) than other setae (19–24, Table 1). Lenticulus not observed. Porose areas usually oval (four pairs), but Aa may be of different shape (Figs 1a, 2a, b), Aa larger than other porose areas, lyrifissure im posterolateral to seta lm, opisthonotal gland opening (gla) and other lyrifissures in normal position. Gnathosoma. Subcapitular setae h and m similar in size and longer (23) than a (12), all smooth (Figs 4a). Chelicera (length 96–99) with seta cha longer (34) than chb (18); all barbed (Fig. 5a). Palp (length 55–58) with smooth setae (Fig. 5b), inf longer than sup; formula of palp setae (trochanter to tarsus + solenidion ω): 0-2-1-3-9(1). Epimeral and lateral regions. Central epimeral setae (1a, 2a, 3a) short (5), other setae longer (18–20), 1c inserted on lateral part of pedotectum 1 (Pd1), 4c longest (39) and thickest, and inserted on apophysis directed anteriorly (Figs 4a, b); all smooth. Tutorium triangular, relatively long (145) and wide in distal part, with two or more dens (Figs 3b, c). Pedotectum I (Pd1) large, spade-like (110–112 x 45–48), with longitudinal striae (Fig. 3a), pedotectum II small. Genal tooth (gt) triangular, long (42–45), curved ventrally and pointed, custodium (cus) shorter (16), pointed, discidium (dis) well-developed, triangular, circumpedal carina absent. Anogenital region. Genital setae (six pairs) and aggenital (one pair) slightly longer (15–18) than adanal (three pairs) and anal setae (two pairs, 13–15, Fig. 4a). Ovipositor with seven pairs of short coronal setae k (10), setae τ1, τ2 and τ3 (20, 7 and 7, respectively, τ1 and τ2 unpaired) inserted on two anterior lobes and pairs ψ1 and ψ2 (18 and 4, respectively) positioned on posterior lobe (Fig. 3). Lyrifissure iad short, located close to anterior end of anal opening. 2017


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FIGURES 4–5. Cerachipteria iturrondobeitiai sp. nov., adult. 4. (a) Ventral aspect, legs partially drawn, scale bar 50 μm, (b) location of some epimeral setae (enlarged). 5. Mouthparts of adult, antiaxial aspect, right side, scale bar 20 μm. (a) Chelicera, (b) palp.

Legs. Trochanters II and IV and all femora flattened and with ventral carina; porose areas present on inner side of these segments. Genua I and II without anteroventral projection, genu IV clearly shorter than tibia IV, but longer than genu III; other genua shorter than respective tibiae (Fig. 6). Solenidion ω1 on tarsus I longer and thicker than ω2 and strongly curved outwards; seta ft" and famulus ε short (Fig. 6a). Seta s on tibia II thicker than other setae and with long barbs on outer side (Fig. 6b). Tarsi monodactyle, claws smooth. 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—1-2-23(1)-12. Description of juvenile stages Larva stocky, light brown, cuticle plicate with granular cerotegument (Fig. 7). Prodorsum subtriangular, prodorsal setae minute, except for short ro; all smooth. Mutual distance between setae of pairs le and in more than twice longer than that of pair ro. Pair le inserted approximately midway between pairs ro and in. Opening of bothridium oval, with outer curved addition, bothridial seta clavate, with narrow, barbed head. Ridge present anteromedial to bothridium. Gastronotum of larva (Fig. 7) with 11 pairs of setae, including h2 inserted laterally to medium part of anal valves (Figs 8a, 9a). Setae of c-series minute, other setae short or medium sized (Table 1) and thickly barbed (Fig. 7), except for short and smooth h1 and h2 (Fig. 8a). Length of setae of dand l-series increasing from anterior to posterior. Cupules not observed in plicate cuticle, except for 228


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ih located lateral to anterior part of anal opening. Gland opening gla anterolateral to h2 (Fig. 9A). Paraproctal valves (segment PS) glabrous. Leg segments stocky, trochanter III and all femora flattened, cuticle of most segment reticulate, well observed on femora (Fig. 10a). Some leg setae (d on all femora, l-series on genua and tibiae) slightly thicker than other setae, most tarsal setae thin, with short barbs or smooth, except for one seta thicker and with long barbs on ventral side. Solenidion φ1 on tibia I on large apophysis, solenidion ω1 on tarsus I thick and curved outwards, seta ft" and famulus ε short. Leg claws smooth (Figs 10a, b, c). TABLE 1. Measurements of some morphological characters of juvenile stages of Cerachipteria iturrondobeitiai sp. nov., (mean measurements of 10 individuals per instar in μm); Nd—not developed. Morphological characters

Larva

Protonymph

Deutonymph

Tritonymph

Adult

Body length

221

271

358

442

455

Body width

120

147

191

254

281

Lengh of prodorsum

78

74

89

101

60

Length of:seta bs

44

52

61

74

52

seta ro

16

10

13

14

56

seta le

2

2

2

3

32

seta in

2

2

2

3

105

seta ex

1

1

1

2

6

seta c1

2

2

3

5

Lost

seta c2

2

2

3

4

35

seta c3

3

3

3

4

Lost

seta da

8

6

8

10

Lost

seta dp

12

15

21

25

Lost

seta la

4

3

3

4

34

seta lp

12

13

15

18

24

seta h1

3

11

17

25

25

seta h2

2

2

4

6

20

seta h3

Nd

13

19

20

19

seta p1

Nd

2

2

3

19

genital opening

Nd

12

30

34

49

anal opening

52

63

75

89

76

Prodorsum and prodorsal setae and bothridial seta of protonymph as in larva, but ridge between bothridium and seta le well-developed. Gastronotum with 15 pairs of setae due to appearance of seta h3 and p-series (Fig. 8b), retained by subsequent nymphs (Figs 11a, b); all short and smooth, except for barbed h3. Opisthonotal gland opening gla lateral to seta lp (Fig. 9b). In protonymph one pair of genital setae appears on genital valves, and two pairs are added in deutonymph and two pairs in tritonymph (Figs 11a, b). In deutonymph one pair of aggenital setae and three pairs of adanal setae appear, retained by subsequent instars; all short and smooth. In tritonymph two pairs of short and smooth anal setae appear on anal valves (Fig. 11b). Cupules not observed in plicate cuticle, except for ips in deutonymph and iad in tritonymph, located lateral to anterior part of anal opening. Length of setae in tritonymph increasing from anterior to posterior (Fig. 12a). Leg segments stocky, especially femora and distant part of tibiae and basal part of tarsi I and II. Trochanters III and IV and

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all femora flattened, cuticle of most leg segments reticulate, well observed on femora (Fig. 13a). Most setae on femora, genua and tibiae slightly thicker and barbed (d on all femora, l-series on genua and tibiae, ev' on femora III and IV and v' on tibiae III and IV), tarsal setae thin, with short barbs or smooth, except for one seta on ventral side of legs I and II (Figs 13a, b). Solenidion φ on tibia I on large apophysis. Solenidion ω1 thick, directed anteriorly and strongly curved outwards, seta ft" and famulus ε short (Fig. 13a). Tibia IV with seta d at solenidion φ (Fig. 13d), other leg solenidia without seta d. All genua shorter than tibiae, leg claws smooth.

FIGURE 6. Cerachipteria iturrondobeitiai n. sp. legs of adult: femur, genu, tibia and tarsus shown from right side, setae on the opposite side which are not illustrated indicated in brackets in the legend, scale bar 50 μm. (a) Leg I, genu (l'), tibia (l'), tarsus (pl'); (b) leg II, genu (l'), tarsus (pv'); (c) leg III; (D) leg IV.

Summary of ontogenetic transformations Ontogeny of C. iturrondobeitiai sp. nov. is generally similar to that of A. gigantea studied by Seniczak & Seniczak (2016), except for seta h3 in the larva, which is absent in the former species and present in the latter species, and the shape of some setae slightly differs. In all juvenile instars of C. iturrondobeitiai the prodorsal setae are minute, except for short ro, and gastronotal setae of c-series

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are short, whereas other setae on the dorsal part of gastronotum are short or medium sized and barbed. In the adult seta in becomes long, ro and le are of medium size, and ex remains short. The bothridial seta is clavate in all instars, with narrow, long head, but in the juveniles barbs are longer than in the adult, and in the larva the head is slightly thicker than in the nymphs. The larva has 11 pairs of gastronotal setae, including h2, and the nymphs have 15 pairs due to the appearance in the protonymph of setae h3 and p-series, which are retained by other nymphs. The notogaster of adult loses setae c1, c3 and d-series, so that 10 pairs of gastronotal setae remain. The formula of gastronotal setae in C. iturrondobeitiai is 11-15-15-15-10 (from larva to adult), whereas the formulae of epimeral, genital and aggenital setae and that of segments PSAN are the same as in A. gigantea. The number of setae and solenidia on legs increases during the ontogeny as in A. gigantea (Seniczak & Seniczak 2016), except for seta l' on femora III, which is lacking in C. iturrondobeitiai. Tibia IV of deutonymph and tritonymph has seta d at solenidion φ, which is lost in the adult. In all instars the leg claws are smooth.

FIGURES 7–8. Cerachipteria iturrondobeitiai sp. nov. 7. Larva, dorsal aspect, legs partially drawn, scale bar 50 μm. 8. (a) Anal region of larva, (b) anogenital region of protonymph, legs partially drawn, scale bar 50 μm.

Ecology and biology We found C. iturrondobeitiai sp. nov. common and abundant in Gorbea Natural Park (Basque country, northern Spain), where it occurs in moss and coniferous and deciduous litter, together with Achipteria coleoptrata (Linnaeus, 1758), A. magna (Sellnick, 1928) and Pseudachipteria floresiana Pérez-Íñigo, 1992. However, the proportions of C. iturrondobeitiai differ in particular samples and habitats. For example, in June in spruce litter (spruce forest near Saldropo), C. iturrondobeitiai 2017


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dominated, achieving a density of 1973 individuals per 1 dm3, while A. coleptrata was not abundant (six larvae and one deutonymph). In this microhabitat, the juveniles of C. iturrondobeitiai highly dominated (88% of all individuals), and the stage structure of this species was the following: 900 larvae, 524 protonymphs, 111 deutonymphs, 206 tritonymphs and 232 adults. In moss on beech rut on lower altitude (43°03'19"N, 02°42'81"E, 632 m a. s. l.), P. floresiana dominated (4378 individuals per 1 dm3), and C. iturrondobeitiai and A. magna were not abundant (25 and 171 individuals per 1 dm3, respectively). In this sample, the juveniles C. iturrondobeitiai made 92% of all individuals of this species, and the stage structure was: 125 larvae, 29 protonymphs, two deutonymphs, one tritonymph and 14 adults. The sex ratio of C. iturrondobeitiai (females to males) was 1:1.1 and 52% females were gravid, carrying two or rarely one large egg (188 x 101).

FIGURE 9. Cerachipteria iturrondobeitiai sp. nov., lateral aspect, right side, legs partially drawn, scale bars 50 μm. (a) Larva, (b) tritonymph.

Type deposition The holotype and 6 paratypes (in 70% ethanol) of C. iturrondobeitiai sp. nov. are deposited in the University Museum of Bergen (Norway) and 6 paratypes are deposited in the Department of Ecology, UTP University of Science and Technology, Bydgoszcz, Poland. Etymology The species is named in honour of prof. Juan Carlos Iturrondobeitia (Department of Zoology and Cellular Animal Biology, University of the Basque Country, Leioa, Spain) for his wide input in our knowledge of the oribatid mites.

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FIGURE 10. Cerachipteria iturrondobeitiai sp. nov., legs of larva: femur, genu, tibia and tarsus shown from right side, setae on the opposite side which are not illustrated indicated in brackets in the legend, scale bar 20 μm. (A) Leg I, tarsus (pl', pv'); (B) leg II, tarsus (pv'); (C) leg III.

Comparison of the morphology of Cerachipteria iturrondobeitiai with some species of Achipteria We compare the morphological ontogeny of C. iturrondobeitiai sp. nov. with that of A. gigantea and A. bella (Sellnick, 1928) investigated by Seniczak & Seniczak (2016) and Seniczak & Seniczak (2007, as Parachipteria bella), respectively. Since the ontogeny of these species are relatively well known, the differences between Cerachipteria and Achipteria can easily be demonstrated. We also compare the morphology of the adult of C. iturrondobeitiai with congeners, such as C. digita Grandjean, 1935, C. jugata Mihelčič, 1956; C. franzi Willmann, 1953; C. minuscula (Berlese, 1902) and C. pyrenaica Pérez-Íñigo jr., 1990. The adults of C. iturrondobeitiai, A. gigantea and A. bella differ from one another by the following important morphological characters: (1) Shape of lamellar complex on the prodorsum: in C. iturrondobeitiai the lamellae are separated and cover about half of the prodorsum in dorsal aspect, whereas in other species they are fused medially and cover almost the whole prodorsum. In C. iturrondobeitiai the translamella can be of different shape or absent. (2) Location of epimeral seta 1c: in C. iturrondobeitiai this seta is located on lateral part of pedotectum I, whereas in other species it is inserted on ventral part of this pedotectum. (3) Shape and location of epimeral seta 4c: in C. iturrondobeitiai this seta is clearly longer and thicker than other epimeral setae and is located on apophysis, whereas in other species this seta is thin and the apophysis is absent. (4) Legth of genu IV: in C. iturrondobeitiai this segment is shorter than tibia IV, whereas in other species it is longer than tibia IV. (5) Presence of seta l' on femur III: in C. iturrondobeitiai this seta is absent, whereas in other species it is present. (6) Length of seta ev' on femur IV: in C. iturrondobeitiai this seta is short, whereas in other species it is long. 2017


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The juveniles of C. iturrondobeitiai, A. gigantea and A. bella are strikingly similar, but differ from one another amongst others by the body size, presence of seta h3 in the larva, pattern of prodorsal cuticle, shape of most gastronotal setae and leg claws and characters with specific value (Table 2). The ontogeny of leg setae of C. iturrondobeitiai is similar to that of A. gigantea, but the shape of some leg setae differ. In the latter species some setae on femora, genua and tibiae (d on all femora, l-series on genua and tibia, bv'' on femur II, ev' on femur III) are thicker than in the former species. Moreover, the femora of nymphs of A. gigantea have relatively denser reticulation of cuticle than those of C. iturrondobeitiai. The adult of C. iturrondobeitiai differs from those of A. gigantea and A. bella by more morphological characters than their juveniles, and the most interesting seems to be the unique location of epimeral setae 1c and 4c, relatively short genu IV, presence of seta l' on femur III and length of seta ev' on femur IV. TABLE 2. Selected morphological characters of Cerachipteria iturrondobeitiai sp. nov., Achipteria gigantea and A. bella. Morphological characters

C. iturrondobeitiai

A. giganteaa

A. bellab

Formula of gastronotal setae

11-15-15-15-10

12-15-15-15-10

11-15-15-15-10

Body size μm

390–462

913–1053

390–450

Shape of lamellae

Separated

Joined

Joined

Shape of sensillus

Clavate

Clavate

Fusiform

Location of seta in

On lower part of La

1/3 of length of La

On lower part of La

Adult

Octotaxic system

Porose areas

Sacules

Porose areas

Length of seta c2c

Long

Medium size

Long

d

Long

Medium size

Short

Length of seta lp

Location of seta 1c

Lateral part of Pd1

Ventral part of Pd1

Ventral part of Pd1

Seta 4c located on

Apophysis

Epimere IV

Epimere IV

Apophysis on genua I and II

Absent

Present

Absent

Seta ev' on femur IV

Short

Long

Long

Length of seta bv'' on femur II

Shorter than v''

As long as v''

Shorter than v''

Presence of seta l' on femur III

Absent

Present

Present

Number of leg claws

1

3

3

Shape of leg claws

Smooth

Serrate

Smooth

Pattern of prodorsal cuticle

Indistinct

Indistinct

Reticulate

Shape of leg claws

Smooth

Serrate

Serrate

Juveniles

Length of seta c1

As short as c3

Longer than c3

Longer than c3

Length of seta dp

Longer than da

As short as da

As short as da

Presence of seta h3

Absent

Present

Absent

Shape of gastronotal setae

Thickly barbed

Thickly barbed

Barbed

Larva

a

—according to Seniczak & Seniczak (2016); b— according to Seniczak & Seniczak (2007) and completed by the authors; c— medium size means about half of the distance between insertions of setae c2 and la, long means nearly as long as this distance in dorsal aspect; d—medium size means about half of the distance between insertions of setae lp and h3, long means nearly as long as this distance in dorsal aspect.

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FIGURES 11–12. Cerachipteria iturrondobeitiai sp. nov. 11. Anogenital region, legs partially drawn, scale bars 50 μm. (a) Deutonymph, (b) tritonymph. 12. Tritonymph. (a) Dorsal aspect, legs partially drawn, scale bar 50 μm, (b) opening of bothridium and seta ex. TABLE 3. Selected morphological characters of adults of Cerachipteria species. Body length in μm

Species C. digita Grandjean, 1935

395–400

Translamella Absent

Length of seta c 2a

lpb

No

No

C. franzi Willmann, 1953

525

Absent

No

No

C. iturrondobeitiai sp. nov.

390–462

Present/absent

Yes

No

C. jugata Mihelčič, 1956

560

Present

No

No

C. minuscula (Berlese, 1902)

350

?

?

?

C. pyrenaica Pérez-Íñigo jr., 1990

418–445

Absent

Yes

Yes

a

— longer than the distance between insertions of setae c2 and la in dorsal aspect; b —longer than the distance between insertions of setae lp and h3 in dorsal aspect.

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FIGURE 13. Cerachipteria iturrondobeitiai sp. nov., legs of tritonymph: femur, genu, tibia and tarsus shown from right side, setae on the opposite side which are not illustrated indicated in brackets in the legend, scale bar 50 μm. (a) Leg I, femur (l'), genu (l'), tibia (l'), tarsus (pl'); (b) leg II, femur (l'), genu (l'); (c) leg III; (d) leg IV.

The comparison of the adult of C. iturrondobeitiai with congeners shows that the shape of the translamella varies in Cerachipteria, which is unique in this genus (Table 3). Generally, most Cerachipteria species are poorly described, except for C. digita described by Grandjean (1935). The adult of C. iturrondobeitiai is similar to that of C. digita, but differs from it by the following morphological characters: (1) Location of porose area A2: in C. digita A2 is located medially to seta h2, whereas in C. iturrondobeitiai A2 is placed anterolaterally to this seta. (2) Location of porose area A3: in C. digita A3 is located anteromedially to seta h2, whereas in C. iturrondobeitiai A2 is placed laterally to this seta. (3) Location of notogastral seta lm; in C. digita this seta is inserted more medially than in C. iturrondobeitiai, e. g. the distance between seta lm and porose area A1 is clearly longer than between A1 and seta h3, whereas in C. iturrondobeitiai A1 is placed aproximatelly midway between these setae. (4) Length of notogastral setae: in C. digita these setae are shorter than in C. iturrondobeitiai, especially c2 and la. 236


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(5) Shape of bothridial seta: in C. digita the head of bothridial seta is shorter than in C. iturrondobeitiai. The shape of the translamella and porose areas Aa varies in C. iturrondobeitiai, but the location of other porose areas and the length of setae are relatively stable and differ from those of C. digita.

Discussion Cerachipteria is a well-defined genus of Achipteriidae (Grandjean 1935), which have monodactyle legs and separate prodorsal lamellae in the adults. There are other potential morphological characters that can be of diagnostic value for this genus, but more investigation is needed. For example, Grandjean (1935) showed the unusual position of two epimeral setae in C. digita, 1c inserted on the lateral part of pedotectum I and 4c long and thick and inserted on apophysis, similar to C. iturrondobeitiai. In Achipteria and Anachipteria seta 4c has no apophysis, and in the former genus seta 1c is inserted on the ventral part of pedotectum I, whereas in the latter genus this seta is inserted either on the lateral part of pedotectum I (Root et al. 2008) or on the ventral part (Ermilov et al. 2011). The ovipositor of C. iturrondobeitiai is unique because neotrichy of coronal setae k occurs; seven pairs of these setae are present, instead of three pairs in other species of Achipteriidae (Abd el Hamid 1964; Lindo et al. 2008; Ermilov et al. 2011; Ermilov & Anichkin 2014). Neotrichy of coronal setae is probably rare in Brachypylina (Circumdehiscentiae) and was observed in Hungarobelba pyrenaica Miko & Travé, 1996, in which the ovipositor has over 30 setae k. Therefore, more investigations are required on the morphology of the ovipositor in Cerachipteria species to know if the neotrichy of coronal setae concerns other species. The shape of some leg segments and setae may also be diagnostic. For example, in C. iturrondobeitiai genu IV is distinctly shorter than tibia IV, whereas in Anachipteria this segment is either shorter (Abd el Hamid 1964; Root et al. 2008; Ermilov et al. 2014) or longer than tibia IV (Lindo et al. 2008). In Achipteria genu IV is approximately as long or longer than tibia IV (Hirauchi & Aoki 1997; Palacios-Vargas & Socarrás 2001; Maruyama 2003; Ermilov et al. 2011; Bayartogtokh & Ryabinin 2012; Seniczak & Seniczak 2016). In C. iturrondobeitiai genua I and II lack anteroventral projection, in Anachipteria this projection is present (Abd el Hamid 1964; Root et al. 2008), whereas in Achipteria it is present or absent (Hirauchi & Aoki 1997; Palacios-Vargas & Socarrás 2001; Maruyama 2003; Ermilov et al. 2011; Maruyama et al. 2016). In C. iturrondobeitiai seta v'' on femur II is distinctly longer than bv'', as in Anachipteria (Root et al. 2008) and some species of Achipteria (Maruyama et al. 2016), whereas in most species of the latter genus both setae are long (Hirauchi & Aoki 1997; Maruyama 2003; Ermilov et al. 2011; Seniczak & Seniczak 2016). In C. iturrondobeitiai, seta l' on femur III is absent, whereas in Achipteria and Anachipteria this seta may be present (Lindo et al. 2008; Ermilov et al. 2011; Seniczak & Seniczak 2016) or absent (Maruyama 2003; Root et al. 2008; Bayartogtokh & Ryabinin 2012; Maruyama et al. 2016). In C. iturrondobeitiai seta ev' on femur IV is short, whereas in Achipteria and Anachipteria it can be short or long. In most species of Cerachipteria the shape of leg segments and setae is unknown, and therefore more investigations are required. In contrast, the deutonymphs and tritonymphs of all genera have seta d at solenidion φ on tibia IV, but this seta is lost in the adult and only solenidion φ remains. In Phenopelopidae, which juveniles also have plicate cuticle, the presence of seta d or solenidion φ on tibia IV has generic value (Seniczak et al. 2014, 2015a, c). In C. iturrondobeitiai the shape of the translamella varies between individuals, being distinct, thin as a line, incomplete or absent. Similar variability of the translamella is observed in Oribatula Berlese, 1895 (Seniczak et al. 2012) and some species of Sphaerozetinae (Ceratozetidae) sensu

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Shaldybina (1975), in which the presence or absence of the translamella has no diagnostic value (Seniczak et al. 2015d, 2016c). The shape of the chelicera of the adult of C. iturrondobeitiai is generally similar to that of other species of Achipteriidae, but the size depends on the body length. For example, in this species the chelicera constitutes 21.7% of the total body length, whereas in A. gigantea, which is approximately twice as long as C. iturrondobeitiai, the chelicera is relatively smaller and constitutes 19.0% of the total body length (Seniczak & Seniczak 2016). Similar dependencies between the body length and chelicera are observed in some species of Ceratozetidae (Seniczak et al., 2016a–d). The juveniles of C. iturrondobeitiai are strikingly similar to other species of Achipteriidae, but the shape of some gastronotal setae can be diagnostic. For example, in C. iturrondobeitiai all setae of c-series are minute, in A. bella and A. coleoptrata these setae are short (Seniczak 1978; Seniczak & Seniczak 2007), whereas in other species seta c1 is longer than other setae of c-series (Seniczak 1977, 1978; Seniczak & Seniczak 2007, 2016; Lindo et al. 2008). In C. iturrondobeitiai and A. willmanni van der Hammen, 1952, all setae of p-series are short, whereas in A. gigantea and A. bella seta p1 is longer than other setae of p-series (Seniczak & Seniczak 2007). Some of these species also differ by the shape of posterior gastronotal setae. Subías (2016) consider C. jugata Mihelčič, 1956 as subspecies of C. digita but this proposal is debatable because the former species is larger than the latter species. Cerachipteria jugata also differs from C. digita by having the translamella, which in the latter species is absent, but this character varies in Cerachipteria. This author included A. bella and A. willmanni in Campachipteria, which is incorrect because these species are heterotridactylous, whereas species in Campachipteria are monodactyle (Aoki 1995). Cerachipteria iturrondobeitiai commonly occurs in the Gorbea Natural Park (Basque country, northern Spain), and in spruce litter achieved a density of 1973 individuals per 1dm3 (= 98 650 individuals per 1m2 x 5cm deep), mainly due to many juveniles (88% of all individuals of this species) in June. However, the number of large eggs inside the gravid females was small (one or two eggs), which suggests that the great abundance of species, especially the larvae (47% of all individuals) in some microhabitats was possible due to activity in egg laying. Oribatella arctica is also a medium sized species (length 420–450), females carry one or two large eggs, and the maximal density of this species was also high (98,000 individuals per 1m2, Seniczak et al. 2015b). Other species of Achipteriidae are also rich in juveniles. For example, in larch litter the juveniles of A. gigantea (Seniczak & Seniczak 2016) constituted 80% of all individuals in June, whereas the medium sized Anachipteria geminus (Lindo et al. 2008) dominated in canopy lichens (coniferous temperate and montane forests, Pacific Northwest of North America), achieving the maximum density 1450 individuals /100 g dry mass of lichen. In November, the juveniles of this species were several times more abundant than the adults. These data indicate that the juveniles of Achipteriidae are important components of oribatid mite agglomerations in soil and arboreal habitats, and knowledge of their morphology is helpful in ecological studies.

Acknowledgements We thank two anonymous reviewers for thorough review of this manuscript and valuable suggestions, and Dr. Lizel Hugo-Coetzee, a section editor of Systematic & Applied Acarology for all editorial suggestions.

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