Two new species of the family Nippobodidae (Acari, Oribatida), including a description of the leg-folding process Author Fernandez, Nestor Author Theron, Pieter Author Leiva, Sergio text ZooKeys 2018 781 109 139 http://dx.doi.org/10.3897/zookeys.781.27389 journal article http://dx.doi.org/10.3897/zookeys.781.27389 1313-2970-781-109 C7FA94CF666A438A8AA7EEF1C1E0CE4F C7FA94CF666A438A8AA7EEF1C1E0CE4F Nippobodes panemorfis sp. n. Figures 1-10, 11-13, 14-22, 23-27, 28-41, 42-47, 48-53 Etymology. The specific epithet "panemorfis" is derived from "panemorfi" (πανεμορφη in Greek) meaning beautiful, due to the aesthetic features of the cuticle and setae. Diagnosis (adult female). Prodorsum. Complex shape; triangular in dorsal view with rounded central posterior zone; double hook-shaped, interlocking posterior prodorsal condyle and anterior zone humeral apophysis; rounded rostrum, with groove and large hump; deep, easily discernible round-ovoid prodorsal posterior depression; tutorium strongly curved, large lamina, connected to prodorsal wall, determining a pocket structure. Reticulate-foveate microsculpture on tutorium, pedotectum I, pedotectum II. Polyhedral bothridium situated under zone where humeral part, overlaps with anterior prodorsal zone; bothridial opening ovoid, located at bottom of U-shaped structure. Notogaster: deep, round-ovoid anterior notogastral depression present; ten pairs of setae c, la, lm, lp, h1, h2, h3, p1, p2, p3; setae c looped, dentate, sharply tipped; marginal setae h3, p3 on conspicuous promontories, followed by deep v-shaped incision; circumgastric depression completely surrounding notogaster, originating before setae la, running between setae la, h1, h2 and h3, p3, p2, p1; setae 1c, 3c, 4b situated marginally; setae 1b largest; genital opening on elevated zone; lateral genital zone locking structure with, longitudinal cuticular elevation, promontories with parallel furrow; genital plate smaller than anal plate; adanal setae ad1, ad2 inserted on elevated zone; ad3 setae smallest. Material examined. Holotype: ♀♀ Female. Label details: "Thailande. Khao Yai National Park (nord-est de Bangkok) Khao Khieo au-dessous d'Air Force check point; 1150 m; versant nord, foret assez seche ; tamisage debris . 28/XI/1985. Leg: D.H. Burckhardt et L. Loebl" . Paratypes: Two adult females, same locality and date as Holotype; deposited in Collection of NHMG; preserved in 70 % ethanol. Additional material studied using SEM: six specimens, not deposited. "Thailande. Khao Yai National Park (nord-est de Bangkok) Khao Khieo au-dessous d'Air Force check point; 1150 m; versant nord, foret assez seche ; tamisage debris . 28/XI/1985. Leg: D.H. Burckhardt et L. Loebl" Description. Measurements. SEM: 597 (542-720) x 368 (332-401) (n = 6). Light microscopy: 610 x 360 (n = 1); all specimens female. Shape. Rounded-ovoid (dorsal view) (Figure 1). Elongate oval (lateral view) (Figures 11, 23). Figures 1-10. Nippobodes panemorfis sp. n. Adult female, SEM. 1 dorsal view 2 detail of notogastral lm setae 3 detail of notogastral lm setae microsculpture, high magnification 4 frontal view 5 prodorsal lamellar (le) setae, detail 6 rostral (ro) setae, detail 7 dorsal anteroposterior view 8 prodorsal posterior condyle interlocked with humeral apophysis anterior zone 9 anterior view, humeral apophysis and interlamellar seta 10 interlamellar seta, detail. For abbreviations: see "Material and methods". Scale bars: 200 μm (1); 100 μm (4, 7); 50 μm (5, 6, 8); 25 μm (9); 17μm (2); 3 μm (10); 2 μm (3). Figures 11-13. Nippobodes panemorfis sp. n. Adult female, Optical observations. 11 lateral view 12 dorsal view 13 ventral view. Scale bars: 200 μm (11); 270 μm (12); 180 μm (13). Colour. Black, slightly shiny when observed in reflected light; rarely dark brown. Cerotegument. Not observed; small particles, similar to rest of cerotegumental layer on circumgastric depression (s.c) lateral zone (Figure 16); the layer may have existed. Integument. Microsculpture complex, varying according to body region. Smooth: prodorsum (on Figure 4 indicated by ❄); notogaster (on Figures 1, 7 indicated by ❄); on most epimeral surface (on Figures 28, 41 indicated by ❄); genital, anal, aggenital zones. Tuberculate: rostum (Figure 14, indicated by ⌘); infracapitulum near setae h (Figure 42, indicated by ⌘); epimeral zone between setae 1a, 1b (Figure 28, indicated by ⌘). Figures 14-22. Nippobodes panemorfis sp. n. Adult female, SEM observations. 14 rostrum, frontal view 15 subcapitulum, frontal view 16 notogaster, partial lateral view 17 sensillus, lateral view 18 apical zone, sensillus 19 seta c, dorsal view 20 pedotectum II, lateral view 21 lamellar seta 22 lamellar seta, detail. Scale bars: 50 μm (15, 17, 20); 20 μm (14, 16); 10 μm (18, 19); 5 μm (21); 2 μm (22). Reticulate-foveate: Tutorium (Tu) (on Figure 26 indicated by ❖); Pedotectum I (Pd I) (on Figures 26, 27 indicated by ❖), Pedotectum II (Pd II) (Figure 20, indicated by ❖). Rugose: external zone of humeral apophyse (h.ap). (Figures 1, 4, 7, 8, 9, 23, indicated by ⛭). Favulariate: bothridial zone (Figure 24a, indicated by ✱). Sulcate: zone of bothridial opening (Figure 24a, b indicated by ➡). Punctate: Discidium (dis) (Figure 30 indicated by ➧); epimeral zone surrounding setal insertion (Figures 30, 43, indicated by ➧). Figures 23-27. Nippobodes panemorfis sp. n. Adult female, SEM observations. 23 lateral view 24a bothridial zone 24b bothridial opening 25 lateral view, zone of h3, p3, setae 26 prodorsum, lateral view 27 pedotectum I, lateral view. Scale bars: 200 μm (23, 26); 30 μm (24, 27); 20 μm (25); 5 μm (24). Prodorsum. Complex shape: in dorsal view, more or less triangular with central posterior zone rounded (Figures 1, 4); lateral posterior zone with interlocking double hook-shaped posterior prodorsal condyle (p.pr.co) (indicated by ◙) and anterior zone humeral apophysis (h.ap) (Figures 8, 9); lateral view: triangular; with complex lateral posterior zone, with double hook and sigmoid lamellae (Figure 23). Rostrum round, with a conspicuous groove parallel to margin (Figure 23 indicated by ➧) large hump visible in front of groove (Figures 14). Setae: sigmoid, with small dentitions and thin parallel longitudinal ridges: Rostral (ro), interlamellar (in), lamellar (le) setae (Figures 1, 4, 5, 6, 9, 10, 21, 22): length: 123 (106-141) (n = 12); setae le, in, inserted each on large tubercle, setae ro inserted on small tubercle (Figures 1, 4, 9, 26); setae ro situated behind region of tuberculate microsculpture, marginally to depression created by Tu and lateral prodorsal wall (Figures 4, 26); le setae inserted on anterior end of lamellar zone (Figure 26), situated posteriorly and to the exterior of ro setal insertion alignment (Figures 1, 4, 7); setae in situated behind and externally to le setal insertion level (Figures 1, 4, 7), inserted near the double hook (Figures 8, 9); deep, rounded-ovoid prodorsal posterior depression (p.p.d) clearly discernible between dorsosejugal furrow (d.sj) and notogastral anterior depression (n.a.d) (Figure 1). Lamellae (Lam) clearly visible in lateral view (Figures 11, 23) (see Lateral Region). Bothridium (bo) (Figures 23, 24) situated under double hook zone (See: Lateral view). Sensillus (si) (Figures 17, 18) sickle-shaped, strongly curved, directing upward with long stalk followed by a swollen zone, plentifully scattered with small asperities and with small barbs (Figure 18 indicated by ➡); long sharp apical tip; length 152 (149-160) (n = 6). Tu well developed, sharply tipped; lacking free extremity, welded to lateral prodorsal wall, determining pocket-like structure (Figures 4, 26). Pocket structure conceals leg I when leg-folding process is activated (See: Leg-Folding Process). Interlocking double hook. The interlocking double-hook zone is complex, formed by p.pr.co and anterior zone h.ap (Figures 1, 4, 7, 8, 9, 23); where h.ap situated externally (indicated by ✱), grips on to p.pr.co (indicated by ◙) on the interior. Cuticular surface of p.pr.co smooth with some irregular depressions (Figure 8); cuticular surface of h.ap rugose externally (Figures 1, 4, 7, 8, 9 indicated by ⛭), internally smooth. Notogaster. Dorsal view, notogaster polyhedral-rectangular shape (Figure 9); d.sj) convex, clearly delimited (Figure 1). Deep, round-ovoid n.a.d present, extending posteriorly from d.sj. Ten pairs of setae c, la, lm, lp, h1, h2, h3, p1, p2, p3; setae c situated on lateral margin of n.a.d (Figures 1, 4, 7); setae c: looped, dentate, sharply tipped (Figure 19). Length: 150 (144-175); la, lm, lp, h1, h2, h3, p1, p2, p3 (Figures 1, 2, 16, 23, 25): simple, small dentitions, with parallel longitudinal ridges, sharply tipped (Figure 3); four pairs situated laterally: h3, p1, p2, p3; three pairs (la, h2, h1) situated internally to s.c; two pairs (lm, lp) situated internally to la, h2 (Figures 1, 7, 23). Setae h3, p3 inserted on conspicuous promontories (Figure 1, 23, 25, 42); deep v shaped incision observed behind each seta (Figure, 25, indicated by ➧), determining a scalloped notogastral margin in this region (Figure 1). Setal lengths: la 102 (97-112); lm 134 (127-142); lp 125 (118-132); h1, h2 70 (68-76); h3 91 (86-98); p3 59 (56-64); p2, p1 43 (41-47); s.c completely surrounding notogaster, originating slightly in front of setae la, running between setae la, h1, h2 and h3, p3, p2, p1 (Figures 1, 7 trajectory indicated by ●). Posterior notogastral view (Figure 7). Deep ovoid p.p.d as well as n.a.d clearly visible; setae c situated on paraxial zone of h.ap. Trajectory of s.c indicated by ●; externally to s.c, flat surface of notogaster extending from s.c to notogastral margin; scalloped zone (behind setae h3, p3), some distance from s.c, not interrupting its trajectory. Lateral region.Tu strong, large lamina, together with prodorsal wall and lamellae determining a pocket structure; anterior Tu ending in sharp angle, with interior part welded to prodorsal wall (Figure 1 indicated by ⇣); behind le setal insertion level, pocket structure internally delimited by the Lam (Figures 23, 26) (See Leg-folding process). Pd I, prominent extended lamina; (Figures 26, 27). Pd II, small lamina, rounded apex (Figures 20, 23); Tu, Pd I, Pd II with reticulate-foveate cuticular microsculpture (Figures 20, 26, 27 indicated by ❖). Complex, polyhedral bo situated under the zone where h.ap overlaps the anterior prodorsal zone (Figure 23). Bothridial opening observed at the bottom of a long U-shaped structure (Figures 24a, 24b), with sulcate microsculpture on inferior zone (Figure 24a, 24b indicated by ➡); smooth bothridial ring (bo.ri) surrounding ovoid botridial opening; Lam sigmoid, lacking sharp cuspis (Figures 23, 26); setae le inserted on promontories at Lam apical zone; s.c clearly visible, originating in zone anterior to la setal insertion level (Figure 23, trajectory indicated by ●); v-shaped incision observed behind h3, p3 setae (Figure 25, indicated by ➧); b.ng convex (Figure 23). Ventral region. Epimeral chaetotaxy 3 -1-3- 2 (Figures 13, 42); setae 1c, 3c, 4b situated marginally (Figure 30); setae 1b largest (Figure 41); epimeral borders easily observed; bo.2, b.sj traversing medial plane; bo.3 small; apodemes apo.1, apo.2, apo.dj, apo.3 clearly visible (Figure 13); small setae, many small barbs (Figure 30); length: 9 (5-18). Genital plate ovoid, with four pairs of setae (Figure 35, 36, 37); genital setae: with small barbs, variable in shape (Figures 36, 37); length: 5 (4-7); genital opening on elevated zone (Figures 28, 35); setae ag in margin of elevated zone (Figures 28, 32, 42); medium sized setae ag with small dentitions, sharp tip (Figures 32) ag: 20 (8-21). Complex structure involved in leg-folding process (see Locking structure), situated laterally to setae ag (Figures 28, 31, 42); constituted by longitudinal cuticular elevation, with parallel furrow and lateral to it a cuticular promontory and opposite, a polyhedral plate (in Figures 28, 42 indicated by ) (see Leg-folding process); genital plate smaller than anal plate (Figures 28, 42). Anal opening with elevated zone posterior to h3 insertion level (well visible in ventral posterior view) (Figure 42); anal plate more or less rectangular with rounded anterior and posterior zones; two pairs of anal setae (Figures 28, 32, 38, 42). Figures 28-41. Nippobodes panemorfis sp. n. Adult female, SEM observations. 28 ventral view 29 tuberculate zone 30 discidium with epimeral seta 3c31 ridge associated with leg folding 32 aggenital setae 33 adanal seta 34a subcapitulum 34b subcapitular setae a35 genital zone 36 genital seta type 1 37 genital seta type 2 38 anal zone 39 ornamentation of anal setae 40 epimeral zone 41 epimeral seta 1a. Scale bars: 200 μm (28); 50 μm (35, 38, 40); 20 μm (31, 34); 10 μm (29, 30, 33); 5 μm (41); 3 μm (34); 2 μm (32, 36, 37, 39). Setae: an small dentitions, parallel ridges (Figure 39), length 14 (15-20); three pairs of adanal setae (Figures 28, 33, 38, 42); setae ad1, ad2 inserted on elevated zone; ad3 setae smallest (Figures 38, 42); adanal setae: medium sized, small dentitions, sharply tipped (Figures 32, 33), length 32 (30-36). Three pairs of subcapitular setae, h, m and a: h, m simple, finely barbate (Figures 34a, 44), a elongated leaf-shaped, with some narrow, shallow longitudinal furrows (Figure 34b); setae h situated in margin of tuberculate zone (Figure 42 indicated by ⌘); setae m curving, lengths: h 10 (9-12); m 30 (28-32); a 11 (10-12). Figures 42-47. Nippobodes panemorfis sp. n. Adult female, SEM. 42 ventral view 43 epimeral seta 2a44 subcapitulum anterior zone 45 palp, lateral view 46 palp, anterior zone 47 porose area, detail w solenidium. Scale bars: 42 200 μm ; 44 20 μm ; 45 10 μm ; 5 μm (46); 2 μm (43); 0.1 μm (47). Palp (Figure 34): the first four segments display normal setation (0 -2-1- 3); tarsus particular, presenting only: cm barbate, (vt) barbate, w solenidion and eupathid acmx, sulx, ul'x , ul"x. Solenidion unusually shaped (Figures 45, 46), with porous surface (Figure 47); eupathid sulx, ul'x , ul"x with an obvious apical perforation (Figure 46). Legs. See Figures 48-53, Table 1. All legs with very small genu and long tibia. Femur leg IV with large round porose area (Figure 53). Femur III with large femoral groove (f.g) (Figures 51, 52) (see Leg-folding process). Setal formulae I (1 -4-2-4-16- 1) (1 -2- 2); II (1 -4-3-2-15- 1) (1 -1- 2); III (2 -3-1-2-15- 1) (0 -1- 0); IV (1 -2-1-2-14- 1) (0 -1- 0). Table 1. Nippobodes panemorfis sp. n. setae and solenidia.

Femur	Genu	Tibia	Tarsus	Claw
Leg I
dp,da,l",v	d,v	d,(l), v	(ft)(tc),(it),(p),(u),(a),s,(pv)
1,2	1,2
Leg II
dp,da,l",v	d,v,l	l'v	(pv)(ft),(tc),(it),(p),(a),(u),s
1,2
Leg III
d, l' ,v	d	l' ,v	(pv),s,(a),(u),(p),(it),(tc),(ft)
Leg IV
d,v	l	l",v	ft",(tc),(it),(p)(u),(a),s,(pv)

Figures 48-53. Nippobodes panemorfis sp. n. Legs. Adult female, optical observations. 48 leg I antiaxial 49 leg I, detail solenidia and famulus 50 leg II antiaxial 51 leg III, antiaxial 52 femur III ventral, detail of femoral groove 53 leg IV antiaxial. Scale bars: 100 μm (48, 50, 51, 53); 15 μm (49); 10 μm (52). Remarks. Future ontogenetic studies are necessary in order to confirm nomination of notogastral setae. As only the adult stase was available for study, we used standard, previously used notation (see Morphological terminology). We were unable to locate information on the palp in previous studies. Setae l' of genu II were indicated by Chen and Wang (2007) as bifurcate; however, in our studies only one instance of bifurcate setae l' was observed. Another particularity is the presence of (it) on tarsus IV. The femoral groove was observed, though not indicated in any previous study.