Revision of the family Carabodidae (Acari, Oribatida) XII. Yoshiobodescamerunensis sp. n. and Rugocepheuscostaricensis sp. n. Author Fernandez, Nestor Author Theron, Pieter Author Leiva, Sergio Author Tiedt, Louwrens text ZooKeys 2017 706 31 50 http://dx.doi.org/10.3897/zookeys.706.14807 journal article http://dx.doi.org/10.3897/zookeys.706.14807 1313-2970-706-31 6FF97BDA9D4A43B3AD3A94F7C68DE0DC 6FF97BDA9D4A43B3AD3A94F7C68DE0DC Rugocepheus costaricensis sp. n. Figures 36-39, 40-42, 43-52, 53-61, Table 2 Etymology . The specific epithet is derived from Costa Rica, country of origin of the type material. Material examined. Holotype. Adult female "CCR 0978 Tu 11 Costa Rica Turrialba foret naturelle du catie alt. 560 m. Triage d'humus cote est surface nid d'Atta au pied de Castilla elastica 1.IX. 1978. Leg. P.WERNER" Deposited in the Ccllection of the MHNG, Switzerland, preserved in 70% ethanol. Paratypes. 2 adult females, same locality and date of holotype, deposited in the Ccllection of the MHNG, Switzerland, preserved in 70% ethanol. Figures 36-39. Rugocepheus costaricensis sp. n. Adult (with cerotegument), SEM. 36 dorsal view 37in setae 38 fontal inclined view 39 frontal view. Scale bars: 100 μm (36); 5 μm (37); 50 μm (38); 100 μm (39). Diagnosis (adult female). Body and legs entirely covered by simple porous cerotegumental layer. Integumental microsculpture over entire body: irregular, small tuberculate. Prodorsum. More or less triangular in dorsal view; deep low lamellar furrow delineating Y-shaped structure; elevated interlamellar process with superior flat zone; CSO present; rostrum beak-shaped; tutorium and Pedotecta I expanded laterally; small triangular discidium; supratutorial depression deep, with several rounded depressions. Lamellae without lamellar tip, forming bridge concealing le setae. Smooth ring-shaped bothridium; bothridial tooth present; sensillus barbate. Fourteen pairs of notogastral setae c1, c2, da, dm, dp, la, lm, lp, h1, h2, h3, p1, p2, p3. Four notogastral furrows present: paired central longitudinal furrow; one lateral unpaired semicircular furrow; another unpaired semicircular furrow delimiting an unpaired elevated central area (devoid of setae); a pair of elevated areas (with c1, da, dm, dp); unpaired semicircular elevated area (with c2, la, lm, lp, h1, h2). Anterior genital furrow clearly observed; epimere 4 borders elevated; genital plate situated in elevated zone surrounded by furrow. Epimeral chaetotaxy 3-1-3-3; long epimeral setae. Four pairs of genital setae in a single line; crescent-shaped structure anterior to anal plate; anal plate with small sharp tip; aggenital and adanal setae more or less similar in length. Subcapitulum diarthric, three pairs of highly different setae a, m, h. Mentum complex. Description. Measurements. SEM: females 501 μm (489-515) x 270 μm (267-286). Light microscopy: females 506 μm (490-518) x 282 μm (276-301). Colour. Specimens without cerotegument: females light brown to brown. Cerotegument . Simple layer ( +/- 0.7 μm ) (Figures 47, 49, 50); uniformly covering entire body and legs. Slightly irregular surface (Figure 50). Large number of pores observable on the surface, porous (0.4-0.7 μm ) diameter (Figures 44, 46, 47, 50). Integument. Microsculpture simple, covering entire body: irregular, small tuberculate (Figure 49); tubercules (1-2.5 μm ). Only lateral anterior lamellar zone presenting different microsculpture: round to ovoid depressions (Figures 38, 39, 43). Setation. Setae in lanceolate (resembling leaf of Salix spp.), length 30 μm (28-34) (Figure 37); ro setae lanceolate, 13 μm (11-14) (Figure 44). Setae le lanceolate, slightly curved, basally and medially serrate, 23 μm (21-25) (Figure 45). Notogastral setae; c1, c2, da, dm, dp, la, lm, lp,h1,h2 aciculiform, 51 μm (41-61) (Figure 46); h3, p1, p2, p3, 25 μm (23-27) (Figures 36, 46). Simple: ag, 20 μm (17-22) (Figure 61); ad 20 μm (17-22) (Figure 61); ge 17 μm (15-19) (Figure 60); epimeral 18 μm (15-21) (Figure 53). Spiniform: an 10 μm (11-8) (Figure 55); m 3.5 μm (3-4) (Figure 57); Setae a setiform, 7 μm (5-9) (Figure 56); h setae L-shaped, barbate, 19 μm (18-21) (Figure 58). Figures 40-42. Rugocepheus costaricensis sp. n. Adult (with cerotegument) optical microscopy. 40 dorsal view 41 ventral view 42 lateral view. Prodorsum. Very complex. For proper understanding of structures, descriptions from various angles/views are included. Dorsal view (Figure 36). More or less triangular with lateral polyhedral expansion at level of bo and in setal level insertion; anterior expansion of Tu (Figure 36) clearly visible. Deep l.l.f delineates Y-shaped structure (Ys); posterior of Ys with depressed rounded zone (p.Ys) extending laterally in p.p.d *; ro setae and CSO clearly visible. Beak-shaped rostrum; sensillus barbate. Frontal view (Figure 39). More or less triangular; e.i.p elevated with flat superior medial zone (Figure 38); conspicuous l.l.f running to posterior zone of e.i.p; from ro setal zone, l.l.f delineates a Y-shaped structure. On posterior of e.i.p the l.l.f delimiting a large ear shaped structure where in setae are situated. Lamellae (lam), running laterally, internal margin delimited by l.l.f; le setae on the anterior zone of lam; le setae inserted behind ro setal insertion level; small transversal depression posterior to ro setae (Figure 38 indicated by thick arrow); CSO present anterior to ro setal insertion. Rostral zone extended to rounded beak-shape with several transversal semicircular furrows (Figure 38 indicated by dashed arrow). Lateral inclined view (Figures 38, 43). Elevated e.i.p with flat superior zone; lam clearly delimited by conspicuous l.l.f; particular cuticular microsculpture of round to ovoid depressions externally to lam. Elevated ear-shaped structure where in setae are situated; ro setae, CSO, and beak-shaped rostral zone, easily observed. Tu expanded laterally and anteriorly; Pd I: large expanded ovoid structure; several depressions (p.tu.d, p.tu.d1) between Tu and Pd I; s.tu.d a conspicuous depression, running parallel between lam and Tu, with internal round depression (a.tu.d); le setae inserted on anterior zone of lam; lam zone anterior to le insertion, lacking lamellar tip, forming a bridge concealing le setae. Bothridium cup-shaped, smooth bothridial ring, incomplete, with bothridial tooth. Notogaster (Figure 36). Oval, with fourteen pairs of setae: c1, c2, da, dm, dp, la, lm, lp, h1, h2, h3, p1, p2, p3. Four furrows present: paired central longitudinal (c.fu) furrows; one lateral unpaired semicircular furrow (l.fu), and one unpaired semicircular (s.c) furrow; an unpaired elevated central area (i.e.a) is defined by paired c.fu. A pair of elevated areas (p.e.a) defined by c.fu and l.fu; an unpaired semicircular elevated area (i.s.e.a) defined by l.fu and s.c. The i.e.a is devoid of setae; p.e.a with c1, da, dm, dp; i.s.e.a with c2, la, lm lp, h1, h2. Setae h3, p1, p2, p3 situated between s.c and b .ng. Setae c1, c2, da, dm, dp, la, lm, dp, h1, h2 situated on dorsal protuberances (d.pr), while h3, p3, p2, p1 are inserted on lateral thickenings (Figure 43); lyrifissure im and gla clearly visible (Figure 40). Lateral region (Figures 43, 48). Lam (Figure 48) with elevated zone bearing in setae; towards anterior of le setae, lacking lamellar tip, forming a bridge, permitting concealment of setae; s.tu.d a deep depression; tu clearly delimited by prominent thickening; a.tu.d.,p.tu.d1, and p.tu.d2 between tu and Pd I. Rostrum beak-like. Inferior curved margin of lamella continuous with inferior bothridial part; both structures related to s.tu.d, permitting concealment of tarsus, tibia and dorsal area of genu and femur of leg I during leg-folding (protection mechanism). Pd I: large curved extended lamina. Pd II: small rectangular to polyhedral lamina. Humeral apophysis (h.ap): large polyhedral structure, conspicuous oblique posterior furrow on surface (s.fu); anterior h.ap. zone overlapping posterior part of bothridial zone. Discidium (dis): small triangular structure. Several large ovoid depressions behind acetabulum IV and posterolateral to genital and anal openings. Figures 43-52. Rugocepheus costaricensis sp. n. Adult, (with cerotegument) SEM. 43 lateral view 44 rostral zone 45le setae; 46 notogastral setae 47 notogastral cerotegumental layer, ventral view 48 anterior prodorsal zone 49 notogastral cerotegumental layer, ventrolateral view 50 cerotegumental layer 51 bothridial zone 52 ventrolateral inclined zone. Scale bars: 100 μm (43); 5 μm (44); 10 μm (45); 5 μm (46); 10 μm (47); 20 μm (48);10 μm (49); 2 μm (50); 20 μm (51); 50 μm (52). Ventral region. Epimeral zone more or less smooth with large elevations and depressions. Paraxial zone of epimera 1 and 2 with longitudinal furrow; large paraxial depression behind bo.sj. Epimere 4 posterior border elevated. Anterior genital furrow (a.g.f) well visible (Figure 53); genital plate situated on elevated zone surrounded by furrow (Figure 60). Epimeral chaetotaxy 3-1-3-3 (Figure 53); long epimeral setae (Figure 59). Four pairs of genital setae in a single line (Figure 60). Crescent-shaped structure anterior to anal plate (Figure 55 indicated by large dot); anal plate with small sharp tip. Aggenital and adanal setae more or less similar in length (Figures 54, 61). Subcapitulum diarthric (Figure 57); three pairs of highly differing setae a, m, h (Figure 57). Mentum complex. Legs (Table 2). I(1-3-3-4-16-1) (1-2-2); II(1-4-2-3-16-1) (1-1-2); III(2-3-1-2-14-1) (1-1-0); IV(1-2-2-2-12-1) (legs similar to other species, therefore not illustrated). Figures 53-61. Rugocepheus costaricensis sp. n. Adult, (with cerotegument) SEM. 53 ventral view 54ad2 setal zone 55 anal zone 56a subcapitular setae 58h subcapitular setae 57 subcapitulum 59 epimeral 3b setae 60 genital zone 61 aggenital, adanal setae. Scale bars: 100 μm (53); 5 μm (54); 20 μm (55); 2 μm (56); 5 μm (57); 5 μm (58); 5μm (59); 20 μm (60); 10 μm (61). Table 2. Leg setae and solenidia of Rugocepheus costaricensis sp. n.

Legs	Femur	Genu	Tibia	Tarse	Claw
v (l)	(v) d	(l),v	(pv); s; (a); (u); (p); (it); (tc); (ft)
σ	1,2	1,2
(l) d	v l	v (v)	d Ad; (pv); s; (a); (u); (p); (it); (tc); (ft).
σ	1,2
l	d	v (l)	(v) ft; (tc); (it); (p); (u); (a); s; (pv).
σ
d	v d	(l).	v ft; (tc); (p); (u); (a); s; (pv

Remarks. Rugocepheus costaricensis sp. n. displays important differences to Rugocepheus joffrevillei Fernandez, Theron & Rollard, 2013 and R. formosus Mahunka, 2009. Principal differences: beak-shaped rostrum; distribution of furrows and elevated areas on dorsal zone of notogaster, central elevated area without setae; ventral zone with discidium differing in shape; genital and anal zone very different. Discussion. Using SEM allows significant progress in detailed descriptions, as the small body size, morphological characteristics, and complex topology makes Yoshiobodes a difficult genus to study. This complexity is compounded by brief, somewhat cryptic original descriptions and illustrations. Reeves (1997) , contributed much to our understanding of this genus, specifically due to studies of both adults and immatures. Reeves also originally pointed out the following characters with reference to the adult prodorsum of Yoshiobodes : "Dorsosejugal depression deep, slit-like, widest medially" (page 316) (in our series of papers on the revision of the family Carabodidae , this depression is designated as the "posterior prodorsal depression (p.p.d)" Fernandez et al. 2013 ), but this structure was not noted again until this present paper. The analysis by Reeves of the work done by Bellido (1978) is noteworthy as he analyses the depression observed on the prodorsum in protonymphs, deutonymphs and tritonymphs of Carabodes . Reeves (1997) indicates: "The scalloped edged depression on the prodorsum of protonymphs, deutonymphs and tritonymphs appears similar to the foveate sclerite found in immatures described by Bellido (1978) of Carabodes willmanni Bernini, 1975." The most recent generic diagnosis by Ermilov et al. 2014 is based on data from Mahunka (1986) and additions by authors, but the type specimen, Y. irmayi (Balogh & Mahunka, 1969) does not seem to have been studied. SEM and optical microscopy studies by Reeves (1997) on adults as well as ontogenetic studies, were also not discussed. Reeves 1997 indicated that, on comparison, "a specimen of Y. irmayi from St. Lucia (on loan from the Hungarian Natural History Museum) to North American material showed them to be conspecific". For the purpose of this present paper, Yoshioiodes is considered only on the basis of Balogh and Mahunka (1969) ( Carabodes irmayi ) and Reeves (1997) . We await further studies on type specimens of the following subgenera: Yoshiobodes (Yoshiobodes) Mahunka, 1986, type species Carabodes irmayi Balogh & Mahunka, 1969; Yoshiobodes (Berndobodes) Mahunka, 1986 type species, Berndobodes spiculifer Mahunka, 1986; Yoshiobodes (Dongnaiobodes) subgen. n. type species Yoshiobodes hexasetosus Ermilov, Shtanchaeva, Subias & Anichkin, 2014. As part of the ongoing revision of the Family Carabodidae (started in 2013), we have studied the type material of Berndobodes spiculifer Mahunka, 1986, and further information on this genus will be included in an upcoming revisionary paper.