KEY WORDS Acari, Oribatida, Lohmanniidae, Vietnam, new genus, new species.

Transcription

1 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam: the family Lohmanniiae (Acari: Oribatia), with the escriptions of Beoslohmannia anneae n. gen., n. sp., an Paulianacarus vietnamese n. sp. Nestor FERNANDEZ National Council of Scientific an Technological Research (CONICET), Subtropical Biological Institut (IBS), Evolutive Genetic Laboratory FCEQyN, Misiones National University, Felix e Azara 1552, 3300 Posaas Misiones (Argentina) nestorfernan51@yahoo.fr Pieter THERON Research Unit for Environmental Sciences an Management, North-West University, 2520 Potchefstroom Campus (South Africa) pieter.theron@nwu.ac.za Christine ROLLARD Muséum national Histoire naturelle, Département Systématique et Évolution, UMR 7205 case postale 53, 57 rue Cuvier F Paris ceex 05 (France) chroll@mnhn.fr Elio Rorigo CASTILLO National Council of Scientific an Technological Research (CONICET), Subtropical Biological Institut (IBS), Evolutive Genetic Laboratory FCEQyN, Misiones National University, Felix e Azara 1552, 3300 Posaas Misiones (Argentina) castillo.eliororigo@gmail.com Fernanez N., Theron P., Rollar C. & Castillo E. R Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam: the family Lohmanniiae (Acari: Oribatia), with the escriptions of Beoslohmannia anneae n. gen., n. sp., an Paulianacarus vietnamese n. sp. Zoosystema 36 (4): KEY WORDS Acari, Oribatia, Lohmanniiae, Vietnam, new genus, new species. ABSTRACT In this first work on the oribati mite fauna of Hòn Chông limestone hills, Vietnam, we escribe Beoslohmannia anneae n. gen., n. sp. an Paulianacarus vietnamese n. sp., in both cases stuie with optical microscopy. Beoslohmannia n. gen. presents a very interesting leg-foling process, unusually shape legs an significant notogastral neotrichy. Paulianacarus Balogh, 1960 is newly reporte from Vietnam an P. vietnamese n. sp. is most similar to P. rugulosus Mahunka, 1995 from Thailan. Publications Scientifiques u Muséum national Histoire naturelle, Paris

2 Fernanez N. et al. MOTS CLÉS Acariens Oribates Lohmanniiae Vietnam genres nouveaux, espèces nouvelles. Résumé Les acariens Oribates es sols profons es collines calcaires e Hòn Chông, Vietnam: la famille es Lohmanniiae (Acari: Oribatia), avec la escription e Beoslohmannia anneae n. gen., n. sp., et Paulianacarus vietnamese n. sp. Dans ce premier travail sur la faune Oribatologique es collines calcaires e Hòn Chông u Vietnam, nous écrivons Beoslohmannia anneae n. gen, n. sp. et Paulianacarus vietnamese n. sp., tous eux étuiés en microscopie optique. Beoslohmannia n. gen. présente un système très intéressant pour replier ses pattes, une forme e pattes très particulière ainsi qu une importante néotrichie notogastral. Paulianacarus Balogh, 1960 est nouveau pour le Vietnam et P. vietnamese n. sp. ressemble à P. rugulosus Mahunka, 1995 e Thaïlane. INTRODUCTION The Hòn Chông limestone hills of Kien Giang Province, locate in the extreme southwestern part of Vietnam, harbours cave an soil invertebrate communities exceptional in richness an micro-enemics (Deharveng et al. 2009). These tiny hills (the largest one has an area of less than one km 2 ) are currently uner intense pressure ue to quarrying, an a large part of their fauna is critically threatene, with high risk of species extinctions. In spite of the large number of genera an species escribe from this area uring the last two ecaes, plenty of remarkable taxa are yet to be escribe. The most iversifie soil mites, principally oribatis, are also the least stuie. A single species of mite (Actinotrichia, Eutrombiiiae) has been escribe so far from the area (Makol & Gabrys 2005) an another has been ientifie (Deharveng et al. 2009), whereas hunres of species remain to be examine in the samples at han. Stuying the bioiversity of Hòn Chông limestone hills is a very important consieration in the impact assessment of this small an fragile ecosystem. The extreme isolation of these karst hills can generate enemism among arthropos (Deharveng et al. 2009) as in other isolate karsts of tropical regions. The number of arthropo species collecte in Hòn Chông is approaching 500, most of them unescribe. Actually, this number strongly unerestimates the total bioiversity of the area, because ata resolution is extremely uneven from one group to the other, ranging from species or morphospecies (Collembola, Pauropoa, cave groups ) the family (many millipees an centipees) or even orer (many spiers, mites, aquatic fauna). Mites in particular have harly been stuie though they woul certainly provie the largest iversity of all soil arthropos (Deharveng et al. 2009: 37). Beoslohmannia anneae n. gen., n. sp., escribe in the present paper isplays a very particular type of setae covering the boy an legs. At the moment of eployment of the protection mechanism (Fernanez et al. 2013a), the legs are conceale behin the large umbrella-shape setae present on all legs. The notogaster, ventral region an legs also show interesting particularities. In Paulianacarus vietnamese n. sp., all setae are small an the anal an aanal plates are similar to those in Paulianacarus rugulosus Mahunka, 1995 from Thailan. Due to the limite number of specimens, their rarity an particularities, we i not perform Scanning Electron Microscopy stuies, which woul result in the loss of the coate material, thus epositing the largest possible number of specimens in the MNHN. 772

3 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam MATERIAL AND METHODS The material was obtaine by Louis Deharveng an Anne Beos using Berlese-funnel extractors (see particulars uner each species) in Specimens were macerate in lactic aci, an observe in the same meium using the open-mount technique (cavity slie an cover slip) escribe by Granjean (1949) an Krantz & Walter (2009). Drawings were mae by means of an Olympus BHC compoun microscope (Rungis, France), equippe with a rawing tube. Measurements taken: total length (tip of rostrum to posterior ege of notogaster) an with (wiest part of notogaster), in micrometres (μm). Setal formulae of the legs inclue the number of soleniia (in parentheses); tarsal setal formulae inclue the famulus (ε). In orer to stuy several structures, principally notogaster, subcapitulum an legs, an particular setae, specimens were issecte an monitore uring the lactic aci maceration process (in warm 70% lactic aci), before being staine with chlorazol black E (Coineau 1974). Abbreviations Morphology Morphological terms an abbreviations use are those evelope by Granjean (1950); Norton & Behan-Pelletier (2009); Fernanez et al. (2013a-c). Institutions MNHN Muséum national Histoire naturelle, Paris. SYSTEMATICS Family Lohmanniiae Berlese, 1916 Genus Beoslohmannia n. gen. Type species. Beoslohmannia anneae n. sp. Etymology. The generic prefix Beos an the specific epithet are in homage to Dr Anne Beos, MNHN. Diagnosis (ault) Notogastral neotrichy Genital plate univie, ten pairs of setae; anal an aanal plate fuse, with six pairs of setae, four pairs antiaxial an two pairs paraxial situate in the mile posterior zone; preanal plate broaly triangular, posterior part covering the anterior anal-aanal plate zone; preanal plate, anterior part fuse on posterior genital opening margin. Leg ventral blaes: femora I an II with two prominences; femora III an IV with teeth; all femora presenting umbrella-shape setae that conceal legs uring foling process. Beoslohmannia anneae n. sp. (Figs 1-4; Table 1) Type material. Holotype: Vietnam, Kien Giang Province; Kien Luong; Nui Hon Chong sommet est, flanc nor, 27.I.2003, seconary forest, 1, Berlese funnel extraction of eep mineral soil. Louis Deharveng an Anne Beos leg. (VIET-929) X ; Y , alt. 120 m. Paratypes: same ata, 2. MNHN, preserve in 70 % ethanol. Diagnosis (ault female) Setation Umbrella-shape an simple. Umbrella-shape: with two variations, one with roune an more or less smooth ege, the other with spiky ege. Smooth ege: ro, le, exa, exp; notogastral, subcapitular h, m 2 ; epimeric; genital antiaxial; aanal; spiky ege: in setae. Simple setae: subcapitular a, m 1 ; paraxial genital; paraxial aanal. Umbrella-shape setae covering almost entire boy an legs. Proorsum Triangular to polyheral in orsal view, anteriorly truncate in lateral view; postbothriial transverse ban sb, poorly evelope. Rostrum roune; bothriium large, orso-laterally opene; sensillus setiform, particularly curve, short barbs place istally; in inserte between bothriia; Notogaster. Transverse bans not observe; setae pairs: twelve pairs primary; pairs aitional neotrichial setae; all umbrella-shape; c 1, c 2, c 3, 1, 2, cp, e 1, e 2, f 1, f 2, h 1, h 2 large; neotrichal setae small. Five pairs of lyrifissures present, ia, ip, below unsclerotize lateral longituinal line; ips on aanal fol ban; im above e 2 setae an ih posterior to h 2 setae. 773

4 Fernanez N. et al. Ventral region Subcapitular setae, four pairs. Aoral setae: or 1 largest, with apical bea; or 2 elongate; or 3 small, sharply tippe. Chelicera: fixe igit, four teeth; chb seta well evelope, cha seta only represente by insertion. Apoemata short, clearly visible; epimeric setal formulae Anal setae two pairs; aanal setae four pairs. Lateral region Peofossa housing retracte legs I-IV clearly visible; unsclerotize longituinal line, terminating at level of ih lyrifissure. Legs All claws monoactyle, with tooth antiaxially. Femur I, II with three umbrella-shape setae; III, IV, with one only. Setal formulae I ( ) (2-1-2); II ( ) (1-1-2); III ( ) (0-1-1); IV ( ) (1-0-0). Soleniia setiform; setae present on tibia an genu I, II, III an only on genu IV. Description Measurements 530 μm ( ) 243 μm ( ) (Holotype 1 an 2 paratypes measure). Shape Oval (Fig. 1A, B). Colour Specimens without setae: yellow to light brown; slightly shiny, when observe in reflecte light. Cerotegument Almost nonexistent; only small amount present near insertion on first an secon legs. Integument Polyheral microsculpture on proorsum, zone aroun bo, only visible in transmitte light. Setation (legs not inclue; see Legs an Foling process) Two types: umbrella-shape an simple setae. Two variants of umbrella shape: one with roune an more or less smooth ege, the secon with spiky ege. Smooth-ege variant foun on: proorsum (Fig. 1B, ro, le, exa, exp setae); notogaster (Fig. 1B, all setae); subcapitular (Fig. 2E, h, m 2 ); epimeric (Fig. 2B, all setae); genita (Fig. 2D, (antiaxial setae); aanal setae (Fig. 2D). Spiky-ege variant: only in setae (Fig. 1B). Simple setae on: subcapitulum, a, m 1 (Fig. 2E); paraxial zone of genital plate (Fig. 2D); paraxial setae aanal plate (Fig. 2D). Umbrella-shape setae presenting central zone with peicel; upper part umbrella-shape, with irregular margin an protuberances of ifferent size an nerves (Figs 1B; 2B, D, E). Proorsum Triangular to polyheral in orsal view (Fig. 1D); triangular, but anteriorly truncate in lateral view (Fig. 2A). Rostrum broaly roune; chitinous rige present on either sie of proorsal area (Fig. 1D). Each ro seta inserte near the anterior ege of this rige; le setae place near rige, short istance posteriorly an paraxially to ro; bo large, orsolaterally opene; si setiform, particularly curve (Figs 1A, B, D; 2A) with a few short barbs; in inserte between bothriia; exa situate near chitinous rige. Umbrella-shape setae covering almost entire proorsum (Fig. 1B); shape ifference between in an other proorsal setae is notable (Fig. 1B). Faint polygonate pattern present anterior an posterior bo. Postbothriial transverse ban sb harly iscernible, only visible in specimens without setae (Fig. 1D). Dorsal view, ue transparency (l.) observe between exa an le (Fig. 1D). Notogaster Transverse bans not observe. Twelve pairs of primary notogastral setae: c 1, c 2, c 3, 1, 2, cp, e 1, e 2, f 1, f 2, h 1 an h 2 well iscernible, an between pairs of aitional neotrichial setae (Fig. 1C), situate behin h 2. Umbrella-shape setae covering almost entire notogaster (Fig. 1B). Five pairs of lyrifissures present, ia, ip, situate below the unsclerotize lateral longituinal line 774

5 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam ro A ro B le le exa in c1 exp c1 c3 c e1 h2 3 e1 e2 f1 h1 si bo exp c2 1 exa in e2 f2 f2 h2 h1 C D ro le E F chb exa cha in bo si exp chb cha Fig. 1. Beoslohmannia anneae n. gen., n. sp. ault: A, orsal view, without setae; B, orsal view with setae; C, setal neotrichy isposition; D, proorsum; E, chelicera; F, fixe igit. Abbreviations: see Material an methos. Scale bars: A, B, 100 μm; C, E, 50 μm; D, F, 150 μm. (Fig. 2A) (see Lateral region); ips situate on the aanal fol ban (Fig. 2C [BPAD]); im above e2 setae an ih behin h2 (Fig. 2C). Lateral region Proorsal margin present on either sie of cavities where legs I an II are house when retracte; 775

6 Fernanez N. et al. Table 1. Beoslohmannia anneae n. gen., n. sp., setae an soleniia. Abbreviations: see Material an methos. Leg I Femur Genu Tibia Tarsus Claw Seta va, vp, l, (l),, (l), v (p), (it), (tc), (ft), ε, p, (pv), s, (a), (u) 1 Soleniia σ 1, σ 2 -j ω 1, ω 2 - Leg II Seta,, va, vp, lp, (l), (l), v (ft), tc, (it), (p), (u), (a), s, (pv) 1 Soleniia σ- -j- - ω 1, ω 2 - Leg III Seta (l), lp,,,, v (u), (p), (tc), (ft), a, s, (pv) 1 Soleniia σ- -j Leg IV Seta lp,, la,, (u), (p), (tc), (it), a, s, (pv) 1 Soleniia σ clearly visible proorsal posterior margin (b.p) an attachment lines of articular skin λ 1 an λ 2 (with proorsum an notogaster respectively) (Fig. 2A). Anterior notogastral zones presenting conspicuous tectum (Fig. 2A) an clearly efine unsclerotize lateral longituinal line, terminating almost at level of ih lyrifissure an elimiting notaspis (unpaire orsal) an pleuraspis (paire narrow lateral zones) (Fig. 2A); in posterior notogastral zone, when unsclerotize line oes not exist, notaspis an pleuraspis not elimite (Fig. 2A). Anteriorly each pleuraspis presenting a roune lobe between legs II an III, where lyrifissure ia is present; posteriorly at level of 3 an e 2 setae, well-marke ege forming roof of cavities in which legs III an IV are house when retracte, with a protruing angle between them (Fig. 2A). Ventral region Subcapitulum anterior zone more or less triangular, posterior zone ovoi. Four pairs of subcapitular setae (Fig. 2E) (see Setation). Aoral setae characteristic: or 1 largest, with apical bea; or 2 elongate; or 3 with small sharp tip (Fig. 2F). Chelicera fixe igit with four teeth, one of them paraxial (inicate in Fig. 1F with otte shaing); chb setae well evelope; only insertion of cha setae visible, situate laterally an posterior to chb (Fig. 1E). Coxisternal region ivie into two parts by the ventrosejugal groove. Apoemes short an clearly visible; epimeric setal formulae , all seta of similar shape (Fig. 2B) (See Setation). Genital plate univie with ten pairs of setae (Fig. 2D); six simple setae aligne paraxially, four umbrellashape setae antiaxially in semi-circle, following shape of genital plate margin. Preanal plate more or less triangular: anterior margin fuse to posterior margin of genital opening, impossible to remove without breaking (Fig. 2C); posterior margin covering anterior part of ano-aanal plate, internally excavate to receive them (Fig. 2C). Anal plate fuse to aanal, resulting in single ano-aanal plate with six pairs of setae (Fig. 2C, D). Paraxial alignment of two simple setae situate in the mile posterior zone of the plate; antiaxial alignment with four umbrella-shape setae (Fig. 2C, D). Ban BPAD clearly visible in specimens immerse in lactic aci for a lengthy perio; lyrifissure ips present near margin of this ban (Fig. 2C, D). Legs Stuying the legs is very complex, an ue to limite number of aults available for stuy, it is necessary to consier our stuy preliminary (see Remarks). Legs are particular by their shape, principally at femur level (Fig. 3A-D), an ue to the presence of the very large umbrella-shape setae. Each claw bearing a single prominent tooth situate antiaxially (Fig. 3A-D). Between tarsus an genu, all legs have a classic appearance, but femora present particular charac- 776

7 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam A h 1 f 1 e 1 1 λ 2 c 1 si ih h 2 f 2 ip im e 2 2 bp 3 c 3 λ 1 c 2 exp in exa le ia IV III II I ro A, B B C, D 1b 1c 1a 2a C G D 3a 3b 3c ia pr ips 4b 4c 4 l. 4a E a BPAD pr m 2 m 1 F or 2 or 1 h or 3 LL Fig. 2. Beoslohmannia anneae n. gen., n. sp. ault: A, lateral view; B, ventral view (genital an ano-aanal zone, semi schematic); C, genital opening, preanal plate an aanal plate without setae; D, genital opening, preanal plate an aanal plate with setae; E, subcapitulum; F, aoral setae. Abbreviations: see Material an methos. Scale bars: A-D, 100 μm; E, 60 μm; F, 20 μm. teristics. Two types of femur can be istinguishe. Femora legs I an II isplaying large ventral blae (Fig. 3A, B), with two prominences, carrying three umbrella-shape setae. Femora legs III an IV has ventral blae, with teeth, carrying only one umbrella-like seta (Fig. 3C, D). Femur I robust, wie an long; ventral blae situate on proximal part, presenting two more or less 777

8 Fernanez N. et al. (it) σ 2 A φ (l) ω 1 (ft) ε v p ω 2 ω 1 ω 2 (ft) tc (pv) (tc) (it) (p) (u) s (a) σ B φ (l) (l) v (pv) (p) (a) s (u) σ 1 (l) va vp va σ φ lp vp C (l) lp v (pv) (ft) a (p) s (tc) (u) σ D la (ft) (pv) (tc) (p) (u) a s lp Fig. 3. Beoslohmannia anneae n. gen., n. sp. ault: A, leg I, antiaxial; B, leg II, antiaxial; C, leg III, antiaxial; D, leg IV, antiaxial. Abbreviations: see Material an methos. Scale bar: 70 μm. roune prominences, the istal largest; insertion of one umbrella-shape seta on istal one. Secon umbrella-like seta situate at the same level, but more orsally, the thir towars posterior near articulation with trochanter. Femur II iffering in shape; narrower an less robust, ventral blae situ- 778

9 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam A B C D Fig. 4. Beoslohmannia anneae n. gen., n. sp. ault, leg foling: A, leg I; B, leg II; C, leg III; D, leg IV. Scale bar: 70 μm. ate further back than in femur I. Proximal prominences small, polyheral, an carry insertion of one umbrella-shape seta; istal prominence roune an larger than the previous. Secon umbrellashape seta situate istal to first, the thir istal near articulation with trochanter. Femur III presenting ventral blae with three teeth, umbrella-shape seta situate towars istal; orsal leaf-shape setae present, small in relation to umbrella-shape setae. Femur IV isplaying more or less roune blae with only two teeth; umbrella-shape setae situate istal. Setal formulae I ( ) (2-1-2); II ( ) (1-1-2); III ( ) (0-1-1); IV ( ) (1-0-0). All soleniia setiform; setae present on tibia an genu I, II, III an genu IV, situate near soleniia (Fig. 3A-D). 779

10 Fernanez N. et al. Leg foling process Depressions housing legs (l.) are epresse zones concave-arc shape, irecting slightly obliquely, situate externally to epimeric zone (inicate by Granjean 1950). At the moment of leg-foling, this zone partially conceals legs. Eviently the paraxial sie is the unique protecte zone. In this instance, epressions l. show a complementary association with umbrella-shape setae situate on the antiaxal surface of legs an femurs. During leg-foling, tarsus an tibia are irecte towars trochanter an femur, an the genu functions as a hinge (Fig. 4A-D). Legs are house in epressions an large umbrella-shape setae cover the entire antiaxial sie. This protection mechanism is very simple, but effective. Umbrella-shape epimeric setae assisting with camouflage, resulting in legs being very ifficult to etect in specimens with all setae intact. In Figure 4A-C, legs are not completely fole, permitting an unerstaning of positioning of legs. When fully fole, as seen in several cases, only extremities of long setae an the part of soleniia are visible (see Discussion). Remarks Observation by optical microscopy is ifficult ue to orsal an ventral covering of specimens by umbrella-shape setae, which have to be remove for stuy. Legs are challenging to stuy, because in all specimens legs were situate in leg-housing epressions, an obtaining whole legs by issection with all setae intact was almost impossible. Preparations containing lactic aci resulte in margins of umbrella-shape setae becoming torn. Setal shapes were similar in all cases, but sizes varie; in several cases, uring observation of setae slightly from the sie, they appeare to be fan-shape structures. Neotrichous setae are small in relation to others. Determining the number of setae affecte by neotrichy necessitate removal of all, an observing insertions. A similar situation arose on the epimeric zone where all setae were remove an insertions stuie. One of the particularities of B. anneae n. gen., n. sp. is the leg-foling process involving umbrella-shape setae on femora, an the leg housing epressions situate in the lateroventral zone. Processes observe in Bovicaraboes an Beoslomannia are very ifferent an involve issimilar structures an strategies. Granjean (1950: 101) inicate that the leg housing epression, Niches latérales pour les pattes, permits receiving of legs when the specimen retracts them against the boy. This observation by Granjean is very important, but etaile stuies on this aspect of Lohmanniiae were not mae (Norton 2010 cite in page 9 an 14 this structures with the term peofossae, as a characteristic of Lohmanniiae). Our initial observations create the impression that these mites ha no legs as none were visible. The problem was that specimens were covere by umbrella-shape setae an soil particles, together impeing aequate observation. Soil particles were eliminate an legs were foun completely conceale behin umbrellashape setae on legs, an legs retracte in the leg housing epressions. Epimeric umbrella-shape setae contribute to the protection mechanism. On legs I an II we foun three umbrella-shape setae an on III an IV only one. In both cases, when legs are retracte to their maximum, only the apical zone of the longest setae an the soleniia are partially visible. Femoral shape an the number of umbrella-shape setae vary among legs I-IV, possibly relate to the larger number of sensorial organs to protect (in an evolutionary sense). Genus Paulianacarus Balogh, 1960 Paulianacarus vietnamese n. sp. (Figs 5-7; Table 2) Etymology. The specific epithet is eicate to the country Vietnam, where the specimens were collecte; vietnamese (Latin = to Vietnam). Type material. Holotype: Vietnam, Kien Giang province; Kien Luong; Hon Chong, Nui Hon Chong: nor-est e la colline, bosquet en bas e pente, terre noire fine assez sèche, 27.I.2003, seconary forest, 1, Berlese extraction: Louis Deharveng an Anne Beos leg. (VIET-930). X ; Y alt. 5 m. Paratypes: same ata, 2. MNHN, preserve in 70% ethanol. 780

11 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam A l ro B a l. m 2 h m 1 exp l. exa II I 1c 1b 1a sb 2a c 3 ia 3c III 3b 3a 3 4 IV 4c 4b 4a e 2 ip G ips f 2 ih s 8 p an 2 h h1 p 1 p 2 an Fig. 5. Paulianacarus vietnamese n. sp. ault: A, orsal view; B, ventral view. Abbreviations: see Material an methos. Scale bar: 170 μm. Diagnosis Complex microsculpture: punctillate, polyheral. Three types of setae: simple, small: proorsum, notogaster, epimeric area, genital paraxially, aggenital, anal, aanal; simple, large, thick: subcapitular a, m 1, m 2 ; barbate: subcapitular h, genital antiaxially. Proorsum. triangular, polyheral; rostrum bilobate; ro setae inserte istant from rostrum; le seta posteriorly 781

12 Fernanez N. et al. an laterally to ro setal insertion level; bo cup-shape, orsally opene; si, setiform, small barbs istally; in setae posterior to bothriium an near transverse ban; exa, exp positione marginally. Postbothriial transverse ban sb clearly iscernible, posterior to le setae an anterior to orsosejugal furrow. Notogaster Sixteen pairs of setae: c 1, c 2, c 3, 1, 2, cp, e 1, e 2, f 1, f 2, h 1, h 2, h 3, p 1, p 2, p 3. Transverse ban S 2 situate posterior to c setal alignment, only ban crossing sagittal plane; S 3, S 4 between c, setal alignment; S 5 small, behin setal alignment; S 6 curving, posterior to e 2 setal insertion; S 8 visible in ventral view, anterior to p 3 ; S 9 posterior to h 2. Five pairs of lyrifissures: ia, ih, im, ip, ips. Lateral region Pleuraspis anterior zone roune lobe; between legs I, II, zone small teeth; unsclerotize longituinal line exceeing level ih lyrifissure. Ventral region Four pairs of subcapitular setae; epimeric setal formulae ; genital plate univie, 10 pairs of setae paraxially, four antiaxially; ano-aanal plate incompletely fuse; anterior part of plate, presenting a very faint line, permitting ientification of both plates in this zone; six pairs of setae; lyrifissure ips present as roune structure on aanal fol ban. Ault escription Measurements 788 μm ( ) 441 μm ( ) (three specimens; eposite in MNHN). All specimens female. Shape Ovoi elongate (Fig. 5A, B). Colour Yellow to light brown; slightly shiny when observe in reflecte light. Cerotegument Nonexistent; or isappearing uring observation in lactic aci. Integument Complex microsculpture; punctillate cuticular surface an a little below polyheral (Fig. 7D), microsculpture very ifficult to observe on cuticular structures for example transverse ban. Setation (legs not inclue) Three types: a) simple, small: proorsum, notogaster, epimeric, genital paraxially, aggenital, anal, aanal, (Figs 5A, B; 6A); b) simple, large, thick: subcapitular a, m 1, m 2 (Fig. 5B); c) barbate: subcapitular h, genital antiaxially (Fig. 5B). Proorsum Triangular to slightly polyheral in orsal view (Fig. 5A); triangular in lateral view (Fig. 6A). Rostrum with meial roune incision, efining a bilobate rostrum (Fig. 6A; inicate by arrow), only visible in orsal or ventral views (Figs 5A, B). Depression housing legs l. (for legs I an II) (Fig. 5A) well visible as concave arc-shape zones, situate laterally (arrows inicating the course in Fig. 5); ro setae inserte far from rostrum at level of anterior arche margin of l. (observe by transparency) (Fig. 5A); le setae place posteriorly an laterally to ro setae insertion level; le setal insertion foun at level of more arche zone of l. (Figs 5A; 6A) (observe by transparency); bo cup-shape, orsally opene (Fig. 6C); si setiform, with few small barbs istally (Figs 5A; 6B); in inserte posterior to bothriium an near transverse ban sb (Figs 5A; 6A); exa an exp well visible, situate marginally (Figs 5A; 6A). Transverse postbothriial ban sb clearly iscernible in orsal an lateral views; situate posteriorly to le setae an anteriorly to sj furrow; lateral en at bothriial level (Figs 5A; 6A). Notogaster Sixteen pairs of notogastral setae: c 1, c 2, c 3, 1, 2, cp, e 1, e 2, f 1, f 2, h 1, h 2, h 3, p 1, p 2, p 3 clearly iscernible (Figs 5A; 6A). Transverse bans: S 2 clearly visible (Figs 5A; 6A), situate posterior to c setal alignment, only ban crossing sagittal plane; meial limit inistinct an variable among specimens. In orsal view (Fig. 5A) the limit is between setal insertions c 1 an c 2. In lateral view (Fig. 6A) the same limit almost forming 782

13 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam e 1 1 c 1 si in le A h 3 f 1 s 6 s 5 im s 4 s 3 2 s 2 c 2 bp sb bo exa ** ro h 1 s9 h 2 f 2 e 2 3 c 3 exp ** p 2 p 3 ih ** IV ** III ia vsj II I σ φ ft (tc) B la v p pv (a) (p) pv s (u) lp D φ v C si σ σ bo (ft) (a) (tc) (p) E v pv s (u) Fig. 6. Paulianacarus vietnamese n. sp. ault female: A, lateral view; B, leg III antiaxial; C, bothriium, lateral view; D, genu, tibia III, orsal view; E, leg IV, antiaxial. Abbreviations: see Material an methos. Scale bars: A, 350 μm; B, E, 270 μm; C, 23 μm; D, 45 μm. 783

14 Fernanez N. et al. Table 2. Paulianacarus vietnamese n. sp., setae an soleniia. Abbreviations: see Material an methos. Leg I Femur Genu Tibia Tarsus Claw Seta lp, bv, v,,, (l) (l), x, v (pv), m, n, s, (a), (u), (p), (tc), (ft), ε 1 Soleniia σ 1, σ 2 -j ω 1, ω 2 - Leg II Seta, v, vp, lp, (l), (l), (l), (pv), s, (a), (u), (p), (tc), (ft) 1 Soleniia σ- -j- ω Leg III Seta, lp, la, v, ft, p, (pv), s, (a), (u), (tc), (p) 1 Soleniia σ- -j Leg IV Seta,, v,, (ft), (a), pv, (tc), (p), (u), s 1 Soleniia σ a line terminating near c 3 setal insertion; S 3 an S 4 foun between c an setal alignment (Figs 5A; 6A); S 5 small, situate posteriorly to setal alignment an anteriorly to gla (Fig. 5); S 6 curving, situate posterior to e 2 setal insertion; S 8 only visible in ventral view (Fig. 5B) anteriorly to p 3 setal level; S 9 situate posteriorly to h 2 setal level (Figs 5A; 6A). Five pairs of lyrifissures present: ia, ih, big size, situate below unsclerotize lateral longituinal line; ia anteriorly on frontal lobe of pleuraspis (see Lateral region); ih situate between h 2, f 2 setal level, well visible in lateral an ventral views (Figs 5B; 6A), in ventral view, near S 8 ban, but in variable position situate either in front of or behin this ban; im large, situate above unsclerotize lateral longituinal line, near S 6 ban, clearly visible in lateral view (Fig. 6A); ip small; visible in ventral view (Fig. 5B); ips roune, situate on BPAD fol ban (Fig. 5B). Lateral region Proorsal margin presenting epression housing legs l., clearly visible (inicate with asterisks Fig. 6A) for legs I, II; bp well visible. Anterior zone of pleuraspis presenting roune lobe with lyrifissure ia. Laterally between legs I an II, zone with small teeth (Fig. 6A, inicate with ouble arrow). Anterior notogastral zone presenting conspicuous tectum (Fig. 6A). Unsclerotize longituinal line easily iscernible, exceeing the level of ih lyrifissure an clearly elimiting notaspis an pleuraspis (Fig. 6A). Situate posteriorly on same pleuraspis, l. of legs III, IV. Posterior zone (where unsclerotize line oes not exist) very small (Fig. 6A). Several muscular insertions visible (Fig. 6 inicate with black arrow). Ventral region Subcapitular setae, four pairs (Fig. 5B, see Setation). Apoemes well visible; epimeric setal formulae All setae of similar shape (Fig. 2B, see Setation). Genital plate univie with ten pairs of setae (Fig. 5B); six aligne paraxially an four antiaxially in semi-circle following shape of the genital plate margin. Preanal plate typically shape, characteristic of the genus (Fig. 5B). Anal plate fuse with aanal, elimiting single plate with six pairs of setae (Fig. 5B), anterior part of both plates with faint line, representing vestige of fusion (see Remarks). BPAD clearly visible after lengthy soaking in lactic aci; lyrifissure ips roune, positione near ban margin (Fig. 5B). Legs Setal formulae I ( ) (2-1-2); II ( ) (1-1-1); III ( ) (1-1-0); IV ( ) (1-0-0). Remarks Microsculpture complicating observation: limits of transverse ban S 2 very har to etect in orsal view; in lateral view, easier, as this limit almost 784

15 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam A la σ φ ft (p) (tc) lp B p pv s pv cm ω acm (u) (a) D φ C (l) (l) σ σ (ulζ) su (l) v v v ω 1 v (ft) ε ω 2 (a) (pv) m v lp (tc) (p) (u) s n bv Fig. 7. Paulianacarus vietnamese n. sp. ault: A, leg I, antiaxial; B, palp, antiaxial; C, leg II, antiaxial; D, microsculpture. Abbreviations: see Material an methos. Scale bars: A, C, 350 μm; B, 100 μm; D, 2 μm. becomes a line. Number of genital antiaxial setae variable, on one specimen three setae on one sie an four on the other. Anal an aanal plates prouce a very interesting situation: plates are fuse, but anterior part with line partially elimiting anal an aanal plates; particular microsculpture of aligne points on anal plate, situate parallel, allowing ifferentiation of plates espite being fuse, somewhat similar to Paulianacarus rugulosus Mahunka, 1995 from Thailan. Genital setae number in P. rugulosus is nine, in P. vietnamese n. sp. asymmetric variations in the number of genital an aanal setae were observe, for example with nine-ten genital setae an fourfive aanal setae. Aanal setae show variation in one specimen: one sie with five an another with four. The generic iagnosis of Paulianacarus was mae at the same time the author institute another new 785

16 Fernanez N. et al. genus, Millotacarus Balogh, 1960, both from Maagascar. Several authors inicate that the ifferences between these genera were problematic (Mahunka 1985; Coetzee 2001). Balogh & Balogh 1987 establishe the ifferences between them, aing several characters to the iagnosis of Millotacarus, but this i not solve the problem. The possibility exists that Millotacarus may be a synonym of Paulianacarus. The stuy of type material is neee. DISCUSSION We compare Beoslohmannia n. gen. with all genera presenting the following combination of traits: univie genital plates; fuse anal an aanal plates; two pairs of anal an four pairs of aanal setae; broa preanal plates; gastronotic neotrichy an type an shape of legs. Beoslohmannia n. gen., showe particular characteristics: shape an isposition of preanal plate; isposition of anal an aanal setae; gastronotic neotrichy; leg shape an chaetotaxy, permitting easy ifferentiation of the new genus. Setal types, the leg foling process an other important characteristics observe in the new genus were not inclue in our comparison ue to the lack of comparable stuies. The genus appearing most similar to Beoslohmannia n. gen. is Paulianacarus, for this reason, in the present paper, species from both genera were inclue. In Bovicaraboes eharvengi Fernanez, Theron & Rollar, 2013, B. levyi Fernanez, Theron & Rollar, 2013 an B. fortauphini Fernanez, Theron & Rollar, 2013 (Fernanez et al 2013a) a very interesting protection mechanism was escribe. Several boy an leg structures are involve in this process an a series of particular aaptations are observe. Eviently Caraboiae an Lohmanniae present very ifferent characteristics an are evolutionarily istant, but in both cases a protection mechanism exists. One of the particularities of B. anneae n. gen., n. sp. is the leg-foling process involving umbrellashape setae on femora, an the leg housing epressions situate in the lateroventral zone. Processes observe in Bovicaraboes an Beoslomannia are very ifferent an involve issimilar structures an strategies. Granjean (1950: 101) inicate that the leg housing epression, Niches latérales pour les pattes permits receiving of legs when the specimen retracts them against the boy. On legs I an II we foun three umbrella-shape setae an on III an IV only one. In both cases, when legs are retracte to their maximum, only apical zone of the longest setae an the soleniia are partially visible. Femoral shape an the number of umbrella-shape setae vary in legs I-IV, possibly relate to the larger number of sensorial organs to protect (in an evolutionary sense). Acknowlegements This work is base on research supporte in part by the National Research Founation of South Africa (UID) Any opinion, finings an conclusions or recommenations expresse in the material are those of the authors an therefore the NRF oes not accept any liability in regar thereto. The authors wish to thank the reviewers for their constructive comments. REFERENCES Balogh J Oribates (Acari) nouveaux e Maagascar (1 re série). Mémoires e Institut scientifique e Maagascar, Série A, vol. 14: Balogh J. & Balogh P A new outline of the Family Lohmanniiae Berlese, 1916 (Acari, Oribatei). Acta Zoologica hungarica 33 (3-4): Coetzee L New species of the family Lohmanniiae (Acari, Oribatia) from South Africa. Navorsinge van ie Nasionale Museum Bloemfontein 17 (3): Coineau Y Éléments pour une monographie morphologique, écologique et biologique es Caeculiae (Acariens). Mémoires u Muséum national Histoire naturelle, 22 Zool., 81: 299. Deharveng L., Beos A., Le Cong Kiet, Le Cong Man & Truong Quang Tam Enemic arthropos of the Hon Chong hills (Kien Giang), an unrivalle bioiversity heritage in Southeast Asia, in Le Cong Kiet, Truong Quang Tam & Ly Ngoc Sam (es), Beleaguere Hills: Managing the Bioiversity of the Remaining Karst Hills of Kien Giang, Vietnam. Nha Xuat Ban Nong Nghiep, TP. Ho Chi Minh: Fernanez N., Theron P. & Rollar C. 2013a. The family Caraboiae (Acari: Oribatia) I. Description 786

17 Oribati mites from eep soils of Hòn Chông limestone hills, Vietnam of a new genus, Bovicaraboes with three new species, an the reescription of Haryboes mirabilis Balogh, International Journal of Acarology 39 (1): Fernanez N., Theron P. & Rollar C. 2013b. Revision of the family Caraboiae (Acari:Oribatia) IV. Aftiboes anjaviilavai gen. nov., sp. nov., Rugocepheus joffrevillei sp. nov., an reefinition of the genus Rugocepheus Mahunka, International Journal of Acarology 39 (6): org/ / Fernanez N., Theron P. & Rollar C. & Tie L. 2013c. Family Caraboiae (Acari: Oribatia) V. The genus Congocepheus Balogh, 1958 (secon part), with a reescription of Congocepheus involutus Mahunka, 1997, an escriptions of two new species. Zoosystema 35 (4): Granjean F Observation et conservation es très petits Arthropoes. Bulletin u Muséum Histoire naturelle 21 (2): Granjean F Étue sur les Lohmanniiae (Oribates, Acariens). Archives e zoologie expérimentale et générale 87: Krantz G. W. & Walter D. E A manual of acarology (3r e). Texas Tech University Press, Lubbock, 807 p. Mahunka S Neue un interessante Milben aus em Genfer Museum LIV. Oribatis from South Inia I (Acari: Oribatia). Revue suisse e Zoologie 92 (2): Mahunka S New oribatis (Acari:Oribatia) from Thailan. Acta Zoologica Acaemiae Scientarum Hungaricae 41 (2): Makol J. & Gabrys G Caecothrombium eharvengi sp. nov. (Acari: Actinotrichia Eutrombiiiae) from Vietnam, with a proposal of Caecothrombiinae subfam. nov. Zoologischer Anzeiger, A Journal of Comparative Zoology (4), 243: Norton R. A Systematic relationships of Lohmanniiae (Acari: Oribatia), in Sabelis M. W. & Bruin J. (es), Trens in Acarology. Springer Netherlans: Norton R. & Behan-Pelletier V Suborer Oribatia, in Krantz G. W. & Walter D. E. (es), A manual of acarology (3r e). Texas Tech University Press, Lubbock: Submitte on 20 May 2014; accepte on 31 July 2014; publishe on 26 December

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