mixacarus(phyllolohmannia)pectinatussp. n. (acari, oribatida, lohmanniidae), a new species of...

ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 5, pp. 766–776. © Pleiades Publishing, Inc., 2014. Original Russian Text © S.G. Ermilov, S. Kalúz, 2014, published in Zoologicheskii Zhurnal, 2014, Vol. 93, No. 6, pp. 709–719.

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Mixacarus (Phyllolohmannia) pectinatus sp. n. (Acari, Oribatida, Lohmanniidae), a New Species of Oribatid Mites from India

S. G. Ermilova and S. Kalúzb

aTyumen State University, Tyumen, 625003 Russia e-mail: [email protected]

bInstitute of Zoology, Slovak Academy of Sciences, Bratislava, 84506 Slovakia e-mail: [email protected]

Received March 1, 2013

Abstract—A new species of oribatid mites (adult and juvenile instars), Mixacarus (Phyllolohmannia) pectinatus sp. n., is described from India. The new species differs from the majority of species of the subgenus in the absence of distally dilated setae and narrower leaf-shaped setae on dorsal and ventral body sides. DOI: 10.1134/S0013873814050091

A new species of the subgenus Mixacarus (Phyllo-lohmannia) J. Balogh et P. Balogh, 1987 was revealed by us in the material from India; this species is de-scribed below.

The oribatid genus Mixacarus Balogh, 1958 (Acari, Oribatida, Lohmanniidae) comprises 2 subgenera and 22 species distributed in the tropics and subtropics (Subías, 2004, online version 2012). Six representa-tives of this subgenus are distributed in the Oriental region (Vietnam, Indonesia, Thailand, and southeast-ern China) (Subías et al. 2012): M. (P.) foliifer Golos-ova 1984, M. (P.) hamanni Balogh, 1961, M. (P.) lui-seae Mahunka, 2008, M. (P.) tenasserimensis Fuan-garworn et Chaisuekul, 2011, M. (P.) variatus Fuan-garworn et Chaisuekul, 2011, and M. (P.) yinae (Hu et Aoki 1993). A key for the above species was com-posed by Fuangarworn and Chaisuekul (2011). The main morphological characters of Mixacarus (Phyl-lolohmannia) were described by Balogh and Balogh (1987) and included by them into keys (Balogh and Balogh, 1987, 1992).

The present work describes the adults and all the juvenile stages (larva, proto-, deuto-, and tritonymph) of the new species. In the subgenus M. (Phylloloh-mannia), the juvenile stages are examined only in two species, (P.) tenasserimensis Fuangarworn et Chaisue-kul, 2011 and M. (P.) variatus Fuangarworn et Chai-suekul 2011 (Fuangarworn and Chaisuekul, 2011). A review of the morphology of developmental stages in mites of the family Lohmanniidae was performed by Ermilov et al. (2012).

The terminology generalized by Norton and Behan-Pelletier (2009) and Norton (2010) was used in the present study. The temporary slides in lactic acid were prepared for morphological studies. The body length (from the rostrum to the posterior margin of the noto-gaster) and the length of the setae was measured in lateral view and the width, in the widest part of the notogaster in dorsal view. Some specimens were dis-sected for the detailed study of the mouthparts and leg chaetom. All the measurements are given in microme-ters.

Mixacarus (Phyllolohmannia) pectinatus Ermilov et Kalúz sp. n.

Material. Holotype, 11 paratypes, and 30 juvenile stages (8 larvae, 6 protonymphs, 4 deutonymphs, and 12 tritonymphs), India, Arunachal Pradesh, 28°19'32"N, 95°57'31"E, 1300 m above sea level, soil in forest, collected by L. Dembicky and O. Sausa, 26.V–01.VI.2012. Mites are stored in a 96% ethyl alcohol with added glycerol. The holotype is deposited at Zoological Institute, Russian Academy of Sciences, St. Petersburg; 4 paratypes, at the Siberian Zoological Museum, Novosibirsk; 7 paratypes and juvenile stages, in the private collection of S.G. Ermilov.

Adult (Figs. 1–3)

Diagnosis. Body length 1078–1261, width 713–780; trichobothria pectinate, with 7 setiform em-branchments; notogaster with 16 pairs of narrow leaf-shaped setae, dilated in median part; subcapitulum

Mixacarus (Phyllolohmannia) pectinatus sp. n.

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with simple (unmodified) setae (including 2 pairs of setae m); epimeral formula 3–1–3–4; 2 pairs of anal and 4 pairs of adanal setae. All ventral setae simple, excluding epimeral setae 1с widened in median part and adanal setae.

Description. Dimensions. Body length of holotype 1078, width 713; of five paratypes, 1078–1261 × 730–780, on average, 1148 × 750.

Integument (Fig. 1, 1, 2). Coloration red-brown. In-tegument of body and legs with micro-pits (visible only under immersion objective, × 1000), forming polygonal ornament. Prodorsum, notogaster, and ano-genital area with rounded porous areas up to 20 in diameter, forming fine dotted transverse strips on notogaster.

Prodorsum (Figs. 1, 1; 2, 1). Rostrum widely rounded. Rostral (ro, 184–192), lamellar (le, 246–250), interlamellar (in, 254–266), anterior exoboth-ridial (exa, 184–192) and posterior exobothridial (exp,

184–192) setae narrow leaf-shaped, widened in me-dian part, pubescent with straight, densely adjacent, needle-shaped cilia. Trichobothria (ss, 168–176) pect-inate, with 7 long setae.

Notogaster (Fig. 1, 1, 2) rounded. Notogastral setae constituting 16 pairs (180–217); all these setae narrow leaf-shaped, pubescent, similar in shape (Fig. 2, 2) to prodorsal setae. Lyrifissurae im situated medially in relation to setae e2; other lyrifissurae (ia, ip, ih, ips) in positions typical of representatives of genus.

Gnathosoma (Fig. 2, 3–5). Subcapitulum longer than wide (295–319 × 254–270). Subcapitular setae simple, represented by 4 pairs: h (77–86), m1 (98–106), and m2 (77–86) weakly pubescent; a (65–73) smooth. Adoral setae smooth: or1 (32–41) leaf-shaped, strongly dilated in median-proximal part; or2 (61–49) leaf-shaped, dilated in median part; or3 (41–49) thick-ened, setiform. Palps (139–143) four-segmented, setal formula 0–1–3–10(+1ω). Chelicerae (303–315) with

Fig. 1. Mixacarus (Phyllolohmannia) pectinatus sp. n., adult: (1) dorsal view; (2) ventral view. Scale 300 µm.

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Fig. 2. Mixacarus (Phyllolohmannia) pectinatus sp. n., adult: (1) interlamellar seta; (2) notogastral seta f2; (3) subcapitulum, left part; (4) palp; (5) chelicerae, anterior part; (6) epimeral seta 1b; (7) epimeral seta 1c; (8) epimeral seta 3c; (9) anterior-median genital seta; (10) posterior-lateral genital seta; (11) anal seta an2; (12) adanal seta ad2; (13) adanal seta ad4. Scale (µm): (1–3, 5) 50; (4, 6–13) 20.



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Fig. 3. Mixacarus (Phyllolohmannia) pectinatus sp. n., adult: (1) tibia and tarsus of right leg I in dorsal view; (2) femur, genu, and tibia of leg II in dorsal view; (3) genu, tibia, and tarsus of leg III, right, in ventral view; (4) trochanter, femur, genu, and tibia of leg IV, right, in ventral view. Scale 100 µm.

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two setae: chb (77–86) simple, weakly pubescent; cha (8–12) spine-shaped, smooth.

Epimeral area (Figs. 1, 2; 2, 6–8). Epimeral formula 3–1–3–4. Setae 1a, 1b, 2a, 3a, 3b, 4a, 4b, 4c (65–82) simple, pubescent; 3с and 4d (32–49) fine, smooth; 1с (41–53) spindle-shaped, pubescent.

Anogenital area (Figs. 1, 2; 2, 9–13). Anogenital area with 10, 2, and 4 pairs of genital, anal (an1, an2) and adanal (ad1—ad4) setae. Median genital setae (6 pairs) short (57–65), smooth; lateral setae longer (98–114) and thicker, pubescent (excluding pair of smooth anterior lateral setae). Anal setae (69–82) weakly thickened, pubescent. Adanal setae ad1 (164–168) noticeably dilated in median part, pubescent; ad2

(114–131), ad3 (98–110), and ad4 (98–110) weakly dilated and less pubescent. Lyrifissurae iad not found.

Legs (Fig. 3, 1–4). Tarsus with single claw, all claws smooth. Femur with ventral carinae. Formulae of setae and solenidia represented in Table 1, their designations, in Table 2. Majority of setae simple, some outer (on legs I, II), inner (on legs III, IV), and dorsal setae wide, concave-leaf-shaped, rounded dis-tally and tubular at base. Famulus (e) conical, short. Solenidia ω1 on tarsus I, ω on tarsus II, φ on tibia III thickened, blunt distally; other solenidia longer and finer. Solenidia φ on tibiae I and II, σ" on genu I, σ on genua II–IV forming double group with seta d.

Juvenile Stages (Figs. 4–6)

Description. Dimensions. Body size in larva 498–564 (mean value 531, n = 5), protonymph 614–664 (651, n = 4), deutonymph 780–830 (809, n = 4), trito-nymph 962–1062 (1012, n = 4). Body with in larvae

Table 1. Formulae of leg setae and solenidia in Mixacarus (Phyllolohmannia) pectinatus sp. n. at different developmental stages

Stage Formula of setae Formula of solenidia Leg I

Larva 0–2–3–4–14 2–1–1 Protonymph 0–2–3–4–16 2–1–2 Deutonymph 0–5–3–5–16 2–1–2 Tritonymph 0–5–3–5–17 2–1–2 Adult 0–5–3–5–17 2–1–2

Leg II Larva 0–2–3–4–13 1–1–1 Protonymph 0–3–3–4–13 1–1–1 Deutonymph 0–5–3–5–13 1–1 –1 Tritonymph 0–6–3–5–13 1–1–1 Adult 0–6–3–5–13 1–1–1

Leg III Larva 0–2–2–3–12 1–1–0 Protonymph 1–3–2–3–12 1–1–0 Deutonymph 2–3–2–3–12 1–1–0 Tritonymph 2–3–2–3–12 1–1–0 Adult 2–3–2–3–12 1–1–0

Leg IV Protonymph 0–0–0–0–6 0–0–0 Deutonymph 1–2–2–3–11 1–0–0 Tritonymph 2–3–2–3–12 1–0–0 Adult 2–3–2–3–12 1–0–0



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282–332 (298, n = 5), protonymph 398–415 (402, n = 4), deutonymph 498–531 (514, n = 4), tritonymph 664–730 (680, n = 4).

Integument (Fig. 4, 1, 5) semitransparent (in larva) or pale yellowish (in nymphs). Integument similar to that in adult mites: body and integument with densely arranged micro-pits pits (visible only under immersion objective, × 1000), forming polygonal ornament. Pro-dorsum, notogaster, and anogenital area with porous areas virtually indistinct in larva and protonymph. Larval prodorsum with several short transverse folds also present in anterior part of gastronotal area. Post-bothridial (Sb) and notogastral transverse strips (S1–S7) distinct, subsequent stripes weakly distinct.

Prodorsum (Fig. 4, 1, 5) triangular, its length consti-tuting approximately two thirds of length of notogas-tral area. Rostrum rounded. Prodorsal setae weakly (in larva) or significantly (in nymphs) dilated in median part, pubescent. Ratio between length of setae similar in all immature stages: in > ro ~ le ~ exa ~ exp. Trichobothria pectinate, with 9 (in larva) or 7 (in nymphs) setae.

Gastronotal area (Figs. 4, 1, 5, 6; 6, 1–4). Anterior margin distinct, convex. Posterior part rounded. Larva with 14 pairs of simple notochaetae, h4 and rudimental p1–p3 (not included into number of setae in larva) pre-sent; nymphs with 16 pairs of long, dilated in median, median-proximal, or median-distal part of notogastral

Table 2. Changes in leg chaetom of Mixacarus (Phyllolohmannia) pectinatus sp. n. at different developmental stages

Stage Trochanter Femur Genu Tibia Tarsus Leg I

Larva – d, bv" d, (l), σ', σ" (l), v', φ (ft), (tc), (p), (u), (a), s, (pv), e, ω1, Protonymph – – – – m, n, ω2 Deutonymph – (l), v" – v" - Tritonymph – – v' – it' Adult – – – – –

Leg II Larva – d, b, v'' d, (l), σ d, (l), v', φ (ft), (tc), (p), (u), (a), s, (pv), ω Protonymph – – – – – Deutonymph – l1', v'' – v'' – Tritonymph – l2" – – – Adult – – – – –

Leg III Larva – d, ev' d, l', σ d, (l), v', φ (ft), (tc), (p), (u), a', s, (pv) Protonymph v' l' – – – Deutonymph l' – – – – Tritonymph – – – – – Adult – – – – –

Leg IV Protonymph – – – – ft", p',(u),(pv) Deutonymph v' d, ev' d, l', σ d, (l), v' ft', tc', p", a', s Tritonymph l' l' – – tc" Adult – – – – –

Notes: Setae are given only for stages where they appear. Latin letters designate setae [(e) famulus]; Greek letters designate solenidia; (dσ and dφ) Seta and solenidion are united into a common group ; (') inner setae; (") outer setae; pairs of setae are given in paren-theses.

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Fig. 4. Mixacarus (Phyllolohmannia) pectinatus sp. n., juvenile stages: (1) larva, dorsal size; (2) larva, subcapitulum, left part; (3) larva, palpal tibia and tarsus; (4) chelicerae, right part; (5) deutonymph, dorsal side; (6) deutonymph, dorsal side; (7) tritonymph, subcapitulum, left part. Scale (µm): (1) 100; (2, 4, 6, 7) 50; (3) 10; (5) 200.



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setae. Setae h4 and rudimental p4 reduced in proto-nymph; p1–p3 similar in structure to other notogastral setae starting from protonymphal stage. Lyrifissurae ia, im, ip situated similarly to lyrifissurae in other ju-venile stages of Lohmanniidae.

Gnathosoma (Fig. 4, 2–4, 7). Subcapitulum longer than wide. Subcapitular setae simple, smooth in larva and pubescent (excluding smooth setae a) in nymphs. Second pair of setae m2 appearing in protonymphs. Adoral setae structurally similar to setae of adult mites: or1 and or2 leaf-shaped; or3 thickened, setiform, appearing in protonymphs. Palps four-segmented, setal formula 0–0–3–10(+1ω) in larva and protonymph, 0–1–3–10(+1ω) in deuto- and tritonymph. Chelicerae with long, simple, smooth seta chb and very short and smooth seta cha.

Epimeral area (Fig. 5, 1–5). Epimeral formula in larva, protonymph, deutonymph, and tritonymph con-

stituting 3–1–2, 3–1–2–1, 3–1–2–2, and 3–1–3–3, respectively. Setae 1с weakly dilated in median-distal part, 3с and 4с simple and smooth, other setae slightly thicker and pubescent. Claparede’s organ (Cl) with long neck visible in lateral view in larva.

Anogenital area (Fig. 6, 1–4). Genital, adanal, and anal formulae in ontogenesis (from larva to trito-nymph) looking like 0–1–5–8, 0–4–4–4, and 0–0–2–2, respectively. Simple and smooth adanal setae appear-ing in protonymph; starting from stage of deutonymph, adanal setae pubescent (ad1–ad2 dilated in median part); in tritonymph, setae ad1–ad3 dilated. Fine anal setae appearing in deutonymph; these setae becoming pubescent in tritonymph. Genital setae simple, pubes-cent (except for pair of smooth anterior-lateral setae in deuto- and tritonymph). Genital folds without trans-verse suture. Lyrifissurae ih present starting from lar-val stage; ips, since protonymphal stage; setae iad not found.

Fig. 5. Mixacarus (Phyllolohmannia) pectinatus sp. n., juvenile stages: (1) epimeral area of larva; (2) epimeral area of protonymph; (3) epimeral area of deutonymph; (4) epimeral area of tritonymph; (5) Claparede’s organ, lateral view. Scale (µm): (1, 2) 50; (3, 4) 100; (5) 20.

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Legs (Fig. 7, 1–5). Morphology of segments and se-tae similar to that in adult mites. Formulae and desig-nations of leg setae and solenidia in ontogenesis repre-sented in Tables 1 and 2.

Differential diagnosis. In the structure of the nar-row leaf-shaped notogastral setae, Mixacarus (P.) pe-ctinatus sp. n. is similar only to M. (P.) hamanni, known from Java (Indonesia) (Balogh, 1961). The new species, however, is characterized by the absence of strongly dilated and distally rounded setae on the pro-dorsum and in the epimeral area that are present in M. (P.) hamanni; by the presence of pectinate trichobothria, in contrast to the setiform and bilaterally pubescent trichobothria of the latter species; by inter-

rupted stripes on the notogaster (in M. (P.) hamanni, these stripes are continuous and distinct), and by other small characters. All the other representatives of the subgenus are characterized by strongly dilated distally and widely leaf-shaped setae on the prodorsum, sub-capitulum, notogaster, subcapitulum, and in epimeral and anogenital areas.

The juvenile stages of the new species differ from the corresponding stages of M. (P.) tenasserimensis and M. (P.) variatus (Fuangarworn and Chaisuekul, 2011) in the following characters: the absence of strongly dilated distally setae on the body; the pres-ence of 14 and 16 pairs of notogastral setae in nymphs and larvae, respectively (in M. (P.) tenasserimensis

Fig. 6. Mixacarus (Phyllolohmannia) pectinatus sp. n., anogenital area in juvenile stages: (1) larva; (2) protonymph; (3) deutonymph; (4) tritonymph. Scale (µm): (1, 2) 50; (3, 4) 100.



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Fig. 7. Mixacarus (Phyllolohmannia) pectinatus sp. n., juvenile stages: (1) larva, right leg I in dorsal view; (2) larva, left leg II in dorsal view; (3) larva, right leg III in ventral view; (4) protonymph, femur of right leg II in ventral view; (5) tritonymph, genu of left leg III in ventral view. Scale 50 µm.

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and M. (P.) variatus, these setae constitute 13 and 17 pairs, respectively); the epimeral formulae of deuto- (3–1–2–2) and tritonymphs (3–1–3–3) (in M. (P.) tenasserimensis and M. (P.) variatus, these formula look like 3–1–3–3 and 3–1–3–4, respectively).

Etymology. The species name pectinatus points to pectinate trichobothria of the new species.

ACKNOWLEDGMENTS

We are grateful to U.Ya. Shtanchaeva (Universidad Complutense de Madrid, Madrid, Spain) for consulta-tions, critical review of the manuscript, and valuable comments, and to L. Dembicky and O. Sausa for col-lecting of the material.

REFERENCES 1. Balogh, J., “An Outline of the Family Lohmanniidae

Berlese, 1916 (Acari: Oribatei),” Acta Zool. Acad. Sci. Hungar. 7 (1–2), 19–44 (1961).

2. Balogh, J. and Balogh, P., “A New Outline of the Fam-ily Lohmanniidae Berlese, 1916 (Acari: Oribatei),” Acta Zool. Hungar. 33 (3–4), 327–398 (1987).

3. Balogh, J. and Balogh, P., The Oribatid Mites Genera of the World. Vol. 1 (Hungar. Nat. Mus., Budapest, 1992).

4. Ermilov, S.G., Shtanchaeva, U.Ya., Subías, L.S., and Anichkin, A.E., Morphology of Juvenile Instars of Meristacarus sundensis Hammer, 1979 and Cryptacarus promecus Grandjean, 1950 (Acari, Oribatida, Lohman-niidae),” Syst. Appl. Acarol. 17 (3), 281–300 (2012).

5. Fuangarworn, M. and Chaisuekul, C., “Two New Spe-cies of the Oribatid Mite Subgenus Phyllolohmannia (Oribatida: Lohmanniidae: Mixacarus) from Thailand,” Intern. J. Acarol. 37 (1), 114–128 (2011).

6. Norton, R.A., “Systematic Relationships of Lohmannii-dae (Acarina: Oribatei), in Trends of Acarology: Pro-ceedings of the 12th International Congress (Dordrecht, 2010), pp. 9–16.

7. Norton, R.A. and Behan-Pelletier, V.M., “Oribatida,” in A Manual of Acarology (TX) (Texas Tech Univ., Lub-bock, 2009), pp. 430–564.

8. Subías, L.S., “Listado sistemático, sinonímico y bio-biogeográfico de los á caros oribátidos (Acariformes: Oribatida) del mundo (excepto fosiles),” Graellsia 60 (número extraordinario), 3–305 (2004). Online version accessed in February 2011; http://www.ucm.es/info/zoo/ Artropodos/Catalogo.pdf.

9. Subías, L.S., Shtanchaeva, U.Ya., and Arillo, A., “Lis-tado de los á caros oribátidos (Acariformes, Oribatida) de las diferentes regions biogeográficas del mundo,” Monograf. electron. S.E.A. 4, 1–815 (2012).

mixacarus(phyllolohmannia)pectinatussp. n. (acari, oribatida, lohmanniidae), a new species of...

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