Acari, Oribatida, Otocepheidae - BioOne

Systematic & Applied Acarology 21(2): 209–217 (2016) http://doi.org/10.11158/saa.21.2.5 Article

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

http://zoobank.org/urn:lsid:zoobank.org:pub:F2B922B3-F0E4-449F-8EE6-FEEF4BF711B1

A new species of Pseudotocepheus (Acari, Oribatida, Otocepheidae) from Chile, with a key to species of the genus from the Neotropical region and discussion on taxonomic status of Constrictocepheus SERGEY G. ERMILOV Tyumen State University, Tyumen, Russia. E-mail: [email protected]

Abstract A new species of oribatid mite of the genus Pseudotocepheus (Oribatida, Otocepheidae) is described from Sphagnum-mosses on swamps in the vicinity of Punta Arenas of Chile. Pseudotocepheus puntaarenasensis sp. nov. is morphologically most similar to P. sexdentatus (Trägårdh, 1931), however, it differs from the latter by the smaller body size, parallel costulae basally, interlamellar setae distanced from costulae and similar length of rostral and lamellar setae. The taxonomic status of the genus Constrictocepheus Grobler, 1998 is discussed; it is considered as a junior synonym of Pseudotocepheus. The initial systematic placement of eight species of Pseudotocepheus, which has been combined with Constrictocepheus, is supported. An identification key to all known species of Pseudotocepheus from the Neotropical region is given. Key words: oribatid mites, new species, Pseudotocepheus, Constrictocepheus, systematics, new synonym, key, mosses, Chile

Introduction At present, the oribatid mite fauna (Acari, Oribatida) of Chile is poorly known (Trägårdh 1931; Hammer 1962; Covarrubias 1986, 2009; Martínez & Casanueva 1999; Olszanowski & Norton 2002; Colloff 2009; Ermilov & Pešić 2011). In the course of taxonomic identification of oribatid mites from Sphagnum-mosses in the vicinity of Punta Arenas of Chile, we found one new species, belonging to the genus Pseudotocepheus Balogh, 1960 (family Otocepheidae—see Norton & BehanPelletier 2009; Schatz et al. 2011). The main goal of this paper is to describe and illustrate this species. Pseudotocepheus (=Nesotocepheus Hammer, 1972) is a genus of oribatid mites that was proposed by Balogh (1960) with Pseudotocepheus paulinai Balogh, 1960 as type species. It comprises about 40 species (see different opinions in Grobler 1998; Balogh & Balogh 2002; Subías 2004, updated 2015), which are distributed in the tropics and subtropics (Subías 2004, updated 2015). The main generic characters are (based on Balogh 1960; Balogh & Balogh 1992; including our additions): body of normal length (as in most Otocepheidae); pedotecta I and II represented by small laminae; prodorsal and notogastral condyles present or sometimes absent; medial notogastral condyles never fused in one medial unpaired condyle; 10 pairs of setiform or straight, thickened notogastral setae; three pairs of genital, one pair of aggenital, two pairs of anal and three pairs of adanal setae; aggenital condyles not developed; location of adanal setae ad3 and adanal lyrifissures varied. The identification keys to selective species of Pseudotocepheus were given by Balogh & Balogh (1992) and Grobler (1998). Below, we present an identification key to all known Neotropical species of this genus. © Systematic & Applied Acarology Society

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Also, we discuss the taxonomic status of the otocepheid genus Constrictocepheus Grobler, 1998, which is morphologically very similar to Pseudotocepheus.

Materials and methods The collection locality and habitat for the new species are given in the "Material examined" section. Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. The body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the ventral plate. Notogastral width refers to the maximum width in dorsal aspect. Lengths of body setae were measured in lateral aspect. All body measurements are presented in micrometers. Formulas for leg setation are given in parentheses according to the sequence trochanter–femur– genu–tibia–tarsus (famulus included). Formulas for leg solenidia are given in square brackets according to the sequence genu-tibia-tarsus. General terminology used in this paper follows that of Wallwork (1962), Aoki (1967), and Norton & Behan-Pelletier (2009). Drawings were made with a drawing tube using a Carl Zeiss transmission light microscope “Axioskop-2 Plus”.

Description of Pseudotocepheus puntaarenasensis sp. nov. (Figs 1–11) Diagnosis Body size: 464–564 × 199–249. Costulae not reaching the insertions of lamellar setae. Rostral and lamellar setae setiform, barbed unilaterally. Interlamellar, notogastral, anal and adanal setae thickened, straight, slightly barbed. Bothridial setae short, clavate, barbed. Exobothridial setae minute. All prodorsal and notogastral condyles present, rounded distally. Notogaster with concavity posterior to setae c. Epimeral, genital and aggenital setae setiform. A pair of strong longitudinal ridges present laterally to genital aperture. Lyrifissures iad in inverse apoanal position. Leg seta l’ on trochanters III long, thick, heavily barbed. Description Measurements. Body length: 514 (holotype, male), 464–498 (five paratypes, all males), 564 (one paratype, female); notogaster width: 232 (holotype, male), 199–232 (five paratypes, all males), 249 (one paratype, female). Integument. Body color light brown. Body surface with slightly visible microfoveolae (their diameter up to 2), covered by dense microgranular cerotegument. Lateral body sides (between lateral condyles and acetabula II) with tubercles (tub, their diameter up to 6). Prodorsum. Rostrum broadly rounded. Costulae (cos) half as long as prodorsum, parallel, not reaching the insertions of lamellar setae, located dorso-laterally. Rostral (ro) and lamellar (le) setae similar in length (53–61), setiform, curving antero-medially, barbed unilaterally. Interlamellar setae (in, 28–36) thickened, straight, slightly barbed. Bothridial setae (bs, their length out of bothridia 28– 36) clavate, with short stalk (8–12) and longer (20–24) head rounded distally, barbed. Exobothridial setae (ex, 6–8) minute, thin, smooth. Lateral carinae (car) short, slightly visible. Medial prodorsal condyles (co.pm) tubercle-like, located separately. Lateral prodorsal condyles (co.pl) strong, triangular, rounded distally. Notogaster. Medial (co.nm) and lateral (co.nl) notogastral condyles tubercle-like, rounded distally. Medial condyles separate. Distance between condyles co.nm–co.nm slightly shorter than co.pm–co.pm. Notogaster with 10 pairs of setae of medium length (c, 53–57; la, lm, lp, h2, h3, 41– 210

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49; h1, p1–p3, 28–36), thickened, straight, slightly barbed. Small concavity (con) present posteriorly to setae c. Lyrifissures (ia, im, ip, ih, ips) and opisthonotal gland openings (gla) distinct.

FIGURES 1–2. Pseudotocepheus puntaarenasensis sp. nov.: 1 — dorsal view (legs except left trochanter III not illustrated); 2 — ventral view (legs except trochanters IV not illustrated). Scale bar 100 μm.

Gnathosoma. Subcapitulum longer than wide (94–102 × 69–77). Subcapitular setae setiform, slightly barbed; h (36–41) longer, thicker and more densely barbed than a (18–20) and m (28–32). Adoral setae and their alveoli absent. Palps (57–61) with setation 0-2-1-3-8(+ω). Solenidion of each palp half as long as palptarsus, pressed to surface of the latter, connected to distal seta in anterior part. Chelicerae (98–102) with barbed setae; cha (32–36) longer than chb (14–16). Trägårdh’s organ (Tg) tapered. 2016

ERMILOV: A NEW SPECIES OF PSEUDOTOCEPHEUS (OTOCEPHEIDAE) FROM CHILE

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Epimeral and lateral podosomal regions. Epimeral setal formula: 3-1-3-3. All setae setiform, thin, slightly barbed. Setae 1a, 1c, 2a, 3a, 4a, 4b, 4c (18–28) shorter than 1b, 3b, 3c (36–41). Pedotecta I (Pd I) and II (Pd II) well developed, represented by small lamina. Discidia (dis) triangular, broadly rounded distally.

FIGURES 3–7. Pseudotocepheus puntaarenasensis sp. nov.: 3 — proterosoma, lateral view (legs I, II not illustrated); 4 — medio-posterior part of hysterosoma, lateral view; 5 — subcapitulum, left half, ventral view; 6 — palp, right, antiaxial view; 7 — chelicera, left, paraxial view. Scale bars 100 μm (3, 4), 50 μm (5–7).

Anogenital region. A pair of strong longitudinal ridges (r) present laterally to genital aperture. Three pairs of genital setae (g1–g3, 16) thin, smooth. One pair of aggenital setae (ag, 20–24) setiform, slightly barbed. Three pairs of adanal (ad1, ad2, 24–28; ad3, 28–32) and two pairs of anal (an1, an2, 20–24) setae thickened, straight, slightly barbed. Adanal setae ad3 located in preanal position. Distance between setae ad3–ad3 similar to ad2–ad2 and longer than ad1–ad1. Lyrifissures iad in inverse apoanal position. 212


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Legs. Claw of each tarsus indistinctly serrate on dorsal side. Tarsi without teeth, but one to two small tubercles present. Formulas of leg setation and solenidia: I (1-4-3-4-16) [1-2-2], II (1-4-3-315) [1-1-2], III (2-3-0-2-15) [1-1-0], IV (1-2-1-2-12) [0-1-0]; homology of setae and solenidia indicated in Table 1. Famulus short, indistinctly dilated distally, slightly triangular basally. Leg setae barbed (except p on tarsi I and s on tarsi I, II smooth). Seta l’ on trochanters III long, thick, heavily barbed. Leg setae u setiform on all tarsi. Solenidia φ1 and φ2 on tibiae I, and φ on tibiae II setiform, other solenidia thickened, blunt-ended distally.

FIGURES 8–11. Pseudotocepheus puntaarenasensis sp. nov.: 8 — leg I, right, antiaxial view; 9 — genu, femur and trochanter of leg II, right, antiaxial view; 10 — leg III, without tarsus, right, antiaxial view; 11 — leg IV, right, antiaxial view. Scale bars 50 μm. 2016


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TABLE 1. Leg setation and solenidia of Pseudotocepheus puntaarenasensis sp. nov. Leg

Trochanter

Femur

Genu

Tibia

Tarsus

I

v'

d, (l), bv''

(l), v', σ

(l), (v), φ1, φ2

(ft), (tc), (it), (p), (u), (a), s, (pv), ɛ, ω1, ω2

II

v'

d, (l), bv''

(l), v', σ

l', (v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv), ω1, ω2

III

l', v'

d, l', ev'

σ

(v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv)

IV

v'

d, ev'

d

(v), φ

ft'', (tc), (p), (u), (a), s, (pv)

Roman letters refer to normal setae, Greek letters to solenidia (except ɛ = famulus). Single prime (') marks setae on anterior and double prime (") setae on posterior side of the given leg segment. Parentheses refer to a pair of setae.

Material examined Holotype (male) and six paratypes (one female and five males): Chile, vicinity of Punta Arenas, 53о38’02.8”S, 65o31’01.6”W, swamp, in Sphagnum sp., 18 November 2014, collected by V.A. Stolbov. Type deposition The holotype is deposited in the collection of the Senckenberg Institution Frankfurt, Germany; six paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. Etymology The specific name puntaarenasensis refers to the vicinity of Punta Arenas, where the new species was collected. Remarks In having the costulae not reaching the insertions of lamellar setae, short and clavate bothridial setae, all prodorsal and notogastral condyles present, and thickened and straight interlamellar and notogastral setae, Pseudotocepheus puntaarenasensis sp. nov. is most similar to P. sexdentatus (Trägårdh, 1931) from Chile. However, it differs from the latter by the smaller body size (464–564 × 199–249 vs. 890 × 420), parallel costulae basally and interlamellar setae distanced from costulae (vs. costulae slightly convergent basally and interlamellar setae inserted near to costulae), and similar length of rostral and lamellar setae (vs. lamellar setae half as long as rostral setae). Discussion on taxonomic status of Constrictocepheus Grobler, 1998 Grobler (1998) proposed a new genus, Constrictocepheus with Pseudotocepheus geminatus Balogh & Mahunka, 1969 as type species, and combined eight species with this genus from Pseudotocepheus. The main generic diagnostic characters were listed: 1) discidia (as pedotecta IV in Grobler 1998) broad, visible in dorsal view (vs. smaller, not visible in dorsal view in Pseudotocepheus); 2) prodorsum narrowed basally, notogaster narrowed anteriorly (vs. widened in Pseudotocepheus); 3) lamellae and bothridia located dorsally on prodorsum (vs. dorso-laterally in Pseudotocepheus); 4) medial prodorsal condyles usually represented by one pair (vs. represented by one pair or tubercles in Pseudotocepheus); 5) medial prodorsal and notogastral condyles—if present —connected (vs. well separated in Pseudotocepheus). In our opinion, these listed characters cannot be used as diagnostic traits for the independence of Constrictocepheus. The reasons are as follows: 1) The visibility or invisibility of discidia in dorsal view, morphology of basal part of prodorsum and anterior part of notogaster, localization of lamellae and bothridia on prodorsum, 214


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localization of medial prodorsal and notogastral condyles are species characters, because other otocepheid genera encompass species with these different traits (for example, such selective traits present in Dolicheremaeus, Eurostocepheus, Fissicepheus, Megalotocepheus, Otocepheus, Papillocepheus). 2) We also noted that different sizes of discidia (broad in Constrictocepheus vs. smaller in Pseudotocepheus) are probably illusory, because it is caused by the width of the anterior part of the notogaster. If notogaster narrows anteriorly, discidia are well visible in dorsal aspect as large (broad) structures, but if the notogaster is widened anteriorly, discidia are not visible or hardly visible in dorsal aspect and seen as smaller structures. 3) The presence of one pair of medial prodorsal condyles is not useful, because it cannot distinguish Constrictocepheus from Pseudotocepheus, species of the latter genus also sometimes have one pair of these condyles. Hence, if some other genera of Otocepheidae include representatives with visible or invisible discidia in dorsal view, narrowed or widened prodorsum basally and notogaster anteriorly, localization of lamellae and bothridia dorsally and dorso-laterally on prodorsum, there is no sense to use these characters to distinguish the genus Constrictocepheus from Pseudotocepheus. Therefore, based on our observations, we propose the following taxonomic change: Constrictocepheus Grobler, 1998 is a junior subjective synonym of Pseudotocepheus Balogh, 1960 (syn. nov.). Also, we support the initial systematic placement of eight species of Pseudotocepheus (P. coarctatus P. Balogh, 1985; P. curtisetus Hammer, 1966; P. geminatus Balogh & Mahunka, 1969; P. gobletus Chakrabarti, Mondal & Kundu, 1978; P. monteithi J. Balogh & P. Balogh, 1983; P. orientalis Mondal & Kundu, 1984; P. punctatus Hammer, 1966; P. tenuisetus Hammer, 1966), which was combined with Constrictocepheus by Grobler (1998). Thus, the genus Pseudotocepheus currently comprises 43 species, which are distributed in the tropics and subtropics (Australian, Ethiopian, Neotropical and Oriental regions) (Subías 2004, updated 2015). Key to species of Pseudotocepheus from the Neotropical region 1. Bothridial setae rod-like; medial prodorsal and medial notogastral condyles connected medially; body size: 1040 × 549 . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudotocepheus geminatus Balogh & Mahunka, 1969. Brazil - Bothridial setae with well-developed head; medial prodorsal and medial notogastral condyles separated medially (if present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Bothridial setae with stalk shorter than rounded distally head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 - Bothridial setae with stalk longer than fusiform or lanceolate head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3. Costulae reaching insertions of lamellar setae; body size: 777–864 × 341–415 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudotocepheus australis (Mahunka, 1980). Neotropical region - Costulae distinctly not reaching insertions of lamellar setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Costulae parallel basally; rostral and lamellar setae similar in length; body size: 464–564 × 199–249 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudotocepheus puntaarenasensis sp. nov. Chile - Costulae slightly convergent basally; lamellar setae half as long as rostral setae; body size: 890 × 420 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudotocepheus sexdentatus (Trägårdh, 1931). Chile 5. Medial prodorsal condyles absent; basal part of prodorsum tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . 6 - Medial prodorsal condyles present; basal part of prodorsum not tuberculate . . . . . . . . . . . . . . . . . . . . . . . 7 6. Medial notogastral condyles absent; posterior notogastral setae long, with curved, flagellate tip; body size: 556–794 × 238–375 … Pseudotocepheus amonstruosus Mahunka, 1973 (see also Balogh & Mahunka 1978; Mahunka 1978; Pérez-Íñigo & Baggio 1980; Grobler 1997). Ethiopian and Neotropical regions and Ceylon - Medial notogastral condyles present; posterior notogastral setae of medium size, straight, without flagellate tip; body length: 900–1325 … Pseudotocepheus andinus (Fernandez, Martinez & Eguaras 1990). Argentina 7. Medial notogastral condyles absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 - Medial notogastral condyles present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 2016


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8. Lateral notogastral condyles absent; costulae indistinctly developed; body size: 623 × 267 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudotocepheus sturmi P. Balogh, 1984. Colombia - Lateral notogastral condyles present; costulae strong, well visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Costulae reaching insertions of lamellar setae; adanal setae ad3 in paraanal position; body size: 745–950 × 317–420 . . . . . . . . . . . . . . . . . . . . . . . . .Pseudotocepheus sexidimorphus (Vasiliu & Călugăr, 1977). Cuba - Costulae distinctly not reaching insertions of lamellar setae; adanal setae ad3 in preanal position; body length: 1060 . . . . . . . . . . . . . . . . . . . . Pseudotocepheus punctatus Hammer, 1966. New Zealand and Chile 10. Posterior notogastral setae long, with curved, flagellate tip; body size: 720 × 356 . . . . . . . . . . . . . . . . . . . . . . . Pseudotocepheus pauliensis Pérez-Íñigo & Baggio 1993 (see also Grobler 1997). Neotropical region - Posterior notogastral setae of medium size, straight, without flagellate tip . . . . . . . . . . . . . . . . . . . . . . . 11 11. Costulae reaching insertions of lamellar setae; body size: 505–603 × 201–254 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudotocepheus hauseri (Mahunka, 1980). Neotropical region - Costulae distinctly not reaching insertions of lamellar setae; body size: 880–900 × 340–370 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudotocepheus curtipilus (Trägårdh, 1931). Chile

Acknowledgements The author thanks Dr. Elizabeth A. Hugo-Coetzee (National Museum, Bloemfontein, South Africa) and two anonymous reviewers for the valuable comments, and Dr. V.A. Stolbov, I.V. Kuzmin, S.A. Ivanov and R.V. Latyntcev (Tyumen State University, Tyumen, Russia) for collecting mites from Chile. The present study was supported by the Russian Science Foundation (project 14-14-01134).

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