Зоологический журнал, 2020, T. 99, № 12, стр. 1375-1385

A New Species of Pseudotocepheus Balogh 1960 from Chile and a Redescription of Leptotocepheus sexidimorphus (Vasiliu et Călugăr 1977) comb. n. from Cuba (Acari, Oribatida, Otocepheidae)

S. G. Ermilov *

Tyumen State University
625003 Tyumen, Russia

* E-mail: ermilovacari@yandex.ru

Поступила в редакцию 28.05.2019
После доработки 28.07.2019
Принята к публикации 4.08.2019

Полный текст (PDF)

Аннотация

A new oribatid mite species of the family Otocepheidae is described from Chile. Pseudotocepheus parafoveolatus Ermilov sp. n. differs from P. foveolatus Hammer 1966 in the presence of stiff interlamellar setae, as well as stiff medium-sized notogastral setae lm, lp, h3 and p3; and the absence of a foveolate body surface. A detailed supplementary description of the otocepheid species, Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977) comb. n., newly transferred from the genus Pseudotocepheus, is provided. This redescription is based on specimens from Cuba.

Keywords: otocepheid mites, Pseudotocepheus, Leptotocepheus (Longocepheus), systematics, morphology, Neotropical region

This work is based on a random set of previously unstudied oribatid mites (Acari, Oribatida) of the family Otocepheidae from the Neotropical region, which were stored in the collections of the Tyumen State University Museum of Zoology (Tyumen, Russia). In the course of taxonomic identification, I found two species of the genus Pseudotocepheus Balogh 1960. One of these species (from Chile) is new to science. The other species (from Cuba) is Pseudotocepheus sexidimorphus (Vasiliu et Călugăr 1977) (it is known from this country only).

The primary goal of the paper is to describe and illustrate a new species under the name Pseudotocepheus parafoveolatus Ermilov sp. n. Pseudotocepheus was proposed by Balogh (1960), with Pseudotocepheus paulinai Balogh 1960 as type species. The genus comprises 45 species, which are distributed in the Australasian, Ethiopian and Neotropical regions (Subías, 2019). The main generic traits were summarized by Ermilov (2016). The identification keys to some species of Pseudotocepheus were presented by Grobler (1998), Balogh and Balogh (2002), Ermilov (2016), and Ermilov and Minor (2019).

The secondary goal of the paper is to discuss the systematic placement of P. sexidimorphus: namely, to transfer it to the genus Leptotocepheus Balogh 1961 (subgenus Longocepheus Balogh et Mahunka 1966) and to present its supplementary description (this species has been briefly described and poorly figured). In particular, this paper presents new information about some morphological structures and provides their measurements, identifies leg setae and solenidia, as well as describes the morphology of gnathosoma.

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 notogaster. Notogastral width refers to the maximum 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. Morphological terminology used in this paper follows that of F. Grandjean: see Travé and Vachon (1975) for references, Norton (1977) for leg setal nomenclature, and Norton and Behan–Pelletier (2009), for overview. Drawings were made with a camera lucida attached to the transmission light microscope, Leica DM 2500.

The following abbreviations are used: cos – costula; ppc – posterolateral prodorsal carinae; ro, le, in, bs, ex – rostral, lamellar, interlamellar, bothridial and exobothridial setae, respectively; cpm, cpl – medial and lateral prodorsal condyles, respectively; cnm, cnl – medial and lateral notogastral condyles, respectively; c, la, lm, lp, h, p – notogastral setae; ia, im, ip, ih, ips – notogastral lyrifissures; gla – opisthonotal gland opening; a, m, h – subcapitular setae; v, l, d, cm, acm, ul, sul, vt, lt – palp setae; ω – palp and leg solenidion; cha, chb – cheliceral setae; Tg – Trägårdh’s organ; Pd I, Pd II – pedotecta I, II, respectively; aps – sternal apodeme; 1a, 1b, 1c, 2a, 3a, 3b, 3c, 4a, 4b, 4c – epimeral setae; dis – discidium; g, ag, an, ad – genital, aggenital, anal and adanal setae, respectively; iag – aggenital lyrifissure; iad – adanal lyrifissure; cvr – circumventral ridge; σ, φ – leg solenidia; ɛ – leg famulus; v, ev, bv, l, d, ft, tc, it, p, u, a, s, pv – leg setae.

Pseudotocepheus parafoveolatus Ermilov sp. n. (Figs 1–3)

Fig. 1.

Pseudotocepheus parafoveolatus Ermilov sp. n., adult: a – dorsal view (legs not shown), b – ventral view (gnathosoma and legs not shown). Scale bar 100 µm.

Fig. 2.

Pseudotocepheus parafoveolatus Ermilov sp. n., adult: a – anterior part of body, lateral view (legs not shown); b – posterior part of body, lateral view; c – subcapitulum, ventral view; d – palp, right, antiaxial view; e – chelicera, right, antiaxial view. Scale bar (µm): a, b – 100; c, e – 50; d – 20.

Fig. 3.

Pseudotocepheus parafoveolatus Ermilov sp. n., adult: a – leg I, without trochanter, right, antiaxial view; b – femur and genu of leg II, right, antiaxial view; c – trochanter, femur and genu of leg III, left, antiaxial view; d – leg IV, right, antiaxial view. Scale bar 50 µm.

Material. The holotype (♂) and 2 paratypes (2♂♂): Chile, Región de Magallanes y de la Antártica Chilena, Provincia de Magallanes, 30 km south-east of Punta Arenas, before Laguna Parrillar National Park, 53°23′00″ S, 071°13′34″ W, swamp, moss Sphagnum magellanicum, Berlese funnel extraction, 10.XI.2014 (V.A. Stolbov, S.A. Ivanov).

The holotype is deposited in the collection of the Senckenberg Institute, Görlitz, Germany; two paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All in ethanol with drop of glycerol.

Diagnosis. Body size: 780–879 × 348–381. Body ratio (length/width): 2.2–2.3. Body surface microgranulate. Costulae distinctly not reaching insertions of lamellar setae. Tutoria present. Rostral and lamellar setae long, setiform, barbed. Interlamellar setae long, stiff, barbed. Exobothridial setae minute. Bothridial setae short, clavate. Lateral and medial prodorsal condyles and lateral notogastral condyles tubercle-like; medial notogastral condyles very small. Notogaster with 10 pairs of barbed setae; lm, lp, h3 and p3 of medium size, stiff, others very long, flagellate. Epimeral setal formula: 3–1–3–3. Epimeral and anogenital setae setiform, barbed. Adanal setae ad3 in anterolateral position. Adanal lyrifissures removed from anal aperture. Leg setae u setiform on all tarsi.

Description. Measurements. Body length: 780 (holotype), 795, 879 (paratypes); notogaster width: 348 (holotype), 348, 381 (paratypes). Body ratio (length/width): 2.2–2.3.

Integument (Figs 1a, 1b; 2a, 2b). Body color light brown to brown, but genital plates and legs dark brown. Body surface granulate (diameter of granules less than 1). Foveolate ornamentation absent. Lateral parts of body between bothridia and pedotecta I–III densely tuberculate (diameter of tubercles up to 10).

Prodorsum (Figs 1a, 2a). Rostrum broadly rounded. Costulae lineate, distinctly not reaching insertions of lamellar setae. Anterolateral prodorsal carinae absent, posterolateral prodorsal carinae present. Tutoria well developed. Rostral (102–110) and lamellar (110–118) setae setiform, barbed, curving anteromedially. Transverse ridge located anterior to insertions of lamellar setae. Interlamellar setae (106–114) stiff, barbed. Exobothridial setae (2) very short, setiform, smooth. Bothridial setae (45–49) with short stalk and longer, clavate, roughened head. Lateral and medial prodorsal condyles tubercle-like, located separately.

Notogaster (Figs 1a, 1b, 2a, 2b). Anterior notogastral margin straight. Lateral notogastral condyles tubercle-like, distinctly distanced from each other. Medial notogastral condyles present, but very small, poorly developed. Dorsal circummarginal ridge not visible. Notogaster with 10 pairs of barbed setae; lm, lp, h3, and p3 (149–182) stiff, others (298–365) flagellate. Lyrifissures distinct; ia located posterolateral to c, im posterolateral to lm, ip between p2 and p3, ips lateral to h3, ih anterior to h3. Opisthonotal gland openings located posterolateral to lm and anterior to im.

Gnathosoma (Figs 2c–2e). Subcapitulum longer than wide (164–168 × 102–106). Subcapitular setae setiform, barbed, m and h (49–61) longer than a (24–32). Adoral setae and their alveoli absent. Palps (90–98) with setation 0–2–1–3–8(+ω). Postpalpal setae (10) spiniform. Chelicerae (168–172) with two setiform, barbed setae, cha (61) longer than chb (28–32). Trägårdh’s organ of chelicerae narrowly triangular.

Epimeral and lateral podosomal regions (Figs 1b, 2a). Unpaired sternal apodeme reduced to small part. Epimeral setal formula: 3–1–3–3. Setae setiform, barbed, 1a, 2a, 3a, and 4b (20–24) shorter than 1c, 4a and 4c (32–36) and 1b, 3b and 3c (65–73). Pedotecta I represented by small lamina. Discidia elongate triangular, rounded distally.

Anogenital region (Figs 1b, 2a, 2b). Aggenital lyrifissures located lateral to genital aperture. Three pairs of genital (24–28), one pair of aggenital (57–65), three pairs of adanal (69–77), and two pairs of anal (an1, 41–49; an2, 61–65) setae setiform, barbed. Adanal setae ad1 in posterior, ad2 in lateral, ad3 in anterolateral positions. Adanal lyrifissures removed from anal aperture. Circumventral ridge present.

Legs (Figs 3a–3d). Claw of each leg strong, slightly barbed on dorsal side. Dorsoparaxial porose area on femora I–IV and on trochanters III, IV poorly visible. Tarsi I–IV with one pair of small teeth. Femora rounded ventroanteriorly. Formulas of leg setation and solenidia: I (1–4–3–4–16) [1–2–2], II (1–4–3–3–15) [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. Leg setae u setiform on all tarsi.

Table 1.  

Leg setation and solenidia of Pseudotocepheus parafoveolatus Ermilov sp. n.

Leg Tr Fe Ge Ti Ta
 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 (except ɛ – famulus), Greek letters refer to solenidia. Single apostrophe (') marks setae on the anterior, and double apostrophe ('') – setae on the posterior sides of a given leg segment. Parentheses refer to a pair of setae.

Remarks. Pseudotocepheus parafoveolatus sp. n. is morphologically most similar to Pseudotocepheus foveolatus Hammer 1966 from New Zealand (Hammer, 1966) in having short, clavate bothridial setae and some long, flagellate notogastral setae. The former species differs from the latter in the presence of stiff interlamellar setae and stiff notogastral setae lm, lp, h3 and p3 of medium size (versus these setae being long and flagellate in P. foveolatus). Also, the body of the new species does not have a foveolate body surface (versus the body being foveolate in P. foveolatus).

Etymology. The specific name parafoveolatus refers to the similarity between the new species and Pseudotocepheus foveolatus Hammer 1966.

Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977) comb. n. (Figs 4–6)

Fig. 4.

Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977), adult: a – dorsal view (legs not shown); b – ventral view (gnathosoma and legs not shown). Scale bar 100 µm.

Fig. 5.

Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977), adult: a – anterior part of body, lateral view (legs not shown); b – posterior part of body, lateral view; c – subcapitulum, ventral view; d – palp, right, antiaxial view; e – chelicera, left, paraxial view. Scale bar (µm): a, b – 100; c, e – 20; d – 10.

Fig. 6.

Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977), adult: a – leg I, without trochanter, right, antiaxial view; b – femur and genu of leg II, right, antiaxial view; c – trochanter, femur and genu of leg III, left, antiaxial view; d – leg IV, right, antiaxial view. Scale bar 50 µm.

Material. Sixteen specimens (8♂♂, 8♀♀): Cuba, Pinar del Rio Province, Baños de San Juan, Las Terrazas, 22°49′24.00″ N, 82°55′36.00″ W, leaf litter in tropical forest (date and collector unknown; collection of the Tyumen State University Museum of Zoology, Tyumen, Russia).

Supplementary description. Measurements. Body length: 464–531 (males), 581–647 (females); notogaster width: 182–215 (males), 249–282 (females). Body ratio (length/width): 2.2–2.4.

Integument (Figs 4a, 4b, 5a, 5b). Body color light brown, but genital plates and legs dark brown. Body surface granulate (diameter of granules less than 1) and foveolate (diameter of foveoles up to 8), foveoles on prodorsum larger than on notogaster. Lateral parts of body between bothridia and pedotecta I–III densely tuberculate (diameter of tubercles up to 6).

Prodorsum (Figs 4a, 5a). Rostrum broadly rounded. Costulae lineate, slightly not reaching insertions of lamellar setae. Anterolateral and posterolateral prodorsal carinae and tutoria absent. Rostral (45–49 in males; 53–57 in females) and lamellar (49–53 in males; 61–65 in females) setae setiform, barbed, curving anteromedially. Interlamellar setae (41–45 in males; 49–53 in females) stiff, barbed. Exobothridial setae (16 in males; 20 in females) setiform, slightly barbed. Bothridial setae (57–61 in males; 69–73 in females) with long stalk and shorter, fusiform, roughened head; sometimes heads in females narrower than in males. Lateral and medial prodorsal condyles tubercle-like, located separately.

Notogaster (Figs 4a, 5a, 5b). Anterior notogastral margin straight. Lateral notogastral condyles tubercle-like, distinctly distanced from each other. Unpaired medial notogastral condyle broad, without fusions, distinctly developed or, rarely, reduced to simple ridge. Dorsal circummarginal ridge not visible. Notogaster with 10 pairs of stiff, barbed setae (36–41 in males; 53–61 in females). Setae h2 specifically directed lateral. Lyrifissures distinct; ia located anterolateral to c, im posterolateral to la, ip between p2 and p3, ips between h3 and p3, ih anterior to h3. Opisthonotal gland openings located anterolateral to la.

Gnathosoma (Figs 5c–5e). Subcapitulum longer than wide (94–98 × 77–82 in males; 102–110 × 86–90 in females). Subcapitular setae setiform, barbed, m and h (28 in males; 32 in females) longer than a (16 in males; 20 in females). Adoral setae and their alveoli absent. Palps (49–53 in males; 57–61 in females) with setation 0–2–1–3–8(+ω). Postpalpal setae (6–8) spiniform. Chelicerae (102–110 in males; 114–123 in females) with one (cha) setiform, barbed seta (30–32 in males; 36 in females), chb not found. Trägårdh’s organ of chelicerae narrowly triangular.

Epimeral and lateral podosomal regions (Figs 4b, 5a). Unpaired sternal apodeme reduced to small part. Epimeral setal formula: 3–1–3–3. Setae setiform, slightly barbed, 1a, 2a, 3a and 4b (16 in males; 20 in females) shorter than 1c, 4a and 4c (20–24 in males; 24–32 in females) and 1b, 3b and 3c (32–36 in males; 41–45 in females). Pedotecta I represented by small lamina. Discidia elongate triangular, rounded distally.

Anogenital region (Figs 4b, 5a, 5b). Aggenital lyrifissures located lateral to genital aperture. Three pairs of genital (g1, 16; g2, g3, 12 in males; g1, 20; g2, g3, 16 in females), one pair of aggenital (36–41 in males; 45–49 in females), three pairs of adanal (41–45 in males; 49–53 in females) and two pairs of anal (32–36 in males; 41–45 in females) setae setiform, barbed. Adanal setae ad1 in posterior, ad2 and ad3 in lateral positions. Adanal lyrifissures removed from anal aperture. Circumventral ridge present.

Legs (Figs 6a–6d). Claw of each leg strong, slightly barbed on dorsal side. Dorsoparaxial porose area on femora I–IV and on trochanters III, IV poorly visible. Tarsi I–IV with one pair of small teeth. Femora pointed ventroanteriorly. Formulas of leg setation and solenidia: I (1–4–3–4–16) [1–2–2], II (1–4–3–3–15) [1–1–2], III (2–2–1–2–15) [1–1–0], IV (1–2–2–2–12) [0–1–0]; homology of setae and solenidia indicated in Table 2. Leg setae u setiform on tarsi I and thorn-like on tarsi II–IV.

Table 2.  

Leg setation and solenidia of Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977)

Leg Tr Fe Ge Ti Ta
 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, ev' l ', σ (v), φ (ft), (tc), (it), (p), (u), (a), s, (pv)
IV v' d, ev' d, l ' (v), φ ft '', (tc), (p), (u), (a), s, (pv)

See Table 1 for explanations.

Remarks. (1) The species Pseudotocepheus sexidimorphus was described by Vasiliu and Călugăr (1977) based on material from Cuba and included in the genus Nesotocepheus Hammer, 1972 (at present, it is a junior synonym of Pseudotocepheus). However, the taxonomic analysis of Otocepheidae indicated an incorrect systematic placement of this species, which fits the definition/diagnosis of the genus Leptotocepheus (see Ermilov, Minor, 2018) based on the presence of unpaired medial notogastral condyle (versus the medial notogastral condyle being either absent or represented by one pair in Pseudotocepheus). Vasiliu and Călugăr (1977) did not discuss the morphology or the presence/absence of medial notogastral condyles in P. sexidimorphus. However, their Figs 6, 8 (especially Fig. 8 ) clearly show a developed unpaired medial notogastral condyle. All our specimens also have this condyle. Thus, P. sexidimorphus should be transferred to the genus Leptotocepheus, subgenus Longocepheus: Leptotocepheus (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977) comb. n.

(2) Regarding combining L. (Longocepheus) sexidimorphus, I will note that it is morphologically most similar to L. (Longocepheus) neozealandicus Ermilov et Minor 2018 from New Zealand in having long, fusiform bothridial setae and comparatively long, stiff interlamellar and notogastral setae. The former species differs from the latter in the localization of adanal setae ad3 in the lateral (versus anterolateral) position and the presence of foveolate body surface (versus such surface being absent in L. neozealandicus). Also, in L. sexidimorphus, costulae are long (versus short) and the teeth on leg tarsi are minute (versus strong).

(3) My Cuban specimens of L. (Longocepheus) sexidimorphus are morphologically similar in their general appearance to those from the original description (Vasiliu, Călugăr, 1977). In particular, body size is medium to large; females larger than males; body surface foveolate; rostrum broadly rounded; costulae slightly not reaching insertions of lamellar setae; rostral and lamellar setae long, setiform; interlamellar setae long, stiff; exobothridial setae short, but well visible; bothridial setae long, fusiform; lateral and medial prodorsal and lateral notogastral condyles tubercle-like, separated; unpaired medial notogastral condyle broad; 10 pairs of notogastral setae comparatively long, stiff, setae h2 specifically directed laterally; epimeral setal formula: 3–1–3–3; three pair of genital setae; adanal setae ad3 in lateral position; adanal lyrifissures removed from anal aperture.

However, two important differences are present. Firstly, specimens of L. (Longocepheus) sexidimorphus from the original description are clearly larger than my specimens: 745 × 317 (male), 950 × 420 (female) versus 464–531 × 182–215 (males), 581–647 × 249–282 (females). Also, the dorsal and ventral setae are smooth in male and barbed in female (versus the dorsal and ventral body setae being barbed in my males and females).

First of all, the difference in the body size of L. (Longocepheus) sexidimorphus is not critical due to the fact that among representatives of Otocepheidae there is a well-known significant variation of this parameter. In particular, this trait is evident in some species of Leptotocepheus (Longocepheus), such as L. (L.) australis (Balogh et Mahunka 1966), L. (L.) youngai (Mahunka 1984), and L. (L.) globosus (Grobler 1995). Secondly, the original description of L. (Longocepheus) sexidimorphus (Vasiliu et Călugăr 1977) is based only on two specimens (the holotype and one paratype). Therefore, the general conclusion about the differences in the morphology of the dorsal and ventral body setae in females and males for this species looks premature.

Thus, due to the lack of significant morphological differences (excluding barbulation of body setae in males and females) between Vasiliu and Călugăr’s specimens on the one hand and my specimens on the other, I identify all of these specimens as L. (Longocepheus) sexidimorphus. However, additional research (for example, genetic) is necessary.

Список литературы

  1. Balogh J., 1960. Oribatides (Acari) nouveaux de Madagascar (1re série) // Mémoires de l’Institut Scientifique de Madagascar, Série A. V. 14. P. 7–37.

  2. Balogh J., Balogh P., 2002. Identification keys to the oribatid mites of the Extra-Holarctic regions. V. 1. Miskolc: Well-Press Publishing Limited. 453 p.

  3. Ermilov S.G., 2016. 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 // Systematic and Applied Acarology. V. 21. № 2. P. 209–217.

  4. Ermilov S.G., Minor M., 2018. Contribution to the knowledge of the oribatid mite genus Leptotocepheus (Acari, Oribatida, Otocepheidae) // Systematic and Applied Acarology. V. 23. № 11. P. 2269–2278.

  5. Ermilov S.G., Minor M., 2019. New Pseudotocepheus (Acari, Oribatida, Otocepheidae) from New Zealand, with a key to known species of the genus from the Australian region // International Journal of Acarology. V. 45. № 3. P. 131–140.

  6. Grobler L., 1998. Subfamily Pseudotocepheinae (Acari; Oribatida; Otocepheidae): Constrictocepheus gen. nov. and new Pseudotocepheus species from South Africa // Navorsinge van die Nasionale Museum, Bloemfontein. V. 14. № 1. P. 1–44.

  7. Hammer M., 1966. Investigations on the oribatid fauna of New Zealand. Part I // Det Kongelige Danske Videnskabernes Selskab Biologiske Skrifter. V. 15. № 2. P. 1–108.

  8. Norton R.A., 1977. A review of F. Grandjean’s system of leg chaetotaxy in the Oribatei (Acari) and its application to the family Damaeidae // Dindal D.L., ed. Biology of oribatid mites. Syracuse, SUNY College of Environmental Science and Forestry. P. 33–61.

  9. Norton R.A., Behan-Pelletier V.M., 2009. Oribatida // A Manual of Acarology (TX). Lubbock: Texas Tech University Press. P. 430–564.

  10. Subías L.S., 2019. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles). Online version accessed in March 2019, 536 pp.; http://bba.bioucm.es/cont/docs/RO_1.pdf

  11. Travé J., Vachon M., 1975. François Grandjean. 1882–1975 (Notice biographique et bibliographique) // Acarologia. V. 17. № 1. P. 1–19.

  12. Vasiliu N., Călugăr M., 1977. Oribates nouveaux de la Sierra de la Gran Piedra – Cuba // Resultats des expeditions biospeleologiques cubano-roumaines a Cuba. V. 2. P. 241–246.

Дополнительные материалы отсутствуют.