Acari, Oribatida - BioOne

Systematic & Applied Acarology 22(7): 948–961 (2017) http://doi.org/10.11158/saa.22.7.4 Article

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

http://zoobank.org/urn:lsid:zoobank.org:pub:AB5A0154-DC5D-423A-9A21-A0048611AE25

New faunistic and taxonomical findings of oribatid mites (Acari, Oribatida) of the family Otocepheidae from Myanmar SERGEY G. ERMILOV Tyumen State University, Tyumen, Russia. E-mail: [email protected]

Abstract The first data on the oribatid mites of Myanmar are presented and these belong to the family Otocepheidae. A new species of the genus Eurostocepheus (Oribatida, Otocepheidae) is described from litter of disturbed primary forest of Nat Ma Taung National Park in the Chin State. Eurostocepheus (Eurostocepheus) peterjaegeri sp. nov. is morphologically most similar to E. (E.) mahunkai Mondal & Kundu, 1999, but differs by the shorter notogastral setae c and la compared to the other notogastral setae, setae la pressed to notogastral surface, welldeveloped lateral prodorsal carinae, smooth genital plates and by the presence of diagonal ridges on the anal plates. A supplementary description of Otocepheus (Otocepheus) heterosetiger Aoki, 1965 which was originally described from Thailand, is given in detail on the basis of specimens from Myanmar. The main morphological traits for this species are summarized. Key words: otocepheid mites, Eurostocepheus, Otocepheus, new species, supplementary description, morphology, systematics, new record, Oriental region

Introduction The oribatid mite fauna (Acari, Oribatida) of Myanmar is still completely unknown. This work is part of a study of oribatids from this country and includes data on the family Otocepheidae. It is based on materials which were received from the collection of the Senckenberg Museum of Natural History (Görlitz, Germany) and collected during a two-week expedition of the German zoologists in May 2015 in forests on Mt. Victoria of Nat Ma Taung National Park in the Chin State. In the course of taxonomic identification I found two representatives of the family Otocepheidae, one species belonging to the genus Eurostocepheus Aoki, 1965 is new to science, and the other known species, Otocepheus heterosetiger Aoki, 1965, both are from the nominative subgenera. The primary goal of the paper is to describe and illustrate a new species under the name Eurostocepheus peterjaegeri sp. nov. The genus Eurostocepheus includes two subgenera and eight species; the generic revision, including new generic and subgeneric diagnoses, identification key, distribution and ecology of known taxa, was presented by Ermilov & Starý (2017). The secondary goal of the paper is to present a supplementary description of O. heterosetiger, adding new information about some morphological structures and their measurements, identification of leg setae and solenidia, and morphology of the gnathosoma. This species was previously only known from Thailand (Aoki 1965; Mahunka 2008).

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Material, methods, terminology and abbreviations Material examined Eurostocepheus (Eurostocepheus) peterjaegeri sp. nov. (holotype: female; two paratypes: two males) and Otocepheus (Otocepheus) heterosetiger Aoki, 1965 (six specimens: three females and three males): Myanmar (Burma), Chin State, Nat Ma Taung National Park, road Kampetlet-Mindat, 21°12'33.8''N, 94°01'26.8''E, 2150 m a.s.l., disturbed primary forest, litter, Winkler extraction, 11.V.2014 (collected by Peter Jäger). Methods Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. 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 of the ventral plate. 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. Drawings were made with a camera lucida using a Leica transmission light microscope “Leica DM 2500”. Images were obtained with an AxioCam ICc3 camera using a Carl Zeiss transmission light microscope “Axio Lab.A1”. Terminology Morphological terminology used in this paper follows that of F. Grandjean: see Travé & Vachon (1975) for references, Norton (1977) for leg setal nomenclature, and Norton & Behan–Pelletier (2009), for overview. Abbreviations The following abbreviations are used on the figures: cos—costula; apc—anterolateral prodorsal carinae; ppc—posterolateral prodorsal carinae; ro, le, in, bs, ex—rostral, lamellar, interlamellar, bothridial and exobothridial setae, respectively; bo—bothridium; co.pm, co.pl—medial and lateral prodorsal condyles, respectively; co.nm, co.nl—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; dmr—dorsal circummarginal ridge; h, m, a —subcapitular setae; v, l, d, cm, acm, ul, sul, vt, lt—palp setae; cha, chb—cheliceral setae; cht—cheliceral tooth; Tg— Trägårdh’s organ; Pd I, Pd II—pedotecta I, II, respectively; st—sternal apodeme; 1a, 1b, 1c, 2a, 3a, 3b, 3c, 4a, 4b, 4c—epimeral setae; dis—discidium; vr—ventral ridge; g, ag, an, ad—genital, aggenital, anal and adanal setae, respectively; iag—aggenital lyrifissure; anr—ridge on anal plate; iad—adanal lyrifissure; cvr—circumventral ridge; ω, σ, φ—solenidia; ɛ—leg famulus; v, ev, bv, l, d, ft, tc, it, p, u, a, s, pv—leg setae.

Description of new species Eurostocepheus (Eurostocepheus) peterjaegeri sp. nov. (Figs 1–10) Diagnosis Body size: 1294–1344 × 614–680. Body ratio (length/width): 1.9–2.1. Body surface densely microgranulate. Rostrum undulate. Rostral, lamellar and interlamellar setae long, setiform, barbed. 2017

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Bothridial setae lanceolate, slightly roughened. Lateral prodorsal condyles tubercle-like, medial prodorsal condyles broadly rounded distally, lateral notogastral condyles triangular. Notogastral setae setiform, barbed, c and la shortest, la pressed to notogaster surface. Subcapitular setae a shortest, h longest. Epimeral setal formula: 3-1-3-3; setae setiform, barbed, 2a, 3a, 4b shortest, 1b longest. Ventral plate with a pair of longitudinal ridges posterior to epimeral setae 4a. Anal plates with diagonal ridge in medioposterior parts. Anogenital setae setiform, barbed. Anal setae inserted in longitudinal position. Adanal setae ad1 in posterolateral, ad2 and ad3 in paraanal positions. Adanal lyrifissures located in diagonal position and close to anal aperture. Some leg setae it and tc on tarsi dilated medio-distally; u setiform on tarsi I and thorn-like on tarsi II–IV.

FIGURES 1–2. Eurostocepheus peterjaegeri sp. nov., adult: 1—dorsal view (legs not shown); 2—ventral view (gnathosoma and legs not shown). Scale bar 200 μm.

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Description Measurements. Very large species. Body length: 1344 (holotype: female), 1294, 1310 (two paratypes: two males); notogaster width: 680 (holotype), 614, 647 (two paratypes). One female distinctly larger than two males in body size. Body ratio (length/width): 1.9–2.1. Integument (Figs 1–3). Body color light brown to brown, but genital plates and legs dark brown. Body surface densely microgranulate (except genital plates), granules rounded in dorsal view and conical in lateral view (diameter and length of granules less than 1). Foveolate ornamentation absent except slight foveolae (their diameter up to 4) on lamellae. Pedotecta II and lateral parts of body between bothridia and pedotecta I, II slightly tuberculate (diameter of tubercles up to 12).

FIGURES 3–6. Eurostocepheus peterjaegeri sp. nov., adult: 3—lateral view (legs not shown); 4— subcapitulum, ventral view; 5—palp, without trochanter, left, antiaxial view; 6—anterior part of chelicera, right, antiaxial view. Scale bars 200 μm (3), 50 μm (4–6).

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FIGURES 7–8. Eurostocepheus peterjaegeri sp. nov., adult: 7—leg I, right, antiaxial view; 8—trochanter, femur and genu of leg III, left, antiaxial view. Scale bar 100 μm.

Prodorsum (Figs 1–3). Rostrum slightly undulate. Costulae strong, broad, located dorsolaterally, rounded distally. Anterolateral and posterolateral prodorsal carinae present. Rostral and lamellar setae similar in length (188–209), setiform, barbed, curving antero-medially. Interlamellar setae (180–192) setiform, erect, barbed. Exobothridial setae (36–41) setae setiform, thin, slightly roughened. Bothridial setae (their length out of bothridia 118–131) with long, smooth stalks and short, hardly barbed, lanceolate heads. Lateral prodorsal condyles tubercle-like, rounded distally. Medial prodorsal condyles broadly rounded distally, located separately. Notogaster (Figs 1–3). Anterior notogastral margin slightly convex medially. Lateral notogastral condyles strongly distanced from each other, triangular, rounded distally, covered partially by lateral prodorsal condyles. Dorsal circummarginal ridge well visible. Notogaster with 10 952


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pairs of setae long, setiform, barbed, c, la (155–172) shorter than h3, p1 (205–282) and other setae (266–348). Setae la pressed to notogaster surface. Lyrifissures distinct, ia located laterally to c, im anterolaterally to lp, ip between p2 and p3, ips between h3 and p3, ih anterior to h3 and distanced from them. Opisthonotal gland openings located posterior and very close to im. Gnathosoma (Figs 4–6). Subcapitulum longer than wide (250–266). Subcapitular setae setiform, a (36–45) slightly barbed, h (118–127) and m (86–98) barbed. Adoral setae and their alveoli absent. Palps (139–155) with setation 0-2-1-3-8(+ω). Solenidia of palptarsi connected with setae ul’, ul” medioanteriorly. Postpalpal setae (12–14) erect, slightly barbed. Chelicerae (250–266) with two setiform, barbed setae, cha (86–102) longer than chb (28–36). Dorsal sides of chelicerae with one distinct tooth. Trägårdh’s organ narrowly triangular.

FIGURES 9–10. Eurostocepheus peterjaegeri sp. nov., adult: 9—trochanter, femur and genu of leg II, right, antiaxial view; 10—leg IV, left, antiaxial view. Scale bar 100 μm.

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Epimeral and lateral podosomal regions (Figs 2, 3). Apodemes I, II and sejugal apodemes welldeveloped, apodemes III slightly visible. Unpaired sternal apodeme reduced to small part between epimeral setae 1a. Epimeral setal formula: 3-1-3-3. Setae setiform, barbed, 2a, 3a, 4b (53–61) shorter than 1a (61–69), 1c, 3b, 3c, 4a, 4c (90–102) and 1b (131–139). Pedotecta I and II of typical morphology for the genus. Discidia elongate triangular, rounded distally. Anogenital region (Figs 2, 3). Aggenital lyrifissures located close and anterolateral to genital aperture. A pair of long, longitudinal ridges posterior to epimeral setae 4a. Anal plates with one strong diagonal ridge in medioposterior parts. Four pairs of genital (g1, g4, 45–49; g2, g3, 53–57), one pair of aggenital (53–57), three pairs of adanal (ad1, 127–145; ad2, ad3, 114–127) and two pairs of anal (an1, 94–102; an2, 77–94) setae setiform, barbed. Anal setae inserted in longitudinal position, an1 on ridges. Adanal setae ad1 in posterolateral, ad2 and ad3 in paraanal positions. Adanal lyrifissures located in diagonal position and close to anal aperture. One circumventral ridge present. Legs (Figs 7–10). Claw of each leg strong, slightly serrate on dorsal side. Dorso-paraxial porose areas on femora I, II and dorso-antiaxial porose areas on femora III, IV and trochanters III, IV poorly visible. Tarsi I, II with one to two small teeth. Genua pointed ventrally. Femora with broadly rounded expansions ventroanteriorly. Formulas of leg setation and solenidia: I (1-4-3-4-16) [1-2-2], II (1-43-3-15) [1-1-2], III (2-3-1-2-15) [1-1-0], IV (1-2-2-2-12) [0-1-0]; homology of setae and solenidia indicated in Table 1. Solenidia φ1 on tibiae I setiform, other solenidia thickened, blunt-ended. Famuli short, erect, blunt-ended, inserted posterior to solenidia ω1. Majority of leg setae setiform; it’ and tc’ on tarsi I, II, it” on tarsi III and tc” on tarsi III, IV slightly dilated medio-distally; l” thorn-like on genua I–III; p and s simple on tarsi I; u setiform on tarsi I and thorn-like on tarsi II–IV. Setae l’ setiform, very long on trochanters III. TABLE 1. Leg setation and solenidia of adult Eurostocepheus peterjaegeri sp. nov. and Otocepheus heterosetiger Aoki, 1965. 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’

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)

Note: Roman letters refer to normal setae, Greek letters to solenidia (except ɛ = famulus). Single prime (’) marks setae on the anterior and double prime (”) setae on the posterior side of a given leg segment. Parentheses refer to a pair of setae. Tr—trochanter, Fe—femur, Ge—genu, Ti—Tibia, Ta—tarsus.

Type deposition The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany. Two paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. Etymology The species name is dedicated to the well-known German zoologist Dr. Peter Jäger (Senckenberg Research Institute, Frankfurt am Main, Germany), for his extensive contributions to our knowledge of Araneae. Remarks The new species is morphologically most similar to Eurostocepheus (Eurostocepheus) mahunkai Mondal & Kundu, 1999 from India in having medial prodorsal condyles and relatively long interlamellar and notogastral setae, but differs by the shorter notogastral setae c and la compared to 954


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the other notogastral setae (vs. all notogastral setae approximately equal in length), setae la pressed to notogastral surface (vs. la erect, not pressed), well-developed lateral prodorsal carinae (vs. lateral prodorsal carinae rudimentary), genital plates smooth (vs. genital plates striate), and by the presence of a diagonal ridge on each anal plate (vs. ridges on anal plates absent). Based on the main generic traits (Ermilov & Starý 2017), E. peterjaegeri sp. nov. is included in Eurostocepheus, however, this species has well-developed anterolateral and posterolateral prodorsal carinae and long notogastral setae (vs. anterolateral and posterolateral prodorsal carinae not developed, and notogastral setae short or medium sized). Thus, these additional morphological traits must be included in the future generic diagnosis, or the systematic placement of the species should be reconsidered.

Supplementary description Otocepheus (Otocepheus) heterosetiger Aoki, 1965 (Figs 11–25) Measurements. Very large species. Body length: 1045–1260 (six specimens: three females and three males); notogaster width: 365–531 (six specimens). Females larger than males in body size: 1094– 1260 × 415–531 vs. 1045–1079 × 365–398. Body ratio (length/width): 2.3–2.9. Integument (Figs 11–13, 25). Body color light brown, genital plates and legs dark brown. Body surface densely microgranulate (except genital plates) and sparsely foveolate (diameter of foveolae up to 4), granules rounded in dorsal view and conical in lateral view (diameter and length of granules less than 1), interrupted by spaced slight reticulate ornamentation in anogenital and epimeral regions. Prodorsum (Figs 11–13, 21, 22). Rostrum rounded. Costulae strong, narrow, located dorsally, rounded distally. Anterolateral and posterolateral prodorsal carinae present. Rostral and lamellar setae similar in length (151–264), setiform, barbed, curving antero-medially. Interlamellar setae (65– 73) dilated in median parts, erect, barbed. Exobothridial setae (12–16) setae setiform, thin, smooth. Bothridial setae (their length out of bothridia 77–94) with long, smooth stalks and short, slightly barbed, clavate heads. Lateral prodorsal condyles tubercle-like, rounded distally. Notogaster (Figs 11–13, 23–25). Anterior notogastral margin straight. Lateral notogastral condyles located close to each other, quadrangular, covered partially by lateral prodorsal condyles. Dorsal circummarginal ridge well visible. Notogaster with 10 pairs of barbed setae, c, la, lm, lp, h2, sometimes h1 (106–118) dilated in median parts, shorter than thickened p1, p2, sometimes h1 (118– 139), setae p3, h3 (36–41) setiform, thin. Setae c and la pressed to notogaster surface. Lyrifissures distinct, ia located laterally to c, im laterally to lm, ip between p2 and p3, ips between h3 and p3, ih anterior to h3. Opisthonotal gland openings located posterior and very close to im. Gnathosoma (Figs 14–16). Subcapitulum longer than wide (225–233). Subcapitular setae setiform, a (28–32) slightly barbed, h (82–94) and m (61–73) barbed. Adoral setae and their alveoli absent. Palps (118–139) with setation 0-2-1-3-8(+ω). Solenidia of palptarsi connected with setae ul’, ul” medioanteriorly. Postpalpal setae (10) erect, slightly barbed. Chelicerae (225–233) with two setiform, barbed setae, cha (73–90) longer than chb (32–36). Dorsal sides of chelicerae with one distinct tooth. Trägårdh’s organ narrowly triangular. Epimeral and lateral podosomal regions (Figs 12, 13). Apodemes I, II and sejugal apodemes well-developed, apodemes III not visible. Unpaired sternal apodeme of medium size. Epimeral setal formula: 3-1-3-3. Setae setiform, barbed, 1a, 2a, 3a (16–20) shorter than 1c, 3c, 4b, 4c (57–73) and 1b, 3b, 4a (110–118). Pedotecta I and II of typical morphology for the genus. Discidia elongate triangular, rounded distally. 2017


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FIGURES 11–12. Otocepheus heterosetiger Aoki, 1965, adult: 11—dorsal view (legs not shown); 12—ventral view (gnathosoma and legs not shown). Scale bar 200 μm.

Anogenital region (Figs 12, 13). Aggenital lyrifissures located close and anterolateral to genital aperture. Four pairs of genital (g1, g4, 32–36; g2, g3, 36–41), one pair of aggenital (49–53), three pairs of adanal (ad1, 61–65; ad2, ad3, 45–53) and two pairs of anal (an1, 61–69; an2, 41–44) setae setiform, barbed. Adanal setae ad1 in posterolateral, ad2 and ad3 in paraanal positions. Adanal lyrifissures located parallel and close to anal aperture. Two circumventral ridges present. Legs (Figs 17–20). Claw of each leg strong, slightly serrate on dorsal side. Dorso-paraxial porose areas on femora I, II and dorso-antiaxial porose areas on femora III, IV and trochanters III, 956


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IV poorly visible. Tarsi I, II with one to two small teeth. Genua pointed ventrally. Femora with broadly rounded expansions 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 (2-3-1-2-15) [1-1-0], IV (1-2-2-2-12) [0-1-0]; homology of setae and solenidia indicated in Table 1. Solenidia φ1 on tibiae I setiform, other solenidia thickened, bluntended. Famuli short, erect, blunt-ended, inserted posterior to solenidia ω1. Majority of leg setae setiform; setae (or some of these) it’ and tc’ on all tarsi slightly dilated medio-distally; l” thorn-like on genua I, II; p and s simple on tarsi I; u setiform on tarsi I and thorn-like on tarsi II–IV. Setae l’ setiform, very long on trochanters III.

FIGURES 13–16. Otocepheus heterosetiger Aoki, 1965, adult: 13—lateral view (legs not shown); 14— subcapitulum, ventral view; 15—palp, without trochanter, right, ventroparaxial view; 16—chelicera, left, paraxial view. Scale bars 200 μm (13), 50 μm (14, 16), 20 μm (15).

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FIGURES 17–18. Otocepheus heterosetiger Aoki, 1965, adult: 17—leg I, right, antiaxial view; 18—trochanter, femur and genu of leg III, left, antiaxial view. Scale bar 50 μm.

Remarks The specimens of O. heterosetiger from Myanmar and Thailand (Aoki 1965; Mahunka 2008) are morphologically very similar. Only one difference is present: adanal setae ad1 clearly longer than ad2, ad3 in specimens from Myanmar vs. all adanal setae similar in length in specimens from Thailand. Thus, this addition must be considered in the future identification of the species.

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FIGURES 19–20. Otocepheus heterosetiger Aoki, 1965, adult: 19—trochanter, femur and genu of leg II, right, antiaxial view; 20—leg IV, left, antiaxial view. Scale bar 50 μm.

Based on the original description (Aoki 1965) and supplementary descriptions of this paper and Mahunka (2008), the following diagnostic morphological traits for O. heterosetiger are proposed: Body size: 1038–1302 × 365–582. Body ratio (length/width): 2.3–2.9. Body surface densely microgranulate and sparsely foveolate. Rostrum rounded. Rostral and lamellar setae long, setiform, barbed. Interlamellar setae dilated in median parts, barbed. Bothridial setae clavate, slightly barbed. Lateral prodorsal condyles tubercle-like, lateral notogastral condyles quadrangular. Notogastral setae barbed, c, la, lm, lp, h2, sometimes h1 dilated in median parts, p1, p2, sometimes h1 longest, thickened, p3, h3 shortest, setiform, thin, c and la pressed to notogaster surface. Subcapitular setae a shortest, h longest. Epimeral setal formula: 3-1-3-3; setae setiform, barbed, 1a, 2a, 3a shortest, 1b, 3b, 4a longest. Anogenital setae setiform, barbed. Adanal setae ad1 in posterolateral, ad2 and ad3 in paraanal positions. Adanal lyrifissures located parallel and close to anal aperture. Some leg setae it and tc on tarsi dilated medio-distally; u setiform on tarsi I and thorn-like on tarsi II–IV.

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FIGURES 21–25. Otocepheus heterosetiger Aoki, 1965, adult, microscope images: 21—interlamellar seta, bothridium, lateral prodorsal condyle and basal part of bothridial seta; 22—bothridial and exobothridial setae; 23—lateral notogastral condyle and notogastral seta c; 24—notogastral setae h1 and h2; 25—notogastral setae lp, h3 and p3, lyrifissures ih and ips, part of dorsal circummarginal ridge and basal part of notogastral seta h2.

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Acknowledgements I cordially thank Dr. Elizabeth A. Hugo-Coetzee (National Museum, Bloemfontein, South Africa), Dr. Leonila Corpuz-Raros (Crop Protection Cluster, College of Agriculture and Museum of Natural History, University of Los Baños, Los Baños, Philippines) and one anonymous reviewer for their valuable comments; Dr. Peter Jäger (Senckenberg Research Institute, Frankfurt am Main, Germany), who collected materials from Myanmar and transferred it for my study from the collection of the Senckenberg Research Institute, and Prof. Dr. Jochen Martens (Johannes Gutenberg Universität, Mainz, Germany) for support in the field.

References Aoki, J. (1965) A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata). I. Subfamily Otocepheinae. Bulletin of the National Museum of Natural Science, 8(3), 259–341. Ermilov, S.G. & Starý, J. (2017) Contribution to the knowledge of the oribatid mite genus Eurostocepheus (Acari, Oribatida, Otocepheidae). Systematic and Applied Acarology, 22(5), 640–652. https://doi.org/10.11158/saa.22.5.4 Mahunka, S. (2008) More oribatids from Thailand (Acari: Oribatida). Revue suisse de Zoologie, 115(4), 623– 649. https://doi.org/10.5962/bhl.part.80450 Mondal, B.K. & Kundu, B.G. (1999) A new species of the genus Eurostocepheus (Acari: Oribatei, Otocepheidae) from Darjeeling, India. Records of the Zoological Survey of India, 97(1), 179–186. 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. In: Dindal, D.L. (Editor), Biology of oribatid mites. SUNY College of Environmental Science and Forestry, Syracuse, pp. 33–61. Norton, R.A. & Behan-Pelletier, V.M. (2009) Suborder Oribatida. Chapter 15. In: Krantz, G.W. & Walter, D.E. (Editors), A Manual of Acarology. Texas Tech University Press, Lubbock, pp. 430–564. Travé, J. & Vachon, M. (1975) François Grandjean. 1882–1975 (Notice biographique et bibliographique). Acarologia, 17(1), 1–19. Submitted: 12 May 2017; accepted by Lizel Hugo-Coetzee: 31 May 2017; published: 27 Jun. 2017

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