Two New Circumpolar Mite Species of the Genus ... - Springer Link

ISSN 0013–8738, Entomological Review, 2011, Vol. 91, No. 8, pp. 1054–1072. © Pleiades Publishing, Inc., 2011. Original Russian Text © E.E. Lindquist, O.L. Makarova, 2011, published in Zoologicheskii Zhurnal, 2011, Vol. 90, No. 8, pp. 923–941.

Two New Circumpolar Mite Species of the Genus Arctoseius Thor (Parasitiformes, Mesostigmata, Ascidae) E. E. Lindquista and O. L. Makarovab a

Invertebrate Biodiversity, Research Branch, Agriculture & Agri-Food Canada, Ottawa, Ontario K1A 0C6, Canada e-mail: [email protected], [email protected] b Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, 119071 Russia e-mail: [email protected] Received April 20, 2011

Abstract—We describe two new species of Arctoseius, A. sexsetus sp. n. and A. haarlovi sp. n. These descriptions provide additional data on the circumpolar distribution and morphological variation of these species, which were previously presented under the same names by Lindquist (1964) in his unpublished dissertation. The relationships of these species with other circumpolar species that may form a species-group, including the type-species of the genus Arctoseius, A. laterincisus Thor 1930, and its questionable identity are discussed. DOI: 10.1134/S0013873811080100

The genus Arctoseius Thor 1930 at present numbers about 50 species, among which approximately half inhabit Arctic landscapes, including no less than 12 species that have a circumpolar arctic geographic range (Makarova, 2011). In his unpublished dissertation of 1963 on North American material, the first author described four species belonging to this genus (Lindquist, 1964), two of which (“Arctoseius sexsetus Lindquist 1963”, ms. and “Arctoseius haarlovi Lindquist 1963”, ms.) are now known to be broadly distributed in the Russian part of the Arctic. In this communication we present detailed descriptions of the above-named species, including the first found male of A. sexsetus sp. n. and information about new records. Previous confusion between Arctoseius laterincisus Thor 1930 and A. haarlovi is reviewed, and the relationships between these species along with A. sexsetus and A. multidentatus Evans 1955, which may form a species-group, are considered. The setal nomenclature follows Lindquist & Evans, 1965 with some modification (Lindquist, 1994). The leg and palpal chaetotaxy complies with Evans, 1963, 1964. The poroidotaxy and adenotaxy are given according to Johnston and Moraza, 1991, with small addition (Makarova, 2003). The length of all shields was measured along the mid-line, and the width at the broadest part, except the sternal shield, whose length was measured from the level of setae st1 to the posterior margin, and the width at the broadest part between levels of setae st2 and st3 (Makarova, 2000a). The

length of the epigynal shield was measured to include the anterior flap, the width at the posterior part. The length of all legs and tarsi are given excluding the ambulacrum, and also its pedicel on leg I. The subcapitulum length was measured from its anterior margin excluding projections (internal malae, corniculi). The measurements were carried out on all available specimens and their limits are stated in micrometers (μm). The following ratios were used in the description (Makarova, 2000a): lD/wD—the length-to-width ratio of the dorsal shield; J(1–4)min/jmax—the ratio of the length of dorsocentral setae inserted on the podonotal and opisthonotal regions, notably the shortest among J1–4 and the longest in the series j; (J4–J4)/(J3–J3)—the ratio of transverse distances between bases of opisthonotal setae J4 and J3 in pairs; Z5/(Z5–Z5)—the ratio of the setae Z5 length to the distance between them; lSt/wSt, lAn/wAn, lVA/wVA—the length-to-width ratios of the sternal, anal, and ventrianal (in males) shields respectively; lCo/lD—the ratio of the lengths of the corniculus and the dorsal shield, %; lCh/lD—the ratio of the lengths of the movable cheliceral digit and the dorsal shield, %;

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lCh/lCo—the ratio of the lengths of the movable cheliceral digit and the corniculus; lTIV/lD—the ratio of the lengths of tarsus IV and the dorsal shield; lTIII/wTIII—the length-to-width ratio of tarsus III. In the male description, the features common with females were omitted. DESCRIPTIONS OF NEW SPECIES Arctoseius sexsetus Lindquist et Makarova sp. n. (Figs. 1, 2, 3, 8) Arctoseius sexsetus Lindquist 1963 pp. 100–102, figs. 10–14 (nomen nudum).

(1964),

Material. Holotype, ♀: RUSSIA, TAYMYR, northern coast of Taymyr Lake, Cape Blizhnii, lemming hill in polygonal tundra on ridge, 10.VIII.1994, O.L. Makarova. Paratypes: 5 ♀, 3 ♂, 1 deutonymph with same data as for holotype; 1 ♂, vicinity of Norilsk town, at foot of Talnakh Mts., Krasnye Kamni site, tundra in nival valley, 8.VII.2000, O.L. Makarova; FAR EAST, 1 ♀, Primorskii Territory, Amurskaya Province, vicinity of Ust-Urkima Village, alder forest litter, 1977, A.A. Lyashchev; ARKHANGELSK PROVINCE, 1 ♀, vicinity of Lomovoe Village, cutover land, VIII 1984, N.M. Klyuyeva; 1 ♀, Pinega Reserve, karst ravine Maseyev, short grass gramineous-forb meadow, 13.VIII.2005, A.B. Babenko; USA, ALASKA, 2 ♀, 1 deutonymph, Cape Thompson, Flag Hill, from willow, 30.VII.1960, W.C. Hanson; Other material: EAST SIBERIA, 3 ♀, delta of Yana River, Shirokostan Peninsula, vicinity of Ledyanoe Lake, Dryas community on south slope of valley, 4– 6.VIII.1994, A.B. Babenko; 1 deutonymph, vicinity of Yakutsk, birch forest with alder and wild rosemary, 14.VII.1992, M.B. Potapov; SOUTHERN SIBERIA, 7 ♀, 1 ♂, North-Eastern Altai, Turochakskii District, vicinity of the Obogo lodge, spruce-fir-cedar mountainous taiga, litter and soil, 17.VIII.2007, I.I. Marchenko; 4 ♀, 5 ♂, North-Eastern Altai, Altyn-Tu Range, Archa Mt., dwarf birch mountainous tundra, litter and soil, 19.VIII.2007, I.I. Marchenko. Holotype is deposited in Zoological Institute, Russian Academy of Sciences, St. Petersburg; 1 paratype ♀ from Alaska, in United States National Museum of Natural History, Washington, DC; 1 paratype ♀ and 1 deutonymph from Alaska, in Canadian National Collection of Insects and Arachnids, Ottawa; other paratypes, in Zoological Institute, St. Petersburg. ENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011

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Diagnosis. Dorsal shield narrow, its length more than twice its width, podonotal region with 16–17 pairs of setae, opisthonotal region with 11–13 pairs of setae (z3, S1 and S2 always absent, J3 sometimes absent, S3 and in male s6 sometimes on soft cuticle). Peritreme shortened, not exceeding mid-level of coxa II anteriorly. Gnathotectum in general with three processes, which sometimes bifurcated. Corniculi large, longer than cheliceral digits. Fixed digit of chela slightly abbreviated, terminating in a trident or quadrident, slightly more posteriorly with pilus dentilis and 1 denticle. Movable digit slightly longer than fixed one. Tibia IV with 6 setae. Ambulacrum I on pedicellate base, claws small. In female, sternal shield united with endopodal extensions; anal shield small, variably roundish-oval, circumanal setae of similar size. In male, entire ventrianal shield present; metapodal platelets free; spermatodactyl simple, groove-like; legs without dimorphically modified setae. Description. Middle-sized yellowish mites with narrow idiosoma. Idiosomal shields very finely punctate, reticulate ornamentation developed only on lateral (endopodal) parts of sternal (sternitigenital) shield. Many setae of body and appendages with fine, hair-like tips, which often are broken off. Lateral incisions of dorsal shield rather broad, extending far within longitudinal level of seta Z1 (incision length usually 30–46), in some females slightly oblique, in some males shortened (up to 16). Female. Dorsal shield 444–552 × 184–248, narrow, lD/wD 2.10–2.57, its maximal width at level of setae s3 (Fig. 1, 1). Podonotal region with 16–17 pairs of simple setae (z3 always absent, sometimes symmetrically or asymmetrically s2 absent). Opisthonotal region with 11–13 pairs of setae (S1 and S2 always absent, often at least asymmetrically J3 omitted, rarely J4 missed, S3 sometimes inserted off of shield). Among podonotal setae z1, s1 and z6 distinguished by slightly shorter length (16–25), length of other setae 19–38. On opisthonotal region setae Z4 (44–57) and Z5 (50–71) clearly longer than others, setae of series J shortest (J1–4 16–28, J5 11–15), length of setae Z1–3 increasing posteriad (20–34), length of setae S3–5 18–30; J(1–4)min/jmax 0.53–0.80, (J4–J4)/ (J3–J3) 1.41–1.87, Z5/(Z5–Z5) 0.78–1.27. Dorsal shield with 2 pairs of cutaneous glands gdZ3 and gdZ4. Marginal setae on soft cuticle, their length 16–32; in series r 4 (rarely 2, 3, 5) setae, in series R 5 (rarely 4) setae.

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Fig. 1. Arctoseius sexsetus sp. n., female: (1) idiosoma, dorsal view; (2, 3) gnathotectum; (4) peritrematal shield; (5) chelicera, lateral view; (6) palp. Scale (μm): (1) 100; (2, 3, 5) 25; (4, 6) 50.

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Fig. 2. Arctoseius sexsetus sp. n.: (1–10) female; (11) male [(1) idiosoma, ventral view; (2) tritosternum; (3) subcapitulum; (4–6) variants of posterior margin form of sternal shield; (7, 8) epigynal shield; (9,10) anal shield; (11) tip of tarsus II with ambulacrum]. Scale (μm): (1) 100; (2–10) 50; (11) 25.


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Fig. 3. Arctoseius sexsetus sp. n.: (1, 2) female; (3–9) male [(1) subcapitulum and chelicera; (2, 4, 5) gnathotectum; (3) apex of hypostome; (6) ventrianal shield; (7) idiosoma, ventral view; (8) chelicera, ventral view; (9) chelicera, lateral view]. Scale (μm): (1, 3–6) 50; (2, 8, 9) 25; (7) 100; (1, 2) from Lindquist, 1964.



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Fig. 4. Arctoseius haarlovi sp. n.: (1–5, 7) female; (6) male [(1) idiosoma, dorsal view; (2) idiosoma, ventral view; (3) tritosternum; (4) tip of tarsus III with ambulacrum; (5, 7) anterior region of sternal shield; (6) anterior region of sternitigenital shield]. Scale (μm): (1–3, 6, 7) 50; (4, 5) 25; (1) from Lindquist, 1964.


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Fig. 5. Arctoseius haarlovi sp. n.: (1, 2) female; (3–9) male [(1–4) chelicera; (5) idiosoma, ventral view; (6–8) gnathotectum; (9) leg II without basal segments]. Scale (μm): (1–4, 6–9) 25; (5) 50; (5) from Lindquist, 1964.



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Fig. 6. Arctoseius haarlovi sp. n.: (1, 4, 8) female; (2, 3, 5–7, 9–11) male [(1) internal malae; (2, 3) posterior region of sternitigenital shield; (4) variants of posterior margin form of sternal shield; (5–7, 9–11) variant forms of ventrianal shield; (8) palp]. Scale (μm): (1) 10; (2–4, 8) 25; (5–7, 9–11) 50.


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Fig. 7. Arctoseius haarlovi sp. n.: (1) male; (2–19) females [(1–3) lateral incision of dorsal shield; (4–6) epigynal shield; (7–9) anal shield; (10) subcapitulum; (11–13) peritreme region, with variant lengths of peritreme: (11) Novaya Zemlya, (12) SW Greenland, (13) Finnish Lapland; (14) prosoma, lateral view; (15–19) variant forms of gnathotectum]. Scale (μm): (1–3, 10, 15–19) 25; (4–9, 11–14) 50.



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Fig. 8. Collection records of Arctoseius sexsetus sp. n.

Base of tritosternum narrow (29–36 × 15–22); lacinia broadest in middle of their length, with sparse large barbs (Fig. 2, 2); length of lacinia free part 60–76. Presternal platelets indistinct, granulate, confluent with sternal shield (Fig. 2, 1). Sternal shield longer (122–153) than wide (99–133), lSt/wSt 1.06– 1.30, consolidated with endopodal platelets between coxae I–II and coxae II–III. Endopodal projections between coxae I–II narrow, stick-like; their lateral margin concave, apices encompassing opening of gland gvb. Anterior margin of sternal shield often weakly sclerotized, straight, slightly concave or convex; its posterior margin slightly concave or angulate (Figs. 2, 4–6); lateral parts with reticulate ornamentation; sternal shield with typical setae st1–3 and lyrifisENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011

sures iv1–3. Setae st4 on soft cuticle. Length of setae st1–4 20–36. Endopodal strips between coxae III and IV free, narrow. Epigynal shield (124–156 × 71–96) of varying form (Figs. 2, 1, 7, 8), in general flask-shaped with small hyaline flap not extending to sternal shield, and posterior margin often truncate; setae st5 (20–28) and paragenital poroids iv5 on soft cuticle. Opening of gland gv2 medially to coxa IV usually poorly visible. One pair of postgenital platelets present. Opisthogastric setae (7 pairs) of moderate length, setae JV1–4, ZV1,2 18–32, only JV5 longer, 34–42. Two pairs of metapodal platelets present; anterior ones small, transversal, situated on level of posterior margin of epigynal shield; posterior metapodal platelets narrow-oval, longitudinal, 18–32 × 4–8. Anal shield of varying form

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Fig. 9. Collection records of Arctoseius haarlovi sp. n. (finds from Ellesmere Island and Baffin Island from Behan, 1978).

(Figs. 2, 1, 9, 10), usually rounded with truncate anterior margin; its length (72–92) approximately equal to width (72–100), lAn/wAn 0.84–1.15; paranal setae (32–40) slightly shorter than postanal seta (38–48); opening of gland gv3 poorly noticeable; cribrum narrow. Exopodal platelets between coxae II–III and III– IV rudimentary. Peritrematal shield of uneven width (Fig. 1, 4), its anterior end free, its posterior edge connecting with exopodal platelet enveloping coxa IV posteriorly; lyrifissures ip1–3 and gland gp present. Peritreme shortened (104–144 × 8–12), not extending beyond mid-level of coxa II anteriorly. Spermathecal apparatus not visible. Gnathotectum (width 40–56) usually with three long, thin processes (Figs. 1, 2, 3; 3, 2), middle projec-

tion (34–44) longer than lateral ones (23–31). Subcapitulum longer (108–132) than wide (88–116). Deutosternum with 7 (rarely 8) rows of small denticles (6–20 denticles in each row); groove width 11–16 (Figs. 2, 3; 3, 1). Hypostomatic pairs of setae hp1 (26– 34) and hp3 (24–32) longer than hp2 (15–20) and basal pair pc (20–26); all setae simple, attenuate. Corniculi large, narrow, 55–64 × 15–19; lCo/lD 11.5– 13.1%. Internal malae much shorter than corniculi, smooth, gradually tapering to tip. Chelicera not large, its length without basal segment 132–160; cheliceral digits of moderate size (45–56, lCh/lD 9.4–10.8%), shorter than corniculus (lCh/lCo 0.74–0.93). Fixed digit of chela ending in a trident, including apical hook (Fig. 1, 5) or a quadrident if apical hook bifid; posteriorly, setiform pilus dentilis and 1 small denticle preENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011


sent. Movable digit slightly longer than fixed one, bidentate. Palp length 152–176; internal seta of trochanter (36–42) longer than external seta (26–30); apart from common specialized setae on femur (al, thick with oblique tip) and genu (al1 spatulate, al2 thick with oblique tip), two strong anterolateral setae present on tibia, in some aspects looking like “another apotele” (Fig. 1, 6). Legs of moderate length (I 448–584, II 328–392, III 264–332, IV 392–468); leg I slightly longer than dorsal shield. Length of tarsi I 120–168, II 80–104, III 70–100, IV 112–144. Tarsi generally of moderate length (lTIV/lD 0.23–0.27) and width (lTIII/wTIII 2.9– 3.6). Leg chaetome deficient in comparison with setation typical for genus; seta pl on tibia IV always absent; also seta av2 on tibia I and seta av on genua II and III sometimes symmetrically or asymmetrically absent. Setae of legs simple, generally of moderate length, seta pd2 on genu II slightly shorter (28–37) than width of segment (36–40). Length of two elongated setae on basitarsus and telotarsus IV (62–80 and 60–76 respectively) usually slightly exceeding length of tibia IV (60–72). Ambulacrum I on pedicellate base, claws I (11–13) smaller than claws II–IV (15–18). Tarsus I distally with 5 rod-like solenidia, 3 of them inserted apically; length of sensillum with lanceolate apex 35–42. Ambulacra of legs II–IV (length 30–35) with short paradactyli (12–16). Apical setae on tarsi II–IV ad1 and pd1 (as on Fig. 2, 11) of moderate length (22–28). Four subapical setae on tarsi II–IV evenly distant from apex, almost of equal length, ventral setae av1 and pv1 (20–30) slightly shorter than lateral setae al1 and pl1 (24–34). Male. Dorsal shield 404–464 × 168–224, lateral incisions often smaller than in females (16–32). Podonotal region with 16–17 pairs of setae (z3 always absent, s6 sometimes on soft cuticle). Opisthonotal region with 11–13 pairs of setae (S1, S2 always absent, J3 often absent, S3 sometimes absent or inserted on soft cuticle). Among podonotal setae, z1, s1 and z6 (all 15– 24 in length) as a rule shorter than others (22–30). In opisthonotal region, length of setae Z1–5 increasing posteriad (22–58), setae J1–4 15–24, S3–5 15–29, J5 10–13. Marginal setae lengths 16–27; in series r 3–4 setae, in series R 3–4 setae. Sternitigenital shield united with endopodal platelets developed between coxae I–II and coxae II–III, rarely also with platelets between coxae III–IV (Fig. 3, 7); length of shield 216–251, width between levels of setae st2 and st3 92–121, width on level of ENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011

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setae st5 45–68. Sternitigenital shield with setae st1–5 (18–31) and lyrifissures iv1–3. Ventrianal shield broadly oval, with anterior margin convex or undulate (Fig. 3, 6), with 5 pairs of opisthogastric setae (ZV1 and JV5 on soft cuticle); length of shield (128–158) almost equal to width (124–160); paranal setae (27–31) slightly shorter than postanal seta (33–44). Caudal setae JV5 (24–35) longer than other opisthogastric setae (17–26). Apices of gnathotectum similar to female but sometimes shortened (up to 14–20) and variously subdivided (Figs. 3, 4, 5). Corniculi large (51–60 × 15–19), lCo/lD 11.9–13.4%. Internal malae, as opposed to female, abruptly tapered in the middle, their anterolateral margin with numerous microvilli (Fig. 3, 3). Cheliceral digits (40–47, lCh/lD 9.3–10.1%) shorter than corniculus (lCh/lCo 0.74–0.85). Fixed digit with apical trident and one denticle behind acicular pilus dentilis. Movable digit with one large denticle near tip and simple gutter-like spermatodactyl (length 20–26), hardly protruding beyond tip of digit (Fig. 3, 8, 9). Length of legs I–IV 440–488, 296–320, 252–268, and 352–384, respectively. Legs without dimorphically modified setae. Differential diagnosis. Adults of this species are similar to those of Arctoseius wisniewskii Gwiazdowicz et Kamczyc 2009 (Sudety, Poland). A. sexsetus sp. n. can be easily distinguished by the form of the chela - in both sexes the fixed digit terminates in a trident (in A. wisniewskii the tip of this digit is of the typical form); in the male the denticle of the movable digit is subapical in position, not more distant as in the male of A. wisniewskii. Distribution. At present known from tundra and forest (mainly boreal) districts of Alaska, Eastern Europe, northern Siberia, Far East and Altai Mts., Southern Siberia (Fig. 8). Ecology. Recorded from meadows including zoogenic ones (e.g. lemming colony outcrops, bird of prey outlooks, etc.), dry and humid tundra sites, smallleaved woods, mountainous taiga. Variability. Mites from mountain taiga and tundra sites of North-Eastern Altai, in comparison with more northern or arctic populations, are generally smaller— in females the lengths of dorsal shields are 444–524 (on average 476), not 504–552 (518) as in the more northern populations; their idiosoma more tapering— in female lD/wD 2.21–2.62 (2.40), not 2.10–2.44

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(2.23); and their setae collectively shorter—in female most of podonotal setae 19–34, not 24–38, Z5 50–61 (56), not 56–71 (62). In Altaic specimens of A. sexsetus sp. n. dorsal setae s2, J3 and J4 always present, setae S3 always inserted on dorsal shield; leg chaetome complete for this species (only pl absent on tibia IV); in male projections of gnathotectum are simple or scarcely divided apically, not deeply bifurcate. Etymology. The species name indicates the chaetome deficiency of tibia IV, with only 6 setae. Arctoseius haarlovi Lindquist et Makarova sp. n. (Figs. 4, 5, 6, 7, 9) Arctoseius laterincisus (?), Haarløv, 1942 (not Thor 1930), pp. 23–27. Arctoseius laterincisus, Weis-Fogh, 1947, p. 257. Arctoseius laterincisus, Chant, 1963, pp. 248–249. Arctoseius laterincisus, Evans, 1955, p. 288. Arctoseius ?laterincisus, [Haarløv, 1942, not Thor 1930] Lindquist, 1961, pp. 320–321, 335. Arctoseius laterincisus, Karg, 1971, p. 268; Karg, 1993, p. 267. Arctoseius haarlovi Lindquist 1963 pp. 103–109, figs. 15–17 (nomen nudum).

(1964),

Material. Holotype, ♀: SPITSBERGEN, Advent Bay, near Longyearbyen, drier substrate sparsely covered by Carex subspathacea, 7.V.1998, G. Sovik. Paratypes: 3 ♀, 5 ♂, 4 deutonymphs, with same data as for holotype, 7.V.1998 and 9.VI.1998; 1 ♀, 2 ♂, 3 deutonymphs, 1 protonymph, Advent Bay, moister substrate covered by Carex subspathacea, 30.VII.1998, G. Sovik; 2 ♀, 1 ♂, Horsund, tundra behind storm bank, together with ♀ and ♂ of Arctoseius multidentatus, 10.VIII.1989, A.V. Uvarov and Yu.B. Byzova; CANADA, NUNAVUT, 6 ♀, 2 ♂, 3 protonymphs, Ellef Ringnes Island, bird perch, 29.VII.2005, O.L. Makarova; NORTHWEST TERRITORIES, 1 ♀, Melville Island, Bailey Point, winter nest of lemming, 21.VII.1965, J.E.H. Martin; USA, ALASKA, 1 ♂, 3 miles SW Point Barrow, 14.VI.1968, R.W. Strandtmann; GREENLAND, 2 ♀, Mørkefjord Station, Cassiope heaths, 25.VIII.1939, N. Haarløv; 1 ♀, Nedre Midsommer So, Oxyria, 18.VI.1966, J.E.H. Martin; SEVERNAYA ZEMLYA ARCHIPELAGO, 4 ♀, 8 ♂, 3 deutonymphs, 3 protonymphs, Bolshevik Island, Solnechnaya Bay, lower reaches of Shumnaya River, owl outlook on cliff,

17.VIII.2000, O.L. Makarova. Other material: GREENLAND, 1 ♀, Søndre Strømfjord, dry slope of hill above airport, 30.VII.2003, A. Fjellberg; 1 ♀, Søndre Strømfjord, along road to inland icecap, shrub heath, 29.VII.2003, A. Fjellberg; 1 deutonymph, 3–year East Greenland Expedition, Scoresbysundfjord, 15.VIII.1933; FINNISH LAPLAND, 3 ♀, Kilpisjärvi, 69o05'N, 20o49'E, Jehkats fell, mountain birch stand, 8.X 2008, V. Huhta; ARKHANGELSK PROVINCE, 1 ♂, Pinega Reserve, karst ravine Maseyev, wet large-herb meadow, 19.VIII.2005, A.B. Babenko; KOLGUEV ISLAND, 2 ♀, Goltsovoe Lake, 69o12'N, 49o30'E, bird of prey outlook on hill tip, thick grassy turf, 27–28.VIII.1994, A.B. Babenko; 1 deutonymph, the same site, and date, herb-grass meadow on valley slope, A.B. Babenko; 1 ♂, the same site and date, moss-lichen-willow turf under Betula nana, A.B. Babenko; PECHORA BAY, 2 ♀, 2 ♂, Kuznetskoe Lake, 68o50'N, 53o45'E, dry sandy hill, gramineous vegetation, 25–26.VIII.1994, A.B. Babenko; 1 ♀, 3 ♂, polar fox hill, Cornus suecica, 25–26.VIII.1994, A.B. Babenko; dry sandy ridge, lichens, cowberry, 25– 26.VIII.1994, A.B. Babenko; NOVAYA ZEMLYA, 1 ♀, Northern Island, W. Barents stand, 13.VIII.1992, V.I. Bulavintsev; 2 ♀, Northern Island, Krestovaya Bay, VIII 1993, V.I. Bulavintsev; VAIGATCH ISLAND, 1 ♀, 19.VII.1984, V.I. Bulavintsev; YAMAL PENINSULA, 1 ♀, lower reach of Murtyyakha River, 70o18'N, 67o53'E, willow bed with cloudberry, Sphagnum spp., 22–23.VIII.1994, A.B. Babenko; 1 ♀, lower reach of Yakhadyyakha River, lemming hill with Poa arctica, 20–21.VIII.1994, A.B. Babenko; 4 ♀, Seyakha Lake, tundra, litter, 9.VII.1986, V.I. Bulavintsev; EAST SIBERIA, 1 ♀, delta of Kolyma River, tundra, VIII.1991, O.V. Starikova; 4 ♀, Suntar-Khayata Range, 63o13'N, 139o36'E, 1430 m a.s.l., forb meadow, 7.VII.2002, O.L. Makarova; WRANGEL ISLAND, 6 ♀, 1 ♂, Mamontovaya River, south slope of valley, lemming hill with Salix arctica, 23.VII.1994, A.B. Babenko; 1 ♂, Mamontovaya River, at foot of cliff, herbaceous meadow, 23.VII.1994, A.B. Babenko. Holotype and paratypes from Advent Bay, Spitsbergen, Alaska, and Greenland are deposited in Canadian National Collection of Insects and Arachnids, Ottawa; other paratypes—in Zoological Institute, Russian Academy of Sciences, St. Petersburg. Diagnosis. Dorsal shield unornamented, narrow; its length more than twice its width, podonotal region with 17–18 pairs of setae (z3 present, s2 on soft cuticle or at least asymmetrically on shield margin); opisthoENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011


notal region narrowing posterolaterally, with 12–13 pairs of setae (S2 always absent, only one pair of setae present in area usually occupied by J4 and Z4, S1 rarely asymmetrically absent, S3 sometimes off shield). Peritreme much shortened, usually not exceeding mid-level of coxa III anteriorly. Gnathotectum in general with three processes, the mid-tine very slender. Corniculi of common proportions, slightly shorter than cheliceral digits. Fixed digit of chela fully developed, with row of 7 to 11 teeth. Tibia IV with 7 setae. Ambulacrum I on pedicellate base, claws small. In adults sternal or sternitigenital shield united with endopodal extensions I–II and II–III, apart from small fragment behind coxa I which usually separated. Postanal seta about 1.3 longer than paranal setae. In female, anal shield small, variably oval or roundish. In male, ventrianal shield entire, slender, with metapodal platelets free; spermatodactyl strongly sinuate in lateral view, not tapering apically; leg II with dimorphically modified ventral setae on femur, genu, tibia and tarsus, setae v2 on femur and mv on tarsus especially enlarged, swollen proximally and inserted on raised base. Description. Relatively small yellowish mites with narrow idiosoma. Idiosomal shields very finely punctate, reticulate ornamentation clearly developed only on lateral (endopodal) parts of sternal (sternitigenital) shield and on anal (male ventrianal) shield. Dorsal shield narrow, not covering lateral and posterior parts of idiosoma. Setae of idiosoma and appendages often with exceedingly fine, hair-like tips. Lateral incisions of dorsal shield often extending to longitudinal level of seta Z1 (incision length usually 20–36), sometimes rather wide, incisions asymmetrically shortened or obliterated in some females and males (Figs. 7, 1–3). Female. Body length 440–496. Dorsal shield 388– 465 × 176–214, narrow, lD/wD 2.03–2.36, its maximal width at level of setae s3 (Fig. 4, 1). Podonotal region with 17–18 pairs of simple setae (z3 present, s2 on soft cuticle or at least asymmetrically on shield margin). Opisthonotal region with 12–13 pairs of setae (S2 always absent, only one pair of setae present in area usually occupied by J4 and Z4, S1 rarely asymmetrically absent, S3 sometimes off shield). Among podonotal setae z1 (9–13) and s1–2 (14–22) distinguished by clearly shorter length, length of other setae 22–38. On opisthonotal region, setae Z5 (34–46) slightly longer than others, setae of series J shortest (J1–3 18–30, J5 8–13), length of setae Z1–3 and S1, ENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011

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S3–5 (24–35); J(1–3)min/jmax—0.61–0.88, Z5/(Z5– Z5) 0.77–1.18. Dorsal shield with 1 pair of clearly cutaneous glands gdZ3, another pore-like pair anterolaterad setae J5 doubtfully glandular. Marginal setae of r-R series on soft cuticle, r3 (26–36) often slightly longer than others (17–34); 4 setae in series r, 5 (rarely 4) in series R. Base of tritosternum narrow (18–26 × 11–19); laciniae with sparse thin barbs of different lengths and hair-like tip (Fig. 4, 3); length of lacinia free part 55–74. Presternal platelets indistinct, granulate, lineate, confluent with sternal shield (Fig. 4, 2). Sternal shield usually slightly longer (110–126) than wide (103–123), lSt/wSt 0.98–1.14, consolidated with endopodal platelets between coxae I–II and coxae II–III. Endopodal projections between coxae I–II narrow, rod-like, often partly or fully separated, their apices encompassing opening of gland gvb (Figs. 4, 5–7). Sternal shield with anterior margin often weakly sclerotized, straight or slightly concave; its posterior margin more or less truncate, irregular (Fig. 6, 4); lateral (endopodal) parts of shield lineate-reticulate; sternal shield with typical setae st1–3 and lyrifissures iv1–3, vestigial glands gv1 usually visible on posterior margin (Fig. 6, 4). Setae st4 on soft cuticle. Sternal setae st1–2 (29–36) slightly longer than st3 (24–30), in turn slightly longer than st4 (20–26). Endopodal strips between coxae III and IV free, narrow. Epigynal shield (104–120 × 49–68) of varying form, usually tongue- or narrowly flask-shaped, with small hyaline flap not extending to sternal shield, posteriorly weakly reticulate with posterior margin slightly to strongly convex (Figs. 7, 4–6); setae st5 (18–28) and paragenital poroids iv5 on soft cuticle. Opening of gland gv2 medially to coxa IV usually poorly visible. One pair of postgenital platelets present. Opisthogastric setae (usually 7 pairs) of moderate length, JV3–4, ZV1 shortest (18– 24), JV1–2, ZV2 slightly longer (22–34), JV5 longest (28–38). Two pairs of metapodal platelets present; anterior ones small, transversal, situated on level of posterior margin of epigynal shield; posterior metapodal platelets narrow-oval, often rod-like, longitudinal, 15–26 × 2–6. Anal shield of varying form (Figs. 7, 7–9), usually oval with rounded anterior margin, weakly reticulate; its length (56–87) usually greater than its width (52–67), lAn/wAn 0.95–1.40; paranal setae (24–36) slightly shorter than postanal seta (30–44); opening of gland gv3 small, poorly discernible; cribrum of usual width (12), but often looking narrow if being folded caudally. Exopodal plate-

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lets between coxae II–III and III–IV rudimentary. Peritrematal shield of uneven width, becoming a very narrow strip alongside coxa II, then much expanding into triangular projection between coxae I–II, and dorsally into area where its anterior end free or narrowly connected to dorsal shield beside seta z1 (Fig. 7, 14); posterior end of peritrematal shield connecting with exopodal platelet enveloping coxa IV posteriorly; lyrifissures ip1–3 and gland gp present. Peritreme much shortened (usually 32–52 × 6–8), not extending beyond mid-level of coxa III anteriorly (Figs. 4, 2; 7, 11), rarely ending at level of anterior margin of coxa III (Fig. 7, 12) or posterior quarter of coxa II (Fig. 7, 13), in such cases its length 64–92. Spermathecal apparatus weakly discernible as a long (ca 100 when fully extended), slender (0.5–1.0) simple duct, widened (ca 2) along its terminal 5–10 below level of epigynal shield. Gnathotectum (width 28–37) usually with three tines of similar length (17–26), middle tine (17–20) often thinner and inserted slightly more ventrally than lateral ones (Fig. 7, 16), rarely dilated apically (as in Figs. 5, 7; 7, 17). Subcapitulum about as long (88–107) as wide (83–107). Deutosternum with 7 rows of small denticles (8–20 denticles in each row); groove width 9–12 (Fig. 7, 10). Hypostomatic pairs of setae hp1 (23–35) and hp3 (27–38) longer than hp2 (13–20) and basal pair pc (20–28); all setae simple, attenuate. Corniculi of common proportions, moderate-sized (31–39 × 11–14), lCo/lD 6.90–9.14%. Internal malae subequally as long as corniculi, sparsely fimbriate, gradually tapering to tip (Figs. 6, 1; 7, 10). Chelicera not large, its length without basal segment 100–120; cheliceral digits of moderate size (38–47, lCh/lD 8.60– 11.35%), longer than corniculus (lCh/lCo 1.17–1.44). Fixed digit of chela fully developed, with a row of 7–12 denticles in paraxial position (of which the most apical one is larger), and ending with two offset subapical teeth and the apical hook (Figs. 5, 1, 2); setiform pilus dentilis minute. Movable digit (39–43) similar in length to fixed one, bidentate. Palp length 144–172; internal seta of trochanter (29–33) longer than external seta (20–25); of the usual specialized setae, femur al thick with oblique tip and genu al1 and al2 thick, spatulate, with oblique tip (Fig. 6, 8). Legs of moderate length (I 378–472, II 272–330, III 236–305, IV 360–456); leg I similar in length to dorsal shield. Length of tarsi I 104–136, II 70–96, III 69–92, IV 104–141. Distal margin of coxa I dorsally and coxa III posteriorly serrate (Fig. 7, 14). Tarsi generally of moderate length (lTIV/lD 0.25–0.31) and width

(lTIII/wTIII 3.2–4.0). Leg chaetome typical for genus and stable in specimens studied; seta pl on tibia IV present. Setae of legs simple, generally of moderate length, seta pd2 on genu II slightly shorter (26–34) than width of segment (35–40). Length of two more erect and attenuated setae pd3 and pd4 on telotarsus and basitarsus IV, respectively, subequal in length (60–74), slightly exceeding length of tibia IV (52–62); tarsus IV setae av2 and mv (50–60) and md (48–55) also moderately long. Ambulacrum I on pedicellate base, claws I (7–10) smaller than claws II–IV (11–13). Tarsus I apically with 5 rod-like solenidia; length of sensillum with lanceolate apex 28–35. Ambulacra of legs II–IV (length 20–26) with short paradactyli (10–15). Apical setae on tarsi II–IV ad1 and pd1 of moderate length (20–24). Four subapical setae on tarsi II–IV evenly distant from apex or ventral pair inserted slightly more distally than lateral pair; ventral setae av1 and pv1 (15–21) slightly thicker and shorter than lateral setae al1, pl1 (22–28) (Fig. 4, 4). Male. Dorsal shield 348–420 × 160–196, lateral incisions (usually 15–30) similar to those on females. Podonotal and opisthonotal regions with same setation as on female (z3 present, S2 absent, only one pair of setae present in area usually occupied by J4 and Z4). Other than being about 10–15% shorter, relative lengths of dorsal shield setae as in female; Z5/(Z5–Z5) 0.87–1.36. Marginal setae lengths 15–28 (r3 longest); in series r 4 setae, in series R 4–5 (rarely asymmetrically 3) setae. Sternitigenital shield united with endopodal platelets developed between coxae I–II and coxae II–III, but separate from (Fig. 5, 5) or sometimes touching anterior tip of endopodal strip between coxae III–IV (Figs. 6, 2, 3); length of shield 156–192, width between levels of setae st2 and st3 96–116, width on level of setae st5 33–41. Sternitigenital shield with setae st1–5 (st5 sometimes asymmetrically off shield corner) and lyrifissures iv1–3; setae st1–3 (20–27) longer than st4–5 (14–20). Ventrianal shield partly or entirely reticulated, usually elongate-oval, with anterior margin convex (Fig. 5, 5), with 3–5 pairs of opisthogastric setae (ZV1 sometimes asymmetrically rudimentary or absent, JV4 and JV5, rarely also JV3 and ZV2 on soft cuticle) (Figs. 6, 5–7, 9–11); ventrianal shield slender, often extending laterally only to longitudinal midlevel of coxal cavities IV, not approaching free posterior metapodal plates (11–22 × 2–7); length of shield (128–156) about 1.5 (1.26–1.89) times width ENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011


(72–100); paranal setae (24–28) clearly shorter than postanal seta (31–36). Caudal setae JV5 (24–30) usually longer and ZV1–2 (15–21) often shorter than other opisthogastric setae (17–28). Gnathotectum with tines similar to female (Fig. 5, 6) but somewhat shorter (middle tine 13–22) and sometimes variously irregular in different ways (Figs. 5, 7, 8). Corniculi proportionately similar to those of female (26–32 × 10–12) but apices more splayed, lCo/lD 7.1–8.6%. Internal malae similar in form to female (Fig. 6, 1). Chelicera length without basal segment 90–94, cheliceral digits (34–40, lCh/lD 9.0–10.3%) longer than corniculus (lCh/lCo 1.12– 1.34). Fixed digit of chela fully developed, with a row of 7–10 denticles, but lacking offset subapical teeth near apical hook (Fig. 5, 4). Movable digit with one large denticle well removed from apical hook and with sinuate spermatodactyl (length 20–22), protruding 12–14 beyond tip of digit and not tapering to apex (Figs. 5, 3, 4). Length of legs I–IV 380–408, 260–280, 232–264 and 340–376, respectively. Length of tarsi I–IV 100– 116, 64–80, 67–74 and 100–116, respectively. Leg II slightly stouter than on female, and with four dimorphically modified, opposable setae ventrally (Fig. 5, 9); femur with v2 stout basally (width 5–7) and attenuate apically, on elevated base; telotarsus with mv spinelike, on elevated base; genu and tibia with av usually slightly spinelike, shorter than on female. Other leg structure and setation as in female. Differential diagnosis. Adults of this species are very similar to those of another arctic species, Arctoseius multidentatus Evans 1955, in the sinuate form of spermatodactyl, narrow ventrianal shield, and dimorphism of leg II in males and the multidenticlate fixed cheliceral digit of both sexes. Adults of A. haarlovi sp. n. are easily distinguished by having shorter peritremes than in any other known species of Arctoseius. They differ from A. multidentatus in having shorter corniculi, lacking one of the two pairs of opisthonotal setae in the area usually occupied by J4 and Z4, and in being only two-thirds as large in body size. Distribution. Circumpolar, arctic, throughout tundra zone, in polar deserts and subarctic mountains (Fig. 9). Ecology. A. haarlovi sp. n. inhabits the broad spectrum of biotopes in all subzones of the tundra zone. It was found in dry lichen heaths, gramineous and herbENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011

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grass meadows, in sedge-moss and shrub tundra communities, and wet willow beds. Records within the polar desert zone are associated with bird perches, whereas this species has not been found in lemming colonies having similar modified vegetation. These observations possibly point to the bringing of mites to the northernmost arctic islands by birds. Outside the Arctic, A. haarlovi sp. n. has been found in the northern taiga subzone in a treeless karst ravine with cold soil and in the subarctic highlands—in the upper timberline and subalpine meadow. Variability. Variational peculiarities among specimens at hand include asymmetrical irregularities of the gnathotectum (Figs. 5, 8; 7, 19), asymmetrical obliteration or extension of the lateral incision of dorsal shield of adult female and male (Figs. 7, 1–3), asymmetrical losses of individual dorsal setae (j4, z1, s1, s2, J3, J5, S1, S5) or one or another of the posterior marginal setae and opisthogastric seta ZV1, and the presence of an unpaired opisthonotal seta in the J4 position. Some variation in length of peritreme (Figs. 7, 11–13), in form of the female epigynal (Figs. 7, 4–6) and anal shields (Figs. 7, 7–9) and the male ventrianal shield (Figs. 6, 5–7, 9–11) are also evident. In one instance the presence of a rudimentary tritosternum was observed on a female (Fig. 4, 5). The leg chaetome is generally stable, with only one record of the symmetrical omission of seta av on genu III in a female. Etymology. The species is named after the Danish acarologist, Niels Haarløv, who first collected material of this species during the autumn and winter of 1939– 40 from Greenland (Haarløv, 1942). TAXONOMICAL REMARKS The status of the type species of the genus Arctoseius, A. laterincisus Thor 1930, is yet unclear. The type material of this species no longer exists (Lindquist, 1964; Winston, 1999). Subsequent to its original description, this species has not been found in a variety of material collected from Svalbard, although 8 other species of Arctoseius are known already from the archipelago (Byzova et al., 1995; Makarova, 2000b; Coulson, Refseth, 2004; Coulson, 2007). Confusion of Arctoseius haarlovi with A. laterincisus began when Haarløv (1942) described and illustrated a species questionably under the name “A. laterincisus (?)” that he collected from Greenland. His discussion of morphological differences between his

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specimens and those described by Thor (1930) also supported his doubt. Lindquist (1961) reviewed the possibility that Thor and Haarløv were not dealing with the same species, but Haarløv’s initial element of doubt in the correct identification was increasingly ignored in subsequent references to A. laterincisus, such that keys to species of Arctoseius distinguish A. laterincisus primarily on two adult attributes, the great size of the dorsal shield, based on Thor’s measurements, and the reduced length of the peritreme, based on Haarløv’s illustration (Bernhard, 1963; Bregetova, 1977; Karg, 1971, 1993). However, the differences noted by Lindquist (1961) among data presented by Thor and Haarløv alone sufficed to assume that two species were involved. Thor’s description and illustrations were of specimens uniquely large in size of idiosoma (750–800 long, 380–400 wide), with elongated posterior body setae (Z5 and JV5 distinctively longer than adjacent paired setae), less reduced peritremes (in the photo seeming to reach the anterior margins of coxae III), and with the female epigynal shield pictured as broad, somewhat wedge-shaped, with nearly straight lateral margins. In contrast, Haarløv dealt with adults scarcely more than half as large (idiosoma 420– 496 long, ca 240 wide), without elongated posterior body setae, with more reduced peritremes (not extending beyond the mid-level of coxae III), and with the female epigynal shield more bulb- or vase-shaped, with convex posterior margin and concave lateral margins. Although Thor’s primary type material of A. laterincisus no longer exists, the male ventrianal shield was described as large, nearly reaching the lateral margins of the body (which is typical of many species of the genus), whereas in males now available of Haarløv’s species, this shield is notably narrow, not even approaching the level of the freely standing metapodal plates. Thor noted the corniculi to be strongly formed in A. laterincisus, and one may suppose that they perhaps are similar to the elongated form in A. sexsetus and A. multidentatus, whereas the cornuculi are of the usual shorter form in Haarløv’s species. Although remaining enigmatic in many morphological respects, Thor’s Arctoseius laterincisus may be a member of a species group that includes Arctoseius multidentatus Evans 1955, A. sexsetus sp. n., one undescribed new arctic species (Arctoseius sp. aff. multidentatus, its description in preparation), and perhaps A. wisniewskii Gwiazdowicz & Kamczyс 2009, based on the following attributes: dorsal shield not reticu-

lated, rather narrow (lD/wD 1.9–2.6); a few posterior idiosomatic setae elongated (at least Z5 and JV5 longer than adjacent setae, and the paranal setae nearly as long as the postanal one); peritremes shortened, not exceeding mid-level of coxae II; and corniculi more or less elongated (lCo/lD usually 8–13%). In other respects, however, it is A. haarlovi that may have a sister-relationship with A. multidentatus, based on other apomorphies: fixed cheliceral digit multidentate, with row of 7 to 12 teeth; male spermatodactyl sinuate in lateral view; male leg II with dimorphically modified setae ventrally on femur, tibia, tarsus; male ventrianal shield slender, extending laterally only to longitudinal midlevel of coxal cavities IV, about 1.3–1.5 longer than wide. The local gamasid mite faunas within the Arctic usually number 8–13 species of Arctoseius and many of them may be very similar superficially. Moreover, quite often one can reveal the co-existence of several (up to 7) species of Arctoseius in the same biotope, and even up to 5 species in one soil sample (Lindquist, 1961; Makarova, 1999, 2000a, 2011). For this reason, although the genus Arctoseius was established by S. Thor based on a presumed monotypy, one may wonder whether the type series included specimens of two (or several) morphologically similar species, e.g. A. multidentatus and Arctoseius sp. aff. multidentatus. These species in some tundra areas were recorded coexistent with A. haarlovi, in one soil core by us. The males of Arctoseius sp. aff. multidentatus possess the rather wide form of ventrianal shield that incorporates the metapodal plates. However, until detailed investigations of gamasid mites on Svalbard are conducted, the question of the identity of Arctoseius laterincisus Thor 1930 will remain open. Seven specimens of A. haarlovi (6 females and 1 male) from three areas of the Atlantic sector (SW Greenland, Finnish Lapland, and Arkhangelsk Region, Russia) are distinguished from other adults by some divergent morphological features. Their peritremes are significantly longer (in females averaging twice as long, 64–92 instead of 32–52), and extending to the transverse level of the anterior margin of coxae III (Fig. 7, 12) or even to the level of the posterior quarter of coxae II (Fig. 7, 13). Dorsal setae s2, usually situated at least asymmetrically on the cuticle (Fig. 4, 1), in these specimens are inserted symmetrically on the dorsal shield margin. Mites collected from the north and east of Greenland do not present these peculiarities. One may speculate whether these distinctions ENTOMOLOGICAL REVIEW Vol. 91 No. 8 2011


support the hypothesis of J.A. Downes (1988), that the faunas of the northern and southern parts of Greenland have been derived mainly from different sources, namely from Queen Elizabeth Islands and from boreal North America and boreal Europe, respectively.

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ACKNOWLEDGMENTS Numerous colleagues provided us with material from which additional specimens of new species were found—A.B. Babenko, A. Fjellberg, J. Böcher, V.I. Bulavintsev, I.I. Marchenko, M.B. Potapov, G. Sovik, A.V. Uvarov, or with information— V.M. Behan-Pelletier, H.V. Danks. Tables of pictures and maps have been prepared for publication with the invaluable participation of K.V. Makarov. The accuracy of phrases in a translation of the Russian and English versions of this article was checked by V.V. Grebennikov. The Carlsberg Foundation (Denmark), the Russian Fund of Basic Research, and the Programs “Biosphere Origin and Evolution” and “Biodiversity” of the Russian Academy of Sciences supported the second author during certain aspects of this work. We are very grateful to all.

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