Zootaxa 2309: 30–42 (2009) www.mapress.com / zootaxa/
ISSN 1175-5326 (print edition)
Article
Copyright © 2009 · Magnolia Press
ZOOTAXA ISSN 1175-5334 (online edition)
A revision of the genus Sierraphytoptus Keifer 1939 (Eriophyoidea, Phytoptidae) PHILIPP E. CHETVERIKOV1 & SOGDIANA I. SUKHAREVA Department of Invertebral Zoology, Saint-Petersburg State University, Faculty of Biology and Soil Science; Universitetskaja nab., 7/9, 199034, St. Petersburg, Russia. E-mail:
[email protected]
Summary Comparative analysis of the eriophyoid mites of the genus Sierraphytoptus Keifer 1939 (= Fragaricoptes Roivainen, 1951) (Phytoptidae: Sierraphytoptinae: Sierraphytoptini) living on strawberry (Fragaria spp.) found that they are represented by two species: Sierraphytoptus setiger (Nalepa, 1894) (= Phyllocoptes setiger Nalepa, 1894) and Sierraphytoptus ambulans sp. n. Sierraphytoptus setiger forms red galls on the leaves of Fragaria viridis Duch., whereas the vagrant species S. ambulans causes no visible damage and lives mostly on the lower leave surface of Fragaria vesca L. but sometimes on F. viridis also. A supplementary description of S. setiger from North-West Russia and a key to all species of the tribe Sierraphytoptini are given. A new combination Austracus taiwanensus (K.-W. Huang 2006) comb. n. (removed from the genus Sierraphytoptus) is proposed. Key words: eriophyoid mites, new species, taxonomy, Sierraphytoptus setiger, Sierraphytoptus ambulans
Introduction The evolution of mites of the superfamily Eriophyoidea is strongly related with the evolution of their hostplants (Sukhareva 1992, 1994; Oldfield 1996; Lindquist & Oldfield 1996; Bagnjuk et al. 1998). The family Phytoptidae Murray, 1877 includes the most ancient representatives of Eriophyoidea, which retain the plesiomorphies of more than two setae on the prodorsal shield (ve and sc) and having the subdorsal seta (c1) and solenidion (φ) on tibia I. According to the concept of Sukhareva (1992, 1994) and Bagnjuk et al. (1998), this family represents an early evolutionary lineage of eriophyoids on Angiosperm plants and includes equally annulated (subfamilies Phytoptinae Murray, 1877 and Novophytoptinae Roivainen, 1953) and diversely annulated forms (subfamily Sierraphytoptinae Keifer, 1944). The equally annulated phytoptine mites have an elongated worm-like body and a relatively small and rounded prodorsal shield. Critical analysis of the subfamily Phytoptinae and some data concerning morphology and biology of the mites of the genus Phytoptus from sedges and endoparasitic mites from the subfamily Novophytoptinae were given in our previous papers (Tchetverikov 2004; Chetverikov & Sukhareva 2007; Petanović et al. 2007; Chetverikov et al. 2009). The diversely annulated Sierraphytoptinae are characterized by their compact fusiform body and relatively large prodorsal shield. This subfamily includes about 20 species, grouped in two tribes: Mackiellini Keifer, 1939 (inhabit Arecaceae [palms]) and Sierraphytoptini Keifer, 1944 (Amrine et al. 2003; Navia et al. 2007), representing the evolutionary lineage of Phytoptidae found on dicotyledonous plants. According to Amrine et al. (2003), the tribe Sierraphytoptini includes three monotypic genera — Neopropilus Huang, 1992, Austracus Keifer, 1944 and Fragariocoptes Roivainen, 1951 plus Sierraphytoptus Keifer, 1939, with the type species S. alnivagrans Keifer, 1939 and S. taiwanensus K.-W. Huang, 2006, recently described from Taiwan. Up to now, mites of this tribe were found on plants from four orders: Trochodendrales, Malpighiales, Fagales and Rosales (Sukhareva 1994; Amrine et al. 2003; Huang 2006). Only two species of the tribe Sierraphytoptini [Fragariocoptes setiger (Nalepa, 1894) from Fragaria spp.
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Accepted by O. Seeman: 18 Nov. 2009; published: 11 Dec. 2009
and S. alnivagrans from Alnus glutinosa (L.)], have been recorded in Europe (Roivainen 1951; Petanović 1988; Amrine et al. 2003). In this paper, we present our analysis of published data concerning the species “setiger”, and new data from original material from North-West Russia and South Siberia of Russia. We show that the F. setiger of previous authors (Nalepa 1894; Liro 1941; Roivainen 1951; Farkas 1965; Amrine et al. 2003) comprises two separate species of Sierraphytoptus.
Material and methods Mites were collected during May–October 2008 and May–September 2009 in North-West Russia (Leningrad Province), Ural region (Sverdlovsk Province) and in South Siberia (Altajskiy kraj and Zabaykalsky krai of Russia) from leaves of Fragaria vesca and Fragaria viridis using a fine pin and mounted on slides in Berlese medium. The morphological terminology and systematic classification follows that of Lindquist (1996) and Amrine et al. (2003). Measurements of mites were made according to Amrine and Manson (1996). All measurements are given in micrometers (µm) and, unless stated otherwise, refer to the length of the structure. Each measurement of the holotype precedes the corresponding range for paratypes (given in parentheses). The nomenclature of host plants follows Kamelin (2001) and Sennikov (2006). We also collected and reared mites in the laboratory (18–20°C, 70 % air humidity, natural illumination). About 30 plants of Fragaria vesca infested by Sierraphytoptus were dug out with a lump of soil on 1 May 2009 near the village of Vyritza (Leningrad Prov., Russia) and transferred to laboratory. On 25 July 2009, we did the same with 20 plants F. viridis infested by Sierraphytoptus from Luga district of Leningrad Prov. (Russia). Every 10 days we collected and slide-mounted mites from these plants to provide specimens for comparative morphological analysis.
Family Phytoptidae Murray, 1887 Subfamily Sierraphytoptinae Keifer, 1944 Tribe Sierraphytoptini Keifer, 1944 Key to genera and species of the tribe Sierraphytoptini 1. 2. 3. 4. 5.
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Solenidion on tibia I present ......................................................................................................................................... 2 Solenidion on tibia I absent (Sierraphytoptus Keifer, 1939) ........................................................................................ 4 Prodorsal shield with one pair of setae ve (Neopropilus Huang, 1992)................................ N. jatrophus Huang, 1992 Prodorsal shield with two pairs of setae, ve and sc (Austracus Keifer, 1944) ............................................................ 3 Empodium 4-rayed, 16–18 dorsal annuli, 2 annuli between tubercles of c1 and posterior shield margin ..................... ........................................................................................................................................ A. havrylenkonis Keifer, 1944 Empodium 8-rayed, 37 dorsal annuli, 5–7 annuli between tubercles of c1 and posterior shield margin ....................... ............................................................................................................... A. taiwanensus (K.-W. Huang 2006) comb. n. Empodium 2-rayed, 16–17 dorsal annuli ..........................................................................S. alnivagrans Keifer, 1939 Empodium 4-rayed, more than 20 dorsal annuli.......................................................................................................... 4 Suboral plate before coxae I with three longitudinal lines (Fig. 2 CG), 35 (32–38) dorsal annuli, 5 annuli between tubercles of c1 and posterior shield margin, c1 31 (22–36) long, frontal lobe minute, not more than 1 μm ................. ................................................................................................................................................ S. setiger (Nalepa, 1894) Suboral plate before coxae I with two longitudinal and one transverse lines (Fig. 3 CG), 24 (22–27) dorsal annuli, 3 annuli between tubercles of c1 and posterior shield margin, c1 16 (13–19) long, frontal lobe 4 (3–5) long ................ ......................................................................................................................................................... S. ambulans sp. n.
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Genus Neopropilus Huang, 1992:228 Neopropilus jatrophus K.-W. Huang, 1992:228, fig. 29–34 Type host: Jatropha curcas L. (Euforbiaceae) Relation to host: vagrant on the lower leaf surface Distribution: Taiwan
Genus Austracus Keifer, 1944:20 Austracus havrylenkonis Keifer, 1944:20, fig. 182 Type host: Nothofagus dombeyi (Mirb.) Blume (Fagaceae) Relation to host: mites cause galls on fruits Distribution: South America (Argentina) Austracus taiwanensus (K.-W. Huang 2006) comb. n. Sierraphytoptus taiwanensus K.-W. Huang 2006:63, fig.1 Type host: Trochodendron aralioides S. et Z. (Trochodendraceae) Relation to host: vagrant on the lower leaf surface, no damage observed Distribution: Taiwan Comparative material. One female and one nymph of A. taiwanensus comb. n. from T. aralioides. The mites are vagrant on the lower leaf surface, causing no visible damage. Slide # 030416-01 (TAIWAN: Nantou Renai, 16.IV.2003, leg. K.W. Huang) and slide # 961031-17 (TAIWAN: Taichung Anmashan, 31.X.1996, leg. C.F.Wang).
Genus Sierraphytoptus Keifer, 1939:151 Sierraphytoptus Keifer, 1939:151, fig. LI; 1965:8, fig. 4; Newkirk, Keifer 1975:567; Amrine et al., 2003:17, fig. 18. Fragariocoptes Roivainen, 1951:51 [Synonymy by Roivainen (1953, p.85)]; Farkas, 1965:8
Type species: Sierraphytoptus alnivagrans Keifer, 1939, by monotypy. Diagnosis: opisthosoma diversely annulated; prodorsal shield with four setae – ve and sc; a pair of setae c1 on the dorsal part of opisthosoma; solenidion on leg I, II missing. Species included: Sierraphytopyus alnivagrans Keifer, 1939, S. ambulans sp. n., S. setiger (Nalepa, 1894) Distribution and hosts. Mites of the genus are recorded from North America, Asia (Armenia, Ural region and South Siberia of Russia, China) and Europe. Host-plants of these mites include dicots from the orders Fagales (Alnus spp.) and Rosales (Fragaria spp.). Remarks. The species S. taiwanensus from T. aralioides is removed from the genus because it possesses a solenidion on tibia I, while in Sierraphytoptus it is missing. The tibial solenidion was not measured and figured in the description of S. taiwanensus given by K.W. Huang (2006), but the female (slide # 030416-01, Taiwan, Nantou Ren-ai 2003/04/16, coll. K.W. Huang) and nymph (slide # 961031-17, Taiwan, Taichung Anmashan, 1996/10/31, coll. C.F. Wang) of this mite do have a tibial solenidion (Fig. 1). Therefore, the morphological features of S. taiwanensus correspond to the diagnosis of the genus Austracus Keifer, 1944 and we transfer it to this genus: Austracus taiwanensus (K.-W. Huang 2006) comb. n. Mites of the genus Sierraphytoptus were first found by А. Nalepa (1894) inside red galls on leaves of F.
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viridis and described under the name Phyllocoptes setiger (Table 1). Later, Liro (1941) found similar mites on F. vesca in Finland. He identified them as P. setiger and suggested that there were two subspecies, which remained unnamed: one formed red galls on leaves of F. viridis and lived inside them, but the other lived on the lower surface of leaves on F. vesca, causing no visible damage. Liro (1941) also wrote that these subspecies differed in the number of dorsal annuli (Table 1) and that they could be separate species. The same suggestion was made by Bagdasarian (1981), who gave detailed descriptions of vagrant nymphal and female S. setiger from Fragaria sp. in Armenia. TABLE 1. Some characteristics of the “setiger” species from Fragaria spp. according to different authors. Characteristics
Reference Nalepa, 1894
Liro, 1941
Roivainen, 1951
Farkas, 1965
Bagdasarian, 1981
Genus
Phyllocoptes
Phyllocoptes
Fragariocoptes
Sierraphytoptus
Sierraphytoptus
Location
Austria
Finland
Finland
Hungary
Armenia
Date of collection not stated
22.VIII.1940
12.IX.1949
not stated
8.VI.1965; 9,30.III.1966, 10.IV.1966; 12,23.VI.1966; 4.VIII.1966; 22.VII.1967
Host-plant
Fragaria viridis
Fragaria vesca
Fragaria viridis
Fragaria viridis (= F. collina)
Fragaria sp.
Relation to host
cause red galls on the leaves
vagrant on the lower leaf surface
vagrant on the lower cause red galls on the leaf surface leaves
vagrant on the lower leaf surface, mainly along the central vein
Number of dorsal annuli
35*
21–24
25*
34
23–25
3*
2*
5*
3*
Number of annuli 6* between rear shield margin and tubercles of c1
* The data are given according to the figures presented in the cited papers.
FIGURE 1. Austracus taiwanensus (K.-W. Huang, 2006) comb. n. L1-f—leg I of female, L1-n—leg I of nymph. Scale: 30 µm.
Ten years after Liro found phytoptid mites on Fragaria, these mites were recorded in Finland again by Roivainen (1951). He thought these mites were very close to the vagrant mites P. setiger referred to by Liro, but differed from the mites collected by Nalepa from galls (Table 1). Roivainen (1951) also noted that both forms (gall-forming and vagrant) of the species “setiger” have four setae on their prodorsal shield and a pair
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of subdorsal setae (c1) on the dorsal surface of opisthosoma that did not fit to the characteristics of the genus Phyllocoptes. On this basis he established a new monotypic genus, Fragariocoptes Roivainen, 1951, for the species “setiger”. However, by that time the genus Sierraphytoptus Keifer, 1939 had already been established, to which this species could be placed. Therefore Roivainen (1953, p.85) synonymised the genus Fragariocoptes with Sierraphytoptus and transferred the species “setiger” into it. Therefore, previous authors have considered the possibility that S. setiger comprises two species, one forming galls and the other vagrant, but have never split the species. Afterall, another possibility is that both forms represent the deutogyne and protogyne of the same species. For this to be true, we would be able to rear deutogynes from the protogynes. We would also expect to find both forms of mites throughout the year, especially summer. In North-West Russia there are three species of Fragaria: two widespread (F. vesca and F. moschata Duch.) and one much rarer (F. viridis) (Tzvelev 2000). It is remarkable that during 2008–2009 we looked for Seirraphytoptus on F. moschata many times and never found them. Our field collections found Sierraphytoptus only from lower surface of leaves on F. vesca and from red leaf galls on F. viridis during spring, summer and autumn. The vagrant and gall-forming mites distinctly differed in their number of dorsal annuli, length and form of body and design of prodorsal shield (Fig. 2, 3; Tables 1, 2). During summer, both forms were numerous and laid eggs intensively. Furthermore, during our field work they were never found simultaneously together either on the leaf surface or in galls. Moreover, the mites reared in our laboratory revealed no morphologically distinguishable forms of females living on the same species of Fragaria: all the mites on F. vesca from Vyritza were of the vagrant type, whereas on F. viridis from Luga were of the gall-making type. Therefore, the above evidence shows that the two forms are not deutogyne and protogyne. After morphological analysis and comparison of our original data (Table 2) with published data we conclude that the vagrant mites and mites forming galls from Fragaria spp. are the two separate species: S. ambulans sp. n. and S. setiger.
Sierraphytoptus alnivagrans Keifer, 1939 Sierraphytoptus alnivagrans Keifer, 1939:152, fig. LI; Newkirk, Keifer 1975:567 Host: Alnus incana (L.) Moench. subsp. tenuifolia (Nutt.) Breitung; A. glutinosa (L.) (Betulaceae) Relation to host: vagrant on the lower surface of leaves along the veins and on the midrib Distribution: North America (USA, California), Bosnia (former Yugoslavia) (Petanović 1988) and Serbia (19°25'11'' E, 43°54'51'' N; Mt. Tara, 13.06.2007) (Petanović, personal communication).
Sierraphytoptus ambulans sp. n., Fig. 2 Phyllocoptes setiger Liro, 1941:42, fig. 31 Fragariocoptes setiger Roivainen, 1951:51, fig. 22 Sierraphytoptus setiger Bagdasarian and Pogosova, 1976:223–226, fig.1; Bagdasarian 1981:26–29, fig. 4
Female (holotype). Body fusiform, light orange, 178 (155–201), 56 (52–59) wide, 57 (54–59) thick. Prodorsal shield with two curved admedian lines at the end connected with four short lines. Median line missing. Thin transverse line near posterior shield margin (Fig. 2 D, AD). Prodorsal shield 40 (36–44), 49 (46–52) wide, frontal lobe 4 (3–4). External vertical setae ve 9 (8–10), their tubercles 27 (26–28) apart, situated in a small hollow under the antero-lateral shield margin; scapular setae sc 6 (6–8), directed upward and centrally, their tubercles 18 (15–19) apart. Gnathosoma 20 (19–21), directed downward. Dorsal pedipalp genual setae d simple.
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TABLE 2. Measurements of females of two Sierraphytoptus species from Fragaria (original data from Leningrad Prov., Russia). Characteristics
S. setiger (n=10)
S. ambulans (n=10)
Min-max
Mean ± SD
Min–max
Mean ± SD
length of body
185–247
220 ± 20.0
155–201
174 ± 14.2
width of body
63–70
67 ± 3.6
52–59
57 ± 2.5
thickness of body
62–72
67 ± 4.1
54–59
57 ± 2.5
length of prodorsal shield
36–40
38 ± 1.8
36–44
41 ± 2.1
width of prodorsal shield
41–46
44 ± 2.7
46–52
50 ± 2.1
length of gnathosoma
18–21
19 ± 1.1
19–21
20 ± 1.0
length of ve (s.d.1)
7–9
8 ± 0.7
8–10
9 ± 0.7
distance between tubercles of ve
24–26
25 ± 1.2
26–28
26 ± 0.9
length of sc (s.d.2)
4–6
5 ± 0.6
6–8
7 ± 0.7
distance between sc
15–20
18 ± 2.1
15–19
17 ± 1.0
distance between tubercles of ve and sc
14–16
15 ± 0.8
14–16
15 ± 0.7
length of c1
22–36
31 ± 4.3
13–19
16 ± 1.6
number of annuli between rear shield margin and tubercles of c1
4–5
5 ± 0.5
3–3
3 ± 0.0
length of leg I
31–33
32 ± 1.0
27–31
30 ± 1.5
length of tibia I
5–6
6 ± 0.5
5–5
5 ± 0.0
length of l' (s. tib. I)
3–5
4 ± 0.8
2–5
3 ± 1.2
length of tarsus I
5–7
6 ± 0.7
6–7
7 ± 0.5
length of ω (claw I)
10–11
11 ± 0.4
9–11
10 ± 0.6
length of leg II
29–31
29 ± 0.7
25–29
27 ± 1.1
length of tibia II
4–5
5 ± 0.3
4–6
5 ± 0.6
length of tarsus II
5–6
6 ± 0.5
5–7
6 ± 0.7
length of ω (claw II)
9–10
10 ± 0.5
9–10
10 ± 0.5
length of epigynium
11–14
13 ± 1.1
12–16
14 ± 1.2
width of epigynium
20–21
20 ± 0.7
16–20
17 ± 1.1
length of 3a (genital seta)
14–19
16 ± 1.3
14–19
16 ± 1.5
length of 1b (s.cox. I )
6–10
8 ± 1.5
7–10
9 ± 1.6
distance between tubercles of 1b
9–10
10 ± 0.7
9–10
10 ± 0.5
length of 1a (s.cox. II )
15–26
20 ± 3.9
12–23
16 ± 3.2
distance between tubercles of 1a
9–10
10 ± 0.7
8–9
9 ± 0.5
length of 2a (s.cox. III)
24–41
34 ± 6.1
31–38
34 ± 2.8
distance between tubercles of 2a
23–24
23 ± 0.7
18–22
20 ± 1.7
length of sternal line
11–13
12 ± 0.9
9–11
10 ± 0.8
length of c2 (s.l)
26–35
31 ± 2.4
31–37
34 ± 2.3
length of d (s.v.1)
26–31
28 ± 1.7
30–36
32 ± 1.9
length of e (s.v.2)
21–26
23 ± 2.3
16–23
20 ± 1.9
length of f (s.v.3)
23–31
28 ± 2.6
24–27
26 ± 0.9
length of h1 (s.acc.)
2–3
2 ± 0.5
2–4
3 ± 0.7
number of dorsal annuli
32–38
35 ± 2.0
22–27
24 ± 1.3
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FIGURE 2. Sierraphytoptus ambulans sp. n., female. D—female in dorsal view, AD—prodorsal shield, CG— coxogenital region, em—empodium and tarsal solenidion (enlarge), L1—leg I, L2—leg II. Scale: D= 60 µm; AD, CG, L1, L2 = 40 µm; D = 20 µm.
Leg I 30 (27–31), tibia 5 (5–5), l' - 3 (2–5), tarsus 6 (6–7), ω - 10 (9–11), empodium simple, 4-rayed; leg II 26 (25–29), tibia 5 (4–6) long, l' absent, tarsus 6 (5–7) long, ω - 9 (9–10) long, without knob, empodium simple, 4-rayed (Fig. 2 L1, L2, em). Setae bv present. Coxae with numerous thin short lines. Sternal line 9 (9– 11), bifurcated anteriorly. Rounded plate with two longitudinal lines and one transverse line situated in front of coxae I (Fig. 2 CG). Setae 1b 8 (7–10), 10 (9–10) apart; 1a 17 (12–23), 8 (8–9) apart; 2a - 33 (31–38), 21
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(18–22) apart. Genitalia 13 (12–16), 17 (16–20) wide; genital coverflap with 8–12 longitudinal ridges situated anteriorly (Fig. 2 CG); setae 3a 16 (14–19). Opisthosoma with 23 (22–27) slightly serrated smooth dorsal annuli, 53 (48–55) microtuberculate ventral annuli, 5–6 coxogenital annuli. Setae c1 16 (13–19) on 3 (3) annulus; setae c2 35 (31–37) on 9 (8–11) annulus; setae d 30 (30–36) on annulus 20 (18–24); setae e 19 (16–23) on annulus 32 (28–36); setae f 26 (24– 27) on 4th ventral annulus from rear. Seta h1 present. Male: not found. Type material. Female holotype (slide # 53), 9 females paratypes (on the same slide) from Fragaria vesca L. (Rosaceae) [lower leaf surface, no visible damage observed], RUSSIA: Leningrad Province, Gatchina district, fir-forest on the right bank of river Oredezh near station Vyritza, 59°23.6' N, 30°18.3' E, 21 September 2008, collected by Ph. E. Chetverikov. Type material is deposited in the collection of the Laboratory of Phytoacarology (Department of Invertebrate Zoology, Biological and Soil Sciences Faculty, Saint Petersburg State University, Russia). Additional material. 2 deutogyne female and 11 protogyne females (slide # 3-09) from F. vesca [lower leaf surface, no visible damage observed], RUSSIA: Leningrad Province, Gatchina district, fir-forest on the right bank of river Oredezh near station Vyritza, 59°23.6' N, 30°18.3' E, 19 May 2009, leg. Ph. E. Chetverikov; 1 deutogyne female, 3 protogyne females and 2 nymphs (slide # 3/1-09), the same host, locality and leg., 2 June 2009; 16 protogyne females and 8 nymphs (slide # 39-09), the same host, locality and leg., 26 July 2009; 38 protogyne females and 12 nymphs (slides # 32-08 and # 53-08), the same host, locality and leg., 21 September 2008. 2 protogyne females (slide # 6-09) from F. vesca. [lower leaf surface, no visible damage observed], RUSSIA: Leningrad Province, Kingisepp district, fir-forest on the bank of Kopanskoye Lake, 59°70.0' N, 28°70.8' E, 10 June 2009, leg. Ph. E. Chetverikov; 29 females and 12 nymphs (in slides ## 35-08, 36-08, 37-08) from F. vesca [lower leaf surface, no visible damage observed], RUSSIA: Leningrad Province, Vyborg district, pine-forest near station Yappilya, 62°38'03'' N, 27°43'08'' E, 7 September 2008 leg. Ph. E. Chetverikov; 22 females and 7 nymphs (in slides ## 42-08, 48-08, 49-08, 50-08 and 51-08) from F. vesca [lower leaf surface, no visible damage observed], the same host, locality and leg., 11 September 2008; 4 overwintered females (slide # 1-09) from F. vesca [lower leaf surface, no visible damage observed], RUSSIA: Leningrad Province, Luga district, pine-forest on the right side of the highway M-20 near village Zheltzy, 58°82'34'' N, 29°91'04'' E, 4 May 2009, leg. Ph. E. Chetverikov; 5 overwintered females (slide # 2-09) from F. viridis [lower leaf surface, no visible damage observed], RUSSIA: Altajskiy kraj, left bank of river Loktevka near village Kurya, 51°59'41'' N, 82°28'43'' E, 2 May 2009, leg. P. G. Efimov; 1 protogyne female, 1 male and 3 nymphs (slide # 39-09) from F. viridis [lower leaf surface, no visible damage observed], RUSSIA: Zabaykalsky Krai, Aginskoye district, birch-forest on the right side of the highway A166, 51°12'75'' N, 114°39'56'' E, 20 July 2009, leg. A.N.Baranova. 5 protogyne females, 1 male and 2 nymphs (slide # 42-09) from F. viridis [lower leaf surface, no visible damage observed], RUSSIA: Ural region, Sverdlovsk Province, suburb of Pervouralsk, 59°98'06'' N, 56°94'03'' E, 07 September 2009, leg. N.Bannikov. Etymology. The specific name is derived from the Latin verb ambulo (“wander”) and is a Partisipium Praesentis Activi (Active Present Participle). Differential diagnosis. The vagrant mite S. ambulans sp. n. is very close to the gall-making mite S. setiger. They are distinguished by the number of dorsal annuli (22–27 in S. ambulans versus 32–38 in S. setiger), length of the setae c1 (13–19 versus 22–36) and length (155–201 versus 185–247) and form of the body (compact fusiform versus elongated, almost worm-like). Besides these species have a different life-style: S. ambulans are vagrant on the lower leaf surface whereas S. setiger form red galls on leaves. Distribution. Finland, North-West Russia, Ural region and South Siberia (Altayskiy Kray and Zabaykalsky Krai) of Russia, Armenia. Lastly, it was realized that the species previously identified as S. REVISION OF THE GENUS SIERRAPHYTOPTUS
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setiger, found 21.VIII.1988 as vagrant on the lower leave surface of F. vesca on Mt. Durmitor, Montenegro (Petanović et al. 1996) belongs to S. ambulans (Petanović, personal communication). Hosts. In North-West Russia and Fennoscandia S. ambulans mainly live on F vesca. To the south of these territories mites S. ambulans also live on F. viridis and one more unidentified species of Fragaria. Deuterogyny of S. ambulans. During 1965–1967 from March till August Bagdasarian and Pogosova (1976) collected this species from Fragaria sp. in Armenia (Table 1) and described deutogyne and protogyne females. According to their data the deutogyne and protogyne females slightly differ only in length and width of the body and length of setae ve and sc (Table 3). From May till August 2009 we observed S. ambulans from F. vesca collected on 1 May and transferred to laboratory in order to get new data concerning their biology. In the beginning of May only single, bright orange, egg-laying deutogyne females were found in the folds of young leaves. Protogyne females then appeared after mid-May. All protogynes were morphologically identical to the deutogynes but they were light orange and it was easier to clarify them in Berlese medium: overwintered females usually were more turbid in slides than summer ones, and the design on their prodorsal shields was less distinct. In June, July and August all mites fed on and laid eggs only on the lower leaf surface; no visible damage was registered on leaves. TABLE 3. Some morphological characteristics of deutogyne and protogyne females of S. ambulans from Fragaria sp. from Armenia (according to Bagdasarian and Pogosova 1976). Characteristics
Deutogyne females
Protogyne females
Date of collection
9, 30 March 1966
8 June 1965
Length of body
150–175
180–220
Width of body
50–60
60–70
Length of ve (s.d.1)
7–8
6–7
Length of sc (s.d.2)
8–9
7–8
Sierraphytoptus setiger (Nalepa, 1894), fig. 3 Phyllocoptes setiger Nalepa, 1894:311, fig. 3,4 Sierraphytoptus setiger Farkas, 1965:8, fig.4; Fragariocoptes setiger Amrine et al., 2003:17, fig.17
Protogyne female (n=10). Body elongated, whitish or pallid pink, 226 (185–247), 67 (63–70) wide, 68 (62– 72) thick. Prodorsal shield with two wavy admedian lines. Median line missing or indistinct. Three short lines forming trident fork-like figure in posterior part of shield (Fig. 3A D). Prodorsal shield 38 (36–40), 43 (41– 46) wide, frontal lobe minute (not more than 1 μm). Setae ve 9 (7–9), their tubercles 25 (24–26) apart, situated in a small hollow immediately under the antero-lateral shield margin; scapular setae sc 5 (4–6), directed upward and centrally, their tubercles 19 (15–20) apart. Gnathosoma 19 (18–21), directed downward. Dorsal pedipalp genual setae d simple. Leg I 32 (31–33), tibia 6 (5–6), l' 4 (3–5), tarsus 6 (5–7), ω 11 (10–11), without knob, empodium simple, 4-rayed; leg II 29 (29–31), tibia 5 (4–5), l' absent, tarsus 6 (5–6), ω 10 (9–10), without knob, empodium simple, 4-rayed (Fig. 3 L1, L2, em). Setae bv present. Sternal line 12 (11–13), bifurcated anteriorly. Coxae with numerous thin short lines. Rounded plate with three longitudinal lines situated before coxae I (Fig. 3 CG). Setae 1b 8 (6–10), 10 (9–10) apart; 1a 21 (15–26), 10 (9–10) apart; 2a 37 (24–41), 23 (23–24) apart. Genitalia 12 (11–14), 20 (20–21) wide, genital coverflap with 8–10 longitudinal ridges situated anteriorly (Fig. 3 CG); setae 3a 15 (14–19). Opisthosoma with 36 (32–38) dorsal annuli and 53 (51–54) ventral annuli both with microtubercles (Fig. 3 LM), 5–6 coxo-genital annuli. Setae c1 32 (22–36) on 5 (4–5) annulus; setae c2 28 (26–31) on 10 (9–11)
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annulus; setae d 27 (26–31) on 21 (18–23) annulus; setae e 23 (21–26) on 34 (29–37) annulus; setae f 27 (23– 31) on 4th ventral annulus from rear. Setae h1 2 (2–3). Male: not found. Material examined. 10 protogyne females (slide # 1) from F. viridis (inside red hairy galls on leaves), RUSSIA: Leningrad Prov., Luga district, pine forest near Lake Beloye, 58°48.4' N, 30°29.7' E, 6 July 2008, leg. T.G. Chetverikova; 30 protogyne females, 4 deutogyne females and 22 nymphs (slides # 37-09 and # 3809) from red galls, the same host, locality, 25 July 2009, leg.Ph. E. Chetverikov; 12 deutogyne females (slide # 40-09) from the lower surface of dry leaves, the same host, locality and leg., 15 August 2009; 24 protogyne females and 18 nymphs (slides # 40-09 and # 41-09) from red galls, the same host, locality and leg., 15 August 2009; 4 deutogyne females (slide # 44-09) from the lower surface of dry leaves, the same host, locality and leg., 9 September 2009; 17 deutogyne females, 5 protogyne females and 3 nymphs (slide # 43-09) from red galls, the same host, locality and leg., 9 September 2009. Deuterogyny of S. setiger. In contrast to the whitish or pallid pink protogyne females of S. setiger the deutogyne females of this species are bright orange. Morphologically these females do not differ. According to our field collections and observations in the laboratory during July numerous egg-laying protogyne females, plus nymphs and single deutogyne females, are present inside galls. During August the number of deutogyne females increases. By the end of August some leaves of their host-plants become dry. On these leaves only deutogyne females occur inside dry galls and on the lower leave surface. In the beginning of September we found numerous deutogyne females with single protogynes and nymphs (but no eggs) inside red galls on the leaves which still remained green and deutogyne females on dry leaves and inside dry galls. TABLE 4. Eriophyoid mite species with a plate before coxa I. Subfamily
Tribe, species
Nalepellinae
Nalepellini: Nalepella tsugifoliae Keifer, 1953; Setoptus jonesi (Keifer, 1938); Phantacrus lobatus Keifer, 1965
Sierraphytoptinae
Mackiellini: Palmiphytoptus oculatus Navia & Flechtmann, 2002 Sierraphytoptini: Sierraphytoptus setiger (Nalepa, 1894)
Phyllocoptinae
Phyllocoptini: Epitrimerus pyri (Nalepa, 1894); Monotrymacus Mohanasundaram, 1982; Arectus bidwillius Manson, 1984; Caliphytoptus quercilobatae Keifer, 938; Neodicrothrix tiliacorae Mohanasundaram, 1984; Platyphytoptus salinianae Keifer, 1938; Petanovicia cerberae Boczek, 1996 Anthocoptini: Tegolophus colifraxini (Keifer, 1938) Tegonotini: Glabrisceles euterpis Navia & Flechtmann, 2002; Scolocenus spiferus Keifer, 1962
Eriophyinae
Eriophyini: Eriophyes pyri (Pagenstecher, 1857); Proartacris pinivagrans Mohanasundaram, 1984
Cecidiphyinae
Cecidophyini: Achaetocoptes ajoensis (Keifer, 1961); Kolacarus bambusae Boczek, 1998; Neserella decora Meyer & Ueckermann, 1989
Nothopodinae
Nothopodini: Nothopoda rapanae Keifer, 1951; Anthopoda jonstoni Keifer, 1959 Colopodacini: Apontella bravasiae Boczek & Nuzzaci, 1988; Colopodacus africanus Keifer, 1960
Diptilomiopinae
Neodiptilomiopus vishakantai Mohanasundaram, 1982; Diptilostatus nidipalpus Flechtmann, 2003
Remarks. Both described species of Sierraphytoptus from Fragaria spp. possess a suboral plate before coxa I (Fig. 2 CG, 3 CG). The third species of this genus, S. alnivagrans from Alnus spp., has a weakly granulated suboral plate which was not drawn in Keifer's (1939) description (Petanovi ć , personal communication). Other species of the tribe Sierraphytoptini apparently do not have a suboral plate. The nature
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of this structure is unclear, but is probably a protuberance of coxae I, and a generally accepted term for this plate does not exist (Lindquist 1996). In the 20th century it was figured many times in the descriptions of new species and named differently, e.g.: “suboral plate” (Kiefer 1953) and “anterior coxal approximation” (Kiefer 1965; Manson 1984). In the monograph “Revised keys to world genera of Eriophyoidea” (Amrine et al. 2003) we found 26 figures of type species having a similar plate (Table 4). Besides, many eriophyid mite species from asteraceous plants and grasses (Aceria spp., Aculodes spp. and Abacarus spp.) have this structure (Petanovi ć et al. 2000; Skoracka 2001, 2002; Petanovi ć , personal communication). We think that the presence/absence of this plate, its form and design is of taxonomic significance.
FIGURE 3. Sierraphytoptus setiger (Nalepa, 1894), female. LM—lateral view, AD—prodorsal shield, em— empodium and tarsal solenidion, L1—leg I, L2—leg II, CG—coxogenital region. Scale: LM= 90 µm; AD, CG = 40 µm; em = 25; L1, L2 = 40; CG = 65 µm.
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Acknowledgments We are grateful to Drs. Radmila U. Petanović (Faculty of Agriculture Belgrade-Zemune, Belgrade, Serbia) and Sergey V. Mironov (Zoological Institute, Russian Academy of Sciences) for their critical notes and advice. We sincerely thank Dr. Owen Seeman (Queensland Museum, Australia) and anonymous reviewers for their comments on the manuscript, Dr. K.-W. Huang (Department of Zoology, National Museum of Natural Science, Taichung, Taiwan) for the possibility to work with type materials from Taiwan and Dr. P.G. Efimov, Dr. G.Y. Konechnaya (Komarov Botanical Institute, Saint Petersburg, Russia) and Mrss. T.G. Chetverikova, A.N. Baranova, O.V. Pereloma and Mr. N. Bannikov for their help in determining plant species and collecting mites.
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Nalepa, A. (1894) Beiträge zur Kenntniss der Phyllocoptiden. Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum Verhandlungen der kaiserlichen Leopoldinische-Carolinische Deutschen Akademie der Naturforscher (Halle). 61(4), 289–324. Navia, D., Gondim, M.G.C. & De Moraes, G.J. (2007) Eriophyoid mites (Acari: Eriophyoidea) associated with palm trees. Zootaxa, 1389, 1–30 pp. Newkirk, R.A. & Keifer, H.H. (1975) Appendix 3. Synoptic keys to groups and genera. Eriophyoidea. In: Jeppson L.R., Keifer H.H. & Baker E.W. (Eds.), Mites injurious to economic plants. University of California Press Berkeley, USA: 562–587. Oldfield, G.N. (1996) Diversity and host plant specificity. In: E.E. Lindquist, M.W. Sabelis and J. Bruin (eds.). Eriophyoid Mites - Their Biology, Natural Enemies and Control. Elsevier Science Publ., Amsterdam, The Netherlands, pp. 199–216. Petanović, R., Boczek, J. & Shi, A. (2000) Four new Aceria species (Acari: Eriophyoidea) from Serbia. Acta entomologica serbica, 5(1/2), 119–129. Petanović, R. (1988) Eriofidne grinje u Jugoslaviji. Naučna knjiga, Beograd,. str. 159. Petanović, R., Chetverikov, Ph. E. & Smiljani, D. (2007) Eriophyoid mites (Acari: Eriophyoidea) from sedges (Carex spp., Cyperaceae) in Serbia. Acta entomologica serbica, 12(2), 107–113. Petanović, R., Boczek, J., Jovanović, S., & Stojnić, B. (1996) Eriophyoidea (Acari: Prostigmata). Fauna Durmitora, 5, Crnogorska akademija nauka i umjetnosti, Posebna izdanja, knjiga 32, Odjeljenje prirodnih nauka, knjiga 18, Podgorica, 5–42. Roivainen, H. (1951) Contributions to the knowledge of the eriophyids of Finland. Acta Entomologica Fennica, 8, 1–72. Roivainen, H. (1953) Some gall mites (Eriophyidae) from Spain. Publicado en los Archivos del Instituto de Aclimatacion, 3, 9–43. Sennikov, A.N. (2006) Family Rosaceae Juss. In: Budantzev, A.L. & Jakovlev, G.P (Eds), Illustrated keys to plants of Leningrad province. KMK-press, Moscow, pp. 278–319 [in Russian]. Skoracka, A. (2001) A new species of eriophyoid mite from Festuca altissima ALL. (Poaceae) in Poland (Acari: Eriophyoidea). Genus, 12 (3), 385–394. Skoracka, A. (2002) Two new species of eriophyoid mites (Acari: Eriophyoidea) from grasses in Poland. Zootaxa, 54, 1– 15. Sukhareva, S. I. (1992) Four-legged mites from grasses. Saint-Petersburg State University Press, 232 pp. [in Russian]. Sukhareva, S. I. (1994) Family Phytoptidae Murray, 1877 (Acari: Tetrapodili), its consisting, structure and suggested ways of evolution. Acarina, 2(1–2), 47–72. Tchetverikov, F. E. (2004) Some features of morphology and biology of eriophyid mite Novophytoptus rostratae Roivainen, 1947. Vestnik SPbGU, 3, 8–17 [in Russian]. Tzvelev, N.N. (2000) Manual of the vascular plants of North-West Russia. St.-Petersburg State Chemical-Pharmaceutical Academy Press, Saint-Petersburg, Russia, 781 pp.
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