See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/259655788

A revision of the genus Sierraphytoptus Keifer

1939 (Eriophyoidea, Phytoptidae)

Article in Zootaxa · November 2009

CITATIONS READS

3 94

1 author:

Philipp E Chetverikov
Saint Petersburg State University
72 PUBLICATIONS 210 CITATIONS

SEE PROFILE

All content following this page was uploaded by Philipp E Chetverikov on 06 October 2014.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document
and are linked to publications on ResearchGate, letting you access and read them immediately.
 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

Zootaxa 2309: 30–42 (2009) ISSN 1175-5326 (print edition)

www.mapress.com / zootaxa/ Article ZOOTAXA
Copyright © 2009 · Magnolia Press 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: 1philipp-chetverikov@yandex.ru

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 host-
plants (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.

30 Accepted by O. Seeman: 18 Nov. 2009; published: 11 Dec. 2009

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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. Solenidion on tibia I present ......................................................................................................................................... 2

- Solenidion on tibia I absent (Sierraphytoptus Keifer, 1939) ........................................................................................ 4
2. 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
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.
4. Empodium 2-rayed, 16–17 dorsal annuli ..........................................................................S. alnivagrans Keifer, 1939
- Empodium 4-rayed, more than 20 dorsal annuli.......................................................................................................... 4
5. 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.

REVISION OF THE GENUS SIERRAPHYTOPTUS Zootaxa 2309 © 2009 Magnolia Press · 31

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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 Ren-
ai, 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.

32 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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 Fragaria Fragaria viridis Fragaria viridis (= F. Fragaria sp.
viridis vesca collina)
Relation to host cause red galls on vagrant on the vagrant on the lower cause red galls on the vagrant on the lower
the leaves lower leaf surface leaf surface leaves leaf surface, mainly
along the central vein
Number of dorsal 35* 21–24 25* 34 23–25
annuli
Number of annuli 6* 3* 2* 5* 3*
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

REVISION OF THE GENUS SIERRAPHYTOPTUS Zootaxa 2309 © 2009 Magnolia Press · 33

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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.

34 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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 4–5 5 ± 0.5 3–3 3 ± 0.0
and tubercles of c1
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

REVISION OF THE GENUS SIERRAPHYTOPTUS Zootaxa 2309 © 2009 Magnolia Press · 35

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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

36 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

(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 Zootaxa 2309 © 2009 Magnolia Press · 37

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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)

38 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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 # 38-
09) 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

REVISION OF THE GENUS SIERRAPHYTOPTUS Zootaxa 2309 © 2009 Magnolia Press · 39

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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.

40 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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.

Reference

Amrine, J. W. Jr. & Manson, D. C. M. (1996) Preparation, mounting and descriptive study of eriophyoid mites. pp. 383–
396. In: Lindquist, E. E., M. W. Sabelis and J. Bruin (Eds.). Eriophyoid Mites. Their Biology, Natural Enemies and
Control. Elsevier Science Publ., Amsterdam.
Amrine, J.W. Jr., Stasny, T. A., & Flechtmann, C.H.W. (2003) Revised keys to the world genera of the Eriophyoidea
(Acari: Prostigmata). Indira Publishing House, Michigan, USA, 244 pp.
Bagdasarian, A.T. (1981) Eriophyoid mites of the fruit trees and bushes from Armenia. Armenian SSR Academy of
Sciences Press, Erevan, 200 pp. [in Russian]
Bagdasarian, A.T. & Pogosova, A.R. (1976) On the fauna of eriophyoid mites on berries in Armenia (Acarina,
Eriophyoidea). Institute of zoology, Academy of Sciences of Armenian SSR, Zoological papers, XVII, 223–235 pp.
[in Russian].
Bagnjuk, I.G., Sukhareva, S.I. & Shevchenko, V.G. (1998) Major trends in the evolution of four-legged mites as a
specialized group (using families Pentasetacidae Shev., Nalepellidae Roiv. and Phytoptidae Murray (Acari:
Tetrapodili) as examples). Acarina, 6(1–2), 59–76.
Chetverikov, Ph. E. & Sukhareva, S.I. (2007) Supplementary descriptions and biological notes on eriophyid mites
(Acari: Eriophyidae) of the genus Novophytoptus Roivainen, 1947. Acarina, 15(1), 261–268.
Chetverikov, Ph. E., Petanović, R., Sukhareva, S.I. (2009) Systematic remarks on eriophyoid mites from the subfamily
Phytoptinae Murray, 1877 (Acari: Eriophyoidea: Phytoptidae). Zootaxa, 2070, 63–68.
Farkas, H. (1965) Spinnentiere Eriophyidae (Gallmilben). In: Brohmer, P., Ehrmann, P., Ulmer, G. (Eds), Die Tierwelt
Mitteleuropas, verlag von Quelle & Meyer, Leipzig, band 3, 155 s.
Huang, K.-W. (1992) Some new eriophyid mites from Taiwan. Bulletin of the National Museum of Natural Science, 3,
225–240. [check full title]
Huang, K.-W. (2006) Eriophyoid mites (Acari: Eriophyoidea) on Trochodendron aralioides (Trochodendraceae) from
Taiwan. Zootaxa, 1141, 63–68.
Kamelin R.V. (2001) Genus Fragaria L. In: Tzvelev N.N. (Eds), Flora of East Europe. Saint-Petersburg State Chemical-
Pharmaceutical Academy Press, X, pp. 452–456 [in Russian].
Keifer, H.H. (1939) Eriophyid studies III. Bulletin-Department of Agriculture State of California, 28, 144–163.
Keifer, H.H. (1944) Eriophyid studies XIV. Bulletin-Department of Agriculture State of California, 33, 18–38.
Keifer, H.H. (1953) Eriophyid studies XXI. Bulletin-Department of Agriculture State of California, 42, 65–79.
Keifer, H.H. (1965) Eriophyid studies B-16. Bureau of Entomology California Department of Agriculture, 20 pp.
Keifer, H.H. (1975) Eriophyoidea, Chapter 12. In: Jeppson L.R., Keifer, H.H. & Baker E.W. (Eds), Mites injurious to
economic plants. University of California Press Berkeley, USA, pp. 327–396.
Lindquist, E.E. (1996) External anatomy and notation of structures. 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. 301–327.
Lindquist, E.E. & Oldfield G.N. (1996). Evolution of eriophyoid mites in relation to their host plants. 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. 277–300.
Liro, J.I. (1941) Neue und selten Eriophyiden (Acarine). Annales Zoologici Societatis Zooogicae-Botanicae Fennica
Vanamo, 8(7), 1–54.
Manson, D.C.M. (1984) Eriophyoidea except Eriophyinae (Arachnida: Acari). Fauna of New Zealand, 4, 1–142.
Murray, A. (1877) Economic Entomology, Aptera. So. Kensington Mus. Handbooks, Chapman & Hall. London. 433 pp.

REVISION OF THE GENUS SIERRAPHYTOPTUS Zootaxa 2309 © 2009 Magnolia Press · 41

 TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

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.

42 · Zootaxa 2309 © 2009 Magnolia Press CHETVERIKOV & SUKHAREVA