Research Article |
Corresponding author: Fedor V. Konstantinov ( fkonstaninov@gmail.com ) Academic editor: Nikolay Simov
© 2024 Fedor V. Konstantinov, Reza Hosseini.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Konstantinov FV, Hosseini R (2024) Review of the genus Salicarus (Hemiptera, Heteroptera, Miridae). ZooKeys 1211: 57-100. https://doi.org/10.3897/zookeys.1211.129660
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The genus Salicarus Kerzhner, 1962 is revised, with differential diagnoses and redescriptions provided for nine species. Three distinct species groups were recognized within the genus: S. nitidus group (S. cavinotum, S. genistae, S. nitidus, S. perpusillus), S. roseri group (S. concinnus, S. roseri, S. urnammu), and S. fulvicornis group (S. halimodendri, S. fulvicornis). A key to species, illustrations of dorsal habitus, male and female genitalia, and selected diagnostic structures are included. Additionally, available host information and distributional records are summarized. Phoenicocoris qiliananus Zheng, 1996 is synonymized with Salicarus halimodendri V. G. Putshkov, 1977.
Distribution, hosts, new synonymy, Palearctic, phylogeny, taxonomy
Salicarus Kerzhner, 1962 belongs to the subfamily Phylinae of the hyperdiverse family Miridae, or plant bugs, the second largest family of insects with incomplete metamorphosis (
The present paper serves as a supplement to the mentioned revision by
Observations, measurements, and habitus images were made using a Zeiss Stemi 508 stereomicroscope equipped with a Canon EOS 2000D digital SLR camera. Partially focused images of each specimen or structure were stacked using Helicon Focus software. Images of the selected structures, including male and female genitalia, were taken with a Keyence VHX–500F digital microscope at the University of Hamburg. Illustrated structures were macerated in potassium hydroxide, cleared in distilled water, and then transferred to glycerin jelly for proper orientation. Scanning electron micrographs of selected structures were taken using a Quanta 250 scanning electron microscope.
Unless otherwise stated, all measurements are in millimeters. Measurements shown in Table
Measurements (mm). Abbreviations. Cun–Clyp – distance between apex of clypeus and apex of corium in dorsal view, Head Length – distance between apex oSf clypeus and the highest point of vertex, AntSeg1, AntSeg2 – length of antennal segments I and II, InterOcDi – width of vertex between inner margins of eyes in dorsal view.
Species | Length | Width | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
Body | Cun–Clyp | Pronotum | Tibia3 | AntSeg2 | Head | InterOcDi | Pronotum | |||
Salicarus cavinotum | ||||||||||
♂♂ (n = 5) | Mean | 2.16 | 1.86 | 0.37 | 0.98 | 0.49 | 0.65 | 0.36 | 0.83 | |
SD | 0.25 | 0.27 | 0.03 | 0.05 | 0.03 | 0.03 | 0.02 | 0.05 | ||
Range | 0.60 | 0.70 | 0.06 | 0.14 | 0.07 | 0.07 | 0.05 | 0.10 | ||
Min | 2.00 | 1.60 | 0.34 | 0.90 | 0.46 | 0.61 | 0.34 | 0.78 | ||
Max | 2.60 | 2.30 | 0.40 | 1.04 | 0.53 | 0.68 | 0.39 | 0.88 | ||
♀♀ (n = 5) | Mean | 2.15 | 1.91 | 0.39 | 1.02 | 0.49 | 0.68 | 0.40 | 0.89 | |
SD | 0.09 | 0.08 | 0.03 | 0.07 | 0.03 | 0.03 | 0.01 | 0.04 | ||
Range | 0.23 | 0.22 | 0.09 | 0.15 | 0.07 | 0.06 | 0.04 | 0.09 | ||
Min | 2.05 | 1.83 | 0.35 | 0.95 | 0.45 | 0.65 | 0.38 | 0.85 | ||
Max | 2.28 | 2.04 | 0.44 | 1.10 | 0.52 | 0.71 | 0.41 | 0.94 | ||
Salicarus concinnus | ||||||||||
♂♂ (n = 5) | Mean | 3.21 | 2.78 | 0.62 | 1.51 | 0.71 | 0.80 | 0.42 | 1.25 | |
SD | 0.33 | 0.21 | 0.03 | 0.04 | 0.04 | 0.02 | 0.01 | 0.07 | ||
Range | 0.75 | 0.50 | 0.08 | 0.13 | 0.10 | 0.05 | 0.01 | 0.18 | ||
Min | 2.95 | 2.63 | 0.58 | 1.45 | 0.65 | 0.78 | 0.41 | 1.18 | ||
Max | 3.70 | 3.13 | 0.65 | 1.58 | 0.75 | 0.83 | 0.43 | 1.35 | ||
♀♀ (n = 5) | Mean | 3.24 | 2.79 | 0.59 | 1.56 | 0.71 | 0.81 | 0.44 | 1.25 | |
SD | 0.18 | 0.11 | 0.06 | 0.11 | 0.09 | 0.03 | 0.01 | 0.09 | ||
Range | 0.45 | 0.25 | 0.15 | 0.20 | 0.20 | 0.06 | 0.03 | 0.23 | ||
Min | 3.00 | 2.70 | 0.53 | 1.45 | 0.65 | 0.79 | 0.43 | 1.15 | ||
Max | 3.45 | 2.95 | 0.68 | 1.65 | 0.85 | 0.85 | 0.45 | 1.38 | ||
Salicarus fulvicornis | ||||||||||
♂♂ (n = 5) | Mean | 3.82 | 3.08 | 0.57 | 1.54 | 0.85 | 0.80 | 0.40 | 1.20 | |
SD | 0.10 | 0.07 | 0.05 | 0.06 | 0.05 | 0.02 | 0.00 | 0.12 | ||
Range | 0.25 | 0.15 | 0.10 | 0.18 | 0.10 | 0.05 | 0.00 | 0.30 | ||
Min | 3.70 | 3.00 | 0.53 | 1.45 | 0.80 | 0.78 | 0.40 | 1.00 | ||
Max | 3.95 | 3.15 | 0.63 | 1.63 | 0.90 | 0.83 | 0.40 | 1.30 | ||
♀♀ (n = 5) | Mean | 3.29 | 2.82 | 0.53 | 1.41 | 0.73 | 0.84 | 0.45 | 1.18 | |
SD | 0.14 | 0.10 | 0.02 | 0.01 | 0.05 | 0.02 | 0.02 | 0.05 | ||
Range | 0.35 | 0.25 | 0.05 | 0.03 | 0.10 | 0.04 | 0.05 | 0.13 | ||
Min | 3.15 | 2.70 | 0.50 | 1.40 | 0.68 | 0.83 | 0.43 | 1.13 | ||
Max | 3.50 | 2.95 | 0.55 | 1.43 | 0.78 | 0.86 | 0.48 | 1.25 | ||
Salicarus genistae | ||||||||||
♂♂ (n = 5) | Mean | 2.36 | 2.10 | 0.46 | 1.18 | 0.58 | 0.75 | 0.39 | 1.04 | |
SD | 0.11 | 0.08 | 0.02 | 0.03 | 0.02 | 0.02 | 0.01 | 0.01 | ||
Range | 0.28 | 0.22 | 0.04 | 0.06 | 0.04 | 0.05 | 0.02 | 0.04 | ||
Min | 2.20 | 2.00 | 0.44 | 1.14 | 0.56 | 0.72 | 0.38 | 1.02 | ||
Max | 2.48 | 2.22 | 0.48 | 1.20 | 0.60 | 0.77 | 0.40 | 1.06 | ||
♀♀ (n = 5) | Mean | 2.72 | 2.43 | 0.49 | 1.27 | 0.59 | 0.79 | 0.43 | 1.06 | |
SD | 0.09 | 0.10 | 0.01 | 0.03 | 0.03 | 0.02 | 0.01 | 0.03 | ||
Range | 0.20 | 0.24 | 0.03 | 0.08 | 0.08 | 0.05 | 0.02 | 0.06 | ||
Min | 2.60 | 2.30 | 0.47 | 1.22 | 0.54 | 0.76 | 0.42 | 1.04 | ||
Max | 2.80 | 2.54 | 0.50 | 1.30 | 0.62 | 0.81 | 0.44 | 1.10 | ||
Salicarus halimodendri | ||||||||||
♂♂ (n = 5) | Mean | 3.71 | 3.11 | 0.57 | 1.42 | 0.78 | 0.89 | 0.45 | 1.32 | |
SD | 0.17 | 0.11 | 0.05 | 0.02 | 0.02 | 0.03 | 0.02 | 0.05 | ||
Range | 0.33 | 0.25 | 0.10 | 0.05 | 0.05 | 0.05 | 0.05 | 0.13 | ||
Min | 3.55 | 2.95 | 0.50 | 1.40 | 0.75 | 0.85 | 0.43 | 1.25 | ||
Max | 3.88 | 3.20 | 0.60 | 1.45 | 0.80 | 0.90 | 0.48 | 1.38 | ||
♀♀ (n = 5) | Mean | 3.34 | 2.96 | 0.58 | 1.41 | 0.69 | 0.92 | 0.49 | 1.36 | |
SD | 0.15 | 0.12 | 0.02 | 0.03 | 0.03 | 0.03 | 0.01 | 0.03 | ||
Range | 0.35 | 0.33 | 0.05 | 0.08 | 0.08 | 0.08 | 0.03 | 0.08 | ||
Min | 3.13 | 2.83 | 0.55 | 1.38 | 0.65 | 0.88 | 0.48 | 1.33 | ||
Max | 3.48 | 3.15 | 0.60 | 1.45 | 0.73 | 0.95 | 0.50 | 1.40 | ||
Salicarus nitidus | ||||||||||
♂♂ (n = 2) | Min | 2.30 | 2.00 | 0.40 | 1.06 | 0.52 | 0.74 | 0.40 | 0.94 | |
Max | 2.60 | 2.28 | 0.46 | 1.22 | 0.53 | 0.75 | 0.42 | 0.96 | ||
♀♀ (n = 2) | Min | 2.18 | 1.98 | 0.42 | 1.07 | 0.56 | 0.70 | 0.40 | 0.96 | |
Max | 2.40 | 2.16 | 0.44 | 1.14 | 0.57 | 0.74 | 0.40 | 0.98 | ||
Salicarus perpusillus | ||||||||||
♂♂ (n = 3) | Mean | 2.24 | 1.88 | 0.40 | 1.09 | 0.50 | 0.69 | 0.38 | 0.87 | |
Min | 2.14 | 1.84 | 0.38 | 1.02 | 0.49 | 0.66 | 0.37 | 0.84 | ||
Max | 2.37 | 1.95 | 0.41 | 1.14 | 0.52 | 0.71 | 0.39 | 0.88 | ||
♀♀ (n = 5) | Mean | 2.27 | 1.97 | 0.42 | 1.04 | 0.53 | 0.70 | 0.39 | 0.90 | |
SD | 0.11 | 0.14 | 0.02 | 0.02 | 0.01 | 0.03 | 0.01 | 0.03 | ||
Range | 0.24 | 0.33 | 0.04 | 0.04 | 0.03 | 0.07 | 0.02 | 0.07 | ||
Min | 2.16 | 1.83 | 0.40 | 1.02 | 0.51 | 0.66 | 0.38 | 0.86 | ||
Max | 2.40 | 2.16 | 0.44 | 1.06 | 0.54 | 0.73 | 0.40 | 0.93 | ||
Salicarus roseri | ||||||||||
♂♂ (n = 5) | Mean | 3.80 | 3.31 | 0.68 | 1.65 | 0.74 | 0.86 | 0.41 | 1.40 | |
SD | 0.16 | 0.20 | 0.02 | 0.10 | 0.04 | 0.03 | 0.02 | 0.06 | ||
Range | 0.38 | 0.50 | 0.05 | 0.25 | 0.10 | 0.08 | 0.05 | 0.15 | ||
Min | 3.58 | 3.13 | 0.65 | 1.58 | 0.68 | 0.83 | 0.39 | 1.33 | ||
Max | 3.95 | 3.63 | 0.70 | 1.83 | 0.78 | 0.90 | 0.44 | 1.48 | ||
♀♀ (n = 5) | Mean | 3.76 | 3.25 | 0.67 | 1.61 | 0.71 | 0.89 | 0.41 | 1.42 | |
SD | 0.22 | 0.21 | 0.06 | 0.07 | 0.07 | 0.02 | 0.05 | 0.07 | ||
Range | 0.55 | 0.50 | 0.15 | 0.20 | 0.18 | 0.05 | 0.13 | 0.18 | ||
Min | 3.38 | 2.88 | 0.58 | 1.50 | 0.60 | 0.85 | 0.33 | 1.30 | ||
Max | 3.93 | 3.38 | 0.73 | 1.70 | 0.78 | 0.90 | 0.45 | 1.48 | ||
Salicarus urnammu | ||||||||||
♂♂ (n = 3) | Mean | 3.61 | 3.04 | 0.67 | 1.53 | 0.73 | 0.85 | 0.43 | 1.28 | |
Min | 3.45 | 2.88 | 0.63 | 1.50 | 0.70 | 0.83 | 0.43 | 1.25 | ||
Max | 3.75 | 3.20 | 0.73 | 1.55 | 0.75 | 0.89 | 0.43 | 1.35 | ||
♀♀ (n = 3) | Mean | 3.34 | 2.94 | 0.63 | 1.48 | 0.66 | 0.85 | 0.43 | 1.29 | |
Min | 3.20 | 2.83 | 0.60 | 1.45 | 0.60 | 0.83 | 0.40 | 1.25 | ||
Max | 3.45 | 3.08 | 0.65 | 1.53 | 0.70 | 0.88 | 0.45 | 1.38 |
The material examined for this study is retained in the following collections:
UGNHM Natural History Museum of the University of Guilan, Rasht;
Bar code labels, uniquely identifying each specimen, were attached to all examined specimens listed in the “Material examined” sections. Further information, such as additional photographs of habitus and genitalia structures, georeferenced coordinates, specimens dissected, notes, and collecting methods, can be obtained from the Heteroptera Species Pages (http://research.amnh.org/pbi/heteropteraspeciespage/), which assembles available data from a specimen database (
Salicarus Kerzhner, 1962: 381. Type species by original designation: Capsus roseri Herrich-Schaeffer, 1838.
Salicarus:
Salicarius
[sic!]:
Body broadly oval, with short appendages (Figs
Male. Coloration. Dorsum and venter uniformly chestnut to dark brown with whitish antennal segments III and IV in S. nitidus species group (Fig.
Surface and vestiture. Dorsum shiny to moderately shiny, head and pronotum usually smooth, scutellum and hemelytron weakly rugose. Vestiture composed of dense simple setae intermixed with one of two types of scale-like setae: wide, apically serrate, densely covering dorsum, thoracic pleura, abdomen, and sometimes appendages (S. nitidus species group, Fig.
Structure. Body elongate-oval to oval, total length 2.0–4.0. Head: Flattened and strongly sloping, barely protruding beyond anterior margin of eyes (Figs
Genitalia. Genital capsule cone-shaped, without distinctive ornamentation, as long as or slightly longer than wide at base. Sclerotized apical part of phallotheca narrow, beak-shaped, somewhat constricted at base (Fig.
Female. Coloration, surface, and vestiture. As in male. Structure. Similar to male, usually smaller on average (Table
Genitalia. Dorsal labiate plate with large and wide, broadly oval or apically tapering sclerotized rings (Fig.
Three distinct groups of species can be recognized within Salicarus, and the species treatments below are arranged alphabetically within each group:
Salicarus nitidus species group. This group includes S. cavinotum, S. genistae, S. nitidus, and S. perpusillus. Species in this group are characterized by their uniformly dark color, dorsum densely covered with wide, apically serrate scale-like setae (scales type 2 sensu
Salicarus roseri species group. This group includes S. concinnus, S. roseri, and S. urnammu. Species in this group are characterized by their highly variable color pattern, relatively large, oval body with a total length of 3.0–4.0. The dorsum vestiture consists of short, adpressed simple setae, while narrow, apically acuminate scale-like setae (scales type 1 sensu
Salicarus fulvicornis species group. This group includes S. halimodendri and S. fulvicornis. Species in this group are variable in coloration, with elongate males (3.6–4.0) and more ovoid females (3.5–3.9). The entire dorsum, except the head, is clothed with a mixture of silvery narrow, apically acuminate scale-like setae, and dense, comparatively long simple setae that are approximately 1.5× as long as the scales. The apical blades of the vesica are very long, thin, gradually curved, and abruptly furcate. Species of this group feed exclusively on Caragana spp. (Fabaceae: Hedysareae) and are mainly distributed in Central Asia and Mongolia.
1 | Smaller than 2.8. Antennal segment II distinctly swollen, fusiform. Dorsum and venter, including head, pronotum and abdomen, with dense, wide, apically serrate scales (Fig. |
2 |
– | Larger than 3.0. Antennal segment II thin. Scale-like setae on dorsum, if present, narrow and apically acuminate (Fig. |
5 |
2 | Femora with scale-like setae | 3 |
– | Femora without scale-like setae | 4 |
3 | Apical blades of vesica long, gradually curving, closely located but separate, not adjoining each other (Fig. |
S. nitidus |
– | Apical blades of vesica gradually curved and tightly adjoining each other along their entire length (Fig. |
S. perpusillus |
4 | Antennal segment II distinctly fusiform in both sexes, 1.6–1.8× as wide at midpoint as segment I at apex (Fig. |
S. cavinotum |
– | Antennal segment II in male swollen along entire length, slightly fusiform, with midpoint width subequal to apical width of segment I (Fig. |
S. genistae |
5 | Hemelytron without scale-like setae, clothed with short, strongly adpressed, simple silvery setae only (Fig. |
S. roseri |
– | Hemelytron clothed with a mixture of simple setae and narrow, apically acuminate scale-like setae (Fig. |
6 |
6 | Dorsum clothed with short, subequal in length to scale-like setae on hemelytron, adpressed simple setae; narrow scale-like setae scarce and present on hemelytron only (Fig. |
7 |
– | Pronotum, scutellum, and hemelytron clothed with a mixture of silvery narrow scale-like setae and dense, long, simple setae ~ 1.5× as long as scales (Fig. |
8 |
7 | Vesica with almost parallel apical blades (Fig. |
S. concinnus |
– | Vesica with gradually diverging apical blades (Fig. |
S. urnammu |
8 | Coloration of dorsum variable, ranging from almost entirely dark brown to pale yellow, but vertex always dirty to whitish yellow along posterior margin (Fig. |
S. halimodendri |
– | Dorsum uniformly dark brown to brown (Fig. |
S. fulvicornis |
Heterocapillus cavinotum Wagner, 1973: 121.
Heterocapillus cavinotum:
Salicarus cavinotum:
Holotype
: ♂ Greece • Dodecanese Islands: Petaloudes, Rhodos, 36.444°N, 28.222°E, 01 Jun 1972, Eckerlein (AMNH_PBI 00184018) (
Other specimens examined: Greece • Dodecanese Islands: Petaloudes, Rhodos, 36.444°N, 28.222°E, 01 Jun 1972, Eckerlein, 1♂ (AMNH_PBI 00240965) (
Recognized by the small size, body length 2.0–2.6; antennal segment II fusiform in both sexes, wider in female; dorsum uniformly dark brown, with dense, wide and apically serrate silvery scale-like setae (Figs
Head, pronotum, and vestiture A–C Salicarus concinnus in dorsal view: A ♂ paratype, head and pronotum B ♀ paratype, head and pronotum C ♀ paratype, vestiture on hemelytron D–F S. fulvicornis in dorsal view: D ♂ head and pronotum E ♀ head and pronotum F ♂ vestiture on hemelytron G–I S. halimodendri in dorsal view: G ♂ paratype H ♀ paratype I ♂ paratype, vestiture on hemelytron J–L S. roseri: J ♂ head and pronotum K ♀ head and pronotum L head and thoracic pleura in lateral view M–O S. urnammu in dorsal view: M ♂ head and pronotum N ♀ head and pronotum O ♀ vestiture on hemelytron.
Salicarus cavinotum is most similar to S. perpusillus in general appearance, size, and vesica structure, which appear indistinguishable between these species (Fig.
Male. Coloration. Dorsum and venter uniformly brown to dark brown (Fig.
Surface and vestiture. Smooth, moderately shiny; dorsum, thoracic pleura, and abdomen with dense, silvery, broad and apically serrate scale-like setae and adpressed to semierect, long, almost twice as long as scales, simple setae, dark on cuneus and apex of corium, yellowish elsewhere; legs and antennae without scale-like setae; sides of pronotum and hemelytron at base with robust, long, erect to semierect, black bristle-like setae.
Structure. Body 2.4–3.0× as long as posterior width of pronotum; total length 2.0–2.6; vertex 2.3–2.7× as wide as eye; antennal segment I short, swollen, widest at apex, about twice as long as width at apex; segment II fusiform, 1.6× as wide at midpoint as segment I at apex, 4.9–5.3× as long as wide, 0.6× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; segments III and IV filiform; pronotum 2.1–2.4× as wide as long, 1.2–1.3× as wide as head.
Genitalia. Right paramere with oval body about twice as long as wide, basally broadly rounded and expanded well proximally beyond basal process, apical process long and straight, apically rounded. Left paramere similar to those of S. genistae (Fig.
Female. Coloration, surface, and vestiture. As in male (Fig.
Structure. Body 2.2–2.5× as long as posterior width of pronotum; total length 2.1–2.3; vertex 2.5–2.9× as wide as eye; antennal segment I short, swollen, widest at apex, about twice as long as width at apex; segment II fusiform, wider than in male, 1.7–1.8× as wide at midpoint as segment I at apex, 4.1–4.3× as long as wide, 0.5–0.6× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; pronotum 2.1–2.4× as wide as long, 1.3× as wide as head.
Genitalia. Dorsal labiate plate with large and wide, broadly oval, but apically tapering sclerotized rings.
Currently this species is documented exclusively in Greece, spanning Thessaly, the Peloponnese and Attic peninsulas, as well as Crete and Rhodes Island (
Genista sp. (
Atractotomus genistae Lindberg, 1948: 53.
Heterocapillus genistae:
Salicarus genistae:
Paralectotypes: Cyprus • Ayios Hilarion, 35.3125°N, 33.28333°E, 07 Jun 1939, Hakan Lindberg, 1♂ (AMNH_PBI 00336958) (
Other specimens examined: Cyprus • Kakomallis Mt., 34.83333°N, 33.03333°E, 914 m, 13 Jun 1965, G. Mavromoustakis, 2♂ (AMNH_PBI 00336959, AMNH_PBI 00336960), 2♀ (AMNH_PBI 00336966, AMNH_PBI 00336967) (
Recognized by the relatively small, stumpy, dark brown body, total length male 2.2–2.5, female 2.6–2.8 (Fig.
Head, pronotum, and vestiture A, B paratype of Salicarus cavinotum, head and pronotum in dorsal view: A ♂ B ♀ C, G S. genistae, head and pronotum in dorsal view: C ♂ G ♀ D–F paratypes of S. perpusillus: D ♂ head and pronotum in dorsal view E ♀ head and pronotum in dorsal view F ♀ vestiture on dorsum and legs H–I S. nitidus, head and pronotum in dorsal view: H ♂ I ♀.
Salicarus genistae is most similar in habitus, coloration, size, and male genitalia structure to S. cavinotum, S. nitidus, and S. perpusillus. It differs habitually from these species by its sexually dimorphic antennal segment II: in males it is slightly fusiform with the width at the midpoint being subequal to the apical width of segment I; in females it is distinctly fusiform, being 1.5–1.6× as wide at the midpoint as segment I at the apex. Consequently, antennal segment II is 6.4–6.7× as long as wide in males of S. genistae, being 4.9–5.2× as long as wide in female. In other three closely related species this ratio ranges 4.1–5.3× in males and 3.9–4.3× in females. However, these ratios should be used with caution due to observed polymorphism and the extremely dense vestiture of antennal segment II, which can affect the measurements. Salicarus genistae further differs from both S. nitidus and S. perpusillus in the absence of scales on femora. In vesica structure, with the apical blades tightly adjoining each other, it is most similar to S. cavinotum (Fig.
Scanning electron images of selected structures A–C Salicarus fulvicornis: A pretarsus in lateral view B hind tarsus in lateral view C vestiture on hemelytron D–F S. roseri: D pretarsus in lateral view E head and thoracic pleura in lateral view F metathoracic spiracle and scent gland evaporative area.
Male. Coloration. Dorsum and venter uniformly chestnut to dark brown (Fig.
Surface and vestiture. Smooth, moderately shiny; dorsum, thoracic pleura, and abdomen with dense, silvery, broad and apically serrate scale-like setae and adpressed to semierect, long, almost twice as long as scales, simple setae, dark on cuneus and extreme apex of corium and goldish yellow elsewhere (Fig.
Structure. Body stumpy, oval, 2.1–2.4× as long as posterior width of pronotum; total length 2.2–2.5; vertex flat, 2.0–2.2× as wide as eye; segment I short, swollen, widest at apex, about twice as long as width at apex; segment II swollen along entire length, somewhat fusiform, with midpoint width subequal to apical width of segment I, 6.4–6.7× as long as wide, 0.5–0.6× as long as basal width of pronotum, 0.7–0.8× as long as width of head; segments III and IV filiform; labium reaching metacoxa; pronotum 2.2–2.4× as wide as long, 1.4–1.5× as wide as head; mesonotum only slightly exposed.
Genitalia. Right paramere oval, approximately twice as long as wide, basally broadly rounded, and well expanded proximally beyond basal process, with long, straight, gradually tapering apical process (Fig.
Female. Coloration, surface, and vestiture. As in male (Fig.
Structure. Body 2.5–2.7× as long as posterior width of pronotum; total length 2.6–2.8; vertex 2.2–2.6× as wide as eye; antennal segment I short, swollen, widest at apex, about twice as long as width at apex; segment II somewhat shorter than in male, strongly swollen, fusiform, 1.5–1.6× as wide at midpoint as segment I at apex, 4.9–5.2× as long as wide, 0.5–0.6× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; pronotum 2.1–2.2× as wide as long, 1.3–1.4× as wide as head.
Genitalia. Dorsal labiate plate with large and wide, broadly oval, but apically tapering sclerotized rings (Fig.
Originally described and still known from Cyprus. A record from Manavgat (Antalya province of Turkey) based on a single specimen of unknown sex (
Genista fasselata Decne. (
Refer to the corresponding section in the redescription of S. nitidus.
Atractotomus nitidus Horváth, 1905: 275.
Heterocapillus nitidus:
Salicarus nitidus:
Holotype: Spain • Castile-La Mancha: ♀ Pozuelo de Calatrava 38.91°N, 3.84°W, Collection date unknown, José María de la Fuente (
Other specimens examined: France • Corse (Corsica): Costa, 42.0333°N, 8.95°E, 19 Jul 1963, Unknown collector, 1♀ (AMNH_PBI 00336969) (
Recognized by the small, stumpy, uniformly dark brown body, total length male 2.3–2.6, female 2.2–2.4; dorsum with dense, wide, and apically serrate scale-like setae, femora also clothed with scales (Fig.
Most similar to S. perpusillus in size, body proportions, and the distinctly fusiform antennal segment II in both sexes, as well as the presence of scale-like setae on femora (although scales are absent on the bases of tibiae and antennae). However, it differs from this species, as well as from S. cavinotum and S. genistae in the vesica structure with apical blades that are not tightly adjoining. Additionally, S. nitidus is the only one of the closely related species mentioned above that feeds on Calicotome rather than Genista spp. Refer to the diagnosis of S. genistae for additional discussion.
Male. Coloration. Dorsum and venter uniformly brown to dark brown (Fig.
Surface and vestiture. Smooth, moderately shiny; dorsum, thoracic pleura, femora, and abdomen with dense, silvery, broad and apically serrate scale-like setae and adpressed to semierect, long, almost twice as long as scales, simple setae, dark on cuneus and apex of corium, yellowish elsewhere; series of long simple setae on fore coxa silver; sides of pronotum and hemelytron at base with robust, long, erect to semierect, black bristle-like setae.
Structure. Body 2.5–2.7× as long as posterior width of pronotum; total length 2.3–2.6; vertex 2.3–2.6× as wide as eye; antennal segment II distinctly fusiform, 1.4–1.5× as wide at midpoint as segment I at apex, 4.1–4.3× as long as wide, 0.5–0.6× as long as posterior width of pronotum, 0.7× as long as width of head; pronotum 2.1–2.3× as wide as long, 1.3× as wide as head.
Genitalia. Right paramere spoon-shaped, with long, straight, and blunt apical process (Fig.
Female. Coloration, surface, and vestiture. As in male (Fig.
Structure. Body 2.3–2.5× as long as posterior width of pronotum; total length 2.2–2.4; vertex 2.4–2.6× as wide as eye; antennal segment II somewhat wider than in male, 1.6–1.7× as wide at midpoint as segment I at apex, 4.0–4.1× as long as wide, 0.6× as long as posterior width of pronotum, 0.7× as long as width of head; pronotum 2.2–2.3× as wide as long, 1.3–1.4× as wide as head.
Genitalia. Dorsal labiate plate with large and wide, broadly oval at base, apically tapering sclerotized rings.
Ciudad Real province of Spain (
Calicotome villosa (Poir.) Link. (
Salicarus nitidus (Horváth, 1905) belongs to a group of four hardly distinguishable species with a complex taxonomic history, which also includes S. cavinotum (Wagner, 1973), S. genistae (Lindberg, 1948), and S. perpusillus (Wagner, 1960). These species inhabit the North Mediterranean, from central Spain in the West to Cyprus in the East.
In his monographic treatment of Mediterranean plant bugs
Heterocapillus genistae (Cyprus, on Genista sp.): relatively large, body length male 2.5 mm, female 3.0 mm, antennal segment II slightly spindle-shaped in male, distinctly spindle-shaped in female.
Heterocapillus cavinotum (Rhodos Island, on Genista spp.): body length male 2.35 mm, female 1.9 mm, pronotum with a pair of pits, antennal segment II distinctly spindle-shaped in both sexes, 3.8–4.0× as long as wide at middle.
Heterocapillus nitidus (central Spain, Corsica, on Calicotome villosa): body length male 2.5 mm, female 2.2–2.3 mm, antennal segment II spindle-shaped in both sexes, male 4×, female 3.9× as long as wide at middle.
Heterocapillus perpusillus (southern Spain, southern France, on Genista sp.): body length male 2.1 mm, female 2.3–2.4 mm, antennal segment II spindle-shaped in both sexes, male 4.2×, female 4.0× as long as wide at middle.
Since then, H. cavinotum was additionally reported from Peloponnese peninsula and Crete (
Atractotomus (Heterocapillus) perpusillus Wagner, 1960: 81.
Heterocapillus perpusillus:
Salicarus perpusillus:
Holotype: Spain • Andalucia: ♂ Northern Slopes of Veleta Peak [Veleta -Nordhang], Sierra Nevada, 37.07°N, 3.37°W, 2500 m, 03 Aug 1959, E. Wagner, (AMNH_PBI 00184020) (
Other specimens examined: Spain • Catalonia: Campllong, Bergueda, 41.88333°N, 2.81667°E, 15 Jul 1984, E. Ribes, 1♂ (ZISP_ENT 00011719), 1♀ (ZISP_ENT 00011719) (NMPC) • Seros, Segria, 41.462°N, 0.412°E, 27 Jun 1971, J. Ribes, Ulex sp. (Fabaceae), 1♂ (AMNH_PBI 00338308) (
Recognized by the small and stumpy body, total length 2.1–2.4; dorsum uniformly dark brown with dense, wide and apically serrate scale-like setae (Fig.
Salicarus perpusillus is habitually most similar to S. nitidus in body proportions, the distinctly fusiform antennal segment II in both sexes, and the presence of scale-like setae on the femora. However, the latter species differs from S. perpusillus in its distinctive vesica structure with separate, not tightly adjoining apical blades. Indistinguishable from S. cavinotum in vesica structure but differs from that species by the presence of dense scale-like setae on the femora.
Male. Coloration. Dorsum and venter uniformly brown to dark brown (Fig.
Surface and vestiture. Smooth, moderately shiny; dorsum, thoracic pleura, and abdomen with very dense, silvery, broad and apically serrate scale-like setae and adpressed to semierect, long, almost twice as long as scales, simple setae, dark on cuneus and apex of corium, yellowish elsewhere; antennal segments I and II, all femora and basal parts of tibiae clothed with scale-like setae; sides of pronotum and hemelytron at base with robust, long, erect to semierect, black bristle-like setae.
Structure. Body 2.1–2.3× as long as posterior width of pronotum; total length 2.1–2.4; vertex 2.2–2.5× as wide as eye; segment II distinctly fusiform, 1.3–1.5× as wide at midpoint as segment I at apex, 4.3–4.6× as long as wide, 0.5–0.6× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; pronotum 2.1–2.2× as wide as long, 1.2–1.3× as wide as head.
Genitalia. Right paramere spoon-shaped, with long, straight apical process. Right paramere with short and straight apical process and broadly rounded sensory lobe. Vesica with gradually curved and comparatively long apical blades tightly adjoining each other along their entire length, with length of larger blade distinctly exceeding distance between its base and secondary gonopore (Fig.
Female. Coloration, surface, and vestiture. As in male (Fig.
Structure. Body 2.4–2.6× as long as posterior width of pronotum; total length 2.2–2.4; vertex 2.3–2.7× as wide as eye; segment II distinctly somewhat wider than in male, 1.6–1.7× as wide at midpoint as segment I at apex, 3.9–4.2× as long as wide, 0.6× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; pronotum 2.1–2.2× as wide as long, 1.3× as wide as head.
Genitalia. Dorsal labiate plate with large and wide, broadly oval at base, apically tapering sclerotized rings.
Spain (
Genista versicolor Boiss. (
Salicarus (Salicarus) concinnus V. G. Putshkov, 1977: 365.
Salicarus concinnus:
Holotype: Tajikistan • ♂ Kondara Canyon, Valley of Varzob River, 38.83333°N, 68.83333°E, 1100 m, 08 Jul 1955, Lopatin, (AMNH_PBI 00233863) (
Other specimens examined: Tajikistan • 6 km W Kuibyshevsk, Valley of Vakhsh River, 37.96666°N, 68.75°E, 14 Jul 1943, A. N. Kiritshenko, 1♀ (AMNH_PBI 00233774) (
Recognized by the following combination of characters: Body oval, total length 3.0–3.7; antenna uniformly pale yellow, with thin segment II (Fig.
Salicarus concinnus is most similar in size, body proportions, vestiture, and vesica structure to S. roseri and S. urnammu. The vestiture of the dorsum in all three species is mainly composed of short, adpressed simple setae, with the addition of scarce, narrow, apically acuminate scale-like setae on the hemelytron in the case of S. concinnus and S. urnammu. In S. roseri, scale-like setae are present on the thoracic pleura only. The color pattern of the dorsum in these species is highly variable, although in S. concinnus, it tends to be more uniform, frequently being either dark brown with a yellowish vertex or whitish yellow with a darkened posterior margin of the pronotum. In contrast, in dark specimens of S. urnammu and pale specimens of S. roseri, the anterior part of the pronotum is most frequently darkened, and the hemelytron usually has a more or less darkened endocorium (Figs
Male. Coloration. Variable, ranging from entirely or largely brown to pale yellow with darkened basal margin of pronotum (Fig.
Surface and vestiture. Dorsum smooth, shiny. Pronotum, scutellum, and hemelytron with short, subequal in length to scale-like setae on hemelytron, adpressed simple setae, usually dark brown, sometimes yellowish; hemelytron additionally with silver scale-like setae; thoracic pleurites clothed with dense scale-like setae only (Fig.
Structure. Body oval, 2.4–2.8× as long as posterior width of pronotum, total length 3.0–3.7; head vertical, rather vide, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly attenuate and covering anterior margin of pronotum, 2.1–2.3× as wide as eye; frons weakly convex; clypeus flat, not visible in dorsal view; antennal segment II thin and short, 0.5–0.6× as long as posterior width of pronotum, 0.9× as long as width of head; pronotum with broadly rounded anterior and posterior corners, 1.9–2.1× as wide as long, 1.5–1.6× as wide as head.
Genitalia. Right paramere oval, ~ 1.9× as long as wide, with basal part broadly rounded and expanded proximally beyond basal process; apical process comparatively short, subrectangular (Fig.
Female. Coloration, surface and vestiture. As in male (Fig.
Genitalia. Dorsal labiate plate with large and broadly oval sclerotized rings (Fig.
Central Asia. Known from Southern Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan (
Feeds on fructiferous Salix spp. (
Capsus roseri Herrich-Schaeffer, 1838: 78.
Capsus geniculatus
Stål, 1858: 355 (synonymized by
Capsus saliceticola
Stål, 1858: 355 (synonymized by
Sthenarus vittatus
Fieber, 1858: 339 (synonymized by
Sthenarus roseri:
Plagiognathus (Sthenarus) roseri:
Lygus roseri: Vollenhoven (1875): 93 (description).
Sthenarus (Phoenicocoris) roseri:
Salicarus roseri:
Sthenarus
(Salicarius (sic!)) roseri:
Belarus • Korolevo nr Vitebsk, 55.13333°N, 30.5°E, 08 Jul 1905, Birulya, 1♂ (AMNH_PBI 00233591) (
Recognized by the following characters: body oval, total length 3.4–4.0; antennal segment I brown, segment II thin, brown or at least with darkened base and apex (Fig.
Salicarus roseri easily differs from congeners by the absence of scale-like setae on dorsum. It further differs by having short, robust, straight, and slightly diverging apical blades of the vesica, being most similar to S. urnammu in this respect, although the blades in the latter species are shorter.
Male. Coloration. Highly variable, dorsum ranging from uniformly dark brown to pale yellow, with somewhat darkened head (Fig.
Surface and vestiture. Dorsum shiny, head and pronotum smooth, scutellum and hemelytron weakly rugose (Fig.
Structure. Body oval, 2.6–2.9× as long as width of pronotum at base, total length 3.6–4.0; head vertical, rather vide, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly attenuate and covering anterior margin of pronotum, 1.8–1.9× as wide as eye; antennal segment II at base distinctly thinner than segment I, slightly dilating apically, comparatively short, 0.5–0.6× as long as basal width of pronotum, 0.8–0.9× as long as width of head; pronotum with broadly rounded anterior and posterior corners, 2.0–2.2× as wide as long, 1.5–1.7× as wide as head.
Genitalia. Right paramere elongate-oval, ~ 2.5× as long as wide, with long, straight and apically blunt apical process (Fig.
Female. Coloration, surface and vestiture. As in male (Fig.
Genitalia. Sclerotized rings of dorsal labiate late large, broadly oval (Fig.
Widely distributed in the Palearctic, including almost the entire Europe, extending eastward to the Khabarovsk and Kamchatka territories in Russia, and southward to Spain, Italy, Greece, Turkey, Transcaucasia, Iran, Turkmenistan, Kazakhstan, Mongolia, and Inner Mongolia of China. To the north, it extends to the central Fennoscandia, Karelia, Arkhangelsk and Komi Provinces, the southern Yamalo-Nenets district, southern Krasnoyarsk Territory, southern and central Yakutia, and Magadan Territory (
Confined to Salix spp. (
Salicarius [sic!] urnammu Linnavuori, 1984: 51.
Salicarus urnammu:
Armenia • Aralykh, 40.11722°N, 44.27055°E, 07 Jun 1911, K. Satunin, 3♂ (AMNH_PBI 00233861, AMNH_PBI 00233862, AMNH_PBI 00233858) (
Recognized by the oval body, total length male 3.5–3.8; female 3.2–3.5; antenna typically yellow, in dark specimens segments I and II partly brown, segment II thin (Fig.
Salicarus urnammu is most similar to S. concinnus and S. roseri but can usually be distinguished from these species by the color pattern. It further differs from S. roseri by the presence of scale-like setae on the hemelytron, and from S. concinnus by the diverging apical blades of the vesica. Refer to the diagnoses of these species for additional discussion.
Male. Coloration. Variable, ranging from yellow, frequently with orange tinge, sometimes with partly brown pronotum, scutellum, and endocorium to almost unofrmly dark brown, with yellow base of hemelytron and cuneus (Fig.
Surface and vestiture. Dorsum shiny, head and pronotum smooth, scutellum and hemelytron weakly rugose; clothed with short, subequal in length to scale-like setae on hemelytron, strongly adpressed, simple silvery setae, scarce on vertex and pronotum, dense on scutellum and hemelytron; hemelytron additionally with scarce, silvery, narrow, apically acuminate scale-like setae (Fig.
Structure. Body oval, 2.8–2.9× as long as posterior width of pronotum, total length 3.5–3.8; head vertical, rather vide, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly attenuate and covering anterior margin of pronotum, 1.8–2.1× as wide as eye; antennal segment II at base distinctly thinner than segment I, slightly dilating apically, comparatively short, 0.5–0.6× as long as posterior width of pronotum, 0.8–0.9× as long as width of head; pronotum with broadly rounded anterior and posterior corners, 1.9–2.0× as wide as long, 1.5× as wide as head.
Genitalia. Right paramere elongate-oval, ~ 2.4× as long as wide, with straight, comparatively short, and blunt apical process (Fig.
Female. Coloration, surface and vestiture. As in male (Fig.
Genitalia. Sclerotized rings of dorsal labiate plate large, broadly oval.
Originally described from Iraq, this species was subsequently found in Turkey, Transcaucasia, Iran, and Turkmenistan (
Salix spp. (
Agalliastes fulvicornis Jakovlev, 1889: 348.
Chlamydatus fulvicornis:
Phoenicocoris flagellatus
Salicarus fulvicornis:
Lectotype of Agalliastes fulvicornis Jakovlev, 1889 • ♀ Mongolia: Selenge Aimak: Between Khara and Boroiin [Boro], 48.83°N, 106.195°E, Yakovlev coll. (AMNH_PBI 00233377) (
Paratypes of Phoenicocoris flagellatus
Other specimens examined: Mongolia • Central Aimak: Nr Songiin [Songino], SW of Ulaanbaatar, steppe, 47.81666°N, 106.66666°E, 18 Jun 1967, I. M. Kerzhner, Caragana bungei (Fabaceae), 7♂ (AMNH_PBI 00233373-AMNH_PBI 00233376), 8♀ (AMNH_PBI 00233374, AMNH_PBI 00233447-AMNH_PBI 00233450) (
Recognized by the following combination of characters: Body in male elongate, almost parallel sized, 3.1–3.6× as long as posterior width of pronotum, total length 3.7–4.0 (Fig.
Salicarus fulvicornis is a distinctive species that can be easily distinguished from its congeners. Females of this species may be confused with dark specimens of S. concinnus and S. roseri. However, S. fulvicornis is easily differentiated by the presence of flattened scales on the pronotum and scutellum, as well as by the contrastingly long simple vestiture. It further differs from both species by having long and thin apical blades of the vesica that slightly diverge from each other.
Male. Coloration. Uniformly dark brown to brown (Fig.
Surface and vestiture. Dorsum smooth; pronotum, scutellum, and hemelytron clothed with a mixture of silvery, narrow, apically acuminate scale-like setae and dense, long, ~ 1.5× as long as scales, adpressed, goldish yellow simple setae, these setae on corium sometimes dark brown (Fig.
Structure. Body elongate, almost parallel-sided, 3.1–3.6× as long as posterior width of pronotum; total length 3.7–4.0; head vertical, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly distinctly attenuate and covering anterior margin of pronotum, 1.9–2.1× as wide as eye; frons weakly convex; clypeus flat, barely visible in dorsal view; antennal segment II rod-shaped, slightly thinner than segment I, comparatively long, 0.7–0.8× as long as posterior width of pronotum, 1.0–1.1× as long as width of head; pronotum with broadly rounded anterior and posterior corners, 2.0–2.4× as wide as long, 1.5–1.6× as wide as head.
Genitalia. Right paramere elongate-oval, not expanded proximally beyond basal process, with long, straight, apically blunt apical process (Fig.
Female. Coloration, surface and vestiture. As in male (Fig.
Genitalia. Dorsal labiate plate with large and broadly oval sclerotized rings.
Known from Mongolia, adjacent steppe regions of Russia (Altai Rep., Buryatia Rep., Zabaykalsky Terr.), and Inner Mongolia in China (
Feeds on Caragana spp. (Fabaceae), including Caragana bungei Ledeb. and Caragana spinosa (L.) Vahl ex Hornem.
Salicarus (Salicarus) halimodendri
V. G.
Salicarus halimodendri:
Phoenicocoris qiliananus
Zheng, 1996 in
Salicarus qiliananus:
Holotype: Kazakhstan • East Kazakhstan Prov.: ♂ Bazarskiy Picket, Zaysan, 47.45°N, 84.86666°E, 22 Jun 1930, F.K. Lukjanovitsh, (AMNH_PBI 00233844) (
Paratypes: Kazakhstan • Almaty Prov.: Iliyskiy on Ili River, 43.52194°N, 76.82972°E, 05 Jun 1969, Seitova, 2♂ (AMNH_PBI 00233855) (
Other specimens examined: Kazakhstan • Zhambul Prov.: Karatau Mts., 4 km S of Karabastau, 42.88722°N, 70.80667°E, 557 m, 18 May 2015, F. Konstantinov & N. Simov, 1♀ (AMNH_PBI 00343015) (
Recognized by the oval body, total length: male 3.6–3.9, female 3.1–3.5; antennal segment I brown, segment II thin, basally or entirely darkened, remining segments dirty yellow (Fig.
Dark specimens of Salicarus halimodendri are somewhat similar to S. fulvicornis in having long simple setae and the arrangement of flattened scale-like setae on the dorsum. However, the latter species can be distinguished by the exceptionally long and thin, subapically bifurcate apical blades (Fig.
Male. Coloration. Variable, ranging from almost entirely dark brown to pale yellow (Fig.
Surface and vestiture. Dorsum weakly rugose, head smooth, shiny. Pronotum, scutellum, and hemelytron with a mixture of silvery scale-like setae and dense, comparatively long, ~ 1.5× as long as scales, adpressed, silvery simple setae; mesopleuron clothed with scale-like setae alone, while vertex, antenna, legs, metapleuron, and abdomen covered exclusively with adpressed silvery simple setae; tibial spines black to dark brown.
Structure. Body oval, 2.7–2.9× as long as posterior width of pronotum; total length 3.6–3.9; head vertical, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly distinctly attenuate and covering anterior margin of pronotum, 2.0–2.2× as wide as eye; frons weakly convex; clypeus flat, barely visible in dorsal view; antennal segment II rod-shaped, slightly thinner than segment I, 0.5–0.6× as long as posterior width of pronotum, 0.8–0.9× as long as width of head; pronotum with broadly rounded anterior and posterior corners, 2.3–2.5× as wide as long, 1.4–1.5× as wide as head.
Genitalia. Right paramere elongate-oval, ~ 1.7× as long as wide, with long, slightly narrowing and apically rounded apical process (Fig.
Female. Coloration, surface and vestiture. As in male. Structure. Similar to male but body smaller and more stumpy, 2.4–2.5× as long as posterior width of pronotum, total length 3.1–3.5; head with slightly more convex frons and clypeus, vertex 2.2–2.4× as wide as eye; antennal segment II distinctly thinner than segment I, 0.5× as long as posterior width of pronotum, 0.7–0.8× as long as width of head; pronotum 2.3–2.4× as wide as long, 1.4–1.5× as wide as head.
Genitalia. Sclerotized rings of dorsal labiate plate large, broadly oval. Posterior wall weakly sclerotized, with indistinctly bordered longitudinal sclerotized bands at sides (Fig.
This species inhabits plains and foothills of Central Asia within the area of its host plant, spanning from Uzbekistan and southwestern and southern Kazakhstan to Mongolia.
Salicarus halimodendri is known to feed exclusively on Caragana halodendron (Pall.) Dum. Cours. (Fabaceae), commonly known as the common salt tree. This distinctive shrub is primarily found in saline deserts and semideserts and was long classified within the monotypic genus Halimodendron.
Phoenicocoris qiliananus Zheng, 1996 was described in
Frank Wieland (Zoological Museum, University of Hamburg) and Mike Wilson (National Museum of Wales, Cardiff) provided access to collections and a supportive environment during the stay of the first author in these museums. Attilio Carapezza (Università degli Studi di Palermo) and Franco Faraci (Bardolino, Verona) offered helpful comments on the relevant literature and specimens. We are also indebted to András Orosz (Hungarian Natural History Museum) for sending pictures of the holotype of S. nitidus from the collection of
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was funded by the Russian Foundation for Basic Research (RFBR) (grant 20-54-56011) and the Iran National Science Foundation (INSF) (grant 98025845).
Conceptualisation: FK. Data curation: FK and RH. Formal analysis: FK. Writing-original draft: FK. Visualisation:FK and RH. Funding acquisition: FK and RH. Writing-review and editing: FK and RH.
Fedor V. Konstantinov https://orcid.org/0000-0002-7013-5686
Reza Hosseini https://orcid.org/0000-0002-6556-8401
All of the data that support the findings of this study are available in the main text or Supplementary Information.
USI numbers of figured specimens
Data type: pdf
Explanation note: Label data.