A new species of Salicarus (Hemiptera: Miridae: Phylinae) from Morocco

Received: 9 October 2025; accepted: 27 November 2025 · Editor: Mihail Kechev 385 Historia naturalis bulgarica 47 (12): 385–391 ISSN 2603-3186 (online) | ISSN 0205-3640 (print) · nmnhs.com/historia-naturalis-bulgarica https://doi.org/10.48027/hnb.47.123 Publication date [online]: 1 December 2025 Research article A new species of Salicarus (Hemiptera: Miridae: Phylinae) from Morocco Fedor Konstantinov1, 2, Armand Matocq3 (1) National Museum of Natural History, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel Blvd, 1000 Sofia, Bulgaria, [email protected] ✉; https://orcid.org/0000-0002-7013-5686 � (2) Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia (3) Muséum national d’Histoire naturelle, Département Systématique & Evolution, UMR 7205 MNHN/CNRS, 45 rue Buffon, F-75005 Paris (France), [email protected] ✉; https://orcid.org/0000-0002-2159-9097 � https://zoobank.org/34407320-35CA-4039-882A-30AB2841090E � Abstract: Salicarus maroccanus sp. nov. is described from Morocco, representing the first record of the genus Salicarus in North Africa. A differential diagnosis is provided, together with illustrations of the male and female genitalia and photographs of the dorsal habitus of the new species. A modified key couplet is also provided to incorporate the new species into the key to Salicarus species by Konstantinov & Hosseini (2024). Keywords: North Africa, taxonomy, plant bugs Introduction This paper provides a description of a new species of plant bug (Miridae), the most species-rich family of true bugs (Heteroptera), many of which are hostspecific. The family is particularly diverse in Mediterranean ecosystems (Cassis & Schuh, 2012) and remains insufficiently studied even within the Palaearctic region, both in terms of generic and higher-level classification and in the discovery of new species. The genus Salicarus Kerzhner, 1962 has a complex taxonomic history and was redefined by Konstantinov (2023) to include nine species. More recently, Konstantinov & Hosseini (2024) provided a comprehensive revision of the genus, including an illustrated key to species, redescriptions, and recognition of three species groups: the S. nitidus group, the S. roseri group, and the S. fulvicornis group. The first group comprises four small (2.0–2.8 mm) stumpy species densely clothed with broad silvery scales, and with distinctly swollen antennal segments I and II. These species are distributed in Mediterranean Europe, from Spain to Cyprus, and feed on legumes of the tribe Genisteae. The S. roseri species group includes three species with variable colouration, large oval body (3.0–4.0 mm) and narrow scalelike setae which are sparsely distributed only on the hemelytra and/or thoracic pleura. All three feed exclusively on Salix spp. and occur in Central and Western Asia, with S. roseri, the type species of the genus, having a broader Euro-Siberian distribution (Konstantinov & Zinovyeva, 2017). The S. fulvicornis species group comprises two species distributed mainly in Central Asia and Mongolia, and feeding on Caragana spp. They are variable in colouration, with the dorsum densely covered with narrow scales. Males are elongate and parallel-sided (3.6–4.0 mm), while females are shorter and ovoid (3.5–3.9 mm). Recent collecting in Morocco revealed a new species belonging to the S. fulvicornis species group based on vesica structure and body shape. However, it clearly differs from all Salicarus spp. in several characters, including a vestiture composed solely of Fedor Konstantinov, Armand Matocq 386 Historia naturalis bulgarica 47 (2025) simple setae, without scales on dorsum or thoracic pleurites. In addition, the discovery of this species substantially expands the known distribution of the genus, which was previously undocumented in North Africa. Materials and methods Photographs of the habitus were taken using a Keyence VHX-5000 digital microscope. Gentilic structures were examined and photographed using Zeiss Axioscope 5 equipped with a Canon EOS 700D digital camera. Partially focused images of each specimen or structure were stacked using Helicon Focus software. All measurements were taken using a calibrated eyepiece graticule and are given in millimetres. Terminology for the male genitalia follows Konstantinov (2019), with additional details illustrated in Fig. 13 of Konstantinov (2008), while terminology for the female genitalia follows Davis (1955) and Schwartz (2011). The holotype and one paratype of the new species are deposited in the Muséum national d’Histoire naturelle, Paris (MNHN), six paratypes are in the personal collection of A. Matocq, and two paratypes are housed in the National Museum of Natural History, Sofia (NMNHS). Taxonomic account Salicarus maroccanus sp. n. (Figs 1–3) urn:lsid:zoobank.org:act: A60D67A3-848E-4945-AB30-381F0E63C237 � Type material. Holotype: ♂, Morocco: Souss-Massa region, Environs of Aqesri, N 30° 37′, W 9° 30′, 207 m a.s.l., 06 May 2009, A. Matocq leg., MNHN (EH 31947). Paratypes: 1 ♀, same locality label as holotype, MNHN (EH 31948). 6 ♀♀, same locality label as holotype. Matocq coll. (Paris). 1 ♂, same locality label as holotype, NMNHS. Morocco: MarrakeshSafi region: 1 ♀, Route S501, Ijoukak, Caidat de Talat-n-Yakoub, N 30° 59′, W 08°10′, 02 May 2000, A. Matocq leg., NMNHS. Diagnosis. Recognised by the following combination of characters: Body in male elongate, almost parallel sized, total length 3.6–4.0 (Fig. 1A), female more stumpy, 2.8–3.2 (Fig. 1B); dorsum and appendages uniformly dark brown to black; segment II thin, rod-shaped; vestiture composed of adpressed dense simple setae only, without scales on dorsum or thoracic pleura; vesica strongly coiled at middle, with long, thin, and almost straight apical blades closely adjoined along their entire length (Fig. 2A, B). Salicarus marroccanus sp. n. is a distinctive species that can be easily distinguished from its congeners. It is most similar in body colouration, size, and body proportions of both sexes to the predominantly Mongolian S. fulvicornis. However, the latter is easily separated by the presence of dense flattened scales on the dorsum and a comparatively smaller secondary gonopore. Salicarus roseri, the only species of the genus lacking scales on the dorsum, differs from the new species in several characters including the presence of scale-like setae on the thoracic pleura, a yellow pattern even in the darkest-coloured specimens (e.g., yellow tibiae, apices of femora, and the last two antennal segments), and an ovoid body shape in males. All species of S. fulvicornis and S. roseri species groups further differ from the new species by having diverging apical blades of the vesica (fig. 7 in Konstantinov & Hosseini, 2024). In contrast, the vesica of S. maroccanus, equipped with long, thin, apical blades closely appressed along their entire length, is somewhat similar to that of the S. nitidus group (fig. 8 in Konstantinov & Hosseini, 2024), but differs in the straight shape of the apical blades and the smaller secondary gonopore. Furthermore, species of the S. nitidus group greatly differ from the new species in all other respects, including smaller overall size, stouter body, incrassate antennal segments I and II, and a dorsum densely clothed with broad scales. Description. Male. Colouration. Uniformly dark brown to black (Fig. 1). Head: Dark brown to black; antenna dark brown, with slightly paler segments III and IV, labium dark brown with black segment IV. Thorax: Pronotum, scutellum, and thoracic pleurites uniformly dark brown to black, hemelytron uniformly brown to dark brown, with semitransparent pale brown membrane; coxae, femora, tibiae, and tarsi dark brown, without additional markings. Abdomen: Uniformly dark brown to black. Surface and vestiture. Dorsum shining, smooth, with weakly rugose hemelytron; pronotum, scutellum, and hemelytron clothed with dense brown A new species of Salicarus (Hemiptera: Miridae: Phylinae) from Morocco 387Historia naturalis bulgarica 47 (2025) Fig. 1. Habitus of S. maroccanus sp. nov.: A – male dorsal view; B – female dorsal view; C – male head and thorax in lateral view. simple setae, semierect and longer at sides of pronotum and hemelytron, shorter and adpressed elsewhere; antenna, legs, and venter covered with similar but shorter and sometimes dirty whitish setae, scarce on thoracic pleurites, dense elsewhere; tibial spines black. Structure. Body elongate, almost parallel-sided, 3.1–3.4× as long as posterior width of pronotum; total length 3.6–4.0; Head: Vertical, slightly protruding beyond eyes anteriorly and ventrally; vertex flat, posteriorly distinctly attenuate (Fig. 1C), 1.6–2.0× as wide as eye; frons weakly convex; clypeus flat, barely visible in dorsal view; antenna inserted near ventral margin of eye; antennal segment short, cylindrical; segment II rod-shaped, slightly thinner than segment I, 0.7× as long as posterior width of pronotum, subequal to width of head; segments III and IV filiform; labium reaching mesocoxa. Thorax: Pronotum trapezoidal, with indistinct calli and broadly rounded anterior and posterior corners, 2.4× Fedor Konstantinov, Armand Matocq 388 Historia naturalis bulgarica 47 (2025) as wide as long, 1.4–1.6× as wide as head; metathoracic scent gland evaporatory area broadly triangular, peritreme oval, apically rounded. Legs: Comparatively short, femora swollen, broader medially, tibia cylindrical, tarsal segment short, segment II 1.3 × as long as segment I, segment III 1.5 Fig. 2. Male genitalia of S. maroccanus sp. nov.: A – vesica in left lateral view; B – apex of vesica, magnified, in right lateral view; C – right paramere; D – left paramere; E – apex of phallotheca. A new species of Salicarus (Hemiptera: Miridae: Phylinae) from Morocco 389Historia naturalis bulgarica 47 (2025) × as long as I and 1.2 × as long as II; claw (Fig. 3D) gradually bent from midpoint, pulvillus distinctly surpassing midpoint of claw, attached along its entire length; parempodium setiform, not apically spatulate. Genitalia. Right paramere small, broadly oval, with apically blunt apical process (Fig. 2C). Left paramere with straight, comparatively short apical process and thin, roughly triangular, and apically rounded sensory lobe (Fig. 2D). Vesica strongly coiled at middle, with two long and thin, gradually tapering apical blades closely adjoined along their entire length; larger blade twice as long as smaller one; secondary gonopore comparatively small, located on membrane far from apex, with small gonopore sclerite (Fig. 2A, B). Female. Colouration, surface and vestiture as in male (Fig. 1B). Structure. Similar to male but body shorter and oval, 2.6–2.9× as long as posterior width of pronotum, total length 2.8–3.2; head with slightly more convex frons and clypeus, and with smaller Fig. 3. Female genitalia and male claw of S. maroccanus sp. nov.: A – dorsal labiate plate of the bursa copulatrix; B – bursa copulatrix on ventral view, showing sclerites encircling vulva and vestibulum; C – posterior wall of bursa copulatrix; D – male hind claw. Fedor Konstantinov, Armand Matocq 390 Historia naturalis bulgarica 47 (2025) eyes, vertex 2.0–2.2 × as wide as eye; antennal segment II somewhat thinner than in male and distinctly thinner than segment I, 0.6–0.7× as long as posterior width of pronotum, 0.8–0.9× as long as width of head; pronotum 2.3–2.5× as wide as long, 1.3–1.4× as wide as head. Genitalia. Dorsal labiate plate with large and broadly oval sclerotised rings (Fig. 3A). Sclerites encircling vulva triangular, symmetrical (Fig. 3B). Vestibulum characteristically long, S-shaped (Fig. 3B). Posterior wall clothed with dense fine spinules and longitudinal sclerotised bands at sides (Fig. 3C). Distribution. Known from two localities in Souss-Massa and Marrakesh-Safi regions of Morocco. Host. Unknown. Discussion. Salicarus maroccanus sp. n. is assigned to the respective genus based on the diagnosis outlined in Konstantinov (2023) and a combination of characters including the strongly flattened and sloping head with the clypeus barely visible from above, the posteriorly carinate vertex, the contrastingly large, S-shaped vestibulum of the bursa copulatrix, the broadly rounded right paramere, the vesica strongly coiled at middle, with two gradually tapering apical blades that remain fused along most of their length, and the secondary gonopore with a small gonopore sclerite. However, two characters set this species, otherwise belonging to the S. fulvicornis species group, from congeners: the absence of scalelike setae and the setiform, rather than apically spatulate parempodium. The latter feature was documented in detail by Schwartz & Stonedahl (2004) and has previously been considered one of the non-unique synapomorphies uniting the Salicarus + Phoenicocoris clade (Konstantinov, 2023). Spatulate parempodia have also been reported in otherwise unrelated taxa, such as Campylomma spp. (Konstantinov et al., 2015), and Mesopsallus spp. (Schwartz & Stonedahl, 2004; Konstantinov, 2023). Broader comparative studies of pretarsal structures are therefore required to clarify the distribution and evolutionary significance of this character. Phoenicocoris vidali (Lindberg, 1940), a poorly known species, was originally described from nine males and a single female collected in Ras-el-Ma, Atlas Moutains, Morocco, and placed in the genus Sthenarus. The species is still apparently known only from the type series, some specimens of which were later studied by Wagner, who provided a redescription and schematic drawings of the male genitalia, including the vesica (fig. 2k in Wagner, 1960, reprinted as fig. 699b1 in Wagner 1975). According to Wagner (1960: 11), the vesica is S-shaped, twisted, slender, of simple structure, distally with two unequal, slender chitinous rods, alongside a membranous area. Wagner (1975: 105) further described it as very slender, with the apical part strongly curved and bearing two closely adjoining long chitinous rods. In their comprehensive revision of Phoenicocoris, Schwartz & Stonedahl (2004) supported the placement of P. vidali within the genus based on these descriptions. However, direct examination of the vesica of this species is still highly desirable for unequivocal generic assignment, given the close relationship between Phoenicocoris and Salicarus and their reliable separation based on male genitalia. In any case, P. vidali is clearly distinct from S. maroccanus sp. n. in several characters, including its smaller body size (2.2–2.4 mm), short antennal segment II (subequal to head width and distinctly swollen, spindle-shaped in males and somewhat clubshaped, distally dilated in females), and yellow fore and middle legs with darkened apices of femora. The new species may be incorporated into the key presented by Konstantinov & Hosseini (2024) by modifying couplet 5 and adding a new couplet 6 as follows: 5. Hemelytron without scale-like setae, clothed only with simple setae . . . . . . . . . . . . . . . . . . . . . 6 — Hemelytron clothed with a mixture of simple setae and narrow, apically acuminate scale-like setae (sometimes scarce and easily obliterated) (fig. 3C, F, I in Konstantinov & Hosseini 2024) . . . . . . . 7 (6 in Konstantinov & Hosseini 2024) 6. Body of male ovoid; colouration variable in both sexes, but even the darkest specimens with dirty yellow tibiae and apices of femora; dorsum often with additional yellow markings (fig. 2F–I in Konstantinov & Hosseini 2024). Thoracic pleura clothed with scale-like setae. Vesica with straight, robust, diverging apical blades (fig. 7G, H in Konstantinov & Hosseini 2024). Widely distributed in the Palearctic. On Salix spp. ............................. S. roseri — Body of male parallel-sided; both sexes uniformly dark brown to brown. Thoracic pleura without scale-like setae. Vesica with thin, straight apical blades closely adjoined along their entire length (fig. 2A, B). Morocco .................... S. maroccanus sp. n. A new species of Salicarus (Hemiptera: Miridae: Phylinae) from Morocco 391Historia naturalis bulgarica 47 (2025) related genera (Hemiptera: Miridae: Phylinae). Arthropod Systematics & Phylogeny 81: 845– 879. https://doi.org/10.3897/asp.81.e104396 � Konstantinov F.V., Neimorovets V.V., Korzeev A.I. 2015 The genus Campylomma Reuter, 1878 (Hemiptera: Heteroptera: Miridae: Phylinae): two new synonyms and discussion of the diagnosis. Zootaxa 3974 (2): 203–216. https://doi.org/10.11646/zootaxa.3974.2.5 � Konstantinov F.V., Zinovyeva A.N. 2017 New records of the plant bug subfamilies Orthotylinae and Phylinae (Heteroptera, Miridae) from the European part of Russia. Entomological Review 97 (6): 775–793. https://doi.org/10.1134/S0013873817060070 � Schwartz M.D. 2011 Revision and phylogenetic analysis of the north American genus Slaterocoris Wagner with new synonymy, the description of five new species and a new genus from Mexico, and a review of the genus Scalponotatus Kelton (Heteroptera: Miridae: Orthotylinae). Bulletin of the American Museum of Natural History 354: 1– 290. https://doi.org/10.1206/354.1 � Schwartz M.D., Stonedahl G.M. 2004 Revision of Phoenicocoris Reuter with descriptions of three new species from North America and a new genus from Japan (Heteroptera: Miridae: Phylinae). American Museum Novitates 3464: 1–55. https://doi.org/10.1206/00030082(2004)464<0001:ROPRWD>2.0.CO;2 � Wagner E. 1960 Neuer Beitrag zur Systematik der Gattungen Sthenarus Fieber und Psallus Fieber (Hem. Het. Miridae). Mitteilungen der Deutschen Entomologischen Gesellschaft 19: 9–13. Wagner E. 1975 Die Miridae Hahn, 183l, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Teil 3. Entomologische Abhandlungen 40 Suppl., [ii +] 483 pp. https://doi.org/10.1515/9783112653241 � Acknowledgements We are grateful to Jean-Claude Streito (INRA) for taking dorsal habitus pictures for this paper. We also thank the two anonymous reviewers for their constructive comments. Funding for the laboratory work of F. Konstantinov in this study was partly provided by the Bulgarian National Science Fund (project KP-06-PN71/13, BG-175467353-2023130067 “Exploring the evolution of the plant bug subfamily Phylinae (Insecta, Hemiptera, Miridae): insights from phylogenetic reconstructions, host associations, and distributional patterns”). Conflicts of interest No potential conflict of interest has been reported by the authors, reviewers, or subject editor. As a member of the editorial board Fedor Konstantinov recused himself from discussions and decisions on the manuscript. References Cassis G., Schuh R.T. 2012 Systematics, biodiversity, biogeography, and host associations of the Miridae (Insecta: Hemiptera: Heteroptera: Cimicomorpha). Annual Review of Entomology 57: 377–404. https://doi.org/10.1146/annurev-ento-121510133533 � Davis N.T. 1955 Morphology of the female organs of reproduction in the Miridae (Hemiptera). 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