First report of Phymatinae and Holoptilinae from the Middle East, with a description of a new species of Putoniola Bergroth (Heteroptera: Reduviidae: Holoptilinae) from Israel

Israel Journal of entomology, Vol. 54, pp. 45–94 (30 october 2025) DoI: 10.5281/zenodo.17464510; Issn (online) 2224-6304 received: 6 July 2025 / revised: 30 July 2025 / accepted: 1 september 2025 urn:lsid:zoobank.org:pub:403CB1C6-B6D0-4248-BC4B-65fB1eaD5f1e First report of Phymatinae and Holoptilinae from the Middle East, with a description of a new species of Putoniola Bergroth (Heteroptera: Reduviidae: Holoptilinae) from Israel Tanya Novoselska 1* & Tamar Feldstein-Farkash 1,2 1The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies, Tel Aviv University, Tel Aviv, 6997801 Israel. E-mail: [email protected] 2School of Zoology, Tel Aviv University, Tel Aviv, 6997801 Israel. E-mail: [email protected] *Corresponding author: [email protected] ABSTRACT this study presents the first record and a comprehensive review of the reduviidae subfamilies Phymatinae and Holoptilinae in Israel. the paper provides up dated diagnoses, detailed genitalia illustrations and a revised identification key for species of the genus Putoniola Bergroth, 1898 (Holoptilinae), along with description of a new species, Putoniola asvadurovi sp. n. the findings of Phymata (Phymata) monstrosa (fabricius, 1794) (Phymatinae) in Israel confirm an old middle eastern record of the species from ‘syria’ and considerably extend its previous West mediterranean distribution. Information on distribution of both species and their biology and host plant associations is provided. molecular phylogenetic analyses employing CoI and 16s mitochondrial genes confirm the broad “Phymatine complex” and successfully integrate the new Putoniola species. We contribute to addressing a critical gap in knowledge of the phylogenetic relationships of tribes within the subfamilies Holoptilinae and Phymatinae, which are not yet fully understood and require further investigation. this is the first molecular study to include Phymata (P.) monstrosa (Phymatinae) and to add Putoniola asvadurovi sp. n. (Holoptilinae), both originating from the Pa laearctic region. KeyWorDs: assassin bugs, biodiversity, Dasycnemini, Hemiptera, identification key, Israel, molecular taxonomy, new species, zoogeography. INTRODUCTION the reduviidae (assassin bugs) are a highly diverse group of predatory insects with a wide range of prey specializations and associated morphological diversity. the family encompasses at least 25 subfamilies, around 981 genera and 7000 de scribed species (maldonado Capriles 1990; moulet et al. 2014; Henry 2017; Weirauch et al. 2019; schuh & Weirauch 2020; ye et al. 2021). Kerzhner (1992) amended maldonado Capriles’s (1990) catalogue of the reduviidae after comparing it with unpublished data of P.V. Putshkov & V.g. Putshkov. a revised classification by masonick et al. (2025), based on molecular data and morphological characters, re cognizes 19 subfamilies and 40 tribes. the knowledge of the fauna of Israeli reduviidae is currently limited, with probably numerous species yet to be discovered. representatives of nine subfamilies of the reduviidae have been so far documented in Israel (Bodenheimer 1937; 46 Israel Journal of entomology, Vol. 54, 2025 Bytinski-salz & sternlicht 1967; Dispons 1964a; Halperin 1990; linnavuori 1952, 1960, 1961, 1973; Wygodzinsky 1952; Hoberlandt1952 ; moulet 2005, 2008; van der Heyden 2018). However, there has been no comprehensive review of these taxa. to date, the Israel national Collection at the steinhardt museum of natural History contains 11 subfamilies of reduviidae, constituting one of the most inclusive collections of these insects in the region. two subfamilies—Phymatinae and Holoptilinae—are recorded in Israel for the first time, thus significantly expanding the distribution of these insects in the middle east. the study provides the first documentation of the Phymatinae subfamily, particularly of the Phymatini tribe, in Israel, based on the first recorded occurrence of the species Phymata (Phymata) monstrosa (fabricius, 1794). It also documents the first record of the Holoptilinae subfamily from Israel based on the description of a new species of the genus Putoniola Bergroth, 1898. this species is the first such species to receive a molecular barcode sequence in its genus. the phylogenetic trees were reconstructed from the CoI and 16s sequences. the resulting expanded 16s tree will serve as a foundational element for future research, particularly concerning the Holoptilinae subfamily. MATERIALS AND METHODS the following institutional acronyms are used in the paper: amnH – american museum of natural History, new york, usa; HnHm – Hungarian natural History museum, Budapest, Hungary; nCBI – the national Center for Biotechnology Information, Bethesda, usa; nHmW – naturhistorisches museum in Wien, Vienna, austria; nHm – the natural History museum, london, uK; smnHtau – the steinhardt museum of natural History, tel aviv, Israel; tuZ – university of tartu, Zoological Collections, estonia. the present study is based on pinned specimens deposited in the collection of the smnHtau. In the smnHtau accession numbers (e.g., smnHtau In.326234), “In.” refers to the Insect Collection catalogue. the following species have been avai lable for examination only on photographs: Putoniola vaulogeri equatorialis Dispons, 1966 and Putoniola vaulogeri vaulogeri (montandon, 1897) (amnH); and Rudebeckocoris fabiani Vásárhelyi, 1980, with identification labels by D. rédei (HnHm). the following species are known to us only from the original and subsequent descriptions and drawings, and from the nHm online resourses: Putoniola arabica, P. atakorensis, P. kermana, P. v. vaulogeri and P. v. angusticeps. the nomenclature follows Putshkov & moulet (2009) and masonick et al. (2017, 2025). the terminology follows Putshkov & moulet (2009), forero et al. (2010), rédei & tsai (2011), masonick & Weirauch (2020a, b) and masonick et al. (2017, 2025). for morphological identification of the veins, the forewings were separated from the body by cutting through the pleural sclerites and mounted on a slide. noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 47 species identification required examination of both male and female genitalia. the parameres were not entirely exposed; although they could partly be examined without dissection, they had to be dissected to observe the base of the parameres. the dissected genitalia were placed in glycerol gelatin for studying and imaging. the total length was measured from the clypeus to the tip of the abdomen; the length of the forewing was measured from the base of the clavus to the forewing apex; the length of the abdomen (dorsal view) was measured from the medial area of urotergite I to the apex of urotergite VII. the last measurement is crucial for the species identification based on the ratio of forewing length to abdomen length. all specimens were examined under a leica m125 stereo microscopes. figures 43–47 were taken using a Canon eos 6D camera, with the images stacked manually. all other photographs were taken with the touptek XCam4K8mPB camera attached to a leica mZ16 binocular microscope, using mII ImageView 3.7 software. the images were touched with lightroom CC 2015, stacked with Helicon focus and edited with adobe Photoshop Cs6 to produce the final plates. the specimen smnHtau In.326234 was cleaned, dried and sputter coated prior to examination and photographing under a Quanta 200feg environmental scanning electron microscope at Center for nanoscience and nanotechnology, tel aviv university, Israel). the distribution maps were prepared using google earth Pro images. the plant nomenclature follows Wfo (2025). Weather conditions were sourced from the Israel meteorological service (2025) website. Dna was extracted from specimens smnHtau In.326234, 326236, 454881 (Putoniola asvadurovi sp. n.) and smnHtau In.370391, 370462 (Phymata (P.) monstrosa) using different commercial kits for genomic Dna extraction from tissue (geneaid, new england Biolabs and Qiagen). the only Dna that amplified in PCrj, however, was a combination of tissue lysis with atl buffer from Dneasy kit (Qiagen) followed by extraction with phenol:chlorophorm:isoamyl alcohol (25:24:1) and ethanol precipitation, for specimens smnHtau In.370391 and smnHtau In.454881. the 16s gene was amplified and sequenced with the 16sar-l and 16sbr-H primers (Palumbi et al. 1991). Cytochrome c oxidase subunit I sequence of Phymata (P.) monstrosa was obtained using degenerate primers jglCo1490 and jgHCo2198 (geller et al. 2013). CoI for Putoniola asvadurovi sp. n. could not be amplified. sequences were deposited in genBank under accession numbers PV188576-7 (16s) and PV179415 (CoI). for the pylogenetic reconstruction sequences of the 16s rrna gene from the Phymatinae and Holoptilinae were downloaded from genBank (table 3). sequen ces of the genus Neocentrocnemis miller, 1956 were also downloaded as the Centrocnemidinae (reduviidae) was the closest subfamily to those analyzed by masonick et al. (2025). sequences of Microtomus Illiger, 1807 (reduviidae: Ham macerinae) were used to root the tree, following masonick et al. (2017). sequences were aligned using mafft 7.304 (Katoh et al. 2019) applying the e-Ins-i 48 Israel Journal of entomology, Vol. 54, 2025 parameters. ambiguous positions were removed, resulting in a final 497-bp-long alignment. the phylogenetic tree was reconstructed under the maximum likelihood criterion with Phyml 3.0 (Dereeper et al. 2008; guindon et al. 2010) using the gtr model of sequence evolution. Bootstrap percentages were computed based on 100 replicates. similarly, a phylogenetic tree of CoI was reconstructed to verify the position of Phymata (P.) monstrosa within the genus. RESULTS Taxonomy family reduviidae latreille, 1807 subfamily Holoptilinae amyot & audinet-serville, 1843 Holoptilinae amyot & audinet-serville, 1843: 318 (Holoptilides). Type genus: Holoptilus le Peletier & audinet-serville, 1825: 280. the subfamily Holoptilinae, also known as feather-legged assassin bugs or ant wolves, comprises almost 80 species in 16 genera (including three fossil ones), divided among three tribes: aradellini Wygodzinsky & usinger, 1963, Dasycnemini Wygodzinsky & usinger, 1963, and Holoptilini le Peletier & audinet-serville, 1825. the majority of Holloptilinae are known from the Palaearctic and oriental regions, although they are also significantly represented in the afrotoropical, australasian and neotropical regions (Wygodzinsky & usinger 1963; malipatil 1985, 2018; maldonado Capriles 1990; Poinar 1991; Putshkov & Putshkov 1996; Putshkov & moulet 2009; rédei & tsai 2011; moulet et al. 2014; shah et al. 2019). the Holoptilinae are characterized by the very long and often stiff pilosity on the body and appendages, the 1st antennomere short and thick, much shorter than setae on the 2nd joint, the head often as long as or shorter than wide and the forewings with a reduced and narrowed corium and a diminished clavus (rohdendorf 1949; Putshkov & moulet 2009; Weirauch et al. 2010; moulet et al. 2014). the Holoptilinae are considered rare. their apparent rarity may be due to their secretive nature and hard-to-find habitats. they are known to survive for extended periods without food (Jacobson 1911; Priesner & alfieri 1953). the adults can be nocturnal or diurnal, are often found on plants, often on trees, but also on the ground, in plant debris and in the bird plumage, where they prey on various arthropods, particularly on ants. the best-known predatory behaviour is that of the Javanese Ptilocerus venosus (Walker, 1873) that feeds on the ants Dolichoderus thoracicus (smith, 1860), paralyzed by the fluid secreted by a special abdominal gland called a ‘trichome’ (Jacobson 1911). the Holoptilinae species exhibit a distinctive egg attachment mechanism, each egg being individually cemented to the substrate (sahayaraj & Hassan 2023). the behaviour and habitats of various Holoptilinae species are detailed by Jacobson (1911), Villiers (1948), Priesner & alfieri (1953), Dispons (1964b), Putshkov (1987), murugan (1988), Weirauch et al. (2014), and moulet et al. (2014). noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 49 tribe Dasycnemini Wygodzinsky & usinger, 1963 Dasycnemini Wygodzinsky & usinger, 1963: 49. Type genus: Dasycnemus Bergroth, 1898: 186. Diagnosis: the setae of the antennae and legs are longer than the diameter of the respective segments. the membrane has two or three free longitudinal veins that are not connected beyond the middle of the membrane. additionally, the trichome is absent, and the species are typically shorter than 5 mm. Distribution: five extant genera are known from sub-saharan africa (including socotra I.), western north africa and arabian Peninsula (montandon 1897; Villiers 1945, 1956, 1957, 1960, 1986; Wygodzinsky & usinger 1963; linnavuori 1964, 1974, 1986; Dispons 1966; maldonado Capriles 1990; Putshkov & Putshkov 1996; Wranik 2003; moulet 2006; moulet et al. 2014). the present communication is the first record of this tribe for Israel and the levant. genus Putoniola Bergroth, 1898 Putoniella montandon, 1897: 102. (Junior homonym of Putoniella Kieffer, 1896 (Diptera)) Putoniola Bergroth, 1898: 186. (replacement name for Putoniella montandon, 1897) Rudebeckocoris miller, 1956: 434; Villiers 1986: 216; Kerzhner 1992: 59; Putshkov & Putshkov 1996: 170. type species: Rudebeckocoris pretoriae miller, 1956, by orig. des. (s. africa, transvaal) Type species: Putoniella vaulogeri montandon, 1897. Diagnosis: the antennae and legs are densely covered with setae; the setae are notably longer than the diameter of the respective segments; the scutellum features a continuous row of long setae along its posterior margin. the projection of the second antennal segment extends beyond the base of the third antennomere. the pronotum is approximately twice as wide as long. the membrane with two or three free longitudinal veins, not joined beyond the middle of the membrane; the middle vein may occasionally fork basally. the abdominal trichome is absent. We propose the most important distinguishing features for the genus Putoniola as follows: the ratio of forewing length to apex of abdomen length; head and pronotum shape; the ratio of head width to collar width; the shape of the labium shape segments and intercalary sclerites; paired setae; the veining of the forewings; the pygophore and the shape of the paramere. Distribution: the known distribution of the genus Putoniola includes the af rotropical region and the southern Palaearctic (north africa and middle east) (see below for a list of species and subspecies). It is noteworthy that this genus has not been previously reported in countries bordering Israel. Species included: Putoniola arabica Dispons, 1966: yemen (linnavuori 1986; Putshkov & Putshkov, 1996; Putshkov & Pluot-sigwalt 2008); Putoniola asvadurovi sp. n.: Israel; P. atakorensis Villiers, 1952: senegal (Villiers 1952), Cameroon, Central african republic, senegal (linnavuori 1986); P. kermana Dispons, 1964: Iran (Dispons 1964, 1966; linnavuori 1986; Putshkov & Putshkov 1996; Putshkov 50 Israel Journal of entomology, Vol. 54, 2025 & Pluot-sigwalt 2008); P. vaulogeri angusticeps linnavuori, 1986: yemen, uae (moulet et al. 2014), saudi arabia (linnavuori 1986; Putshkov & Putshkov 1996); P. vaulogeri equatorialis Dispons, 1966: sudan (Villiers 1956, 1957, 1960, 1986; Dispons 1966; linnavuori 1986, 1974; maldonado Cariles 1990; moulet et al. 2014), eritrea, somalia, sudan (Putshkov & Putshkov 1996), yemen, uae (moulet et al. 2014); P. vaulogeri vaulogeri (montandon, 1897): algeria, eritrea, somalia, sudan (linnavuori 1986), Chad (linnavuori 1986; Putshkov & Putshkov 1996), niger (Villiers 1952), Iran (ghahari et al. 2024), tunisia (montandon 1897; linnavuori 1986; Villiers 1952, 1986; Putshkov & Putshkov 1996). Notes: there has been some confusion regarding synonymy of Rudebeckocoris with Putoniola, based on a mistake. maldonado Capriles (1990) ignored the synonymy established by Villiers (1986). therefore, the text of maldonado Capriles cannot be interpreted as a rejection of Villier’s synonymy. Kerzhner (1992) reported correctly Villiers (1986) and the synonymy of Rudebeckocoris with Putoniola, implicitly accepting it. Putshkov and Putshkov (1996) listed Rudebeckocoris as a synonym of Putoniola. In conclusion, nobody has ever objected to the synonymy established by Villier (1986). as to the identification labels by D. rédei (HnHm), they cannot be interpreted as a taxonomic action, probably he simply meant that those specimens belonged to the taxon described by Vásárhely as Rudebeckocoris fabiani. moulet (2014) suggested that P. v. angusticeps might not be a valid subspecies, as the male genitalia appear the same as of the nominotypical species. Putoniola asvadurovi sp. n. figs 1–42, 68a, 70; table 1 LSID: urn:lsid:zoobank.org:act:8e4e4f6f-D11f-418a-B757-f915BC158f6a. Etymology: the species is named in honour of the first author’s son, artem asvadurov, who assisted in collecting of the type material. Figs 1–3. Putoniola asvadurovi sp. n., holotype smnHtau In.326233: (1) dorsal view, (2) ventral view, (3) specimen labels. 1 2 3 noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 51 Diagnosis: the collected specimens can be placed in the genus Putoniola Bergroth, 1898 based on the following combination of characters: the elongated setae on the antennae and legs, exceeding the diameter of the re s pective segments, distinguish P. asvadurovi sp. n. from the members of the aradellini. the absence of the abdominal trichomes distinguishes the new species from the Holoptilini. the fore wing with two distinct cells, the veins of the corium with a single row of the long setae and the 2nd antennal segment with the apical projection separate P. asvadurovi sp. n. from the members of the Dasymothini. the new species can be clearly distinguished from congeners based on the following combination of morphometric and structural characteristics: In Putoniola arabica, the forewings only extend slightly beyond the apex of the abdomen, whereas in P. asvadurovi sp. n. the forewings are longer than the apex of the abdomen; In Putoniola atakorensis, P. vaulogeri equatorialis, P. vaulogeri angusticeps and P. vaulogeri vaulogeri, the forewing length to abdomen length ratio is 1:1.9, body larger, with a male body length of 3.0–3.5 mm, whereas P. asvadurovi forewing to abdomen ratio is 1:1.68–1.69 and male body length of 2.53–2.54 mm; In Putoniola kermana, the ratio of forewing length to abdomen length is 1:1.2 compared to 1:1.68–1.69 in P. asvadurovi sp. n., the length ratio of the antennal segments aII:aIII+aIV is 1:1.46 compared to 1:1.70–1.73 in P. asvadurovi sp. n., and the male body is longer of 3.0–3.5 mm compared to 2.54–2.55 mm for Figs 4–9. sem micrographs of the habitus of Putoniola asvadurovi sp. n., paratype smnHtau In.326234: (4) dorsal view, (5) anterodorsal view, (6) anterodorsolateral view, (7) posteroventral view, (8) posteroventrolateral view, (9) posterolateral view. 4 9 87 6 5 52 Israel Journal of entomology, Vol. 54, 2025 P. asvadurovi sp. n. It should also be noted that, despite Dispons’ (1964) asser tion that the aII projection is extremely variable in shape, length and thickness, in all of the examined specimens a cylindrical, apically rounded the aII projection was found. the paramere in P. asvadurovi sp. n. is broadened at base, bearing a dis tinct ventromedial keel. the last character is indicated for the first time for Puto niola and is unique to P. asvadurovi sp. n. Description: Male. macropterous (figs 1, 2, 4–9, 21–25). Colour (figs 1, 2, 21, 26): general colouration pale-golden in shade; eyes redviolet; ocelli, apical part of 3rd segment of antenna, epipleurite, tarsi pale to dark brown; clavus at apical ⅓ and at base pale to dark brown; middle part hyaline; veins pale to dark brown; cells of corium hyaline; membrane semitransparent with veins and anterior margin light pale to grey; hindwings milky-white. Figs 10–13. sem micrographs of the head of Putoniola asvadurovi sp. n., paratype smnHtau In.326234: (10) anterior view, (11) dorsal view, (12) anterolateral view, (13) anterodorsolateral view. 10 1312 11 noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 53 Body (figs 1, 2, 4–25, 30–35), excluding antennae and legs, with round granules of varying densities of coverage across different body parts. Body covered with two types of setae: greyish downy, felt-like setae, and long, stiff, toothed setae originating from the suckers, slightly tapering at the apex and notably prominent. Colar, pronotum, thorax, trochanters and coxae covered with greyish downy, feltlike setae. antennae, head, rostrum, pronotum, femora and tibiae, scutellum, corium veins, pygophore and paramere with scattered long, stiff, toothed setae. abdominal venter with a few scattered long, stiff, toothed setae. legs covered with serrate setae; apex of hind femora with brush-like tuft of serrate setae (fig. 23); tarsi covered with scattered long, stiff, toothed setae. Figs 14–17. sem micrographs of the head and antennae of Putoniola asvadurovi sp. n., paratype smnHtau In.326234: (14) anterolateral view showing labrum (lm), 3-segmented labium (lII–lIV), stylet fascicle (sf), and labial groove (lg); (15) anterior view; (16) antennomere I (aI), lateral view; (17) antennomeres II–III (aII, aIII) and projection of aII. 17 14 15 16 60 Israel Journal of entomology, Vol. 54, 2025 dorsally with serrate setae; t8 forming transverse ridge, with inward curved margins. Dorsal abdominal scent glands (Dag1, Dag2) located on anterior margin of tergites t5 and t6 (figs 26–29). Male genitalia. Pygophore ovoid (figs 2, 7–9, 23–36, 38), smooth; ventral rim forming thick transverse ridge posteriorly, median projection (mp) narrowly spatulate with noticeably convex dorsal side. Parameres (figs 23, 24, 29, 30–35) symmetrical, strongly curved and broadened at base that bears distinct ventro-medial keel, loose screw-shaped in distal half, and with markedly flattened tip; with serrate setae and sensilla; tightly wrapped around pygophore, crossing each other in resting position. the position of the paramere can vary, with right paramere wrapping around left one or vice versa. Phallus (figs 36–41) distinctly articulated, elongated vertically. articulatory apparatus small. Basal plates (bp) short, twisted rope-like, joined at base, V-divided into 1+1 straight and smooth branches, each of which articulates with chitinous protrusions dorsoapical portion of struts (strda); support bridge prolongation (sbp) twisted rope-like; strda ribbon-shaped, tapering in length towards apex becoming sinuous and filiform (fig. 41). Phallotheca unsclerotized, flattened, fairly shapeless. Female. unknown. Measurements: table 1. Holotype: Israel: ♂, macropterous adult, SMNHTAU In.“326233.Israel:/ 'avrona nature reserve,/ tree #37, Control nW,/ without oil spill year 2014/ 29°40'46.9"n, 34°59'59.3"e/ 41m. 04.iv.2017/ noVoselsKa, t./ asVaDuroV, a./ on Vachellia tortilis subsp./ raddiana/ Beating vegetation”. the specimen is pinned and marked with the printed red label (fig. 3). Paratypes: Israel: 'Arava Valley: 1♂, macropterous adult, SMNHTAU In.326234, same as holotype, pinned; SEM micrographs (Figs 4–25, 30–35); 1♂, macropterous adult, SMNHTAU In.335902, same as holotype [this specimen was collected dead and mostly damaged, its parts have been glued to a cardboard]; 1♂, macropterous adult, SMNHTAU In.326235, 'Avrona Nature Reserve, 29°40'43.8"N 35°00'03.2"e, 41 m a.s.l., 15–16.v.2018; n. segev, pitfall trap under Vachellia tortilis, tree #32 (control, without oil spills), pinned; 1♂, macropterous adult, SMNHTAU In.326236, 'Avrona Nature Reserve, 29°40'43.8"n 35°00'03.2"e, 41 m a.s.l., 23–25.v.2018, g. sinaiko, pitfall trap under Vachellia tortilis, Tree #32 (control, without oil spills), pinned; 1♂, macropterous adult, SMNHTAU In.454881, 'Avrona nature reserve, 29°40'46.9"n 34°59'59.3"e, 41 m a.s.l., 23.v.2020, t. novoselska, a. asvadurov, beating Vachellia tortilis raddiana, tree #37 (control, without oil spills). Ecology: the 'arava Valley region experienced significant Holocene tectonic acti vity, resulting in substantial alterations to its geomorphology (rinat et al. 2014). the entire region, and the 'avrona nature reserve in particular, is characterised by extreme arid conditions (amit et al. 1993). Climate indicators for the 'arava Valley region, including a notable 0 mm rainfall during the study days, reveal distinct daily and seasonal variations. Precipitation data supplied by the Israel meteorological service (2025) reflect varying rainfall totals in the study area between 2017 and 2020. specifically, a total of 13.6 mm of rainfall was recorded in 2017, followed by a significant increase to 52.9 mm in 2018. In 2019, the precipitation decreased to 11.8 mm, and the period covering late 2020 (30.xi–31.xii.2020) showed a combined total of 30.7 mm. noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 61 Table 1. measurements of Putoniola asvadurovi sp. n., in mm. Characters Holotype, ♂ Paratypes, ♂ (n=5) Body length to apex of hemelytron 2.54 2.53–2.54 Body length to apex of abdomen 2.46 2.45–2.46 Head length 0.34 0.33–0.34 Head width across the eyes 0.65 0.62–0.65 ratio head length to width 0.52 0.52–0.53 labrum length 0.05 0.05 labium total length 0.51 0.50–0.52 lII 0.28 0.27–0.28 lIII 0.11 0.11 lIV 0.12 0.12–0.13 labium segments length ratio, lII:lIII:lIV 1:0.39:0.43 1:0.39–0.41:0.44–0.46 Interocular distance 0.35 0.35 Distance between ocelli 0.28 0.28 Collar length 0.31 0.30–0.31 Pronotal disk region length across middle line 0.39 0.39–0.40 Pronotum width across humeral angles 0.83 0.83–0.84 scutellum length 0.14 0.13–0.14 scutellum width 0.40 0.38–0.40 antennae, total length 1.64 1.57–1.65 aI 0.14 0.13–0.14 aII 0.95 0.91–0.95 aIII+aIV 0.55 0.53–0.56 apical projection on the aII 0.06 0.06–0.07 length ratio aI:aII:aIII+aIV 1:6.79:3.93 1:6.79–7.00:4.00–4.08 length ratio aII:aIII+aIV 1:1.73 1:1.70–1.72 abdomen length 1.12 1.12–1.13 abdomen width, maximum 1.03 1.02–1.03 genital capsule length 0.36 0.36 genital capsule width 0.32 0.32 forewing length 1.88 1.89–1.90 ratio forewing length to abdomen length 1.68 1.68–1.69 fore leg, total length 1.51 1.52–1.56 Coxa 0.10 0.10–0.11 trochanter 0.17 0.17–0.18 femur 0.48 0.49–0.50 tibia 0.52 0.52 tarsus 0.21 0.21–0.22 tarsomere I 0.04 0.04–0.05 62 Israel Journal of entomology, Vol. 54, 2025 all specimens of P. asvadurovi sp. n. were collected on and under the acacia trees Vachellia tortilis subsp. raddiana (savi) Kyal. & Boatwr. (syn. Acacia raddiana savi) and Vachellia tortilis subsp. tortilis (forssk.) galasso & Banfi (syn. Acacia tortilis (forssk.) Hayne), either by beating or in pitfall traps. The ants (Hymenoptera: Formicidae) comprise 90 % of the insect population and are therefore most common, nearly dominant insects in the study area (segev 2021; gavish-regev et al. 2022). Based on this fact and on the previous records of ant-predation we can assume that the ants are the main potential primary prey of the P. asvadurovi sp. n. the association of P. asvadurovi sp. n. with Vachellia spp. can implify that they probably prey on the arboreal ant species. the P. asvadurovi sp. n. specimens collected were not covered in sand particles or dust, which is unusual for many terrestrial or subterranean desert species. this further supports their arboreal (tree-dwelling) existence or may also indicate an (unknown) mechanism enabling them to remain clean. our repeated attempts to collect specimens of P. asvadurovi sp. n. using a light trap have been unsuccessful, mainly due to persistent winds averaging 15–20 km/s. Characters Holotype, ♂ Paratypes, ♂ (n=5) tarsomere II 0.17 0.17–0.18 Claw 0.03 0.03 mid leg total length 1.95 1.93–1.98 Coxa 0.11 0.11–0.12 trochanter 0.18 0.18–0.19 femur 0.63 0.62–0.63 tibia 0.69 0.68–0.69 tarsus 0.28 0.28–0.29 tarsomere I 0.04 0.04–0.06 tarsomere II 0.24 0.24–0.23 Claw 0.06 0.06 Hind leg total length 3.70 3.67–3.76 Coxa 0.19 0.18–0.20 trochanter 0.24 0.23–0.24 femur 0.69 0.69–0.71 tibia 2.21 2.21–2.22 tarsus 0.30 0.29–0.31 tarsomere I 0.05 0.04–0.06 tarsomere II 0.25 0.25–0.25 Claw 0.07 0.07–0.08 noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 63 Revised key to males of all known species and subspecies of the genus Putoniola (based on Dispons 1966) 1 Forewings extend slightly beyond apex of abdomen. Veins chocolate brown, appear obtusely careniform. Paramere straight. Body length ♂ 2 mm ................ ............................................................................................................. P. arabica – Forewings widely surpassing apex of abdomen. Veins testaceous. Paramere straight or curved ...............................................................................................2 2 Length ratio of forewings to abdomen less than 1:2. Paramere curved and broadened at base with or without ventro-medial keel ......................................3 – Length ratio of forewings to abdomen at least 1:2. Paramere straight ..............4 3 Length ratio of forewings to abdomen 1:1.2 in male. Paramere without ventromedial keel. Projection of aII apically variable. length ratio aII: aIII+aIV 1:1.46. Body length ♂ 3.0–3.5 mm, ♀ 2.7 mm ................................ P. kermana – Length ratio of forewings to abdomen is at 1:1.68–1.69 in male. Paramere with ventro-medial keel (fig. 32). Projection of aII cylindrical, apically rounded. Length ratio AII: AIII+AIV 1:1.70–1.73. Body length ♂ 2.54–2.55 mm ........... ..............................................................................................P. asvadurovi sp. n. 4 At least partly testaceous ...................................................................................5 – Overall darker, or only pronotum testaceous .....................................................6 5 Completely testaceous. Pronotal calluses hemispherical. Paramere straight and flat. Body length ♂ 3.5 mm..............................................P. vaulogeri vaulogeri – Only forewings and femora testaceous. Pronotal calluses triangular, with two concave sides and one side strongly arcuate. Body length ♂ 3.5 mm ................. ......................................................................................P. vaulogeri angusticeps Fig. 42. Biotope and habitat of Putoniola asvadurovi sp. n., ‘avrona nature reserve: (a) vegetation in the hyper-arid area shrubland and trees with dominant Vachellia tortilis raddiana (savi) Kyal. & Boatwr. and Vachellia tortilis (forssk.) galasso & Banfi; (B) collecting event under Vachellia tortilis by artem asvadurov in 2017. A B 64 Israel Journal of entomology, Vol. 54, 2025 6 Forewings and femora black, entire tibiae, tarsi and last segment of antennae very dark brown, shiny. Pronotum black. Body length ♂ 3.0 mm ....................... ......................................................................................P. vaulogeri equatorialis – Forewings brownish grey, femora, apex of tibiae, and tarsus dark brown. Pronotum testaceous. Body length ♂ 3.5 mm ...............................................P. atakorensis subfamily Phymatinae laporte de Castelnau, 1832 Type genus: Phymata latreille, 1802. the subfamily Phymatinae, commonly known as ambush bugs, is distributed globally, across both tropical and temperate regions, apart from australia and new Zealand. all members of the subfamily are predators, preying on the large variety of insects (Hernandez et al. 2019). Phymatinae are characterised by the high, partially hiding the rostrum laterally bucculae; the antennae shorter than the head and the pronotum together; the head and the prothorax with an inferior antennal groove where the antenna rests in the lateral view; the raptorial forelegs; the 4-segmented antennae; the absence of the abdominal trichome. Body length ♂ 6.3–9.5 mm, ♀ 9.6–10.7 mm. Phymatinae are divided into four tribes. Carcinocorini Handlirsch, 1897 includes ambush bugs with a distinctive crab-like appearance, enlarged forelegs and a broad, flattened body, distributed in the oriental region. macrocephalini Handlirsch, 1897 is characterized by elongated heads and large scutellum, which covers most of the abdomen, with cosmopolitan distribution, except to the West Palaearctic. the nearly cosmopolitan Phymatini laporte de Castelnau, 1832 is the largest and most diverse tribe of ambush bugs, encompassing a wide range of species with varying morphologies. themonocorini Carayon, usinger & Wygodzinsky, 1958 is a relatively small tribe containing one extant afrotropical genus (with a few species), and one fossil neotropical monotypic genus (masonick et al. 2017). Hereby the subfamily Phymatinae is recorded from Israel and possibly for the levant for the first time. Note. maldonado (1990) considered the Phymatidae as a distinct family and excluded them from his systematic Catalogue of the reduviidae of the World. tribe Phymatini laporte de Castelnau 1832 the Phymatini tribe is the largest and most diverse of the five tribes of Phymatinae, comprising 115 species and 42 subspecies in five genera: Anthylla stål, 1876, Kelainocoris Kormilev, 1963 and Paraphymata Kormilev, 1962; Neoanthylla Kormilev, 1951; Phymata latreille, 1802. Diagnosis: the members of this tribe display a considerable range of body shapes and sizes, from small and slender to large and robust. the head above the eye and propleura just ventral to the lateral margin with a distinct longitudinal antennal groove for reception of antennae at rest is a unique character among reduviidae noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 65 (Handlirsch 1897; froeschner & Kormilev 1989; forero 2004; gil-santana et al. 2015). Distribution: native to tropical and subtropical regions, Phymatini show remar kable adaptability, colonizing temperate zones in the nearctic, Palaearctic and orien tal regions (Kormilev 1962; froeschner & Kormilev 1989; Cui et al. 2003). genus Phymata latreille, 1802 Acanthia fabricius, 1775: 696. type species: Acanthia crassipes fabricius, 1775, by monotypy. Phymata latreille, 1802: 247; fieber 1961: 109; Puton 1879: 126 (synopsis, description, key); Handlirsch 1897: 127–230; Bianchi 1899: 222 (systematic review); gulde 1940: 17 (review, europe); Kiritshenko 1951: 237 (review, ussr); Dispons & stichel 1959: 184 (review, europe); Kormilev 1962: 308 (key); Kerzhner & Jaczewski 1964: 1024 (review, key, european ussr); Wagner 1967: 33 (review, europe); Cmoluchowa 1978: 38 (key, Poland); Putshkov 1987: 215 (review, key, biology, ukraine); Putshkov & moulet 2009 (key, france); schuh et al. 2009 (phylogeny); masonick & Weirauch 2020a (taxonomy, key, molecular data, nearctic); masonick & Weirauch 2020b (phylogenetic analysis). Syrtis fabricius, 1803: 121 (syn. latreille, 1804: 244). type species: Cimex erosus linnaeus, 1758, by subsequent designation by Westwood (1842: 19) (suriname). Discomerus laporte de Castelnau 1832: 14 (syn. Burmeister, 1835: 251). type species: Cimex erosus linnaeus, 1758, by monotypy (suriname). Figs 43, 44. Phymata (P.) monstrosa (fabricius, 1794), male smnHtau In.370457, dorsal (43) and lateral (44) views. (Photo by oz rittner) 43 44 66 Israel Journal of entomology, Vol. 54, 2025 Type species: Acanthia crassipes fabricius, 1775. the genus Phymata latreille, 1802 comprises approximately 110 species of ambush bugs that are primarily distributed in the new World. the genus is further subdivided into four subgenera: Euryphymata Kormilev, 1962; Neophymata Kormilev, 1962; Phymata latreille, 1802; and Phymatispa Kor milev, 1951. the subgeneric placement of three species remains uncertain: Phymata albimana Zayas, 1966, Phymata grilloi Zayas, 1988 and Phymata isabelae rivero-aragón & grilloravelo, 2005. Diagnosis: the genus Phymata is characterized by the following characters: the pronotal disc with two submedian keels. the lateral margins of pronotum with irregular and sharp teeth. the lateral edge of connexival segments is dentate. the fore femur is subtriangular, swollen, without the fossula spongiosa, middle and hind tibiae with upper side carinate laterally and sulcate medially. sexual dimorphism is present (froeschner & Kormilev 1989; Putshkov & moulet 2009). Distribution: of some 110 described species of the genus Phymata, the majority occur in the neotropical region. only five species have been recorded from the Palaearctic region, all belonging to the subgenus Phymata latreille, 1802 (masonick & Weirauch 2020). Hereby the genus Phymata is recorded from Israel and possibly for the levant for the first time. subgenus Phymata latreille, 1802 Phymata (Phymata) Kormilev, 1962: 309. Diagnosis: the subgenus Phymata is characterized by the posterior border of the Figs 45–47. Phymata (P.) monstrosa (fabricius, 1794), female smnHtau In.370461, dorsal (45), ventral (46) and lateral (47) views. (Photo by oz rittner) 45 46 47 noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 67 abdomen without notch, the parameres with one point only, the straight struts of the phallus and the slender basal plates and struts of phallus (froeschner & Kormilev 1989). following species of P. (Phymata) are recorded in the Palaearctic region: P. (P.) chinensis Drake, 1947 (Putshkov & Putshkov 1996); P. (P.) crassipes (fabricius, 1775) (Putshkov & Putshkov 1996; tanco 2022; ghahari et al. 2024), P. (P.) griseipennis Horváth, 1907 (Putshkov & Putshkov 1996; Carapezza 1997), P. (P.) monstrosa (fabricius, 1794) (Putshkov & Putshkov 1996; Putshkov & moulet 2009; tanco 2022); P. (P.) subinermis Horváth, 1907 (Kormilev 1962; Putshkov & Putshkov 1996). Figs 48–51. Phymata (P.) monstrosa (fabricius, 1794), female smnHtau In.370461: (48, 49) head and pronotum, anterior (48) and dorsal (49) views; (50) head and thoracic pleura, left lateral view; (51) head, pronotum and scutellum, dorsolateral view. 49 50 48 51 68 Israel Journal of entomology, Vol. 54, 2025 Fig. 52. Phymata (P.) monstrosa (fabricius, 1794), dorsal view showing pterothorax and scutellum with carina, male smnHtau In.241329. abbreviations: Dag1 and Dag2 – dorsal abdominal scent glands on abdominal tergites 5 and 6, cs2–cs7 – connexivum segments, mls – median longitudinal sulcus, scm – scutum. Putshkov and moulet (2009) were uncertain whether the names monstrosa and griseipennis refered to two different species or to different forms of the same species, considering that griseipennis had been originally described as a colour form of monstrosa. carina of scutellum mls damage noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 69 Phymata (Phymata) monstrosa (fabricius, 1794) figs 43–70, table 2 Acanthia monstrosa fabricius, 1794: 74. Syrtis monstrosa fabricius 1803: 122; Hahn 1835: 57; Burmeister 1835: 251; Herrich-schäffer 1835: 57; germar 1836: 21; rambur 1842: 168. Phymata coarctata flor, 1860: 404 (syn. Puton, 1879: 127, with Phymata crassipes; Putshkov, 1987: 219, with Phymata monstrosa); Putshkov & Putshkov 1996: 185 (Palaearctic); Putshkov & moulet 2009: 586–588 (key, description, ecology, france). Phymata monstrosa (fabricius, 1794): Dohrn 1859: 41 (distribution); fieber 1961: 110; Handlirsch 1897: 150 (distribution); oshanin 1908: 502 (catalogue, distribution, Palaearctic); Pic 1915: 170 (list, algeria, tunisia); Dispons 1960: 305 (key); Kormilev 1962: 372–373 (description), 1966: 277 (description); sienkiewicz 1964: (list, tunisia); servadei 1967: 251 (catalogue, Italy); Josifov 1968: 29 (key). Redescription: male and female macropterous (figs 43–47); body pear-shaped expanded posteriorly; edge of abdomen expands beyond margins of wings. Colour (figs 43–59) matte, variable with some elements of camouflage, dark brown to black in both sexes, some specimens lighter yellow-brown. Pronotal anterior lobe darkened with variable pale markings. legs: males with darkened femora, females paler or with darkened femora. Connexivum II–IV segments with small pale spots; segment V darkened laterally and pale basally; segments VI–VII pale with small darkened spots. Corium darkened. forewing membrane hyaline. Body granulated, covered with tubercles and with setae of types differing in texture and length: hard or soft, short or longer, but not longer than tubercles, some setae recurved. Head: frontal process V-shaped, well developed; ocellar process and preocellar process long; eyes rounded in lateral view; ocelli located on posterior sides of head, Figs 53–55. Wing venation of Phymata (P.) monstrosa (fabricius, 1794): (53, 54) forewings, male smnHtau In.241329 (53) and female, smnHtau In.370391 (54), dorsal (a) and ventral (B) views; (55) hindwing, male smnHtau In.370457, ventral view. 53A 54A 55 55 B B 76 Israel Journal of entomology, Vol. 54, 2025 Fig. 68. Distribution of (a) Putoniola asvadurovi sp. n. and (B) Phymata (P.) monstrosa (fabricius, 1794) in Israel. green dots indicate collecting localities. A B noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 77 monstrosa specimen (pers. comm. Dr Herbert Zettel, nHmW). the specimen now housed at the nHmW bears three labels reflecting Handlirsch’s direct involvement in its identification: “syria Coll. signoret,” “monstrosa det. signoret,” and “monstrosa det. Handlirsch.” Despite Handlirsch’s initial reservation, subsequent citations by oshanin (1908: 502, 1912: 48) and Kormilev (1966: 314, 373) uncritically repeated the syrian record. However, later research challenged this claim: Putshkov and Putshkov (1996) treated the syrian provenance with a question mark, and Putshkov and moulet (2009) implicitly dismissed the record by excluding this genus and its species from their treatment of the syrian fauna. this exclusion was further supported by ghahari et al. (2024). fairmaire (1889: 505) stated the following in his necrology for signoret: “Des voyages en Italie, en grèce, en turquie et en asie mineure lui avaient permis de réunir une collection intéressante et nombreuse.” the historical term “asia minor” could encompass regions that are currently part of syria. notwithstanding, it is impossible to establish exactly in which country the specimen was collected because in the late XIX century (and during signoret’s lifetime) ‘syria’ included the present territories of Jordan, lebanon, Israel, syria and turkey’s province of Hatay. In this study, we definitively establish the presence of P. (P.) monstrosa in Israel. the finding of P. (P.) monstrosa in Israel implicitly confirms the middle-eastern origin of the signoret’s specimen, whereas the distribution of the species was previously regarded as strictly West-mediterranean (Putshkov & Putshkov 1996; Putshkov & moulet 2009; ghahari et al. 2024). Habitat and plants associations: the specimens of Phymata monstrosa were collected in Israel in samaria (shomeron), the Judean Desert and the Central negev. these regions are characterized by mountainous topography and arid or semi-arid conditions. most of the specimens were found in the Judean Desert, characterized by an extremely arid climate, and higher and therfore somewhat more humid and less hot mountainous ridge of the Central negev. the specimen from samaria, that is mainly characterized by more temperate mediterranean climate, was found in a more arid part, on the eastern slopes of the samarian mountains, at an edge of the extremely arid Jordan Valley. Phymata monstrosa is known along its distributional range from various alti tudes, including significant heights, e.g. lindberg documented collecting of speci mens at an altitude of 2000 m in the moroccan atlas (lindberg 1932). However, in Israel P. monstrosa is found at a wide range of altitudes from 110–1000 m. at least some Phymata species, as obligatory phytobionts, are strongly associated with bushy habitats; particularly P. monstrosa is recorded on Ficus sp. (moraceae), Daphne sp. (thymelaeaceae), Cneorum sp. (rutaceae), Quercus sp. (fagaceae) (Perrier 1937) and Retama sphaerocarpa (fabaceae) (ribes et al. 1997). In Israel, se veral specimens of P. monstrosa have been found on Ephedra sp. (ephedraceae) and Hirschfeldia incana (Brassicaceae), which are new plant associations for this species. 78 Israel Journal of entomology, Vol. 54, 2025 Contributions to the phylogeny of Holoptilinae and Phymatinae the “Phymatine complex” comprises four subfamilies: Centrocnemidinae, Holoptilinae, Phymatinae and elasmodeminae. the subfamily Phymatinae is the most extensively studied phylogenetically, while Centrocnemidinae and Holoptilinae have partial molecular data, and elasmodeminae has as yet no data. While the first three are considered monophyletic and are believed to be sister taxa within the extant reduviidae, the phylogenetic relationships of elasmodeminae remain to be determined. the representation of the subfamily Holoptilinae in molecular records is scarce, with only 27 records pertaining to five genera. the tribe aradellini, which includes Aradelloides malipatil, 1983 and Aradellus Westwood, 1874, is the worst phylogenetically represented. additional sequencing is crucial to fully clarify its phylogeny. the tribe Dasycnemini is represented by the sequences of two species: Locoptiris Villiers, 1943 (sequence noted as Lisarda inornata (Walker, 1873) reduviidae: salyavatinae in nCBI), and Neolocoptiris villiersi Wygodzinsky & usinger, 1963, while Dasycnemus Bergroth, 1898 still awaits sequencing. the genus Putoniola is discussed in the present study. the largest tribe Holoptilini, with its eight genera, is, nevertheless poorly re presented in the genBank. representatives of only three genera—Ptilocerus gray, 1832, Ptilocnemus Westwood, 1840 and Thysanopus Bergroth, 1893—have been se quenced. this leaves five genera—Holoptiloides miller, 1956, Holoptilus le Peletier & audinet-serville, 1825, Orthocnemis Westwood, 1845, Ptilocoris montan don, 1907, Smiliopus Bergroth, 1909—out of the analysis. Interestingly, Orthocnemis possesses unique morphological characteristics: the absence of trichomes (replaced by a noticeable swelling and a fringe of setae). this contrasts with most other Holoptilini genera, which have clearly visible trichomes. these distinctions suggest that Orthocnemis might warrant placement in a separate tribe, a hypothesis that presents a promising avenue for future research. the phylogenetic analysis presented here provides support to the relationships within the Phymata, previously described by masonick et al. (2017), who performed an extensive phylogenetic study of Phymatinae and revealed that the genus Phymata originated in the neotropical region and dispersed to the nearctic and Palaearctic regions during the last 30 million years. out of some 110 species of Phymata, only five are known from the Palaearctic, as noted above. Phymata crassipes, the only Palaearctic species sampled, diverged from the new World taxa roughly around the oligocene–miocene boundary. the present study provides molecular data for a second Palaearctic species, P.(P.) monstrosa (appendix 1, p. 89). In the analysis the two Palaearctic species cluster together, as expected in relation to the study by masonick et al. (2017). In the CoI analysis (fig. 69) they form a cluster separated from all other neotropical species, while in the 16s (fig. 70) analysis they are nested within other Phymata. this may be explained by the fact that the evolution rate of CoI is double that of 16s in Hemiptera (li et al. 2012). noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 79 Fig. 69. reconstructed phylogenetic CoI-based tree for the Phymatinae (reduviidae). 80 Israel Journal of entomology, Vol. 54, 2025 Fig. 70. reconstructed phylogenetic 16s-based tree for the Holoptilinae and Phymatinae (reduviidae). noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 81 regarding the Holoptilinae, the phylogenetic analysis (fig. 70) comprises 16s sequences of Dasycnemini and Holoptilini tribes. this analysis, however, is too con servative to resolve the relationships between the tribes. Clustering of genera did not result in significant bootstrap support. the present study adds a new species to the genus Putoniola, accompanied by much-sought-after molecular data that are crucial for resolving the tribal re la tionships within the Holoptilinae. In addition, the sequence data of the Palaearctic Phymata (P.) monstrosa support a previous hypothesis on the divergence and dispersal of the genus (masonick et al. 2017). the new findings emphasize the importance of combined molecular and morphological studies to establish the phylogeny of the “Phymatine complex”. ACKNOWLEDGEMENTS the authors are especially grateful to Dr Pierre moulet (museum requien, avignon, france), Dr Pavel V. Putshkov (schmalgausen Institute of Zoology, Kyiv, ukraine), Dr David g. furth (smithsonian Institution, Washington, usa), Dr Dimitri forero (Pontifical Xavierian university, Department of Biology, Bogotá, Colombia), Dr Petr Kment (Department of entomology, national museum, Praha, Czech republic), and Dr Zoya efremova (smnHtau) for their assistance in providing PDf reprints of significant publications. We appreciate the generous assistance with getting images of specimens from the curators of the following institutions: Dr ruth salas and Dr Jessica l. Ware (amnH), Dr anna Ágnes somogyi and Dr Zoltán Vas (HnHm), Dr Villu soon (tuZ), and Dr Herbert Zettel and Dr alice laciny (nHmW). mr oz rittner (smnHtau) is thanked for expertly taking pictures and processing them for figures 43–47, and Dr Zahava Barkay (Center for nanoscience and nanotechnology, tel aviv university, Israel) is thanked for her kind assistance with sem. the authors are grateful to Dr gideon Pisanty (smnHtau) and Dr tal levanony (the yehuda naftali Botanic garden, tel aviv university, Israel) for their explanation of the distribution of Vachellia trees in Israel. 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Buchdruckerei der Kaiserlichen akademie der wissenschaften, st Petersburg. 586 pp. https://doi.org/10.5962/bhl.title.12423 92 Israel Journal of entomology, Vol. 54, 2025 Species Subfamily Voucher 16S Locality 16S COI References Phymata granulosa Phymatinae r_CW_1007 mexico: Chiapas Ky510662 mn136979 masonick et al. 2017; masonick & Weirauch 2020a Phymata granulosa Phymatinae r_CW_5393 mexico mn148969 mn136981 masonick & Weirauch 2020a Phymata inconspicua Phymatinae r_CW_4037 argentina: la rioja Ky510659 mn136982 masonick et al. 2017; masonick & Weirauch 2020a Phymata lindigiana Phymatinae r_CW_2278 Columbia: Cundinamarca Ky510657 masonick et al. 2017 Phymata luteomarginata Phymatinae r_CW_5099 usa: California mn136984 masonick & Weirauch 2020a Phymata luxa Phymatinae r_CW_3083 usa: texas Ky510680 mn136985 masonick et al. 2017; masonick & Weirauch 2020a Phymata minuta Phymatinae r_CW_3339 Honduras Ky510654 mn136986 masonick et al. 2017; masonick & Weirauch 2020a Phymata monstrosa Phymatinae SMNHTAU In.370391 Israel: 'Arad PV188576 PV179415 Present study Phymata pacifica hainesi Phymatinae r_CW_910 mexico: Baja California Ky510678 mn136990 masonick et al. 2017; masonick & Weirauch 2020a Phymata pacifica hainesi Phymatinae r_CW_907 mexico: Baja California mn136989 masonick & Weirauch 2020a Phymata pacifica pacifica Phymatinae r_CW_4160 usa: California KX512314 mn136998 masonick et al. 2017; masonick & Weirauch 2020a Phymata parva Phymatinae r_CW_5401 mexico mn148984 mn137003 masonick & Weirauch 2020a Phymata parva Phymatinae r_CW_5414 mexico mn148986 mn137005 masonick & Weirauch 2020a Phymata pennsylvanica Phymatinae Kt231816 Zhang et al. 2016 Phymata pennsylvanica Phymatinae r_CW_4676 Canada: on: niagara Ky510676 mn137007 masonick et al. 2017; masonick & Weirauch 2020a Appendix 1. list of sequences used for the reconstruction of phylogenetic trees based on 16s and CoI mitochondrial sequences. noVoselsKa & felDsteIn-farKasH: PHymatInae anD HoloPtIlInae of Israel 93 Species Subfamily Voucher 16S Locality 16S COI References Phymata rossi Phymatinae r_CW_4704 usa: arizona Ky510663 mn137009 masonick et al. 2017; masonick & Weirauch 2020a Phymata saileri Phymatinae r_CW_4705 usa: arizona Ky510666 mn137011 masonick et al. 2017; masonick & Weirauch 2020a Phymata salicis Phymatinae r_CW_5539 usa: arizona mn148990 masonick & Weirauch 2020a Phymata salicis Phymatinae r_CW_5540 usa: nevada mn148991 masonick & Weirauch 2020a Phymata severini Phymatinae r_CW_5476 mexico mn148992 mn137012 masonick & Weirauch 2020a Phymatinae sp. Phymatinae uCr<usaCa>:ent:00004817 JQ888471 Zhang & Weirauch, unpubl. Themonocoris endroedyi Phymatinae r_CW_827 or911766 or909599 masonick et al. 2025 Themonocoris sp. Phymatinae r_CW_824 south africa gu188458 Weirauch et al. 2011 Locoptiris taiwanensis Holoptilinae r_CW_5602 thailand or911677 or909546 masonick et al. 2025 Neolocoptiris villiersi Holoptilinae r_CW_1986 french guiana: Pararé Ky510647 masonick et al. 2017 Ptilocerus sp. Holoptilinae uCr_ent 00001974 gu188453 Weirauch et al. 2011 Ptilocerus sp. Holoptilinae r_CW_689 laos: Vientiane Prov. gu188454 Weirauch et al. 2011 Ptilocnemus femoralis Holoptilinae r_CW_220 australia: sa fJ230431 Weirauch & munro 2009 Ptilocnemus lemur Holoptilinae mW540749 mW540749 ye et al. 2021 Putoniola asvadurovi Holoptilinae SMNHTAU In.454881 Israel: 'Avrona Nature Reserve PV188577 Present study Appendix 1. list of sequences used for the reconstruction of phylogenetic trees based on 16s and CoI mitochondrial sequences. 94 Israel Journal of entomology, Vol. 54, 2025 Species Subfamily Voucher 16S Locality 16S COI References Homalocoris erythrogaster Hammacerinae r_CW_5541 or909540 masonick et al. 2025 Microtomus cinctipes Hammacerinae CW 141 nicaragua fJ230411 or909552 Weirauch & munro 2009; masonick et al. 2025 Microtomus sp. Hammacerinae CW 030 french guiana fJ230395 Weirauch & munro 2009 Microtomus sp. Hammacerinae yB-BCI148659 Panama mn621014 Basset, unpubl. Neocentrocnemis sp. Centrocnemidinae uCr_ ent_00118998 or911696 or909558 masonick et al. 2025 Neocentrocnemis stali Centrocnemidinae China KC887530 KC887530 gao & Cai, unpubl. Neocentrocnemis stali Centrocnemidinae ZglC164-12 KJ629592 Zhao & Cai, unpubl. Appendix 1. list of sequences used for the reconstruction of phylogenetic trees based on 16s and CoI mitochondrial sequences.