Saltbush-associated Asphondylia species (Diptera: Cecidomyiidae) in the Mediterranean Basin and their chalcidoid parasitoids (Hymenoptera: Chalcidoidea)

Numerous species of gall midges (Diptera: Cecidomyiidae) have been recorded from saltbush (Chenopodiaceae: Atriplex) around the world but only 11 of them belong to the large cecidomyiid genus Asphondylia. Of these, two species were de-scribed in the late 19th century from complex bud galls on Atriplex halimus in the Mediterranean Basin. In the present study Asphondylia punica is redescribed, A. conglomerata is synonymized with it, and Asphondylia scopuli is described from Atriplex lanfrancoi, an endemic plant to the Maltese Islands. Descriptions are accompanied by information about the galls and life history of the gall midges, and a review of the parasitic Hymenoptera associated with A. scopuli is provided. Four species of parasitoids were found and attributed to the families Eurytomidae, Pteromalidae, Eupelmidae and Eulophidae, of which the pteromalid Mesopolobus melitensis is described as new.peer-reviewe

Hidden Diversity—A New Speciose Gall Midge Genus (Diptera: Cecidomyiidae) Associated with Succulent Aizoaceae in South Africa

Aizoaceae (Caryophyllales) constitute one of the major floral components of the unique Greater Cape Floristic Region (GCFR), with more than 1700 species and 70% endemism. Within succulent Aizoaceae, the subfamily Ruschioideae is the most speciose and rapidly diversifying clade, offering potential niches for the diversification of specialized herbivorous insects. Nevertheless, insect diversity on these plants has not been studied to date, and knowledge of gall-inducing insects in the Afrotropics is generally scarce. Our recent observations indicate that succulent Aizoaceae in the GCFR support a rich and largely unstudied community of gall midges (Diptera: Cecidomyiidae). Here, we provide a first report of their diversity with a description of a new genus, Ruschiola Dorchin, and ten new species, based on morphological and molecular analyses of material collected during a three-year targeted survey across major GCFR vegetation types. A high degree of morphological uniformity in Ruschiola suggests recent diversification and necessitated the use of molecular data and laboratory rearing from host plants to verify species boundaries and host ranges

Stefaniola Kieffer 1913

Genus Stefaniola Kieffer, 1913 Stefaniola is the largest genus of Lasiopterini, with 108 described species (Gagné 2004) and many undescribed species, all of which develop in plants of the Chenopodiaceae in Central Asia and the Mediterranean basin. The usually small and stout adults superficially resemble those of the related genus Careopalpis, from which they differ in the morphology of the ovipositor. The shape of the ovipositor in Stefaniola is highly diverse, but the aculeus is always curved dorsally rather than ventrally, often conspicuously long, and the apical lamella is variously shaped, sometimes with additional protruding lobes. The shape of the lateral plate and its setae are likewise diverse among species. Apical antennal flagellomeres are sometimes fused, and number of flagellomeres may vary within the same species and individual. Palpus is one-segmented. Larvae lack a spatula. Many species are known only from their galls and larvae (Möhn 1971).Published as part of Dorchin, Netta & Freidberg, Amnon, 2008, The Chenopodiaceae-feeding gall midges (Diptera: Cecidomyiidae) of the Na'aman salt marsh, Israel, pp. 1-22 in Zootaxa 1937 on page 18, DOI: 10.5281/zenodo.18498

Schizomyia botellus Dorchin and Freidberg, new species

Schizomyia botellus Dorchin and Freidberg, new species (Figs. 30 –36, 39–40, 44, 46, 48, 54 –55, 62– 64) Adult: Head (Fig. 30): Eye facets circular to hexagonal. Gap between eyes on vertex 0-1 facet wide. Palpus 4 -segmented, segments successively longer; first segment round, about as long as wide; consecutive segments much longer than wide. Labellum slightly tapered, with few long setae. Antenna: 12 flagellomeres in both sexes, covered by microtrichia. Female flagellomeres 1–9 cylindrical; flagellomeres 1–2 partially fused; flagellomeres 5 and on successively shorter; flagellomere 10 as long as wide; flagellomere 11 rounded, slightly wider than long; flagellomere 12 rudimentary, partially fused with preceding; flagellomeres 1–10 each with two whorls of anastomosing circumfila, strong setae posterior to basal whorl and between whorls (Fig. 31); segments 11–12 with only one whorl of circumfila. Male flagellomeres cylindrical; flagellomeres 1–2 longer than succeeding, flagellomeres 3–11 about the same length; flagellomere 12 slightly shorter than preceding, rounded apically; all flagellomeres covered by spacious interconnected whorls of winding circumfila and bear relatively few strong setae mostly at flagellomere bases (Fig. 32). Thorax: General color gray, covered by sparse, white hair. Wing: hyaline; length 2.26–2.70 mm in female (n= 15), 2.04–2.52 mm in male (n= 8); R 1 joins C at about mid-length of wing, R 5 joins C posterior to wing apex, M present as very weak fold, Cu forked. Haltere stalk orange, knob white. Legs: general color orange, covered by short black hairs dorsally, white hairs ventrally; coxae with long, sparse, white hairs; tarsal claws evenly curved, untoothed; empodia longer than bend in claws (Fig. 33). Female abdomen (Fig. 34): General color brownish-orange; sclerites black; entire abdomen covered by sparse white hairs. Tergites 1–5 rectangular, less sclerotized anteriorly than elsewhere, with posterior and median rows of strong setae, and a pair of tiny sensory setae anteriorly; tergite 6 similar but more setose; tergite 7 longer than preceding, distal half covered by strong setae, proximal half with several shorter setae and pair of tiny sensory setae; tergite 8 shorter than preceding, with short, tapered projection posterioly, bare except for pair of anterior sensory setae. Sternites 2–6 each comprising two sclerites, anterior rectangular, with several strong setae, posterior narrow band-like, with row of strong setae; anterior sensory setae not visible. Sternite 7 at least three times as long as preceding, posterior part shovel-like, strongly sclerotized; posterior two thirds covered by long setae. Base of ovipositor posterior to sternite 7 forms setose invagination. Ovipositor curled inside abdomen, distal part needle-like, slightly sclerotized ventrally, with several short setae on and towards apex (Fig. 35). Male abdomen: General color and hairiness as in female. Tergite 1 with narrow band-like sclerotized area medially, posterior row of long setae, and few long setae elsewhere; tergites 2–7 rectangular, with posterior row of long setae, several long setae mesally, and evenly scattered scales; tergite 8 hardly differentiated from surrounding membrane, with small amorphous sclerotized patch and no long setae. All tergites with pair of small, anterior sensory setae. Sternites 2–6 rectangular, with posterior row of long setae separated from anterior area of sternite by weakly sclerotized band; anterior area of sternite with several strong setae. Sternite 7 entire, without weakly sclerotized band and with more numerous, evenly scattered strong setae. Sternite 8 hardly differentiated from surrounding membrane, with small but strongly setose sclerotized patch. Anterior sensory setae not vsible. Terminalia (Fig. 36): Gonocoxal apodeme comprises two strongly sclerotized, wide lobes connected by concave bridge. Gonocoxite angular proximally, with long apical lobe extending far beyond gonostylus, ridged medially; most surface proximal to insertion of gonostylus covered by short, strong setae; apical lobe rounded apically, ridged and densely covered by long, strong setae especially around apex. Gonostylus stout, rectangular except for slight apical projection pointed anteriorly, evenly covered by microtrichia on both dorsal and ventral surfaces; dorsally with several strong setae mostly on distal half, ventrally with row of strong setae along apical denticles and group of strong setae along posterior margin; unfused denticles extend almost to tip of projection. Aedeagus tapered, notably longer than hypoproct and cerci. Hypoproct narrow rectangular, slightly widened just before notched apex; setulose, with few longer setae apically. Cerci separated by deep triangular notch, considerably wider than hypoproct, setulose throughout, jagged and strongly setose around apex. Larva (third instar) (Fig. 39): Orange. length 2.6–3.2mm (n= 6). Integument covered entirely by pointed spicules. Antennae about twice as long as wide. Cephalic apodemes about as long as head capsule. Spatula (Fig. 40) with wide shaft and two triangular anterior lobes widely separated. On each side of spatula one asetose sternal papilla and one group of three lateral papillae, two with short setae, one asetose. Pleural and dorsal papillae with strong setae. Thoracic segments 2–3 and all abdominal segments each with two small mesoventral bulges. Terminal segment with two tiny, barely perceptible papillae. Pupa (Figs. 44, 46, 48): Antennal bases developed into prominent horns, each divided into two tips close to apex; arched along dorsal contour. Cephalic seta about as long as antennal horn, originating from elevated base. Frons on each side with pair of papillae at mid-posterior area, one with long seta, the other asetose and group of three tiny lateral papillae, two setose, one asetose. Mid facial area with conspicuous blunt bulge on each side. Prothoracic spiracle long and blunt; trachea ends at apex. Dorsal and lateral parts of abdominal segments (Fig. 48) each evenly covered by short, pointed spinules except for short crinkled posterior area and anterior area of striking, complex spines. Holotype: 3, Israel, Mishor Paran, 17.iii. 1995, N. Dorchin, reared from Deverra triradiata gall. Paratypes: 2 Ƥ, 23, same data as holotype, TAUI: 7 Ƥ, 9 exuviae (on two microscope slides), Nahal Lavan 24.iii. 2000, N. Dorchin (4 Ƥ, exuviae TAUI, 3 Ƥ ZFMK); 1 Ƥ, 23, 6 larvae (on one microscope slide), Israel, Nahal Yamin, 8.iii. 2010, A. Freidberg, TAUI; 13, 1Ƥ, Sede Boqer, 18.iii. 2010, N. Dorchin and E. Morgulis, TAUI. All material from Deverra triradiata. Distribution. Tunisia (Matmata), Egypt (Sinai, Wadi El Raha), Israel (Negev desert: Nahal Lavan, Mishor Paran, Nahal Paran, Nahal Yamin, Sede Boqer, Zomet Mash’abim, Nahal Raham). Etymology. The species name is Latin for “small sausage”, a noun in apposition, with reference to the conspicuous jointed galls. Biology. Galls of this species were recorded from Deverra scoparia (in Tunisia) and Deverra triradiata (in Egypt and Israel). The species is univoltine, with galls developing from late winter (February) to spring (April). Pupation takes place inside the gall and adults emerge from early March to mid April. Other life-history aspects are unknown, but eggs are presumably laid on the plant and first-instar larvae probably spend the summer in diapause until they become active the next winter. The galls (Figs. 54 –55, 62– 64) develop from axillary buds into elongate, jointed structures, 2–8 cm long, depending on the number and size of joints. They are light green, often with fine longitudinal striation, and are very conspicuous on the background of the virtually leafless stems. After adult emergence, the gall turns yellow and then gray when completely dry. Such old, dry galls remain on the plants for more than a year so that old and new galls are often found in spring side by side (Fig. 55). The gall is hollow (Fig. 64 b) and the spaces within joints are often connected to each other. Larvae feed gregariously inside the gall, several in each joint. A large gall may contain more than 20 larvae. Before pupation, the larvae prepare holes in the gall’s walls, which remain sealed by a thin layer of plant tissue through which the pupae will break out. The larvae then spin thin, transparent sheaths of silk that divide the space inside the gall into separate compartments in which they pupate with or without spinning a cocoon. The gall does not contain visible mycelia. Larvae can be heavily attacked by external hymenopteran parasitoids, each of which feeds on several larvae and may kill all the larvae in a large gall. Endoparasitoids were also observed. Remarks. Frauenfeld (1859) found the galls of this species while on the Novara expedition, during which he also collected and described the closely related Schizomyia buboniae. He stated that he was unable to determine the identity of the host plant or rear the insects because the plant was too young, but his accurate description of fully developed galls and the mention of emergence holes suggest that the galls were already empty. This species has therefore been known so far only from its distinctive galls, which were beautifully illustrated by Frauenfeld (1859) (Fig. 62) and later by Houard (1912) (Figs. 63-64). The latter reported Deverra tortuosa as the host of Frauenfeld’s galls, but it is clear from Frauenfeld’s original drawing (Fig. 62) that the host was in fact D. triradiata (the two plant species differ strikingly). Houard (1912) found galls of variable sizes in Matmata (Tunisia) and, like Frauenfeld, failed to rear the adults although he mentioned that the larvae pupated inside the galls. For some reason he found larvae only in smaller galls (Fig. 64), whereas larger galls (Fig. 63) were occupied by coccids, which no doubt invaded the galls once the original inhabitants left. Although galls of all sizes are very similar in structure and despite illustrating them in detail, Houard (1912) did not recognize that they were all caused by the same insect and erroneously attributed the large galls to the coccids. Schizomyia botellus and S. buboniae are the only Schizomyia species that are known to pupate inside their galls, which explains the well developed structures on the pupal head and dorsum, and possibly also the reduced number of terminal papillae in the larvae. Schizomyia botellus is larger than S. buboniae but adults of the two species are otherwise very similar morphologically. In contrast, the immature stages are easily distinguishable. The larval spatula in S. buboniae has smaller teeth that are not as widely and deeply separated as those of S. botellus, and the pupae of the two species differ strikingly in their head and abdominal structures; the pupa of S. botellus has two conspicuous facial protuberances that are absent in S. buboniae, significantly shorter and not as broadly separated antennal horns, and a dorsal field of complex spines on each abdominal segment that is absent in S. buboniae. The relation of these two species to other described species of Schizomyia is unknown.Published as part of Dorchin, Netta & Freidberg, Amnon, 2011, The gall midges (Diptera: Cecidomyiidae) of Apiaceae in Israel, pp. 28-48 in Zootaxa 3044 on pages 40-43, DOI: 10.5281/zenodo.20344

Houardiella Kieffer 1912

Genus Houardiella Kieffer, 1912 Houardiella comprises two described species in the tribe Asphondyliini (Gagné 2004). It is closely related to the genus Asphondylia, with its needle-like, piercing ovipositor that is accompanied by a pair of basal lobes at its base, and the progressively shorter flagellomeres in the female. It differs from Asphondylia in having onesegmented (as opposed to three-segmented) palpi, a single tooth rather than two teeth on the male gonostylus, and an apical, posterior extension of the gonocoxite. Immature stages resemble those of Asphondylia, with well developed cephalic horns and abdominal spines in pupae, and a massive spatula in larvae.Published as part of Dorchin, Netta & Freidberg, Amnon, 2008, The Chenopodiaceae-feeding gall midges (Diptera: Cecidomyiidae) of the Na'aman salt marsh, Israel, pp. 1-22 in Zootaxa 1937 on page 12, DOI: 10.5281/zenodo.18498

Stefaniola crispa Dorchin and Freidberg, new species

Stefaniola crispa Dorchin and Freidberg, new species Adult. – Head (Fig. 54): Eye facets circular, gap between eyes on vertex 0.5–1.5 times as wide as facet. Palpus 1 -segmeted, 1.0– 1.5 times as long as wide, rounded apically, with long fine setae. Labellum well developed, with several scales and long setae. Antenna (Fig. 55): similar in both sexes; scape wide conical, pedicel globular, flagellomeres: 10 in both sexes, barrel-shaped, about 1.5 times as long as wide, each with two whorls of connected circumfila, whorl of strong setae between two circumfila, and whorl of smaller setae proximal to proximal circumfilum, evenly setulose; apical flagellomere tapered; flagellomeres 1–2 sometimes partially fused. Thorax: Wing: length 1.3 –2.0 mm in females (n= 15), 1.5–1.8 mm in males (n= 7); transparent, veins other than Sc and R 5 barely visible, R 5 joining C at wing mid-length, M present as fold, Cu unforked. C, Sc and R 5 brownish, with sparse hairs. Legs (Fig. 56): covered by dark, fusiform scales; tarsal claws strong, evenly curved, with small, strongly curved teeth close to base of claw; empodia extend slightly beyond bend of claw, with long hairs; pulvilli well developed, about half as long as claw. Female abdomen (Fig. 57): General color brownish-orange. Covered by white scales; scale pattern on dorsum comprising three black spots on anterior part of each tergite; medial spot triangular, smaller than two lateral semi-spherical spots. Tergites 1–7 more or less rectangular, less sclerotized anteriorly than elsewhere, with posterior row of setae; tergite 7 wider than preceding tergites, with 2–3 posterior rows of setae, and evenly covered by scales; trichoid sensilla not detectable. Tergite 8 mostly undifferentiated from surrounding membrane, with only small sclerotized patch, without setae. Sternites 2–6 rectangular, less sclerotized in mid part, with posterior row of setae and several evenly scattered setae; sternite 7 more setulose than preceding sternites; sternite 8 completely undifferentiated from surrounding membrane. Area anterior and posterior to lateral group of setae on segment 8 with longitudinal wrinkles and grooves. Ovipositor (Fig. 58): Lateral group of setae comprising more than 100 slightly curved setae. Sclerotized rods widened toward lateral plate. Lateral plate sheathing entire base of cercus, bearing 25–31 long, slightly and evenly curved setae, more than ten times as long as wide. Aculeus very thick at base, narrowed abruptly at distal third, tip pointed anterodorsally; base of aculeus with 6 erect and extraordinarily long, hooked setae, curled posteriorly. Apical lamella globular, slightly longer than aculeus. Male abdomen (Fig. 59): General color and scale pattern as in female. Tergites 1–7 rectangular, with 1–2 posterior rows of setae, otherwise evenly covered by scales; trichoid sensilla not detectable. Tergite 8 mostly undifferentiated from surrounding membrane, sclerotized only medially, without strong setae, evenly covered by scales. Sternites 2–7 unsclerotized both anteriorly and posteriorly, with 1–2 posterior rows of setae and several additional setae at mid length. Sternite 8 less sclerotized but more setulose than preceding sternites, entirely covered by setae. Terminalia (Fig. 60): Gonocoxite massive, evenly covered by strong setae on sclerotized parts; mediobasal lobe prominent, setose. Gonostylus stout and robust, cylindrical, slightly curved, almost same width throughout length, covered by fine setulae on entire surface both dorsally and ventrally, with wide, blunt tooth. Cerci wide, almost completely fused, distally separated by small notch, evenly setulose. Hypoproct considerably narrower than fused cerci, with very shallow apical notch. Paramere wide and robust, evenly covered by strongly curved setae, with apical pair of setae on elevated bases. Aedeagus considerably longer than paramere, slender, cylindrical, blunt, hardly narrowed toward tip. Larva. – Unknown. Pupa. – Unknown. Holotype – Ψ, Israel, Akko, 20.VII. 1996, N. Dorchin, reared from Arthrocnemum macrostachyum stem. Paratypes – All material from Israel, Akko, reared by N. Dorchin from Arthrocnemum macrostachyum stems unless otherwise noted. 6 Ψ, 3 ɗ, same data as holotype; 5 Ψ, 1 ɗ, 26.VII. 1997; 1 Ψ, 29.VIII. 1998; 1 ɗ, 12.VIII. 2002, N. Dorchin and A. Freidberg; 1 Ψ, 3 ɗ, 20.VIII. 2002, N. Dorchin and A. Dorchin. Distribution. – Israel (Akko salt marsh). Etymology: The name crispa, an adjective, is Latin for “curly”, referring to the striking setae on the aculeus of the female ovipositor. Biology: The species develops in single chambered, inconspicuous stem infestation in Arthrocnemum macrostachyum. As in Houardiella gracilis, all adults were reared from material in which no infestations were observed, hence the immature stages are unknown. Remarks. – Based on ovipositor morphology, this new species belongs to a small group within the genus Stefaniola, whose very short aculeus differs greatly from the long and dorsally curved aculeus typical of most described Stefaniola species. This attribute is shared by S. defoliata Dorchin, S. rufa Dorchin, and S. siliqua Dorchin, all reared from Suaeda monoica (Dorchin 2001), and a yet undescribed species from other Suaeda species in Israel. However, S. crispa differs from these species in its longer ovipositor and in the strikingly long setae on the aculeus. The only other species in the subtribe Baldratiina, in which similar, but relatively shorter, curled setae are found are Izeniola bassiae and I. potanini. The new species is currently placed in Stefaniola, although a much-needed genetic analysis of the Baldratiina may reveal that the five Stefaniola species mentioned above form a distinct group that does not belong in this genus.Published as part of Dorchin, Netta & Freidberg, Amnon, 2008, The Chenopodiaceae-feeding gall midges (Diptera: Cecidomyiidae) of the Na'aman salt marsh, Israel, pp. 1-22 in Zootaxa 1937 on pages 18-20, DOI: 10.5281/zenodo.18498

Lasioptera umbelliferarum Kieffer 1909

Lasioptera umbelliferarum Kieffer 1909 (Figs. 3, 12–13, 20, 23, 26, 42 –43, 52– 53) Lasioptera umbelliferarum Kieffer 1909: 28 Neotype of Lasioptera umbelliferarum designated here – Ƥ, Israel, Giv’at Olga, 19.III. 1996, A. Freidberg, reared from Bilacunaria boissieri stem gall. Lasioptera umbelliferarum was originally recorded (Rübsaamen 1895) and subsequently described only from its gall (Kieffer 1909) from Seseli sp. from Russia (near the Caspian Sea) but the host plant could have been misidentified. The type is considered lost together with the majority of Kieffer’s collection. Möhn (1968) later described the larvae of this species in his revision of the Palaearctic species of Lasioptera based on material that he recovered from herbarium-preserved galls. He noted that he never found in these herbaria galls of L. umbelliferarum on Seseli but did find them on three species of the closely related genus Hippomarathrum. We found the galls in Israel on Bilacunaria, a closely related genus to Seseli and Hippomarathrum that is sometimes treated under the latter (Mabberley 2008). The recorded distribution of L. umbelliferarum fits the distribution of Bilacunaria and the published descriptions of the galls and larvae leave little doubt that the species we found in Israel is indeed L. umbelliferarum. Pupae and adults of this species are described here for the first time. The neotype is designated in order to fix the identification of the species and maintain the current usage of its name. It is deposited in the National Collection of Insects, Zoological Museum, Tel Aviv University, Israel (TAUI). Adult: Head: Frons, gena, scape, pedicel, and area along posterior margins of eyes densely covered by white scales and hairs. Eye facets circular. Gap between eyes on vertex 0–1 facet wide. Palpus 4 -segmented; palpiger usually well developed; segments 1-3 successively longer; first segment as long as or slightly longer than wide; fourth segment same length or shorter than third. Labella slightly tapered, with few long setae. Antenna: Dark brown. Scape and pedicel covered by white scales. Flagellomeres: 17–19 in female (n= 13), 13–16 in male (n= 10), number occasionally varies between antennae of same individual; flagellomeres except for apical as long as wide or slightly wider than long, especially in proximal half of antenna; first two flagellomeres partially fused, last often longer than preceding, consisting of two fused flagellomeres; each flagellomere except for apical with two whorls of circumfila connected by 1–2 longitudinal branches, row of short, curved setae proximal to proximal circumfilum, and several strong and very long setae between circumfila whorls; otherwise evenly covered by microtrichia (Fig. 3). Apical flagellomere often with 3–4 circumfila whorls. Circumfilar whorls on flagellomeres 1–2 closer to each other and often with more longitudinal connections than those on other flagellomeres. Thorax: Dark brown, covered laterally by white scales and hairs, dorsally with two longitudinal lines of long white hairs. Wing: hyaline; length 2.00– 2.53 mm in female (n= 12), 1.84–2.37 mm in male (n= 8); R 1 and R 5 join C at about 0.4 and 0.6 of wing length, respectively; M present, Cu unforked. R 1, R 5 and C along anterior wing margin densely covered by dark brown scales except for white spot at merging point of R 5 and C. Posterior wing margin with long, fine hairs. Haltere whitish-yellow. Legs: coxae with long white hairs, remaining parts covered by white scales dorsally, short black scales with sparse white scales ventrally. Tarsal claws evenly curved, each with short, thick basal tooth. Empodia longer than bend in claw. Female abdomen: As in L. carophila except for the following: densely covered by silvery-white scales ventrally and laterally; tergites each with wide band of dense black scales occupying half to two thirds of tergite length, with small extension in mid-posterior part. Tergite 7 as in Fig. 12. Sclerites of tergite 8 long and slender, widened anteriorly and posteriorly (Fig. 13). Sternites evenly sclerotized. Ovipositor: long (segments 9 + 10 of abdomen 4.29–5.33 times as long as tergite 6 (n= 12)). Male abdomen: as in L. carophila except for the following: color and scale pattern as in female. Terminalia (Fig. 20): gonostylus evenly narrow. Aedeagus blunt to slightly rounded apically. Hypoproct entire. Cerci separated by deep notch. Larva (third instar): bright orange with dark brown spatula; length: 2.36–3.56 mm (n= 20). Integument covered entirely by rounded to slightly pointed bumps. Antennae about twice as long as wide. Cephalic apodemes 1.5–2.8 times as long as head capsule (n= 11). Spatula (Fig. 23) with long shaft and two apically rounded anterior teeth separated by evenly rounded notch; shaft conspicuously widened immediately posterior to teeth and slightly widened at posterior end. On each side of spatula one sternal papilla and four lateral papillae, all asetose. Median pleural papilla asetose. Other pleural papillae and dorsal papillae with short setae. Terminal segment with two groups of three or four terminal papillae with short setae (Fig. 26). Pupa (Figs. 42–43): orange. Antennal bases developed into short, pointed horns. Cephalic seta long, situated on elevated base. Frons without papillae. Prothoracic spiracle thin and elongate; trachea ends at apex. Abdominal segments covered by short, pointed spinules except for short crinkled posterior area on each segment. Neotype: Ƥ, Israel, Giv’at Olga, 19.iii. 1996, Freidberg A., reared from Bilacunaria boissieri stem gall. Mounted on permanent microscope slide. Deposited in TAUI. Other material examined. 3 larvae, Georgia, Elisabethpol, vii–viii. 1934, unspecified collector, ex. Hippomarathrum crispum, Möhn collection number 9382 C, (mounted on permanent microscope slide by N. Dorchin), SMNS; 2 larvae, Israel [Palästina], unspecified date and collector, ex. Hippomarathrum crispum, Möhn collection number 9382 B, (mounted on permanent microscope slide by N. Dorchin), SMNS; 5 Ƥ, 2 3, same data as Neotype (1 Ƥ ZFMK, others TAUI); 2 Ƥ, Giv’at Olga, 12.iii. 1996, Dorchin N., ex. Bilacunaria boissier i (1 Ƥ TAUI, 1 Ƥ ZFMK); 4 Ƥ, 6 3, Park Hasharon, 22.i. 2000, Freidberg A. and Freidberg P., ex. Bilacunaria boissier i (3 Ƥ, 4 3 TAUI, 1 Ƥ, 2 3 ZFMK); 2 Ƥ, 1 3 (pinned), Park Hasharon, 22.i. 2000, Freidberg A., ex. Bilacunaria boissier i TAUI; 12 larvae (on two microscope slides), 3 pupal exuviae (on one microscope slide), Hadera, 20.ii. 2010, Freidberg A., ex. Bilacunaria boissieri, TAUI; 1 Ƥ, 2 3, Tel Baruch, 8.iii. 2011, Freidberg A., ex. Bilacunaria boissieri (pinned), TAUI. Distribution. South-west Asia, around the Caspian Sea (Russia [Petrowsk], Iran [Kurdistan], Georgia, Azerbaijan). The discovery of this species in Israel suggests that it has a continuous distribution in the Irano-Turanian region. In Israel it is distributed along the coastal plain, presumably corresponding to the distribution range of its host plant. Biology. Lasioptera umbelliferarum is univoltine. Galls develop into conspicuous swellings in stems and flower stalks (umbels) that may reach several centimeters in length and more than 2 cm in diameter (Figs. 52–53). Galls in umbels often prevent proper development of flowers. In Israel, the galls begin to develop in late winter (February), are green, sometimes with darker longitudinal stripes, and reach their final size in March–April, when the larvae inside them are still first instars. Each gall may contain several dozen larval chambers embedded in woody tissue so that galls are extremely hard and difficult to dissect. At later stages of gall development, a thick layer of spongy tissue develops around the larval chambers. By summer, the galled stems dry out and turn grayishbrown but remain attached to the bases of the green plants. The fully grown larvae remain inactive inside the galls until the following winter. They pupate and emerge in February-March. The inside walls of the larval chambers in fully developed galls are covered by a conspicuous white layer of mycelia. The galls frequently contain inquilinous moth larvae that feed on the gall tissues and dig tunnels in them, thus killing some of the gall inducers indirectly. The gall-midge larvae are also attacked by several species of parasitic wasps. Collecting the galls while they are still green results in the death of larvae inside them, thus the only way to rear this species is to collect the dried galls that contain pupae or diapausing larvae in February, shortly before adult emergence. This might be the reason why adults of this species have not been described to date. The above described phenology is in agreement with the data given by Rübsaamen (1895), who noted that galls collected in Russia in mid June contained third-instar larvae. By contrast, Möhn (1968), who examined galls from various localities in Western Asia, noted that galls that were collected in July-August contained fully developed pupae. These pupation dates do not match our findings and appear to be erroneous, in particular Möhn’s record from Israel. Remarks. The closest relative of L. umbelliferarum appears to be L. eryngii, which induces similar galls in several Eryngium species in Europe and the Mediterranean region, but we never found L. eryngii in Israel. According to Möhn (1968), the two species differ in the length of setae on the larval ventral papillae, and the teeth of the larval spatula are usually not as widely separated in L. eryngii as in L. umbelliferarum. Lasioptera umbelliferarum has a longer ovipositor and a significantly longer and slenderer sclerite 8 (compare Figs. 10–11 to Figs. 12–13). For additional differences between L. umbelliferarum and related species see the remarks under L. carophila above.Published as part of Dorchin, Netta & Freidberg, Amnon, 2011, The gall midges (Diptera: Cecidomyiidae) of Apiaceae in Israel, pp. 28-48 in Zootaxa 3044 on pages 36-37, DOI: 10.5281/zenodo.20344

Stefaniella brevipalpis Kieffer 1898

<i>Stefaniella brevipalpis</i> Kieffer, 1898 <p> <i>Stefaniella brevipalpis</i> Kieffer, 1898: 56</p> <p> Neotype of <i>Stefaniella brevipalpis</i> designated here – Ψ, Israel, Akko, 7.IX.2002, A. Freidberg, reared from <i>Atriplex portulacoides</i> stem gall. The neotype is designated in order to clarify the taxonomic status of <i>S. brevipalpis</i> Kieffer. Material representing this species is not found among specimens in the USNM or in the Muséum National d’Historire Naturelle in Paris, France (MNHN) (R. Gagné, personal communication). <i>Stefaniella brevipalpis</i> was described from an undisclosed locality in Italy, and is the only <i>Stefaniella</i> species known from <i>Atriplex portulacoides</i>. The galls were later described and illustrated by Houard (1908). The individuals we reared in Israel developed in the same host plant and the same type of galls from which the original name-bearing type had been described. Based on this information and on the original morphological description of the gall midges, we consider our specimens to be adequate for neotype designation. The neotype is deposited in the National Collection of Insects, Zoological Museum, Tel Aviv University, Israel (TAUI). The following description ensures recognition of the designated specimen, and characters differentiating the species from other congeners are detailed below under the remarks section.</p> <p> Adult. – <i>Head</i> (Fig. 45): Eye facets circular, less ordered and farther apart on vertex than laterally, gap between eyes on vertex 0.5–1.5 times as wide as facet. Palpus 2-segmented; second segment somewhat thinner than first, about as long. Antenna similar in both sexes; scape conical, pedicel globular, 9–10 flagellomeres, barrel-shaped, about 1.3 times as long as wide, each with two closely appressed whorls of circumfila, one whorl of setae proximal to basal circumfilum, and group of stronger setae distal to it. Apical flagellomere sometimes comprised of 2–3 fused segments, slightly tapered.</p> <p> <i>Thorax</i>: Wing: length 1.2–1.4 mm in females (n=5), 1.0– 1.2 mm in males (n=3), transparent. Vein R5 joins C at wing mid length, M present, Cu unforked. C, Sc and R5 with dark scales. Legs (Fig. 46): Tarsal claws toothed, evenly curved, teeth very short, strongly curved at base. Empodia reaching bend of claw or slightly shorter.</p> <p> <i>Female abdomen</i> (Fig 47): Tergites 1–6 rectangular, not extended ventrally, tergite 7 extending ventrally, each tergite with 1–2 posterior rows of setae, anterior trichoid sensilla, and evenly covered by scales; tergite 8 smaller than preceding tergites, tapered ventrally, with anterior trichoid sensilla, without setae or scales. Sternites 2–7 rectangular, with row of posterior setae and several additional, scattered setae; sternite 8 completely undifferentiated from surrounding membrane. Area between tergite 8 and lateral group of setae covered by minute hyaline setae on small bumps. Ovipositor (Figs. 48–49): Lateral group of setae comprising about 70 relatively short, straight or curved setae. Sclerotized rods widened posteriorly to form weakly sclerotized triangular plate covered by minute spines. Lateral plate sheathing almost entire base of cercus, bearing about 40 straight, short and thick setae that are narrowed abruptly at tip. Aculeus straight and thick, evenly narrowed towards tip, with two rows of minute fine setae; tip of aculeus slightly curved ventrally. Apical lamella rectangular, as long as aculeus, evenly setose.</p> <p> <i>Male abdomen</i>: Tergites 1–7 rectangular, setation and scales as in female. Tergite 8 completely undifferentiated from surrounding membrane. Sternites 2–7 as in female; sternite 8 shorter and smaller than preceding sternites, without setae or scales. Terminalia (Fig. 50): Gonocoxite cylindrical, with several strong setae on sclerotized parts, more numerous on ventral part than on dorsal part; mediobasal lobe prominent, evenly setulose. Gonostylus slender, strongly arched, slightly angular at distal third, evenly setulose with several strong setae both dorsally and ventrally. Cerci fused, notched posteriorly, evenly setulose, with several strong setae. Hypoproct entire, distally rounded, evenly setulose, with two strong apical setae. Paramere divided longitudinally into dorsal and ventral lobes; ventral lobe smooth, with apical seta on elevated base; dorsal lobe densely covered by curved setae. Aedeagus slender, truncated at tip, slightly longer than paramere.</p> <p>Larva (Figs. 52–53) – White to pale yellow; cylindrical; length: 1.9–2.2 mm (n=4). Integument bumpy on entire surface, bumps rounded on dorsum, tapered on ventrum. Antenna about 1.5 times as long as wide. Posterolateral apodeme 2.2 times longer than head capsule. Spatula with slender shaft and two triangular anterior lobes separated by wide notch. On each side of spatula asetose sternal papilla, two inner lateral papillae with strong setae, and one asetose outer lateral papilla. Pleural and dorsal papillae with long setae. Terminal segment with six papillae, two situated ventrally next to anus, all with strong setae.</p> <p>Pupa. – Unknown.</p> <p> Other material examined – All material from Israel, Akko, collected by N. Dorchin from stem galls of <i>Atriplex portulacoides</i> unless otherwise noted. 2 Ψ, 3.III.1995; 4 larvae, 9.III.1999; 2 Ψ, 2 ɗ, 7.IX.2002, A. Freidberg; 3Ψ, 1ɗ, 10.IX.2008, A. Freidberg.</p> <p>Distribution. – Israel (Na’aman salt marsh), Italy, UK, Portugal, Czech Republic.</p> <p> Etymology: The name <i>brevipalpis</i> is Latin for short palpus. Kieffer (1898) indicated that the second segment of the palpus is about half as wide as the first, an attribute that allegedly sets the species apart from its congener, <i>S. atriplicis</i>.</p> <p> Biology: The species induces single or multi-chambered galls in stems, leaf petioles or midribs of <i>Atriplex portulacoides</i> (Figs. 9–10). Galls are not common and are sometimes very difficult to find, although they can be locally abundant on several adjacent plants. Although the host is abundant in other localities in Israel (e.g., in the Atlit salt marsh and along the banks of the Yarkon river in Tel Aviv), galls were found only in the Akko salt marsh. The galls constitute 1–3cm wide swellings of variable shapes and are mostly green, but sometimes have a touch of red. First and second instar larvae are found in chambers lighter in color than the remainder of the gall tissue. Third instar larvae are found individually in rigid capsules inside the galls, where they spin a white cocoon before pupating. The chamber is sealed inside the gall by a round, thin, whitish disk that appears to be derived from silk. Pupae were not found and adults were extremely difficult to rear, both due to high parasitism rates and to a very short emergence season. However, the fact that adults were reared both in March and in August-September suggests that the species has at least two generations a year.</p> <p> Remarks. – The genus <i>Stefaniella</i> includes 9 described species, all but one are restricted to saltbushes (<i>Atriplex</i> spp.). Three species, <i>S. atriplicis</i> Kieffer, <i>S. brevipalpis</i>, and <i>S. trinacriae</i> Stefani, were described from salt-marsh habitats in the Mediterranean area (Kieffer 1898, De Stefani 1900), whereas the remaining six were described from hosts that typically grow in arable or disturbed habitats in central Asia or in Italy and Eastern Europe (Gagné 2004).</p> <p> <i>Stefaniella brevipalpis</i> is the only species recorded from <i>A. portulacoides</i>. Although Houard (1908) lists <i>A. halimus</i> as a host, we never found <i>S. brevipalpis</i> galls on this plant, so this record may be erroneous. <i>Stefaniella brevipalpis</i> galls are much smaller and more regular in shape than the amorphous galls of <i>S. trinacriae</i> on <i>A. halimus</i>, which often exceed 5 cm in length (N. Dorchin, unpublished data). The larvae of <i>S. brevipalpis</i> are orange, whereas those of <i>S. trinacriae</i> and <i>S. altiplicis</i> (both on <i>A. halimus</i>) are white to pale yellow (Kieffer 1898, 1912, De Stefani 1900). Kieffer (1898) stated that the larval spatula differs between <i>S. brevipalpis</i> and <i>S. atriplicis</i> but did not elaborate. Another alleged difference between these species is the thinner second segment of the palpus in <i>S. brevipalpis</i>, but we did not find the palpus in this species to be different from that of congeners we examined. We did find a good morphological difference between the ovipositors of <i>S. brevipalpis</i> and <i>S. trinacriae</i>, namely, a posterior extension of the lateral plate in the latter that is absent in the former (Fig. 51).</p> <p> While <i>S. trinacriae</i> appears to stand out from the other two species both with regard to adult and gall morphology, <i>S. brevipalpis</i> and <i>S. atriplicis</i> are possibly synonymous. We examined adults reared by Fedotova from <i>Atriplex longifolia</i>, which allegedly belong to <i>S. atriplicis</i>, and could not find any morphological differences between them and the adults we reared from <i>A. portulacoides</i>. Examination of type material of two central-Asian species, <i>S. usjurtensis</i> Fedotova and <i>S. gigantea</i> Fedotova, did not reveal any morphological differences as well, other than the larger size of <i>S. gigantea</i> adults. Given the remarkable morphological uniformity of <i>Stefaniella</i> adults, a study of the immature stages is highly warranted, but it is likely that only a molecular analysis would allow determining the validity of species and the relationships among them. Until such analysis is conducted, the best characters for distinguishing among <i>Stefaniella</i> species are those of their galls.</p>Published as part of <i>Dorchin, Netta & Freidberg, Amnon, 2008, The Chenopodiaceae-feeding gall midges (Diptera: Cecidomyiidae) of the Na'aman salt marsh, Israel, pp. 1-22 in Zootaxa 1937</i> on pages 15-18, DOI: <a href="http://zenodo.org/record/184984">10.5281/zenodo.184984</a&gt

Lasioptera foeniculi Dorchin and Freidberg, new species

Lasioptera foeniculi Dorchin and Freidberg, new species (Figs. 2, 4, 7, 15–17, 21, 24, 27–29, 51) Adult: Head (Fig. 2): General color dark brown, densely covered by white scales. Eye facets circular, gap between eyes on vertex 0–0.5 facet wide. Palpus 4 -segmented; palpiger usually well developed; segment 1 as long as or slightly longer than wide; segments successively longer or segments 3–4 same length. Antenna: flagellomeres 11– 13 in both sexes (n= 9 Ƥ, 4 3); flagellomeres 1–3 longer than wide, remaining flagellomeres except for apical, rectangular, wider than long; first two flagellomeres partially fused, last flagellomere often longer than preceding, rounded apically, often consisting of 2 fused flagellomeres; each flagellomere except apical with two whorls of circumfila connected by 1–2 longitudinal branches, row of short, curved setae proximal to proximal circumfilum, and several strong setae between circumfila whorls; otherwise evenly covered by microtrichia (Fig. 4). Apical flagellomere often with more than 2 circumfila whorls. Thorax: General color dark brown, covered by white scales. Wing (Fig. 7): length 0.95–1.27 mm in female (n= 11), 0.91–1.16 mm in male (n= 4); covered by dark brown hairs, veins faint; C covered by dark brown scales to merging point with R 5, M present, Cu unforked. Legs covered by dark brown scales dorsally, white scales ventrally. Tarsal claws toothed; teeth evenly curved. Empodia longer than bend in claws (Fig. 6). Female abdomen: general color pinkish, covered by white scales; scale pattern on each tergite comprises wide band of black scales on proximal part and thin band of white scales along lateral and posterior margins. Tergites 1– 6 with two sensory setae at mid-anterior part and row of setae along posterior margin; tergite 7 unsclerotized mesolaterally, with two sensory setae anteriorly and 1–2 rows of strong setae along posterior margin (Fig. 17); sclerite 8 divided into two elongate sclerites, each widened posteriorly, with one sensory seta at proximal third and few setae posteriorly (Fig. 15). Sternites weakly sclerotized along posterior half, with two closely adjacent sensory setae anteriorly, few setae mesally, and row of strong setae along posterior margin; sternite 7 less sclerotized laterally and more setose than preceding; sternite 8 undifferentiated from surrounding membrane. Ovipositor (Fig. 15): long (segments 9 + 10 of abdomen 4.48–5.16 times as long as tergite 6 (n= 9)); lateral group of setae on segment 8 comprising relatively few (15-20) short, apically rounded, scale-like setae; lateral plate virtually absent; dorsal part of fused cerci bearing two long, hooked, apically rounded dark setae and at least two fine strongly curved setae anterior to the hooked setae; fused cerci otherwise setulose and with numerous straight, long, fine setae on entire surface. Male abdomen (Fig. 16): Scale pattern as in female. Tergites 1–6 with two sensory setae anteriorly and row of setae posteriorly; sclerotization of tergites 7–8 reduced to narrow anterior band, with two anterior sensory setae and no posterior setae. Sclerites of sternites 2–7 divided into two bands, anterior longer than posterior, with two closely adjacent sensory setae anteriorly, row of strong setae posteriorly and several strong setae mesally; Sclerotization of sternite 8 reduced to single median band, with anterior sensory setae and row of posterior setae. Terminalia (Fig. 21): gonocoxite with numerous, evenly distributed setae; mediobasal lobe long and narrow, sheathing aedeagus almost to apex, covered by curved setae. Gonostylus narrowed abruptly at distal third, setulose on proximal third, ridged on distal two thirds, with several setae; tooth well developed. Aedeagus blunt apically. Hypoproct entire. Cerci separated to base, setose and setulose. Larva (third instar): Ovoid, orange, with dark brown spatula; length: 1.53–1.62 mm (n= 6). Integument evenly bumpy except for two narrow anteroventral patches on each segment. Cephalic apodeme about twice as long as head capsule. Antennae about twice as long as wide. Spatula (Fig. 24): with long slender shaft and two pointed anterior teeth separated by triangular notch. On each side of spatula one sternal papilla and four lateral papillae, all asetose. Median pleural papilla asetose. Other pleural papillae and dorsal papillae with long, well developed setae. Terminal segment with two groups of three terminal papillae with long, conspicuous setae (Fig. 27). Pupa (Figs. 28–29): Brown. Antennal bases developed into very short, blunt horns. Cephalic seta long, situated on elevated base. Prothoracic spiracle thin and elongate, trachea ends at apex. Frons on each side with one pair of lateral and one pair of median papillae, one papilla of each pair with very short seta. Dorsum of abdominal segments covered by tiny, pointed spinules. Holotype: Ƥ, Israel, Kefar Hahoresh, 19.x. 2009, N. Dorchin, reared from fruit gall on Foeniculum vulgare, mounted on permanent microscope slide, deposited in TAUI. Paratypes: 4 Ƥ, 3 3, Israel, Kefar Hahoresh, 3.x. 1997, N. Dorchin (3 Ƥ, 2 3 TAUI, 1 Ƥ, 1 3 ZFMK); 6 larvae (on one microscope slide), Israel, Kefar Hahoresh, 11.x. 1997, N. Dorchin (TAUI); 4 Ƥ, 1 3, 4 pupal exuviae (on one microscope slide), same data as holotype (3 Ƥ, 1 3, exuviae TAUI, 1 Ƥ ZFMK). All material from Foeniculum vulgare. Other material examined. 2 Ƥ, 2 3, Israel, Kefar Hahoresh, 3.x. 1997, N. Dorchin, ex. Foeniculum vulgare. Distribution. This species is currently known only from Israel. Galls were observed in the Upper and Lower Galilee and the Shfela (Nahal Amud, Sede Ya’acov, E’n Alva, Kefar Hahoresh, and Tel Keshet). Etymology. The species name refers to its host-plant genus, Foeniculum. Biology. Galls develop in fruits of Foeniculum vulgare (Fig. 51). Thus, this is the only Lasioptera species from Apiaceae that does not develop in stems or umbels. A galled fruit is 2–5 times as large as a normal fruit, contains a single chamber, and larvae develop in it gregariously (up to 12 larvae observed in dissected galls). Mycelia were not observed in the galls. The galls are uncommon but not rare and usually only one or few fruits are galled within a single umbel. They constitute inflated, distorted fruits, usually green but sometimes purplish, and were observed only in September-November. The larvae leave the galls and pupate in the soil. Adults emerged within 10 days. It therefore appears that this species is univoltine or bivoltine - with two generations within a very short time during the fall. Other life-history aspects of this species are unknown. The galls are often attacked by hymenopteran ectoparasitoids that each prey on several gall-midge larvae and may kill all the larvae in a gall. Remarks. This species is most similar morphologically to Microlasioptera flexuosa (Winnertz 1853) that develops in stems of Phragmites australis (Skuhravá and Skuhravý 1981). The latter species was originally described under Lasioptera but transferred to Microlasioptera by Skuhravá and Skuhravý, who created the genus for it. We do not see the need for a separate genus for the two species and consider them as belonging to Lasioptera. The small number of hooked setae on the female cerci and the absence of the lateral plate represent an extreme situation in the genus, but intermediate states exist between it and the more common situation of an ovipositor with a well developed lateral plate and numerous hooked setae as in L. carophila, for example. An example for such an intermediate state is found in L. hungarica Möhn that is redescribed by Skuhravá and Skuhravý in the same work (1981), including a detailed discussion and illustration of the morphological variability of the ovipositor. Among the distinguishing characters of Microlasioptera, the authors mention the weak wing venation and reduced number of palpal segments (mostly 3 rather than the usual 4). We consider both characters insufficient to justify a new genus. Lasioptera foeniculi differs from M. flexuosa in the smaller number of antennal flagellomeres in females (11– 13 versus 17–20, respectively), the constant number of 4 palpal segments (as opposed to usually 3 in M. flexuosa), and the number of hooked setae on the female fused cerci (always 2 in L. foeniculi, 2–4 in M. flexuosa), although these characters vary often within the same species in the Lasiopterini. Larvae of the two species exhibit better differences: in M. flexuosa, the group of lateral papillae closest to the spatula (basically numbering 3 papillae in Lasiopteridi but usually only 1 in Lasioptera) has been completely lost whereas in L. foeniculi one of these is still present. It is noteworthy that according to Skuhravá and Skuhravý (1981), all lateral papillae in M. flexuosa are asetose but Möhn (1968, under L. flexuosella) showed that two of them are setose. Both descriptions agree that the spatula is very weakly sclerotized, often without a visible shaft. By contrast, in L. foeniculi the spatula is well developed. Apart from developing in different host plants and galls, the two species differ also in their life history, as L. foeniculi pupates in the soil whereas M. flexuosa pupates inside the gall.Published as part of Dorchin, Netta & Freidberg, Amnon, 2011, The gall midges (Diptera: Cecidomyiidae) of Apiaceae in Israel, pp. 28-48 in Zootaxa 3044 on pages 37-39, DOI: 10.5281/zenodo.20344

Careopalpis Marikovskij 1961

Genus Careopalpis Marikovskij, 1961 Careopalpis is an Old World genus of the tribe Lasiopterini, currently comprising 14 species, all associated with Chenopodiaceae in Eastern Europe and Central Asia. Adults are small, stout, and very uniform morphologically. The female ovipositor is characterized by a ventrally curved aculeus, a fusiform apical lamella, and a lateral plate bearing relatively long and curved setae. The ovipositor is shorter than in the related genus Baldratia, the number of antennal flagellomeres rarely deviates from 10 in both sexes, and flagellomeres are never fused. Palpus is one-segmented. Larvae do not have a spatula.Published as part of Dorchin, Netta & Freidberg, Amnon, 2008, The Chenopodiaceae-feeding gall midges (Diptera: Cecidomyiidae) of the Na'aman salt marsh, Israel, pp. 1-22 in Zootaxa 1937 on page 9, DOI: 10.5281/zenodo.18498