Ita Tournier, 1878

Meregalli, Massimo & Borovec, Roman, 2011, Radiation in the halophytic coenoses of the Peri-Tethys: taxonomy and biogeography of the genus Ita (Coleoptera: Curculionidae), Journal of Natural History 45 (21 - 22), pp. 1331-1401 : 1335-1344

Ita Tournier, 1878

( Figure 2 )

Ita Tournier, 1878 : XVIII.

Rhinospathus Desbrochers, 1895: 57 .

Xeronoma Iablokov-Khnzorian, 1964: 173 , syn. n.

Type species

Ita crassirostris Tournier, 1878 , designated by Alonso-Zarazaga and Lyal (1999).

Redescription

Size small, between 1.50 and 2.30 mm, body elliptical. Integument black, rostrum smooth and glossy, often ferruginous, particularly in apical half, more or less deeply punctured before antennal insertion and smooth, glossy, finely punctulate in apical part; head, pronotum and elytra very densely punctured; interspaces of punctures glossy, smooth, without any granules or raised sculpture; legs and antennae often at least in part reddish or yellowish. Hind wings fully developed. Vestiture: rostrum almost completely devoid of vestiture, except near its base, with four long sensilli at the epistoma, two near the middle and two laterally, with two sensilli on the sides; underside with a few downward-directed setae in apical quarter of prementum; head, pronotum, scutellum, elytra, ventral parts and coxae very densely covered with oval or oblong, matt, light greyish or yellowish to nearly whitish scales, inserted in the punctures, three to five times as long as wide, rounded on both extremes, seldom subacute or subtruncated at apical margin, smaller and narrower on head, transversely inserted and directed towards median part on pronotum, longitudinally aligned on elytra, not or very scarcely imbricate and generally appressed to integument, seldom slightly raised posteriorly; sternum nearly as densely covered as pronotum and elytra, with scales of similar shape, abdominal ventrites with usually elliptical, more slender scales; legs with slender scales on femora and tibiae, apex of tarsal segments with scales and a few setae. Rostrum robust, short, at most as long as head and pronotum together, usually shorter, distinctly broadened from base to apex, with maximum width at apex and shortest width at or near base, generally slightly or more distinctly broadened at antennal insertion; apex conically prominent at sides of epistoma; epistoma distinct, nearly truncate; dorsum from base to antennal insertion more or less deeply rugose or punctured, punctures progressively shallower and more sparse, scattered towards apex; in lateral view rostrum of equal thickness from base to apex, seldom very slightly narrowed towards apex, more or less uniformly curved, often more curved and weakly gibbous basad; underside furrowed medially, roughly punctured, ventral keels more raised, subparallel basally, lowered and scarcely distinct, strongly divergent laterally; scrobes often nearly as broad as rostrum thickness, very short, rarely extended near to eyes, upper and lower margins not keeled, curved downwards, upper margin directed below the eye; integument of bottom of scrobes deeply rugose, matt. Mouth parts ( Figure 2C ): maxilla with moderately curved cardo, stipe subcylindrical, feebly broadened apicad, palpifer broad, cylindrical, with a strong lateral seta, mala semicircular, with about 10 strong, lateral, curved teeth, and long, dense setae; maxillary palpi three-segmented, robust, prominent from rostrum, segments 2 and 3 visible from above in frontal view of apex, segments 1 and 2 transverse, segment 1 distinctly broader than 2, segment 3 shortly conical, with few short setae at apex; labium narrowly rectangular, with two groups of apical setae; labial palpi bisegmented, short; mandibles completely exposed in frontal view of rostrum, with two fully developed main teeth and the third, lower tooth very minute. Antennae slender, inserted in basal part of rostrum, in some species inserted near to base, up to slightly before mid-length in other species; scape narrow, moderately and regularly thickened apicad, slightly curved forwards, as short as rostrum width at antennal insertion or longer, up to nearly twice as long as rostrum width at antennal insertion; funicle seven-segmented, segment 1 thicker than the others, segments 2–7 of variable length, often segments 2–4 longer than wide; club oblong, segment 1 shorter than the length of segments 2 + 3 together, margin of each segment and annulus of segment 3 with a ring of short and dense setae. Head rounded, short, vertex regularly convex, eyes large, slightly oval, with maximum diameter in basal-frontal direction, moderately prominent from the convexity of head, interocular space usually narrower than rostrum at base. Pronotum short, transverse, base straight, sides often scarcely broadened at mid-length, seldom more broadly rounded and with maximum width in basal half; anterior margin straight, postocular lobes not developed, upper part not prominent above head; dorsum weakly convex, occasionally with a narrow glossy median line on anterior part. Scutellum visible, triangular. Elytra oblong, at base slightly broader than pronotal base, humeri prominent, sides usually nearly linearly and moderately broadened from base to the point of maximum width, beyond mid-length, distinctly rounded in apical part and very regularly jointly united at apex; in lateral view dorsum flattened or weakly convex in posterior half, declivity regularly and moderately curved; all intervals flat, suture not distinct from the other intervals; whole surface covered with dense uniform punctures, striae 10, undifferentiated in dorsal view, barely distinct also on internal face of elytra. Hind wings with only radial and cubital veins distinctly sclerotised, cubital vein longer than radial vein, anal vein 3A barely distinct, other anal veins indistinct, postradial stripe weakly differentiated, hind margin with long setae. Legs slender. Fore coxae conical, slightly longer than wide, adjacent to each other, middle coxae shorter, transversely oval, with a small raised lateral prominence, slightly separated, hind coxae transverse, widely separated; trochanters small, triangular, with a long downward-directed seta; femora scarcely broadened at middle, not toothed; tibiae straight, slightly broadened from base to apex, inner side linear, apical uncus present, big, connected to median part of apex; inner margin not toothed, premucro indistinct also in females; fore and middle tibiae nearly perpendicularly truncated at apex; hind tibiae slightly obliquely truncated, apex simple, lacking differentiated corbels. Tarsi elongate, segments 1 and 2 long conical, very slightly broadened from base to apex, segment 2 as long as or shorter than segment 1, segment 3 of various shapes, approximately as long as segment 2, with lobes round, broadly expanded or small and scarcely developed, usually slightly asymmetrical, with inner lobe smaller; lobes of fore tarsi sometimes broader than those of middle and hind tarsi; onychium slender, longer than segment 3, claws ( Figures 2G, H ) short and strong, widely open, connate basally at margin of onychium, distinctly thickened and sub-denticulate at base, inner side crenulate and with a seta; underside of tarsal segments 1 and 2 with sparse, narrow setae, underside of segment 3 with a small pad of fine, whitish setae on apical part of lobes. Prosternum without apical emargination, not impressed medially, in front of coxae shorter than diameter of coxae, posterior margin ciliate; mesosternum very narrow, much shorter than diameter of middle coxae; metasternum as long as length of middle coxae, greatly broadened laterally, metepisternum subrectangular, moderately broadened anteriad. Ventrite I slightly longer than length of metacoxae, intercoxal process very broad, apex rounded, suture between ventrites I and II continuous, slightly sinuate in median part, not curved forwards at lateral margin, ventrite II at sides as long as ventrite I, shorter than ventrite I in front of metacoxae, ventrites III and IV narrow, suture straight; ventrite V shorter than ventrites III + IV together, apex sublinear. Proventriculus not differentiated. Genitalia: aedeagus ( Figure 2A ) slender, slightly and regularly curved, dorsum membranous, sides strongly sclerotized, sclerotization more or less thickened medially, sides sublinearly converging towards lamella; lamella very small, narrow, acutely pointed, generally curved upwards, struts of median lobe as long as median lobe; internal sac lacking large sclerites, with a stiff, narrow, single median sclerotized flagellum protruding from internal tube and nearly as long as struts of median lobe; tegmen lacking a dorsal plate, strut elongated, ring broad, narrow, with very feeble sclerotization, parameroid lobes not sclerotized, barely visible; hemisternum VIII triangular, expanded, with few apical setae; sternum IX scarcely curved, shortly bifurcate ( Figure 2B ). Sternum VIII of female ( Figure 2D ) with narrow, elongate strut, lacking differentiated arms; lamina broad, as long as wide or transverse, weakly sclerotized in basal and lateral part; hemisternum IX cylindrical ( Figure 2E ), not broadened basad, styli apical, slender, slightly conical; spermatheca ( Figure 2F ) large, as long as hemisterna, nodulus thickened and enlarged, cornu short and very narrow, sharply curved, acutely pointed.

Systematic position

As summarized above, the systematic position of the genus Ita has been debated among authors and an attempt towards a more solid classification is required. Unfortunately, the framework on which the infrafamily-rank classification of the Curculionoidea is based is still quite weak, and synapomorphies for the various subfamilies and tribes have never been properly defined. It is true that any discussion lacking a cladistic analysis on a broad spectrum of taxa is purely speculative, but on the other hand it is also true that, in the absence of standardized sets of characters, the inclusion of Ita in an analysis based on character-states of uncertain significance in selected taxa of various subfamilies may not result in a more reliable placement of the genus. Thus, the classification here proposed is based on a critical analysis of some key traits, on which the present-day composition of the Curculionoidea is founded, and is thus consistent with the current system, pending an analysis of far larger scale than can not be carried out in this study.

The original placement of the genus in the Tanyrhynchini ( Entiminae , cf. Alonso- Zarazaga and Lyal 1999) does not seem to be justified, since no shared characters ascribed phylogenetic significance between the taxa referred to that tribe and Ita . The attribution to the Molytinae by Alonso-Zarazaga (2002), possibly following Iglesias Iglesias (1922), can be excluded on the basis of several discriminating traits. Ita differs from the Molytinae in the morphology of the apex of the hind tibiae, lacking corbels (cf. Thompson 1992), in the different structure of the male sternum VIII ( Fig. 2B ), that in the Molytinae is composed of a pair of broad subtriangular hemisterna, with setae on their apical part, near the joining part (cf. Thompson 1992), and in the sclerotization of the hind wings, limited to the radial and cubital veins, whereas in the Molytinae several other veins are sclerotised (cf. Zherikhin and Gratshev 1995). The undifferentiated proventriculus is another state that distinguishes Ita from the Molytinae and also from other apparently more derived Curculionidae ( Kissinger 1963; Calder 1989; Kuschel 2008), that have a proventriculus composed of eight plates with setae and sclerites. The absence of this structure would probably suggest a more basal position of Ita with respect of these subfamilies, even though there is faint indication that in rare and occasional cases the proventriculus is secondarily undifferentiated ( Molytinae Cryptoplini, Kushel 2008 ; but the affinities of Cryptoplini are questionable). Finally, the flagellum in the internal sac of the aedeagus of Ita ( Figure 2A ) is usually absent from the Molytinae , although it has been observed in some extrapalaearctic genera, such as some Trypetidini ( Platynanus Aurivillius , Phronira Broun ), Trigonocolini ( Trigonocolus Lacordaire ), Ithyporini ( Ectatorhinus Lacordaire ) and Pissodini ( Orthorhinus Schoenherr ) (C. Lyal, personal communication). Such a flagellum is frequently expressed in “primitive” taxa, such as the Belinae (R. Oberprieler, personal communication) and some, but not all, species of Ocladius Schoenherr (Brachyceridae) ( Meregalli and Colonnelli 2006); it can be added that it is usually present in taxa with an aedeagus of the pedotectal type, that is, with a dorsal plate covering the whole median lobe (cf. Thompson 1992). This aedeagal type is found in most of the taxa of Curculionoidea that, in a cladistic analysis, clustered in basal clades ( Marvaldi et al. 2002). However, the dorsal plate is lacking in Ita , whose aedeagus is of the pedal type, with short parameroid lobes connected with the ring of the tegmen. Among the weevils with an aedeagus of the pedal type, the flagellum is sporadically and randomly expressed in genera belonging to unrelated taxa, such as several Curculioninae (i.e. tribe Mecinini : R. Caldara, personal communication), Tanyrhynchini (R. Oberprieler, personal communication), the previously mentioned Molytinae , etc. In these taxa the flagellum seems thus to be a plesiomorphic remnant, occasionally conserved, and therefore its phylogenetic information for Ita should be lightly weighted, it being simply an indicative trait and not a synapomorphy.

Most authors had placed Ita in the subfamily Erirhininae , now upgraded to family-rank as Erirhinidae ( Thompson 1992; Alonso-Zarazaga and Lyal 1999). Within this family, Ita was generally left in the monotypic tribe Itini . However, in the typical Erirhinidae the aedeagus is of the pedotectal type and this would exclude Ita from being part of that subfamily.

The form of the rostrum in lateral view, that of the scrobes and the mouthparts, the short and round head, the broadly divergent, sub-appendiculate claws ( Figures 2G, H ), the aedeagus of the pedal type and the structure of the male sternum VIII ( Figure 2B ) suggest that Ita belongs to the subfamily Curculioninae , where it was placed by Alonso-Zarazaga and Lyal (1999). The presence of a flagellum in the internal sac of the aedeagus, shared with many Curculioninae , and the common absence of a differentiated proventriculus, although apparently plesiomorphic states and thus only indicative, reinforce this classification. Within this subfamily the genus cannot be included in the tribe Styphlini , near to which it has often been listed ( Reitter 1913; Klima 1934), since all the genera of this tribe (with the exception of Geranorhinus Chevrolat, 1860 , which is to be excluded from the Styphlini ) share a peculiar shape of the rostrum: curved, with the antennae inserted near the apex, and with narrow and deeply impressed scrobes, which follow the dorsal curvature along its whole length, and whose upper margin is directed above the eyes. Ita , in contrast, has very short scrobes and the antennae are inserted near or before the mid-length of the rostrum. Also, the genitalia of both the sexes show significant differences. An attribution of Ita to any of the other tribes of the Curculioninae would equally be scarcely justified, and could result in a polyphyletic taxon. In the position of the antennae and the shape of the scrobes Ita approaches Geranorhinus , a genus that is also associated with salty habitats, although its species feed on Tamarix rather than on the Salicornioideae . Clearly the general appearance is relatively distinct: the rostrum of Geranorhinus is not broadened dorsally and does not show sexual dimorphism, and the aedeagus has no sclerotized flagellum; however, the shape of the male sternum VIII is quite similar in the two genera, and other possible homologies are seen in the wing venation, with only radial and cubital veins distinctly sclerotized and hind margin with long setae (see Zherikhin and Gratshev 1995), and in the structure of the female genitalia, mainly sternum VIII and spermatheca. For these reasons, we propose maintaining Ita in the tribe Itini of the Curculioninae , also tentatively including Geranorhinus in the same tribe.

Key to the species

N.B. The main characters discriminating the species of Ita are the shape of the rostrum and the ratio between some of its parts. The length was always measured with the rostrum in lateral view; the rostrum length is the space from the anterior margin of the eye to the apex of the epistoma; the other length measurements are self explanatory; the antennal insertion refers to the anterior margin of the scrobe; the width was always measured in dorsal view, perpendicularly to the rostrum (see also Figure 1 ). All measurements are reported in table 1.

1. Species from the western Mediterranean region and Macaronesia......... 2

Species from western and central-western Asia........................... 9

2. Trochanters and at least basal part of femora dark ferruginous to blackish. Usually at least antennomeres 4–7 brownish to dark brownish. Rostrum in female at apex feebly widened, 1.1–1.2 times as wide as at antennal insertion, with almost straight sides. Scales on elytra narrow, not covering integument. Sclerotized margins of aedeagus as wide as median membranous part. Spain............................................... Ita hispanica sp. nov.

Trochanters and basal part of femora at most dark reddish; at most antennomeres 6 and 7 dark brownish to blackish, seldom also segments 4–5 dark ferruginous. Insects not showing the complex of characters of rostrum and aedeagus as above...................................................... 3

3. Rostrum very short, ratio length/width at apex <2.0 in male, <2.5 in female.................................................................. 4

Rostrum relatively or very slender, ratio length/width at apex> 2.0 in male,> 2.5 in female......................................................... 6 4. Sides of rostrum in apical part straight. Rostrum in male very short, ratio length/width at apex <1.8. Sclerotized margins of aedeagus narrow, central membranous part broader than sclerotized margins. Sicily, northern Tunisia.......................................... Ita crassirostris Tournier

Sides of rostrum in apical part feebly or strongly concave. Rostrum in male slightly longer, ratio length/width at apex 1.8–1.9. Structure of aedeagus variable................................................................ 5

5. Apical part of rostrum in female strongly broadened, sides nearly linear, width at apex> 0.20 mm. Antennal insertion in male in basal half of rostrum. Sclerotized margins of aedeagus narrower than median membranous part. Canary Islands: La Graciosa..................... Ita gratiosa sp. nov.

Apical half of rostrum in female moderately broadened, sides strongly concave, width at apex <0.20 mm. Antennal insertion in male at mid-length of rostrum, or slightly beyond. Sclerotized margins of aedeagus as broad as median membranous part. Southern Algeria.... Ita chobauti (Desbrochers)

6. Elytral scales slender, elliptical, four times as long as wide. Rostrum in female very slender, scarcely broadened at apex, ratio length/width at apex almost 3.00. Algeria: Blida ................................... Ita gracilis Tournier

Elytral scales oval, three times as long as wide, densely covering integument. Rostrum in female robust, ratio length/width at apex 2.45–2.80........... 7

7. Rostrum length> 0.45 mm in male and> 0.60 mm in female. Sclerotized sides of aedeagus as broad as median membranous part. South-eastern Morocco........................................... Ita chavanoni sp. nov.

Rostrum length <0.45 mm in male and <0.60 mm in female............. 8

8. Sclerotized margins of aedeagus broad, expanded onto central part and nearly touching at mid-length, central membranous part very narrow. Rostrum of female in basal half deeply and densely punctured, nearly matt. Antennal insertion in male at mid-length of rostrum. Segment 3 of fore tarsus twice as wide as segment 2. North-western Algeria............................................... Ita berbera sp. nov.

Sclerotized margins of aedeagus narrow, central membranous part broader than sclerotized margins. Rostrum of female in basal half shallowly punctured, glossy. Antennal insertion in male before mid-length of rostrum. Segment 3 of fore tarsus less than twice as wide as segment 2. Tunisia................................................ Ita punica sp. nov.

9. Segment 3 of tarsus narrow, at most 1.3 times as wide as segment 2...... 10

Segment 3 of tarsus wide, broadly bilobed, 1.5 times as wide as segment 2............................................................. 11

10. Antennomeres 3–5 longer than wide, fore tarsal segment 1 nearly three times as long as wide. Antennal insertion, at least in female, less basal: ratio length from base to antennal insertion/length from base to apex> 0.40; ratio length from base to antennal insertion/width at antennal insertion> 1.10. Turkmenistan, Uzbekistan........................... Ita turkmena sp. nov.

Antennomeres 3–5 globose, fore tarsal segment 1 twice as long as wide. Antennal insertion, at least in female, basal: ratio length from base to antennal insertion/length from base to apex <0.40; ratio length from base to antennal insertion/width at antennal insertion 1.00. Armenia........................... ................... Ita latirostris (Iablokov-Khnzorian)

11. Antennal insertion near base of rostrum, ratio length from base to antennal insertion/width at antennal insertion <1............................... 12

Antennal insertion near mid-length of rostrum; ratio length from base to antennal insertion/width at antennal insertion>1...................... 13

12. Elytra flattened; rostrum thickened in lateral view; ratio length from antennal insertion to apex/width at apex 1.20–1.25 in male, 1.60–1.70 in female. Armenia, Azerbaijan, Georgia......................... Ita kirschi Tournier

Elytra distinctly convex; rostrum slender in lateral view; ratio length from antennal insertion to apex/width at apex 1.35–1.40 in male, 1.80–1.95 in female. Central Turkey.............................. Ita korotyaevi sp. nov.

13. Elytral scales broad, three times as long as wide, completely covering integument. Central Turkey................................... Ita caldarai sp. nov.

Elytral scales slender, four times as long as wide, integument between scales clearly visible......................................................... 14

14. Rostrum more strongly broadened towards apex, width at apex in male> 0.20 mm; scales on striae dense, imbricate, slightly raised and partly forming regular rows. Israel.................................. Ita friedmani sp. nov.

Rostrum scarcely broadened towards apex, width at apex in male 0.18 mm; scales uniformly distributed on whole elytral surface, without any trace of longitudinal rows. “ Syria ” (according to the political boundaries of the late nineteenth century).................................... Ita marina sp. nov.

Figures

Figure 1. Measurements. (A) Body length (1), rostrum length (2) and elytra length (3); (B) rostrum length from base to antennal insertion (1); (C) rostrum width at base (1); rostrum width at antennal insertion (2); rostrum width at apex (3).

Figure 2. Morphology of the genus Ita. (A) Aedeagus of I. hispanica; (B) male sternum VIII and IX of I. hispanica; (C) mouth parts of I. hispanica; (D) female sternum VIII of I. chavanoni; (E) hemisternites IX and styli of I. chavanoni; (F) spermatheca of I. chavanoni; (G) onychium, claws and (H) detail of claw of I. chavanoni. Scale bar: (A) 250 µm; (B–E; G) 100 µm; (F) 50 µm.