Measurements (in mm). Body length: 2.20–8.70. Body width (metathorax or abdominal segments I–II) 0.73–2.44. Head width: 0.35–1.16.

General. Body elongated, slender, rounded in cross section.

Colouration. From yellow to pale brown head. All thoracic and abdominal segments from distinctly white to slightly yellow.

Vestiture. Setae on body thin, in different colouration, distinctly different in length; piliform, often with some asperities.

Head capsule. Head oval or suboval, slightly or more flattened laterally, endocarinal line present and very distinct, more than half the length of frons. Frontal sutures on the head in different sizes, and ever extended to antennae. One or two stemmata (st), anterior stemma in the form of a pigmented spot with convex cornea behind the antenna. Dorsum of the epicranium with five setae; des₃ located anteriorly on epicranium close border with frontal suture. Frons with four setae; fs₂ absent; fs₄; and fs₅ subequal. Head also with two les and two ves. Epicranial area with three pes and 2–3 sensilla.

Antennae located at the end of the frontal suture on each side, membranous and distinctly convex basal article bearing 3–4 sensilla and one conical sensorium, the later elongated, narrow.

Clypeus transverse-shaped, approximately 2.5–3 times as wide as long with two cls, and one sensillum (clss) between setae; all very close to margin with frons.

Mouthparts. Labrum with three piliform lms; anterior margin bisinuate. Epipharynx with three finger-like als; with 2–3 ams; and 0–1 mes; labral rods (lr) distinct, elongated. Mandibles distinctly broad, bifid, teeth of unequal height; slightly truncate; both setae piliform. Maxilla stipes with one stps, two pfs and one mbs and one sensillum; mala with six finger-like dms; five vms; all vms distinctly shorter than dms. Maxillary palpi with two palpomeres; basal palpomere with one short mxps and two sensilla; distal palpomere with one sensillum and a group of micro cuticular apical processes. Prelabium oval-shaped, with one prms; ligula with sinuate margin and 1–2 ligs; premental sclerite well sclerotized but without anterior and posterior extensions, U-shaped. Labial palpi with two palpomeres (partially appears as one palpomere); each of the palpomeres with one sensillum, distal palpomere with cuticular apical processes. Postlabium with three pms, all located laterally.

Thorax. Prothorax slightly smaller than meso- and metathorax. Spiracle bicameral, placed between the pro- and mesothorax (see, e.g., Skuhrovec et al. 2015). Prothorax with 9–10 prns; two ps; and one eus. Mesothorax with one prs, three pds; one as; two long and one short ss; one eps; one ps; and one eus. Chaetotaxy of metathorax almost identical to that of mesothorax. Each pedal area of thoracic segments well separated, with 5–6 pda.

Abdomen. Abdominal segments I–III of almost equal length, next abdominal segments decreasing gradually to the terminal parts of the body. Abdominal segment X reduced to four anal lobes of unequal size, the lateral lobes being distinctly the largest, the dorsal and the ventral lobes being very small. Anus located terminally. Eight spiracles, bicameral, all spiracles functional, close to the anterior margin. Abdominal segments I–VII with one prs; three pds, pds₂ the longest one; one long and one minute ss; two long eps; one ps; one lsts; and two eus. Abdominal segment VIII with one prs; 2–3 pds, if there are three setae, then pds₂ the longest one; one long and one minute ss; two long eps; one ps; one lsts; and two eus. Abdominal segment IX with four ds; 1–2 ps; and 1–2 sts. Abdominal segment X with one minute seta present or absent.

17 L3 larvae: 7 exx., 29.07.2010, Gródek ad Hrubieszów, CE Poland, leg. E. Szwaj, det. J. Łętowski; 10 exx., ex seed capsules of Campanula cervicaria L., 05.07.2017, Stara Planina, Babin Zub, east Serbia, leg. I. Toševski, all collected in association with adults det. R. Caldara. Accession numbers of sequenced specimens: MH558546.

Measurements (in mm). Body length: 4.43–5.57 (mean 4.90). Body width (metathorax or abdominal segments I–II) up to 1.37. Head width: 0.70–0.84 (mean 0.71).

General. Body elongated, slender, curved, rounded in cross section (Fig. 1).

Colouration. Black head (Fig. 1). All thoracic and abdominal segments from distinctly white to slightly yellow (Fig. 1).

Vestiture. Setae on body thin, light yellow to greyish, distinctly different in length (minute to very long).

Head capsule (Fig. 2). Head oval, slightly flattened laterally. Frontal sutures distinct, seem as pallid stripes. Anterior stemma (st), in the form of a small pigmented spot. Des_1–3 and des₅ long; des₄ short (Fig. 2). Fs₁ long; fs₂ absent; fs₃ very short; fs₄ long; and long fs₅ (Fig. 2). Les₁ and les₂ as long as des₅; both ves medium to very short. Epicranial area with three pes and two sensilla in line with des₂.

Antennae bearing one relatively elongated conical sensorium; and basal membranous article with four sensilla equal in length, and two pores (Fig. 3).

Clypeus (Fig. 5) approximately three times as wide as long with two cls of medium size, equal in length, and one sensillum; anterior margin sinuate.

Mouthparts. Labrum (Fig. 5) almost two times as wide as long, with three piliform lms, almost equal in the length; all located more or less anteromedially, lms₂ and lms₃distinctly reach labral margin. Epipharynx (Fig. 6) with three medium sized finger-like als, all similar in length; with two rather short, equal in length ams; and one medium size, finger-like mes; labral rods (lr) distinct, elongated, slightly convex. Mandibles (Fig. 4) bifid; cutting edge with a blunt tooth; bearing with two setae in medium size, piliform, and aligned longitudinally. Maxilla (Fig. 7) stipes with long stps and both pfs, minute mbs, and one sensillum close to mbs; mala with six medium sized finger-like dms; five vms, three medium size, two very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomeres with medium, cuticular apical processes; length ratio of basal and distal palpomeres 1:1. Prelabium (Fig. 7) with one short prms; ligula with one minute ligs; premental sclerite narrow, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:1.2; each of the palpomeres with one sensillum, distal palpomeres with medium, cuticular apical processes. Postlabium (Fig. 7) with long pms₁ located basally, very long pms₂ located medially and long pms₃ located apically; membranous area basolaterally sparsely and finely asperate.

Thorax. Prothorax (Fig. 8) with nine very long prns, weakly pigmented dorsal sclerite present with six long prns, this sclerite subdivided in two triangular plates medially; two long ps; and one short eus. Meso- and metathorax (Fig. 8) with one medium prs, three long pds; one long as; two very long and one minute ss; one long eps; one long ps; and one very short eus. Each pedal area of the thoracic segments with five very long pda.

Abdomen. Abdominal segments I–VII (Figs 9–10) with one medium prs; one long and two medium size pds (order: medium, long, medium); one very long and one minute ss; two very long eps; one medium ps; one medium lsts; and two very short eus.Abdominal segment VIII (Fig. 10) with one very short to minute prs; one short and two long to relatively long pds (order: short, long, relatively long); one long and one minute ss; two very long eps; one medium ps; one medium lsts; and two very short eus. Abdominal segment IX (Fig. 10) with four short ds; one medium ps; and two short sts. Abdominal segment X (Fig. 10) without seta.

This species lives on various species of Campanula (C. glomerata L., C. incurva Auch., C. latifolia L., C. persicifolia L., C. rapunculus L., C. rhomboidalis L., C. thyrsoides L., C. trachelium L.) in central Europe (Hoffmann 1958; Smreczyński 1976; Caldara and Legalov 2016). It was never reported to feed on C. cervicaria L., a species widely distributed in Europe. Larvae are seed feeders developing inside seed capsules.

This is one of the most variable species and with the widest Palaearctic distribution in the genus (Europe and central and northern Asia to the Russian Far East) (Caldara and Legalov 2016; Jiang et al. 2018). The three most variable characters in adults are the colour of the dorsal vestiture, which varies from whitish grey to light brown, the density of the elytral scales, which sometimes completely cover the integument, and the length of the rostrum, especially in the female and Anatolian populations. It is clear that it would be very interesting to perform a detailed molecular study of these populations. Apart from the characters of the shape of the rostra, the uncus of the male metatibiae and that of the penis, this species differs from C. graminis and related species also by the more angulate shape of the elytral base. Also the immatures of C. distinctus can easily be separated from those of C. graminis by several characters in larvae: postlabium with medium size pms₁ and pms₃, a very long pms₂ (Fig. 7) and a membranous area of postlabium basolaterally finely asperate as well as in pupae: Vs and sos absent (or as microsetae) (Fig. 82), pronotum with one sls (Fig. 83), and abdominal segments I–VII without ventral setae (Fig. 82). Finally, we could confirm that these two species are well separated molecularly as previously reported (Vahtera and Muona 2006; Hendrich et al. 2015; Horecka et al. 2017).

11 L3 larvae: 6 exx., 18.07.2010, Wólka ad Lublin, CE Poland, leg. E. Szwaj, det. J. Łętowski; 5 exx., ex seed capsules of Campanula macrostachya Waldst. et Kit. ex Willd., Dobra, Iron Gate, east Serbia. 13.07.2015, leg. I. Toševski, all collected in association with adults, det. R. Caldara. Accession numbers of sequenced specimens: MH558545.

Measurements (in mm). Body length: 3.75–6.27 (mean 4.80). Body width (metathorax or abdominal segments I–II) up to 1.63. Head width: 0.65–0.78 (mean 0.71).

General. Body elongated, slender, curved, rounded in cross section (Fig. 11).

Colouration. Pale brown head (Fig. 11). All thoracic and abdominal segments from distinctly white to slightly yellow (Fig. 11).

Vestiture. Setae on body thin, slightly from orange to pale brown, distinctly different in length (minute to very short or long to very long). Cuticle distinctly asperate.

Head capsule (Fig. 12). Head oval, slightly flattened laterally. Frontal sutures narrow, but distinct. Anterior stemma (st), in the form of a large pigmented spot. Des_1–3 and des₅ long; des₄ short to very short (Fig. 12). Fs₁ long; fs₂ absent; fs₃ very short; fs₄ long; and long fs₅ (Fig. 12). Les₁ and les₂ as long as des₅; both ves very short. Epicranial area with two sensilla and three minute pes in line with des₂.

Antennae bearing one medium size conical sensorium, and basal membranous article with three sensilla different in length, two behind conical sensorium, and one ahead of it (Fig. 13).

Clypeus (Fig. 15) approximately 2.5–3 times as wide as long with two short cls, cls₁ slightly shorter than cls₂, and one sensillum; anterior margin sinuate.

Mouthparts. Labrum (Fig. 15) less than two times as wide as long, with three piliform lms, different in the length; lms₁ located anteromedially, very close to margin of clypeus, lms₂ located in the middle, and lms₃ located anterolaterally; lms₁ and lms₂ of medium size, and lms₃ distinctly shorter than the previous two; only lms₂ distinctly reaches labral margin. Epipharynx (Fig. 16) with three long finger-like als, all of identical in length; with three ams in different length, ams₁ and ams₂ piliform of medium size, finger-like short ams₃ and enlarged in middle, and also located more close to lr; without mes; labral rods (lr) distinct, elongated, oval. Mandibles (Fig. 14) bifid; bearing with two setae in medium size, piliform, and aligned longitudinally, mds₁ located basally; mds₂, located distinctly apically. Maxilla (Fig. 17) stipes with very long stps and pfs₂, medium pfs₁, very short to minute mbs, and sensillum close to mbs; mala with six medium sized finger-like dms; five vms, different in length, three setae medium size, and two setae very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with some cuticular apical processes; length ratio of basal and distal palpomeres 1:0.8. Prelabium (Fig. 17) with one short prms; ligula with two very short to minute ligs; premental sclerite broad, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.8; each of the palpomeres with one sensillum, distal palpomere with cuticular apical processes. Postlabium (Fig. 17) with short pms₁ located basally, very long pms₂ located medially and short pms₃ located apically; membranous area basolaterally distinctly asperate.

Thorax. Prothorax (Fig. 18) with nine very long and one very short to minute prns, small pigmented dorsal sclerite present with five long prns, this sclerite subdivided in two triangular plates medially; two very long to long ps; and one short eus. Meso- and metathorax (Fig. 18) with one long prs, three very long pds; one very long as; two very long and one very short to minute ss; one very long eps; one very long ps; and one short to very short eus. Each pedal area of the thoracic segments with 5–6 very long pda.

Abdomen. Abdominal segments I–VII (Figs 19–20) with one medium prs; one short and two very long to long pds (order: short, very long, long); one very long and one minute ss; two long eps; one very long ps; one long lsts; and two short to very short eus. Abdominal segment VIII (Fig. 20) with one very short to minute prs; one short and two long to relatively long pds (order: short, long, relatively long); one long and one minute ss; two very long eps; one long ps; one long lsts; and two short to very short eus. Abdominal segment IX (Fig. 20) with three relatively long and one short to very short ds; one relatively long and sometimes one minute ps; and one relatively long to short and one short to very short sts. Abdominal segment X (Fig. 20) with one very short seta (ts).

Larvae were collected while feeding on the seeds of several species of Campanula, mainly C. glomerata, C. persicifolia, and C. rotundifolia L. (Hustache 1932; Hoffmann 1958; Smreczyński 1976; Lohse and Tischler 1983; Caldara and Legalov 2016) without producing galls. The species was not previously reported on Campanula macrostachya Waldst. and Kit. ex Willd., a taxon distributed from Ukraine along the Balkans until Anatolia. Pupae, as well as immatures of M. ajugae, were also collected on Adenophora liliifolia (L.) A. DC, although in another Serbian locality (see below). This genus, however, is very closely related to Campanula (Cano-Maqueda and Talavera 2011).

This is a very common and variable species with a wide European and Asian distribution from the Iberian Peninsula to eastern China (Caldara and Legalov 2016; Jiang et al. 2018). The two most variable characters in adults are the colour of the dorsal vestiture, which varies from whitish grey to light brown, and the density of the elytral scales, sometimes completely covering the integument. The rostrum varies somewhat in length and curvature, especially in the female. Cleopomiarus graminis is very closely related to C. longirostris as demonstrated by our data on the molecular fragment COI (I Toševski, unpublished data). Therefore, the differences between these two taxa found in the study of the immature stages, especially in the larvae – antennae with a very long conical sensorium and three sensilla (Figs 13, 33), dorsal setae (except des₄) long (Figs 12, 32), prothorax with nine very long and one very short to minute prns (Figs 18, 38) – are very important in order to confirm the specific rank of both taxa. On the other hand, the larva of C. longirostris is distinctly longer than the larva of C. graminis. With regard to the differences from C. distinctus, another widespread sympatric species sometimes confused with C. graminis, see the Remarks for the former taxon.

11 L3 larvae: south-eastern France, Menton, July 2007, ex capsules of Campanula trachelium L., leg. and det. R. Caldara, all determined by association with reared adults.

Measurements (in mm). Body length: 6.60–8.70 (mean 8.3). Body width (abdominal segments I–III) up to 2.44. Head width: 1.05–1.16 (mean 1.10).

General. Body elongated, slender, curved, rounded in cross section (Fig. 21).

Colouration. Pale brown head with indistinct pattern around frontal sutures (Fig. 21). All thoracic and abdominal segments from distinctly white to slightly yellow (Fig. 21).

Vestiture. Setae on body thin, orange, distinctly different in length (minute to very short or long to very long). Cuticle slightly asperate.

Head capsule (Fig. 22). Head oval, slightly flattened laterally. Frontal sutures medium width, distinct. Two pairs of stemmata (st), anterior one in the form of a large pigmented spot; and posterior one in form of a very small pigmented spot, located on each side close des₅. Des_1–3 long; des₄ short and des₅ long to very long (Fig. 22). Fs₁ long; fs₂ absent; fs₃ short; fs₄ long; and long fs₅ (Fig. 22). Les₁ and les₂ as long as des₅; both ves very short. Epicranial area with three pes and two sensilla in line with des₂.

Antennae bearing one medium size conical sensorium, and basal membranous article with four sensilla different in length, three behind conical sensorium, and one ahead of it (Fig. 23).

Clypeus (Fig. 25) approximately 2.5 times as wide as long with two short cls, cls₂ distinctly longer than cls₁, and one sensillum.

Mouthparts. Labrum (Fig. 25) less than 2.5 times as wide as long, with three piliform lms, different in the length; lms₁ located anteromedially, close to margin, lms₂ located in the middle, and lms₃ located posterolaterally; lms₁ and lms₂ of medium size, and lms₃ distinctly shorter than the previous two; only lms₂ distinctly reaches labral margin. Epipharynx (Fig. 26) with three long finger-like als, two als of identical in length, and the third one distinctly shorter and also located more close to labral rods (lr); with three ams in different length, ams₁ and ams₂ piliform and short, finger-like ams₃ and enlarged in middle, and also located more close to lr; without mes; labral rods (lr) distinct, elongated, oval. Mandibles (Fig. 24) bifid; mds₁ relatively long, piliform, located basally; mds₂ medium size, piliform, located distinctly apically and laterally. Maxilla (Fig. 27) stipes with very long stps and both pfs; very short to minute mbs, and sensillum close to mbs; mala with six medium sized finger-like dms; five vms, different in length, three setae medium size, and two setae very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with, cuticular apical processes; length ratio of basal and distal palpomeres 1:0.8. Prelabium (Fig. 27) with one relatively long prms; ligula with two very short to minute ligs; premental sclerite broad, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.8; each of the palpomeres with one sensillum, distal palpomere with short, cuticular apical processes. Postlabium (Fig. 27) with very short pms₁ located basally, very long pms₂ located medially and short to medium size pms₃ located apically; membranous area basolaterally distinctly asperate.

Thorax. Prothorax (Fig. 28) with eight very long to long and one very short to minute prns, small pigmented dorsal sclerite present with four long prns, this sclerite subdivided in two triangular plates medially; two very long to long ps; and one short to very short eus. Meso- and metathorax (Fig. 28) with one long prs, three medium to long pds; one very long to long as; two very long and one very short to minute ss; one long eps; one long ps; and one short to very short eus. Each pedal area of the thoracic segments with 5–6 very long to long pda.

Abdomen. Abdominal segments I–VII (Figs 29–30) with one short prs; one long and two short to very short pds (order: short, long, short); one long and one minute ss; two very long to long eps; one relatively long ps; one short lsts; and two very short eus. Abdominal segment VIII (Fig. 30) with one very short to minute prs; one relatively long and two very short pds (order: very short, long, very short); one relatively long and one minute ss; two relatively long eps; one short ps; one short lsts; and two very short eus. Abdominal segment IX (Fig. 30) with three relatively long and one very short to minute ds; one relatively long and sometimes one minute ps; and two short to very short sts. Abdominal segment X (Fig. 30) with one very short seta (ts).

We can confirm that larvae of this species feed on seed capsules of Campanula trachelium L., where they pupate without producing galls. It is noteworthy that adults did not exit by making a hole in the capsules but remained inside with the rostrum folded in the ventral canal until these opened spontaneously and forcefully, blowing up the seeds. On the other hand, it would be impossible, especially for the female, to straighten up the very long rostrum inside the capsule due to the limited available space. This is a more advantageous behaviour and apparently opposite to that of Rhopalapion longirostre (Olivier, 1807), another species where the female rostrum is more than twice as long as the stout male rostrum. In this species, Wilhelm et al. (2011) argued that the long rostrum is presumably an advantage for this weevil because its larvae can feed on plant parts with high energy density into buds (i.e., pollen grains) and that natural selection favours rostrum elongation. However, these authors reported that the elongated rostrum of females also bears a high risk when metamorphosed weevils attempt to leave their site of pupal development, which is the dry seed chambers, and therefore mortality during escaping may counteract selection for rostrum elongation, thus placing a limit on rostrum exaggeration. It is noteworthy that R. longirostre does not possess a ventral canal, which allows it to retain the folded rostrum.

This species is only known from France, Italy, and Switzerland, where it is quite common. The adult is very closely related to C. graminis, as also supported by preliminary molecular studies (I Toševski, unpublished data), from which it differs only by the very long rostrum especially in the female and usually by the larger size (Caldara and Legalov 2016). Therefore, the larval differences between these two taxa, in C. longirostris antennae bearing one medium size conical sensorium and four sensilla (Fig. 23), dorsal setae (except des₄) extremely long (Fig. 22), prothorax with eight very long and one very short to minute prns (Fig. 28), are very important since they allow easy separation of these two species.

13 L3 larvae, ex seed capsules of Campanula lingulata Waldst. and Kit., 26.06.2017, Staničenje, Pirot, east Serbia, leg. I. Toševski, all collected in association with adults, det R. Caldara. Accession numbers of sequenced specimens: MH558547.

Measurements (in mm). Body length: 5.10–7.30 (mean 5.67). Body width (metathorax or abdominal segments I–II) up to 2.02. Head width: 0.83–0.96 (mean 0.91).

General. Body elongated, slender, weakly curved, rounded in cross section (Fig. 31).

Colouration. Pale brown or almost yellow head (Fig. 31). All thoracic and abdominal segments from white to slightly yellow (Fig. 31).

Vestiture. Setae on body thin, slightly from orange to brown, distinctly different in length (minute to very short or long). Cuticle distinctly asperate.

Head capsule (Fig. 32). Head suboval. Frontal sutures distinct. Endocarina distinctly widened in the middle of the length. Two small stemmata (st), located close to des₅. Des_1–2 and des₅ very long; des₃ medium size; des₄ short (Fig. 32). Fs₁ long; fs₂ absent; fs₃ medium; fs₄ long; and fs₅ very long (Fig. 32). Les₁ and les₂ as long as des₅; both ves very short. Epicranial area with two sensilla and three pes in line with des₂.

Antennae bearing one very long conical sensorium, and basal membranous article with three sensilla almost equal in length (Fig. 33).

Clypeus (Fig. 35) approximately 4.25 times as wide as long with two almost equal in length cls: cls₂ some longer than cls₁, and one sensillum; anterior margin sinuate.

Mouthparts. Labrum (Fig. 35) two times as wide as long, with three piliform lms, different in the length; lms₁ and lms₂ located medially, and lms₃ located anterolaterally; lms₁ very long and reaches distinctly the labrum margin, lms₂ long, and lms₃ short, three times as shorter than lms₁. Epipharynx (Fig. 36) with three medium sized finger-like als, and three medium to short ams in different shape; labral rods (lr) distinct, elongated. Mandibles (Fig. 36) bifid; bearing with two setae in short to medium size, piliform, and aligned longitudinally, mds₁ located basally; mds₂, located distinctly apically. Maxilla (Fig. 34) stipes with long stps and equal in length pfs₁ and pfs₂, very short to minute mbs, and two sensilla close to mbs; mala with six medium sized finger-like dms; five vms, different in length, three setae medium size, and two setae very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with cuticular apical processes; length ratio of basal and distal palpomeres 1:0.8. Prelabium (Fig. 37) with one short prms; ligula with two short ligs; premental sclerite broad, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.9; each of the palpomeres with one sensillum, distal palpomere with short, cuticular apical processes. Postlabium (Fig. 37) with short pms₁ located basally, very long pms₂ located medially and short pms₃ located apically; membranous area basolaterally distinctly asperate.

Thorax. Prothorax (Fig. 38) with nine long and one very short prns; two long ps; and one short eus. Meso- and metathorax (Fig. 38) with one short prs, three long pds; one long as; two long and one very short to minute ss; one long eps; one long ps; and one short eus. Each pedal area of the thoracic segments with six different in length pda.

Abdomen. Abdominal segments I–VII (Figs 39–40) with one very short prs; two short and one long pds (order: short, long, short); one long and one minute ss; two long eps; one long ps; one long lsts; and two short eus. Abdominal segment VIII (Fig. 40) with one very short prs; three pds (order: short, long, short); one long and one minute ss; two long eps; one long ps; one long lsts; and two short eus. Abdominal segment IX (Fig. 40) with one medium long and three very short ds; two short ps; and two short sts. Abdominal segment X (Fig. 40) without seta.

Previously, the unique biological datum on this species was reported by Weill et al. (2011), who collected adults in Syria on Michauxia campanuloides L’Hér., a small genus of Campanulaceae distributed in the Middle East, possibly a synonym of Campanula (Crowl et al. 2014). Therefore, the observation that this species feeds on Campanula lingulata Waldst. and Kit. is unpublished. Moreover, adults were recently observed feeding on flowers of Campanula sibirica L. in eastern Serbia (I Toševski, pers. obs.), but larval development on this plant species is not confirmed. Like C. distinctus and C. graminis, larvae are seed feeders inside capsules of the host plant without producing galls.

This species was previously known from Anatolia, Syria and many countries of the Balkans but not from Serbia. The adults of this species are characterized by a very long rostrum in the females. This character, however, is not uncommon in the Palaearctic Cleopomiarus. For example, this character is shared with C. longirostris and C. distinctus, two other taxa presented in this paper. It is distinguishable from these species by the less globose and moderately elongate elytra, and moreover by the shape of the male and female genitalia (Caldara and Legalov 2016). Other characters of the immatures allow easy separation of C. medius from these two species as well as from C. graminis (see keys to larvae and pupae). There are also substantial molecular differences between C. medius and other species (I Toševski, unpublished data).

10 L3 larvae: south-eastern France, Castellar (Menton), Juin 2005, ex seed capsules of Campanula rapunculus L., leg. and det. R. Caldara all collected in association with adults.

Measurements (in mm). Body length: 2.20–3.15 (mean 2.8). Body width (metathorax or abdominal segments I–II) up to 0.73. Head width: 0.35–0.51 (mean 0.45).

General. Body elongated, slender, curved, rounded in cross section (Fig. 41).

Colouration. Pale brown or almost yellow head (Fig. 41). All thoracic and abdominal segments from distinctly white to slightly yellow (Fig. 41).

Vestiture. Setae on body thin, slightly from orange to pale brown, distinctly different in length (minute to very short or long to very long). Cuticle distinctly asperate.

Head capsule (Fig. 42). Head suboval, distinctly flattened laterally. Frontal sutures narrow, but distinct. Two stemmata (st), anterior one in the form of a small pigmented spot; and posterior one in form of a very small pigmented spot, located on each side close des₅. Des_1–3 and des₅ very long; des₄ relatively long (Fig. 42). Fs₁ long to very long; fs₂ absent; fs₃ long medium, laterally to fs₄; fs₄ very long; and fs₅ very long (Fig. 42). Les₁ and les₂ as long as des₅; both ves medium size. Epicranial area with two sensilla and three pes in line with des₂.

Antennae bearing one very long conical sensorium, and basal membranous article with three sensilla different in length, two behind conical sensorium, and one ahead of it (Fig. 43).

Clypeus (Fig. 45) approximately three times as wide as long with two medium size cls, cls₁ distinctly longer than cls₂, and one sensillum; anterior margin sinuate.

Mouthparts. Labrum (Fig. 45) less than 2.5 times as wide as long, with three piliform lms, different in the length; lms₁ located posteromedially, very close to margin of clypeus, lms₂ located in the middle, and lms₃ located laterally; lms₁ very long and reaches distinctly the labrum margin, lms₂ long, and lms₃ medium size, more than twice times as short as lms₁. Epipharynx (Fig. 46) with three medium sized finger-like als, two als of identical in length, and the third one distinctly shorter and also located close to labral rods (lr); with three short ams in different shape, ams₁ and ams₂ piliform, finger-like ams₃ and enlarged in middle, and also located more close to lr; without mes; labral rods (lr) distinct, elongated, oval. Mandibles (Fig. 44) bifid; bearing with two setae in short to medium size, piliform, and aligned longitudinally., mds₁ located basally; mds₂, located distinctly apically. Maxilla (Fig. 47) stipes with very long stps and pfs₂, medium size pfs₁, very short to minute mbs, and sensillum close to mbs; mala with six medium sized finger-like dms; five vms, different in length, four setae medium size, and one seta very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with short, cuticular apical processes; length ratio of basal and distal palpomeres 1:0.8. Prelabium (Fig. 47) with one very short prms; ligula with two very short to minute ligs; premental sclerite broad, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.9; each of the palpomeres with one sensillum, distal palpomere with short, cuticular apical processes. Postlabium (Fig. 47) with short pms₁ located basally, long pms₂ located medially and short pms₃ located apically; membranous area basolaterally only a partly and finely asperate.

Thorax. Prothorax (Fig. 48) with nine very long and one very short to minute prns, small pigmented dorsal sclerite present with four long prns, this sclerite subdivided in two triangular plates medially; two very long to long ps; and one short eus. Meso- and metathorax (Fig. 48) with one long prs, three very long to long pds; one long as; two very long and one very short to minute ss; one long eps; one very long to long ps; and one short to very short eus. Each pedal area of the thoracic segments with 5–6 very long pda.

Abdomen. Abdominal segments I–VII (Figs 49–50) with one long prs; two relatively long to short and one very long to long pds (order: relatively long, very long, short); one very long to long and one minute ss; two very long eps; one very long to long ps; one relatively long to short lsts; and one short to very short and one relatively long eus. Abdominal segment VIII (Fig. 50) with sometimes one very short to minute prs; one short and one long to relatively long pds (order: short, long); one long and one minute ss; two very long eps; one very long to long ps; one relatively long to short lsts; and one short to very short and one relatively long eus. Abdominal segment IX (Fig. 50) with two relatively long and two short to very short ds; one relatively long and one minute ps; and one relatively long sts. Abdominal segment X (Fig. 50) with one very short seta (ts).

Adults of this species are usually collected on the flowers of Campanula rapunculus L., and we can confirm that larvae feed on seeds of this plant as previously reported by Hoffmann (1958).

This species is widely distributed and common in southern Europe, whereas it appears rare in North Africa and the Middle East. Adults can be confused with some related species such as C. plantarum (Germar, 1823), C. micros (Germar, 1821) and C. reitteri (Caldara & Legalov, 2016), from which they differ by some external characters and the shape of their genitalia (Caldara and Legalov 2016). In contrast, this species is poorly related morphologically to the other species of Cleopomiarus studied here. This difference is confirmed also by the larval morphology, which differs from all of the other species mainly by a longer fs₃ that is almost as long as fs₄.

Measurements (in mm). Body length: 3.80–8.39. Body width (metathorax or abdominal segments I–II) 1.55–2.04. Head width: 0.57–0.83.

General. Body slender, C-curved, rounded in cross section.

Colouration. From black to dark brown head. All thoracic and abdominal segments yellowish, with some asperities.

Vestiture. Setae on body thin, in different colouration, distinctly different in length; piliform.

Head capsule. Head almost rounded, sometimes slightly flattened laterally, endocarinal line present and distinct, more than half the length of frons. Frontal sutures on the head narrow and loosened, but distinct, and ever extended to the antennae. One stemma (st), in the form of a pigmented spot with convex cornea. Dorsum of the epicranium with four or five setae; des₃ located anteriorly on epicranium close border with frontal suture. Frons with three or four setae; fs₂ absent. Head also with two les and two ves. Epicranial area with two or three pes and more or without sensilla.

Antennae located at the end of the frontal suture on each side, membranous and distinctly convex basal article bearing one very long conical sensorium; basal membranous article with 1–4 sensilla.

Clypeus transverse-shaped, approximately 2.5–3.5 times as wide as long with two cls, and one sensillum (clss) between setae; all very close to margin with frons.

Mouthparts. Labrum with three piliform lms; anterior margin bisinuate. Epipharynx with three finger-like als; with two ams; and 0–2 mes; labral rods (lr) elongated. Mandibles distinctly broad, bifid, teeth of unequal height; slightly truncate; both setae piliform and located apically. Maxilla stipes with one stps, two pfs and one mbs and one sensillum; mala with six finger-like dms, in two sizes, first or first and second dms very long as pfs, next medium length; five vms; all vms distinctly shorter than dms. Maxillary palpi with two palpomeres; basal palpomere with one short mxps and two sensilla; distal palpomere with one sensillum and a group of micro cuticular apical processes. Prelabium oval-shaped, with one prms; ligula with sinuate margin and 2–3 ligs; premental sclerite feebly visible. Labial palpi with two palpomeres; each of the palpomeres with one sensillum, distal palpomere with cuticular apical processes. Postlabium with three pms, all located laterally.

Thorax. Prothorax slightly smaller than meso- and metathorax. Spiracle bicameral, placed between the pro- and mesothorax (see, e.g., Skuhrovec et al. 2015). Prothorax with ten prns; two ps; and one eus. Mesothorax with one prs, three pds; one as; two long and one short ss; one eps; one ps; and one eus. Chaetotaxy of metathorax almost identical to that of mesothorax. Each pedal area of the thoracic segments well separated, with 5–6 pda.

Abdomen. Abdominal segments I–III of almost equal length, next abdominal segments decreasing gradually to the terminal parts of the body. Abdominal segment X reduced to four anal lobes of unequal size, the lateral lobes being distinctly the largest, the dorsal and the ventral lobe very small. Anus located terminally. Eight spiracles, bicameral, all spiracles placed medially or anteromedially and functional. Abdominal segments I–VIII with one prs (sometimes abdominal segment VIII without); three pds, pds₂ the longest one; one long and one minute ss; two long eps; one ps; one lsts; and two eus. Abdominal segment IX with 3–4 ds; 1–3 ps; and two sts. Abdominal segment X with one minute seta present or absent.

5 L3 larvae: north-eastern Italy, Venezia Giulia, Duino (Trieste), Rilke path, August 2017, ex galls on capsules of Campanula pyramidalis L., leg. E. Tomasi, all collected in association with adults, det. R. Caldara.

Measurements (in mm). Body length: 3.50–4.75 (mean 3.9). Body width (abdominal segment II) up to 1.65. Head width: 0.57–0.65 (mean 0.60).

General. Body moderately elongated, rather stout, curved, rounded in cross section (Fig. 51).

Colouration. Almost black head (Fig. 51). All thoracic and abdominal segments from greyish-white to yellowish; prodorsum with brownish dorsal sclerite; all abdominal segments covered with fine spiculation (Fig. 51).

Vestiture. Setae on body thin, brown, rather short or minute, piliform.

Head capsule (Fig. 52). Head oval, slightly flattened laterally. Endocarinal line present and very distinct. Stemma placed below des₅. Des_1–3 and des₅ long; des₄ medium size (Fig. 52). Fs₁ long; fs₂ absent; fs₃ and fs₄ medium size; and fs₅ long (Fig. 52). Les₁ and les₂ as long as des₅; one ves very short. Epicranial area with three pes (in line with des₂), and also two sensilla.

Antennae located at the end of the frontal suture on each side, membranous and distinctly convex basal article bearing one conical sensorium, relatively elongated; basal membranous article with four sensilla (styloconica) equal in length, and one (ampullacae) (Fig. 53).

Clypeus (Fig. 55) trapezium-shaped, approximately 3.3 times as wide as long with two medium size, equal in length cls, and one sensillum (clss) between setae; all very close to margin with frons; anterior margin of clypeus rounded to inside.

Mouthparts. Labrum (Fig. 55) 2.5 times as wide as long, with three piliform lms, lms₁ twice longer than (equal in the length) lms₂ and lms₃; all located more or less anteromedially, all reach labral margin; anterior margin double sinuate. Epipharynx (Fig. 56) with three medium sized finger-like als, all similar in length; with two rather short, different in length ams; and one medium size, finger-like mes; labral rods (lr) distinct, kidney-shaped. Mandibles (Fig. 54) distinctly broad, bifid, teeth of unequal height; slightly truncate; cutting edge with a blunt tooth; bearing with two setae in short size, piliform, and aligned longitudinally. Maxilla (Fig. 57): stipes with long stps, two long pfs, one minute mbs and two sensillae close to mbs; mala with six finger-like dms (first and second elongated, forth to sixth medium size); five vms (two medium size and three very short); all vms shorter than dms. Maxillary palpi with two palpomeres; basal palpomere with one short mxps and two sensilla; length ratio of basal and distal palpomeres almost 1:1; distal palpomere with one sensillum and a group of microcuticular processes apically. Prelabium (Fig. 57) oval-shaped, with one medium prms; ligula with sinuate margin and three minute ligs; premental sclerite narrow, ring-shaped, well visible. Labial palpi with one palpomere (partially seems as two palpomere); palpomere with one sensillum and medium, cuticular apical processes. Postlabium (Fig. 57) with three pms, all located laterally; pms₁ and pms₃ short, pms₂ medium size; membranous area basolaterally finely asperate.

Thorax. Prothorax smaller than meso- and metathorax. Spiracle bicameral, placed between the pro- and mesothorax. Prothorax (Fig. 58) with ten prns (two minute and eight long), well pigmented dorsal sclerite with four long prns); two medium ps; and one short eus. Meso- and metathorax (Fig. 58) with one short prs, three medium pds; one medium as; two medium and one minute ss; one medium eps; one medium ps; and one very short eus. Chaetotaxy of metathorax (Fig. 58) almost identical to that of mesothorax. Each pedal area of the thoracic segments with six medium length pda (four of them placed on well-separated pedal areas, next two setae outside).

Abdomen. Abdominal segments I–VII (Figs 59, 60) with one short prs; three medium size pds (equal in length); one medium and one minute ss; two medium eps; one medium ps; one medium lsts; and two very short eus. Abdominal segment VIII (Fig. 60) without prs; three medium pds; one medium and one minute ss (sometimes absent); two medium eps; one medium ps; one medium lsts; and two very short eus. Abdominal segment IX (Fig. 60) with three medium ds; one relatively long ps; and two short sts. Abdominal segment X (Fig. 60) without seta.

The only detailed biological data are reported by Tomasi (2002), who observed that this species lives in Friuli-Venezia Giulia (Italy) on Campanula pyramidalis L., where larvae cause a distinct swelling of the calix of the flowers, which remain closed.

This species has a well-delimited distribution (south-eastern Poland, Austria, north-eastern Italy, Slovenia, Croatia, Serbia, Montenegro, Macedonia). It is easily distinguishable from all other species of Miarus by the shape of the body of the penis, which is characterized by the presence of two lateral flanges at its apex. However, for the external morphology, the M. abnormis adults are very similar to several other species, such as M. ajugae and M. campanulae, from which they can be distinguished only by the characters of the male ventrite five (fovea less deep, teeth less robust). Unfortunately, the females of these three species appear not to be distinguishable (Caldara 2007), and the molecular fragment COI poorly differentiates these species. Therefore, the differences between the immatures of these species are much important for the separation of these three species. According to the larval morphology, M. abnormis appears more closely related to M. ajugae than to M. campanulae due to several features (mala with six finger-like dms, different in length: two setae elongated, and four setae of medium length; epipharynx with 1–2 mes, and finally des₄ and fs₃ present), confirming what was suggested by the adult morphology (Caldara 2007). Moreover, larvae of M. abnormis differ from other Miarus species here studied mainly by an epipharynx with one finger-like mes.

26 L3 larvae: 9 exx., 12.07.2009, Bychawa ad Lublin, CE Poland, leg. E. Szwaj, det. J. Łętowski; 12 exx, ex galls on capsules of Adenophora liliifolia, 30.06.2017, Kaludjerske Bare, Mt. Tara, Central Serbia, leg. I. Toševski, det. R. Caldara; 5 exx, ex galls on capsules of Campanula bononiensis L., 14.07.2017, Zavojsko jezero, Pirot, east Serbia, leg. I. Toševski, all collected in association with adults, det. R. Caldara. Accession numbers of sequenced specimens: MH558548.

Measurements (in mm). Body length: 4.50–8.39 (mean 5.70). Body width (metathorax or abdominal segments I–II) up to 2.04. Head width: 0.68–0.83 (mean 0.70).

General. Body slender, C-curved, rounded in cross section (Fig. 61).

Colouration. Head dark brown to black (Fig. 61). All thoracic and abdominal segments yellowish with fine speculation, with clearly separated dark pigmented pedal areas (Fig. 61).

Vestiture. Setae on body very thin, piliform, distinctly different in length (minute to very short or long to very long).

Head capsule (Fig. 62). Head almost rounded. Frontal sutures narrow and loosened, but distinct. One stemma (st), in the form of a large pigmented spot. Des_1–3 and des₅ long; des₄ very short (Fig. 62). Fs₁ long; fs₂ absent; fs₃ short; fs₄ and fs₅ long (Fig. 62). Les₁ and les₂ as long as des₅; both ves very short. Epicranial area with three very short pes and also three sensilla.

Antennae bearing one very long conical sensorium, and basal membranous article with three sensilla and one pore (Fig. 63).

Clypeus (Fig. 65) approximately 3.5 times as wide as long with two short, almost equal in length cls, and one sensillum between them; anterior margin sinuate.

Mouthparts. Labrum (Fig. 65) 1.6 times as wide as long, with three piliform lms, rather equal in length; lms₁ located medially, lms₂ located anteromedially, and lms₃ located anterolaterally; all of them reaches labral margin. Epipharynx (Fig. 66) with three long finger-like als, all of identical in length; with two medium size ams; and two mes, first finger-like, second sharp and more slender; labral rods (lr) elongated, broad, slightly convergent posteriorly. Mandibles (Fig. 64) bifid; cutting edge with small blunt tooth; bearing with two setae in medium to long size, piliform, located apically and aligned longitudinally. Maxilla (Fig. 67) stipes with long stps and both pfs, very short to minute mbs, and one sensillum close to mbs; mala with six finger-like dms, different in length: first and second very long, forth to sixth medium size; five vms, different in length, three setae medium size, and two setae very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with cuticular apical processes; length ratio of basal and distal palpomeres 1:0.9. Prelabium (Fig. 67) with one medium prms; ligula with two minute ligs; premental sclerite narrow, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.7; each of the palpomeres with one sensillum, distal palpomere with cuticular apical processes. Postlabium (Fig. 67) with short pms₁ located basally, long pms₂ located medially and short pms₃ located apically; membranous area without any asperities.

Thorax. Prothorax (Fig. 68) with ten prns (nine very long and one minute), small pigmented dorsal sclerite present with five long prns, this sclerite subdivided in two triangular plates medially; two long ps; and one short eus. Meso- and metathorax (Fig. 68) with one medium prs, three long pds; one very long as; two long and one minute ss; one long eps; one long ps; and one short eus. Chaetotaxy of metathorax (Fig. 68) almost identical to that of mesothorax. Each pedal area of the thoracic segments with six very long pda.

Abdomen. Abdominal segments I–VII (Figs 69–70) with one medium prs; two medium and one long to very long pds (order: medium, very long, medium); one very long and one minute ss; two long eps; one medium ps; one medium lsts; and two short eus. Abdominal segment VIII (Fig. 70) with one short prs; two short and one long pds (order: short, long, short); one long and one minute ss; two medium eps; one medium ps; one medium lsts; and two short eus. Abdominal segment IX (Fig. 70) with two relatively long and two short ds; two different in length ps; and two short sts. Abdominal segment X (Fig. 70) with one very short seta (ts).

Miarus ajugae was collected on various species of the genus Campanula (C. carpathica Jacq., C. glomerata L., C. latifolia L., C. macrorrhiza Gay ex DC, C. media L., C. patula L., C. persicifolia L., C. rapunculoides L., C. rapunculus L., C. rhomboidalis L., C. rotundifolia L., C. trachelium L.) and Phyteuma (P. orbiculare L., P. spicatum L.) (Caldara 2007). However, it was never reported previously to feed on Campanula bononiensis L. and Adenophora liliifolia (L.) A. DC. (see also Biology of C. graminis).

This species with large Palaearctic distribution (from France and north-western Africa along all Europe to the Far East) is very closely related to M. campanulae, from which it differs mainly by the shape of the apex of the body of the penis (Caldara 2007). Unfortunately, molecular studies on the fragment COI revealed poor differences between these two species (Vahtera and Muona 2006; Hendrich et al. 2015, I. Toševski, unpublished data). Therefore, the consistent differences which we found in larval morphology between C. ajugae and C. campanulae are very important in order to confirm the validity of these two taxa at species level. Larvae of M. ajugae differ from all other species mainly by an epipharynx with two mes, first finger-like, second sharp and slender.

9 L3 larvae: 30.07.1939, ex Campanula, Store Dyrehave, Denmark, leg. J.P. Kryger, collected in association with adults, det. Van Emden, coll. British Museum of Natural History (London).

Measurements (in mm). Body length: 3.80–5.96 (mean 5.20). Body width (metathorax or abdominal segments I–II) up to 1.61. Head width: 0.58–0.64 (mean 0.61).

General. Body slender, C-curved, rounded in cross section (Fig. 71).

Colouration. Brown to dark brown head (Fig. 71). All thoracic and abdominal segments yellowish with distinct asperation (Fig. 71).

Vestiture. Setae on body thin, slightly from dark orange to brown, distinctly different in length (minute to very short or long to very long) distinct asperate.

Head capsule (Fig. 72). Head almost round, very slightly flattened laterally. Frontal sutures narrow and loosened, but distinct. One stemma (st), in the form of a large pigmented spot. Des_1–3 and des₅ long; des₄ absent (Fig. 72). Fs₁ long; fs₂ and fs₃ absent; fs₄ and fs₅ long (Fig. 72). Les₁ and les₂ as long as des₅; both ves short. Epicranial area with two pes.

Antennae bearing one very long conical sensorium, and basal membranous article with four sensilla different in length, three behind conical sensorium, and one ahead of it (Fig. 73).

Clypeus (Fig. 75) approximately 2.5–3 times as wide as long with two short to very short cls, localized posterolaterally, cls₁ slightly longer than cls₂, and one sensillum between them; anterior margin sinuate.

Mouthparts. Labrum (Fig. 75) less than two times as wide as long, with three piliform lms, different in the length; lms₁ located medially, lms₂ located anteromedially, and lms₃ located anterolaterally; lms₁ long, lms₂ medium size, and lms₃ distinctly shorter than the previous two; no one distinctly reaches labral margin. Epipharynx (Fig. 76) with three long finger-like als, all of identical in length; with two ams in different length, ams₁ piliform of medium size, and finger-like short ams₂; without mes; labral rods (lr) indistinct, slightly elongated, oval. Mandibles (Fig. 74) bifid; bearing with two setae of medium to long size, piliform, both located apically. Maxilla (Fig. 77) stipes with very long stps and pfs₁, long pfs₂, very short to minute mbs, and one sensillum close to mbs; mala with six finger-like dms, different in lengths: first very long (as long as pfs₁), next medium size; five vms, different in length, two setae medium size, and three setae very short. Maxillary palpi: basal palpomere with one short mxps and two sensilla; distal palpomere with short, cuticular apical processes; length ratio of basal and distal palpomeres 1:0.9. Prelabium (Fig. 77) with one short prms; ligula with two very short to minute ligs; premental sclerite narrow, ring-shaped. Labial palpi with two palpomeres; length ratio of basal and distal palpomeres 1:0.8; each of the palpomeres with one sensillum, distal palpomere with short, cuticular apical processes. Postlabium (Fig. 77) with short pms₁ located basally, long pms₂ located medially and short pms₃ located apically; membranous area without any asperities.

Thorax. Prothorax (Fig. 78) with nine very long and one very short to minute prns, small pigmented dorsal sclerite present with seven long prns, this sclerite subdivided in two triangular plates medially; two very long to long ps; and one medium eus. Meso- and metathorax (Fig. 78) with one medium prs, three very long pds; one very long as; two very long and one very short to minute ss; one long eps; one long ps; and one medium to long eus. Each pedal area of the thoracic segments with 5–6 very long pda.

Abdomen. Abdominal segments I–VII (Figs 79–80) with one medium to long prs; two medium to short and one very long to long pds (order: medium/short, very long, medium); one very long and one minute ss; two long eps; one medium to short ps; one medium to short lsts; and one medium to short and one very short to minute eus. Abdominal segment VIII (Fig. 80) with one very short prs; two short and one long to relatively long pds (order: short, long, short); one long and one minute ss; two very long eps; one medium to short ps; one medium to short lsts; and one medium to short and one very short to minute eus. Abdominal segment IX (Fig. 80) with two relatively long and two short to very short ds; two relatively long and sometimes one minute ps; and one relatively long to short and one short to very short sts. Abdominal segment X (Fig. 80) with one very short seta (ts).

Larvae live and pupate in the capsules of several species of Campanula (C. cochleariifolia Lam., C. patula L., C. persicifolia L., C. rapunculoides L., C. rapunculus L., C. rotundifolia L., C. scheuchzeri Vill. C. trachelium L.), and Phyteuma spicata L. where they cause distinct swelling (Hoffmann 1958; Buhr 1964)

This species has a wide European distribution and is very similar to M. ajugae. Only the examination of the penis allows easy separation of these two taxa. Therefore, as reported in the Remarks of C. ajugae, the discovery of clearly distinctive characters between the larvae of these two species is extremely important. Moreover, larvae of M. campanulae differ from the two other species studied mainly by an epipharynx without mes, and des₄ and fs₃ absent.

Measurements (in mm). Body length: 3.00–6.50. Body width: 1.50–3.80. Head width: 0.65–1.10.

Body moderately slender or stout. Smooth, dark brown or black spotted cuticle. Rostrum long or very long, from four up to five times as long as wide, reached up to meso- or metacoxae. Antennae elongated. Pronotum from 2.2 up to 2.9 times as wide as long. Mesonotum distinctly shorter than metanotum. Abdominal segments I–V of equal length; abdominal segments VI and VII diminish gradually; abdominal segment VIII almost semicircle; abdomen segment IX distinctly reduced. Spiracles on abdominal segments placed dorsolaterally: on abdominal segments I–V functional; and on segment VI atrophic on next ones invisible. Urogomphi (ur) stout and short, conical, each of them with sclerotized apex.

Chaetotaxy: setae piliform, in a different size. Head capsule with 0–1 vs, 0–1 sos, and two os. Rostrum with 0–1 pas and one rs (placed medially and apically). Pronotum with: two as, one ds, 1–2 sls, 0–1 ls and three pls. Setae on head, rostrum and pronotum very thin, light and relatively short. Dorsal parts of meso- and metathorax with 2–3 setae placed medially. Apex of each femora with one fes. Abdominal segments I–VIII with two setae laterally and sometimes 2–3 short setae ventrally. Dorsal parts of abdominal segments I–VII with 4–5 setae, and abdominal segment VIII with 3–4 setae dorsally. Abdominal segment IX with 2–4 micro-setae ventrally.

12 specimens: 2 ♂; 2 ♀, 03.08.2010, Gródek ad Hrubieszów, CE Poland, leg. E. Szwaj, det. J. Łętowski; 2 ♂; 6 ♀, ex seed capsules of Campanula cervicaria L., 05.07.2017, Stara Planina, east Serbia, leg. I. Toševski, det. R. Caldara.

Measurements (in mm). Body length: 3.00–4.10 (mean 3.20). Body width: 1.50–2.60 (mean 1.80). Head width: 0.65–0.77 (mean 0.67).

Body moderately slender (Figs 81–84). Rostrum long, approximately 4.0 times as long as wide, reached up to mesocoxae. Antennae moderately elongated. Pronotum 2.2 times as wide as long. Urogomphi (ur) stout (Figs 81–84).

Chaetotaxy: setae very thin, greyish, piliform, medium size to short. Head capsule with only two os of different in length (second pair placed on eye spots). Rostrum with one pas and one rs (Figs 82, 83). Pronotum with: two as, one ds, one sls, and three pls (Figs 83, 84). All setae of prothorax almost equal in length (Fig. 83). Dorsal parts of meso- and metathorax with three setae placed medially. Apex of each femora with one fes (Figs 82–84). Abdominal segments I–VIII with two setae laterally. Dorsal parts of abdominal segments I–VII with four setae: d₁–₃ placed postero-medially, d₄ postero-laterally, abdominal segment VIII with only three setae dorsally. Abdominal segment IX with two micro-setae ventrally.

15 specimens: 2 ♂, 3 ♀, 27.07.2010, Wólka ad Lublin, CE Poland, leg. E. Szwaj, det. J. Łętowski; 4 ♂, 2 ♀, ex seed pods of Adenophora liliifolia, Dobra, Iron Gate, east Serbia, leg. I. Toševski, det. R. Caldara; 4 ♀, ex seed pods of Campanula macrostachya, 13.07.2015, Dobra, Iron Gate, east Serbia, leg. I. Toševski, det. R. Caldara.

Measurements (in mm). Body length: 3.60–4.10 (mean 3.70). Body width: 2.10–2.25 (mean 2.15). Head width: 0.65–0.70 (mean 0.67).

Body moderately slender (Figs 85–88). Rostrum long, approximately four times as long as wide, reached up to mesocoxae. Antennae slender and elongated. Pronotum 2.9 times as wide as long. Urogomphi (ur) moderately stout (Figs 85–88).

Chaetotaxy: setae very thin, greyish, piliform, medium size to short. Head capsule with one vs, one sos, and two os of different length (second pair placed on eye spots). Rostrum with one pas and one rs (Figs 86, 87). Pronotum with: two as, one ds, two sls, one ls, and three pls (Figs 87, 88). All setae of prothorax equal in length (Fig. 88). Dorsal parts of meso- and metathorax with three setae placed medially. Apex of each femora with one fes (Figs 86, 87). Abdominal segments I–VIII with two setae laterally and three micro setae ventrally. Dorsal parts of abdominal segments I–VII with four setae: d₁–₃ postero-medially, and d₄ postero-laterally. Abdominal segment VIII with four setae dorsally. Abdominal segment IX with four micro-setae ventrally.

18 specimens: 8 ♂, 10 ♀, south eastern France, Menton, July 2007, ex seed capsules of Campanula trachelium L., leg. and det. R. Caldara

Measurements (in mm). Body length: 5.20–6.50 (mean 5.60). Body width: 3.00–3.80 (mean 3.30). Head width: 0.90–1.10 (mean 1.00).

Body stout (Figs 89–92). Rostrum very long, almost five times as long as wide, reached almost up to metacoxae. Antennae slender and moderately elongated. Pronotum 2.5 times as wide as long. Urogomphi (ur) short (Figs 90–92).

Chaetotaxy: sparse, setae very thin, short, piliform. Head capsule with one vs, one sos, and two os of different length (second pair placed on eye spots). Rostrum with one rs and one es (Figs 90, 91). Pronotum with: two as, one ds, one ls, one sls, and three pls (Figs 91, 92). All setae of prothorax equal in length (Fig. 92). Dorsal parts of meso- and metathorax with two setae placed medially. Apex of each femora with one fes (Figs 90–92). Abdominal segments I–VIII with two setae laterally and three medium long setae ventrally. Dorsal parts of abdominal segments I–VII with five setae: d₁ placed antero-medially, d₂–₄ postero-medially, d₅ postero-laterally. Abdominal segment VIII with only three setae dorsally. Abdominal segment IX with two micro-setae ventrally.

4 specimens: 2 ♂, 2 ♀, ex seed capsules of Campanula lingulata Waldst. and Kit., 26.06.2017, Staničenje, Pirot, east Serbia, leg. I. Toševski, det R. Caldara.

Measurements (in mm). Body length: 4.40–4.60. Body width: 2.40–2.50. Head width: 0.70–0.80.

Body moderately stout (Figs 93–96). Rostrum very long, approximately 4.5 times as long as wide, reached beyond mesocoxae. Antennae slender and moderately elongated. Pronotum 2.4 times as wide as long. Urogomphi (ur) stout (Figs 94–96).

Chaetotaxy: sparse, setae short to very short, light, piliform. Head capsule with one very short vs, and two os of different length, both placed between eye spots. Rostrum with one rs (Figs 94, 95). Pronotum with: two as, one ds, two sls, one ls and three pls (Figs 95, 96). All setae of prothorax equal in length (Fig. 96). Dorsal parts of meso- and metathorax with three setae placed medially. Apex of each femora with one fes (Figs 94, 95). Abdominal segments I–VIII with two setae laterally and three medium long setae ventrally. Dorsal parts of abdominal segments I–VII with four setae: d₁–₃ postero-medially, d₄ postero-laterally. Abdominal segment VIII with only three setae dorsally. Abdominal segment IX with four micro-setae ventrally.

Measurements (in mm). Body length: 3.20–4.00. Body width: 1.90–2.70. Head width: 0.55–0.70.

Body moderately slender or stout. Smooth, dark brown or black spotted cuticle. Rostrum long or very long, from 2.3 up to 3.5 times as long as wide, reached up to meso- or metacoxae. Antennae elongated. Pronotum from 2.1 up to 2.5 times as wide as long. Mesonotum distinctly shorter than metanotum. Abdominal segments I–V of equal length; abdominal segments VI and VII diminish gradually; abdominal segment VIII almost semicircle; abdomen segment IX distinctly reduced. Spiracles on abdominal segments placed dorsolaterally: on abdominal segments I–V functional; and on segment VI atrophic on next ones invisible. Urogomphi (ur) slender and elongated, conical, each of them with sclerotized apex.

Chaetotaxy: setae piliform, in a different size. Head capsule with one vs, one sos, and 3–4 os. Rostrum with one pas and three rs (placed medially and apically). Pronotum with: two as, two ds, two sls, one ls and three pls. All setae of prothorax almost equal in length. Dorsal parts of meso- and metathorax with three setae placed medially. Apex of each femora with one fes. Abdominal segments I–VIII with two setae laterally and sometimes 3–4 short setae ventrally. Dorsal parts of abdominal segments I–VII with 4–5 setae, and abdominal segment VIII with 3–4 setae dorsally. Abdominal segment IX with 4–6 micro-setae ventrally.

3 specimens: 1 ♂, 2 ♀, north-eastern Italy, Venezia Giulia, Duino (Trieste), Rilke path, August 2017, ex galls on capsules of Campanula pyramidalis L., leg. E. Tomasi, det. R. Caldara.

Measurements (in mm). Body length: 3.55–3.60. Body width: 1.90–2.00. Head width: 0.55–0.60.

Body moderately slender (Figs 97–100). The cuticle densely covered with asperities. Rostrum medium length, approximately 2.3 times as long as wide, reached up to mesocoxae. Antennae moderately elongated. Pronotum 2.1 times as wide as long. Urogomphi (ur) slender and elongated, conical, each of them with sclerotized apex (Figs 98–100).

Chaetotaxy: setae brownish to dark brown, piliform, short to medium size. Head capsule with one vs, one sos, and three os of equal length; rostrum with three rs (rs₁ and rs₂ placed medially, rs₃ more apically) and one es (Figs 98, 99). Pronotum with: two as, two ds, two sls, one ls and three pls (Figs 99. 100). Setae on head and rostrum slightly shorter than those on prothorax. All setae of prothorax almost equal in length (Fig. 100). Mesothorax with three setae placed medially; metathorax with three setae placed laterally. Apex of each femora with one fes (Figs 98–100). Abdominal segments I–VIII with two short, thin setae laterally and three short setae ventrally. Dorsal parts of each abdominal segments I–VIII with four setae: d₁–₃ placed postero-medially, d₄ postero-laterally. Abdominal segment IX with six microsetae ventrally.

12 specimens: 2 ♂, 1 ♀, 24.08.2009, Ciechanki, CE Poland, leg. E. Szwaj, det. J. Łętowski; 3 ♂, 2 ♀, 30.06.2017, ex galls on capsules of Adenophora liliifolia, 30.06.2017, Kaludjerske Bare, Mt. Tara, Central Serbia, leg. I. Toševski, det. R. Caldara; 3 ♂, 1 ♀, ex galls on capsules of Campanula bononiensis, 14.07.2017, Zavojsko jezero, Pirot, east Serbia, leg. I. Toševski, det. R. Caldara.

Measurements (in mm). Body length: 3.20–4.00 (mean 3.40). Body width: 1.90–2.70 (mean 2.10). Head width: 0.65–0.70 (mean 0.65).

Body rather stout (Figs 101–104). Rostrum moderately elongated, approximately 3.5 times as long as wide, reached almost up to metacoxae. Antennae rather stout and moderately elongated. Pronotum 2.5 times as wide as long. Urogomphi (ur) slender and elongated (Figs 102–104).

Chaetotaxy: setae piliform, greyish to dark brown, different in length, short to medium size. Head capsule with one vs, one sos, and four os of different length (fourth placed on eye spots). Rostrum with one pas, and three rs (rs₁ and rs₂ placed medially, rs₃ more apically) and one es (Figs 102, 103). Pronotum with: two as, two ds, one ls, two sls, and three pls (Figs 103, 104). All setae of prothorax equal in length (Fig. 104). Dorsal parts of meso- and metathorax with three setae placed medially. Apex of each femora with one fes (Figs 102, 103). Abdominal segments I–VIII with two setae laterally and four medium long setae ventrally. Dorsal parts of abdominal segments I–VII with five setae: d₁ placed antero-medially, d₂–₄ postero-medially, d₅ postero-laterally, and abdominal segment VIII with only three setae dorsally. Abdominal segment IX with four micro-setae ventrally.