Habroloma ( Parahabroloma ) taxillusi Kato & Kawakita, 2023

Tamadera, Yutaka, 2025, New or little-known jewel beetles on mistletoe, Taxillus Tiegh. (Loranthaceae), in Japan: Unusual morphology and biology in Habroloma Thomson, 1864 (Coleoptera: Buprestidae), Zootaxa 5725 (1), pp. 55-88 : 67-75

Habroloma ( Parahabroloma) taxillusi Kato & Kawakita, 2023

[Japanese name: Ôbayadorigi-hirata-chibi-tamamushi]

Figs. 2C, D , 8B , 9–12 , 13C, D

Habroloma ( Parahabroloma) taxillusi : Kato & Kawakita, 2023a: 140 ( nomen nudum and unavailable name, according to Article 16.4.2 of ICZN, 1999; host: Taxillus yadoriki ).

Habroloma ( Parahabroloma) taxillusi Kato & Kawakita, 2023b: 196 (designation of holotype depository; type locality: “Yakukachi, Amami-shi, Kagoshima Pref. ”, Japan); Otobe, 2024: 52 (Honshu, Japan).

Additional description. Male. Body ovate, moderately convex dorsally ( Fig. 9C, D ). LB 2.77–2.96 mm (mean 2.87 mm); WB 1.68–1.79 mm (mean 1.74 mm); LB/WB 1.63–1.67 (mean 1.65) (n = 10 for all measurements except terminalia).

Integument above black, sometimes with weak golden-bronze or faint purplish tints; underside, antennae, and legs black, sometimes with faint golden-bronze reflections, except for tarsal pads pale brown. Dorsal and ventral surfaces moderately shiny. Vestiture mainly consisting of yellowish-brown, white, and black setae; yellowish-brown setae predominant, sometimes becoming more strongly yellowish or orangish ( Fig. 9H–M ). Head densely clothed with short, recumbent yellowish-brown setae. Pronotum clothed with short semirecumbent, yellowish-brown, white, and black setae which are arranged as follows: 1) four black round patches across middle, rather inconspicuous, with several white spots occurring around the patches; 2) yellowish-brown setae in remaining space, often becoming slightly lighter and denser apicolaterally. Elytra clothed with yellowish-brown, white, and black setae which are arranged on each elytron as follows: 1) yellowish-brown setae predominant, becoming denser around white bands and spots, occurring along sutural margin throughout; 2) white spots scattering in basal half; 3) first white transverse bands at apical 1/3, strongly wavy, zigzag-shaped, with setae relatively denser; 4) second white transverse band at subapical part, weakly wavy, with setae relatively denser; 5) one white patch at apical part; and 6) four to five inconspicuous black patches: one patch at just before basal 1/3, one or two patches just anterior to first transverse band, and the remaining one patch between the first and second transverse bands, the patches with very sparse setae, and sometimes with brownish tints. Underside sparsely setate, with fine, recumbent, whitish to yellowish setae, becoming slightly longer and denser on lateral parts of metacoxal plates.

Head, when viewed from above, relatively narrow in width, very shallowly, faintly bisinuately concave on frons, with oculofrontal margin ridged and very weakly produced anteriorly. Eyes, when viewed from above, narrowly visible, weakly convex laterally. Vertex coarsely variolate-punctate; frons widely, weakly concave, faintly grooved along upper side of midline, with surface coarsely variolate-punctate; supra-antennal pores round, distance between the pores distinctly a little larger than the diameter of the pore ( Fig. 10C ); clypeus extraordinarily narrow, without elevated basal margin and boundary with frons (as a result LC cannot be measured), WC/LSC 0.20–0.30 (mean 0.27) [LSC/WC 3.33–5.00 (mean 3.83)], weakly round at apical margin which is continuously arcuately connected with infra-antennal ridges, with disk strongly declivous apicad from narrowest part to subapical part, and then becoming flattened in apical part ( Fig. 10C ). Antennae ( Fig. 10D ) short, not reaching level of widest point of pronotum when laid laterally; scape stout claviform, slightly longer than pedicel; pedicel ovate, longer than antennomere 3; 3 subrectangular, slightly longer than 4; 4–6 subrectangular, subequal to each other in length or becoming longer toward apical antennomeres; 7–10 triangular, moderately enlarged; 11 subtriangular to sublingulate. Mouth parts and anteclypeus invisible in both ventral and frontal sides at rest.

Pronotum widest at base, WP/LP 3.05–3.30 (mean 3.13), BMP/AMP 2.07–2.26 (mean 2.16), almost as wide as elytra; lateral margins evenly, strongly arcuately narrowed apicad; apicolateral angles nearly right; basolateral angles broadly acute; apical margin moderately deeply, arcuately emarginate, with faint median lobe; basal margin trisinuate, with median lobe moderately produced; disk rather evenly convex, faintly depressed along basal median lobe, with ill-defined apicolateral depressions bearing small round pore at the middle of each, which is mostly hidden by dense pronotal setae; surface rather densely variolate-punctate, becoming sparse in apical half of median part except near apical margin. Scutellar shield small, triangular, glabrous.

Elytra widest at base, LE/WE 1.22–1.24 (mean 1.23), LE/LP 3.74–4.06 (mean 3.81); humeral calli weakly developed; lateral margins subparallel-sided in about basal 1/4, then weakly narrowed to around middle, then slightly more strongly narrowed to subapical part, and finally arcuately convergent to conjointly rounded apices, serrated with two to four minute denticles at subapical part ( Fig. 10B ); epipleura distinctly delimited by distinct marginal carinae occurring from base to subapical part; lateral carinae occurring from just behind humeri to subapical part, weakly sinuate behind basal parts; disk weakly depressed along lateral half of basal margin on each elytron, constricted behind humeri; surface irregularly, sparsely sculptured with round punctures and minute ones, of which the latter is associated with elytral vestiture. Hind wing as shown in Fig. 14C , with reduced anal field which is delimited by short anal fold, without anal embayment; veins MP 3, MP 4, and CuA 2 absent, but a short trachea branching from basal part of vein CuA 3+4 present ( Fig. 14D ).

Underside. Prosternum ( Fig. 10G ) with arcuate prosternal lobe rather straight in middle of apical margin which is weakly margined; prosternal process wide; trapezoidal plate as long as wide, WTP/LTP 1.33–1.53 (mean 1.42), WTP/BTP 1.52–1.67 (mean 1.58), with rounded apical corners, with sides which are arcuately dilated posteriorly from narrowest base to widest point around apical corners, with apex broadly arcuate; disk flattened on trapezoidal plate bearing distinctly arcuate marginal striae along sides, evenly convex on prosternal lobe, with a pair of deep, transverse grooves above procoxal cavities; surface on prosternal lobe finely strigulate except posterior part, and on trapezoidal plate with setiferous pin-prick punctures and ocellate sculptures. Hypomera with transversely oval hypomeral markings consisting of linearly incised punctures ( Fig. 10F ). Metaventrite moderately variolate-punctate on median portion surrounded by katepisternal suture and moderately variolate on outside of the median portion. Legs with femora and tibiae stout; metacoxal plates with an arcuate notch on each medial posterior margin, weakly produced posteriorly at each lateral angle which is broadly acute and rounded, with surface moderately variolate; metafemora obtusely angulate at about distal 1/3 of each outer margin; metatibiae without a fringe of spines on each outer margin, but with sparse short setae throughout the margin; inner tooth of each claw small. Abdominal ventrites with sternal groove only on ventrite 5, which has round apex; surface variolate, ventrite 1 with linearly confluent sculptures near median part.

Male terminalia (n = 5 for measurement). Sternite 9 ( Fig. 11A ) rather wide, SL/SW 0.81–0.93 (mean 0.86), round at apical margin, without setae. Tegmen ( Fig.11B ) rather wide; parameres PL/PW 2.79–3.10 (mean 2.94), bearing a pair of setae on apicolateral margins, with sides which are subparallel-sided in basal 1/5, then gradually dilated to widest part at apical 1/4, then weakly narrowed to subapical part, and finally arcuately convergent to apices, with rather shallow dorsal notch and deeper ventral notch; phallobase PbL/PbW 1.75–2.11 (mean 1.93), more than 1/5 length of tegmen. Penis ( Fig. 11C ) wide, PeL/PeW 3.93–4.72 (mean 4.44), shorter than tegmen; dorsal plate with sides (not including sides of median struts) subparallel in basal half, and then convergent apicad, with distinct broad apex which is roundly truncate, basally with median struts more than 1/3 length of penis, laterodorsally with enlarged sclerotized part in basal 1/3 of ventral plate to basal median struts; ventral plate membranous, but weakly sclerotized as a linear patch in middle of apical 2/4.

Female. Sexual dimorphism: Abdominal ventrite 5 with broad apical comb consisting of 15–21 short blunt denticles which become longer medially, the comb facing downwards ( Figs. 9F, G , 10H, I ); antennae with serrate antennomeres which are slightly smaller than male ( Fig. 10E ). Measurements (n = 10): LB 2.81–3.20 mm (mean 3.02 mm); WB 1.74–1.91 mm (mean 1.84 mm); LB/WB 1.58–1.72 (mean 1.64); WC/LSC 0.24–0.33 (mean 0.29) [LSC/WC 3.00–4.20 (mean 3.52)]; WP/LP 3.06–3.41 (mean 3.2); BMP/AMP 2.12–2.28 (mean 2.19); LE/WE 1.19– 1.25 (mean 1.22); LE/LP 3.66–4.14 (mean 3.89); WTP/LTP 1.40–1.54 (mean 1.46); WTP/BTP 1.52–1.68 (mean 1.58).

Female terminalia (n = 4 for measurement). Ovipositor ( Fig. 11D–F ) short in external part; proctiger with a pair of short, straight baculi at base; coxites transversely subtrapezoidal, with arcuate apical margin, sparsely setate on each side of apical margin with rather long setae; styli subtriangular, short, wide, SlL/SlW 1.52–2.33 (mean 1.90), bearing several rather long setae apically; ventral valve without setae; vagina sack-shaped (under incompletely inflated state), expanded near opening of spermatheca; spermatheca membranous, rather wide tubular, rather short in length, widest at about apical 2/3, without spermathecal gland.

Differential diagnoses. Habroloma taxillusi is readily distinguished from other congeners by the following combination of characters: 1) clypeus prominently narrowed, about 3.5 times as long as wide, with round apical margin; 2) elytral punctures round, which are not associated with setae; and 3) female abdominal ventrite 5 with apical comb (absent in male).

In Japan, H. taxillusi is somewhat similar in general appearance to Habroloma ( Parahabroloma) nixilla insulicola Kurosawa, 1959 , and their distribution ranges overlap at Amami-Ôshima Island in the Ryukyus, Japan. However, H. taxillusi is distinguished from H. nixilla insulicola by the diagnostic character states mentioned above and by the following two points: trapezoidal plate wider, about 1.4 times as wide as long; and host association with T. yadoriki ( H. nixilla insulicola : Lagerstroemia subcostata Koehne var. subcostata [ Lythraceae ]).

Specimens examined. Holotype ( ♀, see the Remarks for sex; missing abdomen). “ NIPPON: Kagoshima-ken | Yakukachi.Amami-shi | 23-V-2009. A. Kawakita | ex. Habroloma taxillusi [!] [white card; printed] || 09. 5. 23. YKC | オオバヤドリギ LM→ | BM. in. p. 6. 2 e [white card; handwritten] || Holotype | Habroloma taxillusi | des. Kato & Kawakita, 2023 [red card; printed] || k39 [pale yellow card; handwritten] || NSMT-I-C-200348 | National Museum of | Nature and Science | MK-BP-k40 [white card; printed]”.

Paratype ( 1 ♂, figured in Kato & Kawakita 2023a: figs. 1F, 2F, 3F; missing left hind leg). “ NIPPON: Kagoshima-ken | Yakukachi. Amami-shi | 23-V-2009. A. Kawakita | ex. Habroloma taxillusi [!] [white card; printed] || 09. 5. 23. YKC | オオバヤドリギ BM | 6. 7 e [white card; handwritten] || Paratype | Habroloma taxillusi | des. Kato & Kawakita, 2023 [yellow card; printed] || a328 [pale yellow card; handwritten] || k40 [pale yellow card; handwritten] || NSMT-I-C-200349 | National Museum of | Nature and Science | MK-BP-k39 [white card; printed]” .

Non type specimens ( 29 ♂♂, 28 ♀♀). JAPAN: [Honshu: Mie]— 6 ♂♂, 2 ♀♀, Kobune, Kiwa-chô, Kumanoshi, 4.XI.2023, H. Fukutomi leg., captured on Taxillus yadoriki (HFCI; YTJ); [Honshu: Wakayama]— 1♂ ( Fig. 9M ), Higashi-kônogawa, Minabe-chô, Hidaka-gun, 7.X.2023, Y. Tamadera leg., captured on T. yadoriki parasitizing Quercus phillyreoides A.Gray [ Fagaceae ] (YTJ); [Kyushu: Ôita]— 3 ♂♂ ( Fig. 9L ), Kamaeura, Kamae, Saiki-shi, 23.VII.2023, Y. Tsutsumiuchi leg., captured on T. yadoriki (YTJ) ; [Kyushu: Miyazaki]— 8 ♂♂ ( Fig. 9J ), 14 ♀♀, Heiwadai-kôen, Shimokitakata-chô, Miyazaki-shi, 16, VII.2023, Y. Tamadera leg., captured on T. yadoriki (YTJ) ; 1 ♂, ditto, emerged on 29.VII.2023 from T. yadoriki , TaY-136 (YTJ); 1 ♀, ditto, emerged on 2.VIII.2023 from T. yadoriki , TaY-136 (YTJ); 1 ♂, ditto, emerged on 5.VIII.2023 from T. yadoriki , TaY-136 (YTJ); 1 ♂, ditto, emerged on 9.VIII.2023 from T. yadoriki , TaY-136 (YTJ); 2 ♂♂ ( Figs. 2C , 9A, C, E ), 6 ♀♀ ( Figs. 2D , 9B, D, E, G ), same locality, 25.VII.2023, Y. Tamadera leg., captured on T. yadoriki (YTJ) ; 1 ♀ ( Fig. 9K ), Aya-minami-gawa, Aya-chô, Higashimorokata-gun, 25.VII.2023, Y. Tamadera leg., emerged on 8.VIII.2023 from T. yadoriki parasitizing Castanopsis sieboldii , TaY-145 (YTJ); [Ryukyus: Ôsumi Islands: Yakushima Is.]— 1 ♂, 1 ♀, Miyanoura-rindô, Miyanoura, Yakushima-chô, Kagoshima-ken, 25.VII.2023, T. Saeki leg., captured on T. yadoriki parasitizing Styrax japonicus Siebold et Zucc. [ Styracaceae ] (YTJ); [Ryukyus:Amami Islands:Amami-Ôshima Is.]— 3 ♂♂ ( Fig.9I ), 1 ♀, Kuba, Tatsugô-chô, Kagoshima-ken, 9.V.2023, H. Fukutomi leg. (HFCI; YTJ); 1 ♀ ( Figs. 8B , 9H ), near Nagakumotôge, Tatsugô-chô, Kagoshima-ken, 25.IX.2023, Y. Tamadera leg., captured on T. yadoriki parasitizing Castanopsis sieboldii subsp. lutchuensis (Koidz.) H.Ohba [ Fagaceae ] (YTJ); 1 ♀, Aminoko, Setouchi-chô, Kagoshima-ken, 8.V.2023, H. Fukutomi leg. (HFCI); 2 ♂♂, Amurogama, Setouchi-chô, Kagoshima-ken, 16.VII.2023, T. Saeki leg., captured on T. yadoriki (YTJ) .

Distribution. Japan: Honshu, Kyushu, and Ryukyus (Yakushima Is. [Ôsumi Islands]; Amami-Ôshima Is. [Amami Islands]). See also Fig. 15 .

Biology. Host plant. Loranthaceae : Taxillus yadoriki . Adults are leaf surface feeders; that is, they feed on the leaves from the adaxial surface. Characteristic feeding scars are left on the surface ( Figs. 12M , 13D ).

The adults were collected on T. yadoriki parasitizing the following various trees: Cryptomeria japonica (L.f.) D.Don [ Cupressaceae ]; Cinnamomum camphora (L.) J.Presl [ Lauraceae ]; Castanopsis sieboldii subsp. sieboldii , C. sieboldii subsp. lutchuensis , Quercus glauca , and Q. phillyreoides ; and Styrax japonicus .

Leaf-mining habit. Mines of H. taxillusi were found on Taxillus yadoriki parasitizing Castanopsis sieboldii subsp. sieboldii , C. sieboldii subsp. lutchuensis , and Quercus glauca by the author (in Wakayama, Miyazaki, and Amami-Ôhsima Is.; Figs. 12 , 13C ). Infested leaves were relatively young and sometimes still covered with trichomes on either side of the adaxial or abaxial surfaces ( Fig. 12C ). Each mine was of the full-depth type and formed a linear-blotch occurring along the leaf-margin, but it is almost invisible from the abaxial side of the leaves because the abaxial leaf surface of T. yadoriki was always covered with dense brownish trichomes. When full-grown, the mine occupied less than 1/4 of whole the leaf blade. Eggs, which were ochreous (remaining eggshell + varnish-like substances) and covered with trichomes of the host leaf ( Fig. 12E ), were laid singly on the adaxial surface of the leaf blade near the base (n = 20; Figs. 12A, C , 13C ) or occasionally near the apex (n = 6; Fig. 12F–I ), with one exception in which the egg was laid singly on the middle along the leaf-margin (n = 1). Each hatched larva linearly mined along the leaf-margin in the periods of first to second instars, then the last (third) instar makes a blotch mine and finally a delta-like portion at the terminal part of the blotch mine ( Fig. 12B, F–I ). The first and second instars left granular frass inside mines, and then the last instar made an oval pupal chamber inside the center of the blotch mine with its granular frass ( Fig. 12K, L ). The frass pupal chamber was made simultaneously while feeding on the parenchymal tissue, thus some posterior segments of the larval abdomen was usually toward the center of the blotch mine ( Fig. 12J ), and when full grown, the pupal chamber was completely closed. Adults always emerged from the abaxial surface of the leaves with oval exit holes ( Fig. 12D ). See also Kato & Kawakita (2023a).

Hibernating individuals have never been found in H. taxillusi , but probably the adults hibernate in various gaps at least in mainland Japan.

Remarks. Kato & Kawakita (2023a) described H. taxillusi as a new species but failed to mention the type depository for the holotype. This is therefore a nomen nudum and unavailable name, according to Article 16.4.2 of ICZN (1999). Later, Kato & Kawakita (2023b) designated the type depository for the holotype, making the name available.

This study found mistakes of labeling in the holotype and paratype of H. taxillusi .According to Kato & Kawakita (2023b), the holotype of this species was designated on the basis of a male individual, and one paratype female individual as follows:

“ Holotype: JAPAN: ♂ (MK-BP-k40), Yakukachi, Amami-shi , Kagoshima Pref., NSMT-I-C-200348” ;

“ Paratype: JAPAN: ♀ (MK-BP-k39), same data as holotype, NSMT-I-C-200349”.

Additionally, Kato & Kawakita (2023a), in which the name H. taxillusi was a nomen nudum, shows the data as follows:

“ Holotype: JAPAN: ♂ (MK-BP-k40), Yakukachi, Amami-shi , Kagoshima Pref. ( 28.228°N, 129.347°E, 40 m above sea level), 23-V-2009 (as larva on Taxillus yadoriki collected by A. Kawakita), emerged on 7-VI-2009, NSMT-I-C-200268” ;

“ Paratype: JAPAN: ♀ (MK-BP-k39), same data as holotype, emerged on 2-VI-2009, N NSMT-I-C-200269” .

However, the author confirmed that the actual specimen pinned with a holotype label is a female (NSMT-I-C-200348) and the specimen pinned with a paratype label is a male (NSMT-I-C-200349) (see Specimens examined). Moreover, all habitus images of this species in Kato & Kawakita (2023a: figs. 1F, 2F, 3F) are definitely based on the male specimen, but the figure captions do not state that the specimen illustrated is the holotype. These facts suggest that the type labels and depository serial number labels for the holotype and paratype were reversed. The labels remain as they were originally placed and were not moved as a result of this study.

This study also found an error in the original description of H. taxillusi in Kato & Kawakita (2023a). These authors described the clypeus of this species as follows: “clypeus transverse, ~ 2.6× as wide as long, with the anterior margin somewhat arcuately emarginate”. These character states, however, do not match those of the examined specimens in this study (see Additional description). Habroloma taxillusi has a prominently narrowed clypeus (with round apical margin; Fig. 10C ). This study, thus, concludes that the original description of the clypeus is in error. This is almost certainly a copy and paste from the corresponding part of the description of Habroloma ( Parhabroloma) elaeocarpusi , which is also described in Kato & Kawakita (2023a), because identical wording was used for the clypeus description in both H. taxillusi and H. elaeocarpusi . Examination of the type specimens shows that the clypeus description of Kato & Kawakita (2023a) applies only to H. elaeocarpusi .

The apical comb of the female abdominal ventrite 5 in H. taxillusi ( Figs. 9F , 10H, I ) is not described in Kato & Kawakita (2023a). This character state was confirmed in all the non-type female specimens in this study, although the author could not examine the abdomen of the female type specimen owing to the missing abdomen.

Figures

FIGURE 2. Dorsal habitus of mistletoe-associated Habroloma spp. A, B: Habroloma (Parahabroloma) tsutsumiuchii sp. nov. (A, male holotype; B, female paratype, Miyazaki); C, D: H. (P.) taxillusi Kato & Kawakita (C, male, Miyazaki; D, female, Miyazaki). Scale bar = 1.00 mm.

FIGURE 8. Elytral punctures in Habroloma spp. Black triangular marks indicate several non-setiferous punctures.A: Habroloma (Parahabroloma) tsutsumiuchii sp. nov. [paratype: Nara, Honshu]; B: H. (P.) taxillusi Kato & Kawakita [Amami-Ôshima Is., Ryukyus]; C: H. (P.) eximium eximium (Obenberger) [Miyazaki, Kyushu]. Scale bars = 0.25 mm.

FIGURE 9. Habitus of Habroloma (Parahabroloma) taxillusi Kato & Kawakita. A, B: ventral view, male (A) and female (B) [Miyazaki, Kyushu]; C, D: lateral view, male (C) and female (D) [Miyazaki, Kyushu]; E–G: apex of ventrite 5, caudal view (E, male; F, female) and lateral view (G, female, a black triangular mark indicates an apical comb); H–M: variations in body and setae coloration [H, I, Amami-Ôshima Is., Ryukyus; J, K, Miyazaki, Kyushu; L, Ôita, Kyushu; M, Wakayama, Honshu]. Scale bars = 1.00 mm for A–D, H–M; = 0.25 mm for E, F.

FIGURE 10. Habroloma (Parahabroloma) taxillusi Kato & Kawakita [Amami-Ôshima Is., Ryukyus]. A: Outline of head, pronotum, and elytron; B: serration of elytral lateral margin, obliquely ventral view; C: clypeus and its adjacent parts, frontal view; D: male antenna, outer side; E: female antenna, outer side; F: hypomeron (part), oblique view; G: prosternum; H, I: apical comb of female abdominal ventrite 5, caudal view (H) and ventral view (I). Abbreviations: hm, hypomeral marking; ir, infra-antennal ridge; sg, sternal groove; sp, supra-antennal pore; tp, trapezoidal plate (see the Materials & methods for its definition).

FIGURE 11. Male and female terminalia (part) of Habroloma (Parahabroloma) taxillusi Kato & Kawakita [Amami-Ôshima Is., Ryukyus]. A: Male sternite 9, dorsal view; B: tegmen, dorsal view; C: penis, ventral view; D, E: external part of ovipositor omitting proctiger, dorsal view (D) and ventral view (E); F: internal part of ovipositor, laterodorsal view. Abbreviations: co, common oviduct; cx, coxites; dp, dorsal plate; ms, median strut (s); pa, parameres; pb, phallobase; spm, spermatheca; st, stylus; ts, tegminal strut; vg, vagina; vn, ventral notch; vp, ventral (membranous) plate.

FIGURE 12. Biology of Habroloma (Parahabroloma) taxillusi Kato & Kawakita. A: full-grown mine, TaY-021 [Amami- Ôshima Is., Ryukyus]; B: blotch part of A (a black triangular mark indicates the delta-like portion), TaY-021; C: a mine on a younger leaf, TaY-136; D: adult exit hole, abaxial side of leaf [Miyazaki, Kyushu]; E: egg (indicated by a black triangular mark), TaY-487; F–I: growing process of blotch area of a mine from F to I, TaY-136 (a black triangular mark in J indicates the delta-like portion); J: a third instar larva in the mine, TaY-136; K, L: a frass pupal chamber (K, adaxial side of leaf, TaY-021; L, horizontal section, TaY-487); M: an adult feeding on adaxial leaf surface [Miyazaki, Kyushu]. Scale bars: B, J–M = 3 mm; E = 0.5 mm.

FIGURE 13. Taxillus yadoriki leaves infested by Habroloma spp. A: Full-grown four mines of Habroloma (Parahabroloma) tsutsumiuchii sp. nov., with five eggs, four adult exit holes, and adult feeding scars (indicated by black triangular marks) [Nara, Honshu]; B: two mines of H. (P.) tsutsumiuchii sp. nov., with two eggs [Nara, Honshu]; C: mine of H. (P.) taxillusi Kato & Kawakita, with a single egg [Amami-Ôshima Is., Ryukyus]; D: adult feeding scar of H. (P.) taxillusi [Miyazaki, Kyushu]. Asterisks (*) near A and C indicate the abaxial side of the infested leaf, respectively.

FIGURE 14. Hind wings. A, B: Habroloma (Parahabroloma) tsutsumiuchii sp. nov. [paratype; Nara, Honshu]; C, D: H. (P.) taxillusi Kato & Kawakita [Amami-Ôshima Is., Ryukyus]; B, D: trachea branching from basal part of vein CuA3+4, indicated by white triangular marks and traced above with white lines. Scale bars in B, D = 0.10 mm. Abbreviations: CAS, Cubitoanal Spur (vein CuP + AA3); MSP, Medial Spur; RML, Radiomedial Loop.

FIGURE 15. A distribution map of the two Habroloma species, with distribution ranges of the host mistletoe Taxillus yadoriki (broken line) and its Japanese relative T. pseudochinensis (solid line). Study sites in mainland Japan: a, b) Chiba; c) Kanagawa; d) Nara; e) Mie; f) Wakayama; g) Fukui; h) Hyôgo; i) Tottori; j) Kôchi; k) Ôita; l) Miyazaki.