Tonza citrorrhoa Meyrick, 1905

Kobayashi, Shigeki, Matsuoka, Haruka, Kimura, Masaaki, Sohn, Jae-Cheon, Yoshiyasu, Yutaka & Lees, David C., 2018, Designation of a new family group name, Tonzidae fam. nov., for the genus Tonza (Lepidoptera, Yponomeutoidea), based on immature stages of Tonza citrorrhoa, European Journal of Taxonomy 443, pp. 1-32 : 18-29

Tonza citrorrhoa Meyrick, 1905

Figs 1–13

Tonza citrorrhoa Meyrick, 1905: 614 – Heppner (1992: 74) (species list) – Kobayashi et al. (2015: 69–72, figs 1–2).

Diagnosis

See Kobayashi et al. (2015: 69) and diagnosis of the family.

Material examined (4 ³³, 7 ♀♀, 10 unsexed)

JAPAN:2³³, 5♀♀ adults, Ryukyus, Okinawa Prefecture,Mt.Kubura-dake,Yonaguni Is., 3–6May2016, M. Kimura leg. (host: Putranjiva matsumurae ), 22 Apr. 2016 (larva) (OPU-IN-LE 2018IV 0003–0009); material in NHMUK, see above, has not been critically re-examined; 2 ³³, 2 ♀♀, 3 unsexed, pupae, same data as for preceding, but May 2016 (OPU-IN-LE 2018IV 0018–0022); 7 unsexed larvae, same data as for preceding, but May 2016 (OPU-IN-LE 2018IV 0026–0028|SK600, 0025|SK603, 0026|SK604, 0035|SK543(exuvia)).

Type locality

SRI LANKA: Hantane.

DNA barcodes

GenBank accession no. LC 310893 , voucher no. SK-097. BIN: BOLD:ACX8102, voucher no. USNMENT00657266.

Additional description

ADULT. Wingspan range 11.0– 14.4 mm (mean 13.7 mm, n = 9): forewing length range 5.2–7.1 mm (mean 6.6 mm, n = 9).

MALE GENITALIA AND FEMALE GENITALIA. See Kobayashi et al. (2015: fig. 2).

MATURE LARVA ( FIGS 4–6 , 9 ). Range: 11.5–12.5 mm in length (mean 12.0 mm, n = 4). Integument varying from yellowish green to yellow in colouration, full mature larva yellow in colouration, marked with numerous purple blotches and white dots ( Fig. 4 ). Pinacula conspicuous, dark brown in colouration ( Fig. 4A, J ). Primary setae long and secondary setae absent ( Fig. 4 A–E).

HEAD ( FIG. 5 ). Hypognathous. Head capsule marked with numerous ochreous spots ( Fig. 5A, C ), 0.90– 1.00 mm in width (mean 0.94 mm, n = 5). Frontclypeus wide, extending a half to epicranial notch. Mandible about 0.15 mm in length, with three large teeth, two smaller teeth, and three small teeth at inner portion, with M 1 and M 2 setae much shorter ( Fig. 5F ). Six stemmata arranged in an arc except for S 1 and S 6; S 1 more posterior and S 6 ventrad ( Fig. 5 A–C).

CRANIAL SETAE ( FIG. 5 A–C). MD 1 very long; A 1, A 2 and A 3 as obtuse triangle with A 2 most distant from stemmata; P 1 below Af2– P 2 line; pore not found.

THORAX ( FIG. 6 A–B). T 1 shield indistinct, yellow in colouration, marked with dark brown patches. Jugular gland (adenosma) absent, but like structure present ventrally on T 1 ( Fig. 4F ), anterior to the legs and antero-medial to the long setae SV 1 and SV 2; the structure not eversible and a tubular or sacciform gland at inner side of body not found. D1 and D2 approximated on T 2 and T 3; SD 1 and SD 2 approximated on T 2 and T 3; L-group trisetose, L 1 and L 2 on the same pinacula on T 1 and separated on T 2 and T 3; SV 1 and SV 2 on the same pinacula on T 1 and on separate pinacula on T 2 and T 3; Seta V 1 present on T 1, absent T 2 and T 3. Thoracic legs pale ochreous to brown in colouration; pale brown claws elongate, slightly curved to inner side ( Figs 4F , 6B ).

ABDOMEN ( FIG. 6 C–F). D1 above level of D2 except for A 9; SD 2 very small, separated from pinaculum of SD 1; L-group trisetose on A 1–7, bisetose on A 8 and unisetose on A 9; L 2 minute; SV-group trisetose on A 3 and A 4, bisetose on A 5 and A 6, unisetose on A 1, A 2, and A 7– A 9; Seta D1 of A 9 segment markedly more slender than D2. Anal shield indistinct as in figure 6 F. Prolegs present on A 3, A 4, and A 10 ( Fig. 4A, G ). Ventral prolegs elongate, about 2× length of width of proleg base; crochets uniordinal, arranged in a circle, being usually 12 in number ( Figs 4 H–I, 9L–N). Crochets of the anal prolegs arranged in a semicircle ( Fig. 4K ).

PUPA ( FIGS 7, 10–12 ). General: long and slender, yellowish green in colouration, 10–11 mm in length, 1.0–2.0 mm in diameter. Maxillary palpi concealed ( Fig. 10E ). Dorsum of A 2– A 9 with two pairs of minute spiny setae from dorsal side ( Figs 7B, C, 11). Other characters as for family diagnosis.

Distribution

INDIA: Khasia/Khasi Hills; Darmsala, Punjab; Coimbatore (NHMUK): new record. SRI LANKA: Colombo, Kandy, Bogawantalawa, Puttalam, Maskeliya, Bentota, Gulla, Dondanduwa, Hikkaduwa, Nawalatipiya (NHMUK): new record, Hantane [Hanthana] ( Meyrick 1905). CHINA: Taiwan ( Heppner 1992; USNM, on BOLD). JAPAN: Kagoshima Prefecture: Amai-Ohshima Is. ( Seino 2016), Okinawa Pref.: Okinawa Is. ( Kobayashi et al. 2015), Iriomote Is. ( Umetsu 2016), Yonaguni Is.: [new record]. INDONESIA: Telawa, Java (NHMUK; abdomen missing, identity not certain but included among 13 identified specimens constituting Meyrick’s collection); Sulawesi (NHMUK), identity not certain. PHILIPPINES (NHMUK): identity not certain.

Hostplants

Putranjivaceae : Putranjiva matsumurae : new larval hostplant record. The adult has been recorded feeding on ‘Marygold flowers’ [sic; Tagetes L., Asteraceae Bercht. & J. Presl ] at Coimbatore, S. India (NHMUK).

Life history

The detailed biology of this species is unknown. The third author (Kimura) observed a number of late instar larvae on the young leaves of Putranjiva matsumurae . The larva is a leaf webber tying together several leaves loosely with silk threads ( Fig. 3 A–C). The full grown larva is suspended from the tree by a silk lifeline spun out from the head spinnerets. Pupation takes place at the intersection of some cross silken threads ( Fig. 3H ). A number of the mature larvae occurred on tall trees of the hostplant in Yonaguni Is., Okinawa Pref. ( Fig. 3A ). Young larvae were not found in our study. Given that no larval mine was observed, the young larvae are probably external feeders like the later instars.

Remarks

The resting posture of the adult moth with head end lowered and abdomen lifted is similar to certain Argyresthiidae , Ypsolophidae and several genera in Yponomeutidae ( Fig. 1B ). However, when at repose in nature the suspension may be different and rather unusual involving only the prothoracic and mesothoracic legs in contact with the lower surface of a leaf with the wing and body tending to hang vertically below a leaf. In this position, the antennae relatively long with respect to wing length may accentuate a potential false head-like appearance.

Leaf webbing larvae occur in several yponomeutoid families, Yponomeutidae , Scythropiidae , Plutellidae , Ypsolophidae (Ypsolophinae) and Attevidae ( Sohn et al. 2013) . However, larvae of Tonza form rather looser webs than other yponomeutoid leaf-webbers. We observed that the larvae move on silk threads using the claws of the thoracic legs and the prolegs, but cannot move on flat surfaces without silk.

ADULT Male genitalia

Molecular analysis

The COI DNA barcoding region (COI-5P) was sequenced from a Japanese specimen of Tonza . We performed sequence comparison to check species variation for T.citrorrhoa , using the BOLD Identification System (IDS) from the BOLD website (http://www.barcodinglife.org/) [accessed 1 Jul. 2016]. The DNA barcode sequence of T. citrorrhoa ( Fig. 2 , sample ID: SK-97) was clearly distinguished from that of T. purella registered in BOLD with more than 3% pairwise divergence ( Fig. 2 ) and to a sample from Madagascar belonging to BIN BOLD:ACU1104 by 2.91% pairwise divergence. The nearest neighbour of T. citrorrhoa in the BOLD database which is 0.35% pairwise divergent is a Taiwanese conspecific specimen ( Fig. 2 ), which belongs to the same BIN (BOLD:ACX8102).

Figure 13 shows a maximum likelihood tree of the DNA barcoding region from 23 species of Yponomeutoidea and Gracillarioidea Stainton, 1854, including our sequence, SK-97. This tree based on a single locus could not resolve all the higher-level relationships within the groups, except for Attevidae Mosher, 1916 (85% bootstrapping support). The traditional sense of Plutellidae , which would comprise Plutella Schrank, 1802 , Orthenches Meyrick, 1885 , and Tonza , among other taxa not represented here, did not form a monophyletic group. Two species, T. citrorrhoa and Glyphipterix equitella Scopoli, 1763 , were grouped together with 87% bootstrapping support.

Figures

Fig. 1. Female adult of Tonza citrorrhoa Meyrick, 1905 (OPU-IN-LE 2018IV0005), host: Putranjiva matsumurae Koidz., from Yonaguni Is., Okinawa Pref., Japan. A. Adult specimen. B. Resting posture of adult, dorso-lateral view. Scale bar: 2 mm.

Fig. 2. A tree estimated by neighbor-joining analysis under Kimura’s two-parameter distance model using DNA barcode sequence, retrieved from BOLD identification system (IDS). Branch lengths are proportional to distances. Sequence with ID started from “SK-” is obtained from the present study, and the others are from the BOLD database. The letters within square brackets refer to the collection site of specimens as follows: Au = Australia; Ca = Canada; Cr = Costa Rica; Ec = Ecuador; Eu = Europe; Gh = Ghana; Ja = Japan; Ma = Macedonia; Pa = Pakistan; Pg = Papua New Guinea; Sk = South Korea; Ta = Taiwan; Un = Unknown; Us = United States; Ve = Venezuela. Asterisks show the superfamily Yponomeutoidea.

Fig. 3. Larvae and pupae of Tonza citrorrhoa Meyrick, 1905 on Putranjiva matsumurae Koidz. A. Tree of hostplant and larval webs. B. Later larva forming web on young leaves. C. Later larva, lateral view. D. Mature larva, dorsal view. E. Same, ventral view. F. Pupa, ventral view. G. Same, lateral view. H. Pupa at the joint of some cross silken threads, lateral view.

Fig. 4. Mature larva of Tonza citrorrhoa Meyrick, 1905 (OPU-IN-LE2018IV0031|SK600). A. Gene- ral, lateral view. B. Same, ventral view. C. Head and prothorax, dorsal view. D. Same, ventral view. E. Same, lateral view. F. Jugular glands, lateral view. G. A3, 4. H. Proleg on abdominal segment 3. I. Same, on abdominal segment 4. J. Spiracle on abdominal segment 8. K. Anal proleg, ventral view. Scale bar: 1 mm.

Fig. 5. Mature larval head of Tonza citrorrhoa Meyrick, 1905 (OPU-IN-LE2018IV0031|SK600, 0025|SK603, 0026|SK604). A. Head capsule, frontal view. B. Same, rear view. C. Same, lateral view. D. Labrum, frontal view. E. Same, rear view. F. Mandible, inner view. Abbreviations: A1, A2, A3 = anterior setae; Af1, Af2 = adfrontal setae; Afa = pore between setae Af1 and Af2; at = antenna; C1, C2 = clypeal setae; F1 = frontal seta; Fa: pore on the frons; L1 = lateral seta; lb = labium; lp = labial palpus; M1, M2 = mandibular setae; MD1, MD2 = microdorsal setae; MG1 = microgenal seta; mp = maxillary palpus; P1, P2 = posteriodorsal setae; S1, S2, S3 = stemmatal setae; spr = spinneret; SS1, SS2, SS3 = substemmatal setae; stm = stemma.

Fig. 6. Mature larval thorax and abdominal segment of Tonza citrorrhoa Meyrick, 1905 (OPU-IN- LE2018IV0031|SK600, 0025|SK603, 0026|SK604). A. Thorax segment 1–3. B. Thoracic leg on thorax segment 1. C. A 1–4. D. A5–9. E. A9–10. F. Anal shield. Abbreviations: ans = anal shield; apl = anal proleg; Ba = claw, lateral view; Bb = same, frontal view; D1, D2 = dorsal setae; L1, L2, L3 = lateral setae; MD1 = microdorsal seta; MV3 = microventral seta; SD1, SD2 = subdorsal setae; sp = spiracle; SV1, SV2, SV3 = subventral setae; V1 = ventral seta; XD1, XD2 = XD group, dorsal setae on the anterior margin of the T1.

Fig. 9. SEMs of the mature larval structure of Tonza citrorrhoa Meyrick, 1905 (OPU-IN- LE2018IV0035|SK543). A. Mandible and maxillary palpus. B. Jugular gland-like structure. C. Prothoracic legs. D. Meso- and metathoracic legs. E. Right mesothoracic leg. F. Left mesothoracic leg. G. Same, base of claw. H. Right metathoracic leg. I. Same, base of claw. J. Left metathoracic leg. K. Same, claw and tarsus. L–M. Crochets of proleg on A3. N. Same on A4. O. Crochets of anal prolegs.

Fig. 10. SEMs of the pupal head and abdomen of Tonza citrorrhoa Meyrick, 1905 (OPU-IN-LE2018IV0018–0020). A. Frontal process, ventral view. B. Head, lateral view. C. Frontal process, dorsal view. D. Same, lateral view. E. Face, ventral view. F. A5, ventral view. G. Setae of abdominal tergum 6. H. Same, A7. I. A5–A7, dorsal view.

Fig. 13. Maximum Likelihood tree of 23 yponomeutoid/gracillarioid species based on the DNA barcoding region. Numbers on nodes indicate bootstrap support values. Tonza citrorrhoa Meyrick, 1905 is indicated in bold.

Tables

Table 2. (Continuation.)

PUPA
	General form	Frontal process	Labial palpus	Maxillary palpus	Thoracic spiracle	Cremaster	Cocoon	Lateral ridges on abdomen
Tonzidae fam. nov.	fusiform	present	present	small	present	present	absent	absent
Plutellidae ( Plutella group)	fusiform	absent	present	large	present	absent	present	absent
Yponomeutidae	fusiform	absent	present	large	present	absent	present	present
Ypsolophidae	compact	absent	present	large	present	absent	present	absent
Glyphipterigidae	fusiform	present, small	present	large	present	absent	present	present
Argyresthiidae	compact	absent	present	large	absent	absent	present	absent
Lyonetiidae	fusiform ( Lyonetiinae ) compact ( Cemiostominae )	present ( Lyonetiinae )/ absent	present	small	absent	present	present	present
Attevidae	fusiform	absent	absent	small	present	present	absent	absent
Praydidae	compact	absent	absnet	small	present	absent	present	absent
Heliodinidae	compact	absent	present	large	present	absent	present	present
Bedelliidae	fusiform	present	absent	small	present	absent	absent	present
Scythropiidae	fusiform	present	present	small	absent	absent	absent	absent

Table 2. (Continuation.)

ADULT
	Ocelli	Chaetosema	No. of segments of maxillary palpus	2nd segment of labial palpus	Antenna compared with forewing	Scape of antennae	Pterostigma	No. of R veins on forewing	Transverse costa of A2 on forewing
Tonzidae fam. nov.	absent	absent	3	smooth	long (1.0–1.1 ×)	smooth	absent	3	present
Plutellidae ( Plutella group)	present	absent	3	scale brush	short	smooth	present	5	present
Yponomeutidae	absent	absent	1–4	smooth	short	with scale flap or pecten	present	5	present
Ypsolophidae	present	absent	4	scale brush	short	with scale flap	present absent	5	absent
Glyphipterigidae	present	absent	4	smooth	short	smooth	present	5	present
Argyresthiidae	absent	absent	1	smooth	short	with pecten	present	5	present
Lyonetiidae	absent	absent	absent	absent	long (0.8–1.5 ×)	with eye- cap	absent	2–4	present/ absent
Attevidae	absent	present	3	smooth	short	smooth	absent	5	present
Praydidae	absent	absent	1	smooth	short	smooth	present	5	present
Heliodinidae	present	absent	2	smooth	short	smooth	absent	4	present
Bedelliidae	absent	absent	1	smooth	long (1.0 ×)	with pecten	absent	4	present
Scythropiidae	absent	absent	4	smooth	short	with pecten	present	5	present

Table 2. (Continuation.)

	Spiniform setae of tergum A2–A7	Sternite	Pleural lobes	Coremata	Uncus	Socii	Teguminal processes
Tonzidae fam. nov.	absent	normal	present	present	small with a pair of long processes	small with long terminal setae	large, with dense setae
Plutellidae ( Plutella group)	absent	normal	present	present/ absent	absent	absent	sclerotized setose lobes
Yponomeutidae	present	sclerotized	present	present	semielliptical to oblong	long hairly lobe	absent
Ypsolophidae	absent	normal	present	present, long	small with short setae	oblong, rounded	absent
Glyphipterigidae	absent	forming conical lobe	absent	absent	absent	absent	absent
Argyresthiidae	absent	strongly sclerotized	present	present	oblong	small with special scales at apex	absent
Lyonetiidae	present/absent	forming processes‡	absent	present/ absent	absent	absent	absent
Attevidae	absent	sclerotized	present	present	small with a pair of short processes	slender with special scales	absent
Praydidae	absent	strongly sclerotized	present	present	absent	hairly process	absent
Heliodinidae	absent	normal	present	absent	absent	absent	absent
Bedelliidae	absent	normal	absent	absent	absent	absent	absent
Scythropiidae	absent	sclerotized	present	?	oblong	small with unscaled	absent