Tischeria Zeller, 1839

Stonis, Jonas R., Diškus, Arūnas, Remeikis, Andrius, Orlovskytė, Svetlana, Solis, Alma, Paulavičiūtė, Brigita, Xu, Jiasheng & Dai, Xiaohua, 2023, Genera of Tischeriidae (Lepidoptera): a review of the global fauna, with descriptions of new taxa, Zootaxa 5333 (1), pp. 1-131 : 50-51

4. Genus Tischeria Zeller, 1839

( Figs 195–264 )

Tischeria Zeller, 1839: 219 . Type species: Tinea complanella Hübner, 1817 : pl. 64, fig. 428, junior syn. of Tischeria T. ekebladella ( Bjerkander, 1795) .

Philodoxa Gistel, 1848 : xi. Type species: Tinea complanella Hübner, 1817 : pl. 64, fig. 428, junior syn. of Tischeria ekebladella ( Bjerkander, 1795) .

Evexia Gistel, 1850: 486 . Type species: Tinea complanella Hübner, 1817 : pl. 64, fig. 428, junior syn. of Tischeria ekebladella ( Bjerkander, 1795) .

Diagnosis. External characters of the adults and wing venation characters are not informative and, therefore, insufficient for the differentiation of Tischeria because of their general uniformity. However, the majority of the species have a bright ochreous yellow forewing speckled with darker scales apically or, occasionally, also with a small black tornal spot. In the male genitalia, species of Tischeria are characterized by the presence of a juxta and medium wide to very wide phallus with usually long lateral processes, but without carinae. In the female genitalia, the genus is characterized by the presence of antrum and a slender, short ductus spermathecae with only 2–4 coils. Leaf mines of Tischeria are mostly on Fagaceae host plants, blotch-like, with a distinctive round nidus inside the leaf. See Tabs 2 , 3 for occurrence of these diagnostic characters in other genera of Tischeriidae .

Notes. Molecular data provided relatively strong support for the distinctness of this genus. In all cases Tischeria appears as a sister taxon to Rytietia gen. nov. (described below) ( Figs 62 , 63 , 67 ).

Adult. Head: frontal tuft overlapping the frons, comprised of long, slender or relatively wide lamellar scales; pecten distinctive; collar distinctly paired, comprised of relatively wide lamellar scales. Forewing usually bright ochreous yellow speckled with darker scales in apically half, occasionally also with a small black tornal spot. Hindwing slender, without androconia.

Male genitalia ( Figs 195–208 ). Uncus with two long and basally wide lateral lobes which look triangular in ventral view. Socii membranous, often usually distinctly paired (occasionally unpaired), with numerous tiny spines. Tegumen large to medium large, usually without spines on the diaphragm, occasionally diaphragm little spinose; pseudognathos absent. Valva simple-shaped, slightly sinuous in lateral view, wide basally, gradually tapering towards apex; basal process of valva short. Transtilla and anellus absent. Juxta present, well-developed, comprised of either one or two pairs of horn-like processes, well demarcated from the phallus. Vinculum large to small; ventral plate usually triangular, with a short anterior process or rounded distally. Phallus usually wide to moderately wide, occasionally slender, with well-developed lateral processes, without spines or carinae.

Female genitalia ( Figs 209–213 ). Ovipositor lobes usually large or very large; the gap between ovipositor lobes wide; second pair of ovipositor lobes twice smaller in comparison to main ovipositor lobes. Lateral lobes usually short and wide, occasionally indistinctive. Anterior apophyses usually distinctly shorter than posterior apophyses. Prela with three pairs of rod-like projections; these projections sometimes slightly or strongly widened basally. Caudal sclerite strongly developed, inverted V-shaped, with slender and pointed projection caudally.Antrum present, variously developed.Accessory sac absent; ductus spermathecae short, slender, with 2–4 coils; vesicle usually large, rounded. Corpus bursae long to medium long, usually gradually narrowing towards caudal end, with short spines proximally.

Bionomics. Generally, the species of the genus are associated with Fagaceae (mainly known from Quercu s L.). However, one species has been recorded from an Ulmaceae host plant in China ( Xu et al. 2018), three other species of uncertain taxonomic position (placement) have been reared from Rhamnaceae host plants in the USA ( Braun 1972) (see Discussion), and one species has been recorded from Lythraceae in Indonesia ( Meyrick 1936). Larvae mine leaves and produce blotch-like leaf mines with no frass deposited; a nidus is distinctly visible throughout the mine.

Species diversity and geographical distribution. The genus is widespread in Europe, temperate Asia, and North America. Species were believed to occur in South Africa ( Puplesis & Diškus 2003; Puplesis et al. 2004) until recently when these African species were transferred to another genus, Manitischeria ( Stonis et al. 2019a) . In Central America only two Tischeria species, including T. elongata Walsingham ( Walsingham 1914; Stonis & Diškus 2007) and T. neokristenseni Diškus & Stonis , sp. nov., were recognized. The genus is not yet known to occur in South America (notably from Colombia where Quercus , a potential host plant, occurs).

In total, it consists of twenty described species worldwide. This number includes the recently described Caucasian-Iranian T. caucasica Klasínski & Stonis ( Klasínski et al. 2020; Alipanah et al. 2022), Central American T. neokristenseni Diškus & Stonis sp. nov. (described below), the hitherto little known Mexican T. elongata Walsingham (see our documentation of the holotype; Figs 214–217 ), North American T. quercitella Clemens (see documentation of a specimen deposited at NHMUK; Figs 218–223 ), Japanese T. quercifolia Kuroko (see our new documentation of the male genitalia of the species; Figs 224–229 ), the East Asian T. lvoskyi Kozlov (see the first photographic documentation of the holotype; Figs 230–236 ), and East Asian T. siorkionla Kozlov. The latter was described as an allopatric East Asian subspecies of the European T. decidua Wocke, 1876 ( Kozlov 1986) and subsequently treated as a subspecies ( Kozlov 1997; Stonis et al. 2014). In the current publication, we elevate the taxonomic rank of siorkionla to species for the following reasons: 1) T. siorkionla Kozlov, 1986 (stat. nov.) differs from the European T. decidua Wocke , at least in some morphological characters of the male genitalia ( Figs 200, 201, 205–208 ); 2) there is a significant, well-known gap between East Asia and Europe in the distribution range of Quercus (e.g., Menitsky 2005); 3) to avoid usage of subspecies category.

Here, we also report a new distribution record of T. dodonaea Stainton, 1858 from the Caucasus, Armenia, feeding on the Caucasian oak (=Persian oak) Quercus macranthera Fisch. & C.A.Mey. ex Hohen. : 1 Ô, 1 ♀, ARMENIA, Aragatsotn Province, southern slope of the Mount Aragats, Antarut, 40°21’50”N, 44°16’25”E, the Caucasian oak forest, 1780 m, 2.viii.2022, leg. J.R. Stonis, genitalia slide no. AD1125Ô (MfN); 1 ♀, ARMENIA, Yerevan, Jrvezh Forest Park, 21.viii.2022, 40°11’06”N, 44°37’11”E, 1514 m, at light, leg. J.R. Stonis (MfN).

Currently, the genus includes six species with a doubtful placement. The taxonomic position at least of three North American species previously attributed to Tischeria need an in-depth revision (see Discussion).

Figures

FIGURE 62. The Maximum Likelihood tree of 11 randomly selected sequences of species representing 11 genera of Tischeriidae (Table 1 + downloaded Paratischeria ferruginea HM244395, Neotischeria neotropicana HM244391, Dishkeya bifurcata LNAUT3265-15, Coptotrichoides serjaniphaga OK017167) based on the 608 bp long mtDNA CO1-5’ sequences). The Kimura 2-parameter model (1.000 replicates) was used to calculate divergence. The best supported and resolved dichotomies are highlighted. The morphological differences of genitalia are shown to the right of the tree

FIGURES 195–208.Male genitaliaof Tischeria.195, juxtaof T. ekebladella(Bjerkander);196, same,lateral view;197,T.ekebladoides Puplesis & Diškus; 198, same, lateral view; 199, T. caucasica Klasínski & Stonis; 200, juxta of T. decidua Wocke, Crimea; 201, same, T. siorkionla Kozlov, 1986, slide no. AD589 (ZIN); 202, phallus, T. ekebladoides; 203, same, T. caucasica; 204, same, T. ekebladella; 205, same, T. decidua; 206, same, T. siorkionla; 207, valva, lateral view, T. decidua, slide no. AD590; 208, same, T. siorkionla slide no. AD589 (195–198, after Diškus & Puplesis 2003; 199, 203, Klasínski et al. 2020; 200, 201, 205–208 Stonis et al. 2014)

FIGURES 209–213. Details of antrum in the female genitalia of Tischeria. 209, T. ekebladoides Puplesis & Diškus, Tunisia, paratype, slide no. Diškus074 ZMUC (after Diškus & Puplesis 2003); 210, 211, T. ekebladella (Bjerkander), Poland, slide no. Diškus073 ZMUC (after Diškus & Puplesis 2003); 212, T. caucasica Klasínski & Stonis, Georgia, paratype, slide no. RA1082 MfN (after Klasínski et al. 2020); 213, T. sichotensis Ermolaev, Far Eastern Russia, slide no. AD587 MfN (after Stonis et al. 2014)

FIGURES 214–217. Re-study and re-documentation of holotype of Tischeria elongata Walsingham, Mexico, holotype (NHMUK). 214, male adult; 215, male genitalia, slide no. 15283 BMNH, capsule with phallus removed; 216, same, juxta; 217, same, phallus

FIGURES 218–223. Documentation of Tischeria quercitella Clemens, USA, Kirkwood, St. Louis, 18.vii.1883 (NHMUK). 218, male adult; 219, male genitalia, slide no. 28955 BMNH, socii; 220, same, valva; 221, same, capsule with phallus removed; 222, same, ventral view of phallus; 223, same, lateral view of phallus

FIGURES 224–229. Tischeria quercifolia Kuroko, Japan, Honsyu, Nagana-ken, Hukusima, 4.v.1976, T. Kumata (ZMUC). 224, male adult; 225, male genitalia, slide no. Diškus087 ZMUC, capsule with phallus removed; 226, same, lateral view of juxta; 227, same, ventral view of juxta; 228, phallus; 229, lateral view of valva

FIGURES 230–236. First photographic documentation of Tischeria lvovskyi Kozlov, Far Eastern Russia. 230, male holotype; 231, 232, male genitalia, holotype, slide no. 13838 (ZIN), capsule with phallus removed; 233, juxta in glycerol of a non-type specimen collected at the type locality by R. Puplesis, 4.vii.1982 (ZIN); 234, holotype, slide no. 13838 (ZIN), phallus; 235, same, basal processes of valvae; 236, same, juxta

FIGURES 237–243. Adults of Tischeria neokristenseni Diškus & Stonis, sp. nov., Guatemala. 237, male, holotype without abdomen, general view; 238, male head, paratype, frontal view; 239, same, dorsal view; 240, paratype, female; 241, 242, paratypes, males; 243, paratype, female (MfN)

FIGURES 244–249. Male genitalia of Tischeria neokristenseni Diškus & Stonis, sp. nov., Guatemala. 244, capsule with phallus removed, holotype, genitalia slide no. AD1077; 245, same, juxta; 246, same, phallus; 247, capsule with phallus removed, paratype, genitalia slide no. AD521; 248, same, juxta; 249, same, phallus; (MfN)

FIGURES 250–257. Female genitalia of Tischeria neokristenseni Diškus & Stonis, sp. nov., Guatemala, paratype, genitalia slide no. AD1076 (MfN). 250, general view; 251–257, details of genitalia

FIGURES 258–264. Bionomics of Tischeria neokristenseni Diškus & Stonis, sp. nov. 258–260, habitat and host plant, Quercus sp. (section Lobatae), montane mixed forest at elevation 2020 m, Santa Cruz del Quiché, El Quiché Department, Guatemala; 261–264, leaf mines

FIGURES 63, 64. The Maximum Likelihood trees of Tischeriidae based on the 608 bp long mtDNA CO1-5’ sequences. The Kimura 2-parameter model (1.000 replicates) was used to calculate divergence. The best supported and resolved dichotomies are highlighted. 63, using the same (as in the Fig. 62) 11 randomly selected sequences of species representing 11 genera of Tischeriidae; the families of host plants are shown to the right of the tree; 64, same, but omitting Rytietia uncinata sp. nov.; the distribution of species is shown to the right of the tree

FIGURES 66, 67. The Neighbor-Joining tree based of the 608 bp long mtDNA CO1-5’ Tischeriidae sequences. The Kimura 2-parameter model was used to calculate the divergence. 66, using the sequences of Coptotriche, Dishkeya, Rytietia gen. nov., Pafazaria gen. nov., Manitischeria, and Tischeria species; 67, using sequences of genera with a juxta in the male genitalia (Remark: this was also performed applying Maximum Likelihood method and the results were similar)