Phyllocnistis, Zeller, 1848

Brito, Rosângela, Lopez-Vaamonde, Carlos, Gonçalves, Gislene L., Becker, Vitor O., Mielke, Olaf H. H. & Moreira, Gilson R. P., 2017, Taxonomic revision of Neotropical Phyllocnistis Zeller, 1848 (Lepidoptera: Gracillariidae), with descriptions of seven new species and host plant associations, Zootaxa 4341 (3), pp. 301-352 : 305-306

PHYLLOCNISTIS Zeller, 1848

Phyllocnistis ; Zeller 1848: 250 (key), 264–266; species included: P. suffusella (Zeller) , P. saligna (Zeller) . Herrich-Schäffer 1853 –1855: 341. Stainton 1854a: 285. Stainton 1854b: 161. Frey 1856: 314 –315. Herrich-Schäffer 1857: 56. Clemens 1859: 317, 327. Stainton 1859: 424 –425. Wocke 1861: 127. Stainton 1863: 605 –606, 608. Chambers 1871: 206. Wocke 1871: 333. Clemens 1872: 82 –83. Zeller 1873: 314. Chambers 1875: 106. Heinemann & Wocke 1877: 708. Zeller 1877: 450. Meyrick 1895: 757. Busck 1900: 252 –253. Rebel 1901: 218, 278. Kirby 1903: xxxviii–xxxix. van Deventer 1904: 87. Meyrick 1906: 62. Spuler 1910: 421. Meyrick 1915a: 347, 618. Hampson 1918: 387. Forbes 1923: 149, 153, fig. 112. Turner 1923: 175. Braun 1927: 194. Meyrick 1928: 782. Davis & Miller 1984: 27. Davis 1987: 374. Davis & Robinson 1998: 114. De Prins & Kawahara 2009: 113 –117. Sohn et al. 2012: 39. Brito et al. 2016: 276, 281, figs. 7–8. De Prins et al. 2016: 34.

Phylloenistis [sic]; Chambers 1875: 303; species included: P. populiella Chambers , P. ampelopsiella Chambers. Phylloetis [sic]; Chambers 1876: 19.

Type species. Opostega suffusella Zeller, 1847 . Secondary designation by Hampson (1918).

Systematic history. A detailed revision of the history of the genus Phyllocnistis was provided by De Prins & Kawahara (2009), from which the main points are reproduced here. The genus was proposed by Zeller, 1848, with P. suffusella (= unipunctella (Stephens, 1834)) and P. saligna (Zeller, 1839) included. In this work, Zeller referred to four images as representative of the genus (see original description figs. 31–34), illustrates the head, fasciae, strigulae and forewing venation of P. suffusella . However, it is noteworthy that these drawings are not present in the same work that describes the genus; rather, such drawings are found in the description of P. suffusella proposed by Zeller (1847), one year before the original description of Phyllocnistis .

The taxonomic position of Phyllocnistis remained uncertain for a long time, and the genus was allocated to different groups and families by different authors. Herrich-Schäffer (1853 -1855) was the first to include the genus in Tineidae; and Stainton (1854a, 1854b, 1859) the first to associate Phyllocnistis with Lyonetidae [sic], followed by other contemporaneous authors (e.g. Frey 1856; Wocke 1861, 1871). At the same period, Herrich-Schäffer (1857) proposed a separate group inside Lyonetidae [sic] named Phyllocnistina, which was composed of the genera Phyllocnistis , Bucculatrix Zeller and Cemiostoma Zeller. However , based on wing venation characters, Clemens (1859) transferred Phyllocnistis to Lithocolletidae, grouping the genus this time with Lithocolletis Zeller, Tischeria Zeller and Leucanthiza Clemens. Analyzing the characters of leaf mines from 20 genera of Lepidoptera, Stainton (1863) regrouped Phyllocnistis with Bucculatrix and Cemiostoma, also adding Lithocolletis, Nepticula Heyden and Lyonetia Hübner. According to Stainton, all these genera, with the exception of Bucculatrix , exhibit an endophyllous larval stage that remains inside the mine during development. Chambers (1871) also observed a few similarities between Phyllocnistis and Lithocolletis larvae, and in the same study he compared the adults of these genera. Many authors of that period allocated Phyllocnistis to Tineina, a group which included various Microlepidoptera genera ( Zeller 1873, 1877; Chambers 1875; van Deventer 1904). Nevertheless, Heinemann & Wocke (1877) treated Phyllocnistis as a separate family, grouping this genus with Cemiostoma and Bucculatrix in Phyllocnistidae. The grouping of these genera in the family was not recognized by some authors. Busck (1900) extended the definition of Phyllocnistis in his work, reallocating the genus once again in Tineidae, a classification that was also used by Meyrick (1895, 1906). Rebel (1901) regrouped the genus with Bucculatrix and Cemiostoma. He also added Opogona Zeller and Opostega Zeller to Phyllocnistinae , and assigned this subfamily to the Lyonetiidae . However, a few years later, Spuler (1910) re-transferred Phyllocnistis to Phyllocnistidae. Meyrick (1915a) and Turner (1923) continued to allocate the genus to Lyonetiidae . But later on and independently, Braun (1927) and Meyrick (1928), the latter analyzing the morphology of immature stages, assigned the genus for the first time to Gracillariidae , a classification mostly used by present-day authors ( Davis 1987; Davis & Robinson 1998; De Prins & Kawahara 2009; De Prins & De Prins 2016; De Prins et al. 2016). According to latest review of Gracillariidae ( Kawahara et al. 2017) , Phyllocnistinae is a monotypic, derived lineage formed by the genus Phyllocnistis , and sister to Oecophyllembiinae and Marmarinae . These three subfamilies share hypermetamorphic larvae, which present sap-feeding and spinning morphs, and thus lack a tissue-feeding larva.

Diagnosis. Adults: They can be distinguished from those of other genera by the light gray scales covering the body, and by the presence of a set of fasciae and strigulae on the forewings ( Figs. 3 , S1; Tab. 2). Head: reduced compound eyes; labial palpi slender and well developed. Thorax: forewing lanceolate, usually presenting five distinct fasciae: one longitudinal (lf), and four transversal (tf1–tf4), that may vary in color among species; presence of oblique strigulae: three costal (a–c) emerging from tf2, tf3 and tf4 and four apical (d–g); on the distal portion of forewing, the presence of well-marked black blotch, the apical spot (as), from where the apical strigulae emerge. Venation reduced, forewing with one discoidal cell and no accessory cells; R1 emerging from discoidal cell, and M3 and CuA2 absent; hindwing with open cell.

Pupa: set of tergal spines on abdominal segments, varying in size and shape among species; cremaster modified, with a pair of caudal processes.

Larva: presenting hypermetamorphosis, initial three instars sap-feeding, followed by a final spinning stage, shared only with Marmarinae and Oecophyllembiinae ; pupal cocoon endophyllous.

Figures

FIGURE 3. Forewings of Neotropical Phyllocnistis. (A) P. abatiae holotype (left forewing reversed); (B) P. aurilinea lectotype (left, reversed); (C) P. baccharidis holotype (left, reversed); (D) P. bourquini holotype (right); (E) P. citrella lectotype (left, reversed); (F) P. dorcas holotype (right); (G) P. drimiphaga holotype (right); (H) P. helios holotype (right); (I) P. jupiter paratype (left, reversed); (J) P. kawakitai holotype (left, reversed); (K) P. maxberryi holotype (right); (L) P. meliacella holotype (right); (M) P. norak holotype (right); (N) P. ohshimai holotype (right); (O) P. ourea paratype (right); (P) P. perseafolia holotype (left, reversed); (Q) P. petronellii paratype (right); (R) P. phoebus paratype (left, reversed); (S) P. puyehuensis holotype (right); (T) P. rotans lectotype (right); (U) P. sciophanta holotype (left, reversed); (V) P. selene specimen (left, reversed); (W) P. sexangula lectotype (left, reversed); (X) P. tethys specimen (right); (Y) P. tropaeolicola holotype (right); (Z) P. wygodzinskyi holotype (left, reversed); (AA) P. xylopiella holotype (right). Scale bars: 1 (A – H, K, L, O, P, R – Z), 0.5 mm (I, J, M, N, Q, AA).