Membracis foliata ( Linnaeus, 1758 )

Sakakibara, Albino M. & Evangelista, Olívia, 2010, Membracis foliata (Linnaeus) (Hemiptera: Membracidae: Membracinae) and allied species: an effort towards their correct determination, Journal of Natural History 44 (35 - 36), pp. 2131-2148 : 2132-2135

Membracis foliata ( Linnaeus, 1758)

(Figures 1–4)

Cicada foliata Linnaeus, 1758, p 435 (type-locality: “ India ” [ Surinam]). (Fig. ult.) Merian, 1705, pl 5.

Membracis foliata ; Fabricius, 1775, p 675 (partim); Fabricius, 1787, p 262; Fabricius, 1794, p 9 partim); Fairmaire, 1846, p 245 (partim); Funkhouser, 1927, p 46 (cat.); Funkhouser, 1951, p 48, pl I, Figure 1; Metcalf and Wade, 1965, p 1233 (cat.); Sakakibara, 1971, p 138, Figure 2; McKamey, 1998, p 210 (cat.).

[Cigale feuillée à taches noires and blanches] Stoll, 1788, p 17, pl I, Figure 2.

Cicada flaveola Gmelin, 1789, p 2092 (type-locality: “America Meridionali”); Funkhouser, 1927, p 46 (cat.) (syn.).

Membracis flaveola ; Buckton, 1901, p 31, pl I, Figure 4; pl II, Figure 2; Metcalf and Wade, 1965, p 1230 (cat.); Sakakibara, 1971, p 138, Figure 1; McKamey, 1998, p 210 (cat.).

Membracis expansa Walker, 1851, p 475 (type-locality: Venezuela); Buckton, 1901, p 31 (syn.); Funkhouser, 1927, p 46 (cat.); Metcalf and Wade, 1965, p 1231 (cat.) (= M. flaveola ); Broomfield, 1971, p 349; McKamey, 1998, p 210 (cat.).

Membracis surgens Walker, 1851, p 475 (type-locality: Brazil); Funkhouser, 1927, p 46 (cat.) (syn.); Metcalf and Wade, 1965, p 1231 (cat.) (= M. flaveola ); Broomfield, 1971, p 379; McKamey, 1998, p 210 (cat.).

Membracis jessica Goding, 1920a, p 158 (type-locality: Ecuador); Goding, 1920b, p 34, Figure 1; Funkhouser, 1927, p 47 (cat.) (syn.); Metcalf and Wade, 1965, p 1231 (cat.) (= M. flaveola ); McKamey, 1998, p 210 (cat.).

Membracis maculata Goding, 1928, p 204 ; Metcalf and Wade, 1965, p 1232 (cat.) (= M. flaveola ); McKamey, 1998, p 210 (cat.).

Some of the latest works on Membracis taxonomy were published by Sakakibara (1971, 1992), in an attempt to discuss the identity of M. foliata (Linnaeus) , M. flaveola (Gmelin) and M. lunata Fabricius and enable their accurate identification. The controversy regarding the determination of those taxa, however, could not be completely resolved at that time and the identity of these species still remained uncertain, especially that of M. foliata .

Linnaeus (1758) described Cicada foliata as “foliacea rotundata, nigra arcu albo [rounded foliaceous, black with white arc]” and indicated a figure at plate 5 of Merian’s work ( Merian 1705) (Figure 1). The indicated illustration thus represents the taxon he intended to describe. Nevertheless, he commented (in other words) that other examined specimens exhibited a different arrangement of the white fasciae on the pronotum. He considered them conspecific and attributed these variations to sexual dimorphism. According to him, the forms varied among specimens, some having only one fascia in form of arc and others with two fasciae: one vertical, followed by another in C-shape before apex. He also cited “DeGeer”, the collection to which the examined specimens likely belonged.

Among those specimens there were, in fact, more than one species, as DeGeer (1773) demonstrated later on. Probably examining the same material used by Linnaeus, DeGeer separated the specimens into three different species: the aforementioned C. foliata L., plus C. foliata-fasciata and C. foliata-arcuata , which he described as new. He considered C. foliata as diagnosed by Linnaeus and also indicated Merian’s illustration; C. foliata-fasciata would be that with two white fasciae, one vertical and the other in C-shape; and C. foliata-arcuata , that with only one curved fascia. He also commented that Linnaeus erroneously considered all of them as sexual dimorphic variants of C. foliata . Unfortunately, the work of DeGeer (1773) was overlooked or not well understood by subsequent taxonomists, who continued interpreting the identity of M. foliata in different ways, based on Linnaeus’ description.

Fabricius (1775) erected the genus Membracis and included, among others, Membracis foliata , repeating the description given by Linnaeus (l. cit.) “…thorace foliaceo, rotundato, atro: arcubus albis [thorax foliaceous, rounded, black: white arc]”. However, on this occasion, he actually had to hand a specimen different from that described by Linnaeus, so he added, at the end, the following remarks: “in meo specimine fasciae tres flavae in thorace [in my specimen with three yellow fasciae on thorax]”.

Later on, Fabricius (1787) defined M. foliata in a different way, in comparison to the diagnosis given by Linnaeus (l. cit.): “thorace foliaceo rotundato flavo: fascia maculaque atris [pronotum foliaceous rounded yellow: with band and spot black]”

Figures 1–7. Membracis species , lateral view. (1) Merian’s illustration (plate 5, 1705), showing M. foliata (L.) on a manihot root on the right, and in detail on the left; (2) M. foliata (L.) (lectotypus ♀, LC); (3) M. expansa Walker (holotype ♀, BMNH), junior synonym of M. foliata (L.); (4) M. surgens Walker (holotype ♀, BMNH), junior synonym of M. foliata (L.); (5) M. curvilinea Walker (lectotype ♀, BMNH), junior synonym of M. foliataarcuata (DeGeer) ; (6, 7) M. lunata Fabricius (♀♀ syntypes, ZMUC), syn. nov. of M. foliatafasciata (DeGeer) .Note: Scale bar, 2 mm.

and indicated “Stoll, Cicad. tab. I. figure 2.”, which is similar to Merian’s illustration. He might have realized that his latter remarks were incorrect and, therefore, described M. lunata from the specimens that probably were previously identified as M. foliata . He defined M. lunata as follows: “thorace foliaceo rotundato atro, lunulis tribus albis [pronotum foliaceous, rounded, black, with three white moon-shaped bands]”. In the description of M. lunata, Fabricius also said that the two posterior fasciae could be sometimes united – “lunulis tribus albis, quarum posteriores interdum connatae”.

Gmelin (1789), on the other hand, considering all those specimens yet as Cicada foliata , included the following species as synonyms: M. lunata Fabricius and C. foliatafasciata DeGeer and also cited Merian’s figure. Also, he presented C. flaveola as a new name for M. foliata sensu Fabricius, 1787 (based on Stoll’s pl I, Figure 2).

Fabricius (1794) redescribed M. foliata with the same words used in his previous work, but among the references he erroneously cited DeGeer’s Table 32, Figures 9 and 10, which illustrate two other Membracis species , M. foliatafasciata and M. foliataarcuata , respectively.

In the Linnean Collection there is one female specimen labelled “ foliata ” (certainly a syntype of Cicada foliata L.) (Figure 2), which corresponds to that illustrated by Merian (1705). This confirms the determination of Membracis foliata (Linnaeus) in the sense that we present here.

Up to now, Membracis foliata (Linnaeus) has the following synonyms: Membracis expansa Walker, 1851 (Figure 3, holotype ♀, BMNH), Membracis surgens Walker, 1851 (Figure 4; holotype ♀, BMNH) and Membracis jessica Goding, 1920 . Although the holotype of the latter species was not examined, it could be readily determined with the illustration provided in Goding (1920b, figure 1). Furthermore, the description of M. jessica ( Goding 1920a) perfectly matches with M. foliata : “Nearly identical in form and size with foliata , from which it differs in being yellow with a broad vertical band extending from lateral angles to middle of dorsum and a large quadrangular spot before apex of posterior process, sooty black. Long. 15 mm.; alt. 8 mm.”

Lectotypus ♀, here designated for Cicada foliata Linnaeus, 1758

As stated previously, it is clear that Linnaeus based his description of Cicada foliata on specimens from more than one species. His remarks on different patterns of white fasciae on the pronotum, which he attributed to sexual dimorphism, have prevented accurate identification of this taxon until now. As the type-species of the family’s type-genus, the correct determination of its identity is crucial to nomenclatural stability in the group. Therefore, we designate as lectotype the female specimen in the Linnean Collection labelled “foliata”, which perfectly matches the figure presented by Merian, cited by Linnaeus in the original description. Additional labels are placed in the same pin, as follows: “ Cicada foliata Linnaeus, 1758 ”, “ LECTOTYPUS ”(red, square card), “ Lectotype designated by Sakakibara and Evangelista 2010”.

Material examined

Cicada foliata Linnaeus (lectotypus ♀, LC); Membracis expansa Walker (holotype ♀, BMNH) , Membracis surgens Walker (holotype ♀, BMNH) . Additional non-type material from ECUADOR: Sucumbios ( Shushufindi : 1♀, MNHN) ; GUYANA: “ Courantyne River ” (13, BMNH) ; “ Trinidad ” (1♀, BMNH) and “ Amazon ” (1♀, BMNH) .