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McKamey2022 Membracis Enchenopa
McKamey2022 Membracis Enchenopa
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Stuart H. Mckamey
The subfamily Membracinae contains having a lateral carina that attains the lateral
five tribes: Aconophorini, Hoplophorionini, margin, well posterior to the humeral angle
Hypsoprorini, Membracini, and Talipedini (Fig. 3). Strümpel and Strümpel (2006)
(Deitz 1975, Wallace and Deitz 2004). Of revised Enchophyllum and restricted it to
these, Membracini is the most morpho- those species with only one lateral carina
logically diverse. Dietrich and McKamey and without any carinae on the metopidium
(1995) investigated the phylogeny of the (Figs. 4, 5); those with metopidial carinae
subfamily and provided a key to the gen- were relegated to Enchenopa. In redefining
era of Membracini. They distinguished these two genera, they grouped together two
the morphology of Enchophyllum Amyot almost independent sets of species in terms
and Serville from Enchenopa Amyot and of immature morphology: those current and
Serville traditionally: Enchophyllum hav- former Enchophyllum, with nymphs having
ing a pronotum with a lateral carina on the abdominal scoli all subequal in length
each side that terminates before or above and with a uniform or mostly uniform gray
the humeral angle (Fig. 1), and Enchenopa or white waxlike covering (Figs. 2, 4, 6), and
Figs. 1, 2. Enchenopa adult, and fifth instar. 1, Enchenopa sp. near Enchenopa amazonensis Strümpel and
Strümpel, adult with lateral carinae terminated above humeral angle and also with metopidial carinae. 2, Enchenopa
sp. nymph with abdominal scoli all subequal in size. (Photos courtesy of Robert Oelman and Milan Kozenek,
respectively.).
the former and some current Enchenopa but not others (Figs. 13, 14) and stated [in
whose nymphs have the abdominal scoli Spanish]:
increasing in length posteriorly and with a
patchy white or a non-white or gray coat- “Following the characteristics of the
ing, or no waxlike coating at all (some genus Membracis the lateral carina
Enchenopa binotata [Say], Enchenopa should not be found, but finding it
curvata [Fabricius], Enchenopa latipes is not important when it passes only
[Say], Enchenopa luizae Lencioni-Neto above the humeral [angles] or that
and Sakakibara, and Enchenopa pilosa almost reaches it . . .”
Strümpel and Strümpel; Fig. 3). Although
the former group closely resembles the Richter also said it should be added
nymphs of Membracis and Phyllotropis, it is to the features of Membracis. Later in
not yet known if either of these Enchenopa the same work, Richter (1947) described
groups are monophyletic. M. paullula Richter and M. robiginosa
Adults of Membracis Fabricius spe- Richter, which are small species with one
cies such as M. foliata (Linnaeus) and lacking a lateral carina, and described a
M. foliatafasciata (DeGeer) (photographs series of new, low-crested Membracis
provided in Sakakibara and Evangelista that bridge the gaps between Membracis,
2010) are immediately recognized as belong- Enchenopa, and Enchophyllum. These
ing to that genus by having a large, foliaceous species have a less foliaceous pronotum
pronotum that lacks any trace of lateral cari- with one or two dorsal white maculae
nae. Phyllotropis Stål species are similar, (as occur in many species of Enchenopa
but differ in having an oblique carinae on and Enchophyllum; as in Fig. 1) and
the pronotum (see Sakakibara 1992). Adults M. carinulata Richter (Fig. 12) has one
of most Enchophyllum and Enchenopa or two short metopidial carinae as in
are also recognized as such, in having a Enchenopa. Some of Richter’s other
less foliaceous pronotum, at least one lat- species of Membracis, most notably
eral carina, and a frontal horn. Folicarina M. caquetensis Richter, even have an
Sakakibara (1992) was established for a anterior white macula as in Enchenopa
species with a greatly compressed prono- albifrons Strümpel and Strümpel,
tum bearing several metopidial carinae as Enchenopa auridorsa Sakakibara and
in Enchenopa, but without a frontal horn. Marques, Enchenopa grandis Strümpel
A close relationship of these five genera is and Strümpel, Enchenopa longicollum
suggested by the gradation among them and, (Olivier), and Enchenopa pittieri Strümpel
as Dietrich and McKamey (1995) noted, and Strümpel.
these genera are separated only by a combi- Sakakibara (2012) redefined Talipedini
nation of characters rather than a single char- to include Erechtia Walker; described the
acter. The nymphs of most Enchophyllum new, related genus Pseudoerechtia; rein-
(Figs. 4, 6), Folicarina (Fig. 7), Membracis stated Tropidoscyta Stål; and rejected
(Fig. 8), and Phyllotropis (Fig. 9) share the the synonymy of Talipes Deitz (1975)
uniform or patchy white waxlike covering under Trinarea Goding. Sakakibara jus-
of some Enchenopa. tified his synonymy of Trinarea under
Richter (1947) modified the limits of Erechtia by citing “several diagnostic
Membracis. In his description of M. flava characters” between them but only men-
Richter, which he wrote was close to M. tioned the presence of a very weak trans-
mexicana Guérin-Méneville, he observed versal carina of the pronotal metopidium
a short lateral carina in some females, in Trinarea, in contrast to a prominent
Fig. 3. Enchenopa pilosa. Adult pronotal lateral carina attains lateral margin, nymph abdominal scoli increasing
in size posteriorly. (Photo courtesy of Ted C. MacRae.).
one in other Erechtia and the lack of basis of these synapomorphies and cla-
one in Talipes. However, if membrac- distic analyses, I accept the Membracinae
ids are known for anything, it is for the tribes and Membracini genera as defined
astounding variation in their prono- by Dietrich and McKamey (1995) except
tum, which weakens Sakakibara’s argu- for Campylenchia and their traditional,
ment. More importantly, Talipedini as broader definition of Enchophyllum.
defined by Deitz (without Erechtia and In Strümpel and Strümpel’s (2006)
Tropidoscyta) is supported by three revision of Enchophyllum, they excluded
non-pronotal unique synapomorphies: eight species “because they have been
adults with reduced hind tarsi, nymphs transferred to other genera.” Three of
with a unique abdominal scoli pattern these transfers were made by Strümpel
intermediate between Hoplophorionini and Strümpel (2014) in their revision of
and Membracini (McKamey and Deitz Enchenopa, with Campylenchia as a new
1996, their Fig. 91; Lencioni-Neto and junior synonym. The five Enchophyllum
Sakakibara 2016); and females guarding species not treated in either revision were
egg masses by flicking their hind legs at Enchophyllum imbelle Stål, Enchophyllum
intruders (McKamey and Deitz 1996). melaleucum (Walker), Enchophyllum
Furthermore, Trinarea appendiculata of peruanum (Schmidt), Enchophyllum
Talipedini was found to be the sister-group schmidti (Funkhouser) and Enchophyllum
of Hoplophorionini in the cladistic anal- ensatum intermedium Fitch. Also, they
yses of Dietrich and McKamey (1995) did not treat a species that was formerly
and McKamey and Deitz (1996). On the in Campylenchia: Enchenopa minans
Figs. 4–9. Membracini. 4, Enchophyllum quinquemaculatum (Fairmaire) adult and nymph from Severiano Moraes
Filho, Garanhuns, Pernambuco, Brazil (courtesy of Caiu Flat). 5, Enchophyllum sp. near Enchophyllum markantum
Strümpel and Strümpel 2006) from Guyana, Mazaruni-Potaro, Bartica, 94m (SHM collection). 6, Same, nymph.
7, Folicarina sp. adult and nymphs from Tambopata, Peru (Photo courtesy of Richard C. Hoyer/Birdernaturalist.).
8, Membracis mimica Walker from Vale do Sol, Parnamirim, Rio Grande do Norte, Brazil (Photo courtesy of Víctor
de Paiva.). 9, Phyllotropis fasciata (Fabricius) teneral adult, exuvia, and eclosing adult from Puerto Francisco de
Orellana (Coca), Ecuador (Photo courtesy of Chan [from Hong Kong]).
Figs. 10–14. Selected species described by Richter under Membracis. 10, Leioscyta paullula (Richter),
new combination. 11, Leioscyta robiginosa (Richter), new combination. 12, Membracis carinulata Richter.
13, Membracis flava Richter female. 14, Same. (All illustrations from Richter 1947, Revista de la Academia
Colombiana de Ciencias Exactas, Físicas y Naturales, with permission).
(Fairmaire) and they listed Enchenopa 244 arm. The light was diffused using
varians (Walker) as a junior synonym a simple, lampshade-style cone of trans-
of Enchenopa monoceros. Here the lucent paper between the specimen and
identities of Enchophyllum imbelle, light sources. Images were focus-stacked
Enchenopa minans, Enchenopa tatei, using Zerene Stacker, version 1.04, Build
and Enchenopa varians are ascertained T201706041920. Stacked images were
and placed. For clarity, the holotype of enhanced and edited in Adobe Photoshop
Enchenopa monoceros (Germar), which CSS Extended version 12.0.
is the type species of Enchenopa, is also The pinned specimens figured are
shown here for the first time. located in the following institutions:
Figs. 15–18. Pinned adults of Membracini incertae sedis (Both species exhibit lateral carinae originating on the
metopidium.). 15, Enchophyllum imbelle Stål syntype, frontal view. 16, Same, lateral view (NHRS-GULI000075675
©2020. Photos courtesy of Gunvi Lindberg, CC-BY 4.0 license.). 17, Erechtia trinotata Funkhouser holotype
(USNM), frontal view. 18, Same, lateral view.
Figs. 19–21. Enchenopa monoceros. 19, Holotype male (Photo courtesy of Ihor Shidlovsky, LVIV.). 20, Male
(Photo courtesy of Ricardo A. Palonsky). 21, Female (From Strümpel and Strümpel 2014, with kind permission of
the CeNak, Univ. Hamburg.).
Membracini, a more thorough study of time the identity of the type species of
their relationships is required, ideally Enchenopa and the sexual dimorphism
including genomic characters. within it.
Only some basic taxonomic changes The identities of Enchophyllum peru-
can be made now: referring two spe- anum (Schmidt), Enchophyllum schmidti
cies to Leioscyta Fowler; clarifying the (Funkhouser), and Enchophyllum ensa-
identity of four species of Enchenopa tum intermedium, the last of which were
not treated in either of Strümpel and described in a footnote (Fitch 1856: 465)
Strümpel’s revisions (2006, 2014); refer- and not present in Fitch’s collection in the
ring selected species to Membracini NYSM (Timothy McCabe, pers. comm.),
incertae sedis; and verifying for the first could not be determined.
Figs. 22–24. Enchenopa species not treated by Strümpel and Strümpel (2014). 22, Enchenopa varians Walker,
holotype (BMNH). 23, Enchenopa minans (Fairmaire), syntype (NHMW), frontal view. 24, Same, lateral view
(Photos courtesy of O. Evangelista, CSIRO.).
Stål, C. 1869b. Bidrag till membracidernas Mitteilungen aus dem Zoologischen Museum
kannedom. Öfversigt af Kongliga Svenska Hamburg 17(191): 1–137. https://www.
Vetenskaps-Akademiens Förhandlingar. 26 (3): zobodat.at/pdf/Ent-Mitt-Zool-Mus-Hamburg_
231–300. 17_0001-0137.pdf.
Strümpel, H. and R. Strümpel. 2006. Revision of Walker, F. 1858. Homoptera, pp. 1-117. In In-
the Neotropical treehopper genus Enchophyl- secta saundersiana: or characters of unde-
lum (Hemiptera: Membracidae, Membraci- scribed insects in the collection of William
nae). Mitteilungen aus dem Zoologischen Wilson Saunders (no editor given), Esq. John
Museum Hamburg 14(175): 335–371. https:// Van Voorst. London 4: 1-117. https://www.
www.zobodat.at/pdf/Ent-Mitt-Zool-Mus- biodiversitylibrary.org/item/25015#page/69/
Hamburg_14_0335-0371.pdf. mode/1up.
Strümpel, H. and R. Strümpel. 2014. Revision Wallace, M. S. and L. L. Deitz. 2004. Phylogeny
der amerikanischen Membracidengattung and systematics of the treehopper subfamily
Enchenopa (Hemiptera: Auchenorrhyncha: Centrotinae (Hemiptera: Membracidae). Mem-
Cicadomorpha: Membracidae) mit Besch- oirs on Entomology, International 19: i–iv,
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