Notes on the New World Treehopper Genera Membracis

Fabricius, Enchenopa Amyot and Serville, and Closely

Related Taxa (Hemiptera: Membracidae: Membracinae)
Author: Mckamey, Stuart H.
Source: Proceedings of the Entomological Society of Washington,
124(1) : 85-97
Published By: Entomological Society of Washington
URL: https://doi.org/10.4289/0013-8797.124.1.85

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 PROC. ENTOMOL. SOC. WASH.
124(1), 2022, pp. 85–97

NOTES ON THE NEW WORLD TREEHOPPER GENERA MEMBRACIS

FABRICIUS, ENCHENOPA AMYOT AND SERVILLE, AND CLOSELY
RELATED TAXA (HEMIPTERA: MEMBRACIDAE: MEMBRACINAE)

Stuart H. Mckamey

Systematic Entomology Laboratory, Agricultural Research Service, U.S.

Department of Agriculture, c/o National Museum of Natural History, P.O. Box
37012, Washington, DC 20013, USA (e-mail: stuart.mckamey@usda.gov,
urn:lsid:zoobank.org:author:E6BD01B8-0684-464A-8A86-812701DA05C7)

Abstract.—The similarities and differences in adults and nymphs among Enchenopa

Amyot and Serville, Enchophyllum Amyot and Serville, Folicarina Sakakibara,
Membracis Fabricius, and Phyllotropis Stål are discussed. Leoiscyta pallula (Richter),
new combination and Leoiscyta robiginosa (Richter), new combination, are both
referred from Membracis. Enchophyllum imbelle Stål, Membracis carinulata Richter,
and Membracis flava Richter are referred to Membracini incertae sedis, new placements.
Enchenopa melaleuca Walker, reinstated combination, is referred back to its
original genus from Enchophyllum. The holotype and sexual dimorphism are shown
for Enchenopa monoceros (Germar). Enchenopa varians Walker is reinstated as a
valid species. Because earlier workers placed Campylenchia Stål as a junior synonym of
Enchenopa, but did not list all species, Enchenopa minans (Fairmaire) new combination
and Enchenopa tatei Goding, reinstated combination, are proposed. For the first time,
nymphs of Phyllotropis and Folicarina are figured.
Key Words: Membracini, new combination, immatures, Neotropical
DOI: 10.4289/0013-8797.124.1.85

The subfamily Membracinae contains
five tribes: Aconophorini, Hoplophorionini,
Hypsoprorini, Membracini, and Talipedini
(Deitz 1975, Wallace and Deitz 2004). Of
these, Membracini is the most morpho-
logically diverse. Dietrich and McKamey
(1995) investigated the phylogeny of the
subfamily and provided a key to the gen-
era of Membracini. They distinguished
the morphology of Enchophyllum Amyot
and Serville from Enchenopa Amyot and
Serville traditionally: Enchophyllum hav-
ing a pronotum with a lateral carina on
each side that terminates before or above
the humeral angle (Fig. 1), and Enchenopa
having a lateral carina that attains the lateral
margin, well posterior to the humeral angle
(Fig. 3). Strümpel and Strümpel (2006)
revised Enchophyllum and restricted it to
those species with only one lateral carina
and without any carinae on the metopidium
(Figs. 4, 5); those with metopidial carinae
were relegated to Enchenopa. In redefining
these two genera, they grouped together two
almost independent sets of species in terms
of immature morphology: those current and
former Enchophyllum, with nymphs having
the abdominal scoli all subequal in length
and with a uniform or mostly uniform gray
or white waxlike covering (Figs. 2, 4, 6), and

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 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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.).

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 VOLUME 124, NUMBER 1 87
the former and some current Enchenopa
whose nymphs have the abdominal scoli
increasing in length posteriorly and with a
patchy white or a non-white or gray coat-
ing, or no waxlike coating at all (some
Enchenopa binotata [Say], Enchenopa
curvata [Fabricius], Enchenopa latipes
[Say], Enchenopa luizae Lencioni-Neto
and Sakakibara, and Enchenopa pilosa
Strümpel and Strümpel; Fig. 3). Although
the former group closely resembles the
nymphs of Membracis and Phyllotropis, it is
not yet known if either of these Enchenopa
groups are monophyletic.
Adults of Membracis Fabricius spe-
cies such as M. foliata (Linnaeus) and
M. foliatafasciata (DeGeer) (photographs
provided in Sakakibara and Evangelista
2010) are immediately recognized as belong-
ing to that genus by having a large, foliaceous
pronotum that lacks any trace of lateral cari-
nae. Phyllotropis Stål species are similar,
but differ in having an oblique carinae on
the pronotum (see Sakakibara 1992). Adults
of most Enchophyllum and Enchenopa
are also recognized as such, in having a
less foliaceous pronotum, at least one lat-
eral carina, and a frontal horn. Folicarina
Sakakibara (1992) was established for a
species with a greatly compressed prono-
tum bearing several metopidial carinae as
in Enchenopa, but without a frontal horn.
A close relationship of these five genera is
suggested by the gradation among them and,
as Dietrich and McKamey (1995) noted,
these genera are separated only by a combi-
nation of characters rather than a single char-
acter. The nymphs of most Enchophyllum
(Figs. 4, 6), Folicarina (Fig. 7), Membracis
(Fig. 8), and Phyllotropis (Fig. 9) share the
uniform or patchy white waxlike covering
of some Enchenopa.
Richter (1947) modified the limits of
Membracis. In his description of M. flava
Richter, which he wrote was close to M.
mexicana Guérin-Méneville, he observed
a short lateral carina in some females,
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but not others (Figs. 13, 14) and stated [in
Spanish]:
“Following the characteristics of the
genus Membracis the lateral carina
should not be found, but finding it
is not important when it passes only
above the humeral [angles] or that
almost reaches it . . .”
Richter also said it should be added
to the features of Membracis. Later in
the same work, Richter (1947) described
M. paullula Richter and M. robiginosa
Richter, which are small species with one
lacking a lateral carina, and described a
series of new, low-crested Membracis
that bridge the gaps between Membracis,
Enchenopa, and Enchophyllum. These
species have a less foliaceous pronotum
with one or two dorsal white maculae
(as occur in many species of Enchenopa
and Enchophyllum; as in Fig. 1) and
M. carinulata Richter (Fig. 12) has one
or two short metopidial carinae as in
Enchenopa. Some of Richter’s other
species of Membracis, most notably
M. caquetensis Richter, even have an
anterior white macula as in Enchenopa
albifrons Strümpel and Strümpel,
Enchenopa auridorsa Sakakibara and
Marques, Enchenopa grandis Strümpel
and Strümpel, Enchenopa longicollum
(Olivier), and Enchenopa pittieri Strümpel
and Strümpel.
Sakakibara (2012) redefined Talipedini
to include Erechtia Walker; described the
new, related genus Pseudoerechtia; rein-
stated Tropidoscyta Stål; and rejected
the synonymy of Talipes Deitz (1975)
under Trinarea Goding. Sakakibara jus-
tified his synonymy of Trinarea under
Erechtia by citing “several diagnostic
characters” between them but only men-
tioned the presence of a very weak trans-
versal carina of the pronotal metopidium
in Trinarea, in contrast to a prominent
 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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
one in Talipes. However, if membrac-
ids are known for anything, it is for the
astounding variation in their prono-
tum, which weakens Sakakibara’s argu-
ment. More importantly, Talipedini as
defined by Deitz (without Erechtia and
Tropidoscyta) is supported by three
non-pronotal unique synapomorphies:
adults with reduced hind tarsi, nymphs
with a unique abdominal scoli pattern
intermediate between Hoplophorionini
and Membracini (McKamey and Deitz
1996, their Fig. 91; Lencioni-Neto and
Sakakibara 2016); and females guarding
egg masses by flicking their hind legs at
intruders (McKamey and Deitz 1996).
Furthermore, Trinarea appendiculata of
Talipedini was found to be the sister-group
of Hoplophorionini in the cladistic anal-
yses of Dietrich and McKamey (1995)
and McKamey and Deitz (1996). On the
basis of these synapomorphies and cla-
distic analyses, I accept the Membracinae
tribes and Membracini genera as defined
by Dietrich and McKamey (1995) except
for Campylenchia and their traditional,
broader definition of Enchophyllum.
In Strümpel and Strümpel’s (2006)
revision of Enchophyllum, they excluded
eight species “because they have been
transferred to other genera.” Three of
these transfers were made by Strümpel
and Strümpel (2014) in their revision of
Enchenopa, with Campylenchia as a new
junior synonym. The five Enchophyllum
species not treated in either revision were
Enchophyllum imbelle Stål, Enchophyllum
melaleucum (Walker), Enchophyllum
peruanum (Schmidt), Enchophyllum
schmidti (Funkhouser) and Enchophyllum
ensatum intermedium Fitch. Also, they
did not treat a species that was formerly
in Campylenchia: Enchenopa minans

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 VOLUME 124, NUMBER 1 89

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]).

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 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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
varians (Walker) as a junior synonym
of Enchenopa monoceros. Here the
identities of Enchophyllum imbelle,
Enchenopa minans, Enchenopa tatei,
and Enchenopa varians are ascertained
and placed. For clarity, the holotype of
Enchenopa monoceros (Germar), which
is the type species of Enchenopa, is also
shown here for the first time.
244 arm. The light was diffused using
a simple, lampshade-style cone of trans-
lucent paper between the specimen and
light sources. Images were focus-stacked
using Zerene Stacker, version 1.04, Build
T201706041920. Stacked images were
enhanced and edited in Adobe Photoshop
CSS Extended version 12.0.
The pinned specimens figured are
located in the following institutions:

BMNH England, London, The Natural

Materials and Methods History Museum
Morphological terms follow Dietrich DZUP Brazil, Curitiba, Universidade
and McKamey (1995). Images of Figs. 5 Federal do Paraná, Departamen
and 6 were taken with a Canon EOS to do Zoologia, Coleção Ento­
5DS R DSLR camera with an adjust- mológica Pe. Jesus Santiago
able 65mm lens. Photos were taken Moure
using Capture One Pro version 10.1.2, LVIV Ukraine, L’Viv, Ivan Franko
64 Bit, Build 10.1.2.23 imaging soft- National University of L’viv,
ware, aided by CamLift version 2.9.7.1. Biological Faculty, Zoological
The specimen was lit using two adjust- Museum, Germar Collection
able Dynalite MH2050 RoadMax flash NHMW Austria, Vienna, Naturhistorisches
heads, each attached to a Manfrotto Museum Wien

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 VOLUME 124, NUMBER 1 91

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.

NHRS Sweden, Stockholm, Naturhisto­ Results and Discussion

riska Riksmuseet The current concepts of Enchenopa,
NYSM USA, New York, Albany, New Enchophyllum, Folicarina, Membracis,
York State Museum and Phyllotropis are still based solely
USNM USA, Washington DC, U.S. on the form of the pronotum. Given the
National Museum of Natural gradation of pronotal forms among them,
History, Smithsonian Institution and the species here moved to unplaced

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 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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
their relationships is required, ideally
including genomic characters.
Only some basic taxonomic changes
can be made now: referring two spe-
cies to Leioscyta Fowler; clarifying the
identity of four species of Enchenopa
not treated in either of Strümpel and
Strümpel’s revisions (2006, 2014); refer-
ring selected species to Membracini
incertae sedis; and verifying for the first
time the identity of the type species of
Enchenopa and the sexual dimorphism
within it.
The identities of Enchophyllum peru-
anum (Schmidt), Enchophyllum schmidti
(Funkhouser), and Enchophyllum ensa-
tum intermedium, the last of which were
described in a footnote (Fitch 1856: 465)
and not present in Fitch’s collection in the
NYSM (Timothy McCabe, pers. comm.),
could not be determined.

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 VOLUME 124, NUMBER 1 93

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.).

Membracis Fabricius http://n2t.net/ark:/65665/3aef7a5c8-bc71-

Restricting the definition of Membracis
to only those species with an at least
somewhat foliaceous pronotum and lack a
lateral carina in both sexes, which do not
fit in any of the aforementioned genera,
Richter’s (1947) species M. pallula, M.
robiginosa, M. flava, and M. carinulata
must be removed from the genus. The first
two are here referred to Leioscyta Fowler,
but the last two cannot be assigned to
any current genus and must therefore
be placed in Membracini incertae sedis.
Enchophyllum imbelle Stål (Figs. 15, 16),
which Strümpel and Strümpel (2006,
2014) excluded from treatment, and
Erechtia trinotata Funkhouser (Figs. 17,
18) also fall into this category.
Leioscyta Fowler
The species of this genus are
much smaller than any Membracis,
Enchenopa, Enchophyllum, Folicarina, and
Phyllotropis (for example, L. pallula
is 3 mm and L. robiginosa is “a little
larger” [Richter, 1947]). Additionally,
Leoiscyta species usually have a pair
of lateral carinae running all along
the pronotum (as in Fig. 10), such as
L. pulchella Funkhouser (see holotype at
421d-90d3-979e0bdb6b05). However, the
genus has other small species that have
these carinae absent or weak, such as
L. robiginosa (Fig. 11) and L. ferruginea
Funk­houser (see holotype at https://collections.
nmnh.si.edu/search/ento/?ark=ark:/65665/
36973a650d9ef4f70823067fde87676ac). The
shape of the lateral carina of L. pallula
and the presence of two rather than three
discoidal cells in the forewing (i.e., having
only one m-cu crossvein) of both L. pal-
lula and L. robiginosa distinguish them
from most other genera of Membracini
with species about 3 mm or less: Eunusa
Fonseca, Paragara Goding, Bolbonota
Amyot and Serville, and some Erechtia
Walker. The sole species of Pseudoerechtia
Sakakibara is at least 4.4 mm in length,
so not included among these genera with
smaller species. Therefore, two Membracis
species described by Richter fall within the
current bounds of Leioscyta, resulting in
the following new combinations:
Leoiscyta pallula (Richter), new combina-
tion (Fig. 10).
Membracis pallula Richter 1947: 398.
Leoiscyta robiginosa (Richter), new com-
bination (Fig. 11).
Membracis robiginosa Richter 1947: 398.

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 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Enchenopa Amyot and Serville
Enchenopa melaleuca Walker, rein-
stated combination.
Enchenopa melaleuca Walker 1858: 59.
Enchophyllum melaleucum; Fowler 1894:
7 [error]
As Walker originally placed it,
Enchenopa melaleuca does appear to
belong to Enchenopa, based on the illus-
tration by Fowler (1894), which shows a
black metopidium and two pairs of cari-
nae. Buckton’s (1902) less reliable tax-
onomy also suggests Enchenopa, but
the carinae cannot be deciphered from
his poor illustration, which shows a
metopidial white spot. Fowler (1894) did
not justify his placement of the species
in Enchophyllum, but under Enchenopa
referred to the distinguishing character
between the two genera being “the cessa-
tion or continuation of the lateral carina,”
which corresponds to the traditional, out-
dated concept of used by Dietrich and
McKamey (1995). Fowler’s illustration
shows that the species is most similar to
Enchenopa. longimaculatum Strümpel
and Strümpel (2014) and Enchenopa sin-
gularis Strümpel and Strümpel (2014).
Enchenopa monoceros (Germar) (Figs. 19–21)
In their revision of the genus, Strümpel
and Strümpel (2014) had not found the
holotype of Enchenopa monoceros (Fig.
19) and therefore accepted identifications
found in ten other collections. Because
this is the type species of Enchenopa, it
merits clarification. The species is sexu-
ally dimorphic, as shown by Lencioni-
Neto (2011, 2018); the female (Fig. 21)
has a longer frontal horn than the male
(Fig. 20), as illustrated by Strümpel and
Strümpel (2014). One of the most distinc-
tive characteristics of Enchenopa monoc-
eros is the distoposterior clear area of the
forewing. This is not a diagnostic feature
for the species, however, because there
are several other species that share this
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clear area of the forewing: Enchenopa
beebei (Haviland), Enchenopa costari-
censis Schmidt, Enchenopa euniceae
Rothea and Creão-Duarte, Enchenopa
luizae, and Enchenopa varians. Strümpel
and Strümpel (2014) reported the full dis-
tribution of Enchenopa. monoceros as
Argentina, Bolivia, Brazil, Costa Rica,
Ecuador, Mexico, Paraguay, Peru, and
Venezuela.
Enchenopa varians Walker, revalidated,
reinstated combination (Fig. 22).
Enchenopa varians Walker 1858: 62.
To Enchenopa monoceros (Germar);
Goding 1928: 210 [new synonym; error].
Note: Goding (1928) was the first to treat
Enchenopa varians as a junior syno-
nym of Enchenopa monoceros. This
erroneous synonymy was followed by
all later workers, including Strümpel
and Strümpel (2014). The holotype of
Enchenopa varians (BMNH) has a fron-
tal horn that is even longer than that of
the female of Enchenopa monoceros but
shares the clear area of the forewing.
In treating Campylenchia as a junior
synonym of Enchenopa, Strümpel and
Strümpel (2014: p. 5) neglected to provide
new combinations for, place in synonymy,
or otherwise mention two valid species of
Campylenchia. Therefore, two new combi-
nations are proposed here:
Enchenopa minans (Fairmaire 1846:
252), new combination (Figs. 23, 24).
Note: The illustrated syntype of
Enchenopa minans has a broken ante-
rior horn but otherwise conforms to
Strümpel and Strümpel’s (2014) defini-
tion of the genus.
Enchenopa tatei Goding, 1928: 211,
reinstated combination.
Note: The Enchenopa tatei holotype
(USNM; images available at http://
n2t.net/ark:/65665/m360d3402c-e351-
4c12-ac3f-9ba670a73278) is too broken
 VOLUME 124, NUMBER 1 95

to evaluate possible synonymy, but has carinae as in Enchenopa, but lacks a

the metopidial carinae and a lateral
carina, both of which are characteristic
for the genus.
Membracini incertae sedis
Enchophyllum imbelle Stål 1869: 271, new
placement (Figs. 15, 16).
Note: Enchophyllum imbelle resembles
Membracis but has a lateral carina, as in
Phyllotropis. It is significantly larger than
any Leioscyta (see scale bar). Until the
relationships between Enchophyllum,
Membracis, and Phyllotropis are
resolved, Enchophyllum imbelle cannot
be placed to genus.
Erechtia trinotata Funkhouser 1930: 412
(Figs. 17, 18).
Note: Erechtia trinotata (Fig. 4) is
most similar to the low-crested species
Membracis carinulata Richter (1947)
and M. caquetensis Richter (1947), the
former differing in its lack of an ante-
rior white macula and the latter in its
shorter first dorsal macula. It is curious
why Funkhouser placed this species in
Erechtia, which by the traditional defi-
nition (Dietrich and McKamey 1995)
being used at the time, and even by
the revised definition by Sakakibara
(2012), the pronotum of Erechtia is not
at all foliaceous and has multiple sharp,
raised lateral carinae. Sakakibara
(2012) referred Erechtia trinotata to
Membracini incertae sedis but did
not provide an illustration or photo-
graphs, which are included here for
clarity and confirmation that it should
be considered as Membracini incer-
tae sedis.
Membracis carinulata Richter 1947: 399,
new placement (Fig. 12).
As noted above, M. carinulata Richter
(Fig.12) has one or two short, metopidial
frontal horn.
Membracis flava Richter 1947: 391, new
placement (Figs. 13, 14).
Note: Richter reported that most
M. flava lacked a short lateral carina
anteriorly (Fig. 13) while others had one
(Fig. 14). Dimorphism of this kind has
never been reported for other species, so it
is possible that Richter mistook interspe-
cific variation for intraspecific variation.
Acknowledgments
I thank Ihor Shidlovsky (LVIV) for the
photograph of the Enchenopa monoceros
holotype, Olivia Evangelista (CSIRO) for
photographs of the Enchenopa varians
and Membracis minans types and Herbert
Zettel and Harald Bruckner (NHMW) for
assisting her, Gunvi Lindberg (NHRS)
for the photographs of the Enchophyllum
imbelle holotype, Timothy McCabe
(NYSM) for checking Fitch’s collection,
the CeNak, Univ. Hamburg, for permis-
sion to reproduce an image from Strümpel
and Strümpel’s 2014 article, Dr. Carlos
E. Sarmiento for permission to repro-
duce images from Richter’s 1947 article,
Robert Oelman, Milan Kozanek, Ted
MacRae, Caiu Flat, Richard C. Hoyer,
Víctor de Paiva, Ricardo Palonsky, and
Chan for their photographs of live spec-
imens, Ben Proshek (USDA Systematic
Entomology Laboratory [SEL]) for pho-
tographing Figs. 5, 6, Alyssa Seemann
(SEL) for producing all plates, and Chris
Dietrich (Illinois Natural History Survey)
and Olivia Evangelista (CSIRO) for com-
ments on earlier drafts of the manuscript.
Mention of trade names or commercial
products in this publication is solely for
the purpose of providing specific infor-
mation and does not imply endorsement
by the USDA. The USDA is an equal
opportunity provider and employer.

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 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Literature Cited
Buckton, G. B. 1902. A monograph of the
Membracidæ. L. Reeve & Co., Limited.
pp. 93-180. https://www.biodiversitylibrary.
org/item/238423#page/7/mode/1up.
Deitz, L. L. 1975. Classification of the higher
categories of the New World treehoppers
(Homoptera: Membracidae). North Carolina
Agriculture Experiment Station Technical
Bulletin 225: i–ii, 1–177.
Dietrich, C. H. and S. H. McKamey. 1995. Two
new Neotropical genera and investigation of
the phylogeny of the subfamily Membracinae
(Homoptera: Membracidae). Proceedings of
the Entomological Society of America 97(1):
1–16. https://www.biodiversitylibrary.org/page/
16152215/#page/7/mode/1up.
Fairmaire, L. M. H. 1846. Revue de la tribu des
membracides. Annales de la Société
Entomologique de France (2) 4: 235–320.
https://www.biodiversitylibrary.org/page/
8261565#page/241/mode/1up.
Fitch, A. 1856. Third report on noxious and other
insects of the State of New York. Transactions
of the New York State Agricultural Society
16: 315–490. http://www.biodiversitylibrary.
org/page/4584660#page/539/mode/1up.
Fowler, W. W, 1894. Order Rhynchota: sub-
order Hemiptera-Homoptera, pp. 1–24. In
Godman, F. D. and O. Salvin (eds.). Bio-
logia Centrali-Americana; or, Contributions
to the knowledge of the fauna and flora of
Mexico and Central America. 2. [i]-xi, [a
page errata], [1]-339.Taylor and Francis,
London. https://www.biodiversitylibrary.org/
item/14633#page/349/mode/1up.
Funkhouser, W. D. 1927. Membracidae, pp.
1-581. In G. Horváth, G. and H. M. Parshley
(eds.). General catalogue of the Hemiptera.
Smith College. Northampton, Mass. Fasc. 1.
xii + 581 pp. https://www.biodiversitylibrary.
org/item/29906#page/3/mode/1up.
Funkhouser, W. D. 1930. New genera and spe-
cies of Neotropical Membracidae. Journal
of the New York Entomological Society
38: 405–421. https://www.biodiversitylibrary.
org/page/5076965#page/449/mode/1up.
Goding, F. W. 1928. Membracidae of South
America and Antilles, III. Subfamily Mem-
bracinae. Journal of the New York Entomo-
logical Society 36: 201–234.
Lencioni-Neto, F. 2011. Riqueza de Especies e
Distribução Espacial dos Membracidae
(Hemiptera, Auchenorrhyncha) emu ma Peg-
uena Área do Campus Urbanova da Universi-
dade do Vale do Paraíba – SP, Brasil. Revista
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 16 Sep 2022
Terms of Use: https://bioone.org/terms-of-use Access provided by United States Department of Agriculture National Agricultural Library (NAL)
Univap, São José dos Campos-SP 17(29):
80–98. https://doi.org/10.18066/revunivap.
v17i29.14.
Lencioni-Neto, F. 2018. Membracideos no Vale
do Paraíba. Privately published. 185 pp.
Lencioni-Neto, F. and A. M. Sakakibara.
2015. A new species of Enchenopa (He-
miptera: Membracidae) from the Brazilian
Atlantic forest. Zoologia (Curitiba) 32(3):
257–259. http://dx.doi.org/10.1590/S1984-
46702015000300009.
Lencioni-Neto, F. and A. M. Sakakibara. 2016.
Description of the last nymphal instar of
Talipes appendiculatus (Hemiptera: Mem-
bracidae: Membracinae), and notes on the
biology of the immatures and adults. Zoo-
logia 33(6): 1–5. https://doi.org/10.1590/
s1984-4689zool-20160103.
McKamey, S. H. and L. L. Deitz. 1996. Generic
revision of the New World tribe Hoplophori-
onini (Hemiptera: Membracidae: Membraci-
nae). Systematic Entomology 21: 295–342.
Olivier, G. A. 1792. Membracis, pp. 657–669. In
Encyclopédie méthodique: Histoire naturelle
des animaux: Insectes [no editor given].
Panckoucke, Paris 7(2). 807 pp. https://www.
biodiversitylibrary.org/item/78652#page/299/
mode/1up.
Richter, L. 1947. Membracidae Colombianae.
Revisión de las espécies Colombianas del
género Membracis. Revista de la Academia
Colombiana de Ciencias Exactas, Físicas y
Naturales 7: 382–403.
Sakakibara, A. M. 1992. Sobre alguns Membracini
(Homoptera, Membracidae): notas taxonômi-
cas e descrições de gênero e espécies novos.
Revista Brasileira de Entomologia 36:
93–100.
Sakakibara, A. M. 2012. Taxonomic reassessment
of the treehopper tribe Talipedini with no-
menclatural changes and descriptions of new
taxa (Hemiptera: Membracidae: Membraci-
nae). Zoologia 29(6): 563–576. https://doi.
org/10.1590/S1984-46702012000600008.
Sakakibara, A. M. and O. Evangelista. 2010.
Membracis foliata (Linnaeus) (Hemiptera:
Membracidae: Membracinae) and allied spe-
cies: an effort towards their correct determi-
nation. Journal of Natural History 44(35–36):
2131–2148.
Stål, C. 1869a. Hemiptera Fabriciana. Fabricians-
ka Hemipterarter, efter de i Köpenhamn och
Kiel förvarade typexemplaren granskade och
beskrifne. 2. Svenska Vetenskaps-Akademiens
Handlingar 8(1): 1–130. https://www.biodiver-
sitylibrary.org/item/105468#page/9/mode/1up.
 VOLUME 124, NUMBER 1 97

Stål, C. 1869b. Bidrag till membracidernas
kannedom. Öfversigt af Kongliga Svenska
Vetenskaps-Akademiens Förhandlingar. 26 (3):
231–300.
Strümpel, H. and R. Strümpel. 2006. Revision of
the Neotropical treehopper genus Enchophyl-
lum (Hemiptera: Membracidae, Membraci-
nae). Mitteilungen aus dem Zoologischen
Museum Hamburg 14(175): 335–371. https://
www.zobodat.at/pdf/Ent-Mitt-Zool-Mus-
Hamburg_14_0335-0371.pdf.
Strümpel, H. and R. Strümpel. 2014. Revision
der amerikanischen Membracidengattung
Enchenopa (Hemiptera: Auchenorrhyncha:
Cicadomorpha: Membracidae) mit Besch-
reibungen neuer Arten. Entomologische
Mitteilungen aus dem Zoologischen Museum
Hamburg 17(191): 1–137. https://www.
zobodat.at/pdf/Ent-Mitt-Zool-Mus-Hamburg_
17_0001-0137.pdf.
Walker, F. 1858. Homoptera, pp. 1-117. In In-
secta saundersiana: or characters of unde-
scribed insects in the collection of William
Wilson Saunders (no editor given), Esq. John
Van Voorst. London 4: 1-117. https://www.
biodiversitylibrary.org/item/25015#page/69/
mode/1up.
Wallace, M. S. and L. L. Deitz. 2004. Phylogeny
and systematics of the treehopper subfamily
Centrotinae (Hemiptera: Membracidae). Mem-
oirs on Entomology, International 19: i–iv,
1–377.

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