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Phylogeny and biogeography of Neotropical spittlebugs (Hemiptera:

Cercopidae: Ischnorhininae): Revised tribal classification based on
morphological data

Article  in  Systematic Entomology · August 2014

DOI: 10.1111/syen.12091

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Systematic Entomology (2015), 40, 82–108 DOI: 10.1111/syen.12091

Phylogeny and biogeography of Neotropical spittlebugs

(Hemiptera: Cercopidae: Ischnorhininae): revised tribal
classification based on morphological data
A N D R E S S A P A L A D I N I 1, D A N I E L A M . T A K I Y A 2,
R O D N E Y R . C A V I C H I O L I 1 and G E R V Á S I O S . C A R V A L H O 3
1
Programa de Pós-graduação em Entomologia, Departamento de Zoologia, Setor de Ciências Biológicas, Universidade Federal do
Paraná, Curitiba, Brazil, 2 Laboratório de Entomologia, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Rio de
Janeiro, Brazil and 3 Programa de Pós-Graduação em Zoologia, Faculdade de Biociências, Pontifícia Universidade Católica do Rio
Grande do Sul, Porto Alegre, Brazil

Abstract. The spittlebug family Cercopidae is currently divided into two subfamilies:
the paraphyletic Old World Cercopinae and the monophyletic New World Ischnorhin-
inae. The most recent classification scheme proposed by Fennah in 1968 divided the
New World Cercopidae into four tribes: Tomaspidini, Ischnorhinini, Hyboscartini and
Neaenini. Herein we present a phylogenetic analysis of Ischnorhininae using 108 mor-
phological characters and including 53 of the 59 recognized genera, to evaluate the
tribal-level classification and understand the processes underlying the current distri-
butional patterns of these genera. We found significant support for the monophyly of
many Neotropical genera, but Fennah’s tribal classification is revised because tribes
Neaenini, Ischnorhinini and Tomaspidini were recovered as polyphyletic. Hyboscartini
was synonymized with Tomaspidini. A taxonomic key to tribes and genera of Neotropi-
cal spittlebugs is provided based mostly on recovered apomorphies. The biogeographical
analysis suggests a Neotropical origin of ischnorhinines, more specifically in northwest-
ern South America. This was possibly coincident spatially and temporally with the origin
of grasses, with ancestral range expansions southward to the Amazonian and Paraná
regions, and posterior vicariant events, possibly related to the expansion of forests in the
Chacoan region, the South America diagonal of open formations. Dispersals to the Cha-
coan region and to the Nearctic region are hypothesized to have occurred only within
genera. In the Chacoan region it is associated with more recent events, such as the diver-
sification of C4 grasses and establishment of the savannas and seasonally dry forests.

Introduction the Indo-Malayan, Afrotropical and Neotropical regions; 70%

Cercopoidea (spittlebugs), Cicadoidea (cicadas) and Membra-
coidea (leaf- and treehoppers) together comprise the hemipteran
infraorder Cicadomorpha (Hemiptera: Auchenorrhyncha). The
monophyly of each superfamily is strongly supported by mor-
phological (Evans, 1963; Blocker, 1996; Hamilton, 1999) and
molecular (Cryan, 2005; Cryan & Urban, 2012) evidence. With
approximately 3000 described species classified among 320
genera, 34 tribes and 5 families, spittlebug diversity is highest in
Correspondence: Andressa Paladini, Universidade Federal do Paraná,
CxP. 19020, 81531–980 Curitiba – PR. E-mail: andri_bio@yahoo.
com.br
of spittlebug species have a tropical distribution. Spittlebug
nymphs develop inside spittle masses on their host plants (as pro-
tection from desiccation, predation and/or parasitism), whereas
the adults are free-living. Characteristics (consistency, viscos-
ity, physical placement, etc.) of the spittle masses seem to vary
across spittlebug families (Rakitov, 2002). Spittlebugs are also
known as froghoppers, because of their prodigious jumping abil-
ity, reaching heights of 115× the body length (Burrows, 2006).
Members of the Cercopidae, the largest cercopoid family,
are distributed worldwide and currently in the New World
there are 431 species in 59 genera (Carvalho & Webb, 2005;
Paladini & Cryan, 2012). Cercopids feed on the xylem sap

82 © 2014 The Royal Entomological Society


of a large variety of plants, and in the New World they feed
principally on grasses (Carvalho & Webb, 2005). Due to this
habit, several species of Aeneolamia, Deois, Isozulia, Kanaima,
Mahanarva, Maxantonia, Notozulia, Prosapia, Sphenorhina,
Tunaima and Zulia are responsible for significant economic
damage to pastures and sugarcane crops throughout South and
Central America (Valério & Koller, 1992). Adult feeding causes
phytotoxemia evidenced by leaf chlorosis and severe spittlebug
outbreaks can result in the yellowing of the entire aboveground
portion of the plant (Carvalho & Webb, 2005).
Several classifications for spittlebugs have been proposed
through the years by Lallemand (1912, 1949), Lallemand &
Synave (1961), Metcalf (1961, 1962), and Fowler & Cockerell
(1894–1909), but the currently most widely adopted classifica-
tion was proposed by Fennah (1968, 1979) in his revisionary
study. Fennah (1968) divided Cercopidae into two subfamilies:
the Old World Cercopinae, characterized by having a single lat-
eral spine on the posterior tibia, and the New World Tomaspidi-
nae, with two lateral spines on the posterior tibia. Tomaspidinae
was further divided into four tribes: Tomaspidini, characterized
by an inflated postclypeus (Fig. 1L) and strong apical reticula-
tion of the forewing venation (Fig. 2A); Ischnorhinini, charac-
terized by strong lateral compression of the postclypeus, which
bears one lateral carina extending towards the mandibular plate
(Fig. 2I); Hyboscartini, characterized by a median furrow or
weak depression on the postclypeus bordered by a small pair
of prominences, weak apical reticulation of the forewing vena-
tion (Fig. 2C), ovoid basal body of the flagellum, and hind
basitarsus with about 11 apical spines; and Neaenini [originally
described as a subfamily of Aphrophoridae (Metcalf, 1939)],
characterized by a longitudinal groove on the postclypeus with
a pair of prominences, weak apical reticulation in the forewing
venation, subglobose basal body of the flagellum, and basitar-
sus with fewer than 11 apical spines. In their recent checklist
of New World Cercopidae, Carvalho & Webb (2005) proposed
the usage of Ischnorhininae Schmidt, 1920 instead of Tomas-
pidinae Schmidt, 1922 based on the principles of priority and
coordination (ICZN, 1999), given that Schmidt (1920) first pro-
posed the family-group name Ischnorhininae. However, in this
checklist, the authors declined to adopt a tribal classification,
citing inconsistencies in characters used by previous researchers
(Lallemand, 1949; Fennah, 1968; Costes, 1971; Carvalho, 1992)
to define tribes.
Recently, a molecular phylogeny of Cercopoidea (Cryan &
Svenson, 2010) confirmed the monophyly of Ischnorhininae,
concluding that the New World lineage arose from a paraphyletic
assemblage of Old World clades. Based on their molecular dat-
ing estimates, the origin of Cercopidae dates from the Early Cre-
taceous (122.78–168.34 Ma), with the first fossil record from
the mid-Cretaceous (Shcherbakov & Popov, 2002; Shcherbakov,
2002), and the divergence of Ischnorhininae from the related
Old World lineage occurred during the mid-Cretaceous, between
91.06 and 134.02 Ma. During this time period, continental
Africa would have been separating from northern South Amer-
ica, isolating ischnorhinines from the remaining cercopid lin-
eages (Cryan & Svenson, 2010; Ronquist & San Martín, 2011).
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
Neotropical spittlebug phylogeny and biogeography 83
However, their taxon sampling did not allow for rigorous test-
ing of tribal classifications proposed by earlier workers (Metcalf,
1961; Fennah, 1968). Thus, tribal-level classification of Cercop-
idae remains problematic.
Based on morphological data, a phylogenetic hypothesis is
proposed for the genera of New World spittlebugs to evaluate
their current tribal-level placement as proposed by Fennah
(1968) and to reconstruct ancestral areas of distribution based
on the current distribution of terminals, in order to infer possible
vicariance or dispersal events in between the major endemism
areas of South and Central America.
Material and methods
Taxon sampling
We used species as terminals for the phylogenetic analy-
sis and included representatives of most of the 59 genera of
Ischnorhininae. Representative species were selected based on
the availability of exemplars, giving preference, when avail-
able, to the type species (Table 1). Six monotypic genera of
Ischnorhininae were not included, Bradypteroscarta, Chinana,
Hyalotomaspis, Lujana, Hemiplagiophleboptena and Plagio-
phleboptena, because they are known only from holotypes of
type species. At least two species of each genus were sam-
pled, in order to sample the morphological variation within
each genus (Prendini, 2001). The inadequacy of including a
single exemplar species from each higher taxon, rather than
all or even a few species per higher taxon, has been demon-
strated by Wheeler (1992). Inclusion of at least two exemplars
may better distinguish character states that are autapomorphic or
homoplasious from those that are synapomorphic for the higher
taxa which the exemplars represent (Wheeler et al., 1993).
Three species of uncertain position within Ischnorhininae were
included: Tomaspis biolleyi, Tomaspis handlirschi and Hemit-
omaspis minuscula. Based on the single phylogenetic estimate
of cercopoids, Cercopinae is paraphyletic in relation to a mono-
phyletic Ischnorhininae (Cryan & Svenson, 2010). An attempt
to include few representatives of this subfamily was hindered by
the difficulty in hypothesizing homologies between them and
the ingroup. Given that Cercopinae includes more than 1000
species, is understudied and in need of a modern taxonomic
treatment, it may be misleading to include just a small sam-
ple of terminals of this lineage without proper selection, based
on the complete morphological variation. Thus, only species of
Aphrophoridae were included as outgroups and used to root the
cladograms (Nixon & Carpenter, 1993).
Morphological study
The morphological study was carried out through the examina-
tion of dry pinned specimens of both sexes using a stereoscopic
microscope. Genitalia were detached from the abdomen and
cleared in hot 10% KOH solution for approximately 5–10 min
and washed in 70% alcohol and acetic acid, and subsequently
84 A. Paladini et al.

Table 1. Neotropical Cercopidae (=Ischnorhininae) tribes, according to Fennah (1968), and genera, including their number of species in parenthesis,
geographic distribution and species sampled in this study.

Tribe Genus Distribution Species sampled

Hyboscartini Chinana (1) Argentina –

Hemiplagiophleboptena (1) Bolivia –
Hyboscarta (4) Ecuador, Brazil, Bolivia H. melichari
Plagiophleboptena (2) Bolivia, Ecuador P. smaragdina
Ischnorhinini Baetkia (3) Brazil, French Guiana, Suriname B. compressa*, B. maroniensis
Homalogrypota (3) Brazil, Peru, Bolivia H. coccinea*, H. interrupta
Ischnorhina (7) Brazil, Bolivia, Ecuador I. amazonica, I. ephippium*, I. hamiltoni,
I. surinamensis
Laccogrypota (18) Brazil, Peru, Colombia, Ecuador, French L. valida*, L. inca
Guiana
Neolaccogrypota (1) Ecuador N. brunnea*
Neosphenorhina (5) Brazil N. ocellata*, N. schombergi
Schistogonia (5) Brazil S. bidentata, S. sanguinea*
Sphenoclypeana (2) Brazil S. haematina*, S. parana
Tiodus (6) Panama, Colombia, Peru T. nigricans
Typeschata (3) Brazil T. marginata*
Tomaspidini Aeneolamia (8) Brazil, Mexico, Honduras, Guatemala, A. flavilatera, A. colon
Trinidade, French Guiana, Colombia,
Granada
Aracamunia (1) Bolivia A. dimorpha*
Bradypteroscarta (1) Ecuador –
Carachata (1) Brazil C. dimorphica*
Carpentiera (1) Bolivia, Peru C. insignis*
Catrimania (5) Brazil, Peru C. semivitrea*, C. longula
Choconta (5) Colombia, Bolivia, Venezuela, Peru C. elliptica*
Deois (24) Brazil, Argentina D. flavopicta, D. terrea*, D. flexuosa,
D. schach
Deoisella (2) Brazil D. fasciata*, D. picklesi
Ferorhinella (2) Brazil F. brevis*, F. balatra
Hemitomaspis (3) Peru, Bolivia, Panama H. minuscula
Huaina (1) Mexico, Costa Rica H. inca*
Hyalotomaspis (1) Mexico –
Iphirhina (5) Mexico, Guatemala, Costa Rica I. quota*, I. perfecta
Isozulia (6) Paraguay, Bolivia, Argentina, Colombia, I. astralis, I. soluta
Ecuador, Peru
Kanaima (3) Brazil, Argentina K. katzensteinii*, K. fluvialis
Korobona (1) Brazil K. lineata*
Lujana (1) Brazil –
Mahanarva (46) Brazil, Uruguay, Ecuador, Peru, Colombia, M. quadripunctata, M. liturata,
Costa Rica, Nicarágua, New Granada, M. rubripennis, M. integra
Paraguay, Gorgona Island, Panama,
Bolivia, Venezuela
Makonaima (2) Peru, Ecuador M. lucifera
Maxantonia (36) Brazil, Ecuador, Colombia, Peru, Bolivia, M. rubescens, M. quadriguttata*
Panama
Monecphora (9) Brazil, French Guiana, Peru, Costa Rica M. semilutea, M. cingulata*, M. nigritarsis,
M. sp.n., M. longitudinalis
Neomonecphora (6) Brazil, French Guiana N. laurentana, N. apicalis
Notozulia (1) Argentina, Brazil, Paraguay N. entreriana*
Ocoaxo (30) Mexico, Costa Rica, Ecuador, Panama, O. lineatus*, O. assimilis
Colombia, United States
Olcotomaspis (1) Guatemala –
Orodamnis (1) Venezuela O. rynchosporae*
Pacachantocnemis (1) Costa Rica, Colombia, New Granada P. bella*
Panabrus (1) Dominica P. dominicanus*
Prosapia (15) Mexico, Costa Rica, Guatemala, Cuba, P. bicincta*, P. ignifera
Jamaica, Panama, United States
Sinopia (1) Brazil, French Guiana S. signata*

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108

 Neotropical spittlebug phylogeny and biogeography 85

Table 1. Continued.

Tribe Genus Distribution Species sampled

Sphenorhina (60) Ecuador, Brazil, Bolivia, Peru, Paraguay, S. rubra*, S. coronata, S. parambae
Costa Rica, Colombia, Venezuela,
Tomaspis (1) Brazil T. furcata*
Tomaspisina (3) Colombia, Peru –
Tropidorhinella (4) Bolivia, Brazil, Colombia T. onorei
Tunaima (13) Brazil, Colombia T. brunneorubra*, T. semiflava, T. insignifica
Urubaxia (1) Brazil U. tricolor*
Vorago (5) Venezuela, Brazil, Guyana, Ecuador V. boxi
Zulia (8) Paraguay, Bolivia, Ecuador, Brazil, Z. birubromaculata, Z. morosae,
Venezuela Z. pubescens*
Neaenini Marcion (1) Venezuela M. equestris*
Neaenus (1) Mexico N. varius*
Simorhina (1) Bolivia, Peru S. sciodes*
Tapaiuna (3) Venezuela, Costa Rica, Ecuador T. pehlkei
Tomaspisinella (2) Panama, Ecuador T. apicifasciata
Zuata (30) Bolivia, Venezuela, Peru, Colombia, Z. seguyi
Ecuador

Species marked with an asterisk (*) are type species.

stored in vials with glycerin. Species were identified using taxo-
nomic keys (Fennah, 1968; Paladini & Carvalho, 2008; Paladini
et al., 2008b, 2010), original descriptions and photographs of
type-specimens.
Illustrations of relevant structures were made with a camera
lucida and transformed into digital vectors with Corel Draw
vX5. Photographs were taken with a Leica DFC-550 digital
camera attached to the stereoscopic microscope (Leica MZ16).
Images were captured with IM50 software (Image Manager;
Leica Microsystems Imaging Solutions Ltd, Cambridge, UK)
and stacked with the Auto-Montage software of Taxonline (Rede
Paranaense de Coleções). Photographs were also taken with
the JVC KYF75U digital camera attached to the stereoscopic
microscope LEICA Z16APO and mounted with the software
Automontage PRO 54040013 in the New York State Museum
(Albany, NY). Scanning electron microscope images (SEM unit
Philips model XL 30) were taken at the Centro de Microscopia e
Microanálises (CEMM) of the Pontifícia Universidade Católica
do Rio Grande do Sul (PUCRS).
Character coding
Characters were coded to include most of the morphologi-
cal variation. Some characters were taken from the literature
(Distant & Fowler, 1881–1905; Fennah, 1968; Paladini et al.,
2008b); other characters were based on our own observations
of adult males and females. A total of 108 characters were
coded, including features of the external morphology (characters
1–50), male genitalia (characters 51–99) and female genitalia
(characters 100–108).
Each character was considered a hypothesis of grouping (taxic
homology sensu Patterson, 1982). Primary homologies were
proposed by similarity or topological correspondence (de Pinna,
1991). We used contingent coding when novel features appeared
and evolved, and this feature shows variation (Sereno, 2007).
Multistate characters were treated as unordered (nonadditive)
under Fitch parsimony (Fitch, 1971). Character state polarity
was determined by rooting cladograms with an outgroup (Nixon
& Carpenter, 1993).
Character statements were composed of four fundamental
functional components following Sereno (2007): locator, vari-
able, variable qualifier and character state. Missing data were
coded as ‘-’ and nonapplicable characters were coded as ‘?’. The
data matrix was built using Winclada v1.00.08 (Nixon, 2002).
Cladistic analysis
Searches for the most parsimonious trees were conducted
using two character-weighting schemes: equal weights and
implied weights. In the implied weights scheme, characters
are analysed with weights inversely proportional to their level
of homoplasy (Goloboff, 1993). The constant K determines
how strongly characters will be weighted against homoplasy;
however, there is no well-justified criterion by which to
choose particular values of K, and this decision is probably
matrix-dependent (Goloboff, 1993). Therefore, there is a need
to explore different concavities and only groups present in
the results with all values explored for the concavity constant
should be used (Goloboff et al., 2008b). For the analysis with
implied weights we used 20 values of K (1–20).
The choice for the best topological hypotheses generated
from the 20 different implied weighting schemes was based on
their stability. A tree is considered more stable if nodes are
less parameter-dependent (Giribet, 2003), and therefore best
hypotheses were chosen if their topology were more similar
to remaining ones (Mirande, 2009). Measurements used to
compare all topologies were the distortion coefficient (Farris,
1989), which compares the retention index of each tree to every

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108

86 A. Paladini et al.
other tree, and SPR distance (Goloboff et al., 2008a), which
calculates the minimum number of SPR movements required to
transform each tree in every other tree.
Analyses were made using TNT v1.1 (Goloboff et al., 2008a)
using strategies of Traditional and New Technology Searches
to find most parsimonious trees. Tree search strategies used
were: (1) Traditional search with 1000 replications with TBR
algorithm, 99 trees saved per replication; (2) Traditional search
with 10 000 replications with TBR, nine trees saved per repli-
cation; (3) New technology search with Ratchet (Nixon, 1999)
with perturbation phase adjusted to 25 and 5 for up-weighting
and down-weighting and 20 iterations; Tree drifting (Goloboff,
1999) with 20 cycles and Tree Fusing (Goloboff, 1999) with 100
rounds; and (4) New technology search with 200 iterations of
Ratchet (Nixon, 1999) with the perturbation phase adjusted to 8
for both up-weighting and down-weighting, Sectorial search in
default mode, 20 cycles of Tree drifting, and 10 rounds of Tree
fusing, until the best fit was obtained three times. Each round
of analysis was performed until the minimum tree length was
hit 100 times. Strategies 1–3 were used in analyses with equal
weights and strategy 4 only for implied weights.
Branch support was calculated using the relative Bremer
support or decay index (Bremer, 1994) implemented on TNT
(Goloboff et al., 2008b) under implied weighting schemes.
Nonparametric bootstrap support values were computed under
a maximum likelihood framework based on the Mkv model
for morphological characters (Lewis, 2001) with GARLI v2.01
(Zwickl, 2006) by running 100 000 generations for each 100
bootstrap pseudoreplicate.
Biogeographic areas
Distributional data for species were taken from the literature
and directly from specimen label data. Eight major areas
(Fig. 11A) of the Nearctic and Neotropical regions defined in
previous biogeographic analyses (Morrone, 2006, 2013) were
used in this analysis.
The Nearctic region (A) comprises cold temperate areas of
North America and northern Mexico. The five Latin Ameri-
can provinces in the Nearctic region are in Mexico with the
exception of the Baja California province that extends north-
wards into the United States. The Mexican transition zone
(B) is a complex and variable area where the Nearctic and
Neotropical biotic elements overlap and comprises the south-
western U.S.A., Mexico and a large part of Central America,
extending into the Nicaraguan lowlands. The Neotropical region
was further divided into three dominions of the Caribbean sub-
region, and the three remaining subregions. The Mesoamerica
dominion (C) covers the lowlands of the Mexican Gulf and
Pacific coast; the Antillean dominion (D) extends to areas of
the Caribbean islands; and the Northwestern South America (E)
dominion comprises the Pacific coast of Ecuador, Colombia and
Panama; northern Colombia, Venezuela and Peru; and Trinidad
and Tobago. The Amazonian subregion (F) includes lowland
tropical forest regions of northern Brazil, Guyana, Venezuela,
Colombia, Ecuador, Peru, Bolivia and Argentina. The Chacoan
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
subregion (G) comprises the diagonal of open formations of
South America (an area extending across South America, from
northeastern Brazil to central north Argentina, basically com-
prising the Caatinga, Cerrado and Chaco) and occupies north-
ern and central Argentina, southern Bolivia, western and central
Paraguay, Uruguay, and central and northeastern Brazil. The
Parana subregion (H) includes areas of tropical and subtrop-
ical forests and is situated in northeastern Argentina, eastern
Paraguay, and southern and eastern Brazil. Finally, the South
America transition zone (I) extends along the highlands of the
Andes between western Venezuela, northern Chile and west cen-
tral Argentina.
Biogeographical analysis
Biogeographical methods based on reticulate discrete models
are often being used in historical biogeographical studies (Ron-
quist & San Martín, 2011) given that they allow vicariant events
between all combinations of areas of potential ancestral distri-
butions. An event-based parsimony method was chosen herein
to reconstruct the ancestral areas of distribution of Ischnorhini-
nae and infer possible important vicariant and dispersal events.
A distinctive characteristic of event-based methods is the pro-
posal of explicit models for the different types of processes (e.g.
vicariance, dispersal and extinction) that affect the geographic
distribution of living organisms (Crisci et al., 2003). Ances-
tral distributions of Ischnorhininae were reconstructed using
a Dispersal-Vicariance Analysis (DIVA) (Ronquist, 1997), but
incorporating phylogenetic uncertainty by evaluating ancestral
areas of all possible most stable parsimonious trees recovered by
the weighted analysis (Nylander et al., 2008). S-DIVA analyses
were run in Reconstruct Ancestral State in Phylogenies (RASP)
(Yu et al., 2013) with a maximum of six areas at each node and
frequencies of an ancestral range at a node are averaged over all
trees, and each alternative ancestral range at a node is weighted
by the frequency of which the node occurs (Yu et al., 2010).
Results
Morphological characters
Genera of Ischnorhininae are defined by structures of the head,
pronotum, number of apical spines of the posterior tibia and
wing venation. These characters are present in the key of Fennah
(1968) and are the basis for the classification of tribes proposed
here. Unfortunately, most of these characters are homoplasious
and some are polymorphic within species or even in individuals.
Polymorphic characters were not used in the analysis. According
to Fennah (1968) characters of the genitalia are considered less
influenced by parallel evolution, but even these characters are
extremely homoplastic.
The postclypeus may be strongly compressed laterally with
one lateral carina on the inferior angle in Ischnorhinini (Fig. 2I)
or it may be inflated as in Tomaspidini, sometimes swollen as
in Tomaspis furcata (Fig. 1L). In profile the postclypeus may be
 Neotropical spittlebug phylogeny and biogeography 87

Fig. 1. Ischnorhininae head, lateral view: (A) Tomaspis biolleyi; (B) Vorago boxi; (C) Tropidorhinella onorei; (D) Carpentiera insignis; (E) Prosapia
bicincta; (F) Zuata carvalhoi; (G) Homalogrypota coccinea; (H) Sphenorhina rubra; (I) Maxantonia quadriguttata; (J) Simorhina sciodes; (K) Kanaima
katzensteinii; (L) Tomaspis furcata.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108

88 A. Paladini et al.

Fig. 2. Characters of the head and thorax. Forewing: (A) Maxantonia sp.; (B) Huaina inca; (C) Simorhina sciodes. Hindwing: (D) Maxantonia sp. Head
and pronotum, dorsal view: (E) Zuata carvalhoi ; (F) Mahanarva liturata; (G) Deois incompleta; (H) Hemitomaspis minuscula. Head and pronotum,
lateral view: (I) Typescata marginata; (J) Sphenoclypeana parana. Head and postclypeus, ventral view: (K) Schistogonia bidentata; (L) Sphenoclypeana
parana; (M) Simorhina sciodes; (N) Deois flexuosa; (O) Hyboscarta melichari; (P) Maxantonia rubescens.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108


convex (Fig. 1C) or angulate (Fig. 1H), if angulate, the angle
may vary from straight, as in Ischnorhinini (Fig. 1A, G), to
obtuse, as in Mahanarva, Maxantonia and Prosapia (Fig. 1E,
I). The median carina of the postclypeus varies from narrow,
as in most of the genera of Cercopidae that have a median
carina (Fig. 2L), to wide, as in Hyboscarta (Fig. 2O), and the
number of carinae may also vary. For example, both sexes of
Maxantonia have three longitudinal carinae on the postclypeus
(Fig. 2P), a synapomorphy for the genus. The postclypeus may
also have one pair of small protuberances in the inferior margin.
Fennah (1968) assigned this character to the tribes Neaenini
and Hyboscartini, but it is also present in Schistogonia which
is placed in Ischnorhinini. Fennah assumed that the postclypeus
characters are useless for Cercopidae taxonomy because of
the occurrence of intermediary conditions or similar forms
resulting from parallel evolution. However, we choose to use
these characters in our phylogeny because some of them, even
if homoplasious, have a high retention index and are useful for
clade support.
The rostrum varies in length, and this variability may be
found between genera or subgenera. For example, in Mahanarva
(Mahanarva) the rostrum reaches the mesocoxae, whereas in
Mahanarva (Ipiranga) the rostrum is shorter and does not reach
the mesocoxae.
The ocelli are always situated nearer to each other than to
the eyes. However, the distance between them (measured here
based in the diameter of one of the ocelli) and their position
with respect to the posterior margin of the head (comparing the
position of the ocelli related to the imaginary line that passes
through the eyes) vary among the genera.
Although the antennae of Cercopidae always have a long
flagellum and an arista situated above the flagellum (Figs 8A,
9A–C), there are variations in antennal structure among gen-
era and in some cases among species. The characters included
here are: the shape of the basal body of the flagellum, the size
and presence of the arista (a basiconic sensillum described by
Liang & Fletcher, 2002, and Liang, 2001), and the number
of setae on the scape. Setae (trichoid sensilla; Fig. 9A) may
vary from 40 to 60 μm, may be straight or slightly curved,
and are located near the base of the pedicel or rarely on
other areas of the antenna. The basal body of the flagellum
varies in shape (it may be subcylindrical, conic or ovoid)
and has many sensilla (coeloconic and campaniform) vary-
ing in number and arrangement (Fig. 9C–E). According to
K.G.A. Hamilton (personal communication) characters per-
taining to the sensilla are very important to the phylogeny
of Cercopidae.
The pronotum of Cercopidae is hexagonal. Variation is
found in the shape of the antero-lateral margins, which can
be convex or straight. In most species the posterior mar-
gin is grooved, but in some cases it is straight. The pres-
ence of a median carina exhibits interspecific variation. A
pronotum with a curved profile is characteristic of some gen-
era: Tapaiuna, Simorhina and Makonaima. This condition is
usual in Old World Cercopids (Cercopinae), as can be seen
in Cosmoscarta.
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
Neotropical spittlebug phylogeny and biogeography 89
With rare exceptions, the posterior tibia of Ischnorhininae
has two lateral spines, diagnostic characteristic used by Fennah
(1968). This character is difficult to use, because some genera
(e.g. Makonaima lucifera, Tapaiuna and Neomonecphora) of
Neotropical Cercopidae have only one spine or the basal spine
can be small and inconspicuous (almost imperceptible) as in
most Ischnorhinini. The basitarsus has an apical crown of spines.
Fennah (1968) used the number of these spines as a character,
but they exhibit considerable intraspecific variation, so we chose
instead to use as characters the number of rows of spines on the
basitarsus and the number of setae.
The tegmina vary in the ratio of length to width, and devel-
opment of the venation and the apical plexus of veins. In some
genera (e.g. Mahanarva, Prosapia, Kanaima and Maxantonia)
many species are polymorphic. Some species have mimetic
forms. For example, the colour pattern of Laccogrypota valida
is identical to that of Maxantonia mimica yet these two species
belong to different genera and tribes.
Many of the characters used are from the male genitalia.
According to Fennah (1968), the pygofer and seventh and eighth
sternites of the male abdomen in Ischnorhininae are less variable
than in Cercopinae. In Ischnorhininae the pygofer is short and
the subgenital plates are fused with it along the basal margin
and consequently are immobile. The aedeagus varies in shape,
number of processes, insertions of these processes, denticulation
and curvature; the paramere also varies in shape, number of
spines and shape of the dorsal margin. The male genitalia have
interspecific variation but the general pattern is mostly stable at
the generic level.
Finally, in the female genitalia the basal process of the first
valvulae of the ovipositor varies in size, shape and orientation.
In the second valvulae of the ovipositor the extent of dentition
on the dorsal margin also varies. In some genera (e.g. Tomaspis
and Korobona) the first valvulae of the ovipositor has teeth on
the ventral margin.
Head
(1) Tylus, median carina: (0) absent (Fig. 3B), (1)
present (Fig. 3A). CI = 0.05, RI = 0.63
(2) Vertex, median carina: (0) absent (Fig. 3B), (1)
present (Fig. 3A). CI = 0.04, RI = 0.43
(3) Vertex, shape: (0) narrow (rectangular) (Fig. 3B), (1)
wide (quadrangular) (Fig. 3A). CI = 0.07, RI = 0.6
(4) Ocelli, distance between them measured in ocellar
diameters: (0) one (Fig. 2E), (1) two (Fig. 2H), (2) more.
CI = 0.14, RI = 0.4
(5) Ocelli, position relative to imaginary line through
the middle of the eyes: (0) anterad, (1) on, (2) posterad.
CI = 0.09, RI = 0.59
(6) Antennae, shape of the basal body of flagellum: (0)
conical (Fig. 9D), (1) ovoid (Fig. 9C), (2) subcylindrical.
CI = 0.11, RI = 0.69
(7) Antennae, size of basal body of flagellum relative
to pedicel: (0) larger (Fig. 8B), (1) smaller (Fig. 8A), (2)
equal. CI = 0.33, RI = 0.8
90 A. Paladini et al.

Fig. 3. Characters of the head, thorax and abdomen: Head, dorsal view: (A) Kanaima fluvialis; (B) Sphenorhina rubra. Subgenital plate, ventral view:
(C) Baetkia maroniensis; (D) Simorhina sciodes; (E) Neomonecphora apicalis; (F) Ocoaxo sp.; (G) Tunaima insignifica. Aedeagus and paramere: (H)
Simorhina sciodes; (I) Hyboscarta melichari; (J) Deois flexuosa, posterior leg, lateral view.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108


(8) Antennae, length and density of setae on pedicel:
(0) long and abundant (Fig. 9A), (1) short and scarce
(Fig. 9B). CI = 0.2, RI = 0.7
(9) Antennae, arista of basal body of flagellum:
(0) absent (Fig. 9D), (1) present (Fig. 9C). CI = 0.16,
RI = 0.54
(10) Antennae, length of arista related to pedicel:
(0) larger, (1) equal (Fig. 9B), (2) smaller. CI = 0.18,
RI = 0.72
(11) Postclypeus, shape in ventral view: (0) inflated
(Fig. 2N), (1) compressed (Fig. 2L), (2) flattened.
CI = 0.33, RI = 0.82
(12) Postclypeus, lateral compression: (0) strong
(Fig. 2L), (1) weak (Fig. 2K). CI = 1, RI = 1
(13) Postclypeus, shape in frontal view: (0) rectangular
(Fig. 2K), (1) quadrangular (Fig. 2M), (2) rounded.
CI = 0.66, RI = 0.9
(14) Postclypeus, recession in profile: (0) receding
backward (Fig. 1K), (1) not receding (Fig. 1B), (2)
projected forward (Fig. 1C, D). CI = 0.2, RI = 0.46
(15) Postclypeus, shape of profile: (0) angulate
(Fig. 1A), (1) convex (Fig. 1L). CI = 0.07, RI = 0.69
(16) Postclypeus, shape of angle: (0) acute or right
(forming a vertex) (Fig. 1A), (1) obtuse (Fig. 1I).
CI = 0.05, RI = 0.92
(17) Postclypeus, swelling: (0) absent (Fig. 1J), (1)
present (Fig. 1L). CI = 0.14, RI = 0.57
(18) Postclypeus, inferior margin, lateral carina: (0)
absent (Fig. 2J), (1) present (Fig. 2I). CI = 0.33, RI = 0.9
(19) Postclypeus, inferior margin, lateral carina:
(0) slightly delimited, (1) strongly delimited (Fig. 2I).
CI = 0.5, RI = 0.8
(20) Postclypeus, longitudinal carina in frontal view:
(0) absent (Fig. 2M), (1) present (Fig. 2K). CI = 0.25,
RI = 0.84
(21) Postclypeus, longitudinal carina: (0) prominent
(Fig. 2K), (1) not prominent (Fig. 2N). CI = 0.2, RI = 0.5
(22) Postclypeus, thickness of longitudinal carina: (0)
wide (Fig. 2O), (1) narrow (Fig. 2K). CI = 1, RI = 1
(23) Postclypeus, number of longitudinal carinae: (0)
one (Fig. 2K), (1) three (Fig. 2P). CI = 1, RI = 1
(24) Postclypeus, lateral grooves: (0) absent, (1)
present. CI = 0.33, RI = 0.33
(25) Postclypeus, lateral grooves: (0) slightly delimited
(Fig. 1G), (1) strongly delimited (Fig. 1K). CI = 0.08,
RI = 0.62
(26) Postclypeus, triangle formed in apex with tylus:
(0) absent (Fig. 2N), (1) present (Fig. 2K). CI = 0.25,
RI = 0.83
(27) Postclypeus, pair of lateral prominences on infe-
rior margin: (0) absent (Fig. 2L), (1) present (Fig. 2K).
CI = 0.2, RI = 0.42
(28) Rostrum, length relative to mesothoracic coxae:
(0) barely reaching base of mesocoxae, (1) reach-
ing mesocoxae, (2) surpassing mesocoxae. CI = 0.12,
RI = 0.61
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
Neotropical spittlebug phylogeny and biogeography 91
Thorax
(29) Pronotum, proportion of length of anterolateral
margin to half-length anterior margin: (0) smaller
(Fig. 2E, H), (1) larger, (2) equal (Fig. 2F, G). CI = 0.14,
RI = 0.76
(30) Pronotum, shape of anterolateral margin: (0) con-
vex (Fig. 2F), (1) straight (Fig. 2E). CI = 0.12, RI = 0.63
(31) Pronotum, posterior margin: (0) slightly grooved
(Fig. 2E), (1) strongly grooved (Fig. 2F), (2) straight.
CI = 0.04, RI = 0.44
(32) Pronotum, median carina: (0) absent (Fig. 2H), (1)
present (Fig. 2F). CI = 0.05, RI = 0.55
(33) Pronotum, median carina: (0) complete, (1)
reduced. CI = 0.1, RI = 0.55
(34) Pronotum, shape of profile in lateral view: (0)
curved (Fig. 1J), (1) not curved (Fig. 1K). CI = 0.14,
RI = 0.4
(35) Pronotum, shape of the humeral angles: (0)
obtuse, (1) rounded. CI = 0.08, RI = 0.5
(36) Tegmina, ratio of length/width: (0) narrow (L/W
= >3) Fig. 2C, (1) wide (L/W = <2.9) Fig. 2B. CI = 0.05,
RI = 0.56
(37) Tegmina, venation: (0) visible but not elevated, (1)
prominent. CI = 0.06, RI = 0.41
(38) Tegmina, vein A2 : (0) indistinct, (1) distinct.
CI = 0.06, RI = 0.42
(39) Tegmina, apical plexus: (0) present (Fig. 2A), (1)
absent. CI = 0.14, RI = 0.45
(40) Tegmina, development of the apical plexus: (0)
developed (Fig. 2A, B), (1) reduced (Fig. 2C). CI = 0.14,
RI = 0.57
(41) Posterior wings, thickness of base of vein Cu1 :
(0) thickened, (1) not thickened (Fig. 2D). CI = 0.09,
RI = 0.55
(42) Posterior leg, number of lateral spines on tibia:
(0) one, (1) two (Fig. 3J). CI = 0.25, RI = 0.61
(43) Posterior leg, size of basal spine of tibia compared
to those of apical crown: (0) larger, (1) equal, (2) smaller.
CI = 0.11, RI = 0.64
(44) Posterior leg, size of apical spine of tibia com-
pared to those apical crown: (0) equal, (1) larger.
CI = 0.1, RI = 0.3
(45) Posterior leg, spine on inner face of femur: (0)
present, (1) absent. CI = 0.11, RI = 0.61
(46) Posterior leg, size of spine on inner face of femur:
(0) conspicuous, (1) inconspicuous. CI = 0.06, RI = 0.58
(47) Posterior leg, number of rows of spines on
the basitarsus: (0) one, (1) two, (2) three. CI = 0.11,
RI = 0.52
(48) Posterior leg, density of setae on the basitarsus:
(0) sparse, not concealing spines of apical crown, (1)
dense, concealing spines of apical crown. CI = 0.12,
RI = 0.56
(49) Posterior leg, subungueal process: (0) present
(Fig. 9F, G), (1) absent. CI = 0.06, RI = 0.42
92 A. Paladini et al.
(50) Posterior leg, shape of subuengueal process: (0)
clawlike (Fig. 9G), (1) triangular, (2) rectangular (Fig. 9F)
CI = 0.25, RI = 0.77
Male abdomen
(51) Pygofer, process on lateral margin between anal
tube and subgenital plates: (0) absent (Fig. 4P), (1)
present (Fig. 4O). CI = 0.2, RI = 0.42
(52) Pygofer, number of process on lateral margin
between anal tube and subgenital plates: (0) one
(Fig. 4C), (1) two (Fig. 4E). CI = 0.2, RI = 0.5
(53) Subgenital plates, length compared to pygofer:
(0) short (Fig. 4O), (1) long (Fig. 4I). (2) very long
(Fig. 4A). CI = 0.25, RI = 0.79
(54) Subgenital plates, fusion: (0) fused only at base
(Fig. 3C), (1) free for most of length (Fig. 3F, G), (2) fused
for most of length (Fig. 3D, E). CI = 0.14, RI = 0.65
(55) Subgenital plate, basal process: (0) absent
(Fig. 4C), (1) present (Fig. 4B). CI = 0.11, RI = 0.52
(56) Subgenital plate, shape of basal process: (0)
elongate (Fig. 4D), (1) short (Fig. 4L). CI = 1, RI = 1
(57) Subgenital plate, shape of basal process: (0)
acuminate (Fig. 4A), (1) not acuminate (Fig. 4D).
CI = 0.33, RI = 0.71
(58) Subgenital plate, shape of dorsal margin: (0)
rounded (Fig. 4I), (1) straight (Fig. 4K), (2) with acute
process (Fig. 4M). CI = 0.1, RI = 0.62
(59) Subgenital plate, spine at apex: (0) absent (Fig. 4H,
Q), (1) present (Fig. 4B). CI = 0.07, RI = 0.45
(60) Subgenital plate, shape of apex: (0) finger-like
(Fig. 4M), (1) acute (Fig. 4R), (2) truncated (Fig. 4O),
(3) rounded (Fig. 4F, J). CI = 0.12, RI = 0.56
(61) Subgenital plate, denticuli on inner face: (0)
absent (Fig. 4G), (1) present (Fig. 4H). CI = 0.07,
RI = 0.48
(62) Subgenital plate, position of denticuli on inner
face: (0) spread around, (1) only on the dorsal margin.
CI = 0.33, RI = 0.6
(63) Paramere, position relative to subgenital plates:
(0) surpassing (Fig. 4O), (1) not surpassing (Fig. 4N).
CI = 0.25, RI = 0.4
(64) Parameres, orientation relative to subgenital
plate: (0) directed forward (Fig. 5P), (1) directed back-
ward (Figs 3G, 5R), (2) directed downward (Fig. 5I).
CI = 0.5, RI = 0.5
(65) Paramere, spine: (0) present (Fig. 5K), (1) absent
(Fig. 5U, S). CI = 1, RI = 1
(66) Paramere, number of spines: (0) one (Fig. 5F), (1)
two (Fig. 5B). CI = 0.12, RI = 0.5
(67) Paramere, position of primary spine: (0) apical
(Fig. 5N), (1) subapical (Fig. 5H). CI = 0.05, RI = 0.5
(68) Paramere, insertion of primary spine: (0) lateral,
(1) mesal. CI = 0.5, RI = 0.66
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
(69) Paramere, orientation of primary spine: (0) hor-
izontal (backward, forward) (Fig. 5L, M), (1) vertical
(upward, downward) (Fig. 5R). CI = 0.09, RI = 0.16
(70) Paramere, apex of primary spine: (0) acute
(Fig. 5T), (2) blade-shaped (Fig. 5D), (3) rounded.
CI = 0.28, RI = 0.28
(71) Paramere, position of secondary spine: (0) apical,
(1) subapical (Fig. 5A). CI = 0.33, RI = 0.01
(72) Paramere, insertion of secondary spine: (0) lat-
eral, (1) mesal (inside). CI = 1, RI = 1
(73) Paramere, shape of apex of secondary spine: (0)
acute (Fig. 5B), (1) truncated, (2) blade-shaped (Fig. 5G).
CI = 1, RI = 1
(74) Paramere, dorsal process: (0) reduced (Fig. 5J), (1)
developed (Fig. 5C). CI = 0.04, RI = 0.57
(75) Paramere, shape of dorsal process: (0) rounded
(Fig. 5D), (1) acutely produced (Fig. 5S). CI = 0.1,
RI = 0.25
(76) Paramere, shape of apex: (0) long and narrow
(Fig. 5T, O), (1) short and wide (Fig. 5E, G, Q), (2) long
and not narrow (Fig. 5J). CI = 0.12, RI = 0.7
(77) Aedeagus, shape: (0) cylindrical (Fig. 6H), (1)
flattened (Fig. 6A). CI = 0.11, RI = 0.55
(78) Aedeagus, flatness: (0) anterior-posterior, (1) lat-
eral. CI = 0.5, RI = 0.75
(79) Aedeagus, apex: (0) simple (Fig. 7J), (1) with
processes (Fig. 7L). CI = 0.08, RI = 0.76
(80) Aedeagus, number of apical processes: (0) one pair
(Fig. 7E, I), (1) two pairs (Fig. 7K), (2) more. CI = 0.12,
RI = 0.22
(81) Aedeagus, shape of apical processes: (0) long and
slender (Figs 6R, 7R), (1) spine-shaped (Figs 6D, 7D), (2)
fin-shaped (Figs 6L, 7L). CI = 0.21, RI = 0.6
(82) Aedeagus, orientation of apical processes (rela-
tive to long axis):(0) horizontal (Fig. 7C), (1) vertical
(Fig. 7F). CI = 0.25, RI = 0.57
(83) Aedeagus, insertion of apical processes: (0) dorsal
(Fig. 7D), (1) ventral (Figs 6A, 7A), (2) lateral (Figs 6E,
7E), (3) both (lateral and dorsal) (Figs 6O, 7O). CI = 0.23,
RI = 0.52
(84) Aedeagus, shape of apical processes: (0) cylindri-
cal (Figs 6R, 7R), (1) flattened (Figs 6C, 7C). CI = 0.12,
RI = 0.63
(85) Aedeagus, shape of apex: (0) bifid (Figs 6P, Q, 7P,
Q), (1) truncated (Figs 6N, 7N), (2) rounded (Figs 6K,
7K), (3) acute. CI = 0.09, RI = 0.47
(86) Aedeagus, size of the apex relative to shaft: (0)
enlarged (Fig. 7O), (1) not enlarged (Figs 6G, 7G).
CI = 0.07, RI = 0.45
(87) Aedeagus, denticuli on surface: (0) present (Figs
6M, 7M), (1) absent (Figs 6F, 7F). CI = 0.09, RI = 0.58
(88) Aedeagus, position of denticuli on surface: (0)
ventral, (1) dorsal (Figs 6A, 7A), (2) both (Figs 6M, 7M).
CI = 0.28, RI = 0.5
(89) Aedeagus, localization of denticuli teeth: (0)
sparse (all over the aedeagus surface), (1) not sparse (lim-
ited to a specific area). CI = 0.25, RI = 0.62

(90) Aedegus, quantity of denticuli: (0) numerous
(Fig. 7A), (1) few (Fig. 7M). CI = 0.33, RI = 0.75
(91) Aedeagus, shaft: (0) simple (Fig. 6C, I), (1) with
processes (Figs 6B, 7B). CI = 0.05, RI = 0.48
(92) Aedeagus, shape of the shaft: (0) straight (Fig. 6I),
(1) C-shaped (Fig. 6K), (2) S-shaped (Fig. 6G, Q).
CI = 0.2, RI = 0.8
(93) Aedeagus, insertion of the shaft processes: (0)
dorsal (Fig. 6J), (1) ventral (Fig. 6A), (2) lateral (Fig. 6L).
CI = 0.66, RI = 0.85
(94) Aedeagus, number of shaft processes: (0) one
(Fig. 6D), (1) one pair (Fig. 7H). CI = 0.33, RI = 0.81
(95) Aedeagus, shape of shaft processes: (0) elon-
gated (Fig. 6N), (1) spiniform (Fig. 7D), (2) fin-shaped
(Fig. 7L). CI = 0.5, RI = 0.84
(96) Aedeagus, shape of shaft processes: (0) cylindrical
(Fig. 7B), (1) flattened (Fig. 7D). CI = 0.33, RI = 0.81
(97) Aedeagus, orientation of shaft processes (relative
to long axis): (0) horizontal (Fig. 7H), (1) vertical
(Fig. 7A). CI = 0.14, RI = 0.45
(98) Aedeagus, size of shaft processes relative to shaft
length: (0) longer than half length of shaft (Fig. 6J),
(1) shorter than half shaft length (Fig. 6N). CI = 0.5,
RI = 0.92
(99) Aedeagus, size of base: (0) enlarged, (1) not
enlarged. CI = 0.05, RI = 0.25
Female abdomen
(100) Ovipositor, margin of first valvulae: (0) with teeth
(Fig. 8H), (1) smooth (Fig. 8I). CI = 0.2, RI = 0.6
(101) Ovipositor, basal process of first valvulae: (0)
absent, (1) present (Fig. 8J). CI = 0.33, RI = 0.8
(102) Ovipositor, basal process of first valvulae:
(0) developed (Fig. 8G), (1) not developed (Fig. 8H).
CI = 0.09, RI = 0.56
(103) Ovipositor, number of basal processes of first
valvulae: (0) one (Fig. 8I), (1) two (Fig. 8G). CI = 0.2,
RI = 0.69
(104) Ovipositor, orientation of basal process of first
valvulae: (0) directed downward (Fig. 8I), (1) directed
backward (Fig. 8H), (2) directed forward (Fig. 8C).
CI = 0.22, RI = 0.41
(105) Ovipositor, shape of basal process of first valvu-
lae: (0) rounded (Fig. 8H), (1) elongated (Fig. 8I).
CI = 0.2, RI = 0.57
(106) Ovipositor, shape of second valvulae of ovipos-
itor: (0) short and wide (Fig. 8F), (1) long and slender
(Fig. 8E). CI = 0.16, RI = 0.7
(107) Ovipositor, teeth on dorsal margin of second
valvulae: (0) present (Fig. 8D), (1) absent (Fig. 8C).
CI = 0.08, RI = 0.68
(108) Ovipositor, position of teeth on dorsal margin
of the second valvulae: (0) limited to beginning of
apical third (Fig. 8E), (1) extended beyond apical third.
CI = 0.12, RI = 0.63
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
Neotropical spittlebug phylogeny and biogeography 93
Taxonomic key for Ischnorhininae genera (except
for incertae sedis species)
1. Small and delicate insects of about 5 mm length; post-
clypeus inflated and quadrangular, in frontal view (Figs. 1F,
2M), usually with a pair of small protuberance in the inferior
margin … Neaenini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1′ . Large and robust insects of >5 mm length; postclypeus
rectangular or rounded (Fig. 2P, N), in frontal view . . . . . . . . . 2
2. Pronotum with the anterior lateral margin longer than half
length of the anterior margin; postclypeus laterally compressed
with apex acutely angled, inferior lateral carina (in most of the
genera) apparent and strongly marked . . . . Ischnorhinini . . . . 8
2′ . Pronotum with the anterior lateral margin shorter or equal to
half length of the anterior margin; postclypeus inflated, rounded
not forming an acute angle at apex, inferior lateral carina absent
if present slightly marked … Tomaspidini . . . . . . . . . . . . . . . 21
3. Paramere with spines (Fig. 3I) . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3′ . Paramere without spines (Fig. 3H) . . . . . . . . . . . . . . . . . . . . . 5
4. Postclypeus swollen; vertex with a median carina; small
insects (length about 5 mm), black body with two yellow round
spots on the tegmina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neaenus
4′ . Postclypeus not swollen, flattened in ventral view, vertex
without a median carina; small insects (length < 5 mm), black
body with one white transversal stripe slightly marked located
in the apical third of the tegmina or dotted stripe slightly
marked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tomaspisinella
5. Pronotum curved (Fig. 1J); paramere with apex short
(Fig. 5U) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5′ . Pronotum not curved (Fig. 1K); paramere with apex long and
narrow (Fig. 5S) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6. Small and rounded insects (length about 5 mm), body black
with an orange stripe in the middle of the tegmina; tegmina
smooth and shiny with venation not visible . . . . . . . . . . Marcion
6′ . Elongated insects with length >5 mm ; tegmina with distinct
venation and apical plexus developed . . . . . . . . . . . . . Simorhina
7. Really small insects (length <2 mm ), with a black body;
vertex narrow; postclypeus, in frontal view, flattened; aedeagus
without processes . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitomaspis
7′ . Insects with regular size (length about 5 mm); vertex wide;
postclypeus, in frontal view, inflated; subgenital plates with a
spine-like apex (Fig. 4R) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zuata
8. Pronotum anterolateral margin straight; parameres directed
backward in relation to the subgenital plates; dorsal margin of
the first ovipositor valvulae with teeth . . . . . . . . . . . . . . . . . . . . . 9
8′ . Pronotum anterolateral margin convex; parameres directed
forward in relation to the subgenital plates; dorsal margin of the
first ovipositor valvulae smooth . . . . . . . . . . . . . . . . . . . . . . . . . 10
9. Large insects (length about 15 mm), black body with two
tranverse red stripes on the tegmina, humeral angle, anterolateral
margins and postclypeus red; vertex wide; pedicel with an
abundant density of long setae; postclypeus rounded in profile;
tegmina with apical plexus strongly developed . . . . . . . Huaina
9′ . Large insects (length <15 mm), black body with bright red
stripes or spots on the tegmina, vertex narrow, pedicel without
long setae; postclypeus slightly angled in profile; tegmina with
apical plexus developed . . . . . . . . . . . . . . . . . . . . . . . . . . Iphirhina
94 A. Paladini et al.

Fig. 4. Pygofer and subgenital plates, lateral view: (A) Sphenorhina rubra; (B) Panabrus dominicanus; (C) Pacachanthocnemis bella; (D) Aeneolamia
colon; (E) Ferorhinella balatra; (F) Choconta elliptica; (G) Monecphora semilutea; (H) Mahanarva (I.) integra; (I) Korobona lineata; (J) Isozulia
astralis (K) Kanaima katzensteinii; (L) Neomonecphora apicalis; (M) Ocoaxo sp. (N) Simorhina sciodes; (O) Baetkia compressa; (P) Cosmoscarta sp.;
(Q) Iphirina quota; (R) Zuata carvalhoi.

10. Insects usually red with black marks; postclypeus angulated 10′ . Insects with other colours; postclypeus rounded
in profile, compressed or strongly compressed with a lateral rarely angulated in profile, inflated or flattened, lateral
carina in the inferior margin present and strongly marked (rarely carina in the inferior margin absent, if present slightly
slightly delimited) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11 delimited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

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 Neotropical spittlebug phylogeny and biogeography 95

Fig. 5. Parameres, lateral view: (A) Sphenorhina rubra; (B) Aeneolamia colon; (C) Panabrus dominicanus; (D) Pachacanthocnemis bella; (E)
Monecphora semilutea; (F) Monecphora cingulata; (G) Ferorhinella balatra; (H) Mahanarva (I.) integra; (I) Huaina inca; (J) Isozulia astralis; (K)
Korobona lineata; (L) Homalogrypota coccinea; (M) Ischnorhina unifascia; (N) Kanaima katzensteinii; (O) Baetkia marionensis; (P) Neomonecphora
apicalis; (Q) Ocoaxo sp.; (R) Iphirina quota; (S) Zuata carvalhoi.; (T) Tiodus nigricans; (U) Marcion equestris.

11. Pronotum anterolateral margin convex, curved in profile; 11′ . Pronotum anterolateral margin straight, not curved in
postclypeus angulated, slightly compressed in frontal view, with profile; postclypeus angulated, strongly compressed in frontal
lateral grooves slightly delimited; tegmina with apical plexus view, with lateral grooves strongly delimited; tegmina with
poorly developed (Fig. 2C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 apical plexus developed (Fig. 2A) . . . . . . . . . . . . . . . . . . . . . . . 13

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96 A. Paladini et al.

Fig. 6. Aedeagus, lateral view: (A) Sphenorhina rubra; (B) Aeneolamia colon; (C) Monecphora semilutea; (D) Panabrus dominicanus; (E)
Pachacantocnemis bella; (F) Monecphora cingulata; (G) Ferorhinella balatra; (H) Mahanarva (Ipiranga) integra; (I) Huaina inca; (J) Isozulia astralis;
(K) Korobona lineata; (L) Homalogrypota coccinea; (M) Hyboscarta melichari; (N) Kanaima katzensteinii; (O) Ocoaxo sp.; (P) Tiodus nigricans; (Q)
Ferorhinella balatra; (R) Tomaspis handlirschi.

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 Neotropical spittlebug phylogeny and biogeography 97

Fig. 7. Aedeagus, dorsal view: (A) Sphenorhina rubra; (B) Aeneolamia colon; (C) Monecphora semilutea; (D) Panabrus dominicanus; (E)
Pachacantocnemis bella; (F) Monecphora cingulata; (G) Ferorhinella balatra; (H) Mahanarva (Ipiranga) integra; (I) Huaina inca; (J) Isozulia astralis;
(K) Korobona lineata; (L) Homalogrypota coccinea; (M) Hyboscarta melichari; (N) Kanaima katzensteinii; (O) Ocoaxo sp.; (P) Tiodus nigricans; (Q)
Ferorhinella balatra; (R) Tomaspis handlirschi.

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98 A. Paladini et al.

Fig. 8. Characters of the head and abdomen. Antennae, basal body of the flagellum: (A) Maxantonia rubescens; (B) Sphenoclypeana parana. Female
genitalia: (C) Maxantonia rubescens, pygofer. Second ovipositor valvulae: (D) Tomaspis furcata; (E) Mahanarva dubia; (F) Neosphenorhina ocellata.
First ovipositor valvulae: (G) Sphenorhina rubra; (H) Tomaspis furcata; (I) Mahanarva rubrovenata; (J) Neosphenorhina ocellata.

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 Neotropical spittlebug phylogeny and biogeography 99

Fig. 9. Scanning electron micrographs of characters of the head, thorax and abdomen. Antennae: (A) Sphenorhina rubra; (B) Kanaima fluvialis; (C)
Notozulia entreriana, basal body of the flagellum; (D) Sphenoclypeana parana, basal body of the flagellum; (E) Notozulia entreriana, sensilla of the
basal body of the flagellum. Subungueal process: (F) Deois flexuosa; (G) Sphenoclypeana parana. Second ovipositor valves: (H) Kanaima fluvialis; (I)
Kanaima fluvialis, teeth on the dorsal margin.

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100 A. Paladini et al.
12. Large (length >15 mm) and dark brown insects, aedea-
gus with more than two pairs of apical processes; basal
process of the first valvulae of the ovipositor directed
backwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neolaccogrypota
12′ . Large insects (length about 15 mm) tegmina black with
red marks, aedeagus with one pair of apical processes; basal
process of the first valvulae of the ovipositor directed downward,
three processes on lateral margin between pygofer and anal tube
(Fig. 4F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Choconta
13. Tylus and vertex without a median carina; postclypeus with-
out a lateral carina in the inferior margin; subungueal process
absent; process on lateral margin of the pygofer hook-like;
paramere apex long and flattened with one spine turned down-
ward (Fig. 5T) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tiodus
13′ . Tylus and vertex with a median carina; postclypeus with a
lateral carina in the inferior margin; subungueal process present;
subgenital plates short in relation to pygofer (Fig. 4O); aedeagus
with apical processes inserted laterally . . . . . . . . . . . . . . . . . . . 14
14. Antenna with basal body of flagellum without an arista
(Fig. 8B); postclypeus with one pair of lateral prominences
on inferior margin (Fig. 2K); vein A2 of tegmina distinct;
subgenital plates free for most of their length . . . . Schistogonia
14′ . Antenna with basal body of flagellum with an arista
(Fig. 8A); postclypeus without lateral prominences in the infe-
rior margin; aedeagus shaft straight . . . . . . . . . . . . . . . . . . . . . . 15
15. Large insects (length >15 mm); antennae with basal body of
the flagellum conic (Fig. 8B), longer than pedicel; postclypeus
with lateral carina on the inferior margin slightly delimited,
triangle formed in the apex with tylus absent; rostrum not
surpassing mesocoxae . . . . . . . . . . . . . . . . . . . . . Sphenoclypeana
15′ . Medium insects (length ≤15 mm); antenna with basal body
of the flagellum with the same length as pedicel; postclypeus
strongly flattened with lateral carina on the inferior margin
strongly delimited (Fig. 2I); rostrum reaching or surpassing
mesocoxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Aedeagus without shaft processes; second valvulae of the
ovipositor long and slender (Fig. 8E) . . . . . . . . . . . . . . . . . . . . . 17
16′ . Aedeagus with one pair or one isolated shaft process; second
valvulae of the ovipositor short and wide (Fig. 8F) . . . . . . . . 18
17. Paramere surpassing subgenital plates, with one apical spine
turned backwards; second valvulae of the ovipositor with two
basal processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baetkia
17′ . Paramere not surpassing subgenital plates, with one sub-
apical spine turned backwards (Fig. 4O); aedeagus laterally flat-
tened with denticuli on the surface . . . . . . . . . . . . . Laccogrypota
18. Large insects (length about 15–20 mm), tegmina
red with black marks; two rows of apical spine on the
basitarsus; aedeagus with one spiniform process on the
shaft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Typeschata
18′ . Medium to small insects (length about 8–15 mm); one row
of apical spines on the basitarsus; aedeagus with one pair of
processes on the shaft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Aedeagus with one pair of spiniform apical
processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosphenorhina
19′ . Aedeagus with one pair of fin-shaped apical processes
(Fig. 7L) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
20. Aedeagus with apex not enlarged; paramere with dorsal
process developed but not swollen . . . . . . . . . . . Homalogrypota
20′ . Aedeagus with an enlarged apex; paramere with dorsal
process developed and swollen (Fig. 5M) . . . . . . . . Ischnorhina
21. Pronotum with a metallic brightness, subgenital plates with
two lobes (Fig. 3E); aedeagus cylindrical, long and slender,
C-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
21′ . Pronotum without a metallic brightness, subgenital plates
without two distinct lobes (Fig. 3F); aedeagus with different
shapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22. Small insects (length about 5 mm); pronotum, length of
anterolateral margin smaller than half-length of anterior margin
(Fig. 2E); tegmina with apical plexus reduced . . . . . Orodamnis
22′ . Medium insects (length about 10 mm); pronotum, length of
anterolateral margin equal than half-length of anterior margin
(Fig. 2F); tegmina with apical plexus developed . . . . . . . . Deois
23. Subgenital plates without or with one elongated basal
process (Fig. 4A); aedeagus if with shaft processes these ones
with an elongated shape; base of vein Cu1 thickened . . . . . . . 24
23′ . Subgenital plates without or with a short basal pro-
cess; aedeagus without shaft processes; base of vein Cu1 not
thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
24. Tylus and vertex with a median carina; pronotum with one
developed median carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24′ . Tylus and vertex without a median carina; pronotum with
one reduced median carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
25. Vertex narrow (Fig. 3B); posterior leg strong, tibia with basal
spine larger and stronger than those located on the apical crown;
subgenital plates fused at the base; aedeagus with a rounded apex
with flattened apical processes (Fig. 7C); second valvulae of the
ovipositor short and wide . . . . . . . . . . . . . . . . . . . . . . Monecphora
25′ . Vertex wide (Fig. 3A); posterior tibia with basal spine equal
or smaller than those located on the apical crown; subgenital
plates free for most of it length; aedeagus without apical
processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26. Postclypeus rounded in profile (Fig. 1K); aedeagus without
a shaft process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
26′ . Postclypeus angulated in profile (Fig. 1H); aedeagus with a
process on the shaft inserted ventrally . . . . . . . . . . . . . . . . . . . . 28
27. Small insects (length about 7 mm); aedeagus straight, cov-
ered with spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catrimania
27′ . Medium insects (length >7 mm); aedeagus barrel-shaped,
covered with spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tunaima
28. Insects with a variable coloration; antennae with arista longer
than pedicel; subgenital plates with an elongated and acuminated
basal process (Fig. 4A); paramere with two subapical spines
(Fig. 5A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphenorhina
28′ . Insects black with red spots; antennae with arista shorter or
with equal length of pedicel; subgenital plates with an elongated
and rounded basal process (Fig. 4B); paramere two spines: one
apical and the other one subapical (Fig. 5C) . . . . . . . . Panabrus
29. Tegmina narrow; aedeagus with short shaft processes,
shorter than half of the shaft (Fig. 6N) . . . . . . . . . . . . . . . . . . . .30
29′ . Tegmina wide; aedeagus with apical processes, if with
shaft processes these ones longer than half of the shaft
(Fig. 6J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

30. Insects red with apex of tegmina translucid; postclypeus
angulated in profile, not receding backwards . . . . . Aracamunia
30′ . Insects black with red marks, totally black
or brown; postclypeus convex in profile, receding
backwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kanaima
31. Aedeagus without processes covered with little denticuli;
paramere with two spines . . . . . . . . . . . . . . . . . . . . . . . . . Urubaxia
31′ . Aedeagus with shaft processes without denticuli; paramere
with one subapical spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32. Postclypeus rounded in profile; rostrum not reaching meso-
coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mahanarva (Ipiranga)
32′ . Postclypeus, in females, angulated (obtuse angle) in profile;
rostrum reaching mesocoxae . . . . . . . Mahanarva (Mahanarva)
33. Tylus with a median carina; paramere with dorsal process
rounded; aedeagus C-shaped rarely straight . . . . . . . . . . . . . . . 34
33′ . Tylus without a median carina; paramere with dorsal
process acutely produced (Fig. 5S) rarely rounded; aedeagus
straight rarely with another shape . . . . . . . . . . . . . . . . . . . . . . . . 39
34. Postclypeus rounded in profile if angulated with a slightly
marked carina in the inferior margin . . . . . . . . . . . . . . . . . . . . . 36
34′ . Postclypeus angulated in profile, forming an obtuse angle
(Fig. 1E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35. Tegmina with apical plexus reduced; subungueal process
rectangular; apex of subgenital plates rounded; second valvulae
of ovipositor without teeth on the dorsal margin . . . . . . . . . . . . . .
Zulia (Neozulia)
35′ . Tegmina with apical plexus developed; subungueal process
triangular; apex of subgenital plates finger-like; second valvulae
of ovipositor with teeth on the dorsal margin . . . . Zulia (Zulia)
36. Postclypeus projecting forward (Fig. 1C) without lateral
grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ferorhinella
36′ .– Postclypeus not projecting forward with lateral
grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37. Small insects (length about 5 mm); postclypeus with a
narrow longitudinal carina . . . . . . . . . . . . . . . . . . . . . . . . Deoisella
37′ . Medium insects (length >5 mm); postclypeus with a wide
longitudinal carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38. Small and pale insects (length about 7 mm); paramere with
an apical spine always turned backward; aedeagus with an
enlarged and anteroventral flattened apex . . . . . . . . . . . Tapaiuna
38′ . Larger insects with bright colours (length >7 mm);
paramere with an apical spine turned forward or backward;
aedeagus with the apex not enlarged . . . . . . . . . . . . . Hyboscarta
39. Distance between ocelli of two ocellar diameters; tegmina
with apical plexus reduced; dense layers of setae on the
posterior basitarsus concealing spines of the apical crown;
second valvulae of ovipositor without spines on the dorsal
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
39′ . Distance between ocelli of one ocellar diameter; tegmina
with apical plexus developed; sparse layers of setae on the pos-
terior basitarsus not concealing spines of the apical crown;
second valvulae of ovipositor with spines on the dorsal
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
40. Elongated insects, black with white or yellowish marks on
tegmina; paramere with one spine; aedeagus straight with a large
amount of denticuli on its surface . . . . . . . . . . . . . . . . . Notozulia
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Neotropical spittlebug phylogeny and biogeography 101
40′ . Rounded insects, with bright coloration; paramere with two
spines; aedeagus S-shaped . . . . . . . . . . . . . . . . . . . . . Ferorhinella
41. Postclypeus with lateral grooves strongly delimited;
pronotum with posterior margin slightly grooved; posterior
leg, basitarsus with two rows of spines; subungueal process
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
41′ . Postclypeus with lateral grooves slightly delimited;
pronotum with posterior margin strongly grooved; posterior
leg, basitarsus with one row of spines; subungueal process
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
42. Postclypeus with one longitudinal carina . . . . . . . . . Ocoaxo
42′ . Postclypeus with three longitudinal carinas
(Fig. 2P) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maxantonia
43. Antennae with arista longer than pedicel; basal body of
flagellum conical . . . . . . . . . . . . . . . . . . . . . . . Pachacanthocnemis
43′ . Antennae with arista shorter or with the same length as
pedicel; basal body of flagellum with another shape . . . . . . . 44
44. Insects brownish with two orange stripes on the tegmina;
postclypeus with a longitudinal carina; rostrum surpassing
mesocoxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vorago
44′ . Insects with other colour pattern; postclypeus without a
longitudinal carina; rostrum not surpassing mesocoxae . . . . . 45
45. Postclypeus compressed anteriorly in males; not
swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carachata
45′ . Postclypeus not compressed anteriorly; swollen . . . . . . . 46
46. Pronotum with anterolateral margin straight; spine on the
inner face of femur absent . . . . . . . . . . . . . . . . . . . . . . . . Tomaspis
46′ . Pronotum with anterolateral margin convex; spine on the
inner face of femur present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
47. Insects pale with postclypeus yellowish and two round black
spots in the tegmina apex; rostrum not surpassing mesocoxae;
antennae with long and abundant setae on pedicel . . . . Sinopia
47′ . Insects with other coloration pattern; rostrum surpassing
mesocoxae; antennae with short and scarce setae on
pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
48. Posterior leg, tibia with two lateral spines . . . . . . Korobona
48′ . Posterior leg, tibia with one lateral spine . . . . . . . . . . . . . 49
49. Postclypeus angulated in profile; pronotum curved in lateral
view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Makonaima
49′ . Postclypeus convex in profile; pronotum not curved in
lateral view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomonecphora
Cladistic analysis
The data matrix included 100 taxa and 108 characters (Table
S1). The equal weights analysis produced 196 equally parsimo-
nious trees, with length of 1044 steps. Multiple analyses were
done under different search strategies, but although the num-
ber of trees found varied, in all the analyses the strict consensus
was stable. The strict consensus had 56 nodes collapsed and the
monophyly of most cercopid genera was corroborated with a
high value of support. Relationships among genera and tribes
of Ischnorhininae were poorly resolved.
In the analysis with the implied weighting scheme for each
one of the 20 K values chosen (K1–K20) one cladogram with
the highest fit value was obtained. The best K range for the data
102 A. Paladini et al.
matrix presented here was 8–13; this range was chosen based
on the comparison and selection of the best comparison values
of the distortion coefficient (Farris, 1989), and the similarity
coefficient (SPR) (Table S2). A strict consensus of the six
trees of the same length obtained by the best K range was
made and the resultant cladogram (Fig. 10) will be used as the
hypothesis for the New World Cercopidae relationships in the
discussion. The trees obtained with the other K values were
taken into consideration to know if the clades are present in all
the concavity values and the main differences found (Fig. 10).
Revised classification of Ischnorhininae
All Neaenini (Figs 10, S1–S2) taxa, except of Tapaiuna,
which is transferred herein to Ischnorhinini, were recovered
as monophyletic in all concavity values (but placement within
Zuata was variable). The tribe is supported by one synapomor-
phy, postclypeus quadrangular in frontal view (character 131 )
and one homoplasy, second valve of the ovipositor without teeth
on the dorsal margin (character 1070 ). The Simorhina+ clade is
defined by the absence of the paramere spine (character 651 ).
Hemitomaspis minuscula is nested within Zuata.
All genera included in Ischnorhinini sensu Fennah (1968)
were recovered in a monophyletic clade (Tiodus+ ; Figs 10,
S1–S2) in the equal weighting scheme and in most K values,
except of Neolaccogrypota brunnea. Herein, Ischnorhinini is
redefined in a broader sense (Huaina+ ; Figs 10, S1–S2) to
include all representatives of Fennah’s Ischnorhinini and three
other genera, Huaina, Choconta, and Iphirhina), which are
herein transferred from Tomaspidini. Ischnorhinini, as currently
defined, was recovered as monophyletic with concavity values
of K8–K13 and is supported by one homoplasy: pronotum
with proportion of length of anterolateral margin larger than
half-length of anterior margin (character 291 ).
Finally, the representative of the type genus of Hyboscartini
was recovered nested well within Tomaspidini and is herein
synonymized with it. Although H. melichari is not the type
species of Hyboscarta, it is very similar to it and would be
coded differently in only a few characters of the male genitalia
(Carvalho & Webb, 2005). Tomaspidini as herein defined,
includes besides Hyboscarta, Tapaiuna, previously placed in
Neaenini, and excludes the three above-cited genera transferred
to Ischnorhinini. This tribe was recovered as monophyletic
and is supported by five homoplasies: postclypeus convex in
profile (character 151 ); posterior leg with basitarsus with one
row of apical spines (character 470 ); apex of aedeagus without
processes (character 790 ); aedeagus shaft with one pair of
processes (character 941 ) and aedeagus shaft processes fin-like
shaped (character 952 ).
Tomaspidini was recovered divided into five main lineages
(named T1–T5 in Fig. 10). Clade T1 (Figs 10, S1–S2)
includes all four subgenera of Deois plus Orodamnis, which was
described by Fennah (1953) and treated by Metcalf (1961) as
another subgenus of Deois. Deois and Orodamnis are present in
trees obtained with all concavity values except K1 and K2–K3
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
and share two homoplasies: vertex wide (31 ) and aedeagus shaft
C-shaped (character 921 )
Tomaspidini clade T2 (Figs 10, S1–S2) includes Zulia,
Deoisella, Tropidorhinella, Hyboscarta, and Tapaiuna and
it is present in three concavity intervals (K4, K5–K7 and
K14–K16). Within this clade, the last three genera were recov-
ered as a monophyletic group supported by, among other apo-
morphies, a single lateral spine on the posterior tibiae (character
421 ). Notably, Fennah (1968) considered this character state to
be exclusive of Cercopinae (Old World Cercopidae) using that
feature as one of the main diagnostic characters for the division
of the two subfamilies of Cercopidae. However, further obser-
vations have documented this character state to be present in
some Neotropical genera, reducing the utility of this charac-
ter for subfamily-level definition. In addition, Zulia is currently
divided into two subgenera, Neozulia and Zulia and together
they were never recovered as a monophyletic group, suggesting
the possibility of elevating these subgenera to genera.
Tomaspidini clade T3 (Figs 10, S1–S2) comprises 12 gen-
era, including Tomaspis and several monotypic ones. It was not
recovered in analyses with other concavity values, but was sup-
ported by three homoplasies: tegmina (proportion length/width)
narrow (character 360 ); aedeagus with apical processes (charac-
ter 791 ) and teeth of the dorsal margin of the second valvulae of
the ovipositor present (character 1070 ).
Tomaspidini clade T4 (Figs 10, S1–S2) includes Monecphora,
Catrimania, Tunaima, Panabrus, and Sphenorhina, and is sup-
ported by one synapomorphy: processes of the aedeagus shaft
inserted laterally (character 931 ) and one homoplasy: median
carina of the pronotum reduced (character 331 ). This clade is
present in concavities K5–K7 and K14–K17. Tunaima and
Catrimania were recovered as polyphyletic and this result is
observed in all topologies obtained with different concavities
values, suggesting that both genera need a taxonomic review and
a phylogenetic analysis including more species.
Tomaspidini clade T5 (Figs 10, S1–S2) includes seven genera.
It was not recovered in all concavity values, but it was supported
by five homoplasies: median carina on tylus present (character
11 ), median carina on vertex present (character 21 ), aedeagus
shaft with processes (character 911 ), processes of the aedeagus
shaft cylindrical (character 906 ), and teeth of the dorsal margin
of the ovipositor limited to the beginning of the apical third
(character 1080 ).
Finally, the present analysis was not able to recover a sta-
tistically supported placement of Tomaspis biolleyi (originally
described in Sphenorhina) and T. handlirschi and these species
are of uncertain generic placement as stated by Carvalho &
Webb (2005), as well as incertae sedis within Ischnorhininae.
Biogeographical analysis
Ischnorhinines originated and started to diversify during the
mid- to Late Cretaceous (Cryan & Svenson, 2010), when
angiosperms were already important elements of the vegetation
(McLoughlin, 2001). Although the Neotropical cercopids are
currently distributed throughout South and Central America,
 Neotropical spittlebug phylogeny and biogeography 103

Fig. 10. Strict consensus of cladograms for the Neotropical cercopids resulting from the analysis of morphological data with the implied weighting
scheme using the optimal concavity interval of K8-K13. Dashed clades were also recovered with the equal weights analysis. For the main clades
discussed, information on whether it was also recovered with other concavity values is represented with filled or white rectangles (see legend of values
on top left). Relative Bremer support is indicated above each node and maximum likelihood bootstrap support is indicated below each node. Taxa and
branch colours refer to Fennah (1968) tribal classification (see legend on lower left), whereas the revised classification given herein is indicated by a
dashed line on the right of taxa names.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108

104 A. Paladini et al.
with a few species extending north to Ontario (Canada) in North
America (Hamilton, 1977), the S-DIVA analysis recovered the
ancestral area of distribution of Ischnorhininae (Fig. S3) as
the Northwestern South America dominion, an area located
in the middle of the tropical belt of the Americas during the
Ischnnorhinae divergence time. Due to scarce fossil data, is
difficult to obtain a picture of the tropical vegetation during
these time period; there is little evidence of tropical rainforests,
but it was possibly composed of savanna woodlands (composed
mainly of cycadophytes, ferns, and brachyphyllous conifers) or
angiosperm shrublands (Saward, 1992).
The restricted distribution of ancestral cercopids in North-
western South America was expanded by putative disper-
sal events to the Amazonian subregion by the ancestor of
Choconta + Laccogrypota+ (D1 in Fig. 11) in ischnorhinines
and to Amazonia or Amazonia + Paraná subregions by the
ancestor of tomaspidines (D2 in Fig. 11). Both areas cur-
rently harbour tropical rainforests that were already diverse in
the Eocene (Burnham & Johnson, 2004) and were partially
connected at different times at least until the late Pleistocene
(between 33 000 and 25 000 yr ago; Ledru, 1993), which would
facilitate dispersal between areas. Based on a study of forest avi-
fauna, connections between the Amazonian and Atlantic Forest
appear to have occurred through different routes in South Amer-
ica and at different periods: during the mid- to late Miocene
routes were associated with geotectonic events related to the
Andean uplift, and during the Pliocene to Pleistocene routes
were associated with climate change and expansion of humid
forests (Batalha-Filho et al., 2013). Without a divergence date
estimate for the Neotropical cercopid lineages, it is impossible to
infer when these dispersals may have occurred; however, consid-
ering that Pliocene–Pleistocene climate change was possibly a
major cause for inferred vicariant events of descendants of these
lineages (see below), these dispersal events (D1 and D2) proba-
bly occurred during or prior to the late Miocene.
Within Ischnorhinini, a vicariance event (V1 in Fig. 11) can
be postulated between Choconta + Neolaccopgrypota and Tio-
dus+ separating the ancestor of these clades in the Northwest-
ern South America and Amazonia subregions, respectively. The
Amazonian ancestor further dispersed to the Paraná or Cha-
coan subregions (D3 in Fig. 11), right before the vicariance
event separating the ancestor of Sphenoclypeana and Laccogry-
pota + Baetkia+ (V2 in Fig. 11) in the Paraná or Chacoan sub-
regions and Amazonian subregion, respectively.
Within the Tomaspidini, a vicariance event (V3 in Fig. 11)
can be postulated for the ancestor of two major clades (T2 + T3
and T4 + T5 in Fig. 11) limiting them to Northwestern South
America and Paraná, respectively. The former ancestor further
extended its range to the Paraná subregion (D4 in Fig. 11),
right before another vicariance event (V4 in Fig. 11) limiting
Ferorhinella + Notozulia and Maxantonia + Ocoaxo+ ancestors
in Paraná and Northwestern South America, respectively.
The analysis recovered two additional vicariance events
involving the ancestor of Tomaspidini and the ancestor of Vor-
ago+ . These events are not further discussed, because ances-
tral distributions recovered may have been biased due to the
species sampling of this analysis. For the case of the ancestor
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
of Tomaspidini, extant Deois are also recorded from the Andean
subregion, with D. morialis occurring in Chile, whereas in the
other case, Vorago radicis occurs in the Amazonian subregion
of both Brazil and Guyana. For the same reason, biogeograph-
ical events postulated for ancestors of generic lineages are also
not further discussed.
Several species of the Neotropical cercopids are currently
associated with grasses (Carvalho & Webb, 2005), most prefer-
ably nitrogen-fixing C4 grasses (Thompson, 2004). A recent
molecular estimate (Bouchenak-Khelladi et al., 2010) places
the origin of Poaceae between 120 and 94 Ma (95% confi-
dence interval) in northern Gondwana (either South America or
Africa), probably in moist and forested environments – a time,
place and habitat consistent with the origin of the Neotropical
cercopids. Interestingly, by the Eocene, grasses had dispersed
and achieved a nearly worldwide distribution, whereas, as far
as the single phylogenetic study conducted for cercopoids sug-
gest (Cryan & Svenson, 2010), no Old World cercopid lineage
has originated from within the Ischnorhininae, the South Amer-
ican lineage. Conversely, the South American lineage seems to
have originated from a paraphyletic assemblage of Old World
cercopids with origins dating back from 168.34 to 122.78 Ma,
substantially preceding the origin of grasses.
However, in South America, although open and dry habitats
were present by the early Miocene (23.3–16.3 Ma), grasses were
still a rare component of these habitats and became more preva-
lent only around the late Miocene (∼14 Ma), the onset of arid
grassland habitats with C4 grasses becoming dominant around
7 Ma (Strömberg, 2011; Ab’Saber, 2000). The diversification of
these South American grasses is somewhat contemporaneous
with the development of the Chacoan subregion. This subre-
gion currently includes areas covered with seasonally dry trop-
ical forests, where grasses are a minor element, and savannas,
which have grasses as their dominant component (Pennington
et al., 2000). The development of the Chacoan subregion is seen
a major vicariance event between the humid rainforests found
in the Amazonian and Paraná subregions (Morrone, 2006). This
could also explain some of the vicariance events discussed here
for cercopids (V2, V3 and V4) and others occurring between and
within genera studied. In addition, the recent expansion of these
drier forests in South America explains why the vast majority of
dispersals to the Chacoan subregion occur within ischnorhinine
genera and are not more common in more basal lineages of this
much older group.
Independent dispersals northwards to northern Central and
North America are seen in at least five genera. Species of
Ocoaxo and Neaenus are found in the Mesoamerican domin-
ion, more specifically in the Mexican Pacific Coast, Mexican
Gulf and Yucatan Peninsula. The Mesoamerican distributional
pattern usually includes Neotropical taxa that evolved in mon-
tane humid habitats (Morrone, 2006). Finally, dispersals towards
the Nearctic Region are seen within three genera: two of them
occupying the Continental Nearctic dominion, with Huaina inca
specifically occurring in the Sonora province and some species
of Sphenorhina occurring in the Mexican Plateau province; two
species of Prosapia, P. bicincta and P. ignifera, are known to
 Neotropical spittlebug phylogeny and biogeography 105

Fig. 11. Biogeographical analysis of the Neotropical cercopids showing relative probabilities of ancestral area alternative distributions as recovered
by S-DIVA. Distributions of terminal taxa are given as coloured circles next to genera in a simplified cladogram based on strict consensuses obtained.
Each colour represents a single area defined in the South and Central American map to the left. The main biogeographical events hypothesized and
discussed are given as D (dispersal leading to ancestral area expansion) and V (vicariance).

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108

106 A. Paladini et al.
have a wide range of distribution in the Nearctic, from Miami
(Florida) to Ontario (Canada).
Future taxonomic research in Cercopidae
Many phylogenetic and nomenclatural questions regarding
this conspicuous group of herbivores remain unanswered. Fur-
ther investigations with an expanded taxonomic sampling,
specially of speciose genera, and different kinds of data (molec-
ular and behavioural), are needed to better understand questions
regarding the generic and tribal relationships within Ischnorhin-
inae. The present paper is regarded as the first attempt to
reconstruct phylogenetic relationships of members of the fam-
ily Cercopidae with a wide generic sampling and morphologi-
cal data. Aiming for a revised classification of the Neotropical
cercopids (Table S3), the phylogenetic analysis was helpful in
uncovering synapomorphies that were useful to define the lim-
its of the revised tribes. As stated by Paladini & Cryan (2012),
generic and tribal concepts within Neotropical Cercopidae are
often not well supported by synapomorphic characters. All of
Fennah’s tribes were recovered as nonmonophyletic, except for
Hyboscartini, for which a single terminal was included. This
type genus was nested well within the Tomaspidini clade and
a synonymy was proposed. Also, based on these apomorphies,
a dichotomous key to tribes and genera of this economically
important group are given.
In order to make further progress into the systematics of
Ischnorhininae, and cercopids in general, it is important to sam-
ple Old World representatives. The Old World Cercopidae cur-
rently includes approximately 1100 species (Metcalf, 1961), and
very few higher-level taxonomic revisions and no phylogenetic
hypothesis with adequate sampling have been conducted. By
including an adequate taxon and character sampling of these
related lineages, we may uncover with better support the sis-
ter lineage of Ischnorhininae, information crucial to determining
divergence times and confirming the biogeographical scenario
proposed in the present paper.
Supporting Information
Additional Supporting Information may be found in the online
version of this article under the DOI reference:
10.1111/syen.12091
Figure S1. Optimization of unambiguous character state
changes over the strict consensus of cladograms for the
Neotropical cercopids, resulting from the analysis of mor-
phological data with the implied weighting scheme using
the optimal concavity interval K8–K13. Characters were
mapped using Winclada (Nixon, 2002). Black squares rep-
resent nonhomoplastic apomorphies, white squares homo-
plastic ones. Character numbers are indicated above the
squares and character states below: (A) Neaenini clade; (B)
Ischnorhinini clade; (C) Tomaspidini clade T1; (D) Tomas-
pidini clade T2.
© 2014 The Royal Entomological Society, Systematic Entomology, 40, 82–108
Figure S2. Optimization of unambiguous character state
changes over the strict consensus of cladograms for the
Neotropical cercopids, resulting from the analysis of mor-
phological data with the implied weighting scheme using
the optimal concavity interval K8–K13. Characters were
mapped using Winclada (Nixon, 2002). Black squares repre-
sent nonhomoplastic apomorphies, white squares homoplas-
tic ones. Character numbers are indicated above the squares
and character states below: (A) Tomaspidini clade T3; (B)
Tomaspidini clade T4; (C) Tomaspidini clade T5.
Figure S3. Biogeographical analysis of the Neotropical cer-
copids showing relative probabilities of ancestral area alter-
native distributions as recovered by S-DIVA. Distribution of
terminal taxa are given as coloured circles next to genera
in complete cladogram based on strict consensus obtained.
Each colour represents a single area defined in South and
Central American map in Fig. 11.
Table S1. Data matrix of 108 morphological characters for
100 taxa of Ischnorhininae and outgroups. Characters: (?) not
observed, (-) inapplicable.
Table S2. Index values of single best-fit topologies obtained
with different K values, where: K, concavity constant; F, fit;
L, number of steps with equal weights; RI, retention index;
CI, consistency index; SPR, average values of SPR compar-
isons to every other topology; Ric, distortion coefficient.
Table S3. New classification proposal for Ischnorhininae.
Acknowledgements
We thank Vinton Thompson (Metropolitan College of New
York) for valuable comments on the draft of this manuscript,
for his assistance during the senior author’s visit to the collec-
tion of the American Museum of Natural History, New York,
and his helpful advice and encouragement for this project. We
also thank Wayne Mathis and Stuart McKamey for assistance
and specimen loans from the National Museum of Natural His-
tory, Washington, D.C. Chris Dietrich (Illinois Natural Histroy
Survey), Jason Cryan (North Carolina Museum of Natural Sci-
ences) and Kazunori Yoshizawa (Hokkaido University) also
provided valuable comments on a previous version of this
manuscript. Several other people helped during the develop-
ment of this paper with helpful comments: Augusto Ferrari,
Marcel Gustavo Hermes and Olivia Evangelista. This work is
the result of a doctoral thesis (Universidade Federal do Paraná,
Programa de Pós-Graduação em Entomologia), it was sup-
ported by a doctoral fellowship from the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq process num-
ber 140228/2008-0) and a postdoctoral grant (process number
150163/2013-4) to the senior author. This research is also par-
tially funded by the advisor’s grant (RRC) from PROTAX/CNPq
(processes number 561298/2010-6 and 303127/2010-4). This
paper is the contribution number 1907 of the Departamento de
Zoologia, Universidade Federal do Paraná.

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Accepted 9 May 2014
First published online 26 August 2014