An extraordinary tribe of Tropiduchidae from the Eocene Baltic amber (Hemiptera: Fulgoromorpha: Fulgoroidea)

Abstract

The new tribe Patollini trib. n. of the Tropiduchidae with the extinct genus Patollo gen. n., comprising two species Patollo natangorum sp. n. and P. aestiorum sp. n. from Eocene Baltic amber is described. Taxonomic placement of some fossil taxa ascribed to Tropiduchidae is discussed. The classification of Tropiduchidae is discussed, as well as phylogenetic position and fossil record of Tropiduchidae and related taxa.

Full text

Accepted by A.P. Liang: 19 Mar. 2013; published: 9 May 2013
ZOOTAXA
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Copyright © 2013 Magnolia Press
Zootaxa 3647 (2): 371–381
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http://dx.doi.org/10.11646/zootaxa.3647.2.8
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An extraordinary tribe of Tropiduchidae from the Eocene Baltic amber
(Hemiptera: Fulgoromorpha: Fulgoroidea)
JACEK SZWEDO1 & ADAM STROI SKI
Museum and Institute of Zoology, Polish Academy of Sciences, Department of Palaeozoology, 64 Wilcza Street, PL 00-679 Warszawa,
Poland. E-mail: szwedo@miiz.waw.pl; adam@miiz.waw.pl
1Corresponding author. E-mail: szwedo@miiz.waw.pl
Abstract
The new tribe Patollini trib. n. of the Tropiduchidae with the extinct genus Patollo gen. n., comprising two species Patollo
natangorum sp. n. and P. aestiorum sp. n. from Eocene Baltic amber is described. Taxonomic placement of some fossil
taxa ascribed to Tropiduchidae is discussed. The classification of Tropiduchidae is discussed, as well as phylogenetic
position and fossil record of Tropiduchidae and related taxa.
Key words: Patollini trib. n., Patollo gen. n., Patollo natangorum sp. n., Patollo aestiorum sp. n., Baltic amber,
Palaeogene, classification, phylogeny, taxonomy, fossil record, Baltic amber, new tribe, new genus, new species.
Introduction
Fossil representatives of the family Tropiduchidae were described from both imprints and fossil resins. The oldest
descriptions of taxa ascribed to this family comes from Germar & Berendt (1856) monograph of inclusions from
the Eocene Baltic amber (Emeljanov 1983; Shcherbakov 2006). Other taxa from the Baltic amber were added by
Szwedo (2000) and Szwedo & Stroi ski (2010), and from imprints from the Eocene deposits of the Green River
(Shcherbakov 2006).
The family Tropiduchidae Stål, 1854, is one of the smaller planthopper families comprising 575 described
species in 164 genera (Yang et al. 1989; Shcherbakov 2006; Gnezdilov 2007; Bourgoin 2012). Tropiduchidae are
distributed in the warmer regions of the world, feed on shrubs and trees, and some are crop pests. (Fennah 1982;
Wilson et al. 1994; O’Brien 2002). The most recent higher classification of the family was provided by Fennah
(1982) who recognized 15 tribes (three of them divided into subtribes), based on body structure and tegminal
venation. This subdivision was primarily based on diagnostic characters without discussion of homology and
evolutionary trends (Asche & Wilson 1989). The tribes and subtribes recognized by Fennah (1982) are as follows:
Trypetimorphini Melichar, 1914, Neomatissini Fennah, 1982, Tambiniini Kirkaldy, 1907, Turneriolini Fennah,
1982, Paricanini Melichar, 1914, Isporisini Fennah, 1982, Eporini Fennah, 1982 (subtribes Eporina Fennah, 1982,
Clardeina Fennah, 1982), Catullini Melichar, 1914, Cyphoceratopini Fennah, 1945, Tangiini Melichar, 1914
(subtribes Tangiina Fennah, 1982, Neotangiina Fennah, 1982), Alcestini Melichar, 1914, Remosini Fennah, 1982,
Tropiduchini Melichar, 1914, Eutropistini Kirkaldy, 1906 (subtribes Duriina Fennah, 1982, Kazerunina Dlabola,
1977, Eutropistina Fennah, 1982) and Cixiopsini Fennah, 1982. Later, the extinct tribe Jantaritambiini Szwedo,
2000, described from Baltic amber was added (Szwedo 2000). Shcherbakov (2006) added another extinct tribe
Emilianini, based on the imprint of tegmen from the Eocene of the Green River, Colorado. Subsequently, Szwedo
& Stroi ski (2010) added another extinct tribe from the Eocene Baltic amber – Austrini Szwedo et Stroi ski, 2010.
Gnezdilov (2007) transferred the Gaetuliina Fennah, 1978 from Nogodinidae to Tropiduchidae giving it tribal
status, and Trienopini Fennah, 1954 from Issidae.

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Material and methods
Specimens were examined under normal and polarized light, using an OLYMPUS SZH10 stereoscopic
microscope. Drawings were made using an attached cameral lucida. Photographs were taken with use of
OLYMPUS C-5060W digital camera attached to the stereoscopic microscope, and then readjusted using Combine
ZP and Adobe Photoshop Elements 5.0 software.
Vein nomenclature follows the interpretation proposed by Szwedo & yła (2009) and Nel et al. (2012).
Systematics
Order Hemiptera Linnaeus, 1758
Suborder Fulgoromorpha Evans, 1946
Superfamily Fulgoroidea Latreille, 1807
Family Tropiduchidae Stål, 1854
Tribe Patollini trib. n.
Type genus: Patollo gen. n., here designated.
Diagnosis. The new tribe differs from other Tropiduchidae by the combination of characters as follows. Head with
vertex wider than long in midline. Brachypterous, with costal area present, not reaching apex of clavus; nodal line
absent, common portion of M and CuA present. Hind tibiae with three lateral spines. Male genital styles fused at
base.
Description. Head with compound eyes wider than pronotum. Vertex distinctly wider than long in midline;
with margins strongly carinate; vertex anteriorly widely angulate. Frons and clypeus smooth, lacking pustules or
setae. Frons with median carina, lateral margins of frons carinate. Antennal pedicel longer than wide; plate organs,
microsetae and macrosetae present only in distal half. Postclypeus without carinae. Apical segment of rostrum
distinctly longer than broad. Pronotum short, with two distinct anterolateral carinae fused medially. Mesonotum
widely rhomboid, lateral angles at about half of mesonotum length, close to the posterior margin of pronotum;
mesonotum tricarinate, lateral carinae not connected anteriorly; scutellum separated from mesonotum by transverse
groove.
Tegmen ovoid, with full venation developed, brachypterous; anterior margin strongly convex, anteroapical
angle widely angulate, apex rounded, posteroapical angle widely rounded, clavus with apex reaching to about of
tegmen length. Costal area about as wide as costal cell, subequal throughout, with apex not exceeding apex of
clavus, with transverse veinlets. Costal cell with transverse veinlets. Basal cell elongate, stems Sc+R and M leaving
basal cell with a common stalk, separately from CuA stem. Forking pattern of stems Sc+R, M and CuA not regular,
variable on left and right tegmina, but with common portion of branches M and CuA present. Nodal line absent,
transverse veinlets not forming regular pattern or cells. Clavus with transverse veinlets.
Hind tibiae with three lateral spines in distal half and apical row of teeth. Basitarsomere longer than second
tarsomere, with apical row of teeth and subapical setae. Second tarsomere with median portion arcuate, covered
with setae. Basitarsomere about as long as combined length of second and third tarsomeres.
Abdominal segments III-VIII subequal in length. Pygofer narrow, about 3 times as high as long in the middle
at ventral margin. Genital styles lobate, wide, with subapical processes, near the margin of pygofer, fused at base.
Anal tube not exceeding length of genital styles, with anus in the middle.
Composition. Patollo gen. n.

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Genus Patollo Szwedo et Stroi ski gen. n.
Type species: Patollo natangorum Szwedo et Stroi ski sp. n., here designated.
Composition. Two species. Patollo natangorum sp. n., P. aestiorum sp. n.
Age and occurrence. Eocene, Baltic amber.
Etymology. The generic name is derived from the name of the Old Prussian (Baltic Prussian) god of war, the
underworld, and the dead – Patollo. Gender: masculine.
Diagnosis. Head with vertex wider than long in mid line. Metatibio-tarsal formula 8 : 2+(7) : 2 (0); Anal tube
narrow, about 5 times as long as wide. Male genital style in lateral aspect, about as long as high, with small spine
on dorsal margin at level of pygofer apex.
Description. Vertex with margins strongly carinate, anterior aspect of vertex broadly angulate, lateral margins
subparallel, elevated, posterior margin broadly concave. Disc of vertex concave, smooth, with incomplete carinae
(median or lateral ones). Frons and clypeus smooth, lacking pustules or setae; frons with median carina reaching
frontoclypeal suture, lateral margins of frons carinate. Compound eye bulging, about as broad as long,
posteroventrally delimited by narrow callus. Antennal fovea placed below the compound eye, elevated at posterior
margin; pedicel club-like, longer than wide, arista long, placed centrally on pedicel. Postclypeus without carinae.
Rostrum with apex slightly exceeding mesocoxae, apical segment of rostrum distinctly longer than wide.
Mesonotum in apical portion delicately wrinkled. Stem of M forked apicad of fork of stem of Sc+R.
Pro- and mesofemora not flattened, subequal in length; pro- and mesotibiae subtrapezoid in cross section,
subequal in length. Profemora and protibiae and mesofemora and mesotibiae subequal in length, respectively.
Metafemora slightly flattened, distinctly shorter than metatibiae. Metacoxae very well developed, enlarged, with
meracantha distinct, subtriangular with acute apex. Metatibiae dilated apically, flattened to concave in apical
portion.
Male. Pygofer in lateral view wider in apical portion with posterodorsal angle widely rounded, without
processes. Anal tube narrow, without processes, apical portion, apicad of anus, in lateral view narrower than basal
one. Genital styles with rounded apex, lower margin with small tooth-like expansion near base.
Female unknown.
Patollo natangorum Szwedo et Stroi ski, sp. n.
(Figs 1–17)
Diagnosis. Disc of vertex without median carina; two incomplete and weak lateral carinae from anterior margin of
disc of vertex directed posteriorly, these are not reaching half of vertex length measured in the middle.
Description. Total length 4.9 mm, head with compound eyes about 2.16 mm wide, length of vertex in mid line
0.34 mm, 0.4 mm at lateral margin, width of vertex 1.1 mm; length of frons in mid line 1.28 mm, 1.2 mm at lateral
margin; apical segment of rostrum 0.33 mm long; antennal scape 0.11 mm long, pedicel 0.23 mm long, arista 0.86
mm long.
Length of pronotum 0.32 mm, width of disc of pronotum 1.52 mm, length of mesonotum+scutellum 0.84 mm,
width of mesonotum at base 1.28 mm.
Pronotum slightly shifted from mesonotum, hence the arcuate anterior of mesonotal ridge is visible. Anterior
mesonotal ridge strongly arcuate, reaching posterior margin of mesonotum.
Length of tegmen about 2.37 mm, width of tegmen about 1.06 mm; length of clavus 1.52 mm.
Profemur about 1 mm long, protibia 1.04 mm, protarsus with claws 0.43 mm, first tarsomere 0.16 mm, mid
tarsomere 0.16 mm, apical tarsomere 0.21 mm, tarsal claws 0.07 mm. Mesotibia 1.18 mm, mesotarsus with tarsal
claws 0.33 mm, basitarsomere 0.17 mm, mid tarsomere 0.17 mm, apical tarsomere with tarsal claws 0.33 mm.
Metafemur 1.15 mm long, metatibia 1.82 mm long, 0.26 wide at apex, metatarsus with tarsal claws 0.66 mm long,
basitarsomere 0.37 mm long, about 0.15 mm wide at apex, midtarsomere 0.21 mm long, 0.13 mm wide at apex,
apical tarsomere 0.21 mm long.
Abdomen with genital segment 2.5 mm long. Anal tube 1.07 mm long, 0.2 mm wide. Pygofer 1.61 mm high,
0.63 mm long at upper margin 0.43 mm long in the narrowest point, 0.5 mm near ventral margin. Genital style 0.91
mm long, 1 mm wide.

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FIGURES 1–8. Patollo natangorum gen. et sp. n. 1. anterior portion of body; 2. head in left lateral view; 3. left tegmen
venation; 4. right tegmen venation; 5. left hind leg; 6. apex of hind tibia; 7. anal tube in dorsal view; 8. apex of male abdomen,
left lateral view.
Left tegmen. Stem Sc+R forked in basal half of tegmen, basad of stem M forking; apicad of claval veins Pcu
and A1 junction. Branch ScRA1 forked basad of stem M forking, RA forked basad of fork of RP; stem RP forked
slightly basad of fork of branch M1+2. Stem M forked at about half of tegmen length; branch M3+4 fused with branch
CuA1, which extends to the margin. Stem CuA forked slightly apicad of stem M forking, slightly basad of costal
area apex. Claval veins Pcu and A1 connected slightly basad of half of clavus length.
Right tegmen. Stem Sc+R forked slightly apicad of stem CuA forking, well basad of stem M forking. Branch
RA probably with three terminals, stem RP forked apically at level of claval apex. Stem M forked basad of claval
apex; branch M1+2 forked apically, branch M3+4 fused with CuA1 for a long distance, then terminal M3 separated near

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apex. Stem CuA forked well basad, at basal of tegmen length, basad of claval veins junction. Claval veins Pcu
and A1 connected at half of tegmen length. The details are difficult to discern because of a milky veil obstructing
the specimen.
Etymology. The specific epithet is derived from the Natangi—the Old Prussian tribe inhabiting the area
between the rivers Pregel (now Pregolya) and Alle (now Łyna) during the Middle Ages.
FIGURES 9–13. Patollo natangorum gen. et sp. n. 9. specimen in amber, left lateral view; 10. anterior portion of body; 11.
anterior portion of body, left laterodorsal view; 12. head in left lateral view; 13. hind legs.

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FIGURES 14–17. Patollo natangorum gen. et sp. n. 14. left tegmen; 15. mid and hind legs; 16. left hind tarsus; 17. apex of
male abdomen.
Age and occurrence: Baltic amber, Middle Eocene. This fossil resin is aged within the range of 38–47 Ma
(Ritzkowski 1997; Perkovsky et al. 2007). Absolute dating analyses of glauconites from Sambia Peninsula showed
that the “blue earth” formation (amber bearing Prussian Formation) is allocated to the Middle Eocene (Lutetian:
44.1 ± 1.1 Ma) and is thus significantly older than previously assumed (Wappler 2003, 2005). Limnic sediments of
Eckfeld Maar, aged 44.3 ± 0.4 Ma, correlate with K-Ar radiometric data from the Sambia Peninsula and contain

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insect genera known only from Baltic amber (Wappler 2005). However, assumptions on the Middle Eocene age of
Baltic amber was discussed by Perkovsky et al. (2007), and the Upper Eocene (Bartonian/Priabonian: 37.7 ± 3 Ma)
age of the Prussian Formation is preferred by these authors. Weitschat & Wichard (2010) mentioned that two
additional amber-bearing horizons in the underlying beds of “blue earth” indicate that amber had already been
transported to secondary deposits about 50 million years ago.
Material examined. Holotype, male. Specimen No. MIBUG 5419 (AUF 084JS), Coll. Jacek Serafin,
Piaseczno, deposited in the Museum of Amber Inclusions, University of Gda sk, Poland. Right side of the
inclusion covered with a milky veil. Syninclusions: fragment of unidentified plant, a few bubbles of gas.
Patollo aestiorum Szwedo et Stroi ski, sp. n.
(Figs 18–30)
Diagnosis. Disc of vertex with incomplete and weak median carina posteriorly, not reaching half of vertex length.
Description. Total length 5.33 mm, head with compound eyes 1.57 mm wide, length of vertex in mid line 0.28
mm, 0.35 mm at lateral margin, width of vertex 0.86 mm; length of frons in mid line 1 mm, 1.14 mm at lateral
margin, length of postclypeus in mid line 0.67 mm, length of anteclypeus in mid line 0.5 mm, length of clypellus
0.14 mm, length of rostrum 0.93 mm, subapical segment 0.41 mm, apical segment 0.4 mm; length of second
antennal joint 0.29 mm, length of antenna 0.78 mm.
Length of pronotum 0.28 mm, width of disc of pronotum 0.86 mm between anterolateral carinae at posterior
margin, about 1.37 mm, length of mesonotum+scutellum 0.71 mm, width of mesonotum at base 1.2 mm.
Length of tegmen 2.05 mm, width of tegmen 1.05 mm; length of clavus 1.5 mm.
Profemur 1 mm long, fore tibia 0.86 mm, protarsus with claws 0.43 mm, first tarsomere 0.17 mm, mid
tarsomere 0.17 mm, apical tarsomere 0.24 mm, tarsal claws 0.07 mm. Mesofemur 1 mm, mid tibia 1.14 mm,
mesotarsus with tarsal claws 0.5 mm, basitarsomere 0.2 mm, mid tarsomere 0.2 mm, apical tarsomere 0.29 mm.
Metacoxa 0.22 mm long, metatrochanter 0.22 mm long, metafemur 1 mm long, metatibia 1.8 mm long, 0.25 wide
at apex, metatarsus with tarsal claws 0.64 mm long, basitarsomere 0.36 mm long, 0.16 mm wide at apex,
midtarsomere 0.23 mm long, 0.13 mm wide at apex, apical tarsomere 0.23 mm long.
Abdomen with genital segment 3.33 mm long. Anal tube 1.07 mm long, 0.2 mm wide. Pygofer 1.43 mm high,
0.64 mm long at upper margin 0.4 mm long in the narrowest point, 0.47 mm near ventral margin, 0.36 mm in mid
line at ventral margin. Genital style 1.07 mm long, 1.14 mm wide.
Tegmen length : width ratio 1 : 1.95, claval apex reaching 0.73 of tegmen length. Clavus with veinlet between
stem CuA and CuP, apicad of claval veins junction, free portion of vein Pcu measured from anterior claval angle to
claval veins junction 1.8 times longer than common claval vein Pcu+A1. Metatibio-tarsal formula 8 : 2+(7) : 2 (0);
basitarsomere longer dorsally than wide between tips of outermost teeth (2.45 : 1); mid tarsomere longer dorsally
than wide between tips of outermost teeth (1.6 : 1). Anal tube narrow, about 5 times as long as wide, with lateral
margins subparallel, apical margin straight. Male genital style in lateral aspect, about as long as high, with small
appendix on dorsal margin at level of pygofer apex, with indistinct lateral keel; apical margins thickened; covered
with short, thick setae.
Terminal system of branches RA, RP and M forking disorganized, vary on left and right tegmen; veinlets on
corium and membrane between longitudinal branches not regular, vary on left and right tegmen. Variability in
tegminal venation and veinlets system of left and right tegmina as in figures.
Left tegmen. Stem Sc+R forked slightly anteriad of stem M forking and anteriad of stem CuA forking, branch
ScRA1 forked at level of stem M forking; branch RA forked twice reaching margin with three terminals; branch RP
forked slightly basad of level of claval apex. Stem M forked at level of claval veins junction, branch M1+2 forked at
level of contact of fused claval veins with claval margin, branch M1 reaching apex with two terminals, branch M2
reaching margin with two terminals; branch M3+4 fused for a long distance with branch CuA1, basad of claval apex,
then common branch forked slightly before apex. Stem CuA forked slightly apicad of claval veins junction, branch
CuA1 fused with M3+4 then with M4, branch CuA2 not forked before apex. Claval vein Pcu and A1 fused apicad of
half of length of clavus.
Right tegmen. Stem Sc+R forked slightly apicad of claval veins junction, branch ScRA1 forked at level of stem
M forking; ScRA1 forked before reaching the margin; branch RA forked before reaching the margin; branch RP

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forked twice before the margin, reaching it with three terminals. Stem M forked slightly apicad of claval veins
junction, branch M1+2 reaching the margin with as single terminal (but see discussion); branch M3+4 forked at level
of apex of costal area, branch M3 forked just after the apex of clavus; branch M4 fused with CuA1 for a distance,
reaching margin together with CuA1. Stem CuA forked just before margin, slightly apical of claval apex. Claval
veins Pcu and A1 fused at about half of length of clavus.
Remarks. The venation of the right wing of P. aestiorum can be interpreted as described above or,
alternatively, the stem M1+2 is forked distinctly basad of the claval apex, with blind branch M2. Another equally
probable interpretation is that branch M2 is “broken” and the distal portion of the branch M2 is fused with M3 for a
distance with branch M3 re-emerging at the level of claval apex.
Age and occurrence. Eocene, Baltic amber, for details see comments in the section on P. natangorum.
Etymology. The specific epithet is derived from the name of the people inhabiting the eastern Baltic Coast
during the Antiquity – Aestii.
FIGURES 18–23. Patollo aestiorum gen. et sp. n. 18. anterior portion of body; 19. venation of left tegmen; 20. venation of
right tegmen; 21. right hind leg; 22. right hind tarsus; 23. apex of male abdomen.

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FIGURES 24–30. Patollo aestiorum gen. et sp. n. 24. specimen in amber, right lateral view; 25. specimen in amber, left lateral
view; 26. anterior portion of body, dorsal view; 27. anterior portion of body, right laterodorsal view; 28. hind legs; 29. hind
tarsi; 30. apex of male abdomen.

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Material examined. Holotype, male, Nr. 373 B; collection of Mr. Carsten Gröhn, Glinde, Germany, to be
deposited in the Geologisch-Paläontologisches Institut, Hamburg Universität, Germany. Left side covered with
milky veil, abdominal segment exposed, very probably due to the activity of decaying gases. Syninclusions: a
bunch of stellate hairs, very tiny apterous aphid.
Discussion
The tribe Patollini from Eocene Baltic amber clearly differs from any other tribe of the Tropiduchidae. Brachyptery
combined with the presence of a short costal area are the most striking features of the group. A short costal area
with transverse veinlets, with the apical portion not reaching the apex of the clavus is present in the fully winged,
recent tribe Alcestini. Also, the ovoid shape of the wing resembles that of Alcestini, as well as lack of a nodal line
(however, venation in the Alcestini is more regular and well developed). The short tegmina, lack of a nodal line and
presence of veinlets in the costal cell somewhat resembles the Austrini, and the head characters of the two tribes
are similar, i.e. vertex wider than long in midline. The costal area with veinlets and the head with vertex wider than
long are also present in the Gaetuliini (both extant species and fossils from Eocene Baltic amber). Meracantha
[(post-coxal process of Fennah (1982)] of the Patollini is stout and spine-like, as in the Austrini, however, the
metacoxa is enlarged and together with metepimeron covers a large area. Fennah (1982: 632) stated that the form
of the process is useful for separating genera, but of limited value for distinguishing tribes. The metatibia with three
lateral spines is present in the extant tribes Alcestini, Cyphoceratopini, Remosini and Tangiini, as well as in extant
and fossil Gaetuliini. The bilaterally symmetrical genital styles fused at base are present in most extant tribes of
Tropiduchidae.
The diversity of Tropiduchidae known from Eocene Baltic amber is quite high. There are three extinct tribes
known, Jantaritambiini, Austrini and Patollini, and representatives of Gaetuliini, which include extant species,
were reported as well (Szwedo & Stroi ski 1999, 2010; Szwedo 2000). This suggests rapid diversification of the
Tropiduchidae during the Palaeogene.
Most planthoppers are macropterous, but brachyptery is common in several families (O’Brien & Wilson
1985). Brachyptery was defined by Metcalf (1950) as tegmina with reduced venation, covering only the basal
segments of the abdomen. Capinera (2008) defined brachyptery as having short wings that do not cover the
abdomen or nearly so. Asche (1985) discussed pterygopolymorphic species of Delphacidae that had various types
of tegmina and hind wings. O’Brien & Wilson (1985) stated that brachyptery in planthoppers may range from a
partial to an almost complete reduction in the size of the hind wings with a relatively long tegmina, to the apparent
absence of hind wings and great shortening of the tegmina. The problem of wing reduction in the Hemiptera was
discussed by Kerzhner (1981) based on his observations of Nabidae. He defined macropterous forms as those
having normally developed tegmina and hind wings with complete venation, more or less exceeding the apex of the
abdomen. Submacropterous forms are those with hind wings distinctly shorter than tegmina and tegmina often
slightly shorter and narrower than those of macropters. Subbrachypterous forms are those with the tegmina merely
reaching the apex of the abdomen and with strongly reduced hind wings. Brachypterous (micropterous) forms are
these with the tegmina reaching to of the body length, reduced membrane of the tegmen, and eubrachypterous
(micropterous) are those with the tegmina shorter, membrane reduced to narrow appendix or missing, clavus and
corium fused and hind wings in the form of short and small scales or completely absent. Metcalf (1950) defined
macropterous tegmina as usually longer than the abdomen, frequently much longer, with fully developed venation,
and koeliopterous tegmina as those of moderate length, covering most of the abdomen and with fairly developed
venation. The latter term (used as coeliopterous) was subsequently used by Fennah (1982) to describe tegmina of
some tropiduchid tribes. The tegmina of Patollini trib. n. match the definitions of brachyptery, in terms of their
length, however, they have fully developed venation and the membrane is not shortened or reduced.
Further study of the external morphological structures, venation patterns of extant and extinct taxa, external
and internal genital characters of both males and females is necessary in order to understand the evolutionary
relationships of this family and its relatives.
Acknowledgements
The paper partly results from the research grant of the Ministry of Science and Higher Education of Poland No. NN
303 2979 37 awarded to JS.

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References
Asche, M. (1985) Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha). Marburger
Entomologische Publikationen, 2(1), 1–910.
Asche, M. & Wilson, M.R. (1989) The palm-feeding planthopper genus Ommatisus (Homoptera: Fulgoroidea: Tropiduchidae).
Systematic Entomology, 14, 127–147.
http://dx.doi.org/10.1111/j.1365-3113.1989.tb00271.x
Bourgoin, T. (2012) FLOW (Fulgoromorpha Lists on the Web): a world knowledge base dedicated to Fulgoromorpha. Version:
8. Last update: 16 December 2012. Available from http://hemiptera-databases.org/flow (Accessed 17 December 2012)
Capinera, J.L. (Ed.) (2008) Encyclopedia of Entomology. 2nd ed., Springer Science+Business Media B.V., Dordrecht,
LXIII+4346 pp.
Emeljanov, A.F. (1983) Cretaceous planthopper from Taimyr (Homoptera, Dictyopharidae). Paleontologicheskiï Zhurnal 3,
79–85. Published in English as: Yemel’yanov, A.F. (1984) Dictyopharidae from the Cretaceous deposits on the Taymyr
Peninsula (Insecta, Homoptera). Paleontological Journal, 17(3), 77–82. [In Russian]
Emeljanov, A.F. (1999) Notes on delimitation of families of the Issidae group with description of a new species of Caliscelidae
belonging to a new genus and tribe (Homoptera, Fulgoroidea). Zoosystematica Rossica, 8(1), 61–72.
Fennah, R.G. (1982) A tribal classification of the Tropiduchidae (Homoptera: Fulgoroidea), with the description of a new
species on tea in Malaysia. Bulletin of Entomological Research, 72, 631–643.
http://dx.doi.org/10.1017/S0007485300008658
Germar, E.F. & Berendt, G.C. (1856) Die im Bernstein befindlichen Hemipteren und Orthopteren der Vorwelt. In: Berendt, G.C.
(Ed.), Die im Bernstein befindlichen organischen Reste der Vorwelt. Commission der Nicholai’schen Buchhandlung,
Berlin, pp. ii + 1–40.
Gnezdilov, V.M. (2007) On the systematic positions of the Bladinini Kirkaldy, Tonginae Kirkaldy, and Trienopidae Fennah
(Homoptera, Fulgoroidea). Zoosystematica Rossica (2006), 15(2), 293–297.
Kerzhner, I.M. (1981) Poluzhestkokrylye semeïstva Nabidae. Nasekomye khobotnye. Fauna SSSR. [Heteroptera of the family
of Nabidae. Rhynchota. Fauna of the USSR], Leningrad, 13, 2, 1–326. [In Russian]
Metcalf, Z.P. (1950) Phylogeny of the Homoptera Auchenorrhyncha. Proceedings: Eighth International Congress of
Entomology, Stockholm 1948, 561–565,
Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, D.-Y., Roques, P., Guilbert, E., Dostál, O., Szwedo, J. (2012) Traits and
evolution of wing venation pattern in paraneopteran insects. Journal of Morphology, 273(5), 480−506.
http://dx.doi.org/10.1002/jmor.11036
O’Brien, L.B. (2002) The wild wonderful world of Fulgoromorpha. In: Holzinger, W. (Ed.), Zikaden - Leafhoppers,
Planthoppers and Cicadas (Insecta: Hemiptera: Auchenorrhyncha). Denisia 4, zugleich Kataloge des OÖ.
Landesmuseums, Linz, Neue Folge Nr. 176, pp. 83–102.
O’Brien, L.B. & Wilson, S.W. (1985) Planhopper systematics and external morphology. In: Nault L. R. & Rodriguez J. G.
(Eds.), The Leafhoppers and Planthoppers. John Wiley & Sons, New York, pp. 61–102.
Perkovsky, E.E., Rasnitsyn, A.P., Vlaskin, A.P. & Taraschuk, M.V. (2007) A comparative analysis of the Baltic and Rovno
amber arthropod faunas: representative samples. African Invertebrates, 48(1), 229–245.
Ritzkowski, S. (1997) K-Ar-Altersbestimmung der Bernsteinführenden Sedimente des Sammlandes (Paläogen, Bezirk
Kaliningrad). Metalla (Sonderheft), 66, 19–23.
Shcherbakov, D.E. (2006) The earliest find of Tropiduchidae (Homoptera: Auchenorrhyncha), representing a new tribe, from
the Eocene of Green River, USA, with notes on the fossil record of higher Fulgoroidea. Russian Entomological Journal,
15(3), 315–322.
Szwedo, J. (2000) First fossil Tropiduchidae with a description of a new tribe Jantaritambiini from Eocene Baltic amber
(Hemiptera: Fulgoroidea). Annales de la Société entomologique de France (N.S.), 36(3), 279–286.
Szwedo, J. & Stroi ski, A. (1999) Redescription of Tritophania patruelis Jacobi, 1938 from Eocene Baltic amber (Hemiptera:
Nogodinidae). Annales Zoologici, 49(3), 203–207.
Szwedo, J. & Stroi ski, A. (2010) Austrini – a new tribe of Tropiduchidae planthoppers from the Eocene Baltic amber
(Hemiptera: Fulgoromorpha). Annales de la Société entomologique de France, 46(1–2), 132−137
Szwedo, J. & yła, D. (2009) New Fulgoridiidae genus from the Upper Jurassic Karabastau deposits, Kazakhstan (Hemiptera:
Fulgoromorpha: Fulgoroidea). Zootaxa, 2281, 40−52
Wappler, T. (2003) Die Insekten aus dem Mittel-Eozän des Eckfelder Maares, Vulkaneifel. Mainzer Naturwissenschaftliches
Archiv, 27, i–viii + 1–234.
Wappler, T. (2005) The age of Baltic amber: could Eckfeld resolve this problem? Fossils X3, Insects – Arthropods – Amber.
Programme and Abstracts, 3rd International Congress of Palaeoentomology with 2nd International Meeting on
Palaeoarthropodology and 2nd World Congress on Amber and its Inclusions, 7th to 11th February 2005, Pretoria, South
Africa, pp. 53.
Wetischat, W. & Wichard, W. (2010) Baltic amber. In: Penney, D. (Ed.), Biodiversity of Fossils in Amber from Major World
Deposits. Siri Scientific Press, Manchester, UK, pp. 80–115.
Wilson, S.W., Mitter, C., Denno, R.F. & Wilson, M.R. (1994) Evolutionary patterns of host plant use by delphacid planthoppers
and their relatives. In: Denno, R.F. & Perfect, T.J. (Eds.), Planthoppers. Their Ecology and Management. Chapman &
Hall, New York, London, pp. 7–113.
Yang, J.T., Yang, C.T. & Wilson, M.R. (1989) Tropiduchidae of Taiwan (Homoptera: Fulgoroidea). Taiwan Museum Special
Publication, Series 8, 65–115.