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The Pink Katydids of Sabah (Orthoptera: Tettigoniidae: Phaneropterinae: Eulophophyllum ) with Description of Two New Species

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Two new species of the previously monotypic genus Eulophophyllum Hebard, 1922 are described. All species of the genus known up until now occur in forested areas in Sabah, Borneo. The genus is unique for the strongly widened media field of the tegmen, in which all branches of the media anterior plus radius sector are strongly curved and run anteriorally. There is also a striking color difference between the sexes, with males uniformly green and females pink. The two new species E. lobulatum Ingrisch & Riede sp. n. and E. kirki Ingrisch & Riede sp. n. have large leaf-like expansions of the hind tibiae that are absent in E. thaumasium Hebard, 1922. They differ from each other in the number of main vein branches in the media field of the tegmen. Stridulation of E. lobulatum sp. n. consists of short double-clicks ranging from 6.5 to 8.5 kHz, repeated at longer intervals.
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S. INGRISCH, K. RIEDE, G. BECCALONI 67
Journal of orthoptera research 2016, 25(2)
Abstract
Two new species of the previously monotypic genus Eulophophyl-
lum Hebard, 1922 are described. All species of the genus known up
until now occur in forested areas in Sabah, Borneo. The genus is
unique for the strongly widened media field of the tegmen, in which
all branches of the media anterior plus radius sector are strongly
curved and run anteriorally. There is also a striking color difference
between the sexes, with males uniformly green and females pink.
The two new species E. lobulatum Ingrisch & Riede sp. n. and E. kirki
Ingrisch & Riede sp. n. have large leaf-like expansions of the hind
tibiae that are absent in E. thaumasium Hebard, 1922. They differ
from each other in the number of main vein branches in the media
field of the tegmen. Stridulation of E. lobulatum sp. n. consists of
short double-clicks ranging from 6.5 to 8.5 kHz, repeated at longer
intervals.
Key words
crypsis, tegminal venation, stridulation
Introduction
The genus Eulophophyllum Hebard, 1922 was previously known
from a single species, Eulophophyllum thaumasium Hebard, 1922,
described from one female collected in "Labuan, North Borneo"
(Federal Territory of Labuan, Malaysia: an island 8 km off the coast
of Sabah state). No additional specimens of either sex have been
collected since.
The genus is unique within Phaneropterinae in that the tegminal
venation with all main branches in the strongly widened medial
field are curved dorso-anteriorally, i.e., their ends are pointing to-
wards the head, while the normal case in Phaneropterinae is that
those veins run more or less oblique posteriorally. These unique
features were probably the reason for Hebard's epithet thaumasium
(θαυμάζειν [thaumazein] – amazed).
During more recent investigations by KR in 1991 (Riede 1997)
and Peter Kirk in 2013, more specimens of the genus differing from
E. thaumasium by conspicuous lobes of the hind tibia and showing a
spectacular color polymorphism have been collected, photographed,
or sound recorded. We have also noticed a photograph of a juvenile
Eulophophyllum from montane forest on Mount Kinabalu, Sabah
on a website (Lee 2016). Since this forest type is different from the
forest types in which the known species of Eulophophyllum were
collected it is possible that it represents an undescribed species.
In the present paper the strange tegminal venation will be dis-
cussed and the relation of Eulophophyllum to other Phaneropterinae
genera considered. Two new species from Sabah are described and
differentiated. Stridulation of one of the species is documented.
Material and methods
Recording of stridulation was done in the field (Silau Silau) with
a Sony Digital Audio Tape-corder TCD-D100, microphone Telinga
EM-3, frequency range up to 24 kHz (sampling frequency 48 kHz)
in darkness (starting 20:24 h, temperature 20°C). The singing male
sat more than 2.5 m above ground. Recordings were digitised using
a Sony Linear PCM recorder PCM-M10, using a sampling rate of 96
kHz, and stored in wav format.
Sound analysis was done using Amadeus Pro Version 2.2.2 on
a Mac Mini. In order to get a clear overview of the stridulation pat-
tern over time, noise and sounds of crickets and another tettigoniid
singing in the background were suppressed using an Audio Unit
bandpass filter: 20 Hz to 6 kHz -20 db, 6.3 kHz +20 db, 8 to 10 kHz
-20 db, 12 to 16 kHz +20 db, 20 kHz -20 db. For detailed analysis
of the song, the unfiltered sound was used. For spectral analysis a
Blackman window with 1024 dots was used.
Images of the stridulatory apparatus were produced using a
Canon A6 camera mounted to a Motic M3 microscope. The images
were processed with CaptureOne and stacked using ZereneStacker.
Images and a copy of the sound file will be available via OSF
(Cigliano et al. 2016).
Abbreviation of veins: Cu Cubitus, MA Media anterior, MP Media
posterior, R Radius, Sc Subcosta.
Morphology of tegminal venation
The venation of the strongly widened tegmina of Eulophophyllum
is unique. The genus does not seem to be closely related to any other
Phaneropterinae, although one might speculate that its peculiar
tegminal venation may have evolved from an ancestor which had
similar tegmina to Dysmorpha Brunner, 1878 or Leptoderes Serville,
1838. The widened dorsal margins of the hind tibiae possessed
by the species of Eulophophyllum described below are superficially
similar to those of Xantia Brunner, 1878, but this is probably due
to convergence as that genus has a quite different appearance in all
other respects. In his description of Eulophophyllum Hebard (1922)
noted that it keys out (in Brunner's 1891 key) to the Holochlorae
(now Holochlorini) close to the genus Liotrachela Brunner, 1878
[which mainly occurs on the Philippines (OSF 2016)]. However he
also pointed out that there are big differences in the shape of vertex,
pronotum, tegmina, legs and ovipositor.
The shape and venation of the tegmen in Eulophophyllum re-
sembles Dysmorpha obesa Brunner, 1878, in having a widened media
The pink katydids of Sabah (Orthoptera: Tettigoniidae: Phaneropterinae:
Eulophophyllum) with description of two new species
sigfrid ingrisch, Klaus riede, george Beccaloni
[SI, KR] Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D- 53113 Bonn, Germany. Email: s.ingrisch@zfmk.de,
k.riede@zfmk.de
[GB] The Natural History Museum, Cromwell Road, London, SW7 5BD, UK
Journal of Orthoptera Research 2016, 25(2): 67-74
Journal of orthoptera research 2016, 25(2)
S. INGRISCH, K. RIEDE, G. BECCALONI
68
field with dorso-anteriorally curved cross veins within it. But in
Dysmorpha the subcosta and radius are fused to a single broad vein,
only at their base and tip are they free (Fig. 1A). Moreover, there
is a distinct radius sector, the media anterior does not run parallel
to radius, and the cross veins in the media field are connected by
secondary veins, which is not the case in Eulophophyllum. In contrast,
in both Leptoderes and Eulophophyllum the subcosta and radius run
parallel for the greater part of the wing, but in Leptoderes they are
sinuate (Fig. 1B) while in Eulophophyllum they are simply curved
(Fig. 1C-F).
With regard to the widening of the tegmen and the course of
its veins, Eulophophyllum seems to be most closely related to the
genus Leptoderes Serville, 1838 [distributed in tropical SE Asia; see
OSF (2016)] although in the latter genus the tegmina are widen-
ing towards the apex, and the head and pronotum are different.
Indeed the course of the radius, radius sector and media anterior
veins in Leptoderes show an intermediate position between that
found in Eulophophyllum and other genera of Phaneropterinae with
rather broad wings e.g., Holochlora, in which the branches of radius
and media anterior run dorso-posteriorally into the dorsal (hind)
margin. In Leptoderes, both branches of radius sector and also the
end of the radius stem are strongly curved while the media anterior
and its branches run vertically into the dorsal margin (the posterior
margin in spread wings) (Fig. 1B). In Eulophophyllum all branches of
the radius and media anterior are strongly curved and run oblique-
anteriorally into the dorsal margin (hind margin in spread wings,
Fig. 1C-D), but there are differences in details between the species.
The line drawings in Hebard's (1922) description of E. thauma-
sium, show a distinct radius sector and a second branch of the radius
[images also available in OSF (2016)]. The anterior branch of the
media runs close to and parallel to the radius over almost the whole
length, while the media posterior is fused with the cubitus anterior
after a short distance from base, and that vein reaches the dorsal
margin of the tegmen in lateral view (when the wings are spread the
hind margin) (Fig. 1C-F). On photographs of Eulophophyllum from
the Danum Valley and Kinabalu, and also on the photograph of
the holotype of E. thaumasium made by Piotr Naskrecki and avail-
able in OSF (2016), it seems as though the media anterior is fused
with radius sector shortly after it gives rise to the last curved cross
vein. This is striking as the meshwork of veinlets of the tegmina of
Eulophophyllum is indistinct. Thus the main part of the tegmen is
formed by the greatly widened media field. All branches of the media
anterior and radius are strongly curved and run dorso-anteriorally.
The number of branches of the media anterior plus radius differs
between species, however.
Taxonomy
Eulophophyllum Hebard, 1922
Hebard. 1922. Proc. Acad. Nat. Sci. Philad. 74:160
urn:lsid:Orthoptera.speciesfile.org:TaxonName:12499
Diagnosis.—The genus has a unique tegminal venation as discussed
above. Apart from this it differs from Leptoderes by a more robust
and in frontal view shorter head and by a rather short and stout
pronotum, instead of an elongate pronotum with a laterally concave
disc. The tibial tympana are conchate internally and are open on
the external side.
Apart from the tegminal venation and the modified hind tibiae,
the two new species are notable for their color polymorphism
comprising two color variants, (1) a variant with striking pink and
red-brown body color, with wings and legs contrasting with the
pastel green tegminal veins and other marks of the same color and
(2) a uniform green variant so far only observed in males. Color
pattern of E. thaumasium have been described as ochreous buff with
yellowish pattern (Hebard 1922), but this description was based
on a dead female specimen whose color would have altered after
death.
Key to species
1. Hind tibia simple. Female tegmen with six dorsal branches of
MA plus R. Labuan Island . . . . . . . . . E. thaumasium Hebard, 1922
Dorsal angles of hind tibia forming widened lobes . . . . . . . . 2.
2. Tegmen with five dorsal branches of MA plus R. Dorsal area
of tegmen behind R in female semi-oval, in male strongly raised,
almost semi-circular with concave anterior area along stridulatory
area of dorsal field in lateral view (Fig. 1C-D). Male cercus thick
at base, narrowing towards and gradually transforming into apical
third; moderately curved with spinule at tip Fig. 1G). Mt. Kinabalu
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. lobulatum sp. n.
Tegmen with seven to eight dorsal branches of MA plus R. Dorsal
area of female tegmen behind R nearly semi-oval in shape, in male
slightly higher (Fig. 1E-F). Male cercus moderately thickened at base,
narrowing towards apical third; apical third markedly thinner, slightly
curved Fig. 1H). Danum Valley. . . . . . . . . . . . . . . . . . . E. kirki sp. n.
Eulophophyllum lobulatum Ingrisch & Riede sp. n.
urn:lsid:Orthoptera.speciesfile.org:TaxonName:493860
Holotype (male): East Malaysia: Sabah, Mt. Kinabalu NP, headquar-
ters, Silau-Silau trail near Bukit Tupei trail (N 6° 0’ 40’’, E 116° 32’
29’’, 1600 m a.s.l.), 25.v.1993, leg. Hoffmann (Fig. 2). Depository:
Zoological Research Museum Koenig (ZFMK), Bonn, Germany.
Other material (photograph of a female, Fig. 3G): East Malaysia:
Sabah, Mt. Kinabalu, viii.2011, photographed by Mark Eller (http://
www.whatsthatbug.com/2012/02/05/unknown-katydid-from-
borneo/).
Diagnosis.—The new species differs in both sexes from E. thauma-
sium by its tegminal venation having only five instead of six dorsal
branches of MA plus R and more strikingly by the dorsal margins
of the hind tibia being expanded into a pair of lateral lobes that
are absent in the female holotype of E. thaumasium, which has the
hind tibia of normal shape. From E. kirki sp. n., E. lobulatum sp. n.
differs by the more rounded instead of oval, shape of the tegmen,
which has only five instead of seven to eight dorsal branches of MA
plus R. The apical area of the male cercus is wider than in E. kirki.
Description (male holotype).—Head with vertex sloping between
ocelli; fastigium verticis with a medial furrow bordered by a parallel-
sided rectangular carina opened behind, separated from fastigium
frontis by a deep transverse furrow; lateral ocelli situated on both
sides at base of that carina. Frons distinctly higher than wide, with
shallow and wide subocular grooves, bulging in middle; antennae
inserted between lower half of compound eyes (Fig. 2C). Pronotum
slightly widening from anterior to posterior margin; anterior mar-
gin subtruncate, very slightly concave in middle, posterior margin
broadly rounded; disc flat with a faint medial carina in posterior
area, lateral margins straight and subangular (Fig. 2B); paranota
S. INGRISCH, K. RIEDE, G. BECCALONI 69
Journal of orthoptera research 2016, 25(2)
slightly longer than high, posterior area in situ covered by a project-
ing flap of tegmen. Tegmina leaf-like, strongly widened; anterior
margin moderately convex near both ends, almost straight in middle;
posterior margin strongly convex, nearly semi-circular (Fig. 2A).
Tegminal venation: Costa normal, costal field moderately
widened with oblique cross veins; subcosta and radius parallel
and closely approaching each other, little diverging before apex of
tegmen; radius without distinct branches but connected by numer-
ous weak veinlets to media anterior, two slightly stronger veinlets
in posterior half of tegmen running more oblique than the other
veinlets in this area might be regarded as remains of radius branches
that are fused with media anterior; media forked shortly behind
base; media anterior running parallel and closely approached to
radius, both veins connected by numerous faint cross veinlets; media
posterior fused with cubitus and the fused veins forming the hind
margin of tegmen; media field (area between media anterior and
media posterior) extremely widened with all cross veins within this
field curved dorso-craniad, the last two of those cross veins seem
to have a twofold base where the more distad might be a branch
of radius; dorsal area of tegmen narrow, triangular and short (Figs
1C, 2A-B).
Legs: Fore tibia normal, in cross-section quadrangular, tibial
tympana with conchate cover on anterior (internal) side, open
on posterior (external) side; mid tibia with dorsal margins little
widened in basal half; hind femur widened in basal half, narrow
in apical half. Hind femur on ventral margins with four internal
and seven external spinules in apical area; hind knees bi-spinose.
Hind tibia with both dorsal margins strongly expanded conferring
a leaf-like appearance (Fig. 2A-B); with numerous distinct spines
on lateral margins of expanded area, on internal side nearly from
Fig. 1. A-F, Comparison of venation of left (A, F) or right tegmina (B-E) between Dysmorpha, Leptoderes and Eulophophyllum species
(only main veins and stronger cross veins, most drawings without dorsal field): A, male of Dysmorpha sp. (Myanmar, Tenasserim); B,
female of Leptoderes sp. (Thailand, Tak, Mae Salid); C, male (holotype) and D, female of E. lobulatum sp. n. (Sabah, Kinabalu); E, fe-
male (holotype) and F, male of E. kirki sp. n. – G-H, Left male cercus of E. lobulatum sp. n. (G, latero-dorsal view) and E. kirki sp. n. (H,
dorso-lateral view). Left tegmina (A, F) mirror inverted to ease comparison. Abbreviations for veins: C Costa, Ma Media anterior, Mp
Media posterior, R Radius, Rs Radius sector, Sc Subcosta. Not to scale. All after photographs. Stippled line in F indicates part of tegmen
hidden by hind tibia.
Journal of orthoptera research 2016, 25(2)
S. INGRISCH, K. RIEDE, G. BECCALONI
70
Fig. 2. Eulophophyllum lobulatum sp. n. male (holotype): A, lateral view original setting; B, dorsal view after spreading wings of both sides;
C, frontal view of head, pronotum and fore legs; D, stridulatory file on underside of left tegmen; E, stridulatory area of right tegmen;
F, abdominal apex dorso-apical view; G, subgenital plate and cerci ventral view. Scales 10 mm (A-B), 1 mm (C-D, F-G), 5 mm (E). For
color version, see Plate II.
S. INGRISCH, K. RIEDE, G. BECCALONI 71
Journal of orthoptera research 2016, 25(2)
base, on external side behind basal quarter; ventral area of hind
tibia compressed with a single, rounded medial margin carrying
three minute spinules; only in apical area a little widened with two
angular margins carrying three external and two internal spines; at
tip with one dorsal and three ventral apical spurs on both sides.
Stridulatory file curved, 3.93 mm (diagonally measured and
disregarding basal tubercles) with 61 teeth and four minute tu-
bercles at very base; stridulatory teeth in about basal third dense
and narrow (37 teeth per 1.535 mm), afterwards wide and spaced
(24 teeth per 2.51 mm; Fig. 2D). Behind stridulatory vein with a
triangular semi-transparent window. Mirror in stridulatory field of
right tegmen also triangular (Fig. 2E). Tenth abdominal tergite with
apical margin wide and shallowly concave in middle, very faintly
convexly projecting on both sides of concavity. Epiproct tongue-
shaped, flattened, tip broadly rounded. Cerci rather stout at base
but gradually narrowed and little curved towards subparallel-sided,
nearly cylindrical posterior area, towards apex more strongly curved,
in apical area strongly setose and at tip provided with a small acute
spinule (Figs 1G, 2F). Subgenital plate with lateral margins slightly
approaching from base to hind margin; ventral surface with a pair of
longitudinal furrows separating a medial carina from lateral bulges;
hind margin subtruncate or very faintly concave in middle, on both
sides forming a roundish groove from which the small styli arise
(Fig. 2G). Phallus membranous.
Coloration (male holotype): Tegmen and hind leg (especially
lobes of hind tibia) yellowish brown in the dried specimen, prob-
ably green when alive; hind wings transparent. Body, fore and mid
legs discolored dirty brown. Face including bases of antennae light
yellowish brown; compound eyes reddish brown; lateral margins
of clypeus black. Disc of pronotum along fore and lateral margins
light yellowish brown, centre of disc until hind margin and paranota
dark brown.
Female. The female on the photograph agrees with the male
holotype in general characters but its coloration resembles that of
the pink variant of E. kirki sp. n. Subgenital plate not clearly visible
on the photographs. Ovipositor sickle-shaped.
Coloration (based on photograph of the pink color variant of
female, Fig. 3G). Head, pronotum, thorax, abdomen, and ovipositor
light pink. Antennal flagellum dark brown with spaced white and
black annulation. Head and pronotum pink with lighter ornaments.
Pronotum with light green lateral angles. Tegmen pink to medium
red in general color with anterior margin, veins and main cross
veins and area between radius and media anterior pastel green;
dorsal margin of tegmen in more than anterior half reddish, then
light green. Fore and mid legs pink; fore tibia at base, mid tibia in
basal half green; tarsi green. Hind femur pink in basal half, api-
cal half green; posterior tibia green at base and very tip, the larger
expanded central area pinkish red on ventral, dark red on dorsal
side; posterior tarsus green.
Measurements (male holotype).— (In mm.) Body w/ wings: 33; body
w/o wings: 20; pronotum: 5.8; tegmen: 28; tegmen width: 17.5;
anterior femur: 7; hind femur: 23; hind tibia: 23.
Etymology.—Named for the strongly widened dorsal margins of
the hind tibiae.
Stridulation.—The calling song of the male of E. lobulatum consists
of short double-clicks ("zic-zic") of together about 280 - 352 ms
(mean 308.65 ms ± 22.63 ms (s.d.), median 309 ms, n = 26). The
time from the beginning of the first to the beginning of the second
click was rather constant, between 160 - 169 ms (mean 163.96 ms
± 2.07 ms, median 164 ms (s.d.), n = 26). The song is rather loud
even to the human ear, with a peak frequency between 6.5 and 8.5
kHz and a second quieter maximum between 13.5 and 14.5 kHz
(Fig. 4A-D).
Eulophophyllum kirki Ingrisch & Riede sp. n.
urn:lsid:Orthoptera.speciesfile.org:TaxonName:493861
Holotype (female): East Malaysia: Sabah, Danum Valley (N 4° 57’
55’’; E 117° 41’ 25’’, ca 170 m), 6.vi.2013, photographed by Peter
Kirk (Fig. 3B-F).
Other material (photograph of a male, Fig. 3A): East Malaysia:
Sabah, Danum Valley field centre, night walk, 24.i.2009, photo-
graphed by Paul Bertner (https://www.flickr.com/photos/rainfor-
ests/3430798861).
Diagnosis.—This species is very similar to E. lobulatum sp. n. with
regard to the strongly widened tegmina, lobate hind tibiae and
green and pink color polymorphism. It differs by the more semi-
oval and relatively longer tegmina with seven or eight instead of
five transverse veins in the medial field. The apical area of the male
cerci is narrower than in E. lobulatum.
Description (female holotype).—Pronotum with concave anterior and
convex posterior margin; disc flat, lateral margins straight and sub-
angular; paranota a little longer than high, posterior area covered by a
projecting flap of tegmen. Tegmen strongly widened; anterior margin
moderately convex near both ends, nearly substraight in middle;
posterior margin strongly convex, nearly semi-circular (Fig. 3C-F).
Tegminal venation: costa normal, costal field widened with
oblique cross veins; subcosta and radius parallel and closely ap-
proaching each other, little diverging before apex of tegmen; radius
without distinct branches but connected by numerous weak veinlets
to media anterior, two of those veinlets in posterior half of tegmen
that are slightly stronger and run more obliquely than the other
veinlets in this area, can be regarded as branches of radius that
are fused with media anterior; media forked shortly behind base;
media anterior running parallel with and close to radius, both
veins connected by numerous faint cross veinlets; media posterior
fused with cubitus and the fused veins forming the hind margin
of tegmen; media field (area between media anterior and media
posterior) extremely widened with all cross veins within this field
curved (in situ) dorso-craniad, the last of those cross veins with a
twofold base; dorsal area of tegmen narrow, triangular and short
(Fig. 1E).
Legs: Fore tibia normal (quadrangular), tibial tympana con-
chate on anterior (internal), open on posterior (external) side; mid
tibia with dorsal margins little widened in basal half; hind femur
widened in basal half, narrow in apical half; hind tibia with both
dorsal margins strongly expanded conferring a leaf-like appearance
(Fig. 3B).
Coloration (living female, pink color variant, Fig. 3B-F). Head,
pronotum, thorax, abdomen, and ovipositor light pink. Antennae
pink at base, otherwise dark brown with white and black spaced
annulation. Head with a white band from base of mandibles to
compound eyes, running along and continued behind eyes; com-
pound eyes light green, ocelli white; tips of mandibles and maxillary
palpi black. Pronotum with pale green lateral angles. Tegmen pink
at very base, later red with anterior margin, veins and main cross
veins and area between radius and media anterior pastel green;
Journal of orthoptera research 2016, 25(2)
S. INGRISCH, K. RIEDE, G. BECCALONI
72
Fig. 3. Eulophophyllum species in habitat (A, D, G) and sitting on red leaves (B-C, E-F): A, E. kirki sp. n. male (Danum); B-F, E. kirki sp.
n. female (Danum); G, E. lobulatum sp. n. female (Kinabalu). – A, C, F, G, lateral view; B, apical view of hind legs and ovipositor; D,
oblique lateral view. Photographs: A, Paul Bertner; B-E, Peter Kirk; F, Mark Eller. For color version, see Plate III.
S. INGRISCH, K. RIEDE, G. BECCALONI 73
Journal of orthoptera research 2016, 25(2)
subcosta, radius and fused media - cubitus posterior for the greatest
part pink or red. Fore and mid legs pink; fore tibia at base and mid
tibia in basal half green, both with a black subapical mark; tarsi
dirty green. Hind femur pink at base, getting darker posteriorally,
apical half and ventral margin except at base green; posterior tibia
green at base and apex, the larger expanded central area dark red
on ventral, brown on dorsal side, margins green; posterior tarsus
green.
Coloration (living male, green color variant, Fig. 3A). Rather
uniformly green; thorax, fore and mid legs pale green; abdomen
for the greater part yellowish green. Head green with a white band
bordering compound eyes on posterior side and running down to
clypeus; antennal flagellum blackish, narrowly annulated and with
spaced white rings; maxillary palpi with black tips. Pronotum green;
disc with white lateral bands. Tegmen green; anterior margin, most
veins and main cross-veins are a little lighter yellowish green; along
radius in the area between radius and media there is a dark greyish
green band. Fore tibiae with a black spot near ventral end.
The male cerci have the apical area distinctly narrowed setose
before tip.
Etymology.— Named after the photographer of the holotype, Peter
Kirk.
Discussion
Up until now, the remarkable Phaneropterinae genus Eulopho-
phyllum was based on the description of only one specimen of one
species, a female from the island Labuan off the west coast of Sabah,
Malaysia, Borneo (Hebard 1922). The male of this species is still
unknown. Here we describe two additional species, E. lobulatum sp.
n. and E. kirki sp. n., based on collections and photographs of males
and females. Unfortunately E. kirki sp. n. could not be collected
due to strict conservation rules in the Danum Valley Conservation
Area (SEARRP 2016).
At present, the IZCN lacks clear rules and formally does not
Fig. 4. Stridulation of Eulophophyllum lobulatum sp. n. (holotype) in field: A, oscillogram showing time pattern of double-click echemes
of undisturbed male; B, single double-click in greater time resolution; C, sonogram of a single double-click; D, spectrum analysis of
sound. A, after applying a bandpass filter 6-16 kHz; B-D, from unfiltered recordings. Recordings by P. Hoffmann.
Journal of orthoptera research 2016, 25(2)
S. INGRISCH, K. RIEDE, G. BECCALONI
74
exclude a photograph as a holotype. There are some instances of
description of species based on photographs alone (Marshall &
Evenhuis 2015), but this approach has been discussed critically
by Cianferoni & Bartolozzi (2016). These authors fear that "this
[photographs as holotypes] will lead ...to severe problems....",
which is particularly true for insects, in which genitalia or other
hidden structures are used for species differentiation. This is not
the case for the katydids described here. Therefore, we decided to
include and name E. kirki sp. n., because (1) wing venation is the
main diagnostic feature and is clearly visible on photographs, (2)
to stimulate the search for and capture of a few individuals of E.
kirki sp. n. that may also occur outside protected areas, and (3) to
stimulate the search for the male of E. thaumasium at Labuan, at
least using photography. However, for all species mentioned here we
recommend collection, designation and institutional deposition of
more material. This includes the unknown male of E. thaumasium.
Since we found no major dimorphism between sexes apart from
sex specific differences, we predict that the general aspect of the
unknown male of E. thaumasium is similar to that of the female.
The recent discovery and collection of the tetrigid Arulenus miae
Skejo & Caballero, 2016 from a mountain forest in Mindanao was
triggered by an amateur Facebook photograph. In this case, the
authors organised a dedicated search for specimens. In contrast,
rainforest habitats of the species described here are either under
severe pressure (Labuan Island), or protected by strict law prohibiting
collection of any organisms (Danum Valley). Therefore, we opted
for a timely description of both species, hoping that our publication
will lead to further photographic and/or acoustic detections, that
it will convince authorities to grant permits for collection of type
material, and that it may stimulate habitat protection measures for
these enigmatic species.
We hypothesize that Eulophophyllum consists of an allopatric
species complex of at least three species: E. lobulatum from montane
rainforest, E. kirki from lowland forest, and E. thaumasium from
an island. Additional species probably exist on other islands (e.g.,
Bangi) and the genus might even occur in Palawan (Philippines).
On mainland Borneo, distinct species will probably be found along
the major mountain ranges (Crocker range) and micro-geographic
regions. Hybridisation could occur along contact zones between
montane and lowland forests. Detection of these highly cryptic
species will be facilitated by their conspicuous song, which might
be similar between the species, but with distinct diagnostic differ-
ences. Besides photographs, song recordings are another valuable
contribution which can be easily made. Many Phaneropterinae
species have a communication system between male and female
where both sexes stridulate, whilst in others only the males sing
(Heller et al. 2015). Up to now we only have information about
the acoustic behaviour of the male of E. lobulatum sp. n. described
here.
Pink coloration.— The pink color variant in Eulophophyllum has so
far only been documented for females of E. lobulatum sp. n. and
E. kirki sp. n. The photographer of E. kirki sp. n. placed the pink
female on leaves of a plant with young red leaves to demonstrate
the cryptic appearance of the pink individual when sitting on such
leaves (Fig. 3B-C, F). However, when discovered in the field, neither
this female nor the pink female of E. lobulatum sp. n. were resting
on red leaves (Fig. 3D, G). In fact, the pink female of E. kirki sp. n.
was resting on a branch with green leaves when discovered (Fig.
3D) and was very conspicuous to a human observer.
Possible reasons for the pink coloration of the females are as
follows: (1) Against a background of variable vegetation a pink
female with green veins might merge optically with the background
when seen from a distance (Fig. 3G); (2) It is possible that the pink
individuals which have so far been found may have simply been
resting on an atypical background - making them more obvious to
human observers. Their 'usual' resting site may be reddish first-flush
leaves, common in many rainforest plants; (3) the pink coloration
might serve as warning coloration, although we consider this to be
unlikely. Further field observations will be necessary to shed light
on the function of the pink coloration of these species and it may
reveal something unexpected.
Acknowledgements
Field work of Klaus Riede and Peter Hoffmann was supported by
the German Research Foundation (DFG). Peter Hoffmann recorded
and captured the male holotype specimen of E. lobulatum. We wish
to express our gratitude to the photographers for their permission
to publish their specimen photographs in this publication and in
OSF: Peter Kirk recorded the female holotype of E. kirki sp. n., Paul
Bertner photographed the male of E. kirki sp. n. (previously published
in www.flickr.com), and Mark Eller photographed the female of E.
lobulatum sp. n. (previously published in www.whatsthatbug.com).
References
Bertner P. 2016. https://www.flickr.com/photos/rainforests/3430798861
(accessed 2016-08-16).
Cianferoni F., Bartolozzi L. 2016. Warning: potential problems for taxonomy
on the horizon? Zootaxa 4139: 128-130. http://doi.org/10.11646/
zootaxa.4139.1.8.
Cigliano M.M., Braun H., Eades D.C., Otte D. 2016. Orthoptera Species File.
Version 5.0/5.0 http://Orthoptera.SpeciesFile.org [accessed 2016-10-17].
Eller M. 2016. http://www.whatsthatbug.com/2012/02/05/unknown-
katydid-from-borneo/ 3430798861 (accessed 2016-08-16).
Hebard M. 1922. Studies in Malayan, Melanesian and Australian Tettigoniidae
(Orthoptera). Proceedings of the Academy of Natural Sciences of
Philadelphia 74: 121-299, pls 11-22.
Heller K.-G., Hemp, C. Ingrisch S., Liu C. 2015. Acoustic communication
in Phaneropterinae (Tettigonioidea) - a global review with some new
data. Journal of Orthoptera Research 24: 7-18.
Lee C.C. 2016. www.wildborneo.com.my/photo.php?f=cld1303478.jpg
(accessed 2016-08-16).
Riede K. 1997. Bioacoustic monitoring of insect communities in a Bornean
rain-forest canopy, pp. 442-452. In: Stork N.E., Adis J., Didham R.K.
(Eds) Canopy Arthropods. Chapman & Hall, London.
Marshall S.A., Evenhuis N.L. 2015. New species without dead bodies: a case
for photo-based descriptions, illustrated by a striking new species of
Marleyimyia Hesse (Diptera, Bombyliidae) from South Africa. ZooKeys,
525: 117-127. http://dx.doi.org/10.3897/zookeys.525.6143
OSF. 2016. = Cigliano et al. 2 016
Santos C.M.D., Amorim D.S., Klassa B., Fachin D.A., Nihei S.S., De Carvalho
C.J.B., Falaschi R.L., Mello-Patiu C.A., Couri M.S., Oliveira S.S., Silva
V.C., Ribeiro G.C., Capellari R.S., Lamas C.J.E. 2016. On typeless species
and the perils of fast taxonomy. Systematic Entomology 41: 511-515.
doi:10.1111/syen.12180
SEARRP 2016. The South East Asia Rainforest Research Partnership http://
www.searrp.org/danum-valley/ (accessed 2016-09-03).
Skejo J., Caballero J.H.S. 2016, A hidden pygmy devil from the Philippines:
Arulenus miae sp. nov. — a new species serendipitously discovered in
an amateur Facebook post (Tetrigidae: Discotettiginae). Zootaxa 4067:
383-393.
... Describing species from photographs or illustrations is not a common practice, but it has been done a number of times (Fricke and Kacher 1982, Welch et al. 1986, Jonkers and Roersma 1990, Wallach and Jones 1992, Jones et al. 2005, Sinha et al. 2005, Robb et al. 2013, Cheng et al. 2015, Marshall and Evenhuis 2015, Ingrisch et al. 2016, Lonsdale and Marshall 2016, Nardelli 2016. Most of the names based on photographs are valid, but exceptions do exist (for an extensive historical overview of this practice, see Krell and Marshall (2017)). ...
... It is impossible to acquire permits for those areas, as the process is arduous and (understandably) discouraging (SEARRP 2021). Ingrisch et al. (2016) faced this exact problem. They decided to describe a new katydid species based on photographs taken in a protected area. ...
... They correctly concluded the following: "Therefore, we opted for a timely description of both species, hoping that our publication will lead to further photographic and/or acoustic detections, that it will convince authorities to grant permits for collection of type material, and that it may stimulate habitat protection measures for these enigmatic species." By describing a new species, Ingrisch et al. (2016) illustrated the problem and made steps not only to inspire further research but also to grasp the elusive limits of modern species research. The paper was published in this very journal, proving that exceptions do exist. ...
Article
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A heated debate on whether a new species should be described without a physical specimen, i.e., by designating a photographed specimen to serve as a holotype, has been ongoing for a long time. Herewith, without nomenclatural actions, a new species of the Batrachidein pygmy grasshoppers belonging to the genus Scaria Bolívar, 1887 is identified from the Andean rainforest in Peru. This species is clearly different from all its congeners by morphology and coloration. Two individuals of this peculiar species are known only from the photographs found on iNaturalist. The species has not been observed since 2008 when the photographs were taken. A short historical overview of the topic is given, illustrating the pros and cons of photograph-based species description. The concepts of names, holotypes, research effort, and conservation are discussed and related to the problem at hand. The current state of the taxonomic community's beliefs regarding this issue is reflected by the authors' three unsuccessful attempts to name this new species.
... Faced with the decision between ignoring it, naming it informally, or giving it a proper name they chose to give it a proper name. Ingrisch et al. (2016) found themselves in a similar situation upon completion of their revision of the genus Eulophophyllum, a group of spectacular pink katydids, for which species defining characters are clearly visible on the photographs used in their revision. They were unable to obtain specimens of a distinctive species associated with threatened and strictly protected rainforest habitats and opted to name it using a photo as a proxy with the expressed hope that it would 'lead to further photographic and/or acoustic detections, that it will convince authorities to grant permits for collection of type material [i.e., a neotype], and that it may stimulate habitat protection measures for these enigmatic species'. ...
... It is then up to the taxonomist's judgment whether to proceed with a formal description and naming without a preserved specimen. Some decide that the characters depicted in the photographs are sufficient for a formal description and naming (Hecq and Larsen 1998, Marshall and Evenhuis 2015, Ingrisch et al. 2016, Lonsdale and Marshall 2016, others decide against it (Linke 2008a,b, Edwards et al. 2009, Madika et al. 2011, van der Heyden 2015. ...
... A major point of criticism in the current debate was the fear that encouraging photo-based descriptions by example could open the floodgates for descriptions without preserved type material , Ceríaco et al. 2016, Cianferoni and Bartolozzi 2016. Not much time has passed, but so far, the feared flood of 'fast and sloppy' photo-based descriptions has not manifested itself, and we know of only four recent examples of insects described using a photo as a proxy for a lost or unpreserved type (Hecq and Larsen 1998, Marshall and Evenhuis 2015, Ingrisch et al. 2016, Lonsdale and Marshall 2016. The number and worldwide provenance of signatories of the Ceríaco et al. petition opposing photo-based descriptions indicates that many taxonomists expect that they will never need or want to describe a species using a photo as a proxy type. ...
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The option of describing new taxa using photographs as proxies for lost or escaped ('unpreserved') type specimens has been rarely used but is now undergoing renewed scrutiny as taxonomists are increasingly equipped to capture descriptive information prior to capturing and preserving type specimens. We here provide a historical perspective on this practice from both nomenclatural and practical points of view, culminating in a summary and discussion of a new Declaration of the International Commission of Zoological Nomenclature containing recommendations about descriptions without preserved specimens. We conclude that although descriptions using photographs as proxy types are Code-compliant and occasionally justified, the conditions under which such descriptions are justified are likely to remain relatively rare. Increasing restrictions on specimen collecting, which we deplore because of the centrality of collecting and collections to all of biodiversity science, could lead to more 'proxy type' descriptions in those taxa in which photographs can provide sufficient information for descriptions, but we predict that such cases will remain infrequent exceptions. Most of us were taught in our academic infancy that the description of a new species requires a published description and a designated type, and most entomologists involved with the discovery and description of new species realize that many other criteria should be met in order to justify a species description. We do not all agree on exactly where the bar should be set, but there is general agreement that new species must at a minimum be justified in the context of related and similar species and must be diagnosable. Descriptions of species that cannot be subsequently recognized are detrimental, and descriptions of species outside the context of the rest of their clade are usually of little use. The requirements for a published description and a designated type are embedded in the International Code of Zoological Nomenclature (ICZN 1999) and are governed by specific rules; the criteria for justified species descriptions and 'good' taxonomy are matters of taxonomic judgement. While new species have occasionally been described and formally named without a preserved type specimen, a recent paper by Marshall and Evenhuis (2015) explicitly discussing and executing such a procedure sparked a heated worldwide debate on social media and in a still growing number of formal publications. This discussion has sometimes drifted into emotional or uninformed arguments and is becoming increasingly repetitive but demonstrates that the issue is a controversial one. The coincidence of the launch of Insect Systematics and Diversity and the publication of a Declaration by the International Commission on Zoological Nomenclature (ICZN) in March 2017 on descriptions without preserved types (ICZN 2017) led to the drafting of this paper. Our aim is to calm the waves by giving a historical perspective of the issue, clarifying the rules of nomenclature, and emphasizing the centrality of taxonomic judgment and good practice.
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Whether or not a species might reasonably be described without the preservation of a type specimen is a matter of ongoing discussion among taxonomists (Dubois & Nemesio 2007; Minteer et al. 2014; Krell & Wheeler 2014; Lobl et al. 2016; Marshall & Evenhuis 2016; Santos et al. 2016). Here, we attempt to make our own contribution to the topic.
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Arulenus miae Skejo & Caballero sp. nov. is described from Buknidon and Davao, Mindanao, the Philippines. The species was serendipitously found in an amateur photo posted in Orthoptera Facebook group by Leif Gabrielsen. Holotype and paratype are deposited in Nederlands Centrum voor Biodiversiteit in Leiden, the Netherlands. Detailed comparison with Arulenus validispinus Stål, 1877 is given. A new diagnosis of the genus and A. validispinus is given. The paper is part of the revision of the subfamily Discotettiginae. This study provides a good example of how social networks can be used as a modern tool of discovering biodiversity if the regulations of the International Code of the Zoological Nomenclature are followed. A brief insight into habitat and ecology of this rainforest and mountainous species is presented.
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A new bombyliid species Marleyimyia xylocopae Marshall & Evenhuis, sp. n., an apparent mimic of the carpenter bee Xylocopa flavicollis (De Geer), is described from South Africa on the basis of photographs only. The pros and cons of species descriptions in the absence of preserved type specimens are discussed.
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Phaneropterinae is the largest subfamily within the bush-crickets/ katydids (Tettigonioidea), with about 2451 species, and with a world-wide distribution. Its acoustic communication differs from all other tettigonioid groups in that females primarily and typically respond to the male calling song with their own acoustic reply, a behaviour referred to as duetting. This type of response seems to have been lost only in a few species with wingless females. According to our literature review, information about the song patterns of about 330 species of Phaneropterinae have been published world-wide. Included in this number are ca 170 species of Barbitistini, a flightless West Palearctic tribe, which are treated separately. In the present study we summarize information from the above 330 species. We examine the morphology of stridulatory and hearing organs, and analyze the acoustic signals for frequency, number of syllables and number of interval types. We also have examined if and how responding by sound may have influenced other aspects of the acoustic communication system, especially the structure of the male calling song. Overall, the songs of male Phaneropterinae are similar to those of other tettigonioids. However, some Phaneropterinae species with very long and complex songs are found on all continents, exceeding in these characters nearly all other Ensifera species. These songs contain several different types of syllables and intervals of various duration. Because of this high interspecific variability (reaching from very simple to extremely complex), male phaneropterine songs are by far more variable than those of other tettigonioid families. However, since there are so few data on the behaviour of most Phaneropterinae species, and especially for females, we still are limited in our understanding of the reasons behind the song variability. Sexual selection by females choosing to respond preferentially to certain song types could be an important evolutionary force, but probably only in combination with some unknown ecological and behavioural factors.
Orthoptera Species File
  • M M Cigliano
  • H Braun
  • D C Eades
  • D Otte
Cigliano M.M., Braun H., Eades D.C., Otte D. 2016. Orthoptera Species File. Version 5.0/5.0 http://Orthoptera.SpeciesFile.org [accessed 2016-10-17].
  • M M Cigliano
  • H Braun
  • D C Eades
  • D Otte
Cigliano M.M., Braun H., Eades D.C., Otte D. 2016. Orthoptera Species File. Version 5.0/5.0 http://Orthoptera.SpeciesFile.org [accessed 2016-10-17].
On typeless species and the perils of fast taxonomy
  • C M D Santos
  • D S Amorim
  • B Klassa
  • D A Fachin
  • S S Nihei
  • De Carvalho
  • C J B Falaschi
  • R L Mello-Patiu
  • C A Couri
  • M S Oliveira
  • S S Silva
  • V C Ribeiro
  • G C Capellari
  • R S Lamas
Santos C.M.D., Amorim D.S., Klassa B., Fachin D.A., Nihei S.S., De Carvalho C.J.B., Falaschi R.L., Mello-Patiu C.A., Couri M.S., Oliveira S.S., Silva V.C., Ribeiro G.C., Capellari R.S., Lamas C.J.E. 2016. On typeless species and the perils of fast taxonomy. Systematic Entomology 41: 511-515. doi:10.1111/syen.12180