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Alma B. Mohagan1,2, Romeo R. Patano Jr.1,2*, Emmanuel P. Leaño1,2,
Merced G. Melencion1,2, Fulgent P. Coritico1,2, and Victor B. Amoroso1,2
Morphology and Morphometrics of Diotarus verrucifer
(Stål, 1877), a Pygmy Grasshopper Endemic to Mindanao
*Corresponding Author: romeonojrpatano@gmail.com
1Center for Biodiversity Research and Extension in Mindanao
2Department of Biology, College of Arts and Sciences
Central Mindanao University, Musuan, Maramag, Bukidnon 8710 Philippines
Diotarus Stål, 1887 (Tetrigidae: Cladonotinae) is a genus of pygmy grasshoppers with four
species, all of which are inhabiting the Philippines with one species known also from Sulawesi.
Diotarus verrucifer Stål, 1877 was originally described from Semper’s collection and from an
unknown location from the Philippines. The species has been reported once since the description
by Gůnther in 1938. In this study, we report species diagnostic characteristics and describe
its natural habitat. The brief morphological description and, more importantly, the updated
distribution map are firsts for this species. The species is similar to Diotarus galeatus and is still
not clear if they represent separate species. We present detailed measurements of our specimens
in order to make comparison in the future possible.
Keywords: Diotarus verrucifer, groundhopper, Marilog District, measurements, Mt. Agad-agad,
pronotum, tectiform
Philippine Journal of Science
149 (3): 571-579, Sept 2020
ISSN 0031 - 7683
Date Received: 15 Apr 2020
INTRODUCTION
The genus Diotarus Stål, 1877 belongs to the family
Tetrigidae and subfamily Cladonotinae. Members of this
family are characterized mainly by having filiform or
thicker antennae, and the medial ocellus is situated below
the lower margin of the eyes. The dorsal margin of the
anterior and middle femora has a well-developed carina,
with the facial carina of the frontal costa considerably
widened in which they enclose a widened scutellum
(wider than the scapus) (Tumbrinck 2014). However,
the subfamily seems not to be monophyletic (Tumbrinck
2014; Skejo et al. 2019). The genus includes four species,
three of which are endemic in the country while Diotarus
pupus is also known from Sulawesi. The genus Diotarus
is unique through its pronotum by having semicircular to
truncated apex and elevated anterior margin in relation to
the head (Tumbrinck 2014). Those species have crested,
tectiform pronotum with truncated apex, giving the
animal a leaf-like pattern. This is only one of the genera
with leaf-like mimicry (Skejo et al. 2019). The holotype
originated from the Semper’s collection, and most of
the grasshoppers’ material lack detailed locality data
(Stål 1877; Bolívar 1887; Tumbrinck and Skejo 2017;
Muhammad et al. 2018).
Based on Semper’s collections as shown by Tumbrinck
(2014), except for D. verrucifer, some specimens of
species from the genus Diotarus were collected in Luzon –
specifically from Norzagaray, Bulacan and Mount Bulusan
in Sorsogon (D. galeatus) – while other species were
collected in Camarines Sur (D. pupus) and Los Baños,
Laguna (D. ikonnikovi).
New distribution records and habitat of D. verrucifer are
described in this paper. We aim to provide its morphological
571
description, as we have observed a lack of actual images in
its natural environment online or in literature. The specific
localities on which this species can be found were also
illustrated. We provide habitat description with supporting
images. This study provides new pygmy grasshoppers
records from the area in which we recently recorded
Spartolus pugionatus (Mohagan et al. 2020).
MATERIALS AND METHODS
Entry Protocol and Permits
Necessary permits such as an approved gratuitous permit
(GP) from the Department of Environment and Natural
Resources, in compliance with the Philippine Republic
Act No. 9147 for the collection of the specimens, and
the Institutional Animal Care and Use Committee permit
for animal handling were obtained prior to the conduct
of the research project.
Photography and Measurements
Documentation of the specimens was conducted using a
Canon™ digital single-lens reflex camera with the use of
a stereomicroscope. Small body parts such as the antennae
were measured using an ocular micrometer. The studies of
Skejo and Berner (2017), Tan and Artchawakom (2015),
Tumbrinck and Skejo (2017), and Muhammad et al.
(2018) were used as guides for the measurements. The
following measurements were taken:
BL – body length (length from vertex of head to apical
margin of subgenital plate); PL – pronotum length (from
the anterior margin (FM included if present) to the tip);
PW – pronotum lobe width (in its widest part); PH –
pronotum height (from the lowest part of the lateral lobes
to the highest part of dorsum); FFL – fore femur length;
FFW – fore femur width (in the middle); MFL – mid
femur length; MFW – mid femur width (in the middle);
HFL – hind femur length (from the base to the tip of the
knee); HFW – hind femur width (in its widest parts);
VW – vertex width; EW – compound eye width (in
frontal view between the eyes, or in dorsal view between
the place of the supraocular lobes); SW – scutellum (or
frontal ridge) width (between the facial carinae); and IAH
– infrascapular area height (in its highest part).
The following measurements were also made:
MAL – middle segment of antenna length (7th or 8th
segment); MAW – middle segment of antenna width (7th
or 8th segment); 1stAW – first antennal segment width
(in the middle of the segment) or the scapus width; HTL
– hind tibia length (from knee to base of apical spine);
n(HTIS) – number of inner spine on hind tibia; n(HTOS)
– number of outer spine on hind tibia; 1stTL – first tarsal
segment (basal) length; 3rdTL – third tarsal segment
(apical) length (without claws); SGPL – subgenital
plate length; SGPW – subgenital plate width; OVDL –
ovipositor dorsal valve length; OVDW – ovipositor dorsal
valve width (maximum, including denticle); OVVL –
ovipositor ventral valve length; and OVVW – ovipositor
ventral valve width (maximum, including denticle).
All measurements are shown in millimeters. All of the
collected specimens examined in the study were mounted
and then deposited to the Central Mindanao University,
University Museum, Zoological Section, Tetrigidae
Collection.
Studied area. The study was conducted in Mt. Agad-agad,
Sitio Langinlanon, Barangay Pugaan, Iligan City, Lanao
del Norte and Marilog District, Davao City Philippines –
as shown in Figure 1 and are listed in Table 1. Other than
these localities, there are no other georeferenced localities
for this species. We give approximate coordinates for
Davao and Zamboanga, the localities for which Günther
reported this species. Table 1 shows all the known
(published) localities together with their accompanying
references, as well as new localities present in this study.
RESULTS AND DISCUSSION
Family Tetrigidae Rambur, 1838
Subfamily Cladonotinae Bolívar, 1887
Genus Diotarus Stål, 1877
Species included: Diotarus galeatus Bolívar, 1887,
endemic to Luzon (Angat and Norzagaray); Diotarus
ikonnikovi Bey-Bienko, 1935, also endemic to Luzon
(Los Baños); Diotarus pupus Bolívar, 1887, known from
the Philippines and with records from Sulawesi that need
to be checked.
Species verrucifer Stål, 1877 by original monotypy.
Type locality: Philippines, no specified type locality in
Semper’s collection.
Distribution: Philippines. Mindanao: Lanao del Norte:
Iligan City (new locality record); Davao Region: Marilog
District (new locality record)
Material examined in this study. Philippines, Mindanao
Island, Lanao del Norte, Iligan City, 08°27’31.30”N,
125°35’43.20”E; 325 masl, 24-ii-2020, 1 male and 2
females, coll. A.B. Mohagan, R.R. Patano, F.P. Coritico,
and V.B. Amoroso [repository (CMU-UM)]; 1 male and
5 females, same locality: Davao del Sur: Davao City,
Marilog District, Marilog Forest Reserve, 07°27’13.74”N,
Mohagan et al.: Morphology and Morphometrics of
Diotarus verrucifer
Philippine Journal of Science
Vol. 149 No. 3, September 2020
572
Table 1. Localities in from which Diotarus verrucifer is known, together with information on coordinates, elevation, date of research, and
reference for each.
Locality Coordinates Elevation Date Reference
Unknown region, Philippines Unknown Unknown 1887 Stål 1877
Mindanao: Davao* 07°11’26.55”N,
125°27’19.23”E
Unknown 1938 Günther 1938
Mindanao: Zamboanga* 06°55’17.19”N,
122°04’44.45”E
Unknown 1938 Günther 1938
Mt. Agad-agad, Sitio Langinlanon, Barangay
Pugaan, Iligan City, Lanao del Norte
08°27’31.30”N,
125°35’43.20”E
216–420 masl February 2020 Present study
Lawi-lawi Forest Reserve, Sitio Maharlika and
Sitio Calinan, Barangay Baganihan
07°27’13.74”N,
125°15’1.12”E
1220–1240 masl February 2018 – July
2019
Present study
Mt. Malambo, Barangay Datu Salumay 07°29’87”N,
125°15’22.23”E
1,151–1,178
masl
February 2018 – July
2019
Present study
Lola Mommy’s Rainforest, Sitio Epol,
Barangay Baganihan
07°27’19.73”N,
125°14’33.37”E
1,197–1,345
masl
February 2018 – July
2019
Present study
Mt. Ulahingan, Sitio Tagumpay, Barangay Datu
Salumay
07°28’29.89”N,
125°16’36.77”E
1,280–1,320
masl
February 2018 – July
2019
Present study
*Günther’s records were identified with uncertainty, but they likely belong to D. verrucifer, as D. galeatus seems to endemic to Luzon Island.
Figure 1. Map of the Philippines (A) and Mindanao (B) showing the distribution of Diotarus verrucifer Stal, 1877. The study was conducted
from months of February 2018 to February 2020. Localities are shown in Figure 1 and in Table 1.
Philippine Journal of Science
Vol. 149 No. 3, September 2020
Mohagan et al.: Morphology and Morphometrics of
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573
125°15’1.12”E, 1,171 masl, 18-ii-2020, coll. A.B.
Mohagan, R.R. Patano Jr., E.P. Leaňo, M.G. Melencion,
and V.B. Amoroso [repository (CMU-UM)].
Diagnosis. The species can be easily distinguished from
the other three species of the genus Diotarus by the
shape of the pronotum (Figure 2). Diotarus ikonnikovi
and Diotarus pupus both have smooth pronotum,
while Diotarus galeatus and D. verrucifer have warty.
D. verrucifer is the most similar to D. galeatus from
Luzon Island. The two species probably had the same
ancestor and represent geographical varieties of it. Head
morphology is very similar, if not the same between the
two. D. verrucifer has a somewhat stouter pronotum, and
the general outline is a bit different. D. galeatus has more
uneven pronotum, and the coloration is different – for
example, the carina is striped.
Updated Description of Diotarus verrucifer Based on
Mindanao Specimens
(nine specimens altogether: two males, seven females)
Head. Head situated at the bottom of the anterior apex
of the tectiform pronotum in the lateral view. Vertex,
Figure 2. Diotarus verrucifer Stål, 1877 female: (A) dorsal view, (B) lateral view, and (C) ventral view. Scale bar:
10 mm.
Mohagan et al.: Morphology and Morphometrics of
Diotarus verrucifer
Philippine Journal of Science
Vol. 149 No. 3, September 2020
574
seen from above, has a blunt or clearly rounded process
produced before the eyes. Facial carinae of the frontal costa
are considerably widened and are enclosing a broadened
scutellum (broader than the scapus). Supraocular lobes
are absent (dorsal view). Vertex is almost twice as wide
as a compound eye (dorsal view). Lateral ocelli are
situated in the lower third of the compound eyes height.
Frontal costa are not strongly projected in lateral view.
Scutellum is wide, which is almost 1 mm (Table 2). Scape
is considerably narrower than scutellum. Compound eyes
are also touching the anterior margin of pronotum, while
the occipital area is narrow (Figure 3).
Antennae. Antennae are filiform and thicker. Antennal
Table 2. Measurements (in mm) and counts of Diotarus verrucifer Stål, 1877. All the measurements are in millimeters.
Body
parameters
Females Males Average
female
Average
male
BL 15.3 15.8 14.8 14.6 15.4 14.8 15.3 14.1 13.7 15.14 13.9
PL 14.6 14.5 14.3 14.2 14.4 14.2 14 13 12.6 14.31 12.8
PW 6.1 6 5.8 5.6 6.1 5.8 6 5.5 5.2 5.91 5.35
PH 6.1 6 5.7 5.5 5.6 5.6 5.55 5.1 4.6 5.72 4.85
HFL 7.4 7 7.2 7.2 7.1 7 7.2 7 6.6 7.16 6.8
HFW 3.7 3.7 3.65 3.5 3.45 3.55 3.5 3.2 3 3.58 3.1
VW 2.5 2.6 2.5 2.4 2.4 2.5 2.5 2.1 2.2 2.49 2.15
EW 1.2 1.2 1.1 1.1 1.2 1.2 1.15 1.1 1.1 1.16 1.1
SW 1 1 0.9 1.1 1 1 1.1 0.9 0.8 1.01 0.85
IAH 2.5 2.5 2.4 2.5 2.45 2.5 2.4 2.1 2 2.46 2.05
MAL 0.28 0.3 0.29 0.28 0.29 0.29 2.29 0.21 0.18 0.57 0.195
MAW 0.11 0.14 0.13 0.11 0.11 0.14 0.15 0.09 0.09 0.13 0.09
1ST AW 0.09 0.1 0.09 0.09 0.09 0.09 0.1 0.08 0.07 0.09 0.08
FFL 4.6 4.6 4.6 4.5 4.5 4.6 4.5 4.2 4.1 4.56 4.15
FFW 1.1 1.2 1.1 1.1 1.1 1.2 1.1 1.1 0.7 1.13 0.9
MFL 4.5 4.6 4.4 4.4 4.4 4.5 4.45 4.2 3.2 4.46 3.7
MFW 1.4 1.45 1.3 1.4 1.3 1.4 1.2 1.2 1 1.35 1.1
HTL 7.4 7.4 7.2 7.2 7.3 7.45 7.3 6.6 4.8 7.32 5.7
n(THIS)R 5 5 5 5 5 5 5 5 5 5.00 5
n(THIS)L 5 5 5 5 5 5 5 5 5 5.00 5
n(THOS)R 0 0 0 0 0 0 0 0 0 0 0
n(THOS)L 0 0 0 0 0 0 0 0 0 0 0
1ST TL 1.5 1.6 1.5 1.45 1.4 1.55 1.4 1.2 1.1 1.49 1.15
3rd TL 1.1 1.15 1.1 1.1 1 1.1 1 0.9 0.8 1.08 0.85
SGPL 0 0 0 0 0 0 0 1 0.9 0.00 0.95
SGPW 0 0 0 0 0 0 0 0.9 0.8 0.00 0.85
OVDL 2 2.1 2 1.9 2 1.9 2 0 0 1.99 0
OVDW 0.7 0.8 0.8 0.8 0.75 0.75 0.8 0 0 0.77 0
OVVL 1.8 1.8 1.8 1.9 1.8 1.8 1.8 0 0 1.81 0
OVVW 0.6 0.7 0.7 0.6 0.6 0.65 0.7 0 0 0.65 0
grooves are situated just next to the lower margins of
the compound eye. Antennae have 15 segments in males
and 16 in females: 1st scape, 2nd pedicel, and 3rd and 4th
basal segment (which is almost of the same size with the
first segment) are present; 5th–9th central segments are
much longer and thicker than the 1st segment; 10th–12th
apical segments are almost of the same size with the
central segments in which the last segment (14th, or 15th
in females) is reduced and pointed in shape (Figure 3).
Pronotum. Dorsum of pronotum is granulated and rough;
it exhibits variable coloration from black to pale yellow; it
is always with strong median carina. It ranges 14.0–14.6
mm in length and 5.6–6.1 mm in width for females (Table
Philippine Journal of Science
Vol. 149 No. 3, September 2020
Mohagan et al.: Morphology and Morphometrics of
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575
2). Pronotum is elevated in the anterior part, humpy, and
with one or more short, upper multi-peaked ridges on
the median carina, or irregular elevations. Median carina
descends towards the pronotal apex – the process is on
a lower level than the anterior part. Pronotum covers
almost the entire abdomen but sometimes not the tip of
the abdominal apex and not surpassing the hind femora.
Anterior margin is projected forwards. Median carina is
continuous and more elevated in the anterior part than in
the posterior. Extralateral carinae are strong, composed of
tubercles. Interhumeral carinae are absent. Humeroapical
carinae fuses with external lateral carinae obtuse, forming
a right angle. Humeral angles are absent. Paranota has no
spine-like projections along its length (Figure 4).
Wings. Wings are absent, as it is a flightless species.
Abdomen. The abdomen is colored black to yellow with
a spine-like subgenital plate (in females). The width of
each of the 8 abdominal segment segments is about 0.85
mm (Figure 4).
Figure 3. Diotarus verrucifer morphology: (A) antenna, (B) head in dorsal view, (C) head in frontal view, and (D)
head and paranota in lateral view. Scale bars: 2.4 mm.
Legs. Dorsal margin of the anterior and middle femora
has an evident carina. Legs are variable in color, from
black to pale yellow. Dorsal margin of fore femora has
three strong undulations, both on the dorsal and ventral
margins. Dorsal and ventral margins of mid femora
are straight or bearing weak undulations. All the tibiae
bear spines in the distal parts (Figure 5). Hind femora
are about twice as long as wide. External outer area
has transverse dark-colored ridges. Genicular teeth and
antigenicular teeth are small and blunt. Hind tibiae are
variable in color, from black to yellow. Fore, mid and
hind tarsi are black to yellow in color. Middle segments
are much longer compared to 1st and 3rd segments of the
fore, mid, and hind tarsi (Table 2). Pulvilli of the hind
tarsi rounded to slightly right-angled, but not sharp or
angulate (Figure 5).
Local name. There are no known local names reported
on this species according to the local people in Marilog
District, Davao City and Pugaan, Iligan City, Lanao del
Norte, Philippines. With that, we propose English name
“Mindanao helmed pygmy grasshopper.” “Helmed”
Mohagan et al.: Morphology and Morphometrics of
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Philippine Journal of Science
Vol. 149 No. 3, September 2020
576
Figure 4. Pronotum and abdominal morphology of Diotarus verrucifer Stål, 1877 female: (A) dorsal view and
(B) lateral view. Scale bar: 10 mm.
Figure 5. Leg morphology of Diotarus verrucifer Stål, 1877: foreleg (A – outside, and B – inside), middle leg
(C – outside, and D – inside), and hind leg (E – outside, and F – inside). Scale bar: 7 mm.
Philippine Journal of Science
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Figure 7. Habitat of Diotarus verrucifer Stål, 1877: (A) view on Mt. Agad-agad (216–420 masl) and (C) view
on Mt. Malambo (1,151–1,345 masl); habitat of the species in agroforest of (B) Mt. Agad-agad and
(D) montane forest in Marilog District.
refers to their anterior margin of pronotum that looks
like a helmet, while “Mindanao” refers to the island the
species is inhabiting.
Habitat and ecology. D. verrucifer was observed perching
in ferns and shrubs in the forest patches Marilog District,
Davao City, together with Arulenus miae Skejo et
Caballero, 2016 or the Mia’s pygmy devil and Spartolus
pugionatus Stål, 1877 or the Mindanao yellow striped
giraffehopper (Figure 6). In Mt. Agad-agad, Pugaan, Iligan
City, Lanao del Norte Philippines, they are mostly observed
in leaf litters. Both areas are considered secondary forests
and some are agroforest areas, with remnants of loggings
and other anthropogenic disturbances such as agricultural
activities. The species was observed mostly in dense
forest patches in Marilog District, which means it prefers
untouched rainforest habitats, just as Arulenus miae (Skejo
and Caballero 2016) and Spartolus pugionatus Stål, 1877
while two individuals were just observed in forest patch of
Mt. Agad-agad. Marilog District is mostly composed of a
montane forest with an elevation of 1,151–1,345 m, while
Mt. Agad-agad is can be considered as an agroforest as it
is composed of forest patches (Figure 7). Altogether, nine
individuals (two males and seven females) were examined
in the present study, but much more were observed in the
field (approximately more than 40).
Variability. Variability of Diotarus verrucifer can be
noted on four morphological characters mostly observed
in the pronotum – (1) purely black to pale yellow in color
throughout the body; (2) having tectiform pronotum in
which its anterior part is extending above the head with
a truncated apex, or more flat pronotum; (3) pronotum
with a leaf-like pattern in the middle down to the apex,
Figure 6. Diotarus verrucifer Stål, 1877 (A and B) in its natural habitat perching on shrubs and ferns.
Mohagan et al.: Morphology and Morphometrics of
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Philippine Journal of Science
Vol. 149 No. 3, September 2020
578
which is formed by three rows or ridges in the pronotal
process; and (4) having various undulations on fore and
mid femora with a rough like textures.
ACKNOWLEDGMENTS
This study would not be possible without the help of
the Commission on Higher Education, as the funding
agency of our research in Marilog District, as part of
DARE TO (Discovery-Applied Research and Extension
for Trans/Interdisciplinary Opportunities) program
entitled “Saving Terrestrial Biodiversity: Inventory,
Assessment, Conservation and Capability Building in
Marilog Forest Reserve, Southern Mindanao, Philippines”
and to the National Research Council of the Philippines
for also funding our research in Mt. Agad-agad entitled
“Biodiversity, Inventory, Assessment and Conservation
for Ecotourism Development in Mt. Agad-agad, Lanao del
Norte”; to the MAMATRIPCEDI (Matigsalug-Manobo
Tribal People Council of Elders Davao Inc.) of Marilog
District; to the local researchers and also to stakeholders
for collaboration; to the Department of Environment and
Natural Resources for the issuance of the GP; and to the
College of Veterinary Medicine of Central Mindanao
University, Musuan, Bukidnon for the issuance of IACUC
permit. The research was also not possible without the
valuable help of our University President Dr. Jesus
Antonio G. Derije in implementing our research. Finally,
the authors would like also to thank and acknowledge
Josip Skejo for the identification of the specimens and for
sharing his knowledge on pygmy grasshoppers.
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