Dtsch. Entomol. Z. 71 (2) 2024, 265-287 | DOI 10.3897/dez.71.125877

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Integrative taxonomic revision of the grasshopper genera Parapetasia
Bolivar, 1884, and Loveridgacris Rehn, 1954 (Orthoptera,
Pyrgomorphidae), with description of a new species of Loveridgacris

Jeanne Agrippine Yetchom Fondjo!*, Martin Husemann*, Armand Richard Nzoko Fiemapong?®,
Alain Didier Missoup', Martin Kenne!, Maurice Tindo', Oliver Hawlitschek*,
Tarekegn Fite Duressa*°, Sheng-Quan Xu®, Wenhui Zhu®°, Claudia Hemp’

1 Zoology Unit, Laboratory of Biology and Physiology of Animal Organisms, Graduate School in Fundamental and Applied Sciences, University of
Douala, Douala, Cameroon

Staatliches Museum fiir Naturkunde Karlsruhe, Karlsruhe, Germany

University of Neuchatel, Neuchatel, Switzerland

Department of Evolutionary Biology and Environmental Studies, Universitat Ziirich, Ziirich, Switzerland

School of Plant Sciences, College of Agriculture and Environmental Sciences, Haramaya University, Dire Dawa, Ethiopia

College of Life Sciences, Shaanxi Normal University, Xi’an, China

N DOD oO fF W PY

Senckenberg Biodiversity and Climate Research Center, Frankfurt, Germany
https://zoobank. org/8 108C5 BO-40C9-40CA-A 38B-8805F'173900D

Corresponding author: Jeanne Agrippine Yetchom Fondjo (jayetchomfondjo@gmail.com)

Academic editor: Susanne Randolf # Received 21 April 2024 # Accepted 19 September 2024 @ Published 21 October 2024

Abstract

The taxonomic status of the Pyrgomorphid genera Parapetasia Bolivar, 1884, and Loveridgacris Rehn, 1954 is complex and
challenging. Here, we use a combination of morphological, distributional, and genetic data to revise the two genera and provide new
information on their diversity. We describe a new species, Loveridgacris tectiferus sp. nov., from Tanzania and formally resurrect
the status of Parapetasia rammei as a valid species within Parapetasia, resulting in two species in Parapetasia (P. femorata and
P. rammei) and two in Loveridgacris (L. impotens and L. tectiferus sp. nov.). We also sequenced the COI and 16S genes of 10
Pyrgomorphidae species and provided the first phylogeny of the group. Our data show that all species are clearly distinct and
represent molecular operational taxonomic units (mOTUs), with the exceptions of L. impotens and L. tectiferus sp. nov., which
are morphologically clearly distinct but for which the concatenated sequence alignments of the two individual gene datasets (COI
and 16S) do not provide sufficient information. In addition, high interspecific distances were found between Parapetasia and
Loveridgacris. Moreover, the complete mitogenomes of L. impotens and L. tectiferus sp. nov. were sequenced using next-generation
sequencing technology. The total lengths of the assembled mitogenomes were 15,592 bp and 15,737 bp, representing 13 protein-
coding genes, 22 transfer RNA genes, two ribosomal RNA genes and one D-loop region, respectively. To aid in identification, we
present a key for the two genera, including a key to species. This study provides insights into the morphology, distribution, and
phylogeny of Pyrgomorphidae in Africa.

Key Words

Afrotropical areas, cytochrome oxidase I, DNA barcoding, mitogenome, phylogeny

Copyright Jeanne Agrippine Yetchom Fondjo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

266 Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Introduction

The family Pyrgomorphidae, which has the type genus
Pyrgomorpha Seville, 1838, is easily identifiable due to its
unique phallic complex, which is relatively uniform with-
in the family, as described by Dirsh in 1961. Members of
this family are commonly referred to as gaudy grasshop-
pers and are renowned for their strikingly vivid coloration,
which serves as a warning to predators. Pyrgomorphidae
can sequester and accumulate plant secondary compounds,
such as cardiac glycosides, from the toxic plants on which
they feed. This accumulation leads to many species dis-
playing aposematism, signaling their toxicity through con-
spicuous coloration. The Pyrgomorphidae family is the
only member of the superfamily Pyrgomorphoidea and
is closely related to the superfamily Acridoidea (Song et
al. 2015). The presence of a groove in the fastigium and
distinctive male phallic structures, including the cingulum
that extends around the ventral side, medially directed en-
dophallic apodemes, an ejaculatory sac that opens to the
genital chamber, and undivided valves of the penis, differ-
entiate the Pyrgomorphidae from other families, as not-
ed by Kevan and Akbar (1964) and Dirsh (1961). Eades
(2000) also described an ejaculatory sac opening to the
genital chamber as a distinguishing characteristic.

The Pyrgomorphidae include 31 tribes, 149 genera, and
487 species (Marifio-Pérez and Song 2018) and are glob-
ally distributed. Most species of Pyrgomorphidae occur
in tropical and subtropical countries of Africa, Asia, and
Australia (Kevan and Akbar 1964; Marifio-Pérez and Song
2018). The tribe Dictyophorini, which includes only five
genera with a small number of species each, namely, Dic-
tyophorus Thunberg, 1815; Maura Stal, 1873; Camoensia
Bolivar, 1882; Parapetasia Bolivar, 1884; and Loveridgac-
ris Rehn, 1954, is distributed in Africa south of the Sahara
(Kevan et al. 1974). The classification of the genera Para-
petasia and Loveridgacris, as well as the species included
in them, is complex. Parapetasia was first established by
Bolivar in 1884 with Parapetasia femorata Bolivar, 1884,
as the type by monotypy. Loveridgacris was first described
by Karsch in 1888, with Petasia impotens Karsch, 1888, as
the type that was later transferred to Parapetasia (Bolivar,
1904). Many researchers followed this classification and
referred to Petasia impotens as Parapetasia. Parapetasia
rammei Sj6stedt, 1923, the second species of Parapetasia,
was described approximately 40 years later. Rehn (1953)
conducted a partial revision of the genus Parapetasia in
1953 and separated the genus into two subgeneric entities
based on the morphology of the pronotum and the size of
the tegmina: Parapetasia (s.str) and Loveridgea (designat-
ed as a new subgenus). Rehn also distinguished two spe-
cies within the subgenus Parapetasia (Parapetasia) based
on female morphological features: the type species Par-
apetasia femorata and the newly described Parapetasia
calabarica Rehn, 1953. In the subgenus Loveridgea, Rehn
included two species, P. (L.) impotens and P. (L.) uluguren-
sis. Additionally, Rehn (1953) placed P. rammei Sj6stedt,
1923, in the subgenus Parapetasia (Loveridgea). Shortly

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thereafter, Rehn (1954) elevated the subgenus Parapetasia
(Loveridgea) Rehn to Loveridgacris Rehn. Dirsh (1956),
Johnston (1956), and Kevan (1962) maintained P. ram-
mei in the subgenus Loveridgacris until Akbar and Kevan
(1964) later raised the subgenus Parapetasia (Loveridgac-
ris) to the generic status of Loveridgacris on the basis of
phallic structures, with L. impotens and L. ulugurensis as
the two species of this genus. Dirsh (1965), in his work on
the genus Parapetasia, included five species distributed in
Central, East, and West African forests, based on Rehn’s
descriptions: Parapetasia femorata, which was recorded
in the lowlands of West African forest areas of the less el-
evated parts of Cameroon (Rehn 1953); Parapetasia cal-
abarica, known only from southern Nigeria (Rehn 1953);
Parapetasia impotens from Tanzania and southeastern
Kenya; Parapetasia ulugurensis Rehn, 1953 from the Ulu-
guru Mountains of Tanzania; and P. rammei, which was
restricted to the more elevated areas of Cameroon. Shortly
thereafter, Kevan et al. (1974) identified L. u/ugurensis as a
synonym of L. impotens and regarded P. calabarica and P.
rammei as “almost undoubtedly mere forms of P. femora-
ta.” Later, Kevan (1977) effectively designated P. calabar-
ica and P. rammei as true synonyms of P. femorata, with P.
calabarica being the micropterous form and P. rammei be-
ing the brachypterous form. Mestre and Chiffaud (2009),
based on Kevan’s (1977) analysis, concluded that the ge-
nus Parapetasia is a monotypic Afrotropical genus.

The taxonomic status of species belonging to the gen-
era Parapetasia and Loveridgacris remains challenging
despite Kevan’s (1977) attempt to synonymize some
of the species. Some researchers, including Hochkirch
(1998) and Seino and Njoya (2018), still consider certain
species, such as L. impotens, L. ulugurensis, P. femora-
ta, and P. rammei, to be distinct. Additionally, P. ram-
mei shares many similarities with L. impotens in terms
of morphology, coloration, and ecological preference,
making it difficult to clearly differentiate between the two
genera. To resolve this taxonomic confusion, molecular
data may be useful. While some studies have generated
DNA sequence data for P. femorata, none have focused
on all species of these genera. Thus, we have conducted
a comprehensive revision of the genera Parapetasia and
Loveridgacris using an integrative approach that com-
bines morphological studies, particularly of the phallic
complex, with molecular analysis. Our study includes the
description of a new Loveridgacris species and proposes
a key to distinguishing valid species in both genera.

Materials and methods

Materials

Field collections and observations were made between
2020 and 2022 at five localities in three regions of Cam-
eroon: Bekob and Iboti (Ebo forest in the littoral region),
Manengouba Mountain (littoral region), Fotouni in the
western highlands of Cameroon, and Somalomo in the Dyja

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Biosphere Reserve (eastern region). Furthermore, addition-
al field trips were made at several locations in Tanzania.
Individuals were collected by sight and hand using a sweep
net. The collected specimens were deposited in the collec-
tions of the Karlsruhe Natural History Museum (SMNK)
and the private collection of Claudia Hemp (CCH). In ad-
dition, some historical specimens belonging to the follow-
ing collections have also been examined: the Hamburg Zo-
ological Museum, Germany (ZMH), and the Museum fir
Naturkunde Berlin, Germany (MfN). Paratypes of P. ram-
mei deposited at MfN Berlin and erroneously labeled as
type (male) and allotype (female) were also examined.

Depositories

MfN: Museum fiir Naturkunde, Leibniz-Institut ftir Evo-
lutions- und Biodiversitatsforschung, Berlin, Germany;
SMNK: Staatliches Museum ftir Naturkunde Karlsruhe,
Karlsruhe, Germany; ZMH: Zoologisches Museum
Hamburg, Leibniz-Institut fir Analyse des Biodiversi-
tatwandels, Hamburg, Germany; CCH: Collection of
Claudia Hemp.

Morphological analysis

Observations of external and internal morphological fea-
tures were made with a Leica M165 C binocular micro-
scope. Photographs of whole specimens were made with
a high-resolution DUN Inc. stacking system (DUN Inc.,
California, USA).

Measurements were obtained using a digital caliper
(at a scale of 0.01 mm). All measurements are given in
millimeters (mm). For all measurements, males and fe-
males were measured separately. For each species, the
following characters were examined: HeadL: length of
head; HeadW: width of head; AntenL: length of anten-
na; 1.0.D.: interocular distance; FastigL: length of fas-
tigium of vertex; PronotL: pronotum length; PronotW:
pronotum width; TegL: length of tegmina; TL: hind tibia
length; FL: hind femur length; fW: hind femur width; and
BodyL: body length, measured from the tip of the frons
to the hindmost tip of the abdomen. The measurements of
the specimens correspond to the average value of the dif-
ferent body parts plus the standard deviation (SD) of all
newly collected samples, as well as all historical samples
held by the ZMH and MfN (Germany).

Dissections and preparations of male and female
genitalia followed the standard methods of Kevan et al.
(1969) and Martinelli et al. (2017). The extracted internal
genitalia were placed in a 1.5 mL microcentrifuge tube
containing a solution of 5 uL proteinase K (20 mg/mL)
and 25 uL buffer (pH 8.0, 10 mM Tris-Cl, 25 mM EDTA,
100 mM NaCl, 0.5% SDS) and were kept overnight in an
incubator at 55 °C. The genitalia were gently separated
from the digestion solution and then kept at 95 °C for
10 minutes to inactivate the enzyme. Preparations were

267

then washed with double-distilled water (ddH,O). Photo-
graphs of male and female genitalia were obtained with
a Keyence VHX-7000 digital microscope (Clustermarket
London, United Kingdom). The terminology for male
genitalia and female spermatheca followed Dirsh (1956,
1957, 1970), Kevan et al. (1974), and Rowell (2013).

Distribution of species

Distributional data were obtained from geographical co-
ordinates recorded during field observations, from local-
ity records taken from specimen labels in different col-
lections of museums, and from records available in the
literature. A distribution map of the species was made
using QGIS 3.28.3 “Firenze” (2023).

DNA extraction, PCR amplification,
sequencing, and data depository

DNA was extracted from the femoral muscle tissue of 24
specimens stored in 96% ethanol. The species considered in
this study are large; hence, only fragments of femoral mus-
cles were used for DNA extraction at the Museum der Natur
Hamburg. A high-salt extraction method was used (Paxton
et al. 1996). The primer pair LCO and HCO (Folmer et al.
1994) was used for amplification of the COI gene, while the
primer pair 16S-F and 16S-R (Palumbi et al. 1991) was used
to amplify the 16S markers. The thermocycling conditions
consisted of an initial denaturation at 94 °C (3 min), 35 cy-
cles at 94 °C (denaturation, 30 s), 50 °C (annealing, 45 s),
72 °C (extension, | min), and a final extension at 72 °C (10
min). Samples were run on a 1% agarose gel stained with
GelRed (Biotium, Remont, CA, USA) to test for amplifica-
tion. Successfully amplified samples were purified with an
ExoSap Enyzme cocktail (VWR, Pennsylvania, USA). The
purified PCR products were then sequenced in both direc-
tions by Macrogen Europe (Amsterdam, Netherlands).

Phylogenetic analyses

Sequences were aligned and checked in GENEIOUS
PRO (Kearse et al. 2012) using the MUSCLE algorithm
(Edgar 2004). The aligned sequences included each one
sequence of Parapetasia femorata (MT011522) and Zo-
nocerus elegans (MT011544), which were downloaded
from GenBank. We checked for pseudogenes (Numts) by
translating sequences into amino acids using the inverte-
brate mitochondrial code and checking for frame shifts.
Furthermore, the National Center for Biotechnology Infor-
mation (NCBI) Blast and BOLD databases were used to
check for species identity (a few related taxa are available
in the databases). We used MRBAYES 3.2 (Ronquist et
al. 2012) to reconstruct the phylogeny. For this, we used
the reversible jump model. Zonocerus elegans was defined

dez.pensoft.net

268

as the outgroup. Analyses were run for 1 million gener-
ations, sampling every 100 generations for a total of 10
000 trees. The first 25% of the samples were discarded as
burn-in. The average split frequencies were less than 0.01,
indicating convergence of the analyses. The final tree was
visualized with FIGTREE v.1.4.2 (Rambaut 2010). The net
evolutionary divergence between groups of sequences was
estimated using MEGA11 (Tamura et al. 2021).

Complete mitochondrial genome assembly,
annotation, and analysis

For L. impotens and L. tectiferus, for which the phyloge-
ny was not resolved when using only COI, the complete

Results

Taxonomy

Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

mitochondrial genome was sequenced by Novogene,
China. Thereafter, the sequences were checked and as-
sembled using MitoZ (Meng et al. 2019). All mitochon-
drial genes were further adjusted and corrected using
GENEIOUS 10.1.3 (Kearse et al. 2012) with the refer-
ence mitogenome of Oxya sinensis (Thunberg, 1815).
The base composition, codon distribution, and length of
the protein-coding genes were calculated in Geneious
Prime 2023.1.2 (Kearse et al. 2012). Nucleotide compo-
sitional differences (composition skew) were measured
using the formula (A — T)/(A + T) for AT skew and (G
— C)/(G+ C) for GC skew (Perna & Kocher, 1995). The
genetic distances for different PCGs among the two
Loveridgacris species were estimated using MEGA 11
(Tamura et al. 2021).

Key to Parapetasia Bolivar and Loveridgacris Rehn

The genus Parapetasia can be easily distinguished from the genus Loveridgacris by several characters:

1 (2)

2 (1)

Fastigium of vertex triangular; eyes small, hemispherical, prominent; pronotal disc sellate; posterior part of metazona
raised, swollen, with median margin slightly or strongly emarginate (Fig. 2A, B, G, H); hind femora upper-median margin
distinctly raised; elytra shortened, or, if brachypterous, slightly reticulated; male subgenital plate with slightly incised
and parallel margins (Fig. 2E); epiphallic bridge narrow; appendices subparallel; ectophallus short; ventral process of
cingulum broadly triangular (Fig. 6A, B, D, E, G, H, J, K) ...... eee Parapetasia Bolivar, 1884 (West and Central Africa)
Fastigium of vertex rounded; eyes of moderate size, oval, not prominent; pronotal disc not sellate; posterior part of
metazona not notably raised nor swollen, its median margin not emarginate (Fig. 2C, F, 1); hind femora upper-median
margin flat, not raised; elytra brachypterous, strongly reticulated; male subgenital plate with fused margins (Fig. 5A, B);
epiphallic bridge wide or broad; appendices divergent; ectophallus elongate; ventral process of cingulum subtriangular

(iS CE ale rre a epee ren eroray em eee renee eg

Genus Parapetasia Bolivar, 1884

Parapetasia Bolivar, 1884 (type species: Parapetasia femorata Bolivar,
1884b, by monotypy).

Parapetasia (Parapetasia) Rehn, 1953

Parapetasia (Loveridgea) Rehn, 1953

Parapetasia (Loveridgacris) Rehn, 1954

Diagnosis of the genus Parapetasia Bolivar. Fastigium
of vertex triangular; tegmina vestigial, or if brachypterous,
strongly reticulated; the eyes small, hemispherical, and
prominent; posterior part of metazona raised, swollen, with
median margin slightly or strongly emarginate; hind femo-
ra upper-median margin distinctly raised; male subgenital
plate with slightly incised and parallel margins; epiphallic
bridge narrow; appendices subparallel; ectophallus short;
ventral process of cingulum broadly triangular.
Differential diagnoses for species of Parapetasia.
Parapetasia femorata can be easily distinguished from
Parapetasia rammei by the following characters: tegmina
dark brown, strongly reduced, vestigial, (yellow—brown
with brown veins, shortened or brachypterous, semilobed

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h SSA REISS ao tee ia ee eee Loveridgacris Rehn, 1954 (East Africa)

in P. rammei), first and/or second abdominal segments
with a lateral black band behind the insertion points of
the femora (absent in P. rammei); anterior projections of
epiphallus large (small in P. rammei); lateral plates sub-
parallel (oblique or divergent in P. rammei); lophi large,
strongly curved (very small, slightly curved in P. ram-
mei), Suprazygomal plate widely rounded (U-shapedin
P. rammei); apodemal lobes only slightly produced ven-
trally (strongly produced ventrally in P. rammei); basal
emargination of cingulum shallow (very deep in P. ram-
mei), endophallic apodemes short (strongly elongate or
slender in P. rammei).

Parapetasia femorata Bolivar, 1884
Figs 1A, B, 2A, D, G, 6A, D, G, J, 7A, D, G

Holotype. GAaBon * 9; 6687; Natural History Museum
Vienna, Austria.

Synonyms. Parapetasia (Parapetasia) calabarica
Rehn, 1953: 121, 122-124, pl. 2: f. 17, pl. 3: f. 26. Kevan
et al. (1974): 229; Kevan (1977): 318 (new synonym).

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

269

Figure 1. Habitus images of Parapetasia and Loveridgacris species. A. P. femorata nymph; B. P. rammei nymph; C. P. femorata
adult; D. P. rammei adult; E. L. impotens nymph; F. L. impotens adult (the black arrow points at the yellowish foamy secretion on

the abdominal segment).

Parapetasia rammei Sjostedt, 1923, p. 10-11, pl. 1: f.
1, 2. Kevan et al. (1974): 229; Kevan (1977): 318 (new
synonym).

Material examined. CAMEROON * | male, 1 female;
Iboti in the Ebo Forest; 4.450°N, 10.450°E; 736 m; 07
Jan. 2022; J.A. Yetchom Fondjo leg. and A.R. Nzoko
leg.; SMNK; CMJ244. ¢ 1 female; Iboti in the Ebo Forest;
4 450°N, 10.450°E; 736 m; 07 Jan. 2022: J.A. Yetchom
Fondjo leg.; SMNK; CMJ245. « 1 female; Somalomo in
the Dja Biosphere Reserve; 3.371°N, 12.733°E; 06 Jun.
2022; A.R. Nzoko leg.; SMNK; CMJ1439. « 1 male; Be-
kob in the Ebo Forest; 4.350°N, 10.420°E; 936 m; 20
Mar. 2021; J.A. Yetchom Fondjo leg.; SMNK; CMJ598.
° 6 females; Mukondje Farm, Mundame-Mungo Fluss;
25 Nov. 1904; R. Rohde leg.; ZMH. « 1 male, 4 females
and 1 nymph; Esosung, Bakossi-Gebirge; 10 Sep. 1909;
C. Rathke leg.; ZMH. ¢ 3 males, 4 females, 4 nymphs;

Esosung, Bakossi-Gebirge; 01 Nov. 1912; R. Rohde leg.;
ZMH. ¢ 1 male; Esosung, Bakossi-Gebirge; 1913; ZMH.
¢ 2 females; Esosung, Bakossi-Gebirge; 1930; O. Krober
leg.; ZMH. * 3 males; Buea, south-West; 1891; S. Preuss
leg.; MfN. * 1 male, 3 females; South; 1891; S. Preuss
leg.; MfN. « 1 male, 1 female; Station Jaunde [ Yaoundé],
Centre; Mar. 1997; V. Carnap S.G. leg.; MfN. « 1 male, 1
female; Dibongo of Sanaga, Littoral; Ld. Kam leg.; MfN.
¢ 1 male, 1 female; Lolodorf, South; L. Conradt S. leg.;
MfN. ¢ 1 female; Victoria [Limbe], south-West; S. Pre-
uss leg.; MfN. * 1 female; Barombi station, south-West;
Preuss S. leg.; MfN. * 1 male; Duala [Douala], Littoral;
Dr Schafer leg.; MfN. ¢ 1 female; Nlobe-Ndunge; 500—
700 m from Edea-Douala, Littoral; Dr Schafer leg.; MfN.
¢ 1 female; Longi; Jun. 1904; MfN. « 1 nymph; Japoma,
Littoral; Dr Schafer leg.; MfN. * 1 nymph; Victoria [Lim-
bé], south-West; Jan. 1898-1899; MEN. ¢ 2 females; north

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270 Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

2mm

Figure 2. Frontal, dorsal and lateral views. A—-C. Frontal view; A. P. femorata;, B. P. rammei;, C. L. impotens; D—-F. Dorsal view;

D. P. femorata; E. P. rammei, F. L. impotens, G—I. Lateral view; G. P. femorata;, H. P. rammei; I. L. impotens.

Mundame, Elephantensee; 21 Jan.—15 Feb. 1996; S. Con-
radt leg.; MfN. * 1 female; Buea, south-West; MEN. « 1
female; Mundame; 1896; MfN. * 2 males; Bissika, Span.
Guinea; Dr Escherich leg.; MfN.

Redescription. Male. Body: robust, depressed, with
very finely or moderately rugose and tuberculated in-
tegument. Head (Figs 1A, 2A, D, G): acutely conical;
fastigium of vertex slightly curved upwards, flat, slight-
ly concave in basal part, distinctly triangular, narrow-
ing toward apex (Fig. 2D); frontal carina hardly visi-
ble; antennae thick, shorter, or only slightly longer than
head and pronotum together, with short transverse or
subtransverse segments, the last apical segment being

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distinctly longer than others. Thorax (Fig. 2D, G):
pronotum with large inflation in front of first sulcus,
strongly tuberculated in anterior part of prozona and
posterior part of metazona, with the posterior part of
the prozona and anterior part of the metazona being
very finely tuberculated; median carinae inconspicuous
and interrupted, lateral carinae absent; inferior margins
of lateral lobes of pronotum straight; prozona shorter
than metazona; posterior margin of metazona strong-
ly emarginate; prosternal process very short, subacute
or obtuse-angular; mesosternal interspace wider than
long. Legs (Fig. 2D, G): hind femur slender, its external
area not expanded, its upper-median margin distinctly

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Figure 3. A-E. L. tectiferus sp. nov. A. Male in frontal view;
B. Female in frontal view; C. Male head and pronotum;
D. Female head and pronotum; E. Male in ventral view; F. Male
of L. impotens in ventral view.

raised; obliquely expanded area at the base of hind fe-
mur strongly pronounced; external apical spine of hind
tibiae present; hind tarsal segments not elongate. Elytra
(Fig. 2G): strongly reduced, micropterous, not reaching
point of insertion of metathoracic legs, with rounded
posterior margins. Abdomen (Fig. 2D): often annulat-
ed; abdominal tergites each with a trigonal medio-dorsal
tubercle; male supra-anal plate subtriangular; male sub-
genital plate compressed toward apex above, margins
slightly incised, parallel; male cerci conical (Fig. 7A).
Epiphallus (Fig. 6A): bridge narrow, its anterior margin
emarginate; anterior projections large, fairly prominent,
not broadly rounded; appendices broad, subparallel,
with apical lobes having smaller and broader processes,
attached marginally to the basal part of the lateral plates;
lateral plates subparallel, almost straight, directed poste-
riorly, with external margins not expanded; lophi large,
strongly curved upward, anteriorly directed with acute
apex. Ectophallus (Fig. 6D, G, J): central membrane
fairly narrow, rather triangular or subtriangular, marked

271

at its lateral margins by furrows; zygoma broadly trans-
verse, not extending halfway along the cingulum; su-
prazygomal plate widely rounded, moderately wide,
highly shorter than the zygoma; apodemal lobes only
slightly produced ventrally, the apices fairly wide apart;
valves of cingulum small, narrow, and divergent in dor-
sal view; rami of cingulum rather broad in dorsal view,
extending into sheath; dorsal cleft of cingulum rather
narrow, ventral cleft small; suprarami well developed;
basal emargination of cingulum shallow; sheats rather
well developed; ventral process of cingulum short, not
reaching the apex of endophallic apodemes nor the bas-
al thickening of cingulum in ventral view. Endophallus
(Fig. 6D, G, J): endophallic apodemes broad or stout,
rather short, not reaching the basal emargination of cin-
gulum in ventral; aedeagal valves narrow, slender with
button-like apices; aedeagal sclerites stout and shorter,
ventrally directed; spermatophore sac small, ovoid, not
extending beyond the lateral limits of endophallic apo-
demes; gonopore at the middle.

Female. As in male, but larger. Abdomen (Fig. 7D):
subgenital plate in female without carina or keel, its pos-
terior margin rounded and smooth; egg guide prominent,
conical, and slightly elongated; ovipositor valves large,
not sinuate. Genitalia (Fig. 7G): spermatheca thick, lack-
ing an apical pocket, with a laminated appearance in the
apical part; median longitudinal groove of genital cham-
ber reduced; spermatheca duct short, with an elongate,
terminally thickened region; secondary diverticulum of
spermathecal appendage of varying shape.

Color. Predominantly brownish, sometimes with or-
ange or red markings; eyes entirely black in adults; an-
tennal scape black; head brownish, margin of vertex, an-
tennae light brown or dark brown in some parts in adults;
sternum light brown and black in some parts; dorsal part
of mesothorax with a broad black band bordered later-
ally by the elytra; elytra dark brown; first and/or second
abdominal segments with a lateral black band behind the
insertion points of the femora; lower external, lower in-
ternal, and medial internal hind femoral areas blackish;
fore and middle femora, outer-medial, upper-external and
upper-internal areas of hind femora dark-brown; hind tib-
iae light brown; tarsi light brown; cerci black.

Nymph (Fig. 1A). Eyes dark-red; antennae predomi-
nantly black with yellow apex; hind knee predominantly
yellow with black median mark.

Measurements. Male. Body length 37.80-40.45 mm;
Female. Body length 45.98-—62.69 mm. Adult P. femorata
individuals exhibit very large size variations in both sex-
es. Additional information on the measurements is given
inLable*l::

Geographical distribution (Fig. 8). Parapetasia
femorata has been recorded from Gabon, Cameroon, Ni-
geria, and Equatorial Guinea. In Cameroon, P. femorata
was discovered in two localities within the proposed Ebo
Forest, namely, Bekob and Iboti, as well as in Somalomo,
a location within the Dja Biosphere Reserve, and Ngout-
adjap and Zamakoe.

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272

Table 1. Measurements in millimeters (mm) of the examined Parapetasia and Loveridgacris species; n: number of individuals; FastigL:
length of fastigium of vertex; PronotL: pronotum length; PronotW: pronotum width; TegL: length of wings; TL: hind tibia length; FL:
hind femur length; Fw: hind femur width; and BodyL: body length, measured from the tip of the frons to the hindmost tip of the abdomen.

Species
Parameters

HeadL
HeadW
AntenL
1.0.D.
FastigL
PronotL
PronotW
TegL
TL

FL

FW
BodyL

Species
Parameters

HeadL
HeadW
AntenL
1.0.D.
FastigL
PronotL
PronotW
TegL
TL

FL

FW

BodyL

The measurements represent the average value of the different body parts plus the standard deviation (SD). The range reffers to the minimum and
maximum values.

Ecology. Parapetasia femorata 1s distributed through-
out the lowlands of West and Central Africa and is exclu-

Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Parapetasia femorata Bolivar, 1884

Parapetasia rammei Sjéstedt, 1923

Male Female Male Female
(Mean + SD) (Range) (Mean + SD) (Range) (Mean + SD) (Range) (Mean + SD) (Range)
6.06 + 0.43 5.69-6.54 7.30 + 0.98 5.50-9.11 5.18 + 0.32 4.95-5.40 6.23 + 0.57 5.32-6.81
(n=5) (n= 15) (n= 2) (n= 7)
5.03 + 0.42 4,53-5.58 6.22+0.31 5.53-6.61 4.51 +0.05 4.47-4.54 5.33 + 0.31 5.09-5.96
(n=5) (n= 15) (n= 2) (n=7)
14.99 + 1.33 13.13-16.49 16.68 + 1.56 12.84-18.61 11.65 + 0.53 11.27-12.02 13.85+0.61 13.00-14.52
(n=5) (n= 15) (n= 2) (n= 7)
2.69+0.14 2.51—2.82 3.34+0.17 3.06-3.63 2.77 + 0.02 7.75-2.78 3.30+0.17 3.15-3.66
(n=5) (n= 15) (n= 2) (n=7)
2.24+0.27 1.83-2.54 3.02 + 0.39 2.46-3.80 1.88 + 0.01 1.87-1.89 2.78 + 0.30 2.41-3.04
(n=5) (n= 15) (n= 2) (n=7)
10.94 + 0.73 10.08-11.64 13.84 + 0.92 12.41-15.37 10.37+0.11 10.29-10.44 13.50 + 0.77 12.63-14.64
(n=5) (n= 15) (n= 2) (n=7)
9.59 + 0.58 8.73-10.25 12.81 40.79 11.41-14.22 0.81 £0.59 8.39-9.23 11.28+0.61 10.26-12.11
(n=5) (n= 15) (n= 2) (n=7)
2.71 £0.82 1.96-3.86 414+0.89 2.53-5.48 10.29 + 0.76 9.75—10.83 15.03 £1.11 13.15-16.60
(n=5) (n= 15) (n = 2) (n=7)
16.01 0.87 15.15-17.27 19.69 + 1.26 17.10—21.26 13.07 + 0.83 12.48-13.65 17.23 + 0.81 15.86-18.04
(n=5) (n= 15) (n = 2) (n=7)
17.62 + 1.04 16.47-19.12 21.89+4 1.31 19.15—23.47 15.10+0.21 14.95-15.25 19.64 + 0.58 18.83-20.31
(n=5) (n= 15) (n= 2) (n=7)
3.65 + 0.07 3,553.71 4.47+0.37 3.77-5.08 3.55 40.27 3.36-3.74 4.1740.21 3.87-4.49
(n=5) (n= 15) (n= 2) (n=7)
38.60 + 1.06 37.80—-40.45 54.00 + 5.06 45.98-62.69 31.10 + 1.80 32.83-35.37 45.75 + 2.40 43.22-49.73
(n=5) (n= 15) (n= 2) (n= 7)
Loveridgacris impotens (KKarsch, 1888) Loveridgacris tectiferus Hemp sp. nov.
Male Female Male Female
(Mean + SD) (Range) (Mean + SD) (Range) (Mean + SD) (Range) (Mean + SD) (Range)
8.324 0.42 8.02-8.62 8.63 + 0.48 8.19-9.12 7.41 (n=1) NA 6.87 + 0.60 6.44-7.29
(n=2) (n=4) (n= 2)
7.11 + 1.09 6.34-7.88 7.10+0.47 6.40-7.41 8.95 (n= 1) NA 9.18+40.53 9,558.81
(n= 2) (n=4) (n= 2)
19.41 43.13 17.19-21.62 20.42 + 1.21 19.21-22.10 21.30 (n=1) NA 19+ 00 19.00-19.00
(n= 2) (n=4) (n= 2)
3.55+0.17 3.43-3.67 4.17+0.22 3.99-4.49 410 (n=1) NA 4.20+0.14 4.10-4.30
(n= 2) (n=4) (n= 2)
3.67 + 0.34 3.43-3.91 3.89 + 0.30 3.46-4.15 3.50 (n= 1) NA 3.05 + 0.07 3.00-3.10
(n= 2) (n=4) (n= 2)
16.05 + 1.44 15.03-17.06 16.41 + 1.75 14.54-18.70 13.6 (n= 1) NA 21.60+ 0.14 21.50-21.70
(n= 2) (n=4) (n= 2)
13.39+4 1.22 12.52-14.25 13.53 + 1.34 11.89-15.11 9.30 (n= 1) NA 9.50 + 00 9.50-9.50
(n= 2) (n=4) (n= 2)
14.96 + 2.02 13.53-16.38 17.88 + 1.54 16.59-19.80 29.00 (n= 1) NA 27.70 + 0.71 27.20-28.20
(n= 2) (n=4) (n= 2)
21.62+41.11 20.83-22.40 24.20 + 1.12 22.99-25.69 20.67 (n= 1) NA 23.194 00 (n 23.19-23.19
(n= 2) (ni=4) =2)
24.12 + 0.69 23.63-24.60 27.29 + 0.75 26.19-27.85 25.00 (n= 1) NA 24.25 + 0.49 23.90-24.60
(n= 2) (n=4) (n= 2)
4.25+0.22 4.09-4.40 4.82+0.34 4.40-5.11 3.44 (n= 1) NA 4.65+0.21 4.80-4.50
(n= 2) (n=4) (n= 2)
55.544 6.58 50.88-60.19 58.66 + 6.77 48.72-63.69 51.20 (n= 1) NA 50.65 + 4.17 47.70-53.60
(n= 2) (n=4) (n= 2)

Parapetasia rammei Sjéstedt, 1923
Figs 1C, D, 2B, E, H, 6B, E, H, K, 7B, E, H

sively found in forest habitats with a closed canopy and

close proximity to marshy areas, where litter is abundant.
Within forest habitats, the species is geophilous. Para-
petasia femorata is present throughout the year in Camer-
oon, with the highest abundance observed during the dry
season from November to January. This species is known

to produce foamy secretions on tergites 3 and 4.

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Holotype. CAMEROON °* 9; Bare-Dschang, [Stockholm]

Paratypes. CAMEROON * | 4, Bamenda; Adametz
S.G. leg.; MfN URL: http://cool.mfn-berlin.de/u/bb659e
[MfN]. * 1 2: Bangwe; 1000 m; Mitte V-Mitte VI. 99
[mid-May—mid-Jun. 1999]; G. Conrau S. leg.; MfN

URL: http://cool.mfn-berlin.de/u/d4c8af [MEN].

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

213

Figure 4. A, B. L. tectiferus sp. nov. A. Male; B. Female; C, D. L. impotens, C. Male; D. Female.

Material examined. CAMEROON. ° | male; Bamen-
da; Adametz S.G. leg.; URL: http://cool.mfn-berlin.de/u/
bb659e (MfN). * 1 female; Bangwe; 1000 m; Mitte V—
Mitte VI. 99 [mid-May—mid-Jun. 1999]; G. Conrau S.
leg.; URL: http://cool.mfn-berlin.de/u/d4c8af (MfN). * 1
female; Fotouni, West; 5.362°N, 10.246°E; 15 Jun. 2020;
J.A. Yetchom Fondjo; SMNK; CMJ678. « 1 female; Fo-
touni, West; 5.362°N, 10.246°E; 13 Aug. 2020; J.A. Ye-
tchom Fondjo leg.; SMNK; CMJ61. * 2 males, 2 females;
Fotouni, West; 5.362°N, 10.246°E; 16 Jan. 2021; J.A.
Yetchom Fondjo; SMNK; CMJ679. « 1 female; Fotount,
West; 5.362°N, 10.246°E; 14 Mar. 2022; J.A. Yetchom
Fondjo; SMNK; CMJ63. * 2 females; Fotouni, West;
5.362°N, 10.246°E; 15 Mar. 2022; J.A. Yetchom Fondjo;
SMNK; CMJ64.

Redescription. Male. Body: robust, depressed, with
strongly rugose and tuberculated integument. Head
(Fig. 2B, E, H): acutely conical; fastigium of vertex slight-
ly curved upwards, flat, slightly concave in basal part,
distinctly triangular and narrowing toward apex; anten-
nae thick, shorter than head and pronotum together, with
short transverse or subtransverse segments, the last apical
segment being distinctly longer than others. Thorax (Fig.
2E, H): pronotum with large inflation in front of first sul-
cus, strongly and intensely rugose with more pointed tu-
bercles; median carinae interrupted, lateral carinae absent;

inferior margins of lateral lobes of pronotum straight; pro-
zona shorter than metazona; posterior margin of metazona
slightly emarginate in the middle; prosternal process very
short, subacute; mesosternal interspace wider than long.
Legs (Figs 1D, 2E, H): hind femur slender, its external area
not expanded; upper-median margin of hind femora dis-
tinctly raised; obliquely expanded area at the base of hind
femur strongly pronounced; external apical spine of hind
tibiae present; hind tarsal segments not elongate. Elytra
(Fig. 1D, 2E, H): less reticulated, shortened or brachypter-
ous, oval, semilobed, reaching dorsally the third abdominal
tergite, anterior margins rather curved, with very broadly
rounded posterior margins. Abdomen (Figs 1D, 2E, H):
often annulated; abdominal tergites each with a trigonal
medio-dorsal tubercle; male subgenital plate compressed
toward apex above, margins slightly incised, parallel;
male supra-anal plate conical; male cerci (Fig. 7B) coni-
cal, straight. Epiphallus (Fig. 6B): bridge narrow, its an-
terior margin curved emarginate, posterior margin almost
straight; anterior projections small; lateral plates oblique or
divergent, its external margins fairly expanded: appendices
of epiphallus narrow, subparallel, with apical lobes having
only broader terminal processes, smaller processes absent,
attached marginally to the basal part of external expansion
of lateral plates, lying marginally to the external expansion
of lateral plates; lophi very small or short, slightly curved,

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274 Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Figure 5. L. tectiferus sp. nov. A. Male semidorsal; B. Male in lateral view; C. Female semidorsal; D. Female in lateral view;

E. L. tectiferus sp. nov. mating pair; F. Epiphallus.

and anteriorly directed with acute apex. Ectophallus
(Fig. 6E, H, K): central membrane fairly narrow, subtri-
angular, marked at its lateral margins by furrows; zygoma
broadly transverse, not extending halfway along the cingu-
lum; suprazygomal plate rather U-shaped, slightly short-
er than the zygoma; apodemal plates strongly produced
ventrally, the apices fairly close to each other; valves of
cingulum of smaller size, narrow, divergent in dorsal view;
rami of cingulum narrow in dorsal view; dorsal cleft of
cingulum large, ventral cleft narrow; suprarami well de-
veloped, large; sheats of moderate size, the inner margins
separated from each other; ventral process of cingulum
broadly triangular, slender, almost exceeding beyond en-
dophallic apodemes in ventral view; basal emargination
of cingulum very deep; Endophallus (Fig. 6E, H, K): en-
dophallic apodemes of medium size, strongly produced

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forwards ventrally, exceeding beyond the basal emargin-
ation of cingulum; aedeagal valves small, short with but-
ton-like apices, with ventrolaterally directed process in its
distal part; aedeagal sclerites narrow and of moderate size;
pseudoarch small; spermatophore sac small, ovoid, not ex-
tending beyond the lateral limits of endophallic apodemes;
gonopore at the middle.

Female. Similar to male but larger. Abdomen (Fig. 7E):
ovipositor valves large, not sinuate; subgenital plate with-
out carina, narrowed posteriorly, slightly emarginate at
apex; egg guide prominent, conical, and slightly elongat-
ed; median longitudinal groove of genital chamber slen-
der. Genitalia (Fig. 7H): spermatheca thick, lacking an
apical pocket, with a laminated appearance in the apical
part; spermatheca duct short, secondary diverticulum of
spermathecal appendage of varying shape.

Dtsch. Entomol. Z. 71 (2) 2024, 265-287 2/5

Figure 6. Phallic structures. A-C. Epiphallus dorsal view; A. P. femorata; B. P. rammei; C. L. impotens, D-F. Ectophallus + En-
dophallus dorsal view; D. P. femorata, E. P. rammei; F. L. impotens; G—I. Ectophallus + Endophallus ventral view; G. P. femorata;
H. P. rammei; I. L. impotens;, J-L. Ectophallus + Endophallus lateral view; J. P. femorata; K. P. rammei; L. L. impotens. A: appen-
dices; AC: apodemal plate of cingulum; AE: aedeagus (aedeagal valves); AP: anterior projection of epiphallus; B: bridge of epiphal-
lus; BC: basal thickening of cingulum; BE: basal emargination of cingulum; CM: central membrane of epiphallus; CV: valve of
cingulum; DC: dorsal cleft of cingulum; EA: endophallic apodeme; F, marginal furrow separating suprarami and rami of cingulum;
L: lophus of epiphallus; LP: lateral plate of epiphallus; RC: ramus of cingulum; S: sheath of ectophallus; SR: supraramus of cingu-
lum; SZ: suprazygomal plate of cingulum; VC: ventral cleft of cingulum; VP: ventral process of cingulum; Z: zygoma of cingulum.

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2/6 Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Figure 7. A-C. Male cerci; A. P. femorata; B. P. rammei; C. L. impotens,; D-F. Female subgenital plates; D. P. femorata; E. P. ram-
mei. F. L. impotens. G-I. Female spermatheca; G. P. femorata; H. P. rammei; 1. L. impotens. AB: apical bulb of the spermathecal
appendage; EG: egg-guide; FP: floor pouch of the female genital chamber; MLG: median longitudinal groove of the female genital
chamber; PVS: postvaginal sclerite of the female genital chamber; SB: secondary diverticulum of the cecum of the spermatheca;
SC: cecum of the spermatheca; SD: spermathecal duct; SP: spermathecal vesicle; TD: terminal dilatation of the spermathecal duct.

Color. Predominantly grayish; eyes red dark or entire-
ly black in adults; labium, labrum, and mandibles red;
hind knee entirely black in adults; elytra yellow-brown
with brown veins; cerci yellowish or blackish; abdominal
segments all separated by red lines; lower external, low-
er internal, mid internal, and upper internal hind femoral
areas bright black.

Female. Pronotum testaceous brown with blood-like
colored tubercles, more or less blackish below; antennae
and legs reddish brown, more or less outlined with red; the
outer-medial and upper-external area slightly variegated

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with yellowish brown, the margins more or less red, the
tips of the femora black; hind tibiae all reddish brown,
sometimes outlined with a blood-like color.

Male. Apex of antennae, apical 2/3 of hind tibiae,
and ankles with blood-like colored tubercles; margin of
vertex, lower part of antennae, fore and median femora,
upper-external and medio-external areas of hind femora,
anterior and median tibiae, and basal part of hind tibiae
marked with yellow, especially on ribs.

Nymph with entirely black antennae, hind knees com-
pletely yellow.

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Measurements. Male. Body length 32.83—35.37 mm;
Female. Body length 43.22-49.73 mm. Adult individu-
als of P. rammei exhibit very large size variations in both
sexes (Table 1).

Geographical distribution. Parapetasia rammei
(as shown in Fig. 8) is limited to Cameroon. The spe-
cies has been primarily observed in highland zones and
grass fields, specifically in Bare-Dschang (the type lo-
cality), Bamenda, Bangwe, Fotouni, and Mt. Manen-
gouba.

Key to the species of Parapetasia

277

Ecology. Parapetasia rammei is typically associated
with highland ecosystems. The species is commonly found
on bare ground, low vegetation, and sometimes on shrubs
in open vegetation, such as cultivated farms and fallows, as
well as in steppe habitats. In its natural habitat, it can be ob-
served throughout the year, with adults being particularly
abundant during the rainy season. Nymphs are more com-
monly found during this period as well. Foamy secretions
on tergites 7 and 8 are notable characteristics of the species
and are also common in other pyrgomorphid species.

1(2)

2(1)

Tegmina dark brown, strongly reduced, vestigial; first and/or second abdominal segments with a lateral black band
behind the insertion points of the femora; anterior projections of epiphallus large; lateral plates subparallel; lophi large,
strongly curved; suprazygomal plate widely rounded; apodemal lobes only slightly produced ventrally; basal emargina-
HOR Of ClPOlirin SitallGwe ericlOpal ile fapOcennes SORT ts Ferre. cs See eR wa eR, Sy en eek Sata oe eye Pee a eee uses
een. sia Aiea ey SNe pe eN S| een Dee Se Parapetasia femorata Bolivar, 1884 (Gabon, Cameroon, Equatorial Guinea, Nigeria)
Tegmina yellow-brown with brown veins, shortened or brachypterous, semilobed; first and/or second abdominal seg-
ments without a lateral black band behind the insertion points of the femora; anterior projections of epiphallus small;
lateral plates oblique or divergent; lophi very small, slightly curved; suprazygomal plate U-shaped; apodemal lobes
strongly produced ventrally; basal emargination of cingulum very deep; endophallic apodemes strongly elongate or

SOHC eee PL ee See a cee el AL

Genus Loveridgacris Rehn, 1954

Parapetasia (Loveridgea) Rehn (1953)

Parapetasia (Loveridgacris) Rehn (1954)

Parapetasia (Loveridgeacris) Kevan (1962) (subsequent misspelling)
Parapetasia (Loveridgacris): Akbar & Kevan, (1964)

Diagnosis of the genus Loveridgacris. Fastigium of ver-
tex rounded apically; tegmina brachypterous and slight-
ly reticulated; eyes ovate and not prominent; posterior
part of metazona not notably raised nor swollen, its me-
dian margin not emarginate; hind femora upper-median
margin flat, not raised; male subgenital plate with fused
margins; epiphallic bridge wide or broad; appendices
divergent; ectophallus elongate; ventral process of cin-
gulum subtriangular.

Loveridgacris impotens (Karsch, 1888)
Figs 1E, F; 2C, F, I, 3F, 4C, D, 6C, F,1, L, 7C, EI

Petasia impotens Karsch, 1888

Parapetasia impotens Karsch

Holotype. DEUTSCH-OsTAFRIKA [German East Africa].
¢ 1 male; Uzigna [Usegna]; MfN.

Synonyms. Loveridgacris ulugurensis Rehn (1953):
124, 126, pl. 2: f. 18 and 19, pl. 3: f. 23 and 24. Kevan et
al. (1972) 293.229:

Material examined. DEUTSCH-OSTAFRIKA [German
East Africa]. * 1 male; Uzigna [Usegna]; MfN. « 1 male;
Amani; 26 Nov. 1906; S.G. Vosseler leg.; MfN. * 1 male;

RP Ee Pee Parapetasia rammel Sjostedt, 1923 (Cameroon)

Amani; 1 Nov. 1906; S.G. Vosseler leg.; MfN. * 1 male,
1 female; Amani; Nov. 1906; Vosseler S.G. leg.; MfN.
¢ 2 males; Amani; Nov. 1907; S.G. Vosseler leg.; MEN.
¢ 1 female; Amani; 16 Jan. 1906; S.G. Vosseler leg.; MfN.
¢ 1 male; Amani; 30 Nov.—5 Dec.1906; S.G. Vosseler leg.;
MEN. ¢ 1 male; Amani; S.G. Vosseler leg.; MfN. * 1 male;
Amani; 20 Nov— 5 Dec. 1906; S.G. Vosseler leg.; MfN.
¢ 1 male, 6 females, 1 nymph; Amani; S.G. Vosseler leg.;
MEN. TANZANIA. * 1 male; Uluguru-Berge; 11 Dec. 1998;
S. Gotze leg.; MfN. DEUTSCH-OSsTAFRIKA [German East
Africa]. « 1 female; Sigital; Jul. 1903; S. Gotze; MEN.
¢ | female; Muoa, Bez.langa; S. Fischer; MfN. « 1 female;
V. Karger leg.; MfN. * 2 males, 3 females; 1903, vend.1
Apr. 1911; Dr F. Eichelbaum; ZMH. TANzania. « 1 fe-
male; Usambara Nguelo; S. Heinsen; MfN. « 1 female;
Usambara Nguelo; 14 Jun. 1905; H. Rolle leg.; ZMH.
DEUTSCH-OSTAFRIKA [German East Africa]. * 4 males;
1908: S.G. Vosseler; MEN.

Redescription. Male. Body: robust, depressed, with
strongly rugose and tuberculated integument. Head
(Fig. 1F, 2C, F, I): acutely conical; fastigium of vertex
slightly curved upward, flat, slightly concave in basal part,
with rounded apex in dorsal view; antennae thick, shorter
than head and pronotum together, with short transverse
or subtransverse segments, the last apical segment being
distinctly longer than others; eyes oval, of moderate size.
Thorax (Figs 2F, I, 3F, 4C, D): pronotum less testaceous,
not deeply and concavely saddle-shaped, with large infla-
tion in front of first sulcus, moderately rugose with slight-
ly pointed tubercles; posterior part of metazonal disc not
swollen or raised dorsally; median carinae interrupted,
lateral carinae absent; lower margins of lateral lobes of

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2tG

pronotum rather angular; prozona shorter than metazo-
na; median posterior margin of metazona not emargin-
ate; prosternal process very short, triangular, expanded at
its base with angular apex: mesosternal interspace wid-
er than long. Legs (Figs 2F, I, 3F, 4C, D): hind femur
slender; upper-median margin of hind femora flat, not
raised, almost of equal height to upper-external margin;
obliquely expanded area at the base of hind femur less
pronounced; external apical spine of hind tibiae present;
hind tarsal segments not elongate. Elytra (Figs 1F, 2F,
I, 4C, D): shortened or brachypterous, slightly elongated
and strongly reticulated, reaching dorsally the third ab-
dominal tergite, with evenly rounded posterior margins.
Abdomen (Figs 2F, I, 3F, 4C, D): often annulated; ab-
dominal tergites each with a trigonal medio-dorsal tuber-
cle; male subgenital plate compressed toward apex above,
margins fused but not separated; male supra-anal plate
conical; male cerci conical (Fig. 7C). Epiphallus (Fig.
6C): bridge wide or broad, its anterior margin emarginate,
and posterior margin almost straight; anterior projections
large, fairly prominent; lateral plates almost divergent, its
external margins slightly expanded; lophi of larger size,
upcurved and anteriorly directed with acute apex; ap-
pendices divergent, with broad apical lobes bearing long
terminal processes, attached submarginally to the anteri-
or projections and lying close to the lateral plates. Ecto-
phallus (Fig. 7F, I, L): elongate, stout; central membrane
broad, marked at its lateral margins by furrows; zygoma
narrow; suprazygomal plate narrow, slightly shorter than
the zygoma, with rather widely rounded apex; apodemal
plate broad and rounded in lateral view, lobes slightly
produced ventrally, the apices fairly wide apart, without
anterior blunt points; valves of cingulum of smaller size,
divergent in dorsal view; rami of cingulum extending into
sheath; sheath wide, inner margins fairly close to each
other; dorsal cleft of cingulum large; ventral cleft of cin-
gulum of moderate size; supraram1 well developed, large;
sheats well developed: ventral process of cingulum rather
subtriangular, broadly covering the endophallic apode-
mes, slender, but not reaching or extending beyond en-
dophallic apodemes in ventral view; basal emargination
of cingulum shallow. Endophallus (Fig. 7F, I, L): en-
dophallic apodemes moderately produced forward ven-
trally, rather broad, and reaching the basal emargination
of cingulum; aedeagal valves of smaller size, broad, with
button-like apices, and with ventrolaterally directed pro-
cess in its distal part; aedeagal sclerites stout, slender, and
curved; pseudoarch small, distinct, broad; spermatophore
sac small, ovoid, extending beyond the lateral limits of
endophallic apodemes; gonopore distally placed.

Female. As for male, but larger. Abdomen (Figs 4D,
7F): ovipositor valves large, not sinuate; subgenital
plate without a carina, its posterior margin rounded and
smooth; egg-guide prominent, conical, and highly elon-
gated; median longitudinal groove of genital chamber
slender. Genitalia (Fig. 71): spermatheca thick, lacking an
apical pocket, with a laminated appearance in the apical
part; spermatheca duct slender, secondary diverticulum
of spermathecal appendage of varying shape.

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Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Color. General coloration brownish or reddish; eyes
entirely black; head dark-red or brownish; labium, la-
brum, and mandible blackish; elytra light brown with
dark-brown veins; lower-external, lower-internal, and
medial-internal areas of hind femora blackish; hind tibiae
sometimes brown in basal half and black in apical half.

Female. Antennae reddish brown; pronotum brown
with dark-red tubercles; fore and middle femora, out-
er-medial, upper-external, and upper-internal areas of
hind femora dark-red; fore and middle tibiae dark red;
hind tibiae all brown in basal 2/3 and black in apical
parts; tarsi blackish or dark-red; abdomen brownish; the
posterior margins of the segments marked by red lines.

Male. Antenna light-brown; pronotum brownish with
light-red tubercles in male; fore and middle femora, out-
er-medial, upper-external, and upper-internal areas of
hind femora light-red; fore and middle tibiae light-red;
hind tibiae all light-red in outer area and black in inner
area; tarsi blackish or brownish.

Measurements. Male. Body length 50.88—60.19 mm;
Female. Body length 48.72-63.69 mm. Adults of L. im-
potens exhibit significant size variation in both males and
females. Table 1 provides detailed measurements of vari-
ous body parts for this species.

Geographical distribution (Fig. 8). Loveridgacris im-
potens is a species that is found tn East Africa. The species
is known only from some of the Eastern Arc Mountains
of Tanzania, on Zanzibar, and the Shimba Hills of Kenya.

Ecology. Loveridgacris impotens is a geophilous spe-
cies found in lowland wet forests. The species produces
the toxic foams (see the black arrow on Fig. 1F) by com-
bining haemolymph with air through the spiracles.

Loveridgacris tectiferus Hemp, sp. nov.
https://zoobank.org/4A3C74E4-1002-458F-AC93-7F882A DOF4B5
Figs 3A-H, 4A, B, 5C, D

Holotype. TANZANIA. * male; Udzungwa Mountains,
Mang’ula; in disturbed lowland wet forest at border to
National Park; Sep. 2022; Claudia Hemp leg.; Deposito-
ry. CGH,

Paratypes. TANZANIA. * 2 females; same data as for
holotype. Depository: CCH.

Measurements. (mm) Males (n = 1): Body length:
51.20; Median length of pronotum: 13.60; length of hind
femur: 25.00. Females (n= 2): Body length: 47.70—53.60;
Median length of pronotum: 21.50—21.70; length of hind
femur: 23.90—24.60.

Diagnosis. Loveridgacris tectiferus sp. nov. can be
distinguished from L. impotens by the coloration of the
antennae and hind tibiae. In L. impotens, the antennae are
light or reddish brown, while in L. tectiferus sp. nov., seg-
ments alternate between black and orange (Fig. 4C, D).
Similarly, the hind tibiae are uniformly brown and be-
come darker at their apical parts in L. impotens, but are
black with a median dull orange part in L. tectiferus sp.
nov. The most noticeable difference between the two spe-
cies 1s the shape of the tegmina, which are lobe-like and

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Legend

© Parapetasia femorata
@ Parapetasia rammei

4 Loveridgacris impotens
4 Loveridgacris tectiferus

0 500 1000Km

|

2g9

Figure 8. Geographical distribution of species in the genera Parapetasia and Loveridgacris (P. femorata, unfilled circle; P. rammei,
filled circle; L. impotens, unfilled triangle; L. tectiferus sp. nov., filled triangle).

attached to the abdomen in L. impotens, while tectiform
in L. tectiferus sp. nov. Both species are very similar in
the overall shape of the head, pronotum, legs, and body,
as well as in the tegminal pattern of darker veins on a
light brown ground and their size. The epiphallus of both
species is also very similar, consisting of hooked lophi
and elongate appendices with bulbous end parts. How-
ever, in L. tectiferus sp. nov., the lophi are slender, and
the hooks are slightly longer than those in L. impotens.
Additionally, the appendices of the epiphallus are stouter
in L. impotens, and the bridge of the epiphallus is slightly
longer than that in L. tectiferus sp. nov. (see Fig. SF and
6C for comparison).

Description. Male. General coloration. Overall, col-
or brown with antennae with conspicuously black and
orange colored segments. Eyes and labrum black. Hind
femora ventrally black, hind tibiae black with dull orange
middle part (Fig. 4A). Head and antennae. Antennae
thick, rod-like with basal segments black, and then alter-
nating one or two segments orange and black (Fig. 3A,
C). Antennae slightly shorter than head and pronotum
together. Fastigium of vertex upcurved, slightly concave
in the middle, with acute-angular apex. Frons incurved.
Frontal ridge narrow, constricted between antennae (Fig.
3A). Pronotum and wings. Pronotum in front of first
sulcus with large hump, remaining pronotum strongly ru-
gose with elevated ridges and tubercles. Median carina
irregular, obtuse, lateral carinae absent. Posterior margin
of pronotum with ridge-like tubercles along the edge.
Prosternal process low, subpyramidal. Mesosternal inter-
space wider than long, with deep grooves (Fig. 3E). Teg-
mina tectiform, shortened, slightly exceeding abdominal
segment 3. Hind wings absent. Abdomen. All abdominal
segments with dorsal tubercles. Male supra-anal plate tri-

angular (Fig. 5A), cerci laterally compressed, black, with
blunt apex. Subgenital plate obtusely conical (Fig. 5B).
Internal morphology. Epiphallus typical for Dictyophori-
ni. Lophi of epiphallus strongly sclerotized with well-de-
veloped dorsolaterally directed apical hooks. Appendix
of the epiphallus with angular externolateral sclerotized
processes running parallel to the lophi; apices knob-like
with ventral directed dent. Bridge little sclerotized and
comparatively narrow.

Female. Larger and stouter than the male, with the
same coloration as male; predominantly brown with an-
tennae with black and orange segments, black eyes, and
labrum. Underside of hind femora black, hind tibia black
with median dull orange part (Fig. 4B). Antennae, head,
and pronotum as male. Supra-anal plate triangular with
blunt apex (Fig. 5C). Cerci laterally compressed, black
(Fig. 5C, D). Ovipositor valves black, straight, rounded,
with blunt apices (Fig. 5C, D).

Etymology. From Latin: -tectum = roof, because of the
tectiform-shaped tegmina.

Habitat. A geophilous species of lowland wet forest.

Ecology. In captivity, individuals have the same pref-
erence for monocotyledonous plants as observed for Dic-
tyophorus griseus (Rowell et al. 2015). Even when offered
various other plants, they preferred to feed on Liliaceae
leaves and flowers. Mating took approximately half an
hour, and the male sat on top of the female, bending its ab-
domen under that of the female for copulation (Fig. 5E).
Even when roughly handled, no reflex bleeding was ob-
served, as is common in other Dictyophorini species and
also observed in L. impotens (Fig. 1F).

Nymphs. Unknown but are probably similar to nymphs
of L. impotens (Fig. 1E).

Distribution. Tanzania, Udzungwa Mountains.

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280

Key to the species of Loveridgacris

Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

1(2) Antennae light or reddish brown; hind tibiae uniformly brown, their apical parts darker; tegmina lobe-like and attached

tovthe abaomen(Fis. AC DY 2 secte ch aber cnhtees Setteet Seat dee

Jaded 252 Loveridgacris impotens (Karsch, 1888) (Tanzania, Kenya)

2(1) Antennal segments alternate between black and orange; hind tibiae black with a median dull orange part; tegmina

TECTTORM Gia Et ae Re hates weed Ne Ae eR res coe tease ae

Phylogenetic analysis

In total, 47 DNA barcode sequences belonging to 10 Pyr-
gomorphidae species were analyzed. The locations of
collection of the samples used are presented in Table 4.
For the genera Parapetasia and Loveridgacris, two spe-
cies representing all known species were analyzed. Six
additional species defined as outgroups included Zonoce-
rus elegans (Thunberg, 1815), Phyteumas purpurascens
(Karsch, 1896), Phymateus viridipes (Stal, 1873), Tapro-
nota calliparea calliparea (Schaum, 1853), Dictyophorus
spumans (Thunberg, 1787), and Dictyophorus griseus
(Reiche & Fairmaire, 1849). One phylogenetic tree based
on the concatenated sequence alignments of the two in-
dividual gene datasets (COI = 565 bp, 16S = 376 bp)
was constructed with the BI method (Fig. 9). The con-
catenated sequence alignment included 941 bp. The most
basal clusters of the phylogenetic tree comprised the sub-
tribes Zonocerina (Zonocerus), Phymateina (Phymateus,
Phyteumas), and Taphronotina (Taphronota). The tribes
Taphronotini and Phymateini were well resolved, and
all members of these tribes clustered together; similarly,

Zonocerus elegans (N=5)

@® >0.95
Oo >0.85

063

Rul seh inchs Beat Loveridgacris tectiferus Hemp, sp. nov. (Tanzania)

members of the subtribes Zonocerina and Phymateina
were well resolved. The tribe Dictyophorini represented
a separate clade relative to Taphronotini and Phymateini.
The tribe Dictyophorini was divided into three groups,
representing three different genera. We found strong
support for the monophyly of the genera Loveridgacris,
Parapetasia, and Dictvophorus. The species tree inferred
using the BI approach (Fig. 9) clustered. Loveridgacris
near Parapetasia with high posterior probability support
(score > 0.95). The cluster, including only the members
of Parapetasia, was divided into two groups, and Para-
petasia rammei was the sister to Parapetasia femorata.
Both had relatively large interspecific distances (3.75%).
The two Loveridgacris species (L. impotens and L. tecti-
ferus sp. nov.) showed substantial sequence divergence
from the other genera. The distances between L. impo-
tens and P. femorata (6.70%) and between L. impotens
and P. rammei (7.58%) were relatively large. At the spe-
cies level, L. impotens and L. tectiferus sp. nov. are not
completely resolved, but L. tectiferus sp. nov. is mono-
phyletic with high support; the species show low genetic
distance (0.33%).

Phyteumas purpurascens (N=3)

Phymateus viridipes (N=4)

Taphronota calliparea (N=8)

A Dictyophorus griseus (N=8)

Dictyophorus spumans (N=1)

CM-Fotouni

CM-Fotouni

CM-Fotouni Parapetasia rammei
CM-Fotouni
CM-Iboti
CM-Iboti
CM-Bekob
MT011522

Parapetasia femorata

Loveridgacris tectiferus sp. nov.
TZ-Udzungwa

TZ-Nilo

TZ-Nilo

TZ-Nilo

Loveridgacris impotens
22Z-Jozani

ZZ-Jozani
2Z-Jozani
Z2Z-Jozani

ZZ-Jozani

Figure 9. Bayesian inference (BI) tree built from the concatenated sequence alignment of mtDNA COI/16S gene fragments. The
Bayesian posterior probabilities (PPs) are shown close to the nodes.

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Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Mitochondrial genome organization, structure,
and base composition in Loveridgacris

The mitogenomes of Loveridgacris impotens and Love-
ridgacris tectiferus sp. nov. are 15,592 bp and 15,737 bp
long, respectively (see Suppl. 1). Both are organized in
the typical metazoan mitochondrial gene set consisting
of 37 genes, namely, 13 protein-coding genes, 22 transfer
RNA genes, two ribosomal RNA genes (rmL and rrnS),
and one A+T-rich control region (Table 2). A comparison
of the whole mitogenomes, rrnL genes, rrnS genes, and
A+T-rich regions of L. impotens and L. tectiferus sp. nov.
are shown in Table 4. The nucleotide composition of the
entire mitochondrial genome of both species is A+T-bi-
ased, with contents ranging from 70.0% in L. impotens to
74.1% in L. tectiferus sp. nov. (Table 3). The skew met-
rics of the protein-coding genes within L. impotens and
L. tectiferus sp. nov. showed a positive AT-skew and a neg-
ative GC-skew, indicating that base C was more abundant
than base G in the mitogenomes of both species (Table 3).

The pairwise genetic distances (see Suppl. 2) inferred
from all 13 protein-coding genes showed that the in-
terspecies genetic distance ranged from 0% (ATP8) to
0.8% (ND4), indicating relatively low genetic distances
between L. impotens and L. tectiferus sp. nov., regard-
less of the gene considered. Among the 13 PCGs, ND2,
COI, COII, ATP8, ATP6, CONTI, ND3, and ND6 CYTB
were encoded on the majority strand (J-strand), while
NDI, ND4, ND4L, and ND5 were encoded on the mi-
nority strand (N-strand) (Table 2). Transfer RNA genes
are located on the J strand, except for tRNA-Gln, tR-
NA-Cys, tRNA-Tyr, tRNA-Phe, tRNA-His, tRNA-Pro,
tRNA-Leul, and tRNA-Val, which are located on the N
strand (Table 2). Most of the PCGs (ND2, COI, ATP6,
COI, ND4L, ND6, and CYTB) have ATG as the start
codon; ATP8 has ATC as the start codon; COI has ACT
as the start codon; ND3 and ND5 have ATT as the start
codon, and ND1 has ATA as the start codon. All PCGs use
complete TAA as the stop codon, except COHI, NDS, and
ND4, which use ACT, ATT, and TAG, respectively, as the
stop codon (Table 2).

Discussion

Taxonomy

This study was conducted to investigate the genera Par-
apetasia and Loveridgacris using a combination of mor-
phological, distributional, and molecular data. Parapeta-
sia rammei has been used by Seino and Njoya (2018);
hence, this name is currently considered valid in the OSF
(Cigliano et al. 2023). In this study, Parapetasia rammei
was formally resurrected.

We present herein the description of a new species,
Loveridgacris tectiferus sp. nov., discovered in Tanza-
nia. Despite exhibiting distinguishable morphological
traits, our genetic analysis reveals minimal differences

281

Table 2. Distribution of protein-coding genes (PCGs), transfer
RNA genes, ribosomal RNA genes, and A+T-rich regions in the
mitogenome of Loveridgacris impotens.

Species Loveridgacris impotens
Genes Length Position Start Stop Strand Anticodon
(bp) codon codon
ND2 1,023 197-1219 ATG TAA J
COX1 1545 1415-2959 ACT TAA J
COX2 684 3029-3712 ATG TAA J
ATP8 159 3858-4016 ATC TAA J
ATP6 678 4010-4687 ATG TAA J
COX3 820 4694-5513 ATG ACT J
ND3 354 5548-5901 ATT TAA J
ND5 1717 6311-8027 ATT ATT N
ND4 1335 8112-9446 GTG TAG N
ND4L 294 9440-9733 ATG TAA N
ND6 522 9871-10392. ATG TAA J
CYTB 1143. 10396-11538 ATG TAG J
ND1 945 11625-12569 ATA TAG N
tRNA-Ile 64 1-64 AAT TAA J GAT
tRNA-GIn 69 62-130 N TTG
tRNA-Met 67 130-196 J CAT
tRNA-Trp 71 1224-1294 J TCA
tRNA-Cys 63 1287-1349 N GCA
tRNA-Tyr 69 1354-1422 N GTA
tRNA-Leu2 65 2955-3019 J TAA
tRNA-Asp 64 3711-3774 J GTC
tRNA-Lys 71 3775-3845 J CIT
tRNA-Gly 64 5484-5547 J TCC
tRNA-Ala 66 5902-5967 J TGC
tRNA-Arg 65 5967-6031 J TCG
tRNA-Asn 68 6043-6110 J
tRNA-Ser 70 11537-11606 J TGA
tRNA-Glu 64 6184-6247 J TTC
tRNA-Phe 64 6246-6309 N GAA
tRNA-His 70 8043-8112 N GTG
tRNA-Thr 68 9736-9803 J TGT
tRNA-Pro 65 9804-9868 N TGG
tRNA-Ser2 68 6111-6178 J GCT
tRNA-Leul 65 1273-12637 N TAG
tRNA-Val 68 13939-14006 N TAC
L-rRNA 1252 13897-12646 J
S-rRNA 785 14017-14801 N
A+T-rich 791 14802-15592 J N/A

region

Table 3. Nucleotide composition of the complete mitogenome
for each Loveridgacris species examined.

Species Loveridgacris impotens __Loveridgacris tectiferus
sp. nov.
Accession number OR730795 OR730794
Length (bp) 15592 15737
Whole A 6869 (40.0%) 6947 (44.1%)
mitogenome T 4679 (30.0%) 4709 (29.9%)
G 1539 (9.9%) 1543 (9.8%)
C 2507 (16.1%) 2538 (16.1%)
AGT 11546 (70.0%) 11656 (74.1%)
G+C 4046 (26.0%) 4081 (25.9%)
AT-skew 0.14 0.19
GC-skew -0.24 -0.24

between Loveridgacris tectiferus sp. nov. and L. impo-
tens. This suggests that Loveridgacris tectiferus sp. nov.
is likely a recently evolved species, indicative of its youth
within the taxonomic hierarchy. Indeed, L. impotens and

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282

Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Table 4. Taxon sampling and GenBank accession numbers.

Species

Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus griseus (Reiche & Fairmaire, 1849)
Dictyophorus spumans (Thunberg, 1787)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)
Loveridgacris impotens (Karsch, 1888)

Loveridgacris tectiferus sp. nov.
Loveridgacris tectiferus sp. nov.

Parapetasia femorata Bolivar, 1884
Parapetasia femorata Bolivar, 1884
Parapetasia femorata Bolivar, 1884
Parapetasia femorata Bolivar, 1884
Parapetasia rammei Sjostedt, 1923
Parapetasia rammei Sjostedt, 1923
Parapetasia rammei Sjostedt, 1923
Parapetasia rammei Sjostedt, 1923
Phymateus viridipes (Stal, 1873)
Phymateus viridipes (Stal, 1873)
Phymateus viridipes (Stal, 1873)
Phymateus viridipes (Stal, 1873)
Phyteumas purpurascens (Karsch, 1896)
Phyteumas purpurascens (Karsch, 1896)

Phyteumas purpurascens (Karsch, 1896)

dez.pensoft.net

Country
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon

South A frica

Tanzania

Tanzania

Tanzania

Tanzania

Zanzibar

Zanzibar

Zanzibar

Zanzibar

Zanzibar

Tanzania

Tanzania

NA
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon
Cameroon

Tanzania

Tanzania

Tanzania

Tanzania

Tanzania

Tanzania

Tanzania

Locality
Fotouni
Fotoun1
Fotouni
Fotoun1
Fotouni
Fotouni
Fotouni
Fotouni
Western
Cape
Nilo
Nilo

Nilo

Nilo

Jozani

Jozani

Jozani

Jozani

Jozani

Specimen codes

CMJ65 (non-type
specimen voucher)
CMJ66 (non-type
specimen voucher)
CMJ67 (non-type
specimen voucher)
CMJ70 (non-type
specimen voucher)
CMJ72 (non-type
specimen voucher)
CMJ814 (non-type
specimen voucher)
CMJ816 (non-type
specimen voucher)
CMJ817 (non-type
specimen voucher)
ORTH48 (non-type
specimen voucher)
TZC1351 (non-type
specimen voucher)
TZC1379 (non-type
specimen voucher)
TZC1380 (non-type
specimen voucher)
TZC1381 (non-type
specimen voucher)
ZZC1434 (non-type
specimen voucher)
ZZC1435 (non-type
specimen voucher)
ZZC1436 (non-type
specimen voucher)
ZZC1437 (non-type
specimen voucher)
ZZC1438 (non-type
specimen voucher)

Udzungwa TZC1336 (holotype)
Udzungwa TZC1352 (paratype)

NA

Iboti

Iboti

Bekob

Fotouni

Fotouni

Fotouni

Fotouni

Neguru

Neguru

Neuru

Neguru

Neuru

Wikwescho

Wikwescho

NA

CMJ244 (non-type
specimen vouvher)
CMJ245 (non-type
specimen vouvher)
CMJ598 (non-type
specimen vouvher)
CMJ61 (non-type

specimen vouvher)
CMJ62 (non-type

specimen voucher)
CMJ63 (non-type

specimen voucher)
CMJ64 (non-type

specimen voucher)
TZC1339 (non-type
specimen voucher)
TZC1340 (non-type
specimen voucher)
TZC1354 (non-type
specimen voucher)
TZC1355 (non-type
specimen voucher)
TZC1343 (non-type
specimen voucher)
TZC1388 (non-type
specimen voucher)
TZC1389 (non-type
specimen voucher)

GenBank Accession number

COI
OR583878

OR583879

OR583880

OR583881

OR583882

OR583885

OR583886

OR583887

NA

NA

OR578932

NA

OR578933

NA

NA

NA

NA

NA

OR730794

OR583893

MT011522

OR583883

OR578931

OR583884

OR583875

OR583876

OR578930

OR583877

NA

OR583890

NA

OR583894

OR583892

OR578937

OR578938

16S
PP552786

PP552787

PP552788

PP552789

PP552790

PP552791

PP552792

PP552793

PP552794

PP552822

PP552823

PP552824

NA

PP552980

PP552825

PP552826

PP552827

PP552828

PP552820
PP552821
NA

PP552818

PP552979

PP552819

PP552814

PP552815

PP552816

PP552817

PP552803

PP552804

PP552805

PP552806

PP552807

PP552808

PP552809

GenSeq
Nomenclature
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 16S
genseq-4 16S
genseq-4 COI, 16S
genseq-4 16S
genseq-4 COI
genseq-4 16S
genseq-4 16S
genseq-4 16S
genseq-4 16S
genseq-4 16S
genseq-1 COI, 16S
genseq-2 COI, 16S
genseq-4 COI
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 16S
genseq-4 COI, 16S
genseq-4 16S
genseq-4 COI, 16S
genseq-4 COI, 16S
genseq-4 COI, 16S

genseq-4 COI, 16S

References
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Species Country Locality Specimen codes GenBank Accession number GenSeq References
COI 16S Nomenclature
Taphronota calliparea (Schaum, 1853) Tanzania Kimboza TZC1335 (non-type OR583888 PP552795 genseq-4 COI, 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nguru TZC1341 (non-type NA PP552796 genseq-4 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nguru TZC1342 (non-type OR583891 PP552797 genseq-4 COI, 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nilo TZC1383 (non-type OR5S83895 PP552798 genseq-4 COI, 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nilo TZC1384 (non-type OR583896 PP552799 genseq-4 COI,16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nilo TZC1385 (non-type OR578934 PP552800 genseq-4 COI, 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nguru TZC1386 (non-type OR5S78935 PP552801 genseq-4 COI, 16S This study
specimen voucher)
Taphronota calliparea (Schaum, 1853) Tanzania Nguru TZC1387 (non-type OR5S78936 PP552802 — genseq-4 COI, 16S This study
specimen voucher)
Zonocerus elegans (Thunberg, 1815) Tanzania Nguru TZC1337 (non-type NA PP552810 genseq-4 16S This study
specimen voucher)
Zonocerus elegans (Thunberg, 1815) Tanzania Nguru TZC1338 (non-type OR583889 PP552811 genseq-4 COI, 16S This study
specimen voucher)
Zonocerus elegans (Thunberg, 1815) Tanzania Nilo TZC1390 (non-type NA PP552812 genseq-4 16S This study
specimen voucher)
Zonocerus elegans (Thunberg, 1815) Tanzania Nilo TZC1391 (non-type OR578939 PP552813 genseq-4 COI, 16S This study
specimen voucher)
Zonocerus elegans (Thunberg, 1815) - - MT011544 NA genseq-4 COI, 16S Song et al.
(2020)

L. tectiferus sp. nov. inhabit different mountain habitats
in Tanzania, with L. impotens being widely distributed,
while L. tectiferus is restricted to Udzungwa mountain
so far. This suggests a possibility of sympatric speciation
due to habitat isolation, which may cause disruption of
gene flow. Our findings align with prior investigations
of Orthopteran taxa, particularly those inhabiting the
Eastern Arc Mountains. These studies indicate that while
genera within this region have ancient origins, speciation
at the species level appears to be relatively young. This
pattern is attributed to historical climatic fluctuations,
which have intermittently fragmented and interconnected
habitats, facilitating both isolation and subsequent diver-
sification. Similar mechanisms have been documented in
various Orthopteran groups, including Lentulidae (Hemp
et al. 2020), the coptacrine genus Parepistaurus (Hemp
et al. 2015), the hexacentrine genus Aerotegmina (Grzy-
wacz et al. 2021), and the meconematine genus Amytta
(Hemp et al. 2018). These studies collectively underscore
the dynamic interplay between historical environmental
factors and evolutionary processes, shaping the diversity
of Orthopteran fauna in the Eastern Arc Mountains and
beyond. Two species are now included in both Para-
petasia (P. femorata and P. rammei) and Loveridgacris
(L. impotens and Loveridgacris tectiferus sp. nov.).
Akbar and Kevan (1964) used external morphology
and phallic structures to distinguish between the genera
Parapetasia and Loveridgacris. Although they did not
examine the phallic structures of P. rammei, they con-
cluded that Parapetasia has an epiphallus with subparal-
lel lateral plates and appendices, a triangular fastigium of
the vertex, and other anatomical features similar to those
of P. femorata. However, the phallic structures of P. ram-
mei, as illustrated in this study, reveal that the epiphallus

has a narrow bridge but divergent lateral plates, resem-
bling those of Loveridgacris rather than Parapetasia.

Akbar and Kevan (1964) noted that Parapetasia spe-
cies, specifically P. femorata, have epiphallus appendices
with smaller and broader terminal processes. However,
P. rammei has appendices with broader terminal process-
es and lacks smaller terminal processes. The bridge of the
epiphallus is also longer in P. rammei than in P. femo-
rata. Additionally, our findings indicate that the lophi in
P. rammei are smaller than those in Loveridgacris, con-
tradicting Akbar and Kevan’s claim that Loveridgacris
has small lophi.

According to Kevan et al. (1974), P. calabarica and
P. rammei are likely variations of P. femorata with small-
er wings. Later, Kevan (1977) combined all three species
into P. femorata based on their shared geographic range
and morphological similarities. However, DNA evidence
and differences in external morphology and phallic struc-
tures show that P. femorata and P. rammei are distinct
species. Therefore, P. rammei should be recognized as a
separate species within the genus Parapetasia.

The present study highlights the importance of com-
bining multiple sources of information and DNA markers
for the identification of Afrotropical grasshopper species
of the genera Parapetasia and Loveridgacris.

Distribution

The type specimen of P. rammei was found in Bare-
Dschang, a high-altitude location in western Cameroon.
Two specimens at MfN labeled holotype (male) and al-
lotype (female) were actually paratypes and collected
from Bamenda and Bangwe in the northwestern region

dez.pensoft.net

284

of Cameroon. The true holotype, a female specimen,
is housed in the Stockholm Museum. Seino and Njoya
(2018) also collected P. rammei from the northwestern
region of Cameroon, which is known for its high altitude.
We found P. rammei in Fotouni in the western highlands
and in the steppe habitats of the Manengouba Mountains.
Our observations are supported by Rehn (1953), who
suggested that P. rammezi is limited to the highest areas of
Cameroon. Our research, together with museum data, in-
dicates that P._ femorata inhabits forested areas in Western
and Central African countries such as Cameroon, Equa-
torial Guinea, Gabon, and Nigeria. Although the species
is widespread in Cameroon, its distribution in neighbor-
ing countries may be underestimated due to a lack of
sampling. Hence, our results show that P. femorata and
P. rammei occupy different ecological niches.
Loveridgacris impotens and L. tectiferus sp. nov. are
found only in Tanzania and Kenya. Therefore, the genus
Parapetasia is restricted to western and central Africa, while
Loveridgacris is exclusive to East Africa. The new species
L. tectiferus has to be considered a narrow range endemic,
whereas L. impotens is widespread in eastern Africa.

Phylogeny

Although Parapetasia and Loveridgacris have been the
subject of taxonomic discussions since I. Bolivar (1884),
their taxonomy and systematic status have remained com-
plex and challenging; thus, we provide the first attempt at
a molecular phylogeny for the tribe Dictyophorin1.

Our results divided the studied Pyrgomorphidae into
three main tribes (Dictyophorini, Phymateini, and Taph-
ronotini). Taphronotini, which includes a single species,
Taphronota calliparea, is closely related to Dictyophorini,
which includes three genera (Dictyophorus, Loveridgacris,
and Parapetasia). In addition, the genera Zonocerus, Phy-
teumas, and Phymateus are closely related, indicating their
inclusion in the tribe Phymateini. The close relationships
among Pyrgomorphid tribes and genera have previously
been documented by several authors. For instance, Kevan
et al. (1974) suggested a close relationship between Taph-
ronotini and Dictyophorini according to copulatory struc-
tures. Similarly, Marifio-Perez and Song (2018) reported a
close relationship between Taphronotini and Dictyophori-
ni, as well as among Zonocerus, including Zonocerus var-
iegatus, and Phymateus, including Phymateus saxosus.

When comparing our tree with the molecular phy-
logeny of Zahid et al. (2021), we found concordance
regarding the monophyly of Dictyophorini. Similarly,
we found concordance regarding the monophyly of Phy-
mateini when comparing our phylogenetic tree with the
morphological phylogenetic tree by Marifio-Pérez and
Song (2018). In addition, the monophyly of Phymateini
recovered in our tree was not supported by the molec-
ular phylogeny of Zahid et al. (2021), as this tribe was
found to be paraphyletic. In addition, Marifio-Pérez and
Song (2019) reported that Monistria Stal, Dictyophorus,

dez.pensoft.net

Jeanne Agrippine Yetchom Fondjo et al: Integrative taxonomy of two Dictyophorini genera

Phymateus, and Poekilocerus Serville were scattered
throughout the phylogeny, forming paraphyletic groups.
Such discordances in the topologies are not surprising,
as previous studies, e.g., Baker et al. (1998), Friedrich
et al. (2014), Kjer et al. (2016), and Peters et al. (2014),
have demonstrated. However, as our study was limited in
terms of taxon sampling to clarify the taxonomic position
of Parapetasia and Loveridgacris species, we did not in-
clude many species from other tribes. Hence, we are not
able to address these problems with our dataset.

We obtained robust evidence supporting the distinct
status of the genus Parapetasia. Specifically, Parapetasia
rammei has been confirmed as the sister taxon to Para-
petasia femorata, and this distinction is further reinforced
by morphological and phallic structure diagnostic traits.
Furthermore, our phylogenetic tree revealed substantial
genetic differentiation between the genera Loveridgac-
ris and Parapetasia. Consequently, Kevan’s (1977) pro-
posal to synonymize P. rammei and L. impotens with P.
femorata, primarily based on their shared geographic
distribution and some morphological resemblances, is
now questioned. This study marks the first integrative
examination of the phylogeny of Dictyophorini, which
revealed a consistently structured topology. Nonetheless,
to fully resolve the phylogenetic relationships within Dic-
tyophorini, it is imperative to include additional genera
and their respective species in future investigations, as
well as additional genetic markers.

The findings from the maximum likelihood (ML) and
Bayesian inference (BI) analyses and the examination of the
complete mitogenome indicate that it is challenging to estab-
lish a phylogenetic relationship between L. impotens and the
newly described species L. tectiferus sp. nov. using the COI
gene alone. Even the complete mitogenome revealed only
minor divergence in this regard. Generally, it is likely that
L. tectiferus sp. nov. represents a relatively young species
in the early stages of speciation. To gain more conclusive
insights into its taxonomic status and evolutionary trajectory,
broader sampling of specimens 1s needed.

Conclusion

This study offers a reassessment of the Pyrgomorphid
grasshopper genera Parapetasia and Loveridgacris, along
with the first-ever analysis of the mitochondrial genome
in the genus Loveridgacris. Our research unequivocally
demonstrated that the genus Parapetasia comprises two
distinct species, P. femorata and P. rammei, characterized
by both pronounced morphological and genetic dispar-
ities. Additionally, we underscore the substantial differ-
ences, both morphologically and genetically, between the
genera Parapetasia and Loveridgacris. Consequently,
the previous doubts regarding their taxonomic position
are dispelled. Therefore, we confirm that Parapetasia
and Loveridgacris are unequivocally recognized as two
distinct genera. Finally, we describe a new Loveridgacris
species adding to the diversity of the group.

Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Acknowledgments

The authors are grateful to the Alexander von Humboldt
Foundation for its financial support. The authors thank
Cameroon’s Ministry of Scientific Research and Innova-
tion for granting the research permit for field collection (N
0000010/MINRESI/B00/C00/C10/C13). We also thank
the Commission for Science and Technology, Tanzania
and the Tanzania Wildlife Research Institute, Tanzania,
for granting research permits. We thank the Mohamed bin
Zayed Species Conservation Fund for their financial sup-
port. We would like to thank Dr. Ricardo Marifio-Pérez at
the University of Michigan, Prof. Hojun Song at Texas A
& M University, United States, and Mrs. and Katrin Elgn-
er at the Senckenberg museum in Germany for providing
us with the necessary literature. We are thankful to Mrs.
Birgit Jaenicke at the Museum fir Naturkunde Berlin
for granting us permission to examine several samples.
We thank Mr. Aristide Junior Sock Bell and Mr. Sedrick
Junior Tsekane for their valuable assistance during field
investigations in the Ebo Forest, Cameroon. The authors
are grateful to Ms. Eileen Nguyen for her valuable sup-
port when taking the photographs. We would like to thank
Mr. Carsten Bruns and Ms. Lara-Sophie Dey for their as-
sistance in the laboratory. We also thank Mr. Jithin John-
son for the introduction to the mapping. Furthermore,
we are grateful to the Academy of Natural Sciences of
Philadelphia for providing photographs of the types P.
femorata and P. calabarica. The authors are also grate-
ful to the American Journal Experts (AJE) for editing the
manuscript for proper English language.

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Dtsch. Entomol. Z. 71 (2) 2024, 265-287

Supplementary material |

Complete mitochondrial genomes of
Loveridgacris impotens and Loveridgacris
tectiferus sp. nov.

Authors: Jeanne Agrippine Yetchom Fondjo, Martin
Husemann, Armand Richard Nzoko Fiemapong, Alain
Didier Missoup, Martin Kenne, Maurice Tindo, Oliver
Hawlitschek, Tarekegn Fite Duressa, Sheng-Quan Xu,
Wenhui Zhu, Claudia Hemp

Data type: tif

Explanation note: In this supplementary material, the
mitogenomes of Loveridgacris impotens and Love-
ridgacris tectiferus sp. nov. are presented. The leng-
ht of the genomes, their organization and nucleotide
composition are also shown on the file.

Copyright notice: This dataset is made available under
the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided
that the original source and author(s) are credited.

Link: https://do1.org/10.3897/dez.71.125877.suppl1

Supplementary material 2

Genetic distance of the 13 protein-coding
genes between Loveridgacris impotens and
Loveridgacris tectiferus sp. nov.

Authors: Jeanne Agrippine Yetchom Fondjo, Martin
Husemann, Armand Richard Nzoko Fiemapong, Alain
Didier Missoup, Martin Kenne, Maurice Tindo, Oliver
Hawlitschek, Tarekegn Fite Duressa, Sheng-Quan Xu,
Wenhui Zhu, Claudia Hemp

Data type: tif

Explanation note: This supplemtary material shows the
pairwise genetic distances inferred from all 13 pro-
tein-coding genes between Loveridgacris impotens
and Loveridgacris tectiferus sp. nov.

Copyright notice: This dataset is made available under
the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided
that the original source and author(s) are credited.

Link: https://do1.org/10.3897/dez.71.125877.suppl2

287

Supplementary material 3

Localities and coordinates of Parapetasia
and Loveridgacris

Authors: Jeanne Agrippine Yetchom Fondjo, Martin
Husemann, Armand Richard Nzoko Fiemapong, Alain
Didier Missoup, Martin Kenne, Maurice Tindo, Oliver
Hawilitschek, Tarekegn Fite Duressa, Sheng-Quan Xu,
Wenhui Zhu, Claudia Hemp

Data type: docx

Copyright notice: This dataset is made available under
the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided
that the original source and author(s) are credited.

Link: https://do1.org/10.3897/dez.71.125877.suppl3

dez.pensoft.net