Research Article

Journal of Orthoptera Research 2021, 3001): 17-26

Life history of the false flower mantid (Harpagomaniis tricolor Linnaeus, 1758)
(Mantodea: Galinthiadidae) and its distribution in southern Africa

BIANCA GREYVENSTEIN!, HANNALENE DU PLEssis!, JOHNNIE VAN DEN Bere!

1 Unit for Environmental Sciences and Management, North-West University, Potchefstroom, 2520, South Africa.

Corresponding author: Bianca Greyvenstein (biagrey90@gmail.com)

Academic editor: Matan Shelomi | Received 1 April 2020 | Accepted 7 July 2020 | Published 18 February 2021

http://zoobank.org/C30EF79A-5544-450B-B439-8A4BBIDDIE2C

Citation: Greyvenstein B, du Plessis H, Van den Berg J (2021) Life history of the false flower mantid (Harpagomantis tricolor Linnaeus, 1758) (Mantodea:
Galinthiadidae) and its distribution in southern Africa. Journal of Orthoptera Research 30(1): 17-26. https://doi.org/10.3897/jor.30.52816

Abstract

The false flower mantid is the common name for the Mantodea species
Harpagomantis tricolor (Linnaeus, 1758). This species uses camouflage as a
defense mechanism. Limited information (Kaltenbach 1996, 1998) exists
on its distribution in southern Africa or about its life history. This species,
and Mantodea to an extent, are not usually included in biodiversity studies
from this region. The aim of this study was to determine the distribution
of this species in southern Africa based on museum collection records and
to study the biology of Harpagomanitis tricolor under captive breeding condi-
tions. The distribution of Harpagomantis and its morphological variety, i.e.,
discolor, were determined utilising the historical insect collection records of
seven national museums throughout South Africa. Field collected H. tri-
color males and females were mated and reared under laboratory conditions
to record their life history parameters of nymphal duration, oothecae struc-
ture, size and incubation duration, adult longevity, and sex ratio. The re-
sults of this study indicate that the mean duration of the lifecycle of H. tri-
color is 191.33 + 37.96 days. All but three H. tricolor individuals had five
nymphal instars, and the mean duration of the nymphal stage was 140.20
+ 31.03 days. The mean duration of copulation was six hours, while the av-
erage incubation period of oothecae was 144.71 + 9.33 days. These results
indicate that oothecae of H. tricolor probably overwinter under field condi-
tions and that males of this species have evolved various mechanisms to
increase the likelihood of ensuring their own genetic offspring. This study
bridges the gap in rudimental research in which Mantodea, in general, has
been overlooked and establishes a basis on which ecological interactions,
habitat preferences, and imminent threats to H. tricolor can be established.

Keywords

camouflage, copulation, longevity, praying mantis

Introduction

Harpagomantis Kirby is one of four genera in the newly rear-
ranged family of Galinthiadidae (Roy and Stiewe 2014, Svenson et
al. 2015, Schwarz and Roy 2019). Within the Harpagomantis genus,
there were two known species: Harpagomantis tricolor (Linnaeus,
1758) and Harpagomantis discolor (Stal, 1877). However, H. dis-
color was found to be a morphological variety of H. tricolor, and
thus the only species within this genus is H. tricolor. Harpagomantis,

Galinthias, Congoharpax, and Pseudoharpax were previously classi-
fied as Hymenopodidae. However, due to molecular evidence and
the phylogenetic results reported by Svenson and Whiting (2009)
and Svenson et al. (2015), these genera were found to be outside
of Hymenopodidae and were moved to the new family Galinthia-
didae (Roy and Stiewe 2014). Svenson et al. (2015) reported that
the high level of homoplasy in the external morphology of these
mantids contributed to the discrepancies in species identification
based on molecular and morphological characteristics, as these
did not align. Thus, these genera were originally classified within
the Hymenopodidae family. However, morphological characteris-
tics were also used to aid the rearrangement of these families in an
article in which the Mantodea order and its families were revised
and rearranged (Schwarz and Roy 2019).

Harpagomantis have been described as “false flower” mantids
and are pink with green bands and sometimes yellow eyes (Fig. 1).
Harpagomantis are reported to live on flowers where they camou-
flage and wait motionlessly for prey (O’Toole 2003). Camouflage
in mantid species has been reported as the primary defense mech-
anism of these insects (Edmunds 1971), and H. tricolor is no ex-
ception. It has been recorded during biodiversity survey studies in
South Africa, largely in the western Cape region (Grobbelaar et al.
1999, Brand and Samways 2009, Magoba and Samways 2010) and
the Highveld grassland biome (Botha et al. 2018, Greyvenstein et
al. 2020b). However, the distribution of this species is based solely
on studies by Kaltenbach (1996, 1998). The latter study found
this genus widespread in southern Africa and included eight of the
nine South African provinces and records from Botswana, Leso-
tho, Namibia, Zambia, and Zimbabwe.

Cardoso et al. (2020) recently reported being concerned about
the worldwide decline in insect populations and that only 20% of
the total insect diversity has been named. Research is required to
bridge this gap in knowledge and correct the bias in insect studies
that have largely focused on specific taxa such as butterflies and
pollinators (Cardoso et al. 2020). Samways et al. (2020) indicated
that mapping the distribution of specific species could contribute
to determining their range expansion, threat identification, and
habitat favorability. This will aid in bridging the gap in knowledge
regarding the distribution, biology, and ecology of the majority of

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

18

insect species. Similarly, Svenson et al. (2015) reported that the
ecology of most Mantodea species remains unknown. Although
various studies have been done on Mantodea ecology, very few
have been conducted in South Africa and with species of this re-
gion. The information available about species’ ecology, observa-
tions, and biology in South Africa is based on either citizen science
or very old publications that could be outdated.

The aim of this study was to determine the distribution of the
genus Harpagomantis in southern Africa and to study the biology
of H. tricolor under captive breeding conditions.

Materials and methods

Species distribution database.—Distribution records of Harpagoman-
tis spp. were collected during visits to the following institutions
that host curated insect collections in South Africa: Ditsong Mu-
seum of Natural History (Pretoria), Agricultural Research Council
(Biosystematics Division, Pretoria), National Museum (Bloem-
fontein), Albany Museum (Grahamstown), Rhodes University
(Grahamstown), Durban Natural Science Museum, Iziko South
African Museum (Cape Town), and KwaZulu-Natal Museum (Pi-
etermaritzburg). Most specimens in these collections were previ-
ously identified by foreign visiting taxonomists, and many were
sent for identification to the Vienna Museum in Germany, the
University of Drexel in Philadelphia, USA, the Muséum national
d'Histoire naturelle (MNHN) in Paris, France, and the research
collection of Nicolas Moulin in Montérolier, France.
Harpagomantis specimens and distribution labels were photo-
graphed (Canon D1300) and digitized, after which this data was
used to compile a distribution database of the species. The database
contains the following information for each specimen record: ge-
nus and species name (to the available level of identification), col-
lector’s details and collection date where available, and the geo-ref-
erenced locality. Scientific literature (Ehrmann 2002, Svenson 2015,
Schwarz and Roy 2019) was used to determine the current nomen-
clature within the genus. All locality data were georeferenced using
the principles suggested by Wieczorek et al. (2004). Subsequently,
all coordinates were converted from degrees, minutes, and seconds
(DMS) to decimal degrees (DD) (gps-coordinates.net). Decimal de-
grees were used for developing distribution maps for H. tricolor, the
H. discolor variety, and the specimens that have not been morpho-
logically distinguished in southern Africa by means of GIS software
(ArcMaps, Version 10.6.1). The collection dates recorded for each
specimen were used to generate intervals of 11 years (i.e., 1856-
1867 and 1868-1879) to compile a graph indicating the number of
specimens collected over time and during certain intervals.

Rearing and biology.—Specimens were collected in the Grassland
biome in the North West and Free State provinces of South Africa
during the summer of 2016/2017. Adults of these field-collected
individuals were mated, and nymphs that emerged from oothecae
were used to rear a sufficient number of individuals to observe un-
der captive breeding and rearing conditions. A sub-sample of the
field-collected specimens was identified by Nicolas Moulin (hon-
orary associate to MNHN) to confirm the species identification.
For breeding purposes, pairs of males and females were placed
in glass containers. Glass containers (40 cm x 20 cm x 20 cm)
were used to ensure that ample space was available for the male to
avoid sexual cannibalism before, during, or after mating. To fur-
ther limit the likelihood that females would cannibalize the males,
ample food was provided before the male was introduced into
the breeding container. The duration of copulation was recorded

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

per breeding pair (Fig. 1c). After copulation concluded, the male
was removed from the breeding container. The terrariums (15 cm
x 10 cm x 20 cm) in which females were kept after mating were
checked daily for the presence of oothecae that were laid overnight.
Oothecae were removed and put into small containers (5 cm di-
ameter and 5 cm high) inside a desiccator. A humidity level of 68 +
5% was maintained in the closed desiccator, following the method
described by Solomon (1951). The desiccator was kept in an insect
rearing room at a temperature of 27 + 1°C with a 14L:10D photo-
period cycle until nymphs emerged from the oothecae.

Rearing of nymphs was done under controlled conditions
(Fig. 1d). Each specimen was placed into a terrarium (7 cm di-
ameter and 15 cm high) with three holes (each 2 cm in diameter)
covered with gauze to allow air flow. Thin twigs (5 mm x 10 cm)
were placed inside each jar for climbing and hanging purposes, es-
pecially during moults. Food was provided every second day at the
same time as fine water mist was sprayed into each container. Live
aphids (Brevicoryne spp.) (Hemiptera: Aphididae) were provided
as food for first- to third-instar nymphs, after which live crickets
(Acheta sp., Orthoptera: Gryllidae) of different sizes (nymphal in-
stars, i.e., pinheads) were provided. After moulting to the second-
instar, nymphs were removed from the communal terrariums and
placed in separate terrariums to prevent cannibalism. Nymphs
were reared until adulthood, after which males and females were
identified. This was done by counting the number of abdominal
segments and the appearance of the wings. Harpagomanitis tricolor
females have shorter wings (barely covering the abdomen) and six
abdominal segments, while males have eight segments and wings
that are longer than the abdomen (McMonigle 2013, Fatimah et
al. 2016, Brannoch et al. 2017) (Fig. 1a, b).

The following life history parameters were recorded during this
study: size of oothecae, number of egg chambers inside fertilized
and unfertilized oothecae, copulation duration, number of days
between moults, and survival rate (based on nymphs reaching the
adult phase). The mean number of days between moults and days
to adulthood were calculated separately for males and females. The
data discussed in this paper were recorded for 45 individuals (14
males and 31 females) that completed their life cycles. The mean du-
ration of male and female life cycles was calculated, and the hatch
and survival rates were determined. A distinction was also made
between different types of oothecae, i.e., fertilized and unhatched
(produced by field-collected females of which the mating status
was not known). The length, width, and height of each ootheca
were recorded based on descriptions by Brannoch et al. (2017). The
ootheca length was measured along the area of emergence, exclud-
ing the residual process (Brannoch et al. 2017, Greyvenstein et al.
2020a). To determine the number of eggs per ootheca, oothecae
were dorsally dissected along the length and inspected under a
microscope, as was done by Greyvenstein et al. (2020a). Measure-
ments of ootheca parameters were done as indicated in Fig. 1e.

Data analysis.—The descriptive statistics (means and standard error)
and the statistical analyses of the developmental parameters were
done using Statistica Version 13.3 (TIBCO Software Inc., 2017). Sha-
piro-Wilk normality test was used to determine if the data were nor-
mally distributed, and data that were not normally distributed were
log-transformed. T-tests were used to determine if differences exist-
ed between the length, width, height, and the number of eggs per
ootheca between the two types of oothecae (i.e., fertilized and un-
fertilized). T-tests were also used to determine if differences existed
between the mean numbers of days between moults, adult longevity,
and mean number of days required by nymphs to reach adulthood.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

Substrate

Fig. 1. Harpagomantis tricolor female. a. Male; b. Copulating adults; c. Fifth-instar nymph; d. General morphology of the oothecae; e. In-
dicating different parameters and areas of interest as suggested by Brannoch et al. (2017). Photographs by Paul Janse van Rensburg.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

20

Results

Distribution.—The distribution records reported in this paper
were compiled from records available in the seven South Afri-
can institutions that host curated arthropod collections. Results
should be viewed in this context, since no museum records be-
yond those residing in South Africa were included. The results
of this study and the following previously published studies
(Kaltenbach 1996, 1998, Grobbelaar et al. 1999, Brand and
Samways 2009, Magoba and Samways 2010, Botha et al. 2018,
Greyvenstein et al. 2020b), to our knowledge, are the only studies
to include distribution records of this species. The distribution
records included records of Harpagomantis tricolor as well as the
discolor morphological species variety from the following south-
ern African counties: Botswana, Eswatini, Lesotho, Mozambique,
Namibia, and Zimbabwe (Fig. 2).

A total of 290 specimen records were accounted for, of which
272 were collected within the borders of South Africa (this in-
cludes specimens collected in Lesotho and Eswatini). The remain-
ing 18 records of H. tricolor were distributed as follows: two speci-
mens collected in Botswana, six in Namibia, four in Mozambique,
and six in Zimbabwe (Fig. 2). H. tricolor records were collected
throughout South Africa and neighboring countries (Fig. 2). The
distribution of H. tricolor in South Africa seems to be predomi-
nantly towards the eastern region of the country, with a few speci-
men records from the western region, specifically in the Western
Cape Province (Fig. 2). The oldest H. tricolor specimen record
was collected in 1876 in Cape Town. Only four specimens were
collected between 1876 and 1887, while the largest number (37)
were collected between 1912 and 1923 (Fig. 3). Between 1972 and
2019, the average number of specimens collected during the three
11-year intervals was 31 (Fig. 3). Only 48 specimen records (18%)
were collected within protected areas of South Africa, while 224

Harpagomantis morphs in southern Africa

x Harpagomantis tricolor morph (74)

+ Harpagomantis discolor morph (43)

Namibia Botsw:
x

OF
South Africa + x Y
i

ee KX x +
100 200km x

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

records (82%) were collected outside these areas. These 48 speci-
mens were collected in 11 different provincial nature reserves (19
records), four private nature reserves (14 records), two national
parks (9 records), and one specimen was collected in each of a
world heritage site, a protected forest area, and a local nature re-
serve (Fig. 4).

Biology.—The ootheca of H. tricolor is not covered in the usual
foamy sheath that is characteristic of a variety of Mantodea, al-
though exceptions do exist (McMonigle 2013). The oothecae are
usually small, light brown in color, almost rectangular in shape,
and slightly dorsally flattened (Fig. le). The residual process is
not elongated or extended into any shape or point. In cases where
the oothecae of H. tricolor were attached to the stem of a flower-
ing plant in the field, they most likely resembled a thorn. Eggs
were arranged in adjacent rows of between three and five eggs each
(Fig. le). The residual process was also investigated but did not
contain any egg chambers.

In this study, 19 oothecae were produced by field-collected fe-
males mated under captive breeding conditions. Seven of these
did not hatch and 12 oothecae did hatch. In total, 63 nymphs
emerged from the 12 fertile oothecae under captive breeding
conditions. No significant differences were recorded between
the length, width, height, or number of eggs of the fertilized or
unhatched oothecae. Mean ootheca length was 8.5 + 4.11 mm,
containing 17.26 egg + 6.66 chambers per ootheca (Table 1). The
mean width and height of an ootheca was 4.37 + 0.76 mm and
6.15 + 0.83 mm, respectively (Table 1).

Developmental parameters—Of the 63 neonate nymphs that
hatched from the 12 different oothecae throughout this study, 45
completed their lifecycles (14 males and 31 females). The mean
duration between mating and the production of an ootheca was

Mozambique
Zambia

Zimbabwe

ao

ana

t
x
Lesotho

x ‘
x

™ - - =| +}
Esri, USGS | Esri, HERE, Garmin, FAO, NOAA USGS (==

Fig. 2. Distribution records of Harpagomanitis tricolor and its morph (discolor) that occur in southern Africa. Numbers in brackets indi-

cate the number of individual records per variety.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG 21

11.82 + 9.51 days, and the act of copulation itself continued for
approximately six hours (Table 3). The incubation period of an
ootheca was approximately 20 weeks (143 days). The mean hatch
rate was 31%, while the average survival rate was almost 68%
(Table 2). The sex ratio differed between the various oothecae, but
the mean sex ratio (M:F) was 1:1.5. Two of the oothecae produced
only males, while another two produced only females (Table 2).

Number of specimens collected

No significant differences were recorded between the average
duration per instar of females and males. The nymphal period
took approximately 20 weeks to complete (Table 3). However,
females required a longer nymphal period (145.71 + 29.88 days)
than males (128.00 + 31.09 days), even though this difference was
not significant. The mean duration of the lifecycle of H. tricolor
individuals in this study was six months (191.33 + 37.96 days).

37
36
33
29 29
27 27
26
14
13
4
3
a a

No Data 1876-1887 1888-1899 1900-1911 1912-1923 1924-1935

1936-1947

1948-1959 1960-1971 1972-1983 1984-1995 1996-2007 2008-2019

Collection dates

e Harpagomantis spp, outside of Protected areas (224)

@ = Harpagomantis spp. in Protected areas (48)
Protected areas- Catagories

esl Forest Act Protected Area

aa Local Nature Reserve

National Park
i; Private Nature Reserve

= Provincial Nature Reserve

eg World Heritage Site

Sources: Esri, Airbus DS, USGS, NGA, NASA, CGIAR, N Robinson,
NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA,

Fig. 4. Distribution records of Harpagomantis species collected in protected and non-protected areas of South Africa.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

22 B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

Table 1. Mean size and number of egg chambers inside the various types of oothecae of Harpagomantis tricolor reared under captive

breeding conditions. SD = standard deviation.

Oothecae (19) Length (mm) + SD

Width (mm) + SD

Height (mm) + SD Number of eggs/ootheca + SD

T-test t-value 0.573 0.986 0.058 0.267
p-value 0.574 0.338 0.954 0.792
Overall (19) 8.58 + 4.11 4.37 + 0.76 6.15 + 0.83 17.26 + 6.66
Unhatched (7) 7.86 + 2.24 4.14 + 0.69 6.14+ 1.21 16.71 + 7.20
Fertilized (12) 9.00 + 4.63 4.50 + 0.80 6.17 + 0.58 17.58 + 6.63

Table 2. Mean duration (in days) of each of the life stages of Harpagomantis tricolor and differences between male and female develop-
ment under captive breeding and rearing conditions. Three of the females developed to the sixth-instar and were not included in the

table below.
Life stage Mean duration (days + SD) t-value p-value
Overall Males Females

Ootheca (incubation period) 144.71 + 9.33 142.51 + 10.90 145.68 + 8.55 1.057 0.297
First instar 26.62 + 11.07 25.36 +9.91 27.19 11.674 0.373 0.710
Second instar 24.67 + 15.63 28.57 + 17.51 22.90 + 14.66 -1.110 0.273
Third instar 27.67 + 13.06 33.38 + 18.5 25.42 + 9.37 -1.784 0.082
Fourth instar 41.55 + 22.91 49.00 + 29.63 39.24 + 20.47 -0.838 0.407
Fifth instar 51.50 + 13.28 54.00 + 9.00 51.06 + 14.06 -0.499 0.624
Copulation to oothecae (days) * 11.82 + 9.51 12.27 + 8.67 11.63 + 9.99 -0.135 0.894
Copulation duration (hours) ** 06:10 + 0.04 06:15 + 0.04 06:08 + 0.034 -0321 0.750
Total nymphal period (days) *** 140.20 + 31.03 128.00 + 31.09 145.71 + 29.88 1.776 0.082
Adult longevity (days) * *** 51.11 + 39.76 31.57 + 29.72 59.93 + 40.97 -0.509 0.613
Period from hatch to death (days) 191.33 + 37.96 161.71 + 20.47 204.71 + 36.58 -0.509 0.613

* duration of period between male and female copulation and production of ootheca; ** duration of male and female copulation; *** from ootheca hatch to final moult

(first-instar to fourth/fifth-instar); **** duration of adult phase.

Table 3. The mean hatch rate, survival rate, and gender dynamics that resulted from each of the field-collected H. tricolor females (12
individuals) that were kept in the laboratory and each of their associated fertile oothecae (12).

Ootheca number Oothecae No. of eggs per Hatch rate (%)
incubation (days) ootheca

Ootheca 1 123 16 81.25
Ootheca 2 149 14 28.57
Ootheca 3 145 19 31.58
Ootheca 4 155 18 33.33
Ootheca 5 127 16 25.00
Ootheca 6 138 18 16.67
Ootheca 7 145 15 20
Ootheca 8 156 9 44.44
Ootheca 9 153 36 13.89
Ootheca 10 147 12 25.00
Ootheca 11 143 21 14.29
Ootheca 12 139 17 35.29
Mean + (SD) 143.33 + 10.31 17.58 + 6.63 30.78 + 18.35
Discussion

Distribution patterns of Harpagomantis tricolor in southern Afri-
ca.—Although there have been some discrepancies in the past
regarding the species within this genus, it is assumed that two
morphological varieties exist: H. tricolor and H. discolor. Speci-
men records throughout the museum collections in South Af-
rica exist for both morphological varieties (morphs). According
to Stal (1877) and Giglio-Tos (1927), H. discolor males do not
have a brown spot on the hindwings, and the species is gener-
ally larger than H. tricolor. Rehn (1927) reported that H. tricolor
was a much smaller species with limited distribution (mostly in
the Western Cape region of South Africa), while H. discolor oc-
curs throughout South Africa but predominantly in the northern
region. The latter species is also believed to be larger and have

Survival (%) Male (%) Female (%) Sex Ratio (¢:9)
69.23 33.33 66.67 1:2
100 0.00 100.00 0:4
100 83.33 16.67 1:0.2
50 33:33 66.67 12
100 0.00 100.00 0:4
100 33.33 66.67 1:2
33.33 100.00 100.00 1:1
50 100.00 0.00 2:0
60 33.33 66.67 12
66.67 0.00 100.00 0:2
66.67 50.00 50.00 1:1
16.67 60.00 40.00 1:0.67
67.71 + 28. 04 43.89 + 36.15 64.44 + 33.46 speshet5

elongated processes on the eyes (non-visual elongations that do
not contain ommatidia).

However, Karny (1908) indicated that H. discolor could be a
variety of H. tricolor. This view was shared by Beier (1955), who
stated that H. discolor was a “pigment-poor” variety of H. tricolor.
A similar conclusion was drawn by Kaltenbach (1996), and this
species was therefore considered an intra-species variety (in size
and color) of H. tricolor and, according to Kaltenbach (1996),
H. discolor is a synonym of H. tricolor. Ehrmann (2002) agreed
with Beier (1953, 1955) and Kaltenbach (1996) and noted that
H. discolor was a synonym for H. tricolor.

Rehn (1927) indicated the possibility of a clinal North-South
differentiation between the two Harpagomantis morphs. Similarly,
Kaltenbach (1994) noted a clinal variation of the subspecies of
Bisanthe Stal, 1876, also in southern Africa. Kaltenbach (1994) de-

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

scribed B. menyharthi menyharthi (Brancsik, 1895) to have a north-
ern distribution (towards Zambia), while B. menyharthi raggei was
recorded largely in the south towards Zimbabwe and Botswana
(South of the Zambezi river). Although Kaltenbach (1994) indi-
cated that temperature could have played a role in the clinal dif-
ferentiation of the subspecies of Bisanthe in southern Africa, other
factors could also be responsible for such observed differentia-
tions. For example, Lombardo (1995) noted that the Great Rift
Valley tectonic plate could be the dividing line between the clinal
differentiations noted in the subspecies of Popa spurca Stal, 1856
(Popa spurca spurca Stal, 1856 and Popa spurca crassa Giglio-Tos,
1917) in Africa. The Great Rift Valley has been shown to be an im-
portant natural barrier for various invertebrate species, especially
Busseola fusca (Fuller) (Lepidoptera: Noctuidae) and Sesamia cre-
tica Lederer (Lepidoptera: Noctuidae), and is suspected to be the
cause of their genetic differentiation into different clades towards
the south, east, and west of the Rift Valley (Sezonlin et al. 2006,
Assefa et al. 2015, Goftishu et al. 2016). This, as well as tempera-
ture (as indicated by Kaltenbach 1994), could be contributing fac-
tors to the differentiation reported in some African mantid species
as well and should be further investigated. Despite this possible
differentiation, the morphological variety, discolor, of H. tricolor re-
mains a synonym, and thus only one species is known within the
Harpagomantis genus.

The discolor variety of H. tricolor was last recorded in 1977 in
Harkerville in the western Cape, while H. tricolor (except for the
specimen collected during this study) was also collected in the
western Cape during 2015 at Stellenbosch. It should further be
noted that 176 H. tricolor specimens were recorded during this
study; however, the morphological variety of these specimens
could not be identified.

Literature on H. tricolor is somewhat scarce, but some studies
have reported on the distribution of this genus. For example, in
1999, H. tricolor was collected on an indigenous plant species, De-
lairea odorata (Asteraceae) (Cape Ivy), which occurs along the east
coast of South Africa (Grobbelaar et al. 1999). This mantid species
was also recorded in fynbos and native vegetation that were cleared
of alien invasive trees (Magoba and Samways 2010) as well as in
the De Hoop Nature Reserve, a World Heritage site in the western
Cape (Brand and Samways 2009). Harpagomantis specimens were
also recorded in the Highveld grassland biome of South Africa (Bo-
tha et al. 2018, Greyvenstein et al. 2020b). Beyond these studies,
the distribution of this genus is recorded to be throughout South
Africa, but predominantly in Western Cape, KwaZulu Natal, and
Transvaal (Beier 1955). Patel et al. (2016) listed the distribution
of this genus to include Botswana, Namibia, Mozambique, and
Zimbabwe, which is similar to the distribution of the genus de-
scribed by Kaltenbach (1996). As the results of this study indicate
(Fig. 2), various regions in South Africa (i.e., the Northern Cape,
Free State, and parts of the Eastern Cape) had very few to no distri-
bution records. Thus, these regions should be the priority of future
investigations to determine if the extent of the distribution of this
mantid species as presented in this study is a true representation
with regards to H. tricolor in South Africa. This will also shed some
light on the fact that these areas might have been underrepresented
or under-investigated in previous collection efforts and, thus, few
museum specimens from within these regions exist.

Gillon and Roy (1968) reported that a similar species,
Pseudoharpax virescences Serville, 1839 (Mantodea: Galinthiadidae),
occurs from Senegal to Cameroon, while more recently Moulin et
al. (2017) and Moulin (2018) reported Congoharpax aberrans La
Greca, 1954, and various other Galinthiadidae species to be found

23

in the western tropical African countries: the Congo Republic, DR
Congo, Cameroon, Ghana, Ivory Coast, and Gabon. These regions
should be investigated in the future, as they could be a possible
habitat of H. tricolor beyond that recorded in this study. Similarly,
museum specimen records in European countries should also be
incorporated, and citizen science platforms, such as iNaturalist and
iSpot, should be used to establish the distribution of this species.

Ecomorphs (morphologically similar characteristics that align
with particular habitats, such as certain camouflage capabilities)
of mantids such as Harpagomantis are suspected to have evolved
several times in different geographic regions due to similar habi-
tats and ecological pressures (Svenson and Whiting 2009, Wieland
2013, Svenson et al. 2015). The morphological foundation of
Mantodea taxonomy has caused inconsistencies, since the bio-
geographical distribution of ecomorph species was not previously
considered and because a range of species that are morphologi-
cally similar occur on other continents. For example, H. tricolor in
South Africa and species of the genus Creobroter Audinet (Manto-
dea: Hymenopodidae) from India (More and Prashant 2018) are
morphologically similar. Thus, future investigations could shed
some light on the extent of differentiation between the North and
South H. tricolor varieties.

Many specimens were collected in Gauteng province, which
is the region in South Africa with the highest human popula-
tion density. This high population density could explain the large
numbers of specimens collected in this region (Greyvenstein et al.
2020a). However, as reported by Grytnes and Romdal (2008), this
could also be due to ease of access to natural areas where speci-
mens can be collected outside of protected areas. In this study,
most specimens were collected in provincial nature reserves. Davis
et al. (2005) indicated that the Department of Agriculture, Conser-
vation, Environment and Land Affairs has focused on protecting
as much local flora and fauna as possible in provincial and local
nature reserves in highly populated areas throughout South Africa.

This study suggests that provincial nature reserves, more
so than national parks, may create refuge areas for species in a
mosaic of disturbed and highly populated areas. An example of
a provincial and/or local area that can be regarded as a refuge
for birds in highly developed areas was reported by Wang et al.
(2013) in China, where the Hengshui Lake nature reserve, close
to the city of Jizhou, was created as a safe place for migratory
and endangered bird species. However, more research is needed
to determine if protected areas serve as refuges for less mobile
species such as mantids. Future investigations should, therefore,
be conducted to determine if Harpagomantis species are still pre-
sent within provincial nature reserves, as suggested by the historic
specimen records.

Biology.—Since no previous information on the biology of H. tri-
color could be found, comparisons of its biology are made with
species such as Ephestiasula pictipes Wood-Mason, 1879 (Mantodea;
Hymenopodidae) (Vanitha et al. 2016), which is in the Hymeno-
podidae family, from where H. tricolor was moved based on mo-
lecular evidence (Svenson et al. 2015). Due to the lack of literature
about the particular biological parameters measured in this study,
the investigations reported by Vanitha et al. (2016) will be used for
comparison purposes. The study by Gillon and Roy (1968) indicat-
ed various biological parameters of the related species Pseudoharpax
virescences Serville, 1839 (Mantodea: Galinthiadidae) and will thus
be also be used as a comparison to H. tricolor. The shapes of the
oothecae of H. tricolor and P. virescences are similar, but H. tricolor
oothecae are longer and wider than those of P. virescences.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

24

Larsen (2002) suggested that the structure and morphology of
mantid oothecae provide them with the ability to survive harsh
environmental conditions. The function of the shell shape of some
mantid oothecae, for example that of Gongylus Thunberg, 1815
(Mantodea: Empusidae) and Empusa Illiger (Mantodea: Empusi-
dae), is to divert heat (Larsen 2002). Another explanation for the
unique shape and color of the oothecae of some species is that
they aid in crypsis (Thomann 2002). The shape and color of H. tri-
color oothecae resemble, to an extent, the tubercle or auxiliary buds
of plants. This could be an adaptation of this mantid species to
blend into its environment, which is suggested to be predominant-
ly on flowering plants, thus allowing the oothecae to be more in-
conspicuous and limiting unwanted investigation from potential
predators. The ootheca of H. tricolor is an example of the wide va-
riety of structural diversity and cryptic adaptations that are found
throughout the oothecae of Mantodea (Rivera and Svenson 2016).
Their small size (length, width, and height) contributes to their
inconspicuousness, especially on thorny vegetation. The frequency
of H. tricolor oothecae attached to various plant species, as well as
the placement of the oothecae on the plants themselves, should
be investigated in the future to assess the possibility of oothecae
structures as cryptic adaptations and their effectivity as such.

The number of eggs within the oothecae of P. virescences was re-
ported to be between 12 and 13 hatchlings per oothecae (Gillon
and Roy 1968). Unfortunately, no dissections were made by Gillon
and Roy (1968); thus, the number of eggs was not reported. Suck-
ling (1984) reported an average of 34 eggs per ootheca for Orthodera
ministralis (Fabricius, 1775) (Mantodea: Mantidae), which is close to
the maximum number of eggs recorded for H. tricolor in this study.
No differences were observed between any of the size parameters
of fertilized and unhatched oothecae in this study. This is in con-
trast to the significant differences in size of fertilized and unhatched
oothecae of Galepsus lenticularis (Saussure, 1872) (Mantodea: Tara-
chodidae) (Greyvenstein et al. 2020a). Similarly, Greyvenstein et al.
(2020a) reported differences with regards to the number of eggs in-
side fertilized and unhatched oothecae. This was not the case for H.
tricolor, as no differences in this regard were recorded in this study. It
was noted that no oothecae were laid by the captively reared adult
females, which is also in contrast to results reported by Greyvenstein
et al. (2020a) for G. lenticularis. The oothecae of G. lenticularis have
been described as “primitive” and resembling that of the Blattodea
(Ene 1964, Greyvenstein et al. 2020a), those of H. tricolor could sug-
gest a more advanced species based on their evolutionary history. To
this point, different behaviors and biological adaptations could also
be a result of their different evolutionary histories. The difference in
evolutionary traits/ages of the species, environmental stimuli, food-
related resources, or survival strategies could have been the reason
that the captively reared H. tricolor females did not oviposit unferti-
lized oothecae (oothecae produced without mating).

Developmental parameters.—The extended incubation period of
H. tricolor oothecae (145 days) recorded in this study was much
longer than that reported by Vanitha et al. (2016) for E. pictipes. It is
possible that under natural environmental conditions, the oothecae
of H. tricolor undergoes diapause during winter, but under captive
rearing conditions at a constant temperature and humidity, this in-
cubation period was shorter. Overwintering of oothecae has been
reported for some Mantodea species, such as Brunneria borealis Scud-
der, 1896 (Mantodea: Coptopterygidae); Tenodera aridifolia sinensis
Saussure, 1871 (Mantodea: Mantidae); and Mantis religiosa Linnaeus,
1758 (Mantodea: Mantidae) (Kaltenbach 1963, Berg et al. 2011, Mc-
Monigle 2013, Maxwell 2014, Svenson et al. 2015, Hurd et al. 2019).

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

A high hatch rate and low survival rate were reported by Vani-
tha et al. (2016) for E. pictipes, while the opposite was recorded for
H. tricolor in this study. Hatch and survival rates can be influenced
by frequency of feeding, food resources, genetics, and temperature,
depending on the survival strategy of the species (Matthews and
Matthews 1978, Hurd and Eisenberg 1984, Suckling 1984, Iwasaki
1996, Vanitha et al. 2016, Christensen and Brown 2018). The average
duration of the period between mating and production of an oothe-
ca in this study was 12 days, while E. pictipes only required a week to
produce the first ootheca after females mated (Vanitha et al. 2016).

The average duration of copulation for H. tricolor was six houts.
McMonigle (2013) reported that sperm transfer occurs within 30
minutes of the initial copulation action of mantids. The extended
copulation period is suggested to be a form of safeguarding of the
genetic prodigy of the male since this behavior results in decreased
competition with other males (Prokop and Vaclav 2005). Beyond
decreasing sperm competition, males in a better condition (fit-
ness) were also reported to copulate longer with females (Prokop
and Vaclav 2005, Holwell 2006). Strategic ejaculation and adjust-
ment of developmental duration has also been reported in males
of Pseudomantis albofimbriata (Stal, 1860) (Mantodea: Mantidae),
when these males were reared in a male-dominated environment
(Allen et al. 2011). The latter authors reported that male develop-
ment was slower under conditions where many males were pre-
sent, while the opposite was observed when many females were
present. Allen et al. (2011) indicated that male mantids took longer
to mature and suggested that this could indicate more investment
in the development of testes. These males also copulated for a sig-
nificantly longer time and transferred more sperm per copulation
event (Allen et al. 2011). Multiple paternity has been documented
for T. aridifolia by Watanabe et al. (2011) who suggested the exist-
ence of competition between males of the same mantis species for
copulation or mating opportunities. It should be noted that the
artificiality of the mating conditions within this study could have
contributed to the mating behavior results; however, the possible
stress induced by the artificiality was reduced as much as possible
by, for example, ensuring the container in which mating occurred
was ample, that individuals collected from the field were only bred
with after two days to allow the insects to acclimate to the artificial
conditions of the container, and that the container was not moved
or disturbed when the breeding pair were introduced to the single
container but only observed to record the data.

Sexual dimorphism in size where males are smaller than fe-
males has been observed in various mantid species (Wieland
2013). Some examples of mantid species with size sexual dimor-
phism are Creobroter sp. Stal, 1877 (Hymenopodidae), Polyspilota
aeruginosa (Goeze, 1778) (Mantidae), Parasphendale sp. (Mio-
mantidae), and Theopropus elegans (Westwood, 1832) (Mantidae)
(McMonigle 2013). Differences in size and color between sexes
were noted for H. tricolor in this study. However, the duration of
the adult stages did not differ significantly between males and fe-
males of H. tricolor. Differences in antennal morphology were also
noted between male and female T: aridifolia from the sixth instar
onwards (Carle et al. 2014). Pseudomantis albofimbriata (Stal, 1860)
and G. lenticularis exhibit sexual dimorphism, as the wings of
males are fully developed while females are flightless (Holwell et
al. 2006, Greyvenstein et al. 2020a). Sexual dimorphism between
males and females in wings and size could be due to the males
having to find potential reproductive partners. Males of a smaller
size could be more cryptic and more difficult to observe by preda-
tors. The smaller size of males also has dispersal advantages, as it
could increase their ease of flight.

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

Conclusions

This study is the first attempt at mapping the distribution
of Harpagomantis in South Africa and recording the biology of
H. tricolor. The distribution of false flower mantids in South Africa
seems to be predominantly towards the northeastern region, in the
savanna and grassland biomes. Extended copulation duration of
this species could be a by-product of males trying to decrease sperm
competition, which could also have led to the short duration of
the male nymphs compared to female nymphs of H. tricolor. It is
suggested that this species goes into diapause in the ootheca phase.

Acknowledgements

We would like to thank the following people at each of these
institutions for allowing us to access the collections: Audrey Nda-
ba at Ditsong Museum of Natural History (Pretoria), Vivienne Uys
at the Agricultural Research Council (Biosystematics Division),
Asley- Kirk Springs and Burgert Muller at the National Museum
(Bloemfontein), Helen James and Musa Mlambo at the Albany
Museum (Grahamstown), Martin Hill and Thabisa Mdlangu at
Rhodes University (Grahamstown), Kirstin Williams at the Dur-
ban Natural Science Museum, Tricia Pillay at KwaZulu Natal Mu-
seum (Pietermaritzburg), and Aisha Mayekiso at Iziko Museum
of South Africa (Cape Town). We also thank Simon van Noort at
Iziko Museum of South Africa, Entomology Specify. The National
Research Foundation of South Africa contributed funding to this
project (grant number: 101176). Lastly, we thank Nicolas Moulin
for his expertise in identifying the sub-sample of species used for
the biological aspect of this study.

References

Allen LE, Barry KL, Holwell GI, Herberstein ME (2011) Perceived risk of
sperm competition affects juvenile development and ejaculate ex-
penditure in male praying mantids. Animal Behaviour 82: 1207-1206.
https://doi.org/10.1016/j.anbehav.2011.09.009

Assefa Y, Conlong DE, van den Berg J, Martin LA (2015) Ecological genetics
and host range expansion by Busseola fusca (Lepidoptera: Noctuidae).
Environmental Entomology 44: 1265-1274. https://doi.org/10.1093/
ee/nvv079

Beier M (1953) Some new and interesting South African mantids from the
Transvaal Museum. Annals of the Transvaal Museum 22: 255-262.

Beier M (1955) Chapter VIII: Mantidea. In: B Hanstr6m, P Brinck, G Rudebeck
(Eds) South African animal life. Results of the Lund University Expedi-
tion in 1950-1951 (Vol. 2). Almqvist & Wiksell, Stockholm, 234-265.

Berg MK, Schwarz CJ, Mehl JE (2011) Die Gottesanbeterin, Mantis religiosa:
Die Neue Brehm-Bicherei. Westarp Wissenschaften, Hohenwarsle-
ben, 33 pp.

Botha M, Siebert SJ, van den Berg J, Ellis S, Greyvenstein BM (2018) Diver-
sity patterns of selected predaceous arthropod groups in maize fields
and margins in South African Highveld grassland. Agricultural and
Forest Entomology 20: 461-475. https://doi.org/10.1111/afe.12277

Brand MR, Samways MJ (2009) Comparative impact of invasive alien trees
and vineyards on arthropod diversity in the Cape Floristic Region,
Western Cape. MSc dissertation. University of Stellenbosch, Stellen-
bosch, South Africa, 296 pp.

Brannoch SK, Wieland EF Rivera J, Klass K, Bethoux O, Svenson GJ (2017)
Manual of praying mantis morphology, nomenclature and practices
(Insect, Mantodea). ZooKeys 696: 1-100. https://doi.org/10.3897/
zookeys.696.12542

Cardoso P, Barton PS, Birkhofer K, Chichorro EF Deacon C, Fartmann
T, Fukushima CS, Gaigher R, Habel JC, Hallmann CA, Hill MJ,
Hochkirch A, Kwak ML, Mammola S, Noriega JA, Orfinger AB, Pe-

25

draza F, Pryke JS, Roque FO, Settle J, Simaika JP, Stork NE, Suhling
F, Vorster C, Samways MJ (2020) Scientists’ warning to humanity on
insect extinctions. Biological Conservation 242: 1-13. https://doi.
org/10.1016/j.biocon.2020.108426

Carle T, Yamawaki Y, Watanabe H, Yokohari F (2014) Antennal develop-
ment in the praying mantis (Tenodera aridifolia) highlights multitudi-
nous processes in Hemimetabolous insect species. PLoS ONE 9: 1-14.
https://doi.org/10.1371/journal.pone.0098324

Christensen T, Brown WD (2018) Population structure, movement
patterns, and frequency of multiple matings in Tenodera sinensis
(Mantodea: Mantidae). Environmental Entomology 47: 676-683.
https://doi.org/10.1093/ee/nvy048

Davis ALV, Scholtz CH, Deschodt C (2005) A dung beetle survey of
selected Gauteng nature reserves: implications for conservation of the
provincial scarabaeine fauna. African Entomology 13: 1-16. https://
hdl.handle.net/10520/EJC32629

Edmunds M (1971) Defensive behaviour in Ghanaian praying mantids.
Zoological Journal of the Linnean Society 51: 1-32. https://doi.
org/10.1111/j.1096-3642.1972.tb00771.x

Ehrmann R (2002) Mantodea: Gottesanbeterinnen der Welt. Natur und
Tier Verlag: Munster, 519 pp.

Ene JC (1964) The distribution and _ post-embryonic develop-
ment of Tarachodes afzelii (Stal), (Mantodea: Eremiaphilidae).
Annals and Magazine of Natural History 7: 493-511. https://doi.
org/10.1080/00222936408651488

Fatimah S, Sultana R, Wagan MS (2016) Study on the gender identification
of praying mantids (Dictyoptera: Mantodea). Journal of Entomology
and Zoology Studies 4: 529-531.

Giglio-Tos E (1927) Mantidae. In: Das Tierreich: Eine Zusamenstellung
und Kennzeichnung der rezenten Tierforrnen Lieferung (eds. FE
Schulze & W Kukenthal). Walter de Gruyter & Co.: Berlin, 650-592.

Gillon Y, Roy R (1968) Les Mantes de Lamto et des savanes de Cote d'Ivoire.
Bulletin de l'IFAN. Série A - Sciences Naturelles 30: 1038-1151.

Goftishu M, Assefa Y, Fininsa C, Niba A, Capdevielle-Dulac C, Le Ru BP
(2016) Phylogeography of Sesamia cretica Lederer (Lepidoptera:
Noctuidae). Phytoparasitica 44: 641-650. https://doi.org/10.1007/
$12600-016-0556-8

Greyvenstein B, Du Plessis H, Moulin N, van den Berg J (2020a) Distribu-
tion of Galepsus spp. in Southern Africa and life history of Galepsus
lenticularis (Mantodea: Tarachodidae). Insects 11: 1-17. https://doi.
org/10.3390/insects 11020119

Greyvenstein B, Siebert SJ, van den Berg J (2020b) Effect of time of day
on efficacy of sweep net sampling of arthropod predators in maize
agro-ecosystems in the North West Province, South Africa. African
Entomology 28: 150-163. https://doi.org/10.4001/003.028.0150

Grobbelaar E, Neser S, Neser OC (1999) Biological control of Delairea
odorata in California: a survey for its insect natural enemies in South
Africa. A report to the Agricultural Research Service, United States
Department of Agriculture. Agricultural Research Council: Pretoria,
South Africa, 64 pp.

Grytnes J, Romdal TS (2008) Using museum collections to estimate
diversity patterns along geographical gradients. Folia Geobotanica 43:
357-359. https://doi.org/10.1007/s12224-008-9017-6

Holwell GI, Barry KL, Herberstein ME (2006) Mate location, antennal
morphology, and ecology in two praying mantids (Insecta: Manto-
dea). Biological Journal of the Linnean Society 91: 307-313. https://
doi.org/10.1111/j.1095-8312.2007.00788.x

Hurd LE, Eisenburg RM (1984) Experimental density manipulations of the
predator Tenodera sinensis (Orthoptera: Mantidae) in an old-field com-
munity. I. Mortality, development and dispersal of juvenile mantids.
Journal of Animal Ecology 53: 269-281. https://doi.org/10.2307/4356

Hurd LE, Cheng KX, Abcug J, Calhoun LV, Geno ME, Merhige RR, Rosenthal
IH (2019) Climate change, succession, and reproductive success of
a praying mantid. Annals of the Entomological Society of America
saz070: 1-5. https://doi.org/10.1093/aesa/saz070

Iwasaki T (1996) Comparative studies on the life histories of two praying
mantises, Tenodera aridifolia (Stoll) and Tenodera angustipennis Saus-
sure (Mantodea: Mantidae) I. Temporal pattern of egg hatch and nym-

JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1)

26

phal development. Applied Entomology and Zoology 31: 345-356.
https://doi.org/10.1303/aez.31.345

Kaltenbach AP (1963) Kritische Untersuchungen zur Systematik, Biologie
und Verbreitung der europdischen Fangheuschrecken (Dictyoptera -
Mantidae). Zoologische Jahrbiicher Abteilung Systematik 90: 521-598.

Kaltenbach AP (1994) Bisanthe menyharthi (Brancsik, 1895): Klinale Vari-
ation, Subspecies-Bildung und geographische Verbreitung (Insecta:
Mantodea: Mantidae). Annalen des Naturhistorischen Museums in
Wien - Serie B fiir Botanik und Zoologie 96: 59-67.

Kaltenbach AP (1996) Unterlagen fiir eine Monographie der Mantodea
des siidlichen Afrika: 1. Artenbestand, geographische Verbreitung und
Ausbreitungsgrenzen (Insecta: Mantodea). Annalen des Naturhis-
torischen Museums in Wien 98: 193-346.

Kaltenbach AP (1998) Unterlagen fiir eine Monographie der Manto-
dea (Insecta) des siidlichen Afrika: 2. Bestimmungstabellen ftir die
hdheren Taxa, Nachtrage zum Artenbestand. Annalen des Naturhis-
torischen Museums in Wien 100: 19-59.

Karny H (1908) Orthoptera (I.) Blattaeformia Oothecaria. In: Schultze L
(Ed.) Zoologische und anthropologische Ergebnisse einer Forschun-
gsreise im westlichen und zentralen S’dafrika—Denkschriften der
medizi—ch - naturwissenschaftlichen Gesellschaft zu. Jenaische
Denkschriften, Berlin, 335-390.

Larsen T (2002) The wandering violin: spotlight on Gongylus gongyloides.
Invertebrate 1: 4-9.

Lombardo F (1995) A review of the genus Popa Stal, 1856 (Insecta Manto-
dea). Tropical Zoology 8: 257-267. https://doi.org/10.1080/039469
73:4.995. 10539285

Magoba RNN, Samways MJ (2010) Comparative impact of invasive alien
trees and vineyards on arthropod diversity in the Cape Floristic Region,
Western Cape. PhD thesis, University of Stellenbosch, Stellenbosch.

Matthews RW, Matthews JR (1978) Insect Behaviour. John Wiley & Sons:
New York, 507 pp.

Maxwell M (2014) Developmental patterns in Stagmomantis limbata (Mantodea:
Mantidae): variation in instar number, growth, and body size. Journal of
Orthoptera Research 23: 49-58. https://doi.org/10.1665/034.023.0104

McMonigle O (2013) Keeping the Praying Mantis. Coachwhip Publica-
tions, Greenville, 202 pp.

More SV, Prashant MS (2018) Diversity of praying mantids from Tilari for-
est, Chandgad, Kolhapur district of Maharashtra India. International
Journal of Entomology Research 3: 57-64.

Moulin N, Decaéns T, Annoyer P (2017) Diversity of mantids (Dictyoptera:
Mantodea) of Sangha-Mbaere Region, Central African Republic, with
some ecological data and DNA barcoding. Journal of Orthoptera Re-
search 26: 117-141. https://doi.org/10.3897/jor.26.19863

Moulin N (2018) Liste commentée et catalogue illustré des Mantodea du
Gabon. Les cahiers de la fondation Biotope 24: 2-60.

O'Toole C (2003) Preying for dinner-time. Nature 422: 564-565. https://
doi.org/10.1038/422564b

Patel S, Singh G, Singh R (2016) Global distribution of Empusidae,
Eremiaphilidae, Galinthiadidae and Iridopterygidae (Mantodea: Dic-
tyoptera: Insecta): A Checklist. International Journal of Zoological
Investigations 2: 219-236.

Prokop P, Vaclav R (2005) Males respond to the risk of sperm competition
in the sexually cannibalistic praying mantis, Mantis religiosa. Ethology
111: 836-848. https://doi.org/10.1111/j.1439-0310.2005.01113.x

Rehn JAG (1927) Contributions to our knowledge of the Dermaptera and
Orthoptera of the Transvaal and Natal — Part I]: Mantidae. Annals of
the Transvaal Museum 12: 1-54.

Rivera J, Svenson GJ (2016) The Neotropical ‘polymorphic earless praying
mantises’ — Part I: molecular phylogeny and revised higher-level sys-
tematics (Insecta: Mantodea, Acanthopoidea). Systematic Entomol-
ogy 41: 607-649. https://doi.org/10.1111/syen.12178

B. GREYVENSTEIN, H. DU PLESSIS AND J. VAN DEN BERG

Roy R, Stiewe M (2014) Révision du genre Galinthias Stal, 1877 (Mantodea,
Galinthiadidae). Bulletin de la Société Entomologique de France 119:
199-215.

Samways MJ, Barton PS, Birkhofer K, Chichorro F Deacon C, Fartmann
T, Fukushima CS, Gaigher R, Habel JC, Hallmann CA, Hill MJ,
Hochkirch A, Kaila L, Kwak ML, Maes D, Mammola S, Noriega JA,
Orfinger AB, Pedraza E Pryke JS, Roque FO, Settele J, Simaika JP, Stork
NE, Suhling FE, Vorster C, Cardoso P (2020) Solutions for humanity on
how to conserve insects. Biological Conservation 242: 1-15. https://
doi.org/10.1016/j.biocon.2020.108427

Schwarz CJ, Roy R (2019) The systematics of Mantodea revisited: an up-
dated classification incorporating multiple data sources (Insecta:
Dictyoptera). Annales Societe Entomologique de France 55: 101-196.
https://doi.org/10.1080/00379271.2018.1556567

Sezonlin M, Dupas S, Le Ri B, Faure N, Le Gall P, Silvain J (2006) Phyloge-
ographic pattern and regional evolutionary history of the maize stalk
borer Busseola fusca (Fuller) (Lepidoptera: Noctuidae) in sub-Saharan
Africa. Annales de la Société entomologique de France 42: 339-351.
https://doi.org/10.1080/00379271.2006.10697466

Solomon ME (1951) Control of humidity with potassium hydroxide, sul-
phuric acid or other solutions. Bulletin of Entomological Research 42:
543-554. https://doi.org/10.1017/S0007485300028947

Stal C (1877) Systema Mantodeorum. Essai d’une systématisation nou-
velle des Mantodées. Bihang till Kongliga Svenska Vetenskaps Akad-
emiens Handlingar 4: 1-91.

Suckling DM (1984) Laboratory studies on the praying mantis Orthodera
ministralis (Mantodea: Mantidae). New Zealand Entomologist 8: 96-
101. https://doi.org/10.1080/00779962.1984.9722478

Svenson GJ, Whiting MF (2009) Reconstructing the origins of praying
mantises (Dictyoptera, Mantodea): the roles of Gondwanan vicari-
ance and morphological convergence. Cladistics 25: 468-514. https://
doi.org/10.1111/j.1096-0031.2009.00263.x

Svenson GJ, Hardy NB, Wightman HMC, Wieland F (2015) Of flowers and
twigs: phylogenetic revision of the plant-mimicking praying mantises
(Mantodea: Empusidae and Hymenopodidae) with a new suprage-
neric classification. Systematic Entomology 40: 789-834. https://doi.
org/10.1111/syen.12134

Thomann CH (2002) Central-southeastern U.S. mantid Stagmomantis caro-
lina Johansson 1763). Invertebrate 4: 1-3.

TIBCO Software Inc (2017) Statistica (data analysis software system),
version 13.3. http://statistica.io

Vanitha K, Bhat PS, Raviprasad TN, Srikumar KK (2016) Biology and be-
haviour of Ephestiasula pictipes (Wood-Mason) (Hymenopodidae:
Mantodea) under captive breeding. International Journal of Pest
Management 62: 308-318. https://doi.org/10.1080/09670874.2016.
1195026

Wang PEY, Xiaomei Deng X, Marcucci DJ, Le Y (2013) Sustainable Devel-
opment Planning of Protected Areas near Cities: Case Study in China.
Journal of Urban Planning and Development 139: 133-143. https://
doi.org/10.1061/(ASCE)UP.1943-5444.0000133

Watanabe E, Adachi-Hagimori T, Miura K, Maxwell MR, Ando Y, Take-
matsu Y (2011) Multiple paternity within field-collected egg cases
of the praying mantid Tenodera aridifolia. Annals of the Entomo-
logical Society of America 104: 348-352. https://doi.org/10.1603/
AN10035

Wieczorek J, Guo Q, Hijmans RJ (2004) The point-radius method for
georeferencing locality descriptions and calculating associated uncer-
tainty. International Journal of Geographical Information Science 18:
745-767. https://doi.org/10.1080/13658810412331280211

Wieland F (2013) The phylogenetic system of Mantodea (Insecta: Dic-
tyoptera). Species, Phylogeny and Evolution 3: 3-222. https://doi.
org/10.17875/gup2013-711

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