Documenting Mantodea species in South African museum collections and an updated species list

Abstract Background The previous species list of South African Mantodea, published in 1998, was largely compiled from the literature and did not incorporate data from the many insect museum collections available in the country. It is estimated that approximately 120 species of Mantodea occur in South Africa; however, since no historical museum records were previously incorporated, the current information is considered to be outdated and not a true reflection of the Mantodea fauna within this region. A checklist of species is an important benchmark for any insect group, especially in light of the worldwide declines of insect diversity reported over the last decade. Checklists that provide accurate information on insect diversity, especially for groups, such as the Mantodea which could be under threat and thus could provide important information that can be used in determining the threat status of species, as well as to aid in their conservation in general. New information This paper provides an updated checklist of the praying mantids (Insecta, Mantodea) species of South Africa. While 120 species were previously reported to occur in South Africa, this paper reports 157 species in 64 genera that represent eight different superfamilies, 14 families and 22 subfamilies. Additionally, five species are reported for the first time to occur in South Africa. This species list was generated from the approximately 4000 specimen records of which 3558 records reside within South Africa. The remaining 732 records represent 14 other African countries. Occurrence records from two citizen-science platforms (iNaturalist and Gbif.org), were also incorporated in this study, adding 1880 species records in South Africa. The low number of specimens in the national collections indicate that this group of insects is poorly collected and highlights the lack of knowledge about South Africa’s mantid fauna, as well as a lack of taxonomic expertise as 1532 museum specimens remain unidentified to species level.


Introduction
Until recently, the Mantodea Order consisted of approximately 24 families with 2400 species (Wieland and Schutte 2012, McMonigle 2013, Wieland 2013, Green 2014).The classification system has recently been revised and the Mantode Order now composed of 16 superfamilies, 29 families and 436 different genera (Schwarz and Roy 2019).
Approximately 120 species of Mantodea were reported to occur in South Africa (Schoeman 1985a, Schoeman 1985b), when the previous species list was compiled between 1996 and 1998 (Kaltenbach 1996, Kaltenbach 1998).Kaltenbach (1996) estimated that there were approximately 131 Mantodea species in South Africa and that 19% of these species were endemic.Ehrmann (2002) estimated a total of 125 species within the region.Beyond the abovementioned checklist information, very little is known of South African Mantodea biology and ecology.A 2023 Scopus (www.scopus.com)internet search of published scientific papers indicated that between 1927 and 2023, 792 papers were published on Mantodea worldwide.However, only 15 of these publications were from institutions in South Africa, seven of which belong to the authors of this paper and two papers were the previous checklist from Kaltenbach (1996), Kaltenbach (1998).The remaining papers conducted on Mantodea in South Africa all addressed molecular and genetic aspects and, in some of these cases, it was actually Blattodea that were investigated.Studies on the biology and distribution of mantids throughout the world are limited and, in South Africa, largely absent.
It is possible that many mantid species in the southern African region have not been documented yet.The only surveys of Mantodea in South Africa were done by Kaltenbach (1996) and the Mantodea Project which was done in collaboration with the Cleveland Museum of Natural history in Ohio, USA (Svenson et al. 2012).However, no species list from the latter survey was published.The latter survey was done during 2005 and only included three regions within South Africa (Cape floristic region, Richards Bay in KwaZulu-Natal Province, and the Kruger National Park in Mpumalanga Province).This paper, compiled from museum records and previous checklists by Kaltenbach (1996), Kaltenbach (1998), contributes to the information on Mantodea in South Africa and identifies the knowledge gaps with regards to mantids in South Africa.An example of Mantodea museum specimens in a collection which were identified (including genitalia plates if applicable per species) by various taxonomists.These specimens were amongst those used to compile the species list in this paper.

All
Documenting Mantodea species in South African museum collections and an ... identified by means of the literature and through assistance from a taxonomist who specializes in African Mantodea (Nicolas Moulin).Many Mantodea specimens in South African collections have only been identified to genus level.These "ignota specimens" (approximately 1600) were, therefore, not included in this checklist.However, they are included in the database itself (available in Suppl.material 1).
In order to compile this database, all of the Mantodea specimens and distribution labels were photographed and the label information documented.This database contains the following information for each specimen record: genus and species name, collector's details, collection date, if available, and locality.The website mantodeaspeciesfile.org (Otte et al. (2022) as well as other literature on specific species, such as those by Ehrmann (2002), Roy (2004), Roy (2006), Roy (2009), Roy (2010), Roy (2013), Roy (2018), Roy (2022) were used to determine the current nomenclature.The reclassification of the Order Mantodea by Schwarz and Roy (2019) was also applied during the updating of this checklist.The updated species list was compared to that provided in publications by Kaltenbach (1996), Kaltenbach (1998) after which similarities and differences were highlighted.
To our knowledge, this paper provides the most comprehensive list of Mantodea in South African collections.Since only a limited number of Mantodea specimens of 14 other African countries were present in South African museum collections, these records were not included in this paper.The scope of this study, did however not allow for vistits to museums residing outside of South Africa, which is required when the latter information is used to compile comprehensive Mantodea species lists for these African countries.However, to increase the comprehensiveness of this checklist, various European and American museum collections were contacted and provided information on South African Mantodea in their collections.These museums were: The Natural History Museum, (Italy); Smithsonian National Museum of Natural History (USA), Museum für Naturkunde (Germany), Natural History Museum (UK), Staatliches Museum für Naturkunde (Germany), Royal Belgian Institute of Natural Sciences (Belguim) and the Muséum national d'Histoire naturelle (France).Data from Mantodea specimens in a private collection in Germany (Christian Schwarz), as well as records relevant Gbif.org records within South Africa were also included in this study.
This species list includes information on the taxonomists who identified the species in the South African museum collections and is indicated for each species by the "ID" tag, as well as the year in which the specimen was identified (if available).Furthermore, the hosting museum collection of each specimen is also included in brackets ().A list of abbreviations for the various institutions and collections are provided in Table 1.Specimens that are not held locally, or for which only literature records exist, are indicated under the ID tag column in the checklist as with the abbreviation "Lit" with the reference to the relevant publications.The number of records in the various collections in South Africa, as well as the number of Research Grade observation records of the species listed on the two citizen-science platforms, are provided in Suppl.material 1.It should be noted that no details of the persons who provided identifications of species listed on the citizen-science platforms are listed in the checklist.This will be addressed in the Discussion section of the paper.
The geographical distribution of species beyond South Africa, is based on information provided by Roy (1967), Kaltenbach (1996), Kaltenbach (1998), Ehrmann (2002), Roy (2004), Roy (2006), Roy (2009), Roy (2010), Roy (2013), Roy (2018), Roy (2022) as well as museum records indicated below, after the tag "Distribution".The abbreviations used for the different countries are listed in Table 2.It should be noted that all the species listed below are present in South Africa and, thus, this country is not listed under the distribution.Where there is no "Distribution" tag associated with a species, the species has only been reported from South Africa.The specimen type, i.e., Holotype, Paratype or DNA barcode, sex, and museums in which they are kept, is provided in the Suppl.material 1.
List of abbreviations for the museums and collections in which the Mantodea specimen are hosted.
Documenting Mantodea species in South African museum collections and an ...

Abbreviation Country
Abbreviation Country

Results
This updated checklist includes information on species of the Mantodea that were not previously listed in South African checklists.The known species richness has increased from approximately 120 species in 1998 to 157 species (this report).The South African Mantodean fauna have eight superfamilies, 14 families, 22 subfamilies, 19 tribes, 14 subtribes and 15 genera (Suppl.material 2).A summation of the number of records and species within these 14 families is presented in Fig. 2.
Abbreviations of country names listed in the section describing the distribution of different Mantodea species recorded in South Africa.
This checklist encompasses 157 Mantodea species that occur, or are reported to occur in South Africa, including the first report of five species within the region (indicated with two asterisks ** in the notes section of each species).However, some anomalies were recorded (indicated by # in the notes section of each species).These anomalies are addressed in the Discussion section of this paper.Native status: Suspected to be endemic to southern Africa (Kaltenbach 1996) Distribution: MOZ Dystactula natalensis Kaltenbach, 1996 Native status: Suspected to be endemic to southern Africa (Kaltenbach 1996) Notes: ID: Lit (Kaltenbach 1996, Ehrmann 2002) Tribe Chroicopterini Giglio-Tos, 1915

Analysis
This checklist was compiled from a database that has been generated after recording all the available details of specimens (approximately 4000 records over 170 years) in eight South African museum collections.An additional 1945 Mantodea records from private collections, several museums outside of South Africa, and two citizen-science platforms were included.Although all specimens were identified to family level, a large number (1600) were only identified to genus level.All records within the database could, therefore, be used to generate distribution maps for the 14 Mantodea families (Fig. 3).Despite only a few distribution records being available for specimens of some of the families, distribution patterns indicate that all families occur in the hotspots indicated in this study.These hotspots are the north-eastern parts of the Savannah biome (towards the Kruger National Park), along the eastern coast in the Indian Ocean coastal belt (KwaZulu-Natal province), southern coastal region in the thicket biome (Eastern Cape -Gqeberha), and fynbos biome in the south-western Cape region.The families with the lowest species richness and the lowest number of records were Amorphoscelidae, Dactylopterygidae, Nanomantidae, Rivetinidae and Toxoderidae (Fig. 3).The three most species-rich families were Mantidae, Miomantidae and Eremiaphilidae (Fig. 3), which were also the families with the highest numbers of specimen records.The latter three families made up 50.2% of the total number of specimens in the surveyed collections, with those in the Eremiaphilidae having the highest representation (853 specimens).The distribution maps do, however, elude that some areas of the country have either been under-represented or the abundance of mantids in these regions, for example the Northern Cape Region, is very low.Interestingly, the distribution of Empusidae, which is only represented by seven species, indicates a region-wide distribution and more than 200 records of this family were recorded during this study.

Discussion
This paper illustrates the value of museum data, although it was only after the documentation thereof, that these data allowed us to update the Mantodea species list of South Africa.Historic data encompasses many years of collected specimens and, as suggested by Hill et al. (2012), it remains as data that are invaluable and irreplaceable.
Although the more modern methods of observational data collection through citizenscience platforms are easily accessible (Moulin 2020), it remains problematic as the identification of these specimens are difficult, especially for groups such as arthropods.This is due to the vast numbers of insect species and because, in many cases, identification requires microscopic investigation of the genitalia and wing venation.Due to these difficulties, as well as data only at genus level, records sourced from the citizenscience platforms were only used to compile the distribution maps and they were not included in the supplementary information.Although citizen-science platforms may provide valuable information (Moulin 2020), care should be taken when data are used.For example, 11 Miomantis species were recorded from the museum collections during this study, while nine other species are also suggested to occur in South Africa (Kaltenbach 1996, Kaltenbach 1998).However, all 467 Miomantis observations from iNaturalist and GBIF.org were listed as Miomantis caffra Saussure, 1871.Since no other Miomantis species has been recorded on the latter platforms, it is highly likely that some of the Miomantis species level identifications were incorrect.Despite this type of error, the data recorded on citizen-science platforms can be helpful to determine distribution patterns.These platforms are becoming increasingly important, but there is no substitution for taxonomic expertise and investigation of specimens, especially for the many species of Mantodea that have not yet been added to publicly accessible DNA databases.For example, of the 157 species listed in this checklist, only 50 are represented on DNA databases such as GenBank, NCBI and BOLD.
Another example which illustrates the value of data from citizen-science platforms (Gbif.org in this case), as well as the caution needed in interpreting such data, is that of Pseudocreobotra ocellata.This species, according to literature, is native to North Africa and does not occur in South Africa (Ehrmann 2002).However, a DNA barcode of the latter species was found on the BOLD database (Ratnasingham and Hebert 2007) during this study which, according to the locality of the specimen used for the DNA analysis, it was collected in KwaZulu-Natal in South Africa (Suppl.material 1).The sequence provided on BOLD is 99.23% similar to that of an unpublished DNA sequence of Pseudocreobotra wahlbergi that is available on GenBank.The latter sequence has never been published and should, thus, be treated with caution.The specimens identified as P. occellata on the BOLD system could either be new distribution records or be due to a lack of reliable P. wahlbergi sequences.
One of the species, Galepsus centralis, which has not previously been reported from South Africa, was collected in the Grassland biome in the north-western region of South Africa and identified by a mantid taxonomist (Nicolas Moulin) during this study.This species was previously reported to occur only in Tanzania and the Democratic Republic of the Congo.It could also be possible that this species has always occurred in southern Africa, but was not detected or it has expanded its range to southern Africa, similar to what has been reported for other mantid species in recent years (Schwarz and Ehrmann 2018).Interestingly, Kaltenbach (1996) listed some species that may be endemic to southern Africa, including Miomantis caffra.The latter species has expanded its range to New Zealand and Australia (Ramsay 1984, Connors et al. 2022).
This species list indicates that the diversity of Mantodea in South Africa is high and that approximately 6% of the known Mantodea species worldwide occurs in this region.A few areas within South Africa seem to be "hotspots" or regions with high diversity and should be investigated further.These areas may be related to the biomes within the country since insect communities tend to be closely correlated to plant communities (Schaffers et al. 2008).It is, therefore, suggested that the "hotspot" areas identified in this study be priority areas for future research.The conglomeration of distribution records on the maps in areas such as Pretoria may be due to the ease of access (Grytnes and Romdal 2008) of this museum as it resides within a large city with a large surrounding human population.
The current state of knowledge suggests that South Africa could have a high level of Mantodea endemicity, i.e. 38% of the species from this study are suggested by Kaltenbach (1996) to be endemic to southern Africa.Furthermore, "priority species" (for example, the 24 species of which only one record exists) were identified and should be investigated first to address the severe lack of knowledge regarding these species.For example, Calamothespis oxyops has a single distribution record (Baberton 1910), i.e. the Holotype specimen in the Ditsong National Museum of Natural History.One other record of this species was recorded on iNaturalist (which could indicate that it only occurs in South Africa).This citizen-science record has, however, not been verified (not Research Grade) and was, thus, not included in the Suppl.material 1. Lastly, the aim of this paper was not only to update the Mantodea checklist of South Africa, but also to develop a dataset that can guide future research on Mantodea diversity within the region.The absence of taxonomic expertise to identify Mantodea in South Africa provided a challenge during the compilation of this checklist.This was addressed through collaboration with the international Mantodea scientist community who assisted with identifications, provided taxonomic keys and shared old literature regarding the Mantodea of the region.
of the National insect collections and museums throughout South Africa were visited during this study.The following seven institutions constitutes the national insect collections in the country: Ditsong Museum of Natural History (Pretoria) (DNMNH), Agricultural Research Council (Biosystematics Division, Pretoria) (ARC), National Museum (Bloemfontein) (NMB), Albany Museum (Makhanda) (AMG), Rhodes University (Makhanda) (RU), Durban Natural Science Museum (DNSM), Iziko South African Museum (Cape Town) (Iziko) and KwaZulu-Natal Museum (Pietermaritzburg) (NMSA).Specimens in these collections where mostly identified by visiting taxonomists during previous visits to these institutions.Many of the museum specimens were also previously identified by taxonomists at the departments of Dr. Max Beier at the Vienna museum in Germany, Dr. James Rehn at the Academy of Natural Sciences in Philadelphia, USA, Dr. Alfred Peter Kaltenbach at the Natural History Museums in Wien, Austria and Dr. Roger Roy at the Muséum national d'Histoire naturelle (MNHN) in France (Fig. 1).Furthermore, a small subset of South African Mantodea were identified by Nicolas Moulin, Honorary Associate at Muséum national d'Histoire naturelle in France, during 2019.Unidentified specimens that were encountered in the abovementioned museums were Figure 1.

Figure 2 .
Figure 2. Summary of the number of species and number of records per Mantodea family recorded in South Africa.

Figure 3 .
Figure 3. Distribution maps of the Mantodea families that occur within South Africa.The number of specimen records per family is provided in brackets.