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New data on the presence of Hierodula transcaucasica Brunner von Wattenwyl, 1878 on Crete/Greece (Mantodea: Mantidae)

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Three years ago, a finding of a mantis on the Greek island of Crete was reported and discussed in this journal. At that time, the species was not identified without any doubt. Actually, the specimen was a female of Hierodula transcaucasica Brunner von Wattenwyl, 1878. Recent data on the presence of this species on Crete are presented, including a distribution map. The nomenclatural issue concerning the taxonomic identity of this species is discussed.
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New data on the presence of Hierodula transcaucasica Brunner von Wattenwyl,
1878 on Crete/Greece (Mantodea: Mantidae)
Nuevos datos sobre la presencia de Hierodula transcaucasica Brunner von Wattenwyl, 1878, en
Creta/Grecia (Mantodea: Mantidae)
Torsten van der Heyden 1, Christian J. Schwarz 2
1. Member of the editorial board of BV news Publicaciones Científicas Hamburg (Germany)
tmvdh@web.de
2. Ruhr University Bochum, Department of Biology and Biotechnology, ND 1/31 Bochum
(Germany)
ChristianSchw@gmx.de
ABSTRACT: Three years ago, a finding of a mantis on the Greek island of Crete was reported and
discussed in this journal. At that time, the species was not identified without any doubt. Actually, the
specimen was a female of Hierodula transcaucasica Brunner von Wattenwyl, 1878. Recent data on the
presence of this species on Crete are presented, including a distribution map. The nomenclatural issue
concerning the taxonomic identity of this species is discussed.
KEY WORDS: Hierodula tenuidentata Saussure, 1869, Hierodula transcaucasica Brunner von Wattenwyl,
1878, Sphodromantis viridis (Forskål, 1775), Mantidae, Mantodea, distribution, Crete, Greece.
RESUMEN: Hace tres años se presentó y discutió en esta revista un hallazgo de una mantis en la isla griega
de Creta. En ese momento, la especie no se pudo identificar con absoluta precisión. Realmente, el espécimen
era una hembra de Hierodula transcaucasica Brunner von Wattenwyl, 1878. Se presentan datos recientes
sobre la presencia de esta especie en Creta, incluyendo un mapa de distribución. Se discute la cuestión de la
nomenclatura relativa a la identidad taxonómica de esta especie.
PALABRAS CLAVE: Hierodula tenuidentata Saussure, 1869, Hierodula transcaucasica Brunner von
Wattenwyl, 1878, Sphodromantis viridis (Forskål, 1775), Mantidae, Mantodea, distribución, Creta, Grecia.
Introduction
On 24-VII-2018, a specimen of a mantis was observed and photographed in Kouses on the Greek
island of Crete (Fig. 1).
At the time, the species was not identified without any doubt. It was assumed that the specimen
might be a female of Hierodula transcaucasica Brunner von Wattenwyl, 1878 or of Sphodromantis viridis
(Forskål, 1775) (VAN DER HEYDEN, 2018b). The specimen turned indeed out to be a female of H.
transcaucasica, an invasive species which has recently become established in the Mediterranean and the
Balkans (BATTISTON et al., 2018, 2020b; CIANFERONI et al., 2018; VAN DER HEYDEN, 2018c; ROMANOWSKI
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et al., 2019; SCHWARZ & EHRMANN, 2019; ZLATKOV et al., 2020; PINTILIOAIE et al., 2021; VUJIĆ et al.,
2021). Recently, more specimens of H. transcaucasica were found on Crete. They are presented here.
Fig. 1: Female of Hierodula transcaucasica Brunner von Wattenwyl, 1878, Kouses, Crete, Greece, 24-VII-
2018, (VAN DER HEYDEN, 2018a).
https://www.biodiversidadvirtual.org/insectarium/Paramantini-img1019338.html
Material and methods
The distribution map was created in SimpleMappr (SHORTHOUSE, 2010). We included new
specimens encountered and photographed by the first author as well as data from the online platform
INATURALIST (2021).
Abbreviations
MHNG: Muséum d’histoire naturelle de la Ville de Genève
NHMW: Naturhistorisches Museum Wien
Recent findings of Hierodula transcaucasica Brunner von Wattenwyl, 1878 on Crete
On 9-VII-2021, a nymph of H. transcaucasica was observed and photographed in Kalamaki, a small
village located at the southern coast of Crete (Fig. 2).
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Fig. 2: Nymph of Hierodula transcaucasica Brunner von Wattenwyl, 1878, Kalamaki, Crete, Greece, 9-VII-
2021. (Photo: Torsten van der Heyden)
On 11-VII-2021 and on 12-VII-2021, single nymphs of H. transcaucasica were found at the same
location in Kalamaki. Additionally, an exuvia of the species was found just a few meters away on 13-VII-
2021 (Fig. 3).
Fig. 3: Exuvia of Hierodula transcaucasica Brunner von Wattenwyl, 1878, Kalamaki, Crete, Greece, 13-
VII-2021. (Photo: Torsten van der Heyden)
Furthermore on 13-VII-2021, another nymph of H. transcaucasica was observed and photographed
in Neo Kalamaki, about 600 m southeast of the location of the findings mentioned before (Fig. 4).
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Fig. 4: Nymph of Hierodula transcaucasica Brunner von Wattenwyl, 1878, Neo Kalamaki, Crete, Greece,
13-VII-2021. (Photo: Torsten van der Heyden)
Additionally, photos of five specimens of H. transcaucasica recently found on Crete by several
individuals and identified as Hierodula tenuidentata Saussure, 1869 were uploaded to INATURALIST (2021):
Single specimens were found near Chania (northwestern part of Crete), near Agios Nikolaos (northeastern
part of the island) and in Agios Ioannis and Matala (both villages are located in the same area as Kouses,
Kalamaki and Neo Kalamaki) (Fig. 5).
Fig. 5: Distribution of Hierodula transcaucasica Brunner von Wattenwyl, 1878 on Crete, Greece (red dots).
Discussion
Taking the recently recorded specimens into account, it appears that H. transcaucasica is well
established on the island. Since specimens were found in different locations in various areas of Crete, it is
very likely that the species has colonized most parts of the island. Currently, records on Crete are
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concentrated around urbanized areas, which is in concordance with most records in other parts of the
colonized range (BATTISTON et al., 2018; CIANFERONI et al., 2018; VAN DER HEYDEN, 2018c;
ROMANOWSKI et al., 2019; SCHWARZ & EHRMANN, 2019; ZLATKOV et al., 2020; PINTILIOAIE et al., 2021;
VUJIĆ et al., 2021).
This bias has to be regarded as both distributional and methodological. On the one hand, SCHWARZ
& EHRMANN (2019) pointed out that there were no records in western Anatolia at the time, which could have
had provided a migration pathway between the native range of the species in Crimea viz. the eastern shores
of the Black Sea, and the Balkans. It can viably be assumed, therefore, that H. transcaucasica reached and
subsequently spread on the Balkan Peninsula and the islands of the Aegean Sea via human trade routes
(likely via oothecae deposited on cargo).
On the other hand, specimens from urban habitats are more likely to be accidentally recorded than
specimens from natural ecosystems, obscuring the true distribution of the species. Studies on the distribution
and ecology of the species in Crete’s natural habitats are still lacking.
Nomenclatural notes
As pointed out by ZLATKOV et al. (2020), there has been considerable disagreement in the recent
literature regarding the identity of this species. The considerable morphological variability (in particular
regarding the shapes of stigma and pronotum, and the color of discoidal spines) observed in specimens from
Central Asia (native range of H. tenuidentata) and both native and colonized European areas has led
SCHWARZ et al. (2018) and BATTISTON et al. (2018) to suggest a possible synonymy between H.
tenuidentata and H. transcaucasica. Both were described after females, the types of which are housed in the
MHNG and NHMW, respectively.
However, the two species have never been formally synonymized. Nevertheless, several authors
prematurely accepted the synonymy (BATTISTON et al., 2018, 2019, 2020b; MOULIN, 2020; PINTILIOAIE et
al., 2021; VUJIĆ et al., 2021), while others continued to use H. transcaucasica (CIANFERONI et al., 2018;
VAN DER HEYDEN, 2018c; ROMANOWSKI et al., 2019; SCHWARZ & EHRMANN, 2019; ZLATKOV et al., 2020).
This unfortunate development may lead to the false assumption that we are dealing with two different
species in Europe instead of one. This is all the more worrying since the very similar Hierodula patellifera
(Audinet-Serville, 1838) has also become established in Europe (BATTISTON et al., 2020a; MOULIN, 2020)
and might lead to further confusion.
All three taxa belong to a subset of Hierodula species exhibiting rather uniform genital morphology,
while external morphology is characterized by relatively similar habitus but high intraspecific variability. For
this reason, SCHWARZ et al. (2018) pointed out that the investigation of a considerable number of specimens
from across the whole range of H. tenuidentata and H. transcaucasica is necessary to validate or refute the
synonymy. Until the two species receive a proper taxonomic treatment, we suggest to only use H.
transcaucasica for the European populations. The differences to its East Asian congener H. patellifera,
currently known in Europe from southern France and northern Italy, were worked out by BATTISTON et al.
(2020a).
Acknowledgements
We like to thank José Ramón Correas for useful comments on the manuscript.
References
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The Giant Asian Mantis (Hierodula tenuidentata) is reported for the first time in the Veneto region and the survival of adult individuals in Northern Italy till winter is confirmed. Despite the already well documented ability of this alien species to survive the harsh winters of Northern Italy, new data on its ecology in these areas allows one to better understand the potential of this species to spread in Europe. This species was able to remain viable in persistent minimum temperatures below 0°C and with extremes of -5 °C. The current distribution dynamics suggest Veneto as a possible meeting point between the westward expanding Balkan populations and those of the lower Po Valley already known for some years.
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Hierodula tenuidentata is reported for the first time in Romania, from several places, with strong evidence on the existence of stable populations in this country. Preliminary data regarding the ecology of the species and its relations with indigenous fauna are presented, as well as differences between this species and its closest congeners from Romania and Europe.
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An alien mantis species, Hierodula tenuidentata, has expanded its areal in Europe in the last few years. It was registered in several European countries, including some on the Balkan Peninsula. Herein are presented for the first time records of H. tenuidentata on the territory of Serbia, several from the capital city of Belgrade, and one from Novi Kneževac, a town in the northern part of country. The species was recognized for the first time in photos posted in a Facebook group of insect lovers and enthusiasts in 2019, but positive identification only took place in 2020 after group members collected specimens. Social networks can be a very important tool for the monitoring of invasive species in general, and invasive mantids in particular, especially due to their size and attractiveness.
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Originally from Asia, Hierodula patellifera (Serville, 1839) occurs several Mediterranean countries, such as Italy. These arrivals could come from many factors: new pets or commercial human transport. The presence of Hierodula patellifera (Serville, 1839) is here reported for the first time in France. A well settled and probably widespread population of this species is here discussed as its adaptability to the Mediterranean climate. Some considerations on the potential impacts on the local ecosystems and its future spreading in Europe as an invasive species are given.
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Hierodula transcaucasica Brunner von Wattenwyl, 1878 is a mantid native to the Caucasus and southern and western Asia. Here we present the first records of the species for the Balkans represented by findings of the species in Northern Greece in two localities (Thasos in 2015 and Charopo in 2017) and in Bulgaria in 2017. These recent findings implicate the potential of the species to colonize larger area of the Balkan Peninsula. The adaptability of H. transcaucasica to different environments and ecological corridors may facilitate the spread of this species in the Balkans and other Euro-Mediterranean countries. The key to the species of the subfamily Mantinae of the Mediterranean Basin is provided.
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This paper summarizes the current knowledge about invasive praying mantids (Mantodea) in Europe. The continent has been spared invasive mantodeans for a long time, but reports on allochthonous populations increased recently. We differentiate between native species increasing their range, true neobiota and single specimens which failed to establish viable populations. The first group is represented by Ameles spallanzania (Rossi, 1792), Mantis religiosa Linnaeus, 1758, Sphodromantis viridis vischeri (Werner, 1933), and Hierodula transcaucasica Brunner von Wattenwyl, 1878. The latter is now established on the Balkan Peninsula. True neobiota are the Nearctic Brunneria borealis Scudder, 1896 in southern Spain, the afrotropical Miomantis caffra Saussure, 1871 and M. paykullii Stål, 1871 in Portugal, as well as Tenodera sinensis Saussure, 1871 in Germany. Single in-dividuals or oothecae of the following species have also been recorded: Iris oratoria (Linnaeus, 1758), Hymenopus coronatus (Olivier, 1792), Empusa pennata (Thunberg, 1815), Idolomantis diabolica (Saussure, 1869), Tenodera sinensis Saussure, 1871, Hierodula sp., and Stagmomantis carolina (Johansson, 1763). The immigration pathways and potential risks for the native mantodean fauna are discussed.
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Hierodula transcaucasica Brunner von Wattenwyl, 1878 is recorded from Skopelos island (Sporades), giving the first exact locality of this species for Greece. Moreover new records based on photos of Hierodula sp. for Kalmykia (Russia), Ukraine, and Greece and the first ones for Macedonia and Italy are given. The problem of the specific attribution of these specimens through photos and the question of eventual allochthony of one or more species of Hierodula in Europe are discussed.
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The checklist of the praying mantids (Mantodea) occurring in Nepal is updated after the study of new and re-evaluation of previously studied material. Nine taxa are removed from the checklist: Amantis bolivarii Giglio-Tos, 1915, Deiphobe infuscata (Saussure, 1870), Creobroter apicalis Saussure, 1869, Creobroter urbanus (Fabricius, 1775), Hierodula transcaucasica Brunner de Wattenwyl, 1878, Humbertiella indica Saussure, 1869, Mantis religiosa religiosa Linnaeus, 1758, Memantis fuliginosa (Thunberg, 1815), and Memantis gardneri Werner, 1931. Three species are new for Nepal: Amantis fumosa (Giglio-Tos, 1915), Creobroter laevicollis, and Gonypeta humbertiana Saussure, 1869. Five species, namely A. bolivarii, C. apicalis, D. infuscata, Euthyphleps rectivenis Wood-Mason, 1889, and M. gardneri, have records from neighbouring states of India and possibly occur in Nepal but have escaped detection so far. The genus Rhombodera Burmeister, 1838 is removed from the Nepalese checklist, as its only species previously recorded from the country is moved to Rhombomantis Ehrmann & Borer, 2015 as Rhombomantis tectiformis (Saussure, 1870) (comb. nov.). There are 17 additional taxonomic changes. Creobroter elongatus Beier, 1929 is a new synonym of C. apicalis (Saussure, 1869) (syn. nov.). Bolivaria moseri (Saussure, 1871) is removed from synonymy with Bolivaria xanthoptera (Olivier, 1792), which is itself transferred back to the genus Deiphobe Stål, 1877 (stat. rev. and comb. rev.). Deiphobe yunnanensis Tinkham, 1937 is synonymized with D. mesomelas (Olivier, 1792) (syn. nov.). Deiphobe incisa Werner, 1933 is synonymized with D. mesomelas (Olivier, 1792) (syn. nov.), but the male and female paratypes of D. incisa are assigned to D. infuscata (Saussure, 1870) and D. brunneri (Saussure, 1871), respectively. Ephestiasula pictipes (Wood-Mason, 1879), E. amoena (Bolivar, 1897), and E. intermedia Werner, 1930 are all synonymized with Pachymantis rogenhoferi Saussure, 1872 (syn. nov.), which is transferred to the genus Ephestiasula as E. rogenhoferi (Saussure, 1872) (comb. nov.). Gonypeta humbertiana is removed from synonymy with G. punctata De Haan, 1842. Gonypeta punctata lateralis Werner, 1933 is a new synonym of G. humbertiana (syn. nov.). Paula micra Liana, 2009 is removed from synonymy with Gonypetyllis semuncialis Wood-Mason, 1891 and reinstated as Gonypetyllis micra (Liana, 2009) (stat. rev. & comb. nov.). Mantis orientalis Saussure, 1870 is removed from synonymy with Statilia nemoralis (Saussure, 1870) and placed into synonymy of Statilia maculata (Thunberg, 1784) (syn. nov.). Mantis nobilis Brunner de Wattenwyl, 1893 is removed from synonymy with Statilia nemoralis and reinstated as Statilia nobilis (Brunner de Wattenwyl, 1893) (stat. rev.). Mantis indica Mukherjee, 1995 is removed from synonymy with Statilia nemoralis (Saussure, 1870) and placed into synonymy with Statilia nobilis (Brunner de Wattenwyl, 1893) (syn. nov.). The revised Nepalese checklist now contains 42 species, and will likely be supplemented by additional species with increased sampling efforts.
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The presence of the Giant Asian Mantis, Hierodula tenuidentata Saussure, 1869 (Insecta Mantodea), is here reported for the first time in Italy. A well settled and probably wide spread population of this species is here described in detail, evaluating the increase in the number of individuals in the recent years and its adaptability to the European continental climate and to anthropized environments. The synonymy of this species with H. transcaucasica Brunner von Wattenwyl, 1878 is here discussed and some considerations on the potential impacts on the local ecosystems and its future spreading in Europe as an invasive species are here given.
SimpleMappr, an online tool to produce publication-quality point maps
  • D P Shorthouse
SHORTHOUSE, D. P. (2010). SimpleMappr, an online tool to produce publication-quality point maps. Available from: http://www.simplemappr.net. [Accessed on 19-VII-2021].