The Afrotropical Miomantis caffra Saussure 1871 and M. paykullii Stal 1871: first records of alien mantid species in Portugal and Europe, with an updated checklist of Mantodea in Portugal (Insecta: Mantodea)

Eduardo Marabuto

doi:10.3897/BDJ.2.e4117

Biodiversity Data Journal : Taxonomic paper

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Taxonomic paper

The Afrotropical Miomantis caffra Saussure 1871 and M. paykullii Stal 1871: first records of alien mantid species in Portugal and Europe, with an updated checklist of Mantodea in Portugal (Insecta: Mantodea)

Eduardo Marabuto ^{‡,
§}

‡ Computational Biology and Population Genomics Group, Centro de Biologia Ambiental, Faculdade de Ciências, Universidade de Lisboa, Campo Grande, 1749-016, Lisboa, Portugal

§ Centro de Estudos do Ambiente e do Mar, Departamento de Biologia, Universidade de Aveiro, Portugal, Aveiro, Portugal

Corresponding author:

Academic editor: Edward Baker

Received: 28 Sep 2014 | Accepted: 30 Oct 2014 | Published: 12 Nov 2014

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

Citation: Marabuto E (2014) The Afrotropical Miomantis caffra Saussure 1871 and M. paykullii Stal 1871: first records of alien mantid species in Portugal and Europe, with an updated checklist of Mantodea in Portugal (Insecta: Mantodea). Biodiversity Data Journal 2: e4117. https://doi.org/10.3897/BDJ.2.e4117

Abstract

The recent growing interest on the Mantodea fauna of southern Europe and Portugal in particular, has enabled the discovery of two geographically separated populations of hitherto unknown species in Europe. Analysis of specimens shows that they belong to two Afrotropical mantids: Miomantis caffra Saussure, 1871 and Miomantis paykullii Stal, 1871, thus raising the number of known species in Europe to 39 and in Portugal to 11.

While these are remarkable findings, they also represent the first alien mantis species recorded from this continent. As yet, these species appear to be confined to artificial humanised gardened areas but call for more attention to the problem of biological invasions and the need for better bio-security measures for the conservation of natural ecosystems.

In the absence of recent revisionary work on the Mantodea of Portugal and given the need to provide an accessible identification tool, both a checklist and a key to species are provided for all species in the country.

Keywords

New records, Western Europe, Biological invasion, biotic homogenisation, mantis, key to species

Introduction

Human impact on biological diversity can occur at different levels and magnitudes. The effect on ecosystems can be direct, by their removal or alteration, or manifest themselves indirectly such as through climate change or the translocation of species between different biogeographic regions. Generally, due to an incomplete knowledge of ecosystems by researchers and policy-makers, many introduction cases may be overlooked, even more rarely tracked down precisely as they happen. Therefore, their effect has mostly been under-evaluated. Competition with natives for resources and other environmental constraints limits the successful establishment of most alien species (Elton 1958), and if they do establish there is a chance the effect is not visible right away, especially if crops are not involved.

The term "alien species" generally includes those naturally exogenous to a given habitat, ecosystem or biogeographical area which have established themselves outside their natural range either unaided or with human assistance. Several paths and vectors for introduction of such species have been identified (e.g. Hulme et al. 2008). These include deliberate or unintentional ways, the prevalence of which differs according to the biological group. Introductions, particularly if mediated by man are fundamentally different from natural range expansions both quantitatively and qualitatively. In contrast to natural dispersal, which usually involves highly mobile organisms with a broad ecological plasticity, these may include species with reduced ability to overcome important physiological barriers (e. g. slugs or amphibians arriving on oceanic islands) or have distant origins (e. g. the introduction of reindeer in sub-Antarctic islands) (Wilson et al. 2009).

In Europe there are over 1500 established alien arthropod species, of which 1390 are insects (Roques et al. 2009). According to the DAISIE database, the largest joint project to identify alien species in the European territory (DAISIE 2009 and www.europe-aliens.org), the major pathway of entrance and establishment of species in the territory is unintentional direct introduction (1341, 86%). Of these, introduction with horticultural and ornamental items accounts to 468 species (29%). 'Only' 218 species have been supposedly intentionally introduced into the European territory, mostly as biological control agents (Rabitsch 2010). It is estimated that 24.2% have an economic impact (Vilà et al. 2010) and that the majority of aliens settle preferably in humanised, altered habitats (65%, 1040 species), i.-e. that depart from the natural local assemblages (Lopez-Vaamonde et al. 2010). Parks and gardens (31.4%, 500 species), houses and built areas (31%, 493 species) and then agricultural land (29.7% or 472 species) are the preferred allochthonous habitats.

While mantises have so far not been reported as alien species in Europe, the worldwide scenario is quite different. From as early as 1899, Mantis religiosa L. was introduced into USA and Canada with nursery plants (McLeod 1962, Cannings 2007), alongside the Oriental Tenodera sinensis (Helfer 1987, Hurd 1999). Iris oratoria (Linnaeus, 1758), another Euro-Mediterranean species arrived in California during the thirties (Maxwell and Eitan 1998) from where it seems to be spreading. Finally, the south African Miomantis caffra has been established in New Zealand for almost forty years (Ramsay 1984).

The European Mantodea species list (Heller and Bohn 2013) includes around 38 species belonging to four distinct families (Amorphoscelidae, Empusidae, Mantidae and Tarachodidae). Mantidae is by far the richest family with 30 species (Battiston et al. 2010) and all but the genus Ameles are of straightforward identification. As for now, Ameles is still under scientific scrutinity as authors debate the status of several taxa (Agabiti et al. 2010, Battiston and Fontana 2005, Battiston et al. 2010, Obertegger and Agabiti 2012, Wieland et al. 2014). In the meantime, Fauna Europeaea (Heller and Bohn 2013) seems not to be up to date with recent findings and the number of Ameles spp. in the region may be around eight (A. assoi, A. decolor, A. fasciipennis, A. heldreichi, A. insularis, A. paradecolor, A. picteti and A. spallanzania).

Irrespective of the criteria for the definition of species and biogeographical structure, the most up-to-date and synthetic overall work on Euro-Mediterranean Mantodea, Battiston et al. 2010, gave 127 species from a wide area from the Canary Islands to the Caucasus, all Europe, North Africa and Mediterranean islands. Diverse and heterogeneous landscapes, different land-uses, rugged topography and the character of refugia-expansion cycles and contact zones during and after past glaciations allow for an interesting biological diversity in this crossroad between three different continents. However, there is a great south-north disparity and because of their typical thermophilous profile, the Mantodea attain highest diversity in warmer areas of the southern Mediterranean. On the other hand, the aridity and lack of many suitable habitats renders the Sahara desert unsuitable and an effective barrier to interchangeability of biota with the Afrotropical region. The only exception is the corridor represented by the Nile river valley, promoting the contact and migration of species from deep in east Africa to the Mediterranean. Therefore, Egypt has the greatest diversity of mantis species in the region, some of which are clearly of Afrotropical origin like the genera Heterochaeta and Miomantis.

In the summer of 2014, unexpected findings of two mantis species in Portugal triggered a revision of the species occurring in the country, alongside the investigation of their identity. In fact, the specimens studied do not key to any of the known European species of Mantodea. A combination of characters such as complete development of wings, simple cerci, no foliaceous leg extensions, small size and an elongated pronotum (in contrast to all the Ameles spp.) among others place the available specimens in the genus Miomantis.

Miomantis Saussure, 1870 currently includes 70 Afrotropical species (Ehrmann 2002) and had not yet been recorded in Europe. The only available key for the genus is that of Giglio-Tos (1927) but this is largely based on morphological characters not necessarily relevant for phylogenetic species delimitation or are based on coloration and isometric scaling. M. paykullii Stal, 1871 is the most widespread species in the genus, almost ubiquitous south of the Sahara and present in Egypt and Mauritania (Ehrmann 2002, Battiston et al. 2010). This is the species found in Portugal first although followed closely by a second observation of a conspecific individual suggesting that there is an established population. Almost a month after this finding, five other males were independently located 200km to the north of the first records. These key out to the same genus but the slightly different morphology places them in the closely related species, Miomantis caffra Saussure, 1871 which is found naturally in South Africa and as an alien invasive in New Zealand (Ramsay 1984, Ramsay 1990). Both species, the genus Miomantis and tribe Miomantini are faunistic novelites to Portugal and the European continent.

Materials and methods

The first specimen of M. paykullii was collected by hand at a garden light around which it was flying. A second specimen was observed 3 days later in the same place by beating the vegetation (Arbutus unedo and Phyllostachys aurea bamboo hedge) but flew off to an inaccessible area. The third and fifth to seventh specimens, belonging to M. caffra were attracted to a porch light during the night and intercepted hunting insects attracted by it. The fourth specimen, also M. caffra, was located in a different locality. All six collected specimens of Miomantis spp. were maintained alive till further analysis. Later, these were prepared and mounted dry, remaining in the author's personal reference collection. The captured specimen of M. paykullii and the first of M. caffra were photographed by the author.

Taxon treatments

Miomantis paykullii Stal, 1871

Materials Download as CSV

taxonRank:
species
; vernacularName:
Egyptian pygmy mantis
; genus:
Miomantis
; specificEpithet:
paykullii
; scientificNameAuthorship:
Stal, 1871
; continent:
Europe
; country:
Portugal
; countryCode:
PT
; stateProvince:
Faro
; county:
Loulé
; municipality:
Quarteira
; locality:
Vila Sol
; verbatimElevation:
40
; decimalLatitude:
37.090
; decimalLongitude:
-08.093
; samplingProtocol:
ad hoc observation
; eventDate:
2014-08-05
; habitat:
garden
; individualCount:
1
; sex:
male
; recordedBy:
Eduardo Marabuto
taxonRank:
species
; vernacularName:
Egyptian pygmy mantis
; genus:
Miomantis
; specificEpithet:
paykullii
; scientificNameAuthorship:
Stal, 1871
; continent:
Europe
; country:
Portugal
; countryCode:
PT
; stateProvince:
Faro
; county:
Loulé
; municipality:
Quarteira
; locality:
Vila Sol
; verbatimElevation:
40
; decimalLatitude:
37.090
; decimalLongitude:
-08.093
; samplingProtocol:
vegetation beating
; eventDate:
2014-08-08
; habitat:
garden
; individualCount:
1
; sex:
male
; recordedBy:
Eduardo Marabuto

Description

Adapted from Giglio-Tos 1927, Ehrmann 2002 and Agabiti et al. 2010: Small or medium sized species. Green or light-brown coloured, patternless except occasional obscured humeral vein. Head broader than the pronotum, especially in the male. Eye prominent, slightly conical, more apparent in male but not ending in a spine. Pronotum slender with weakly developed supracoxal dilation, as long as or longer than fore coxa and smooth in male, finely toothed in female. Forewing hyaline in male, more opaque in female, distally dilated and slightly exceeding the tip of the abdomen in the former, only reaching the base of the cerci in the latter. Male hindwing hyaline, female yellowish, crossed by yellow veins. Supra-anal plate longer than broad, triangular. Foreleg unpatterned, coxa finely toothed with 5-6 spaced small spines stronger in female. Femur with 4 discoidal spines and 4 external. Fore tibia with 7 external spines. Body length: 36-39mm; pronotum length: 11-12mm in male, to 14mm in female; forewing length 23mm in male, 19-21mm in female.

Portuguese specimens:

Both observed specimens are adult-stage males conforming well with the descriptions of the species available (Ehrmann 2002, Battiston et al. 2010). The first is illustrated in Fig. 1. This specimen is a straw-coloured and patternless mantis with a darker forewing radial vein. Both forewings and hindwings are hyaline, forewings being slightly less transparent and brownish. Head is short with prominent conical eyes. The raptorial forelegs bear the typical spine scheme of the genus and species: 5-6, 4, 7. Biometrics: pronotum length: 10mm; whole body length: 38mm; forewing length: 26mm; fore-leg length measurements - coxa: 6.6mm, femur: 8.2mm, tibia: 4.7mm; ratio forewing/ pronotum length: 2.6.

Figure 1.

Miomantis paykullii, observed and photographed in Algarve, Portugal.

a: Dorso-lateral habitus view. Note hyaline forewings with obscured radial vein.
b: Dorsal view. Note the smooth lateral side of relatively long pronotum, straight frons, simple cerci and hyaline wings.
c: Head frontal view. Note the conical, prominent and striped eyes.
d: Detail of foreleg. Characteristic are the coxa bearing very small spines (5-6), femora crenulate between the 4 external spines and tibiae with 7 external spines.

Distribution

An Afrotropical species cited throughout the biogeographical area, with some island populations. Countries where it has been found are: Burkina Faso, Cameroon, Chad, Egypt, Ghana, Israel, Ivory Coast, Kenya, Mauritania, Mauritius, Mozambique, Niger, Senegal, Togo, Uganda and Zimbabwe (Ehrmann 2002, Battiston et al. 2010).

Recorded habitats and distribution in Portugal:

The area is a private property garden of the resort Vila Sol where many exotic plant species are planted. Among them, hedges of Phyllostachys aurea and Lantana camara are typical while stands of Pennisetum alopecuroides, Cyperus papyrus and garden turf are widespread with smaller amounts of other exotic species. The only natives are isolated Pinus pinea trees, remnants from the open woodland pre-resort and a hedge of Arbutus unedo. At dusk, several garden lights at ground level are regularly lit and during the night, an automatic irrigation system maintains humidity levels high, even during the summer. Prior to the installment of the resort, the whole area was a dry thermomediterranean open Pinus pinea woodland with mild winters and warm summers on an arenite substrate. The biogeographic province is the Gaditano-Onubo-Algarviense in its Algarviense coastal sector with some notable endemics (Rivas-Martinez et al. 1990, Costa et al. 1999) and seriously under threat from urban development. This particular site is one already lost as a natural habitat and is home to only the most resilient and human-adapted species.

Ecology

Ecological plasticity in this species over a broad temperature range accounts for its wide distribution. According to Prete et al. (1999), in Ghana M. paykullii mostly inhabits grasslands, the colour of which (a proxy for humidity levels) determines the final coloration and proportion between brown and green morphs. Presumably, a brown morph develops in a less moist environment. In Ghana, adults display no deimatic behaviour and attempt to fly or walk away from danger (Edmunds 1972). Adults are also presumably very mobile and active at night and especially sensitive to bat echolocation high frequency sounds (80-100 KHz), thus being able to evade predation (Prete et al. 1999). This phenomenon has been experimentally tested with the closely related M. natalica Beier, 1930 by Cumming (1996).

Conservation

Showing a wide distribution centred in the subsaharan African continent, M. paykullii should not be at risk of any kind. However, in the Euro-Mediterranean area, where it has only been found along the Nile valley and nearby areas of Israel, this species has recently been evaluated as at "Potential risk", because of sparse observations for a long time (Agabiti et al. 2010).