Research Article |
Corresponding author: Dávid Selnekovič ( david.selnekovic@uniba.sk ) Academic editor: Pavel Stoev
© 2023 Dávid Selnekovič, Katarína Goffová, Ján Šoltýs, Eva Kováčová, Ján Kodada.
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:
Selnekovič D, Goffová K, Šoltýs J, Kováčová E, Kodada J (2023) Mordellistena platypoda, a new species of tumbling flower beetle from the island of Ischia in Italy (Coleoptera, Mordellidae). ZooKeys 1148: 41-63. https://doi.org/10.3897/zookeys.1148.86845
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Mordellistena A. Costa, 1854, the most species-rich genus of tumbling flower beetles comprises more than 800 species worldwide and more than 150 reported from Europe. Here, a new species Mordellistena (s. str.) platypoda is described from the island of Ischia in Italy. The species hypothesis is based primarily on morphological characters which are visualised using scanning electron microscopy images, high-resolution photographs, and drawings. The species hypothesis is supported by analysis of a 658 bp fragment of cytochrome c oxidase subunit I (COI). Divergences in the COI gene are evaluated using maximum likelihood and Bayesian inference analyses. The species delimitation is assessed using Assemble Species by Automatic Partitioning (ASAP) and Poisson Tree Processes (PTP) methods. Genetic distances are visualised using multidimensional scaling. Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov. is recovered as a well-separated species by both molecular and morphological analyses. Our results show that M. platypoda Selnekovič, Goffová & Kodada, sp. nov. is most closely related to M. tarsata Mulsant, 1856, although the two species differ significantly in vestiture colouration, presence of lateral ctenidia on the third metatarsomere, and presence of sexual dimorphism on the protibia. The results indicate that such morphological differences, which were traditionally used to distinguish between species groups, may in fact be present between closely related species. Interestingly, examination of the numerous museum material did not reveal additional specimens of the new species, and therefore M. platypoda Selnekovič, Goffová & Kodada, sp. nov. is currently known only from the Italian island of Ischia.
Cytochrome c oxidase subunit I (COI), DNA barcoding, integrative taxonomy, morphology, species delimitation
More than 800 species are currently classified within Mordellistena A. Costa, 1854, making it the most species-rich genus of tumbling flower beetles. It is reported from every continent except Antarctica. However, the generic placement of many species, especially Indomalayan and Neotropical, needs to be reassessed according to the current generic classification. In Europe, the genus is represented by more than 150 species (
A recent collecting trip to the island of Ischia in Italy in June 2019 yielded more than 1,000 Mordellidae specimens representing 12 species. Within this material, we recognised a series of 52 specimens belonging to the new species described herein, Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov. (Fig.
This study is based on the examination of more than 400 specimens of the genus Mordellistena deposited at the following institutions:
MNHU Museum für Naturkunde der Humboldt Universität, Berlin, Germany;
SNSD Senckenberg Naturhistorische Sammlungen, Dresden, Germany.
The type series of M. platypoda consist of 52 specimens collected on the island of Ischia in Italy. The specimens of the additional species included in the phylogenetic analyses were acquired from several localities in Italy, Slovakia, Spain, Cyprus, and Bulgaria (Table
Specimen ID | GenBank | Locality | Coordinates |
---|---|---|---|
Mordellistena austriaca DSBS 60 | OM680976 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena confinis DSBS 243 | OP586774 | Italy, Firenze, Cinipetta | 43.570000°N, 11.420556°E |
Mordellistena confinis DSBS 244 | OP586776 | Italy, Firenze, Cinipetta | 43.570000°N, 11.420556°E |
Mordellistena confinis DSBS 245 | OP586769 | Italy, Firenze, Cinipetta | 43.570000°N, 11.420556°E |
Mordellistena confinis DSBS 250 | OP586770 | Italy, Firenze, Cinipetta | 43.570000°N, 11.420556°E |
Mordellistena confinis DSBS 329 | OP586775 | Italy, Firenze, Cinipetta | 43.570000°N, 11.420556°E |
Mordellistena hirtipes DSBS 205 | OM681007 | Cyprus, Limassol env. | 34.755278°N, 33.093333°E |
Mordellistena hirtipes DSBS 207 | OM681008 | Cyprus, Limassol env. | 34.755278°N, 33.093333°E |
Mordellistena hirtipes DSBS 208 | OM681009 | Cyprus, Limassol env. | 34.755278°N, 33.093333°E |
Mordellistena lindbergi DSBS 144 | OM680979 | Cyprus, Limassol env. | 34.755278°N, 33.093333°E |
Mordellistena lindbergi DSBS 270 | OP586772 | Cyprus, Akamas | 35.057586°N, 32.345527°E |
Mordellistena lindbergi DSBS 271 | OP586767 | Cyprus, Akamas | 35.057586°N, 32.345527°E |
Mordellistena lindbergi DSBS 280 | OP586771 | Cyprus, Foinikaria env. | 34.766272°N, 33.100258°E |
Mordellistena minima DSBS 79 | MT232550 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena minima DSBS 172 | OM680982 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena minima DSBS 173 | OM680983 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena minima DSBS 215 | OP586765 | Italy, Sardinia, Castiadas | 39.206812°N, 9.562685°E |
Mordellistena minima DSBS 218 | OP586768 | Italy, Sardinia, Castiadas | 39.206812°N, 9.562685°E |
Mordellistena minima DSBS 219 | OP586766 | Italy, Sardinia, Castiadas | 39.206812°N, 9.562685°E |
Mordellistena platypoda DSBS 83 | OM680977 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 118 | OM680978 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 194 | OM680997 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 195 | OM680998 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 199 | OM681002 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 233 | OM681019 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 235 | OM681020 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena platypoda DSBS 237 | OM681021 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena pseudorhenana DSBS 11 | OP586773 | Cyprus, Foinikaria env. | 34.755278°N, 33.093333°E |
Mordellistena pseudorhenana DSBS 12 | MT232533 | Cyprus, Foinikaria env. | 34.755278°N, 33.093333°E |
Mordellistena pseudorhenana DSBS 43 | MT232539 | Slovakia, Chotín env. | 47.806389°N, 18.198056°E |
Mordellistena purpurascens DSBS 82 | MT232552 | Italy, Ischia, Serrara env. | 40.721389°N, 13.883056°E |
Mordellistena purpurascens DSBS 111 | MT232554 | Spain, Tossa de Mar | 41.736667°N, 2.935000°W |
Mordellistena purpurascens DSBS 117 | MT232555 | Italy, Ischia, Serrara env. | 40.721389°N, 13.883056°E |
Mordellistena purpurascens DSBS 182 | OM680985 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 183 | OM680986 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 186 | OM680989 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 187 | OM680990 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 188 | OM680991 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 192 | OM680995 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 200 | OM681003 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 201 | OM681004 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpurascens DSBS 226 | OM681016 | Italy, Sardinia, Castiadas env. | 39.206812°N, 9.562685°E |
Mordellistena purpurascens DSBS 227 | OM681017 | Italy, Sardinia, Castiadas env. | 39.206812°N, 9.562685°E |
Mordellistena purpurascens DSBS 231 | OM681018 | Italy, Ischia, Serrara env. | 40.716667°N, 13.886389°E |
Mordellistena purpureonigrans DSBS 49 | OP575375 | Slovakia, Chotín env. | 47.806389°N, 18.198056°E |
Mordellistena purpureonigrans DSBS 55 | OM680974 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena purpureonigrans DSBS 56 | OM680975 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena purpureonigrans DSBS 71 | OP575374 | Slovakia, Iža env. | 47.748056°N, 18.260556°E |
Mordellistena purpureonigrans DSBS 73 | OP575376 | Slovakia, Iža env. | 47.748056°N, 18.260556°E |
Mordellistena tarsata DSBS 39 | OM680971 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena tarsata DSBS 41 | OM680972 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena tarsata DSBS 42 | OM680973 | Slovakia, Virt env. | 47.760000°N, 18.340556°E |
Mordellistena tarsata DSBS 209 | OM681010 | Cyprus, Skoulli env. | 34.968056°N, 32.446111°E |
Mordellistena tarsata DSBS 210 | OM681011 | Bulgaria, Melnik env. | 41.510000°N, 23.378333°E |
Mordellistena tarsata DSBS 211 | OM681012 | Bulgaria, Melnik env. | 41.510000°N, 23.378333°E |
Mordellochroa abdominalis DSBS 138 | OM681022 | Slovakia, Burda | 47.847778°N, 18.789722°E |
The specimens selected for DNA isolation were killed in 96.3% ethanol and stored at -20 °C. The remaining specimens were killed with the fumes of ethyl acetate. After DNA isolation, the specimens were soaked in 5% acetic acid, dissected, and mounted on cards. The dissected genitalia were cleared in lactic acid for several days, or in 10% KOH overnight, then dehydrated in 96.3% ethanol and mounted on slides in Euparal (Paradox Co., Cracow, Poland). After examination, genitalia were mounted on the card with the respective specimen using dimethyl hydantoin formaldehyde (Entomopraxis, Barcelona, Spain). Specimens were observed under an MZ16 stereomicroscope (Leica, Wetzlar, Germany) with magnification up to 120× with diffused LED light (6400 K). The drawings were prepared using a drawing tube attached to a DM1000 compound microscope (Leica, Wetzlar, Germany) and subsequently inked with the Isograph technical pens (Rotring, Hamburg, Germany). Photographs of habitus were taken with an EOS 5D mark II camera (Canon, Tokyo, Japan) attached to an Axio Zoom V16 stereoscope (Zeiss, Oberkochen, Germany); photographs of genitalia were taken with an Axio Imager 2 (Zeiss, Oberkochen, Germany). The images were stacked in Zerene Stacker 1.4 software (https://zerenesystems.com/cms/stacker) and edited in Adobe Photoshop CC (https://www.adobe.com/products/photoshop.html) and DxO Photolab 5 (https://www.dxo.com/dxo-photolab/). For scanning electron microscopy, the body parts were disarticulated, cleaned in lactic acid for several days, dehydrated in 96.3% ethanol, coated with 25 nm thick gold layer, and examined using Quanta 250 FEG (FEI Europe B.V., Eindhoven, The Netherlands). Measurements were made with an ocular micrometre in a MZ16 stereomicroscope (Leica, Wetzlar, Germany) and are given in the text as the range followed by the arithmetic mean and standard deviation enclosed in parentheses. The measured characters are abbreviated in the text as follows:
EL elytral length from scutellar apex to elytral apices along suture;
EW maximum elytral width;
HL head length from anterior clypeal margin to occipital carina along midline;
HW maximum head width;
LPrL maximum left paramere length;
PL pronotal length along midline;
PW maximum pronotal width;
PygL maximum pygidial length;
RPrL maximum right paramere length;
TL combination of head, pronotal and elytral lengths.
The species description follows the conventional terminology as used in
In total, 56 specimens were used for DNA isolation. Details on the voucher specimens, including sampling localities and GenBank accession numbers, are presented in Table
The consensus sequences, alignment, and final matrix were produced in Unipro UGENE 44.0 software (http://ugene.net/). Pairwise p-distances were calculated using MEGA X (
We generated and analysed a set of 56 sequences of the 658 bp COI gene fragment. The dataset comprised 237 parsimony-informative sites (36%) and 22 singleton sites (3.34%). We recovered almost identical topologies with both maximum likelihood (ML) and Bayesian inference (BI) methods, and therefore only the ML tree is presented with ML bootstrap values and BI posterior probability values (Fig.
Analyses of a 658 bp fragment of the cytochrome c oxidase subunit I gene (COI) in selected Mordellistena A. Costa, 1854 species A combined results of maximum likelihood (ML) and Bayesian inference (BI) analyses; ML bootstrap values and BI posterior probabilities are shown on ML tree. The vertical bars represent results of Assemble Species by Automatic Partitioning (ASAP) and Poisson Tree Processes (PTP) species delimitation analyses B distribution of uncorrected p-distances C multidimensional scaling of uncorrected p-distances.
The uncorrected p-distances within and between species are summarised in Appendix
Italy, Ischia, Serrara env., 40.72138°N, 13.88305°E; steep slopes with grassland communities, ca. 550 m alt. (Fig.
Holotype : italy • male; Ischia, Serrara env.; 40.72138°N, 13.88305°E; ca. 550 m alt.; 30 Jun 2019; D. Selnekovič leg.; steep slopes with grassland communities, on inflorescences of Apiaceae; GenBank: OM680978; SNSD. Paratypes: italy • 9 males, 8 females; same data as for holotype • 23 males, 11 females; Ischia, Serrara env.; 40.71666°N, 13.88638°E; 517 m alt.; 29 Jun 2019; D. Selnekovič leg.; ruderal habitat on road verge, on inflorescences of Daucus carota; GenBank: OM680977, OM680997, OM680998, OM681002, OM681019 to OM681021; SNSD.
Mordellistena platypoda is included in the M. micans species group as defined by
Scanning electron microscope images of diagnostic characters A Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov., male maxilla B M. platypoda, female maxilla C M. purpurascens A. Costa, 1854, male maxilla D M. platypoda, male antenna (basal part) E M. platypoda, male antenna (apical part).
Scanning electron microscope images of diagnostic characters A Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov., male hind leg, lateral view B M. platypoda, male protibia C M. purpurascens A. Costa, 1854, male protibia D M. platypoda, male protarsus E M. platypoda, female protarsus F M. platypoda, mesal portion of male right elytron G M. platypoda, left paramere, mesal view, white triangle points to a cluster of sensilla campaniformia shown in image I; H M. platypoda, right paramere, mesal view I M. platypoda, cluster of sensilla campaniformia on left paramere. Abbreviation: sc = sensillum campaniformium.
A Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov., male protarsus B M. platypoda, female protarsus C M. purpurascens A. Costa, 1854, male protarsus D M. platypoda, phallobase E M. platypoda male sternite VIII F M. platypoda, male tergite VIII G M. platypoda, male sternite IX H M. platypoda, male tergites IX and X I M. platypoda, female sternite VIII J M. platypoda, female tergite VIII K M. platypoda, penis. Scale bars: 0.1 mm.
Mordellistena platypoda most closely resembles a sympatric species M. purpurascens but differs in paler vestiture, weakly expanded male second maxillary palpomere, with few long setae (Fig.
The results of the COI gene analyses show the close relationship of M. platypoda with M. tarsata Mulsant, 1856, with p-distances of 8.21–8.97%. The two species are easily distinguished by the colouration of the dorsal vestiture, which is black with a greenish lustre in M. tarsata compared to yellowish in M. platypoda (Fig.
Measurements (in mm; ♂♂ n = 33, ♀♀ n = 21): TL: ♂♂ 4.58–5.64 (5.25 ± 0.26), ♀♀ 4.84–6.02 (5.56 ± 0.31); HL: ♂♂ 0.81–1.01 (0.92 ± 0.05), ♀♀ 0.84–1.06 (0.97 ± 0.06); HW: ♂♂ 0.94–1.17 (1.07 ± 0.05), ♀♀ 0.95–1.19 (1.10 ± 0.06); PL: ♂♂ 0.98–1.25 (1.14 ± 0.06), ♀♀ 1.06–1.33 (1.22 ± 0.08); PW: ♂♂ 1.10–1.46 (1.32 ± 0.08), ♀♀ 1.19–1.57 (1.42 ± 0.10); EL: ♂♂ 2.75–3.50 (3.18 ± 0.18), ♀♀ 2.92–3.67 (3.37 ± 0.19); EW: ♂♂ 1.13–1.42 (1.31 ± 0.07), ♀♀ 1.25–1.54 (1.45 ± 0.08); RPrL: 0.27–0.32 (0.29–0.01); LPrL: 0.36–0.43 (0.40 ± 0.02).
Body elongated, wedge-shaped, widest in anterior half of elytra (Fig.
Head large, transverse, moderately convex dorsally, with highest point behind middle of eyes (lateral aspect), HW/HL ratio: ♂♂ 1.09–1.23 (1.16 ± 0.03), ♀♀1.01–1.23 (1.13 ± 0.05); occipital carina convex; integument weakly microreticulate, weakly iridescent, with small round setiferous punctures. Eyes broadly oval, vertical diameter ca. 1.3× horizontal diameter; posteriorly reaching to occipital margin; finely faceted; interfacetal setae longer than facet diameter. Anterior clypeal edge weakly convex. Labrum transverse, densely setose, anterior edge weakly convex. Antenna weakly serrate (Fig.
Pronotum 1.1–1.2× as wide as long, widest behind middle, moderately convex; surface microreticulate, densely covered with lanceolate setae, punctures larger than those on head; anterior edge convex in middle, anterolateral angles broadly rounded; lateral carinae sinuate in lateral aspect; posterior edge sinuate, posterolateral angles rectangular in lateral aspect. Scutellar shield triangular, densely setose. Elytra widest between first and second quarter, EL/EW ratio: ♂♂ 2.31–2.64 (2.43 ± 0.07), ♀♀ 2.22–2.40 (2.32 ± 0.05); apices separately rounded; surface microreticulate, densely covered with decumbent lanceolate setae, punctures coarser than those on pronotum. Hindwing as in Fig.
Habitats of Mordellistena platypoda Selnekovič, Goffová & Kodada, sp. nov. near Serrara village, Ischia, Italy A, B type locality, slopes with Mediterranean grassland communities (40.72138°N, 13.88305°E) C, D ruderal communities with Daucus carota Linnaeus along road (40.71666°N, 13.88638°E).
Abdominal ventrite 1 longer than ventrite 2; ventrite 5 with arcuate apical edge. Pygidium long, conical, narrowly truncate at apex, EL/PygL ratio: ♂♂ 1.75–2.03 (1.87 ± 0.06), ♀♀ 2.12–2.35 (2.21 ± 0.06). Male tergite VIII deeply emarginate on posterior edge, setose apically (Fig.
Females are on average slightly larger than males. Males are more slender than females (Fig.
Partial COI gene sequences of holotype and eight paratypes were submitted to GenBank (https://www.ncbi.nlm.nih.gov/genbank/). The accession numbers are listed in Table
The specific epithet is derived from the Greek words πλατύς (platýs), meaning wide, broad, and πόδι (pódi) meaning foot. It refers to the expanded pro- and mesotarsi, an unusual condition that separates M. platypoda from morphologically similar congeners.
The species is known only from the island of Ischia in Italy.
Mordellistena platypoda was sampled in a series of 52 specimens on 29–30 June 2019. The sampling was carried out at two nearby localities (approximately 600 m apart) near Serrara village. The type locality (40.72138°N, 13.88305°E) was characterised by the steep rocky slopes with Mediterranean grassland communities (Fig.
Mordellistena platypoda is morphologically well defined. It is included in the M. micans species group defined by
The morphology-based species hypothesis was evaluated by analysing a 658 bp fragment of the COI gene. This standard DNA barcoding marker has frequently been used in the taxonomy of beetles not only to identify species as originally proposed (
The distribution of pairwise genetic distances (p-distances) shows a distinct gap between the highest intraspecific divergence (1.06%) and the lowest interspecific divergence (8.21%) (Fig.
Mordellistena platypoda was collected in a relatively large series of 52 specimens during one collecting event on the island of Ischia in Italy. Revision of the museum specimens identified as M. micans (Germar, 1817), M. stenidea Mulsant, 1856, and M. grisea Mulsant, 1856 in the Franciscolo collection in
The authors thank Győző Szél (
Uncorrected p-distances between and within the analysed species calculated in Mega X software. Intraspecific divergences are highlighted in bold.
M. austriaca | M. confinis | M. hirtipes | M. lindbergi | M. minima | M. platypoda | M. pseudorhenana | M. purpurascens | M. purpureonigrans | M. tarsata | M. abdominalis | |
---|---|---|---|---|---|---|---|---|---|---|---|
Mordellistena austriaca | N/A | ||||||||||
M. confinis | 0.1505 | 0.0000 | |||||||||
M. hirtipes | 0.1581–0.1596 | 0.1581–0.1596 | 0.0000–0.0015 | ||||||||
M. lindbergi | 0.1672–0.1687 | 0.1702–0.1733 | 0.1550–0.1581 | 0.0000–0.0030 | |||||||
M. minima | 0.1565–0.1596 | 0.1581–0.1596 | 0.1884–0.1915 | 0.1474–0.1505 | 0.0000–0.0046 | ||||||
M. platypoda | 0.1489 | 0.1687 | 0.1535–0.1550 | 0.1717–0.1733 | 0.1763–0.1793 | 0.0000 | |||||
M. pseudorhenana | 0.1565 | 0.1839 | 0.1626–0.1672 | 0.1702–0.1748 | 0.1733–0.1748 | 0.1611 | 0.0000–0.0030 | ||||
M. purpurascens | 0.1641–0.1657 | 0.1581–0.1596 | 0.1170–0.1201 | 0.1657–0.1687 | 0.1793–0.1824 | 0.1672–0.1687 | 0.1733–0.1748 | 0.0000–0.0046 | |||
M. purpureonigrans | 0.1489–0.1535 | 0.1626–0.1672 | 0.1505–0.1550 | 0.1626–0.1702 | 0.1641–0.1687 | 0.1778–0.1824 | 0.1702–0.1733 | 0.1535–0.1581 | 0.0000–0.0046 | ||
M. tarsata | 0.1398–0.1474 | 0.1824–0.1900 | 0.1657–0.1702 | 0.1915–0.1976 | 0.1854–0.1945 | 0.0821–0.0897 | 0.1596–0.1657 | 0.1763–0.1793 | 0.1809–0.1869 | 0.0000–0.0106 | |
M. abdominalis | 0.1991 | 0.2112 | 0.2158–0.2173 | 0.2295–0.2325 | 0.2036 | 0.2158 | 0.2036 | 0.2264–0.2280 | 0.2112–0.2143 | 0.2112–0.2128 | N/A |