Monograph |
Corresponding author: Xiaoshuan Bai ( baixs2007@aliyun.com ) Corresponding author: Ernst Heiss ( aradus@aon.at ) Academic editor: Guanyang Zhang
© 2018 Xiaoshuan Bai, Ernst Heiss, Wanzhi Cai.
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:
Bai X, Heiss E, Cai W (2018) The Oriental flat bug genus Libiocoris Kormilev, 1957 revisited: re-examination, synonymy, and description of a new genus (Heteroptera, Aradidae). ZooKeys 789: 115-137. https://doi.org/10.3897/zookeys.789.26165
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Re-examination of type specimens of Libiocoris
Aradidae , Carventinae , China, Heteroptera , Libiocoris , new combination, new genus, new species, new synonymy, Paralibiocoris
The genus Libiocoris was erected by Kormilev, 1957 for the species poecilus from New Guinea. Usinger & Matsuda, 1959 improved the generic description based on the new species L. antennatus from Papua (New Guinea) and added L. angulatus, also from New Guinea to this genus. Later, L. lobatus Kormilev, 1968 and L. pilicornis Kormilev, 1972 were described again from New Guinea.
The distribution pattern of these eight species within the Indo-Pacific region seems restricted to two biogeographically different areas: the Indo-China region (L. indicus, L. heissi, and L. sinensis) and the Papuasian region (L. poecilus, L. antennatus, L. angulatus, L. lobatus, and L. pilicornis), which raises questions about their assignment to the same genus of apterous Carventinae, all having very limited distribution ranges.
As a result of the re-examination of the genus-type species L. poecilus (holotype male, allotype female, HMHN), of L. angulatus (holotype female,
1 After removal of the waxy incrustation obscuring the dorsal structures and examination of the female holotypes of heissi and sinensis it was evident that both belong to the same taxon and are synonyms.
2 The Chinese species heissi and sinensis differ in essential morphological characters from Libiocoris sensu Kormilev, 1957, for which a new genus Paralibiocoris gen. n. is proposed. A further three new species from China belonging to this genus are recognized and described herein.
3 The single species L. indicus Heiss, 1982 described from north India, tentatively assigned to Libiocoris, is not congeneric with Libiocoris sensu Kormilev, 1957 nor to Paralibiocoris gen. n. erected for the Chinese species.
4 Inconsistencies and remarkable differences in the descriptions of angulatus, antennatus, lobatus, and pilosus raise questions about their congeneric assignment when compared with poecilus.
Depositories of type material examined:
CEHI Collection Ernst Heiss, Tiroler Landesmuseum Innsbruck, Austria
Photographs were taken through Keyence VHX-1000 equipment. Measurements were made using a calibrated micrometre; all measurements are given in millimetres. Abbreviations used as follows:
deltg dorsal external laterotergite (connexivum);
mtg mediotergite;
ptg paratergite;
vltg ventral laterotergite.
Type specimens of L. heissi and L. sinensis are conspecific, thus the following synonymy Libiocoris heissi Bai, Yang & Cai, 2006: 41 = Libiocoris sinensis Bai, Yang & Cai, 2006: 43 syn. n. is here established, heissi having priority.
Libiocoris heissi Bai, Yang & Cai, 2006.
General aspect similar to Libiocoris Kormilev, 1957 but is distinguished from the type species Libiocoris poecilus (characters in brackets) by the following set of morphological characters:
• position of spiracles: II ventral, III–VII lateral and visible from above (II–III ventral IV–V sublateral not visible from above, VI–VII lateral and visible);
• fused deltg II+III shorter, reaching only posterior border of metanotum (Figs
• presence of a smooth oblique callus on vltg VII of male which is independent of spiracle VII (Figure
• fused median longitudinal sclerite reaching from pronotum to tergal plate bottle-shaped along meso- and metanotum, then restricted along mtg I+II and carinate, the fusion line between metanotum – mtg I+II marked by a suture (Figs
• median ridge of abdomen distinctly elevated along midline (flat, not developed), dorsally reflexed vltg VII subrectangular (produced posteriorly, long and acute in male, shorter and acute in female);
• shape of male pygophore pyriform, produced posteriorly (wide and short).
Paralibiocoris gen. n. is very similar to Bruneiaptera Heiss, 2011 from Borneo, sharing basic habitus and dorsal thoracic structures; however, in Brunneiaptera all spiracles (II–VII) are lateral and visible from above.
Apterous, of small size 4.4–5.8 mm; habitus elongate-oval; legs and antennae beset with small setigerous granules; coloration yellowish to reddish or blackish brown.
Head. Subquadrangular, longer or as wide as distance across eyes; clypeus short, genae slightly produced; antenniferous tubercles short with acute apices; antennae long and slender, first and third and second and fourth segments subequal in length, first stout, incrassate, second and third cylindrical, fourth fusiform; eyes small, granulate; postocular tubercles distinct; rostrum arising from a slit-like atrium, not reaching limits of rostral groove.
Thorax. Pronotum short and wide; anterolateral angles produced forward beyond collar forming large blunt or rounded lobes; disc with a median sulcus; separated from mesonotum by a transverse intersegmental furrow; meso- and metanotum separated only laterally, the elevated median ridge smooth without sulcus; lateral sclerites with longitudinal elevations; metanotum separated from fused mtg I+II by a narrow transverse sulcus;
Abdomen. Mtg I and II fused together; mtg III to VI fused into a subquadrangular tergal plate, elevated along midline with usual pattern of large and small callous spots and dots; mtg VII strongly elevated posteriorly in male and slightly elevated in female; pygophore cordate; paratergites VIII clavate or lobiform.
Venter. Prosternum raised and with Y-shaped median carina; meso- and metasternum and sternum II+III fused and flattened medially. Spiracles II ventral, III-VII lateral on dorsally reflexed vltg III-VII and visible from above; spiracle VIII terminal on ptg VIII.
Legs. long and slender, without spine, preapical comb on fore tibia present, femora subcylindrical, claws with fine pulvilli.
From “para-“ close to (Greek) and Libiocoris.
1 | Antennal segment I as long as III | 2 |
– | Antennal segment I longer than III | 3 |
2 | Antennae longer, 2.1 times as long as width of head, anterolateral lobes of pronotum narrow and produced (Figs |
heissi (Bai et al., 2006), comb. n. |
– | Antennae shorter, approx. 1.9 times as long as width of head, anterolateral lobes of pronotum wider and less produced (Figs |
hainanensis sp. n. |
3 | Pronotum narrower 2.86 times wider than long, anterolateral lobes widely rounded (Figs |
roundangulus sp. n. |
– | Pronotum wider, more than three times as wide as long, anterolateral lobes narrower (Figs |
limuensis sp. n. |
Libiocoris
heissi
Bai, Yang & Cai 2006: 41, figs 1, 3–7 (
Libiocoris
sinensis
Bai, Yang & Cai 2006: 43, figs 2, 8–12 (
Holotype (♀): China, Hainan, Baisha, Yinggeling, 1050 m, 10.IX.2005, L. S. Chen leg. (
As both taxa were described on single females and males are now available, the holotype of heissi is redescribed and additional features of the male added.
As generic description.
Apterous female, incrustation removed to recognise dorsal structures.
Head. Slightly longer than wide across eyes (1.0/0.9); clypeus short reaching basal one-third of first antennal segment, strongly raised anteriorly, with tubercle near apex; genae slightly produced over clypeus; antenniferous tubercles short, dilated, apices acute, diverging anteriorly; antennae 2.1 times as long as width of head across eyes, length of antennal segments I to IV = 0.65, 0.30, 0.65, 0.30; eyes small, not protruding; postocular tubercles small but distinct, not reaching outer margin of eyes; postocular borders behind tubercles straight and converging to constricted collar; vertex with Y-shaped granulate carina flanked by two(1+1) large, ovate infraocular callosities; rostrum short, rostral groove wide and deep, closed posteriorly.
Pronotum. 2.8 times as wide as long (1.4/0.5); collar narrow, anterolateral angles produced forward beyond collar as two (1+1) large, blunt, granulate lobes; disc with a longitudinal median furrow flanked by ovate callosities; posterior margin of pronotum slightly convex posteriorly, separated from mesonotum by a deep furrow.
Mesonotum. Wider than pronotum, separated from metanotum by two (1+1) deep furrows laterally; across meso- and metanota medially with an elongate, subpentagonal bottle-shaped ridge, 1.53 times as long as wide (0.87/0.57), subrounded anteriorly and truncate posteriorly, smooth and without longitudinal sulcus; lateral of median ridge disc with four (2+2) longitudinal sclerites, lateral margins granulate.
Metanotum. Wider than mesonotum; separated from fused mtg I+II by a slightly sinuate thin sulcus; lateral of median ridge with two (1+1) large subtriangular callosities, 2 (1+1) longitudinal ridges lateral of callosity discs, lateral margins granulate, similar to those of mesonotum.
Abdomen. Mtg I and II completely fused, depressed at middle, there with a median longitudinal ridge laterally separated by deep furrows from wide oblique lateral plates, sloping posteriorly and sideways, further laterally with two (1+1) large subtriangular depressions; tergal plate with a slightly raised median ridge on mtg III, a pentagonal elevation on mtg IV then tapering posteriorly.
Venter. Sterna III to VI raised along posterior border, depressed along anterior border, and with triangular, smooth spots medially, flanked by two (1+1) large, transversely ovate depressions, these bearing two (1+1) round callous spots; laterally four (2+2) smaller round callous spots present; spiracles II ventral, III–VIII lateral and visible from above.
Legs. Long and slender, without spines, preapical comb on fore tibia present, femora subcylindrical, claws with fine pulvilli.
Male. Morphological features similar to female but of smaller size. Head as long as wide across eyes; median plate of meso- metanotum more elongate 1.82 times as long as wide (0.73/0.40); mtg VII strongly elevated posteriorly; reflexed vltg VII forming triangular lobes posterolaterally, ventral side with a distinct oblique smooth callus, reaching lateral margin; ptg VIII short and clavate much shorter than cordate pyriform pygophore (Figs
[in mm, ♂(n = 2)/♀ (n = 3), holotype in parentheses]. Body length 4.4-4.45/5.1-5.8 (5.6); maximal width of abdomen 2.05-2.2/2.65-3.05 (3.05). Head length 0.8-0.85/0.9-1.05 (1.0), width 0.8/0.8-0.95 (0.9). Pronotum length 0.4/ 0.45-0.5 (0.5), width 1.15-1.2/1.3-1.45 (1.4). Mesonotum width 1.5-1.6/1.7-1.95 (1.8). Metanotum width 1.8-1.9/2.05-2.3 (2.25). Length of antennal segments I–IV = 0.60, 0.25, 0.60, 0.30/0.6-0.7, 0.3-0.35, 0.6-0.7, 0.3-0.35 (0.65, 0.30, 0.65, 0.30).
China (Hainan).
As the generic characters and diagnosis are valid for all hereafter described new congeneric taxa, common features are not repeated except those differing in structure of thoracic median plate, size, and measurements distinctive for the specific taxa.
Holotype: ♂, China, Hainan, Jianfeng, Tianchi, 810 m, 16.VIII.2007, Zhang & Bai (
General aspect similar to Paralibiocoris heissi, but distinguished from it by a wider pronotum, 2.86 times as wide as long (2.80 in P. heissi), anterolateral lobes widely rounded (narrow and more produced), shorter antennae 1.79 times as long as width of head (2.1). Paralibiocoris roundangulus sp. n. differs from P. hainanensis sp. n. and P. limuensis sp. n. by a different shape of the median ridge of meso- and metanotum (Figs
Male. Basic morphological structures as of P. heissi. Head. As long as wide across eyes (0.82/0.82); antennae 1.79 times as long as width of head across eyes, length of antennal segments I to IV = 0.48, 0.24, 0.44, 0.31.
Pronotum. 2.86 times as wide as long (1.23/0.43); collar narrow; anterolateral angles produced forward beyond collar as two (1+1) widely rounded granulate lobes; disc with a longitudinal median furrow flanked by 2 (1+1) large, subtriangular and smaller callosities; lateral margin granulate.
Mesonotum. Wider than pronotum (1.60/1.23); separated from metanotum by two (1+1) deep furrows laterally; across meso- and metanota medially with an elongate, smooth bottle-shaped plate similar to P. heissi, 1.56 times as long as wide (0.67/0.43).
Metanotum. Wider than mesonotum (1.83/1.60); separated from mtg I by a slightly sinuate thin sulcus.
Abdomen. Mtg I and II completely fused, disc with a wide, smooth rectangular plate at middle flanked by two (1+1) large oblique plates, sloping posteriorly and sideways, laterally with two (1+1) small subtriangular depressions; deltg II and III fused, the following separated by fine sulci; posterolateral angles of deltg V to VII progressively angularly protruding; paratergites clavate, short, not reaching beyond posterolateral angles of deltg VII; pygophore elongate cordate, surface rugose (Figs
Venter. Sterna III to VI raised along posterior border, depressed along anterior border, and with triangular, smooth spots medially, flanked by 2 (1+1) shallow, transversely ovate depressions, these bearing 2 (1+1) round callous spots; 4 (2+2) smaller round callous spots present laterally; vltg VII with a small callus near spiracle VII; spiracles II ventral, III–VIII lateral and visible from above.
Female. Morphological features similar to male but of larger size; head slightly longer than wide across eyes (0.93/0.87); length of antennal segments I to IV = 0.48, 0.24, 0.44, 0.31; pronotum wider than long (1.47/0.43); width of mesonotum 1.90; width of metanotum 2.16, anterior lobe of median plate across meso- and metanota truncate, 1.2 times as long as wide (0.78/0.65); mtg VII moderately elevated posteriorly, the posterolateral angles forming triangular lobes; ptg VIII lobiform, reaching basal half of segment IX.
[in mm, ♂(n = 4)/♀ (n = 2), holotype in parentheses]. Body length 4.1-4.15/4.9-5.1 (4.1); maximal width of abdomen 2.05-2.1/2.55-2.75 (2.1). Head length 0.82/0.9-0.93 (0.82), width 0.82/0.8-0.87 (0.82). Pronotum length 0.43/0.43 (0.43), width 1.23/1.4-1.47 (1.23). Mesonotum width 1.55-1.6/1.85-1.90 (1.60). Metanotum width 1.83-1.9/2.1-2.16 (1.83). Length of antennal segments I–IV = 0.48, 0.24, 0.44, 0.31/0.48, 0.24, 0.44, 0.31 (0.48, 0.24, 0.44, 0.31).
The name of species refers to the widely rounded anterolateral angles of pronotum.
China (Hainan).
Holotype (♂): China, Hainan, Jianfeng, Tianchi, 810 m, 16.VIII.2007, Zhang & Bai; (
General aspect similar to Paralibiocoris heissi, but distinguished from the latter by the wider pronotum, 2.91 times as wide as long (2.80 in P. heissi) and more rounded less produced anterolateral lobes (produced and blunt), shorter antennae 1.89 times as long as width of head (2.1) and by posterolateral angles of deltg V to VII slightly protruding and rounded in female. Paralibiocoris hainanensis sp. n. differs from P. roundangulus sp. n. and P. limuensis sp. n. by a different shape of the median ridge of meso- and metanotum (Figs
Male. Basic morphological structures as of P. heissi and other congeners. Head. Slightly longer than wide across eyes (0.8/0.78); antennae 1.89 times as long as width of head across eyes, length of antennal segments I to IV = 0.47, 0.24, 0.47, 0.30.
Pronotum. 2.91 times as wide as long (1.25/0.43); collar narrow; anterolateral lobes produced forward beyond collar as two (1+1) widely rounded granulate lobes; disc with a longitudinal median furrow flanked by 2 (1+1) large, subtriangular and smaller callosities, lateral margin granulate, converging anteriorly.
Mesonotum. Wider than pronotum (1.60/1.25); separated from metanotum by two (1+1) deep furrows laterally; across meso- and metanota medially with an elongate, smooth bottle - shaped plate as P. heissi, 1.63 times as long as wide (0.70/0.43).
Metanotum. Wider than mesonotum (1.80/1.60); separated from mtg I by a slightly sinuate thin sulcus.
Abdomen. Mtg I and II completely fused, disc depressed at middle with a flat rectangular sclerite separated from lateral ovate plates by deep furrows; tergal plate with a slightly elevated granulate ridge which is widest on mtg III, sloping posteriorly; pygophore elongate cordate, surface rugose (Figs
Venter. Vltg VII with a small shiny callus, near spiracle VII; spiracles II ventral, III–VIII lateral and visible from above.
Female. Morphological features similar to male but of larger size. Head slightly longer than wide across eyes (0.80/0.78); length of antennal segments I to IV = 0.46, 0.27, 0.46, 0.33; pronotum wider than long (1.27/0.43); width of mesonotum 1.60; bottle-shaped median thoracic plate 1.56 times as wide as long (0.67/0.43); width of metanotum 1.93.
[in mm, ♂ (n = 9)/♀ (n = 4), holotype in parentheses]. Body length 3.65-4.2/4.2-4.55 (4.2); maximal width of abdomen 1.75-2.1/2.2-2.4 (2.1). Head length 0.7-0.8/0.75-0.8 (0.8), width 0.65-0.78/0.7-0.78 (0.78). Pronotum length 0.35-0.43/ 0.4-0.45 (0.43), width 1.05-1.25/1.2-1.3 (1.25). Mesonotum width 1.4-1.6/1.45-1.6 (1.6). Metanotum width 1.5-1.8/1.65-1.9 (1.8). Length of antennal segments I–IV = 0.44-0.47, 0.24, 0.44-0.47, 0.3/0.47, 0.24-0.27, 0.47, 0.3-0.34 (0.47, 0.24, 0.47, 0.3).
The name refers to the Island of Hainan, the type locality.
China (Hainan).
Holotype (♂): China, Hainan, Limu, Montain, 6.V.2008, Bai, X. S.; (
General aspect similar to Paralibiocoris heissi, but distinguished from the latter by wider pronotum 3.06 times as wide as long (2.80) and more rounded less produced anterolateral lobes (produced and blunt), shorter antennae 1.82 times as long as width of head (2.1) and by antennal segment I longer than III (of same lengths in heissi). Paralibiocoris limuensis sp. n. differs from P. roundangulus sp. n. and P. hainanensis sp. n. by a wider pronotum (3.06 vs. 2.86 and 2.91 respectively) and a leaf-like shape of the median ridge of meso- and metanotum (Figs
Male. Basic morphological structures as of P. heissi and other congeners. Head. Slightly longer than wide across eyes (0.75/0.68); antennae 1.82 times as long as width of head across eyes, length of antennal segments I to IV = 0.40, 0.20, 0.37, 0.27.
Pronotum. 3.06 times as wide as long (1.13/0.37); collar narrow; anterolateral lobes produced forward beyond collar as two (1+1) widely rounded granulate lobes, lateral margins converging anteriorly; structure of disc as in other congeners.
Mesonotum. Wider than pronotum (1.40/1.13); separated from metanotum by two (1+1) deep furrow laterally; across meso- and metanota medially with an elongate, anteriorly tapering leaf-like shaped plate, its surface slightly concave, 1.34 times as long as wide (0.63/0.47).
Metanotum. Wider than mesonotum (1.6/1.40); separated from mtg I by a slightly sinuate thin sulcus.
Abdomen. Mtg I and II completely fused, disc medially with a barrel-shaped sclerite resembling the leaf-stalk of the leaf - shaped ridge, separated from lateral ovate plates by deep furrows; tergal plate with a slightly elevated granulate ridge which is widest on mtg III, sloping posteriorly; pygophore elongate cordate, surface rugose (Figs
Venter. Vltg VII with a glabrous callus near spiracle VII; spiracles II ventral, spiracles III–VIII lateral and visible from above.
Female. Morphological features similar to male but of larger size; head slightly longer than wide across eyes (0.80/0.75); length of antennal segments I to IV = 0.44, 0.24, 0.37, 0.27; pronotum wider than long (1.17/0.40); width of mesonotum 1.6; width of metanotum 1.73; lateral margins of leaf-like median plate across meso- and metanota bisinuous, converging anteriorly to narrow apex, ratio length/width as of male (Figure
[in mm, ♂(n = 3)/♀ (n = 2), holotype in parentheses]. Body length 3.7-3.8/4.2-4.4 (3.8); maximal width of abdomen 1.76-1.9/2.2-2.3 (1.9). Head length 0.7-0.75/0.8 (0.75), width 0.65-0.68/0.7-0.75 (0.68). Pronotum length 0.35-0.37/ 0.4-0.45 (0.37), width 1.02-1.13/1.17-1.2 (1.13). Mesonotum width 1.3-1.4/1.5-1.6 (1.4). Metanotum width 1.5-1.6/1.73-1.8 (1.6). Length of antennal segments I–IV = 0.40, 0.20, 0.37, 0.27/0.4-0.44, 0.20-0.24, 0.37, 0.27 (0.40, 0.20, 0.37, 0.27).
The name of species reflects the locality of this new taxon.
Figure
Libiocoris indicus
The species was described upon two males and a female from Meghalaya State in the north of India, and shares several morphological characters with Libiocoris (habitus, head and antennae, fusion of thoracic segments) and according to the (misleading) redescription of Libiocoris (based on their new species L. antennatus Usinger & Matsuda, 1959) also the position of spiracles II-VII placed laterally and visible from above. Due to this similarity it was “tentatively” assigned to Libiocoris.
Libiocoris, habitus dorsal view. L. poecilus holotype male (81); ditto paratype female (82); L. angulatus holotype female (83); L. indicus holotype male (84)(after Heiss, 1982); L. antennatus holotype female (85) (after Usinger & Matsuda, 1959); L. angulatus holotype female, illustration in Usinger & Matsuda, 1959 with incorrect scale for antennae (86). Scale bar: 1 mm.
Re-examination of the paratype male of indicus has shown that it belongs neither to Libiocoris sensu Kormilev, 1957 (abbreviated L below) nor to Paralibiocoris gen. n. (abbreviated P below) showing following set of characters:
• position of spiracles: II-VII lateral and visible from above (not L, not P);
• fused deltg I+II shorter, reaching laterally only to metanotum (as P, not L);
• distinct glabrous oblique callus on vltg VII of male is lacking (as L, not P);
• fused median thoracic ridge of different shape (not L, not P);
• median thoracic ridge is elevated to sulcus of border metanotum, mtg I+II then sloping posteriorly (as P, not L);
• median ridge of abdomen raised along midline (as P, not L);
• dorsally reflexed vltg VII is subrectangular and not produced (as P, not L);
• shape of pygophore pyriform and produced posteriorly (as P, not L).
The position of spiracles, different from both Libiocoris and Paralibiocoris, is recognized as a diagnostic character used in Aradidae taxonomy for distinguishing genera [e.g., Acaricoris from Kolpodaptera; Parapictinus from Mezira (Usinger & Matsuda, 1959)] which supports a separate generic category for indicus. As other similar specimens from Vietnam and Japan present in our collections need to be included in a separate study, we refrain here from describing a new genus, but this species should be removed from Libiocoris sensu Kormilev, 1957. Inconsistencies and remarkable differences in the descriptions and illustrations of angulatus, antennatus, lobatus, and pilosus compared with poecilus raise questions about their generic assignment.
Libiocoris Kormilev, 1957
Libiocoris Kormilev, 1957 (1956): 390 (original description).
Type species poecilus by original designation, 391: figs 1–5 male HT, female AT, New Guinea (LP)
Libiocoris antennatus Usinger & Matsuda, 1958: 181, fig. 53 female HT, New Guinea (LA)
Libiocoris angulatus Usinger & Matsuda, 1959: 84, fig. 84 female HT, New Guinea (LG)
Libiocoris angulatus Heiss, 1989: 349, figs 12a, b (redescription, correction antennae) (Figs
Libiocoris lobatus
Libiocoris pilicornis Kormilev, 1972: 568, figs 9, 9A, New Guinea (LPC); not seen.
Anterior extension of fused deltg II+III reaching pronotum.
In LP present in types, not shown in Figure
Position of spiracles denoted as + for visible from above and - for not visible. In LP II-III, VI-VII lateral +, IV ventral -, V sublateral -; LA II-VII lateral +; LG II-III lateral +, IV-V sublateral -; LL II-III, V-VIII lateral +, IV-V sublateral -; LPC II-VII lateral +.
Shape of median longitudinal ridge: in LP narrow with furrow; LA narrow with furrow; LG narrow, furrow figured; LL?; LPC posteriorly wider, without furrow.
As none of those female type specimens described after Kormilev’s definition of Libiocoris shares all these characters, it is questionable as to what might be a valid character state and what is due to variability, and whether they belong to the same generic category. This question can be resolved when further material will be available for study and barcoding.
We sincerely thank Tamás Vásárhelyi and David Redei (