identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03CB87AEFF83FFC48FB9FCC9FDCDFF6C.text	03CB87AEFF83FFC48FB9FCC9FDCDFF6C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhizophagus Herbst 1793	<div><p>Taxonomic comparison of Western Palaearctic Rhizophagus of subgenus Eurhizophagus</p><p>Type material examined. Rhizophagus atticus Tozer, 1968, holotype ♂ and 2 ♀ paratypes, Natural History Museum of the United Kingdom (NHMUK).</p><p>Holotype ♂ (Fig. 1), dissected, with the following labels: « Type [round label, white circled in red] / In bark of Pinus / Greece, Parnes Mts, X. 1965, C. Nicolopoulos / Brit. Mus., 1965-598 / Holotype, Rhizophagus atticus sp. n. ♂, E. R. Tozer, 1967 / NHMUK014485018 ».</p><p>Paratype ♀ (Fig. 2), with the following labels: « Paratype [round label, white circled in yellow] / In bark of Pinus / Greece, Parnes Mts, X. 1965, C. Nicolopoulos / Brit. Mus., 1965-598 / Paratype, Rhizophagus atticus sp. n. ♀, E. R. Tozer, 1967 / NHMUK014663279 ».</p><p>Paratype ♀ (Fig. 3), with the same labels as above but barcode « NHMUK014485021 ».</p><p>Other material examined agreeing with the original description of Rhizophagus diaboli . ALPES-DE-HAUTE-PROVENCE (04)—Selonnet (04203), 1060 m, 26.VII.–04.VIII. 2022, 1 ex. HAUTES-ALPES (05)— Veynes (05179), 840 m, 05.VIII.–19.VIII. 2021, 19 ex. ARDENNES (08)—Semide (08410), 120 m, 13.VII.– 05.VIII. 2021, 1 ex. AVEYRON (12)—Saint-Chély-d’Aubrac (12214), 910 m, 05.VIII.–19.VIII. 2021, 3 ex.; Sévérac - le-Château (12270), 820 m, 13.VII.–25.VII. 2022, 1 ex. DSF leg. CHARENTE (16)—Touvérac (16384), 90 m, 24.VIII.–05.IX. 2022, 1 ex. CORRÈZE (19)—Brive - la - Gaillarde (19031), 150 m, 03.VIII.–07.X. 2021, 15 ex. CÔTE-D’OR (21)—Arcenant (21017), 490 m, 24.VI.–05.VII. 2022, 5 ex.; Plombières - lès - Dijon (21485), 350 m, 05.VII.–28.VII. 2022, 7 ex. CÔTES-D’ARMOR (22)—Saint - Brieuc (22278), 20 m, 29.VI.–31.VIII. 2022, 1 ex. DSF leg. DORDOGNE (24)—Montrem (24295), 150 m, 31.VIII.–26.X. 2022, 2 ex.; Lèches (24234), 90 m, 12.VIII.–22.IX. 2022, 3 ex. DOUBS (25)—Rigney (25490), 250 m, 09.VIII.–18.VIII. 2021, 1 ex. DRÔME (26)— Cliousclat (26097), 450 m, 23.VII.–02.VIII. 2021, 2 ex. EURE (27)—Forêt domaniale de Bord-Louviers, 120 m, 26.VII.–16.VIII. 2021, 11 ex. GIRONDE (33)—Escaudes (33155), 20 m, 22.VIII.–06.X. 2022, 2 ex.; Mérignac (33281), 40 m, 13.IX.–23.IX. 2021, 12 ex. ILLE-ET-VILAINE (35)—Gaël (35117), 120 m, 26.VII.–30.VIII. 2022, 5 ex.; Iffendic (35133), 100 m, 22.VII.–26.VII. 2022, 1 ex. JURA (39)—Salins - les - Bains (39500), 360 m, 29.VII.– 18.VIII.2021, 2 ex. LANDES (40)—Bégaar (40031), 20 m, 04.VII.–15.VII. 2022, 1 ex.; Mont - de - Marsan (40192), 50 m, 18.VIII.–30.VIII. 2021: 2 ex., 10.VIII.–23.VIII. 2022: 1 ex. LOIRE-ATLANTIQUE (44)—Joué-sur - Erdre (44077), 20 m, 2022, 4 ex. LOT (46)—Salviac (46297), 220 m, 19.VII.–29.VII. 2021, 2 ex.; Saint - Denis - Catus (46264), 340 m, 29.VII.–09.VIII. 2021, 1 ex. LOT-ET-GARONNE (47)—Fargues - sur - Ourbise (47093), 110 m, 26.VIII.–28.X. 2021, 4 ex. LOZÈRE (48)—Mende (48095), 740 m, 05.VIII.–17.XIII.2021, 3 ex. MAINE-ET-LOIRE (49)—Lézigné (49174), 50 m, 02.VIII.–20.VIII. 2021, 10 ex.; Montreuil - sur-Loir (49216), 30 m, 20.VIII.– 03.IX. 2021, 7 ex. MOSELLE (57)—Volmerange-les-Mines (57731), 320 m, 15.VII.–25.VII. 2022, 1ex. ORNE (61)—Orée-d’Écouves (61228), 400 m, 02.VIII.–10.VIII. 2022, 1 ex. HAUTES-PYRÉNÉES (65)—Ayros - Arbouix (65055), 450 m, 29.VII.–13.IX. 2022, 2 ex. BAS-RHIN (67)—La Vancelle (67505), 430 m, 23.VIII.–09.IX. 2021, 4 ex. HAUT-RHIN (68)—Habsheim (68118), 250 m, 12.VII–29.VII. 2021, 2 ex.; Rixheim (68278), 240 m, 29.VII.– 23.VIII. 2021, 10 ex. SEINE-MARITIME (76)—Arelaune-en-Seine (76401), 50 m, 01.VI.–10.VI. 2022, 1 ex.; Eu (76255), 70 m, 01.VI.–10.VI. 2022, 1 ex. VIENNE (86)—Bonneuil - Matours (86032), 60 m, 12.VIII.–27.X. 2022, 88 ex.; Usseau (86275), 140 m, 11.VIII.–06.X. 2022, 12 ex. YONNE (89)—Chassignelles (89087), 190 m, 26.VII.–16.VIII. 2021, 106 ex.; Gurgy (89198), 50 m, 26.VII.–16.VIII. 2021, 1 ex.</p><p>All material collected by DSF staff. Some of this material has been deposited at the ONF’s National Forest Entomology Laboratory (LNEF-ONF).</p><p>Key to Western Palaearctic Rhizophagus of subgenus Eurhizophagus</p><p>1 Larger, 4.5–5.5 mm; pronotum broader as long; microreticulation longitudinally elongated and sinuate, always clearly visible................................................................................... R. grandis Gyllenhal (Naturally rare species in mountain spruce forests but widely disseminated for biological control in many mountain and lowland stands.).</p><p>- Smaller, &lt;4 mm; pronotum as wide as long or barely longer than wide; polygonal microreticulation or absent............ 2</p><p>2 Antennal club globose, 11 th antennomere clearly visible (Fig. 13); median lobe of genitalia long, curved over the first basal third and then sub rectilinear, with a small tuft of long setae on its ventral side, in the anterior third (Fig. 19); body slender and on average larger (2.8–4.2 mm, median 3. 3 mm) (Fig. 6)....................................... R. depressus (Fabricius) (Common and widely distributed species, especially in lowland and mountain pine forests.).</p><p>- Antennal club subtruncate, 11 th antennomere inconspicuous, embedded in the 10 th (Figs. 11, 12); median lobe of genitalia short, regularly curved, without long tuft of setae on ventral side (Figs. 17, 18); body more compact and on average smaller (2.5–3.6 mm, median 2. 9 mm) (Figs. 1, 2, 3, 5)............................... R. atticus Tozer (= R. diaboli Dodelin, syn. nov.)</p></div>	https://treatment.plazi.org/id/03CB87AEFF83FFC48FB9FCC9FDCDFF6C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Barnouin, Thomas;Vincent, Alexis;Soldati, Fabien	Barnouin, Thomas, Vincent, Alexis, Soldati, Fabien (2025): Rhizophagus atticus Tozer, 1968 in France: synonymy with R. diaboli Dodelin, 2021, distribution and ecology (Coleoptera: Monotomidae). Zootaxa 5691 (2): 329-340, DOI: 10.11646/zootaxa.5691.2.7, URL: https://doi.org/10.11646/zootaxa.5691.2.7
03CB87AEFF85FFC48FB9FEC5FCACF8F4.text	03CB87AEFF85FFC48FB9FEC5FCACF8F4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhizophagus diaboli Dodelin 2021	<div><p>Rhizophagus diaboli compared with R. depressus</p><p>Dodelin (2021) described several characters that distinguish R. diaboli from R. depressus, which have been tested by studying the material examined.</p><p>We confirm that the shape of the antennal club is a valid character to distinguish both species. In R. diaboli, the antennal club is sub-truncate, with the 11 th antennomere embedded in the 10 th (Fig. 12), whereas the club of R. depressus is ovoid, with the 11 th antennomere clearly protruding from the 10 th (Fig. 13). However, this characteristic may sometimes prove ambiguous due to a slight variability in the conformation of the 11 th antennomere in R. diaboli . In addition, the angle of observation can also influence the perception of this morphological character.</p><p>Moreover, we can verify that the characters based on the male genitalia distinguish the two species. Firstly, the median lobe of R. diaboli seen in profile is short and regularly curved up to the apex (Fig. 18), whereas that of R. depressus is clearly longer, curved over the basal third of its length and then sub-rectilinear up to the apex (Fig. 19). This character separates the two species. The arrangement of the setae on the median lobe is also a diagnostic feature. However, while the presence of a small tuft of long setae in the apical third of the ventral face of the median lobe is confirmed in R. depressus (Fig. 19), the representation and description of the arrangement of these setae in R. diaboli is imprecise.Admittedly, only small setae are found on the apical margin of the genitalia, but these setae form a crown along the entire margin and not just on the ventral area (Fig. 18). Finally, the sclerites of the endophallus, visible on the dorsal surface of the median lobe, differ in configuration between the two species. The medial sclerite of R. diaboli is broad and rectilinear while it is fine and strongly arcuate in R. depressus . Moreover, the branches of the anterior sclerite are straight while they are strongly sinuate in R. depressus . Although the representations of these internal parts are somewhat approximate, the configurations we observed are quite different and consistent with the proposed photos (Figs. 15, 16).</p><p>According to Dodelin (2021), the cuticle on the dorsal surface of R. diaboli is smooth and shiny, the microreticulation visible only at very high magnification (x70), whereas it is strongly micro-reticulated in R. depressus, giving it a dull appearance. The microreticulation of the pronotom and elytra is highlighted by the author as one of the main characters distinguishing the two species. However, the study of abundant material from France leads to the invalidation of this character. Indeed, while a fine but clear microreticulation is always present on the pronotum in R. depressus (Fig. 10), the punctation of the pronotum is highly variable in R. diaboli, ranging from a smooth, shiny pronotum with imperceptible microreticulation (Fig. 8) to a dull pronotum with very strong microreticulation (Fig. 9). Moreover, the dull form is more common and more widespread than the shiny form. Both forms can cohabit at the same site. In this case, one of them dominates, with a ratio of more than 10 to 1.</p><p>According to our studied material, there is indeed a subtle difference in the shape of the pronotum between both species. However, the differences described by Dodelin (2021) are exaggerated and, above all, imprecise, as the author does not consider the sexual dimorphism of this character in most of species within the genus (Peacock 1977). The pronotum of the male of R. diaboli is barely longer than wide, with straight, subparallel to divergent margins, narrowing slightly at the apex (Fig. 8), whereas that of the female is as long as wide, with straight subparallel margins narrowing progressively from the anterior fourth (Fig. 9). In R. depressus, the pronotum of the male is as long as wide, with straight, subparallel or barely divergent margins, narrowing progressively over the anterior third, whereas that of the female, with straight, subparallel margins, narrows abruptly in the anterior quarter (Fig. 10).</p><p>Although the use of certain characters has been invalidated (punctation of the pronotum) or clarified (shape of the pronotum and arrangement of the setae on the median lobe of the male genitalia), the diagnosis presented leaves no doubt as to the existence in France of a third species in the subgenus Eurhizophagus, closely related to R. grandis and R. depressus, corresponding approximately to the description of R. diaboli . It may be added that there is an overlap in size between both species, R. diaboli being generally smaller (2.5–3.6 mm; median=2.9 mm) and more compact than R. depressus (2.9–4.2 mm; median=3.3 mm).</p></div>	https://treatment.plazi.org/id/03CB87AEFF85FFC48FB9FEC5FCACF8F4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Barnouin, Thomas;Vincent, Alexis;Soldati, Fabien	Barnouin, Thomas, Vincent, Alexis, Soldati, Fabien (2025): Rhizophagus atticus Tozer, 1968 in France: synonymy with R. diaboli Dodelin, 2021, distribution and ecology (Coleoptera: Monotomidae). Zootaxa 5691 (2): 329-340, DOI: 10.11646/zootaxa.5691.2.7, URL: https://doi.org/10.11646/zootaxa.5691.2.7
03CB87AEFF87FFC78FB9F89DFBCEFC38.text	03CB87AEFF87FFC78FB9F89DFBCEFC38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhizophagus diaboli Dodelin 2021	<div><p>Rhizophagus diaboli compared with R. atticus</p><p>The shape of the antennal club is not a valid character to discriminate between R. diaboli and R. atticus . Although Tozer (1968) and Dodelin (2021) described the club of R. atticus as spherical, the type specimens of R. atticus consistently show a subtruncate antennal club with the 11 th antennomere embedded into the 10 th (Fig. 11), thus falling within the range of variability observed in R. diaboli (Fig. 12). The photos of this character by Nikolopoulos (1969) are consistent with our observations.</p><p>Body size is not a distinguishing feature between both species either. Dodelin (2021) maintains that R. atticus is smaller (2.0–2.8 mm) than R. diaboli (3.0–4.0 mm). However, it was based on an examination of only 10 specimens in total for both species. It is also regrettable that for R. atticus he relied only on the measurements presented by Tozer (1968), omitting the more extensive measurements (2.5–3.8 mm) presented by Nikolopoulos (1969). Our measurements show that the three specimens in the type series vary in size from 2.5–2.8 mm, thus falling within the size range of R. diaboli .</p><p>In the same sex, the shape of the pronotum does not vary significantly between both species. As previously mentioned, the microreticulation of the pronotum of R. diaboli is highly variable (Figs. 8, 9), including totally smooth forms like those observed in type specimens of R. atticus (Fig. 7).</p><p>Considering genitalia, the shape of the median lobe in profile, and the arrangement of setae on it, are perfectly similar between specimens of R. diaboli (Fig. 18) and the holotype of R. atticus (Fig. 17). We do not find the differences in the schemes presented by Dodelin (2021). The genitalia’s internal features of R. atticus holotype are not fully visible, because the genitalia are old and dried out (Fig. 14). Only a chemical treatment would possibly allow us to observe the genitalia in their entirety, but this method has been ruled out to avoid any damage. Nevertheless, the broad, rectilinear medial sclerite and the partially visible anterior sclerite are like those observed in R. diaboli (Fig. 15).</p><p>As a conclusion, considering the similarity in morphology and genitalia, as well as the lack of consistently differing characters between the two species, the following synonymy is established:</p></div>	https://treatment.plazi.org/id/03CB87AEFF87FFC78FB9F89DFBCEFC38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Barnouin, Thomas;Vincent, Alexis;Soldati, Fabien	Barnouin, Thomas, Vincent, Alexis, Soldati, Fabien (2025): Rhizophagus atticus Tozer, 1968 in France: synonymy with R. diaboli Dodelin, 2021, distribution and ecology (Coleoptera: Monotomidae). Zootaxa 5691 (2): 329-340, DOI: 10.11646/zootaxa.5691.2.7, URL: https://doi.org/10.11646/zootaxa.5691.2.7
03CB87AEFF86FFC98FB9FC30FA8EFB18.text	03CB87AEFF86FFC98FB9FC30FA8EFB18.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhizophagus atticus Tozer 1968	<div><p>Rhizophagus atticus Tozer, 1968</p><p>Rhizophagus atticus Tozer, 1968: 59</p><p>= Phizophagus [sic] atticus Nikolopoulos, 1969: 163</p><p>= Rhizophagus diaboli Dodelin, 2021: 134 . New synonymy</p><p>Geographical distribution</p><p>The new synonymy established, and the additional localities brought together with the existing works (Tozer 1968; Nikolopoulos 1969; Dodelin 2021) make it possible to update the distribution of R. atticus . In France, it is now reported from 43 communes in 30 administrative departments (Fig. 4). Contrary to what is assumed by Dodelin (2021), it is therefore widely distributed in France, from lowlands to mid-mountains, although it seems to be absent in the Mediterranean region and Corsica. According to our data, it is most abundant (86% of specimens caught) and most frequent (66% of localities occupied) between 0 and 300 m, its abundance decreasing with altitude (Fig. 20). Although these results are strongly correlated with the distribution of the sampling effort, the relative occurrence of the species confirms that its presence decreases with altitude, with no detection above 1,100 m (Fig. 22). R. depressus, whose distribution of abundance and frequency are like those of R. atticus (Fig. 21), on the other hand, shows a relative occurrence that tends to increase with altitude (Fig. 22).</p><p>Worldwide, R. atticus is currently only known in France and Greece. Given its very wide distribution in France, it should be found in most European countries where it is probably confused with R. depressus . Moreover, its presence in Poland is attested by a photo available on the internet of a specimen identified as R. depressus but that corresponds to R. atticus based on its habitus and antennal club (http://cassidae.uni.wroc.pl/Colpolon/ rhizophagus%20depressus.htm).</p><p>As Galloprotect Pack® apparently has a strong attraction for R. atticus, analysis of the material trapped from the pinewood nematode monitoring scheme set up in Sweden and other European countries in response to implementing decision 2012/535/EU (Schroeder 2014) would provide more precise information on its distribution.</p><p>Biological notes</p><p>Rhizophagus atticus is rarer than R. depressus both in terms of abundance (359 vs 1886 specimens caught) and frequency (13% vs 37% of locality occupancy). Nevertheless, it can be sometimes locally abundant, e.g. Chassignelle (Yonne), with 106 specimens caught in a single trap over a 15-day period in July/August. The coexistence of both species appears to be uncommon, as it was observed in only 13% of localities where at least one of both species was observed. However, about half of the sites occupied by R. atticus are also occupied by R. depressus (Fig. 4).</p><p>More than 90% of R. atticus catches were in pine forests, with the remainder in sawmills and wood piles. At the same time, no detections of this species were made by the LNEF-ONF during various inventories carried out between 2021 and 2024 in more than 40 deciduous and coniferous forest sites. We can therefore deduce that this species is mainly associated with pine ( Pinus species), like R. depressus, and not with silver fir as was suggested (Dodelin 2021). Moreover, this species was collected from under the bark of a pine tree in its type locality in Greece (Tozer 1968; Nikolopoulos 1969). At this stage, however, it cannot be ruled out that it may occasionally develop on other coniferous trees.</p><p>Like the other members of the subgenus Eurhizophagus, R. atticus is probably associated with one or more species of bark beetle, still unknown. R. depressus is mainly associated with the pine bark beetles Tomicus piniperda (Linnaeus), T. minor (Hartig) and Ips acuminatus (Gyllenhal), but also with those of other mountain conifers such as Pityogenes chalcographus (Linnaeus), Trypodendron lineatum (Olivier) and Hylurgops palliatus (Gyllenhal) (Schroeder 1996; Bouget &amp; Moncoutier 2003; Dodelin 2021). In the Scots pine forests of Turkey, it prefers to attack Ips sexdentatus (Boerner) and secondarily I. acuminatus, T. minor and T. piniperda (Akyol &amp; Sarikaya 2017) . It is likely that the associated bark beetles of R. atticus are some of those already identified in R. depressus, although it cannot be ruled out that it preys on other bark beetles. It is also possible that it is less polyphagous than the latter, which could explain its greater rarity.</p><p>Rhizophagus atticus was caught from June to October, with maximum abundance and frequency in August (Fig. 23). Its relative occurrence is high from August to October, with a maximum in August, demonstrating summer dispersal activity extending into early autumn (Fig. 25). Its phenology differs fundamentally from that of R. depressus, whose abundance and frequency are highest in June (Fig. 24) and whose relative occurrence shows maximum activity in late spring, in May and June, with a second peak in autumn, in October (Fig. 25). R. depressus also tends to occur in spring in Belgium, with peaks of activity in May and June (Thieren et al. 2003).</p></div>	https://treatment.plazi.org/id/03CB87AEFF86FFC98FB9FC30FA8EFB18	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Barnouin, Thomas;Vincent, Alexis;Soldati, Fabien	Barnouin, Thomas, Vincent, Alexis, Soldati, Fabien (2025): Rhizophagus atticus Tozer, 1968 in France: synonymy with R. diaboli Dodelin, 2021, distribution and ecology (Coleoptera: Monotomidae). Zootaxa 5691 (2): 329-340, DOI: 10.11646/zootaxa.5691.2.7, URL: https://doi.org/10.11646/zootaxa.5691.2.7
