Mystrium mysticum Roger, 1862

Yoshimura, Masashi & Fisher, Brian L., 2014, A revision of the ant genus Mystrium in the Malagasy region with description of six new species and remarks on Amblyopone and Stigmatomma (Hymenoptera, Formicidae, Amblyoponinae), ZooKeys 394, pp. 1-99 : 35-45

Mystrium mysticum Roger, 1862 Figs 4C, 7, 8C, 9A, 9C, 11A, 12A, 26B, 28A, 28C, 29A, 37B, 38B, 39B, 40B, 41B, 43A, 43C, 44A, 44C, 45A, 45C, 46A, 46C, 47A, 47C, 48A, 48C, 49A

Mystrium mysticum Roger, 1862. MADAGASCAR. Syntypes: two queens [lost: the neotype is designated below].

Mystrium mysticum : Forel 1892, in part (queen); Menozzi 1929.

Mystrium mysticum (Not) (male): Forel 1891; Menozzi 1929 [assumed from descriptions].

Mystrium stadelmanni Forel, 1895. syn. n.

Mystrium stadelmanni : Forel 1895, in part; Menozzi 1929.

Neotype

[here designated]. Alate queen: CASENT0429914, BLF4770, MADAGASCAR, Toliara, Parc National de Kirindy Mite, 16.3 km 127° SE Belo sur Mer (-20.79528°, 44.147°), 80 m alt., 6-10.xii.2001, Fisher-Griswold Arthropod Team leg. [CASC].

Lectotype of Mystrium stadelmanni

[here designated]. Worker: CASENT0104561, MADAGASCAR, Est de I’Imeriua, Sikora leg. [ZMHB: examined].

Worker.

Description. Measurements: lectotype of Mystrium stadelmanni . HL 2.05, HW 1.93, SL 1.16, ML 2.09, HD 1.26, WL 1.99, PnW 0.94, PpW 0.84, PtW 0.80, PtL 0.53, CI 94.3, SI 59.9, MI 108.2, PpI 89.5, PtI 153.0.

HL 1.44-2.27, HW 1.42-2.18, SL 0.86-1.35, ML 1.32-2.31, HD 0.98-1.39, WL 1.63-2.44, PnW 0.77-1.2, PpW 0.69-1.09, PtW 0.6-0.94, PtL 0.4-0.6, CI 96.0-102.0, SI 56-64, MI 91-108, PpI 86-95, PtI 146-168 (10 specimens measured).

Posterolateral corner of head strongly expanding posteriorly. Posterior face of vertex forming slightly blunt angle with its dorsal face on median line of head, so that declivity of vertex on lateral part slightly steeper than on median part. Ventral half of vertex smooth and not sculptured. Eye small to moderately small. Anterior margin of clypeus convex and with short conical setae. Genal tooth of head undeveloped: anterolateral corner of head angulate. Masticatory margin of mandible widely visible in full-face view, difference in width of dorsal surface of mandible relatively large between mandibular shaft and distal portion. Second maxillary palpomere longer than third. First flagellomere (third antennal segment) as long as pedicel (second antennal segment). Shallow and fine longitudinal striae irregularly impressed on central part of pronotal dorsum, sometimes with shallow reticulation around them. On lateral surface of pronotum, shallow, fine longitudinal striae impressed, wide and shallow punctures usually arranged on central horizontal line. Mesonotum differentiated from propodeum in dorsal view, length as long as that of propodeum in large individuals, shorter than propodeum in small individuals. Metanotal groove shallowly impressed in lateral view, mesonotum as high as pronotum in large individuals, higher than pronotum in small individuals. Metanotum weakly developed in largest individuals. Metapleural gland bulla developed, so that propodeal declivity in lateral view weakly convex posteriorly on its ventral portion. Petiole in dorsal view moderately wide, and relatively wider than that of Mystrium rogeri .

Body color brown to black.

Queen.

Description. Measurements: neotype. HL 2.44, HW 2.51, SL 1.49, ML 2.24, HD 1.54, WL 3.25, MnW 1.73, PtW 1.11, PtL 0.66, CI 102.7, SI 59.3, MI 89.2, MnI 69.1, PtI 168.2.

HL 2.14-2.73, HW 2.08-2.84, SL 1.26-1.60, ML 1.85-2.56, HD 1.32-1.72, WL 2.60-3.48, MnW 1.27-1.92, PtW 0.89-1.33, PtL 0.55-0.77, CI 97.2-104.2, SI 56.3-60.7, MI 87.2-98.2, MnI 61.0-71.3, PtI 160.2-188.8 (10 specimens measured).

Wings usually present and well developed; lacking in intercaste. Wing sclerites fully developed even if wings have dropped off in queen; developed to undeveloped in variable degrees in intercaste. Posterolateral corner of head strongly expanding posteriorly; expansion even stronger than that of workers. Posterior face of vertex forming slightly blunt angle with dorsal face on median line of head, so that declivity of vertex on lateral part slightly steeper than on median part. Ventral half of vertex smooth and not sculptured. Eye well developed. Both anterior and lateral ocelli clearly present and developed in most cases, lateral ones rarely smaller; ocelli varied from absent to developed in intercaste. Anterior margin of clypeus convex with small conical setae. Anterolateral portion of head angulate or with short spine. Masticatory margin of mandible widely visible in full-face view, and dorsal surface on distal portion distinctly wider that on mandibular shaft, difference in width much larger than that in Mystrium rogeri . Mandibular teeth (a row of dentition on dorsal side) often lacking on mid-portion of mandibular shaft. A spatulate seta present on basal side of each basal denticle on masticatory margin of mandible. First flagellar segment on antenna as long as pedicel. Setae on pronotum almost simple, narrowing distally with strongly sharpened apex. Propodeal declivity in lateral view almost straight and forming right to slight obtuse angle with dorsal margin, ventral portion with small convexity by metapleural gland. Petiole relatively long in dorsal view, about 0.6 × length of abdominal segment III.

Body color black.

Male.

Description. Measurements: HL 1.13-1.33, HW 1.52-1.81, SL 0.22-0.28, EL 0.62-0.68, WL 2.26-2.86, MnW 1.46-1.89, CI 129.5-136.0, SI 14.3-17.0, EI 49.0-54.8, MnI 96.3-104.4 (7 specimens measured).

Eye relatively small, occupying about 0.5 × length of head. Ocelli relatively distant from dorsal margin of head or just failing to reach dorsal margin in full-face view. Dorsal margin of head in full-face view rounded. Both anterior and lateral ocelli small. Distance between lateral ocellus and eye relatively long, about 3 × longer than diameter of lateral ocellus. Posterior half of vertex clearly differentiated from dorsal half, its dorsal face almost as long as its posterior face. Palpal formula 4,3. First segment of maxillary palp flattened and distinctly wider than second segment. Second maxillary palpomere longer than third. Notauli clearly impressed on mesoscutum. Petiole in dorsal view thin, its length about 0.65 × that of abdominal tergite III. Petiolar dorsum covered with rough, deep punctures. Abdominal tergum VIII roughly and deeply punctured.

Abdominal sternum IX punctured on its distal portion. Basal ring short, not extending basally. Telomere extending slightly further distally than digitus. Basoventral expansion of aedeagus well developed basoventrally, distinctly longer than dorsal extension. Ventral margin of aedeagus strongly curved ventrally in lateral view. Aedeagus weakly narrowing distally, its distal portion widely rounded.

On forewing, cu-a located at junction of Media (M) and Cubitus (Cu).

Body color reddish brown to black.

Distribution.

MADAGASCAR and COMOROS: as in Figure 49A.

Additional material examined.

In addition to the type material, specimens from the following localities were examined in this study: MADAGASCAR. Antsiranana. Forêt d’Ampondrabe, 26.3 km 10° NNE Daraina (-12.97°, 49.7°), tropical dry forest, 175 m alt.; Forêt de Binara, 9.1 km 233° SW Daraina (-13.26333°, 49.60333°), rainforest, 650-800 m alt.; 9.4 km 235° SW Daraina (-13.26333°, 49.6°), montane rainforest, 1100 m alt.; Nosy Faly (-13.3624°, 48.49101°), open secondary vegetation, 15 m alt.; (-13.36435°, 48.49137°), open secondary vegetation, 40 m alt.; Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112° ESE Hellville (-13.41933°, 48.33117°), rainforest, 30 m alt.; Ambondrobe, 41.1 km 175° Vohemar (-13.71533°, 50.10167 °), littoral rainforest, 10 m alt.; Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha (-13.79861°, 48.16167°), rainforest, 600 m alt. R.S. Manongarivo, 10.8 km 229° SW Antanambao (-13.96167°, 48.43333°), rainforest, 400 m alt.; Makirovana forest (-14.17066°, 49.95409°), rainforest, 225 m alt.; Forêt d’Anabohazo, 21.6 km 247° WSW Maromandia (-14.30889°, 47.91433°), tropical dry forest, 120 m alt.; Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina (-14.43667°, 49.775°), rainforest, 450 m alt.; Marojejy R.N.I. #12 (-14.44533°, 49.78564°), rainforest, 375 m alt.; R. Onive (-15.47°, 50.225°), 150 m alt. Toamasina. Nosy Mangabe, 7.43 km S Maroantsetra (-15.4973°, 49.76223°), littoral rainforest, 3 m alt.; Parc National Mananara-Nord, 7.1 km 261° Antanambe (-16.455°, 49.7875°), rainforest, 225 m alt.; Res. Ambodiriana, 4.8 km 306°Manompana, along Manompana river (-16.67233°, 49.70117°), rainforest, 125 m alt.; Manakambahiny, near Vavatenina Forest (-17.46667°, 49.35°); Parc National de Zahamena (-17.73359°, 48.72625°), rainforest, 950 m alt.; Réserve Nationale Intégrale Betampona, Betampona 35.1 km NW Toamasina (-17.91801°, 49.20074°), rainforest, 500 m alt.; Corridor Forestier Analamay-Mantadia, Tsaravoniana (-18.75737°, 48.42302°), rainforest, 1018 m alt.; Tsaravoniana (-18.76124°, 48.42134°), rainforest, 939 m alt.; Ambatoharanana (-18.8024°, 48.40612°), rainforest, 950 m alt.; (-18.80388°, 48.40506°), rainforest, 1013 m alt.; Bevolota 17.1 km N Andasibe (-18.77071°, 48.43164°), montane rainforest, 995 m alt.; Ambatovy, 12.4 km NE Moramanga (-18.83937°, 48.30842°), montane rainforest, 1080 m alt.; Perinet (-18.95°, 48.5°), 800 m alt.; Mahajanga. Réserve d’Ankoririka, 10.6 km 13° NE de Tsaramandroso (-16.26722°, 47.04861°), tropical dry forest, 210 m alt.; Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331° NW Andranofasika (-16.29889°, 46.813°), tropical dry forest, 70 m alt.; Parc National de Namoroka, 16.9 km 317° NW Vilanandro (-16.40667°, 45.31°), tropical dry forest, 100 m alt.; 9.8 km 300° WNW Vilanandro (-16.46667°, 45.35°), tropical dry forest, 140 m alt.; Réserve Spéciale de Bemarivo, 23.8 km 223° SW Besalampy (-16.925°, 44.36833°), tropical dry forest, 30 m alt.; Réserve forestière Beanka, 54.3 km E Maintirano (-18.06009°, 44.54086°), tropical dry forest on tsingy, 262 m alt.; Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova (-18.70944°, 44.71817°), tropical dry forest on tsingy, 150 m alt.; 2.5 km 62° ENE Bekopaka, Ankidrodroa River (-19.13222°, 44.81467°), tropical dry forest on tsingy, 100 m alt.; 3.4 km 93° E Bekopaka, Tombeau Vazimba (-19.14194°, 44.828°), tropical dry forest, 50 m alt.; Bekopaka (-19.14667°, 44.796°), village, 100 m alt. Toliara. Forêt de Kirindy, 15.5 km 64° ENE Marofandilia (-20.045°, 44.66222°), tropical dry forest, 100 m alt.; 48 km ENE Morondava, Kirindy Forest (-20.06667°, 44.65°), tropical dry forest, 30 m alt.; (-20.07444°, 44.67611°), tropical dry forest, 100 m alt.; Parc National de Kirindy Mite, 16.3 km 127° SE Belo sur Mer (-20.79528°, 44.147°), tropical dry forest, 80 m alt.; Makay Mts. (-21.21836°, 45.3106°), gallery forest on sandy soil, 510 m alt.; (-21.21985°, 45.32396°), gallery forest on sandy soil, 500 m alt.; (-21.22284°, 45.32477°), gallery forest on sandy soil, 490 m alt.; (-21.22336°, 45.32628°), gallery forest on sandy soil, 480 m alt.; (-21.25978°, 45.17158°), gallery forest with palm and pandanus, 430 m alt.; Forêt de Beroboka, 5.9 km 131° SE Ankidranoka (-22.23306°, 43.36633°), tropical dry forest, 80 m alt.; 29 km NNW Ranohira, Isalo N.P. (-22.31614°, 45.29625°), gallery forest, 490 m alt.; Parc National de Zombitse, 19.8 km 84° E Sakaraha (-22.84333°, 44.71°), tropical dry forest, 770 m alt.; Réserve Spéciale Kalambatritra, Ampanihy (-23.4635°, 46.4631°), montane rainforest, 1270 m alt.; Forêt de Mite, 20.7 km 29° WNW Tongobory (-23.52417°, 44.12133°), gallery forest, 75 m alt.; Beza-Mahafaly, 27 km E Betioky (-23.65°, 44.63333°), tropical dry forest, 135 m alt.; 10 km NW Enakara, Rés. Andohahela (-24.56667°, 46.81667°), rainforest, 425 m alt.; Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro (-24.7585°, 46.85367°), rainforest, 275 m alt.; Parc National d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro (-24.76389°, 46.75167°), montane rainforest, 900 m alt.; Réserve Privé Berenty, Forêt de Bealoka, Mandraré River, 14.6 km 329° NNW Amboasary (-24.95694°, 46.2715°), gallery forest, 35 m alt.; Forêt de Malaza, Mandraré River, 8.6 km 314° NW Amboasary (-25.00778°, 46.306°), gallery forest, 40 m alt.; Grand Lavasoa, 25.9 km W Tolagnaro (-25.08767°, 46.749°), rainforest, 450 m alt. Fianarantsoa. SE of Fandriana, Korikory, (-20.38444°, 47.66722°), 1670 m alt.; 7 km W Ranomafana National Park (-21.25°, 47.41667°), montane rainforest, 900 m alt.; 8 km E Kianjavato, Vatovavy Forest (-21.38861°, 47.94361°), lowland forest, 145 m alt.; Andrambovato along river Tatamaly (-21.50967°, 47.40762°), cultivated land (tavy), 984 m alt.; Parc National d’Isalo, Sahanafa River, 29.2 km 351° N Ranohira (-22.31333°, 45.29167°), gallery forest, 500 m alt.; 30 km NNW Ranohira, Isalo N.P. (-22.31722°, 45.29333°), gallery forest, 455 m alt.; R.S. Ivohibe 8.0 km E Ivohibe (-22.48333°, 46.96833°), montane rainforest, 1200 m alt.; Forêt d’Analalava, 29.6 km 280° W Ranohira (-22.59167°, 45.12833°), tropical dry forest, 700 m alt.; Forêt de Vevembe, 66.6 km 293° Farafangana (-22.791°, 47.18183°), rainforest, transition to montane forest, 600 m alt.; Réserve Speciale Manombo 24.5 km 228° Farafangana (-23.01583°, 47.719°), rainforest, 30 m alt.

COMOROS. Grande Comore. Grillé (-11.47578°, 43.34669°), montane rainforest, 995 m alt.; Karthala (-11.82699°, 43.4295°), montane rainforest, 1000 m alt. Mohéli. Ouallah (-12.29696°, 43.67392°), rainforest, 680 m alt.; (-12.32717°, 43.65952°), coastal scrub, 10 m alt. Anjouan. (-12.30537°, 44.45031°), along roadside, mango, banana, 500 m alt.

Remarks.

Mystrium mysticum females can be distinguished easily from other Mystrium females by a combination of a distinct longitudinal carina on the central portion of the labrum (Fig. 8A) or a single lateral spine on abdominal sternum VII (Fig. 8C), and the convexity of the anterior margin of the clypeus (Fig. 9C). In addition to these diagnostic characters, a combination of unsculptured and smooth areas on the ventral portion of the vertex (Fig. 11A) and the convexity of the anterior margin of the clypeus (as in Fig. 9C) separates Mystrium mysticum from the other Mystrium species in workers; in larger-sized alate queens (HW>2.00), simple setae on the pronotal dorsum (Fig. 9A) separate Mystrium mysticum from the queens of other species. For males, the small eye and small ocelli that are separated from each other by more than 3 × the maximum diameter of lateral ocellus (Fig. 26B), petiolar dorsum (Fig. 28C) and abdominal ter gum VIII (Fig. 28A) with rough and deep punctures, and well-developed basoventral expansion of aedeagus (Fig. 29A) separate Mystrium mysticum from the other Mystrium males in the Malagasy region.

Several diagnostic characters for Mystrium mysticum have been given in previous papers, (e.g. Forel 1891; Menozzi 1929); however, our comparative study revealed most of those characters could not cover the whole range of the morphological variation observed in Mystrium species. Here we propose a new set of diagnostic characters to identify Mystrium mysticum . The members of the mysticum species group display remarkable morphological variation in body size, relative mandibular length, relative shape of petiolar node in dorsal view, and body color. The available material revealed that the morphological variation within Mystrium mysticum is larger than the difference between Mystrium mysticum and the species it most resembles, Mystrium rogeri . For example, in order to separate Mystrium rogeri from Mystrium mysticum , Menozzi (1929) used the relatively wider shape of the petiolar node in dorsal view and the shape of setae (=hairs). However, in these two species the index of the petiolar node largely overlaps (PtI 153-183 vs. PtI 146-168) and the shape of setae varies gradually from spoon-shaped (small minor worker) to simple (large major worker) in single colony members. The separable characters for workers proposed in Forel (1895, 1899) cannot distinguish species in the mysticum species group. In fact, his syntypes for Mystrium stadelmanni consisted of minor workers of Mystrium mysticum and Mystrium rogeri (see below). In addition to the above variation, Molet et al. (2012) reported the presence of intercaste "mosaic monsters" in Mystrium rogeri , which are intermediate individuals having zero to three ocelli and mesosoma with incompletely developed wing sclerites. We confirmed the same intercastes in Mystrium mysticum as well, and these individuals will be keyed out with either workers or queens in our new identification key.

The redescription of the male of Mystrium mysticum in this study may be the first actual description of the male of this species. Previous male descriptions for Mystrium mysticum have been given in Forel (1891), Emery (1899), and Menozzi (1929). Of these descriptions, Forel (1891) and Menozzi (1929) are assumed to share the same specimen in ZMHB, Berlin. Although we have not examined the actual male specimen in ZMHB, the character description of the specimen does not fit the males in the mysticum species group. For example, the color of the abdomen is light brown in Forel (1891), and the eye occupies almost the entire lateral margin of the head in Menozzi (1929). Rather, those characters fit males in the voeltzkowi species group. We could not find useful information in Emery’s description ( Emery 1899); however, he did not mention any distinct differences from known males at that point. We have already confirmed that the male in the type series of Mystrium rogeri is actually a male of Mystrium oberthueri , so only males of the voeltzkowi species group, i.e. Mystrium voeltzkowi and Mystrium oberthueri were known in 1899. Hence, we consider all male descriptions for Mystrium mysticum in previous studies unreliable.

Clarification of the male-worker association in Mystrium mysticum in this study marks a great advance; however, some problems remain, especially in the level of confidence in diagnosing males. External characters, e.g. strong and rough punctures on the petiolar dorsum (Fig. 28C) and abdominal tergite VIII (Fig. 28A), distinguish the mysticum species group from the other Malagasy species. However, a specific character is neither possible nor practical to distinguish Mystrium mysticum males from Mystrium rogeri males. In fact, although we proposed the aedeagal character to separate Mystrium mysticum and Mystrium rogeri (Fig. 29), this difference cannot be confirmed without dissection. Unexpected remarkable variation could exist in the genital character as well as the other external characters. Further and more thorough taxonomic examinations of males are necessary to find diagnostic characters able to separate these two species.

The neotype of Mystrium mysticum was designated in this study because we concluded that the original type specimens of this "mysterious species" are lost. Our study revealed that no one examined any of the original type specimens after the publication of the original description by Roger (1862). According to that paper, the description was made based on two queens, and at least one of these was alate. The two queens must have been deposited in the museum in Paris; however, we could not find these queens in the MNHN collection. Although the depository of Mystrium mysticum was given again in the original description of Mystrium camillae Emery, 1889, Emery did not provide any further information. Santschi (1914) discussed the differences between Mystrium silvestrii and Mystrium mysticum when he described Mystrium silvestrii as a new species; however, he only provided information about workers, not queens. Forel described five species in the genus Mystrium from 1895 to 1899, and he noted in the latest description for Mystrium rogeri Forel, 1899 that he never examined the actual syntype of Mystrium mysticum . In spite of these difficulties, we still conclude that the species having queens with simple setae is Mystrium mysticum for the following four reasons: (1) we recognize only two species in the mysticum species group from a large number of Mystrium specimens accumulated from the whole range of the Malagasy region; (2) when Forel (1899) described Mystrium rogeri only queens with simple setae were known from Madagascar and were constantly identified as Mystrium mysticum ; (3) as Forel (1899) mentioned, Roger (1862) did not describe any specialized character for the queen’s setae; and (4) the queens of Mystrium rogeri have spatulate setae as Forel (1899) expected. In fact, the oldest queen specimen with spatulate setae, which is found in MNHN, was collected after Forel’s description (collection date is 1919).

A queen was chosen as the neotype because the original syntypes were two queens. The queen caste also offers a greater number of characters to distinguish this taxon from others in this species group. The location of the neotype was not based on a proximity to the original syntype series, but on the availably of a large collection series that included workers, males and queens.

Mystrium stadelmanni Forel, 1895 is synonymized with Mystrium mysticum in this study. Mystrium stadelmanni is described as a species having a longer petiole, narrower mesosoma, and less constricted abdomen compared with Mystrium mysticum ; however, our observations reveal that the boundary of this species is doubtful. None of the diagnostic characters above represent more than intraspecific variation observed in a series of workers from a single colony of Mystrium mysticum . The fact that syntypes of Mystrium stadelmanni consist of two minor workers of Mystrium mysticum and one minor worker of Mystrium rogeri certi fies that Forel’s determination ( Forel 1895) was not sufficient to clarify proper species boundaries in this group. In addition to the above, we should note one more important point. Forel (1895) determined specimens in ZMHB to be the new species Mystrium stadelmanni based on comparisons with workers of Mystrium rogeri that he probably misidentified as Mystrium mysticum . Forel believed his specimens, which were given by Emery ( Forel 1892), were Mystrium mysticum by that time ( Forel 1892; 1895); however, he described Mystrium rogeri in 1899 by using workers from the same collection. Our conclusion of a misidentification of Mystrium rogeri is supported by his description of a "longer petiolar node in Mystrium stadelmanni " which is correct if Forel (1895) studied the two workers of Mystrium mysticum in ZMHB. According to our measurements, Mystrium rogeri has the shorter petiolar node (PtI 153-183) while Mystrium mysticum has the longer node (PtI 146-168). Hence we synonymize Mystrium stadelmanni with Mystrium mysticum by designating a Mystrium mysticum worker in the syntypes of Mystrium stadelmanni as the lectotype. Consequently, the description of Mystrium stadelmanni in Forel (1895) is determined to be the first actual description for the Mystrium mysticum worker.

Updating species boundaries and providing easier identification tools for Mystrium mysticum will help reorganize and connect existing ecological and phylogenetic information to proper species names in Mystrium . This taxonomic revision permits us to update identifications in recent ecological publications ( Molet et al. 2009; Molet et al. 2007a; Molet et al. 2012), changing some of their identification from Mystrium rogeri to Mystrium mysticum : for example, five colonies (BLF00519, BLF04466, BLF04770, BLF06144, and BLF10994) out of 16 colonies listed as Mystrium rogeri in Table 1 of Molet et al. ( Molet et al. 2007a) and out of material for Molet et al. (2009) are those of Mystrium mysticum ; a queen of Mystrium rogeri in Figure 2 of Molet et al. (2009) and in Figure 1 and supplemental Figure A1 of Molet et al. (2012) is that of Mystrium mysticum . A worker of Mystrium mysticum was referred to as Mystrium ‘red’ in Figure 2 of Molet et al. (2009), as was supplemental Figure A1 of Molet et al. (2012) (see also in Mystrium mirror ). On the other hand, " Mystrium mysticum " in the material of three recent phylogenetic papers ( Brady et al. 2006; Kück et al. 2011; Moreau and Bell 2013; Ouellette et al. 2006) are Mystrium voeltzkowi , not Mystrium mysticum . " Mystrium mysticum " in Molet et al. (2009) is Mystrium shadow . Outellette et al. (2006) listed two more related undetermined specimens as Mystrium mysticum 2 and Mystrium mysticum 3; those are Mystrium voeltzkowi and Mystrium mirror , respectively.