Physodeutera (Toxoma) lokobensis, Moravec, 2021

Moravec, Jiří & Trýzna, Miloš, 2021, New or rare Madagascar tiger beetles- 23. Physodeutera (Toxoma) lokobensis sp nov., a new species close to Ph. (T.) conturbata Moravec, Ph. (T.) sulcoprothoracica (W. Horn) and Ph. (T.) dubia (Mařan), with revised type designation of the latter (Coleoptera: Cicindelidae), Zootaxa 5060 (2), pp. 151-182 : 164-173

Physodeutera (Toxoma) lokobensis sp. nov.

( Figs 46–89 , 106–107 )

Type locality. Madagascar Nord : Sambirano , Forest of Lokobe on the island of Nosy Be .

Type material. Holotype ♂ in NMPC, labelled: “N Madagascar, Nosy Be isl. / Lokobe Nat. Park, 26.-30.xi., / circuit Mitsinjo, 2019, / S 13°24´21´´; E 48°18´35´´, / 296 m, M. Trýzna leg.” . Allotype ♀ in CJVB with same label data . Paratypes. 1 ♂, 1 ♀ in CJVB , 2 ♀♀ in CCJM , 2 ♀♀ in CMTD , 1 ♀ in SDEI , 1 ♀ in NMPC , 1 ♀ in COSJ , 1 ♀ in MHCW with same label data. 1 ♂, 1 ♀ in JWCW : “ Madagascar, Nosy Be / Sambirano / forêt de Lokobe, 5.– 9.12.2001, I. Andrew, / V. Dolin & R. Andreeva leg.” [printed] .

Differential diagnosis. Physodeutera (Toxoma) lokobensis sp. nov. is generally largest of the four species treated here. It shares the whitish-testaceous (to yellow-testaceous) area on the antennomere 4 ( Figs 50–51 ) with Ph. (T.) dubia , but the antennae are notably longer towards the body length. Moreover, Ph. (T.) dubia differs in having its aedeagus apex more simply and regularly rounded, while the aedeagus apex of Ph. (T.) lokobensis sp. nov. ( Figs 64–66 ) is almost triangular-topped (dorsally obliquely sloped towards small dorsal emargination); the difference is notably obvious when the aedeagus (the same of the holotype) was slightly turned ( Fig. 65 ). The internal sac is rather similar to that in Ph. (T.) conturbata , yet the satellite piece moved on the opposite side of the arciform piece that is straighter in the new species ( Figs 67–68 ). However, Ph. (T.) conturbata possesses the antennomeres 2–4 metallic black, elytra deep cyaneous-violaceous ( Figs 19–23 ), and its aedeagus has shorter, wider, and regularly rounded apex ( Figs 8–9 ).

Physodeutera (T.) sulcoprothoracica is immediately distinguished from the new species and from the two species treated above by its elytral disc notably shadowed with diffusing, well noticeable (depending on light-angle) velvety black area which extends towards elytral outer margins. Moreover, its elytral punctation on posterior area of the elytral disc is much finer, almost effaced. The only examined and hitherto known male (CCJM ex APCA) has much shorter and wider apical portion of its aedeagus towards wider and rounded apex; moreover, the internal sac contains stick-like (not arcuate) arciform piece and a well-defined voluminous piece with apical tooth ( Figs 100–101 ). Furthermore, the body of Ph. (T.) sulcoprothoracica is notably larger, elytral apex in male slightly emarginated towards small sutural spine, antennomere 4 brownish with blackened apical half, its male labrum has notably anteriad-prolonged anteromedian lobe with more distinct teeth, which are even more acute and of almost equal length in female labrum (see illustrations of the relevant characters in the monograph ( Moravec 2002a, figs 323–329).

Etymology. Named after the Parc National de Lokobe, the type locality of the new species.

Description. Body ( Figs 46–47 , 75–76 ), medium-sized to large, 9.50–10.8 (HT 9.6) mm long, 2.85–3.30 (HT 2.90, AT 3.20) mm wide, rather uniformly bright green-blue, with only lateral areas violaceous.

Head ( Figs 48–49 , 79 ; ventrally Fig. 59 ) with large eyes, almost as wide as body, width 2.90–3.30 mm.

Frons metallic deep blue-green, convex, and almost smooth in median juxtaclypeal area, finely longitudinally striate-wrinkled laterally, fluently passing over blunt frons-vertex fold into vertex; supraantennal plates flat, indistinct.

Vertex black-cyaneous with deep, transverse anterior impression reaching eyes, anteromedian area very finely irregularly rugulose, median surface behind the transverse impression coarsely and irregularly (mostly longitudinally) striate-rugulose, striae wavy, often irregularly fragmented in centre; posterior and occipital areas irregularly vermicular or zigzag rugulose; large juxtaorbital areas with two or three impressions, covered with denser, irregularly parallel rugae, which are deformed by the impressions; sublateral parallel rugae divergent when running onto temples; posteromedian and occipital areas covered with vermicular or irregularly wavy and fragmented rugae.

Genae deep blue with violaceous lustre, finely wrinkled.

Clypeus bright or deep metallic green-blue, often with violet lateral margins, surface almost smooth, indistinctly coriaceous-wrinkled.

Labrum ivory to ochre-testaceous with metallic black-blue basolateral areas of central convexity (more expanded in female); male labrum rather short, length 1.05–1.10 mm, width 1.20–1.25 mm, lateral margins conically attenuated towards indistinct lateral indentations, and prominent, acute anterolateral teeth; tridentate anteromedian lobe in HT short, subtruncate with right-angled teeth and bluntly indicated median tooth ( Fig. 53 ), but in one male PT (JWCW) prolonged anteriad and with rounded lateral margins ( Fig. 81 ); female labrum ( Figs 54–55 , 82 ) almost as long as wide, length 1.25–1.45 mm, width 1.35–1.45 mm; anteriad-prolonged tridentate anteromedian lobe with prominent acute or subacute teeth on either side of protruding medial tooth with obtuse apex.

Mandibles ( Figs 48–49 , 79–80 ) normally shaped, subsymmetrical, with four teeth and basal molar, reddishbrown with blackened margins and apices of teeth; inner teeth gradually smaller towards basal molar, fourth tooth markedly distant from third tooth.

Palpi. Maxillary palpi ( Figs 56 , 96 ) ivory-testaceous with terminal palpomere brownish in male, with last two palpomeres brownish in female; labial palpi in male with brown terminal palpomere, their penultimate (longest) palpomere moderately dilated with subparallel lateral margins, ochre-testaceous in male ( Figs 57 , 97 ), brownish in female ( Figs 58 , 98 ).

Antennae ( Figs 46–47 , 50–52 , 75–78 ) markedly long, almost or entirely reaching anteapical angles of elytra, scape with only apical seta; coloration sexually distinctly dimorphic: male with scape whitish, ivory or ochre-yellow; antennomeres 2–3 metallic black; antennomere 4 black with median area or two basal thirds ivory to ochreyellow, antennomere 5 brownish testaceous, 6–11 brownish, gradually darkened to blackened.

Thorax. Pronotum ( Figs 60–62 , 83–84 ), moderately elongate, length 2.10–2.30 mm, width 1.70–1.85 mm; coloration bright green-blue with violaceous lustre within sulci and on lateral areas, rarely with indistinct bronze lustre in middle; anterior and posterior sulci well pronounced, anterior lobe markedly narrower than posterior lobe and disc; lateral margins of disc convex, usually moderately attenuated anteriad; notopleural sutures almost invisible from above; median line narrow, often partly merging with surface sculpture; surface of disc distinctly but rather shallowly transversely rugulose, rugae irregularly wavy and occasionally anastomosing, coarser on median area, much finer and shallower on sublateral areas, while lateral areas with wider, short, transverse-parallel stria-like rugae; posterior lobe bordered with sharply delineated basal rim, dorsolateral bulges distinctly pulvinate, fluently passing to irregularly and shallowly wrinkled median area; proepisterna variably bright green-blue, deep blue or violet-blue, finely but rather distinctly parallel striate (striae passing over notopleural sutures from lateral margins of pronotal disc); mesepisterna shiny deep blue green with strong violaceous lustre, their surface finely coriaceousasperate in male, mostly nearly smooth in female, female mesepisternal coupling sulci in form of deep longitudinal furrow, which is deeper dorsally, and with variably recognizable, indistinct, rarely more distinct but shallow medioventral pit; metepisterna bright blue-green with chatoyant violet lustre, surface finely asperate; ventral sterna ( Fig. 63 ): iridescent green-blue with violet lustre, prosternum finely transverse striate, mesosternum irregularly wrinkled, metasternum smooth very shallowly wrinkled along middle.

Elytra ( Figs 69–73 , 86–89 ) elongate, 5.50–6.30 mm long; humeral impressions short yet deep, basodiscal convexity and discal impression distinct, the elytral disc becomes distinctly convex below the impression; apical impressions moderate; outer lateral margins slightly dilated above the middle and usually very slightly at arcuate anteapical angles (more often so in female), apices almost subacute in male, rounded in female, mostly without emargination towards indistinct blunt sutural spine; surface punctate throughout, punctures coarser on anterior elytral third, few largest punctures within humeral impressions and on basodiscal convexity, occasionally anastomosing in chains; posterior half of elytral disc and posterior declivity with much finer and spaced punctures; elytral coloration rather bright green-blue, (in some females and the male (JWCW) with violet lustre); central-discal area sometimes with indistinct shade; surface glabrous except for usual, few sensory setae scattered mostly on basodiscal convexity and sparse, short white setae scattered along juxtaepipleural area; elytral maculation consisting of large, whitish or ivory yellow humeral macula in male; female elytra entirely immaculate.

Abdomen ( Fig. 63 ). Ventrites metallic black-blue, with strong green-blue lustre.

Legs sexually dimorphic in coloration; male pro- and mesocoxae ivory to pale ochre with testaceous apices, glabrous except for easily abraded subapical seta, metacoxae metallic green-blue and testaceous apices; female coxae metallic black-blue with greenish lustre; trochanters ivory-yellow; male profemora distinctly bicoloured, blackbrown dorsally, yellow-ochre to ochre-testaceous ventrally and on subapical dorsal area; mesofemora with yellowtestaceous ventral area less expanded and with testaceous subapical spot; metafemora black with testaceous base. Female legs much darker, femora almost entirely black, profemora sometimes with testaceous subapical spot.

Aedeagus ( Figs 64–66 ) elongate, length 2.30–2.40 mm, width 0.40–0.50 mm, almost straight, only moderately dilated in middle (except for more voluminous, 0.55 mm wide aedeagus of the aberrant male in JWCW, Fig. 85 ); apical portion of ventral margin slightly directed ventrad, dorsally conically attenuated towards apex, which is rounded but almost triangular-topped (ventrally moderately arcuate and dorsally obliquely sloped towards indistinct emargination); the difference from other species treated here is notably obvious when the aedeagus (the same of the holotype) was slightly turned ( Fig. 65 ); internal sac ( Figs 67–68 ) somewhat similar to that in Ph. (T.) conturbata yet the satellite piece is placed ventrad from almost straight and thinner arciform piece.

Variability. The dorsal body coloration varies, some females are prevailingly blue. One female (CCJM) has its labial palpi anomalously ivory-testaceous as in the males. The two previously caught adults (JWCW) ( Figs 75–89 ), have more rounded pronotal disc with finer surface sculpture, and the male has much darker antennomere 4 with only paler, brownish basal half. Unfortunately, the aedeagus apex of the male (JWCW) is broken ventrally ( Fig. 85 ), thus its original shape is not exactly known (its internal sac was not cleared in order to avoid further damage of the aedeagus). Despite the above-mentioned differences, we have considered the two JWCW specimens conspecific and included them, with some hesitation, into the paratype series. Nonetheless, as also the aedeagus (apart from its broken apex) is wider and has somewhat different shape, the two JWCW specimens may represent another undescribed species.

Distribution ( Fig. 106 , 107 ). Physodeutera (Toxoma) lokobensis sp. nov. is obviously a very rare species known only from the type locality, the forest of Lokobe on the island of Nosy Be, phytogeographically Sambirano. The primitive evergreen forest is now protected within the Parc National de Lokobe, managed by Madagascar National Parks. The adults caught by the second author in the area of the Mitsinjo Circuit within the national park occurred on larger living trees (> 20 cm in diameter). On sunny days, the adults ran and flew very quickly along the trunk bark of the trees up to four meters above the tree bases, while on the next day, after a heavy overnight rain, they ran and flew rather slowly and only up to two metres above the tree bases. It is interesting that the twelve type specimens were caught by the second author in the forest of Lokobe 20 years after the discovery of the male and female (JWCW). The new species inhabits the forest of Lokobe sympatrically with Physodeutera (Axinomera) rectipenis (W. Horn, 1934) , yet adults of the latter forage on boulders scattered on the forest bed along a semidried brook. As in other species of the genus, developmental stages are unknown.

Physodeutera (Toxoma) lobicornis nosybensis Moravec, 2000 also occurs in Nosy Be but not in the forest of Lokobe; it was described from a degraded forest near Ambatozavavy (also spelled “Ambatozavary” in maps), situated at the Ambatozavavy Bay, northeast of the Parc National de Lokobe (see Moravec 2000, 2002a). Also recently caught specimens come only from the area of Ambatozavavy (Michio Hori, pers. com.).

Figures

FIGURE 106. Distribution of four Physodeutera (Toxoma) species in northern part of Madagascar. Adopted from the map of preserved areas in Madagascar according to Moat & Smith (2007), adapted and published previously (Moravec 2010) under permission of the trustee of the Royal Botanic Gardens, Kew, GB.

FIGURE 107. Above: overall view on the Parc National de Lokobe, Nosy Be, Madagascar. Bellow: biotope of Physodeutera (Toxoma) lokobensis sp. nov., in the forest of Lokobe (Mitsinjo circuit). Photographed by Miloš Trýzna.

FIGURES 46–47. Physodeutera (Toxoma) lokobensis sp. nov., habitus, type locality. 46—♂, 9.6 mm, HT (NMPC); 47—♀, 10.3 mm, AT (CJVB). Bar = 1 mm.

FIGURES 48–59. Physodeutera (Toxoma) lokobensis sp. nov., type locality. 48–49—head: 48—♂, HT (NMPC); 49—♀, AT (CJVB); 50–52—antennomeres 1–5: 50—♂, HT; 51—♂, PT (CJVB); 52—♀, AT; 53–55—labrum: 53—♂, HT; 54—♀, AT; 55—♀, PT (CJVB); 56—apex of lacinia with galea and maxillary palpus, ♂, HT; 57–58—labial palpus: 57—♂, HT; 58—♀, PT (CJVB); 59—head in ventral view, ♂, PT (CJVB). Bars = 1 mm.

FIGURES 60–68. Physodeutera (Toxoma) lokobensis sp. nov., type locality. 60–62—pronotum: 60—♂, HT (NMPC); 61—♀, AT (CJVB); 62—♀, PT (CCJM); 63—ventral view, ♂, PT (CJVB); 64–66—male aedeagus: 64—HT; 65—HT, the same aedea- gus slightly turned; 66—PT (CJVB); 67–68—ditto, showing internal sac in left and right lateral view. Bars = 1 mm.

FIGURES 69–74. Physodeutera (Toxoma) lokobensis sp. nov., type locality. 69–73—elytron: 69—♂, HT (NMPC); 70— ditto, detail of elytral disc; 71—♀, AT (CJVB); 72—♀, PT (CCJM); 73—ditto, detail of posterior elytral area; 74—labels of HT. Bars = 1 mm.

FIGURES 75–78. Physodeutera (Toxoma) lokobensis sp. nov., two aberrant adults, forest of Lokobe, PT (JWCW). 75–76—ha- bitus: 75—♂, 9.5 mm; 76—♀, 10.3 mm; 77–78— antennomeres 1–5: 77—♂; 78—♀. Bars = 1 mm.

FIGURES 79–85. Physodeutera (Toxoma) lokobensis sp. nov., two aberrant adults, forest of Lokobe, PT (JWCW). 79—head, ♂; 80—mandibles, ♂; 81–82—labrum: 81—♂; 82—♀; 83–84—pronotum: 83—♂; 84—♀; 85—male aedeagus (with broken apex). Bars = 1 mm.

FIGURES 86–89. Physodeutera (Toxoma) lokobensis sp. nov., two aberrant adults, forest of Lokobe, PT (JWCW), elytron. 86—♂; 87— ditto, detail of elytral disc; 88—♀; 89—ditto, detail of posterior area of elytral disc. Bars = 1 mm.

FIGURES 19–23. Physodeutera (Toxoma) conturbata Moravec, from type locality, all in SDEI. 19–22—elytron: 19—♂, HT; 20—ditto, detail of elytral disc; 21—♀, PT; 22— ditto, detail of posterior area of elytral disc; 23—labels of the holotype. Bars = 1 mm.

FIGURES 1–9. Physodeutera (Toxoma) conturbata Moravec, from type locality, all in SDEI. 1—habitus ♂, 9.2 mm, HT; 2– 3—Antennomeres 1–5: 2—♂, HT; 3—♀, PT; 4–5—♀, maxillary palpus: 4—dorsal view; 5—ventral view; 6–7—labial palpus: 6—female, PT; 7—male, HT (adopted from Moravec 2002a); 8–9— male aedeagus showing internal sac, HT. Bars = 1 mm.

FIGURES 93–101. Physodeutera (Toxoma) sulcoprothoracica W. Horn. ♂, “Ambre” (CCJM ex APCA). 93—head, ibid.; 94– 95—labrum: 94—♂, ibid.; 95—♀, LT (SDEI), adopted from Moravec (2002a); 96—maxillary palpus, ♂, “Ambre” (CCJM ex APCA); 97–98—labial palpus: 97—♂, ibid.; 98—♀, LT (SDEI), adopted from Moravec (2002a); 99—pronotum, ♂, “Ambre” (CCJM ex APCA); 100–101—aedeagus showing internal sac in its left and right lateral view. Bars = 1 mm.