Terataner nymeria

Terataner nymeria is only known from its type locality, the Analamerana Special Reserve, which is located in the northern tip of Madagascar. The available material was collected in a dry tropical forest habitat at an altitude of 225 m. The new species seems to live on vegetation and nest in dead branches. The strongly elongate and slender body might be an adaptation to the life in sticks and branches resembling other arboreal stick-inhabiting ants, such as Tetraponera. (Hita Garcia et al. 2017)

Identification
Hita Garcia et al. (2017) - The following character combination distinguishes T. nymeria from the remainder of the genus: in full-face view head comparatively thin (CI 87-91) with weakly convex sides, broadest at eye level, conspicuously narrowing towards posterior head margin before significantly broadening again into well-developed angulate posterodorsal corners; petiole and postpetiole without extremely long, dorsal spines; postpetiole in profile strongly rounded cuneiform, anterior face relatively long and straight leading to rounded dorsum, dorsal apex slightly pointing backwards; postpetiole in anterodorsal view arch-like leading towards a rounded apex; dorsum of promesonotum longitudinally porcate; petiole and postpetiole conspicuously sculptured.

This new species is distinguishable from all other Terataner on the basis of the slender and elongate gestalt. There is no other Terataner that comes close, especially when considering the relatively thin and elliptic head shape in full-face view. Some undescribed species from Madagascar have thinner heads compared to most other congeners, but they never have the strongly narrowed posterior head as seen in T. nymeria. The same is true for the strongly elongate waist segments, which are not found in any other Terataner. The phylogenetic affinities of T. nymeria are unknown at present, but on the basis of its peculiar morphology it seems that it is not closely related to any other currently described Terataner species.

Intraspecific variation. Considering that T. nymeria is only known from three collections from the same locality on the same day, it is not surprising that intraspecific variation is neglectable.

Distribution based on Regional Taxon Lists
Malagasy Region: Madagascar.

Ergatoid Queen
No alate or dealate queens occur in any of the Malagasy Terataner species. Colonies are almost always monogynous with a single reproductive, an ergatoid queen, that differs only slightly from corresponding workers. This reproductive lacks both wings and ocelli, and has an almost normal, worker-like thorax. The ergatoid queen is externally distinguishable from workers only by very subtle characters (e.g., small spines or tubercles below the mesonotal groove; a unique pattern of rugae on the sides of the pronotum) that vary among species. Ergatoid queens have ovaries bearing two to three ovarioles.

Nomenclature

 * . Terataner nymeria Hita Garcia, in Hita Garcia, Fischer, et al. 2017a: 21, figs. 1, 3C, 3F-G, 8-10, S3 (w.) MADAGASCAR.
 * Type-material: holotype worker, 11 paratype workers.
 * Type-locality: holotype Madagascar: Antsiranana, Res. Analamerana, 16.7 km. 123° Anivorano Nord, -12.80467°, 49.37383°, 225 m., BLF11298, 3.xii.2004, tropical dry forest, low vegetation (B.L. Fisher); paratypes: 1 workers with same data or BLF11305.
 * Type-depositories: CASC (holotype); BMNH, CASC, HLMD, MCZC (paratypes).
 * Distribution: Madagascar.

Worker
(N = 12). HL 1.08-1.21 (1.13); HW 0.98-1.08 (1.01); FLD 0.36-0.45 (0.41); SL 0.75-0.86 (0.81); EL 0.26-0.30 (0.28); OMD 0.35-0.38 (0.37); WL 1.56-1.76 (1.62); PH 0.52-0.62 (0.55); PW 0.81-0.97 (0.97); PML 0.75-0.86 (0.80); PDH 0.54-0.61 (0.57); MFL 0.98-1.10 (1.02); MFW 0.31-0.36 (0.32); PTH 0.40-0.49 (0.43); PTL 0.47-0.51 (0.49); PTW 0.27-0.33 (0.29); PPH 0.50-0.58 (0.52); PPL 0.47-0.56 (0.50); PPW 0.48-0.58 (0.52); CI 87-91 (90); FLI 36-43 (40); OI 26-29 (28); OMI 70-83 (77); SI 75-82 (80); DMI 52-56 (54); LMI 33-35 (34); PMI 106-113 (110); PPDI 93-101 (97); MFLI 99-104 (101); MFI 306-332 (316); LPeI 105-123 (115); DPeI 55-64 (58); LPpI 91-99 (95); DPpI 99-111 (104).

HEAD. Masticatory margin of mandible with five or six teeth, usually with large, acute apical tooth followed by a smaller, acute subapical tooth, remaining three to four teeth conspicuously smaller and less acute, very often worn down to small, rounded remnants. Palp formula 5,3. Head in full-face view comparatively thin, around 1.1 times longer than wide (CI 87-91), sides of head weakly convex, broadest at eye level, conspicuously narrowing towards posterior head margin before significantly broadening again into well-developed angulate posterodorsal corners; in full-face view posterior head margin straight to very weakly convex; anterior clypeal margin medially with distinct, broad but small impression; frontal carinae very well developed, from level of posterior frontal lobes to shortly before posterior head margin subparallel and moderately raised, posteriorly initially sharply divergent and much more raised, crest-like, merging with strong costate sculpture and separating side of head from posterodorsal head. Antennal scrobes present but very shallow and narrow; antennal scapes moderately long, not reaching posterior head margin (SI 75-82). Eyes of moderate size (OI 26-29), malar distance around 1.2 to 1.4 times longer than maximum eye length (OMI 70-83).

MESOSOMA. In lateral view mesosoma relatively low and elongate (LMI 33-35), promesonotum moderately convex, propodeal dorsum weakly convex, propodeum usually weakly higher than propodeum (PPDI 93-101) and both separated by metanotal groove; promesonotal suture present laterally and completely absent dorsally, metanotal groove well developed, moderately deep but relatively narrow, in profile anterodorsal propodeal margin weakly overhanging metanotal groove. Pronotum strongly marginate from lateral to dorsal mesosoma; anterodorsal corners shaped into sharply rectangular, thin, and transparent lamellae, lamellae becoming weaker and less transparent posteriorly and ending at promesonotal suture; mesonotum not marginate and armed with pair of short, thin, and acute teeth laterally; in dorsal view promesonotum around 1.1 times wider than long (PMI 106-113). Propodeum moderately marginate from sides to dorsum and sides to propodeal declivity; propodeum unarmed, posterodorsal corners angulate; propodeal lobes very well developed, broadly triangular, and blunt; in dorsal view anterodorsal margin of propodeum straight. All femorae strongly swollen medially; length of metafemur around 3.1 to 3.3 times longer than maximum width (HFI 306- 332).

WAIST SEGMENTS & GASTER. In profile petiolar node relatively low and conspicuously triangular cuneiform; in anterodorsal view node with two short but distinct, blunt dorsal spines; in dorsal view approximately barrel-shaped, sides mostly straight and converging anteriorly towards peduncle; in profile petiolar node around 1.1 to 1.2 times longer than high (LPeI 105-123); in dorsal view node between 1.5 to 1.8 times longer than wide (DPeI 55-64). In profile postpetiole strongly rounded cuneiform, around 1.0 to 1.1 times higher than long (LPpI 91-99), anterior face relatively long and straight leading to rounded dorsum, dorsal apex slightly pointing backwards; in anterodorsal view postpetiole arch-like leading towards a rounded apex; in dorsal view around as wide as long to 1.1 times wider than long (DPpI 99-111). Gaster not extremely enlarged.

SCULPTURE. Mandibles unsculptured, smooth, and shiny; clypeus and anterior head margin only weakly sculptured, usually with few weak, longitudinal carinulae; cephalic dorsum between frontal carinae conspicuously longitudinally costate, thickness and height of costae increasing from anterior margin near clypeus to posterior head; antennal scrobe mostly unsculptured, smooth and shiny; lateral head longitudinally costate, weaker developed anteriorly and between eyes and frontal carinae, becoming much stronger posteriorly. Most of mesosoma very conspicuously porcate; lateropronotum and dorsal promesonotum longitudinally porcate; lateral propodeum diagonally porcate, dorsum anteriorly close to metanotal groove transversely porcate, median area of dorsum and propodeal declivity unsculptured, smooth and shiny; procoxae mostly longitudinally rugulose, remainder of legs unsculptured, smooth and shiny. Peduncle unsculptured, smooth and shiny; both waist segments strongly costate/porcate: sides of petiolar node diagonally costate, most costae crossing anterior side of node from one side to another; dorsum of node longitudinally costate/porcate; postpetiole longitudinally costate. Base of first gastral tergite costulate, remainder of tergite with conspicuous reticulate-punctate microsculpture. Apart from first gastral tergite body with very weak to absent microsculpture.

PILOSITY & PUBESCENCE. Whole body covered with abundant, short to moderately long, fine, suberect to erect pilosity; pilosity on gaster noticeably shorter; mandibles with appressed to decumbent hairs; antennal scapes with abundant, shorter, decumbent to suberect hairs on all sides plus row of widely spaced, much longer, erect hairs on anterior margin. Pubescence strongly reduced to absent.

COLORATION. Most of body very dark brown to black.

Type Material
Holotype, pinned worker, Madagascar, Antsiranana, Reserve Analamerana, 16.7 km 123˚ Anivorano-Nord, -12.80467˚, 49.37383˚, 225 m, tropical dry forest, on low vegetation, collection code BLF11298, 3.XII.2004 (B.L. Fisher) (: CASENT0053630).

Paratypes, eleven pinned workers: eight workers with same data as holotype except collected ex dead branch above ground, bamboo and collection code BLF11305 (: CASENT07901 28; CASC: CASENT0107418; CASENT0746053; CASENT0746054; CASENT0746055; CASENT 0790129; : CASENT0790127; : CASENT0790130); and one worker with same data as holotype except collected by beating low vegetation and collection code BLF11306 (CASENT 0107444).

Cybertype, volumetric raw data, 3D PDF, and 3D rotation video of the physical holotype (CASC: CASENT0053630) plus montage images illustrating head in full-face view, profile and dorsal views of the body. The data of the holotype is deposited in Dryad (http://dx.doi.org/10.5061/dryad.sk6s0) and can be freely accessed as virtual representation of the physical holotype.

Etymology
The new species is named after the fictional, female direwolf from Georg R. R. Martin's epic fantasy series “A Song of Ice and Fire” referring to the ferocious, predatory nature of the new species. The species epithet is a noun in apposition and thus invariant.

References based on Global Ant Biodiversity Informatics

 * Hita Garcia F., G. Fischer, C. Liu, T. L. Audisio, G. D. Alpert, B. L. Fisher, and E. P. Economo. 2017. X-Ray microtomography for ant taxonomy: An exploration and case study with two new Terataner (Hymenoptera, Formicidae, Myrmicinae) species from Madagascar. PLoS ONE 12(3): e0172641.