Neoponera Emery, 1901 (Hymenoptera: Formicidae) revisited: 1. The N. laevigata species-group

The Neotropical Neoponera laevigata species-group is revised. This group can be separated from others in the genus by means of a new key to Neoponera species–groups. We propose two new species, N. gojira sp. nov., and N. mashpi sp. nov., the latter being usually confused with its putative sister species, N. laevigata (Smith). This small, termite-feeding species-group now contains five taxa. We provide comprehensive redescriptions for the known castes of the three previously known species, together with images and notes about intraspecific morphological variation, biology, and distribution of the taxa here treated. We reveal the external morphological features of the genitalia of the known males. This is the first contribution en route to a better understanding of the systematics of this charismatic ponerine genus.


Introduction
Neoponera Emery is a behaviorally and morphologically diverse group of ground and mostly arboreal, solitary hunting ants which inhabit exclusively tropical and subtemperate regions of the Americas (Schmidt & Shattuck 2014). A few species, like N. apicalis (Latreille) and N. villosa (Fabricius) are recognizable at a glance, whereas most of the remaining taxa require detailed examination, and little is known about the natural history of most species. Schmidt and Shattuck (2014) provide a good primer about various aspects of their social and reproductive behavior, nesting ecology, A. Troya and J. Lattke / Insect Systematics & Evolution (2022) DOI 10.1163/1876312X-bja10034 A3L Abdominal segment III length. In dorsal view, the distance between the anterior and posterior margin of abdominal tergite III. A3W Abdominal segment III width. In dorsal view, maximum width of tergite of abdominal segment III. A4L Abdominal segment IV length. In dorsal view, the distance between the anterior and posterior margins of abdominal tergite IV. A4W Abdominal segment IV width. In dorsal view, maximum width of tergite of abdominal segment IV. TLa Total accurate length of body: HL + WL + PL + A3L + A4L. TLr Total relative length of body: HL + WL + PL + GL.

Indices
The following ratios were multiplied by 100:

Distribution maps
We used the open-source package Quantum GIS 2.18 to generate maps of distribution, based on coordinates written on specimen labels and from notes recorded in field books from the authors. All localities were transformed to decimal degrees. Google Earth (Google.com 2021), and the online conversion tool NCAT (National Geodetic Survey 2021) were used as information sources to check and confirm locality names. In regards to unusual records without specific localities which were below the administrative circumscription of state, province, or similar, the center of the next most specific site was georeferenced. We also included in the maps, appropriately labeled AntWeb records of N. commutata Roger and N. marginata Roger for which we were confident on their taxonomic and geographic information. Selected AntWeb records belonging to these two common taxa were identified by ant taxonomists, for example, Alex Wild, Bryan Fisher, Flavia Esteves, Jack Longino. Due to their unique morphology within Neoponera these are easily identifiable species, showing low morphological variation across their range of distribution, thus making misidentifications unlikely. Jack Longino (lead researcher in several Central American ant survey campaigns) aided in examining diagnostic characters from representative AntWeb records of N. laevigata from Costa Rica and Panama. This was required since we believe most N. laevigata specimens from these countries are confused with N. mashpi sp. nov. (see below under each species treatment). and "p" -pectinate, for example, 2(1s, 1p) which means that two spurs are present, one simple, and 1 pectinate.

Descriptions and material examined
We used the computer package DELTA (Dallwitz 1980) for encoding the attributes of species (character states) and subsequently translating them into natural language (RTF-rich text files) taxonomic descriptions, requiring only posterior final editing from the user. Transcription of the collection information of examined specimens, from a spreadsheet data matrix to correspondent RTF file, was mostly done using AUTOMATEX (Brown 2013).
Under "status as species" in the taxonomic history of each species, only their first appearance in the taxonomic literature is cited, as well as relevant redescriptions which are usually part of revisionary studies. The user can access the complete history of literature mentions for each species on the Online Catalogue of the Ants of the World -AntCat (AntCat by B. Bolton 2022).
Although the "material examined" section represents the complete set of recorded localities in this work, it does not contain the whole list of specimens/locality. We have included, however, at the beginning of each section the total number of specimens/caste/species examined. The reader can freely download from the Supplementary Material S4 the full list of specimen records containing collection and geographic information, most of which have unique specimen identifiers.
The , and [institution where the material is located, usually represented by a unique specimen identifier composed of capitalized letters and numbers]. The format for "Other material examined" is essentially equal except that unique specimen identifiers are not shown, but museum acronyms are.

Relationships among taxa
In this work we have chosen to add brief comments about the relationships between species within the N. laevigata group, and also between Neoponera and a newly recognized clade (see further below). Our statements are grounded on prior studies by Schmidt (2013) and Branstetter and Longino (2022), as well as on molecular-based preliminary results (A. Troya, J. Longino, M. Branstetter et al. unpublished) which are part of a larger ongoing collaborative study about the phylogeny of the genus Neoponera which will be published separately. This additional information is not to be treated as conclusive, nevertheless we think it adds support to our morphology-based arguments around the species hypotheses shown here.

Species delimitation
We follow the biological species concept sensu Mayr (1942) by considering species as population assemblages which are under presumed continuous gene flow. Since we did not directly assess this process, our criterion for species delimitation was to detect a continuum of expected character gradations lacking discontinuities among distinctive populations. In sympatry, interspecific gene flow can obscure populational distinctness (Arnold 1992;Field et al. 2011). In such case species can be detected by consistently grouping morphologically similar individuals assumed to belong to structured populations, which is a proxy for gene flow (Wild 2005).
The species hypotheses presently proposed, especially the newly described taxa, should be evaluated in further studies using additional information criteria, e.g., population genetics, natural history observations. These will add more objective evidence aiding in the discrimination of species boundaries among cryptic populations which may potentially be under differing selective pressures that are suspected, yet untested in this study.

Morphology
The following few terms are defined in detail to aid in the comprehension of this treatment. In regards to the comparatively complex male external morphological terminology the user can consult Boudinot (2013Boudinot ( , 2015, and HAO. Mesosternal (msp) and metasternal (mtp) processes (Figs 16d,19f,20e,22f,25e): also called articular coxal processes of the meso-and metapectus (see HAO): in ants, especially in the Ponerinae, a pair of ventrally directed extensions of the cuticle, usually with acute or blunt apices, emerge medially from the meso-and metasternon. Each of these emerge either immediately posterior-or just around the mesosternal and metasternal pits, respectively, approximately in between the coxal cavities. When present, these structures may vary considerably in size, external shape and orientation, though probably the most generalized is the triangular-shaped form with acute apices, easily discernible in posteroventral view when the posterior coxae are oriented laterally (not posterad as is usual). In Neoponera, their presence/absence, as well as their size and shape, are considered useful character-states aiding in species and species-groups distinction.
Pheromone venting canal (fvc) at the metapleural gland opening (Fig. 2): a depression of the cuticle located just anteriorly and below the orifice of the metapleural gland. This structure is present in virtually all Ponerinae and is located at the posteroventral margin of the propodeum. It varies in depth, being clearly concave, e.g., Neoponera commutata, to shallowly concave, e.g., Megaponera analis (Latreille). In most Neoponera it is flanged posteriorly by a cuticular flap, and dorsally by an excavated edge of the propodeum.
Anteroventral petiolar carina: a transverse ridge located near the ventral part of the anterior face of tergite of A2 (petiolar node), relatively close to the petiolar condyle which articulates with the propodeal foramen, as seen in dorsal view. In Ponerinae, when well-developed, this carina usually extends anterolaterad forming salient, either acute or blunt projections ("spiracular horns" in . Two variations are recognized: complete and incomplete carina. In the first state, the carina runs horizontally uninterrupted connecting both anterolateral nodal projections, e.g., Leptogenys luederwalti Forel, Loboponera nasica (Santschi). When it is incomplete, it disappears medially on the petiolar tergite, leaving the nodal projections virtually unconnected, in which case these usually are notoriously salient. An incomplete carina is the norm in Neoponera, e.g., N. crenata Roger, N. commutata, N. villosa, as well as in most ponerines. In Neoponera, a complete carina is found for example, in N. apicalis, N. rostrata (Emery).

Morphometrics
The PCA circular plot (Fig. 1,Supplementary Material S2) shows that most variables are positively correlated and also most of them are well-represented in the factor map (= they are close to the circumference margin), which means they are important in explaining the total morphometric variance (ca. 92%) represented in the first two dimensions of the ordination. The quality of representation of the variables is calculated using the square cosine and expressed through the eigenvalues as shown in Fig. 1 for dimensions 1 and 2 (Supplementary Material S4). On the other hand, the NMDS ordination was supported by a robust stress value of 0.01826, which according to the criteria of Kruskal (1964), indicates a strong correlation between ranked observed dissimilarities and ordinated distances. This value also suggests that our observations, i.e., measured specimens, are not grouped randomly. The graph shows a small cluster of convergent observations representing most taxa, except N. commutata, which is suggestive of morphometric similarity as these share a significant fraction of their morphospace. As expected, this is evident between N. mashpi sp. nov. and N. laevigata since their measured characters are more similar in size between each other than with the remaining species (Fig. 3). The PERMANOVA, however, rejected the null hypothesis that all the observations are equal (p < 0.000999; 85% of variance explained, see the full output statistics in Supplementary Material S2), and the pairwise contrast test was significant for some species pairs, including N. mashpi sp. nov. and N. laevigata (p adjusted = 0.03), thus supporting our hypothesis that these belong to distinct, though phylogenetically close lineages (Troya et al. unpubl. data). Except for the dorsal petiolar index (DPI), which is shown in the ordination plot relatively distanced apart from all the species groupings, the remaining indices are comparatively important in explaining the morphometric differences between N. mashpi sp. nov. and the rest of related taxa. Whereas, most of the other variables are important for distinguishing between N. commutata and its congeners. In regards to N. gojira sp. nov., for which a single individual was identified, the ordination shows that it is more similar to N. laevigata than to N. marginata and N. mashpi sp. nov. The morphospecies Neoponera nr. gojira, which is supposed to be closer (or perhaps conspecific) to N. gojira sp. nov., is placed within the morphospace of N. laevigata, N. marginata, and N. mashpi sp. nov. Since the sample size of both N. gojira sp. nov. and Neoponera nr. gojira is currently insufficient for making solid conclusions about the potential place occupied by them in the morphospace of the N. laevigata group, further specimens are required to confirm current assumptions. However, as detailed in its corresponding species treatment, we do have valid reasons to treat N. gojira sp. nov. as a good species.

Worker-and queen-based key to Neoponera species-groups
The following key allows distinguishing species-groups within Neoponera, as well as a newly recognized genus lineage which is apparently sister to Neoponera (Troya et al. unpubl. data). For practical reasons only we here refer to this new genus as "N. emiliae 2", since the three species composing this clade, Neoponera concava Mackay & Mackay, N. schultzi Mackay & Mackay, and N. venusta Forel, are currently assigned to the Neoponera emiliae group sensu . In addition, we have evidence that N. bucki Borgmeier does not belong to Neoponera but it is distantly related to it, probably another new genus lineage. Nevertheless, as we are not proposing new nomenclatural acts in this regard, we have included both N. bucki and the "N. emiliae 2" group in the present key until formalizing their new status in a coming contribution.
Character states in each statement appear in relative order of importance for the identification of a given group.

Neoponera laevigata species-group brief taxonomic notes
Probably based mainly on the presence of the malar carina of Neoponera commutata Roger, Emery (1911) created the "Commutata group" and placed it within Neoponera. Later,  in a treatment of the ecological relations of ants and termites, with a focus on ponerines, created Termitopone to accommodate N. commutata, N. laevigata, and N. marginata Roger. This author based his decision on morphological and ecological observations related to specialized termite predation which is common to all three species. Wheeler's Termitopone remained valid for a relatively long time until its synonymy under Pachycondyla by Brown (1973).  included these three termitophagous species, again under Pachycondyla within the "P. laevigata species-complex", providing a new key to its species, distribution, and brief field and literature notes on their biology and ecology. Schmidt and Shattuck (2014) included these taxa in Neoponera, and according to their molecular phylogeny the N. laevigata group is sister to the N. aenescens group.  A. Troya and J. Lattke / Insect Systematics & Evolution (2022) DOI 10.1163 We found the following combination of morphological and behavioral characters useful for diagnosing this clade of above-ground termite-raiding species. In addition to the strongly ventrally bent mandibles, we consider such behavioral traits as being synapomorphic for the species here treated. The sole exception to confirmed termiteraiding is N. gojira sp. nov., for which we found a single worker and information about its natural history is unknown. Most of the following character states will not be repeated in the general synoptic description, nor in the diagnoses and descriptions of individual species, and apply to all castes except where otherwise stated.
Neoponera laevigata species-group differential diagnosis Known species in the N. laevigata group are easily recognized from other species in the genus by the following: 1) the integument of workers and queens is entirely black and exceptionally lustrous (reflecting objects very much like a mirror), bearing scant piligerous punctae, usually with whitish to pale yellowish tiny appressed pilosity, and erect golden hairs. Although the integument of the males is also black and lustrous, it shows significantly more piligerous punctae than in the other castes within this species-group. Workers and queens in some species of the N. aenescens group (for example, N. carbonaria Smith), N. crenata group (for example, N. moesta Mayr), and N. emiliae group (for example, N. magnifica Borgmeier) also show lustrous integuments but these are covered with abundant piligerous punctae, especially on the head dorsum and mesosoma, and with the exception of N. magnifica, the body surface is on average dark brownish to bluish-greenish, never jet-black as in all species of the N. laevigata group. The integument of N. magnifica is also jet-black and the gaster is lustrous, without sculpture, but this species is easily distinguished from all species in the N. laevigata group since it bears costae on the dorsum of head, pronotum and mesonotum. 2) Except for N. commutata, the distal region of the mandibles in workers and queens is strongly bent ventrad in lateral view, clearly more than in any other Neoponera species. Although the mandibles of workers and queens in N. commutata are similarly bent ventrad as in most Neoponera, the lustrous black integument of the body and their large size (maximum relative body length of workers: 19 mm, and queens: 20 mm, the largest species in the genus), are the easiest clues to separate it from all other Neoponera species. The male caste of N. commutata is the only case within the genus showing well-developed torular lobes (Fig. 14a), and maxillary palps with segments 4 th to 6th fused ( Fig. 18b) (see extended details under the comments section for this species).
Following, is a set of traits which together provide a detailed characterization of the species of the N. laevigata group.
1. Compound eye relatively large, ca. one fourth of head length; in most species flat with its external margin not surpassing lateral margin of head in frontal view. In Neoponera commutata the eye is globose and surpasses the lateral head margin. In males the eye is slightly notched anterodorsally, globose, and occupies ca. twofourths or more of head length; its external margin clearly surpasses the lateral head margin in frontal view.
2. Torular lobe subtriangular with rounded lateral margins, covering approximately 90% of acetabulum of antennal socket in frontal view. Known males of N. mashpi sp. nov. and N. marginata with strongly reduced torular lobe, while the male of N. commutata shows a small, but well-developed lobe covering approximately less than 10% of the antennal socket. 3. Frontal carina surpassing ocular mid-length in frontal view. In N. commutata this carina just reaches ocular mid-length. 4. Malar carina, absent in most species, instead a swelling is present and usually bears foveae and striae. Only N. commutata exhibits a strongly developed malar carina which reaches the anterior eye margin. Males lack this character. 5. Mandibles strongly bent ventrad in lateral view, with external margin weakly sinuous medially in full-face view. Males bear the typical Ponerini spatulate to semitriangular, small, toothless mandibles, always bearing the oval-shaped groove dorsally near the mandibular base. 6. Pronotum lacking a distinct humeral carina, though a very tenuous, blunt, almost indistinguishable border may be present in workers and queens of all species, except for N. commutata and males of all species where the humerus is completely rounded. 7. Notauli absent on mesoscutum (males only). 8. Notopropodeal suture present, forming groove, not depressed though (N. laevigata, N. mashpi sp. nov., and N. gojira sp. nov.), or slightly depressed (N. commutata and N. marginata) in dorsal view. This groove is usually cross ribbed. In males and queens the metanotum is clearly separated from the mesonotum anteriorly, and from the tergum of A1 posteriorly, by well-defined grooves. 9. Mesosternal and metasternal processes well-developed, fang-shaped, anteroposteriorly flattened. 10. Posteroventral cuticular flap at metapleural gland opening and its relative visibility in posterolateral view: strongly reduced, gland orifice entirely visible (worker of N. gojira sp. nov.); well-developed, strongly bent anterad, gland orifice barely visible (worker and queen of N. laevigata and N. mashpi sp. nov.); well-developed, bent dorsad nearly vertically straight, gland orifice almost entirely visible (worker and queen of N. commutata, N. marginata). In males of examined species this structure varies from well-developed, slightly bent dorsad, gland orifice clearly visible (N. mashpi sp. nov., N. marginata), to absent, instead forming a vertically-oriented, salient carina, gland orifice completely visible (N. commutata). 11. Petiolar node cuboid, usually subquadrate in lateral view, or slightly tapering dorsad, the latter state only seen in N. commutata. In males of all examined species the petiolar node also tapers dorsad with both anterior and posterior margins relatively straight in lateral view. 12. Prora either well-developed, with a blunt tip projecting ventrad (N. mashpi sp. nov., N. commutata, N. marginata), or poorly developed with only a small cuticular lip that is usually hardly discernible laterally (N. laevigata, N. gojira sp. nov.). 13. Cinctus moderate (most species) to absent or barely distinguishable (only in N. commutata). All examined males showed a moderate cinctus.
14. Stridulitrum present on presclerite of A4, sometimes poorly developed, faint and narrow. 15. Arolium present, small, roughly half length or less of pretarsal claw. In males the arolium is slightly longer than half-length of pretarsal claw. 16. General body appearance jet-black, surface with sparse erect golden hairs interspersed with few scattered, piligerous punctures from which golden, appressed tiny hairs emerge, most of which do not touch each other. Queens show relatively more erect pilosity than workers, particularly ventrally on head and on abdominal segments 5-7. Males depart from this in showing a body surface with less erect hairs and significantly more piligerous punctures bearing appressed pale-yellowish pubescence. 17. All species show striae on the head, mesosoma and petiolar node, that varies interspecifically in abundance, thickness and complexity. The petiolar node of N. gojira sp. nov., and that of most populations of N. mashpi sp. nov. are almost devoid of striae. Subpetiolar process covered with transverse, strongly impressed, irregular striae. The head, petiole, and subpetiolar process in males are usually significantly less sculptured than in workers and queens. 18. Worker caste either monomorphic (only N. commutata), or polymorphic (N. mashpi sp. nov., N. laevigata, N. marginata). Evidence is lacking for N. gojira sp. nov., though probably also polymorphic. 19. This is the only clade within Neoponera with all constituent species being specialized in termite-raiding. The feeding biology of N. gojira sp. nov. is unknown, but probably it raids on termites too.

Neoponera laevigata species-group description
The following describes the morphological traits shared by all examined species in each caste: worker, queen and male. Species-specific traits are shown under each species description in the "Species accounts" section. The male of N. laevigata, and the queen and male of N. gojira sp. nov. are unknown.

Worker
Head. Frontal view: Mandible triangular, longer than wide, strongly bent ventrad, dorsolateral margin weakly sinuous medially. PF: 4,4. Stipes mostly smooth with strong longitudinal internal groove. Clypeus anteromedially convex, slightly projecting over base of mandibles when closed. Posterior margin of frontoclypeal sulcus just reaching ocular mid-length (except N. commutata where the sulcus barely reaches anterior horizontal ocular length). Frontal carina surpassing ocular mid-length. Eye suboval, maximum length ca. one-fourth of head length, in workers placed anterior to cephalic mid-length while in males is slightly anterior to it (except N. commutata where the eye A. Troya and J. Lattke / Insect Systematics & Evolution (2022) DOI 10.1163 is placed at cephalic mid-length). Mesosoma. Lateral view: Dorsal margin straight. Propodeal spiracle slit-shaped. Ventral view: probasisternum shield-shaped, strongly grooved. Meso-and metasternal processes well-developed, fang-shaped, relatively flattened anteroposteriorly, with internal margins running parallel; space between apices approximately equal to-or shorter than width of each apex. Metasternal process inclined posterad 20°-30° in lateral view (except for N. commutata where the process is inclined ca. 45°). Apices of metasternal processes slightly higher than half height of metacoxal internal margin. Lateral view: Pheromone venting canal at metapleural gland opening usually well-developed with few interspecific variations. Petiole. Lateral view: Subpetiolar process covered with transverse, usually well-impressed, irregular striae. Dorsal view: anteroventral nodal carina incomplete, with feeble concavity medially. Gaster. Lateral view: Anterior margin of tergum of A3 straight. Dorsalmost limit of anterior tergal margin of A3 nearly as high as maximum height of petiolar node margin. Legs. Tibial spur formula 2(1s, 1p), 2(1s, 1p). Internal mid tibial spur halflength of external spur. Internal hind tibial spur close to one-third length of external spur. Arolium present, roughly 0.5 times length of pretarsal claw. Pretarsal claw arched, unarmed (without accessory teeth on its surface). Metabasitarsal gland present on basalanterior face. Pilosity. Body with sparse, flexuous, golden, erect and suberect hairs; appressed pubescence almost absent on entire body (only smallest workers of N. laevigata, and N. mashpi sp. nov.) to slightly more abundant (remaining workers of all species); subpetiolar process always with a combination of fine, relatively abundant whitish (all species except N. commutata) to light golden (only N. commutata) appressed and suberect hairs. Scape hairs mostly erect and suberect with sparse appressed pubescence. Posterior ventrolateral margin of hypopygium (just next to the base of stinger) bearing flexuous hairs. Sculpture. Integument polished, mostly smooth.  Basivolsella setose. Internal and external margins of cuspis nearly straight, its apex very close to posterior margin of digitus. Penial scleritess dorsomedially fused up to approximately one-half or less their length. Pilosity and sculpture. As in worker and queen except for: Appressed pubescence abundant mostly on body dorsum. Erect hairs smaller than maximum eye length. Scape hairs mostly with abundant appressed pubescence, mostly shorter than apical scape width. Meso-and metatarsi bearing abundant stiff setae interspersed with abundant suberect and appressed hairs.

Species accounts
Neoponera commutata (Roger,     Morphology mostly as in worker except for: Head. Frontal view: anterior ocellus maximum diameter subequal to that of posterior ocelli, though sometimes larger. Distance from posterior ocellar margin to posterior margin of head close to twice apical scape width. Distance between anterior and posterior ocellus approximately 1.8 times anterior ocellus maximum length. Mesosoma. Dorsal view: Mesoscutum broader than long, close to twice as long as mesoscutellum. Transscutal line well-developed. Scutoscutellar sulcus strongly impressed, cross-ribbed. Mesoscutellum subtrapezoidal, slightly flat dorsally. Anterodorsal median propodeal sulcus vestigial. Lateral view: Posterior face of propodeal declivity slightly concave. Posterolateral margin of propodeal declivity very similar as in worker though usually less salient and sometimes with crenulae. Ventral view: Mesosternal process with internal margins slightly divergent. Wings. Forewing venation: Rs-M one-third longer than 2M. Forewing evenly covered by fine setose layer, except for subbasal and basal cells which are almost completely devoid of pilosity. Hindwing venation: 1Rs absent or highly reduced. Hindwing bearing 10-14 hamuli. Petiole. Lateral view: anterior nodal margin shows a more pronounced slope than posterior margin. Color, pilosity and sculpture. Mostly as in worker (but see general synoptic description).
Male description. Measurements (n = 3 Head. Frontal view: Mandibular apex, when closed, slightly surpasses lateral margin of labrum. PF: pseudo 4, 4. Labral dorsum mostly rugose. Clypeus anteromedially straight to slightly concave. Area between posterior margin of clypeus and supraclypeal area without depression. Posterior supraclypeal area rounded, slightly protruding from cuticle, followed posteriorly by carinate swelling. Distance between internal margins of antennal sockets 1.5 times socket diameter. Ocellar area positioned slightly posterad to vertex. Distance from posterior ocellar margin to posterior margin of head equal to twice apical scape width. Distance between anterior and posterior ocelli close to one anterior ocellus maximum length. Scape cylindrical, twice as long as pedicel. Ventral view: Prementum with strong transverse carina. Lateral view: Dorsal surface of clypeus relatively straight to slightly convex, not inflated as in remaining known males of the N. laevigata group, always covering anterior clypeal margin. Eye suboval, located at cephalic mid-length, slightly notched in the dorsal third, maximum length slightly shorter than half head length. Mesosoma. Dorsal view: Mesoscutum close to twice as long as mesoscutellum, anterior triangular impression weakly impressed, lacking median longitudinal line. Parapsidal lines strongly impressed, slightly divergent. Scutoscutellar sulcus straight. Anterior subalar area approximately as broad as tegular maximum width. Metanotum surface even, slightly flat. Metapleuropropodeal groove deeply depressed. Anterodorsal median propodeal sulcus present, strongly impressed. Lateral view: Posterior face of propodeal declivity concave. Posterolateral margin of propodeal declivity with strong, sharp carina, running along the posterodorsal propodeal margin. Pheromone venting canal at metapleural gland opening absent, replaced by circular groove surrounded by strong, salient carinae. Ventral view: Probasisternal posterior projection with minute spine. Mesosternal process well-developed, fangshaped, with internal margins running parallel, relatively flattened anteroposteriorly, inclined posterad ca. 45° in lateral view, space between apices approximately equal to width of each apex. Metasternal process inclined posterad, ca. 45° in lateral view, space between apices wider than width of each apex. Wings. Forewing venation: Rs-M onethird longer than 2M. Forewing evenly covered by fine setose layer, except for subbasal and basal cells which are almost completely devoid of pilosity. Hindwing venation: as in queen. Hindwing bearing 10-14 hamuli. Petiole. Lateral view: Anterior and posterior margins of node nearly straight at base and tapering at top. Lateral projection of anteroventral nodal carina well-developed, with acute tip strongly bent dorsad. Dorsal view: Anterior margin of petiolar node slightly shorter than width of posterior margin. Lateral view: Subpetiolar process subtriangular, with rounded anterior cusp, followed by posterior, slightly concave slope (ca. 20° or less), usually with two slightly divergent longitudinal carinae bearing groove in between. Gaster. Lateral view: Meeting of anterior margin of abdominal tergum 3 with its dorsal margin rounded. Cinctus weak. Spine at posterior margin of tergum of A8 strongly acute, slightly flattened laterally, with sharp dorsal carina posteriorly. Abdominal sternum 9 longer than broad. Pygostyles close to four times longer than broad. Dorsal view: Tergum of A3 slightly shorter than A4. Ventral view: Abdominal sternum 7 showing uneven surface, with strong ditch initiating on second third, most surface bearing abundant erect and suberect hairs, ventrolaterally with long barbiculate hairs. Legs. Mesofemur roughly equally thick as metafemur, in dorsal view. Ventral surface of meso-and metafemora almost entirely flattened from base to apex, without longitudinal groove. Arolium roughly 0.7 times (two-thirds) length of pretarsal claw. Pretarsal claw with tiny, barely distinguishable median tooth. Color. Appendages, including antennae and mandibles, black to slightly yellowish black. Genitalia. Dorsal view: Cupula: posterior margin of medial invagination relatively broad. Lobular process well-developed. Gonopod dorsally elongated, with approximately subequal margins from base to apex, elongated in lateral view, with dorsal and ventral margins subparallel along its length, dorsolateral margin rounded medially, smooth surface. Gonostylus subrounded apically, dorsointernal margin mostly evenly continuous. Basivolsella externally convex. Digitus subrectangular, with outer margin nearly straight in ventral view. Penial sclerites, overall outline, subrectangular, dorsal surface mostly flat, though slightly convex posteriorly. Dorsoapical third of penial sclerite surface continuous, not modified. Sculpture. Head feebly rugulose mostly on dorsum; posterolateral pronotal margin slightly striated; axillula, metapleuron, and most of propodeum rugose; striae on petiolar node almost absent.
Natural history notes. This is a relatively common ground surface dweller, found mostly in a variety of habitats from the Amazon-Orinoco watershed, where the majority of the records are known, and have been examined in this study. Individuals of all castes have been collected throughout the year, however, research is still pending in regards to colony life cycle and production of alate forms (but see Schmidt & Overal 2009). Nothing is known in regards to its flight phenology. Several techniques have been used to collect this species including (from the most to the least effective) ground surface pitfall traps, hand collections, Malaise (mostly males), litter sifting and Winkler, and very unusually with canopy fogging. Neoponera commutata nests underground Mill 1984;Schmidt & Overal 2009; A. Troya pers. obs.) with colonies varying in population size from approximately 400-1000 individuals (Mill 1984;Schmidt & Overal 2009). Adrian Troya observed various individuals of N. commutata coming in and out from an (apparently) empty, ca. 1m diameter, dome-shaped, termite nest. The structure was attached to a live tree trunk in a lowland Amazonian forest in Ecuador, and seemed to host an ant colony. After ca. 15 minutes of observation no prey of any kind was seen in the mandibles of the workers. Although no guarding behavior was recognized at what would be their nest entrance located laterally on the dome, the hypothesis of reutilization of termite nests by N. commutata needs further confirmation. Mill (1982aMill ( , 1984, as part of his research on termites, registered the group recruitment, predatory behavior of N. commutata on several termite groups, including the genus Syntermes Holmgren, some of the species of which are considered exclusive prey of these ants, for example S. calvus Emerson, S. molestus (Burmeister), S. spinosus (Latreille) [= S. solidus Emerson] Hermann 1968?;Mill 1984), all of them broadly distributed in Amazonia, except for S. calvus which has only been found in Brazil, French Guiana, and Guyana (Constantino 2020). Most of this data was recorded in a few sites, such as Kartabo and Kaieteur in Guyana, and Vista Alegre, state of Amazonas and Ilha de Maracá state of Roraima, in Brazil. John Lattke observed nocturnal hunting of Syntermes in forests of Cerro Los Pijiguaos, southern Venezuela. A loose column of 6-8 workers making their way through the litter was seen, and a little over an hour later, presumably the same column was seen returning, each worker carrying 2-3 termites in their mandibles. Based on this pattern of specific predatory behavior  proposed that the distribution of N. commutata should follow that of their prey. Thus far, this hypothesis has not been challenged, nor tested. N. commutata is an efficient predator, according to Mill (1984) estimations, only three colonies can consume as much as three times the worker and soldier populations of the above-mentioned termite species per hectare, annually. Mill (1982b) also informed on the nest migration strategy of N. commutata in an Amazonian rain forest.
The venom of N. commutata contains antimicrobial, insecticidal and haemolytic peptides, according to Aili et al. (2016). These proteins are known as ponericins and it is assumed these help prevent diseases originated from the prey organisms brought by the ants inside their colonies (Orivel et al. 2001). Aili et al. (2016) also found in the venom of N. commutata the highest number of proteins as compared to the venoms of the ants Ectatomma brunneum Smith, Myrmecia gulosa (Fabricius), Neoponera apicalis, and Odontomachus hastatus (Fabricius). Schmidt and Overal (2009) found that raiders and nest defenders are equipped with more amounts of venom and that also this is more lethal than in other worker nest mates who do not get involved in foraging activities and colony defense.
Comments. Current morphological evidence allows an unequivocal evolutionary relatedness of N. commutata to the remaining species in the N. laevigata group. However, despite showing some apomorphies with other members in this clade, for example, the mandibles strongly bent ventrad in workers and queens, mesoscutum lacking notauli in males (Fig. 17b), specialized termite-feeding, this lineage is an outlier from a morphological perspective. Traits like the subtriangular antennal scapes (transverse plane), the occipital and malar carinae in workers and queens, the presence of well-developed torular lobes in males, all of which are absent in the remaining species within the clade, provide signs of an evident separation of this lineage with respect to its more derived clade partners which thus far appear to form a closely related group (Troya et al. unpubl. data). This is also clearly depicted by the multivariate analysis in Fig. 3. In addition, we consider a striking feature in this species the (secondary?) fusion of the male maxillary palpomeres 4-6 ( Fig. 18b), which is evidenced by the absence of a complete joint between the 4th and 5th, and the 5th and 6th segments. Thus far, this is the only known case in Neoponera, and to our knowledge unknown in ponerine males. Why and when this loss of segmentation occurred during the course of evolution in this caste, remain open questions.
Distribution notes. Based on our records and the literature, the populations of N. commutata are exclusively South American (Fig. 29b). The elevational range of the species extends from nearly the sea level up to approximately 1300 m. The northern geographic range reaches the Serranía de la Macarena, a Colombian mountainous region lying just at the transition zone between the Amazonian lowlands and the premountainous Andean forests, in the center of that country. The southernmost distribution of N. commutata appear to be marked by the Humid Chaco and the Atlantic Forest in Paraguay, whereas its eastern limit is the state of Paraíba in northeast Brazil, where we examined a single worker from a heavily anthropized zone with remains of stepic savannas, some of which are part of the threatened Caatinga biome. In northwestern South America the Andean Cordilleras may act as the ultimate barrier for N. commutata entering Mesoamerica. Except for an isolated record from the Department of Bolívar in northern Colombia (mined from Antmaps.org, Janicki et al. 2016), no other sightings or collections have been reported from the northern region of that country, nor in Atlantic or Pacific regions in Central America. The Costa Rican record in San José, reported by , is a possible interception of South American origin, as no individuals have been found so far after years of sampling in the region (J. Longino, pers. comm.). Etymology. The specific name honors the French metal band Gojira, in recognition of their altruistic activism for the conservation of nature, as well as for supporting the rights of ancestral indigenous peoples living across Amazonia. Through their lyrics the band promotes an increasing awareness of our paramount, nonetheless usually neglected, biological diversity. The name is a noun in apposition, thus invariable.
Worker diagnosis. Antennal scape surpassing posterior margin of head by ca. one apical scape width; head in frontal view trapezoid, slightly wider posteriorly than anteriorly (Fig. 19d); dorsal masticatory margin of mandible uneven, with shallow, irregular rim (Fig. 19b); masticatory margin of mandible with three comparatively large blunt teeth alternating with blunt denticles (Fig. 19b); posteroventral cuticular flap at metapleural gland opening reduced to narrow carina, orifice of gland completely visible in posterolateral view (Fig. 2d); petiolar node laterally lacking striae, though very tenuous lines are present near its ventral margin (Fig. 19e); prora tiny, barely discernible laterally (Fig. 19e) Head. Frontal view: trapezoidal, with posterior margin wider than anterior margin. Masticatory margin of mandible with four large, blunt teeth interspersed with four denticles, basal margin edentate. Prementum with clear transverse dome. Labral dorsum mostly smooth. Posterolateral margin of torular lobe rounded. Malar carina absent, though swelling is present. Eye convex, relatively flat, not surpassing lateral margin of head, placed anterior to cephalic mid-length. Posterior margin of head concave. Occipital carina absent. Lateral and/or frontal view: Scape when pulled back, surpasses posterior margin of head by close to one apical scape width. Mesosoma. Lateral view: Pronotum with weak, blunt, barely distinguishable humeral carina. Metanotal groove present, though not depressed, usually clearly separating mesonotum and propodeum. Anapleural sulcus present, completely merged with surrounding striae. Posterior face of propodeal declivity flat. Posterolateral margin of propodeal declivity with feeble, blunt carina, almost devoid of crenulae. Pheromone venting canal at metapleural gland opening well-developed, smooth. Propodeal lateral margins subparallel in posterodorsal view. Ventrolateral propodeal declivity with deep transverse groove just dorsad to metapleural gland opening. Ventral view: Probasisternal posterior projection acute. Space between apices of mesosternal process wider than width of each apex. Petiole. Lateral view: Anterior and posterior margins of node nearly straight and semi-parallel, posterior margin slightly convex; posterodorsal margin slightly higher than anterodorsal margin; mid-dorsal margin mostly flat. Lateral projection of anteroventral nodal carina well-developed, with slightly blunt tip moderately bent dorsad.
Dorsal view: Anterior margin of petiolar node roughly as broad as width of posterior margin. Anteroventral nodal carina incomplete, feebly concave medially. Lateral view: Subpetiolar process subtriangular, with acute anterior cusp, keel-shaped, followed by posterior relatively straight slope (ca. 30°). Gaster. Lateral view: Prora present, though poorly developed, similar to a cuticular lip, usually hardly discernible. Meeting of anterior margin of abdominal tergum 3 with its dorsal margin relatively angled. Cinctus moderate. Dorsal view: Tergum of A3 slightly shorter than A4. Posterior dorsum of epipygium smooth, feeble striations and punctae present but hardly discernible and requires rotation of specimen to get better angle of observation. Legs. Mesofemur thickened medially, thicker than metafemur, in dorsal or ventral view. Ventral surface of meso-and metafemora flattened roughly on distal two-thirds, bearing longitudinal shallow groove. Color. Appendages including antennae and mandibles black to ferruginous black, mandibles feebly lighter than remaining integument of body. Pilosity. Erect hairs on body dorsum about as long as maximum eye length. Posterior dorsum of epipygium surrounded by ca. 20, long flexuous hairs. Sculpture. Striae on head present, only dorsally, covering most of surface except posterior third, anepisternum, katepisternum, metapleuron and lateral side of propodeum bearing strong striae; striae on petiolar node almost absent.
Queen and male. Unknown. Natural history notes. Unknown. The holotype was collected in January 2016 in a project aiming to analyze anthropogenic disturbance in the Brazilian Cerrado. The ecosystem where this species was found is composed of tropical savannas, deciduous and semi-deciduous forests, and highly disturbed areas used mostly for agriculture (IBGE 2004, Soterroni et al. 2019García et al. 2021) Comments. Neoponera. gojira sp. nov. is currently known from a single specimen, nonetheless, various morphological features clearly separate it from its potential closest species: N. laevigata and N. mashpi sp. nov. The following characters are useful for separating the workers: the dorsal masticatory mandibular region shows a depressed rim (Fig. 19b) which is absent in N. laevigata and N. mashpi sp. nov., this feature is not to be confused with the strongly excavated mandibular groove of N. marginata, which runs uninterrupted posterad along the entire mandibular base (Fig. 12a); posterior margin of head in full-face view clearly concave, with relatively salient posterolateral corners (Fig. 19d), while in N. laevigata this margin is straight or feebly concave (Figs 20c, 21d), and straight or slightly convex in N. mashpi sp. nov. (Fig. 25c); flap at metapleural gland opening strongly reduced to a narrow, but sharply delineated carina, in posterolateral view, while in both N. laevigata and N. mashpi sp. nov. this flap is welldeveloped and runs uninterrupted along the ventrolateral margin of the metapleuron until reaching its anterior lobe (Fig. 2d); clypeus and internal side of frontal lobes heavily sculpted with foveae and striae (Fig. 12b), in N. laevigata and N. mashpi sp. nov., on the other hand, the clypeus is feebly striated, and the frontal lobes almost always lack sculpture, except for two shallow foveae on the internal side. Heavily impressed clypeal sculpture is also typical in N. marginata, and to a lesser degree in N. commutata. However, impressed sculpture (foveae and striae) on the frontal lobe area has only been seen in N. gojira sp. nov. Compared to the workers of similarly sized species, e.g., N. laevigata (WL= 2.45-3.57 mm; TLa= 7.88-10.03 mm) and N. marginata (WL= 2.75-3.59 mm; TLa= 7.41-10 mm), the holotype of N. gojira sp. nov. is thus far the second largest worker in the N. laevigata group (WL = 3.72 mm; TLa = 10.26 mm), only oversized by workers of N. commutata. It is assumed that the still unknown conspecifics (but see below) may have a similar size showing the characteristic continuous polymorphic size variation that is present in its closest lineages.
Geographic range. Eastern Brazil. Phenotypic variant of N. gojira sp. nov.
In the northeastern Brazilian states of Piauí and Bahia we found ten workers morphologically similar to N. gojira sp. nov. and N. laevigata. This variant shares morphospace with these two species, but also with N. marginata (Fig.. 3). Until additional material is discovered we leave this morphotype as Neoponera nr. gojira which shows the following diagnostic traits: dorsal margin of mandibles, near the masticatory region, with shallow fossulae; antennal scape surpassing posterior margin of head by approximately one apical scape width (Fig. 13b); flap at metapleural gland orifice present, though not strongly curved anterad, so that the orifice is easily visible in posterolateral view (Fig.  2b); petiolar node mostly smooth, though with feebly impressed striae laterally (Fig.  22b); prora tiny (Figs 13d,19e).
Neoponera nr. gojira differs from N. gojira sp. nov. in the following: relatively smaller (WL = 3.24-3.32 mm; TLa = 8.78-8.93 mm); four to five teeth on mid masticatory margin of mandibles clearly smaller and more acute (N. gojira sp. nov. bears three relatively large, blunt teeth, Fig. 19b); posterior margin of head less concave than in Figure 19d). Neoponera nr. gojira differs from N. laevigata in the following: dorsal mandibular surface, near masticatory margin, bearing fused, shallow fossulae (we saw these fossulae in some workers of N. laevigata, though clearly less impressed); petiolar node with tenuously impressed striae (node strongly striate laterally in N. laevigata, especially specimens from Amazonian populations); flap at metapleural gland orifice well-developed but not strongly curved anterad (this flap is also well-developed in N. laevigata but is strongly curved anterad, partially hiding the gland orifice, Fig. 2a   Worker and queen diagnosis. Antennal scape, when pulled back, surpasses posterior head margin by about one apical scape width (Fig. 13b); longitudinal striae present around internal margin of eye and between frontal carinae, diverging posterad and usually vanishing at vertex, but reaching lateral head margin just posterior to eyes (Figs 13b, 21d); dorsum of head in smaller workers significantly less sculpted than larger workers to almost devoid of striae; lateral face of petiolar node always with oblique, usually strongly impressed striae (Fig. 20d); subpetiolar process, in lateral view, subtriangular, with acute anterior cusp, usually followed by relatively flat surface posterad (Figs 13d, 20d); prora absent or very small, lip-shaped (Fig. 13d).  .
Head. Frontal view: subrectangular. Masticatory margin of mandible with four to five teeth interspersed with three to six denticles, basal margin edentate. Prementum with clear transverse dome. Labral dorsum mostly smooth, posterolateral margin rounded. Malar carina absent, though swelling present. Eye convex, relatively flat, not surpassing lateral margin of head, placed anterior to cephalic mid-length. Posterior margin of head straight to slightly concave. Occipital carina absent. Scape when pulled back, surpasses posterior margin of head by close to one apical scape width. Mesosoma. Lateral view: Pronotum with weak, blunt, barely distinguishable humeral carina. Metanotal groove present, though not depressed, usually clearly separating mesonotum and propodeum. Anapleural sulcus present, though usually merged with surrounding striae. Posterior face of propodeal declivity mostly flat. Posterolateral margin of propodeal declivity with feeble, blunt carina, sometimes with crenulae. Pheromone venting canal at metapleural gland opening well-developed, smooth. Propodeal lateral margins subparallel in posterodorsal view. Ventrolateral propodeal declivity with deep transverse groove just dorsad to metapleural gland opening, in posterolateral view. Ventral view: Probasisternal posterior projection acute. Space between mesosternal process lobes approximately equal to width of each. Metasternal process inclined posterad 20°-30° in lateral view. Petiole. Lateral view: anterior and posterior margins of node usually nearly straight and semi-parallel, sometimes posterior margin slightly convex; posterodorsal margin slightly higher than anterodorsal margin; mid-dorsal margin mostly flat. Lateral projection of anteroventral nodal carina well-developed, with acute or slightly blunt tip moderately bent dorsad. Dorsal view: Anterior margin of petiolar node slightly shorter than width of posterior margin. Anteroventral nodal carina incomplete, with feeble concavity medially. Lateral view: Subpetiolar process subtriangular, usually with keel-shaped, acute anterior cusp, followed by posterior straight slope (ca. 20° or less). Gaster. Lateral view: Prora present, though poorly developed, similar to a cuticular lip, usually hardly discernible. Meeting of anterior margin of abdominal tergum 3 with its dorsal margin relatively angled. Cinctus moderate. Dorsal view: Tergum of A3 slightly shorter than A4. Posterior view: Posterior dorsum of epipygium smooth, though feeble striations and punctae may be present. Legs. Mesofemur thickened medially, thicker than metafemur, in dorsal or ventral view. Ventral surface of meso-and metafemora flattened roughly on distad two thirds, usually bearing longitudinal shallow groove. Color. Appendages including antennae and mandibles brown to ferruginous brown, occasionally legs yellowish and mandibles lighter than remaining integument of body. Pilosity. Most erect hairs on body dorsum approximately as long as maximum eye length. Scape hairs about as long as apical scape width. Posterior dorsum of epipygium surrounded by ca. 20, long flexuous hairs. Sculpture. Striae on head present, only dorsally, covering most of surface except posterior third, anepisternum, usually part of katepisternum, metapleuron, lateral and part of dorsal side of propodeum with strong striae, on petiolar node always present, oblique and strongly impressed.
Morphology as in worker except for: Head. Frontal view: Anterior ocellus maximum diameter subequal to that of posterior ocelli. Distance from posterior ocellar margin to posterior margin of head close to twice apical scape width. Distance between anterior and posterior ocelli approximately 1.5 times anterior ocellus diameter. Posterior margin of head straight to slightly concave. Mesosoma. Dorsal view: Mesoscutum about as long as broad, close to 2.5 times as long as mesoscutellum. Transscutal line feebly developed. Scutoscutellar sulcus shallowly impressed, formed by scrobiculate sculpture. Mesoscutellum subquadrate, slightly flat dorsally. Posterior face of propodeal declivity flat. Posterolateral margin of propodeal declivity slightly rounded to barely carinate, not forming crenulae. Anterodorsal median propodeal sulcus present, weakly impressed. Probasisternal posterior projection with spine. Wings. Forewing venation: Rs-M approximately one half larger than 2M. Forewing evenly covered by fine setose layer. Hindwing venation: 1Rs highly reduced. Hindwing bearing 10-13 hamuli. Petiole. Lateral view: anterior and posterior margins of node nearly straight at base and slightly tapering dorsad, with posterior margin slightly convex and with more pronounced slope than anterior margin which is almost perpendicular to horizontal body axis. Gaster. Prora absent. Color. Appendages, including antennae and mandibles, dark brown to black. Pilosity. Meso-and metatarsi bearing abundant spines and stiff setae interspersed with abundant suberect hairs. Sculpture. Striae on head present only dorsally, covering most of surface except posterior third; strong striae on posterolateral margin of pronotum, usually anepisternum and part of katepisternum, axillula, metapectus, and propodeum laterally; petiolar node always with oblique, strong striae laterally.
Male description. Not available, but see , and . These authors possibly revised male material of both N. laevigata and N. mashpi sp. nov. Since these two species are probably closely related, the male of N. laevigata might likely be very similar to that of N. mashpi sp. nov. If the male is finally found in the future we strongly recommend extracting and illustrating its genitalia, in particular the penisvalva which shows interesting differing features among the species here examined (see Fig. 28).
Natural history notes. Since N. laevigata is probably closely related to N. mashpi sp. nov. we consider the possibility that it could display a similar behavior, such as a diet based on termites. Borgmeier (1959) cites the observation "formando fila" (making a row) from P. Telles who collected specimens in Linhares, Espírito Santo, southeastern Brazil. Schmid-Hempel (1998) noted that the phorid fly Ecitomyia juxtaposita Borgmeier is associated with colonies of this species in Brazil.  collected winged gynes in March and October, also in Brazil. These are the only published sources which could potentially be associated with N. laevigata. However, since we could not find the vouchers of these studies no confirmation is currently possible. In regards to Baroni Urbani (1993) who includes this species in an overall analysis of the evolution of recruitment behavior in ants, it is uncertain whether this is in fact N. laevigata as no details about the voucher material is mentioned.
Comments. Workers of this species, which are much more commonly encountered in the field than other castes, are very similar to those of N. mashpi sp. nov. Even for a trained eye it is sometimes uneasy to differentiate both forms. However, upon recognizing the specific body features that vary between these two taxa, the task becomes easier. The queens of both species, however, are not so similar and proper examination allows separation without much trouble. We were fortunate in finding a nest series with close to 300 individuals from Pará, Monte Dourado (MPEG) which allowed us to associate workers with queens, but also to examine size polymorphism among workers, which is less variable than in its closest species N. mashpi sp. nov. (Fig. 3). For instance, the difference in size observed mainly in the variables WL and HL is less extended in the morphospace than in N. mashpi sp. nov.
The morphological characters allowing a differentiation of the known castes of N. mashpi sp. nov. and N. gojira sp. nov. which, based on their examined morphology, we assume are the most closely related species to N. laevigata, are detailed in the 'comments' section of those species. Wheeler (1922) in his work of "Ants of Trinidad", upon studying a specimen collected by P. Whelpley at Caparo, Trinidad and Tobago, mentions for the first time the lateral petiolar node striations and decides to erect Neoponera [= Euponera (Mesoponera)] laevigata var. whelpleyi. Later, in 1936, this author synonymizes it under Termitopone laevigata. We examined the image of Wheeler's (1922) syntype (MCZENT20454) and confirm this is N. laevigata since it matches the diagnostic traits specified in this study, whose populations are mostly distributed in Amazonian habitats. Together, with the record of N. marginata (voucher at American Museum of Natural History) by , these are the only records from the N. laevigata group for a Caribbean island. Although this specimen was collected more than a century ago, the forests of Trinidad and Tobago are relatively well-preserved (Baksh-Comeau et al. 2016) making it likely to find populations of this species in the present. Surprisingly, we could not find a single N. laevigata specimen from Venezuela, yet it is very likely that this species is also present in a number of habitats and regions in that country. AntWeb record CASENT0249152 identified as N. laevigata from the Venezuelan state of Bolívar, represents in fact N. mashpi sp. nov.
Distribution notes. This species is distributed only in South America, with most of the records coming from the Amazonian biome. The remaining scant records belong to the Caribbean forests of Trinidad, the tropical Andean forests of the Cordilleras Central and Occidental of Colombia, and the Pacific forests of the Chocó-Darien, also in Colombia. Although no records of this species were found for any bioregion in Bolivia, Ecuador and Peru, its populations are probably present there. For example, a N. laevigata record in the extremely isolated western Amazonian site of Estirão do Equador in Brazil, lies just at the political border with the northern Peruvian Amazon. Increasing sampling in these regions will likely unveil records for this species.
Morphology as in worker except for: Head. Frontal view: Head usually trapezoidal. Anterior ocellus maximum length subequal to that of posterior ocelli. Distance from posterior ocellar margin to posterior margin of head close to 2.5 times apical scape width. Distance between anterior and posterior ocelli close to 2.5 times anterior ocellus maximum length. Mesosoma. Dorsal view: Mesoscutum about as long as broad, 2.5 times as long as mesoscutellum. Transscutal line well-developed. Scutoscutellar sulcus shallowly impressed, formed by scrobiculate sculpture. Mesoscutellum subtrapezoid, flat dorsally. Posterior face of propodeal declivity flat. Posterolateral margin of propodeal declivity slightly round to barely carinate, not forming crenulae. Anterodorsal median propodeal sulcus present, weakly impressed. Wings. Forewing venation: Rs-M approximately one fifth longer than 2M. Forewing evenly covered by fine setose layer. Hindwing venation: 1Rs highly reduced. Hindwing bearing 10-14 hamuli. Petiole. Lateral view: anterior and posterior margins of node nearly straight at base and slightly tapering at top. Gaster. Dorsoposterior view: Posterior dorsum of epipygium with feeble striations and punctae. Color. Appendages, including antennae and mandibles, mostly black. Pilosity and sculpture. Mostly as in worker (but see general synoptic description).
Head. Frontal view: Mandibular apex, when closed, barely touches lateral margin of labrum. PF: 6,4. Clypeus anteromedially straight to slightly concave, dorsally inflated, with somewhat globular-shaped dome, always covering anterior clypeal margin. Area between posterior margin of clypeus and supraclypeal area with evident depression. Posterior supraclypeal area acute. Supraclypeal area slightly protruding from cuticle, followed posteriorly by tenuously impressed longitudinal carina. Distance between internal margins of antennal sockets approximately one socket diameter. Torular lobe strongly reduced. Eye suboval, slightly notched medially, maximum diameter ca. onethird of head length, placed slightly anterior to cephalic mid-length. Ocellar area positioned at vertex. Distance from posterior ocellar margin to posterior margin of head close to two-thirds apical scape width. Distance between anterior and posterior ocelli approximately equal to 1.5 times anterior ocellus maximum length. Scape trapezoidal, twice as long as pedicel. Mesosoma. As in queen with the following differences: Dorsal view: Mesoscutum close to 2.5 times as long as mesoscutellum, anterior triangular impression vestigial, with faint median longitudinal line. Parapsidal lines weakly impressed, slightly divergent. Scutoscutellar sulcus slightly arched posterad. Dorsolateral view: Anterior subalar area shorter than tegular maximum width. Metanotum surface even, convex. Pheromone venting canal at metapleural gland opening poorly developed, striate. Dorsoposterior view: Metapleuropropodeal groove relatively shallow. Posterolateral margin of propodeal declivity slightly rounded, rugose midventrally. Ventral view: Probasisternal posterior projection round. Mesosternal process vestigial or absent. Wings. As in queen, except for Rs-M approximately as long as 2M. Petiole. Lateral view: anterior and posterior margins of node nearly straight at base and slightly tapering at top. Lateral projection of anteroventral nodal carina well-developed, with acute or slightly blunt tip moderately bent dorsad. Dorsal view: Anterior margin of petiolar node approximately four-fifths (0.8 times) shorter than width of posterior margin. Lateral view: Subpetiolar process subtriangular, with rounded cusp at middle of process, mostly smooth, though few striae may be present. Gaster. Morphology mostly as in worker and queen with the following differences: Lateral view: Abdominal sternum 7 showing even surface with no sulci, anterior half glabrous, posterior half with semicircular patch bearing abundant, appressed, stout hairs. Spine at posterior margin abdominal tergum 8 blunt, slightly flattened dorsoventrally. Pygostyles close to three times longer than broad. Ventral view: Abdominal sternum 9 roughly as long as broad. Color. Appendages, including antennae and mandibles, dark brown to black with slightly light brown tarsi. Genitalia. Ventral view: Cupula: posterior margin of medial invagination narrow. Lobular process vestigial. Basivolsella externally nearly straight. Lateral view: Gonopod subtriangular posterior half strongly directed ventrad, dorsolateral margin relatively straight medially, with medial, oblique, weak incision. Dorsal view: Gonopod subtriangular (subrectangular in lateral view). Gonostylus subacute apically, dorsointernal margin with rounded, tooth-shaped, swelling. Digitus spatulate, with outer margin round in ventral view. Penial sclerites overall outline axeshaped, dorsal surface convex anteriorly, with an abrupt slope posteriorly. Dorsoapical third of penial sclerite surface with strong flap. Sculpture. Striae absent on head; axillula, metapleuron and lateral propodeal surface rugose; posterior and dorsopropodeal surface strongly striated; striae on petiolar node usually feeble and only posterolaterally.
Natural history notes. This is a relatively common, surface ground forager, termite feeding species Leal & Oliveira 1995;. The workers are usually collected through pitfall traps or by hand. Alate females and queens are much less commonly captured, unless searching for underground colonies. Males are quite rare, though they might be present in bulk Malaise samples. Due to their similarity to males of N. mashpi sp. nov., and likely also to males of N. laevigata (which were not examined here), they can be misidentified as such. In warm morning hours the workers can be quite active around the nest entrance on the ground surface: while taking photos of them at a tourist trail, in Reserva Guartelá, southern Brazil, Adrian Troya observed ca. 10-20 workers removing soil particles from underground and piling them some 30 cm near the entrance. None of the ants showed aggressiveness when clearing part of the surrounding grass in order to get a clearer view, nor even when collecting some specimens.
Despite being comparatively easy to identify, and commonly collected in the field, most of what is known about the behavioral biology of N. marginata is basically summarized in three studies. In the first, Leal and Oliveira (1995) provided a comprehensive account on the group-raiding and migratory habits of this species in a semi-deciduous region of the state of São Paulo in southeastern Brazil. The authors recorded more than 200 raids, all of them directed against the termite species Neocapritermes opacus (Hagen). Within the N. laevigata group this "extreme diet specialization" has been documented only in this species, so far (but see Krishna & Araujo 1968;Mill 1982b). Leal and Oliveira (1995) also concluded that N. marginata shows a rudimentary form of the "army ant syndrome", with colonies relocating their nests less frequently than other group-raiding taxa, as for example, some Dorylinae. On the other hand, in regards to the communication system,  found that the pygidial gland of N. marginata produces Citronelal, the main monoterpenoid responsible for triggering trail-recruitment in ants. The third main contribution came from Acosta-Avalos et al. (1999) who found magnetic iron oxides in the head and gaster of N. marginata (but see also Wajnberg et al. 2017). Based on this result, they proposed that colonies of this species use the geomagnetic field during migrations, i.e., nest relocations. Two years later, Acosta-Avalos et al. (2001) linked that hypothesis to their observations of N. marginata colonies displaying a preferred directional migration which is deviated ca. 12° from the magnetic North-South axis.
Comments. Workers and queens of N. marginata are morphologically similar only to those of N. gojira sp. nov., N. mashpi sp. nov., and N. laevigata. These four lineages share traits like, for example, the relatively flattened eyes placed anterior to cephalic mid-length (Figs 20c,21d,22d,23f,25c,26d); absence of malar carina; pronotal humerus feebly angled (not completely rounded as in N. commutata); a cuboid petiolar node, in lateral view; polymorphic workers, and presumably so in N. gojira sp. nov. for whom a single individual is known. However, N. marginata workers and queens are unique among these other lineages in showing a strongly carved, longitudinal groove on the inner side of the mandibular surface (absent in N. mashpi sp. nov. and N. laevigata, while N. gojira sp. nov. shows a shallow rim). Another useful feature, which helps identify queens of N. marginata, is its trapezoidal head which is broader posteriorly than anteriorly, in frontal view, though this character is more evident in the queens. The head in queens of N. mashpi sp. nov. and N. laevigata is subquadrate. In regards to known males, those of N. marginata could only be confused with those of N. mashpi sp. nov. The easiest way to distinguish both forms is using the following combination of traits: a feebly impressed, longitudinal line present medially on the mesoscutal dorsum (Fig. 15a), while in N. mashpi sp. nov. such line is absent; scape trapezoidal ( Fig. 24e), whereas in N. mashpi sp. nov. it is cylindrical; pretarsal claws bearing subapical teeth, while those of N. mashpi sp. nov. lack teeth; blunt tipped spine on posterior margin of tergum of A8, while in N. mashpi sp. nov. the spine tip is acute. The male caste of N. commutata is quite dissimilar to that of N. marginata and N. mashpi sp. nov. Some unique features of the males in the N. laevigata group were already discussed under the treatment for that species, however, the user may also find the following useful: a roughly round head in frontal view (subquadrate in remaining species); dorsoposterior propodeal margin strongly carinate (carina lacking in remaining species); forewing with glabrous subbasal and basal cells (setose in remaining species).
Distribution notes. Our records confirm an exclusively South American distribution for N. marginata, with an elevational range varying between 10 m to around 1700 m. Most of these records lie in the southern limit of its range represented mainly by three biomes: the southern Humid Chaco in Paraguay; the Atlantic Forest in Argentina (Misiones), northern Paraguay, and Brazil (Paraná and São Paulo); and the Cerrado in Brazil (Goias, Mato Grosso do Sul, São Paulo, Minas Gerais). Although highly anthropized (Ribeiro et al. 2009;Da Ponte et al. 2017) the remaining vegetation cover in these regions is mostly represented by ombrophyllous dense and open broadleaf forests, and tropical and subtropical grasslands, savannas, and shrublands (Sena-Souza et al. 2020). We found scant records of N. marginata in Amazonian regions, namely in northwestern Bolivia, western Brazil (Rondônia), and eastern Ecuador. The latter is the northwestern limit range for this species in the continent. However, as there are no large geographic barriers, e.g., mountain ranges, separating Amazonian regions between northern Ecuador and southern Colombia, we might expect N. marginata populations to be present at least in Colombian southern lowland rain forests.  list a record from the northeastern side of the continent, at Arima Reserve (voucher at the American Museum of Natural History), from the island of Trinidad, Trinidad and Tobago. The record of  from Churubamba (not 'Charubamba'), a Bolivian region in the Department of La Paz, is possibly wrong since its elevation surpasses 3400 m. This record is probably related to that from the same author (1925) from Ixiamas at 260 m (voucher at Natural History Museum Sweden), located in the same Bolivian Department.
As in N. mashpi sp. nov. and N. laevigata, N. marginata workers show gradual size polymorphism, though less marked than in the former two. Across its distribution range we did not notice major morphological variations in size, nor in coloration departing from the average ground plan of this species, as diagnosed here. Specimens from Bolivian Amazonian habitats were virtually identical to those examined from wet broadleaf forests in the Atlantic, and from the semi-dry grasslands and savannas in Paraná, southeast Brazil. A. Troya and J. Lattke / Insect Systematics & Evolution (2022) DOI 10.1163 completely lacking striae (Figs 25d, 26b), this is mostly seen in South American and Pacific Central American populations, while in Atlantic Central American populations the node bears slight striae laterally; subpetiolar process subtriangular with anterior acute cusp, in lateral view (Fig. 25d) Propodeal lateral margins subparallel in posterodorsal view. Ventrolateral propodeal declivity with deep transverse groove just dorsad to metapleural gland opening. Probasisternal posterior projection acute. Metasternal process inclined posterad, 20°-30° in lateral view. Petiole. Lateral view: anterior and posterior margins of node usually nearly straight and semiparallel, sometimes posterior margin slightly convex; posterodorsal margin slightly higher than anterodorsal margin; mid-dorsal margin mostly flat. Lateral projection of anteroventral nodal carina well-developed, with acute or slightly blunt tip usually projecting anterad. Dorsal view: Anterior margin of petiolar node slightly shorter than width of posterior margin. Anteroventral nodal carina complete. Lateral view: Subpetiolar process subtriangular, with acute anterior cusp, sometimes keel-shaped, followed by posterior straight slope (ca. 30°). Gaster. Lateral view: Prora well-developed, with blunt tip projecting ventrad. Meeting of anterior margin of A3 tergum with its dorsal margin relatively angled. Cinctus moderate. Dorsal view: Tergum of A3 slightly shorter than A4. Posterior dorsum of epipygium smooth, though feeble striations and punctae may be present. Legs. Mesofemur thickened medially, thicker than metafemur, in dorsal or ventral view. Ventral surface of meso-and metafemora flattened roughly on distal two-thirds, usually bearing longitudinal shallow groove. Color. Appendages including antennae and mandibles brown to ferruginous brown, occasionally legs yellowish and mandibles lighter than remaining integument of body.
Pilosity. Most erect hairs approximately same length as eye maximum length. Scape hairs about as long as apical scape width. Posterior dorsum of epipygium surrounded by ca. 20, long flexuous hairs. Meso-and metatarsi bearing abundant spines and stiff setae interspersed with abundant suberect hairs. Sculpture. Striae on head generally absent, though when present, very feeble and on dorsum only, laterally on clypeus, around frontal carinae, and malar area; anepisternum, metapleuron and lateral side of propodeum with strong striae; petiolar node usually devoid of striae, though when present feebly impressed on lateral face (but see diagnosis and comments Morphology as in worker except for: Head. Frontal view: Anterior ocellus maximum diameter subequal to that of posterior ocelli. Distance from posterior ocellar margin to posterior margin of head close to twice apical scape width. Distance between anterior and posterior ocelli close to 2.5 times anterior ocellus diameter. Mesosoma. Dorsal view: Mesoscutum about as long as broad, close to 2.5 times as long as mesoscutellum. Transscutal line well-developed. Scutoscutellar sulcus shallowly impressed, formed by scrobiculate sculpture. Mesoscutellum subquadrate, flat dorsally. Posterior face of propodeal declivity flat. Posterolateral margin of propodeal declivity slightly rounded to barely carinate, not forming crenulae. Anterodorsal median propodeal sulcus present, weakly impressed. Ventral view: Mesosternal process with internal margins slightly divergent. Wings. Unknown. Petiole. Lateral view: anterior and posterior margins of node nearly straight at base and slightly tapering dorsad. Lateral projection of anteroventral nodal carina slightly acute and moderately bent dorsad. Color. Appendages, including antennae and mandibles, dark brown to black. Sculpture. Striae on head present only dorsally, covering most of surface except posterior third; strong striae on posterolateral margin of pronotum, usually anepisternum and part of katepisternum, axillula, metapectus, and laterally on propodeum; petiolar node lacking striae, (see also general synoptic description).
Male diagnosis. Antennal scape cylindrical (Fig. 27d); mesoscutum lacking median longitudinal line (Fig. 15b); mesosternal process absent; forewing vein Rs-M feebly longer than 2M (Fig. 27f ); hindwing with vein 1Rs incomplete or poorly developed (Fig. 27f )  supraclypeal area acute. Supraclypeal area slightly protruding from cuticule, followed posteriorly by tenuously impressed longitudinal carina. Distance between internal margins of antennal sockets less than one socket diameter. Torular lobe strongly reduced. Eye suboval, slightly notched medially, maximum diameter slightly less than half head length, placed slightly anterior to cephalic mid-length. Ocellar area positioned at vertex. Distance from posterior ocellar margin to posterior margin of head close to twice apical scape width. Distance between anterior and posterior ocelli varying between 1-1.5 times anterior ocellus maximum length. Scape cylindrical, one third longer than pedicel. Mesosoma. Dorsal view: Mesoscutum close to 3 times as long as mesoscutellum, anterior triangular impression weakly impressed, median longitudinal line absent. Parapsidal lines weakly impressed, slightly divergent. Scutoscutellar sulcus straight. Anterior subalar area shorter than tegular maximum width. Metanotum surface even, slightly flat. Metapleuropropodeal groove relatively shallow. Posterior face of propodeal declivity mostly flat. Anterodorsal median propodeal sulcus present, weakly impressed. Lateral view: Posterolateral margin of propodeal declivity slightly rounded to barely carinate, not forming crenulae. Pheromone venting canal at metapleural gland opening poorly developed, sightly striated. Ventral view: Probasisternal posterior projection acute. Mesosternal process vestigial. Metasternal process as in worker and queen. Wings. Forewing venation: Rs-M slightly longer than 2M. Forewing evenly covered by fine setose layer. Hindwing venation: 1Rs present, very short though. Hindwing bearing 8-10 hamuli. Petiole. Lateral view: anterior and posterior margins of node nearly straight at base and slightly tapering dorsad; dorsal margin convex. Lateral projection of anteroventral nodal carina well-developed, with acute or slightly blunt tip feebly projecting anterodorsad. Dorsal view: Anterior margin of petiolar node approximately two-thirds shorter than width of posterior margin. Lateral view: Subpetiolar process subtriangular, with rounded anterior cusp, followed by posterior straight slope (ca. 30°), with two slightly divergent longitudinal carinae separated by groove. Gaster. Lateral view: Meeting of anterior A3 margin with dorsal margin relatively angled. Cinctus moderate. Pygostyles close to three times longer than broad. Dorsal view: Tergum of A3 shorter than A4. Ventral view: Abdominal sternum 7 showing even surface with no ditches, anterior half glabrous, posterior half with semicircular patch bearing abundant, appressed, stout hairs. Spine at posterior margin of tergum of A8 acute, semitubular (not dorsoventrally flattened). Abdominal sternum 9 roughly as long as broad. Legs. Mesofemur thickened medially, thicker than metafemur, in dorsal or ventral view. Ventral surface of meso-and metafemora flattened roughly on distal two-thirds, surface even, without groove. Genitalia. Ventral view: Cupula: posterior margin of medial invagination narrow. Lobular process vestigial. Gonopod dorsally subtriangular, subrectangular in lateral view, with subtriangular posterior half strongly directed ventrad, dorsolateral margin relatively straight medially, with medial, oblique, weak incision. Gonostylus subacute apically, dorsointernal margin with rounded, tooth-shaped, swelling. Basivolsella externally nearly straight. Digitus spatulate, with outer margin round in ventral view. Penial sclerites overall outline axeshaped, dorsally broadly round. Dorsoapical third of penial sclerite surface with strong flap. Color. Appendages including antennae and mandibles brown to ferruginous brown, occasionally legs yellowish and mandibles lighter than remaining integument of body. Sculpture. Striae absent on head, axillula, metapleuron and lateral propodeal surface slightly rugose, on petiolar node usually absent, though when present, feebly impressed on lateral face. Natural history notes. Neoponera mashpi sp. nov. has been collected mostly by hand and Malaise trapping. The elevation records vary from 3 to 746 m, being this species mostly found in preserved mature habitats from lowland and pre-montane rain forests in Central America, to Andean mountainous and Pacific lowland forests from the Chocó-Darien, to dense and open Amazonian and Atlantic ombrophyllous forests in South America. J. Longino (1999) observed this species raiding on unidentified termites in Costa Rica; he also briefly described a possible colony relocation of about 1400 to 1800 workers moving out from an underground nest into the remains of a termite nest, and over its entrance he noted a hovering cloud of unidentified fungus gnats. Adrian Troya observed a few foraging individuals approaching an unidentified termite nest built inside a fallen log in a tropical Andean cloud forest (Reserva de Biodiversidad Mashpi), northwestern Ecuador, the ant colony was located near their potential prey source some 5 m away.
Virtually all published observations on behavioral biology made by previous authors in the island of Barro Colorado (BCI), Panama, which have been attributed to N. laevigata, probably correspond to N. mashpi sp. nov.: : colony activity and diet (termite predation); Borgmeier (1959): diet (termite predation); Downing (1978): foraging and migratory behavior; Hölldobler and Traniello (1980): pheromone recruitment communication; Traniello (1981): behavior against termite deterrence; Longino (1999): foraging and migratory behavior; : sporadic gyne and male collections. Although we could not find vouchers from that specific Central American site, we did examine specimens of N. mashpi sp. nov. from the Panamanian regions of Coclé and Colón, which are in relative proximity to BCI.
Comments. Neoponera mashpi sp. nov. can only be confused with two species in the N. laevigata group: N. laevigata and N. gojira sp. nov. The remaining species, N. commutata and N. marginata are clearly different, with the worker caste in the former being much larger (WL = 5.59-5.68 mm), whereas the latter shows a strongly excavated dorsal groove on the mandible, in frontal view (Fig. 12a), this groove is absent in N. mashpi sp. nov. The following features facilitate separating N. mashpi sp. nov. from N. laevigata and N. gojira sp. nov. in the worker caste: when compared to the first, the scape is smaller, when pulled back it does not reach the posterior head margin by at least one apical scape width, this is usual in the smallest workers and queens, but see also diagnosis ( Fig. 6a), while in N. laevigata the scape always surpasses said margin by close to one apical scape width (Fig. 6b); the lateral nodal face is smooth (Figs 25d, 26b), while that of N. laevigata always bears oblique striae. However, N. mashpi sp. nov. populations from Central American Atlantic forests may also show nodal striations laterally (see distribution notes below), though less strongly impressed than in N. laevigata. The subpetiolar process is subtriangular with an acute anterior cusp, in lateral view (Fig. 25d), in N. laevigata the subpetiolar process is also subtriangular but with an anterior keel-shaped cusp, followed posteriorly by a relatively flat or slightly concave surface (Fig. 13d); prora well-developed (Fig. 15d), while in N. laevigata the prora is either absent or shaped as a tiny cuticular lip (Fig. 13d). Workers of N. mashpi sp. nov. differ from N. gojira sp. nov. in the following features: dorsal masticatory mandibular surface smooth, with few shallow piligerous fossulae medially (Fig. 12c), in N. gojira sp. nov. this region has a distinct longitudinal rim with piligerous foveae and longitudinal striae medially, Fig. 12b); flap at metapleural gland opening well-developed and strongly curved anterad so that the orifice of the gland is barely visible in posterolateral view (Fig. 2a), while in N. gojira sp. nov. the flap is reduced to a narrow, poorly discernible, blunt carina, leaving the gland orifice completely visible in posterolateral view, Fig. 2d; prora well-developed, while in N. gojira sp. nov. the prora is tiny and lip-shaped, Fig. 19e); clearly smaller (WL = 1.66-3.31 mm), whereas the holotype of N. gojira sp. nov. is the largest worker of the N. laevigata group (WL = 3.72 mm); the area of the pheromone venting canal at the metapleural gland opening in lateral view (Fig. 2a) is clearly less impressed than in N. gojira sp. nov. (Fig. 2d).
The queen: in N. mashpi sp. nov. the queen is smaller (WL = 3.67-4.08 mm) than that of N. laevigata , the ocelli are also smaller: ca. 0.085 mm vs ca. 0.14 mm. The scape in N. mashpi sp. nov., when pulled back, fails to reach posterior head margin, whereas in N. laevigata the scape always surpasses said margin by approximately one apical scape width. The petiolar node of the currently known queens of N. mashpi sp. nov. is completely devoid of striae, while in N. laevigata those striae are always strongly impressed and present laterally on the nodal face. The queen of N. gojira sp. nov. is unknown.
The male: we only found two specimens of N. mashpi sp. nov. from the Brazilian Amazon, both bearing a well-developed prora which is one of the diagnostic features for the worker, and according to our observations of the current set of specimens in the N. laevigata group, the form and size of this structure is relatively conserved among all castes in each species. These males are virtually identical to the specimen identified as "Pachycondyla" laevigata by W. Mackay from the Pacific side of Costa Rica, Estación Pitilla (AntWeb record: INBIOCRI001099170, image examined). The Brazilian specimens slightly differ from this Costa Rican specimen in the following: outline of head feebly straight posterad to eyes (rounded in INBIOCRI001099170), and propodeal posterolateral surface more striated than in INBIOCRI001099170. Populations of N. mashpi sp. nov. from the Pacific regions of Central America do not show lateral nodal striae as is the case for INBIOCRI001099170. Thus, we can assign (for now) this Costa Rican male to N. mashpi sp. nov. Since the males of N. commutata greatly differ from the other known males examined in this study, e.g., in bearing torular lobes (absent in remaining known species), those of N. mashpi sp. nov. can only be confused with the males of N. marginata. Among other features, these two species can be distinguished mainly by the presence of a median longitudinal line (but can also be seen as a shallow groove) on the mesoscutum of N. marginata (absent in N. mashpi sp. nov.), and by the spine on the tergum of A8 being dorsoventrally depressed and with blunt posterior tip in N. marginata (conical and acute posteriorly in N. mashpi sp. nov.).
Neoponera mashpi sp. nov. is very similar to N. laevigata and since both occur in sympatry in several ecoregions, mostly in South America (Fig. 29a), they could well be sister lineages as suggested by preliminary molecular analyses (Troya et al. unpubl. data). Additional insights from future studies, for example through population genetics, may potentially evidence a recent divergence. Some of the differential characters presented above, for example, the shape of the subpetiolar process in workers and queens, show certain degree of continuous variation in some examined populations (see distribution notes below). This may cause difficulty in its separation from correspondent castes of N. laevigata, however we consider these variations within the taxonomic limits of N. mashpi sp. nov. In prior treatments of the N. laevigata group,  and , these two forms were treated as a single species. The work of Wheeler partially depicts our definition of N. laevigata, nevertheless, the two workers illustrated in his plates (p. 161) show a developed, ventrally directed prora which we consider diagnostic for N. mashpi sp. nov. We assume that the specimens on which  based his descriptions were a mixture of both N. mashpi sp. nov. and N. laevigata. , on the other hand, represent in their descriptions mostly the morphology of N. mashpi sp. nov., although without providing details on the prora, subpetiolar process, and the flap of the metapleural gland opening, the first two being key diagnostic characters.
Distribution notes. Populations of the Atlantic forests of Central America: Panama, Costa Rica, up to southern Nicaragua, show very tenuous to clear petiolar node striae, though not as strongly impressed as in the Amazonian populations of its known closest species, N. laevigata. The remaining examined populations of N. mashpi sp. nov., including those from the Pacific forests of Costa Rica, south to Amazonia and the Atlantic forests of Brazil (Bahia), lack these striae. This gradual variation in sculpture, i.e., well-impressed striae turning into smooth surfaces, from South American regions north to Central America, has been observed in other ant groups as well (J. Longino, pers. comm.). Despite that, our observations on this trait do not support or suggest a lineage split between Central American and South American populations, though they could represent ongoing hybridization between both species.
Populations of N. mashpi sp. nov. and N. laevigata, are distributed in sympatry in several Amazonian and Chocó-Darien sites. Our current specimen set supports a Central American distribution up to northern Costa Rica only for N. mashpi sp. nov., whereas N. laevigata populations reach up to the mid-western tropical regions of the Cordillera Occidental, up to approximately the Golfo de Cupica in the Chocó-Darien of Colombia. N. mashpi sp. nov. shows a broader range in South America, reaching the Atlantic Forest in northeast Brazil (Rio Grande do Norte and Minas Gerais), to the southern Peruvian Amazon.
Except for the petiolar node striae present in Central American N. mashpi sp. nov., the diagnostic traits of this species are relatively stable across examined specimens from Amazonian and Atlantic forests. Costa Rican records from Guanacaste, to the northwestern Pacific, represent the northernmost distribution range of this species. Thus far no records of any species in the N. laevigata group have been found north of southern Nicaragua (J. Longino, pers. comm.), with the Hess Escarpment potentially acting as a biogeographic barrier not only for this group but for other biota as well (see Mendoza et al. 2019).