﻿Corrieoponenouragues gen. nov., sp. nov., a new Ponerinae from French Guiana (Hymenoptera, Formicidae)

﻿Abstract This study describes the worker and queen castes of the Neotropical ponerine Corrieoponenouraguesgen. nov., sp. nov., an ant from the tropical rainforest in French Guiana. Worker morphology of the taxon is compared with those of other Ponerinae and the similarities between them are discussed, refining the definition of character states for some diagnostic characters at the generic level, providing an identification key to the Neotropical genera, and making some adjustments to the taxonomic framework within the subfamily. Descriptions, diagnosis, character discussion, identiﬁcation key, and glossary are illustrated with more than 300 images and line drawings. Open science is supported by providing access to measurement data for specimens of the new genus, a matrix of character states for all ponerine taxa evaluated in this study, and specimen data for all examined material. The new or revived combinations presented here are Pachycondylaprocidua Emery, comb. rev., Neoponeracuriosa (Mackay and Mackay), comb. nov., Leptogenysbutteli (Forel), comb. nov., and Bothroponeraescherichi (Forel), comb. nov. In addition, Leptogenysbutteli is synonymized with Leptogenysmyops (Emery), syn. nov.


Introduction
Since 2001, 17 Ant Course editions have trained nearly 500 students from 59 countries, included more than 60 international instructors, and offered opportunities to explore the biological diversity in different parts of the globe, from Australia to Southeast Asia, to East Africa, and to North, Central, and South America (www.antcourse.org). Specimens collected in past editions enhanced our understanding of several aspects of ant biology, such as functional morphology (e.g., Peeters et al. 2017;Larabee et al. 2017Gibson et al. 2018), ecology (Kronauer 2004), reproductive biology (Peeters 2017), and natural history (LaPolla et al. 2004;Lattke and Delsinne 2016). The course also created opportunities for remarkable discoveries. For example, two new genera records for USA fauna, Typhlomyrmex Mayr (CASENT0173317) and Fulakora Mann (MCZ-ENT00528501); the first record of the worker caste of the Afrotropical genus Aenictogiton Emery (CASENT0906052); and two genera recorded for the first time in Borneo (Rhopalothrix Mayr and Tyrannomyrmex Fernández;Fisher et al. 2015).
Here we describe a novel ponerine genus and species discovered during the 2018 Ant Course in the Natural Reserve of Nouragues, French Guiana. Ponerinae Lepeletier de Saint-Fargeau (see diagnosis in Fisher and Bolton 2016: 53-54) currently consists of 47 extant genera and 1,263 valid species (AntCat.org). Of these, 17 genera and 673 species are present in the Neotropics (AntWeb.org). Schmidt and Shattuck (2014) recently revised the higher classification of the subfamily and split the former genus Pachycondyla into 19 genera. The authors also described Iroponera odax, a name coined by WL Brown Jr (Cornell University, Ithaca, USA), who unfortunately passed away before describing that unknown genus and species. Iroponera Schmidt & Shattuck is the most recent ponerine genus description based on a new species rather than reclassification.
As part of the process of diagnosing the new genus, we make a few adjustments to the taxonomic framework within Ponerinae, provide a new identification key for workers of the Neotropical genera, and reassess morphological characters across the subfamily.

Study site
Ant Course 2018 was held in August at Nouragues Research Station (4.08796°N, 52.68002°W) in the Natural Reserve of Nouragues in French Guiana (Figs 1, 2). A French overseas territory on the northeastern edge of South America, French Guiana sits on a Precambrian massif known as the Guiana Shield (Grimaldi and Riéra 2001). The Nouragues station's hilly landscape features a granitic inselberg that rises to 430 m (Fig. 2C, E; Grimaldi and Riéra 2001). The station lies in the Guianan Lowland Moist Forests ecoregion, a subset of the Tropical and Subtropical Moist Forests biome (Dinerstein et al. 2017); the climate is equatorial humid, and seasonality is determined by the movements of the Inter-tropical Convergence Zone (Grimaldi and Riéra 2001). The region experiences an average annual temperature of ~26 °C and yearly average precipitation ~2990 mm, with the rainy season occurring from November to August but interrupted in March by a short dry period (Grimaldi and Riéra 2001).

Material examined and specimen records
During this study, we contrasted the morphology of the new genus with 129 species or subspecies representing all 47 currently valid extant Ponerinae genera (AntCat 2021; see Table 1 for taxa list, Suppl. material 1: Table S1 for specimen data). Suppl. mate- Figure 2. Images of the Nouragues Research Station A ant Course 2018 students working in the lab at the Inselberg camp B view of the Inselberg camp kitchen facilities C forested landscape of the reserve viewed from the Nouragues Inselberg D rainforest along the Approuague river E the Nouragues Inselberg (top left of the image) and surrounding landscape F collection site of Corrieopone nouragues. Photography by BL Fisher. rial 3: Table S3 (character matrix) documents characters evaluated that could be unambiguously discretized, and we share it here to disclose our methodology and foster validation, replication, and reinterpretation of our results (see also Suppl. material 2: Table S2 for character statements).
Specimens were examined with a Leica M125 microscope with 187.5 × total magnification power (Leica Microsystems, Switzerland). Minute characters were accessed with scanning electron microscopy (SEM) or through SEM images previously available on AntWeb (www.antweb.org). Each specimen evaluated bears a registered unique identifier (e.g., CASENT0830464) associated with collection and specimen information, images, and identification on AntWeb. Data is accessible on AntWeb through the persistent URL of a given unique identifier (e.g., www.antweb.org/specimen/CASENT0830464).
Maxillary and labial palpal counts are based on direct observation of specimens with protracted maxillolabial complex (N = 41), dissections (N = 47), and existing SEM images of the maxillolabial complex on AntWeb.org (N = 26). For dissected specimens, the maxillolabial complex was removed from the buccal cavity with a pin or forceps. Larger specimens were observed with a Leica M125 microscope; smaller specimens were accessed with SEM. See Suppl. material 3: Table S3 (column Notes_char_9) for specimen data on palps evaluated, including unique specimen identifiers and observation method.
In subsequent sections, we refer to several other species whose morphology was only assessed through extended-focus images of specimens databased on AntWeb. Those species were not included in Table 1 or Tables S1 and S3. Instead, and when necessary, references are accompanied by the unique identifier of relevant specimens, which are hyperlinked to respective webpages on AntWeb.

Images, drawings, and maps
Extended focus montage images were acquired with a Leica DFC 425 camera and LEICA APPLICATION SUITE software (version 3.8;Leica Microsystems,Switzerland). For most SEM images, samples were coated with gold-palladium in a Cressington 108 Sputter Coater (Cressington Scientific Instruments, United Kingdom), and micrographs were taken at high vacuum secondary electron emission (accelerating voltage: 15 kV, spot intensity diameter: 40) in a Hitachi SU3500 microscope (Hitachi High-Technologies, Japan). Uncoated specimens (e.g., primary types, unique specimens) were also imaged at high vacuum secondary electron emission, but with the accelerating voltage set to 1.5 kV and spot intensity to 50. Image enhancement (e.g., contrast, levels, sharpness, darken background) occurred in ADOBE PHOTOSHOP (version 22.4.2; Adobe Inc., United States of America). All images produced in this study are available on AntWeb.
Line drawings were originally traced in ADOBE ILLUSTRATOR (version 25.2.3; Adobe Inc., United States of America) or modified from artwork produced by scientific illustrator Jessica Huppi, whose authorship is credited in figure captions when pertinent. Jessica Huppi line art is part of a work-for-hire agreement that makes the California Academy of Sciences a copyright holder for her artwork. Mapping of the study area occurred on RStudio Desktop (version 1.4.1717; RStudio Team 2021), an integrated development environment for R (version 4.1.0; R Core Team 2021). Spatial data was sourced directly from the following: NASADEM HGT v001 products (NASA JPL 2020), Global Lakes and Wetlands Database (GLWD-2 dataset; Lehner and Döll 2004), Shuttle Radar Topography Mission Water Body models (NASA JPL 2013), Global River Classification dataset (GloRiC version 1.0, Ouellet Dallaire et al. 2018), and Global Tree Cover dataset (Hansen et al. 2013). Alternatively, data were acquired and imported into R with functions of the following R packages: OSMDATA (Padgham et al. 2017), RASTER (Hijmans 2021), RWORLD-MAP (South 2011), and RWORLDXTRA (South 2012); the OSMDATA package imports features from OpenStreetMap (OpenStreetMap.org data were available under the Open Database License). Data geoprocessing was executed with functions of DPLYR (Wickham et al. 2021), GDALUTILS (Greenberg and Mattiuzzi 2020), RAS-TER, and SF (Pebesma 2018) packages. Map projection and mapping were performed with functions available in the TMAP package (Tennekes 2018).

Terminology
Positional and directional terminology references a hypothetical worker seen in profile, with the head oriented to the left and standing with legs slightly spread over the horizontal plane of a multidimensional space (Fig. 3). Mouthparts are retracted, and antennal scapes are directed posteriad, parallel to the head dorsum. We defined three main directional axes from this orientation, which guided the description of direction and relative position of morphological characters in this study. The anteroposterior axis extends horizontally across the space, from the anterior left side to the posterior right; the dorsoventral axis extends vertically, with "dorsal" directed upwards and "ventral" directed downwards. Finally, the

Taxon name and Author(s)year
Determination methodology Promyopias silvestrii (Santschi) Det. B Bolton Psalidomyrmex procerus Emery Det. WL Brown; A Dejean; BL Fisher Pseudoneoponera porcata (Emery) Det. RA Keller Pseudoneoponera tridentata (Smith) Det. BL Fisher Pseudoponera gilberti (Kempf ) Det. FA Esteves, Mackay and Mackay (2010) ID key; compared with Kempf (1960) description Pseudoponera stigma (Fabricius) Type examined Rasopone costaricensis Longino & Branstetter Type examined Rasopone cryptergates Longino & Branstetter Type examined Rasopone cubitalis Longino & Branstetter Type examined Rasopone guatemalensis Longino & Branstetter Type examined Rasopone panamensis (Forel) Type examined Rasopone pluviselva Longino Kempf (1975) ID key mediolateral axis is perpendicular to a median plane that bisects the ant through its bilateral line, with "medial" directed towards the plane and "lateral" diverging from it (Fig. 3). Additionally, we utilized a fourth axis for aiding the description of positions and directions of characters located on appendages, sclerites, processes, or any structure that may project away from the body. The basoapical axis arises from the median plane, with "basal" being close to the plane while "apical" is distant from it (Fig. 4A). Morphological terminology is based on Keller (2011) unless otherwise stated. Wherever possible, terms in the glossary are hyperlinked to correspondent terms in the Hymenoptera Anatomy Ontology portal (Yoder et al. 2010). Terms absent from the glossary below are defined, illustrated, or associated with references to pertinent literature at first use.
Bulbus neck: The constricted portion of the antennal scape, bordered basally by the bulbus (= condylar bulbus in Fisher and Bolton 2016) and apically by the main shaft of the scape. . Anatomical position and coordinate system adopted as framework for positional and directional terminology. Corrieopone nouragues is shown in profile, with the head oriented to the left. The coordinate system is defined by the median plane and three directional axes (anteroposterior, dorsoventral, and mediolateral). Illustration by FA Esteves.
Comb of strigil (sgc, Fig. 4C): Comb-like structure on the ventral face of the basal portion of the probasitarsus; together with the calcar of strigil, forms the antennal cleaning organ.
Epistomal sulcus (es, Fig. 6A, B; as in Richards 1977): Sulcus delimitating the clypeus posteriorly and laterally from the remainder of the head.
Galea (ga, Fig. 6E): The spatulate lobe located at the apical part of the stipes; part of the maxilla.
Galeal comb: Row of setae located on the outer face of the medial margin of the galea, opposite the maxillary comb located near the inner face of the margin.
Helcium (he, Fig. 5C): Structure formed by the specialized presclerites of abdominal segment III, which articulates with the petiole.
Katepisternum (kn, Fig. 5B): Ventral subdivision of the mesopleuron, separated from the anepisternum by the mesepisternal sulcus (= anapleural sulcus in Fisher and Bolton 2016). Note that this usage does not claim homology between the katepisternum in ants and homonymous mesothoracic areas in other insects.
Labrum (lbr, Fig. 6D): Mouthpart appendage that connects to the anterior margin, or the ventral face of the clypeus, and usually folds over the retracted maxillolabial complex.
Labial palps (plb, Fig. 6E): Labial ring-shaped sclerites articulated with the apicolateral portion of the prementum. We refer to each sclerite as a palpomere and use Roman numerals in ascending order to indicate the position of individual palpomeres along the basoapical axis of the palps (e.g., basalmost labial palpomere = labial palpomere I).
Lower and upper metapleuron (as in Snodgrass 1910): Secondary division of the metapleuron into dorsal wing-bearing and ventral leg-bearing portions.
Maxillary palps (pmx, Fig. 6E): Ring-shaped sclerites articulated with the apicomedial or the apicalmost portion of the stipes in the maxilla. We refer to each sclerite as a palpomere and use Roman numerals in ascending order to indicate the position of individual palpomeres along the basoapical axis of the palps (e.g., basalmost maxillary palpomere = maxillary palpomere I).
Mesoscutellar-axillar complex (as in Gibson et al. 1998): In winged ants, the area of the mesonotum that comprises the mesoscutellum and axillae; located posteriad to the transscutal line.
Mesoscutum (as in Gibson et al. 1998): In winged ants, the region of the mesonotum anterior to the transscutal line and mesoscutellar-axillar complex, whose laterals usually bear parapsidal lines.
Mesometapleural suture (as in Fisher and Bolton 2016): Groove-like suture delimiting the mesopleuron from the metapleuron. Mesonotum (Fig. 5A): The tergum of the mesothorax. Mesopleuron ( Fig. 5B; as in Fisher and Bolton 2016): Pleuron of the mesothorax; extends over the lateral and most of the ventral external surfaces of the second thoracic segment.
Mesosternal process (as in Fisher and Bolton 2016): The pair of cuticular projections surrounding the midline of the mesothorax's ventral face, anterior to the mesocoxal cavities.
Metasternal process (as in Fisher and Bolton 2016): The pair of cuticular projections surrounding the midline of the metathorax's ventral face, anterior to the metacoxal cavities.
Metapleuron (mtp, Fig. 5B; as in Fisher and Bolton 2016): Pleuron of the metathorax; extends over the lateral and ventral external surfaces of the third thoracic segment; bears the metapleural gland opening laterally.
Notopleural suture (nps, Fig. 5B): Suture between mesopleuron and mesonotum. Petiolar laterotergite (lt, Fig. 5B, D): Paired, long, narrow, strip-like area of the ventral margin of the petiolar tergite that is separated from the main part of the sclerite by an impression; flanks the petiolar sternite.
Presclerite: Anterior articulatory region of each abdominal sclerite overlapped by the anterior segment. The term pretergite refers to the presclerite of a tergum (ptg, Fig. 5E), and presternite to the presclerite of a sternum.
Pronotum (pn, Fig. 5A): The tergum of the prothorax, which is hypertrophied in workers, and also in queens and males of some ant taxa.
Prora (pr, Fig. 5C; modified from Boudinot et al. 2020): Anteroventral process of abdominal sternite III that contacts the petiole sternite and gives stability to the ventral flexion of the gaster during stinging. It may be located anywhere from the area between the ventral margins of the helcial tergite arch to the anterior face of the poststernite.
Suture and sulcus: A suture is a groove formed by the fusion of two sclerites; sulcus is an impression that corresponds to an apodeme. Figure 7. Glossary of terminology, part IV: vestiture. Individual parts show SEM images taken from Corrieopone nouragues; paratypes, worker caste. Distinct types of seta and microtrichium are highlighted in orange A buoyant seta B elliptic seta C filiform seta D helicoid seta E hook-shaped seta F lanceolate microtrichium G spatulate seta H spatulate microtrichium I spatulate-costate seta J spatulate-bicuspid seta K stout, spine-like seta L tubiform microtrichium. Specimens imaged: CASENT0872031 (A, C, D, K) and CASENT0923158 (B, E-J, L). Images by FA Esteves; available at AntWeb.org. Scale bars: 0.01 mm. decimal places, and presented as minimum and maximum values with holotype measurement within parentheses. Indices are rounded to the nearest integer and expressed as minimum and maximum values with holotype index within parentheses (see original data in Suppl. material 4: Table S4).

HL
Head length (  the anterior face of the pronotum (i.e., anterior inflection point of the pronotum), excluding the neck.

TL
Total length: Sum of HL + WL + length of segments A2 to A7. A2 to A7 is measured as follows: maximum length of the petiole in profile (PL; Fig. 9B) + A3 to A7, or gaster length (GL; Fig. 9B).

Taxonomic changes in Ponerinae
Before describing Corrieopone nouragues gen. nov., sp. nov., we must make some adjustments to the taxonomic framework proposed for Ponerinae by Schmidt and Shattuck (2014), which otherwise would render comments on morphological similarities and differences among genera in subsequent sections cumbersome. The following changes are based on assessing female morphology among specimens examined in this study, with images available on AntWeb, and information gathered from relevant taxonomic literature.

Transfers between Neoponera and Pachycondyla
Neoponera Emery and Pachycondyla Smith are recognized among other Neotropical ponerines by the following combination of characters: The anterior clypeal margin lacks a pair of large teeth-like projections. Torular lobes are closely approximated. Mandibles are triangular to subtriangular, inserted on the anterolateral corner of the head, and armed with numerous teeth. The metapleural gland orifice is closely skirted medially and posteriorly by a well-developed carina. The propodeal spiracle is usually slitshaped; otherwise, the head presents a bilateral carina between the clypeal margin and the anterior margin of the compound eye, and the pretergite of abdominal segment IV presents a stridulitrum. The mesotibia lacks stout, spine-like setae along its dorsal face, and the metatibia presents two spurs. Pretarsal claws are not pectinate. The petiole sternite lacks a posterior spatulate projection that folds posteriad over the remaining sternite; otherwise, the anterior clypeal margin is convex and angulate (see Suppl. material 3: Table S3; Mackay and Mackay 2010;Schmidt and Shattuck 2014).
According to our assessment, Neoponera and Pachycondyla can be set apart from each other by only two characters: the former genus presents distinct arolia between the pretarsal claws and a stridulitrum on the pretergite of abdominal segment IV; the latter does not (see Suppl. material 3: Table S3; Kempf 1961a;Mackay and Mackay 2010;Fernandes et al. 2014;Schmidt and Shattuck 2014). Contrary to Schmidt and Shattuck (2014), the presence or absence of stout, spine-like setae on the posterior portion of the hypopygium is of little importance to distinguish the genera for two reasons. First, the hypopygium bears spine-like setae in some Neoponera species [e.g.,N. bucki (Borgmeier),N. cavinodis,N. crenata,N. striadinodis,and N. unidentata;Fig. 10A,B], which are similar to those found in Pachycondyla (Fig. 10C, D). Second, in every Pachycondyla species examined (N = 6), some specimens presented a hypopygium armed with aristate setae instead (Fig. 10E). Moreover, among those aristate setae, a few had blunt-apices and resembled spines (Fig. 10F). We assume that this intraspecific and intraindividual shape variation is caused by wear. For example, contact with prey during stinging could erode the apices of pristine aristate setae on the hypopygium, which would then resemble spines.
Neoponera curiosa (Mackay & Mackay, 2010) comb. nov. also conforms with the characters shared by Neoponera and Pachycondyla ( Fig. 13; Mackay and Mackay 2010). In addition, the species possesses a stridulitrum on the pretergite of abdominal segment IV (Mackay and Mackay 2010). According to the authors of the taxon, the arolia are underdeveloped. However, note that the adjective was likely used to compare the taxon with other species formerly assigned to Pachycondyla in which the arolium is distinct. For example, Mackay and Mackay (2010) also described the trait as underdeveloped in N. striadinodis; yet we examined one specimen determined by W. P. Mackay that has distinct arolia.

Transfer from Euponera to Leptogenys and new synonymy
The genus Leptogenys Roger occurs in the tropics and subtropics around the world. Most of its species present pectinate or multidentate pretarsal claws, absent elsewhere in Ponerinae (Bolton 1975;Schmidt and Shattuck 2014;Fisher and Bolton 2016). The clypeal medial area usually projects anteriad into a well-developed, angular prominence, generally skirted anteriorly by a translucid lamella; the median area usually bears a longitudinal carina. Torular lobes are small and only conceal the medial portion of the antennal sockets in dorsal view. Mandibles insert on the anterolateral corners of the head. Mandible shapes range from subtriangular to oblique, to falcate, to linear, to bizarre forms in-between. The propodeal spiracle usually presents a round to oval ori- fice. The metatibia bears two spurs; the metabasitarsus does not bear stout, spine-like setae on its dorsal face. The helcium is infra-axial (i.e., helcium positioned ventrad to the midheight of the anterior face of abdominal segment III).
The species originally described as Pseudoponera butteli Forel, 1913, based on specimens collected in Java, Indonesia, was recently assigned to the genus Euponera Forel by Schmidt and Shattuck (2014). However, the above characters setting Leptogenys apart from other ponerines are visible in images of its syntypes ( Fig. 14; specimens CASENT0907293, FOCOL1012, FOCOL1013) or were mentioned in its description (Forel 1913), except for the shape of the pretarsal claws. On our behalf, entomologist and curator Dr B. Landry examined one of the syntypes deposited at the NHMG and confirmed the presence of three preapical teeth on the hindleg claws. Therefore, E. butteli becomes Leptogenys butteli (Forel, 1913) comb. nov.
When we compared type specimen images and evaluated morphological variation among L. processionalis group members, we found several characters distinguishing between most species in the group. However, we could not find any significant differences between L. butteli and L. myops (Fig. 15). Their type specimens (all from Java, Indonesia) present the same head proportions and sculpture; shape of the torular lobes; shape of the clypeal anterior projection; width of the longitudinal protrusion on the clypeal median area; distance from the apex of the antennal scape to the posterior margin of the head; size, shape, and location of compound eyes on the head; sculpture and shape of the mandibles; dorsal outline of the mesosoma in profile; indistinctiveness of the metanotal sulcus; shape of the mesonotum in dorsal view; height of the petiole in comparison with the propodeum; shape of the petiolar tergite; body-color; and approximate length, amount, and inclination of standing setae. Consequently, we synonymize Leptogenys butteli (Forel, 1913), syn. nov., with Leptogenys myops (Emery, 1887).

Transfer from Mesoponera to Bothroponera
In the Afrotropics, the genera Bothroponera Mayr and Mesoponera Emery can be discriminated from other ponerines by a combination of characters in the worker caste (Fisher and Bolton 2016). The anterior portion of the head lacks a dorsolateral carina between the clypeal margin and the anterior margin of the eye. Torular lobes are closely approximated medially. The anterior margin of the eyes is located at or anteriad to the midlength of the head. Mandibles are triangular and inserted on the anterolateral corners of head; masticatory margins bear four or more teeth; basolateral or dorsal faces lack pits of any shape (an oblique dorsolateral sulcus may be present). The dorsoposterior area of the propodeum is devoid of spine-like or tooth-like projections. The mesotibia lacks stout spine-like setae on its dorsal face, and the metatibia bears two distinct spurs on its ventroapical face. The helcium is positioned ventrad the midlength of the anterior face of abdominal segment IV. Note that contrary to Schmidt and Shattuck (2014), only the basal portion of the masticatory margin of the mandibles is edentate in M. subiridescens; the apical portion is armed with four to seven, but sometimes more, teeth (the count includes denticles).
Bothroponera species have a propodeum with a broad dorsal face, a slit-shaped propodeal spiracle, and a nodiform petiole (Schmidt and Shattuck 2014). The metanotal sulcus is usually obliterated; however, intraspecific variation exists, and the sulcus is weakly impressed in some specimens [as B. ilgii (Forel), specimen CASENT0235600; and B. kruegeri (Forel), CASENT0235604]. Regardless, if present, the sulcus does not interrupt the dorsal outline of the mesosoma in profile. On abdominal segment IV, a strongly impressed constriction separates presclerites from postsclerites, as in B. berthoudi (Forel), specimen CASENT0902470; or moderately so, as in B. silvestrii CASENT0235599. Schmidt and Shattuck (2014) assigned Bothroponera species in two groups. The B. sensu stricto group members share a strongly sculptured body, hypertrophied torular lobes, and a metapleural gland orifice closely skirted medially and posteriorly by a well-developed carina; these characters are absent in the B. sulcata group. Notwithstanding the differential diagnosis given by the authors above, several species of the sensu stricto group [e.g., B. berthoudi, CASENT0902470; B. granosa (Roger), CASENT0250375; and B. laevissima (Arnold), CASENT0902471] have the propodeal dorsum as narrow as those of the sulcata group members.
In Mesoponera, the propodeum is tectiform (i.e., roof-shaped), with its lateral surfaces diverging while sloping ventrad from the noticeably narrow dorsum. In most species, the propodeal dorsal face presents slightly or moderately bulging lateral margins, with the medial area slightly concave posteriorly; the lateral margins may be somewhat parallel to one another (as in M. caffraria, CASENT0915251) or diverge continuously posteriad (as in M. ambigua, CASENT0249194). In a few species, the propodeal dorsum is narrower and transversely convex (as in M. subiridescens, CASENT0003151). The propodeal spiracle is usually round to oval [except for M. caffraria and subspecies, M. ingesta (Wheeler), and M. subiridescens; see CASENT0906219]. The petiole is shaped like an upward-pointing wedge in profile, with the anterior and posterior faces of the tergite tapering to a thin dorsal margin. The metanotal sulcus is clearly distinct and usually deeply impressed, and it indents the mesosoma outline in profile. The constriction between the presclerites and postsclerites of abdominal segment IV may be obliterated to moderately impressed. The torular lobes are not hypertrophied. Body is mostly smooth and shiny or densely and uniformly sculptured by fine punctures. These characters are as described here in every Mesoponera species except one.
The species originally described as Pachycondyla (Bothroponera) escherichi Forel, 1910, based on one worker collected in Eritrea (Fig. 16), was recently combined in Mesoponera by Schmidt and Shattuck (2014). However, the species is clearly misassigned. Mesoponera escherichi presents every character that sets Bothroponera and Mesoponera apart from the remaining Ponerinae genera in the Afrotropics, including the absence of a basolateral or dorsal pit on the mandibles (Forel 1910). The shape of the propodeal dorsal face corresponds to that of the B. sulcata group. The propodeal spiracle is slit-shaped. The petiolar tergite is unaligned in the image we evaluated; even so, it shows a somewhat nodiform petiole (clearly not cuneiform) that is reasonably accommodated by the variation seen in the Bothroponera. Moreover, Forel (1910) stated that the anterior and posterior faces of the petiole are vertical surfaces. The metanotal sulcus is present but weakly impressed, and does not interrupt the dorsal outline of the mesosoma in profile. The presclerites and postsclerites of abdominal segment IV are separated by a moderately impressed constriction. The torular lobes are not hypertrophied. Sculpture is not strongly impressed: most of the head and the dorsal face of the mesosoma is densely foveolate (i.e., covered by small pits that are wider than punctures); the lateral face of the mesosoma is mostly costulate; the petiole is somewhat shiny; the gaster is slightly punctate, and is shinier than the head and mesosoma. Therefore, M. escherichi becomes Bothroponera escherichi (Forel, 1910), comb. nov., a member of the B. sulcata species group.

Identification key for the Neotropical genera of Ponerinae based on workers
We did not consider Pachycondyla vieirai Mackay & Mackay in this key, as it was not examined and the taxon description was uninformative for our purposes. The species was considered incertae sedis in Pachycondyla (Schmidt and Shattuck 2014).

1
Mandibles long and linear in full-face view, inserted at the middle of the anterior margin of the head, their bases closely approximate (Fig. 17A)

3
In full-face view, anterior part of the torular lobes widely separated and usually not confluent; separated by a rounded, truncated, or broadly triangular section of the clypeus (Fig. 19A). The lateral margins of the lobes variously shaped, but only rarely with a pinched-in appearance posteriorly (Fig. 19B) . Mandible pitchfork-like; with indistinct basal and masticatory margins; armed with 3 noticeably long, curved teeth; the apical tooth so long and curved that it reaches or surpasses the anterolateral corner of the head opposite from its insertion when the mandible is closed (Fig. 20AA). Metatibia with only one spur, pectinate (Fig. 20BB). Helcium in profile located ventrad to the midheight of the anterior face of the first gastral segment (abdominal segment III), so that the first gastral segment has a long vertical anterior face in profile. Shape of the petiole ranging from a thick, broad scale with sharp lateral margins to a somewhat cuboid node; never subrectangular to subcylindrical with carinate posterolateral margins. Dorsal surfaces of the head and the mesosoma usually with erect/suberect setae, at least partially. Sculpture varying from smooth and shiny to finely shagreened to finely punctate and rugulose .. Hindlegs usually with pectinate pretarsal claws (Fig. 25A), rarely with only 1-2 small preapical teeth; if pretarsal claws not pectinate, then mandible with only 1-2 teeth; if mandibles with > 3 teeth, then pretarsal claws pectinate. Torular lobes distinctly fail to cover the entire antennal sockets in full-face view (Fig. 25B)
4. * Clypeus complex: In dorsal view, clypeus projected anteromedially as a broad, truncated prominence, overhanging the basal margins of the mandibles, and overlapping the basal portion of the masticatory margins of fully closed mandibles; anterior margin of the clypeal projection approximately as wide as the distance between the lateral arches of the toruli, devoid of stout setae or additional protrusions (Figs 37B, 38A, E). In profile, clypeal projection with a broad anteroventral face (avf, Fig. 39A-D), which extends ventroposteriorly (i.e., obliquely) from the clypeal dorsal face in almost 90 degrees; the ventralmost point of this anteroventral face meets the "true" clypeal ventral face (= surface of the clypeal infold in Boudinot et al. 2021;vf, Fig. 39A-D). In anteroventral view, clypeal anteroventral face subrectangular (avf, Fig. 39D). Median area of the clypeus bulging (Fig. 37A, E); seen in profile, it ascends steeply from the clypeal anterior margin to the torular lobes, with posterior portion slightly convex (Fig. 38B, D). 5. In dorsal view, torular lobes closely approximated. 6. In dorsal view, torular lobes medium-to small-sized: not concealing the lateral arches of the toruli (Fig. 38F). Median and lateral arches of the torulus with discontinuous posterior margins (Fig. 38D).
7. In profile, torular lobes located at the dorsalmost part of a prominence formed by the clypeal median area and the frontal carinae (Fig. 38B).
25. Calcar of strigil with a basoventral lamella (Fig. 41A, B). 26. Probasitarsus with anterior and ventral faces densely vested with spatulatecostate setae (Fig. 41A), except for the shorter, spatulate-bicuspid setae present on the area immediately anterior to the comb of strigil (Fig. 41B). 27. Row of stout, spine-like setae present on the posterior face of the probasitarsal notch, parallel to the comb of strigil (Fig. 41B).

Comments on worker characters
The enumeration below corresponds to character numbers presented above. 1. The mandibles articulate with the anterolateral corners of the head in virtually all Ponerinae and are either triangular or subtriangular in most genera (N = 29). Other mandibular shapes with little intrageneric variation are the oblique (Boloponera, Buniapone, Dinoponera, Iroponera, Plectroctena, and Promyopias), pitchfork-like (Belonopelta, Emeryopone, Thaumatomyrmex), elongate sub-oblique (Streblognathus), and scythe-shaped (Harpegnathos). The shape of the mandibles varies from triangular to elongate-triangular in Centromyrmex; from triangular to subtriangular, to oblique in Cryptopone; from subtriangular to oblique, to falcate, to bizarre forms in-between in Leptogenys and Myopias; from sub-oblique to falcate in Psalidomyrmex; and from subtriangular to oblique in Simopelta. Anochetus and Odontomachus are the only ponerines in which the mandibles insert near the midline of the anterior margin of the head.
2. The mandible of Corrieopone is completely edentate (i.e., devoid of any teeth, denticle, or projected apex). To our knowledge, this condition is virtually absent in other Ponerinae, apart from some species of Leptogenys (see Arimoto and Yamane 2018) and Platythyrea (see Fisher and Bolton 2016).
3. Ponerine may present mandibles ornamented with pits and sulci, which are relatively good diagnostic characters to genera.
3.1. The basolateral pit (= basal pit in Fisher and Bolton 2016) is a round to oblong impression on the basal portion of the lateral face of the mandible. It occurs in most Brachyponera species (excluding species from Borneo, Bali, Krakatau, and Sumatra, according to Yamane 2007), Cryptopone [except for C. guianensis and, based on its original description, C. mirabilis (Mackay & Mackay), as far as we know], and Euponera (see Schmidt and Shattuck 2014). The pit is also present in Fisheropone ambigua (Fig. 45A, B), which disagrees with Schmidt and Shattuck (2014) and Fisher and Bolton (2016).
3.2. The dorsal pit resembles the basolateral pit, although more elongated and impressed on the dorsal face of the mandible. In Ponerinae, it is only present in Dolioponera, Euponera fossigera Mayr (see Mayr 1901), Hagensia, and Iroponera ( Fig. 45C; contrary to Schmidt and Shattuck 2014).
3.3. The dorsolateral sulcus runs obliquely along the mandible, from the basal portion of the dorsal face towards the lateral face. It is widespread among the Ponerinae and may be shallowly or deeply impressed, restricted to the basal portion of the mandible, or present along almost the entire lateral face of the mandible. The sulcus is consistently present in species of Asphinctopone, Boloponera, Buniapone, Centromyrmex, most (perhaps all) Ectomomyrmex, Feroponera, Loboponera, Myopias, Odontoponera, Paltothyreus, Phrynoponera, Plectroctena, Promyopias, Psalidomyrmex, Pseudoponera, and Streblogna- thus (see Suppl. material 3: Table S3; Schmidt and Shattuck 2014;Fisher and Bolton 2016). On the other hand, the presence of this character varies among species of other genera, such as Bothroponera (present in B. crassa, absent in B. pachyderma), Dinoponera (weakly present in D. lucida; absent in D. longipes), Leptogenys (see Bolton 1975), Mesoponera (only present in M. subiridescens), Neoponera [present in N. fauveli (Emery), weakly impressed in N. apicalis, absent in N. fisheri], Platythyrea (present in P. punctata, absent in P. turneri), and Pseudoneoponera (present in P. porcata, absent in P. denticulata). Contrary to Schmidt and Shattuck (2014), the sulcus is distinct in all Rasopone species examined here (Fig. 45D); although short and constrained to the basal region of the mandibles, it is also present in Austroponera castanea (Fig. 45E). Among the Pachycondyla species examined, the sulcus is a shallow and short basal impression save in P. lenis, where it is absent. In Euponera sikorae (and in all other Malagasy species, according to Rakotonirina and Fisher 2013), the lateral face of the mandible bears a longitudinal sulcus that runs apicad from the lateral margin of the dorsal mandibular articulation. Whether this sulcus is present in Euponera species occurring in other bioregions remains unknown, but it is absent in at least two Afrotropical species, E. brunoi and E. sjostedti.
3.4. Plectroctena species present a sulcus that skirts the mandibular masticatory margin dorsally (Fisher and Bolton 2016), which we refer to as the dorsomasticatory sulcus.
Among the specimens examined, the following taxa present mandibles devoid of any pit or sulcus,  N. fisheri, N. laevigata, N. luteola, N. verenae, N. villosa, Odontomachus bauri, Ophthalmopone berthoudi, Pachycondyla lenis, Parvaponera darwinii madecassa, Platythyrea turneri, Ponera alpha, P. pennsylvanica, Simopelta oculata, S. transversa, Thaumatomyrmex fraxini, and T. zeteki. 6. We assessed the size of the torular lobes across taxa examined according to the degree of connection between median and lateral arches of torulus (as in Keller 2011, character 08). Among material examined, the lobes are anteriorly and posteriorly continuous with the lateral torular arches in taxa whose antennal sockets are largely exposed in full-face view (Belonopelta, Leptogenys, and Ophthalmopone). On the other extreme, hypertrophied lobes are anteriorly and posteriorly discontinuous with the lateral arches of the torulus (as in Boloponera, Bothroponera sensu stricto group, Loboponera, Platythyrea punctata, Plectroctena, and Psalidomyrmex). Like Corrieopone, most ponerines present intermediate-sized torular lobes that are anteriorly continuous and posteriorly discontinuous with respective lateral arches. In this latter state, the lobes may conceal entirely or partially the lateral arches of the torulus.
18. The metapleural longitudinal flange is a carina that extends along the metapleuron in profile, with its posterior end immediately dorsad the metapleural gland orifice. When well-developed, it projects laterad or ventrolaterad and may overhang the gland orifice (as defined by Keller 2011: character 62). In Simopelta species (ANTWEB1008589), this flange is strongly projected ventrolaterad and overlaps the gland orifice.
19. The propodeal dorsum presents a well-delimited, narrow, median longitudinal groove in Psalidomyrmex (Fisher and Bolton 2016;ANTWEB1008585). In addition, a vestigial longitudinal groove may be present on the propodeal dorsal face of specimens of the Plectroctena minor group (see Bolton and Brown 2002). The propodeal dorsum is transversely concave along its entire length, or only posteriorly, in Hagensia, Mayaponera (CASENT0249137), several Mesoponera (CASENT0249194), and Pseudoponera (CASENT0923115).
24. We classified the metacoxal cavities as closed or open by integrating the definition given by Keller (2011: character 69) with that in Fisher and Bolton (2016;see also Bolton 2003: character 41). If closed, the medial surface of the metacoxal acetabulum does not have a fenestra, and thus, the metacoxal cavity is not connected internally with the propodeal foramen; the annulus externally encircling the cavity is fused. If open, the internal medial surface of the cavity is fenestrate and connects with the propodeal foramen, and the annulus is unfused. In this case, the annulus may encircle the cavity with its free ends overlapping next to the propodeal foramen; or there may be a gap in the annulus.
An unfused cuticular annulus tightly encircles the metacoxal cavities in most Ponerinae we dissected. This condition occurs in Phrynoponera pulchella specimens from Kenya (CASENT0178203, CASENT0178204, CASENT0217125), which is in accord with Bolton and Fisher (2008a). However, a specimen from Tanzania possesses an annular gap in both metacoxal cavities (Fig. 46H); we are uncertain if those were natural or dissection artifacts, as unfortunately, only one specimen from that population was available to us. Metacoxal cavities also present an annular gap in Phrynoponera transversa and Platythyrea punctata. On the other hand, the annulus is fused and uninterrupted in Harpegnathos saltator, Platythyrea cribrinodis ( Fig. 46F; contrary to Fisher and Bolton 2016), and Myopias darioi (Fig. 46G).
30. Most Ponerinae taxa present two metatibial spurs. Here, all specimens examined present a pectinate posterior spur, and in the majority, the anterior spur is simple, as in Corrieopone (see Suppl. material 3: Table S3).
Finally, we found shape variations among the claws of the fore-, mid-, and hindlegs only in Buniapone amblyops and Promyopias silvestrii. These taxa present a propretarsal claw with a long basal tooth, while their meso-and metapretarsi claws are simple (Fig. 48G-I). This condition contradicts Bolton and Fisher (2008b) and Fisher and Bolton (2016), who stated the claws are simple in P. silvestrii.
35. The petiolar node of Corrieopone lacks a spine-like or any other acute projection and is narrow in profile, with its anterior and posterior surfaces tapering to an insignificant dorsal surface.
Thus, according to our definition, the spatulate projection of the petiolar sternite is present in Austroponera, Asphinctopone, Brachyponera, Megaponera analis, Ophthalmopone, Phrynoponera, Platythyrea, Rasopone, and Streblognathus; see also Fisher and Bolton (2016) for the description of the sternite in some of these taxa. The petiolar sternite in Belonopelta deletrix, immediately anterior to its posterior margin, is pro-jected ventrad and slightly posteriad; the same seems to happen in Thaumatomyrmex fraxini. According to the species redescription given by Mackay and Mackay (2010), Neoponera magnifica may also present the character.
40. Like Corrieopone, most ponerine genera present an infra-axial helcium (i.e., positioned ventrad the midheight of the anterior face of abdominal segment III; see Keller 2011, character 114). A few taxa examined present an axial helcium (i.e., positioned at midheight of the anterior face of abdominal segment III): Boloponera, Buniapone, Centromyrmex, Cryptopone (except C. guianensis), Dolioponera,Feroponera,Iroponera,Platythyrea,and Promyopias. 41. As far as we know, the prora is present and well-developed in most Ponerinae. It is usually indistinct in Platythyrea, but contrary to Fisher and Bolton (2016), it is well-developed in some species [as in P. lamellosa (Roger); Fig. 49H], and weakly projected but still visible in others [as in P. pilosula (Smith); Fig. 49I]. According to Schmidt and Shattuck (2014), the prora is absent in Iroponera, but SEM images of I. odax reveal that it is present, albeit weakly projected (Fig. 49J). According to Fisher and Bolton (2016), the trait is absent in Dolioponera, which agrees with what we saw in most specimens. However, SEM images of specimen ANTWEB1008521 show a weak projection on the anterior face of abdominal sternite III; we are unsure whether that detail is natural or an image artifact.
We consider the prora present in Mayaponera and Rasopone, in disagreement with Longino and Branstetter (2020). The trait, although small, is distinct in the profile view of M. becculata, M. conicula, M. constricta, and R. panamensis. In the first two species and R. panamensis, the prora rises from the anterior portion of the abdominal poststernite III (see specimens CASENT0249130, CASENT0644252). In M. constricta, it projects from the area in between the ventral margins of the helcial tergite arch and the anterior portion of the abdominal poststernite III (Fig. 49K) and resembles the condition seen in Dinoponera, Pachycondyla, and Streblognathus (ANTWEB1014000, CASENT0249148, and Fig. 49G, respectively). In M. arhuaca, M. cernua, M. pergandei, and other Rasopone species, the prora is indistinct in the profile view of undissected specimens, for it is a minute prominence located in the area between the ventral margins of the helcial tergite (as in Brachyponera, Fig. 49L), and may be fused entirely with it (as described for Phrynoponera by Bolton and Fisher 2008a).
43. We considered the girdling constriction present if the surface between pre-and postsclerites of abdominal segment IV was interrupted by a shallow or deep impression on the integument. A line, if present, only constituted a constriction if the integument was depressed.
whether they have a stridulatory function in those species is unclear. Finally, the stridulitrum is present in males of Mayaponera pergandei ( Fig. 50G; contrary to Mackay and Mackay 2010) but absent in the worker caste (Fig. 50H, I). Assuming that W. P. Mackay correctly determined the two males we examined, this intercaste variation is interesting because (1) it has not been observed in ants (see Markl 1973), and (2) it suggests that this trait has a reproductive function in M. pergandei.
47. The hypopygium (abdominal sternite VII) is armed with spine-like setae that flank the sting in some Ponerinae. According to previous studies, the setae are present in Dinoponera, Ophthalmopone, Pachycondyla, Paltothyreus, some Leptogenys, and few Ponera (Schmidt and Shattuck 2014;Fisher and Bolton 2016). However, in Pachycondyla, we found that the hypopygial setae may be spine-like or aristate (Figs 10C-F, 12C, D; see details in the preceding subsection "Transfers between Neoponera to Pachycondyla").

Comparison with similar genera
Corrieopone gen. nov. is morphologically distinctive and unlikely to be confused with any other Ponerinae genus. To our knowledge, no other Ponerinae has a clypeal medial area that projects anteriorly as a broad, truncated prominence that overlaps the mandibles dorsally and presents a broad anteroventral face, which is subrectangular in ventro-anterior view (Figs 37B, 38A-E, 39A-D). In addition, no other ant presents the ventral face of the hypopygium with a longitudinal concavity that posteriorly bears a longitudinal carina and stout, hook-shaped setae.
Despite these features, workers of Corrieopone superficially resemble Asphinctopone, Brachyponera, some Hagensia, some Mayaponera, and most Mesoponera species, because they share the following characters: the eyes are located on the anterior part of the head; the mesonotum is dome-shaped; the metanotal sulcus is deeply impressed; the notopleural suture is conspicuous; the metathoracic spiracles are concealed by a cuticular lobe; the dorsoposterior area of the propodeum lacks spines or tubercles; the petiole is unarmed and scale-like; the constriction between the presclerites and postsclerites of abdominal segment IV is shallowly impressed or absent (Fig. 52A-F).

Superscripts:
a Absent in species from Borneo, Bali, Krakatau, and Sumatra (Yamane 2007 M. caffraria and subspecies, M. ingesta, and M. subiridescens); (5) the apicoposterior area of the metatibia presents an apparent metatibial gland cuticular patch (it is absent in Asphinctopone, Brachyponera chinensis, Mayaponera, and Mesoponera save M. caffraria, and likely its subspecies, and M. ingesta); (6) meso-and metatibiae bear two well-developed spurs (Asphinctopone only has one well-developed spur on each tibia); (7) pretarsal claws are simple (the claws present a preapical tooth in Hagensia); (8) the posterior portion of the petiole sternite is not projected (it presents a spatulate projection that folds posteriad over the remaining sternite in Asphinctopone and Brachyponera); (9) the prora is conspicuous in profile (it is indistinct in profile of undissected specimens in Brachyponera, and some Mayaponera species); (10) the pretergite of abdominal segment IV presents a stridulitrum (the stridulitrum is absent in Asphinctopone, Brachyponera sennaarensis, Hagensia, Mayaponera save M. constricta, and Mesoponera ambigua); and (11) the posterolateral portion of the hypopygium lacks stout, spine-like setae (the setae are present in Mayaponera becculata, and Mesoponera save species occurring in the afrotropics). Moreover, Asphinctopone has an oblong, smooth, bright, yellowish cuticular patch on the dorsal face of its subtriangular mandibles (Hawkes 2010); the cuticular patch is absent from the triangular mandibles of Corrieopone. Preliminary phylogenetic analysis based on molecular data inferred a sister relationship between Corrieopone and Asphinctopone (The Ants of the World Project, unpublished data); however, analysis of a more comprehensive dataset is still ongoing and upcoming results may challenge this hypothesis.
Other castes and stages of development Queen (Fig. 53A-F): Winged. Similar to the worker caste but for the slightly greater body length (HL: 1.65; TL: 8.48), larger compound eyes, presence of ocelli, differences in the mesosoma due to the presence of wings, and darker color. Clypeus as described in the definition of the genus. Parapsidal lines present on the mesoscutum (Fig. 53D). In profile, mesoscutum bulging; higher than the pronotum; slightly higher than the mesoscutellum (Fig. 53C). In profile, scutoscutellar sulcus indenting the dorsal outline of the mesoscutum and mesoscutellar-axillar complex. Mesoscutellum dome-shaped in profile; higher than the metanotum and the propodeum (Fig. 53C). Mesopleuron in profile divided into anepisternum and katepisternum. In profile, metapleuron divided i Among species examined (N = 8), the character is present only in Mesoponera caffraria; it is visible in images of specimens identified as M. ingesta on AntWeb.org. j Among species examined (N = 6), the stridulitrum is absent only in Brachyponera sennaarensis, but see Schmidt and Shattuck (2014: 79). k Present in Mayaponera constricta. l Among species examined (N = 8), the stridulitrum is only absent in Mesoponera ambigua. The character was not assessed in M. elisae rotundata; it is unclear in M. caffraria; but see Schmidt and Shattuck (2014: 109). m Among species examined (N = 8), the character is only present in Brachyponera becculata. n Among material examined, the character is absent in the Afrotropical species.
into upper and lower sections; upper metapleuron separated from propodeum by sulcus; lower metapleuron indistinct from propodeum (Fig. 53C). Propodeal spiracle slitshaped. Legs as in the worker caste, including the presence of the metatibial gland cuticular patch on the apicoposterior face of the metatibia (Fig. 53E). Petiole and gaster Color: reddish brown; lateral surfaces of clypeus and appendages slightly lighter; apex of gaster yellowish (Fig. 37).
Head: Head slightly longer than broad (CI: 87-90). In dorsal view, posterior margin of the head slightly concave medially (Figs 37B, 38A). In profile, posteroventral curve of the head without a projecting flange (Figs 37C, 38B). Mandibles triangular, edentate, devoid of basolateral and dorsal pits or dorsolateral and dorsomasticatory sulci (Figs 37B, 38A, E). Basal three-fourths of the mandibular masticatory margin skirted ventrally by two rows of long, flexuous, helicoid setae, with a row of filiform setae present in between. Mandible with a row of filiform setae running obliquely from the basolateral area of the ventral face to the apex of the lateral face; with setae increasing in length towards mandibular apex; apicalmost seta helicoid. Torular lobes closely approximated (Fig. 38A, F); torular lateral arches visible in dorsal view (Fig.  38F). In profile, torular lobes located at the dorsalmost part of a prominence formed by the clypeal median area and the frontal carinae (Fig. 38B). Lateral arch of the torulus projected dorsad, with a somewhat constricted rim (Fig. 38D, F). Twelve antennomeres (Fig. 54A); bulbus hemispherical, notched on the lateral margin (Fig. 54B); bulbus neck tubular (Fig. 54C, D). Antennal scape (antennomere I) surpassing the posterior margin of the head (SI: 119-120); antennomeres III and IV longer than all other preapical antennomeres save the scape; antennomere III almost as long as the apicalmost antennomere (Fig. 54A). Compound eyes present, located just anterior to the midlength of the head, surrounded by a sulcus; widest diameter of compound eyes: six or seven ommatidia (Fig. 38A, B, D). Ocelli absent. Labrum apically bilobed, with lobes broadly rounded apicolaterally (Fig. 39E); long, acute cleft at midpoint of apical margin. Middle of outer face of the labrum bulging transversely, with a short, median carina extending basad from the median cleft; the carina may be a weak, blunt pro- trusion or a somewhat developed ridge (Fig. 55A, B). Palpal formula 4,4 (Fig. 55C); basalmost maxillary palpomere short, bulbous. Outer face of maxillary stipes without a transverse sulcus (Fig. 55C). Galeal comb present (Fig. 55D). Galeal crown armed tance in organizing the course in French Guiana. Financial support for this study was provided by an Investissement d'Avenir grant of the Agence Nationale de la Recherche (CEBA, ANR-10-LABX-25-01). In addition, the research was supported in part by NSF DEB-1932467. We are grateful to the Nouragues Natural Reserve for allowing sampling in the protected area and the Nouragues Research Station and CNRS Guyane for logistical support. A special thanks to the Station team who made the course a success: Bran Leplat, Philippe Gaucher. In addition, we thank the Ant Course 2018 participants: Adam Khalife, Alexandre Ferreira, Cédric Chény, Charlotte Francoeur, Christian Peeters, Cody Cardenas, Frederic Petitclerc, Iryna Ivasyk, Jack Longino, Jason Williams, Jill Oberski, Julie Miller, Lily Leahy, Maria Fernanda Brito de Almeida, Mark Wong, Mélanie Fichaux, Naoto Idogawa, Pamela Eugenia Pairo, Peter Flynn, Phil Ward, Philipp Hoenle, Rachel Wells, and Philippe Gaucher.
We express our gratitude to Bernard Landry (MHNG, Genève, Switzerland) for examining one of the syntypes of Leptogenys butteli deposited in the MHNG. Many, many thanks to Roberto A Keller (Museu Nacional de Hist. Natural e da Ciência, Lisboa, Portugal) for making freely available his innumerous SEM images on AntWeb.org; this study greatly benefited from the exceptional information those images contain. We extend our gratitude to our two reviewers and the editorial team of ZooKeys, whose graceful comments, constructive criticism, and suggestions greatly improved this manuscript.
Finally, we cannot thank enough the extraordinary Michele Esposito (CASC, San Francisco, USA), whose generosity, expertise in photography, design, databasing, and overall support in the lab have contributed to this study in immeasurable ways -thank you!

Table S2
Authors: Flavia A. Esteves, Brian L. Fisher Data type: Мorphological. Explanation note: Here, we list and define morphological characters used to compare the morphology of Corrieopone nouragues gen. nov., sp. nov. with other Ponerinae genera. When producing this list and the associated character matrix (Table  S3), our goals were to document and organize our observations. It only contains characters which we could unambiguously discretize. We share it to disclose our methodology and foster validation, replication, and reinterpretation of our results. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1074.75551.suppl2

Table S3
Authors: Flavia A. Esteves, Brian L. Fisher Data type: Morphological. Explanation note: Table S3 documents states of characters we evaluated across ponerine genera (see character statements in Table S2). When producing this matrix, our goals were to document and organize our observations. It only contains characters which we could unambiguously discretize. We share it to disclose our methodology and foster validation, replication, and reinterpretation of our results. Character numbers correspond to those in Corrieopone genus diagnosis. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1074.75551.suppl3