Corresponding author: Michael G. Branstetter (
Academic editor: M. Engel
The genus
Understanding of the diversity and distribution of
On a global scale, understanding of
The HOC currently consists of 44 extant species distributed across the Nearctic and Palearctic regions, with most Nearctic forms occurring north of Mexico (only a few records from northern Baja California). Species in the MAC occur from the southwestern U.S.A. to northwestern South America (Colombia, Ecuador), with most of the diversity occurring in the wet forests of eastern Mexico and Central America (
Map showing the global distribution of the Middle American clade of
I present here the first comprehensive revision of the Middle American clade of
The first realization that Middle America contained a diverse radiation of previously unknown
Biological information specific to individual species is provided below in the species accounts. Here I present a summary of what is known about the natural history of MAC
Neotropical
An exceptional characteristic of
Most
One of the most intriguing recent discoveries has been the observation that some
Morphological observations were performed at up to 63x magnification using a Leica MZ12.5 stereomicroscope. Measurements were made with Syncroscopy Auto-Montage software by using the measuring tool (calibrated with a stage micrometer) on single images taken at 70× magnification with a JVC KY-F75U digital camera attached to a Leica MZ16A stereomicroscope. Color montage images were created using the same equipment setup used for measurements, except stacks of images were combined with the program Zerene Stacker (
Collection and specimen data for all material examined in this study, along with all color images, have been uploaded to AntWeb (
Collection codes link specimens to collection and locality information. They are essentially “lot numbers,” meaning they apply to many specimens, and should not be confused with specimen codes. When collections are from individual collectors these codes usually are formed by the collector’s initials followed by a number, e.g. MGB1471 for a personal collection. Collections from the ALAS or LLAMA projects have longer codes, in which each part of the code contains information specific to the project, e.g. Wa-B-03-1-32 for a LLAMA collection. All data from the ALAS and LLAMA projects are available from their respective websites (addresses given above). If labels lacked a collection code, then a generic ANTC# code was added to the pin.
Specimen codes, also referred to as “unique specimen identifiers,” were attached to all pins examined in this study. It is usually intended that these codes refer to single specimens only, but some pins studied here included multiple mounted specimens. In these cases, rather than add multiple specimen codes to a single pin, or remove specimens from a pin, I treat the specimen code as a pin code and add text following the code to unambiguously identify which specimen is being referenced (e.g. top specimen).
Specimen and collection data were transcribed primarily from an electronic database and therefore may not exactly match label information. All distances and elevations are provided in metric units, converted from feet or miles when necessary. All latitude and longitude coordinates are provided in decimal degrees, with up to five decimal places, depending on the precision of the measurement. In cases where coordinates were not present on a label, these were estimated using Google Earth (
The material examined section of each species account presents an overview of a species distribution (i.e. map data). It is not an exhaustive list of every specimen examined for a particular species. In general it includes one collection from every site where a species has been collected, with each site separated by at least 2km from other sites. In some cases a series of very close records were included to capture elevational range information at a particular site. The format for this section is the same as for type material, except without the [Error], [Habitat], [Microhabitat], [Collection Code], [Repository], or [Specimen Code] fields. Complete data for all examined specimens are available on AntWeb.
Maps for each species were generated using the software ArcGIS v10.1 (Esri, Redlands, CA). The mountainous basemap used in all maps was accessed within ArcMap, but is attributed to the U.S. National Park Service.
The underlying philosophy driving the separation of species in this study is that of the biological species concept, in which good species represent reproductively isolated entities consisting of one to many populations connected by gene flow (
Morphological variation in
All new species names in this paper should be treated as nouns in apposition and thus invariant. This holds true even if the derivation of a name suggests otherwise.
All measurements presented as a range of values in mm, with the holotype specimen's measurements in parentheses unless otherwise stated (see
Standard measurements used in this study.
Several characters useful in distinguishing
Clypeus structure in
The anterior clypeal margin varies greatly (see
The structure of the basal margin of the mandible usually correlates with the shape of the anterior clypeal margin (see
The clypeus and basal margin of the mandible are very useful in species identification. Consequently, it is highly recommended that specimens be mounted with mandibles open. If all specimens from a series have the mandibles closed, then the mandibles should be carefully pried open (before mounting) by pushing a pin between the mandibles from the ventral side of the head.
Comparison of gastral pilosity among several
Comparison of several different sculpture patterns in
Comparison of the frontal lobes and posterior extension of the clypeus in several
Comparison of petiole and postpetiole shape in several
Comparison of promesonotum and propodeal spines in several
The MAC is distributed from the southwestern U.S.A. (Arizona, New Mexico, Texas) to northwestern South America (Colombia, Ecuador) (
Using just the worker caste it has not been possible to find diagnostic features that adequately distinguish the MAC from the HOC. In general, MAC species exhibit a much greater diversity of phenotypes, resulting in an abundance of autapomorphies. The following features help define the MAC: clypeal carinae usually faint or absent (usually more distinct in HOC); structure of the anterior clypeal margin more variable than in HOC, often forming distinct teeth (usually a simple median emargination in HOC); structure of the basal margin of the mandible more variable, often sinuous, with a distinct basal depression or notch (usually straight in HOC); surface sculpture often reduced, sometimes completely smooth and shiny (HOC species usually completely sculptured); form of pilosity on gastral dorsum more diverse (HOC species usually with pilosity short, bilayered, and not noticeably thickened); gastral tergites and sternites usually smooth and shiny (only sometimes punctate in
A global diagnosis of the worker caste of
Using worker morphology and results from molecular phylogenetic analysis (
List includes 40 species, 33 of which are new. See Supplementary file for a table that links the species names presented here with the code names used in
1 | Anterior constriction of gaster, along with basal striae, distinctly elongate ( |
2 |
– | Anterior constriction of gaster and basal striae not elongate ( |
3 |
2 (1) | Surface sculpture almost entirely smooth and shiny ( |
|
– | Face and much of mesosoma with conspicuous carinulae ( |
|
3 (1) | Anterior clypeal margin in full-face view forming three well-defined teeth (visible even if mandibles closed), with middle tooth formed by median lobe, which projects over true clypeal margin ( |
|
– | Anterior clypeal margin in full-face view variable, not as above ( |
4 |
4 (3) | Face and promesonotal dorsum foveate to coarsely rugoreticulate ( |
5 |
– | Sculpture variable, but not distinctly foveate; other characters variable, not as above | 6 |
5 (4) | Anterior clypeal margin forming four relatively sharp teeth (middle teeth only visible if mandibles open) ( |
|
– | Anterior clypeal margin with a median emargination that has four well-defined to completely effaced blunt teeth ( |
|
6 (4) | Lateral apex of hypostomal bridge projecting ventrally as a subquadrate ( |
7 |
– | Lateral margin of hypostomal bridge tapering to a narrow point, not visible in profile view ( |
11 |
7 (6) | Propodeal spines long and slender (SSL 0.17–0.22, PSI 1.9–2.2) ( |
|
– | Propodeal spines tuberculate or short; face sculpture and petiole variable, not as above | 8 |
8 (7) | Pronotum conspicuously punctate, with longitudinal rugulae faintly present among punctae on dorsum ( |
|
– | Lacking one or more of the above character states | 9 |
9 (8) | Petiole and postpetiole more slender, with postpetiole in profile appearing particularly small and elongate ( |
|
– | Waist segments more robust, with postpetiole in profile sometimes bulging and usually more circular ( |
10 |
10 (9) | Eye usually smaller (EL 0.10–0.16), with 5–8 ommatidia at greatest diameter; body color usually lighter, dark red-brown to brown, but sometimes orange-brown or yellow-brown ( |
|
– | Eye usually larger (EL 0.14–0.19), with 8–11 ommatidia at greatest diameter; body color mostly black ( |
|
11 (6) | Basal margin of mandible straight to slightly sinuous, without a distinct basal notch or depression ( |
12 |
– | Basal margin of mandible sinuous, with a shallow to deep basal depression or notch, but without a basal tooth ( |
30 |
– | Basal margin of mandible sinuous, with a distinct basal depression or notch, and an accompanying small tooth ( |
41 |
12 (11) | Propodeal spines absent; petiole and postpetiole almost completely smooth and shiny, with only faint vestigial punctae sometimes present ( |
( |
– | Propodeal spines present or absent; petiole and postpetiole not as above, either with more conspicuous sculpturing, or with the postpetiole not distinctly larger than petiolar node | 15 |
13 (12) | Face almost completely smooth and shiny, with only some vestigial carinulae near frontal lobes and anterolateral margins of head ( |
|
– | Face with more extensive sculpture, consisting of a fan of carinulae that extend from the area around antennal insertions to at least midpoint of head, but sometimes reaching to posterior margin ( |
14 |
14 (13) | Dorsal and declivitous faces of propodeum in profile flat, and forming a blunt 90° angle ( |
|
– | Propodeum in profile more rounded, with transition between dorsal and declivitous faces less abrupt ( |
|
15 (12) | Face mostly smooth and shiny, except for scattered piligerous punctae and at most a few carinulae, rugulae, and/or punctae around frontal lobes and anterolateral margins of head ( |
16 |
– | Face sculpture more extensive, usually completely sculptured; type of sculpture variable ( |
21 |
16 (15) | Dorsum of promesonotum mostly smooth and shiny, except for scattered piligerous punctae, and at most a few longitudinal rugulae ( |
17 |
– | Dorsum of promesonotum with more extensive sculpturing, either reticulately costate or rugoreticulate ( |
19 |
17 (16) | Mesosoma compact, with promesonotum somewhat bulging ( |
|
– | Lacking one or more of the above character states ( |
18 |
18 (17) | Median lobe of clypeus bicarinate, projecting, and with area between carinae distinctly depressed ( |
|
– | Median lobe of clypeus smooth and with a simple median emargination ( |
|
19 (16) | Promesonotum in profile distinctly asymmetrical, with anterior slope long and gently curving, dorsum nearly flat and at a slight downward angle, and posterior slope short and forming a sharp transition with dorsum ( |
|
– | Promesonotum in profile low domed and roughly symmetrical ( |
20 |
20 (19) | Propodeal spines present, long ( |
|
– | Propodeal spines reduced to sharp right angles, or small upward projecting tubercles ( |
|
21 (15) | Propodeal spines absent, with the dorsal and declivitous faces of the propodeum forming a relatively shallow, blunt angle in profile view ( |
22 |
– | Propodeal spines varying from tuberculate to long, or with the dorsal and declivitous faces of the propodeum forming a sharp, steep angle ( |
23 |
22 (21) | Head and mesosoma strongly sculptured ( |
|
– | Head and mesosoma more faintly sculptured ( |
|
23 (21) | Facial sculpture light, largely effaced, usually not extending all the way to posterior margin of head ( |
|
– | Lacking one or more of the above character states | 24 |
24 (23) | Smaller species (HL ≤ 0.76, HW ≤ 0.68, ML ≤ 1.00); eye usually smaller (EL ≤ 0.12), with 2–7 ommatidia at greatest diameter ( |
25 |
– | Larger species (HL ≥ 0.80, HW ≥ 0.70, ML ≥ 1.02); eye usually larger (EL ≥ 0.11), with 5–11 ommatidia at greatest diameter ( |
29 |
25 (24) | Eye small (EL 0.04–0.08, REL 8–14), subcircular, with 2–5 ommatidia at greatest diameter ( |
26 |
– | Eye larger (EL 0.09–0.12, REL 15–21), more oval-shaped, with 4–7 ommatidia at greatest diameter ( |
27 |
26 (25) | Sculpture on face (and much of mesosoma) consisting of an even distribution of short longitudinal rugulae ( |
|
– | Face densely sculptured and mostly rugoreticulate ( |
|
27 (25) | Anterior clypeal margin either entire, or with a nearly imperceptible median notch ( |
|
– | Anterior clypeal margin not as above ( |
28 |
28 (27) | Anterior clypeal margin with a shallow, but distinct median emargination ( |
|
– | Anterior clypeal margin forming 2 small blunt teeth, which straddle the midline ( |
|
29 (24) | Propodeal spines long and slender ( |
|
– | Propodeal spines tuberculate to short ( |
|
30 (11) | Posterior ¼ or more of face smooth and shiny ( |
31 |
– | Face completely sculptured, mostly rugoreticulate ( |
33 |
31 (30) | Promesonotum in profile with relatively sharp transitions between anterior and dorsal faces, and between pronotum and mesonotum ( |
|
– | Promesonotum in profile more smoothly rounded; pilosity on gastral tergites variable, but without stout setae; face and pronotum sculpture variable | 32 |
32 (31) | Postpetiole in profile bulging and distinctly larger than petiolar node ( |
|
– | Postpetiole smaller, about same size as petiolar node ( |
|
33 (30) | Pilosity on gastral tergites predominately suberect and relatively sparse ( |
34 |
– | Pilosity on gastral tergites shorter, denser, and usually distinctly bilayered, with a layer of suberect setae and an equally dense layer of decumbent setae ( |
35 |
34 (33) | Entire face and most of mesosoma densely punctate, or densely carinulate (longitudinal orientation on face), or intermediate, with carinulae emerging from borders of punctae ( |
|
– | Face less densely sculptured, usually mostly rugoreticulate, but sometimes sculpture more polished, with reticulae indistinct ( |
|
35 (33) | Propodeal declivity in profile forming a broadly sinuous connection between propodeal spine and propodeal lobe ( |
|
– | Propodeal declivity in profile straighter, leaving propodeal spine and lobe separated as distinct features ( |
36 |
36 (35) | Petiole in profile appearing longer and more gracile, with node either dome-like and almost completely smooth and shiny ( |
37 |
– | Petiole in profile usually appearing shorter, more compact, and sometimes more robust, with node only shiny on anterior face and usually of moderate size ( |
38 |
37 (36) | Petiole relatively longer (PL/HW 0.60–0.68); petiolar and postpetiolar nodes mostly smooth and shiny ( |
|
– | Petiole relatively shorter (PL/HW 0.53–0.59); petiolar and postpetiolar nodes smooth only on anterior faces; postpetiole relatively larger ( |
|
38 (36) | Postpetiole in profile bulging and distinctly larger than petiolar node ( |
|
– | Postpetiole in profile smaller, about same size as petiolar node or smaller ( |
39 |
39 (38) | Head and mesosoma dark red-brown to orange-brown, with appendages a distinctly lighter orange- to yellow-brown ( |
|
– | Body and appendage color less contrasting, generally dark to light brown ( |
40 |
40 (39) | Pronotum usually longitudinally rugose on most of dorsum and upper half of side, with small patches of smooth cuticle on middle of dorsum and lower half of side, but sometimes dorsum completely rugose or mostly smooth; propodeal spines tuberculate to short, often relatively longer (PSI 1.2–1.8); petiole compact, with a relatively small node that points slightly posteriad ( |
|
– | Pronotum sculpture variable, but usually not as above; propodeal spines absent or tuberculate, often relatively shorter (PSI 1.0–1.4); petiole variable, but often more elongate, or with the petiolar node distinctly enlarged and pointing vertically ( |
|
41 (11) | Mesosoma in profile somewhat elongate, with metanotal groove wide, shallow, and indistinct, and propodeal dorsum markedly long and flat ( |
|
– | Lacking one or more of the above characters states | 42 |
42 (41) | Eye usually larger (EL 0.09–0.18, REL 18–29), with 6 or more ommatidia at greatest diameter (note other characterisctics of this species highly variable) (Nicaragua to Ecuador) | |
– | Eye usually smaller (EL 0.05–0.14, REL 10–20), with 5 or fewer ommatidia at greatest diameter (southern Mexico to Nicaragua) | 43 |
43 (42) | Anterior 4/5 of face sculptured with short evenly spaced carinulae or rugulae (longitudinal in orientation), remaining posterior surface smooth and shiny ( |
|
– | Face completely sculptured, mostly rugoreticulate ( |
44 |
44 (43) | Pilosity on gastral tergites forming a short layer of dense decumbent to appressed setae and a sparse layer of short suberect setae ( |
|
– | Pilosity on gastral tergites forming a layer of stout suberect setae and a very sparse layer of appressed setae ( |
Anterior constriction of gaster in dorsal view
Face and anterior clypeal margin in full-face view
Comparison of clypeal structure (anterodorsal view) and mesosoma (profile) between
Lateral apex of hypostomal bridge
Mesosoma and waist in profile.
Variation in the structure of the anterior clypeal margin and the basal margin of the mandible
Mesosoma and waist in profile
Face sculpture
Dorsum of promesonotum
Face sculpture
Sculpture on dorsum of promesonotum
Mesosoma and waist in profile view
Clypeus in full-face view
Mesosoma and waist in profile
Face
Mesosoma in profile
Face sculpture
Head and eye in profile view
Faceand clypeus in full-face view
Comparison of the anterior clypeal margin, pronotal dorsum, waist, and gaster
Face sculpture
Face sculpture
Petiole and postpetiole in profile view
Gastral pilosity
Face sculpture
Mesosoma in profile
Mesosoma and waist in profile
Body and appendages in profile view
Face sculpture
Gastral pilosity
Integument mostly black to dark red-brown, with appendages uniformly orange-brown, or mostly dark brown changing to orange-brown at extremities; medium- to large-sized species (see HL, ML, PrW below); anterior clypeal margin with a median emargination; basal margin of mandible straight; propodeal spines absent (PSL 0.10–0.15, PSI 0.7–1.0); petiole and postpetiole almost completely smooth and shiny, with only faint vestigial punctae sometimes present; postpetiole in profile bulging, globular, appearing more voluminous than petiolar node; face with a fan of carinulae extending from frontal lobes to just past midpoint of head or further, sometimes reaching posterior and lateral margins, carinulae when completely covering face, often very dense (almost striate); promesonotum completely smooth and shiny, or with a variable number of transverse striae on dorsal surface; promesonotum in profile domed, symmetrical, and moderately to strongly bulging; eye relatively large (EL 0.14–0.18, REL 18–23), oval-shaped, with 8–10 ommatidia at greatest diameter; setae on gastral dorsum sparse, long, and mostly suberect; frontal lobes of moderate width (FLD 0.20–0.29, FLI 29–32), not obscuring torular lobes in full-face view.
Costa Rica to Ecuador.
(17 measured, paratype JTLC000005880 in parentheses) HL 0.77–0.98 (0.85), HW 0.66–0.88 (0.76), FLD 0.20–0.29 (0.23), PCW 0.05–0.09 (0.07), SL 0.65–0.77 (0.72), EL 0.14–0.18 (0.17), ACL 0.64–0.75 (0.70), ML 1.02–1.30 (1.16), PrW 0.51–0.66 (0.57), PSL 0.10–0.15 (0.13), SDL 0.11–0.15 (0.13), PL 0.37–0.51 (0.41), PH 0.22–0.28 (0.25), PW 0.16–0.22 (0.18), PPL 0.24–0.32 (0.29), PPH 0.23–0.28 (0.25), PPW 0.20–0.28 (0.24), MFL 0.78–1.00 (0.88), MTL 0.64–0.78 (0.70), CI 86–93 (90), SI 83–101 (95), REL 18–23 (22), FLI 29–32 (30), PSI 0.7–1.0 (1.0), MFI 84–94 (87), ACI1 61–65 (62), ACI2 94–101 (98).
Medium- to large-sized species; general body color black to dark red-brown, with appendages uniformly orange-brown (type population), or mostly dark brown changing to orange-brown at joints and extremities; setae golden brown; mandible with 5–7 teeth, consisting of 4 distinct apical teeth, a basal tooth, and 1–2 worn teeth/denticles in between; basal margin of mandible straight, without a basal notch or depression; mandible surface mostly smooth, with scattered piligerous punctae and a few striations on base and lateral surface; anterior clypeal margin with a median emargination; median lobe of clypeus obliquely flattened, mostly smooth and shiny, with a short transverse carinula near anterior margin, remainder of clyepeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions somewhat wide (PCW 0.05–0.09), with sides subparallel to slightly diverging anteriorly; frontal lobes of moderate width (FLD 0.20–0.29, FLI 29–32), not greatly obscuring torular lobes in full-face view; head roughly oval-shaped (CI 86–93), with posterior margin flat, to genetly convex, not depressed medially; eye relatively large (EL 0.14–0.18, REL 18–23), oval-shaped, with 8–10 ommatidia at greatest diameter; face with a fan of carinulae extending from frontal lobes to just past midpoint of head (type population) or further, sometimes reaching posterior and lateral margins, carinulae when completely covering face, often very dense (almost striate); gena with some carinulae; posterolateral and ventral surfaces of head smooth and shiny; scape of moderate length (SI 83–101), reaching, but not distinctly surpassing posterior margin of head in full-face view; scape surface mostly smooth, with scattered piligerous punctae; flagellum with distinct 4-segmented antennal club; promesonotum completely smooth and shiny (type population), or with a variable number of transverse striae on dorsal surface, remainder of mesosoma mostly smooth, except for transverse carinulae on propodeal dorsum, and a few rugulae on side of propodeum and mesopleuron; promesonotum in profile domed, symmetrical, and moderately to strongly bulging; metanotal groove distinct, but often shallow; propodeal spines absent (PSL 0.10–0.15, PSI 0.7–1.0); dorsum of propodeum in profile slightly to strongly convex, usually not flat; petiole and postpetiole almost completely smooth and shiny, with only faint vestigial punctae sometimes present, mostly on venters; postpetiole in profile bulging, globular, appearing more voluminous than petiolar node (PPH/PH 0.89–1.06, PW/PPW 0.73–0.84); petiole in profile appearing of moderate length (PL/HW 0.50–0.58); petiolar node in profile nearly symmetrical, dorsum of node broadly rounded, and pointed vertically to slightly posteriad; gaster smooth and shiny, with scattered piligerous punctae; face with short suberect to decumbent pilosity; setae on remainder of body dorsum sparse, long, and mostly suberect; setae on scapes subdecumbent; setae on legs mostly subdecumbent, with longer suberect setae on femoral venters and coxae.
(5 measured) HL 0.82–0.94 (0.82), HW 0.73–0.87 (0.73), FLD 0.23–0.27 (0.23), PCW 0.07–0.09 (0.07), SL 0.70–0.92 (0.70), EL 0.22–0.23 (0.22), ACL 0.71–0.78 (0.72), ML 1.18–1.42 (1.18), PrW 0.66–0.79 (0.66), PSL 0.13–0.15 (0.14), SDL 0.12–0.15 (0.13), PL 0.44–0.52 (0.44), PH 0.26–0.30, PW 0.19–0.23 (0.19), PPL 0.29–0.35 (0.29), PPH 0.27–0.31 (0.27), PPW 0.25–0.30 (0.25), MFL 0.86–0.99 (0.86), MTL 0.68–0.79 (0.68), CI 89–93 (89), SI 88–108 (96), REL 25–30 (30), FLI 30–32 (31), PSI 0.9–1.1 (1.0), MFI 85–91 (85), ACI1 62–63 (63), ACI2 79–103 (103).
Same as worker except for standard queen modifications and as follows: face sculpture usually slightly longer, and denser; pronotum with faint transverse striations; posterior half of mesoscutum with median patch of longitudinal carinulae; scutellum longitudinally carinulate; wing venation as in
See
The nesting biology of
Colonies include one to five closely spaced nests, but the queen, brood, and most of the workers only occupy one of them. Colonies seem to be continually building new nests and occasionally migrating from one to another. Each nest consists of a horizontally oriented ear-like turret that is sunk into a small alcove. The nest entrance is in the middle of the turret. Next to the entrance the workers always maintain a small clay “door pebble” and when the proper stimulus is applied to the nest entrance, such as an army ant or other predaceous ant, an
Each
As I have circumscribed it here,
Variant 1 includes specimens from Costa Rica to Ecuador, but it does not appear to be a monophyletic entity. The specimens in Costa Rica occur at high elevation, above 1500 m and Longino (pers. comm.) reported finding nests in the ground, rather than in clay banks. One nest was found in a small clay hummock in the middle of a trail in forest. The other was in the ground under leaf litter in forest. There is some variation in how dense the facial carinulae are among sites, with a specimen from Las Alturas having very dense carinulae, similar to variant 2. The variant 1 specimens in Ecuador look nearly identical to those in Costa Rica, with some variation in facial sculpture. They are from lower elevation (800–900 m) and have nests in clay banks like the type form of
Molecular phylogenetic data show that variant 1 specimens from Costa Rica form a clade sister to
Distribution map of
Southern Mexico.
(2 measured) HL 0.60–0.63 (0.63), HW 0.50–0.55 (0.55), FLD 0.12–0.13 (0.13), PCW 0.03 (0.03), SL 0.49–0.52 (0.52), EL 0.08–0.09 (0.09), ACL 0.47–0.49 (0.49), ML 0.76–0.81 (0.81), PrW 0.36–0.39 (0.39), PSL 0.08–0.10 (0.10), SDL 0.06–0.08 (0.08), PL 0.27–0.29 (0.29), PH 0.16–0.19 (0.19), PW 0.12–0.13 (0.13), PPL 0.15–0.16 (0.16), PPH 0.14–0.16 (0.16), PPW 0.16–0.17 (0.17), MFL 0.54–0.57 (0.57), MTL 0.42–0.45 (0.45), CI 85–88 (85), SI 95–98 (95), REL 16–17 (17), FLI 23–24 (23), PSI 1.2–1.3 (1.2), PI 53–54 (53), MFI 93–97 (97), ACI1 68 (68), ACI2 94–96 (94).
Small-sized species; general body color brown to red-brown (note observed specimens are older, fresh material almost certainly darker), with appendages lighter, brown to yellow-brown at extremities; setae golden; mandible with 6 teeth; basal margin of mandible sinuous, with a shallow basal depression, but no basal tooth; mandible mostly smooth, except for some conspicuous basal striae, and scattered piligerous punctae; anterior clypeal margin undulating, forming 2–4 small blunt teeth; median lobe of clypeus smooth, without noticeable carinulae, apex of lobe with a short transverse carinulae, remainder of clypeus smooth and shiny; posterior extension of clypeus between antennal insertions somewhat narrow (PCW 0.03), with sides subparallel; frontal lobes of moderate width (FLD 0.12–0.13, FLI 23–24), not obscuring torular lobes in full-face view; head roughly oval-shaped (CI 85–88), posterior margin with a slight median depression; eye of moderate size (EL 0.08–0.09, REL 16–17), oval-shaped, with 5 ommatidia at greatest diameter; face almost completely smooth and shiny, except for a few carinulae around frontal lobes and on genae, and scattered piligerous punctae; scape of moderate length (SI 95–98), not quite reaching posterior margin when laid back; scape surface mostly smooth, with faint striations, and scattered piligerous punctae; flagellum with a distinct 4–segmented antennal club; all of pronotum and most of mesonotal dorsum smooth and shiny, anepisternum rugose, katepisternum punctate, side of propodeum mostly punctate, with a few rugulae, dorsum and declivity of propodeum with transverse carinulae; promesonotum in profile low-domed, roughly symmetrical, with relatively sharp (and distinctive) transitions between anterior and dorsal faces, and between pronotum and mesonotum; metanotal groove of well-demarcated, of moderate width and depth; propodeal spines tuberculate (PSL 0.08–0.10, PSI 1.2–1.3); petiole appearing of moderate length (PL/HW 0.53–0.54); petiolar node in profile of moderate height (PH/PL 0.60–0.65), roughly symmetrical, with anterior and posterior faces almost equal in length, node dorsum rounded, but somewhat narrow, pointing vertically; postpetiole in profile subspherical, slightly smaller than petiolar node (PPH/PH 0.84–0.86); petiole and postpetiole mostly lightly punctate, with anterior faces of nodes smooth; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with thickened standing pilosity; pilosity on gastral tergites forming a layer of stout suberect setae, and a sparse layer of decumbent setae; setae on scapes dense, decumbent to appressed; setae on legs mostly appressed, with a few suberect to subdecumbent setae on femoral venters and coxae.
Unknown.
Unknown.
This species is recorded from a single Berlese sample of sifted leaf litter collected in cloud forest at 990 m elevation.
Known only from the type locality.
Integument mostly black; medium- to large-sized species (see HL, ML, PrW below); gaster with elongate anterior constriction; entire body almost completely smooth and shiny, with only some faint carinulae and punctae; anterior clypeal margin with a median emargination; basal margin of mandible straight, without a basal notch or deep depression; eye relatively large (EL 0.16–0.20, REL 21–24), oval-shaped, with 9–10 ommatidia at greatest diameter; propodeal spines reduced to very small tubercles (PSL 0.10–0.13, PSI 0.9–1.0); gastral pilosity forming a layer of somewhat stout suberect setae, and a very sparse layer of short decumbent setae; frontal lobes of moderate width (FLD 0.19–0.22, FLI 24–26), not completely obscuring torular lobes in full-face view.
Honduras.
(10 measured) HL 0.88–0.99 (0.94), HW 0.77–0.88 (0.86), FLD 0.19–0.22 (0.22), PCW 0.04–0.06 (0.04), SL 0.77–0.85 (0.83), EL 0.16–0.20 (0.18), ACL 0.66–0.71 (0.69), ML 1.19–1.32 (1.26), PrW 0.52–0.58 (0.55), PSL 0.10–0.13 (0.12), SDL 0.11–0.13 (0.11), PL 0.43–0.47 (0.45), PH 0.25–0.28 (0.27), PW 0.19–0.21 (0.20), PPL 0.27–0.31 (0.30), PPH 0.25–0.27 (0.26), PPW 0.25–0.28 (0.26), MFL 1.00–1.13 (1.08), MTL 0.77–0.87 (0.82), CI 87–92 (92), SI 94–101 (96), REL 21–24 (21), FLI 24–26 (25), PSI 0.9–1.0 (1.0), MFI 71–84 (80), ACI1 62–66 (63), ACI2 82–91 (83).
Medium- to large-sized species; general body color mostly black, with patches of dark brown; appendages black to orange-brown, lighter at extremities; setae dark brown; mandible with 6 teeth, but two teeth nearest basal tooth usually worn and indistinct; basal margin of mandible straight, without basal notch or deep depression; mandible mostly smooth and shining, with scattered piligerous punctae; anterior clypeal margin with a median emargination; median lobe of clypeus smooth and shiny, lacking carinae, remainder of clypeus smooth and shiny; posterior extension of clypeus between antennal insertions of moderate to wide width (PCW 0.04–0.06), sides slightly hourglass-shaped; frontal lobes of moderate width (FLD 0.19–022, FLI 24–26), not obscuring torular lobes in full-face view; head roughly oval-shaped, distinctly longer than broad (CI 87–92), with posterior margin slightly depressed medially; eye relatively large (EL 0.16–0.20, REL 21–24), roughly oval-shaped, with 9–10 ommatidia at greatest diameter; head almost completely smooth and shiny, with short faint longitudinal carinulae around midline of face near antennal lobes; scape relatively long (SI 94–101), just reaching posterior margin of head when laid back; scape surface smooth and shiny, except for scattered piligerous punctae; funiculus with a somewhat distinct antennal club; mesosoma almost completely smooth and shining, except for shallow furrows along metanotal grove, and scattered piligerous punctae; promesonotum in profile low-domed and asymmetrical, with the apex slightly anterior of midpoint; metanotal grove shallow, but distinct; propodeal spines reduced to small tubercles (PSL 0.10–0.13, PSI 0.9–1.0); petiole of moderate length (PL 0.43–0.47, PL/HW 0.52–0.57); petiolar node in profile nearly symmetrical, and of moderate height (PH/PL 0.57–0.60), dorsum smoothly rounded; postpetiole bulging and distinctly wider than petiole (PW/PPW 0.72–0.78), anterior face long and shield-like, posterior face short and truncate; petiole and postpetiole almost completely smooth and shining, with some faint punctae confined mostly to the ventral surfaces; gaster with an elongate anterior constriction and with faint dorsal striae, remainder of gaster smooth and shiny, except for piligerous punctae; most of body dorsum with a layer of moderately long and stout standing pilosity; scape with suberect to decumbent setae; gaster with a layer of suberect setae, and a very sparse layer of short decumbent setae; legs with mostly appressed setae, but some suberect setae on coxae and femoral venters.
(1 measured) HL 0.99, HW 0.89, FLD 0.24, PCW 0.06, SL 0.85, EL 0.26, ACL 0.72, ML 1.53, PrW 0.83, PSL 0.16, SDL 0.15, PL 0.58, PH 0.34, PW 0.25, PPL 0.35, PPH 0.32, PPW 0.33, MFL 1.16, MTL 0.89, CI 90, SI 95, REL 29, FLI 27, PSI 1.1, MFI 77, ACI1 62, ACI2 85.
Same as worker except for standard queen modifications and as follows: face with a few distinct carinulae extending from frontal lobes to ocelli; mesoscutum, near posterior margin, and scutellum, with some longitudinal carinulae/rugulae.
Unknown.
This species is a cloud forest specialist ranging from 1550–2030 m elevation, and is known from one leaf litter sample and one nest collection. The nest was collected underneath the bark of a large log in cloud forest near the edge of dwarf forest. The entire nest was not censused, but it was relatively large, with at least 100 workers, a single queen and brood.
Integument mostly black, red-black, or brown; medium to large-sized species (see HL, ML, PrW below); head and mesosoma foveate to coarsely rugoreticulate; eye relatively small (EL 0.09–0.13, REL 10–14), circular, and slightly bulging, with 5–7 ommatidia at greatest diameter; pilosity on gastral dorsum long, dense and mostly suberect; propodeal spines tuberculate to long and robust, usually of moderate length (PSL 0.15–0.37, PSI 1.3–2.9); anterior clypeal margin forming a single shallow to deep median emargination, or rarely, 4 blunt teeth; basal margin of mandible straight to sinuous, sometimes with a broad basal depression, but without a distinct notch or tooth; 4-segmented antennal club indistinct.
Mexico (Atlantic slope) to Honduras.
(21 measured) HL 0.90–1.20 (1.05), HW 0.77–1.15 (1.00), FLD 0.21–0.32 (0.29), PCW 0.04–0.07 (0.06), SL 0.75–1.01 (0.93), EL 0.09–0.13 (0.12), ACL 0.63–0.78 (0.72), ML 1.15–1.62 (1.45), PrW 0.52–0.78 (0.70), PSL 0.15–0.37, SDL 0.08–0.16, PL 0.45–0.64 (0.56), PH 0.21–0.35 (0.27), PW 0.15–0.24 (0.21), PPL 0.21–0.31 (0.26), PPH 0.18–0.28 (0.24), PPW 0.20–0.30 (0.27), MFL 0.81–1.25 (1.10), MTL 0.66–0.95 (0.88), CI 85–96 (96), SI 85–99 (88), REL 10–14 (12), FLI 25–31 (29), PSI 1.3–2.9 (2.3), MFI 86–105 (91), ACI1 62–65 (64), ACI2 78–89 (82).
Medium to large-sized species; general body color usually red-black (type population) to black, with patches of brown, but some populations more uniformly brown; mandibles and appendages always lighter than body, brown to orange-brown; setae golden brown; mandible with 4–8 teeth (usually 7), consisting of 3–4 distinct apical teeth, a basal tooth, and a variable number of inner teeth, which are often worn and indistinct; basal tooth usually of moderate size (type population), but sometimes more robust and projecting; basal margin of mandible straight to slightly sinuous, sometimes with a shallow, broad basal depression, but without a distinct notch or tooth; mandible mostly smooth and shining, with scattered piligerous punctae and a few lateral striae; median lobe of clypeus usually slightly produced and clearly visible in full-face view (type population), but sometimes becoming obliquely flattened and angled more dorsoventrally, making it less visible; anterior clypeal margin varying from having a shallow to deep median emargination (type population), to forming 4 distinct blunt teeth; median lobe usually with a pair of faint longitudinal carinulae that diverge toward anterior margin, apex of lobe with a faint to strong transverse carinula; posterior extension of clypeus between frontal lobes of relatively moderate width (PCW 0.04–0.07), with sides subparallel to hour-glass-shaped; frontal lobes average to slightly expanded outward (FLD 0.21–0.32, FLI 25–31), with underlying torular lobes always visible in full-face view; head in full-face view roughly oval shaped to subcircular (CI 85–97), with posterior margin slightly to distinctly depressed medially; eyes relatively small (EL 0.09–0.13, REL 10–14), circular, and somewhat bulging, with 5–7 ommatidia at greatest diameter; head foveate to coarsely rugoreticulate, shiny, often with a few short costae extending back from frontal lobes, interstices with piligerous punctae; scape relatively short, not reaching posterior margin of head when laid back (SI 85–99); scape shiny, usually with only scattered piligerous punctae (type population), but sometimes more robust, with punctae deeper and broader, becoming foveolae; flagellum with indistinct 4-segmented antennal club; mesosoma robust, foveate to coarsely rugoreticulate, with foveae most prominent on promesonotal dorsum; propodeal spines varying from short tubercles to long robust spines (PSL 0.15–0.37, PSI 1.3–2.9), which are usually spiniform and project dorsoposteriorly (type population), but sometimes form robust vertical pointing triangles; promesonotum in profile varying from being domed and nearly symmetrical (type population), to domed and asymmetrical, with apex occurring anterior of midpoint, to high-domed and asymmetrical; humeral angles rounded and indistinct, to becoming produced and angulate (type population), the latter occurring when the promesonotal side is scalloped slightly inward; metanotal grove present, but variable in depth and degree of distinctness; anterodorsal margin of propodeum in profile flat to distinctly raised into a welt (type population); propodeal declivity with a variable number of transverse carinae, often mostly smooth and shiny; petiole shape in profile usually appearing relatively long and somewhat gracile (PL/HW 0.53–0.63), with a small distinct node (PH/PL 0.45–0.55) (type population), but sometimes petiole is more robust and strongly wedge-shaped, without a clear distinction between the node and peduncle; postpetiole in profile usually low-domed, nearly symmetrical, and appearing as high or slightly smaller than petiolar node (type population), but sometimes postpetiole distinctly larger than petiolar node; postpetiole in dorsal view elongate, and reaching its widest point near posterior margin; waist sculpture variable, nodes usually mostly smooth and shiny, but sometimes more punctate and/or with longitudinal costae or rugulae, ventral surface punctate, dorsal surface of peduncle punctate and with a variable number of rugulae; gaster mostly smooth and shiny, with scattered piligerous punctae, and short furrows on anterior constriction where gaster inserts into postpetiole; most of body with relatively long standing pilosity; scape either with a single layer of mostly decumbent setae, or bilayered with a sparse layer of longer suberect setae over a denser decumbent layer (type population); gastral pilosity relatively long and somewhat dense, with most setae suberect to subdecumbent; setae on legs suberect to decumbent, with some populations having predominately suberect setae (type population) and others mainly decumbent setae, longer suberect setae always present on femoral venters and coxae.
(5 measured) HL 0.96–1.03 (1.03), HW 0.91–0.99 (0.99), FLD 0.26–0.29 (0.29), PCW 0.05–0.07 (0.06), SL 0.84–0.89 (0.89), EL 0.17–0.19 (0.17) ACL 0.68–0.72 (0.71), ML 1.47–1.57 (1.57), PrW 0.76–0.82 (0.82), PSL 0.18–0.28 (0.28), SDL 0.13–0.16 (0.13), PL 0.59–0.61 (0.60), PH 0.28–0.32 (0.31), PW 0.22–0.25 (0.24), PPL 0.27–0.33 (0.32), PPH 0.23–0.28 (0.28), PPW 0.25–0.31 (0.29), MFL 1.01–1.06 (1.06), MTL 0.79–0.85 (0.85), CI 93–97 (97), SI 86–94 (90), REL 17–20 (17), FLI 28–31 (29), PSI 1.1–2.1 (2.1), MFI 87–95 (93), ACI1 63–64 (63), ACI2 80–85 (80).
Same as worker except for standard queen modifications and the following: Propodeal spines less variable (PSL 0.18–0.28, PSI 1.1–2.1), usually present, of moderate length, and thick at base (only Nahá population with spines tuberculate); setae on scape less variable, usually with a sparse layer of longer suberect setae and a layer of denser decumbent setae (only Nahá population with setae uniformly subdecumbent); wing venation as in
Unknown.
The combination of large size, small eyes, and foveate sculpture make
Variant 1 (
Variant 2 (
Variant 3 (
Some additional populations in Chiapas, Mexico, and all of the populations in Guatemala and Honduras, are most similar to the holotype population (La Unión, Guatemala). However, there is considerable variation among populations and some have character states that are intermediate between the holotype form and the different variants just described. Specimens from higher elevations tend to be larger, darker and more robust. The specimens from Purulhá, Guatemala appear especially robust-looking, with very long, sinuous propodeal spines and coarser sculpturing. Interestingly, these specimens have the petiolar node more wedge-shaped, similar to variant 2. Key character states of the holotype population are indicated in the worker description.
Distribution map of
Integument mostly black; medium-sized species (see HL, ML, PrW below); gaster with an elongate anterior constriction, with anterolateral margins of gaster forming hard shoulder-like angles (best viewed dorsally); median lobe of the clypeus projecting out over mandibles, forming a well-defined, blunt apex; basal margin of mandible sinuous, but without a basal notch or deep depression; basal third of mandible distinctly attenuated (dorsoventrally thinned); face with dense fan of carinulae extending to posterior and lateral margins of head; eye large (EL 0.14–0.19, REL 20–24), oval-shaped, with 9–10 ommatidia at greatest diameter; propodeal spines tuberculate (PSL 0.08–0.10, PSI 1.0–1.4); setae on gastral tergites moderately long and sparse, mostly suberect; frontal lobes of moderate width (FLD 0.16–0.21, FLI 22–27), not covering torular lobes in full-face view.
Guatemala.
(10 measured) HL 0.69–0.93 (0.85), HW 0.60–0.90 (0.82), FLD 0.16–0.21 (0.20), PCW 0.03–0.06 (0.05), SL 0.59–0.78 (0.75), EL 0.14–0.19 (0.18), ACL 0.55–0.66 (0.65), ML 0.93–1.23 (1.13), PrW 0.41–0.56 (0.51), PSL 0.08–0.10 (0.10), SDL 0.06–0.10 (0.10), PL 0.30–0.42 (0.38), PH 0.19–0.26 (0.24), PW 0.41–0.56 (0.51), PPL 0.20–0.27 (0.27), PPH 0.19–0.26 (0.24), PPW 0.21–0.32 (0.28), MFL 0.71–0.98 (0.93), MTL 0.55–0.77 (0.73), CI 86–97 (96), SI 86–98 (91), REL 20–24 (21), FLI 22–27 (24), PSI 1.0–1.4 (1.0), MFI 84–92 (89), ACI1 62–68 (63), ACI2 84–94 (87).
Medium-sized species; general body color black to dark brown, with appendages brown to orange-brown; setae dark brown; mandible with 6–7 teeth, consisting of 3–4 distinct apical teeth, an indistinct basal tooth, and 2–3 inner denticles; basal third of mandible distinctly attenuated (dorsoventrally thinned), with masticatory and basal margins somewhat elongated, attenuated section bordered by an oblique carina; basal margin of mandible sinuous, but without a basal notch or distinct depression; mandible surface mostly smooth and shiny, with scattered piligerous punctae; anterior clypeal margin reduced and mostly hidden underneath the median lobe; median lobe of clypeus projecting out over mandibles, forming a well-defined, blunt apex (almost tooth-like), clypeal carinae absent, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions of moderate to wide width (PCW 0.03–0.06), sides subparallel; frontal lobes of moderate width (FLD 0.16–0.21, FLI 22–27), but not completely obscuring torular lobes in full-face view; head usually robust and somewhat heart-shaped (CI 86–97), with posterior margin broadly depressed medially; eye large (EL 0.14–0.19, REL20–24), oval-shaped, with 9–10 ommatidia at greatest diameter; face densely sculptured with a fan of longitudinal carinulae that extend to the posterior and lateral margins, area between eye and antennal insertion with shorter irregular rugulae, interstices near lateral margins faintly punctate; scape of moderate length (SI 86–98), just reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae and a few striations; flagellum with somewhat distinct 4-segmented antennal club; side and anterior half of pronotum mostly smooth and shiny, with a few scattered rugulae; remainder of promesonotal dorsum longitudinally carinate; mesopleuron and side of propodeum with scattered rugulae and faint punctae; propodeal dorsum and declivity with transverse carinae; promesonotum in profile low-domed and somewhat asymmetrical, with apex occurring anterior of midpoint and the anterior slope longer and steeper than posterior slope; metanotal grove present, somewhat shallow; propodeal spines forming small, but sharp tubercles (PSL 0.08–0.10, PSI 1.0–1.4); petiole of moderate length and form (PL/HW 0.44–0.51); petiolar node in profile somewhat small (PH/PL 0.61–0.67), gently rounded to subquadrate, and slightly angled posteriad, anterior slope distinctly longer than posterior slope; postpetiole bulging, distinctly wider than petiole (PW/PPW 0.60–0.71), and anterior gastral constriction (most noticeable in dorsal view), anterior face of node in profile long and shield-like, posterior face truncate; anterior faces of petiolar and postpetiolar nodes smooth and shiny, posterior faces with a few rugulae and punctae; ventral surface of waist segments faintly punctate; anterior constriction of gaster distinctly elongate and with elongate dorsal striae; gaster in dorsal view with shoulder-like anterolateral corners where anterior constriction begins; remainder of gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with long standing pilosity; setae on scape subdecumbent to appressed, of roughly uniform length; gastral pilosity somewhat stout and mostly forming a sparse layer of suberect to subdecumbent setae; setae on legs mostly decumbent to appressed, with suberect setae on coxae and femoral venters.
(1 measured) HL 0.88, HW 0.81, FLD 0.21, PCW 0.05, SL 0.77, EL 0.25, ACL 0.68, ML 1.38, PrW 0.76, PSL 0.14, SDL 0.12, PL 0.50, PH 0.29, PW 0.21, PPL 0.27, PPH 0.29, PPW 0.35, MFL 0.98, MTL 0.80, CI 92, SI 95, REL 30, FLI 26, PSI 1.2, MFI 83, ACI1 64, ACI2 88.
Same as worker except for standard queen modifications and as follows: mesosoma almost completely carinulate, only mesopleuron mostly smooth; pronotum with transverse carinulae; pronotum with transverse carinulae that wrap around entire surface; mesoscutum and scutellum with longitudinal carinae; anterior of mesoscutum with a short narrow strip of smooth surface from which carinulae arise; gastral setae slightly more dense.
Unknown.
The oddly shaped clypeus and mandible of this species is unique within
Distribution map of
Integument mostly dark red-brown to orange-brown; small-sized species (see HL, ML, PrW below); anterior clypeal margin viewed from anterodorsal angle undulating (straight in full-face view), appearing as 2–4 blunt teeth; basal margin of mandible sinuous, with a shallow, but distinct basal depression; head and mesosoma densely sculptured, mostly rugoreticulae; gastral pilosity short, relatively dense, and usually appearing uniformly suberect to subdecumbent, but sometimes more clearly bilayered; petiole somewhat long and gracile, with node relatively small; petiolar and postpetiolar nodes smooth only on anterior faces; eye relatively small (EL 0.08–0.09, REL 14–16), subcircular to oval-shaped, with 4–5 ommatidia in greatest diameter; frontal lobes moderately expanded (FLD 0.15–0.18, FLI 28–33), mostly to completely covering torular lobes in full-face view; propodeal spines short to medium length (PSL 0.11–0.18, PSI 1.6–2.1).
Guatemala to Honduras.
(9 measured) HL 0.61–0.68 (0.68), HW 0.51–0.59 (0.58), FLD 0.15–0.18 (0.18), PCW 0.01–0.03 (0.03), SL 0.52–0.55 (0.55), EL 0.08–0.09 (0.08), ACL 0.51–0.54 (0.53), ML 0.75–0.82 (0.82), PrW 0.36–0.41 (0.41), PSL 0.11–0.12 (0.12), SDL 0.05–0.07 (0.07), PL 0.28–0.32 (0.32), PH 0.15–0.17 (0.17), PW 0.12–0.13 (0.13) PPL 0.13–0.15 (0.15), PPH 0.13–0.15 (0.15), PPW 0.15–0.16 (0.16), MFL 0.57–0.63 (0.62), MTL 0.47–0.51 (0.51), CI 82–88 (86), SI 92–104 (94), REL 14–16 (14), FLI 28–33 (32), PSI 1.6–2.1 (1.8); MFI 84–95 (93), ACI1 68–71 (68), ACI2 96–102 (96).
Small species; general body color dark red-brown to orange-brown, with patches of dark brown to brown on gaster; appendages lighter, brown to yellow-brown; setae golden brown; mandible with 6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2 middle teeth, which are often worn and indistinct; basal margin of mandible sinuous, with a shallow, but distinct basal depression, accompanied by attenuation of cuticle where mandible fits underneath clypeus; mandible mostly smooth and shining, with scattered piligerous punctae and basal striae; anterior clypeal margin viewed at anterodorsal angle undulating (straight in full-face view), appearing as 2–4 blunt teeth; median lobe of clypeus somewhat flattened, longitudinal carinulae absent or very faint, apex with a short transverse carinula; remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions narrow (PCW 0.01–0.03), sides diverging posteriad; frontal lobes moderately expanded (FLD 0.15–0.18, FLI 28–33), with lateral apices shifted slightly posteriad of torular lobes, which are mostly to completely covered in full-face view; head subrectangular to oval-shaped (CI 82–88), posterior margin slightly depressed medially; eye relatively small (EL 0.08–0.09, REL 14–16), subcircular to oval-shaped, with 4–5 ommatidia in greatest diameter; head mostly rugoreticulate, with a few longitudinal rugae along midline, interstices faintly punctate; scape relatively long (SI 94–104), but variable, either reaching or not quite reaching posterior margin when laid back; scape surface shiny, but somewhat rough, with punctae and faint striae; flagellum with distinct 4-segmented antennal club; mesosoma densely sculptured, except for propodeal declivity, which only has a few faint transverse carinulae; promesonotal dorsum rugose to rugoreticulae, interstices faintly punctate; mesosomal side mostly punctate, with a few rugulae; propodeal dorsum with a few transverse carinae; promesonotum in profile, low-domed, roughly symmetrical; metantoal groove distinct, but somewhat shallow; anterodorsal margin of propodeum in profile with a small welt; propodeal spines present, short to medium length (PSL 0.11–0.18, PSI 1.6–2.1); petiole somewhat long and gracile (PL/HW 0.53–0.59); petiolar node in profile relatively small (PH/PL 0.49–0.59), asymmetrical, with anterior face long and sloping and posterior face short and nearly vertical, dorsum of node rounded, apex pointing posteriad; postpetiole relatively small (PPH/PH 0.85–0.91), somewhat dorsoventrally compressed; anterior faces of petiolar and postpetiolar nodes smooth and shiny, remainder of waist mostly punctate, with a few small rugulae around postpetiolar node; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with short to moderately long standing pilosity; scape with suberect to decumbent setae; gastral pilosity variable, usually short, relatively dense, and uniformly suberect to subdecumbent, but sometimes clearly bilayered, with a longer suberect layer, and a shorter decumbent layer; setae on legs mostly decumbent to appressed, with some suberect setae on coxae and venter of profemur.
(3 measured) HL 0.67–0.69 (0.69), HW 0.59–0.62 (0.61), FLD 0.16–0.21 (0.21), PCW 0.02–0.04 (0.02), SL 0.56–0.58 (0.56), EL 0.16–0.18 (0.17), ACL 0.54–0.57 (054), ML 0.92–1.00 (0.95), PrW 0.50–0.55 (0.53), PSL 0.15–0.16 (0.15), SDL 0.08–0.09 (0.09), PL 0.36–0.38 (0.38), PH 0.18–0.20 (0.19), PW 0.15–0.16 (0.15), PPL 0.15–0.17 (0.15), PPH 0.18–0.19 (0.18), PPW 0.19–0.20 (0.19), MFL 0.63–0.69 (0.63), MTL 0.53–0.58 (0.53), CI 88–90 (88), SI 92–96 (92), REL 27–30 (27), FLI 25–34 (34), PSI 1.6–2.1 (1.6), MFI 88–97 (97), ACI1 67–69 (69), ACI2 96–98 (69).
Same as worker except for standard queen modifications and as follows: mesoscutum mostly longitudinally rugose, but with a central longitudinal carina that is thick at anterior margin; most of katepisternum and part of anepisternum smooth and shiny; gastral pilosity denser, and more distinctly bilayered.
Unknown.
There are some morphological differences unique to the Meambar population (
Molecular phylogenetic data show that
Integument mostly dark brown to brown, rarely black; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, with a distinct basal depression; anterior clypeal margin undulating, forming 2–4 blunt teeth; face completely sculptured, mostly rugoreticulate; mesosoma mostly sculptured, except for pronotum, which is usually lightly carinulate-punctate, with a small smooth patch on dorsum and side, but sometimes pronotum completely punctate or mostly smooth; remainder of mesosoma usually strongly punctate; propodeal declivity in profile usually with distinctive outline, in which propodeal lobe is broadly rounded and makes a smooth, sinuous connection with propodeal spine (degree of sinuosity variable); eye of small to moderate size (EL 0.07–0.11, REL 15–19), oval-shaped, with 4–5 ommatidia at greatest diameter; pilosity on gastral dorsum bilayered, with a layer of longer suberect setae, and a denser layer of decumbent setae; propodeal spines tuberculate, or reduced to sharp angles (PSL 0.07–0.09, PSI 1.1–1.4); geography useful in species determination.
Southern Mexico (Oaxaca, Veracruz).
(20 measured) HL 0.52–0.69 (0.54), HW 0.45–0.60 (0.48), FLD 0.11–0.16 (0.12), PCW 0.02–0.04 (0.02), SL 0.39–0.58 (0.42), EL 0.07–0.11 (0.08), ACL 0.40–0.54 (0.43), ML 0.61–0.87 (0.65), PrW 0.31–0.40 (0.32), PSL 0.07–0.09 (0.08), SDL 0.05–0.07 (0.06), PL 0.22–0.32 (0.23), PH 0.14–0.19 (0.14), PW 0.10–0.16 (0.11), PPL 0.12–0.20 (0.14), PPH 0.12–0.19 (0.13), PPW 0.13–0.19 (0.15), MFL 0.38–0.62 (0.44), MTL 0.31–0.52 (0.35), CI 84–92 (89), SI 85–99 (87), REL 15–19 (16), FLI 24–28 (25), PSI 1.1–1.4 (1.4), MFI 96–117 (109), ACI1 67–72 (70), ACI2 93–105 (102).
Small-sized species; general body color usually dark brown to brown, rarely almost black; appendages brown to yellow-brown, becoming lighter toward extremities; in specimens with dark brown to black body color, flagellum sometimes noticeably bright yellow; setae golden brown; mandible with 6 teeth, with basal tooth well defined; basal margin of mandible sinuous, with a distinct basal depression; mandible surface mostly smooth and shiny, with scattered piligerous punctae, and some striations on base and lateral surface; anterior clypeal margin undulating, forming 2–4 blunt teeth, inner teeth often projecting beyond lateral teeth (type population); median lobe of clypeus with a pair of longitudinal carinulae that diverge anteriorly, apex of lobe with a moderately long transverse carinula, area in between median lobe and anterior margin forming a distinct cavity where mandibles insert, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions of moderate to somewhat narrow width (PCW 0.02–0.04), with sides subparallel to slightly hour-glass shaped; frontal lobes of moderate width (FLD 0.11–0.16, FLI 24–28), not greatly obscuring torular lobes in full-face view; head roughly oval-shaped (CI 84–92), posterior margin with a slight to distinct median depression; eye of small to moderate size (EL 0.07–0.11, REL 15–19), oval-shaped, with 4–5 ommatidia at greatest diameter; face completely sculptured, mostly rugoreticulate, with some longitudinal carinulae extending back from frontal lobes, interstitial areas with light piligerous punctae; one high-elevation population with facial sculpture more polished, becoming smooth toward posterior margin; scape somewhat short (SI 85–99), not reaching posterior margin when laid back; scape surface mostly smooth, with scattered piligerous punctae; flagellum with a distinct 4-segmented antennal club; mesosoma mostly sculptured, except for pronotum, which ranges from completely punctate to almost completely smooth (both extremes rare); pronotum most often lightly punctate on side, and lightly carinulate-punctate on dorsum, with a small smooth patch in middle (type population); remainder of mesosoma mostly punctate, with some longitudinal carinulae, becoming rugoreticulae on mesonotum; propodeal dorsum and declivity sometimes with transverse carinulae; one high-elevation population with mesopleuron rugulose-punctate, and side of propodeum longitudinally carinulate; promesonotum in profile low-domed and roughly symmetrical; metanotal groove well demarcated, of moderate width and depth; propodeum in profile usually with a distinct profile, in which propodeal lobe has a broadly rounded margin that connects smoothly with propodeal spine; connection usually sinuous, but degree of sinuosity variable; propodeal spines usually tuberculate (type population), but sometimes reduced to sharp angles (PSL 0.07–0.09, PSI 1.1–1.4); petiole in profile of moderate length to slightly elongate (PL/HW 0.49–0.55); petiolar node in profile appearing small (PH/PL 0.52–0.64), and asymmetrical, with anterior face longer and more sloping than posterior face, node dorsum broadly rounded (type population) to more angulate, with apex pointing distinctly posteriad; postpetiole in profile subspherical, usually about same size as petiolar node, but sometimes slighly bulging (PPH/PH 0.79–1.01); petiole and postpetiole usually mostly punctate, with only anterior faces smooth and shiny, but sometimes nodes almost completely smooth; most of body dorsum with short standing pilosity; setae on face very short, mostly subdecumbent; pilosity on gastral dorsum bilayered, with a sparse layer of longer suberect setae, and a denser layer of decumbent setae, sometimes lower layer very similar in density to upper layer; setae on scape subdecumbent to appressed; setae on legs mostly decumbent to appressed, with some suberect setae on femoral venters and coxae.
(8 meausred) HL 0.55–0.68 (0.56), HW 0.50–0.62 (0.51), FLD 0.13–0.15 (0.13), PCW 0.03–0.05 (0.03), SL 0.42–0.56 (0.45), EL 0.14–0.17 (0.15), ACL 0.43–0.52 (0.45), ML 0.74–1.00 (0.78), PrW 0.44–0.55 (0.46), PSL 0.09–0.11 (0.11), SDL 0.07–0.09 (0.09), PL 0.28–0.36 (0.29), PH 0.16–0.21 (0.16), PW 0.12–0.17 (0.13), PPL 0.14–0.21 (0.15), PPH 0.16–0.21 (0.16), PPW 0.17–0.22 (0.17), MFL 0.44–0.62 (0.46), MTL 0.38–0.53 (0.39), CI 88–93 (91), SI 83–95 (88), REL 27–30 (29), FLI 25–26 (26), PSI 1.2–1.5 (1.3), MFI 96–115 (111), ACI1 66–69 (68), ACI2 93–106 (102).
Same as worker except for standard queen modifications and as follows (comparsion of type population only): pronotum with transverse carinulae/rugulae; mesoscutum mostly smooth, with piligerous punctae, and some longidutinal carinulae along lateral margin; scutellum smooth in middle, rugulose-punctate on lateral margins; mesopleuron mostly smooth; propodeum carinulate-punctate; lower layer of setae on gastral dorsum very dense, almost pubescent; dorsum of mesosoma and anterior faces of petiolar and postpetiolar nodes with a dense layer of short, decumbent setae; propodeal declivity in profile less sinuous, with propodeal lobe smaller and less evenly rounded; wing venation as in
See
Distinguishing
As currently defined, geography is very useful in separating
Many populations of
Variant 1 (
Variant 2 (
Distribution map of
Integument dark red-brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, usually with a small basal depression; anterior clypeal margin undulating, with two small blunt teeth bordering midline; eye of moderate size (EL 0.07–0.11, REL 15–20), oval-shaped, with 4–5 ommatidia at greatest diameter; face mostly rugoreticulate; mesosoma often mostly sculptured, but pronotum variable, usually rugose with smooth patches on dorsum and side, but sometimes mostly rugose or mostly smooth; propodeal spines tuberculate to short (PSL 0.07–0.10, PSI 1.2–1.8); gastral pilosity usually short, dense and clearly bilayered, with a layer of suberect setae and a denser underlying layer of subdecumbent setae, but sometimes setae more uniformly subdecumbent, or suberect setae thickened; geography useful in species determination.
Southern Mexico (Chiapas, Veracruz?) to Honduras.
(21 measured) HL 0.54–0.65 (0.59), HW 0.45–0.57 (0.51), FLD 0.12–0.16 (0.14), PCW 0.02–0.04 (0.03), SL 0.39–0.51 (0.42), EL 0.07–0.11 (0.08), ACL 0.39–0.48 (0.42), ML 0.62–0.79 (0.70), PrW 0.31–0.40 (0.35), PSL 0.07–0.10 (0.09), SDL 0.05–0.06 (0.05), PL 0.22–0.29 (0.26), PH 0.13–0.18 (0.15), PW 0.11–0.16 (0.14), PPL 0.13–0.18 (0.16), PPH 0.12–0.18 (0.15), PPW 0.13–0.21 (0.16), MFL 0.40–0.54 (0.45), MTL 0.34–0.45 (0.37), CI 84–91 (87), SI 80–89 (81), REL 15–20 (16), FLI 25–29 (26), PSI 1.2–1.8 (1.6), MFI 103–117 (115), ACI1 69–72 (69), ACI2 93–103 (100).
Small-sized species; general body color dark brown to brown or orange-brown, with appendages brown or orange-brown to yellow-brown, becoming lighter toward extremities; setae golden brown; mandible with 6 teeth, inner teeth sometimes worn; basal margin of mandible sinuous, usually with a distinct basal depression, but without tooth; mandible mostly smooth and shining, with scattered piligerous punctae and some basal striae; anterior clypeal margin when viewed from anterodorsal angle undulating, usually forming 2 blunt teeth bordering midline; median lobe of clypeus with a pair of faint longitudinal carinulae that diverge anteriorly, apex of lobe with short transverse carinula; area in between median lobe and anterior clypeal margin forming a shallow concavity where mandibles insert; remaining surface of clypeus mostly smooth; posterior extension of clypeus between antennal insertions somewhat narrow (PCW 0.02–0.04), sides subparallel; frontal lobes of moderate width (FLD 0.12–0.16, FLI 25–29), not greatly obscuring torular lobes in full-face view; head subrectangular to oval-shaped (CI 84–91), posterior margin slightly depressed medially; eye somewhat small (EL 0.07–0.11, REL 15–20), oval-shaped, with 4–5 ommatidia at greatest diameter; head completely sculptured, mostly rugoreticulate, with a few longitudinal carinulae along midline; scape relatively short (SI 80–89), not reaching posterior margin of head when laid back; scape surface mostly smooth, with scattered piligerous punctae; flagellum with distinct 4-segmented antennal club, last segment noticeably bulging; pronotal sculpture variable, dorsum usually longitudinally rugose, with a small smooth patch mesad (type population), side usually rugose on upper half and smooth on lower half (type population), sometimes pronotum completely smooth, or completely rugose, with only a small smooth patch on side; dorsum of mesonotum rugulose punctate; katepisternum and side of propodeum punctate, sometimes with a few rugulae; propodeal dorsum punctate, with a few transverse carinulae; propodeal declivity mostly smooth, with a few transverse carinulae on upper half; promesonotum in profile low-domed, and roughly symmetrical; propodeal spines tuberculate to short (PSL 0.07–0.10, PSI 1.2–1.8); petiole usually somewhat short and stocky (PL/HW 0.47–0.54); petiolar node somewhat small (PH/PL 0.56–0.57), roughly symmetrical, dorsum reaching a defined apex, which points nearly vertical; postpetiole in profile variable, usually small and similar in size to petiolar node (type population), but sometimes bulging (PPH/PH 0.91–1.07); petiole and postpetiole usually mostly punctate, with anterior faces of nodes smooth, and posterior faces of nodes with a few rugulae; most of body dorsum with short standing pilosity; pilosity on gastral dorsum usually distinctly bilayered, with a layer of suberect setae, and a slightly more dense layer of decumbent setae (type population), but sometimes setae more uniformly suberect to subdecumbent and less clearly bilayered; suberect layer of setae sometimes slightly thickened; setae on scapes decumbent to appressed; setae on legs mostly subdecumbent to appressed, with longer suberect setae on femoral venters and coxae.
(7 measured) HL 0.59–0.67 (0.59), HW 0.52–0.61 (0.52), FLD 0.15–0.17 (0.15), PCW 0.04–0.05 (0.04), SL 0.42–0.50 (0.44), EL 0.14–0.17 (0.14), ACL 0.42–0.48 (0.44), ML 0.80–0.95 (0.80), PrW 0.46–0.55 (0.46), PSL 0.10–0.13 (0.11), SDL 0.07–0.08 (0.07), PL 0.28–0.34 (0.30), PH 0.17–0.20 (0.17), PW 0.15–0.17 (0.15), PPL 0.16–0.20 (0.18), PPH 0.17–0.22 (0.17), PPW 0.19–0.24 (0.19), MFL 0.47–0.57 (0.47), MTL 0.39–0.48 (0.41), CI 88–92 (88), SI 79–86 (84) REL 26–28 (28), FLI 26–30 (28), PSI 1.3–1.7 (1.6), MFI 102–113 (110), ACI1 68–70 (68), ACI2 90–100 (100).
Same as worker except for standard queen modifications and as follows: pronotum with transverse carinulae on humeri, becoming smooth mesad; mesoscutum mostly with longitudinal rugulae and foveolae, midline and mesolateral margin smooth; scutellum smooth along midline, and longitudinally carinulate laterad; propodeum with transverse carinulae/rugulae that wrap around surface; mesopleuron mostly smooth; lower layer of setae on gastral dorsum very dense, almost pubescent; wing venation as in
Unknown.
Within the range of
In addition to within site variation, there is considerable among population variation, with almost every population displaying some unique feature. Out of this diversity I describe a couple of variants. Variant 1 (
Distribution map of
Integument mostly red-black; small- to medium-sized species (see HL, ML, and PrW below); anterior clypeal margin in full-face view, with three projecting teeth, two well-defined sharp outer teeth, and a blunt central tooth, formed by a projecting median clypeal lobe; anterior clypeal margin underneath median clypeal lobe with two blunt teeth; basal margin of mandible sinuous, with a distinct basal depression, but no basal tooth; frontal lobes markedly expanded (FLD 0.19–0.22, FLI 24–26), completely obscuring the torular lobes in full-face view; eye of moderate size (EL 0.09–0.11, REL 14–17), with 5–6 ommatidia at greatest diameter; propodeal spines present and of moderate length (PSL 0.16–0.19, PSI 1.8–2.2); gastral setae short, and clearly bilayered, with a layer of suberect setae, and a layer of slightly shorter but denser subdecumbent setae;.
Honduras.
(5 measured) HL 0.74–0.80 (0.78), HW 0.63–0.69 (0.66), FLD 0.24–0.26 (0.26), PCW 0.02–0.03 (0.03), SL 0.54–0.59 (0.59), EL 0.09–0.11 (0.10), ACL 0.54–0.57 (0.57), ML 0.88–0.97 (0.94), PrW 0.45–0.49 (0.48), PSL 0.16–0.19 (0.18), SDL 0.08–0.10 (0.09), PL 0.34–0.37 (0.37), PH 0.18–0.20 (0.20), PW 0.14–0.16 (0.16), PPL 0.18–0.20 (0.20), PPH 0.16–0.19 (0.19), PPW 0.19–0.20 (0.20), MFL 0.60–0.64 (0.63), MTL 0.50–0.52 (0.51), CI 83–87 (86), SI 84–89 (89), REL 14–17 (15), FLI 36–39 (39), PSI 1.8–2.2 (2.1), MFI 104–111 (106), ACI1 66–68 (67), ACI2 96–100 (96).
Small- to medium-sized species; general body color red black, with patches of brown on gaster; mandibles and appendages lighter, mostly brown to orange-brown; setae pale golden brown; mandible usually with 6 teeth, sometimes with 1–2 small denticles, basal tooth well-defined; basal margin of mandible sinuous, with a distinct basal depression, but no basal tooth; mandible mostly smooth and shiny, with scattered piligerous punctae and basal striae; anterior clypeal margin in full-face view, with three projecting teeth, two well-defined outer teeth, and a more blunt central tooth, formed from a projecting median clypeal lobe; anterior clypeal margin underneath median clypeal lobe with two blunt teeth (only visible with mandibles open and from an anterodorsal view); median clypeal lobe surface usually with a single median longitudinal carinula and a variable number of irregular foveolae, remainder of clypeus smooth and shiny; posterior extension of clypeus between antennal insertions hourglass-shaped, with middle of hourglass narrow (PCW 0.02–0.03); frontal lobes noticeably expanded (FLD 0.19–0.22, FLI 24–26), completely obscuring the torular lobes in full-face view; head roughly oval-shaped (CI 83–87), posterior margin with a slight median depression; eye of moderate size (EL 0.09–0.11, REL 14–17), oval-shaped, with 5–6 ommatidia at greatest diameter; head strongly rugoreticulate (not quite foveate), with a few short costae extending back from the frontal lobes and posterior clypeal extension; scape somewhat short (SI 84–89), not reaching posterior margin of head when laid back; scape surface shiny, but somewhat rough, with scattered piligerous punctae, punctulae, and fine striae; flagellum with a somewhat distinct 4-segmented antennal club; mesosoma mostly strongly rugose to rugoreticulate, with finer reticulae, becoming punctae, on pronotal side and anepisternum, rugae near anterior margin of pronotal dorsum usually with a transverse orientation; propodeal dorsum and declivity with transverse carinulae; promesonotum in profile low-domed and roughly symmetrical, but anterior declivity somewhat shorter and steeper; metantoal grove well-demarcated, of moderate width and depth; anterodorsal margin of propodeum with a small welt; propodeal spines present and of moderate length (PSL 0.16–0.19, PSI 1.8–2.2); petiole appearing of moderate length (PL/HW 0.52–0.55); petiolar node relatively small (PH/PL 0.53–0.56), domed, but slightly asymmetrical in profile, with the anterior face longer and more sloping than posterior face; postpetiole subspherical, without a prominent dorsal node; anterior faces of petiolar and postpetiolar nodes smooth and shiny, anterior and lateral sides with a variable number of rugulae and faint punctae; ventral surfaces of waist segments punctate; most of body dorsum with short standing pilosity; setae on scape mostly subdecumbent; gastral setae short, and clearly bilayered, with a layer of suberect setae, and a layer of slightly shorter but denser subdecumbent setae; setae on legs mostly appressed with some suberect setae on coxae and femoral venters.
(1 measured) HL 0.78, HW 0.71, FLD 0.26, PCW 0.04, SL 0.59, EL 0.17, ACL 0.57, ML 1.10, PrW 0.63, PSL 0.21, SDL 0.12, PL 0.41, PH 0.22, PW 0.18, PPL 0.22, PPH 0.21, PPW 0.18, MFL 0.66, MTL 0.55, CI 90, SI 84, REL 24, FLI 37, PSI 1.8, MFI 107, ACI1 66, ACI2 97.
Same as worker except for standard queen modifications and as follows: pronotum with transverse rugae/rugoreticulae; mesoscutum more rugose than rugoreticulate, with rugae longitudinal in orientation; scutellum with a central patch of smooth cuticle surrounded by longitudinal rugae; mesopleuron partly smooth and shiny.
Unknown.
This species has been collected from only a few samples of sifted leaf litter from the forest floor. It inhabits montane wet forest from approximately 1200–1400 m elevation. The shape of the anterior margin of the clypeus is unique among
The structure of the clypeus is unique among known
Although support values are low, molecular phylogenetic results indicate that
See type material.
Integument mostly black and shining; medium-sized species (see HL, ML, PrW below); head mostly smooth and shiny; mesosoma reticulately costate to coarsely rugoreticulate; propodeal spines long and robust (PSL 0.28–0.34, PSI 3.0-3.7); frontal lobes dorsolaterally expanded, obscuring the torular lobes in full-face view (FLD 0.25-0.29, FLI 35-38); eye of moderate to large size (EL 0.11–0.15, REL 16–20), oval-shaped, with 7–9 ommatidia at greatest diameter; anterior margin of clypeus with shallow median emargination; basal margin of mandible straight, without a notch or substantial depression; pilosity on gastral dorsum long, flexuous, and relatively sparse.
Southern Mexico to Nicaragua.
(11 measured) HL 0.74–0.81 (0.76), HW 0.69–0.78 (0.72), FLD 0.25–0.29 (0.27), PCW 0.06–0.09 (0.08), SL 0.58–0.66 (0.62), EL 0.11–0.15 (0.13), ACL 0.52–0.56 (0.54), ML 0.95–1.08 (0.99), PrW 0.55–0.61 (0.56), PSL 0.28–0.34 (0.29), SDL 0.08–0.11 (0.08), PL 0.44–0.50 (0.44), PH 0.22–0.25 (0.23), PW 0.18–0.24 (0.20), PPL 0.18–0.21 (0.19), PPH 0.19–0.22 (0.20), PPW 0.18–0.23 (0.21), MFL 0.71–0.78 (0.72), MTL 0.56–0.63 (0.59), CI 93–97 (95), SI 84–88 (85), REL 16–20 (18), FLI 35–38 (37), PSI 3.0–3.7 (3.5), MFI 96–100 (100), ACI1 65–67 (65), ACI2 85–90 (89).
Medium-sized species; general body color black, with mandibles, clypeus and appendages dark brown to yellow-brown; setae golden; mandible with 6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2–3 inner teeth, which are often worn and indistinct; basal margin of mandible straight, without a distinct notch or depression; dorsal surface of mandible mostly smooth and shining, with scattered piligerous punctae, and several weak striae extending from base; anterior clypeal margin with shallow median emargination; median lobe of clypeus with a pair of faint carinulae that diverge toward anterior margin, apex of lobe with a faint transverse carinula, remainder of clypeus mostly smooth and shining; posterior extension of clypeus between frontal lobes broad, with subparallel sides (PCW 0.06–0.09); frontal lobes expanded dorsolaterally (FLD 0.25-0.29, FLI 35-38), with torular lobes obscured in full-face view; head roughly oval-shaped, slightly longer than broad (CI 93-97), with posterior margin flat or gently curving, never depressed medially; eyes of moderate size (EL 0.11–0.15, REL 16-20), oval-shaped, with 7–9 ommatidia at greatest diameter; face largely smooth and shining, with faint carinulae and punctae on gena, scattered piligerous punctae elsewhere; scape short, not surpassing posterior margin of head when laid back (SI 84-88); dorsal surface of scape striate; funiculus with distinct 4-segmented antennal club; mesosoma compact, shiny, and almost entirely reticulately costate (specimens vary considerably in sharpness, coarseness, and orientation of costae); propodeal declivity smooth; promesonotum in profile dome-shaped, roughly symmetrical; propodeal spines long and robust (PSL 0.28–0.34, PSI 3.0-3.7), usually projecting dorsoposteriorly; petiole relatively long and wedge-shaped (PL/HW 0.60-0.65), node variable, appearing rather robust to slightly more gracile, always angled so that the apex points posteriad; anterior slope of petiole usually long and rising gradually from peduncle, but sometimes shorter and rising more abruptly; posterior slope of petiole short and nearly vertical; dorsum of petiolar node viewed from posterior side flat, to depressed medially, to slightly convex; dorsal portion of petiolar node distinctly wider than ventral portion; postpetiolar node in profile smaller than petiolar node (PPH/PH 0.85–0.94), dome-shaped, slightly asymmetrical, with anterior slope longer and more sloping than posterior slope; petiole and postpetiole generally smooth and shiny, nodes with several deep furrows, ventral surfaces faintly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae, and furrows on anterior constriction where gaster inserts into postpetiole; most of body with a relatively sparse layer of long, flexuous setae; setae on scapes and legs varying from mostly suberect to mostly decumbent; setae on femoral venters and coxae always longer and suberect to subdecumbent.
(5 measured) HL 0.73–0.80 (0.80), HW 0.69–0.78 (0.78), FLD 0.25–0.30 (0.30), PCW 0.07–0.10 (0.10), SL 0.58–0.65 (0.65), EL 0.17–019 (0.18), ACL 0.51–0.59 (0.59), ML 1.05–1.17 (1.17), PrW 0.63–0.71 (0.69), PSL 0.30–0.33 (0.32), SDL 0.09–0.11 (0.09), PL 0.46–0.53 (0.51), PH 0.24–0.26 (0.26), PW 0.21–0.23 (0.23), PPL 0.21–0.25 (0.22), PPH 0.23–0.25 (0.25), PPW 0.23–0.25 (0.25), MFL 0.70–0.79 (0.79), MTL 0.56–0.64 (0.64), CI 94–98 (97), SI 82–86 (84), REL 23–27 (24), FLI 26–38 (28), PSI 2.9–3.4 (3.4), MFI 98–99 (99), ACI1 65–66 (65), ACI2 86–90 (90).
Same as worker except for standard queen modifications and the following: costae on mesoscutum with a decidedly longitudinal orientation, but often wavy, and usually with some reticulation anteriorly; costae on side of propodeum longitudinal in orientation; mesopleuron mostly smooth and shiny; wing venation as in
See
Distribution map of
Integument mostly orange-brown to brown; small-sized species (see HL, ML, PrW below); anterior clypeal margin often with a deep median excision, but sometimes reduced to a shallow median emargination; basal margin of mandible straight; face densely sculptured and mostly rugoreticulate; mesosoma densely sculptured with punctae, rugae, and/or rugoreticulae; eye small (EL 0.04–0.10, REL 8–14), subcircular, with 2–4 ommatidia at greatest diameter; median portion of clypeus angled dorsoventrally, causing it to be mostly hidden in full-face view; posterior extension of clypeus between antennal insertions very narrow (PCW 0.01–0.03), with border of antennal insertions sometimes touching anteriorly; propodeal spines present, short to long (PSL 0.09–0.20, PSI 1.8–5.4); pilosity on gastral tergites distinctly bilayered, with a layer of sparse suberect setae (varying in thickness and length), and a layer of decumbent to appressed setae.
Mexico (Atlantic slope) to Honduras.
(25 measured) HL 0.49–0.73 (0.71), HW 0.43–0.67 (0.63), FLD 0.09–0.14 (0.14), PCW 0.01–0.03 (0.02), SL 0.38–0.61 (0.59), EL 0.04–0.10 (0.06), ACL 0.41–0.62 (0.62), ML 0.57–0.88 (0.88), PrW 0.29–0.45 (0.43), PSL 0.09–0.20 (0.20), SDL 0.04–0.08 (0.08), PL 0.22–0.34 (0.34), PH 0.13–0.19 (0.18), PW 0.10–0.16 (0.14), PPL 0.11–0.20 (0.19), PPH 0.11–0.16 (0.15), PPW 0.14–0.22 (0.19), MFL 0.37–0.68 (0.66), MTL 0.31–0.56 (0.55), CI 86–94, SI 79–97 (94), REL 8–14 (10), FLI 19–24 (22), PSI 1.8–5.4 (2.5), MFI 95–119 (101), ACI1 64–72 (66), ACI2 98–111 (105).
Small-sized species; general body color mostly orange-brown to brown, with appendages orange-brown to yellow-brown, becoming lighter toward extremities; setae golden; mandible with 5–7 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 1–3 smaller inner teeth/denticles, which are often worn and indistinct; basal margin of mandible straight, without a basal notch or depression; mandible mostly smooth and shiny, with scattered piligerous punctae and some basal striae; anterior clypeal margin often with a deep median excision (type population), but sometimes excision reduced to a shallow emargination; median lobe of clypeus flattened, and angled dorsoventrally, causing it to be mostly invisible in full-face view; surface of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions very narrow (PCW 0.01–0.03), sides subparallel, with border of antennal insertions sometimes touching anteriorly; frontal lobes of moderate width (FLD 0.09–0.14, FLI 19–24), not greatly obscuring torular lobes in full-face view; head appearing subrectangular to oval-shaped (CI 86–94), posterior margin flat to slightly depressed medially; eye small (EL 0.04–0.10, REL 8–14), subcircular, with 2–4 ommatidia at greatest diameter; face densely sculptured, usually mostly rugoreticulate and punctate, with some longitudinal rugae along midline (type population), but sometimes mostly rugose; scape of short to moderate length (SI 79–97), usually not reaching posterior margin of head; scape surface mostly smooth, with scattered piligerous punctae; flagellum with a distinct to very distinct 4-segmented antennal club; mesosoma densely sculptured, dorsum of promesonotum ranging from rugoreticulate (type population), to rugose-punctate, to rugose, to mostly punctate, with rugae longitudinal in orientation; side of pronotum usually punctate, with a few rugulae or rugoreticulae (type population), but sometimes mostly rugose, and with a small patch of smooth cuticle; mesopleuron and side of propodeum mostly punctate, with a variable number of rugulae; dorsum and declivity of propodeum with a few transverse carinulae; promesonotum in profile low-domed and roughly symmetrical; metanotal groove usually well-demarcated and somewhat deep; anterodorsal margin of propodeum often raised into a small to large welt, but sometimes average; propodeal spines present, short to long (PSL 0.09–0.20, PSI 1.8–5.4); petiole appearing moderate to slightly elongate (PL/HW 0.45–0.56), usually with a distinct node, but sometimes node less distinct, making petiole look more wedge-shaped; when distinct, petiolar node in profile, average to slightly enlarged (PH/PL 0.48–0.64), and roughly symmetrical, dorsum of node usually gently rounded and pointing vertically (type population), but sometimes broadly rounded, or nearly angulate and pointing slightly posteriad; postpetiole in profile usually subspherical and appearing similar in size to petiolar node (PPH/PH 0.79–0.96), postpetiole in dorsal view often distinctly wider than petiole, sometimes much wider (PPW/PW 0.55–0.83); petiole and postpetiole usually mostly punctate, with only the anterior faces of nodes smooth, but sometimes nodes completely smooth, or completely punctate; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with standing pilosity; pilosity on gastral tergites distinctly bilayered, with a layer of sparse suberect setae (varying in thickness and length), and a layer of decumbent to appressed setae; setae on scapes and legs mostly decumbent to appressed, with some longer suberect setae on femoral venters and coxae.
(5 measured) HL 0.55–0.73 (0.73), HW 0.50–0.69 (0.69), FLD 0.11–0.16 (0.16), PCW 0.01–0.02 (0.02), SL 0.42–0.63 (0.63), EL 0.13–0.16 (0.16), ACL 0.43–0.62 (0.62), ML 0.73–1.01 (1.01), PrW 0.40–0.57 (0.57), PSL 0.12–0.22 (0.22), SDL 0.07–0.10 (0.10), PL 0.27–0.38 (0.38), PH 0.15–0.20 (0.20), PW 0.13–0.17 (0.17), PPL 0.14–0.20 (0.20), PPH 0.15–0.19 (0.19), PPW 0.17–0.23 (0.23), MFL 0.43–0.69 (0.69), MTL 0.37–0.58 (0.58), CI 89–93 (93), SI 81–92 (92), REL 22–27 (23), FLI 22–24 (24), PSI 1.6–2.2 (2.2), MFI 99–118 (99), ACI1 63–74 (63), ACI2 97–107 (98).
Same as worker except for standard queen modifications and as follows (comparison with worker and queen from type population only): pronotum rugoreticulate laterad, and punctate mesad; mesoscutum and scutellum rugoreticulate to foveate; propodeum with transverse carinulae that wrap around surface; katepisternum mostly smooth; petiole more elongate.
Unknown.
A rather uncommon species known almost exclusively from Winkler and Berlese samples of sifted leaf litter, except for one collection from under a stone.
Although quite variable across its range (see below),
Stenamma excisum forms a difficult species complex composed of several allopatric variants, which probably constitute distinct species. The type form is known only from two sites in Honduras, where it is unusually abundant in leaf litter samples. The most important features of this form are the rugoreticulate sculpture on the pronotal dorsum and the nearly triangular excision in the anterior clypeal margin. Other characteristics specific to this form are indicated in the species description above (see parenthetical comments). Specimens from the populations in Honduras and the next closest population are quite divergent from one another. Furthermore, the type form seems to be confined to lower elevations, whereas the other variants usually occur at mid elevations. For these reasons, I highly suspect that there are multiple species within
Variant 1 (
Variant 2 (
Variant 3 (
Variant 4 (
Variant 5 (
Variant 6 (
Variant 7 (
Distribution map of
Integument mostly black to dark red-brown; medium-sized species (see HL, ML, PrW below); petiole and postpetiole almost completely smooth and shiny, with only a few faint vestigial punctae sometimes present; postpetiole in profile bulging, globular, appearing more voluminous than petiolar node; face with a fan of carinae/rugae extending from frontal lobes to approximately ¾ distance to posterior margin of head, remainder of head smooth and shiny; dorsum of promesonotum with distinctive transverse furrows that reticulate toward posterior margin; propodeal spines absent to tuberculate (PSL 0.13–0.14, PSI 1.0–1.1), dorsal and declivitous faces of propodeum in profile flat, forming a blunt 90° angle; eye relatively large (EL 0.13–0.15, REL 19), with 8 ommatidia at greatest diameter; anterior clypeal margin with median emargination; basal margin of mandible straight; gastral setae long, sparse, and uniformly suberect.
Costa Rica.
(3 measured) HL 0.77–0.86 (0.77), HW 0.71–0.81 (0.71), FLD 0.21–0.24 (0.22), PCW 0.07–0.08 (0.07), SL 0.62–0.69 (0.62), EL 0.13–0.15 (0.13), ACL 0.60–0.65 (0.60), ML 1.07–1.20 (1.07), PrW 0.54–0.61 (0.54), PSL 0.13–0.14 (0.13), SDL 0.12–0.13 (0.12), PL 0.40–0.42 (0.40), PH 0.23–0.25 (0.23), PW 0.16–0.17 (0.16), PPL 0.24–0.27 (0.27), PPH 0.21–0.22 (0.21), PPW 0.21–0.22 (0.21), MFL 0.75–0.87 (0.75), MTL 0.62–0.70 (0.62), CI 92–94 (92), SI 85–88 (88), REL 19 (19), FLI 29–31 (31), PSI 1.0–1.1 (1.1), MFI 70–73 (70), ACI1 63–64 (64), ACI2 95–98 (97).
Medium-sized species; general body color black to dark red-brown, with brown patches on gaster; appendages mostly brown, changing to yellow-brown at extremities; setae golden brown; mandible with 5–7 teeth, consisting of 4 distinct apical teeth, a distinct basal tooth, and 1–2 worn denticles in between; basal margin of mandible straight, without a basal notch or depression; mandible surface with scattered piligerous punctae and faint striae; anterior clypeal margin with a median emargination; median lobe of clypeus obliquely flattened, mostly smooth and shiny, with a short transverse carinula near anterior margin, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions somewhat wide (PCW 0.07–0.08), with sides subparallel; frontal lobes somewhat narrow (FLD 0.0.21–0.24, FLI 29–31), not obscuring torular lobes in full-face view, outer margin of frontal lobes nearly parallel, almost without a discernable lateral apex; head subrectangular to oval-shaped (CI 92–94), with posterior margin flat, not depressed medially; eye relatively large (EL 0.13–0.15 REL 19), oval-shaped, with 8 ommatidia at greatest diameter; face with a fan of coarse carinae or costae extending from the frontal lobes to no more than ¾ distance to posterior margin of head; gena with short rugae and rugoreticulae; remainder of head mostly smooth and shiny; scape of moderate length (SI 85–88), not quite reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae; flagellum with distinct 4-segmented antennal club; mesosoma densely sculptured, except for a patch of smooth, shiny cuticle on side of pronotum and katepisternum; dorsum of promesonotum with distinctive transverse furrows, which merge together posteriad and appear reticulate; sculpture on propodeum costate to rugose, with orientation transverse on dorsum, and longitudinal on side; mesopleuron rugose; propodeal declivity with faint transverse carinulae; promesonotum in profile low-domed and roughly symmetrical; metanotal grove somewhat indistinct and wide; propodeal spines absent to tuberculate (PSL 0.13–0.14, PSI 1.0–1.1), appearing in profile as a blunt 90° angle that separates dorsal and declivitous faces of propodeum; petiole of moderate length (PL/HW 0.52–0.56); petiolar node in profile robust and somewhat bulging (PH/PL 0.56–0.59), asymmetrical, with a longer more sloping anterior face and a short almost vertical posterior face; node dorsum flat to gently rounded and pointing posteriad; postpetiole in profile large, bulging, appearing slightly more voluminous than petiolar node, outline asymmetrical, with the anterior face longer and more gently sloping, and the posterior face short and vertical, dorsum of node broadly rounded; petiole and postpetiole mostly smooth and shining; gaster mostly smooth and shiny, with scattered piligerous punctae, and very short furrows around anterior constriction; most of body dorsum with long standing pilosity; scape setae suberect to subdecumbent; gastral setae long, somewhat sparse, and uniformly suberect; setae on legs suberect to subdecumbent, with longer suberect setae on coxae and femoral venters.
Unknown.
Unknown.
I am not completely convinced that
See type material.
Integument mostly black, with legs dark brown, changing to orange-brown only at extremities and joints; medium-sized species (see HL, ML, PrW below); petiole and postpetiole almost completely smooth and shiny, with only a few faint vestigial punctae sometimes present; postpetiole in profile bulging, globular, usually appearing more voluminous than petiolar node; face almost completely smooth and shiny, except for variable number of faint carinulae and punctae; promesonotum almost completely smooth and shiny; eye relatively large (EL 0.13–0.16, REL 18–21), oval-shaped, with 7–9 ommatidia at greatest diameter; anterior margin of clypeus with median emargination; basal margin of mandible straight; propodeal spines absent (PSL 0.12–0.14, PSI 0.9).
Nicaragua to Costa Rica.
(10 measured) HL 0.75–0.93, HW 0.66–0.83 (0.82), FLD 0.20–0.23 (0.23), PCW 0.05–0.08 (0.06), SL 0.63–0.78 (0.76), EL 0.13–0.16 (0.16), ACL 0.63–0.74 (0.73), ML 1.02–1.26 (1.26), PrW 0.51–0.62 (0.61), PSL 0.12–0.14 (0.13), SDL 013–0.15 (0.14), PL 0.39–0.47 (0.47), PH 0.21–0.27 (0.26), PW 0.15–0.19 (0.18), PPL 0.24–0.30 (0.27), PPH 0.20–0.25 (0.24), PPW 0.19–0.24 (0.23), MFL 0.74–0.95 (0.91), MTL 0.58–0.75 (0.73), CI 87–92 (92), SI 91–96 (92), REL 18–21 (19), FLI 27–30 (28), PSI 0.9 (0.9), MFI 85–91 (90), ACI1 60–65 (64), ACI2 92–101.
Medium-sized species; general body color black to dark red-brown, with patches of dark brown on gaster; appendages mostly dark brown with joints and extremities a lighter orange-brown; setae dark golden brown; mandible usually with 6–7 teeth, consisting of 4 distinct apical teeth, a distinct basal tooth, and 1–2 inner teeth/denticles, which are usually worn and indistinct; basal margin of mandible straight, without a basal notch or depression; mandible surface mostly smooth and shiny, with scattered piligerous punctae and a few faint striae; anterior clypeal margin with median emargination; median lobe of clypeus obliquely flattened, mostly smooth and shiny, except for transverse carinula near anterior margin, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions somewhat wide (PCW 0.05–0.08), with sides subparallel to slightly diverging posteriad; frontal lobes of moderate width (FLD 0.20–0.23, FLI 27–30), not covering torular lobes in full-face view; head roughly oval-shaped (CI 87–92), with posterior margin flat, not depressed medially; eye relatively large (EL 0.13–0.16, REL 18–21), oval-shaped, with 7–9 ommatidia at greatest diameter; face completely smooth and shiny, except for variable number of very faint carinulae extending from frontal lobes to about midpoint of head, a few carinulae on gena, and scattered piligerous punctae; scape of moderate length (SI 91–96), usually reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae; funiculus with distinct 4-segmented antennal club; mesosoma almost completely smooth and shiny, except for transverse carinulae on propodeal dorsum, carinae along metanotal grove, and a few scattered rugulae and piligerous punctae; metanotal grove somewhat shallow and wide; propodeal dorsum in profile flat to distinctly convex; propodeal spines absent, at most forming nearly imperceptible nubs (PSL 0.12–0.14, PSI 0.9); petiole of moderate length (PL/HW 0.53–60); petiolar node in profile robust (PH/PL 0.54–0.60), wedge-shaped, with the anterior face long and sloping and the posterior face shorter and nearly vertical; node dorsum rounded, pointing vertically or slightly posteriad; petiolar node in profile robust, more globular than petiolar node, but similarly asymmetrical with long anterior face and short vertical posterior face; petiole and postpetiole mostly smooth and shining, with a few vestigial punctae; gaster smooth and shiny, with only scattered piligerous punctae; most of body dorsum with long standing pilosity; scape setae suberect to subdecumbent; setae on gastral tergites long, sparse, and uniformly suberect; setae on legs suberect to subdecumbent, with longer suberect setae on coxae and femoral venters.
(5 measured) HL 0.87–0.96 (0.90), HW 0.78–0.87 (0.81), FLD 0.24–0.27 (0.25), PCW 0.07–0.08 (0.07), SL 0.72–0.79 (0.76), EL 0.21–0.23 (0.23), ACL 0.70–0.77 (0.71), ML 1.23–1.43 (1.32), PrW 0.67–0.78 (0.74), PSL 0.11–0.15 (0.15), SDL 0.13–0.15 (0.15), PL 0.48–0.53 (0.48), PH 0.27–0.31 (0.29), PW 0.19–0.23 (0.21), PPL 0.30–0.33 (0.30), PPH 0.26–0.30 (0.29), PPW 0.26–0.30 (0.27), MFL 0.87–1.00 (0.93), MTL 0.67–0.79 (0.74), CI 90–92 (90), SI 91–93 (93), REL 26–28 (28), FLI 30–31 (30), PSI 0.9–1.0 (1.0), MFI 86–90 (87), ACI1 60–61 (60), ACI2 94–103 (94).
Same as worker except for standard queen modifications and the following: facial sculpture more developed, with distinct carinulae extending from frontal lobes to ocelli, and some carinulae on gena; pronotum with transverse striae near posterior margin; posterior third of mesoscutum with variable amount of striae/costae, orientation variable, most often longitudinal, but sometimes transverse, or obliquely angled mesad toward posterior margin; scutellum with variable number of longitudinal costae; propodeum with more distinct transverse carinae that extend across the dorsum to upper half of side; wing venation as in
See
Distribution map of
Integument mostly black to dark brown; medium- to large-sized species (see HL, ML, PrW below); anterior clypeal margin with a median emargination; basal margin of mandible straight to slightly sinuous, without a basal notch or depression; head and mesosoma usually densely sculptured, with sharp carinae, rugae, or rugoreticulae; eye relatively large (EL 0.16–0.20, REL 18–22), oval-shaped, with 8–11 ommatidia at greatest diameter; propodeal spines absent, propodeum forming shallow, blunt angles where propodeal dorsum and declivity converge (PSL 0.07–0.11, PSI 0.8–1.1); setae on gastral tergites mostly sparse, long, and suberect, only sometimes with a few short decumbent setae; frontal lobes of moderate width (FLD 0.22–0.26, FLI 25–29), not completely obscuring torular lobes in full face view; metafemur relatively long (MFI 75–88). Similar species:
Mexico (Atlantic slope) to Ecuador.
(10 measured) HL 0.90–1.19 (1.00), HW 0.81–1.04 (0.87), FLD 0.22–0.26 (0.22), PCW 0.04–0.09 (0.05), SL 0.69–0.95 (0.81), EL 0.16–0.20 (0.17), ACL 0.70–0.92 (0.79), ML 1.25–1.62 (1.37), PrW 0.58–0.74 (0.62), PSL 0.07–0.11 (0.09), SDL 0.07–0.13 (0.08), PL 0.41–0.52 (0.45), PH 0.25–0.32 (0.29), PW 0.16–0.23 (0.23), PPL 0.24–0.33 (0.28), PPH 0.21–0.27 (0.23), PPW 0.18–0.27 (0.18), MFL 0.93–1.28 (1.08), MTL 0.71–0.98 (0.81), CI 83–90 (87), SI 84–101 (93), REL 18–22 (20), FLI 25–29 (26), PSI 0.8–1.1 (1.1), MFI 75–88 (81), ACI1 57–62 (59), ACI2 93–102 (97).
Medium- to large-sized species; general body color mostly black to dark brown, with appendages lighter, brown to orange-brown toward extremities; setae dark gold-brown; mandible with 5–6 teeth, consisting of 4 distinct apical teeth, a basal tooth, and 1 tooth in between, which is smaller and often effaced; basal margin of mandible straight to slightly sinuous, without a basal notch or depression; mandible mostly smooth, except for scattered piligerous punctae, and some lateral striations; anterior clypeal margin with a shallow median emargination; median lobe of clypeus with at least a pair of distinct longitudinal carinulae that diverge toward anterior margin, sometimes with a few additional faint carinulae, apex of lobe smooth, or with some faint transverse carinulae; remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between frontal lobes of moderate width (PCW 0.04–0.09), sides subparallel to slightly hour-glass-shaped; frontal lobes of moderate width (FLD 0.22–0.26, FLI 25–29), not greatly obscuring torular lobes in full face view; head usually roughly oval-shaped, but some populations with posterior margin distinctly broader than anterior margin, making head appear more triangular (CI 83–90), posterior margin always depressed medially; eye relatively large (EL 0.16–0.20, REL 18–22), oval-shaped, with 8–11 ommatidia at greatest diameter; face densely sculptured, but sculpture type variable, most often with some longitudinal rugae/carinae along midline, transitioning to rugoreticulae toward lateral margins, but sometimes face almost completely rugoreticulate, or completely carinate, depth and sharpness of sculpture variable; scape of moderate length (SI 84–101), just reaching posterior margin of head when laid back; scape surface mostly smooth, but with distinct carinulae, and scattered piligerous punctures; scape sometimes appearing thickened and more robust; flagellum with distinct 4-segmented antennal club; mesosoma mostly densely sculptured, but sculpture type variable; dorsum of pronotum usually rugose (longitudinal orientation) to rugoreticulate, but sometimes carinate, anterior declivity of pronotum with transverse carinulae; dorsum of mesonotum rugoreticulate to transversely carinate, sometimes intermediate; side of pronotum rugulose to carinate; katepisternum mostly smooth, with some rugulae on upper half; dorsum and declivity of propodeum transversely carinate/carinulate; side of propodeum rugose; promesonotum in profile domed (higher than average), symmetrical to slightly asymmetrical, with location of apex variable; metanotal groove distinct, but narrow; dorsum of propodeum in profile distinctly longer than declivity; propodeal spines absent, propodeum forming shallow, blunt angles where dorsum and declivity converge (PSL 0.07–0.11, PSI 0.8–1.1); petiole of moderate length (PL/HW 0.48–0.55); petiolar node of moderate height (PH/PL 0.60–0.64), subconical in shape, usually pointing vertically to only slightly posteriad, dorsum narrowly to somewhat broadly rounded, posterior margin of petiole, where postpetiole inserts, distinctly bent downwards; postpetiole in profile nearly symmetrical, with anterior face slightly longer than posterior face, postpetiole similar in size to petiolar node (PPH/PH 0.78–0.85); petiole and postpetiole usually mostly punctate, with only anterior faces of nodes smooth, but sometimes nodes mostly smooth, with punctae faint; gaster smooth, with scattered piligerous punctae; most of body dorsum with standing pilosity; setae on gastral tergites mostly sparse, long, and suberect, only sometimes with a few short decumbent setae; setae on scapes subdecumbent to appressed; setae on legs decumbent to appressed, with a few suberect setae on femoral venters and coxae.
(6 measured) HL 1.00–1.17 (1.07), HW 0.85–0.98 (0.89), FLD 0.25–0.27 (0.25), PCW 0.05–0.08 (0.07), SL 0.76–0.92 (0.84), EL 0.23–0.30 (0.29), ACL 0.69–0.91 (0.83), ML 1.45–1.75 (1.57), PrW 0.79–0.91 (0.83), PSL 0.11–0.15 (0.13), SDL 0.12–0.16 (0.14), PL 0.51–0.59 (0.55), PH 0.31–0.36 (0.33), PW 0.21–0.25 (0.23), PPL 0.30–0.37 (0.35), PPH 0.26–0.31 (0.29), PPW 0.26–0.32 (0.31), MFL 0.96–1.27 (1.11), MTL 0.74–0.95 (0.85), CI 83–88 (83), SI 87–97 (94), REL 27–32 (32), FLI 27–30 (28), PSI 0.9–1.0 (0.9), MFI 76–88 (81), ACI1 58–61 (59), ACI2 90–99 (99).
Same as worker except for standard queen modifications and as follows: pronotum transversely carinulate; mesoscutum longitudinally carinulate, or carinate; scutellum rugoreticulate, or longitudinally carinulate to carinate; propodeum with transverse carinulae/carinae that wrap around surface; mesopleuron usually mostly smooth; pilosity on gastral tergites clearly bilayered, with a sparse layer of long, suberect setae, and a dense layer of appressed pubescence; wing venation in
See
This species is rather distinctive with its large size, lack of propodeal spines, and dense sculpturing. It should not be easily confused with any other MAC species.
Over its range,
Distribution map of
Integument mostly dark red-brown to orange-brown; small- to medium-sized species (see HL, ML, and PrW below); anterior clypeal margin undulating, with 4 blunt teeth; basal margin of mandible straight to slightly sinuous, without a basal notch or deep depression; gastral pilosity clearly bilayered, with a layer of suberect setae, and a layer of decumbent setae; petiole distinctly elongate (PL/HW 0.60–0.68); postpetiolar node dorsoventrally flattened and slightly elongate (PPH/PH 0.75–0.84); eye relatively small (EL 0.07–0.11, REL 13–18), subcircular, with 4–5 ommatidia at greatest diameter; median clypeal lobe projecting dorsally outward, resulting in rather distinct dorsal and anterior surfaces (visible in profile); dorsal surface of median lobe with a pair of distinct longitudinal carinulae that strongly diverge around median lobe at anterior margin; frontal lobes slightly to very strongly expanded (FLD 0.19–0.25, FLI 31–46), either completely or mostly covering torular lobes in full-face view; propodeal spines present, short to moderate length (PSL 0.12–0.17, PSI 1.6–1.9).
Southern Mexico to Honduras.
(7 measured) HL 0.65–0.76 (0.70), HW 0.54–0.66 (0.60), FLD 0.19–0.25 (0.24), PCW 0.03 (0.03), SL 0.47–0.56 (0.49), EL 0.07–0.11 (0.11), ACL 0.46–0.56 (0.50), ML 0.81–0.99 (0.87), PrW 0.40–0.48 (0.42), PSL 0.12–0.17 (0.14), SDL 0.07–0.10 (0.08), PL 0.35–0.42 (0.36), PH 0.16–0.19 (0.17), PW 0.11–0.14 (0.13), PPL 0.18–0.20 (0.19), PPH 0.12–0.15 (0.14), PPW 0.14–0.17 (0.16), MFL 0.52–0.64 (0.55), MTL 0.42–0.54 (0.46), CI 82–88 (86), SI 82–89 (82), REL 13–18 (18), FLI 31–46 (41), PSI 1.6–1.9 (1.9), MFI 102–109 (108), ACI1 65–70 (67), ACI2 96–102 (102).
Small- to medium-sized species; general body color dark red-brown to orange-brown, with patches of lighter brown on gaster; appendages orange-brown to yellow-brown; setae pale golden brown; mandible with 5–7 teeth, consisting of 3 distinct apical teeth, a distinct basal tooth, and 1–3 worn or denticulate inner teeth; basal margin of mandible straight to slightly sinuous, without a basal notch or deep depression; mandible mostly smooth and shiny, with scattered piligerous punctae and faint striae; median clypeal lobe projecting dorsally outward, resulting in rather distinct dorsal and anterior surfaces (visible in profile); anterior clypeal margin undulating, with 4 blunt teeth (best viewed at an anterodorsal angle); dorsal surface of median lobe with a pair of distinct longitudinal carinulae that strongly diverge around median lobe at anterior margin; remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions narrow (PCW 0.03), with sides subparallel to hourglass-shaped; frontal lobes slightly to very strongly expanded (FLD 0.19–0.25, FLI 31–46), either completely or mostly covering torular lobes in full-face view; frontal carinae weakly developed, not extending beyond frontal lobes; head subrectangular to somewhat oval-shaped (CI 82–88), posterior margin slightly depressed medially; eye appearing small (EL 0.07–0.11, REL 13–18), subcircular, with 4–5 ommatidia at greatest diameter; head strongly rugoreticulate, with a few longitudinal costae along midline, interstices faintly punctate; scape somewhat short (SI 82–89), not reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae; flagellum with a somewhat distinct 4-segmented antennal club; mesosoma completely sculptured, except for propodeal declivity, which has faint transverse carinulae; promesonotal dorsum rugose to rugoreticulate and punctate, with rugae transverse near anterior margin, becoming arcuate, and then longitudinal posteriad; mesosomal side mostly punctate, with scattered rugulae; anterodorsal margin of promesonotum with a somewhat distinct straight to lightly curving transverse carina, anterolateral margins of pronotum forming distinct shoulders in dorsal view; anterior declivity of pronotum in profile nearly vertical, and forming a sharp transition with the dorsum; promesonotum in profile low-domed, somewhat asymmetrical; metanotal grove distinct, deeper than average; anterodorsal margin of propodeum with a distinct welt; propodeal spines present, short to moderate length (PSL 0.12–0.17, PSI 1.6–1.9); petiole distinctly elongate and gracile (PL/HW 0.60–0.68); petiolar node relatively small (PH/PL 0.45–0.49), subconical and roughly symmetrical in profile, with a rounded dorsum; postpetiolar node dorsoventrally flattened and slightly elongate (PPH/PH 0.75–0.84); petiolar and postpetiolar nodes almost completely smooth and shining, remaining surfaces of waist segments mostly punctate, with a few rugulae surrounding postpetiolar node; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with short standing pilosity; gastral pilosity clearly bilayered, with a layer of suberect setae, and a layer of decumbent setae; scape setae decumbent to appressed; setae on legs mostly decumbent to appressed, with suberect setae on coxae and femoral venters.
Unknown.
Unknown.
The form of the waist segments combined with the projecting median clypeal lobe, make
The most significant form of variation within this species is the extent to which the frontal lobes are expanded. In the Cusuco population, the frontal lobes are greatly expanded laterally and anteroposteriorly, completely covering the torular lobes in full-face view. In the other populations, the frontal lobes are only moderately expanded and they do not completely cover the torular lobes in full-face view, leaving the outer margins somewhat visible. I treat this as intraspecific variation until more material can be gathered.
Molecular phylogenetic data infer that
Distribution map of
Note this species is variable and difficult to characterize globally. See comments section below, discussing population variants. Integument mostly dark brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, with a distinct basal depression, but no tooth; anterior clypeal margin undulating, forming 2–4 blunt teeth; eye of moderate size (EL 0.07–0.12, REL 14–21), oval-shaped, with 4–8 (usually 5–6) ommatidia at greatest diameter; propodeal spines tuberculate to short (PSL 0.06–0.11, PSI 1.0–1.4); face usually completely sculptured, with carinulae, rugoreticulae and punctae, but some populations with posterior ¼ or less of head smooth and shiny; pronotal sculpture variable, often with some carinulae and punctae, but some populations completely smooth; remainder of mesosoma sculptured, with punctae, carinulae and/or rugulae; propodeal lobe in profile usually isolated from propodeal spine and with angulate corners, but some populations with lobe appearing broadly rounded and forming a smooth connection with propodeal spine; geography is useful in species determination.
Southwestern U.S.A. to southern Mexico (Oaxaca).
Small-sized species; general body dark brown to brown (type population), with appendages brown or orange-brown to yellow-brown, usually lighter at joints and toward extremities; setae golden brown; mandible with 6–7 teeth, 2–3 teeth near basal tooth sometimes worn and indistinct; basal margin of mandible sinuous, with a distinct basal depression, but no tooth; mandible mostly smooth and shiny, with some scattered piligerous punctae, and some striations near base and on lateral surface; anterior clypeal margin undulating, forming 2–4 sharp to blunt teeth (4 sharp teeth in type population); median lobe of clypeus with a pair of longitudinal carinulae that diverge toward anterior margin, apex with a short transverse carinula, area in between median lobe and anterior clypeal margin forming a shallow concavity; remaining surface of clypeus mostly smooth; posterior extension of clypeus between antennal insertions somewhat narrow to moderate width (PCW 0.03–0.05), with sides subparallel; frontal lobes of moderate width (FLD 0.11–0.17, FLI 23–29), not greatly obscuring torular lobes in full-face view; head subrectangular to roughly oval-shaped (CI 83–92), posterior margin slightly depressed medially; eye of moderate size (EL 0.07–0.12, REL 14–21), oval-shaped, with 4–8 (usually 5–6) ommatidia at greatest diameter; face sculpture variable, usually completely sculptured, with light rugoreticulae, longitudinal carinulae, and/or punctae, but sometimes sculpture reduced, with posterior 1/4 or less of head becoming smooth and shiny; scape of short to moderate length (SI 81–94), usually not quite reaching posterior margin of head when laid back; scape surface mostly smooth to somewhat rough (type population), with variable density of piligerous punctae; flagellum with a distinct 4-segmented antennal club; pronotal sculpture highly variable, often with some longitudinal carinulae/rugulae and faint punctae (type population), but some populations completely smooth; remainder of mesosoma completely sculptured with punctae and a variable amount of rugulae/carinulae; promesonotum usually low-domed, and asymmetrical, with the anterior face longer and steeper than posterior face (type population), but some populations with promesonotum distinctly domed, and roughly symmetrical; metantoal groove present and distinct, of average width and depth; propodeal lobe in profile usually isolated from propodeal spine and with angulate corners, but some populations with lobe appearing broadly rounded and forming a smooth connection with propodeal spine; propodeal spines tuberculate to short (PSL 0.06–0.11, PSI 1.0–1.4); petiole length and shape variable, often short and stocky, with a somewhat large node that points vertically, and a sinuous venter (type population), but sometimes more elongate, with node smaller and pointing distinctly posteriad, or sometimes anteroposteriorly compressed, making it very narrow (PL/HW 0.46–0.59); postpetiole usually forming a small node, similar in size or smaller than petiolar node (type population) (PPH/PL 0.79–0.97); petiole and postpetiole usually mostly punctate, with only anterior faces of nodes smooth (type population), but some populations with punctae reduced and nodes mostly smooth; gaster usually completely smooth, but some populations with first sternite and tergite lightly to strongly punctate (variable in type population); most of body dorsum with relatively short standing pilosity; gastral pilosity distinctly to indistinctly bilayered, with a layer of longer suberect to subdecumbent setae, and a layer of decumbent setae, density of setae variable, usually relatively dense (type population); setae on scape decumbent to appressed; setae on legs decumbent to appressed with longer setae on femoral venters and coxae.
(8 measured) HL 0.55–0.71 (0.68), HW 0.50–0.65 (0.58) FLD 0.13–0.18 (0.16), PCW 0.03–0.05 (0.04), SL 0.43–0.56 (0.49), EL 0.15–0.18 (0.15), ACL 0.42–0.52 (0.49), ML 0.78–0.98 (0.85), PrW 0.43–0.59 (0.44), PSL 0.09–0.16 (0.12), SDL 0.06–0.10 (0.09), PL 0.29–0.38 (0.34), PH 0.16–0.23 (0.22), PW 0.13–0.18 (0.16), PPL 0.13–0.20 (0.18), PPH 0.15–0.21 (0.20), PPW 0.16–0.23 (0.21), MFL 0.46–0.60 (0.51), MTL 0.41–0.52 (0.45), CI 86–94 (86), SI 82–87 (85), REL 25–31 (25), FLI 23–28 (27), PSI 1.3–1.8 (1.3), MFI 105–114 (113), ACI1 66–71 (68), ACI2 92–100 (99).
Same as worker except for standard queen modifications and as follows (comparison with queens from near type locality only): pronotum with transverse carinulae/rugulae; mesoscutum longitudinally carinulate, with a small patch of smooth cuticle anteromesad; scutellum smooth along midline, with longitudinal rugulae mesad; anepisternum partly smooth, remainder carinulate; katepisternum mostly smooth; propodeum with transverse carinulae that wrap around propodeum; propodeal spines longer than worker; gastral pilosity denser; wing venation as in
Unknown.
There are some key morphological differences separating populations of
Within the
The main features of the type form of
Variant 1 (
Variant 2 (
Variant 3 (
Variant 4 (
Variant 5 (
Variant 6 (
Integument mostly dark brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible straight, never with a basal notch or depression; anterior clypeal margin with a shallow median emargination (best viewed from an anterodorsal angle); face completely sculptured, mostly rugoreticulate; mesosoma mostly sculptured, dorsum of promesonotum with dense longitudinal rugae; eye of moderate size (EL 0.10–0.12, REL 18–21), oval-shaped, with 5–7 ommatidia at greatest diameter; gastral pilosity relatively long, sparse, and mostly suberect, sometimes with a few underlying decumbent setae; petiole in profile appearing somewhat elongate; petiolar node reaching a distinct apex, which points vertically; propodeal spines tuberculate to short (PSL 0.07–0.12, PSI 1.2–2.1); frontal lobes of moderate width (FLD 0.14–0.16, FLI 24–27), not greatly obscuring torular lobes in full-face view.
Southern Mexico to Guatemala.
(18 measured) HL 0.59–0.68 (0.64), HW 0.53–0.61 (0.55), FLD 0.14–0.16 (0.14), PCW 0.02–0.03 (0.02), SL 0.47–0.55 (0.50), EL 0.10–0.12 (0.10), ACL 0.45–0.52 (0.49), ML 074–0.86 (0.74), PrW 0.36–0.43 (0.37), PSL 0.07–0.12 (0.07), SDL 0.05–0.07 (0.05), PL 0.25–0.32 (0.27), PH 0.15–0.18 (0.15), PW 0.11–0.16 (0.12), PPL 0.14–0.18 (0.15), PPH 0.13–0.17 (0.13), PPW 0.15–0.19 (0.15), MFL 0.52–0.62 (0.54), MTL 0.42–0.50 (0.45), CI 87–92 (87), SI 85–96 (89), REL 18–21 (18), FLI 24–27 (26), PSI 1.2–2.1 (1.6), MFI 96–104 (103), ACI1 66–70 (67), ACI2 92–100 (98).
Small-sized species; general body color mostly dark brown to brown, with appendages brown to orange-brown, becoming lighter toward extremities; setae golden brown; mandible with 5–6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 1–2 smaller inner teeth/denticles, which are often worn and indistinct; basal margin of mandible straight, without basal notch or depression; mandible mostly smooth and shiny, with scattered piligerous punctae, and some striations around base and along lateral surface; anterior clypeal margin with a shallow median emargination (best viewed from anterodorsal angle); median lobe of clypeus with a pair of faint longitudinal carinulae that diverge toward anterior margin, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.02–0.03), sides subparallel to slightly hour-glass-shaped; frontal lobes of moderate width (FLD 0.14–0.16, FLI 24–27), not greatly obscuring torular lobes in full-face view; head roughly oval-shaped (CI 87–92), with posterior margin slightly depressed medially; eye of moderate size (EL 0.10–0.12, REL 18–21), oval-shaped, with 5–7 ommatidia at greatest diameter; face mostly rugoreticulate, with some longitudinal rugae along midline; scape of moderate length (SI 85–96), almost reaching posterior margin when laid back; scape surface smooth and shiny, with scattered piligerous punctae; flagellum with distinct 4-segmented antennal club; mesosoma almost completely sculptured except for a small patch of smooth cuticle on side of pronotum; dorsum of pronotum densely rugose (longitudinal orientation), transitioning to rugoreticulae on mesonotum; upper half of side of mesonotum rugose; mesopleuron mostly punctate, with some rugulae; side of propodeum rugulose-punctate; dorsum and declivity of propodeum with transverse carinulae; promesonotum in profile low-domed, slightly asymmetrical with apex shifted anterior of midpoint; metanotal groove of moderate width and depth; propodeal spines tuberculate to short (PSL 0.07–0.12, PSI 1.2–2.1); petiole appearing of moderate length to slightly elongate (PL/HW 0.47–0.54); petiolar node in profile of moderate height (PH/PL 0.55–0.66), and roughly symmetrical, dorsum pointing vertically to slightly posteriad, and usually reaching a well-defined apex; postpetiole in profile slightly asymmetrical, with anterior face longer and more sloping than posterior face, overal size similar to petiolar node (PPH/PH 0.85–0.98); anterior faces of petiolar and postpetiolar nodes mostly smooth and shiny, remaining surfaces faintly punctate; gaster smooth and shiny, with scattered piligerous punctae; most of body dorsum with short to long standing pilosity; setae on gastral dorsum relatively sparse, mostly long and suberect, with a few shorter decumbent setae underneath; setae on scape and legs decumbent to appressed, with some longer suberect setae on femoral venters and coxae.
(5 measured) HL 0.66–0.74 (0.66), HW 0.61–0.67 (0.61), FLD 0.15–0.18 (0.15), PCW 0.02–0.03 (0.03), SL 0.52–0.58 (0.52), EL 0.17–0.20 (0.17), ACL 0.49–0.55 (0.49), ML 0.94–1.09 (0.94), PrW 0.54–0.63 (0.54), PSL 0.13–0.17 (0.13), SDL 0.09–0.10 (0.10), PL 0.37–0.40 (0.37), PH 0.19–0.22 (0.19), PW 0.16–0.19 (0.16), PPL 0.16–0.22 (0.16), PPH 0.18–0.23 (0.18), PPW 0.20–0.24 (0.20), MFL 0.59–0.70 (0.59), MTL 0.50–0.58 (0.50), CI 90–93 (92), SI 85–91 (85), REL 28–30 (28), FLI 25–27 (25), PSI 1.4–1.7 (1.4), MFI 95–103 (1.03), ACI1 64–68 (67), ACI2 94–96 (96).
Same as worker except for standard queen modifications and as follows: pronotum with transverse rugae/carinulae; mesoscutum densely longitudinally carinulae; scutellum rugose to rugoreticulate; propodeum with transverse carinulae that wrap around entire surface; mesopleuron mostly smooth; gastral pilosity denser, especially lower decumbent layer; wing as in
See
This species is very similar to
There is some variation in size and sculpture among populations of
I have sampled a number of populations of
Distribution map of
Integument mottled, pale yellow-brown with patches of darker brown; small-sized species (see HL, ML, PrW below); head and mesosoma almost completely covered with light longitudinal rugulae; eye relatively small (EL 0.06–0.08, REL 10–13), subcircular, with 3–5 ommatidia at greatest diameter; basal margin of mandible slightly sinuous, without a basal notch or deep depression; anterior clypeal margin with a single median emargination; frontal lobes somewhat narrow, not completely obscuring torular lobes in full-face view (FLD 0.15-0.16, FLI 25-28); propodeal spines present, forming short broad triangles (PSL 0.11–0.12, PSI 1.4-1.6); gastral pilosity short and clearly bilayered, with a layer of suberect setae and a layer of decumbent setae.
Northwestern Mexico (Baja California Sur).
(5 measured) HL 0.66–0.70 (0.68), HW 0.55–0.61 (0.58), FLD 0.15–0.16 (0.15), PCW 0.03–0.04 (0.04), SL 0.54–0.56 (0.56), EL 0.06–0.08 (0.06), ACL 0.54–0.58 (0.59), ML 0.79–0.85 (0.83), PrW 0.38–0.40 (0.39), PSL 0.11–0.12 (0.11), SDL 0.05–0.07 (0.06), PL 0.29–0.32 (0.30), PH 0.18–0.19 (0.19), PW 0.12–0.13 (0.13), PPL 0.16–0.17 (0.16), PPH 0.14–0.16 (0.15), PPW 0.15–0.18 (0.17), MFL 0.59–0.60 (0.59), MTL 0.49–0.52 (0.52), CI 83–87 (86), SI 92–98 (95), REL 10–13 (10), FLI 25–28 (26), PSI 1.4–1.6 (1.4), MFI 94–101 (98), ACI1 65–67 (67), ACI2 97–103 (97).
Small-sized species; general body color a mottled pale yellow-brown with patches of darker brown, appendages similar; setae golden brown; mandible with 6 teeth, consisting of 2–3 distinct apical teeth, a basal tooth, and 2-3 middle teeth, which are usually worn and indistinct; basal margin of mandible straight to slightly sinuous, without a basal notch or deep depression; mandible mostly smooth and shiny, with scattered piligerous punctae; anterior clypeal margin with a median emargination, clypeus bordering emargination slightly projecting and attenuate, appearing translucent; median lobe of clypeus obliquely flattened, lacking longitudinal carinulae, apex near anterior clypeal margin with a short transverse carinula, remainder of clypeus smooth and shiny; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.04), sides subparallel; frontal lobes somewhat narrow (FLD 0.15–0.16, FLI 25–28), with underlying torular lobes clearly visible in full-face view; head appearing subrectangular to oval-shaped (CI 83–87), posterior margin flat to very slightly depressed medially; eye relatively small (EL 0.06–0.08, REL 10-13), subcircular, with 3–5 ommatidia at greatest diameter; face completely covered with longitudinal rugulae and piligerous punctae, rugulae sometimes merging, but never becoming truly reticulate, posterior margin of head near occipital foramen mostly smooth and shiny; scape of moderate length (SI 92–98), not quite reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae and fine striations; flagellum with a somewhat distinct 4-segmented antennal club; mesosoma mostly with longitudinal carinulae and rugulae, some faint punctae also present on most surfaces (a few aberrant specimens with transverse carinulae on pronotum); propodeal dorsum and declivity with faint transverse carinulae; promesonotum in profile low-domed, roughly symmetrical; metanotal grove well demarcated, of moderate width and depth; propodeal spines present, forming short broad triangular projections (PSL 0.11–0.12, PSI 1.4–1.6); petiole in profile somewhat compact (PL/PW 51–55), node domed and pointing slightly posteriad, node dorsum flat to gently rounded; postpetiole somewhat dorsoventrally compressed, clearly smaller than petiolar node (PPH/PH 0.77–0.82); petiolar and postpetiolar nodes mostly smooth and shiny, remaining waist surfaces faintly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body with short standing pilosity; setae on scape decumbent to appressed; gastral pilosity short, somewhat dense, and clearly bilayered, with a layer of suberect setae and a layer of decumbent setae (setae on head similar); setae on legs mostly decumbent to appressed, with some suberect setae on venter of profemur.
(1 measured) HL 0.71, HW 0.61, FLD 0.16, PCW 0.04, SL 0.57, EL 0.17, ACL 0.59, ML 0.97, PrW 0.50, PSL 0.15, SDL 0.15, PL 0.34, PH 0.21, PW 0.14, PPL 0.17, PPH 0.17, PPW 0.20, MFL 0.64, MTL 0.55, CI 86, SI 94, REL 29, FLI 26, PSI 1.7, MFI 96, ACI1 62, ACI2 103.
Same as worker except for standard queen modifications and as follows: body color a darker yellow-brown; pronotum with transverse carinulae on sides, smooth in middle; mesoscutum carinulate, with carinulae long and distinct; katepisternum mostly smooth; propodeum with transverse carinulae that wrap around surface; propodeal spines blunt, slightly longer.
Unknown.
Distribution map of
Integument mostly dark brown to brown; medium-sized species (see HL, HW, ML below); lateral margin of hypostomal bridge with a projecting subquadrate to broadly-rounded lobe (usually visible behind mandible in profile); propodeal spines long and rather slender (PSL 00.17–0.22, PSI 1.9–2.2); anterior clypeal margin with a median emargination; basal margin of mandible straight, without a basal notch or depression; head and mesosoma densely carinulate; eye large (EL 0.13–0.15, REL 16–19), with 6–8 ommatidia at greatest diameter; petiolar node in profile subconical, with apex well-defined, almost sharp; postpetiolar node anteroposteriorly compressed; gastral pilosity with a layer of long suberect setae and a sparse layer of shorter decumbent setae; frontal lobes of moderate width (FLD 0.21–0.23, FLI 26–31), not completely obscuring torular lobes in full-face view.
Southern Mexico.
(10 measured) HL 0.80–0.93 (0.91), HW 0.70–0.87 (0.87), FLD 0.21–0.23 (0.23), PCW 0.04–0.08 (0.04), SL 0.69–0.80 (0.75), EL 0.13–0.15 (0.15), ACL 0.62–0.68 (0.68), ML 1.02–1.18 (1.17), PrW 0.47–0.55 (0.55), PSL 0.17–0.22 (0.22), SDL 0.08–0.10 (0.10), PL 0.35–0.43 (0.42), PH 0.22–0.26 (0.26), PW 0.15–0.19 (0.19), PPL 0.19–0.25 (0.25), PPH 0.20–0.25 (0.25), PPW 0.19–0.25 (0.24), MFL 0.79–0.92 (0.89), MTL 0.63–0.74 (0.73), CI 87–96 (96), SI 87–98 (87), REL 16–19 (18), FLI 26–31 (26), PSI 1.9–2.2 (2.2), MFI 88–97 (97), ACI1 63–66 (64), ACI2 85–92 (91).
Medium-sized species; general body color dark brown to brown, with appendages brown to orange-brown; setae dark golden brown; mandible with 6 teeth, with two teeth nearest basal tooth smaller, often worn and indistinct; basal margin of mandible straight, without a basal notch or depression; mandible surface mostly smooth and shiny, with scattered piligerous punctae; anterior clypeal margin with a median emargination; median lobe of clypeus with a pair of faint longitudinal carinulae that diverge anteriorly, apex with a short transverse carinula, remainder of clypeus mostly smooth and shiny, with a few faint striae on median lobe; posterior extension of clypeus between antennal insertions of moderate to somewhat wide width (PCW 0.04–0.08); frontal lobes of moderate width (FLD 0.21–0.23, FLI 26–31), not completely obscuring torular lobes in full-face view; lateral margin of hypostomal bridge with a projecting subquadrate to broadly-rounded lobe that is usually visible behind mandible in profile, but some specimens with lobe reduced; head subcircular to oval-shaped (CI 87–96), posterior margin with a slight median depression; eye large (EL 0.13–0.15, REL 16–19), roughly oval-shaped, with 6–8 ommatidia at greatest diameter; face densely sculptured with a fan of longitudinal carinulae that extend outward from frontal lobes, lateral margins with shorter less densely spaced rugulae, interstices faintly punctate; scape of moderate length (SI 87–98), nearly reaching posterior margin of head when laid back; scape surface mostly smooth and shiny with scattered piligerous punctures; flagellum with a somewhat distinct 4-segmented antennal club; mesosoma densely sculptured; promesonotum mostly longitudinally carinulate, with some transverse carinulae wrapping around anterior declivity; mesopleuron and side of propodeum with irregular rugulae, rugoreticulae, and punctae; propodeal dorsum and part of declivity with transverse carinulae, remainder of declivity smooth and shining; promesonotum in profile low-domed and somewhat asymmetrical, with posterior slope longer and shallower than anterior slope; metanotal grove well demarcated and moderately deep, forming a rather smooth transition with promesonotum; anterior margin of propodeal dorsum in profile sometimes distinctly raised and separated from metanotal grove; propodeal spines long and rather slender (PSL 00.17–0.22, PSI 1.9–2.2), pointing anteroposteriorly and outward from body; petiole of moderate length (PL/HW 0.47–0.51), peduncle in profile somewhat slender; petiolar node in profile somewhat high (PH/PL 0.59–0.69), subconical and roughly symmetrical, with a well-defined apex that points vertically; postpetiolar node in profile usually somewhat anteroposteriorly compressed, with apex rounded and pointing vertically or slightly posteriad; anterior faces of petiolar and postpetiolar nodes smooth and shiny, posterior faces with punctae and a few rugulae, remaining surface of waist punctate; gaster mostly smooth and shiny, with scattered piligerous punctae and a few short striae around anterior constriction; most of body with rather long standing pilosity; scape with a uniform layer of subdecumbent to appressed setae; gaster with a layer of long suberect setae and a sparse layer of shorter decumbent setae; setae on legs mostly subdecumbent to appressed, with suberect setae on coxae and femoral venters.
Unknown.
Unknown.
This species could be confused with
It is worth noting that the lateral hypostomal lobe that groups
Integument black to brown-black; medium-sized species (see HL, ML, PrW below); anterior margin of clypeus with shallow median emargination; basal margin of mandible straight, without notch or substantial depression; head and mesosoma mostly smooth and shining; mesosoma compact, with promesonotum distinctly bulging; petiolar node robust, tall, and distinctly angled posteriad; propodeal spines forming short broad triangles (PSL 0.09–0.14, PSI 1.7-2.5); setae on gastral tergites sparse, long, and suberect; eye of moderate size (EL 0.11–0.15, REL 19-23), oval-shaped, with 6–8 ommatidia at greatest diameter; frontal lobes narrow, not obscuring torular lobe in full-face view (FLD 0.14-0.18, FLI 25-28).
Southern Mexico to Guatemala.
(8 measured) HL 0.59–0.72 (0.70), HW 0.56–0.69 (0.64), FLD 0.14–0.18 (0.17), PCW 0.05–0.07 (0.06), SL 0.46–0.56 (0.55), EL 0.11–0.15 (0.14), ACL 0.45–0.53 (0.52), ML 0.76–0.99 (0.90), PrW 0.41–0.55 (0.50), PSL 0.09–0.14 (0.12), SDL 0.05–0.08 (0.07), PL 0.31–0.41 (0.38), PH 0.23–0.31 (0.31), PW 0.14–0.20 (0.19), PPL 0.17–0.21 (0.21), PPW 0.18–0.26 (0.23), MFL 0.52–0.65 (0.63), MTL 0.39–0.50 (0.48), CI 92–96 (92), SI 80–85 (85), REL 19–23 (21), FLI 25–28 (27), PSI 1.7–2.5 (1.8), MFI 101–113 (101), ACI1 65–70 (68), ACI2 91–98 (95).
Medium-sized species; general body color black to red- or brown-black, with mandibles and appendages lighter, usually dark brown to yellow-brown; setae light brown; mandible with 6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2 inner teeth, which are often worn and indistinct; basal margin of mandible relatively straight, without any notch or significant depression; dorsal surface of mandible mostly smooth and shining, with scattered piligerous punctae and a few short basal striae; median lobe of clypeus with pair of very faint vestigial carinulae that diverge toward anterior margin, apex with a short transverse carinula, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between frontal lobes moderately broad (PCW 0.05-0.07), with subparallel to slightly diverging sides; frontal lobes narrow, not obscuring torular lobes in full-face view; head roughly oval-shaped (CI 92-96), but appearing somewhat tear drop-shaped because of the angled anterior margin of the clypeus and position of eyes; posterior margin of the head flat, never distinctly depressed medially; eye of moderate size (EL 0.11–0.15, REL 19-23), oval-shaped, with 6–8 ommatidia at greatest diameter; face almost completely smooth and shining, with scattered piligerous punctae and a few longitudinal carinulae on gena; scape short (SI 80-85), not reaching posterior margin of head when laid back; scape mostly smooth and shining, with scattered piligerous punctae, and sometimes a few fine striae; flagellum with a distinct 4-segmented antennal club; mesosoma usually mostly smooth and shiny, with some longitudinal carinae in metanotal groove, and a few scattered rugulae on propodeum, but some specimens with more developed sculpture on mesonotum, mesopleuron, and propodeum, consisting of faint carinulae and punctae; promesonotum domed and distinctly bulging upwards above head and propodeum; promesonotal suture usually completely effaced dorsally, but in a few specimens pronotum appears separated from mesonotum; metanotal groove distinct and of moderate depth; propodeal spines short, forming broad triangles (PSL 0.09–0.14, PSI 1.7-2.5); petiole of moderate length (PL/HW 0.52–0.62), node in profile robust and very tall (PH/PL 0.66–0.80), with anterior face longer and more sloping than posterior face, dorsum of node distinctly angled posteriad, and almost reaching a sharp apex; postpetiole in profile subcircular to oval-shaped, somewhat globular, always smaller than petiolar node (PPH/PH 0.70–0.84), dorsum usually with a somewhat distinct longitudinal median lobe; petiole and postpetiole mostly punctate, sometimes with faint rugulae, anterior faces of nodes smooth and shiny; gaster smooth and shiny except scattered piligerous punctae; most of body with moderately long, erect to subdecumbent setae; scapes with subdecumbent to decumbent setae; setae on legs mostly decumbent to appressed, with some suberect setae on femoral venters and coxae.
Unknown.
Unknown.
With its mostly smooth sculpture and bulging promesonotum,
Distribution map of
Integument black to brown-black and shining; medium-sized species (see HL, ML, PrW below); anterior clypeal margin with shallow median emargination; basal margin of mandible relatively straight, without notch or substantial depression; face mostly smooth and shiny; mesosoma with coarse, rugose to rugoreticulate sculpture on most surfaces; promesonotum in profile distinctly asymmetrical, with anterior face gently rounded, dorsal surface flattened, and posterior face short, straight and forming a relatively sharp angle with dorsum; petiolar node, generally robust, and distinctly angled posteriad; postpetiolar node in profile with a longitudinal dorsal lobe that projects out posteriad over postpetiole; eye of moderate size (EL 0.11–0.14; REL 18-22), oval-shaped, with 7–8 ommatidia at greatest diameter; propodeal spines ranging from short tubercles to long spines (PSL 0.10–0.22, PSI 1.3-2.6); setae on gastral tergites relatively sparse, long, and mostly suberect; frontal lobes narrow, not obscuring torular lobe in full-face view (FLD 0.13-0.16, FLI 21-25).
Mexico (Atlantic slope).
(10 measured) HL 0.65–0.76 (0.73), HW 0.57–0.71 (0.67), FLD 0.13–0.16 (0.15), PCW 0.03–0.05 (0.04), SL 0.52–0.62 (0.61), EL 0.11–0.14 (0.14), ACL 0.50–0.58 (0.56), ML 0.84–1.03 (0.97), PrW 0.41–0.51 (0.49), PSL 0.10–0.22 (0.20), SDL 0.06–0.09 (0.08), PL 0.35–0.44 (0.42), PH 0.23–0.29 (0.27), PW 0.14–0.18 (0.18), PPL 0.16–0.22 (0.21), PPH 0.16–0.19 (0.18), PPW 0.16–0.21 (0.20), MFL 0.59–0.71 (0.70) MTL 0.47–0.57 (0.55), CI 89–95 (92), SI 85–92 (90), REL 18–22 (21), FLI 21–25 (23), PSI 1.3–2.6 (2.4), PI 58–65 (62) MFI 96–103 (96), ACI1 64–68 (66), ACI2 90–100 (92).
Medium-sized species; general body color black to brown- or red-black, with mandibles and appendages lighter, usually dark brown to yellow-brown; setae golden; mandible with 6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2 inner teeth, which are often worn and indistinct; basal margin of mandible relatively straight, without a notch or significant depression; dorsal surface of mandible mostly smooth and shiny, with scattered piligerous punctae and a few short basal striae; median lobe of clypeus with a pair of vestigial longitudinal carinulae that diverge toward the anterior margin, apex with a short transverse carinula, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between frontal lobes moderately broad (PCW 0.03-0.05), with subparallel to slightly diverging sides; frontal lobes narrow (FLD 0.13–0.16, FLI 21–25), not obscuring torular lobes in full-face view; head roughly oval-shaped (CI 89–95), but appearing somewhat tear drop-shaped because of the angled anterior margin of clypeus and position of eyes; posterior margin of head flat, never distinctly depressed medially; eye of moderate size (EL 0.11–0.14; REL 18-22), oval-shaped, with 7–8 ommatidia at greatest diameter; face almost completely smooth and shiny, with scattered piligerous punctae and few longitudinal carinulae on genae; scape moderately long (SI 85-92), nearly reaching posterior margin of head when laid back; scape surface largely smooth and shiny, with scattered piligerous punctae, and sometimes with a few faint striae; flagellum with distinct 4-segmented antennal club; mesosoma almost entirely with coarse rugae and rugoreticulae; rugae usually with a predominately longitudinal orientation, but in some specimens rugae on mesonotum have a transverse orientation; katepisternum, metapleuron, and lateral portion of pronotum, with a variably sized patch of smooth cuticle; propodeal dorsum and declivity with transverse carinulae; promesonotum in profile distinctly asymmetrical, with anterior face gently rounded, dorsal surface flattened, and posterior face short, straight and forming a relatively sharp angle with dorsum; promesonotal suture indistinct, but usually discernable, especially when the orientation of the pronotal and mesonotal rugae differ; metanotal grove distinct and of moderate depth; propodeal spines present and often relatively long, but sometimes (usually specimens from lower elevations) becoming reduced to short tubercles (PSL 0.10–0.22, PSI 1.3–2.6); petiole in profile relatively long (PH/HW 0.58–0.65) and very distinctive, having a long peduncle and robust node, which is markedly angled posteriad; petiolar node in profile with long sloping anterior face that begins at about middle of peduncle, posterior face shorter, but usually at a similar angle, dorsum of node broadly rounded to subquadrate; postpetiole in profile asymmetrical, with a long sloping anterior face, and a short nearly vertical posterior face, dorsum with a distinct longitudinal lobe that projects out slightly posteriad over postpetiole, lobe in dorsal view attenuating posteriad, giving it a pinched-in appearance; petiole and postpetiole with variable amount of rugae and punctae; anterior faces of nodes mostly smooth and shiny with remaining surfaces punctatorugose; punctae most visible on ventral surfaces; posterior slope of petiolar node sometimes with a median keel; in a few aberrant populations the petiole and postpetiole lack rugae and are almost completely punctate; gaster smooth and shiny, with scattered piligerous punctae; most of body with sparse layer of moderately long standing seate; setae on scape mostly subdecumbent; setae on remaining appendages suberect to decumbent, with longer suberect setae on femoral venters and coxae.
(5 measured) HL 0.71–0.77 (0.72), HW 0.67–0.73 (0.67), FLD 0.16–0.17 (0.16), PCW 0.03–0.05 (0.03), SL 0.58–0.63 (0.58), EL 0.17–0.19 (0.17), ACL 0.55–0.60 (0.56), ML 1.03–1.16 (1.03), PrW 0.60–0.65 (0.60), PSL 0.17–0.23 (0.17), SDL 0.08–0.11 (0.09), PL 0.41–0.49 (0.41), PH 0.26–0.31 (0.28), PW 0.19–0.22 (0.19), PPL 0.20–0.24 (0.22), PPH 0.20–0.24 (0.21), PPW 0.23–0.26 (0.23), MFL 0.67–0.74 (0.67), MTL 0.53–0.60 (0.53), CI 93–97 (93), SI 84–89 (87), REL 26–27 (26), FLI 23–24 (24), PSI 1.9–2.5 (1.9), MFI 98–102 (100), ACI1 66–69 (66), ACI2 94–96 (95).
Same as worker except for standard queen modifications and as follows: lateral surfaces of pronotum transversely rugose to rugoreticulate, median area smooth and shiny; mesoscutum longitudinally carinate, with carinae fanning out from middle of anterior margin; mesopleuron mostly smooth and shiny; wing venation as in
See
This species is known only from Winkler or Berlese extractions of sifted leaf litter taken from montane, mesophyll forests. Nests have never been collected. It is likely a mid-elevation specialist given its occurrence between 860 to 1620 m.
At the El Cielo reserve in northern Mexico, I found a lone worker that looked like
The populations from Taman in San Luis Potosí and Mirador Grande in Oaxaca are particularly aberrant and I describe them here as variant 1 of
I have decided not to split these variants into a separate species because of lack of sympatry between the two forms and the fact that substantial variation exists along elevational gradients and among sites. Also, the Mirador Grande site, which is where most of the aberrant specimens are from, is very far away from the remaining populations. With that said, there is some evidence to suggest that variant 1 does indeed represent a distinct species. First, there is one record of variant 1 from Taman in San Luis Potosí, which is very near the El Madroño site in Querétaro (< 30 km). Second, the intrapopulation variation at El Cielo is substantial, yet it does not encompass the morphology of the aberrant specimens, even at the same elevation. Lastly, preliminary molecular evidence shows that a specimen from Mirador Grande is as divergent from El Madroño and El Cielo specimens as some clearly distinct (and sympatric) sister species pairs. To solve this problem, more collecting should be done in the region of Taman and El Madroño in order to find an area of sympatry or to provide additional fresh specimens for molecular work.
Integument mostly dark brown to dark red-brown; large-sized species(see HL, ML, PrW below); lateral margin of hypostomal bridge with a projecting quadrate lobe (visible behind mandible in profile); gaster, and most dorsal surfaces, with a layer of thickened suberect to subdecumbent setae, gaster also with sparse layer of thinner appressed setae; mesosoma, waist, and head densely punctate (punctae fainter on head); head and mesosoma with fine rugulae and rugoreticulae, which on the mesosoma merge into the punctae, making them less obvious; eye relatively small (EL 0.13–0.16, REL 13–15), oval-shaped, with 6–7 ommatidia at greatest diameter; anterior clypeal margin with a shallow median emargination; basal margin of mandible straight to slightly sinuous, without a basal notch or depression; propodeal spines short (PSL 0.16–0.19, PSI 1.4–1.6), frontal lobes narrow (FLD 0.24–0.27, FLI 26), not obscuring torular lobes in full-face view.
Southern Mexico.
(4 measured) HL 1.02–1.13 (1.12), HW 0.92–1.05 (1.01), FLD 0.24–0.27 (0.26), PCW 0.05–0.08 (0.08), SL 0.79–0.85 (0.83), EL 0.13–0.16 (0.13), ACL 0.67–0.74 (0.74), ML 1.29–1.41 (1.36), PSL 0.16–0.19 (0.17), SDL 0.10–0.13 (0.10), PrW 0.57–0.64 (0.61), PL 0.46–0.49 (0.49), PH 0.29–0.32 (0.30), PW 0.22–0.25 (0.24), PPL 0.27–0.30 (0.30), PPH 0.26–0.30 (0.29), PPW 0.28–0.31 (0.30), MFL 0.93–1.03 (1.02), MTL 0.57–0.80 (0.57), CI 91–93 (91), SI 81–88 (82), REL 13–15 (13), FLI 26 (26), PSI 1.4–1.6 (1.6), MFI 99–102 (99), ACI1 64–66 (64), ACI2 84–89 (89).
Large species; general body color dark brown (almost black) to dark red-brown, with appendages and some parts of the gaster lighter brown; setae a rich golden brown; mandible with 6 teeth, middle 2 teeth near basal tooth sometimes worn and indistinct; basal margin of mandible straight to slightly sinuous, without a basal notch or significant depression; mandible mostly smooth and shining with scattered piligerous punctae and striae; anterior clypeal margin with a median emargination; median lobe of clypeus with a pair of faint longitudinal carinulae that diverge anteriorly, area in between carinulae slightly depressed, apex of median lobe with a short transverse carinula, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.05–0.08), sides slightly hourglass-shaped; frontal lobes narrow (FLD 0.24–0.27, FLI 26), not obscuring torular lobes in full-face view; lateral margin of hypostomal bridge with a projecting quadrate lobe (visible behind mandible in profile); head robust, somewhat heart-shaped (CI 91–93), with posterior margin broadly depressed medially; eye relatively small compared to HW (REL 13–15), but appearing of moderate size (EL 0.13–0.16), with 6–7 ommatidia at greatest diameter, oval-shaped; face densely sculptured with longitudinal carinulae and rugulae, becoming rugoreticulae toward lateral margins, interstitial areas noticeably punctate; scape relatively short (SI 81–88), not reaching posterior margin of head when laid back; scape surface with scattered carinulae and faint piligerous punctae; flagellum with a somewhat indistinct 4-segmented antennal club; mesosoma densely punctate, and with fine rugulae that merge into surrounding punctae; rugulae on anterior half of pronotal dorsum transversely arcuate across surface; rugulae on remainder of promesonotal dorsum longitudinal in orientation; pronotal side completely punctate; rugulae on mesopleuron somewhat reticulate; part of propodeal dorsum and declivity with transverse carinulae; promesonotum in profile low-domed and asymmetrical, with apex occurring anterior of midpoint; metanotal grove somewhat indistinct with anterior margin merging smoothly with promesonotal declivity; propodeal spines well-developed, short (PSL 0.16–0.19, PSI 1.4–1.6); petiole of moderate length and robust (PL/HW 0.47–0.50), node of moderate height (PH/PL 0.62–0.65), broadly domed, and pointing slightly posteriad; postpetiole in profile nearly symmetrical, somewhat bulging, about as high as petiolar node (PPH/PH 0.90–0.96); petiole and postpetiole completely punctate and with a few rugulae on posterior sides of nodes; gaster mostly smooth and shiny, with scattered piligerous punctae; much of dorsal body surface with a layer of stout standing setae; setae on scape thin, dense, and decumbent (almost pubescent); gaster with a short layer of thickened suberect to subdecumbent setae and a very sparse layer of thinner appressed setae; setae on legs mostly appressed, with a few suberect setae on the coxae and femoral venters.
Unknown.
Unknown.
This species is known from a single collection event made in mesophyll forest at 1680 m elevation. A few workers were found under epiphytes in a treefall. Over 100 leaf litter samples were collected from the same site and no additional specimens were retrieved. Consequently, this species is likely to be arboreal in its habits.
Known only from the type locality.
Note that this species is highly variable. See comments section below discussing population variants. Integument color variable; medium- to large-sized species (see HL, ML, PrW below); lateral apex of hypostomal bridge projecting ventrally as a subquadrate to broadly rounded lobe, which is usually visible behind base of mandible in profile view (sometimes visible only in lateroventral view); propodeal spines tuberculate to short (PSL 0.09–0.19, PSI 1.0–1.6); basal margin of mandible straight; anterior clypeal margin with a single median emargination; face usually completely sculptured, mostly rugoreticulate, with some longitudinal rugulae/carinulae along midline, but sometimes face mostly smooth, with only some longitudinal carinulae; mesosoma usually mostly sculptured with carinae, rugae, rugoreticulae, or punctae, only sometimes with pronotum mostly or completely smooth; eye of moderate size (EL 0.10–0.16, REL 13–21), oval-shaped, with 5–8 ommatidia at greatest diameter; frontal lobes of moderate width (FLD 0.19–0.29, FLI 25–30); first gastral sternite and tergite sometimes punctate.
Mexico to Nicaragua.
(25 measured, lectotype in parentheses) HL 0.81–1.13 (0.91), HW 0.70–1.01 (0.78), FLD 0.19–0.27 (0.23), PCW 0.05–0.08 (0.08), SL 0.68–1.04 (0.75), EL 0.10–0.16 (0.14), ACL 0.62–0.88 (0.67), ML 1.05–1.50 (1.17), PrW 0.45–0.64 (0.53), PSL 0.09–0.19 (0.12), SDL 0.08–0.14 (0.10), PL 0.36–0.54 (0.44), PH 0.21–0.32 (0.25), PW 0.17–0.25 (0.21), PPL 0.21–0.33 (0.25), PPH 0.21–0.33 (0.25), PPW 0.22–0.34 (0.26), MFL 0.77–1.28 (0.85), MTL 0.61–0.97 (0.67), CI 82–91 (86), SI 88–109 (96), REL 13–21 (18), FLI 26–30 (29), PSI 1.0–1.6 (1.2), MFI 75–100 (92), ACI1 61–66 (65), ACI2 84–93 (89).
Medium- to large-sized species; general body color highly variable, ranging from mostly black (type population), to red-brown, to brown, to yellow-brown, with appendages lighter, especially at joints and toward extremities, generally brown or orange-brown to yellow-brown; setae golden brown; mandible with 6–7 teeth (usually 6), consisting of 3 distinct apical teeth, a basal tooth, and 2–3 smaller teeth in between, which are often worn and indistinct; basal margin of mandible straight, without a basal notch or depression; mandible mostly smooth, with scattered piligerous punctae, and a variable number of longitudinal striations, mostly at base and on lateral surface; anterior clypeal margin with a shallow median emargination; median lobe of clypeus often with a pair of faint longitudinal carinulae (type population) that diverge toward anterior margin, but sometimes distinct carinulae replaced or hidden by a variable number of irregular striations, apex of lobe usually with a short transverse carinula, remainder of clypeus mostly smooth; posterior extension of clypeus between frontal lobes of moderate width (PCW 0.05–0.08), with sides subparallel; frontal lobes of moderate width (FLD 0.19–0.27, FLI 26–30), never greatly obscuring torular lobes in full-face view; lateral apex of hypostomal bridge projecting ventrally as a subquadrate to broadly rounded lobe, which is usually visible behind base of mandible in profile view (reduced in type population; sometimes visible only in lateroventral view); head usually roughly oval-shaped (type population), but sometimes slightly elongate, or more often broad, becoming slightly heart-shaped (CI 82–91), posterior margin slightly to distinctly depressed medially; eye of moderate size (EL 0.10–0.16, REL 13–21), oval-shaped, with 5–8 ommatidia at greatest diameter; face usually completely sculptured, mostly rugoreticulate (rarely completely), with some longitudinal rugulae/carinulae along midline extending from frontal lobes to posterior margin, coarseness of sculpture variable (average in type population); face rarely mostly smooth, with only some longitudinal carinulae along midline and on gena; scape usually of moderate length (type population), but sometimes relatively long and slender (SI 88–109), scape when laid back reaching and often surpassing posterior margin of head; scape surface variable, usually with scattered piligerous punctae and some carinulae (type population), but sometimes scape more smooth with carinulae reduced, or scape more robust, with carinulae coarser; flagellum with distinct 4-segmented antennal club; mesosoma sculpture highly variable, usually completely sculptured, without large patches of smooth cuticle, but sometimes pronotum mostly to completely effaced; dorsum of promesonotum carinulate (type population), rugose, or rugoreticulate (often with punctae), almost always with longitudinal orientation (some aberrant specimens with transverse orientation); side of pronotum carinulate (type population), rugulose, or punctate; mesopleuron and side of propodeum punctate to rugulose-punctate, generally with more rugulae on the propodeum; dorsum and declivity of propodeum with transverse carinulae; promesonotum in profile low-domed, and usually slightly asymmetrical, with apex shifted anterior of midpoint, and anterior face steeper than posterior face (some populations roughly symmetrical); metanotal groove usually well-demarcated, width and depth variable (average in type population); propodeal spines tuberculate to short (PSL 0.09–0.19, PSI 1.0–1.6); petiole in profile appearing average (type population) to slightly elongate (PL/HW 0.48–0.62), with peduncle usually thick and robust; petiolar node in profile usually of moderate size (PH/PL 0.51–0.66), with a broadly rounded dorsum that points vertical to slightly posteriad, rarely pointing distinctly posteriad; node sometimes somewhat compressed anteroposteriorly; posterior margin of petiole in profile sometimes distinctly bent downwards, creating a slight concavity below node; postpetiolar node in profile similar in size to petiolar node (type population) or bulging (PPH/PH 0.79–1.05, PW/PPW 0.70–0.86), shape of node subcircular to asymmetrical (slightly asymmetrical in type population); petiole and postpetiole usually mostly punctate, with anterior faces of nodes variably smooth and shiny (type population), sometimes nodes with rugulae, or rarely rugoreticulae; gaster usually smooth and shiny, within scattered piligerous punctae, but sometimes (mostly northern populations in drier habitats) first gastral tergite and sternite lightly to strongly punctate; pilosity highly variable, pilosity on gastral dorsum usually clearly bilayered, with a layer of longer suberect setae, and a layer of shorter decumbent setae, but length and density of each layer variable, lower layer sometimes very dense (almost pubescent), or somewhat sparse and more subdecumbent, causing it to blend in with upper layer, rarely upper layer very long and more dense (type population average), gastral setae never greatly strongly thickened; setae on scapes uniformly suberect to subdecumbent, never with a separate layer of longer suberect setae; setae on legs decumbent to appressed, with longer suberect setae on femoral venters and coxae.
(7 measured) HL 0.92–1.13 (0.98), HW 0.81–1.01 (0.87), FLD 0.23–0.30 (0.25), PCW 0.07–0.10 (0.07), SL 0.81–1.00 (0.82), EL 0.23–0.27 (0.25), ACL 0.72–0.87 (0.75), ML 1.38–1.69 (1.48), PrW 0.82–1.01 (0.82), PSL 0.15–0.29 (0.21), SDL 0.13–0.16 (0.13), PL 0.52–0.68 (0.54), PH 0.28–0.36 (0.29), PW 0.23–0.32 (0.23), PPL 0.27–0.37 (0.28), PPH 0.27–0.39 (0.27), PPW 0.31–0.42 (0.32), MFL 0.98–1.30 (0.99), MTL 0.78–1.01 (0.79), CI 87–92 (89), SI 87–105 (94), REL 26–28 (28), FLI 28–30 (29), PSI 1.1–2.1 (1.6), MFI 78–95 (87), ACI1 60–66 (63), ACI2 83–91 (91).
Same as worker except for standard queen modifications and as follows (comparing queen and worker of type population form only; queen from Rancho Somecla): pronotum with transverse carinulae; mesoscutum and scutellum longitudinally carinulate; propodeum with transverse carinulae that wrap around entire surface; katepisternum mostly smooth; lower layer of setae on gastral dorsum denser; wing venation as in
See
This species is one of the largest and most conspicuous species of
One
As a preface to describing each variant of
Characteristics of the type population (
Variant 1 (
Variant 2 (
Variant 3 (
Variant 4 (
Variant 5 (
Variant 6 (
Variant 7 (
Variant 8 (
Distribution map of
Integument mostly black, or dark brown to brown; medium-sized species (see HL, ML, PrW below); basal margin of mandible usually sinuous, with a distinct basal depression, but without a basal tooth (rarely basal margin appearing nearly straight); anterior clypeal margin undulating, with 2–4 distinct to somewhat blunt teeth; face sculpture variable, usually mostly smooth and shiny, with, at most, faint carinulae extending to about ¾ the distance to posterior margin of head, but some populations with face mostly rugoreticulate, with only area around posterior margin becoming smooth; pronotum sculpture variable, usually mostly smooth and shiny, but sometimes with some longitudinal rugulae on humeri, remainder of mesosoma with rugulae and punctae; postpetiole bulging, appearing distinctly larger than petiolar node (PPH/PH 0.96–1.14; PW/PPW 0.65–0.78); eye of moderate size (EL 0.11–0.16, REL 16–22), oval-shaped, with 6–8 ommatidia at greatest diameter (rarely 9 or 10); propodeal spines usually tuberculate, or at least forming a sharp angle, but sometimes forming a blunt angle, or becoming more developed, short (PSL 0.07–0.12, PSI 1.0–2.0).
Southern Mexico to Honduras.
(23 measured) HL 0.67–0.86 (0.79), HW 0.57–0.76 (0.68), FLD 0.16–0.21 (0.19), PCW 0.02–0.0.05 (0.04), SL 0.53–0.74 (0.68), EL 0.11–0.16 (0.12), ACL 0.49–0.65 (0.59), ML 0.84–1.09 (1.02), PrW 0.39–0.50 (0.46), PSL 0.07–0.12, SDL 0.06–0.09, PL 0.29–0.38 (0.37), PH 0.19–0.24 (0.23), PW 0.15–0.19 (0.18), PPL 0.21–0.27 (0.24), PPH 0.20–0.25 (0.25), PPW 0.22–0.27 (0.26), MFL 0.57–0.82 (0.76), MTL 0.48–0.65 (0.61), CI 83–93 (85), SI 80–101 (100), REL 16–22 (18), FLI 25–30 (28), PSI 1.0–2.0 (1.1), MFI 89–108 (89), ACI1 66–70 (68), ACI2 85–96 (88).
Medium-sized species; general body color mostly black, or dark brown to brown, with appendages dark brown to orange-brown, lighter at joints and toward extremities; setae golden brown; mandible with 6–8 teeth (usually 7), consisting of 3 distinct apical teeth, a basal tooth, and 2–4 smaller teeth/denticles, which are often worn and indistinct; basal margin of mandible usually sinuous, with a distinct basal depression, but without a basal tooth, sometimes basal margin only slightly sinuous or nearly straight; mandible often mostly smooth and shiny, with scattered piligerous punctae and some striations (type population), but sometimes surface completely carinulate, with carinulae extending from base to near teeth; anterior clypeal margin forming 2–4 distinct to somewhat blunt teeth, outer teeth often larger and sharper than inner teeth (best viewed in anterodorsal view); median clypeal lobe with a pair of faint longitudinal carinulae, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth and shiny; area in between median lobe of clypeus and anterior clypeal margin forming a distinct cavity where mandibles insert; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.02–0.0.05), with sides subparallel to slightly hour-glass-shaped; frontal lobes of moderate width (FLD 0.16–0.21, FLI 25–30), not greatly covering torular lobes in full-face view; head roughly oval-shaped (CI 83–93), with posterior margin flat to slightly depressed mesad; face sculpture variable, usually mostly smooth and shiny (type population), with, at most, faint carinulae extending to about ¾ the distance to posterior margin of head, but some populations with face mostly rugoreticulate, with only area around posterior margin becoming smooth; scape of moderate length (SI 80–101), reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae; flagellum with a distinct 4-segmented antennal club; pronotum and mesonotal dorsum sculpture variable, usually mostly smooth and shiny, but sometimes with longitudinal rugulae on humeri, remainder of mesosoma mostly sculptured (contrast between smooth pronotum, and sculptured mesonotum, mesopleuron, and propodeum distinctive); mesopleuron mostly punctate, with a few rugulae; side of propodeum with rugulae and some faint punctae; dorsum and declivity of propodeum with variable number of transverse carinulae; promesonotum in profile low-domed, and roughly symmetrical, but anterior declivity sometimes more sharply angled than posterior face (type population); metanotal groove in profile well-demarcated, anterior slope often smoothly transitioning with promesonotum; propodeal spines usually tuberculate (type population), or at least forming sharp angles, but sometimes forming blunt angles, or more developed and short (PSL 0.07–0.12, PSI 1.0–2.0); petiole of moderate length (PL/HW 0.46–0.56), with peduncle thick, often with anteroventral edge sharp; petiolar node in profile somewhat small (PH/PL 0.58–0.68), wedge-shaped, with anterior face longer and more sloping than almost vertical posterior face, dorsum of node rounded, and distinctly pointed posteriad; postpetiole bulging, globular, appearing distinctly larger than petiolar node (PPH/PH 0.96–1.14; PW/PPW 0.65–0.78), anterior face of node much longer than posterior face; petiolar and postpetiolar nodes mostly smooth and shiny, remaining waist surfaces punctate, with a few rugulae around bases of nodes; gaster smooth and shiny, with scattered piligerous punctae; pilosity on face short, with a layer of sparse suberect setae, and a denser layer of decumbent setae, remainder of body dorsum with longer standing pilosity; pilosity on gastral dorsum usually distinctly bilayered, with a layer of longer suberect to subdecumbent setae, and a layer of shorter decumbent setae, both layers similar in density; setae on scape and legs mostly decumbent to appressed, with some longer suberect setae on femoral venters and coxae.
(10 measured) HL 0.73–0.84 (0.84), HW 0.67–0.76 (0.76), FLD 0.19–0.23 (0.23), PCW 0.03–0.06 (0.06), SL 0.57–0.71 (0.71), EL 0.19–0.22 (0.22), ACL 0.47–0.62 (0.62), ML 1.05–1.34 (1.34), PrW 0.56–0.74 (0.74), PSL 0.13–0.17 (0.14), SDL 0.09–0.10 (0.10), PL 0.37–0.47 (0.47), PH 0.23–0.27 (0.27), PW 0.19–0.22 (0.22), PPL 0.20–0.29 (0.27), PPH 0.24–0.30 (0.30), PPW 0.27–0.33 (0.33), MFL 0.63–0.84 (0.84), MTL 0.55–0.70 (0.70), CI 91–93 (91), SI 82–94 (94), REL 26–30 (29), FLI 27–30 (30), PSI 1.4–1.7 (1.4), MFI 90–108 (90), ACI1 64–68 (65), ACI2 76–93 (87).
Same as worker except for standard queen modifications and as follows (comparison between worker and queen of type population form only): facial sculpture usually more developed (carinulae/rugulae longer and more distinct); mesopleuron mostly smooth and shiny; propodeum with transverse carinulae that wrap around entire surface; lower layer of gastral setae more dense, almost pubescent; propodeal spines (compared to worker from same population) slightly longer, more developed (PSL 0.13–0.17, PSI 1.4–1.7).
Unknown.
The presence of a bulging postpetiole, a sinuous basal margin of the mandible, and a toothed anterior clypeal margin, makes
Distinctive features of the type population form (
Variant 1 (
Variant 2 (
Variant 3 (
Distribution map of
Integument mostly black to red-black; medium- to large-sized species (see HL, ML, PrW below); lateral apex of hypostomal bridge projecting ventrally as a subquadrate to broadly rounded lobe, visible behind base of mandible in profile view; propodeal spines tuberculate (PSL 0.10–0.15, PSI 1.0–1.2); face and mesosoma usually completely sculptured with rugoreticulae and some carinulae (rarely with sculpture reduced); eye of moderate to large size (EL 0.14–0.19, REL 15–20), oval-shaped, with 8–11 ommatidia in greatest diameter; postpetiole usually appearing smaller than petiolar node (PPH/PH 0.85–0.98); basal margin of mandible straight to slightly sinuous; anterior clypeal margin with a median emargination.
Southern Mexico to Nicaragua.
(12 measured) HL 0.93–1.15 (1.07), HW 0.80–1.08 (0.98), FLD 0.22–0.29 (0.25), PCW 0.05–0.10 (0.07), SL 0.79–0.98 (0.91), EL 0.14–0.19 (0.17), ACL 0.70–0.86 (0.78), ML 1.18–1.48 (1.36), PrW 0.53–0.69 (0.63), PSL 0.10–0.15 (0.12), SDL 0.09–0.14 (0.11), PL 0.40–0.54 (0.45), PH 0.25–0.32 (0.30), PW 0.18–0.25 (0.23), PPL 0.23–0.33 (0.26), PPH 0.21–0.31 (0.26), PPW 0.23–0.30 (0.28), MFL 0.97–1.23 (1.14), MTL 0.72–0.93 (0.85), CI 86–96 (91), SI 89–100 (93), REL 15–20 (18), FLI 25–28 (26), PSI 1.0–1.2 (1.1), MFI 82–90 (86), ACI1 61–63 (62), ACI2 84–89 (86).
Medium- to large-sized species; general body color black to red-black, with appendages black or dark brown to orange-brown, becoming lighter toward extremities at joints; setae dark golden brown to golden brown; mandible with 6 teeth, 2 teeth nearest basal tooth sometimes worn and indistinct; basal margin of mandible straight to slightly sinuous, without a basal depression or notch; mandible mostly smooth, with scattered piligerous punctae and a few striations near base and on lateral surface; anterior clypeal margin with a simple median emargination; median lobe of clypeus usually with a pair of longitudinal carinulae (type population), but sometimes surface smooth, or with several irregular carinulae, area in between carinulae often depressed; apex of lobe usually with a single transverse short carinula (type population), but sometimes with several transverse carinulae/rugulae; remainder of clypeus smooth and shiny; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.05–0.10), sides subparallel; frontal lobes of moderate width (FLD 0.22–0.29; FLI 25–28) not greatly obscuring torular lobes in full-face view; lateral margin of hypostomal bridge with a projecting subquadrate lobe, visible in profile view (very rarely reduced and only visible from a lateroventral view); head roughly oval-shaped to more robust, becoming strongly heart-shaped (CI 86–96), posterior margin of head with a distinct median depression (especially in larger specimens); eye moderately large (EL 0.14–0.19, REL 15–20), oval-shaped, with 8–11 ommatidia at greatest diameter; face sculpture variable, usually strongly rugoreticulate, with some carinulae along midline (type population), but sometimes sculpture more polished, with area near posterior margin of head becoming somewhat smooth; scape of moderate length (SI 89–100), reaching posterior margin of head when laid back; scape usually thick and robust, surface carinate to carinulae with piligerous punctae; flagellum with a somewhat indistinct 4-segmented antennal club; mesosoma sculpture variable, often densely rugose, with rugae wavy, almost becoming reticulate (type population), but sculpture often reduced, with promesonotum longitudinally carinulate to nearly smooth, and side of propodeum and mesopleuron more punctate; promesonotum in profile domed, and usually asymmetrical (type population), with apex shifted anterior of midpoint, but sometimes promesonotum symmetrical; metanotal groove well-demarcated, sometimes somewhat wide and with metanotum forming a small welt (type population); propodeal spines tuberculate (PSL 0.10–0.15, PSI 1.0–1.2); petiole of moderate length, usually somewhat robust (PL/HW 0.45–0.52); petiolar node in profile of moderate height (PH/PL 0.59–0.68), dorsum usually broadly rounded and pointing vertically, only rarely forming a sharp apex; postpetiole in profile usually appearing smaller than petiolar node (PPH/PH 0.85–0.98); petiole and postpetiole usually strongly punctate, with a few rugulae on dorsum and anterior face of postpetiolar node (type population), sometimes anterior faces of petiolar and petiolar nodes mostly smooth; gaster mostly smooth, with scattered piligerous punctae; most of body dorsum with relatively long standing pilosity; pilosity on gastral dorsum sparse and mostly suberect (type population), usually with some shorter subdecumbent setae, suberect setae often somewhat stout; setae on promesonotum often noticeably erect; setae on scape uniformly suberect to decumbent; setae on legs mostly decumbent to appressed, with some suberect setae on femoral venters and coxae.
(5 measured) HL 1.01–1.12 (1.11), HW 0.92–1.03 (0.99), FLD 0.27–0.31 (0.31), PCW 0.07–0.10 (0.09), SL 0.87–0.97 (0.95), EL 0.26–0.28 (0.28), ACL 0.76–0.83 (0.83), ML 1.55–1.72 (1.71), PrW 0.93–1.04 (0.99), PSL 0.20–0.23 (0.20), SDL 0.15–0.17 (0.16), PL 0.57–0.60 (0.60), PH 0.33–0.39 (0.36), PW 0.26–0.31 (0.30), PPL 0.30–0.34 (0.34), PPH 0.32–0.39 (0.38), PPW 0.35–0.40 (0.38), MFL 1.13–1.26 (1.26), MTL 0.86–0.96 (0.93), CI 89–92 (89), SI 93–96 (96), REL 28–29 (28), FLI 28–31 (31), PSI 1.2–1.4 (1.2), MFI 78–84 (78), ACI1 60–62 (60), ACI2 83–89 (87).
Same as worker except for standard queen modifications and as follows: pronotum with transverse rugulae; mesoscutum longitudinally costulate; scutellum rugose to rugoreticulate; propodeum rugose to rugoreticulate; postpetiolar node larger, and somewhat anteroposteriorly compressed; mesopleuron mostly smooth; setae on mesoscutum short; setae on gastral dorsum often more dense, and more strongly bilayered, with layer of decumbent setae more dense; anterior quarter of gastral dorsum often with a layer of pubescence under stouter suberect and decumbent setae; wing venation as in
See
The type population was found nesting under rocks along a stream in mesophyll forest. This population was rather intriguing, because the nests were common and foragers were very abundant. In fact,
Nests of this species are large for
As described above under
Within my concept of
Distribution map of
Integument mostly mottled brown to red-brown or orange-brown; medium-sized species (see HL, ML, PrW below); basal margin of mandible with a deep proximal notch containing a small tooth; anterior clypeal margin with a deep unevenly rounded depression that, when viewed from an anterodorsal angle, has two distinct outer teeth and two indistinct inner teeth (all pointing ventrally); mesosoma profile appearing elongate, almost gracile; metanotal grove relatively wide and shallow, not well demarcated by the promesonotal and propodeal margins; propodeal dorsum elongate, distinctly longer than propodeal declivity; gastral pilosity clearly bilayered, with a layer of long suberect setae and a layer of short subdecumbent setae; eye relatively small (EL 0.09–0.11, REL 13–15), with 5–6 ommatidia at greatest diameter; scape and metafemur relatively long (SI 104–120, MFI 70–80); frontal lobes well-developed (FLD 0.16–0.19, FLI 23–25), but not completely covering torular lobes in full-face view; 4-segmented antennal club very distinct.
Honduras to Nicaragua.
(5 measured) HL 0.79–0.94 (0.91), HW 0.63–0.82 (0.78), FLD 0.16–0.19 (0.17), PCW 0.03–0.03 (0.03), SL 0.76–0.86 (0.83), EL 0.09–0.11 (0.11), ACL 0.75–0.80 (0.78), ML 1.03–1.21 (1.15), PrW 0.46–0.55 (0.52), PSL 0.12–0.15 (0.15), SDL 0.11–0.12 (0.11), PL 0.38–0.42 (0.42), PH 0.19–0.23 (0.23), PW 0.14–0.17 (0.15), PPL 0.19–0.20 (0.20), PPH 0.17–0.21 (0.20), PPW 0.18–0.22 (0.21), MFL 0.90–1.03 (1.10), MTL 0.71–0.83 (0.82), CI 80–87 (85), SI 104–120 (107), REL 13–15 (14), FLI 23–25 (23), PSI 1.1–1.3 (1.3), MFI 70–80 (77), ACI1 62–64 (63), ACI2 93–98 (95).
Medium-sized species; general body color a mottled brown to red-brown or orange-brown, with patches of yellow-brown, especially on gaster; appendages generally lighter, orange-brown to yellow-brown; setae golden brown; mandible with 2–3 distinct apical teeth, 1–2 distinct basal teeth, and a variable number of almost imperceptible denticles in between (usually appearing as a long, flat diastema), basal tooth well-defined and robust; basal margin of mandible with a very deep proximal notch containing a small tooth; mandible mostly smooth and shiny, with scattered piligerous punctae and a variable number of striae; anterior clypeal margin with a deep unevenly rounded depression, that, when viewed anteriorly, has two distinct outer teeth and two indistinct inner teeth (all pointing ventrally); median lobe of clypeus somewhat obliquely flattened and concave, without a pair of longitudinal carinulae, only sometimes with a short transverse carinula before anterior margin, remainder of clypeus mostly smooth and shining; posterior extension of clypeus between antennal insertions somewhat narrow (PCW 0.03), sides subparallel; frontal lobes well-developed (FLD 0.16–0.19, FLI 23–25), but not completely covering torular lobes in full-face view; head elongate and roughly oval-shaped (CI 80–87), with posterior margin depressed medially; eye relatively small (EL 0.09–0.11, REL 13–15), subcircular to oval-shaped, with 5–6 ommatidia at greatest diameter; face densely rugoreticulate, with a few longitudinal rugulae medially, interstices with faint piligerous punctae; scape relatively long (SI 104–120), surpassing posterior margin of head when laid back; scape cuticle mostly smooth and shiny, with scattered piligerous punctae and fine striations; flagellum with a very distinct 4-segmented antennal club; mesosoma densely sculptured, promesonotal dorsum mostly with longitudinal rugae and rugoreticulae, but a few transverse rugulae present around anterior declivity; lateral side of mesosoma with irregular rugulae and punctae; propodeal declivity mostly smooth and shining, sometimes with faint transverse striae; promesonotum in profile low-domed and slightly asymmetrical, with the apex occurring anterior of the midpoint, anterior face more strongly defined and longer than posterior face; metanotal grove relatively wide and shallow, not well demarcated by the promesonotal and propodeal margins; propodeal dorsum elongate, distinctly longer than propodeal declivity; mesosomal outline in profile appearing elongate, almost gracile; propodeal spines forming sharp tubercles (PSL 0.12–0.15, PSI 1.1–1.3); petiole somewhat elongate and gracile (PL/HW 0.51–0.60), node small (PH/PL 0.53–0.56), with a long sloping anterior face, and a short vertical posterior face, dorsal surface in profile view rounded, pointing vertically; postpetiolar node slightly smaller than petiolar node (PPH/PH 0.90–0.97), anterior face longer and more sloping angled than posterior face; petiole and postpetiole mostly punctate, with anterior face of postpetiolar node largely smooth and shiny; gaster mostly smooth and shiny, with scattered piligerous punctae; much of body with long suberect to subdecumbent pilosity; scapes with mostly subdecumbent setae; gastral pilosity somewhat sparse and clearly bilayered, with a layer of long suberect setae and a layer of short subdecumbent setae; setae on legs mostly subdecumbent to appressed, with suberect setae on ventral surface of profemur.
Unknown.
Unknown.
Distribution map of
Integument mostly black or dark brown to brown; small- to medium-sized species (see HL, ML, PrW below); lateral margin of hypostomal bridge with a projecting subquadrate lobe, visible behind mandible in profile view; waist segments appearing somewhat slender, with postpetiole particularly small; eye relatively large (EL 0.13–0.16, REL 19–22), with 6–9 ommatidia at greatest diameter; anterior clypeal margin with a median emargination; basal margin of mandible straight; propodeal spines tuberculate (PSL 0.08–0.11, PSI 1.1–1.4); frontal lobes well-developed, but not completely covering torular lobes in full-face view (FLD 0.16–0.19, FLI 24–28); head densely sculptured with carinulae, rugoreticulae, and faint punctae; mesosoma mostly sculptured, but pronotal sculpture variably effaced, sometimes mostly smooth and shiny; gastral setae mostly suberect, long, and relatively sparse.
Honduras to Nicaragua.
(10 measured) HL 0.72–0.83 (0.74), HW 0.63–0.75 (0.67), FLD 0.16–0.19 (0.17), PCW 0.03–0.06 (0.04), SL 0.57–0.67 (0.59), EL 0.13–0.16 (0.13), ACL 0.56–0.62 (0.56), ML 0.89–1.08 (0.90), PrW 0.43–0.50 (0.44), PSL 0.08–0.11 (0.09), SDL 0.07–0.09 (0.08), PL 0.31–0.42 (0.32), PH 0.19–0.24 (0.19), PW 0.13–0.19 (0.15), PPL 0.18–0.24 (0.18), PPH 0.14–0.20 (0.15), PPW 0.17–0.24 (0.18), MFL 0.68–0.79 (0.70), MTL 0.54–0.65 (0.56), CI 87–90 (90), SI 88–95 (88), REL 19–22 (19), FLI 24–28 (25), PSI 1.1–1.4 (1.2), MFI 90–98 (96), ACI2 63–66 (65), ACI2 92–99 (96).
Small- to medium-sized species; general body color black or dark brown to brown, with appendages brown to yellow-brown; setae golden brown; mandible with 6 teeth, but two teeth nearest the basal tooth smaller, often worn and indistinct or denticulate; basal margin of mandible straight, without basal notch or depression; mandible surface mostly smooth and shiny, with scattered piligerous punctae and basal striae; anterior clypeal margin with a median emargination; median lobe of clypeus with a few faint punctae, but lacking longitudinal carinulae, apex with a faint transverse carinula; remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.06), sides subparallel; frontal lobes well-developed (FLD 0.16–0.19, FLI 24–28), but not completely obscuring torular lobes in full-face view; frontal carinae weak, fading into facial sculpture beyond frontal lobes, not surpassing level of anterior margin of eye; head oval-shaped (CI 87–90), posterior margin slightly depressed medially; eye relatively large (EL 0.13–0.16, REL 19–22), oval-shaped, with 6–9 ommatidia at greatest diameter; face densely sculptured with longitudinal carinulae along midline, turning into rugulae and rugoreticulae toward lateral margins, interstices punctate; scape of moderate length (SI 88–95), nearly reaching posterior margin of head when laid back; scape surface with a variable amount of fine punctae and striae; flagellum with distinct 4-segmented antennal club; pronotal sculpture mostly smooth and shiny, or with dense longitudinal rugulae and faint punctae, or with intermediate states with rugulae somewhat effaced (especially on pronotal side); mesonotal dorsum with fine carinulae and punctae; mesopleuron mostly punctate, with a few rugulae; propodeal side with rugulae and some punctae; propodeal dorsum and declivity with transverse carinulae; promesonotum in profile usually low-domed and roughly symmetrical, some specimens with promesonotum more robust, higher; metanotal grove well demarcated, of moderate depth and width; propodeal spines present as small tubercles (PSL 0.08–0.11, PSI 1.1–1.4); petiole and postpetiole appearing somewhat delicate with nodes relatively small (PL/HW 0.48–0.57, PH/PL 0.56–0.63, PPH/PH 0.74–0.84); petiolar node in profile pointing posteriad, with dorsum gently rounded; ventral surface of node concave; postpetiole in profile with a long shield-like anterior face and a short vertical posterior face, anterior half constricted; anterior faces of petiolar and postpetiolar nodes usually mostly smooth and shiny, remaining surface of waist segments mostly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae and a few short striae around anterior constriction; most of body dorsum with moderately long standing pilosity; setae on scape suberect to appressed; gastral setae mostly suberect, long, and relatively sparse, a few shorter decumbent setae also present; setae on legs mostly subdecumbent to appressed, with suberect setae on coxae and femoral venters.
(5 measured) HL 0.76–0.79 (0.79), HW 0.68–0.72 (0.72), FLD 0.19–0.21 (0.20), PCW 0.04–0.06 (0.05), SL 0.62–0.63 (0.63), EL 0.21–0.22 (0.22), ACL 0.59–0.61 (0.59), ML 1.11–1.19 (1.18), PrW 0.63–0.67 (0.67), PSL 0.12–0.13 (0.13), SDL 0.10–0.12 (0.12), PL 0.42–0.45 (0.44), PH 0.23–0.25 (0.25), PW 0.17–0.19 (0.18), PPL 0.21–0.24 (0.22), PPH 0.20–0.23 (0.20), PPW 0.24–0.25 (0.25), MFL 0.74–0.79 (0.77), MTL 0.61–0.65 (0.63), CI 89–91 (91), SI 89–91 (89), REL 30–31 (30), FLI 28–29 (28), PSI 1.1–1.3 (1.1), MFI 90–93 (93), ACI1 63–65 (65), ACI2 94–98 (94).
Same as worker except for standard queen modifications and as follows: pronotum transversely carinulate/rugulose; mesoscutum and scutellum densely carinulate; mesopleuron mostly smooth and shiny; propodeum transversely carinulate, with carinulae wrapping around entire surface; petiole and postpetiole slightly more robust, with dorsum of petiolar node reaching a slightly more defined apex; gastral pilosity more dense.
Unknown.
An interesting phenomenon in
Distribution map of
Integument mostly dark red-brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, with a small basal notch and accompanying tooth; anterior clypeal margin viewed at anterodorsal angle forming 4 blunt teeth (only outer teeth visible in full-face view); pilosity on gastral tergites mostly forming a moderately sparse layer of thickened suberect setae, with only a few thin decumbent setae present; face mostly rugoreticulate; mesosoma mostly rugulose and punctate, without large areas of smooth cuticle; eye of small to moderate size (EL 0.08–0.10, REL 15–18), oval-shaped, with 4–5 ommatidia at greatest diameter; propodeal spines tuberculate (PSL 0.07–0.09, PSI 1.2–1.4); frontal lobes somewhat expanded (FLD 0.14–0.16, FLI 28–31), but not completely obscuring torular lobes in full-face view.
Honduras.
(10 measured) HL 0.56–0.63 (0.61), HW 0.49–0.56 (0.54), FLD 0.14–0.16 (0.16), PCW 0.03 (0.03), SL 0.48–0.53 (0.51), EL 0.08–0.10 (0.10), ACL 0.48–0.53 (0.53), ML 0.68–0.76 (0.74), PrW 0.35–0.40 (0.38), PSL 0.07–0.09 (0.09), SDL 0.05–0.07 (0.07), PL 0.25–0.28 (0.27), PH 0.14–0.17 (0.15), PW 0.11–0.14 (0.13), PPL 0.13–0.16 (0.15), PPH 0.12–0.15 (0.14), PPW 0.14–0.16 (0.15), MFL 0.48–0.55 (0.55), MTL 0.39–0.44 (0.43), CI 87–92 (89), SI 90–99 (95), REL 15–18 (18), FLI 28–31 (30), PSI 1.2–1.4 (1.3), MFI 97–104 (98), ACI1 65–69 (66), ACI2 96–103 (103).
Small-sized species; general body color dark red-brown to brown, with appendages lighter, brown to yellow-brown toward extremities; setae golden brown; mandible with 6 teeth, basal tooth often well-defined, projecting; basal margin of mandible sinuous, with a distinct basal notch and accompanying small tooth; mandible mostly smooth, except for scattered piligerous punctae, and some striae around the basal and along lateral surface; anterior clypeal margin in anterodorsal view undulating, forming 4 blunt teeth (only outer teeth visible in full-face view); median lobe of clypeus with a pair of somewhat distinct longitudinal carinulae, that diverge toward anterior margin, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth; posterior extension of clypeus between antennal insertions somewhat narrow (FLD 0.14–0.16, FLI 28–31), with sides diverging slightly posteriad; frontal lobes somewhat expanded (FLD 0.14–0.16, FLI 28–31), but not completely obscuring torular lobes in full-face view; head roughly oval-shaped (CI 87–92), with posterior margin distinctly depressed medially; eye of small to moderate size (EL 0.08–0.10, REL 15–18), oval-shaped, with 4–5 ommatidia at greatest diameter; face mostly strongly rugoreticulate, with a few longitudinal carinulae along midline near frontal lobes; scape of moderate length (SI 90–99), not quite reaching posterior margin when laid back; scape surface mostly smooth, with faint striations and piligerous punctae; flagellum with a distinct 4-segmented antennal club; dorsum of pronotum with irregular longitudinal rugulae that sometimes merge, dorsum of metanotum more rugoreticulate-punctate, side of pronotum mostly carinulate, with a small patch of smooth cuticle near ventral margin, mesopleuron and side of propodeum rugulose-punctate, propodeal dorsum and upper half of declivity with transverse carinulae; promesonotum in profile low-domed, roughly symmetrical, but anterior face distinctly longer than posterior face; metanotal groove of moderate depth and width, sometimes with a small central welt (metanotum); propodeal spines tuberculate (PSL 0.07–0.09, PSI 1.2–1.4); petiole in profile of moderate length (PL/HW 0.48–0.54), peduncle somewhat slender, narrowing toward body; petiolar node of moderate height (PH/PL 0.52-0.61), asymmetrical, with anterior face slightly longer and more sloping than posterior face, dorsum of node narrowly rounded, almost becoming a well-defined apex, node always pointing distinctly posteriad; postpetiole in profile subspherical to oblong, appearing somewhat compressed anteroposteriorly, roughly symmetrical, and similar in size to slightly smaller than petiolar node (PPH/PH 0.83–0.99); anterior faces of petiolar and postpetiolar nodes mostly smooth, remaining surfaces mostly punctate, with some rugulae on posterior half of postpetiole; gaster mostly smooth and shiny, with scattered piligerous punctae; face pilosity short, bilayered; dorsum of mesosoma and gaster mostly with a sparse layer of thickened, suberect setae, gaster with a few thin decumbent setae; setae on scapes and legs mostly appressed, with some suberect setae on femoral venters and coxae.
(5 measured) HL 0.57–0.63 (0.62), HW 0.51–0.58 (0.57), FLD 0.16–0.18 (0.17), PCW 0.03–0.04 (0.04), SL 0.48–0.52 (0.52), EL 0.14–0.16 (0.16), ACL 0.48–0.52 (0.51), ML 0.75–0.85 (0.85), PrW 0.43–0.49 (0.49), PSL 0.09–0.11 (0.11), SDL 0.07–0.09 (0.08), PL 0.27–0.32 (0.31), PH 0.16–0.17 (0.17), PW 0.12–0.15 (0.15), PPL 0.14–0.15 (0.15), PPH 0.14–0.17 (0.17), PPW 0.15–0.18 (0.18), MFL 0.50–0.56 (0.56), MTL 0.40–0.45 (0.45), CI 90–92 (92), SI 90–97 (91), REL 26–28 (28), FLI 31–32 (31), PSI 1.2–1.4 (1.4), MFI 99–104 (102), ACI1 65, ACI2 99–101 (100).
Same as worker except for standard queen modifications and as follows: pronotum transversely carinulate, with dorsum of cervical shield punctate; mesoscutum longitudinally rugulose, with some irregular reticulations; scutellum longitudinally rugulose to rugoreticulate, usually with a small patch of smooth cuticle mesad; propodeum with transverse carinulae that wrap around surface; mesopleuron mostly smooth; pilosity on gaster distinctly bilayered, with a stout layer of long suberect to subdecumbent setae, and a very dense layer of decumbent to appressed pubescence; wing venation as in
See
This species can be separated from similar forms by its mandible structure, eye size, gastral pilosity, facial sculpture, and overall size. Geography is also useful, as
Given the complexity of the
Distribution map of
Integument mostly dark red-brown (almost black) to brown; small- to medium-sized species (see HL, ML, PrW below); anterior clypeal margin entire, or at most with a nearly imperceptible median notch (only visible from anterodorsal angle); basal margin of mandible straight; face densely sculptured, mostly rugoreticulate; dorsum of promesonotum rugoreticulate, or with many irregular rugulae; pilosity on gastral tergites indistinctly bilayered, forming a somewhat sparse layer of long suberect setae, and a very sparse layer of short decumbent setae; eye of small to moderate size (EL 0.09–0.11, REL 15–19), oval-shaped, with 4–7 ommatidia at greatest diameter; propodeal spines reduced to small triangular tubercles (PSL 0.08–0.14, PSI 1.1–1.4); frontal lobes of moderate width (FLD 0.14–0.19, FLI 25–29).
Southern Mexico to Guatemala.
(17 measured) HL 0.57–0.76 (0.61), HW 0.51–0.68 (0.57), FLD 0.14–0.19 (0.15), PCW 0.03–0.07 (0.03), SL 0.46–0.64 (0.50), EL 0.09–0.11 (0.09), ACL 0.45–0.59 (0.49), ML 0.68–1.00 (0.78), PrW 0.36–0.49 (0.40), PSL 0.08–0.14 (0.11), SDL 0.06–0.11 (0.09), PL 0.26–0.40 (0.31), PH 0.16–0.23 (0.19), PW 0.14–0.19 (0.16), PPL 0.12–0.22 (0.15), PPH 0.13–0.21 (0.16), PPW 0.16–0.22 (0.18), MFL 0.50–0.72 (0.55), MTL 0.41–0.57 (0.45), CI 86–92 (92), SI 88–97 (89), REL 15–19 (16), FLI 25–29 (26), PSI 1.1–1.4 (1.3), MFI 93–104 (103), ACI1 65–69 (67), ACI2 92–99 (97).
Small- to medium-sized species; general body color dark red-brown (almost black) to brown, with appendages lighter, brown to yellow-brown toward extremities; setae golden; mandible with 6–7 teeth (usually 7), consisting of 3 larger, more distinct apical teeth, a basal tooth, and 2–3 smaller denticulate teeth, which are usually more worn, and sometimes indistinct; basal margin of mandible straight, without a basal notch or depression; mandible mostly smooth, except for scattered piligerous punctae, and a few basal striae; anterior clypeal margin entire, or at most with a nearly imperceptible median notch, only visible from anterodorsal view; median lobe of clypeus with faint longitudinal carinulae and some punctations, apex of lobe with a few short transverse carinulae, remainder of clypeus mostly smooth; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.07), with sides subparallel to slightly hour-glass-shaped; frontal lobes of moderate width (FLD 0.14–0.19, FLI 25–29), not obscuring torular lobes in full-face view; head roughly oval-shaped (CI 86–92), posterior margin slightly depressed medially; eye of small to moderate size (EL 0.09–0.11, REL 15–19), oval-shaped, with 4–7 ommatidia at greatest diameter; face densely sculptured, mostly rugoreticulate, with a few longitudinal rugae along midline near frontal lobes; scape of moderate length (SI 88–97), almost reaching posterior margin of head when laid back; scape surface mostly smooth, with scattered piligerous punctae; flagellum with a distinct 4-segmented antennal club; most of mesosoma densely sculptured, dorsum of promesonotum rugoreticulate, or with irregular longitudinal rugae that only sometimes merge, side of pronotum variably rugulose-punctate, sometimes with a large patch of smooth cuticle, mesopleuron mostly punctate, with a few rugulae, side of propodeum rugulose-punctate, propodeal dorsum and upper half of declivity with a few transverse carinulae; promesonotum in profile low-domed, roughly symmetrical, sometimes dorsum slightly flattened; metanotal groove well-demarcated, moderate; propodeal spines tuberculate, often forming somewhat broad triangles (PSL 0.08–0.14, PSI 1.1–1.4); petiole appearing of moderate length (PL/HW 0.50–0.59), peduncle usually somewhat slender; petiolar node of moderate height to somewhat small (PH/PL 0.54–0.65), dorsum broadly rounded, and always pointing distinctly posteriad, area underneath node usually with a small concavity, or notch; postpetiole in profile subspherical to oblong, appearing slightly compressed anteroposteriorly; anterior faces of petiolar and postpetiolar nodes smooth, remaining surfaces mostly punctate, with a few rugulae on posterior half of postpetiole; gaster smooth, with scattered piligerous punctae; face with a mostly uniform layer of short decumbent setae; remainder of body dorsum with longer suberect setae; pilosity on gastral tergites indistinctly bilayered, with a relatively sparse layer of longer suberect to subdecumbent setae, and a sparser layer of decumbent to appressed setae; setae on scapes and legs mostly decumbent to appressed, with some suberect setae on femoral venters and coxae.
(5 measured) HL 0.61–0.76 (0.61), HW 0.56–0.70 (0.57), FLD 0.15–0.21 (0.16), PCW 0.04–0.08 (0.04), SL 0.50–0.64 (0.50), EL 0.15–0.20 (0.15), ACL 0.48–0.61 (0.49), ML 0.84–1.11 (0.86), PrW 0.49–0.63 (0.49), PSL 0.11–0.16 (0.14), SDL 0.09–0.12 (0.10), PL 0.33–0.44 (0.34), PH 0.19–0.25 (0.19), PW 0.17–0.22 (0.18), PPL 0.15–0.22 (0.18), PPH 0.19–0.25 (0.19), PPW 0.18–0.27 (0.21), MFL 0.55–0.73 (0.55), MTL 0.45–0.58 (0.45), CI 89–94 (94), SI 86–96 (86), REL 26–28 (26), FLI 27–31 (27), PSI 1.3–1.4 (1.4), MFI 93–104 (104), ACI1 64–67 (66), ACI2 94–98 (98).
Same as worker except for standard queen modifications and as follows: side of pronotum rugoreticulate to transversely rugose, center of pronotum transversely carinulate; mesoscutum mostly foveolate to longitudinally rugose, usually with some smooth cuticle along midline, foveolae when present piligerous; propodeum with transverse carinulae that wrap around surface; mesopleuron mostly smooth; pilosity on gastral tergites distinctly bilayered, with a sparse layer of longer suberect to subdecumbent setae and a dense layer of decumbent setae, lower layer usually almost pubescent, but sometimes less dense and longer; wing venation as in
Unknown.
This species is known only from extracts of sifted leaf litter collected from the forest floor. It occupies a variety of wet forest habitat types (e.g. rainforest, mesophyll forest, 2° wet forest,
There is some variation among populations of
Integument mostly dark red-brown to orange-brown, with appendages a distinctly lighter, uniform yellow-brown; small- to medium-sized species (see HL, ML, PrW below); anterior clypeal margin undulating, usually with 4 sharp to blunt teeth, outer teeth often larger and sharper; basal margin of mandible usually sinuous, with a shallow to very deep basal depression, but without a basal tooth; gastral pilosity usually appearing somewhat dense, with setae subdecumbent to decumbent and not obviously bilayered, but sometimes more clearly bilayered, with subdecumbent setae becoming suberect; head completely sculptured, mostly rugoreticulate; mesosoma usually mostly sculptured with rugae and punctate, but pronotum sometimes with sculpture largely effaced; petiole of moderate length, somewhat robust; eye relatively small (EL 0.07–0.12, REL 12–17), subcircular to oval-shaped, with 4–6 ommatidia at greatest diameter; propodeal spines tuberculate (PSL 0.07–0.14, PSI 1.0–1.4).
Southern Mexico to Honduras.
(20 measured) HL 0.63–0.83 (0.71), HW 0.54–0.73 (0.60), FLD 0.14–0.20 (0.15), PCW 0.02–0.03 (0.02), SL 0.47–0.69 (0.55), EL 0.07–0.12 (0.08), ACL 0.48–0.66 (0.56), ML 0.76–1.09 (0.86), PrW 0.37–0.50 (0.41), PSL 0.07–0.14 (0.10), SDL 0.06–0.10 (0.08), PL 0.28–0.41 (0.33), PH 0.16–0.22 (0.18), PW 0.13-0.18 (0.14), PPL 0.14–0.21 (0.16), PPH 0.14–0.20 (0.16), PPW 0.16–0.22 (0.17), MFL 0.51–0.77 (0.60), MTL 0.43–0.63 (0.50), CI 83–88 (85), SI 82–95 (92), REL 12–17 (13), FLI 25–31 (26), PSI 1.0–1.4 (1.2), MFI 94–110 (100), ACI1 65–70 (69), ACI2 92–104 (101).
Small- to medium-sized species; general body color mostly dark red-brown (almost black) to orange-brown, with head and pronotum darker, and gaster with lighter patches of brown; appendages mostly a uniform and distinctly lighter orange-brown to yellow-brown; setae golden brown; mandible usually with 6–7 teeth (usually 6), but sometimes with 1–3 additional small denticles, middle 2–3 teeth nearest basal tooth often worn and indistinct; basal margin of mandible usually sinuous (rarely straight), with a shallow to very deep basal depression, but without a basal tooth; mandible mostly smooth and shining, with scattered piligerous punctae and striae (mostly on lateral side); anterior clypeal margin usually with 4 sharp to blunt teeth, outer teeth often larger and sharper, sometimes teeth reduced (especially inner two teeth), almost absent, and clypeal margin strongly emarginate; median clypeal lobe with a pair of very faint longitudinal carinulae, and a short transverse carinula near apex, remainder of clypeus smooth and shiny; posterior extension of the clypeus between the antennal insertions narrow (PCW 0.02–0.03), with sides subparallel; frontal lobes usually of moderate width, but somewhat expanded in Cusuco population (FLD 0.14–0.20, FLI 25–31); torular lobes not completely covered in full-face view; frontal carinae blending into head sculpture, not extending beyond about midpoint level of eye; head subrectangular to oval-shaped (CI 83–88), with posterior margin flat to slightly depressed medially; face mostly rugoreticulate, with a few longitudinal rugae along the midline, sometimes sculpture more polished and posterior quarter of head becoming smooth and shiny, interstices faintly punctate; scape of moderate length (SI 82–95), usually not reaching posterior margin in full-face view; scape surface mostly smooth and shiny, with scattered piligerous punctae; funiculus with a somewhat distinct 4-segmented antennal club (ACI2 65–70); pronotum sculpture varying from nearly completely smooth and shiny to strongly rugose (dorsum) and punctate or rugulose (side), with rugae mostly longitudinal in orientation; mesopleuron and propodeal side mostly punctate, with a few rugulae; propodeal dorsum and declivity (to a lesser degree) with punctae and transverse carinulae; promesonotum in profile low-domed, roughly symmetrical; location of promesonotal suture obscured to well-defined, depending on degree of pronotal sculpturing; metanotal grove well-demarcated, of moderate width and depth; propodeal spines tuberculate (PSL 0.07–0.14, PSI 1.0–1.4); petiole of average length (PL/HW 0.51–0.57); petiolar node of moderate to small size (PH/PL 0.53–0.62) and variable in shape, being either wedge shaped and strongly asymmetrical, or subconical, with the apex gently rounded and pointing only slightly posteriad; postpetiole in profile average-looking, about the same size as petiolar node (PPH/PH 0.83–0.96); anterior faces of petiolar nodes smooth and shiny, remaining surfaces of waist mostly punctate, with a few rugulae around nodes; gaster mostly smooth and shiny, with scattered piligerous punctate and a ring of short striae around the anterior constriction; most of body dorsum with somewhat short standing pilosity; scape decumbent to appressed; gastral pilosity variable, usually appearing somewhat dense, with setae subdecumbent to decumbent and not obviously bilayered, but sometimes more clearly bilayered, with subdecumbent setae becoming suberect; setae on legs decumbent to appressed, with suberect to subdecumbent setae on coxae and profemur venter.
(5 measured) HL 0.72–0.85 (0.75), HW 0.65–0.76 (0.67), FLD 0.17–0.21 (0.17), PCW 0.03–0.05 (0.05), SL 0.53–0.66 (0.59), EL 0.17–0.20 (0.17), ACL 0.55–0.62 (0.57), ML 1.01–1.26 (1.06), PrW 0.55–0.73 (0.57), PSL 0.13–0.17 (0.15), SDL 0.08–0.12 (0.10), PL 0.37–0.48 (0.39), PH 0.20–0.25 (0.21), PW 0.17–0.22 (0.18), PPL 0.18–0.22 (0.19), PPH 0.19–0.26 (0.20), PPW 0.21–0.29 (0.22), MFL 0.60–0.80 (0.65), MTL 0.53–0.69 (0.56), CI 88–90 (90), SI 82–89 (88), REL 25–28 (25), FLI 26––28 (26), PSI 1.5–1.7 (1.5), MFI 95–107 (103), ACI1 64–68 (67), ACI2 93–106 (96).
Same as worker except for standard queen modifications and as follows: pronotum transversely rugose laterad, becoming smooth mesad; mesoscutum ranging from mostly smooth, with some faint foveolae, to mostly carinulate/rugose (strength of sculpture correlates with pronotum sculpture of worker); scutellum longitudinally rugose, often with a central patch of smooth cuticle; most of katepisternum and sometimes lower half of anepisternum smooth; propodeal spines slightly more developed; setae on mesoscutum subdecumbent to appressed, more dense; pilosity in general slightly more dense; wing venation in
Unknown.
The most distinctive feature of this species is its light colored legs, which always make it stand out from other
Across its range there is significant variation in body size, surface sculpture, gastral pilosity and the configuration of the clypeus and basal margin of the mandible. I describe several variants here, but it should be noted that almost every population has some distinctive feature, making separation of this species into discrete forms somewhat arbitrary. In the type population (
Variant 1 (
Variant 3 (
Variant 4 (
I include the specimens from Guerrero, Mexico, in
From the variation described above, there does seem to be a geographic pattern, in which specimens with a smooth pronotum occur in Chiapas, Mexico, both sculptured and unsculptured forms occur in Guatemala, and then mainly sculptured forms occur in Honduras. The degree of reproductive isolation of the different forms/populations is unclear and would be interesting to study. This pattern of morphological variation is similar to that seen in the
Distribution map of
Integument mostly black to brown-black and shining; small to medium-sized species (see HL, ML, PrW below); anterior margin of clypeus with a shallow median emargination; basal margin of mandible straight to slightly curving, but without a distinct basal notch or depression; head and body almost entirely smooth and shiny, except for faint punctae on waist and sometimes on propodeum; pronotum in profile usually distinctly asymmetrical, with a long, steep posterior face, that forms a distinct angle with dorsal surface, and a short anterior face; eye of moderate size (EL 0.11–0.17, REL 19–23), oval-shaped, with 6–8 ommatidia at greatest diameter; propodeal spines reduced to small tubercles or short points (PSL 0.06–0.10, PSI 1.1–1.7); setae on gastral tergites sparse, moderately long, and mostly suberect; frontal lobes narrow (FLD 0.13–0.18, FLI 24–27), not obscuring torular lobe in full-face view.
Mexico (Atlantic slope) to Honduras.
(17 measured) HL 0.58–0.83 (0.66) HW 0.51–0.73 (0.58) FLD 0.13–0.18 (0.15), PCW 0.03–0.05 (0.04), SL 0.49–0.76 (0.54), EL 0.11–0.17 (0.13), ACL 0.47–0.65 (0.54), ML 0.74–1.06 (0.86), PrW 0.38–0.47 (0.44), PSL 0.06–0.10 (0.09), SDL 0.05–0.09 (0.07), PL 0.27–0.39 (0.32), PH 0.18–0.24 (0.22), PW 0.12–0.17 (0.15), PPL 0.14–0.21 (0.18), PPH 0.14–0.19 (0.17), PPW 0.14–0.20 (0.18), MFL 0.52–0.86 (0.62), MTL 0.41–0.67 (0.48), CI 86–92 (89), SI 89–107 (92), REL 19–23 (21), FLI 24–27 (26), PSI 1.1–1.7 (1.2), PI 51–58 (54), MFI 83–105 (94), ACI1 64–71 (66), ACI2 84–99 (99)
Small- to medium-sized species; general body color black to brown-black, with mandibles and appendages brown to yellow-brown, lighter toward extremities; setae translucent brown; mandible with 4–6 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2 inner teeth, which are very often effaced; masticatory margin of mandible sometimes gently curving inward (perhaps due to wear), causing basal tooth to appear more robust; basal margin of mandible straight to slightly curving, without a distinct notch or depression; some specimens with basal section of mandible, where mandible inserts under clypeus, distinctly expanded and thin; dorsal surface of mandible mostly smooth and shining, with scattered piligerous punctae and a few basal striae; median lobe of clypeus somewhat obliquely flattened, and sometimes angled more ventrally than average, creating distinct dorsal and anterior faces; median lobe with faint to absent longitudinal carinulae, middle of lobe often slightly concave, apex of lobe with a short transverse carinula, usually present near anterior margin, remainder of clypeus mostly smooth and shining; posterior extension of clypeus between frontal lobes of moderate width (PCW 0.03–0.05), with subparallel sides; frontal lobes very narrow (FLD 0.13–0.18, FLI 24–27), not obscuring torular lobes in full-face view; head somewhat longer than broad (CI 86–92), roughly oval-shaped, posterior margin flat, never strongly depressed medially, lateral corners gently curving; eyes of moderate size (EL 0.11–0.17, REL 19–23), oval-shaped, with 6–8 ommatidia at greatest diameter; face almost entirely smooth and shiny, except for a variable number of carinulae around frontal carinae and on genae; scape of moderate to somewhat long length (SI 89-107), just reaching or slightly surpassing posterior margin of head when laid back; dorsal surface of scape with scattered piligerous punctae and a few faint striae, otherwise smooth and shining; funiculus with distinct 4-segmented antennal club; mesosoma usually almost entirely smooth and shiny, except for some longitudinal carinulae along metanotal groove, and transverse carinulae on dorsal and declivitous faces of propodeum; in some populations propodeum with more developed rugulae and punctae; promesonotum in profile usually asymmetrically domed and somewhat bulging, with posterior slope distinctly longer and straighter than anterior slope, and forming a more well-defined angle with dorsal surface, apex of promesonotum offset toward posterior margin; lateral and posterior margins of promesonotum in dorsal view, usually well defined, with relatively sharp transitions; one population (from high-elevation) with promesonotum in profile low-domed and roughly symmetrical; propodeal spines forming tubercles, or small projecting points (PSL 0.06–0.10, PSI 1.1–1.7); petiole of moderate length (PL/HW 0.51–0.58), node of variable height (PH/PL 0.59–0.69) and volume, usually pointing slightly posteriad, anterior face longer than posterior face and rising from about midpoint of petiole, posterior face nearly vertical, dorsum in profile usually narrowly rounded, but sometimes forming a defined apex; postpetiole in profile slightly to distinctly smaller than petiolar node (PPH/PH 0.76–0.89), roughly symmetrical, with anterior face slightly longer and more sloping than posterior face; petiole mostly smooth and shining, except for light punctae on venter and lateral portions of peduncle; postpetiole with anterior face smooth, shiny and shield-like, venter and posterior half of postpetiole lightly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body with a thin layer of moderately long standing setae; setae on gastral tergites sparse and uniformly suberect; facial setae mostly decumbent with a few erect hairs; setae on scape subdecumbent to appressed; setae on legs mostly decumbent to appressed, with coxae and ventral surfaces of the femora with some suberect setae.
(3 Measured) HL 0.63–0.69 (0.69), HW 0.58–0.63 (0.63), FLD 0.14–0.16 (0.16), PCW 0.03–0.06 (0.06), SL 0.54–0.56 (0.56), EL 0.17–0.19 (0.18), ACL 0.53–0.56 (0.56), ML 0.91–1.02 (1.02), PrW 0.52–0.56 (0.56), PSL 0.09–0.14 (0.10), SDL 0.08–0.09 (0.09), PL 0.33–0.39 (0.37), PH 0.20–0.24 (0.24), PW 0.17–0.20 (0.19), PPL 0.17–0.20 (0.20), PPH 0.19–0.20 (0.20), PPW 0.19–0.23 (0.22), MFL 0.60–0.67 (0.67), MTL 0.45–0.53 (0.53), CI 91–93 (91), SI 89–92 (89), REL 23–30 (29), FLI 24–26 (26), PSI 1.1–1.7 (1.2), MFI 94–105 (94), ACI1 65–67 (65), ACI2 94–101 (100).
Same as worker except for standard queen modifications and as follows: scutellum with irregular longitudinal rugulae/carinulae; metapleuron with longitudinal carinulae; dorsum of propodeum with transverse carinulae; petiolar node usually same as in workers, but one queen (Huejutla population) with node dorsum in posterior/anterior view forming two sharp points, separated by a steep concavity.
Unknown.
This is a rare species that inhabits montane wet forests from 1000 to 1800 m elevation. Most collections are from Winkler or Berlese extracts of sifted leaf litter. However, at Cusuco National Park in Honduras, I found two nests in a mud bank bordering a stream. The surface of the bank was rocky with an overlying layer of dense mud. The first nest was located by haphazardly cutting into the bank in an area where I saw lots of worker activity. This nest had a single chamber with about 35 workers and some brood. The second nest was located by following a returning forager. This nest had a nearly undetectable entrance (2 mm diameter) consisting only of a small hole, without any surrounding structure or excavated substrate. Behind the entrance was a very short tunnel leading to a single, small chamber. This contained 46 workers, some brood and a single dealate queen. When the nest chamber was disturbed, the queen immediately absconded out onto the clay bank followed by a series of workers carrying brood. Before excavation, foragers outside the nest were observed to be slow moving and solitary, similar to behavior in most other
Molecular phylogenetic data show that
There is minimal morphological variation amongst most populations of
Distribution map of
Integument mostly dark brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible straight, without a basal notch or depression; anterior clypeal margin forming two blunt teeth, which border the midline; face completely sculptured, mostly rugoreticulate; mesosoma completely sculptured, pronotum longitudinally rugose; pilosity on gastral dorsum bilayered, with a layer of long suberect setae, and a layer of very sparse decumbent setae; eye of moderate size (EL 0.10–0.12, REL 17–20), oval-shaped, with 5–6 ommatidia at greatest diameter; propodeal spines tuberculate to short (PSL 0.08–0.10, PSI 1.3–1.6).
Honduras to Nicaragua.
(11 measured) HL 0.59–0.69 (0.69), HW 0.52–0.64 (0.64), FLD 0.14–0.15 (0.15), PCW 0.03–0.05 (0.05), SL 0.46–0.55 (0.55), EL 0.10–0.12 (0.12), ACL 0.45–0.53 (0.53), ML 0.71–0.84 (0.84), PrW 0.37–0.42 (0.42), PSL 0.08–0.10 (0.09), SDL 0.06–0.07 (0.07), PL 0.25–0.30 (0.30), PH 0.16–0.19 (0.19), PW 0.13–0.15 (0.15), PPL 0.14–0.18 (0.18), PPH 0.16–0.18 (0.18), PPW 0.16–0.19 (0.19), MFL 0.48–0.59 (0.59), MTL 0.41–0.51 (0.51), CI 89–94 (93), SI 85–89 (86), REL 17–20 (18), FLI 23–28 (23), PSI 1.3–1.6 (1.5), MFI 104–109 (108), ACI1 67–70 (69), ACI2 96–101 (96).
Small-sized species; general body color mostly dark brown to brown, with appendages brown to orange-brown, lighter at joints and toward extremities; setae golden brown; mandible with 6 teeth, inner two teeth near basal tooth, sometimes worn; basal margin of mandible straight, without a basal notch or depression; mandible surface mostly smooth, with scattered piligerous punctae and some faint striations on base and lateral surface; anterior clypeal margin forming 2 blunt teeth, which border the midline; median lobe of clypeus with a pair of faint longitudinal carinulae, which diverge anteriorly, apex of lobe with a short transverse carinula, area in between lobe and anterior clypeal margin forming a distinct cavity, remainder of clypeal surface smooth; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.05), with sides subparallel; frontal lobes of moderate width (FLD 0.14–0.15, FLI 24–28), not greatly covering torular lobes in full-face view; head subrectangular (CI 89–94), posterior margin with a slight median depression; eye of moderate size (EL 0.10–0.12, REL 17–20), oval-shaped, with 5–7 ommatidia at greatest diameter; face mostly rugoreticulate, with some longitudinal carinulae along midline; scape somewhat short (SI 85–89), not quite reaching posterior margin of head when laid back; scape surface mostly smooth, with scattered piligerous punctae; flagellum with a somewhat distinct 4-segmented antennal club; mesosoma completely sculptured; pronotal dorsum longitudinally rugose, becoming rugoreticulate-punctae on mesonotal dorsum; side of pronotum rugose to rugose-punctate; katepisternum and side of propodeum mostly punctate, with a few rugulae on propodeum; propodeal dorsum mostly punctate, with a few transverse carinulae; propodeal declivity with transverse carinulae; promesonotum in profile slightly asymmetrical, with the anterior face long and steeper than posterior face; metanotal groove relatively distinct, but anterior slope transitioning smoothly into promesonotum; propodeal spines tuberculate (PSL 0.08–0.10, PSI 1.3–1.6); petiole in profile short (PL/HW 0.46–0.50); petiolar node usually small, somewhat dorsoventrally compressed (PH/PL 0.60–0.68), node asymmetrical in profile, anterior face longer and more sloping than nearly vertical posterior face, dorsum of node narrowly rounded, and pointing slightly posteriad; postpetiolar node in profile usually similar in size to petiolar node (type population), but sometimes bulging (PPH/PH 0.90–1.01); petiole and postpetiole usually mostly punctate, with only the anterior faces of the nodes smooth, but sometimes nodes mostly smooth; most of body dorsum with short to moderately long pilosity; pilosity on gastral dorsum bilayered, with a layer of long suberect setae, and a much sparser layer of short decumbent setae; setae on scape subdecumbent to appressed; setae on legs decumbent to appressed, with longer suberect setae on femoral venters and coxae.
(5 measured) HL 0.63–0.71 (0.64), HW 0.60–0.67 (0.61), FLD 0.16–0.17 (0.17), PCW 0.04–0.05 (0.03), SL 0.49–0.56 (0.47 (0.47), EL 0.17–0.20 (0.18), ACL 0.47–0.53), ML 0.91–1.05 (0.93), PrW 0.52–0.61 (0.54), PSL 0.12–0.14 (0.12), SDL 0.08–0.10 (0.09), PL 0.31–0.36 (0.32), PH 0.18–0.21 (0.19), PW 0.15–0.18 (0.16), PPL 0.17–0.21 (0.19), PPH 0.19–0.22 (0.19), PPW 0.20–0.24 (0.21), MFL 0.54–0.65 (0.55), MTL 0.46–0.56 (0.46), CI 94–96 (95), SI 81–84 (78), REL 29–30 (29), FLI 24–27 (28), PSI 1.4–1.6 (1.4), MFI 104–110 (109), ACI1 67–69 (68), ACI2 94–98 (101).
Same as worker except for standard queen modifications and as follows: pronotum with transverse carinulae; mesoscutum and scutellum carinulae; mesopleuron mostly smooth and shiny; propodeum transversely carinulate, with carinulae wrapping around surface; decumbent layer of setae on dorsum of gaster more dense.
Unknown.
Distribution map of
Integument mostly brown to orange-brown and mottled; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, always with a basal notch containing a small tooth; anterior clypeal margin undulating, often forming 4 blunt teeth; face completely sculptured, usually mostly rugoreticulate; gastral pilosity with a dense layer of short decumbent to appressed setae, and a sparse layer of longer suberect setae; eye small (EL 0.05–0.09, REL 10–16), subcircular, with 3–5 ommatidia at greatest diameter; posterior extension of clypeus between antennal insertions narrow (PCW 0.01–0.02), with inner margins of frontal lobes almost touching anteriorly; scape short (SI 81–92), not reaching posterior margin of head when laid back; propodeal spines tuberculate to short (PSL 0.07–0.14, PSI 1.5–2.3).
Southern Mexico to Honduras.
(16 measured) HL 0.52–0.69 (0.65), HW 0.43–0.59 (0.54), FLD 0.11–0.14 (0.14), PCW 0.01–0.02 (0.02), SL 0.36–0.50 (0.50), EL 0.05–0.09 (0.07), ACL 0.37–0.49 (0.47), ML 0.60–0.82 (0.77), PrW 0.30–0.41 (0.38), PSL 0.07–0.14 (0.10), SDL 0.04–0.07 (0.06), PL 0.23–0.31 (0.29), PH 0.14–0.18 (0.17), PW 0.11–0.14 (0.14), PPL 0.12–0.19 (0.16), PPH 0.12–0.17 (0.17), PPW 0.14–0.19 (0.17), MFL 0.39–0.53 (0.51), MTL 0.32–0.44 (0.41), CI 84–89 (84), SI 81–92 (92), REL 10–16 (13), FLI 22–27 (26), PSI 1.5–2.3 (1.6), MFI 107–119 (107), ACI1 69–73 (70), ACI2 94–105 (94).
Small-sized species; general body color a mottled dark brown to light orange-brown, with mandibles and appendages lighter, usually orange-brown to yellow-brown; setae golden brown; mandible with 6–7 teeth (usually 6), consisting of 2–3 distinct apical teeth, a distinct, usually well-defined basal tooth, and 2–3 inner teeth, which are sometimes worn and indistinct; basal margin of mandible sinuous and always with a small basal notch containing a tooth; mandible mostly smooth and shining, with scattered piligerous punctae and basal striae; anterior clypeal margin viewed at an anterodorsal angle weakly to strongly undulating (appearing nearly flat in full-face view), often forming 4 blunt teeth, median undulation (emargination) sometimes narrow and notch-like; median lobe of clypeus lacking a distinct pair of longitudinal carinulae, either completely smooth, or with faint irregular striations (type population), apex of lobe with a transverse carina, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions very narrow (PCW 0.01–0.02), with sides subparallel and inner margins of frontal lobes almost touching anteriorly; frontal lobes of moderate width (FLD 0.11–0.14, FLI 22–27), not obscuring torular lobes in full-face view; head appearing subrectangular (CI 84–89), with posterior margin depressed medially; eyes small (EL 0.05–0.09, REL 10–16), subcircular to slightly oblong, with 3–5 ommatidia at greatest diameter; face densely sculptured, usually mostly rugoreticulate, with longitudinal rugae medially, but sometimes reticulae less distinct and interconnected, interstices lightly punctate; scape relatively short, not reashing posterior margin of head when laid back (SI 81–92), usually of average thickness (type population), but some populations with scape distinctly swollen distally; scape cuticle mostly smooth and somewhat shiny, with scattered piligerous punctae; flagellum with somewhat distinct 4-segmented antennal club, apical segment noticeably enlarged; sculpture on mesosoma variable among populations, lateral surface usually weakly to strongly punctate, with variable number of longitudinal rugulae, dorsal surface of promesonotum variably rugulose-punctate, with pronotum ranging from completely smooth to strongly sculptured, most populations intermediate (type population), pronotum sometimes with a distinct longitudinal carina; propodeal declivity smooth and shiny or with a few transverse carinulae; promesonotum in profile low-domed and roughly symmetrical (type population), or less often flattened and more asymmetrical, with anterior face distinctly longer than posterior; propodeal spines tuberculate or forming short, broad triangular spines (PSL 0.07–0.14, PSI 1.5–2.3); metanotal grove usually well demarcated, of moderate depth and width; petiole of moderate length (PL/HW 0.47–0.56), average-looking; node in profile somewhat small (PH/PL 0.57–0.64), subconical, with anterior face slightly longer and more sloping than posterior face, node dorsum in profile rounded, pointing vertical to slightly posteriad; postpetiole in profile subcircular, usually appearing similar in size to petiolar node (type population), but sometimes slightly larger and more bulging (PPH/PH 0.86–1.08); petiole and postpetiole lightly to somewhat strongly punctate, with nodes variably smooth and shiny; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body dorsum with very short suberect to decumbent pilosity; scapes with a dense layer of decumbent to appressed setae; gastral pilosity consisting of a dense decumbent (type population) to appressed layer of setae, and a much sparser layer of suberect setae, which is sometimes difficult to see among decumbent setae; setae on legs mostly appressed, with a few suberect setae on coxae and femoral venters.
(5 measured) HL 0.56–0.67 (0.66), HW 0.50–0.59 (0.59), FLD 0.12–0.15 (0.15), PCW 0.02–0.03 (0.03), SL 0.41–0.51 (0.51), EL 0.13–0.15 (0.15), ACL 0.41–0.51 (0.51), ML 0.77–0.92 (0.91), PrW 0.43–0.52 (0.52), PSL 0.10–0.11 (0.10), SDL 0.06–0.08 (0.07), PL 0.27–0.32 (0.32), PH 0.16–0.19 (0.19), PW 0.14–0.16 (0.16), PPL 0.15–0.19 (0.19), PPH 0.16–0.19 (0.19), PPW 0.17–0.21 (0.21), MFL 0.44–0.55 (0.54), MTL 0.37–0.46 (0.46), CI 84–91 (89), SI 80–90 (86), REL 25–26 (26), FLI 24–27 (25), PSI 1.3–1.8 (1.5), MFI 103–115 (108), ACI1 69–72 (69), ACI2 95–100 (100).
Same as worker except for standard queen modifications and as follows: pronotum transversely rugose to rugoreticulate laterad, becoming punctate mesad; mesoscutum with narrow strip of smooth cuticle extending from anterior margin to about midpoint along midline; mesopleuron mostly smooth and shiny; propodeum mostly with transverse carinulae that wrap around surface, or less often mostly punctate; propodeal spines smaller (PSL 0.10–0.11, PSI 1.3–1.8).
Unknown.
Based on overall size and form,
Variant 2 (
Distribution map of
Integument mostly dark brown to brown; small-sized species (see HL, ML, PrW below); basal margin of mandible sinuous, with a distinct basal notch and accompanying small inner tooth; anterior clypeal margin forming 4 small teeth (inner teeth only visible from an anterodorsal angle and with mandibles open); gastral setae noticeably bilayered, with a layer of long suberect setae, and a sparse layer of short decumbent setae; face mostly covered with fine longitudinal carinulae and rugulae, posterior margin and posterolateral surfaces smooth and shiny; promesonotal dorsum with fine longitudinal and arcuate carinulae, that usually wrap around a central patch of smooth cuticle, remainder of mesosoma with light rugulae and punctae; eye of moderate size (EL 0.10–0.14, REL 15–20), with 4–5 ommatidia at greatest diameter; propodeal spines reduced to very small tubercles (PSL 0.07–0.09, PSI 1.1–1.3); frontal lobes of moderate width (FLD 0.13–0.16, FLI 20–24), not obscuring torular lobes in full-face view.
Nicaragua.
(10 measured) HL 0.62–0.73 (0.62), HW 0.56–0.70 (0.56), FLD 0.13–0.16 (0.13), PCW 0.03–0.04 (0.03), SL 0.52–0.61 (0.52), EL 0.10–0.14 (0.11), ACL 0.51–0.58 (0.51), ML 0.77–0.89 (0.77), PrW 0.07–0.09 (0.08), PSL 0.07–0.09 (008), SDL 0.06–0.08 (0.07), PL 0.27–0.34 (0.28), PH 0.17–0.20 (0.17), PW 0.13–0.17 (0.13), PPL 0.15–0.19 (0.16), PPH 0.15–0.18 (0.15), PPW 0.17–0.21 (0.17), MFL 0.55–0.65 (0.55), MTL 0.43–0.52 (0.43), CI 90–96 (90), SI 87–94 (94), REL 15–20 (19), FLI 20–24 (24), PSI 1.1–1.3 (1.2), MFI 98–111 (102), ACI1 65–67 (67), ACI2 91–97 (96).
Small-sized species; general body color dark brown to brown, with appendages lighter, brown to yellow-brown toward extremities; setae golden brown; mandible usually with 6 distinct teeth, basal tooth well-defined; basal margin of mandible sinuous, with a distinct basal notch and accompanying small inner tooth; mandible mostly smooth, except for scattered piligerous punctae, and several striae around the base and lateral surface; anterior clypeal margin in anterodorsal view forming 4 small teeth (only outer teeth visible in full-face view); median clypeal lobe with faint longitudinal carinulae that diverge toward anterior margin, apex of lobe with a short and long transverse carinula, area in between transverse carinula and anterior clypeal margin forming a distinct cavity for mandibles, remainder of clypeal surface mostly smooth; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.04), with sides subparallel; frontal lobes of moderate width (FLD 0.13–0.16, FLI 20–24), not covering torular lobes in full-face view; head subcircular to oval-shaped (CI 90–96), with posterior margin distinctly depressed; eye of moderate size (EL 0.10–0.14, REL 15–20), with 4–5 ommatidia at greatest diameter; face mostly covered with many short to long longitudinal carinulae and rugulae, posterior margin and posterolateral surfaces smooth and shiny; scape of moderate length (SI 87–94), not quite reaching posterior margin when laid back; scape surface mostly shiny, with some faint striations, and scattered piligerous punctae; flagellum with a distinct 4-segmented antennal club; promesonotal dorsum with longitudinal and arcuate carinulae, that usually wrap around a central patch of smooth cuticle, but sometimes entire dorsum sparsely carinulate, side of pronotum mostly smooth, mesopleuron and side of propodeum with variably developed punctae and rugulae, sometimes mesopleuron mostly smooth, propodeal dorsum and declivity with some transverse carinulae; promesonotum in profile low-domed, and roughly symmetrical; metanotal groove in profile of moderate width and depth, often with a small central welt (metanotum); propodeal spines reduced to small tubercles (PSL 0.07–0.09, PSI 1.1–1.3); dorsal edge of propodeal lobe in profile markedly elongate; petiole in profile appearing of moderate to long length (PL/HW 0.45–0.51); petiolar node of moderate height (PH/PL 0.58–0.65), slightly asymmetrical, with anterior face longer and more sloping than posterior face, dorsum of node gently rounded, always pointing distinctly posteriad; postpetiole in profile appearing similar in size to petiolar node (PPH/PH 0.83–0.94), subspherical, with anterior face only slightly longer than posterior face; petiole and postpetiole mostly lightly punctate, with some rugulae around nodes, anterior faces of nodes and dorsum of peduncle mostly smooth; gaster mostly smooth and shiny, with scattered piligerous punctae; face pilosity short, with a layer of suberect setae, and a denser layer of decumbent setae; setae on mesosoma and gaster mostly long and suberect, gaster with a sparse layer of decumbent setae; scape setae uniformly subdecumbent; setae on legs mostly subdecumbent, with a few suberect setae on femoral venters and coxae.
(5 measured) HL 0.67–0.70 (0.69), HW 0.62–0.65 (0.63), FLD 0.15–0.16 (0.15), PCW 0.03–0.05 (0.04), SL 0.58–0.59 (0.59), EL 0.17–0.18 (0.18), ACL 0.54–0.56 (0.54), ML 0.96–0.99 (0.99), PrW 0.55–0.56, PSL 0.10–0.12, SDL 0.07–0.09, PL 0.35–0.37 (0.36), PH 0.20 PW 0.16–0.18 (0.18), PPL 0.18–0.19 (0.19), PPH 0.19–0.21 (0.21), PPW 0.21–0.22 (0.22), MFL 0.62–0.65 (0.65), MTL 0.50–0.52 (0.51), CI 92–95 (92), SI 90–93 (93), REL 27–28 (28), FLI 24–25 (24), PSI 1.2–1.4 (1.3), MFI 98–104 (98), ACI1 63–68 (65), ACI2 92–97 (92).
Same as worker except for standard queen modifications and as follows: pronotum with transverse carinulae/rugulae; mesoscutum mostly with longitudinal carinulae/rugulae, sometimes humeral area mostly smooth, with only piligerous punctae; scutellum smooth in center, longitudinal rugulae elsewhere; propodeum with transverse carinulae that wrap around surface; mesopleuron mostly smooth; pilosity on gastral tergites clearly bilayered with a dense layer of short, subdecumbent pubescence, and a relatively sparse layer of longer suberect setae (not as long as in worker); wing venation as in
See
Given the complexity of the
Note that this species is highly variable. See comments section below discussing morphological variants. Integument mostly dark brown, red-brown, or brown; small- to medium-sized species; petiolar node in profile usually broadly rounded and distinctly angled posteriad; postpetiole in profile subspherical; propodeal spines absent to tuberculate (PSL 0.06–0.13, PSI 0.8–1.5); basal margin of mandible usually with a distinct basal notch and small accompanying tooth, but sometimes with only a small notch, or with basal margin sinuous. If basal margin of mandible with notch and tooth then: anterior clypeal margin forming 2–4 sharp to blunt teeth, with outer teeth more projecting; eye of moderate to large size (EL 0.10–0.18, REL 19–29), with 6–10 ommatidia at greatest diameter. If basal margin with small notch, but no tooth then: face completely sculptured, densely rugoreticulate; mesosoma mostly sculptured, punctate-rugulose; pilosity on first gastral tergite sparse, mostly stout and suberect, with only a few decumbent setae. If basal margin of mandible sinuous (without notch and tooth) then: anterior clypeal margin with a simple median emargination; propodeal spines absent, reduced to blunt angles where propodeal dorsum and declivity meet; eye large (EL 0.15–0.18, REL 22–27), with 8–11 ommatidia at greatest diameter; face usually completely sculptured, but sculpture never very dense, mostly carinulate-punctate; carinulae usually longitudinal, but some specimens with transverse carinulae on anterior half of head; pronotum either completely carinulate-punctate, lightly punctate, or completely smooth; carinulae when present usually transverse in orientation; some specimens noticeably long and gracile, with scape, metafemur, and petiole relatively long (SI 107–121, 86–93; PL/HW 0.58–0.63); gastral pilosity mostly sparse and suberect, with only a few decumbent to appressed setae; erect setae often stout.
Nicaragua to Ecuador.
(63 measured) HL 0.55–0.93 (0.66), HW 0.48–0.80 (0.57), FLD 0.11–0.25 (0.16), PCW 0.02–0.06 (0.04), SL 0.46–0.82 (0.55), EL 0.09–0.18 (0.12), ACL 0.47–0.75 (0.55), ML 0.68–1.21 (0.84), PrW 0.34–0.56 (0.40), PSL 0.06–0.13 (0.09), SDL 0.06–0.11 (0.09). PL 0.25–0.45 (0.31), PH 0.16–0.28 (0.19), PW 0.13–0.21 (0.15), PPL 0.13–0.27 (0.17), PPH 0.14–0.26 (0.18), PPW 0.16–0.28 (0.20), MFL 0.50–0.98 (0.59), MTL 0.41–0.76 (0.45), CI 79–94 (86), SI 85–121 (97), REL 18–29 (21), FLI 21–33 (29), PSI 0.8–1.5 (1.1), MFI 71–108 (96), ACI1 62–68 (66), ACI2 86–105 (100).
Small- to medium-sized species; general body color dark brown, to red-brown, to brown, with appendages brown to orange-brown, lighter at joints and toward extremities; setae golden; mandible with 5–7 teeth (usually 6), with basal tooth often appearing bidentate, inner teeth sometimes worn and indistinct; basal margin of mandible usually sinuous, with a distinct basal notch and accompanying small tooth (type population), but sometimes with a basal notch and no tooth, or only sinuous; mandible mostly smooth and shiny, with scattered piligerous punctae and a variable number of basal and lateral striae; anterior clypeal margin usually forming 2–4 sharp to blunt teeth (type population), but sometimes nearly flat, or with a simple median emargination; median clypeal lobe usually rather distinct, and produced slightly over anterior clypeal margin in full-face view (type population), but sometimes obliquely flattened and not produced; dorsal surface of lobe mostly smooth and shiny (type population), or with a variable number or irregular carinulae, apex of lobe with a short to long transverse carina; area in between carina and anterior clypeal margin, usually forming a distinct cavity where mandibles insert (type population); remainder of clypeal surface mostly smooth; posterior extension of clypeus between antennal insertions of narrow to moderate width (PCW 0.02–0.06; type population), with sides subparallel; frontal lobes moderate (type population) to slightly expanded (FLD 0.11–0.25; FLI 21–33), not greatly covering torular lobes in full-face view; head usually roughly oval-shaped (type population) to subcircular, with a distinct median depression in posterior margin, but head sometimes more elongate, with posterior margin flat (CI 79–94); eye of moderate to large size, sometimes relatively very large (EL 0.09–0.18, REL 18–29), oval-shaped, often bulging, with 5–11 ommatidia at greatest diameter; face sculpture highly variable, ranging from mostly smooth and shiny, to densely rugoreticulate (type population with fine longitudinal carinulae on middle of head and on gena, posterior 1/5 of head smooth); scape ranging from relatively short and somewhat thick to long and slender (SI 85–121), not surpassing to distinctly passing posterior margin when laid back (moderate length in type population, just reaching posterior margin); scape surface with numerous piligerous punctae, but mostly shiny; flagellum with a distinct (type population) to very distinct 4-segmented antennal club; mesosomal sculpture highly variable, ranging from mostly smooth, to densely rugose-rugoreticulate (type population with promesonotum mostly smooth and shiny, at most a few longitudinal carinulae dorsally); mesopleuron and side of propodeum mostly smooth, with some faint punctae and carinulae, propodeal dorsum and declivity with light transverse carinulae; promesonotum in profile usually low-domed and roughly symmetrical; metanotal groove variable, usually well-demarcated and of moderate width and depth (type population), but sometimes deeper and better defined, with metanotum forming a small welt, or sometimes shallow and indistinct, with propodeum connecting almost continuously to promesonotum; propodeal spines absent to tuberculate (PSL 0.06–0.13, PSI 0.8–1.5; forming a sharp angle in type population); petiole in profile appearing of moderate length (type population) to somewhat elongate (PL/ML 0.48–0.63); petiolar node usually broadly rounded and pointing distinctly posteriad (type population), but sometimes appearing subquadrate and asymmetrical, with an apex occurring at anterior margin of dorsum; petiolar node similar in size to petiolar node and subspherical; petiole and postpetiole usually mostly smooth and shiny, with only a few faint punctae (type population), but sometimes mostly punctate, rugulae sometimes present on posterior half of postpetiolar node; gaster mostly smooth, with scattered piligerous punctae; most of body dorsum with standing pilosity; gastral pilosity highly variable, sometimes distinctly bilayered, with a sparse layer of stout suberect setae, and a dense layer of short, decumbent pubescence (type population), sometimes pubescence absent, leaving only stout, suberect setae and a few decumbent setae, or sometimes all setae of moderate thickness and with variable density and layering (suberect layer usually always present); setae on scape usually relatively dense, and uniformly subdecumbent to appressed; setae on legs decumbent to appressed, with some longer suberect setae on femoral venters and coxae.
(16 measured) HL 0.59–0.89 (0.81), HW 0.53–0.76 (0.74), FLD 0.13–0.25 (0.22), PCW 0.03–0.07 (0.06), SL 0.48–0.81 (0.65), EL 0.16–0.25 (0.21), ACL 0.49–0.73 (0.65), ML 0.79–1.30 (1.17), PrW 0.45–0.68 (0.60), PSL 0.08–0.15 (0.12), SDL 0.08–0.13 (0.12), PL 0.30–0.47 (0.44), PH 0.18–0.26 (0.24), PW 0.16–0.23 (0.20), PPL 0.15–0.26 (0.24), PPH 0.16–0.24 (0.23), PPW 0.19–0.28 (0.26), MFL 0.53–0.93 (0.77), MTL 0.43–0.71 (0.60), CI 83–94 (91), SI 84–111 (88), REL 26–34 (29), FLI 24–34 (30), PSI 1.0–1.4 (1.0), MFI 78–103 (96), ACI1 63–67 (63), ACI2 90–106 (99).
Same as worker except for standard queen modifications and as follows (only type population queen considered): pronotum with some transverse rugulae laterad; mesoscutum lightly punctate to foveolate, with a central line of smooth cuticle; scutellum with longitudinal carinulae laterad, and a central smooth patch; propodeum with transverse carinulae that wrap around entire surface; mesopleuron mostly smooth; pilosity on mesoscutum bilayered similar to gaster, with a layer of longer erect to suberect setae, and a layer of short, dense pubescence; wing venation as in
See
The
The difficulty in resolving species boundaries in the complex probably stems from several factors, including phenotypic plasticity, morphological convergence, hybridization, and sampling bias. It could also be that if the group represents a recent radiation, there has been insufficient time for some species to reach monophyly. Based on my observations so far, I favor the idea that there are several weakly differentiated forms that occasionally come into contact and hybridize.
An interesting observation about diversification within the
By combining many divergent forms into a single species, it has made characterizing
It will also be hard to adequately separate
Defining features of the holotype’s morphology (
Variant 1 (
Variant 2 (
Variant 3 (
Variant 4 (
Variant 5 (
Variant 6 (
Variant 7 (
Variant 8 (
Variant 9 (
Distribution map of
Body color mostly black to dark brown, sometimes with a bluish glare in specimens with coarsely punctate sculpture; small- to medium-sized species (see HL, ML, PrW below); anterior clypeal margin broadly emarginate, with two blunt inner teeth (best viewed with mandibles open); basal margin of mandible sinuous, but without a basal notch or deep depression; setae on gastral tergites sparse, stout, and suberect, only sometimes with a few underlying short decumbent setae; eye relatively large (EL 0.15–0.18, REL 20–25), oval-shaped, with 8–9 ommatidia at greatest diameter; head and mesosomal sculpture densely punctate, densely carinulate, or carinulate-punctate, with carinulae merging into punctae; propodeal spines reduced to sharp angles or tubercles (PSL 0.09–0.11, PSI 0.9–1.2); frontal lobes well-developed, but not completely obscuring torular lobes in full-face view (FLD 0.15–0.22, FLI 23–28).
Mexico (Atlantic slope) to Honduras.
(9 measured) HL 0.73–0.88 (0.88), HW 0.65–0.78 (0.78), FLD 0.15–0.22 (0.22), PCW 0.03–0.04 (0.03), SL 0.56–0.68 (0.68), EL 0.15–0.18 (0.18), ACL 0.51–0.61 (0.61), ML 0.93–1.11 (1.11), PrW 0.44–0.53 (0.53), PSL 0.09–0.11 (0.10), SDL 0.09–0.11 (0.11), PL 0.34–0.40 (0.39), PH 0.23–0.25 (0.25), PW 0.17–0.19 (0.19), PPL 0.20–0.24 (0.24), PPH 0.19–0.23 (0.23), PPW 0.20–0.23 (0.23), MFL 0.64–0.80 (0.77), MTL 0.51–0.65 (0.63), CI 86–94 (88), SI 79–92 (88), REL 20–25 (23), FLI 23–28 (28), PSI 0.9–1.2 (0.9), MFI 94–115 (101), ACI1 63–66 (63), ACI2 87–94 (89).
Small- to medium-sized species; general body color mostly black to dark brown, with appendages brown to orange-brown; setae dark golden brown; mandible with 6–7 teeth, consisting of 3 distinct apical teeth, a distinct basal tooth, and 2–3 teeth in between, which are often worn and indistinct; basal margin of mandible sinuous, but without a basal notch or deep depression; mandible mostly smooth and shining, with scattered piligerous punctae and striae; anterior clypeal margin broadly emarginate, with two blunt inner teeth (best viewed with mandibles open); surface of median clypeal lobe somewhat rough (irregular depressions, punctae), with a pair of faint longitudinal carinulae that diverge anteriorly, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth and shiny; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.03–0.04), sides subparallel; frontal lobes moderately developed (FLD 0.15–0.22, FLI 23–28), but not completely covering torular lobes in full-face view; head subrectangular to slightly oval-shaped (CI 86–94), with posterior margin slightly to distinctly depressed medially; eye relatively large (EL 0.15–0.18, REL 20–25), oval-shaped, with 8–9 ommatidia at greatest diameter; head coarsely punctate (type population), or carinulate punctate, or mostly carinulate; in specimens with mainly carinulate sculpture, piligerous punctae are present in interstices, especially toward lateral margins; scape somewhat short (SI 79–92), not reaching posterior margin of head when laid back; scape surface mostly smooth and shiny (type population), or rougher, with dense piligerous punctae and carinulae; funiculus with a somewhat distinct 4-segmented antennal club; mesosoma sculpture variable, similar to head sculpture, either coarsely punctate (type population), carinulate punctate, or mostly carinulate, with carinulae mostly longitudinal in direction, but sometimes transverse on pronotal dorsum and propodeal dorsum, or arcuate on pronotal side; propodeal declivity always transversely carinulate; promesonotum in profile low-domed and roughly symmetrical; metanotal grove distinct to somewhat indistinct, of moderate to shallow width and depth; propodeal spines reduced to sharp angles or short tubercles (PSL 0.09–0.11, PSI 0.9–1.2); petiole and postpetiole robust, with nodes somewhat bulging; petiole of moderate length (PL/HW 0.45–0.58); petiolar node in profile usually subconical (type population), roughly symmetrical, with dorsum forming a well-defined apex that points vertically, but sometimes dorsum with a broadly rounded top that points slightly posteriad; postpetiole in profile subspherical, usually similar in same size to petiolar node (PPH/PH 0.79–0.93); petiole and postpetiole punctate, with anterior faces of nodes smooth and shiny; gaster mostly smooth and shiny, with scattered piligerous punctate; most of body dorsum with a layer of medium to short standing pilosity; scape with a uniform layer of subdecumbent to decumbent setae; gaster with a sparse layer of stout suberect setae, and sometimes a few underlying short decumbent setae; leg setae mostly appressed, with suberect setae on coxae and femoral venters.
Unknown.
Unknown.
A rare species,
The type population (20.6km SW Valle Nacional) is the most distinctive, with the head and mesosoma entirely coarsely punctate and without well-defined carinulae (except for the propodeal dorsum and declivity). In the right lighting, these specimens give off a bluish reflectance from the surface sculpturing.
Variant 1 (
Variant 2 (
Variant 3 (
Variant 4 (
Distribution map of
Integument brown (probably darker in recently collected specimens); small to medium-sized species (see HL, ML, PrW below); head, mesosoma, and gaster mostly smooth and shining, with petiole and postpetiole punctate; promesonotum in profile distinctly asymmetrical, with anterior face gently rounded, dorsal surface flat or only slightly curving, and posterior face short, straight and forming a relatively sharp angle with dorsal surface; median lobe of clypeus bicarinate, projecting, and with a broad and rather deep median emargination at anterior margin; basal margin of mandible straight, without notch or substantial depression; petiolar node distinctly angled posteriad; postpetiole in dorsal view, with distinct mesolateral angles and a longitudinal lobe, giving anterior half of postpetiole a distinct pinched-in appearance; eye of moderate size (EL 0.10–0.12, REL 18–19), oval shaped, with 5–6 ommatidia at greatest diameter; setae on gastral tergites of moderate length and density, suberect to subdecumbent; propodeal spines absent, forming at most an obtuse angle in profile view (PSL 0.06–0.07, PSI 1.0-1.3); frontal lobes narrow (FLD 0.14–0.15, FLI 24–25), not obscuring torular lobes in full-face view (FLD 0.14-0.15, FLI 24-25).
Northeastern Mexico.
(4 measured) HL 0.62–0.69 (0.67), HW 0.55–0.63 (0.59), FLD 0.14–0.15 (0.15), PCW 0.03–0.05 (0.05), SL 0.50–0.54 (0.52), EL 0.10–0.12 (0.11), ACL 0.48–0.51 (0.48), ML 0.79–0.86 (0.83), PrW 0.38–0.42 (0.40), PSL 0.06–0.07, SDL 0.05–0.07, PL 0.30–0.32 (0.30), PH 0.19–0.21 (020), PW 0.12–0.14 (0.13), PPL 0.16–0.18 (0.16), PPH 0.16–0.17 (0.16), PPW 0.20–0.21 (0.20), MFL 0.53–0.59 (0.55), MTL 0.44–0.48 (0.46), CI 88–91 (89), SI 86–92 (88), REL 18–19 (19), FLI 24–25 (25), PSI 1.0–1.3 (1.0), PI 51–57 (51), MFI 103–108 (107), ACI1 67–70 (70), ACI2 93–97 (93).
Small- to medium-sized species; general body color brown to light brown, with extremities becoming yellow-brown (note that all studied specimens are over 40 years old, the color of fresh specimens must be darker); setae golden; mandible with 6 teeth, consisting of 3 distinct apical teeth, a somewhat projecting basal tooth, and 2 inner teeth, which are often worn and indistinct; basal margin of mandible relatively straight, without notch or significant depression; dorsal surface of mandible mostly smooth and shining, with scattered piligerous punctae and a few short basal striae; median lobe of clypeus bicarinate, projecting, and with a broad and rather deep median emargination at anterior margin, apex of lobe with a transverse carina, area between carinae distinctly depressed; carinae on median lobe forming a distinctive triangular shape in anterodorsal view; remaining surface of clypeus mostly smooth and shiny; posterior extension of clypeus between frontal lobes somewhat narrow (PCW 0.03–0.05), with subparallel sides; frontal lobes narrow, not obscuring torular lobes in full-face view; head roughly oval-shaped (CI 88–91), posterior margin flat to slightly concave, never greatly depressed medially; eye of moderate size (EL 0.10–0.12, REL 18–19), oval-shaped, with 5–6 ommatidia at greatest diameter; face nearly completely smooth and shiny, with scattered piligerous punctae and a few longitudinal carinulae around frontal carinae and on genae; scape distinctly shorter than HW (SI 86–92), not quite reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, except for scattered piligerous punctures and faint striae; flagellum with distinct 4-segmented antennal club; mesosoma mostly smooth and shining, except metanotal groove with several longitudinal carinae, metapleuron with faint punctae, and propodeal dorsum and declivity with faint transverse carinulae; promesonotum in profile distinctly asymmetrical, with anterior face gently rounded, dorsal surface flat or only slightly curving, and posterior face short, straight and forming a relatively sharp angle with dorsal surface; metanotal groove distinct, but not very deep; propodeal spines essentially absent (PSL 0.06–0.07, PSI 1.0–1.3), forming at most obtuse angles at transition between dorsal and declivitous faces of propodeum in profile; petiole of moderate length (PL/HW 0.51–0.57), node of moderate size (PH/PL 0.62–0.67) and distinctly angled posteriad, with anterior face slightly longer and more sloping than posterior face, node dorsum broadly rounded; postpetiole in profile asymmetrical, with long sloping anterior face and short nearly vertical posterior face; postpetiole in dorsal view with distinctive mesolateral angles and a longitudinal lobe, giving the anterior half of postpetiole a distinct pinched-in appearance; anterior faces of petiole and postpetiole smooth and shiny, remaining surfaces faintly punctate; most of body with a thin layer of short to medium length standing setae; pilosity on gastral tergites somewhat bilayered, with a layer of longer suberect setae, and a layer of equally dense subdecumbent setae; facial setae short and mostly decumbent; setae on scape subdecumbent to decumbent; setae on legs suberect to appressed, with longer suberect setae on femoral venters and coxae.
(1 measured) HL 0.69, HW 0.63, FLD 0.15, PCW 0.05, SL 0.53, EL 0.17, ACL 0.52, ML 0.93, PrW 0.50, PSL 0.09, SDL 0.08, PL 0.34, PH 0.22, PW 0.14, PPL 0.19, PPH 0.19, PPW 0.24, MFL 0.59, MTL 0.49, CI 92, SI 83, REL 27, FLI 24, PSI 1.2, MFI 107, ACI1 68, ACI2 98.
Same as worker except for standard queen modifications. See
Unknown.
The combination of sculpture, clypeus structure, and postpetiole structure make
It should be noted that this species is known from only nine specimens that were collected in 1969. Thus, the coloration described here is certainly different from what fresh specimens probably look like. Further more, this species has not been included in molecular analyses and it is somewhat uncertain to which species it is most closely related. However, because the postpetiole has a distinct dorsal lobe I include
Known only from the type locality.
Integument shining, largely black to red-black; medium-sized species (see HL, ML, PrW below); head mostly smooth and shiny; mesosoma reticulately costate to coarsely rugoreticulate; propodeal spines reduced to small upward projecting points, or sharp right angles at interface between dorsal and declivitous faces of propodeum (PSL 0.14–0.18, PSI 1.4-1.9); eyes large and somewhat bulging (EL 0.15–0.18, REL 23-24), with 7–10 ommatidia at greatest diameter; anterior margin of clypeus with shallow median emargination; basal margin of mandible straight, without notch or substantial depression; pilosity on gastral dorsum long, flexuous, and relatively sparse.
Nicaragua to Panama.
(11 measured) HL 0.71-0.85 (0.84), HW 0.63-0.76 (0.76), FLD 0.22-0.27 (0.26), PCW 0.06-0.07 (0.07), SL 0.60-0.77 (0.77), EL 0.15-0.18 (0.18), ACL 0.54-0.68 (0.68), ML 0.91-1.11 (1.11), PrW 0.48-0.58 (0.57), PSL 0.14–0.18 (0.14), SDL 0.08–0.10 (0.09), PL 0.37-0.46 (0.44), PH 0.21-0.26 (0.26), PW 0.16–0.22 (0.20), PPL 0.19–0.25 (0.23), PPH 0.19–0.25 (0.24), PPW 0.20–0.25 (0.25), MFL 0.72-0.91 (0.91), MTL 0.56-0.71 (0.71), CI 87-93 (91), SI 95-103 (101), REL 23-24 (24), FLI 33-36 (34), PSI 1.4-1.9 (1.4), MFI 82-88 (84), ACI1 63-66 (64) ACI2 84–91 (88).
Medium-sized species; general body color black to red-black, with brown patches on waist and gaster; mandibles, clypeus and appendages dark brown to yellow-brown; setae golden; mandible with 6–7 teeth, consisting of 3 distinct apical teeth, a basal tooth, and 2–3 inner teeth, which are often worn and indistinct; basal margin of mandible straight, without any notch or significant depression; dorsal surface of mandible mostly smooth and shiny, with scattered piligerous punctae and variable number of basal striae; median lobe of clypeus with a pair of vestigial longitudinal carinulae and/or several ill-defined rugulae, apex of lobe with a short transverse carinula, remainder of clypeus smooth and shiny; posterior extension of clypeus between frontal lobes rather broad (PCW 0.06-0.07), with subparallel sides; frontal lobes relatively well developed, but not markedly expanded dorsolaterally (as in
(5 measured) HL 0.71–0.85 (0.79), HW 0.63–0.76 (0.73), FLD 0.22–0.27 (0.24), PCW 0.07–0.09 (0.07), SL 0.64–0.78 (0.69), EL 0.20–0.24 (0.21), ACL 0.53–0.66 (0.60), ML 1.05–1.29 (1.12), PrW 0.57–0.70 (0.60), PSL 0.18–0.21 (0.19), SDL 0.09–0.11 (0.10), PL 0.43–0.53 (0.48), PH 0.24–0.29 (0.25), PW 0.20–0.25 (0.22), PPL 0.20–0.26 (0.23), PPH 0.23–0.29 (0.25), PPW 0.25–0.31 (0.26), MFL 0.77–0.94 (0.81), MTL 0.61–0.73 (0.64), CI 91–93 (93), SI 93–95 (94), REL 28–30 (28), FLI 34–37 (24), PSI 2.0–2.2 (2.0), MFI 87–90 (90), ACI1 63–66 (64), ACI2 82–90 (87).
Same as worker except for standard queen modifications and the following: face usually more sculptured, with light fan of rugulae/carinulae extending from frontal lobes to ocelli, sculpture around antennal sockets more distinct; costae on mesoscutum with a decidedly longitudinal orientation, but often wavy, and usually with some reticulation anteriorly; costae on side of promesonotum longitudinal in orientation; mesopleuron mostly smooth and shiny; propodeal spines always present, short, projecting dorsoposteriad (PSL 0.18–0.21, PSI 2.0–2.2); wing venation as in
Unknown.
Within its range
Distribution map of
Integument mostly black or dark brown to red-brown; small- to medium-sized species (see HL, ML, PrW below); anterior clypeal margin undulating, forming 2–4 blunt teeth; basal margin of mandible usually sinuous, with a slight basal depression, but without a tooth; face completely sculptured, mostly rugoreticulate; pilosity on gastral dorsum predominately suberect and relatively sparse; eye of moderate size (EL 0.10–0.15, REL 18–22), oval-shaped, with 5–8 ommatidia at greatest diameter; propodeal spines tuberculate to short (PSL 0.08–0.16, PSI 1.1–1.8), often robust; petiole in profile often distinctive, appearing somewhat elongate, with node broadly rounded and pointed posteriad, venter under node with a small concavity.
Mexico (Atlantic slope).
(29 measured) HL 0.62–0.78 (0.65), HW 0.55–0.71 (0.58), FLD 0.15–0.20 (0.16), PCW 0.04–0.06 (0.04), SL 0.50–0.67 (0.55), EL 0.10–0.15 (0.12), ACL 0.48–0.60 (0.50), ML 0.77–1.02 (0.85), PrW 0.39–0.49 (0.43), PSL 0.08–0.16, SDL 0.07–0.10 (0.09), PL 0.31–0.40 (0.34), PH 0.17–0.23 (0.19), PW 0.14–0.19 (0.15), PPL 0.15–0.22 (0.20), PPH 0.16–0.21 (0.17), PPW 0.17–0.22 (0.19), MFL 0.53–0.76 (0.60), MTL 0.45–0.62 (0.48), CI 89–96 (89), SI 85–98 (94), REL 18–22 (21), FLI 25–29 (28), PSI 1.1–1.8 (1.6), PI 53–59 (59), MFI 93–107 (97), ACI1 65–71 (68), ACI2 87–98 (92)
Small- to medium-sized species; general body color black, or dark brown to red-brown (type population), with appendages brown to orange-brown or yellow-brown, lighter in joints and toward extremities; setae golden brown; mandible with 6 teeth, 2 teeth nearest basal tooth sometimes more worn and less distinct, basal tooth often well-defined, projecting; basal margin of mandible usually sinuous, with a slight basal depression (type population), but without a tooth, some populations with basal margin almost straight; mandible mostly smooth, with scattered piligerous punctae, and some striations near base and along lateral surface; median lobe of clypeus with a pair of faint longitudinal carinulae that diverge anteriorly, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth; area in between median lobe and anterior clypeal margin forming a distinct concavity where mandibles insert; posterior extension of clypeus between frontal lobes of moderate width (PCW 0.04–0.06), with sides subparallel; frontal lobes of moderate width (FLD 0.15–0.20, FLI 25–29), not greatly covering torular lobes in full-face view; head roughly oval-shaped (CI 89–96), with posterior margin flat to slightly depressed medially; eye of moderate size (EL 0.10–0.15, REL 18–22), oval-shaped, with 5–8 ommatidia at greatest diameter; face usually strongly sculptured, mostly rugoreticulate, with some longitudinal rugae along midline (type population), some populations with sculpture somewhat polished and reticulate less distinct; scape of moderate length (SI 85–98), just reaching posterior margin of head when laid back; surface of scape mostly smooth, with scattered piligerous punctae; flagellum with distinct 4-segmented antennal club; mesosoma usually completely sculptured (type population), but some populations shinier overall and with pronotum mostly smooth and shiny; pronotal dorsum longitudinally rugose, with rugae dense to somewhat sparse and irregular, rugae usually becoming punctae and reticulae on dorsum of mesonotum; side of pronotum mostly punctate or rugose, often with a small to large patch of smooth cuticle on ventral half; mesopleuron and side of propodeum rugulose-punctate; propodeal dorsum with rugoreticulae near anterior margin, changing to transverse carinulae near posterior margin; promesonotum in profile low-domed, slightly asymmetrical, with apex posterior of midpoint, anterior declivity of pronotum sometimes forming a sharp transition with dorsum, sculpture on declivity punctate, contrasting with rugose dorsum; metanotal groove usually well-demarcated, of moderate depth and width; propodeal spines tuberculate to short (PSL 0.08–0.16, PSI 1.1–1.8), often somewhat robust (type population); petiole in profile often distinctive, appearing somewhat elongate (PL/HW 0.53–0.56), with node broadly rounded and pointed posteriad, venter under node often with a small concavity; peduncle sometimes noticeably slender, thickening toward node; postpetiole in profile subspherical, similar in size to petiolar node (PPH/PH 0.85–0.91); anterior face of petiolar and postpetiolar nodes smooth, remainder of waist mostly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae; pilosity on head, short and bilayered, with some longer suberect setae, and many shorter decumbent setae, remainder of body dorsum with longer suberect setae; pilosity on gastral dorsum predominately suberect and relatively sparse; setae on scape mostly subdecumbent; setae on legs mostly subdecumbent to appressed, with some longer suberect setae on femoral venters and coxae.
(11 measured) HL 0.66–0.77 (0.66), HW 0.60–0.70 (0.60), FLD 0.17–0.20 (0.18), PCW 0.05–0.07 (0.05), SL 0.53–0.64 (0.53), EL 0.18–0.20 (0.18), ACL 0.50–0.60 (0.51), ML 0.94–1.13 (0.94), PrW 0.53–0.64 (0.54), PSL 0.14–0.17 (0.15), SDL 0.09–0.12 (0.09), PL 0.37–0.44 (0.38), PH 0.19–0.23 (0.20), PW 0.16–0.19 (0.17), PPL 0.18–0.22 (0.20), PPH 0.18–0.23 (0.19), PPW 0.20–0.24 (0.20), MFL 0.59–0.75 (0.59), MTL 0.49–0.61 (0.51), CI 90–95 (92), SI 84–95 (87), REL 28–30 (30), FLI 26–30 (29), PSI 1.3–1.8 (1.6), MFI 90–104 (102), ACI1 65–68 (68), ACI2 88–98 (96).
Same as worker except for standard queen modifications and as follows: pronotum with transverse rugae and rugoreticulae on lateral surface, changing to carinulae and punctae mesad; mesoscutum densely longitudinally rugose; scutellum rugose to rugoreticulate, sometimes with a smooth patch mesad; side of propodeum rugulose, with faint punctae, dorsum with transverse carinulae; wing venation as in
Unknown.
This species is known only from Winkler and Berlese samples of sifted leaf litter. It occurs from approximately 1000–2000 m elevation and inhabits a variety of montane mesic forest environments (e.g. cloud forest, wet oak forest, oak-pine forest, tropical subevergreen forest,
Several aberrant populations are worth separating out as variants of
Distribution map of
Integument mostly dark red-brown to brown; medium to large-sized species (see HL, ML, PrW below); head and promesonotum mostly foveate to coarsely rugoreticulate; eye relatively small (EL 0.10–0.13, REL 12–16), somewhat bulging, subcircular to slightly oval-shaped, with 6–8 ommatidia at greatest diameter; gastral pilosity long, dense, and mostly suberect; propodeal spines reduced to sharp angles or small tubercles (PSL 0.12–0.16, PSI 1.0–1.3); anterior clypeal margin with a median excavation containing 2 sharp outer teeth and 2 smaller inner teeth (sharp to blunt); basal margin of mandible straight to slightly sinuous, without a basal notch or depression.
Honduras to Panama.
(11 measured) HL 0.87–1.03 (0.99), HW 0.78–0.92 (0.85), FLD 0.26–0.31 (0.29), PCW 0.04–0.08 (0.07), SL 0.67–0.80 (0.74), EL 0.10–0.13 (0.11), ACL 0.58–0.69 (0.60), ML 1.15–1.38 (1.29), PrW 0.55–0.67 (0.63), PSL 0.12–0.16 (0.14), SDL 0.10–0.16 (0.13), PL 0.46–0.56 (0.53), PH 0.25–0.29 (0.29), PW 0.17–0.21 (0.20), PPL 0.27–0.32 (0.32), PPH 0.22–0.27 (0.25), PPW 0.21–0.26 (0.24), MFL 0.82–0.99 (0.93), MTL 0.65–0.80 (0.73), CI 86–92 (86), SI 83–89 (86), REL 12–16 (13), FLI 31–34 (34), PSI 1.0–1.3 (1.1), MFI 91–98 (92), ACI1 65–68 (65), ACI2 82–89 (82).
Medium to large-sized species; general body color mostly dark red-brown, with patches of brown and orange-brown on gaster, appendages lighter; setae golden brown; mandible with 6–9 teeth, consisting of 3 distinct apical teeth, a distinct basal tooth, and 2–5 inner teeth/denticles, which are usually worn and indistinct; basal margin of mandible straight to slightly sinuous, without a basal notch or depression; mandible mostly smooth and shiny, with scattered piligerous punctae and faint striae; anterior clypeal margin viewed from an anterodorsal angle with a median excavation containing 2 sharp outer teeth and 2 smaller inner teeth, which are sharp to blunt, and recessed behind median lobe of clypeus (not visible with mandibles closed); median lobe of clypeus usually with a pair of faint carinulae that diverge toward the anterior margin, apex of lobe with a short transverse carinula, remainder of clypeus mostly smooth and shiny, sometimes with a few additional rugulae on median lobe; posterior extension of clypeus between antennal insertions of moderate width (PCW 0.04–0.08), sides subparallel to slightly diverging posteriad; frontal lobes well-developed, but not completely covering torular lobes in full-face view (FLD 0.26–0.31, FLI 31–34); head appearing subrectangular (CI 86–92), with posterior margin depressed medially; eye relatively small (EL 0.10–0.13, REL 12–16), somewhat bulging, subcircular to slightly oval-shaped, with 6–8 ommatidia at greatest diameter; face usually strongly foveate to coarsely rugoreticulate, with a few longitudinal costae along midline, interstices with piligerous punctae, some high-elevation populations with face sculpture more polished and effaced, especially on side of head, which can be smooth and shiny; scape relatively short (SI 83–89), not reaching posterior margin of head when laid back; scape surface mostly smooth and shiny, with scattered piligerous punctae; flagellum with an indistinct 4-segmented antennal club; mesosoma usually densely sculptured, with promesonotum mostly foveate (especially on dorsum) and remaining surfaces foveate to rugoreticulate or rugose, in some high-elevation populations sculpture is reduced, with foveae on promesonotal dorsum appearing more like transverse furrows, and with side of promesonotum largely smooth and shiny; propodeal declivity mostly smooth and shiny, usually with a few transverse carinulae; promesonotum in profile low-domed, slightly asymmetrical, with the apex shifted anteriorly; anterior face of promesonotum longer and steeper than posterior face and mostly smooth; metanotal groove well-demarcated, of moderate width and depth; propodeal spines forming sharp angles or at most small tubercles (PSL 0.12–0.16, PSI 1.0–1.3); petiole relatively long (PL/HW 0.58–0.63), with node of average size (PH/PL 0.51–0.54); node in profile variable, usually forming a small roughly symmetrical dome with a rounded dorsum, but sometimes dome more asymmetrical with anterior face longer than posterior face, and sometimes with apex more sharp and positioned anterior of midpoint; postpetiole somewhat elongate (PPH/PPL 0.79–0.88) and globular, about the same height and volume as petiolar node (PPH/PH 0.88–0.98), postpetiolar node asymmetrical, with anterior face longer than posterior face; venter of postpetiole in profile usually distinctly sinuous, with anteroventral margin forming a small process; petiolar and postpetiolar nodes mostly smooth and shiny, remaining waist surfaces faintly punctate; gaster mostly smooth and shiny, with scattered piligerous punctae; most of body with a moderately dense layer of long suberect to subdecumbent pilosity; scape with layer of longer sparser suberect setae and layer of shorter subdecumbent setae; gastral setae not distinctly bilayered, forming a moderately dense layer of long suberect to subdecumbent setae; setae on legs suberect to decumbent, with longer suberect hairs on coxae and femoral venters.
(5 measured) HL 0.86–1.01 (0.92), HW 0.78–0.90 (0.84), FLD 0.27–0.32 (0.29), PCW 0.05–0.09 (0.06), SL 0.67–0.78 (0.71), EL 0.16–0.20 (0.17), ACL 0.59–0.67 (0.61), ML 1.25–1.48 (1.33), PrW 0.68–0.79 (0.71), PSL 0.15–0.19 (0.17), SDL 0.13–0.18 (0.14), PL 0.52–0.58 (0.54), PH 0.26–0.31 (0.28), PW 0.19–0.23 (0.21), PPL 0.31–0.33 (0.33), PPH P0.24–0.28 (0.25), PW 0.19–0.23 (0.21), MFL 0.83–0.98 (0.89), MTL 0.67–0.78 (0.69), CI 88–93 (92), SI 83–88 (84), REL 20–23 (20), FLI 33–35 (34), PSI 1.1–1.2 (1.1), MFI 92–94 (94), ACI1 65–67 (65), ACI2 83–89 (86).
Same as worker except for standard queen modifications and the following: mesoscutum foveate, except for a longitudinal strip of completely smooth cuticle, which crosses the entire surface; mesopleuron mostly smooth and shiny; propodeum carinate to rugose, with sculpture transversely wrapping around surface; wing venation as in
Unknown.
Molecular phylogenetic results indicate that
Within my concept of
Preliminary molecular phylogenetic results, which include several populations of each morphotype, show that the two forms separate into sister clades, suggesting that the high-elevation variant may represent a distinct species. However, I choose to recognize one species until variation within the high-elevation form is better understood.
Expedition teams for each year of the Leaf Litter Arthropods of MesoAmerica (LLAMA) project
I owe a great debt of gratitude to many people, institutions, and funding agencies for helping me complete this project. For providing specimens for morphological and molecular work I thank Jack Longino (JTLC), Phil Ward (PSWC, UCDC), Roy Snelling (LACM), Fernando Fernández (ICN), Bill MacKay, Stefan Cover (MCZ), Laura Sáenz, Manuel Solis (INBio), Corrie Moreau (FMNH), James Boone (FMNH), Ted Schultz (USNM), Seán Brady (USNM), David Donoso and Bob Anderson. I am especially grateful to everyone who participated in the Leaf Litter Arthropods of MesoAmerica (LLAMA) project, which provided specimens, field assistance, and financial support for several seasons of fieldwork (
Key to