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Stenamma is a cryptic “leaf-litter” ant genus that occurs in mesic forest habitats throughout the Holarctic region, Central America, and part of northwestern South America (Colombia and Ecuador). The genus was thought to be restricted primarily to the temperate zone, but recent collecting efforts have uncovered a large radiation of Neotropical forms, which rival the Holarctic species in terms of morphological and behavioral diversity. By inferring a broad-scale molecular phylogeny of Stenamma,
Stenammini, systematics, taxonomy, cryptic ants, Neotropics, Central America, cloud forest
The genus Stenamma Westwood comprises a taxonomically challenging but intriguing lineage of myrmicine ants, which occupy mesic forest habitats throughout most of the northern temperate zone, Middle America, and northwestern South America (
Understanding of the diversity and distribution of Stenamma has progressed relatively slowly compared to more conspicuous ant genera. This is because of a general lack of material, poor representation of colony series and reproductive castes in collections, and the tendency for worker morphology to be both conservative and convergent among species. Despite these obstacles, the genus has received considerable attention in the Holarctic region, with the publication of many species descriptions, as well as several large regional revisions (
On a global scale, understanding of Stenamma has been greatly improved by several recent studies that incorporate molecular data. First, two investigations looking at higher-level relationships among ants found that Stenamma is closely related to the genera Aphaenogaster Mayr, Messor Forel, Goniomma Emery, and Oxyopomyrmex André (
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 (Figure 1). The only MAC species to reach the U.S.A. and to co-occur with HOC species is Stenamma huachucanum Smith, which is known from the sky islands of Arizona, New Mexico, and Texas, as well as from many sites in Mexico (Texas record in
Map showing the global distribution of the Middle American clade of Stenamma.
I present here the first comprehensive revision of the Middle American clade of Stenamma, recognizing 40 species, of which 33 are described as new. This work represents the first all-in-one treatment of Neotropical taxa since
The first realization that Middle America contained a diverse radiation of previously unknown Stenamma species came from collections made in Costa Rica by J. T. Longino and the Arthropods of La Selva project (ALAS; http://viceroy.eeb.uconn.edu/ALAS/ALAS.html), which he and R. Colwell organized. Later, it was discovered that several coleopterists (R. S. Anderson, A. F. Newton, S. B. Peck), who used Berlese funnels extensively, had collected many Stenamma specimens as bycatch over the last 50 years from throughout Mexico and Central America. Much of this material was processed at various institutions, but was aggregated at the Los Angeles County Museum (LACM) by the late R. R. Snelling who began to reviseNeotropical Stenamma (Snelling pers. comm.). Additional material from the coleopterists was stored in the Field Museum of Natural History (FMNH) bulk sample collection and reviewed by myself. Over the last six years, a very large amount of material has come from the Leaf Litter Arthropods of MesoAmerica project (LLAMA; https://sites.google.com/site/longinollama/), organized by J. T. Longino and R. S. Anderson. This project has intensively sampled leaf litter insects from tropical wet forests in Chiapas, Guatemala, Honduras, and Nicaragua. Many new species were collected by LLAMA, but just as importantly, the project has provided fresh specimens for the molecular studies described above and on-going work. A significant amount of material has come also from personal collections made during LLAMA expeditions.
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 Stenamma species. The biology of Nearctic and Palearctic species is provided in
Neotropical Stenamma species are found mainly in mesic forest habitats from sea level to around 3000 m elevation, with the highest record reported at 3700 m at Pico Orizaba in Mexico (Stenamma manni Wheeler). Stenamma specieshave been collected mostly in lowland rainforest, montane wet forest, cloud forest, and dwarf forest. A few species (Stenamma huachucanum, Stenamma lagunum, Stenamma manni) also have been found in seasonal woodland habitats in drier areas of Mexico and the southwestern United States. Stenamma has never been collected in tropical dry forest or scrub environments. One hypothesis for the nearly complete geographic separation of the HOC and MAC is the presence of extensive xeric environments in northern Mexico and the southwestern United States (
An exceptional characteristic of Stenamma is that many species seem to be well adapted to cool, wet environments. Although present in the lowlands, the diversity and abundance of Stenamma species peak at mid-elevations between 800–1600 m (Branstetter unpublished data). Also, it has been found that Stenamma can be the most common ant genus in leaf-litter samples collected from very wet and cool, cloud forest localities. These ecological traits are in contrast to the pattern seen in ants generally, in which diversity and abundance decrease with elevation. Biogeographic results indicate that Stenamma originated in the Nearctic, potentially preadapting it to thrive in cool montane forest environments (
Most Stenamma species have very cryptic habits. Nests are usually small, and workers are slow moving, often becoming immobile upon disturbance. Consequently, Stenamma is rarely found by the casual observer and most collections are made by sifting leaf-litter from the forest floor. This fact has given Stenamma its stereotype as a “leaf-litter ant genus.” Although nests of many species do occur in the leaf litter, and foragers are common there, recent collecting efforts have revealed that MAC Stenamma species nest in a variety of microhabitats. I have found nests in large logs, in small rotting branches, in and under bark, in steep clay or mud banks, in and under epiphytes, under rocks, in the ground, and under leaves in leaf litter. At least a few Stenamma species nest arboreally. For example, several variants of Stenamma schmidti Menozzi have been found reliably underneath epiphyte clumps in the canopy and by canopy fogging, and Stenamma longinoi sp. n. is known only from one collection under epiphytes in a treefall. Based on circumstantial evidence, I surmise that a few other species have arboreal nests or at least forage arboreally. Stenamma stictosomum sp. n. and Stenamma felixi Mann have been found in epiphytic orchids inspected at quarantine in the U.S., and Stenamma callipygium sp. n. has been collected most often by beating vegetation.
One of the most intriguing recent discoveries has been the observation that some Stenamma species nest in clay banks.
Stenamma colonies tend to be small and simple, but there is considerable variation in size and structure. Stenamma diversum, for example, has very small colonies. Nests consist of a single chamber and have at most a dozen workers, a few alates, brood, and a single dealate queen. In contrast, colonies of Stenamma manni and Stenamma megamanni are very large. I have found mature colonies of both species in logs and in the ground (usually underneath a log or rock). They tend to have many chambers containing several hundred to perhaps a thousand workers, along with brood and alates (a thorough census has not been performed). Stenamma alas and Stenamma expolitum have colonies of intermediate size, with up to 200 workers, 50 alates, brood, and a single dealate queen (
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 (http://zerenesystems.com/cms/home) rather than Auto-Montage. Adobe Photoshop and Illustrator, both CS5, were used to enhance images and create Figures.
Collection and specimen data for all material examined in this study, along with all color images, have been uploaded to AntWeb (http://www.antweb.org), a site hosted by the California Academy of Sciences. AntWeb subsequently provides all specimen-level data, images, and natural history content to the Global Biodiversity Information Facility (http://www.GBIF.org), the Encylopedia of Life (http://www.EOL.org), and Wikipedia (http://www.Wikipedia.org). The most important linking fields for specimen data are the collection and specimen codes.
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 (http://www.google.com/earth/index.html), the GEOnet Names Server (http://earth-info.nga.mil/gns/html/index.html), the FMNH bulk sample database (http://emuweb.fieldmuseum.org/arthropod/bulksamples.php), and/or data already available on AntWeb. Estimated coordinates are indicated with brackets and often include an error estimate on AntWeb. Data for type material follow the format: [Country], [First Administrative District], [Locality], [GPS Coordinates ±Error], [Elevation], [Collection Date], [Habitat], [Microhabitat], [Collector and Collection Code], [Repository and Specimen Code].
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 (
Stenamma taxonomy is complicated by several phenomena. First, species are almost always represented by morphologically distinct, allopatric populations, in which intrapopulation variation is less than interpopulation variation. If one were to split all of these variants into species, there would be two to three times more MAC species than included here. Second, many species show exceptional variation along elevational gradients (Figure 2). Usually, populations at lower elevations are smaller and more heavily sculptured, while populations at higher elevations increase in size, become darker in color, and have smooth shiny sculpture. If one compares only specimens from low and high elevations they often appear exceptionally divergent, but I have found that it is common for specimens from intervening elevations to have intermediate phenotypes. Consequently, without having thorough geographic coverage, delimiting Stenamma species is a rather arbitrary task. In general, I lump morphological variants into widespread polytypic species if intermediate forms are present and the genetic data are ambiguous, and I split variants into species if they seem “sufficiently” divergent and occur at multiple sites. All species described here should be treated as hypotheses to be tested once more data become available. My reasoning for species delimitation decisions is provided in the comments section of each species account.
Morphological variation in Stenamma connectum along an elevational gradient A 990 m (CASENT0605586) B 1370 m (CASENT0605496) C 1770 m (CASENT06005461) D 1990 m (CASENT0605552).
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 Figure 3).
HL Head length (full-face view): maximum length of the head, measured from the posterior margin of the head to the anterior-most extremity of the clypeus.
HW Head width (full-face view): maximum width of the head, eyes excluded.
FLD Frontal lobes distance (full-face view): the maximum distance separating the outer margins of the frontal lobes.
PCW Posterior clypeus width (full-face view): width of the posterior extension of the clypeus (depressed area between the frontal lobes), measured at the midpoint of the antennal sockets.
SL Scape length (most suitable view): length of the first antennal segment, exclusive of the basal condyle.
EL Eye length (most suitable view): the maximum diameter of the compound eye, including the outer ring of ommatidia, which are often black.
ACL Antennal club length (most suitable view): the combined length of antennal segments 9-12, each measured individually and then summed together.
ML Mesosoma length (profile view): greatest distance from the approximate inflection point, where the pronotum curves into the cervical shield, to the posterior apex of the propodeal lobes.
PrW Pronotum width (dorsal view): maximum width of the pronotum.
PSL Propodeal spine length (profile view): distance from the center of the propodeal spiracle to the tip of the propodeal spine.
SDL Spiracle to declivity length (profile view): minimum distance from the center of the propodeal spiracle to the propodeal declivity.
PL Petiole length (profile view): maximum length of the petiole, measured from the narrowest point of the anterior constriction, to the posterior margin; because the propodeal lobes usually obscure the anterior constriction in lateral view, this point must be approximated.
PH Petiole height (profile view): maximum height of the petiole, measured at a right angle to PL and taken from the dorsal surface of the petiolar node to the ventral surface of the postpetiolar helcium, where it inserts into the petiole.
PW Petiole width (dorsal view): maximum width of the petiole.
PPL Postpetiole length (profile view): maximum length of the postpetiole, measured from the posterior margin of the enlarged portion of the helcium, to the posterior margin of the postpetiole.
PPH Postpetiole height (profile view): height of the postpetiole, measured in a line perpendicular to PPL. If the ventral surface is concave upward, the measurement should be taken from the uppermost portion of the curve.
PPW Postpetiole width (dorsal view): maximum width of the postpetiole.
MFL Metafemur length (most suitable view): maximum length of the metafemur, measured from the distal margin of the trochanter to the metafemur apex. In most cases this measurement was taken from the anterior side of the metafemur.
MTL Metatibia length (most suitable view): maximum distance of the metatibia, measured from the proximal constriction, just before the inserting condyle, to the apex. This measurement is always performed on the dorsal surface of the metatibia so that the proximal condyle is not obscured by the metafemur.
CI Cephalic index: HW/HL × 100.
SI Scape index: SL/HW × 100.
REL Relative eye length: EL/HW × 100.
FLI Frontal lobes index: FLD/HW × 100.
PSI Propodeal spine index: PSL/SDL.
MFI Metafemur index: HW/MFL × 100.
ACI1 Antennal club index: segments 11+12/ACL × 100.
ACI2 Antennal club index: ACL/SL × 100.
Standard measurements used in this study.
CAS California Academy of Sciences, San Francisco, CA, USA.
EAPZ Escuela Agricola Panamericana Zamorano, Tegucigalpa, Honduras.
ECOSCE Colección Entomológica de El Colegio de la Frontera Sur, Unidad San Cristóbal, Chiapas, Mexico.
FMNH Field Museum of Natural History, Chicago, IL, USA.
ICN Insect Collection, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C., Colombia.
INBio Instituto Nacional de Biodiversidad, Costa Rica.
JTLC John T. Longino, personal collection, University of Utah, Salt Lake City, Utah, USA.
LACM Los Angeles County Museum of Natural History, Los Angeles, CA, USA.
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA.
MGBPC Michael G. Branstetter, personal collection.
MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.
NHMB Naturhistorisches Museum, Basel, Switzerland.
PSWC Philip S. Ward, personal collection, University of California, Davis, CA, USA.
UCD Bohart Museum of Entomology, University of California, Davis, CA, USA.
UNAM Universidad Nacional Autonoma de Mexico, Mexico D.F., Mexico.
USNM National Museum of Natural History, Washington, D.C., USA.
UVGC Colección de Artrópodos, Universidad del Valle de Guatemala, Guatemala City, Guatemala.
Several characters useful in distinguishing Stenamma species are discussed below. It should be noted that almost every character type is variable within species and prone to some level of homoplasy among species. Thus, it is difficult to assess which characters are “better” than others for separating species. In MAC Stenamma any variable character can be useful in some cases, but misleading in others. All characters used previously to separate HOC Stenamma species tend to be more variable among MAC species. New characters for distinguishing species include the structure of the basal margin of the mandible, configuration of gastral pilosity, frontal lobe width, promesonotum shape, and propodeal spine length.
Body size. Species of Stenamma are relatively small compared to most other ant genera, but within the genus there is exceptional variability among species, within species, and, in some cases, within single colonies. Even though within-species and within-colony size variation can be substantial, size is still useful as a character in species separation. Using HL, ML, and PrW as proxies for overall size I assign species to the qualitative categories small (HL ≤ 0.73, ML ≤ 0.92, PrW ≤ 0.45), medium (HL 0.74–0.96, ML 0.93–1.27, PrW 0.46–0.62), and large (HL ≥ 0.97, ML ≥ 1.28, PrW ≥ 0.62), with intermediate states possible. One phenomenon that has not been reported previously in Stenamma is the existence of substantial within-colony size variation. For medium- to large-sized species, I have noticed substantial size variation from the smallest nanitics to the largest workers. In some cases, the largest workers have allometrically enlarged heads, making them appear almost soldier-like. In all cases the size variation appears to be continuous rather than discrete.
Clypeus & mandible structure.
Clypeus structure in Stenamma maximon (CASENT0604875) A Clypeus in full-face view B Clypeus in anterodorsal view.
The anterior clypeal margin varies greatly (see Figure 19). It can be entire; have a shallow median emargination; have a deep median excision; have a slight undulation; have two to four sharp to blunt teeth; or have a deep, irregular median depression. It also varies in orientation, usually projecting anteriorly, but sometimes projecting more ventrally, making it difficult to view.
The structure of the basal margin of the mandible usually correlates with the shape of the anterior clypeal margin (see Figure 19). This is because clypeal teeth, if present, usually merge with the basal margin of the mandible when the mandibles are closed. When the anterior clypeal margin is entire or only has a median emargination, the basal margin of the mandible is straight to slightly sinuous. When the anterior clypeal margin is undulating or forming teeth, the basal margin of the mandible is sinuous and usually has a distinct, shallow to deep basal depression. In some species, the basal depression is accompanied by a small tooth that projects inward from the proximal edge of the depression.
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.
Color. Overall body and appendage color and the contrast between the two can be very useful in separating some species. Body color ranges from jet black to pale yellow, with red-black, brown-black, orange-brown, or brown as common intermediate colors. Sometimes species have a mottled appearance, with areas of dark or light cuticle occurring irregularly. Appendages are almost always lighter in color, usually becoming very light toward the extremities. It is important to note that older museum specimens lose their color, often having a faded red-brown color (see Figures 73, 82, 85, 150).
Gastral pilosity. On a given Stenamma specimen, pilosity on the body dorsum is generally similar in form, but it is usually best viewed on the gastral tergites. Among species, pilosity varies in length, density, thickness, and whether or not it is separated into two distinct layers (Figure 5). There is a continuum from being long, sparse, and clearly single layered, to being short, dense, and clearly bilayered, with the bilayered pilosity consisting of a longer layer of suberect to subdecumbent setae and a shorter layer of subdecumbent to appressed setae. The suberect, upper layer of setae can sometimes be distinctly thickened (i.e. stout), and the lower layer can be either very sparse or very dense and thin, becoming pubescent. Using this character is complicated by the fact that some species have intermediate character states and some species show substantial intraspecific variation.
Comparison of gastral pilosity among several Stenamma species A Stenamma diversum (CASENT0606723) B Stenamma monstrosum (CASENT0621327) C Stenamma lagunum (CASENT0622371) D Stenamma schmidti (INB0003665417) E Stenamma longinoi (JTLC000007475).
Sculpture. In principal I adhere to the terminology and philosophy of
Comparison of several different sculpture patterns in Stenamma A Stenamma atribellum (CASENT0622351) B Stenamma tico (CASENT0622571) C Stenamma catracho (CASENT0621306) D Stenamma ochrocnemis (CASENT0603793).
Frontal lobes and posterior extension of the clypeus. There is significant variation in the width of the frontal lobes and the width of the depressed area in between the lobes (Figure 7). In some species the frontal lobes are strongly expanded laterally or dorsolaterally. This variation is characterized quantitatively with the measurement FLD and the index FLI. It is also described qualitatively by whether or not the underlying torular lobes are covered by the frontal lobes in full-face view. The area in between the frontal lobes, referred to here as the posterior extension of the clypeus, can be very narrow to quite wide, with the sides varying in shape from subparallel to strongly hour-glass shaped. An attempt to quantify this variation is made with the measurement PCW. Both of these characters are useful, but most species display intermediate values.
Comparison of the frontal lobes and posterior extension of the clypeus in several Stenamma species A Stenamma llama (CASENT0605479) B Stenamma cusuco (CASENT0622137) C Stenamma diversum (CASENT0606723).
Petiole and postpetiole shape. Many Stenamma species have very distinctive petiole and postpetiole shapes (Figure 8). Categories of qualitative descriptors include overall robustness, symmetry of the nodes (in profile), and sharpness and angle of node apices. I also attempt to capture variation quantitatively with length, width, and height measurements of each segment. Often these are most useful as ratios. For example, if the postpetiole is very wide in relation to the petiolar node, I include an index such as PPW/PW. The actual index used depends on the distinctive feature for a particular species.
Comparison of petiole and postpetiole shape in several Stenamma species A Stenamma crypticum (CASENT0603755) B Stenamma hojarasca (CASENT0622132) C Stenamma lobinodus (CASENT0605658).
Promesonotum shape. The shape of the promesonotum in profile is variable and can be very distinctive in some species (Figure 9). The average form is for it to be low-domed and roughly symmetrical from front to back, but it can become distinctly asymmetrical, with the apex shifted away from the midpoint, or with the anterior or posterior slope much longer than the other. It can also be distinctly bulging, becoming high-domed.
Propodeal spines. The propodeal spines in Stenamma can be absent to quite long and robust (Figure 9). To quantify this character I use the measurement PSL and the index PSI. I also describe them as absent, tuberculate, short, medium, or long. When absent, the juncture of the propodeal dorsum and declivity can be smoothly rounded or it can form a sharp angle.
Comparison of promesonotum and propodeal spines in several Stenamma species A Stenamma alas (CASENT0606832) B Stenamma muralla (CASENT0621311) C Stenamma crypticum (CASENT0603755) D Stenamma lobinodus (CASENT0605658) E Stenamma diversum (CASENT0606723).
The MAC is distributed from the southwestern U.S.A. (Arizona, New Mexico, Texas) to northwestern South America (Colombia, Ecuador) (Figure 1). It is known to co-occur with the Holarctic clade only in the southwestern U.S.A, where Stenamma huachucanum reaches the northern limit of its range. In this area of sympatry it is easy to separate the two clades based on the structure of the anterior clypeal margin and basal margin of the mandible (
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 Stenamma huachucanum and Stenamma manni, and with short basal carinulae in Stenamma atribellum sp. n. and Stenamma callipygium; HOC species commonly with basal carinulae and punctae).
A global diagnosis of the worker caste of Stenamma is presented in
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
Stenamma alas
Stenamma andersoni Branstetter sp. n. Southern Mexico.
Stenamma atribellum Branstetter sp. n. Honduras.
Stenamma brujita Branstetter sp. n. Mexico to Honduras.
Stenamma callipygium Branstetter sp. n. Guatemala.
Stenamma catracho Branstetter sp. n. Honduras.
Stenamma connectum Branstetter sp. n. Southern Mexico.
Stenamma crypticum Branstetter sp. n. Southern Mexico to Nicaragua.
Stenamma cusuco Branstetter sp. n. Honduras.
Stenamma diversum
Stenamma excisum Branstetter sp. n. Mexico to Honduras.
Stenamma expolitico Branstetter sp. n. Costa Rica.
Stenamma expolitum
Stenamma felixi
Stenamma hojarasca Branstetter sp. n. Southern Mexico to Honduras.
Stenamma huachucanum
Stenamma ignotum Branstetter sp. n. Southern Mexico to Guatemala.
Stenamma lagunum Branstetter sp. n. Northern Mexico.
Stenamma leptospinum Branstetter sp. n. Southern Mexico.
Stenamma llama Branstetter sp. n. Southern Mexico to Guatemala.
Stenamma lobinodus Branstetter sp. n. Mexico.
Stenamma longinoi Branstetter sp. n. Southern Mexico.
Stenamma manni
Stenamma maximon Branstetter sp. n. Southern Mexico to Honduras.
Stenamma megamanni Branstetter sp. n. Southern Mexico to Nicaragua.
Stenamma monstrosum Branstetter sp. n. Honduras to Nicaragua.
Stenamma muralla Branstetter sp. n. Honduras to Nicaragua.
Stenamma nanozoi sp. n. Honduras.
Stenamma nonotch Branstetter sp. n. Southern Mexico to Guatemala.
Stenamma ochrocnemis Branstetter sp. n. Southern Mexico to Honduras.
Stenamma pelophilum Branstetter sp. n. Mexico to Honduras.
Stenamma picopicucha Branstetter sp. n. Honduras to Nicaragua.
Stenamma saenzae Branstetter sp. n. Southern Mexico to Honduras.
Stenamma sandinista sp. n. Nicaragua.
Stenamma schmidti
Stenamma stictosomum Branstetter sp. n. Mexico to Honduras.
Stenamma tiburon Branstetter sp. n. Northern Mexico.
Stenamma tico Branstetter sp. n. Nicaragua to Panama.
Stenamma vexator Branstetter sp. n. Mexico.
Stenamma zelum Branstetter sp. n. Honduras to Panama.
| 1 | Anterior constriction of gaster, along with basal striae, distinctly elongate (Figure 10A, B) | 2 |
| – | Anterior constriction of gaster and basal striae not elongate (Figure 10C) | 3 |
| 2 (1) | Surface sculpture almost entirely smooth and shiny (Figure 11A); anterior clypeal margin in full-face view with a shallow median emargination (Figure 11A) (Honduras) | Stenamma atribellum sp. n. |
| – | Face and much of mesosoma with conspicuous carinulae (Figure 11B); median lobe of clypeus projecting over anterior clypeal margin, forming a well-defined median apex (Figure 11B) (Guatemala) | Stenamma callipygium sp. n. |
| 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 (Figure 12A); frontal lobes markedly expanded outward, completely obscuring torular lobes in full-face view (Figure 12A) (FLD 0.24–0.26, FLI 36–39); head and mesosoma densely sculptured with rugoreticulae and punctae; propodeal spines well-developed (Figure 12B) (SSL 0.16–0.19, PSI 1.8–2.2) (Honduras) | Stenamma cusuco sp. n. |
| – | Anterior clypeal margin in full-face view variable, not as above (Figure 19A–I); frontal lobes, body sculpture, and propodeal spines variable | 4 |
| 4 (3) | Face and promesonotal dorsum foveate to coarsely rugoreticulate (Figure 13A, B); pilosity on gastral tergites long, dense, and mostly suberect (Figure 13C); eyes relatively small (EL 0.09–0.13, REL 10–16), with 5–8 ommatidia at greatest diameter; 4-segmented antennal club indistinct; larger species (HL 0.87–1.20, HW 0.78–1.15, ML 1.15–1.62) | 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) (Figure 14A); propodeal spines reduced to sharp angles or small tubercles (Figure 14D) (SSL 0.12–0.16, PSI 1.0–1.3); somewhat smaller species (HL 0.87–1.03, HW 0.78–0.92, ML 1.15–1.38) (eastern Honduras to Panama) | Stenamma zelum sp. n. |
| – | Anterior clypeal margin with a median emargination that has four well-defined to completely effaced blunt teeth (Figure 14B, C); propodeal spines usually present, ranging from short tubercles to long robust spines (Figure 14E–G) (SSL 0.15–0.37, PSI 1.3–2.9); somewhat larger species (HL 0.90–1.20, HW 0.77–1.15, ML 1.15–1.62) (Mexico to western Honduras) | Stenamma brujita sp. n. |
| 6 (4) | Lateral apex of hypostomal bridge projecting ventrally as a subquadrate (Figure 15A, B) to broadly rounded (Figure 15C) lobe, visible behind base of mandible in profile view | 7 |
| – | Lateral margin of hypostomal bridge tapering to a narrow point, not visible in profile view (Figure 15D–F) | 11 |
| 7 (6) | Propodeal spines long and slender (SSL 0.17–0.22, PSI 1.9–2.2) (Figure 16A); face with a dense fan of carinulae extending out from frontal lobes toward posterior and lateral margins of head (Figure 16B); postpetiole appearing somewhat anteroposteriorly compressed (Figure 16A) (southern Mexico) | Stenamma leptospinum sp. n. (part) |
| – | 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 (Figure 17A, B); suberect setae on gastral tergites very stout, somewhat sparse, short (Figure 17C); propodeal spines present, short (SSL 0.16–0.19, PSI 1.4–1.6) (southern Mexico) | Stenamma longinoi sp. n. |
| – | 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 (Figure 18A) (PPH/PH 0.74–0.84, PW/PPW 0.76–0.89); pronotum usually mostly smooth and shiny, with only some vestigial rugulae (Figure 18A); smaller species (HL 0.72–0.83, HW 0.63–0.75, ML 0.89–1.08) (Honduras to Nicaragua) | Stenamma muralla sp. n. |
| – | Waist segments more robust, with postpetiole in profile sometimes bulging and usually more circular (Figure 18B–D) (PPH/PH ≥ 0.79, PW/PPW ≥ 0.87); pronotum sculpture variable, but usually more dense; larger species (HL ≥ 0.81, HW ≥ 0.70, ML ≥ 1.05) (Mexico to Nicaragua) | 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 (Figure 18B, C) (Mexico to Nicaragua) | Stenamma manni (part) |
| – | Eye usually larger (EL 0.14–0.19), with 8–11 ommatidia at greatest diameter; body color mostly black (Figure 18D) (southern Mexico to Nicaragua) | Stenamma megamanni sp. n. |
| 11 (6) | Basal margin of mandible straight to slightly sinuous, without a distinct basal notch or depression (Figure 19A–C); anterior clypeal margin entire (Figure 19A), or with a rounded median emargination (Figure 19B), or with a deep median excision (Figure 19C) (rarely with mandible completely straight and anterior clypeal margin with 2 blunt teeth) | 12 |
| – | Basal margin of mandible sinuous, with a shallow to deep basal depression or notch, but without a basal tooth (Figure 19D–F); anterior clypeal margin undulating, forming 2–4 variably developed teeth (Figure 19D–F) (sometimes teeth very reduced, so clypeal margin is nearly flat) | 30 |
| – | Basal margin of mandible sinuous, with a distinct basal depression or notch, and an accompanying small tooth (Figure 19G–I); anterior clypeal margin usually undulating, forming 2–4 variably developed teeth (Figure 19D–I) | 41 |
| 12 (11) | Propodeal spines absent; petiole and postpetiole almost completely smooth and shiny, with only faint vestigial punctae sometimes present (Figure 20A–D); postpetiole in profile bulging, globular, appearing more voluminous than petiolar node (Figure 20A–D) | (expolitum group) 13 |
| – | 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 (Figure 21A); carinulae around frontal lobes extending to about midpoint of head or less; dorsum of promesonotum smooth and shiny (Nicaragua to Costa Rica) | Stenamma expolitum |
| – | 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 (Figure 21B–D); dorsum of promesonotum smooth and shiny, or with variably developed transverse furrows or striae (Costa Rica to Ecuador) | 14 |
| 14 (13) | Dorsal and declivitous faces of propodeum in profile flat, and forming a blunt 90° angle (Figure 20D); dorsum of promesonotum with distinctive transverse furrows, sometimes reticulate posteriad (Figure 22C) (Costa Rica) | Stenamma expolitico sp. n. |
| – | Propodeum in profile more rounded, with transition between dorsal and declivitous faces less abrupt (Figure 20B, C); dorsum of promesonotum either smooth and shiny, or with few to many transverse striae (Figure 22B, C) (Costa Rica to Ecuador) | Stenamma alas |
| 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 (Figure 23A, B) | 16 |
| – | Face sculpture more extensive, usually completely sculptured; type of sculpture variable (Figure 23C) | 21 |
| 16 (15) | Dorsum of promesonotum mostly smooth and shiny, except for scattered piligerous punctae, and at most a few longitudinal rugulae (Figure 24A, B) | 17 |
| – | Dorsum of promesonotum with more extensive sculpturing, either reticulately costate or rugoreticulate (Figure 24C) | 19 |
| 17 (16) | Mesosoma compact, with promesonotum somewhat bulging (Figure 25A); propodeal spines present as short robust triangles (Figure 25A) (SSL 0.09–0.14, PSI 1.7–2.5); petiolar node robust, relatively tall (PH/PL 0.66–0.80), and distinctly angled posteriad (Figure 25A); scape and metafemur relatively short (SI 80–85, MFI 101–113) (southern Mexico to Honduras) | Stenamma llama sp. n. |
| – | Lacking one or more of the above character states (Figure 25B–D) | 18 |
| 18 (17) | Median lobe of clypeus bicarinate, projecting, and with area between carinae distinctly depressed (Figure 26A); promesonotum in profile asymmetrical, with anterior slope long and gently curving, dorsum nearly flat, and posterior slope short and forming a sharp transition with dorsum (Figure 25B); propodeal spines absent (SSL 0.06–0.07, PSI 1.0–1.3) (northern Mexico) | Stenamma tiburon sp. n. |
| – | Median lobe of clypeus smooth and with a simple median emargination (Figure 26B); promesonotum in profile usually asymmetrical (Figure 25C), with posterior slope distinctly longer than anterior slope, but sometimes promesonotum more evenly domed (Figure 25D); propodeal spines tuberculate to short (SSL 0.06–0.10, PSI 1.1–1.7) (Mexico to Honduras) | Stenamma pelophilum sp. n. |
| 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 (Figure 27A, B); postpetiolar node in profile with a longitudinal dorsal lobe that projects posteriorly over postpetiole (Figure 27A, B) (Mexico) | Stenamma lobinodus sp. n. |
| – | Promesonotum in profile low domed and roughly symmetrical (Figure 27C, D); postpetiole in profile more circular and without a distinct dorsal lobe projecting posteriorly (Figure 27B, C) | 20 |
| 20 (19) | Propodeal spines present, long (Figure 27C) (SSL 0.28–0.34, PSI 3.0–3.7); frontal lobes expanded dorsolaterally, usually completely covering torular lobes in full-face view (Figure 28A) (FLD 0.25–0.29, FLI 35–38); eye smaller (EL 0.11–0.15, REL 16–20) (southern Mexico to Nicaragua) | Stenamma diversum |
| – | Propodeal spines reduced to sharp right angles, or small upward projecting tubercles (Figure 27D) (SSL 0.14–0.18, PSI 1.4–1.9); frontal lobes narrower, with torular lobes partly visible in full-face view (Figure 28B, C) (FLD 0.22-0.27, FLI 33-36); eye larger and somewhat bulging (EL 0.15-0.18, REL 23–24) (Nicaragua to Panama) | Stenamma tico sp. n. (part) |
| 21 (15) | Propodeal spines absent, with the dorsal and declivitous faces of the propodeum forming a relatively shallow, blunt angle in profile view (Figure 29A, B) (SSL 0.08–0.11, PSI 0.8–1.1); eyes large (EL ≥ 0.15, REL ≥ 18) | 22 |
| – | Propodeal spines varying from tuberculate to long, or with the dorsal and declivitous faces of the propodeum forming a sharp, steep angle (Figure 29C–E) (SSL ≥ 0.07, PSI ≥ 1.0); eyes variable | 23 |
| 22 (21) | Head and mesosoma strongly sculptured (Figure 29A, 30A, B); face mostly rugoreticulate or carinate; scape relatively shorter (SI 84–101); eye relatively smaller (REL 18–22); larger, more robust species (HL 0.90–1.19, HW 0.81–1.04, ML 1.25–1.62) (Mexico to Ecuador) | Stenamma felixi |
| – | Head and mesosoma more faintly sculptured (Figure 29B, 30C, D); face with variable amount of carinulae, rugulae, and punctae; scape relatively longer (SI 107–121); smaller, usually somewhat gracile species (HL 0.79–0.88, HW 0.65–0.73, ML 1.04–1.16) (Nicaragua to Ecuador) | Stenamma schmidti (part) |
| 23 (21) | Facial sculpture light, largely effaced, usually not extending all the way to posterior margin of head (Figure 31A); mesosoma mostly reticulately costate (Figure 31B); propodeal spines reduced to sharp right angles, or small upward projecting tubercles (Figure 31B) (PSL 0.14–0.18, PSI 1.4–1.9); eye large, somewhat bulging (EL 0.15-0.18, REL 23–24) (Nicaragua to Panama) | Stenamma tico sp. n. (part) |
| – | 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 (Figure 32A, B); lateral margin of hypostomal bridge tapering to a point, never visible in profile view (Figure 15D–F) | 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 (Figure 32C); lateral margin of hypostomal bridge usually broadly rounded and somewhat projecting, often visible in profile view (Figure 15C) | 29 |
| 25 (24) | Eye small (EL 0.04–0.08, REL 8–14), subcircular, with 2–5 ommatidia at greatest diameter (Figure 32A) | 26 |
| – | Eye larger (EL 0.09–0.12, REL 15–21), more oval-shaped, with 4–7 ommatidia at greatest diameter (Figure 32B) | 27 |
| 26 (25) | Sculpture on face (and much of mesosoma) consisting of an even distribution of short longitudinal rugulae (Figure 33A); body color a mottled yellow-brown (often pale yellow), with patches of darker brown; median clypeal lobe in full-face view visible and well developed (Figure 33A); anterior clypeal margin with a shallow median emargination (northern Mexico) | Stenamma lagunum sp. n. |
| – | Face densely sculptured and mostly rugoreticulate (Figure 33B, C); mesosoma densely sculptured with punctae, rugae, and/or rugoreticulae; body color usually a darker orange-brown; clypeus in full-face view appearing very short, with median lobe nearly invisible due its dorsoventral orientation (Figure 33B, C); anterior clypeal margin often with a deep median excision (Figure 33B), but sometimes only with a very weak median depression (Figure 33C) (Mexico to Honduras) | Stenamma excisum sp. n. |
| 27 (25) | Anterior clypeal margin either entire, or with a nearly imperceptible median notch (Figures 19A, 34A); dorsum of promesonotum either rugoreticulate, or with many irregular rugulae (Figure 34B); petiolar node often broadly rounded and pointing distinctly posteriad (Figure 34C) (southern Mexico to Guatemala) | Stenamma nonotch sp. n. |
| – | Anterior clypeal margin not as above (Figure 25D, G); dorsum of promesonotum with relatively dense longitudinal rugulae/carinulae (Figure 25E, H); petiolar node not as above (Figure 25F, I) | 28 |
| 28 (27) | Anterior clypeal margin with a shallow, but distinct median emargination (Figure 34D); petiole usually appearing more elongate (Figure 34F); petiolar node usually sharper and pointing more vertically; pilosity on gastral tergites longer and mostly forming a relatively sparse layer of suberect setae, only sometimes with decumbent setae (Figure 34F); metafemur relatively longer (MFI 96–104) (southern Mexico to Honduras) | Stenamma ignotum sp. n. |
| – | Anterior clypeal margin forming 2 small blunt teeth, which straddle the midline (Figure 34G); petiole more compact, with the node less sharp and pointing more strongly posteriad (Figure 34I); pilosity on gastral tergites somewhat shorter and usually distinctly bilayered, with a layer of suberect setae, and a layer of decumbent setae (Figure 34G); metafemur relatively shorter (MFI 104–110) (Honduras to Nicaragua) | Stenamma picopicucha sp. n. |
| 29 (24) | Propodeal spines long and slender (Figure 29E) (SSL 0.17–0.22, PSI 1.9–2.2); face with a dense fan of carinulae extending out from frontal lobes toward posterior and lateral margins of head (Figure 35A); postpetiole appearing somewhat anteroposteriorly compressed (Figure 29E) (southern Mexico) | Stenamma leptospinum sp. n. (part) |
| – | Propodeal spines tuberculate to short (Figure 29D) (SSL 0.09–0.19, PSI 1.0–1.6); face sculpture variable, but usually mostly rugoreticulate, and never densely carinulate (Figure 35B, C); postpetiole usually more circular, not anteroposteriorly compressed (Figure 29D) (Mexico to Nicaragua) | Stenamma manni (part) |
| 30 (11) | Posterior ¼ or more of face smooth and shiny (Figure 36A, B) | 31 |
| – | Face completely sculptured, mostly rugoreticulate (Figure 36C, D) | 33 |
| 31 (30) | Promesonotum in profile with relatively sharp transitions between anterior and dorsal faces, and between pronotum and mesonotum (Figure 37B); pilosity on gastral tergites forming a layer of stout suberect setae, and a sparse layer of decumbent setae (Figure 37C); face and pronotum almost completely smooth and shiny (Figures 37A, B) (southern Mexico) | Stenamma andersoni sp. n. |
| – | 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 (Figure 38A) (PPH 0.20–0.25, PPW 0.22–0.27, PPH/PH 0.96–1.14); eye larger (EL 0.11–0.16); face sculpture variable, but usually mostly smooth and shiny, with some carinulae around frontal lobes and anterolateral margins of head (Figure 36A); larger species (HL 0.67–0.86, HW 0.57–0.76, ML 0.84–1.09) (southern Mexico to Honduras) | Stenamma maximon sp. n. (part) |
| – | Postpetiole smaller, about same size as petiolar node (Figure 38B, C) (PPH 0.14–0.19, PPW 0.15–0.20, PPH/PH 0.79–0.97); eye smaller (EL 0.07–0.12); face sculpture usually more developed, with carinulae, punctae, and occasional rugoreticulae (Figure 36B); smaller species (HL 0.55–0.72, HW 0.46–0.64, ML 0.66–0.85) (southern U.S.A. to Mexico) | Stenamma huachucanum (part) |
| 33 (30) | Pilosity on gastral tergites predominately suberect and relatively sparse (Figure 39A, B); decumbent setae if present very 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 (Figure 39C–E) | 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 (Figure 40A–C); eye larger (EL 0.15–0.18, REL 20–25), with 8–9 ommatidia at greatest diameter; propodeal spines absent to tuberculate, shorter (SSL 0.09–0.11, PSI 0.9–1.2) (Mexico to Honduras) | Stenamma stictosomum sp. n. |
| – | Face less densely sculptured, usually mostly rugoreticulate, but sometimes sculpture more polished, with reticulae indistinct (Figure 40D, E); eye smaller (EL 0.10–0.15, REL 18–22), with 5–8 ommatidia at greatest diameter; propodeal spines tuberculate to short, longer (SSL 0.08–0.16, PSI 1.1–1.8) (Mexico) | Stenamma vexator sp. n. |
| 35 (33) | Propodeal declivity in profile forming a broadly sinuous connection between propodeal spine and propodeal lobe (Figure 41A) (southern Mexico [Oaxaca, Veracruz]) | Stenamma connectum sp. n. |
| – | Propodeal declivity in profile straighter, leaving propodeal spine and lobe separated as distinct features (Figures 41B, 42A–F) | 36 |
| 36 (35) | Petiole in profile appearing longer and more gracile, with node either dome-like and almost completely smooth and shiny (Figure 42A), or small and somewhat compressed dorsoventrally (Figure 42B); pronotum mostly sculptured, with punctae on side (Figure 42A, B), and rugae or rugoreticulae on dorsum; frontal lobes often (but not always) distinctly expanded, covering the torular lobes in full-face view (FLD 0.15–0.25, FLI 28–46); propodeal spines present, short to medium length (SSL 0.11–0.17, PSI 1.6–2.1) | 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 (Figure 42C–F); side of pronotum variably sculptured, often mostly smooth, and usually not punctate (Figure 42C–F); frontal lobes almost always of average width, and never completely covering torular lobes in full-face view (FLD 0.11–0.20, FLI 23–31); propodeal spines variable, usually shorter (SSL 0.06–0.14, PSI 1.0–1.8) | 38 |
| 37 (36) | Petiole relatively longer (PL/HW 0.60–0.68); petiolar and postpetiolar nodes mostly smooth and shiny (Figure 42A); postpetiole relatively smaller and appearing more shield-like (PPH/PH 0.75–0.84); scape and metafemur relatively shorter (SI 82–89, MFI 102–109); larger species (HL 0.65–0.76, HW 0.54–0.66, ML 0.81–0.99) (southern Mexico to Honduras) | Stenamma hojarasca sp. n. |
| – | Petiole relatively shorter (PL/HW 0.53–0.59); petiolar and postpetiolar nodes smooth only on anterior faces; postpetiole relatively larger (Figure 42B) (PPH/PH 0.85–0.91); scape and metafemur relatively longer (SI 92–104, MFI 84–95); smaller species (HL 0.61–0.68, HW 0.51–0.59, ML 0.75–0.82) (Honduras) | Stenamma catracho sp. n. |
| 38 (36) | Postpetiole in profile bulging and distinctly larger than petiolar node (Figures 38A, 42C) (PPH 0.20–0.25, PPW 0.22–0.27, PPH/PH 0.96–1.14); eye larger (EL 0.11–0.16), with 6–8 ommatidia at greatest diameter (southern Mexico to Honduras) | Stenamma maximon sp. n. (part) |
| – | Postpetiole in profile smaller, about same size as petiolar node or smaller (Figure 42D–F) (PPH 0.12–0.20, PPW 0.13–0.22, PPH/PH 0.79–1.04); eye smaller (EL 0.07–0.12), with 4–8 ommatidia at greatest diameter (usually ≤ 6) | 39 |
| 39 (38) | Head and mesosoma dark red-brown to orange-brown, with appendages a distinctly lighter orange- to yellow-brown (Figure 43A); basal margin of mandible with basal depression, shallow or deep (Figure 11F); eye often relatively smaller (REL 12–17); larger species (HL 0.63–0.83, HW 0.54–0.73, ML 0.76–1.09 PrW 0.37–0.50) (southern Mexico to Honduras) | Stenamma ochrocnemis sp. n. |
| – | Body and appendage color less contrasting, generally dark to light brown (Figure 43B); basal margin of mandible with basal depression shallow, never deep; eye often relatively larger (REL 14–21); smaller species (HL ≤ 0.72, HW ≤ 0.64, ML ≤ 0.85, PrW ≤ 0.41) | 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 (Figures 42E, 43B) (note geography is easiest way to separate species in this couplet) (southern Mexico [Chiapas] to Nicaragua) | Stenamma crypticum sp. n. |
| – | 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 (Figure 42F) (southern U.S.A. to southern Mexico [Oaxaca]) | Stenamma huachucanum (part) |
| 41 (11) | Mesosoma in profile somewhat elongate, with metanotal groove wide, shallow, and indistinct, and propodeal dorsum markedly long and flat (Figure 44A); anterior clypeal margin in full-face view with a deep uneven median emargination (Figure 44B); basal tooth of mandible very robust and well-defined; eye relatively small (EL 0.09–0.11, REL 13–15), subcircular, with 5–6 ommatidia at greatest diameter (Honduras to Nicaragua) | Stenamma monstrosum sp. n. |
| – | 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) | Stenamma schmidti (part) |
| – | 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 (Figure 45A); pilosity on gastral tergites forming a layer of longer suberect setae and a sparse layer of shorter decumbent setae, all setae of moderate thickness (Figure 46A); larger species (HL 0.62–0.73, HW 0.56–0.70, ML 0.77–0.89) (Nicaragua) | Stenamma sandinista sp. n. |
| – | Face completely sculptured, mostly rugoreticulate (Figure 45B, C); gastral pilosity variable, but not as above (Figure 46B, C); usually smaller species (HL ≤0.69, HW ≤0.59, ML ≤0.82) (southern Mexico to Honduras) | 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 (Figure 46B); propodeal spines tuberculate to short, longer (SSL 0.07–0.14, PSI 1.5–2.3); eye slightly smaller (EL 0.05–0.09, REL 10–16), subcircular, with 3–5 ommatidia at greatest diameter; scape and metafemur relatively shorter (SI 81–92, MFI 107–119); frontal lobes, not expanded outward (FLD 0.11–0.14, FLI 22–27) (southern Mexico to Honduras) | Stenamma saenzae sp. n. |
| – | Pilosity on gastral tergites forming a layer of stout suberect setae and a very sparse layer of appressed setae (Figure 46C); propodeal spines reduced to sharp angles or small tubercles, shorter (SSL 0.07–0.09, PSI 1.2–1.4); eye larger (EL 0.08–0.10, REL 15–18), more oval-shaped, with 4–5 ommatidia at greatest diameter; scape and metafemur longer (SI 90–99, MFI 97–104); frontal lobes slightly expanded outward (FLD 0.14–0.15, FLI 28–31) (Honduras) | Stenamma nanozoi sp. n. |
Anterior constriction of gaster in dorsal view A Stenamma callipygium (CASENT0606207) B Stenamma atribellum (CASENT0622351) C Stenamma manni (CASENT0605527).
Face and anterior clypeal margin in full-face view A Stenamma atribellum (CASENT0622351) B Stenamma callipygium (CASENT0606207).
Stenamma cusuco (CASENT0622137) A Anterior clypeal margin in full-face view B Body in profile.
Stenamma zelum (CASENT0622535) A Face B Dorsum of promesonotum C Gastral pilosity.
Comparison of clypeal structure (anterodorsal view) and mesosoma (profile) between A, D Stenamma zelum (CASENT0622535) B Stenamma brujita (CASENT0126254) C Stenamma brujita (CASENT0604945) E Stenamma brujita (CASENT0126254) F Stenamma brujita (CASENT0604945) G Stenamma brujita (CASENT0604607).
Lateral apex of hypostomal bridge A Stenamma megamanni (CASENT0622853) head in profile view B Stenamma megamanni (CASENT0622853) head in lateroventral view C Stenamma manni (CASENT0604893) in lateroventral view D Stenamma felixi (CASENT0620969) head in profile view E Stenamma felixi (CASENT0620969) head in lateroventral view F Stenamma maximon (CASENT0604675) head in lateroventral view.
Stenamma leptospinum (CASENT0605530) A Mesosoma and waist in profile B Face.
Stenamma longinoi (JTLC000007475) A Mesosoma in profile B Dorsum of promesonotum C Gastral pilosity.
Mesosoma and waist in profile. A Stenamma muralla (CASENT0621311) B Stenamma manni (CASENT0621574) C Stenamma manni (CASENT0605592), D Stenamma megamanni (CASENT0604730).
Variation in the structure of the anterior clypeal margin and the basal margin of the mandible A Stenamma nonotch (CASENT0604711) B Stenamma muralla (CASENT0621311) C Stenamma excisum (CASENT0605563) D Stenamma crypticum (CASENT0603821) E Stenamma maximon (CASENT0604875) F Stenamma ochrocnemis (CASENT0621468) G, H Stenamma schmidti (INB0003665417) I Stenamma saenzae (CASENT0604912).
Mesosoma and waist in profile A Stenamma expolitum (CASENT0600043) B Stenamma alas (JTLC000005880) C Stenamma alas (CASENT0606832) D Stenamma expolitico (INBIO282473).
Face sculpture A Stenamma expolitum (CASENT0600043) B Stenamma alas (JTLC000005880) C Stenamma alas (CASENT0606832) D Stenamma expolitico (INBIO282473).
Dorsum of promesonotum A Stenamma alas (JTLC000005880) B Stenamma alas (CASENT0606832) C Stenamma expolitico (INBIO282473).
Face sculpture A Stenamma lobinodus (CASENT0622422) B Stenamma tico (CASENT0622416) C Stenamma nonotch (CASENT0604711).
Sculpture on dorsum of promesonotum A Stenamma pelophilum (CASENT0605613) B Stenamma llama (CASENT0604952) C Stenamma lobinodus (CASENT0605658).
Mesosoma and waist in profile view A Stenamma llama (CASENT0605236) B Stenamma tiburon (CASENT0620965) C Stenamma pelophilum (CASENT0605613) D Stenamma pelophilum (CASENT0605428).
Clypeus in full-face view A Stenamma tiburon (CASENT0620965) B Stenamma pelophilum (CASENT0606223).
Mesosoma and waist in profile A Stenamma lobinodus (CASENT0605658) B Stenamma lobinodus (CASENT0605814) C Stenamma diversu m (CASENT0606723) D S. tico (CASENT0622571).
Face A Stenamma diversum (CASENT0606723) B Stenamma tico (CASENT0622416) C Stenamma tico (CASENT0600104).
Mesosoma in profile A Stenamma felixi (CASENT0620969) B Stenamma schmidti (INB0003210597) C Stenamma nonotch (CASENT0605789)D Stenamma manni (CASENT0604893) E Stenamma leptospinum (CASENT0605530).
Face sculpture A Stenamma felixi (CASENT0620969) B Stenamma felixi (CASENT0622555) C Stenamma schmidti (INB0003210597) D Stenamma schmidti (INB0002659320).
Stenamma tico (CASENT0600104) A Face B Body in profile.
Head and eye in profile view A Stenamma excisum (CASENT0605563) B Stenamma ignotum (CASENT0603762) C Stenamma leptospinum (CASENT0605530).
Faceand clypeus in full-face view A Stenamma lagunum (CASENT0622371) B Stenamma excisum (CASENT0621834) C Stenamma excisum (CASENT0605441).
Comparison of the anterior clypeal margin, pronotal dorsum, waist, and gaster A–C Stenamma nonotch (CASENT0605789) D–F Stenamma ignotum (CASENT0603762) G–I Stenamma picopicucha (CASENT0606709).
Face sculpture A Stenamma leptospinum (CASENT0605530) B Stenamma manni (CASENT0605592) C Stenamma manni (CASENT0604893).
Face sculpture A Stenamma maximon (CASENT0603886) B Stenamma huachucanum (CASENT0605616) C Stenamma catracho (CASENT0621306) D Stenamma ochrocnemis (CASENT0605129).
Stenamma andersoni (CASENT0604603) A Head B Mesosoma in profile C Gastral pilosity.
Petiole and postpetiole in profile view A Stenamma maximon (CASENT0603886) B Stenamma huachucanum (CASENT0605616) C Stenamma huachucanum (CASENT0605647).
Gastral pilosity A Stenamma stictosomum (CASENT0605499) B Stenamma vexator (CASENT0126485) C Stenamma crypticum (CASENT0605185) D Stenamma hojarasca (CASENT0622132) E Stenamma ochrocnemis (CASENT0621468).
Face sculpture A Stenamma stictosomum (CASENT0605499) B Stenamma stictosomum (CASENT0606221) C Stenamma stictosomum (CASENT012624) D Stenamma vexator (CASENT0604641) E Stenamma vexator (CASENT0605506).
Mesosoma in profile A Stenamma huachucanum (CASENT0605586) B Stenamma crypticum (CASENT0603755).
Mesosoma and waist in profile A Stenamma hojarasca (CASENT0622132) B Stenamma catracho (CASENT0621306)C Stenamma maximon (CASENT0605063)D Stenamma ochrocnemis (CASENT0603793) E Stenamma crypticum (CASENT0603821) F Stenamma huachucanum (CASENT0126556).
Body and appendages in profile view A Stenamma ochrocnemis (CASENT0621468) B Stenamma crypticum (CASENT0603821).
Stenamma monstrosum (CASENT0621327) A Mesosoma in profile B Clypeus and mandibles in full-face view.
Face sculpture A Stenamma sandinista (CASENT0622578)B Stenamma nanozoi (CASENT0621828) C Stenamma saenzae (CASENT0604912).
Gastral pilosity A Stenamma sandinista (CASENT0622578) B Stenamma saenzae (CASENT0603860) C Stenamma nanozoi (CASENT0621828).
http://species-id.net/wiki/Stenamma_alas
Worker: Figures 47, 48; Queen: Figure 49A–D; Male: Figure 49E–G; Map: Figure 50Integument 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. Similar species: Stenamma expolitico, Stenamma expolitum.
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.
Stenamma alas paratype worker (JTLC000005880) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma alas worker variants. Head, profile, and dorsal views A–C Variant 1 (CASENT0600114) D–F Variant 2 (CASENT0606832).
(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 Figure 49D.
Stenamma alas A Queen (CASENT000005886), profile B Same, face C Same, dorsum D Queen (CASENT0623104), wings E Male (CASENT0623103), profile F Same, face G Same, dorsum.
See Figure 49E–G.
The nesting biology of Stenamma alas is described in detail in
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 Stenamma alas worker quickly emerges from the nest and closes the entrance with the pebble. It is hypothesized that all of these complex nesting behaviors evolved to avoid predation by army ants.
Each Stenamma alas nest contains a single small chamber. Colonies are fairly large for Stenamma, with up to 250 individuals. All excavated colonies have contained only one egg-laying queen. Foragers are solitary, slow moving and freeze when disturbed. It is unknown what Stenamma alas forages on primarily, but I have observed workers returning to the nest with cookie baits and small pieces of unidentified organic matter, suggesting that the species might be a generalist scavenger.
Stenamma alas, along with Stenamma expolitico and Stenamma expolitum, belongs to the expolitum species group (a diagnosis of this group is given under Stenamma expolitum below). Stenamma alas is easily separated from Stenamma expolitico and Stenamma expolitum by comparing the sculpturing on the face and promesonotal dorsum. In the field, the holotype form of Stenamma alas (discussed below) can be separated from Stenamma expolitum by the structure of the nest entrance. Stenamma alas nests always have a horizontal turret, whereas Stenamma expolitum nests have vertical turrets.
As I have circumscribed it here, Stenamma alas represents a complex of species, whose boundaries are not clear. The type form of Stenamma alas (Figure 47) occurs only in Costa Rica. It is characterized by the following: facial carinulae extending to about midpoint of head, but not further; pronotum completely smooth and shiny; legs uniformly orange-brown. Variant 1 (Figure 48A–C) differs from the holotype form as follows: facial carinulae more extensive, sometimes very dense and extending all the way to the posterior margin of the head; legs dark brown to brown; dorsum of promesonotum with variably developed transverse striations. Variant 2 (Figure 48D–F) is the same as variant 1 except that the promesonotum in profile is strongly bulging upward, appearing high-domed. Variant 2 is known only from a few localities in the Bocas del Toro and Chiriquí provinces of Panama. It does not occur in sympatry with the other forms and some specimens appear intermediate, with the promesonotum less bulging. There is also variation in how dense and long the facial carinulae appear.
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 Stenamma alas (Donoso, pers. comm.).
Molecular phylogenetic data show that variant 1 specimens from Costa Rica form a clade sister to Stenamma expolitum and Stenamma expolitico (Branstetter unpublished data). Specimens from Ecuador form a clade sister to Stenamma alas. No specimens from Panama have been sampled yet. This result suggests that the variant 1 specimens in Costa Rica are distinct from the type form of Stenamma alas and the specimens in Ecuador, but I cannot tell them apart based on worker or queen morphology. Thus, I treat Stenamma alas as a paraphyletic species, but acknowledge that it could include multiple cryptic taxa. More morphological and molecular data will be needed to resolve this problematic species.
COSTA RICA: Alajuela : 3km E Monteverde, 10.300°N, 84.7833°W, 1400m, 26 Apr 1990 (J. Longino); Río Peñas Blancas, 10.3167°N, 84.7167°W, 800m, 4 Mar 2004 (J. Longino); Heredia : La Selva Biological Station, 10.43047°N, 84.00675°W, 100m, 5 Jun 2007 (M. G. Branstetter); 8km ENE Vara Blanca, 10.20°N, 84.10°W, 1800m, 16 Apr 2002 (ALAS); 9km NE Vara Blanca, 10.233°N, 84.083°W, 1500m, 8 Mar 2005 (J. Longino); 10km NE Vara Blanca, 10.233°N, 84.083°W, 1500m, 12 Feb 2005 (ALAS); 13km NE Vara Blanca, 10.2667°N, 84.0833°W, 1100m, 16 Apr 2001 (ALAS); 11km ESE La Virgen, 10.35°N, 85.05°W, 300m, 15 Apr 2004 (J. Longino); 12km N Vol. Barba, 10.250°N, 84.083°W, 1420m, 10 Jul 1986 (J. Longino); 13km N Vol. Barba, 10.250°N, 84.083°W, 1320m, 10 Jul 1986 (J. Longino); Puntarenas : Las Alturas Biological Station, 8.94997°N, 82.83375°W, 1800m, 26 May 2007 (M. G. Branstetter); Monteverde, 10.30°N, 84.80°W, 1400m, Apr–May 1987 (S. Little); ECUADOR: Pichincha : Otongachi, 0.313°N, 78.950°W, 850m, 6 Aug 2009 (G. Ramón); Río Toachi, 4km W La Palma, 0.3183°N, 78.9533°W, 870m, 25 Jan 2006 (D. A. Donoso); PANAMA: Bocas del Toro : Fortuna-Chiriquí Grande Rd., 8.78333°N, 82.1833°W, 800m, 16 Jul 1987 (D. M. Olson); Sendero Divisa, 15km SSW Chiriquí Grande, 8.783°N, 82.200°W, 1250m, 9 Jul 1987 (D. M. Olson); Chiriquí : El Mirador, Finca Collins, nr Boquete, [ca. 8.813°N, 82.484°W], 1830m, 26 Jun 1976 (A. F. Newton).
Distribution map of Stenamma alas (circles), Stenamma andersoni (squares), and Stenamma atribellum (triangles).
urn:lsid:zoobank.org:act:6EC8C7EC-60F7-47A9-8099-89D349DC73C1
http://species-id.net/wiki/Stenamma_andersoni
Worker: Figure 51; Map: Figure 50Holotype worker. MÉXICO, Oaxaca: 10.9km N. Candelaria, [ca. 16.220°N, 95.891°W], 990m, 12 Jul 1987, cloud forest, ex sifted leaf litter (R. S. Anderson, collection RSA87-15) [USNM, specimen CASENT0604603]. Paratype: same data as holotype [1w, UNAM, CASENT0604602].
Worker diagnosis. Integument brown to red-brown (note that observed specimens are old, fresh specimens probably darker); small-sized species (see HL, ML, PrW below); anterior clypeal margin undulating, forming 2–4 small blunt teeth; basal margin of mandible sinuous, with a shallow basal depression, but without a basal tooth; head and pronotum mostly smooth and shiny; promesonotum in profile with relatively sharp transitions between anterior and dorsal faces, and between pronotum and mesonotum; pilosity on gastral tergites forming a layer of stout suberect setae, and a sparse layer of decumbent setae; eye of moderate size (EL 0.08–0.09, PSL 0.08–0.10), oval-shaped, with 5 ommatidia at greatest diameter; propodeal spines tuberculate (PSL 0.08–0.10, PSI 1.2–1.3); frontal lobes of moderate width (FLD 0.12–0.13, FLI 23–24), not obscuring torular lobes in full-face view. Similar species: Stenamma connectum, Stenamma crypticum, Stenamma huachucanum, Stenamma maximon.
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.
Stenamma andersoni holotype worker (CASENT060460) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Unknown.
Unknown.
This species is recorded from a single Berlese sample of sifted leaf litter collected in cloud forest at 990 m elevation.
Stenamma andersoni should be easy to separate from similar species by its smooth head and pronotum, unique pronotum shape, and thickened gastral setae.
Stenamma andersoni is known from only two specimens collected in 1987. As a result the specimens are somewhat faded in color and were not useable for molecular phylogenetic work. With morphology alone, it is not clear to which species Stenamma andersoni is most closely related, but I hypothesize that it is probably near Stenamma crypticum or Stenamma huachucanum.
Known only from the type locality.
urn:lsid:zoobank.org:act:D989706A-4AFF-47FB-AC6A-6F6C0718E78B
http://species-id.net/wiki/Stenamma_atribellum
Worker: Figure 52; Queen: Figure 53; Map: Figure 50Holotype worker. HONDURAS: Cortés, Parque Nacional Cusuco, 15.50739°N, 88.23373°W ±20m, 2030m, 3 Jun 2010, cloud forest, nest under bark of log (M. G. Branstetter, collection MGB1606) [USNM, specimen CASENT0622351]. Paratypes: same data as holotype [1w, CAS, CASENT0623237], [1w, EAPZ, CASENT0623238], [1w, ECOSCE, CASENT0623239], [1w, FMNH, CASENT0623240], [1w, ICN, CASENT0623241], [1w, INBio, CASENT0623242], [1w, JTLC, CASENT0623525], [1w, LACM, CASENT0623243], [1w, MGBPC, CASENT0623525], [1w, MCZ, CASENT0623244], [1w, MZSP, CASENT0623245], [1w, UCD, CASENT0623246], [1w, UNAM, CASENT0623247], [1dq, 1w, USNM, CASENT0622349], [1w, UVGC, CASENT0623077].
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. Similar species: Stenamma alas, Stenamma callipygium, Stenamma expolitum.
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.
Stenamma atribellum holotype worker (CASENT0622351) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
(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.
Stenamma atribellum paratype queen (CASENT0622349) A Profile B Face C Dorsum.
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.
Stenamma atribellum is a distinctive species that should not be confused with any other MAC species. Molecular phylogenetic data show that it is sister to Stenamma callipygium, which is the only other Stenamma species to have an elongate anterior gastral constriction (Branstetter unpublished data). This character joins Stenamma atribellum and Stenamma callipygium in the atribellum species group. Stenamma atribellum can be separated from Stenamma callipygium by its completely smooth and shiny sculpture and emarginate anterior clypeal margin. Furthermore, these species have not been collected in sympatry and both appear to be narrow endemics.
Stenamma atribellum might be confused with the superficially similar Stenamma alas and Stenamma expolitum, which are both mostly smooth and shiny. However, these latter species do not have the anterior constriction of the gaster elongate and they tend to have more facial sculpture (especially in Stenamma alas). These species are also geographically separated from each another.
HONDURAS: Cortés : Parque Nacional Cusuco, 15.50739°N, 88.23373°W, 2030m, 3 Jun 2010 (M. G. Branstetter); 25km N Cofradia, PN Cusuco, [ca. 15.497°N, 88.227°W], 1550m, 26 Aug 1994 (S. & J. Peck).
urn:lsid:zoobank.org:act:746AA110-6E0F-4B6B-8718-46F8F6ABF231
http://species-id.net/wiki/Stenamma_brujita
Worker: Figures 54, 55; Queen: Figure 56; Map: Figure 57Holotype worker. GUATEMALA, Zacapa: 2km SE La Unión, 14.94706°N, 89.27660°W ±50m, 1550m, 12 May 2009, cloud forest, ex sifted leaf litter (LLAMA, collection Wa-B-03-1-32) [USNM, specimen CASENT0604945]. Paratypes: same data as holotype but 14.94460°N, 89.27726°W ±57m, 1550m, 12 May 2009 (LLAMA, Wm-B-03-1-04), [1w, CAS, CASENT0623248], [1w, EAPZ, CASENT0623249], [1w, ECOSCE, CASENT0623250], [1w, FMNH, CASENT0623251], [1w, ICN, CASENT0623252, [1w, INBio, CASENT0623253], [1w, JTLC, CASENT0623527], [1w, LACM, CASENT0623254], [2w, MGBPC, CASENT0623528, CASENT0623529], [1w, MCZ, CASENT0623255], [1w, MZSP, CASENT0623256, [1w, UCD, CASENT0623257], [1w, UNAM, CASENT0623258], [1dq, 1w, USNM, CASENT0606239, CASENT0606656], [1w, UVGC, CASENT0623259].
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. Similar species: Stenamma zelum.
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.
Stenamma brujita holotype worker (CASENT0604945) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma brujita worker variants. Face, profile, and dorsal views A–C Variant 1 (CASENT0603918) D–F Variant 2 (CASENT0604607) G–I Variant 3 (CASENT0604578).
(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 Figure 56D.
Stenamma brujita A Queen (CASENT0604991), profile B Same, face C Same, dorsum D Queen (CASENT0621749), wings.
Unknown.
Stenamma brujita is known only from Winkler and Berlese samples of leaf litter collected from the floor of wet forest habitats (e.g. lowland rainforest, montane wet forest, cloud forest, pine cloud forest, oak-pine forest). The species has a broad elevational range, occurring from 200–1800 m, but it is most common at mid-elevations (1000–1500 m).
The combination of large size, small eyes, and foveate sculpture make Stenamma brujita a very distinctive species, unlikely to be confused with any other MAC species. However, Stenamma zelum, which is not closely related to Stenamma brujita (
Stenamma brujita is quite variable throughout its range and may comprise a complex of several species. I choose to identify a single species here, because every population exhibits some amount of variation, none of the variants occur in sympatry, and some populations have intermediate phenotypes. I do however, identify three main variants that differ significantly from the holotype population.
Variant 1 (Figure 54A–C) includes all collections from Tamaulipas and Hidalgo, Mexico. It has the following features: body sculpture more rugoreticulate than foveate; propodeal spines long, straight and more slender; petiole with a distinct concavity below node.
Variant 2 (Figure 55D–F), the most distinctive variant, is known from a few collections taken on the wet Atlantic slope of the Sierra Juarez, between Oaxaca and Valle Nacional in Mexico. It has the following features: body very large; general body color very dark, mostly black; petiole wedge-shaped, usually without a distinct node; propodeal spines forming robust, blunt-tipped triangles, which point almost vertically.
Variant 3 (Figure 56G–I) occurs at several localities in Chiapas, México, mainly Nahá and Lago Metzabok. It is characterized by the following: general body color brown; promesonotum in profile high-domed, and asymmetrical; propodeal spines tuberculate; petiole in profile appearing more gracile with node reduced in size; anterior clypeal margin forming 4 blunt teeth.
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.
GUATEMALA: Baja Verapaz : 7.3km E Purulhá, [ca. 15.267°N, 90.132°W], 1700m, 19 May 1991 (R. S. Anderson); Suchitepéquez : Finca Sn. Jerónimo, 14.55914°N, 91.16705°W, 1790m, 11 Dec 2010 (L. Sáenz); Volcán Atitlán, 10km SE Santiago Atitlán, 14.552°N, 91.193°W, 1690m, 10 Sep 2008 (M. G. Branstetter); 4km S Vol. Atitlán, 14.55195°N, 91.192333°W, 1750m, 15 Jun 2009 (LLAMA); Zacapa : 2km SE La Unión, 14.94706°N, 89.27660°W, 1550m, 12 May 2009 (LLAMA); 2km SE La Unión, 14.95463°N, 89.27721°W, 1430m, 12 May 2009 (LLAMA); 3.5km SE La Union, 14.95000°N, 89.26667°W, 1500m, 6 Jun 1991 (R. S. Anderson); HONDURAS: Cortés : Parque Nacional Cusuco, 15.48710°N, 88.23469°W, 1330m, 30 May 2010 (LLAMA); Parque Nacional Cusuco, 15.48940°N, 88.23584°W, 1290m, 30 May 2010 (LLAMA); 25km N Cofradia, PN Cusuco, [ca. 15.497°N, 88.227°W], 1550m, 26 Aug 1994 (S. & J. Peck); MÉXICO: Chiapas : 10km W El Bosque, [ca. 17.0440°N, 92.8612°W], 1475m, 15 Sep 1992 (R. S. Anderson); 10.6km W El Bosque, [ca. 17.043°N, 92.762°W], 1460m, 25–29 Aug 1973 (A. F. Newton); Lago Metzabok, 17.12562°N, 91.63086°W, 570m, 6 Jun 2008 (LLAMA); Nahá, 16.96358°N, 91.59332°W, 985m, 8 Jun 2008 (LLAMA); Nahá, 16.94864°N, 91.59383°W, 930m, 8 Jun 2008 (LLAMA); 12.5km NW Ocosingo, [ca. 16.983°N, 92.183°W], 1400m, 16 Sep 1992 (R. S. Anderson); 19km NW Ocozocoautla, [ca. 16.877°N, 93.458°W], 975m, 4-5 Sep 1973 (A. F. Newton); Hidalgo : 11km SW Chapulhuacán, [ca. 21.147°N, 98.966°W], 1200m, 5 Jul 1976 (A. F. Newton; Oaxaca : 10km S Valle Nacional, [ca. 17.724°N, 96.324°W], 650m, 19 May 1971 (S. B. Peck); 13.2km SW Valle Nacional, 17.65934°N, 96.33426°W, 1360m, 11 Aug 2009 (M. G. Branstetter); 25km S Valle Nacional, [ca. 17.670°N, 96.330°W], 1200m, 21 May 1971 (S. B. Peck); 26km S Valle Nacional, km 71, [ca. 17.645°N, 96.336°W], 1220m, 25 Jun 1983 (S. & J. Peck); Puebla : 24km N Xicotepec de Juarez, [ca. 20.282°N, 97.963°W], 1070m, 17 Jun 1983 (R. S. Anderson); Tamaulipas : El Cielo, nr Alta Cima, 23.06518°N, 99.20433°W, 980m, 21 Aug 2009 (L. Sáenz); 1.8km W Alta Cima, 23.06110°N, 99.21564°W, 1340m, 23 Aug 2009 (M. G. Branstetter); nr Gomez Farias Rancho del Cielo, [ca. 23.063°N, 99.205°W], 1000m, 7 Aug 1983 (S. & J. Peck) Veracruz : Los Tuxtlas, 10km NNW Sontecomapan, 18.583°N, 95.083°W, 200m, 20 Mar 1985 (P. S. Ward); Los Tuxtlas, 10km NNW Sontecomapan, 18.583°N, 95.083°W, 500m, 21 Mar 1985 (P. S. Ward).
Distribution map of Stenamma brujita.
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http://species-id.net/wiki/Stenamma_callipygium
Worker: Figure 58; Queen: Figure 59; Map: Figure 60Holotype worker. GUATEMALA: Baja Verapaz, Biotopo Quetzal, 15.21329°N, 90.21516°W ±110m, 1715m, 8 May 2009, cloud forest, beating vegetation (LLAMA, collection Go-B-02-1-02) [USNM, specimen CASENT0606207]. Paratypes: same data as holotype [1w, CAS, CASENT0606206]; same data but (LLAMA, Go-B-02-1-01) [1w, MCZ, CASENT0606203]; 15.21227°N, 90.21430°W ±50m, 1750m, 7 May 2008 (LLAMA, Wa-B-02-1-37) [1w, MGBPC, CASENT0604923]; 15.21278°N, 90.21552°W ±10m, 1720m, 7–10 May 2009 (LLAMA, Ft-B-02-1-01) [1w, UCD, CASENT0606227]; 15.21202°N, 90.21653°W ±207m, 1735m, 8 May 2009 (LLAMA, Ba-B-02-1-07-05) [1dq, USNM, CASENT0603928]; GUATEMALA: Baja Verapaz, Ranchito El Quetzal, 15.21508°N, 90.22003°W, 1700m, 20 Sep 2008 (R. S. Anderson, RSA2008-139) [1w, UVGC, CASENT0606095].
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. Similar species: Stenamma atribellum.
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.
Stenamma callipygium holotype worker (CASENT0606207) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
(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.
Stenamma callipygium paratype queen (CASENT0603928) A Profile B Face C Dorsum.
Unknown.
Stenamma callipygium has been collected by sifting leaf litter, beating vegetation and baiting with cookie bait. In addition, a single worker was found in a flight intercept trap placed underneath a Malaise trap. This species is a cloud forest specialist known from 1630–1750 m elevation. Very few Stenamma species are known from beating samples, suggesting that this species may nest or at least forage arboreally. The diversity of collecting methods that have retrieved this species suggests that it is an active forager. It is surprising, however, that out of the over 100 leaf litter samples collected at the Biotopo Quetzal by the LLAMA project, only one sample had Stenamma callipygium.
The oddly shaped clypeus and mandible of this species is unique within Stenamma and begs for explanation. Most likely the projecting clypeal tooth is used for the capture or maceration of a specific prey type.
Stenamma callipygium and Stenamma atribellum are sister species and make up the atribellum species group. The diagnostic character state of this group is the elongate gastral constriction. Stenamma callipygium is easy to distinguish from Stenamma atribellum by its median clypeal tooth and carinulate sculpture.
GUATEMALA: Baja Verapaz : Biotopo Quetzal, 15.21329°N, 90.21516°W, 1715m, 8 May 2009 (LLAMA); Biotopo Quetzal, 15.21227°N, 90.21430°W, 1750m, 7 May 2009 (LLAMA); Ranchito El Quetzal, 15.21508°N, 90.22003°W, 1700m, 20 Sep 2008 (R. S. Anderson); 4.5km E Purulhá, [ca. 15.226°N, 90.200°W], 1630m, 21 May 1991 (R. S. Anderson); 7.3km E Purulhá, [ca. 15.2667°N, 90.132°W], 1700m, 19 May 1991 (R. S. Anderson).
Distribution map of Stenamma callipygium (circles) and Stenamma catracho (squares).
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Worker: Figures 61, 62; Queen: Figure 63; Map: Figure 60Holotype worker. HONDURAS: Olancho, Parque Nacional La Muralla, 15.09965°N, 86.74072°W ±20m, 1530m, 2 May 2010, cloud forest, ex sifted leaf litter (LLAMA, collection Wa-C-01-1-36) [USNM, specimen CASENT0621306]. Paratypes: same data as holotype [1dq, 1w, USNM, CASENT0621305, CASENT0621307], [1w, CAS, CASENT0621308]; same data as holotype but 15.09925°N, 86.74063°W ±20m, 1530m, 2 May 2010 (LLAMA, Wa-C-01-1-27) [1w, EAPZ, CASENT0621298], [1w, ECOSCE, CASENT0621299], [1w, FMNH, CASENT0621297], [1w, ICN, CASENT0621300]; 15.09859°N, 86.74216°W ±10m, 1510m, 2 May 2010 (LLAMA, Wm-C-01-1-03) [1w, INBio, CASENT0622022], [1w, JTLC, CASENT0622023], [1w, LACM, CASENT0622024], [1w, MGBPC, CASENT0623530], [1w, MCZ, CASENT0623531], [1w, MZSP, CASENT0623532], [1w, UCD, CASENT0623533]; 15.09694°N, 86.74533°W ±10m, 1460m, 5 May 2010 (LLAMA, Wm-C-01-1-09) [1w, UNAM, CASENT0622048], [1w, UVGC, CASENT0623534].
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). Similar species: Stenamma crypticum, Stenamma cusuco, Stenamma hojarasca.
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.
Stenamma catracho holotype worker (CASENT0621306) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma catracho worker variant (CASENT0621563) A Face B Profile C Dorsum.
(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.
Stenamma catracho paratype queen (CASENT0621305) A Profile B Face C Dorsum.
Unknown.
Stenamma catracho is known only from sifted leaf litter collected from the forest floor. It occurs from approximately 1100–1900 m elevation in montane wet forest habitats.
Stenamma catracho is most likely to be confused with Stenamma cusuco or Stenamma hojarasca, both of which have expanded frontal lobes and somewhat elongate petioles. Stenamma cusuco can be distinguished easily by its distinctive tridentate clypeal margin, and Stenamma hojarasca can be separated by its longer petiole and completely shiny petiolar and postpetiolar nodes.
There are some morphological differences unique to the Meambar population (Figure 62) of Stenamma catracho. Meambar specimens have most pilosity longer, with the gastral pilosity clearly bilayered; they have lighter integument color; and they have narrower frontal lobes. The other populations have the pilosity shorter overall and the gastral pilosity denser and not as clearly bilayered. I treat these differences as intraspecific variation until more material is collected.
Molecular phylogenetic data show that Stenamma catracho most likely belongs to a clade that includes Stenamma crypticum, Stenamma monstrosum, and Stenamma saenzae, with Stenamma monstrosum being its sister species (Branstetter unpublished data).
GUATEMALA: Zacapa : 2km SE La Unión, 14.95344°N, 89.27587°W, 1430m, 12 May 2009 (LLAMA); HONDURAS: Comayagua : PN Cerro Azul Meambar, 14.86960°N, 87.89843°W, 1140m, 20 May 2010 (LLAMA); Olancho : PN La Muralla, 15.09511°N, 86.73925°W, 1420m, 2 May 2010 (LLAMA); PN La Muralla, 15.09965°N, 86.74072°W, 1530m, 2 May 2010 (LLAMA); PN La Muralla, 15.10257°N, 86.73642°W, 1650m, 2 May 2010 (R. S. Anderson); PN La Muralla, 15.09807°N, 86.72047°W, 1860m, 3 May 2010 (LLAMA).
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Worker: Figures 64, 65; Queen: Figure 66; Male: Figure 66E–G; Map: Figure 67Holotype worker. MÉXICO, Oaxaca: 10.8km SW Valle Nacional, 17.68102°N, 96.33026°W ±66m, 1120m, 13 Aug 2009, disturbed mesophyll forest, ex sifted leaf litter (M. G. Branstetter, collection MGB1387) [USNM, specimen CASENT0622438]. Paratypes: same data as holotype [1dq, 1w, CAS, CASENT0623275, CASENT0623261], [1w, EAPZ, CASENT0623262], [1w, ECOSCE, CASENT0623263], [1w, FMNH, CASENT0623264], [1w, ICN, CASENT0623265], [1w, INBio, CASENT0623266], [1w, JTLC, CASENT0623274], [1w, LACM, CASENT0623267], [1w, MGBPC, CASENT0623268], [1dq, 1w, MCZ, CASENT0623276, CASENT0623272], [1w, MZSP, CASENT0623269], [1w, UCD, CASENT0623270], [1w, UNAM, CASENT0623271], [1dq, 1w, USNM, CASENT0622439, CASENT0623260], [1w, UVGC, CASENT0623273].
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. Similar species: Stenamma crypticum, Stenamma huachucanum.
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.
Stenamma connectum holotype worker (CASENT0622438) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma connectum worker variants. Face, profile, and dorsal views A–C Variant 1 (CASENT012648) D–F Variant 2 (CASENT06005461) G–I Variant 3 (CASENT0605552).
(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 Figure 66D.
Stenamma connectum A Paratype queen (CASENT0622439), profile B Same, face C Same, dorsum D Queen (CASENT0126417), wings E Male (CASENT0605585), profile F Same, face G Same, dorsum.
See Figure 66E–G.
Stenamma connectum is found in montane wet forest habitats from 600–2160 m elevation, and is known almost exclusively from Winkler or Berlese samples of sifted leaf litter. Only once has a nest been found: a single dealate queen with brood underneath a moss mat.
Distinguishing Stenamma connectum from Stenamma crypticum and Stenamma huachucanum can be difficult. This is because each species is composed of multiple allopatric populations with no clear evidence of sympatry among divergent forms. Using morphology alone, the best solution would probably be to delimit a single widespread species. However, molecular phylogenetic data strongly suggest the existence of multiple biological species (Branstetter unpublished data). As a result, I have decided to delimit several species guided by the combination of morphological and molecular data. Some phenotypically divergent populations which have not been included in the phylogeny may prove challenging to identify.
As currently defined, geography is very useful in separating Stenamma connectum from similar species. Stenamma connectum occurs in the Mexican states of Veracruz and Oaxaca only. Within Oaxaca, it is found along the Caribbean slope in wet forest habitats. Stenamma huachucanum is distributed from the southwestern U.S.A to Oaxaca. Within Oaxaca, it occurs only in the drier habitats in the central and western parts of the state. I consider specimens from Hidalgo and San Luis Potosí states to be Stenamma huachucanum, but some of these specimens are hard to place. Stenamma crypticum occurs mainly from Chiapas, Mexico to Nicaragua; however, a couple of specimens known from one sample collected in Veracruz at 1600 m (11km N San Andrés Tuxtla) appear most like Stenamma crypticum. These specimens lack the broadly rounded propodeal lobes (in profile) and have the promesonotum mostly smooth. Until more material is collected, I treat these as Stenamma crypticum, but I find it possible that they are actually abberant specimens of Stenamma connectum. One specimen that clearly has the characteristics of Stenamma connectum was collected at nearly the same locality, only slightly lower at 1400 m elevation.
Many populations of Stenamma connectum, including the type population, share a set of distinctive morphological character states that distinguish Stenamma connectum from similar species. Key worker characters for the type population are as follows: body color mostly brown; side of pronotum lightly punctate; dorsum of pronotum carinulate-punctate with a small patch of smooth cuticle in middle; mesopleuron and side of propodeum strongly punctate; propodeal declivity in profile forming a sinuous outline, in which the propodeal lobe is broadly rounded and makes a smooth connection with the propodeal spine; propodeal spines tuberculate. The sinuous outline of the propodeal declivity is the most distinctive feature of this species. It is variable among populations, with specimens from lower elevations having the sinuosity more pronounced. Higher elevation populations begin to lose the character state, making these populations more difficult to separate from similar species. Several variants are described below.
Variant 1 (Figure 65A–C) is known only from the locality Pueblo Nuevo in Veracruz. These specimens differ as follows: smaller overall size; mesosoma completely punctate, with only a few scattered rugulae/carinulae; sinuosity in outline of propodeal declivity pronounced; petiolar node in profile reaching a more angular apex.
Variant 2 (Figure 65D–F) and variant 3 (Figure 65G-I) occur along the same elevational transect as the type population, but at mid (~1770 m) and high (> 1900 m) elevations, respectively. Variant 2 differs from the type form in that it has dark brown body color, reduced pronotal sculpture, and a noticeably yellow flagellum. Variant 3 is a high elevation form of Stenamma connectum. Without the intermediate phenotype present in variant 2, it would be easy to describe variant 3 as a new species as it is very divergent from the type population. It is similar to variant 2 except as follows: larger size; body color almost black; pronotum almost completely smooth and shiny; mesopleuron rugulose-punctate; side of propodeum carinulate; outline of propodeal declivity not strongly sinuous, with the propodeal lobe often more angular and isolated from the propodeal spine; petiolar and postpetiolar nodes mostly smooth; postpetiole bulging.
MÉXICO: Oaxaca : Mirador Grande, 17.89844°N, 96.36263°W, 990m, 14 Aug 2009 (M. G. Branstetter); 10km S Valle Nacional, [ca. 17.724°N, 96.324°W], 650m, 19 May 1971 (S. B. Peck); 10.8 km SW Valle Nacional, 17.68102°N, 96.33026°W, 1120m, 13 Aug 2009 (M. G. Branstetter); 13.2km SW Valle Nacional, 17.65934°N, 96.33426°W, 1360m, 11 Aug 2009 (M. G. Branstetter); 14.8km SSW Valle Nacional, 17.64483°N, 96.33637°W, 1370m, 13 Aug 2009 (M. G. Branstetter); 20.5km SW Valle Nacional, 17.60560°N, 96.38398°W, 1770m, 12 Aug 2009 (M. G. Branstetter); 20.6km SW Valle Nacional, 17.60404°N, 96.378786°W, 1733m, 13 Aug 2009 (M. G. Branstetter); 22.4km SW Valle Nacional, 17.59112°N, 96.39133°W, 1990m, 13 Aug 2009 (M. G. Branstetter); 25km S Valle Nacional, [ca. 17.670°N, 96.330°W], 1200m, 21 May 1971 (S. B. Peck); 26km SW Valle Nacional, 17.58667°N, 96.44948°W, 2160m, 11 Aug 2009 (M. G. Branstetter); 30.1km S Valle Nacional, [ca. 17.627°N, 96.354°W], 1580m, 11–18 Aug 1973 (A. F. Newton); 47.5km SW Valle Nacional, Km 100.5, [ca. 17.595°N, 96.424°W], 2125m, 26 Jul 1992 (R. S. Anderson); 52km S Valle Nacional, [ca. 17.587°N, 96.449°W], 2130m, 22 May 1971 (S. B. Peck); Veracruz : Cordoba, [ca. 18.908°N, 96.958°W], 4 Aug 1969 (S. B. Peck); Cordoba, Paraje Nuevo, Nacimiento, [ca. 18.88°N, 96.86°W], 7 Aug 1969 (S. & J. Peck); park cañon, Hwy150, 3.2km W Fortín, [ca. 18.884°N, 97.019°W, 6–9 Aug 1969 (S. & J. Peck); Fortín, Canyon Río Metlac, [ca. 18.90°N, 97.00°W], 5 Aug 1969 (S. & J. Peck); 7km E Huatusco, [ca. 19.207°N, 96.914°W], 22 Jun 1983 (Peck & Anderson); 7.1km N Huatusco, [ca. 19.262°N, 96.964°W], 1280m, 29 Jul–3 Aug 1973 (A. F. Newton); Paraje Nuevo, Cueva de Ojo de Agua, [ca. 18.877°N, 96.862°W], 7 Aug 1969 (S. & J. Peck) Pueblo Nuevo, nr Tetzonapa, 1 Aug 1953 (E. O. Wilson); 11km N San Andrés Tuxtla, 18.55°N, 96.00°W, 1400m, 23 Mar 1985 (P. S. Ward); 2.7km N Teocelo, [ca. 19.40°N, 96.98°W], 1130m, 22–24 Jul 1934 (A. F. Newton).
Distribution map of Stenamma connectum (circles) and Stenamma cusuco (squares).
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http://species-id.net/wiki/Stenamma_crypticum
Worker: Figures 68, 69; Queen: Figure 70A–D; Map: Figure 71Holotype worker. MÉXICO, Chiapas: 2km SE Custepec, 15.72141°N, 92.93936°W, 1860m, 17 May 2008, oak-pine forest, ex sifted leaf litter (R. S. Anderson, collection RSA2008-013) [USNM, specimen CASENT0604771] Paratypes: MÉXICO, Chiapas: 3km SE Custepec, 15.71485°N, 92.93823°W ±50m, 1700m, 17 May 2008 (LLAMA, Wa-A-02-2-50) [1dq, 1w, CAS, CASENT0623277, CASENT0623280], [1w, EAPZ, CASENT0623281], [1w, ECOSCE, CASENT0623282], [1w, FMNH, CASENT0623283], [1w, ICN, CASENT0623284], [1w, INBio, CASENT0623286], [1w, JTLC, CASENT0623536], [1w, LACM, CASENT0623288], [1w, MGBPC, CASENT0623537], [1dq, 1w, MCZ, CASENT0623278, CASENT0623289], [1w, MZSP, CASENT0623290], [1w, UCD, CASENT0623291], [1w, UNAM, CASENT0623292], [1dq, 1w, USNM, CASENT0623279, CASENT0623293], [1w, UVGC, CASENT0623294].
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. Similar species: Stenamma connectum, Stenamma huachucanum, Stenamma ignotum, Stenamma picopicucha.
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.
Stenamma crypticum holotype worker (CASENT0605185) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma crypticum worker variants. Face, profile, and dorsal views A–C Type population worker with less sculpture (CASENT0603821) D–F Type population worker with more sculpture (CASENT0604745) G–I Variant 1 (CASENT0604839) J–L Variant 2 (CASENT0604997).
(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 Figure 70D.
Stenamma crypticum A Queen (CASENT0604854), profile B Same, face C Same, dorsum D Queen (CASENT0605933), wings.
Unknown.
Stenamma crypticum is a rather common component of the leaf litter in mid- to high-elevation mesic forest habitats in Central America. It has been collected from 900–2800 m, but is most common from 1500–2500 m. Habitat types include cloud forest, mesophyll forest, oak forest, and mixed hardwood forest. Most collections come from samples of sifted leaf litter collected from the forest floor, but some specimens are also known from cookie baits. Nests have never been found, but dealate queens, as well as workers, are common in the leaf litter, suggesting that nests might be in this stratum.
Stenamma crypticum should be distinguished easily from Stenamma ignotum and Stenamma picopicucha using the diagnostic characters given above. As noted under Stenamma connectum above, separating Stenamma crypticum from Stenamma connectum and Stenamma huachucanum is more challenging. This is because each species comprises a complex of multiple divergent populations, with no clear evidence of sympatry among distinct forms. Using morphology alone, it might be best to name a single species; however, molecular phylogenetic data strongly suggest the existence of at least three species (Branstetter unpublished data). Even though some populations/specimens may prove difficult to identify, I have chosen to delimit these species as best as possible. There are a few key morphological characters that separate the type populations of each species from one another, but when considering all populations, geography is useful.
Stenamma crypticum occurs mainly in Nuclear Central America from Chiapas, Mexico to Honduras, whereas Stenamma connectum is found north of Stenamma crypticum in Veracruz, Mexico and on the wet, Caribbean slope of Oaxaca. Complicating this picture, however, are a few specimens collected from Veracruz at 1600 m (11km N San Andrés Tuxtla). These specimens lack the broadly rounded propodeal lobes characteristic of Stenamma connectum and have a mostly smooth promesonotum. I tentatively identify these specimens as Stenamma crypticum, but I find it possible that they could actually be abberant members of Stenamma connectum, which has been collected close by at a slightly lower elevation (1400 m). The phylogenetic position of the putative Stenamma crypticum specimens has not been assessed. Stenamma huachucanum is distributed from the southwestern U.S.A. to Oaxaca, where it occurs only on the drier western side of the state. A wet forest version of Stenamma huachucanum occurs in eastern Mexico from Tamaulipas to Puebla.
Stenamma picopichuca can be separated from Stenamma crypticum using the diagnostic characters listed above, but I am somewhat uncertain about the phylogenetic placement and status of this species. Stenamma picopichucha has the basal margin of the mandible like Stenamma ignotum and the anterior clypeal margin like Stenamma crypticum. Because of several other similarities, I was originally going to include Stenamma picopicucha in Stenamma crypticum, but I later discovered that the two species occur in sympatry at Cusuco in Honduras. At this site, specimens of Stenamma crypticum clearly have a sinuous basal margin of the mandible and specimens of Stenamma picopicucha have a straight margin. The phylogenetic position of Stenamma picopicucha is yet to be tested, and it will be interesting to see if the two species are closely related.
Within the range of Stenamma crypticum there is considerable variation in size, sculpture and gastral pilosity, with some of this variation observable at single sites. At the type locality, for example, I have sampled Stenamma crypticum from 1500 m to about 2200 m elevation. Along this gradient specimens from higher elevation are larger. There is also significant variation in pronotal sculpture within the site, with some specimens having the pronotal dorsum mostly smooth (Figure 69A–C), and others mostly rugose (Figure 69D–F). At first I tried separating these into distinct forms, but I abandoned this scheme after finding specimens with intermediate phenotypes.
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 (Figure 69G–I) occurs at several sites in Guatemala (e.g. Biotopo Quetzal, Purulhá). It is similar to the type form, but has a bulging postpetiole and longer gastral pilosity. Variant 2 (Figure 69J–L) occurs at La Union in Guatemala, with similar-looking specimens at sites in Honduras. Compared to the type form it is smaller and has the suberect layer of gastral pilosity noticeably thickened. Until there is evidence of sympatry, I treat all of this variation as intraspecific.
GUATEMALA: Baja Verapaz : Biotopo Quetzal, 15.21298°N, 90.21510°W, 1750m, 7 May 2009 (LLAMA); Purulhá, Biotopin, 15.21535°N, 90.21618°W, 1700m, 26–30 Mar 2008 (Méndez et al.); 7km E Purulhá, [ca, 15.2667°N, 90.1348°W], 1600m, 25 May 1991 (R. S. Anderson); 4.5km S Purulhá, [ca. 15.226°N, 90.200°W], 1630m, 21 May 1991 (R. S. Anderson); 7km S Purulhá, [ca. 15.194°N, 90.200°W], 1660m, 20 May 1991 (R. S. Anderson); 4km SSE Purulhá, 15.20522°N, 90.22198°W, 2100m, 20 Sep 2008 (M. G. Branstetter); Ranchito El Quetzal, 15.21443°N, 90.22123°W, 1750m, 20 Sep 2008 (R. S. Anderson); Salamá, Cerro Verde, 15.17446°N, 90.19353°W, 1800m, 26–30 Mar 2008 (Méndez et al.); Salamá, Hotel Posada del Quetzal 1, 15.19710°N, 90.21169°W, 1600m, 26–30 Mar 2008 (Méndez et al.) El Progresso : 20km N Estancia de la Virgen, 15.1141°N, 89.8833°W, 1850m, 8 Jun 1991 (R. S. Anderson); Guatemala : 1km SE La Pueblito, [ca. 14.6211°N, 90.5269°W], 1800m, 10 Jun 1991 (R. S. Anderson); nr Las Nubes, 14.53349°N, 90.35941°W, 1850m, 18 Sep 2008 (R. S. Anderson); 5.9km ESE San José Pinula, 14.53349°N, 90.35941°W, 1850m, 18 Sep 2008 (M. G. Branstetter); 7km ESE San José Pinula, 14.53915°N, 90.34933°W, 2060m, 18 Sep 2008 (M. G. Branstetter); Jalapa : Aldea Manzano, 1.8km WNW San José La Sierra, 14.50476°N, 90.25613°W, 1990m, 18 Sep 2008 (M. G. Branstetter); El Manzano, 14.50476°N, 90.25613°W, 2150m, 18 Sep 2008 (R. S. Anderson); San Marcos : La Fraternidad, 14.93604°N, 91.86778°W, 1920m, 11 Sep 2008 (M. G. Branstetter); Rd. Bojonal-Fraternidad, 14.94533°N, 91.88038°W, 1580m, 11 Sep 2008 (R. S. Anderson); 9.8km WSW San Marcos, 14.94427°N, 91.87990°W, 1600m, 11 Sep 2008 (M. G. Branstetter);Suchitepéquez : Finca Sn. Jerónimo, 14.55914°N, 91.16705°W, 1790m, 11 Dec 2010 (L. Sáenz); Volcán Atitlán, 9.5km SE Santiago Atitlán, 14.55828°N, 91.19133°W, 2000m, 10 Sep 2008 (M. G. Branstetter); 4km S Vol. Atitlán, 14.54830°N, 91.19115°W, 1625m, 15 Jun 2009 (LLAMA); 4km S Vol. Atitlán, 14.55112°N, 91.19848°W, 1750m, 15 Jun 2009 (LLAMA); Zacapa : 14km NNE Teculután, 15.1134°N, 89.6781°W, 2270m, 6 Jul 2007 (R. S. Anderson); 2km SE La Unión, 14.94701°N, 89.27594°W, 1550m, 12 May 2009 (LLAMA); HONDURAS: Comayagua : 10km E Comayagua, 14.45973°N, 87.54609°W, 2000m, 15 May 2010 (LLAMA); Cortés : PN Cusuco, 15.50739°N, 88.23373°W, 2030m, 3 Jun 2010 (LLAMA);Lempira : PN Celaque, 8.3km SW Graçias, 14.56132°N, 88.65768°W, 1860m, 30 Sep 2008 (M. G. Branstetter); PN Celaque, 8.7km SW Graçias, 14.5877°N, 88.66117°W, 2100m, 30 Sep 2008 (M. G. Branstetter); Ocotepeque : 13km E Nuevo Ocotopeque, 14.45685°N, 89.06856°W, 2200m, 25 May 2010 (LLAMA); Santa Barbara : 15km SE Santa Barbara, [ca. 14.906°N, 88.100°W], 24 Aug 1991 (S. B. Peck); MÉXICO: Chiapas : Cacahuatan, Las Nubes, [ca. 15.097°N, 92.140°W], 1770m, 1–7 Aug 1950 (Goodnights); Cerro Huitepec (Pico), 5km W San Cristobal, [ca. 16.7500°N, 92.6802°W], 2750m, 18 Sep 1991 (R. S. Anderson); Cerro de Tapalapa, 17.18786°N, 93.12308°W, 2260m, 28 May 2008 (R. S. Anderson); 5km NE Coapilla, 17.17557°N, 93.13187°W, 1990m, 25 May 2008 (LLAMA); 2km SE Custepec, 15.72099°N, 92.95050°W, 1520m, 17 May 2008 (LLAMA); 4km SE Custepec, 15.70658°N, 92.93126°W, 2125m, 20 May 2008 (LLAMA); Lagunas de Montebello, Cinco Lagos, [ca. 15.2667°N, 90.1348°W], 1660m, 23 May 1991 (R. S. Anderson); 7.4km SSW Motozintla de Mendoza, [ca. 15.367°N, 92.233°W], 2000m, 21 Sep 1992 (R. S. Anderson); Najá, 16.97417°N, 91.58592°W, 950m, 14 Jul 2007 (R. S. Anderson); 13km N Pueblo Nuevo Solistahuacán, [ca. 17.211°N, 92.964°W], 1860m, 26–27 Aug 1973 (A. F. Newton); 8.9km E Rayon, 17.20000°N, 92.91633°W, 1500m, 19 Sep 1991 (R. S. Anderson); Sierra Morena, C. Bola, 16.13464°N, 93.60077°N, 1950m, 14 May 2008 (R. S. Anderson); 17km ENE Tonalá, 16.14153°N, 93.60958°W, 1650m, 16 Jul 2007 (M. G. Branstetter); 4km N Union Juarez, Volcan Tacana, lower slopes, [ca. 15.133°N, 92.100°W], 2000m, 19 Sep 1992 (R. S. Anderson); Veracruz : 11km N San Andrés Tuxtla, 18.55°N, 96.00°W, 1600m, 23 Mar 1985 (P. S. Ward).
Distribution map of Stenamma crypticum.
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http://species-id.net/wiki/Stenamma_cusuco
Worker: Figure 72; Queen: Figure 73; Map: Figure 67Holotype worker. HONDURAS, Cortés: Parque Nacional Cusuco, 15.48965°N, 88.23383°W ±35m, 1300m, 31 May 2010, mesophyll forest, ex sifted leaf litter (LLAMA, collection Wm-C-06-1-04) [USNM, specimen CASENT0622137]. Paratypes: same data as holotype [1w, CAS, CASENT0623296], [1w, EAPZ, CASENT0623297], [1w, ECOSCE, CASENT0623298], [1w, FMNH, CASENT0623299], [1w, INBio, CASENT0623300], [1w, LACM, CASENT0623301], [1dq, 1w, MCZ, CASENT0623295, CASENT0623302], [1w, MZSP, CASENT0623303], [1dq, 1w, USNM, CASENT0622136, CASENT0622138]; same data but 15.48940°N, 88.23598°W ±20m, 1290m, 30 May 2010 (LLAMA, Wa-C-06-2-07) [1w, UCD, CASENT0621743], [1w, UNAM, CASENT0621741], [1w, UVGC, CASENT0621742]; 15.49037°N, 88.23402°W ±40m, 1330m, 31 May 2010 (LLAMA, Wm-C-06-1-07) [1w, JTLC, CASENT0623538], [1w, MGBPC, CASENT0623539].
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;. Similar species: Stenamma catracho, Stenamma hojarasca, Stenamma ochrocnemis.
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.
Stenamma cusuco holotype worker (CASENT0622137) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
(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.
Stenamma cusuco paratype queen (CASENT0622136) A Profile B Face C Dorsum.
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 Stenamma species. Most peculiar is the strongly projecting median lobe, which forms a tooth-like protuberance. This trait probably correlates with diet specialization, but nothing is known yet about prey choice in this species.
The structure of the clypeus is unique among known Stenamma species. Thus, Stenamma cusuco should not be confused with any other species.
Although support values are low, molecular phylogenetic results indicate that Stenamma cusuco belongs to a clade that includes Stenamma hojarasca and Stenamma ochrocnemis (Branstetter unpublished data).
See type material.
http://species-id.net/wiki/Stenamma_diversum
Worker: Figure 74; Queen: Figure 75A–D, Male: Figure 75E–G; Map: Figure 76Integument 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. Similar species: Stenamma lobinodus, Stenamma tico.
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.
Stenamma diversum. A, B, E, G, H Worker (CASENT0606723) C, D, F Lectotype worker (USNMCOTYPE24447).
(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 Figure 75D.
Stenamma diversum A Queen (CASENT0606782), profile B Same, face C Same, dorsum D Same, wings E Male (CASENT0622357), profile F Same, face G Same, dorsum.
See Figure 75E–G.
Stenamma diversum occurs in relatively pristine wet forest habitats from sea level to about 1, 100 m elevation. It is a particularly interesting species because it has convergently evolved many of the same nesting behaviors as Stenamma alas and Stenamma expolitum, even though it is not closely related (
Stenamma diversum and Stenamma tico are sister species and together form the diversum species group. This group is defined by the following characters: head mostly smooth and shiny; mesosoma mostly reticulately costate; promesonotum in profile low-domed, roughly symmetrical; postpetiole in profile without a distinct dorsal lobe that projects posteriad over postpetiole (as in Stenamma lobinodus and other lobinodus group species); basal margin of mandible straight; anterior clypeal margin with a median emargination. Based on DNA sequence data Stenamma vexator is likely sister to the diversum species group, but morphological characters could not be found to diagnose the entire clade.
Stenamma diversum is probably the most distinctive Stenamma species, but it does bear some similarity to its sister taxon Stenamma tico and the more distantly related Stenamma lobinodus, both of which share roughly the same color and sculpture pattern. It is easy to separate Stenamma diversum from these other species by comparing the propodeal spines and the frontal lobes. In Stenamma tico, the spines are absent or at most form small dorsally projecting tubercles (PSL 0.14–0.18, PSI 1.4–1.9). In Stenamma lobinodus, the propodeal spines are usually well developed, but they are shorter than those of Stenamma diversum (PSL < 0.23 vs. > 0.27, PSI < 2.6 vs. > 3.0). Stenamma diversum is the only species among these three to have greatly expanded frontal lobes, causing the underlying torular lobes to be covered in full-face view. In the other species, the torular lobes are always visible. The frontal lobes of Stenamma lobinodus are especially reduced. Geography also can help separate these species. Stenamma diversum occurs in sympatry with Stenamma lobinodus only in Oaxaca, Mexico (although never collected together at the same site) and with Stenamma tico only in northern Nicaragua (found in sympatry at Cerro Saslaya and Cerro Musún).
BELIZE: Cayo : Caves Branch, [ca. 17.13°N, 88.70°W], 4-14 Aug 1972 (S. & J. Peck); GUATEMALA: Alta Verapaz : Muchbilhá, 15.86966°N, 90.14254°W, 184m, 9 Nov 2009 (L. Sáenz); Izabal : Carboneras Sn. Gil, 15.63728°N, 88.82826°W, 415m, 19 Nov 2009 (L. Sáenz); 5km NW Morales, 15.51441°N, 88.86459°W, 250m, 17 May 2009 (LLAMA); 16km ESE Morales, 15.41186°N, 88.71671°W, 410m, 19 May 2009 (LLAMA); Finca La Firmeza, 10.7km SE Morales, 15.4067°N, 88.6967°W, 500m, 19 Sep 2008 (M. G. Branstetter); Petén : 13km NW Machaquilá, 16.44603°N, 89.54940°W, 400m, 27 May 2009 (LLAMA); HONDURAS: Comayagua : P.N. Cerro Azul Meambar, 14.86601°N, 87.89955°W, 880m, 21 May 2010 (LLAMA); P.N. Cerro Azul Meambar, 3.4km ESE La Guama, 14.8722°N, 87.9053°W, 760m, 26 Sep 2008 (M. G. Branstetter); Gracias a Dios : Las Marias, 15.72245°N, 84.88078°W, 510m, 10 Jun 2010 (LLAMA); Las Marias, 15.71844°N, 84.87811°W, 370m, 10 Jun 2010 (M. G. Branstetter); Olancho : 11km NNE Catacamas, 14.94493°N, 85.84928°W, 1100m, 12 May 2009 (J. Longino); MEXICO: Chiapas : Lago Metzabok, 17.12622°N, 91.63056°W, 570m, 5 Jun 2008 (LLAMA); Nahá, 16.97444°N, 91.58702°W, 180m, 24 Jun 2008 (LLAMA); Ocosingo, [approx. 16.906°N, 92.099°W], 2 Jun 1969 (J. M. Campbell); Playón de la Gloria, 16.14826°N, 90.89704°W, 180m, 24 Jun 2008 (LLAMA); Oaxaca : 5.7km SW Valle Nacional, 17.73067°N, 96.3304°W, 560m, 13 Aug 2009 (M. G. Branstetter); 7.5km S Valle Nacional, 17.70752°N, 96.30516°W, 680m, 12 Aug 2009 (M. G. Branstetter); NICARAGUA: Jinotega : P.N. Cerro Saslaya, 13.77171°N, 85.01263°W, 1110m, 12 May 2011 (LLAMA).
Distribution map of Stenamma diversum.
urn:lsid:zoobank.org:act:D105B73A-B44E-4420-882E-6C4180D8E149
http://species-id.net/wiki/Stenamma_excisum
Worker: Figures 77–79; Queen: Figure 80; Map: Figure 81Holotype worker. HONDURAS, Atlántida: 12km SW La Ceiba, 15.69150°N, 86.86151°W ±20m, 280m, 19 Jun 2010, tropical rainforest, ex sifted leaf litter (LLAMA, collection Wa-C-09-2-44) [USNM, specimen CASENT0621834]. Paratypes: same data as holotype but 15.69449°N, 86.86330°W ±20m, 200m (Wa-C-09-1-27) [1dq, 1w, CAS, CASENT0621799, CASENT0621801], [1w, EAPZ, CASENT0621800], [1w, FMNH, CASENT0623304], [1w, INBio, CASENT0623305], [1w, LACM, CASENT0623306]; 15.69175°N, 86.86091°W ±20m (LLAMA, Wa-C-09-2-04) [1dq, 1w, MCZ, CASENT0623307, CASENT0623309], [1dq, 1w, USNM, CASENT0623308, CASENT0623310]; 15.69449°N, 86.86344°W ±20m (LLAMA, Wa-C-09-1-24) [1w, UCD, CASENT0621793]; 15.69134°N, 86.86137°W ±20m, 280m, 19 Jun 2010 (LLAMA, Wa-C-09-2-40) [1dq, 1w, MGBPC, CASENT0623541, CASENT0623540].
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. Similar species: Stenamma saenzae.
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.
Stenamma excisum holotype worker (CASENT0621834) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Stenamma excisum worker variants 1. Face, profile, dorsal views A–C Variant 1 (CASENT0605563) D–F Variant 2 (CASENT0605441) G–I Variant 3 (CASENT0605500) J–L Variant 4 (CASENT0605743).
Stenamma excisum worker variants 2. Face, profile, dorsal views A–C Variant 5 (CASENT0126237) D–F Variant 6 (CASENT0126243) G–I Variant 7 (CASENT0622434).
(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.
Stenamma excisum paratype queen (CASENT0621799) A Profile B Face C Dorsum.
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. Stenamma excisum has been collected from 60–2280 m elevation and occurs in a variety of wet forest environments (e.g. tropical rainforest, cloud forest, disturbed mesophyll forest, pine-oak forest, riparian wet forest). The excision in the anterior clypeal margin of most specimens is unique and suggests some sort of diet specialization different from other Stenamma species.
Although quite variable across its range (see below), Stenamma excisum is separable from similar species using the characters in the key and diagnosis. Several species are superficially similar to Stenamma excisum, but no other species has the same unique clypeal structure. Phylogenetic results indicate that Stenamma excisum is sister to Stenamma lagunum and that this clade is sister to the remainder of MAC Stenamma species (
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 Stenamma excisum as it is described here. However, too few collections have been made in intervening areas to adequately assess variation and species boundaries.
Variant 1 (Figure 78A–C) is known mainly from mid-elevation sites along the wet eastern slopes of the mountains in Oaxaca, Mexico, but a single specimen is also known from Nahá in Chiapas, Mexico. This variant is very similar to the type population, but differs as follows: smaller overall body size; body color darker, mostly dark orange-brown; sculpture on pronotal dorsum longitudinally rugose, without reticulae.
Variant 2 (Figure 78D–F) occurs in the same area as variant 1, but the two forms have not been collected together. It is similar to variant 1 except for the following: anterior clypeal margin with a shallow median emargination, lacking a deep excision; propodeal spines in profile longer and somewhat sinuous; anterior margin of propodeal dorsum with a more distinct welt; postpetiole in dorsal view much wider than petiolar node (PW/PPW 0.55–0.61).
Variant 3 (Figure 78G–I) is known from a single aberrant specimen collected near variant 2, but from a slightly higher elevation. It differs from variant 2 as follows: anterior clypeal margin intermediate between variant 1 and 2, with a shallow excision; propodeal spines short, almost tuberculate; antennal segments 11 and 12 noticeably bulging; postpetiolar node mostly smooth; postpetiole in dorsal view not greatly expanded; upper layer of setae on gastral dorsum shorter, sparser, and subdecumbent.
Variant 4 (Figure 78J–L) is found mainly in northern Mexico in Tamaulipas (El Cielo) and Nuevo León (Monterrey de Chipinque) states. It is a very small version of Stenamma excisum and is quite divergent from the type population. I originally intended to describe it as a separate species, but later found intermediate-looking specimens from allopatric populations in Querétaro and Veracruz, Mexico. It differs from the type population as follows: overall body size much smaller; sculpture reduced; face with only light rugoreticulae, punctae, and carinulae; dorsum of pronotum with sparse longitudinal carinulae, interstices shiny; propodeal spines short; decumbent layer of pilosity on gastral dorsum sparse, suberect layer slightly thickened.
Variant 5 (Figure 79A–C) occurs in Veracruz, Mexico. It is intermediate between the type population and variant 4. It differs from variant 4 as follows: head and body densely punctate; dorsum of pronotum with longitudinal rugulae/carinulae, but interstices punctate; petiolar node reaching a sharper apex, which points distinctly posteriad; suberect layer of pilosity on gastral dorsum (and most of body dorsum) longer.
Variant 6 (Figure 79D–F) is known from two localities in Veracruz, Mexico (Paraje Nuevo, Tetzonapa). It is similar to variant 5, but differs as follows: anterior clypeal margin with a median emargination, lacking an excision; body larger and with distinctly elongate appendages; decumbent layer of gastral pilosity very sparse.
Variant 7 (Figure 79G–I) is known from a single locality in Nuevo León, Mexico (38 km SSW Monterrey). It is very similar to variant 4, but differs as follows: anterior clypeal margin with a median emargination, lacking a deep excision; larger overall size; head larger, more robust, with a distinct median depression in posterior margin; petiolar node in profile broadly domed and more asymmetrical, with a longer moresloping anterior face.
HONDURAS: Atlántida : 12km SW La Ceiba, 15.69449°N, 86.86544°W, 200m, 19 Jun 2010 (LLAMA); 12km SW La Ceiba, 15.69150°N, 86.86151°W, 280m, 19 Jun 2010 (LLAMA); Gracias a Dios : Las Marias, 15.70861°N, 84.86211°W, 80m, 8 Jun 2010 (LLAMA); MÉXICO: Chiapas : Nahá, 16.96399°N, 91.59283°W, 985m, 8 Jun 2008 (LLAMA); Nuevo Leon : Monterrey, Chipinque Mesa, [ca. 25.61°N, 100.36°W], 1650m, 22 Jun 1969 (S. & J. Peck); 38km SSW Monterrey, [ca. 25.367°N, 100.467°W], 2280m, 15 Jul 1979 (P. Ward); Oaxaca : Mirador Grande, 17.89844°N, 96.36253°W, 990m, 14 Aug 2009 (M. G. Branstetter); 10.8km SW Valle Nacional, 17.68102°N, 96.33026°W, 1120m, 13 Aug 2009 (M. G. Branstetter); 13.2km SW Valle Nacional, 17.65934°N, 96.33426°W, 1360m, 11 Aug 2009 (M. G. Branstetter); 20.6km SW Valle Nacional, 17.60404°N, 96.37786°W, 1733m, 13 Aug 2009 (M. G. Branstetter); 25km S Valle Nacional, [ca. 17.670°N, 96.330°W], 1200m, 21 May 1971 (S. B. Peck); Querétaro : 29km E Landa de Matamoros, [ca. 21.27°N, 99.16°W], 1600m, 14 Jul 1969 (S. & J. Peck); Tamaulipas : El Cielo, nr Alta Cima, 23.06518°N, 99.20433°W, 980m, 21 Aug 2009 (M. G. Branstetter); El Cielo, 1.2km SE Alta Cima, 23.05005°N, 99.19226°W, 24 Aug 2009 (M. G. Branstetter); 2.5km SSE Alta Cima, 23.03787°N, 99.18941°W, 870m, 24 Aug 2009 (M. G. Branstetter); El Cielo, 1.2km N La Gloria, 23.05717°N, 99.25206°W, 1780m, 23 Aug 2009 (M. G. Branstetter); nr Gomez Farias Rancho del Cielo, [ca. 23.063°N, 99.205°W], 1000m, 7 Aug 1983 (S. & J. Peck); Veracruz : Cordoba, Paraje Nuevo, Nacimiento, [ca. 18.88°N, 96.86°W], 7 Aug 1969 (S. & J. Peck); Fortín, Canyon Río Metlac, [ca.18.90°N, 97.00°W], 5 Aug 1969 (S. & J. Peck); 1.9km S Huatusco, [ca. 19.20°N, 96.95°W], 1344m, 2–5 Aug 1969 (S. & J. Peck); Pueblo Nuevo, nr Tetzonapa, Aug 1953 (E. O. Wilson).
Distribution map of Stenamma excisum (circles) and Stenamma expolitico (squares).
urn:lsid:zoobank.org:act:4CE4E3F9-6FB0-4859-85B4-B4923E513FCE
http://species-id.net/wiki/Stenamma_expolitico
Worker: Figure 82; Map: Figure 81Holotype worker. COSTA RICA, Alajuela Prov.: Rio Peñas Blancas, 10.3167°N, 84.7167°W, 800m, 23 May 1990, wet forest, ex sifted leaf litter (J. Longino, collection JTL2701-s) [INBio, specimen INBIOCRI001282473]. Paratypes: COSTA RICA, Heredia: 16km N Vol. Barba, 10.267°N, 84.083°W, 1020m, 9 Jul 1986 (J. Longino, JTL1340-s) [1w, USNM, CASENT0623094]; same data but 22km N Volcan Barba, 10.333°N, 84.067°W, 500m, 5 Mar 1985 (J. Longino, JTL1340-s) [1w, LACM, CASENT0623093]; 16km SSE La Virgen, 10.267°N, 84.083°W, 1100m, 14–17 Mar 2001 (ALAS, 11/WF/02/28) [1w, UCD, INB0003212462].
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. Similar species: Stenamma alas, Stenamma expolitum.
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.
Stenamma expolitico holotype worker (INBIO282473) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Unknown.
Unknown.
Stenamma expolitico is known from only four specimens, collected from sifted leaf litter in wet forest between 500–1100 m elevation.
Stenamma expolitico belongs to the expolitum species group, which also includes Stenamma alas and Stenamma expolitum (a diagnosis for this group is given under Stenamma expolitum below). Stenamma expolitico can be separated from Stenamma alas and Stenamma expolitum by the presence of transverse furrows on the promesonotal dorsum, and by the shape of the propodeum in profile, which in Stenamma expolitico forms a blunt 90° angle where the dorsal and declivitous faces meet.
I am not completely convinced that Stenamma expolitico is a good species because it is known from only a few workers and is somewhat intermediate in morphology between Stenamma expolitum and Stenamma tico. It could be a hybrid form, or a rare variant of Stenamma expolitum or Stenamma alas. I choose to recognize it as separate species here, because, although rare, it does occur at several sites in sympatry with the other expolitum group species, and it is easily identifiable. Furthermore, many nests of Stenamma alas and Stenamma expolitum have been excavated, and workers with Stenamma expolitico -like morphology have never been found. More material of Stenamma expolitico, especially from nest series, is needed to test this hypothesis and confirm its status as a real biological species.
See type material.
http://species-id.net/wiki/Stenamma_expolitum
Worker: Figure 83; Queen: Figure 84A–D; Male: Figure 84E–G; Map: Figure 85Integument 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). Similar species: Stenamma atribellum, Stenamma alas, Stenamma expolitico.
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.
Stenamma expolitum A, B, E, G Worker (CASENT0600043) C, D, F, H Paratype worker (CASENT0126347).
(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 Figure 84D.
Stenamma expolitum A Queen (CASENT000005582), profile B Same, face C Same, dorsum D Same, wings E Male (JTLC000003501), profile F Same, face G Same, dorsum.
See Figure 84E–G.
Stenamma expolitum is a specialist inhabitant of clay banks. It occurs from 50–1300 m elevation in mature wet forest environments. Nests are generally found in vertical clay banks along streams or cuts along trails. The biology of Stenamma expolitum is reviewed in detail in
Stenamma expolitum, along with Stenamma alas and Stenamma expolitico, belongs to the expolitum species group. This group is defined by the following: propodeal spines absent; petiole and postpetiole almost completely smooth and shiny, with only faint punctae sometimes present laterally; postpetiole in profile bulging, globular, appearing more voluminous than petiolar node; anterior clypeal margin with a median emargination; basal margin of mandible straight. Molecular phylogenetic data firmly show that Stenamma zelum is sister to the expolitum group (
Stenamma expolitum can be separated from other expolitum group species by its nearly completely smooth and shiny face. In the field, Stenamma expolitum can be separated from Stenamma alas by its nest structure, described above. Although they do not co-occur, Stenamma expolitum looks superficially like Stenamma atribellum, which is restricted to Cusuco, Honduras. Both species have completely smooth sculpturing, but the latter species has the anterior constriction of the gaster distinctly elongate.
COSTA RICA: Alajuela : Casa Eladio, Río Peñas Blancas, 10.3167°N, 84.7167°W, 800m, 2 Feb 1994 (J. Longino); 2km N Volcan Arenal, 10.483°N, 84.700°W, 600m, 3 Apr 2005 (J. Longino); Heredia : La Selva Biol. Sta., 10.4333°N, 84.0167°W, 50m, 8 Aug 2004 (J. Longino); P.N. Braulio Carrillo, 10.3378°N, 84.0500°W, 300m, 19 Sep 2006 (TEAM); 11km ESE la Virgen, 10.35°N, 84.05°W, 300m, 5 Nov 2003 (J. Longino); 10km SE La Virgen, 10.3333°N, 84.0833°W, 500m, 16 Feb 2003 (J. Longino); Limón : Colombiana Farm, [ca. 10.167°N, 83.583°W], Apr 1924 (W. M. Mann); NICARAGUA: Jinotega : PN Cerro Saslaya, 13.76650°N, 85.02485°W, 1040m, 12 May 2011 (LLAMA);Matagalpa : RN El Musún, 4.9km NNW Río Blanco, 12.97471°N, 85.23318°W, 1205m, 11 Oct 2008 (M. G. Branstetter).
Distribution map of Stenamma expolitum.
http://species-id.net/wiki/Stenamma_felixi
Worker: Figures 86, 87; Queen: Figure 88A–D; Male: Figure 88E–G; Map: Figure 89Integument 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: Stenamma manni, Stenamma schmidti.
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.
Stenamma felixi A, B, E, G, H Worker (CASENT0620969) C, D, F Lectotype worker (CASENT0126355).
Stenamma felixi variant (CASENT0622555) A Face B Profile C Dorsum.
(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 Figure 88D.
Stenamma felixi A Queen (CASENT0622308), profile B Same, face C Same, dorsum D Same, wings E Male (CASENT0622311), profile F Same, face G Same, dorsum.
See Figure 88E–G.
Stenamma felixi is one of the most widespread and common species of MAC Stenamma. It occurs from approximately 50–1600 m, but is most common above 500 m, and is always found in wet forest environments, ranging from lowland rainforest to cloud forest. Workers have been collected by sifting leaf litter, beating and sweeping vegetation, baiting, using pitfall and Malaise traps, and by general searching. Nests are generally quite large and have been found in rotting logs on the ground, in tree stumps, under bark of logs, and rarely in mud banks. A few lone foundresses have been found under rotting epiphyte clumps in old treefalls, and some workers have been collected from orchids at quarantine in the U.S. All colonies collected so far have been monogynous. Workers seem to be epigeic, solitary foragers, but nothing is known about dietary preference. A very common experience is to find lone, stray workers running across medium- to large-sized logs in forest.
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, Stenamma felixi shows considerable variation in the density and orientation of its sculpturing as well as in petiole shape. However, I have only identified one distinct variant (Figure 87) worth describing in more detail. Specimens from Nicaragua and Costa Rica have very deep carinate sculpturing on the face and mesosoma. The facial carinae are usually longitudinal, but occasionally reticulate laterad. The pronotal dorsum has longitudinal carinae, but the side of the pronotum has arcuate carinae that wrap across the dorsum of the metanotum. The head in profile view has a very distinctive shape, in which the posterior margin of the head is very wide compared to the anterior margin, giving the head a somewhat triangular appearance. Lastly, the petiolar and postpetiolar nodes are noticeably smooth. Specimens from Colombia and Ecuador and from north of Nicaragua lose the carinate sculpture. I treat all of the slight sculpture differences among popluations as intraspecific variation as I have found no evidence of sympatry among forms.
BELIZE: Cayo : Chiquibul N.P., Doyle’s Delight, 16.48972°N, 89.04583°W, 950m, 20–27 Aug 2007 (P. W. Kovarik); Chiquibul N.P., Doyle’s Delight, 16.4833°N, 89.0333°W, 1000m, 19–22 Aug 2007 (P. W. Kovarik); Chiquibul N.P., Doyle’s Delight, 16.49305°N, 89.04694°W, 1100m, 19–28 Aug 2007 (P. W. Kovarik); COLOMBIA: Valle del Cauca : Buenaventura, Bajo Calima, [ca. 3.996°N, 76.974°W], 30m, 16–17 Mar 1967 (Root & Brown); Buenaventura, 3.2km above Río Aguaclara on old rd to Cali, [ca. 3.693°N, 76.925°W], 17–19 Jun 1971 (W. L. Brown); Reserva Forestal Escalerete, [ca. 3.886°N, 77.069°W], 80m, 29 May 2007 (Usma-Aldana); COSTA RICA: Cartago : La Carpintera, [ca. 9.884°N, 83.983°W], 1500m, Apr 1924 (W. M. Mann); Navarro Farm, [ca. 9.8167°N, 83.8833°W, ], 1100m, Mar 1924 (W. M. Mann); Guanacaste : Est. Mengo, SW side Volcan Cacao, [ca. 10.933°N, 85.450°W], 1100m, Feb 1989 (GNP Biodiversity Survey); Est. Pitilla, 9km S Sta. Cecilia, [ca. 10.983°N, 85. 433°W], 700m, Jan 1991 (Curso Microhymenoptera); Heredia : La Selva Biol. St., [ca 10.430°N, 84.007°W], 100m, 16 Jan 1979 (S. P. Cover); 11km ESE La Virgen, 10.35°N, 84.05°W, 300m, 15 Feb 2004 (ALAS); 10km NE Vara Blanca, 10.2333°N, 84.0833°W, 1500m, 12 Feb 2005 (ALAS); 16km SSE La Virgen, 10.2667°N, 84.0833°W, 1100m, 14–17 Mar 2001 (ALAS); Limón : Colombiana Farm, [ca. 10.167°N, 83.583°W], Apr 1924 (W. M. Mann); Río Toro Amarillo, vic. Guapiles, [ca. 10.205°N, 83.789°W], 300m, 25 Feb–9 Mar 1966 (W. L. Brown); Puntarenas : Altamira Biological Station, 9.02922°N, 83.00813°W, 1400m, 30 May 2007 (M. G. Branstetter); 2km W Las Alturas, 8.933°N, 82.850°W, 1260m, 23 Mar 1990 (P. S. Ward); Las Cruces Biological Station, 8.78658°N, 82.95987°W, 1150m, 29 May 2007 (M. G. Branstetter); Monteverde, 0.25km E Lecheria, [ca. 10.307°N, 84.810°W], 1300m, 19 May 1988 (Cover et al.); San José : Bajo La Hondura, Braulio Carillo Nat. Park, [ca. 10.067°N, 83.983°W], 20 Jun 1926 (F. Nevermann); 1km N La Ese, 9.450°N, 83.717°W, 1400m, 5 Aug 1985 (P. S. Ward); Pan-Am Hwy, 23 rd km N San Isidro de General, [ca. 9.466°N, 83.703°W], 1600m, 20–23 Jun 1974 (Harding & Donahue); 2km E San Gerardo, 9.467°N, 83.583°W, 1600m, 4 Aug 1985 (P. S. Ward); ECUADOR: Cañar : 2–6km above Cochancay on Guayaquil-Tambo H’way, [ca. 2.468°S, 79.291°W], 600m, 25 Jul 1973 (W. L. Brown); GUATEMALA: Izabal : 5km NW Morales, 15.51351°N, 88.86647°W, 240m, 18 May 2009 (J. Longino);Petén : 13km NW Machaquilá, 16.44202°N, 89.53495°W, 390m, 28 May 2009 (LLAMA);Zacapa : 2km SE La Unión, 14.94701°N, 89.27594°W, 1550m, 12 May 2009 (LLAMA); Suchitepéquez : 5.5km S Vol. Atitlán, 14.52857°N, 91.19569°W, 1070m, 18 Jun 2009 (LLAMA); HONDURAS: Atlántida : San Juan Pueblo, [ca. 15.583°N, 87.233°W], 100m, Feb–Mar 1920 (W. M. Mann); Cortés : PN Cusuco, 15.48940°N, 88.23732°W, 1290m, 30 May 2010 (LLAMA); Olancho : 9km N Catacamas, 14.93646°N, 85.90488°W, 1340m, 10 May 2010 (M. G. Branstetter); 10km N Catacamas, 14.94125°N, 85.90385°W, 1320m, 10 May 2010 (LLAMA); 11km NNE Catacamas, 14.95031°N, 85.86229°W, 1470m, 12 May 2009 (J. Longino); PN La Muralla, 15.09721°N, 86.73840°W, 1480m, 5 May 2010 (LLAMA); MÉXICO: Chiapas : Lago Metzabok, 17.12360°N, 91.63766°W, 575m, 5 Jun 2008 (LLAMA); Nahá, 16.94885°N, 91.59489°W, 930m, 8 Jun 2008 (LLAMA);Hidalgo : 6.4km SW Chapulhuacán, [ca. 21.155°N, 98.931°W], 1070m, 27 Jun–1 Jul 1973 (A. F. Newton); between Real del Monte and El Chico, [ca. 20.138°N, 98.673°W], Mar–Aug 1913 (W. M. Mann); Oaxaca : Mirador Grande, 17.89844°N, 96.36253°W, 990m, 14 Aug 2009 (M. G. Branstetter);Puebla : 17km NE Teziutlán, [ca. 19.877°N, 97.310°W], 1940m, 7 Jun 1988 (W. P. MacKay); 24km N Xicotepec de Juarez, [ca. 20.282°N, 97.963°W], 1070m, 17 Jun 1983 (R. S. Anderson); Veracruz : Canyon Río Metlac, near Fortín, [ca. 18.90°N, 97.00°W], 975m, 7–28 Aug 1973 (A. F. Newton); 10km S Orizaba, 18.750°N, 97.083°W, 1500m, 19 Mar 1985 (P. S. Ward); 2.7km N Teocelo, [ca. 19.40°N, 96.98°W], 1130m, 22–24 Jul 1973 (A. F. Newton); 11km N San Andrés Tuxtla, 18.55°N, 96.00°W, 1400m, 23 Mary 1985 (P. S. Ward); 10km NNW Sontecomapan, 18.583°N, 95.083°W, 200m, 20 Mar 1985 (P. S. Ward); NICARAGUA: Jinotega : PN Cerro Saslaya, 13.77173°N, 85.01286°W, 1110m, 12 May 2011 (LLAMA);RNDatanlí El Diablo, 13.10410°N, 85.01286°W, 1110m, 12 May 2011 (LLAMA);RNDatanlí El Diablo, 13.08051°N, 85.87462°W, 1170m, 20 May 2011 (LLAMA); PANAMA: Panama : Cerro Campana, [ca. 8.678°N, 79.928°W], 875m, 17 Jan 1960 (Fairchild & Brown).
Distribution map of Stenamma felixi.
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http://species-id.net/wiki/Stenamma_hojarasca
Worker: Figure 90; Map: Figure 91Holotype worker.HONDURAS: Cortés, Parque Nacional Cusuco, 15.48939°N, 88.23678°W ±300m, 1280m, 31 May 2010, mesophyll forest, ex sifted leaf litter (LLAMA, collection Wm-C-06-1-03) [USNM, specimen CASENT0622132]. Paratypes: same data as holotype but 15.48896°N, 88.23439°W ±20m, 1290m, 30 May 2010 (LLAMA, Wm-C-06-1-01) [1w, CAS, CASENT0622124]; 15.48683°N, 88.23422°W ±300m, 1340m, 30 May 2010 (LLAMA, Wm-C-06-1-02) [1w, EAPZ, CASENT0622126]; 15.48723°N, 88.23482°W ±20m, 1330m, 30 May 2010 (LLAMA, Wa-C-06-1-08) [1w, INBio, CASENT0621692]; 15.48717°N, 88.23476°W ±20m, 1330m, 30 May 2010 (LLAMA, Wa-C-06-1-10) [1w, MCZ, CASENT0621693]; 15.48940°N, 88.23695°W ±20m, 1290m, 30 May 2010 (LLAMA, Wa-C-06-2-28) [1w, UCD, CASENT0621752]; 15.48839°N, 88.23592°W ±60m, 1260m, 31 May 2010 (LLAMA, Wm-C-06-2-02) [1w, MGBPC, CASENT0622169].
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). Similar species: Stenamma catracho, Stenamma cusuco, Stenamma ochrocnemis.
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.
Stenamma hojarasca holotype worker (CASENT0622132) A Profile B Face C Dorsum D Anterior clypeal margin in anterodorsal view E Gaster.
Unknown.
Unknown.
Stenamma hojarasca is a rare species known almost exclusively from sifted leaf litter collected from the forest floor. The only exception is a single specimen from a carrion baited pitfall trap in Belize. This species is found in montane mesophyll and cloud forest habitats and has been collected from 1100–1550 m elevation.
The form of the waist segments combined with the projecting median clypeal lobe, make Stenamma hojarasca a very distinctive species. It should not be confused easily with any other MAC species.
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 Stenamma hojarasca belongs to a clade that includes Stenamma cusuco and Stenamma ochrocnemis, with the latter being its sister species (Branstetter unpublished data).
BELIZE: Cayo : Chiquibul N.P., Doyle’s Delight, 16.49305°N, 89.04694°W, 1100m, 19–28 Aug 2007 (P. W. Kovarik); GUATEMALA: Zacapa : 2km SE La Unión, 14.94460°N, 89.27726°W, 1550m, 12 May 2009 (LLAMA); HONDURAS: Cortés : PN Cusuco, 15.48939°N, 88.23678°W, 1280m, 31 May 2010 (LLAMA); PN Cusuco, 15.48683°N, 88.23422°W, 1340m, 30 May 2010 (LLAMA); MÉXICO: Chiapas : 10.6km W El Bosque, 17.043°N, 92.762°W, 1460m, 25–29 Aug 1973 (A. F. Newton).
Distribution map of Stenamma hojarasca (circles) and Stenamma huachucanum (squares).
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http://species-id.net/wiki/Stenamma_huachucanum
Worker: Figures 92–94; Queen: Figure 95; Map: Figure 91Note 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. Similar species: Stenamma connectum, Stenamma crypticum.
Southwestern U.S.A. to southern Mexico (Oaxaca).
Worker description. (28 measured; paratype CASENT0105666 in parentheses) HL 0.52–0.72 (0.72), HW 0.46–0.64 (0.62), FLD 0.11–0.17 (0.17), PCW 0.03–0.05 (0.03), SL 0.43–0.55 (0.55), EL 0.07–0.12 (0.10), ACL 0.43–0.52 (0.52), ML 0.66–0.85 (0.85), PrW 0.33–0.41 (0.41), PSL 0.06–0.11 (0.09), SDL 0.05–0.09 (0.08), PL 0.23–0.35 (0.32), PH 0.15–0.21 (0.21), PW 0.11–0.16 (0.15), PPL 0.13–0.20 (0.17), PPH 0.14–0.19 (0.18), PPW 0.15–0.20 (0.19), MFL 0.44–0.59 (0.59), MTL 0.37–0.48 (0.48), CI 83–91 (86), SI 81–94 (89), REL 14–21 (15), FLI 23–29 (27), PSI 1.0–1.4 (1.1), MFI 101–124 (105), ACI1 67–71 (68), ACI2 92–103 (95).
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.
Stenamma huachucanum A, B, E, G, H Worker (CASENT0126556) C, D, F Paratype worker (CASENT010566).
Stenamma huachucanum worker variants 1. Face, profile, and dorsal views A–C Variant 1 (CASENT0600124) D–F Variant 2 (CASENT0605749) G–I Variant 3 (CASENT0605412) J–L Variant 4 (CASENT0604596).
Stenamma huachucanum worker variants 2. Face, profile, and dorsal views A–C Variant 5 (CASENT0605616) D–F Variant 6 (CASENT0605647).
(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 Figure 95D.
Stenamma huachucanum A Queen (CASENT0600094), profile B Same, face C Same, dorsum D Same, wings.
Unknown.
Stenamma huachucanum is a cryptic leaf litter ant known mostly from Winkler or Berlese samples of sifted leaf litter. As defined here, the species is widely distributed, occupying both relatively dry, seasonal habitats (e.g. tropical deciduous forest with juniper, oak-pine-juniper woodland, oak woodland, oak-pine-douglas fir forest) and tropical wet forest habitats (e.g. mesophyll forest, cloud forest, oak-pine forest). Collections have been made from 1000–2900 m, but the species is most common between 1600–2500 m. In seasonal habitats, workers have been found underneath rocks, in addition to the leaf litter.
Stenamma huachucanum forms a difficult complex composed of many divergent allopatric populations. The complex probably includes several good biological species, but I have chosen to lump most forms into a single entity, because there is no clear evidence of sympatry among forms, and some specimens appear to have intermediate phenotypes that connect distinct forms. The exceptions to this lumping approach are the similar species Stenamma connectum and Stenamma crypticum. These species are morphologically similar to Stenamma huachucanum, but molecular phylogenetic results provide strong evidence that they represent separate lineages (Branstetter unpublished data).
There are some key morphological differences separating populations of Stenamma huachucanum from Stenamma connectum and Stenamma crypticum, but considering all of the variation among populations, the easiest way to identify species is with geography. Stenamma huachucanum occurs from the southwestern U.S.A to Oaxaca, where it is found only in the drier, interior of the state. In eastern Mexico, the species is found from Tamaulipas to Puebla, with no records from Veracruz. Stenamma connectum is found in Veracruz, Mexico and on the wetter, Caribbean slope of Oaxaca. Stenamma crypticum occurs mainly from Chiapas, Mexico to Nicaragua. However, as noted above under both Stenamma crypticum and Stenamma connectum, a few putative Stenamma crypticum specimens are known from one sample taken in Veracruz, nearly in sympatry with Stenamma connectum. If confirmed, this latter case is the only evidence of sympatry among any of these similar-looking species.
Within the Stenamma huachucanum complex there is considerable variation among populations. To help in the identification of Stenamma huachucanum, and to aid future taxonomic efforts, I describe several Stenamma huachucanum variants below.One observation about variation within the complex is that specimens from western and central Mexico, where it is drier, tend to look more like the type population. Specimens from the eastern slope, where it is wetter, become smoother and more aberrant, in general. The most problematic areas are in central Mexico where it transitions from wet to dry. At these localities I find specimens with intermediate features. This is one of the main reasons I have lumped what seem like very different populations together.
The main features of the type form of Stenamma huachucanum are indicated in the species description above (see parenthetical comments). The key characters for the type form are sculpturing and the shape of the petiole. The head is completely sculptured and mostly rugoreticulate-punctate, with some longitudinal carinulae along the midline. The pronotum is lightly carinulate-punctate (longitudinal orientation), with small smooth patches on the dorsum. One characteristic unseen by
Variant 1 (Figure 93A–C) was collected in Sinaloa, Mexico. It is very similar to the type form except as follows: promesonotum more domed; pronotal dorsum strongly carinulate, without punctae.
Variant 2 (Figure 93D–F) was collected from Volcán de Tequila in Jalisco, Mexico. It is similar to the type form except as follows: pronotum mostly smooth; petiolar node in profile broader; petiolar venter in profile straight, not sinuous.
Variant 3 (Figure 93G–I) is from the drier portion of Oaxaca (Asunción Nochixtlan). It varies from the type form as follows: promesonotum distinctly domed and mostly smooth; petiolar node broader, more robust; face sculpture reduced, with posterior margin becoming smooth, and rugoreticulae less visible. Similar-looking specimens also occur in Mexico state (Temascaltepec).
Variant 4 (Figure 93J–L) is from another site in Oaxaca (10.6km N Jct 190/135). It varies from the type as follows: promesonotum in profile low-domed and appearing very long, with the anterior face sloping gently into the metanotal groove; pronotum with many longitudinal carinulae around humeri, and a few on dorsum; gastral pilosity longer.
Variant 5 (Figure 94A–C) is a wet forest version of Stenamma huachucanum. It occurs at several sites on the eastern slope of the mountains in Querétaro. There are similar looking specimens from San Luis Potosí and Hidalgo states. It differs from the type population as follows: overal body size smaller; face sculpture reduced, with posterior ¼ or less of head smooth, remaining sculpture consisting of fine punctae, carinulae, and rugoreticulae; pronotum and dorsum of mesonotum smooth; promesonotum in profile variable, sometimes more flat, with posterior face merging smoothly into metanotum, sometimes more distinctly domed; petiolar node in profile narrow, appearing anteroposteriorly compressed. The specimens with a more domed promesonotum appear intermediate with variant 6.
Variant 6 (Figure 94D–F) is another wet forest version of Stenamma huachucanum. It occurs throughout the El Cielo reserve in Tamaulipas, Mexico, and differs from variant 5 as follows: head more robust, slightly broader, with posterior margin distinctly depressed medially; promesonotum domed, only slightly asymmetrical; propodeal spines forming well-developed, blunt tubercles; outline of propodeum in profile sinuous, with propodeal spine and lobe smoothly connected; petiole appearing elongate; petiolar node small, with a rounded dorsum that points distinctly posteriad.
MÉXICO: Coahuila : H’way 57, pass 25km E Saltillo, [ca. 25.436°N, 100.806°W], 11 Aug 1965 (Cornell Univ. Mexico Field Party); Hidalgo : 6.4km SW Chapulhuacán, [ca. 21.155°N, 98.931°W], 1070m, 27 Jun–1 Jul 1973 (A. F. Newton);11km SW Chapulhuacán, [ca. 21.147°N, 98.966°W], 1200m, 5 Jul 1976 (A. F. Newton); 18km NE Jacala, nr El Alamo, [ca. 21.058°N, 99.052°W], 1700m, 10 Jun 1988 (S. & J. Peck); 16km NE Rancho Viejo, [ca. 21.07°N, 99.02°W], 1550m, 5 Jul 1976 (A. F. Newton); Tlanchinol, 43km SW Huejutla, [ca. 20.988°N, 98.662°W], 1 Aug 1983 (S. & J. Peck); 4–6km N Tlanchinol, [ca. 21.026°N, 98.644°W], 1590m, 6–11 Jul 1973 (A. F. Newton);Jalisco : Volcán de Tequila, 20.82517°N, 103.85509°W, 1840m, 26 Aug 2009 (M. G. Branstetter); Volcán de Tequila, 20.79293°N, 103.85388°W, 2660m, 26 Aug 2009 (M. G. Branstetter); México : 1.6km E Ixtapan de la Sal, [ca. 18.85°N, 99.67°W], 1890m, 10 Aug 1973 (A. F. Newton); 4.8km NE Temascaltepec, [ca. 19.05°N, 100.03°W], 1920m, 14 Sep 1973 (A. F. Newton); 11km NE Temascaltepec, [ca. 19.096°N, 99.983°W], 2100m, 21–15 Sep 1973 (A. F. Newton); 3.2km NE Tenancingo, [ca. 18.985°N, 99.575°W], 2160m, 11 Sep 1973 (A. F. Newton); Oaxaca : 9.4km SE Asunción Nochixtlan, 17.37632°N, 97.19976°W, 2240m, 10 Aug 2009 (M. G. Branstetter); 10.6km N Jct 190/135 (on 135), [ca. 17.282°N, 96.929°W], 1920m, 21 Jul 1987 (R. S. Anderson); 14km NE Oaxaca, km 10 Mex. 175, [ca. 17.145°N, 96.622°W], 1890m, 20 Aug 1973 (A. F. Newton); Puebla : 2.7km S Apulco, nr Zacapoaxtla, [ca. 19.911°N, 97.606°W], 22 Jul 1987 (R. S. Anderson); 7.6km SW La Cumbre, [ca. 20.1416°N, 98.0217°W], 1585m, 23 Jul 1987 (R. S. Anderson); H’way 190, E Río Frío [ca. 19.310°N, 98.638°W], 2900m, 7 Aug 1965 (Cornell Univ. Mexico Field Party); 2km NE Teziutlan, [ca. 19.892°N, 97.296°W], 1220m, 19 Jun 1983 (S. & J. Peck); 24km N Xicotepec de Juarez, [ca. 20.282°N, 97.963°W], 1070m, 17 Jun 1983 (R. S. Anderson); Querétaro : Cerro Zamorano, 20.9328°N, 100.1840°W, 2770m, 26 Jul 2006 (R. S. Anderson); 29km E Landa de Matamoros, [ca. 21.27°N, 99.16°W], 1600m, 14 Jul 1969 (S. & J. Peck); 40km E Landa de Matamoros, [ca. 21.297°N, 99.090°W], 1520m, 14 Jul 1969 (S. & J. Peck); 4.6km SW Pinal de Amoles, 21.16479°N, 99.59419°W, 1960m, 18 Aug 2009 (M. G. Branstetter); 1.9km NE Pinal de Amoles, 21.14974°N, 99.61576°W, 2250m, 18 Aug 2009 (M. G. Branstetter); 7km NE Pinal de Amoles, 21.17601°N, 99.57341°W, 1700m, 18 Aug 2009 (M. G. Branstetter); San Luis Potosí : Taman, 20km SW Tamazunchale, [ca. 21.153°N, 98.947°W], 11 Jun 1983 (S. & J. Peck); 20km33/ W Xilitla, [ca. 21.293°N, 99.194°W], 1600m, 12 Jun 1983 (S. & J. Peck); 22.5km W Xilitla, [ca. 21.300°N, 99.086°W], 1460m, 29 Jun 1973 (A. F. Newton);40km W Xilitla, [ca. 21.259°N, 99.194°W], 1700m, 6 Aug 1983 (S. & J. Peck); Sinaloa : 1.7km NNE El Palmito, 23.5783°N, 105.835°W, 2100m, 15 Jan 2007 (P. S. Ward); Tamaulipas : El Cielo, 1.2km SE Alta Cima, 23.05005°N, 99.19226°W, 920m, 24 Aug 2009 (M. G. Branstetter); El Cielo, 1.8km W Alta Cima, 23.06110°N, 99.21564°W, 1340m, 23 Aug 2009 (M. G. Branstetter); El Cielo, 1.8km NW La Gloria, 23.05871°N, 99.26543°W, 2030m, 23 Aug 2009 (M. G. Branstetter); 0.8km N Conrado Castillo, [ca. 23.491°N, 99.308°W], 4 Sep 1970 (W. Elliott); U.S.A.: Arizona : Cochise Co.: Carr Canyon, 11.6km from jct. 92, [ca. 31.428°N, 110.29°W], 2195m, 17 Aug 2002 (Cover & Deyrup); Hd. Carr Canyon, Huachuca Mts., [ca. 31.432°N, 110.284°W], 2440m, 24 Jul 1950 (W. S. Creighton); Chiricahua Mts, [ca. 31.829°N, 109.267°W], 2 Aug 1954 (A. C. Cole); Chiricahua Mts, trail to Barfoot Lookout from rd., [ca. 31.829°N, 109.267°W], 2560m, 26 Jun 1987 (S. P. Cover); Chiricahua Mtns, 8km WSW Portal, 31.883°N, 109.217°W, 1710m, 12 Aug 2002 (P. S. Ward); Chiricahua Mtns., 11.9km 260°W Portal, 31.917°N, 109.267°W, 2440m, 11 Aug 2003 (P. S. Ward); Chiricahua Mtns., W Turkey Creek, 20km 244° WSW Portal, 31.867°N, 109.350°W, 1800m, 13 Aug 2002 (P. S. Ward); Sunnyside Canyon, 5.1km SE jct. FSR48 on FSR227, [ca. 31.437°N, 110.401°W], 1810m, 18 Aug 2002 (Cover & Deyrup); New Mexico : Bernanillo Co.: 15km E Alameda, 35.200°N, 106.483°W, 1950m, 14 Aug 1997 (P. S. Ward).