Stenamma manni Wheeler

Branstetter, Michael G., 2013, Revision of the Middle American clade of the ant genus Stenamma Westwood (Hymenoptera, Formicidae, Myrmicinae), ZooKeys 295, pp. 1-277 : 127-137

Stenamma manni Wheeler Worker: Figures 110-112; Queen: Figure 113 A–D; Male: Figure 113 E–G; Map: Figure 114

Stenamma manni Wheeler, W. M. 1914: 51. Lectotype worker (here designated): MÉXICO, Hidalgo: on the trail between Real del Monte and El Chico, [ca. 20.138°N, 98.673°W], 10000-11000ft [3050-3350m], spring–summer 1913, pine forest, under large stone in damp spot (W. M. Mann) (MZC, Type 2 -3 8672, pin CASENT0126265, specimen furthest from pin) [examined]. Smith, 1962: 35, worker description. Branstetter, 2009: worker images. Branstetter, 2012: phylogeny.

Stenamma mgb28 [variant 4 below] Branstetter 2012: phylogeny.

Worker diagnosis.

Note that this species is highly variable. See comments section below discussing population variants. Integument color variable; medium- to large-sized species (see HL, ML, PrW below); lateral apex of hypostomal bridge projecting ventrally as a subquadrate to broadly rounded lobe, which is usually visible behind base of mandible in profile view (sometimes visible only in lateroventral view); propodeal spines tuberculate to short (PSL 0.09-0.19, PSI 1.0-1.6); basal margin of mandi ble straight; anterior clypeal margin with a single median emargination; face usually completely sculptured, mostly rugoreticulate, with some longitudinal rugulae/carinulae along midline, but sometimes face mostly smooth, with only some longitudinal carinulae; mesosoma usually mostly sculptured with carinae, rugae, rugoreticulae, or punctae, only sometimes with pronotum mostly or completely smooth; eye of moderate size (EL 0.10-0.16, REL 13-21), oval-shaped, with 5-8 ommatidia at greatest diameter; frontal lobes of moderate width (FLD 0.19-0.29, FLI 25-30); first gastral sternite and tergite sometimes punctate. Similar species: Stenamma felixi , Stenamma leptospinum , Stenamma megamanni , Stenamma muralla .

Geographic range.

Mexico to Nicaragua.

Worker description.

(25 measured, lectotype in parentheses) HL 0.81-1.13 (0.91), HW 0.70-1.01 (0.78), FLD 0.19-0.27 (0.23), PCW 0.05-0.08 (0.08), SL 0.68-1.04 (0.75), EL 0.10-0.16 (0.14), ACL 0.62-0.88 (0.67), ML 1.05-1.50 (1.17), PrW 0.45-0.64 (0.53), PSL 0.09-0.19 (0.12), SDL 0.08-0.14 (0.10), PL 0.36-0.54 (0.44), PH 0.21-0.32 (0.25), PW 0.17-0.25 (0.21), PPL 0.21-0.33 (0.25), PPH 0.21-0.33 (0.25), PPW 0.22-0.34 (0.26), MFL 0.77-1.28 (0.85), MTL 0.61-0.97 (0.67), CI 82-91 (86), SI 88-109 (96), REL 13-21 (18), FLI 26-30 (29), PSI 1.0-1.6 (1.2), MFI 75-100 (92), ACI1 61-66 (65), ACI2 84-93 (89).

Medium- to large-sized species; general body color highly variable, ranging from mostly black (type population), to red-brown, to brown, to yellow-brown, with appendages lighter, especially at joints and toward extremities, generally brown or orange-brown to yellow-brown; setae golden brown; mandible with 6-7 teeth (usually 6), consisting of 3 distinct apical teeth, a basal tooth, and 2-3 smaller teeth in between, which are often worn and indistinct; basal margin of mandible straight, without a basal notch or depression; mandible mostly smooth, with scattered piligerous punctae, and a variable number of longitudinal striations, mostly at base and on lateral surface; anterior clypeal margin with a shallow median emargination; median lobe of clypeus often with a pair of faint longitudinal carinulae (type population) that diverge toward anterior margin, but sometimes distinct carinulae replaced or hidden by a variable number of irregular striations, apex of lobe usually with a short transverse carinula, remainder of clypeus mostly smooth; posterior extension of clypeus between frontal lobes of moderate width (PCW 0.05-0.08), with sides subparallel; frontal lobes of moderate width (FLD 0.19-0.27, FLI 26-30), never greatly obscuring torular lobes in full-face view; lateral apex of hypostomal bridge projecting ventrally as a subquadrate to broadly rounded lobe, which is usually visible behind base of mandible in profile view (reduced in type population; sometimes visible only in lateroventral view); head usually roughly oval-shaped (type population), but sometimes slightly elongate, or more often broad, becoming slightly heart-shaped (CI 82-91), posterior margin slightly to distinctly depressed medially; eye of moderate size (EL 0.10-0.16, REL 13-21), oval-shaped, with 5-8 ommatidia at greatest diameter; face usually completely sculptured, mostly rugoreticulate (rarely completely), with some longitudinal rugulae/carinulae along midline extending from frontal lobes to posterior margin, coarseness of sculpture variable (average in type population); face rarely mostly smooth, with only some longitudinal carinulae along midline and on gena; scape usually of moderate length (type population), but sometimes relatively long and slender (SI 88-109), scape when laid back reaching and often surpassing posterior margin of head; scape surface variable, usually with scattered piligerous punctae and some carinulae (type population), but sometimes scape more smooth with carinulae reduced, or scape more robust, with carinulae coarser; flagellum with distinct 4-segmented antennal club; mesosoma sculpture highly variable, usually completely sculptured, without large patches of smooth cuticle, but sometimes pronotum mostly to completely effaced; dorsum of promesonotum carinulate (type population), rugose, or rugoreticulate (often with punctae), almost always with longitudinal orientation (some aberrant specimens with transverse orientation); side of pronotum carinulate (type population), rugulose, or punctate; mesopleuron and side of propodeum punctate to rugulose-punctate, generally with more rugulae on the propodeum; dorsum and declivity of propodeum with transverse carinulae; promesonotum in profile low-domed, and usually slightly asymmetrical, with apex shifted anterior of midpoint, and anterior face steeper than posterior face (some populations roughly symmetrical); metanotal groove usually well-demarcated, width and depth variable (average in type population); propodeal spines tuberculate to short (PSL 0.09-0.19, PSI 1.0-1.6); petiole in profile appearing average (type population) to slightly elongate (PL/HW 0.48-0.62), with peduncle usually thick and robust; petiolar node in profile usually of moderate size (PH/PL 0.51-0.66), with a broadly rounded dorsum that points vertical to slightly posteriad, rarely pointing distinctly posteriad; node sometimes somewhat compressed anteroposteriorly; posterior margin of petiole in profile sometimes distinctly bent downwards, creating a slight concavity below node; postpetiolar node in profile similar in size to petiolar node (type population) or bulging (PPH/PH 0.79-1.05, PW/PPW 0.70-0.86), shape of node subcircular to asymmetrical (slightly asymmetrical in type population); petiole and postpetiole usually mostly punctate, with anterior faces of nodes variably smooth and shiny (type population), sometimes nodes with rugulae, or rarely rugoreticulae; gaster usually smooth and shiny, within scattered piligerous punctae, but sometimes (mostly northern populations in drier habitats) first gastral tergite and sternite lightly to strongly punctate; pilosity highly variable, pilosity on gastral dorsum usually clearly bilayered, with a layer of longer suberect setae, and a layer of shorter decumbent setae, but length and density of each layer variable, lower layer sometimes very dense (almost pubescent), or somewhat sparse and more subdecumbent, causing it to blend in with upper layer, rarely upper layer very long and more dense (type population average), gastral setae never greatly strongly thickened; setae on scapes uniformly suberect to subdecumbent, never with a separate layer of longer suberect setae; setae on legs decumbent to appressed, with longer suberect setae on femoral venters and coxae.

Queen description.

(7 measured) HL 0.92-1.13 (0.98), HW 0.81-1.01 (0.87), FLD 0.23-0.30 (0.25), PCW 0.07-0.10 (0.07), SL 0.81-1.00 (0.82), EL 0.23-0.27 (0.25), ACL 0.72-0.87 (0.75), ML 1.38-1.69 (1.48), PrW 0.82-1.01 (0.82), PSL 0.15-0.29 (0.21), SDL 0.13-0.16 (0.13), PL 0.52-0.68 (0.54), PH 0.28-0.36 (0.29), PW 0.23-0.32 (0.23), PPL 0.27-0.37 (0.28), PPH 0.27-0.39 (0.27), PPW 0.31-0.42 (0.32), MFL 0.98-1.30 (0.99), MTL 0.78-1.01 (0.79), CI 87-92 (89), SI 87-105 (94), REL 26-28 (28), FLI 28-30 (29), PSI 1.1-2.1 (1.6), MFI 78-95 (87), ACI1 60-66 (63), ACI2 83-91 (91).

Same as worker except for standard queen modifications and as follows (comparing queen and worker of type population form only; queen from Rancho Somecla): pronotum with transverse carinulae; mesoscutum and scutellum longitudinally carinulate; propodeum with transverse carinulae that wrap around entire surface; katepisternum mostly smooth; lower layer of setae on gastral dorsum denser; wing venation as in Figure 113D (cell underneath stigma probably aberrant, not present in other Stenamma manni queens).

Male.

See Figure 113 E–G. Note that the male is from the locality Omilteme in Guerrero, Mexico, not the type locality. Males show some variation across the range of the species.

Biology.

This species is one of the largest and most conspicuous species of Stenamma , which is probably why it was the first Middle American species to be described. As defined here, Stenamma manni occurs from 1200-3700 m elevation, but it seems to be most common between 2000-2500 m. It occurs in wet montane forests like cloud forest, and drier, more seasonal habitats, such as oak woodland. Specimens have been collected in leaf litter samples, at bait cards, in Malaise traps, and by general searching. I have found nests in logs, in the leaf litter, under rocks, and in the ground. In Central America, Stenamma manni is commonly found at the edge of cloud forest in logs and in the ground under logs. All Stenamma manni nests tend to be very large, with hundreds to perhaps over a thousand workers (a complete colony census has not been carried out). Nests usually have brood, alates, and a single egg-laying queen. Stenamma manni is one of the most common Stenamma species to be found at bait cards, suggesting that they are active epigeic foragers. All foragers I have observed have been solitary.

Comments.

Stenamma manni should be separable from most similar species by the combination of presence of a lateral hypostomal lobe; presence of tuberculate to short propodeal spines; and large body size. However, it should be evident from the worker description above that Stenamma manni , as circumscribed here, is highly variable, and thus difficult to characterize in a very satisfactory way. Indeed, the manni complex, with all of its different variants, is a taxonomic nightmare. Nearly every population has unique, often distinctive features, and there is no evidence of sympatry among forms. It is also common to have populations that are intermediate in phenotype between variants. Because of this diversity, I have united most forms, even if very distinct, into one polytypic species. I describe many variants below, providing distinguishing characters for each and approximate geographic ranges. This should help in identification of Stenamma manni as I have defined it, and will provide guidelines for future work on the complex.

One Stenamma manni -like variant that I did recognize as a distinct species is Stenamma megamanni . This species occurs from Chiapas, Mexico to Nicaragua and is sympatric with Stenamma manni at many sites. However, it is a variable species and difficult to recognize from Stenamma manni at a large geographic scale. Within the range where both species occur, I often find two forms in sympatry. What I am calling Stenamma manni always has lighter body color (usually dark red-brown), a smaller eye (usually 5-6 ommatidia at greatest diameter), and a larger, more bulging postpetiole. Stenamma megamanni , in contrast, is always black, has a larger eye (8 or more ommatidia at greatest diameter), and usually has the postpetiole similar in size to the petiolar node. In addition, there seems to be some ecological separation between the species. Even though Stenamma megamanni does occur at high elevation and in leaf litter, I commonly find it nesting at lower elevations in riparian areas. I have found nests both in clay banks and under rocks along streams. I have never found Stenamma manni in these environments. It could be that Stenamma megamanni is an ecomorph of Stenamma manni , but I have found both species together in samples of leaf litter. Molecular phylogenetic data so far show that Stenamma megamanni forms a clade nested within the larger Stenamma manni complex. My current hypothesis is that Stenamma manni and possibly Stenamma megamanni , include more than one species, making it difficult to adequately separate each at a wide regional scale.

As a preface to describing each variant of Stenamma manni , I have noticed several phenotypic trends within the complex that correlate with habitat. Populations from drier areas, especially those from central and western Mexico, tend to be lighter in color, have more developed sculpturing, and have denser pilosity, with the lower layer of gastral pilosity often pubescent. In contrast, specimens from wet forest environments are often shinier and less sculptured and have sparser pilosity. They also tend to be darker in color, but not always.

Characteristics of the type population (Figure 110C, D, F) are indicated within the worker and queen descriptions (see parenthetical comments). It is important to note that although these specimens now appear brown in color, the original description says that they were black, with the tarsi, leg joints, and tips of the mandibles and antennae dark red. Overall, these specimens seem to be rather average looking in size, sculpture, and pilosity, compared with many other Stenamma manni populations. Only the eyes seem somewhat large relative to head size. Most other Stenamma manni populations have one or more characters exaggerated in some way. Another important observation is that these specimens were collected from between 3050-3350 m under a rock in pine forest. This is very high for tropical ants in general, and is one of the highest records for Stenamma . The only report of MAC Stenamma from a higher elevation is from 3700 m at the locality Rancho Somecla on Pico de Orizaba in Veracruz, Mexico. These specimens are also Stenamma manni and they are the most similar in morphology to the type population. I should comment that in color and eye size, the type population might appear more similar to Stenamma megamanni than some of the variants below. However, specimens from the type population are much smaller than Stenamma megamanni and the sculpture and pilosity are different. Also, I see intermediate forms within Mexico that seem to connect the type population with the other variants.

Variant 1 (Figure 110A, B, E, G, H, 111 A–C) is the form that co-occurs with Stenamma megamanni in Central America. It has been collected from Chiapas, Mexico, to Nicaragua, and as mentioned above, can be distinguished by its dark red-brown coloration, smaller eye, and bulging postpetiole, which is usually distinctly larger than the petiolar node in profile view. It occurs almost exclusively above 2000 m. Some populations (e.g. Huitepec) have extremely large workers with allometrically enlarged scapes and metafemurs. Compared to the type population, these larger specimens look like giants. Among populations of this variant within Central America, there is considerable sculpture and size variation.

Variant 2 (Figure 111 D–F) is known only from the Atlantic slope of the Sierra Juarez Mountains between Oaxaca and Valle Nacional in Oaxaca, Mexico. All collections were above 1650 m in cloud forest habitat. I tend to think of this variant as a high elevation, wet forest-adapted version of Stenamma manni . Variant 2 is characterized by the following: general body color brown to yellow-brown, mottled; face mostly smooth and shining, with only some faint carinulae extending back from frontal lobes; pronotum almost completely smooth and shiny; scape slender, somewhat elongate (SI 102-106); metafemur relatively elongate (MFI 82-83); propodeal spines well developed, short (PSL 0.13-0.17, PSI 1.4-1.6); anterodorsal margin of propodeum forming a distinct welt, causing the metanotal groove to appear deep; petiolar and postpetiolar nodes somewhat compressed anteroposteriorly; postpetiolar node enlarged, somewhat bulging; gastral pilosity indistinctly bilayered, with lower subdecumbent layer of setae sparse. The gestalt of this variant is most similar to variant 1. Both have the postpetiole more bulging, and the scape and metafemur longer. The shape of the propodeum in profile is also very similar. Confirming the similarity, molecular phylogenetic data show variant 2 being more closely related to a specimen of variant 1 from Chiapas, Mexico, than to specimens further north in Mexico (Branstetter unpublished data). It is important to note that this variant occurs in sympatry with Stenamma leptospinum . Intriguingly, both share the anteroposteriorly compressed petiolar and postpetiolar nodes.

Variant 3 (Figure 111 G–I) occurs at Nevado de Colima in Jalisco, Mexico, with similar forms at other nearby sites. It seems to be a version of Stenamma manni that is adapted to drier habitats. It is characterized by the following: eye somewhat small, with 6 ommatidia at greatest diameter; dorsum of promesonotum rugoreticulate; side of mesosoma strongly punctate; petiole and postpetiole completely sculptured, mostly punctate, with rugoreticulae on dorsal surfaces of nodes; posterior portion of petiole bent downward, creating a small concavity under node; first gastral sternite strongly punctate; first gastral tergite punctate, but punctae broader, more like small dents, and less dense than sternal punctae; pilosity on gastral dorsum distinctly bilayered, with a layer of long suberect setae, and a layer of very dense (almost pubescent) decumbent setae, dorsum of petiolar and postpetiolar nodes with similar pilosity. There is some evidence for sympatry at Nevado de Colima. Variant 3 was collected at 2070 m. A few specimens of a similar Stenamma manni form were collected at 2440 m (same label as variant 3, but different elevation). These specimens do not have distinct rugoreticulae and the dense pubescence, but seem otherwise similar. Because of the difference in elevation between the specimens, the paucity of material, and the existence of intermediate specimens at other sites, I do not consider this potential sympatry as sufficient evidence for calling variant 3 a new species. However, Nevado de Colima would be a good site to visit to further study species boundaries in this complex.

Variant 4 (Figure 111 J–L) is known only from Pinal de Amoles, Querétaro, Mexico and is similar to variant 3. It could be a mesic forest version of variant 3, as it occurs in wetter oak-pine forest on the Atlantic slope of the Sierra Gorda. It is characterized by the following: general body color dark red-brown to brown; head noticeably broad and thick; eye small, with 5-6 ommatidia at greatest diameter; dorsum of promesonotum rugoreticulate-punctate, but rugoreticulae poorly developed on middle of dorsum, strongest on humeri; side of mesosoma strongly punctate; propodeal spines well-developed, somewhat robust, and distinctively shaped, being broad at the base and then curving outward, appearing somewhat hook-like; petiole noticeably elongate and bent downward at posterior margin; petiolar node in profile broad and subconical, reaching a defined apex; petiole and postpetiole almost completely punctate; first gastral sternite and tergite lightly punctate; pilosity on gastral dorsum somewhat dense, and clearly bilayered, with a layer of longer suberect to subdecumbent setae, and a layer of decumbent setae, both layers similar in density. Specimens with intermediate morphology between variant 3 and 4 occur in Oaxaca on the western, drier side of the state. Among these specimens there is variation in the development of punctae and pubescence on the gaster, which seems to be clearly linked with elevation. One population from 2350 m (15.5km NE Oaxaca) has no punctae on the first gastral sternite and reduced gastral pilosity. In all other respects it is identical to populations from lower elevations within the state.

Variant 5 (Figure 112 A–C) is known from a single collection in Jalisco, Mexico. It is characterized by two main features: pronotal dorsum and face densely rugoreticulate; pilosity on most of dorsal surface of mesosoma, waist, and gaster very dense, long, and flexuous. This variant is probably most similar to variant 3.

Variant 6 (Figure 112 D–F) is known from several sites in Guatemala (Cerro Carmona, Salama, Guatemala City), and Honduras (PN Celaque), ranging from 1500-2000 m approximately. It has the following distinctive features: head oval-shaped, somewhat narrow; pronotal dorsum rugoreticulate; side of pronotum punctate; body in profile appearing somewhat elongate and gracile; petiole long and gracile, node asymmetrical, with a long sloping anterior face and a short posterior face, dorsum usually pointing distinctly posteriad; postpetiole somewhat smaller than petiolar node, not bulging. I suspect this variant could be a distinct species. It occurs in sympatry with variant 1 of Stenamma manni at Cerro Carmona and has been collected near Stenamma megamanni . However, it is very rare (only ten specimens known), and no nests or queens have been found. Molecular phylogenetic data infer it to be closely related to variant 1 and Stenamma megamanni . More sampling is needed to test its status.

Variant 7 (Figure 112 G–I) is known only from RN Datanlí El Diablo in Nicaragua. It is similar to variant 6, except as follows: pronotal sculpture mostly longitudinally carinulate, with a patch of smooth cuticle in middle of dorsum; pilosity more dense; setae on legs noticeably thickened and subdecumbent (best observed in dorsal view).

Variant 8 (Figure 112 J–L) is known from two specimens collected at almost 3000 m at the Sendero Ecologico La Maceta locality in Guatemala. It has the following distinguishing features: eye large, with 9 ommatidia at greatest diameter; pronotum mostly smooth, with some transverse carinulae; postpetiole similar in size to petiolar node; gastral pilosity sparse, and weekly bilayered. This variant is most similar to the type population, in terms of sculpture, body and eye size. Unfortunately, neither specimens from the type population, nor this variant have been included in phylogenetic analyses, so it is unclear how they are related. A few faded specimens from Tajumulco, Guatemala look similar.

Material examined.

EL SALVADOR: Santa Ana: Hacienda Montecristo, 23km NE Metapan, [ca. 14.406°N, 89.372°W], 2300m, 8 May 1871 (S. B. Peck); GUATEMALA:Baja Verapaz: 4km SSE Purulhá, 15.20522°N, 90.22198°W, 2100m, 20 Sep 2008 (M. G. Branstetter); 16.5km N Salama, [ca. 15.2350°N, 90.2833°W], 1660m, 23 May 1991 (R. S. Anderson); Chimaltenango: Finca Chincharas, Rincón Suizo, 14.79775°N, 90.98380°W, 2490m, 17 Sep 2008 (R. S. Anderson); 3.8km NNE Tecpan, 14.79694°N, 90.98171°W, 2500m, 17 Sep 2008 (M. G. Branstetter);El Progreso: Cerro Pinalón, 15.08389°N, 89.94456°W, 2560m, 30 Apr 2009 (LLAMA); 20km N Estancia de la Virgen, [ca. 15.1141°N, 89.8833°W], 1850m, 8 Jun 1991 (R. S. Anderson); Guatemala: Guatemala City, 14.55871°N, 90.46275°W, 1800m, 5 Jul 2007 (R. S. Anderson); Huehuetenango: La Capellania, 15.39867°N, 91.40363°W, 3050m, 14 Sep 2008 (R. S. Anderson); Sendero Ecologico La Maceta, 15.48915°N, 91.55492°W, 2950m, 14 Sep 2008 (R. S. Anderson); Jalapa: 5km ENE Mataquescuintla, 14.53355°N, 90.14323°W, 2550m, 5 Jul 2007 (R. S. Anderson); Miramundo, Pino Dulce, 14.53388°N, 90.15236°W, 2300m, 18 Sep 2008 (R. S. Anderson); Quetzaltenango: Aldea Las Nubes, Volcán Chicabal, 14.80086°N, 91.66803°W, 2280m, 12 Sep 2008 (M. G. Branstetter); nr Roble Grande, 14.92719°N, 91.68271°W, 2750m, 11 Sep 2008 (R. S. Anderson); 11.5km ESE San Marcos, 14.92719°N, 91.68271°W, 2800m, 11 Sep 2008 (M. G. Branstetter); 1.7km SW Santa María, 14.71686°N, 91.53481°W, 1515m, 13 Sep 2008 (L. Sáenz); Volcán Chicabal, 14.78720°N, 91.65839°W, 2700m, 12 Sep 2008 (M. G. Branstetter); 12km SE Zunil, Fuentes Georginas, 14.7488°N, 91.4800°W, 2460m, 27 May 1991 (R. S. Anderson); Quiché: 2.9km SSE Chichicastenango, 14.91861°N, 91.10449°W, 2000m, 17 Sep 2008 (L. Sáenz); San Marcos: 6.3km WSW San Marcos, 14.94686°N, 91.83771°W, 2620m, 11 Sep 2008 (M. G. Branstetter); 9.8km WSW San Marcos, 14.94427°N, 91.87990°W, 1600m, 11 Sep 2008 (M. G. Branstetter); Tajumulco, [ca. 15.044°N, 91.901°W], 2960m, Oct 1934 (F. X. Williams); Sacatepéquez: 5km SE Antigua, 14.54124°N, 90.71026°W, 1740m, 12 Jun 2009 (LLAMA); Cerro Carmona, Finca El Pilar, 14.54115°N, 90.70483°W, 1980m, 9 Sep 2008 (R. S. Anderson); Suchitepéquez: 5km SE Antigua, 14.53581°N, 90.69430°W, 2150m, 10 Jun 2009 (LLAMA); Zacapa: 14km NNE Teculután, 15.11440°N, 89.68047°W, 2270m, 6 Jul 2007 (6 Jul 2007); HONDURAS: Comayagua: 10km E Comayagua, 14.45973°N, 87.54609°W, 2000m, 15 May 2010 (LLAMA); 12km ENE Comayagua, 14.48045°N, 87.53258°W, 2140m, 15 May 2010 (LLAMA); Francisco Morazán: PN La Tigra, 2.6km SW San Juancito, 14.21778°N, 87.09101°W, 1870m, 25 Sep 2008 (M. G. Branstetter); Lempira: PN Celaque, 7.3km SW Graçias, 14.56370°N, 88.64923°W, 1530m, 30 Sep 2008 (M. G. Branstetter); 8.3km SW Graçias, 14.56132°N, 88.65768°W, 1860m, 30 Sep 2008 (M. G. Branstetter); Ocotepeque: 13km E Nueva Ocotepeque, 14.45697°N, 89.06849°W, 2200m, 25 May 2010 (LLAMA); Olancho: 11km N Catacamas, 14.94838°N, 85.91439°W, 2000m, 12 May 2010 (LLAMA); 12km N Catacamas, 14.95787°N, 85.91673°W, 2320m, 9 May 2010 (LLAMA); Santa Barbara: 15km SE Santa Barbara, [ca. 14.906°N, 88.100°W], 24 Aug 1991 (S. & J. Peck); MÉXICO: Chiapas: Cerro Huitepec (Pico), ca. 5km W San Cristobal, [ca. 16.7500°N, 92.6802°W, 2750m, 18 Sep 1991 (R. S. Anderson); 5km NNW Coapilla, 17.18355°N, 93.15222°W, 1915m, 26 May 2008 (LLAMA); 4km E Custepec, 15.71294°N, 92.92761°W, 2180m, 23 May 2008 (M. G. Branstetter); 2km SE Custepec, 15.72188°N, 92.93677°W, 1900m, 19 May 2008 (R. S. Anderson); Huitepec, S. Cristóbal, 16.75181°N, 92.68270°W, 2480m, 29 May 2008 (LLAMA); 7.4km SSW Motozintla de Mendoza, 15.3667°N, 92.2333°W, 2000m, 21 Sep 1992 (R. S. Anderson); 15km San Cristóbal, 16.74707°N, 92.49011°W, 2500m, 29 May 2008 (LLAMA); Colima: 19km NNE Comala, 19.482°N, 103.683°W, 1650m, 25 Dec 1987 (P. S. Ward);Guerrero: vic. Omilteme, [ca. 17.555°N, 99.687°W], 2400m, 29-30 Jul 1965 (Cornell Univ. Mexico Field Party);Jalisco: 10km S Autlán, 19.6833°N, 104.3830°W, 1600m, 20 Dec 1987 (P. S. Ward); E slope Nevado de Colima, [ca. 19.552°N, 103.559°W], 2400m, 21 Sep 1973 (A. F. Newton); Hidalgo: between Real del Monte and El Chico, [ca. 20.138°N, 98.673°W], 3050m, Mar–Aug 1913 (W. M. Mann); Oaxaca: 9.4km SE Asunción Nochixtlan, 17.37632°N, 97.19976°W, 2240m, 10 Aug 2009 (M. G. Branstetter); 33km S Ixtlan de Juarez, [ca. 17.204°N, 96.589°W], 2400m, 10 Aug 1973 (A. F. Newton); 10.6km N Jct 190/135 (on 135), [ca. 17.282°N, 96.929°W], 1920m, 21 Jul 1987 (R. S. Anderson); La Herradura, [ca. 17.221°N, 97.042°W], 1959 (C. Gans); 14km NE Oaxaca, km 10 Mex. 175, 17.145°N, 96.622°W, 1890m, 20 Aug 1973 (A. F. Newton); 15.5km NE Oaxaca, 17.23695°N, 96.56533°W, 2350m, 11 Aug 2009 (M. G. Branstetter); 14.8km SSW Valle Nacional 17.64483°N, 96.33637°W, 1370m, 13 Aug 2009 (M. G. Branstetter); 22.4km SW Valle Nacional, 17.59112°N, 96.39113°W, 1990m, 13 Aug 2009 (M. G. Branstetter); 32km SW Valle Nacional, Km85, [ca. 17.5046°N, 96.3897°W], 1650m, 26 Jul 1992 (R. S. Anderson); 40km SW Valle Nacional, Km93, [ca. 17.5278°N, 96.5834°W], 1900m, 26 Jul 1992 (R. S. Anderson); 5.1km S Suchixtepec, [ca. 16.0833°N, 96.4667°W], 2150m, 25 Jul 1992 (R. S. Anderson); 30.6km S Suchixtepec, [ca. 15.976°N, 96.498°W], 12 Jul 1987 (R. S. Anderson); 47.5km SW Valle Nacional, Km100.5, [ca. 17.350°N, 96.5333°W], 2125m, 26 Jul 1992 (R. S. Anderson); Puebla: H’way 190, E Río Frío, [ca. 19.310°N, 98.638°W], 2900m, 7 Aug 1965 (Cornell Univ. Mexico Field Party); Querétaro: 1.9km NE Pinal de Amoles, 21.14974°N, 99.61576°W, 2250m, 18 Aug 2009 (M. G. Branstetter); Veracruz: Rancho Somecla, Pico Orizaba, [ca. 18.987°N, 97.227°W], 3700m, 24 Aug 1953 (E. O. Wilson); NICARAGUA: Jinotega: RN Cerro Kilambé, 13.56941°N, 85.69479°W, 1540m, 24 May 2011; RN Datanlí El Diablo, 13.10449°N, 85.86774°W, 1400m, 18 May 2011 (LLAMA); Madriz: Apante, 9km S Somoto, 13.40481°N, 86.57944°W, 1550m, 22 Apr 2011 (M. G. Branstetter); 1.9km SE Las Sabanas, 13.33012°N, 86.60723°W, 1650m, 23 Apr 2011 (M. G. Branstetter); 16km S Somoto, 13.32726°N, 86.61001°W, 1730m, 23 Apr 2011 (R. S. Anderson); Nueva Segovia: Cerro Mogotón, 10km NNE Mozonte, 13.75599°N, 86.42062°W, 24 Apr 2011 (M. G. Branstetter); 9km NW Jalapa, 13.98271°N, 86.18933°W, 1700m, 28 May 2011; 14km NNE Ocotal, 13.75409°N, 86.42094°W, 1900m, 24 Apr 11 (R. S. Anderson).