I have only three records of C. distans from Costa Rica: two older museum collections from the Central Valley, and one collection from La Selva Biological Station in the Atlantic lowlands. It is clear from material elsewhere in the range of the species that distans prefers seasonally dry, open or brushy habitats.
In general, distans has very large colonies that are low density. The one La Selva collection I observed was from a recent treefall. A large dead trunk had fallen, bringing a large Bauhinia vine tangle down with it. Crematogaster distans dominated the treefall, with columns of abundant workers spread throughout and streaming back to the canopy along lianas. I did not find any nest center or brood, but I did find two alate queens along with a worker aggregation in a hollow stick. In the Santa Marta region of Colombia I observed a distans nest in a dead stick, and on another occasion I found workers tending Coccoidea on Cnidoscolus (Euphorbiaceae). In a second growth forest in Venezuela I observed columns of workers descending a tree trunk and spreading out onto an Inga sapling, where large numbers of workers were tending scale insects.
Nothing is known of colony founding, but queens exhibit a morphology often associated with social parasitism. Kusnezov (1949) described descolei from a single alate queen and suggested that it was a workerless social parasite because of the small size and shiny integument. Kempf (1968) refuted this by reporting worker-associated colonies. The discovery of a colony of distans at La Selva with queens identical to descolei confirmed the synonymy of descolei under distans. It is possible that Crematogaster distans is a temporary social parasite, requiring colonies of host species to establish new colonies of its own, but it is not workerless.
The species complex occurs from southern USA (Texas) to Argentina (see Comments).
Taxonomic history
Kempf, 1968b PDF: 392 (q.).Combination in Crematogaster (Orthocrema): Emery, 1922c PDF: 134.Combination in Crematogaster (Neocrema): Santschi, 1918d PDF: 182; Kempf, 1972b PDF: 83.Combination in Crematogaster (Orthocrema): Blaimer, 2012c PDF: 55.Senior synonym of Crematogaster distans cordinoda: Kempf, 1968b PDF: 390.Senior synonym of Crematogaster distans paraensis: Kempf, 1968b PDF: 390.Senior synonym of Crematogaster distans parviceps: Kempf, 1968b PDF: 390.Senior synonym of Crematogaster distans pevsnerae: Kempf, 1968b PDF: 390.Senior synonym of Crematogaster distans vanda: Kempf, 1968b PDF: 390.Senior synonym of Crematogaster descolei: Longino, 2003a PDF: 59.Plate 3, 4
Crematogaster distans Mayr , 1870a:402. Lectotype worker: Colombia ( Lindig ) [ NMW ] (examined). Emery, 1922:134: combination in C. (Orthocrema) . Santschi, 1918:182; Kempf, 1972:83: combination in C. (Neocrema) . Kempf, 1968:392: description of queen.
Crematogaster distans r. paraensis Forel , 1904a:37. Syntype workers: Brazil , Para ( Göldi ) [ MHNG ] (examined). Synonymy by Kempf, 1968:390.
Crematogaster distans subsp. parviceps Forel , 1908b:369. Syntype workers: Brazil , Sao Paulo ( von Ihering ) [ MHNG ] (examined). Emery, 1922:135: combination in C. (Orthocrema) . Synonymy by Kempf, 1968:390.
Crematogaster distans r. pevsnerae Forel , 1912:218. Syntype workers: Venezuela , Zig Zag ( Forel ) [ MHNG ] (examined). Emery, 1922:135: combination in C. (Orthocrema) . Synonymy by Kempf, 1968:390 [Kempf suggested probable synonymy; Bolton (1995) interpreted as formal synonymy].
Crematogaster distans var. cordinoda Forel , 1914c:12. Type worker a "variant of the type series of distans " (Kempf 1968:393). Emery, 1922:135: combination in C. (Orthocrema) . Synonymy by Kempf, 1968:390.
Crematogaster distans subsp. vanda Borgmeier , 1929:209. Holotype worker: Brazil , Rio de Janeiro , Niterói ( Bartholdy ) . Synonymy by Kempf, 1968:390.
Crematogaster (Neocrema) descolei Kusnezov , 1949:587, figs. 1-4. Holotype queen: Argentina , Prov. Jujuy , near Ledesma, road to Valle Grande, subtropical mesophyl forest , 2-12-1928 ( KusnezovNo. 3310 ) . NEW SYNONYMY
Range
The species complex occurs from southern USA (Texas) to Argentina (see Comments).
Description of worker (Costa Rica)
Color dark red brown to black.
Mandible largely smooth and shiny, with very faint striae; clypeus emarginate anteriorly, convex, smooth and shiny; face smooth and shiny; scape with moderately abundant, short, fully appressed pilosity and no erect setae; terminal 2-3 segments of antenna sequentially longer and more densely pubescent, especially terminal two, forming 2-segmented club; face and ventral surface of head with sparse short appressed pubescence and no erect setae; no erect setae projecting from sides of head in full face view.
In lateral view, promesonotum compressed, pronotum short but strongly convex, mesonotum projecting above pronotum and propodeum, dropping steeply to propodeal suture; propodeum with short, weakly differentiated dorsal face; propodeal suture deep, V- shaped, but suture partially obscured in lateral view by small lateral carinulae that bridge suture; propodeal spines short, conical, upturned; side and most of dorsum of mesosoma with faint punctation or weak microareolate sculpture, becoming smooth and shiny on posterior face of propodeum; mesonotum with a pair of short, stiff, erect setae, otherwise erect setae lacking on mesosoma; legs with sparse, fully appressed pubescence.
Petiole in side view subtrapezoideal, with faint microareolate surface sculpture; anteroventral tooth a shallow, obtuse gibbosity, not produced or angulate; dorsal face subrectangular, slightly longer than wide, smooth and shiny, anterolateral portions somewhat produced as lobes, anteromedian region depressed, sides approximately parallel anteriorly, converging posteriorly; postpetiole with small acute ventral tooth, postpetiole in dorsal view bilobed, wider than long, with longitudinal median sulcus, posterior margin emarginate; fourth abdominal tergite with faint microareolate sculpture; pair of stiff setae on posterodorsal petiole, two pairs on postpetiole, and about 20 stiff setae on fourth abdominal tergite.
Measurements
HL 0.699, 0.627, 0.698; HW 0.712, 0.649, 0.696; HC 0.671, 0.610, 0.653; SL 0.563, 0.479, 0.564; EL 0.143, 0.157, 0.167; A11L 0.284; A11W 0.135; A10L 0.135; A10W 0.107; A09L 0.069; A09W 0.086; A08L 0.046; A08W 0.069; WL 0.784, 0.686, 0.773; SPL 0.147, 0.140, 0.127; PTH 0.212, 0.187, 0.195; PTL 0.299, 0.258, 0.288; PTW 0.297, 0.295, 0.289; PPL 0.194, 0.180, 0.203; PPW 0.321, 0.286, 0.320; CI 102, 104, 100; OI 20, 25, 24; SI 81, 76, 81; PTHI 71, 72, 68; PTWI 99, 114, 100; PPI 165, 159, 158; SPI 19, 20, 16; ACI 0.56.
Queen (Costa Rica)
In lateral profile dorsal face of propodeum sloping obliquely from postscutellum, such that most of propodeum is posterior to scutellum (in contrast to normal queens, in which dorsal face of propodeum drops steeply from postscutellum and much of propodeum appears ventral to scutellum and postscutellum, Fig. 1); entire body, including scape, mandible, face, mesosoma, petiole, postpetiole, and fourth abdominal tergite extremely smooth and shiny, glass-like; propodeal spines completely absent; petiole and postpetiole robust, generally similar to worker in shape but completely lacking anteroventral petiolar tooth of any kind; remarkably devoid of erect setae or pubescence, funiculus and terminal segments of tarsi with typical subdecumbent pubescence, but no trace of pubescence or erect setae anywhere else on body; size characters as in Figures 4 and 5.
Biology
I have only three records of C. distans from Costa Rica, two older museum collections from the Central Valley, and one collection from La Selva Biological Station in the Atlantic lowlands. It is clear from material elsewhere in the range of the species that distans prefers seasonally dry, open or brushy habitats.
In general, distans has very large colonies that are low density. The one La Selva collection I observed was from a recent treefall. A large dead trunk had fallen, bringing a large Bauhinia vine tangle down with it. Crematogaster distans dominated the treefall, with columns of abundant workers spread throughout and streaming back to the canopy along lianas. I did not find any nest center or brood, but I did find two alate queens along with a worker aggregation in a hollow stick. In the Santa Marta region of Colombia I observed a distans nest in a dead stick, and on another occasion I found workers tending Coccoidea on Cnidoscolus (Euphorbiaceae). In a second growth forest in Venezuela I observed columns of workers descending a tree trunk and spreading out onto an Inga sapling, where large numbers of workers were tending scale insects.
Nothing is known of colony founding, but queens exhibit a morphology often associated with social parasitism (see Natural History Overview). Kusnezov (1949) described descolei from a single alate queen and suggested that it was a workerless social parasite because of the small size and shiny integument. Kempf (1968) refuted this by reporting worker-associated colonies. The discovery of a colony of distans at La Selva with queens identical to descolei confirms the synonymy of descolei under distans . It is possible that Crematogaster distans is a temporary social parasite, requiring colonies of host species to establish new colonies of its own, but it is not workerless.
Comments
This species can at first be confused with crinosa and relatives because of the bare face, general lack of erect setae, and short upturned propodeal spines. The strongly bilobed postpetiole and the robust petiole with subrectangular dorsal face set it apart. There is a subtle mesosomal shape difference that is difficult to define, but involves a relatively compact and elevated promesonotum. These mesosomal and petiolar shape characters, along with the small shiny queens, are more like acuta and relatives than crinosa . Other than the superficial similarity to crinosa relatives, there are no other Costa Rican species with which it can be confused.
The distans complex occurs from southern Texas in the United States south to Argentina. Members of the complex share a characteristic mesosoma, petiole, and postpetiole shape, but they vary in details of surface sculpture and pilosity. The queens are small and with highly polished and smooth integument, but varying in details of pilosity and petiole shape. Kempf (1968) made a start in understanding this group, and a few additional notes are added here, but more material, especially queens associated with workers, is needed to make further progress. In the following paragraphs I outline geographic variation as I currently understand it, based on the few collections I have been able to examine and the published work of Kempf.
A single specimen from Hidalgo Co., Texas, USA, has a punctate face, a pair of setae on face, and no erect setae on mesosoma. A series from San Luis Potosí , Mexico, has a punctate face, a pair of setae on face, a pair on mesonotum, and a seta on each propodeal spine. The two worker syntypes of corvina Mayr 1870, from "Mexico," have punctate faces. One worker has a row of four setae on the pronotum, five on the mesonotum, two on the base of one propodeal spine, and one on base of the other. The other worker has one on the pronotum, seven on the mesonotum, three on one side and one on the other side of the dorsal face of the propodeum. These punctate forms at the northern limit of the range I identify as corvina , but it is clear there is considerable variation in pilosity, and they may grade into forms further south. Punctate faces reappear on two forms from South America: scelerata Santschi from Argentina and scelerata taperensis Borgmeier from Brazil, Pernambuco State. These are nearly identical to corvina but with a more developed anteroventral petiolar tooth. Thus punctate faces seem to occur at the margins of the range of the lineage. Like other amphitropical distributions, it is unknown whether this pattern is due to shared ancestry and some form of centripetal speciation, in which new forms replace older forms in the center of the range, or convergence due to similar selection pressures in marginal habitats.
Two series from Morelos, Mexico have most of face smooth but with distinct patches of punctation posteroventral to eyes and between eyes and antennal insertions. Thus they are intermediate in this character between corvina and other distans-like forms to the south. They have a pair of setae on the face, 2-4 setae on mesonotum, and a seta on each propodeal spine. The types of parviceps , from Sao Paulo state, Brazil, have similar facial sculpture, a combination of smooth and punctate. All remaining forms have the face completely shiny.
The series from Costa Rica and two series from Panama have one pair of setae on the mesonotum and none on the face, pronotum, or propodeum. The dorsal face of the petiole is relatively elongate, proportionally longer than the material from further north. The Costa Rican series from La Selva Biological Station has associated alate queens. These queens are identical in every respect to descolei Kusnezov . Crematogaster descolei was described from a single alate queen from Argentina. The queens are strikingly elegant, like glass figures, with no trace of setae or pubescence on most of body. The dorsal face of the petiole is proportionately quite long.
Material from Colombia and Venezuela, from which I have examined about ten worker collections from a variety of localities, shows a great deal of variation. Some collections from Venezuela appear identical to Costa Rican and Panamanian material. Others from the Santa Marta area of Colombia are also very similar, but have relatively shorter petioles. Finally, some collections from the Santa Marta area and departments of Antioquia, Huila, and Santander are very pilose, with multiple pairs of setae on both the pronotum and mesonotum, and multiple setae on the dorsal face and spine of the propodeum. Two of these collections have queens associated with them. The queens are quite similar in shape to the Costa Rican and descolei queens, but have abundant erect setae on the face and dorsal surfaces, and the petiole is shorter.
Kempf (1968) described his broad view of distans based on many collections, mostly from southern South America. He described worker characteristics and variation that match the less pilose forms from Venezuela, Colombia, Panama, and Costa Rica. In his figure of the dorsal face of a worker petiole, it appears relatively short, like the short-petiole material from Colombia, rather than the long-petiole material from Venezuela and Costa Rica. He described the queen of distans , based on collections from Rio de Janeiro and Sao Paulo states. He described the queen as being nearly identical to descolei , with the same general shape and degree of hairlessness, but with a relatively shorter petiole. He stated that descolei was a probable synonym of distans .
It is possible there are multiple sympatric forms in the Neotropics, with queens being much more differentiable than workers. Putting the above observations together suggests the possibility there are at least three forms: (1) descolei-like , the queens of which are hairless and long petiolate, (2) a widespread form with hairless queens and short petioles, and (3) a setose form, with abundant setae on queens and workers. But much more queen-associated material and some thorough morphometrics will be needed to confirm or modify these patterns. I have followed the "synonym until proven distinct" approach by formally synonymizing descolei under distans . But future work on the complex will require a reevaluation of all the synonymized forms.
