Genus Dolichoderus (Dolichoderinae)

• Dolichoderus Lund, 1831a: 130. Type species: Formica attelaboides Fabricius, by monotypy.
• Hypoclinea Mayr, G. 1855: 377. Type species: Formica quadripunctata Linnaeus, by subsequent designation of Wheeler, W.M. 1911g: 165. Hypoclinea junior synonym of Dolichoderus: Forel, 1878c: 386. Revived from synonymy as subgenus of Dolichoderus: Emery, 1894d: 234; maintained as subgenus Emery, 1913a: 10. Raised to genus: Kempf, 1972b: 118. Note that the correct citation is 'Emery, 1913a' not 'Emery, 1912g' as in Bolton, 1995b: 31. Synonymy: Brown, W.L. 1973b: 181 (Hypoclinea provisional junior synonym of Dolichoderus) ; Shattuck, 1992c: 66 (Hypoclinea junior synonym of Dolichoderus) .
• Monacis Roger, 1862a: 233 . Type species: Formica bispinosa Olivier, by subsequent designation of Wheeler, W.M. 1911g: 167. Junior synonym of Hypoclinea: Mayr, 1862: 704; of Dolichoderus: Forel, 1878c: 386. Revived from synonymy as subgenus of Dolichoderus: Emery, 1894d: 228; maintained as subgenus: Emery, 1913a: 9; Wheeler, W.M. 1922i: 688. Revived status as genus: Brown, W.L. 1950g: 249; maintained as genus: Kempf, 1959c: 227. Note that the correct citation is 'Emery, 1913a' not 'Emery, 1912g' as in Bolton, 1995b: 35. Synonymy: Brown, W.L. 1973b: 182 (Monacis provisional junior synonym of Dolichoderus) ; Shattuck, 1992c: 66 (Monacis junior synonym of Dolichoderus) .
• Diabolus Karavaiev, 1926d: 424 (as subgenus of Dolichoderus) . Type species: Dolichoderus (Diabolus) bifurcatus Karavaiev, by original designation. Junior homonym of Diabolus Gray, 1841 (Mammalia). The replacement name is Karawajewella Donisthorpe, 1944. The type species, Dolichoderus (Diabolus) bifurcatus Karavaiev, is a junior synonym of D. cuspidatus Smith. Synonymy: Shattuck, 1992c: 66.
• Diceratoclinea Wheeler, W.M. 1935c: 69 (as subgenus of Dolichoderus) . Type species: Dolichoderus scabridus Roger, by original designation. Diceratoclinea raised to genus: Brown, W.L. 1950g: 249. Synonymy: Brown, W.L. 1973b: 180 (Diceratoclinea provisional junior synonym of Dolichoderus) ; Taylor, R.W. & Brown, 1985: 93 (Diceratoclinea junior synonym of Dolichoderus) ; Shattuck, 1992c: 66 (Diceratoclinea junior synonym of Dolichoderus) .
• Monoceratoclinea Wheeler, W.M. 1935c: 68 (as subgenus of Dolichoderus) . Type species: Dolichoderus (Hypoclinea) monoceros Emery, by original designation. Monoceratoclinea raised to genus: Brown, 1950g: 249. Synonymy: Shattuck, 1992c: 66.
• Acanthoclinea Wheeler, W.M. 1935c: 69 (as subgenus of Dolichoderus) . Type species: Dolichoderus doriae Emery, by original designation. Acanthoclinea raised to genus: Brown, W.L. 1950g: 249. Synonymy: Brown, W.L. 1973b: 178 (Acanthoclinea provisional junior synonym of Dolichoderus) ; Taylor, R.W. & Brown, 1985: 93 (Acanthoclinea junior synonym of Dolichoderus) ; Shattuck, 1992c: 66 (Acanthoclinea junior synonym of Dolichoderus) .
• Karawajewella Donisthorpe, 1944b: 59. Replacement name for junior homonym Diabolus Karavaiev, 1926. Synonymy: Brown, W.L. 1973b: 181 (Karawajewella provisional junior synonym of Dolichoderus) ; Shattuck, 1992c: 66 (Karawajewella junior synonym of Dolichoderus) .

• OVERVIEW.   In North America, this genus is found only east of the Rocky Mountains. The taxonomy of this genus is relatively well known. Colonies are small to large and nests are constructed in the soil, in curled leaves and hollow stems of plants, and in cartons attached to plants. Workers attend honeydew excreting insects and feed on small arthropods. Reproductives may over winter in the nest. Some forms emit a fluid which has a peculiar smoky or pungent odor (Smith, D.R. 1979: 1415). Two species, D. plagiatus and D. taschenbergi, are believed to be polygynous. Little is known of nest founding in this genus, but there is some evidence (not yet confirmed) of temporary social parasitism of D. taschenbergi on host D. plagiatus (Ortius, 1998). The Nearctic species are all members of the quadripunctatus complex (MacKay, W.P. 1993b: 16).
• ZOOGEOGRAPHICAL REGIONS.  Neotropical, Nearctic, Palearctic, Oriental, Indo-Australian, Australasian.
• NOMENCLATURE.   Prior to 1992, the Nearctic species of Dolichoderus were placed in Hypoclinea.


      
Photo of Dolichoderus mariae Forel from MA, Plymouth Co.. Courtesy of Ant Web, U. of Calif. Davis


• RECOGNITION.   Integument thick, hard, and strongly sculptured. Underside of the head near the base of the mandible with a weak to well-developed flange that is sometimes tooth-like. Metanotal impression distinct. Propodeum with declivious face distinctly concave.
• SIMILAR GENERA.   This genus is unlikely to be confused with any other Nearctic dolichoderines. It is clearly distinguished by the combination of thick, sculptured integument, distinct metanotal impression, and concave declivitous face of propodeum.
• SPECIES IDENTIFICATION.   Identification to species is fairly straght forward. The key to the New World species of Dolichoderus in MacKay, 1993b is the most recent key. Start with couplet 41 on page 22. Also very useful are the descriptions of each species in MacKay, 1993b and especially the discussion on pages 100–101 comparing the four Nearctic species. The figures in Johnson, C. 1989a will be useful.
  

  Dolichoderus Species Recognition

  		mariae	plagiatus	pustulatus	taschenbergi
  Punctures	Head	fine	coarse	fine	fine
  	Pronotum	fine	coarse	coarse	fine
  	Mesonotum & Propodeum	fine	coarse	coarse	coarse
  Erect Hairs	Scape	absent	present	absent	absent
  	Body	absent	abundant	sparse	sparse
  	Mesopleuron	subopaque	subopaque	shining	subopaque
  	Propodeum shape	longer	longer	longer	subquadrate
  	Color	usually bicolored	variable	usually concolorous	concolorous

• TRIBE.  Dolichoderini.
• Literature Overview.   MacKay's revision thoroughly covers the systematics and distribution of this genus and provides a brief introduction to the natural history.
• REVISIONS.   Mayr, G. 1870b: 953–960 .— Mayr, G. 1886d: 434–437 .— Wheeler, W.M. 1905g: 305–319 (Nearctic - out of date) .— Creighton, 1950a: 331–337, pl. 43, figs. 1–4 (Nearctic, disagrees with many points in Wheeler, W.M. 1905g, as Hypoclinea) .— MacKay, W.P. 1993b: 1–148 (New World species) .
• TAXONOMY.   Clark, J. 1930b: 252–268 (key to Australian species, out of date) .— Smith, M.R. 1943e: 310–311 (diagnosis of m., as D. (Hypoclinea)) .— Smith, M.R. 1947f: 593–594, pl. 16, fig. 60 (diagnosis of w., figure (lateral view) of w., as D. (Hypoclinea)) .— Brown, W.L. 1950g: 249 .— Wheeler, G.C. & Wheeler, J. 1951: 172 (larva) .— Kempf, 1959c: 227 (key to Neotropical species formerly in Monacis) .— Wheeler, G.C. & Wheeler, J. 1966: 726–730 (larva) .— Kempf, 1969: 289–292 (key to Neotropical species of Dolichoderus sensu stricto) .— Kempf, 1972g: 251–254 (emended key to Neotropical species formerly in Monacis) .— Wheeler, G.C. & Wheeler, J. 1974b: 396 (correction to generic diagnosis in Wheeler, G.C. & Wheeler, J. 1951) .— Lattke, 1987b: 259–265 (Neotropical, key to bispinosa group) .— Johnson, C. 1989a: 1–9 (North American species) .— Shattuck, 1992c: 66–77, figs. 47–63 (description of w.q.m., diagnosis of l., distribution map, figures (full face and lateral views) w.q.m.) .— Bolton, 1994: figs. 28, 29 (SEMs (full face and lateral view) of w.) .— Xu, Z. 1995a: 33–35 (China (in Chinese)) .—Shattuck, 1999: 68–70, figs. 341–347.

• mariae. Dolichoderus mariae Forel, 1885a: 349 (w.) . [USA, NJ, Cumberland Co.; MHNG, NHMB] Mayr, G. 1886d: 437 (q.) . Combination in D. (Hypoclinea): Emery, 1894d: 229. Note that the correct page number in the citation is 'Wheeler, W.M. 1905g: 308 (m.)' not 'Wheeler, W.M. 1905g: 306 (m.)' as in Bolton, 1995b: 175.
• davisi. Dolichoderus mariae subsp. davisi Wheeler, W.M. 1905g: 308 (w.) . [USA, NJ, Warren Co.] Note that the correct page number in the original citation is 'Wheeler, W.M. 1905g: 308 (w.)' not 'Wheeler, W.M. 1905g: 306' as in Bolton, 1995b: 173. Synonymy: Creighton, 1950a: 334 .
• blatchleyi. Dolichoderus (Hypoclinea) mariae var. blatchleyi Wheeler, W.M. 1917k: 462 (w.) . [USA, IN, Lake Co.] Synonymy: Creighton, 1950a: 334 .

• DISTRIBUTION. Eastern USA (MacKay, W.P. 1993b: 71).
• RECOGNITION. Antennal scapes without erect hairs (except a few at the apex) (occasionally a few present). Mesopleuron subopaque. Basal face of propodeum finely punctate; distinctly longer than wide. Erect hairs virtually absent except for ventral surface of gaster. Color variable; usually bicolored with head, mesosoma, and petiole reddish and gaster black; occasionally head, mesosoma, and petiole brown. Some specimens (especially from the deep South) may be difficult to separate from D. pustulatus which is usually smaller, has coarser punctation on basal face of propodeum, and is often concolorous brown.
• TAXONOMY. Wheeler, W.M. 1905g: 306, 316–319, 387–388, fig. A (description w.q.m., figure (lateral view) of w.) .— Wheeler, W.M. 1916r: 589 (worker, female) .— Cole, A.C. 1940b: 60.— Creighton, 1950a: 333–335, pl. 43, figs. 1–4 (figure (lateral view) w.q.m., wing) .— Wheeler, G.C. & Wheeler, J. 1966: 726–727 (larva) .— Warren & Rouse, 1969: 43.— Johnson, C. 1989a: 2–3, fig. 1, 2 (diagnosis of w., distribution, figure (dorsal and lateral views) of mesoma of w.) .— Shattuck, 1992c: figs 47, 48 (figures (full face and lateral view of mesosoma) of w.) .— MacKay, W.P. 1993b: 75–76, 102, figs. 78, 80, 82, 85, 88, map 17 (description w.q.m., figure (mesosoma) of w., figure (head, wing) of m., distribution, bibliography) .
• MISCELLANEOUS. Smith, M.R. 1924b: 81.—Talbot, 1957: 134–139.— Kannowski, 1959b: 119–120, 129–133 (mating flights) .— Carter, 1962b: 190 (distribution in NC, natural history, most common member of the genus in NC) .— Talbot, 1965: 35–36 (nests, mating flights) .
• PHOTOS.

• plagiatus. Hypoclinea plagiata Mayr, G. 1870b: 960 (w.) . [USA, IL, (no specific locality given); Museum of Stockholm] Wheeler, W.M. 1905g: 311 (q.m.) . Combination in Dolichoderus: Mayr, 1886d: 436; in D. (Hypoclinea): Emery, 1894d: 229.
• borealis. Dolichoderus borealis Provancher, 1888a. [Canada] Synonymy: Emery, 1895d: 330.
• inornatus. Dolichoderus plagiatus var. inornatus Wheeler, W.M. 1905g . [USA, (IL & NJ)] Synonymy: Creighton, 1950a: 335 .

• DISTRIBUTION. Northern and eastern USA and northern Mexico (MacKay, W.P. 1993b: 82).
• RECOGNITION. This is an easily recognized species. Antennal scapes with numerous erect hairs (rarely absent), usually at least 10. Dorsal surface of head with punctures deep and close together forming a rugo-reticulate pattern. Dorsum of mesosoma with coarse punctures; the pronotum is as strongly punctate as the mesosoma and propodeum. Basal face of propodeum distinctly longer than wide. Erect hairs abundant.
• TAXONOMY. Wheeler, W.M. 1905g: 310–312, fig. C (description w.q.m., figure (lateral view) of w.) .— Wheeler, W.M. 1916r: 590 .— Cole, A.C. 1940b: 60.— Gregg, R.E. 1945a: 467 .— Creighton, 1950a: 335 .— Creighton, 1950a: 335–336 (distribution, synonyms) .— Francoeur & Béique, 1966a: 142 (Provancher types) .— Wheeler, G.C. & Wheeler, J. 1966: 727, fig. 1 (larva) .— Johnson, C. 1989a: 3, figs. 1, 2 (diagnosis of w., distribution, figure (dorsal and lateral views) of mesoma of w.) .— MacKay, W.P. 1993b: 81–83, 102, figs. 77, 81, 84, map 18 (description w.q.m., figures w. - head, mesosoma; distribution, bibliography) .— Shattuck, 1994: 61.
• MISCELLANEOUS. Wheeler, W.M. 1905g: 306, 315–316, 388 .— Wheeler, W.M. 1908i: 621–622 .— Wesson & Wesson, 1940: 90, 99.— Kannowski, 1959b: 119–120, 133.— Carter, 1962b: 190 (distribution in NC, brief natural history) .— Wheeler, G.C. & Wheeler, J. 1963: 149–151.— Talbot, 1965: 37 (nests, mating flights) .— Kannowski, 1967: 1246–1252 (population studies) .— Ortius, 1998: 147–150 (finding of a D. taschenbergi queen in a nest of D. plagiatus) .
• PHOTOS.

• pustulatus. Dolichoderus pustulatus Mayr, G. 1886d . [USA, NJ; BMNH, MHNG] Wheeler, G.C. & Wheeler, J. 1966: 727, fig. s (l.) . Combination in D. (Hypoclinea): Emery, 1894d: 229. Subspecies of plagiatus: Wheeler, W.M. 1905g: 313. Revived status as species: Creighton, 1950a: 335. Type locality specified by Creighton, 1950a: 335, but without lectotype designation.
• beutenmuelleri. Dolichoderus plagiatus var. beutenmuelleri Wheeler, W.M. 1904f: 304 (w.) . [USA, NC, (Black Mountains)] Variety of pustulatus: Wheeler, W.M. 1913d: 116. Synonymy: Creighton, 1950a: 335 .
• beutenmuelleri. Dolichoderus plagiatus subsp. pustulatus var. beutenmuelleri Wheeler, W.M. 1905g: 313 . [USA, NC, (Black Mountains)] NEW SYNONYMY

• DISTRIBUTION. Eastern USA. Its occurrence in Brownville, TX is doubtful (MacKay, W.P. 1993b: 85).
• RECOGNITION. Antennal scapes without erect hairs (except a few at the apex) (occasionally a few present). Dorsum of mesonotum and propodeum with coarse, deep punctures that are close together. Basal face of propodeum distinctly longer than wide. Sparse erect hairs present on most body surface except antennal scapes, femora, and tibiae. Usually concolorous dark brown; occasionally bicolored with head and mesosoma dark reddish brown and gaster darker.Some specimens (especially from the deep South) may be difficult to separate from D. mariae whichis usually larger, has finer punctation on basal face of propodeum, and is usually bicolored.
• TAXONOMY. Wheeler, W.M. 1905g: 306, 313 (description of w.q., as D. plagiatus pustulatus) .— Smith, M.R. 1924b: 82.— Smith, M.R. 1931a: 22.— Creighton, 1950a: 335–336 .— Johnson, C. 1989a: 3–4, figs. 1–3 (diagnosis of w., distribution, figure (dorsal and lateral views) of mesoma of w.) .— MacKay, W.P. 1993b: 84–85, 102, figs. 83, 87, map 19 (description w.q.m., distribution) .— Shattuck, 1994: 61–62.
• MISCELLANEOUS. Wheeler, W.M. 1905g: 315–316, 388 .— Wheeler, W.M. 1916r: 462 .— Wheeler, W.M. 1932a: 12 .— Wesson & Wesson, 1940: 89, 99.— Kannowski, 1959b: 133–134, 155.— Carter, 1962b: 190 (distribution in NC, nautral history) .— Talbot, 1965: 36–37 (nests, mating flights) .— Kannowski, 1967: 1246–1252 (population studies) .
• PHOTOS.

• taschenbergi. Hypoclinea taschenbergi Mayr, G. 1866a . [North America (no specific locality given); MNHG] Wheeler, G.C. & Wheeler, J. 1951: 173, pl. I, figs. 1–6 (l.) . Combination in Dolichoderus: Mayr, 1886d: 437; in D. (Hyopclinea): Emery, 1894d: 229.
• gagates. Dolichoderus taschenbergi var. gagates Wheeler, W.M. 1905g: 310 (w.) . [USA, NJ, Ocean Co.] Junior primary homonym of D. gagates Emery, 1890. Replacement name (first): aterrimus Wheeler, W.M. 1915b: 417. Replacement name (second, unnecessary): wheeleriella Forel, 1916: 458. Synonymy: Creighton, 1950a: 336 .
• aterrimus. Dolichoderus (Hypoclinea) taschenbergi var. aterrimus Wheeler, W.M. 1915b: 417 . Replacement name for the junior homonym D. gagates Wheeler. Synonymy: Creighton, 1950a: 336 .
• wheeleriella. Dolichoderus taschenbergi var. wheeleriella Forel, 1916: 458. Unnecessary (second) replacement name for D. gagates Wheeler. Synonymy: Smith, M.R. 1951c: 834.

• DISTRIBUTION. Central and eastern USA.
• RECOGNITION. This is an easily recognized species. Antennal scapes without erect hairs (except a few at the apex) (occasionally a few present). Dorsal surface of pronotum finely punctate; mesosoma and pronotum coarsely punctate. Basal face of propodeum subquadrate - about as wide as long. Erect hairs sparse on mesosoma and gaster. Concolorous dark brown to black.
• TAXONOMY. Wheeler, W.M. 1905g: 306, 309–310, fig. B (description w., figure (lateral view) of w.) .— Smith, M.R. 1924b: 82.— Cole, A.C. 1940b: 60.— Smith, M.R. 1947f: pl. 16, fig. 60 (figure (lateral view) of w.) .— Creighton, 1950a: 336 (distribution, synonyms) .— Creighton, 1950a: 336–227 .— Wheeler, G.C. & Wheeler, J. 1966: 728 (larva) .— Warren & Rouse, 1969: 43–44.— Johnson, C. 1989a: 4–5, figs. 1, 2 (diagnosis of w., distribution, figure (dorsal and lateral views) of mesoma of w.) .— MacKay, W.P. 1993b: 99–102, figs. 79, 86, 89, map 20 (description w.q.m., distribution) .— Shattuck, 1994: 69.
• MISCELLANEOUS. Logier, 1923: 247.— Smith, M.R. 1924b: 81–82.— Dennis, 1938: 292–293.— Wesson & Wesson, 1940: 99.— Kannowski, 1959a (flight activities) .— Carter, 1962b: 190–191 (distribution in NC, natural history) .— Wheeler, G.C. & Wheeler, J. 1963: 151–154.— Bradley & Hinks, 1968: 40–50 (attending aphids on jack pine) .— Bradley, 1972: 245–249 (transplanting colonies in jack pine stands in Man.) .— Bradley, 1973: 1525–1528 (competition with Formica obscuripes) .— Burns, 1973: 97–104 (foraging and tending behavior) .— Burns, 1973: 97–104 (foraging, relationship with tuliptree scale) .— Levings & Traniello, 1982: table 1 (foraging behavior) .— Francoeur & Elias, 1985: 303–307 (natural history, fossil specimens) .— Ortius, 1998: 147–150 (finding of a D. taschenbergi queen in a nest of D. plagiatus) .
• PHOTOS.