Lasius brunneus

This species is widespread in Europe and is found primarily only where broad-leaved trees occur (Rigato & Toni, 2011). In Greece, it is a widespread but rare species. It prefers mountain forests growing on high altitudes. In Achaia and Aetolia-Acarnania, it was observed exclusively in fir forests (Borowiec & Salata, 2021).

Identification
Seifert (2020) - Palaearctic Lasius s. str. species belonging to the Lasius brunneus species complex. The species is rather easily identified by a combination of short scape (SL/CS900 0.874), broad head (CL/CW900 1.041), reduced setae numbers on all body parts (nOcc900 2.0, nGen900 0.1, nGu900 1.3, nSC900 0.0, nHT900 0.1, nSt900 1.1) a very smooth pubescence surface on scape and tibiae, and reduced number of mandibular dents (MaDe900 7.06). The dorsal and posterior profile of propodeum are linear and form a distinct, obtuse angle. The petiole is in anterocaudal view rather rectangular or slightly converging dorsad but typically with straight sides and forming a sharp, weakly emarginate dorsal crest. A typical coloration of medium-sized to large specimens is mesosoma, petiole and appendages light yellowish brown, head a little darker bronze brown and gaster dark to blackish brown. However, small workers from initial colonies may show a homogenously dark coloration as seen in Lasius lasioides or Lasius neglectus but can easily be separated by RAV-corrected NUMOBAT data.

For separation from the eastern sister species Lasius silvaticus and Lasius himalayanus see identification sections for those species.

Collingwood (1979) - Bicoloured with gaster dark brown contrasting with testaceous or pale reddish brown head and alitrunk. Pubescence and body hairs sparse. Occipital hairs restricted to median area of back of head only. Scapes and tibiae never with erect hairs. Back of head flat or feebly concave. Frontal triangle and frontal furrow distinct, ocelli small but always clearly visible. Length: 3.2-4.5 mm.

Wilson (1955) - A distinct species characterized by large males with intermediate sitkaensis-niger type mandibles and workers with short scapes and sparse pilosity.

Distribution
Seifert (2020) - Eurocaucasian, submeridional and temperate. From S. England and Iberia across Central Europe, the Apennine and the Balkans to Asia Minor and the Caucasus. In Scandinavia north to 60°N, absent from Finland. In N. Tyrol ascending to 1410 m and in Anatolia at 37°N to 2000 m.

This is known mostly from mountain forests of all mainland provinces, the Ionian Islands and the Aegean Islands (Borowiec et al., 2022).

Distribution based on Regional Taxon Lists
Oriental Region: India, Pakistan. Palaearctic Region: Albania, Andorra, Armenia, Austria, Belarus, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Finland, France, Georgia, Germany, Greece, Hungary, Iberian Peninsula, Iran, Israel, Italy, Latvia, Lithuania, Luxembourg, Netherlands, Norway, Poland, Portugal, Republic of Korea, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, United Kingdom of Great Britain and Northern Ireland.

Biology
Seifert (1992) - Lasius brunneus is found in all habitats where deciduous trees are present - from dense woodland to open land with scattered trees or hedgerows. Pure, shaded coniferous forests are avoided but light Pinus forests or mixed conifer-deciduous forests are inhabited at lower densities (0.1-2 nests/100 m2). Top densities of 10-23 nests/100 m2 were found in Quercus-Carpinus-Tilia-Acer forests in the Harz mountains in Germany (Seifert 1986). In such sites, big trees are sometimes occupied by two colonies as indicated by intraspecific fightings.

The nests are preferentially constructed in wood and under bark of living or dead deciduous trees, from subterranean parts up to the main branches. Dead parts of a tree are preferred but occasionally galleries may penetrate living wood. Nests containing brood and situated several meters away from the next tree are frequently found under stones or in litter. Possibly such nests are mainly of a temporal nature and represent a seasonal phenomenon. The occurrence of polycalic colonies is very likely but not definitely proved. Collingwood (1979) reports acceptance of dealate queens after swarming. Lasius brunneus is known to occur inside the timbers or stone walls of houses (in Germany, however, much less frequently than Lasius emarginatus).

Trophobiosis with tree aphids, including the big Stomaphis quercus, is obviously the major food source but transport of insects to the nest is frequently observed. Lasius brunneus is very fugitive and quickly escapes after the opening of nests. Workers avoid running over free open surfaces and move as long as possible in the shelter of crevices or grooves. The swarming period begins earlier than in most species of the group and lasts in Central Europe, according to my own records, from 10 June to 19 July (25 June±12 d, n = 14). There seems to be no special time of day for flying but the period near noon (11-14 h) was most frequently noted (own observations; Donisthorpe, Forsslund both cited by Wilson 1955). Early morning flight (5-6 h) is reported by Schenck (Stitz 1939) and even nocturnal flight seems possible. However, the observed light trap catches could be the result of artificial provocation by the light and no result of natural activity.

Collingwood (1979) - This species nests in the interior of old trees, chiefly oak, but has also been recorded from hedgerows. It is fugitive and non-aggressive, rapidly dispersing on disturbance and because of its cryptic habits may be somewhat under-recorded. In Norway and Sweden it has frequently occurred nesting in the timbers of old houses and farm buildings, where its populous colonies may be difficult to dislodge. It chiefly tends tree aphids including the large bark feeding Stomaphis. Single queens initiate colonies in the crevices of old trees but may also be accepted back into the mother nest after the mating flight which occurs in June and early July.

Wilson (1955) - Most European authors agree that brunneus is a timid species adapted to living under the bark and in the wood of tree trunks. Donisthorpe (1927) found a large population of this species in the Windsor Forest of England limited to living trees, which the ants penetrated from the trunk up into the main branches and down into the roots. Various trees were inhabited, including oaks, elm, ash, beech, poplar and maple. It is not clear whether the workers carried on much excavation in the living wood, but this seems unlikely due to the rather unspectacular mandibular apparatus of the species. Forsslund (1949) found brunneus in oaks in dense, undisturbed woodland in several localities in the vicinity of Stockholm. The nests were mostly in dead wood, but occasional galleries penetrated living wood. Scherdlin (1909) found the species in Alsace nesting in the trunks of trees and timber of houses. Clausen (1938) observed a swarm of reproductives inside a house in Zurich. Gosswald (1932) states that in Germany brunneus is found as often under stones as in dead wood; since this observation is divergent from those of other authors, the possibility must be considered that he was erroneously including some alienus in his concept of brunneus.

Donisthorpe (ibid.), who has undertaken the most careful study of this species to date, found workers transporting and tending aphids of the genus Stomaphis. He also observed them carrying psoeids and other small insects to the nests, presumable for use as animal food.

Lasius brunneus appears to hold its nuptial flights earlier in the day and season than other European members of the subgenus. Donisthorpe (ibid.) encountered winged queens and males swarming over the trunk of an oak at noon on June 25, and Forsslund (ibid.) saw the same thing from noon to 1:30 p.m. during the period June 10-16.

Also see Seifert (1992).

Other Insects
Reports of Lasius fuliginosus invading Lasius brunneus nests (Janda et al., 2004) are unlikely based on biology (Seifert, pers. comm., in de la Mora et al., 2021).
 * This species is a host for the temporary parasites and.
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This species is associated with the aphids, , , , , , , , , , , , , , , , , , and  (Saddiqui et al., 2019 and included references).
 * This species is associated with the aphids, , , , , , , , , , , , , , , , , , and  (Saddiqui et al., 2019 and included references).
 * This species is associated with the aphids, , , , , , , , , , , , , , , , , , and  (Saddiqui et al., 2019 and included references).

Nematodes

 * This species is a host for the nematodes, , , and  (Kohler, 2012).

Nomenclature

 *  brunneus. Formica brunnea Latreille, 1798: 41 (w.q.) FRANCE. Mayr, 1855: 358 (m.); Lorite, Chica & Palomeque, 1998: 28 (k.). Combination in Lasius: Mayr, 1861: 50; in Donisthorpea: Donisthorpe, 1915d: 347; in Formicina: Emery, 1916b: 241; in Lasius: Müller, 1923: 127; in Acanthomyops: Kuznetsov-Ugamsky, 1927e: 188; in Lasius (Lasius): Wilson, 1955a: 47. Subspecies of niger: Forel, 1874: 47; Forel, 1892i: 307; Bondroit, 1910: 486; Kuznetsov-Ugamsky, 1927e: 188. Status as species: André, 1881b: 60; Dalla Torre, 1893: 182; Bondroit, 1912: 352; Forel, 1915d: 52; Bondroit, 1918: 26; Müller, 1923: 127; Karavaiev, 1927c: 279; Finzi, 1930d: 316; Menozzi, 1939a: 313; Wilson, 1955a: 47; Bernard, 1967: 358; Kutter, 1977c: 228; Collingwood, 1982: 285; Atanassov & Dlussky, 1992: 238; Seifert, 1992b: 6. Senior synonym of pallida: Latreille, 1802c: 169; Wilson, 1955a: 47; of timida: Smith, F. 1858b: 7; Seifert, 1992b: 6; of alienobrunneus: Stärcke, 1944a: 157; Wilson, 1955a: 47; Kutter, 1977c: 14; Seifert, 1992b: 6; of nigrobrunneus: Wilson, 1955a: 47; Seifert, 1992b: 6. Current subspecies: nominal plus emarginatobrunneus.
 * pallida. Formica pallida Latreille, 1798: 41 (w.q.) FRANCE. Junior synonym of brunneus: Latreille, 1802c: 169. Unrecognisable taxon, incertae sedis in Lasius: Seifert, 1992b: 48. [Note. As the taxa brunneus and pallida were both described by Latreille and the synonymy was also established by Latreille, it seems reasonable to let it stand.]
 * timida. Formica timida Foerster, 1850a: 35 (w.) GERMANY. Schenck, 1852: 54 (q.m.). Junior synonym of brunneus: Smith, F. 1858b: 7; Seifert, 1992b: 6.
 * alienobrunneus. Lasius niger var. alienobrunneus Forel, 1874: 47 (w.) SWITZERLAND . Emery, 1916b: 249 (q.). Junior synonym of alienus: Dalla Torre, 1893: 182. Revived from synonymy as subspecies of brunneus: Ruzsky, 1902d: 17. Raised to species: Bingham, 1903: 340. Subspecies of brunneus: Forel, 1915d: 53; Karavaiev, 1927c: 279. Junior synonym of brunneus: Stärcke, 1944a: 157; Wilson, 1955a: 47; Kutter, 1977c: 14; Seifert, 1992b: 6.
 * nigrobrunneus. Acanthomyops (Dendrolasius) brunneus var. nigrobrunneus Donisthorpe, 1926b: 18 (w.) ITALY. Junior synonym of brunneus: Wilson, 1955a: 47.

Seifert (2020) - Lasius brunneus, Lasius silvaticus and Lasius himalayanus are hypothesized to represent three cryptic species with different geographic distribution.

Worker
Wilson (1955) - (1) Scape shorter relative to head width than in any other member of the subgenus; S1 82-91 in all European series measured; 94 in the himalayanus lectotype and in one specimen from Lahore, Pakistan, both small specimens.

(2) Small individuals (PW 0.50-0.57 mm.), when viewed in perfect full face, with the lateral margins of the eyes not reaching the lateral borders of the head; in Lasius niger and Lasius alienus they reach or exceed it.

(3) Mandibles proportionately shorter, more incurved, and inserted slightly closer to the midline, and head more massive relative to the alitrunk, than other Lasius s. s. (Pl. 1, Fig. 9). Occipital margin viewed in full face flat to feebly convex, as opposed to the typically concave outline of niger and alienus.

(4) Mandible with only two basal teeth in all of seven nest series examined for this character.

(5) Scapes and tibiae completely devoid of standing hairs and nearly devoid of hairs of any inclination. Body pilosity sparse; the curving portion of the occipital angles viewed in full face typically devoid of hairs, rarely with one or two; the latter condition occurs in other members of the subgenus but is highly exceptional.

(6) Alitrunk and petiole homogeneous light reddish brown, rarely medium reddish brown, contrasting against the dark brown gaster. The head usually the same color, occasionally darkening to medium or dark reddish brown to contrast against the alitrunk. (niger and alienus typically concolorous.)

Size range and dispersion probably about the same as in niger. In a sample of 29, with no more than 2 individuals per nest series, PW range 0.50-0.73 mm., mean with standard error 0.630 ± 0.012 mm., standard deviation 0.063 mm. ML less than EW. Anterior margin of median clypeal lobe and structure of the mandibular basal angle essentially as in the niger complex. The greater massiveness of the alitrunk in this species can perhaps best be expressed as a ratio of alitrunk length to the maximum head depth measured perpendicular to the long axis of the head. Several medium-sized brunneus gave such a ratio of 67-69, whereas alienus of comparable size ranged between 70 and 76. Viewed from the side the propodeal angle tends to be more acute and the declivitous face of the propodeum tends to be longer relative to the dorsal face than in other Lasius s. s. The dorsal margin of the petiole was invariably concave in all series examined; occasionally the concavity is so deep as to be nearly semicircular.

Queen
Wilson (1955) - (1) SI low; 68-71 in 9 individuals from 6 localities having HW 1.49-1.64 mm., and 76 in a smaller individual with HW 1.39 mm,

(2) Pilosity and mandibular dentition as in worker.

(3) Frontal suture well marked, set in the middle of a conspicuous broad, shallow trough.

(4) Color distinctive; body uniformly dark reddish brown, appendages a contrasting yellowish brown.

(5) Fore wings infumated in the inner and upper thirds.

Male
Wilson (1955) - (1) Larger than other members of the subgenus, HW 1.04-1.10 mm.

(2) Mandibles of a type intermediate between Lasius pallitarsis and Lasius niger: there is a well marked subapical cleft as in sitkaensis, but it is set more posteriorly than in this primitive species; the basal angle is broadly rounded and the masticatory border lacks denticles, both of which characters are associated with the more advanced niger type.

(3) Frontal suture conspicuous as in queen.

(4) The entire dorsal petiolar margin involved in a deep concavity. In a series from Windsor Forest, Berks, England, secondary lateral convexities are present within this concavity.

(5) Parameres shorter relative to HW than in other members of the subgenus.

SI of several individuals measured 60-64, overlapping part of range of variation of niger (q. v.). ML 0.15-0.17 mm., overlapping part of ranges of niger and emarqinatus.

Type Material
Seifert (2020) - Neotype (top) of three worker specimens collected near Brive, labelled FRA: 44.9125°N, 1.4769°E, Souillac 1.7 km N, 120 m, tree row, leg. Galkowski 2008.09.07 and Neotype (top) Lasius brunneus (Latreille 1798) des. Seifert 2019 ; depository:.

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