Lasius niger

This is one of the commonest European species and is often especially evident at sites of human disturbance including towns, villages and quarries. Nests may occur in walls, pavements, tree stumps in open woodland, pasture and open heath. Occasionally earth mounts are formed and foraging tracks are frequently covered by surface tunnels of earth. This species is aggressive and readily attacks other ants. Nests are single queened and moderately populous with several hundred up to 10,000 workers. Aphids on shrubs and herbs as well as subterranean species are tended. Mating swarms occur from July to late August and in some years huge numbers may fly over a large district on the same date. (Collingwood 1979)

Identification
Wilson (1955) - The worker is best distinguished by its abundant standing appendage pilosity combined with the clypeal outline and mandibular dentition characteristic of the "niger complex" (Lasius niger, Lasius alienus, Lasius emarginatus, Lasius productus). Over most of Europe and in western North America niger has a scape index regression zone intermediate between those of Lasius brunneus and L. emarginatus, but in peripheral Eurasian populations this zone shifts to overlap that of L. emarginatus. The male is best distinguished by the possession of abundant standing pilosity on the appendages combined with the "niger type" mandible described below.

United States: Mackay and Mackay (2002) - [Schar et al. 2017 - The natural North American populations identified as Lasius niger are not conspecific and belong to a currently undescribed species.] These are medium to dark brown ants with large eyes, which have abundant erect and suberect hairs on the scapes (more than 30) and the extensor surface of the anterior tibia (more than 6). The three basal teeth are about equal in size (the teeth adjacent to the penultimate tooth may be smaller in size).

Europe: Collingwood (1979) - Greyish brown to dark brownish black, mid-body occasionally somewhat paler. All appendage surfaces including scapes and tibiae with abundant erect hairs. Length: 3.5-5.0 mm

Distribution
This species has the northernmost range of the four members of the niger complex. In Eurasia it is found from Scotland and southern Fennoscandia south to Morocco, Algeria, and the offshore Atlantic Islands, east through Lebanon, Afghanistan, and the Himalayan plateau to central China and Formosa, then north to northern Russia, southern Yakutsk, and Kamchatka. In North America it ranges from the Pacific Northwest through most of the Great Basin, the Rocky Mountains, and northern Sierra Nevada. (Wilson 1955)

Holarctic from Western United States to Japan, North Africa to Finland (Collingwood 1979).

Distribution based on Regional Taxon Lists
Nearctic Region: Canada, United States. Neotropical Region: Mexico. Oriental Region: India, Maldives, Nepal, Pakistan. Palaearctic Region: Afghanistan, Albania, Andorra, Armenia, Austria, Belarus, Belgium, Bulgaria, Channel Islands, China, Croatia, Czech Republic, Democratic Peoples Republic of Korea, Denmark, Estonia, Faeroe Islands, Finland, France, Georgia, Germany, Gibraltar, Greece, Hungary, Iberian Peninsula, Kazakhstan, Kyrgyzstan, Latvia, Lithuania, Luxembourg, Monaco, Mongolia, Montenegro, 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
Wilson (1955) - Gosswald (1932) has presented a detailed and informative account of the habitat preferences and nesting habits of this species in Germany. He found it to be one of the most abundant and adaptable native ants. It occurs in deciduous, coniferous, and mixed forests, along forest borders, in hedgerows, and in open meadows; it thrives on cultivated land and occasionally enters houses. In forests it tends to nest in rotting tree trunks. In exposed situations with little vegetation cover, it nests mostly under stones. In meadows, along grass-grown paths and forest gardens, and in gardens, it frequently builds earthern mounds, especially where the soil is moist and the ground vegetation dense. Such mounds are irregular in shape and apparently designed to surmount the surrounding vegetation, with the result that they are often quite massive and may exceed 50 cm. in height. In a random field sample, Gosswald found 625 nests under stones, 350 in mounds, and 130 in rotting wood. The species is said to occasionally construct carton nests out of macerated plant material and humus, especially when it nests under stones in pine woods; the carton material closely resembles that characteristically manufactured by Lasius fuliginosus.

The North American population shows a similar latitude in habitat preferences. Field notes supplied me with 16 collections by A. C. Cole and Borys Malkin seem to indicate a general preference for drier and more open situations. The majority of the 16 were taken in open forest of variable composition, while several were taken in meadows and grassy roadstrips. One was taken in a dry rabbitbrush association (Chrysothamnus) near Eagle Nest, N. Mex. Another was taken near timberline in a zone of stunted fir (Bloomington Peak, Wasatch Mts., Idaho; Malkin). The Eagle Nest colony was found in soil at the base of a bush; all of the others were taken under stones.

The previously published Asiatic records seem to indicate broad adaptability also. The Yalung-Yangtse collections recorded by Eidmann (1941) were made both in moist river valleys and in the high grassland of the Tibetan Plateau. All were from under stones. Kuznetzov-Ugamskij (1929a) notes that in the Ussuri region this species occurs under the "widest environmental conditions."

Ecological data accompanying several peripheral collections determined during this revision are noteworthy. On the Canary Islands (Gran Canaria, Teneriffe, La Palma) Wheeler (1927) found niger mostly in moist, shady spots between 1500 and 5000 feet. At Las Mercedes, Teneriffe (2500 feet), it was the only ant species found in a forest of tree-heath (Erica arborea) and laurel (Cerasus lusitanicus). At Ponta Delgada in the Azores niger occurred abundantly with Tetramorium caespitum, nesting under stones and foraging over the ground in files. In Lebanon, Christiansen (in litt.) took workers under rocks in mesophytic forest at Hammana and dealate queens from rotting wood in spruce forest above the Kammuoha Plain. Workers were also taken with the aid of a Berlese funnel from around the roots of grass growing on the banks of the Orontes River two miles south of Hermel. This last locality is surrounded by scrub desert and is ten miles from the nearest well developed woodland, representing an extreme habitat record for Lasius in general and a very extraordinary one for niger in particular.

Lasius niger has generalized food habits. Many authors have observed it gathering insect remains, floral nectar, and the honeydew (excreta) of Homoptera and larval Lycaenidae. Eidmann (1926) and others have suggested that the insect honeydew makes up the bulk of the diet. According to Donisthorpe (1927), in a review of the food habits of this species, niger has been observed occasionally to gather seeds of such plants as Viola, Galium, and Ranunculus; Scott (1926) has seen it collecting seeds of Chelidonium. But apparently no one has checked to see if the seeds are actually used as food. The pastoral habits of niger have been the subject of an excellent study by Eidmann (ibid.). Certain aphids overwinter in the nests of niger and are brought out and placed on the host plants in the spring. At first the aphids are returned to the shelter of the nests each evening. Later, as the nights grow warmer, the herds are left permanently in place. Certain of the workers function as guards during the day, remaining constantly at fixed posts; one worker was observed to return to the same spot each day for eight days. The workers appear to reduce predation by the braconid Trioxys, and on several occasions Eidmann saw them in the act of driving off workers of other ant species. Biisgen (1891) has observed niger workers combating chrysopid larvae in similar fashion. By counting the number of workers returning to the nest gorged with honeydew, Eidmann estimated that a large colony of niger may consume as much as a liter of this material in the course of a summer.

According to Eidmann, niger is principally nocturnal. Its above-ground activity, as measured by the number of ants soliciting honeydew, is highest at midnight and lowest in the early morning. The workers do all they can to avoid light; in order to work above ground they build turret-shaped shelters around the bases of aphid-infested plants and connect these with the nest by means of covered pathways.

Eidmann (1943) has included this species in a general study of overwintering in ants. Most of the workers of the colony move deep into the center of the nest, concentrating in a few chambers. A small number remain with the aphids in the special chambers where these insects are housed. Brood, in the form of small larvae, may be present or lacking.

According to Donisthorpe (1926), who has undertaken a thorough review of the European literature on the subject, the nuptial flights of niger take place in the afternoon and early evening from early July to mid-September. There are no records of nuptial flights in North America, but winged reproductive have been taken in nido from July 3 (Sacramento Mts., N. Mex.) to September 1 (Alta, Utah). Stray males and queens, possibly engaged in a nuptial flight, were found at Heber, Utah, on September 7 (A. W. Grundmann). Eidmann (1926) has studied reproduction and nest-founding in this species in Germany. The pair copulate in flight, fall to the ground, and separate. The queen, without attempting to take flight again, soon drops her wings and seeks cover. The first eggs are laid the following spring, and the first adult brood hatches that summer. One queen was observed to go without food for 382 days in the normal course of founding a colony.

Stroeymeyt et al. (2018) experimentally tested how workers of Lasius niger might alter their behavior after being exposed to a pathogen. They found "pathogen exposure induced behavioral changes in both exposed ants and their nestmates, which helped contain the disease by reinforcing key transmission-inhibitory properties of the colony's contact network. This suggests that social network plasticity in response to pathogens is an effective strategy for mitigating the effects of disease in social groups."

Madsen and Offenberg (2017) - Most colonies of Lasius niger are started by a single claustral queen that uses her stored energy reserves to raise her first brood. However, the high numbers of Lasius queens occurring after mating flights, coupled with the fact that queens avoid areas frequented by workers of established colonies (Sommer and Hölldobler 1995), leads to a high density of new nest foundations in the field. In response to crowding and high inter-colony competition, incipient colonies may use two different strategies to gain competitive advantage. One strategy is pleometrosis, where queens enter facultative associations with non-related queens during colony founding (Bernasconi & Strassmann 1999). A second strategy to improve competition by increasing early colony growth is the sequestration of foreign pupae and larvae through the raiding of brood from neighbouring colonies.

A laboratory study of wild collected queens in Denmark examined the demographic results of pleometrosis and brood raiding. Results showed that both methods had an overall positive effect on colony production, albeit the magnitude at which pupae transplantation boosted colony growth was much larger than pleometrosis. After six weeks, colonies with two and three queens contained, respectively, 26% and 83% more brood than control colonies with only one queen. Queen association also led to overall decreased queen fecundity and death of all resident queens in over 60% of the colonies due to queen fights. After eight weeks, colonies that received 30 or 60 foreign pupae had produced on average 256% more brood than controls and nanitic workers that were 7% longer than those from colonies that did not receive transplanted pupae.

Foraging/Diet
See the general biology discussion above for an overview of diet and foraging. Novgorodova (2015b) investigated ant-aphid interactions of a dozen honeydew collecting ant species in Western Siberia pine and aspen-birch-pine forests (54°7´N, 83°06´E, 200 m, Novosibirsk) and mixed-grass-cereal steppes with aspen-birch groves (53°44´N, 78°02´E, 110 m, near Karasuk) in the Novosibirsk Region and coniferous forests in the northeastern Altai (north end of Lake Teletskoe, 51°48´N, 87°17´E, 434 m). All of the ants studied had workers that showed high fidelity to attending particular aphid colonies, i.e, individual foragers that collect honeydew tend to return to the same location, and group of aphids, every time they leave the nest. Lasius niger showed no specialization beyond this foraging site fidelity. Foragers tended Chaitophorus populeti (Panzer), Aphis craccivora Koch, Aphis viburni Scopoli and Aphis pomi De Geer.

Bles et al. (2018) showed how the dynamics of pheromone trails were important in allowing foragers of this species to travel along straight paths, and therefore spent less time traversing from a food source to the nest, versus bent paths of an equal length.

Guiliani et al. (2019) observed this species foraging on extrafloral nectaries of the invasive Reynoutria x bohemica (Polygonaceae) in Tuscany. The habitats examined were river banks and disturbed habitats.

Reproduction
Chérasse et al. (2019) examined how mating may alter changes in gene expression in queens of Lasius niger. They targeted six genes likely to be important in physiology changes in reproductive function, longevity and immune system function. The specific genes studied were selected based on results found in other studies of newly mated female insects. Their study compared gene expression in virgin queens that had exited their natal nest and were about to fly off for their mating flight to 1-day and 5-day post-mated monandrous and polyandrous queens. All queens were collected from a 2017 mating flight in Belgium. There were 8 queens in each group of the 5 types of queens examined (pre-mated queens, and singly and multiply mated 1 day and 5 day post-mated queens). The inducement of oviposition was found to be followed by an up-regulation of vitellogenin and defensin expression. Vitellogenin is a major yolk protein precursor directly linked to egg production. It was up-regulated in queens mated 5 days earlier. Defensin, an antimicrobial peptide, was up-regulated in 1 day but not 5 day post mated queens. An increase in antimicrobial peptides post mating, and other presumed immune responses, have been observed in other insects. While untested, it has been suggested this could be a response to a copulatory bacterial infection or may serve as a prophylactic measure to prevent such infections.

The overall results were suggested to provide a few conclusions about the effect of mating on queen physiology in insects. First, post mating up-regulation of vitellogenin and antimicrobial peptide expression is a general effect found across a broad range of insects. Second, certain impacts of mating on female insect physiology are generalized across species independent of their mating strategies, while others seem to be species-specific. Lastly, variation in queen-mating frequency does not appear to affect gene regulation. All of these conclusions were offered with the caveat that gene expression changes in newly mated insects has not been extensively examined.

Genetics
Quque et al. (2019) examined and compared proteomes of L. niger workers and queens and foraging and non-foraging workers. Beyond morphological differences, there are also known differences in hormone production and gene expression between queens and workers. Ant proteomes, i.e, the full complement of proteins produced by individuals, have barely been explored in ants. This may be a promising area of study to investigate phenotypic differences between castes and different types of workers.

Of the 2707 proteins found, 255 were assigned to 35 functional groups. Exploratory data analyses showed there were distinctions between the types of individuals in their proteins and, from the assigned functions, in their phenotypes. Queens were mainly characterised by proteins involved in somatic maintenance – mechanisms aiming to avoid or repair damages to macromolecules – cell division, gene expression regulation and trafficking. While workers expressed more proteins related to metabolic pathways (except lipid metabolism) and sensitive nervous system or immunity. Foragers presented more proteins associated with xenobiotic detoxification mechanisms, whereas nest-workers had more proteins involved in digestion metabolism.

These distinctive proteins and their functional expression are speculated upon for each set of individuals and in contrast to one another. For example, one worker sensitive nervous system protein was identified as being linked to olfactory and gustative processing. This may be helpful in deciphering environmental cues related to foraging for food and with the chemically rich communication between nestmates.

Overall, Quque et al. conclude "We showed that proteomics allows assessing fine molecular differences induced by task specialization in a social insect species. The nontargeted screen of the whole proteome highlighted a wide diversity of caste-dependent functions from immunity to reproduction to digestion to insecticide resistance. Our study also raises evolutionary questions about longevity and energy trade-offs in eusocial species, beyond the classical free-radical theory of ageing. Thanks to our exploratory approach, we now have a more global insight into all the functions that can be affected by the division of labour in a eusocial species. Some are well studied (e.g. social immunity), others less, especially in the adult stage (e.g. the ToR pathway, difference in the metabolism of digestion). We therefore hope to pave the way for future experiments to accurately test the numerous and diverse molecular mechanisms induced by a eusocial lifestyle."

New Mexico
Mackay and Mackay (2002) - Occurs in Ponderosa pine-riparian habitat, oak forests, deciduous forest, fir forests, mixed forests and residential areas. This species nests under stones or logs, or in earthen mounds, in rocky loam soils. Reproductives were found in a nest in late August. Workers are primarily nocturnal and feed on insect fragments, nectar and honeydew. It actually moves aphids in and out of its nest and places them on plants, depending on the environmental conditions. Workers guard the aphids when they are on plants. This species also collects seeds, although it is unknown if the seeds are consumed. It occasionally enters houses.

Other Insects

 * This species is a host for the temporary parasites, , , , and.
 * This speices is a host for the encyrtid wasps, and , and the pteromalid wasps ,  and  (Universal Chalcidoidea Database).
 * This ant has been associated with the butterfly, ,  and a species that has recently been split into two species: Polyommatus icarus and Polyommatus celin (Obregon et al. 2015).

Nematodes

 * This taxon is a host for the nematodes (Rubstov, 1970),  (Kaiser, 1986a, 1986b),  (Laciny, 2017),  (Crawley & Baylis, 1921),  (O'Rourke, 1946; O’Grady & Breen, 2011; Laciny, 2021) and  (Gösswald, 1938; Laciny, 2021).

Nomenclature

 *  niger. Formica nigra Linnaeus, 1758: 580 (w.) EUROPE. Latreille, 1798: 43 (q.m.); Wheeler, G.C. & Wheeler, J. 1953c: 148 (l.); Hauschteck, 1962: 219 (k.); Imai & Kubota, 1972: 196 (k.). Combination in Lasius: Fabricius, 1804: 415; in Donisthorpea: Donisthorpe, 1915d: 200; in Formicina (Donisthorpea): Emery, 1916b: 240; in Acanthomyops: Forel, 1916: 460; in Lasius: Menozzi, 1921: 32; Müller, 1923: 124; Kuznetsov-Ugamsky, 1929a: 26; Wilson, 1955a: 59. Senior synonym of pallescens: Mayr, 1861: 49; of nigerrima: Emery, 1892b: 162; of alienoniger, emeryi, nitidus, minimus, transylvanica and material of the unavailable name pilicornis referred here: Wilson, 1955a: 59; of alienoniger: Kutter, 1977c: 14. Current subspecies: nominal plus pinetorum. See also: Imai, 1966: 119; Tarbinsky, 1976: 134; Kutter, 1977c: 227; Yamauchi, 1979: 152; Collingwood, 1982: 287; Kupyanskaya, 1990: 219; Seifert, 1992b: 27.

Type Material

 * Neotype worker plus 2 workers from the neotype nest labelled S:B1. Johannishus 1 km NE k:a, RN-03F6f03, 10.08.74, P. Douwes DATA ZOOTAX; depository ZMLU Lund (Seifert, 2020).

Worker
Wilson (1955) - (1) More than 95 per cent of the workers within HW range 0.61-1.21 mm., exclusive of material from North Africa, the Balearics, Azores, Madeira, and eastern Asia, possess a SI between 95 and 103. This is a strongly allometric character (Fig. 5), with minimas (HW less than 0.61 mm.) ranging up to 109.

(2) As a corollary to (1), ML within this sample exceeds EW.

(3) Mandibular dentition characteristics of the niger complex: basal teeth two to four in number, equal in size, and spaced at even intervals; opposed to the neoniger complex, in which the two or three basal teeth are irregularly spaced and the central one of a set of three is often reduced in size.

(4) Clypeus characteristic of the niger complex: when the mandibles are opened and the head is viewed in perfect full face, the anterior border of the median clypeal lobe describes an even, broad parabolic curve, with the sides at least feebly convex and only occasionally meeting in a point at the midline; opposed to the more angular clypeal border of the neoniger complex. (See Pl. I, Figs. 2 and 3.) The clypeus is usually, but not always, keeled.

(5) Scapes and tibiae always with abundant standing pilosity, except in minimas with PW less than 0.47 mm. The inclination and density of this pilosity show striking geographic variation (see below). In general, niger complex workers with seta counts greater than 25 are almost certainly niger, but those with less might be alienus and should be determined with the aid of the allometry regression zones plotted in Figure 6.

In a sample of 165, with no more than 2 per nest series, PW range, 0.40-0.83 mm., mean with standard error 0.630 ± 0.005 mm., standard deviation 0.069 mm.

Thirty-four individuals each representing a different nest series were examined especially for dentition; 27 had three basal teeth, 5 had four, and 2 had four with the antepenultimate tooth reduced. in size; one lacked the second intercalary tooth.

Petiole outline more variable than in other members of the subgenus. Among 52 nest series examined especially for this character, the dorsal border was gently convex in 14, straight in 11, roundly concave in 22, and angularly concave in 5.

In a strong, reflected, artificial light, the body surface is subopaque to moderately shining. The degree of shininess varies inversely with the density of the pubescence and coarseness of the shagreened sculpturing, both independent and highly variable characters by themselves.

Body nearly or completely concolorous medium to blackish brown. Legs typically medium brown, scapes tending to yellowish brown.

Queen
Wilson (1955) - (1) Within the geographic limits stated in the first worker character above, queens with HW between 1.54 and 1.82 mm. have an SI between 72 and 80.

(2) As a corollary to (I), queens within this sample have a ML between 0.17 and 0.23 mm., with over 90 per cent falling within 0.18-0.21 mm.

(3) Clypeus and mandibular dentition as in worker.

(4) Scapes and fore tibiae with abundant standing hairs; seta count usually over 20 and often over 40. Inclination and density subject to geographic variation as in worker.

(5) Wings hyaline except for a small area distal to the axillary sclerites; this is a general niger complex character.

Male
Wilson (1955) - (1) Within the geographic limits stated in the first worker character above, SI 62-69, ML 0.14-0.16 mm.

(2) Mandible characteristic of the niger complex: primitive preapical cleft of Lasius pallitarsis and Lasius brunneus modified into a shallow angular depression placed centrally on the masticatory border or lacking altogether, so that the masticatory border curves gently inward from the apex and then outward to meet the basal border. The basal angle broadly rounded, the masticatory border curving gradually into the basal border. Denticles lacking on the masticatory border.

(3) The subgenital plate subrectangular, the posterior sclerotized, setiferous lobes not more than one-fourth as wide across their base as the plate itself but protruding past the unsclerotized posterior rim and reaching as far back as the level of the tips of the posterolateral flanges. A single series from Shriek, Belgium (A. Raignier leg.; MCZ), contains individuals with two distinct lobes, a single lobe, and several stages intermediate in the coalescence of two lobes.

(4) Long standing hairs present over most of the surfaces of the scape and fore tibia, but much sparser than in the worker and queen; seta count usually less than 5.

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