Camponotus fedtschenkoi

Camponotus fedtschenkoi Mayr, 1877 is a carpenter ant from the drylands of Asia: sandy deserts, steppes and mountain valleys stretching from the lower Volga to western China. It is a medium-sized, polymorphic, stingless Camponotus with workers of several sizes; in its light form, the body is such a pale yellow that it looks almost white.

It is one of our best-selling species, and also one of the least studied in the wild: almost everything published about it consists of descriptions, catalogues and faunal lists. That is why, as always, we keep apart what has been published, what is known about its genus, what keepers report and what we recommend at Antomas.

Biology

A naturalist, a glacier and an ant

The name honours Alexei Pavlovich Fedchenko (1844–1873), a Russian naturalist born in Irkutsk. Between 1868 and 1871 he travelled across Turkestan with his wife, the botanist Olga Fedchenko: the Zeravshan valley, Samarkand, the Fan Mountains, the Kyzylkum desert and the Alai valley, where he discovered the Trans-Alai range. He returned with large collections of plants and animals, which were shared out among the specialists of the day. The ants went to the Austrian Gustav Mayr, who described this species in 1877 in the Russian account of the journey and published it again in German and Latin in 1880. Its type specimens – workers, queens and males – were collected in those valleys between 1869 and 1871.

Fedchenko never saw the ant that bears his name. He died in September 1873, aged just 29, on a glacier in the Mont Blanc massif, caught in a storm while training for an expedition to the Pamirs. In the very Pamirs he never managed to explore, his name now lives on in the Fedchenko Glacier: some 77 km long, it is considered the longest glacier in the world outside the polar regions. It was discovered by another Russian expedition in 1878, five years after his death.

A century under other names

Mayr described it as a new species, although he defined it with a Latin phrase that says it all: ‘very similar to Camponotus sylvaticus’, the nocturnal carpenter ant of the western Mediterranean that we also stock at Antomas. What set it apart, in his view, were the short, oblique hairs that densely cover its legs and antennae.

For almost a century, specialists could not agree:

• From 1879, Auguste Forel and others treated it as a subspecies of C. sylvaticus, and in 1886 Forel moved it to C. rubripes.
• From 1893, Carlo Emery placed it as a subspecies of C. maculatus, the great catch-all of 19th-century carpenter ants and the same name borne by our giant African carpenter ant, C. maculatus. In 1925 he placed it in the subgenus Tanaemyrmex.
• In 1962, the Soviet myrmecologist Gennady Dlussky restored it to species rank, where it remains.
• In 1997, Alexander Radchenko, in his review of the Camponotus of the Asian Palaearctic, treated C. tichomirovi, described in 1905, and the variety adi, described in 1930, as its synonyms.

Today it is a valid species with no subspecies. Its subgenus, Tanaemyrmex, should be taken with caution: according to the 2016 classification of the formicines, many of the subgenera of Camponotus do not reflect true relationships and are awaiting a thorough revision. And the yellow form kept in Europe is just that, a colour form, with no scientific name of its own.

From the Volga to Xinjiang

Its range covers much of the drylands of Central and Western Asia. It has been recorded in Uzbekistan, Tajikistan, Turkmenistan, Kyrgyzstan, Kazakhstan, Afghanistan, Iran, Armenia, Azerbaijan, Russia and China, between roughly 34 and 45° north. It also appears on the checklist of the ants of Turkey, although Turkish specialists consider that record doubtful, because material from eastern Anatolia closely resembles another species, C. buddhae.

• In China, the first record was published in 2000: small pale workers caught in pitfall traps in the Junggar Basin, in Xinjiang, a region of stabilised dunes with scattered scrub, poplar groves and irrigated land that receives barely 115–164 mm of rain a year.
• In Russia it is known from a single site, the Bogdinsko-Baskunchak Nature Reserve, in the semi-desert of the lower Volga. The entomologists who studied it there describe it as a ‘little-known’ species.
• In the mountains it climbs high: the variety adi, now a synonym, was described from four workers collected at an altitude of 2,800 m by the 1928 Pamir expedition.

Deserts, semi-deserts, steppes and mountain valleys: this is an ant of open, dry ground. The catalogues say little more about where it nests; shops and keepers describe it nesting in the soil and under stones, like its relatives.

A climate of extremes

Where it lives, the climate is continental and dry, and the ant has to cope with both extremes:

• Furnace-like summers. In the low deserts, July averages around 28–32 °C, with highs close to 40 °C, and the surface of the sand exceeds 60–70 °C. There is almost no rain from June to September.
• Proper winters. January averages 1–5 °C in the lowlands and stays below freezing further north and in Xinjiang. There are frosts from December to February and, during cold snaps, lows of −20 to −30 °C. What little rain falls during the year comes between November and April.
• Shelter underground. The nest buffers both. In sandy areas in the north of its range, at a depth of 20–40 cm, the soil reaches about 24–32 °C at midday in July, a far cry from the furnace at the surface. In winter, at nest depth, we estimate that it stays at about 5–13 °C for around three months, without freezing. These are the two figures on which the temperatures and the diapause we recommend for home keeping are based.

Widely sold, little studied

Many claims circulate about this species that we have not been able to back up with any source. There is no published study on:

• Its nuptial flights: when they take place, at what time of day or whether they are mass flights.
• The number of queens in the wild: there are no genetic or field data. Shops and keepers describe it as monogynous, with a single queen, and that is how we keep it at Antomas, but nobody has studied it.
• The size of its wild colonies, or whether a colony occupies several nests or dominates large areas.
• Its daily rhythm: some shops describe it as nocturnal, but nobody has studied it.
• The strength of its formic acid, which is sometimes said to be ‘weak’ without any data to support it.

What is documented is its description, its distribution and its relationships. The rest comes from its genus and from keepers, and we point this out in each case.

Shepherd of aphids and scale insects

Like almost all Camponotus, it lives mainly on sugary liquids: plant nectar and, above all, honeydew, the sweet, sticky liquid excreted by aphids, scale insects and other sap-sucking insects, which the ants ‘milk’ and protect. In their study of the ants of north-western China, Cedric Collingwood and Harold Heatwole note that the region’s Camponotus depend mainly on honeydew from aphids on trees and shrubs, and rarely stray far from them. They supplement this with protein: insects they hunt or find dead and carry back to the larvae.

A common mix-up: you sometimes read that these ants tend ‘aphids and coccinellids’. Coccinellids are ladybirds, which eat aphids and are enemies of the ants that tend them. The correct phrase is aphids and coccids, that is, scale insects. For the keeper, the lesson is simple: sugar must always be available (see ‘Feeding’).

Bacteria that make protein

How does a colony that drinks so much sugar manage to grow? Partly thanks to a lodger. In special cells in their gut, Camponotus and their close relatives harbour Blochmannia, a bacterium that the queen passes on to her daughters inside the eggs. In 2007, Heike Feldhaar’s team showed that this bacterium supplies essential amino acids: on a diet poor in them, colonies treated with an antibiotic that reduced the bacterium reared far less brood, and recovered when those amino acids were added. Two years later, a study of their family tree suggested that the ancestors of these ants acquired it by tending sap-sucking insects, the very ones that provide them with honeydew today.

Nobody has studied it in C. fedtschenkoi specifically, but it is a trait of its whole group. It helps, but it is no substitute: without insects in their diet, the larvae do not grow.

The crop on display

Workers store the liquid they collect in the social stomach (crop), a pouch of the digestive tract inside the gaster that works as a shared larder: they later pass it mouth to mouth to their nestmates and the larvae, a process known as trophallaxis. In a dark ant this is barely noticeable, but in the pale form of this species, with its thin, light cuticle, the full gaster stretches, the pale bands between the segments become visible and the liquid shows through. Some call it the ‘lava lamp’ effect: given dyed nectar, the workers take on its colour.

That does not make it a honeypot ant: it has no repletes hanging from the ceiling as living storage vessels, as in Myrmecocystus. It is the ordinary crop of any Camponotus; it is just that here you can see it.

A queen who founds alone

When she lands after the flight, the young queen sheds her wings, digs a chamber, seals it and raises her first workers without going out to forage. This is claustral founding, the usual method in the genus. Everything she needs she carries with her: the wing muscles she will no longer use and a reserve of fat and proteins. In C. festinatus, a carpenter ant from the North American deserts, these storage proteins, the hexamerins, have been identified: the queen builds them up before the flight and gradually uses them to feed her larvae, until they run out once she has workers.

This has not been measured in C. fedtschenkoi, but the mechanism is the same in all claustral foundresses. That is why a queen with brood can raise her first workers in a test tube without help, although at Antomas we recommend offering her a small snack now and then (see ‘Founding and first workers’).

No sting, but plenty of acid

It belongs to the formicines, the subfamily of ants that lost the sting and traded it for chemistry. The tip of its gaster ends in the acidopore, an opening fringed with tiny hairs through which it sprays formic acid, the very compound named after ants (formica in Latin). It uses it for defence and to alert its nestmates, and the major workers back it up with a bite.

There are no data on the strength of its acid compared with that of other species. What is known about the subfamily, though, is intriguing: in 2015, a team from the University of Texas found that nine formicines from different lineages, after being exposed to fire ant venom, cleaned it off by smearing themselves with their own acid, which neutralises it. A home-made antidote.

Size and morphology

A medium-sized, shiny, polymorphic Camponotus: the workers form a continuous series of sizes, from the smallest to the largest, with no separate soldier caste. The so-called ‘soldiers’ are major workers, larger and with broader heads.

• One name, two colours. Mayr already described two forms: a dark one, shiny and brown, with the legs, petiole and part of the gaster reddish yellow, and a light one, especially among the minor workers, entirely reddish testaceous – that is, reddish yellow – with the head often redder. In his 1997 review, Radchenko characterises it by a body ‘yellow of various shades’.
• The light form. In the lower Volga reserve, workers and queens have a brown to yellowish-brown head, darker than the whitish-yellow thorax and gaster. In Xinjiang, ‘small pale workers’ were recorded. This is the form kept in Europe, and the one we offer: workers of a very pale, almost translucent yellow, and major workers with a head darker or redder than the body.
• The dark forms. Brown or dark colonies turn up in the trade from time to time and are sold as rarities. They are the same species: the colour variation has been noted since the very first description.
• Callow workers. They emerge entirely yellow and their heads darken within a few days. A young colony with many newly emerged workers looks paler than it really is.
• Pilosity. Moderately long erect hairs on the head, the top of the thorax and the gaster, and many short, oblique hairs on the legs and antennal scapes. This is the character Mayr used to separate it from C. sylvaticus, and the one used in keys to identify it.
• Surface and mandibles. Shiny cuticle with a fine, leather-like microsculpture. Mandibles with 6–7 teeth.
• Gaster. Shiny, with sparse pubescence. It visibly swells when the crop is full (see ‘The crop on display’) and ends in the acidopore.
• Queen. Darker than the workers: according to Mayr, head and thorax blackish brown, and legs and sides of the gaster reddish yellow. Before the flight, wings almost transparent, yellowish.
• Male. Brown, and distinguishable from that of C. sylvaticus only by the pilosity of the legs and antennae.

Approximate sizes

• Workers: 5.8–10 mm, from minors to majors
• Queen: 12–13 mm
• Male: 7.3–7.6 mm

These are the measurements of the type specimens given by Mayr (1880). We have found no published head measurements, nor any for particular populations. In the hobby there is talk of major workers up to 12 mm and somewhat larger queens, but these figures do not come from published measurements, so we prefer not to give them. Nor do the figures of 3 mm workers or 1 cm queens that circulate online have any basis: the smallest worker among the types measures almost 6 mm. Nanitics, the first workers of a colony, may be slightly smaller.

Care in captivity

Difficulty: 1/5 – Beginner

At Antomas we consider it one of the best Camponotus to start with: it eats almost anything, copes well with a dry nest once the colony is up and running, has no sting and, once established, is a calm, clean ant. For everything to go smoothly, you only need to bear three things in mind:

• It needs warmth. Keepers agree that without a heat source the brood develops very slowly. A heat cable or heat mat on one side, with a thermostat, makes all the difference.
• The start calls for patience. Queens and small colonies are shy and sensitive. In the first year the colony grows slowly, and this is when water in the tube and leaving it undisturbed matter most.
• It appreciates a winter. It is not essential, but a mild diapause suits its native climate (see ‘Diapause’).

Once past that early stage, the reward is a colony that takes off, with workers of several sizes and that pale yellow that makes it unmistakable. If you are looking for another easy Camponotus from Asia, at Antomas we also have Camponotus nicobarensis, a tropical species that needs no winter.

Recommended parameters

• Nest temperature: 24–28 °C during the active season, with a warm spot of 28–30 °C on one side, always with a thermostat. The colony will settle where it suits it best: you will see the brood piled up next to the heat source.
• Why these figures: in summer, the soil in its range at a depth of 20–40 cm is at about 24–32 °C, and care sheets and keepers agree on 24–28 °C. One keeper noticed that at 25 °C brood development was slow and raised it to 25–27 °C.
• Do not heat the whole nest: in one keeping journal, a heat mat under the entire formicarium killed half the workers of a small colony. Heat should always come from one side, and never stay continuously above about 32 °C.
• Foraging area: at room temperature, about 20–25 °C. It can have a slightly warmer corner.
• Humidity: moisture in the tubes while the colony is small; later, a dry nest (see ‘Humidity: moist for founding, dry later’).
• Water: always, in a test tube with water and cotton wool in the foraging area.
• Light: nest kept dark and foraging area lit, with a normal day–night cycle.
• Diapause: recommended, for about 3 months, from December to February, at 10–15 °C (see ‘Diapause’).

Humidity: moist for founding, dry later

This is an ant from arid lands, but that does not mean the queen and her first brood can live without water. As with almost all Camponotus, at Antomas we do it in two phases:

• Founding and small colony: the test tube with its water reservoir behind the cotton wool, and the formicarium’s humidity systems filled. In one keeping journal, a queen’s brood died off one by one when the tube dried out: this is the costliest mistake at this stage.
• From around 50–100 workers: the nest can be left dry, with water and sugary liquids in the foraging area. One keeper watched his workers gnaw through the water reservoir of an aerated concrete (Ytong) nest and put it out of action, and the colony kept thriving on the water in the foraging area.
• Never waterlogged: excess water brings condensation, mould and mites. If you see droplets on the walls, stop moistening for a few days.

Recommended formicaria

For a queen with brood and small colonies, at Antomas we recommend a plaster formicarium from the Anthouse Qbik Wild range, which holds the humidity that founding requires. The Qbik Wild One P 1C is just the right size to start with:

Qbik Wild One P 1C

Once the colony has more than a few dozen workers and the nest can be kept dry, the natural next step is the Qbik Alpha range: modular, on the drier side and lighter, it lets you keep adding modules as the colony grows. Medium-sized galleries at first, which make it feel secure, and larger ones once the major workers appear. First, a QBIK ONE:

QBIK ONE

And to expand, the QBIK DUO. Expand when the nest is full of brood and workers, not before: in a formicarium that is too large, the colony uses the spare chambers as a rubbish dump. To complete the set-up:

• Mineral oil barrier around the top of the foraging area walls and a tight-fitting lid (see ‘Escapes and barriers’).
• Dry foraging area, with a little sand or fine gravel, and a flat stone or a piece of bark for them to hide under. One keeper saw hundreds of workers take shelter under a log in his foraging area.
• Water in a test tube with cotton wool, and a feeder for sugar, never on cotton wool.
• Heat on one side of the nest, with a thermostat.

Founding and first workers

At Antomas we sell it in two formats, and each starts at a different point:

• Queen with brood: a queen that has already founded her colony in a test tube, with eggs, larvae or pupae. Your job is to see her through to her first workers.
• Queen with 5–15 workers and brood: a colony that has already passed the most delicate phase. The workers look after the brood and go out to forage, and within a few weeks the colony can be moved into a formicarium.

In both cases, founding is claustral and the queen knows what she is doing. Your part is to give her the right conditions:

• Darkness, warmth and peace. Leave the tube just as it arrives, covered with a cloth or a piece of card, in a vibration-free spot at 25–27 °C, with the heat on one side of the tube, not underneath.
• Water, always. Check every so often that the cotton wool is still damp and that there is water left in the reservoir. If it runs out or the tube gets dirty, move the colony to a new one by tipping.
• Short checks. A quick look once or twice a week is enough. They are shy, and stressed queens may eat some of their brood.
• A snack now and then. Although the queen does not need it, at Antomas we recommend offering her a tiny drop of sugary liquid on a small piece of aluminium foil every week or two, never on the cotton wool in the tube, and removing it the next day. Once there are workers, also add a small piece of soft prey once or twice a week.
• Timings. Care sheets mention about 4–6 weeks from egg to worker at 27–30 °C; we have not found any measurements by keepers themselves. At room temperature it takes much longer: in one keeping journal, with a dry tube and little heat, the first worker took ten months to appear.

When should you move it into a formicarium? With about 5–10 workers, or earlier if the tube gets dirty or runs out of water. A queen with 5–15 workers can be moved as soon as she has settled after the journey.

Arrival and moving by tipping

When the colony arrives, let it rest for a couple of days in a dark, mild spot, check that the tube has water and do not move it into the nest straight away. To rehouse it, at Antomas we do it by tipping:

• Set up the formicarium beforehand, with the oil barrier in place, the water ready and, if it is a Qbik Wild, already moistened.
• Work over a tray, with a soft paintbrush to hand, in case any worker climbs up the tube.
• Remove the stopper in the foraging area, next to the nest entrance, and gently tip out the queen, the workers and the brood. Brush in any stragglers.
• Keep the nest dark and the foraging area covered. The workers will soon carry the brood inside, and the queen will follow them.

To move it later into a larger module, use the same method.

Feeding

Sugar, almost daily. It is a heavy consumer of sugary liquids, as befits a shepherd of aphids: keepers give it fresh sugar every day. At Antomas we recommend variety: Anthouse Blue Sugar nectar 4–5 times a week and another nectar or sugary liquid 1–2 times a week. You can also add a little protein to the nectar. Always offer it in a feeder where the drop stays ‘spread out’ within the workers’ reach, and never on cotton wool, which goes mouldy and spoils the liquid. Avoid deep containers: some keepers have lost workers that drowned in sugar water.

Protein: plentiful and varied. The queen needs it to lay and the larvae need it to grow. In our experience at Antomas, the more protein colonies receive, the faster and larger they grow: if you do daily maintenance and remove leftovers, it is better to give too much than too little. If you will not be able to clean often, it is better not to overfeed, because rotting food brings mould and mites. Change the type of insect from one week to the next:

• Thermobia domestica: small, soft prey, perfect for founding queens and young colonies.
• Springtails, Sinella curviseta, and fruit flies: one keeper says his colony loved the flies.
• Cockroaches: the Argentine or dubia roach, Blaptica dubia, or the red runner, Shelfordella lateralis, freshly killed and opened.
• Crickets, Acheta domesticus, cut into pieces while the colony is small.
• Mealworms, Tenebrio molitor, always cut open: whole, the same colony barely paid them any attention.

Better opened. They make far better use of cut-up prey, whose juices they drink and share mouth to mouth, than of a whole insect with a tough exoskeleton. Buy insects from a reliable supplier, free of insecticides, and do not add supplements you are unfamiliar with to the sugar: one keeper lost hundreds of minor workers after trying one containing urea.

Water: always, in the test tube with cotton wool, kept separate from the sugar.

Diapause

Its homeland has a proper winter. In the lowland parts of its range, mean air temperatures stay below 10 °C for about four months, from November to February, with frosts from December to February. Underground, at nest depth, we estimate that the soil stays at about 5–13 °C from mid-December to early March, without freezing.

In captivity, opinions vary. Shops usually recommend a rest period from November to March at 15–18 °C. And some keepers have kept it warm all year round: the author of the most complete keeping journal we have found gave it a diapause at 14–15 °C in the first winter and none in the following ones, and his colonies kept growing. That is why we consider it recommended, not essential. At Antomas we recommend a mild diapause, closer to its real winter: about 3 months, from December to February, at 10–15 °C, ideally around 12–14 °C.

• Gradual start. In November, switch off the heat and leave the colony at 15–18 °C for 2–3 weeks before lowering the temperature further. Reduce protein.
• During diapause: water always, a little sugar and hardly any protein. Disturb it as little as possible.
• Gradual return in March, passing through 15–18 °C again, until it is back in the warm nest. Resume protein as soon as you see activity and brood.
• Queens with brood and very small colonies: in their first winter, a shorter, gentler rest of 6–8 weeks at around 15 °C is better.
• Where: a garage, a cellar or a wine cooler usually works; an unheated room sometimes does not get cold enough. There is no need to go below about 10 °C.

Growth and major workers

There are no published counts of colonies of this species, and the figures given by shops range from a hundred to ten thousand workers. The best data we have come from the most complete keeping journal we have found, kept by a European keeper who followed a colony for three years:

• First year, slow. A colony that arrived with 4 workers had about 36 after just over a year, following a heat-mat mishap and a diapause.
• Second year, take-off. At around 50 workers it began to speed up: more than 400 at twenty months and, by his estimate, about 2,000 at just over two years.
• Third year: about 5,000 workers, also estimated.

These are an amateur’s estimates, not counts, but they show the pattern: a slow start and growth that takes off from about 50 workers. The first major workers appeared in that colony at fourteen months, still small, when it had around 50 workers; over time they emerge larger and larger.

Behaviour and observation

Shy when young, defensive when mature. Keepers describe it as a light-sensitive ant: small colonies hide in the darkest corner and the queen runs for the shade as soon as you shine a light on her. Over the years its character changes: in a large colony, dozens of media and major workers rush to the entrance as soon as they are disturbed, and hundreds stand guard beside it.

Clean and tidy. The author of the most complete journal describes it as a very clean and hygienic species, which makes maintenance easier.

A colourful spectacle. Watch the translucent gasters filling with nectar and the way they pass food mouth to mouth. With coloured nectar, you will see the sharing with the naked eye.

Observe without fear. At Antomas we believe ants are there to be enjoyed, observed and learned from. If you keep the nest dark and the foraging area lit, and watch them calmly, they get used to your presence.

Warning signs:

• Brood that does not develop in the middle of the season: it is almost always a lack of heat or protein.
• Brood dying during founding: check the water in the tube.
• Dead workers next to the heat source: the nest is too hot.
• Mould or droplets on the walls: too much humidity or leftover food.

Escapes and barriers

Like any self-respecting Camponotus, it climbs almost any surface, and shops warn that it needs more care against escapes than other easy species. Keeping journals record all sorts: a worker that crossed a talc barrier and an entire colony that left through an uncovered hole in the foraging area and moved into another formicarium. With this species, at Antomas we use mineral oil:

• A continuous band of mineral oil around the top of the foraging area walls. Renew it when it gets dirty.
• A tight-fitting lid and no uncovered holes: check connections, stoppers and tube passages.
• Nothing that can act as a bridge to the barrier.
• Hard materials: they can gnaw through soft materials, such as aerated concrete, and break out.

Handling and safety

It does not sting, as it has no sting at all. If it feels threatened it bites – the major workers make themselves felt – and it can spray formic acid, which smarts a little on a small wound. Nothing to worry about if you work calmly. Always handle it with tools, a soft paintbrush or soft tweezers, and never with your hands, so as not to harm any worker or, above all, the queen. One colony, one queen: do not combine queens or different colonies. This is an exotic species: it must never be released into the wild.

What not to do

• Let the queen’s tube run out of water.
• Keep it at room temperature all the time, without a heat source: it will barely grow.
• Heat the whole nest from below or keep it continuously above about 32 °C.
• Wet the entire nest or keep moistening it once the colony is large.
• Put sugar on cotton wool or in deep containers where they can drown.
• Always give the same insect, or whole, hard prey to a small colony.
• Leave food remains for more than 24 hours.
• Move it into a formicarium that is too large for its size.
• Combine queens or colonies, or handle it with your hands.

Scientific sources

This care sheet has been compiled from the scientific literature on Camponotus fedtschenkoi and its genus, including Mayr’s original description and the ant catalogues of Central and Western Asia, from climate data for its range and from the experience of Antomas and other keepers. Main references:

1. Mayr, G. (1880). Die Ameisen Turkestan's, gesammelt von A. Fedtschenko. Tijdschrift voor Entomologie 23: 17–40. German and Latin version of the original Russian description (Mayr, 1877). Scan on Internet Archive (open access)
2. Emery, C. (1908). Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes (Hym.). Deutsche Entomologische Zeitschrift 1908: 165–205. PDF on ZOBODAT (open access)
3. Radchenko, A. G. (1997). Review of ants of the subgenera Tanaemyrmex, Colobopsis, Myrmamblis, Myrmosericus, Orthonotomyrmex and Paramyrmamblis of the genus Camponotus in the Asian Palearctic [in Russian]. Zoologicheskii Zhurnal 76: 806–815. AntCat entry
4. Collingwood, C. & Heatwole, H. (2000). Ants from northwestern China (Hymenoptera, Formicidae). Psyche 103: 1–24. doi:10.1155/2000/97127
5. Paknia, O., Radchenko, A., Alipanah, H. & Pfeiffer, M. (2008). A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News 11: 151–159. PDF on ZOBODAT (open access)
6. Kiran, K. & Karaman, C. (2012). First annotated checklist of the ant fauna of Turkey (Hymenoptera: Formicidae). Zootaxa 3548: 1–38. doi:10.11646/zootaxa.3548.1.1
7. Karaman, C. & Aktaç, N. (2013). Descriptions of four new species of Camponotus Mayr (Hymenoptera: Formicidae), with a key for the worker caste of the Camponotus of Turkey. Journal of the Kansas Entomological Society 86: 36–56. doi:10.2317/JKES120516.1
8. Grebennikov, K. A. & Anikin, V. V. (2016). Ecology of the little-known ant species Camponotus fedtschenkoi Mayr, 1877 (Hymenoptera, Formicidae) in Bogdinsko-Baskunchakskii State Nature Reserve. Biology Bulletin 43: 1440–1443. doi:10.1134/S106235901610006X
9. Ward, P. S., Blaimer, B. B. & Fisher, B. L. (2016). A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinomyrmex. Zootaxa 4072: 343–357. doi:10.11646/zootaxa.4072.3.4
10. Feldhaar, H., Straka, J., Krischke, M., Berthold, K., Stoll, S., Mueller, M. J. & Gross, R. (2007). Nutritional upgrading for omnivorous carpenter ants by the endosymbiont Blochmannia. BMC Biology 5: 48. doi:10.1186/1741-7007-5-48
11. Wernegreen, J. J., Kauppinen, S. N., Brady, S. G. & Ward, P. S. (2009). One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology 9: 292. doi:10.1186/1471-2148-9-292
12. Martinez, T. & Wheeler, D. E. (1994). Storage proteins in adult ants (Camponotus festinatus): roles in colony founding by queens and in larval rearing by workers. Journal of Insect Physiology 40: 723–729. doi:10.1016/0022-1910(94)90100-7
13. LeBrun, E. G., Diebold, P. J., Orr, M. R. & Gilbert, L. E. (2015). Widespread chemical detoxification of alkaloid venom by formicine ants. Journal of Chemical Ecology 41: 884–895. doi:10.1007/s10886-015-0625-3