Bee Buzz Box – April 2020 Early forays into Two-queen Hive Management Part I – A Plurality of Queens

Alexander's Apiary of 730 colonies in 1921 operated by his son Franki

Alan Wade and Frank Derwent

It would have been rewarding to have been a New York State beekeeper at the turn of the 19th Century. Any competent apiarist could have harvested almost as much honey as she or he could extract. This bounteous beekeeping region, at present an epicentre of the SARS-CoV-2 pandemic, were then a bees’ Garden of Eden. As the editors of the redoubtable Gleanings in Bee Culture notedii:

Mr A., after his white-honey flow, and shortly following it, had another strong flow from buckwheat and goldenrod. This almost continuous flow for two or three times as long as the flow in most localities made a method or procedure possible with Alexander that was not feasible with the average beekeeper with a short flow.

Among the most successful and inventive apiarists of the day were Gilbert M. Doolittle, famous for his pioneering queen-raising initiatives, and Egbert W. Alexander – the Mr A. – who startled beekeepers by running hives with two or more free ranging queens. In the lingo of the day this proffered the epithet: ‘a plurality of queens’.

But, as we all know, bees operate with just a single queen. Any attempt to run a second queen without first removing the old queen is doomed to failure or so we learnt in Beekeeping 101. The practice of winging it and introducing a new queen without first having dispatched the old queen inevitably results in the loss of the caged queen. The cardinal rule of requeening is to never ever try to requeen a colony unless you are absolutely certain it is queen freeiii.

But Alexander was no ordinary apiarist. He first figured that running a second queen, notably without queen excluders, he would produce many more bees. Secondly, if he were to achieve this seeming impossible feat, he would get even more honey than the bumper crops he had been harvesting for years.

Strong hivesiv in Alexander’s apiary in 1907

But before we go down that path, let’s look at Alexander and his stellar career as an apiarist. H.H. Root, the sub-editor of Gleanings in Bee Culture, knew Alexander well and had published a number of his authoritative findings. So impressed was he that he collated and published articles as Alexander’s writings on practical bee culturev.

In 1904 we began the publication in Gleanings in Bee Culture, an illustrated semi-monthly magazine, a series of articles from the pen of Mr E.W. Alexander, of Delano, Schenectady Co., N.Y. For a period of nearly forty years Mr Alexander had been keeping bees in a large way, producing honey by the carload. He was regarded during a large portion of that time as the most extensive bee-keeper in Northeastern New York; but it was not till later that he began to give to the public the secrets of his success

From this treasure trove we can trace the origins of many of today’s best beekeeping practices. For example Alexander appears to have been the first to advocate introduction of mailed queens to nuclei rather than direct to large colonies. That technique is now recognised as the most reliable way to graft on a new queen to a colony. He was also a strong promoter of brood box reversal, a practice that promotes rapid expansion of colonies in mid to late spring. But before we profile Alexander’s other findings and launch into his efforts to run hives with two or more queens without excluders, we need to do a simple reality check. Left alone, hives run with just a single egg-layer. And for good reason, that is the evolved condition of the honey bee.

The single-queen hive

Most wild, as well as managed honey bee (Apis mellifera and Apis cerana) colonies are, as we are stressing, headed by a single queen. This is as just as true for other species of honey bee – belonging to the Apidae family – if not for stingless bees from the much larger Meliponae family that sometimes run with several queens. By way of example, the fugitive Apis andreniformis (one of the two dwarf honey bees), Apis laboriosa (the giant Himalayan Honey Bee) or any of the lesser known Asian Honey Bee offshoots (e.g. Apis nigrocinta, the Philippine Honey Bee) all sport a single queen. We know this because we’ve all kept bees: we all look for eggs, we all look for brood and we all look for the queen.

Nevertheless, if you have been beekeeping for some time you will more than likely have encountered a second queen, maybe a even third, when doing a thorough spring and autumn hive condition check. The explanation for this is quite simple: colonies are in the routine process of replacing their queen and both mother and daughter(s) may both be present for a short period. Occasionally the ‘plural queen condition’ may extended into months when, however, the colony will revert to its single queen state.

There is a simple take home message. These days we always check all brood frames for that possible extra queen. It has saved us losing the occasional valuable queen when requeening.

The temporary multiple-queen hive condition

So what is the queen and brood status of honey bee colonies under aberrant and transient ‘plural queen’ conditions, either during supersedure (simple queen replacement) or during swarming (reproductive queen replacement) – where large numbers of virgin queens may be present? Simpson, in referring to the swarming scenario, describes the seemingly confusing queen transition process very wellvi:

Apart from changing its queen which, in effect means changing the colony, no treatment seems to be known which, while maintaining its size, will so change the behaviour of a colony that it will not resume swarm preparations when the treatment is discontinued; nor does it appear possible to make a colony prepare to swarm, when it is not already doing so, except by crowding it, increasing its size, or changing its queen.
Note: Much the same can be said of supersedure: remove the supersedure cells and more will appear.

But hang on. There there also times when the queen stops laying, normal in winter or in times of dearth – when the bees run low on stores or there is no nectar and pollen flow. Especially with a sudden cessation in a pollen flow, the queen will immediately stop laying and, if the bees are protein deficient, they will cannibalise colony brood to preserve protein stores.

The no queen scenario

Adjudging whether a queen is actually present when there is no brood can be as perplexing to the novice as to the experienced beekeeper. However a well-seasoned apiarist will tell you that it is a serious mistake to ever assume that a colony is queen free. Under dearth conditions there are often no drones and no brood. Under those conditions a slimmed down queen can be hard to find. Like our chooks, there are times when the shed or hive is simply ‘off the lay’.

So not being able to find the old queen when a new one arrives in the mail is no excuse for simply dropping in the new queen. Message sent, message received: you will almost certainly lose her.

Colon Butlervii states the colony response to queen loss very clearly:

When a colony of honeybees loses its queen, the worker bees soon become aware of the fact that (often within thirty minutes or less) and the behaviour of the colony as a whole tends to change from a state of organised activity to one of disorganised restlessness…

Within a few hours a more definite sign of queenlessness becomes apparent, as one or more worker brood cells containing young larvae will usually have been modified by the bees to from emergency queen cells… The larvae in one of those cells is destined, all being well, to become the new queen of the colony.

Butler’s finding gives a cue to the simplest trick possible to work out if your colony is queenless or queenright. Importantly you don’t need to find the queen.

If you look and there no eggs and no young brood, simply add a well-marked frame of young brood (after checking for disease) from a neighbouring hive. If the colony immediately starts to raise emergency queen cells, you will know that the queen has gone. It’s then safe to put in that newly arrived mailing cage.

There is also the winter condition where a normally functioning colony becomes seemingly queenless. Most beekeepers are familiar with the fact that overwintered (so-called diutinous) bees are long-lived. They, both beekeeper and workers – like their queen and their owner – age remarkable slowly. Since healthy well fed workers are then neither under brood raising stress nor actively foraging, their 6-7 week summer adult life expectancy is extended to many months. Colonies operated in mid-to-high latitude locales tend to be both drone and brood free during winter but nevertheless usually remain queenright.

Enter Bernhard Möbus. He studied brood clusters, opening hives in the depths of Scottish winters to discover more of their survival strategyviii. To his surprise, he found small batches of worker brood were raised intermittently, even in atrocious weather, presumably to supplement an ever diminishing workforce. Judicious raising of small numbers of young bees can be best interpreted as improving the chances of colony expanding rapidly in spring. The workers’ role in protecting the queen during times of dearth is paramount but clearly the queen responds to feeding and can come back into lay whenever the colony needs to replenish its ever dwindling numbers.

The death knell of a queenless colony

The situation is, however, dire if the queen dies and is not replaced or if the queen exhausts her sperm store. These colonies perish once stores or worker bee numbers plummet, a condition signalled by the presence of large numbers of small drones and scattered high capped drone brood in worker comb. Laying workers will deposit eggs on the walls of worker cells, while a surviving drone-laying queen will still deposit an egg at the base of each cell. The result – a colony full of small drones and worn out old worker bees.

Under truly exceptional circumstances, colonies may survive where, quite surprisingly, they raise queens from unfertilised eggs laid by the queen or laying worker (gynacoid queen) in a process called thelytoky. It is a condition that we are unlikely to ever encounter though it is quite common amongst Cape Honey Bee (Apis mellifera capensis) colonies.

Normally, however, bees engage in a well-orchestrated process of replacing their queen. Honey bees invest vast resources in raising both gynes (potential queens) and drones (potential queen mates), a penalty of reliance of the colony on a single queen. This contingency ensures that one well-fertilised queen will remain once the failing or aged queen dies or the colony numbers explode and the colony swarms taking with it the colony queen.

Here we came back yet again to the reality that any hive that has two queens soon reverts to the single queen condition. So the adage that honey bee colonies are headed by a single queen is right on the knocker.

A plurality of queens

Alexander was never one to accept orthodoxy. Despite the maxim of one colony, one queen, he found that he could, nevertheless, establish and run two or more queens in the one colony and, very remarkably, excluder-free. How could this possibly be?

The mood for acceptance of the possibility of running an extra queen in a hive was set when Alexander first aired his finding. Such was the intrigue in the beekeeping community that A.I. Root, the then editor of Gleanings in Bee Culture, was moved to preface Alexander’s 1907 publication with a reserved but supporting statement:

This is the long-expected article for which many of our readers have been anxiously waiting. Now that it has come, some of the statements are so startling that, had they come from any less authority than our correspondent, we should feel inclined to lay it aside to think it over, if not pigeon it altogether. But instead of saying. ‘No, it won’t work’, our readers are requested to try it and report. The immense possibilities that might accrue from the use of two or more queens in one brood-nest are too great to be lightly dismissed… He records that usually but one remains in the fall.

Excerpt from E.W. Alexander’s 1907 publication ‘A plurality of queens without perforated zinc’

So the news of running two queens in a hive came not only with a mixture of scepticism and disbelief but also a nagging optimism that Alexander might have discovered a new way of keeping bees. Despite rapidly declining health and a dissenting beekeeping fraternity, Alexander was not easily dissuaded. He had time enough to demonstrate that bees could be readily induced to accept extra queensix though, had he lived longer, who knows what his findings may have been. Both Macdonald and Averyx review Alexander’s A plurality of queens, Macdonald noting that:

It is only quite recently that any bee-keeper was known to assert that two or more queens can exist in one hive, laying peacefully side by side. The subject is an exceedingly interesting one, and if successfully proved to be feasible it may revolutionise beekeeping. Gleanings has two articles on the subject. Mr Alexander—a high authority—positively asserts that he found five queens in one hive quietly performing their functions without any feeling of rivalry. Further, he gives a method by which he asserts several queens may successfully be given to any hive, and you will not lose one queen in a hundred. Mr J. E. Chambers adopts the opposite view, and maintains that although he has been using more than one queen in a hive to build up, it must be under proper safeguards—something, I take it, like our Wells dummy to divide the two domains. Mr Alexander’s claims he sets aside as impracticable, if not impossible.

To establish an additional queen Alexander used an elaborate engorgement and preconditioning technique. He then introduced mated queens to each of a series of already queenright colonies. The result was stable two-queen – or multi-queen – units, colonies that persisted with two or more queens through to the end of the honey flow.

Such hives, sensibly managed and presumably always operated with young queens, proved to be not only exceptionally productive but almost swarm-proof:

In regard to keeping two or more laying queens in a colony at the same time, and its effect on their swarming, according to our experience so far, it has wholly prevented it, as we have never yet had a colony attempt to swarm that contained two or more laying queens where each had free access to all parts of the hive. We are now wintering a colony with seven two-year-old queens in it, all loose in the cluster of bees. We saw and counted the queens a few days before putting our bees in the cellar; and up to date, January 30, we have not found any dead queens under the cluster of that colony.
Alexander’s writings on practical bee culture, pp.79-82.

His claim that such hives might be routinely maintained year round with more than a single queen was, at least in part, later retracted. That early finding may have been anomalous and is certainly neither the collective experience of later two-queen hive exponents nor our own.

While a number of beekeepers remained doubtful of successfully operating colonies containing two queens, having had difficulty replicating Alexander’s technique (see responses by Hand, by Wright, by Chambers and by Doolittlexi), he was clearly successful in running such colonies, albeit seemingly always under exceptional honey flow conditions ascribed to New York State that existed at the timexii:

He was the only beekeeper in the United States who was ever able to manage from 700 to 800 colonies all in one yard. Others have had as many as 500; but this has always been regarded as an extremely high figure for one place. Mr Alexander’s apiary was located, and is now, in fact (under the management of his son) , on one of the hills near the little town of Delanson, N.Y. It is probably one of the finest, if not the finest, buckwheat bee-ranges in the United States. From the Alexander apiary, when the fields are in bloom, one can see from twenty to thirty white patches down in the valley and on the hills, to which the bees are streaming in countless thousands. Then the locality is remarkable for the large number of asters which possibly furnish almost as much nectar as the buckwheat itself.

As the British Bee Journal recordsxiii, towards the end of his life, Alexander was offering queens for sale from his multiple-queen hives:

Bee-keepers who are interested in the higher branches of the craft are indebted to Mr Alexander for some novel and valuable ideas connected with beekeeping, notably the Alexander method of keeping a great number of queens in one hive for a long period, ready for sending to customers as ordered from each colony.

The acclaimed Everett Franklin Phillipsxiv astutely distilled Alexander’s remarkable finding:

While there is usually but one queen in the colony, it sometimes happens that two are found, usually mother and daughter at the time of supersedure… The specialization which normally permits but one egg-producing female is not well understood nor do we know why a queen usually attempts to kill any rivals (except under swarming conditions). Recently, Alexander has advocated the use of two queens for rapid upbuilding.

Here there is another practical take home message: Modern two-queen colonies, operated with queen excluders, will often persist with a second queen even after removal of queen excluders, that is until autumn or until the main honey flow is over.

Colin Butler who discovered ‘queen substance’, the glue that holds bee colonies together, also found that separation of a queen from brood resulted in emergency queen cell developmentxv. There have been many reports of establishment of a second colony queen where beekeepers have moved brood, say to a top super with a flight entrance, where the colony has nurtured the establishment of a second queen. More pointedly, Farrarxvi reported Alexander’s finding, citing Spojaxvii in Yugoslavia, who in turn had corresponded with Kovtunxviii in the Kharkov area of the then USSR. Collectively these researchers had also explored the conditions under which multiply-queened hives could be established and maintained in the absence of queen excluders confirming Alexander’s findings.

Before we leave this remarkable and largely forgotten beekeeper, it is worth noting that this inventive Alexander made a number of other findings, for example promoting the European practice of overwintering of bees in barns? In one publication, Alexander refers cryptically to a failsafe method of introducing queens, a method with the potential to dispense with queen mailing cages, the technique we usually employ when requeening:

Last summer my son Frank discovered the most practical method of introducing queens that I have ever heard of—a method whereby over 90 per cent are safely introduced and laying within 18 hours from the time the parent queen was removed.

While neither Alexander nor his son ever attempted to publish this finding, he nevertheless provided details of the method in responding to Clarence A. Hall, a beekeeper who was experimenting with this technique of queen introductionxix:

The main and most vital part of this method seems to be in introducing the strange queen to about a quart of bees that are well filled with honey after they have been taken from their parent colony a few hours, and keeping them for a few hours longer with the queen you wish to give to the colony. This gives the queen the odor of the colony, and the bees don’t seem to realise but that it is their mother queen. All the rest is but secondary to this main part.
E.W. Alexander, Delanon, N.Y., Oct. 14.

In exemplifying his claims, Alexander noted:

We can now safely introduce any number of queens to a colony that has a laying queen in a colony that has a laying queen and is in a normal condition.

The fate of the Wells and Alexander schemes?

The 1908 Franco-British Congress of Beekeepers, cognisant of both Alexander’s two-queen and Wells’ doubled hive findings, evaluated the merits and shortcomings of their schemes employing two queensxx. Their prognostications, though not entirely congratulatory, were the prelude to the commercial, and much more practical, two-queen hives that Clayton Leon Farrer, then Floyd Moeller and John Hogg were to later develop.

Despite the perceived impracticability of the Alexander and Wells schemes, reports of new and more reliable methods of operating two-queen colonies very quickly appeared. In 1908 Scottish beekeeper Medicus published clear instructions of how to set up and operate two-queen hives. This scheme, too, had limited ambit: it was constrained by the uncertain weather conditions of southern Scotland and the nature of local honey flows but was avidly practiced by Ellis for several decades. Like the Well’s system, the scheme appears to have only flourished in expert hands so it, too, was soon assigned to the dustbins of history.

Their nevertheless extremely ingenious system, running two queens in the one hive, will the topic of the second and concluding part of this series.

Running bee hives with two queens is not for the feint-hearted. The fire-ridden summer drought just past provided at least one happy dividend, 170 kg of honey from four of our hives coached to operate with that extra queen.

If you were contemplating running a two-queen hive in the manner that Alexander did, we suggest you don’t bother. Doubled hives achieve much the same purpose but you can read up on those in the December 2019 and February 2020 newsletters or simply log into the club website and find them by clicking on the Apiary link.

Readings

iApiary of Alexander (1921). American Bee Journal 41(10):385. https://ia802702.us.archive.org/19/items/americanbeejourn6061hami/americanbeejourn6061hami.pdf
Lloyd, L. (1910). Mr Alexander’s apiary of 730 colonies operated by his son Frank Alexander. Gleanings in Bee Culture 38(19):686. https://babel.hathitrust.org/cgi/pt?id=hvd.32044106180169&view=1up&seq=774

iiMarchant, A.B. (1911). Raising queens above peforated zinc: A plurality of queens for one colony. Gleanings in Bee Culture 39(7):227. https://babel.hathitrust.org/cgi/pt?id=hvd.32044106180151&view=1up&seq=267

iiiHogg, J.A. (1983). Methods for double queening the consolidated broodnest hive. The fundamentals of queen introduction, Part 1 of a two-part series. American Bee Journal 123:383-388. http://www.twilightmd.com/Samples/Hogg/Hogg_Halfcomb___Publications/ABJ_1983_1May.pdf

ivAlexander, W.E. (1907). Gleanings in Bee Culture 35(21):1376-1379. Brood rearing in the spring: How to build up the colonies rapidly: extracting sealed honey in may from the brood-nest to make room for brood-rearing: brood-combs of honey not desirable for spring feeding. https://ia902703.us.archive.org/8/items/gleaningsinbeecu35medi/gleaningsinbeecu35medi.pdf

vRoot, H.H. (1910). Alexander’s writings on practical bee culture, 3rd Edn. A.I. Root Company, Medina, Ohio, 115 pp. https://ia600206.us.archive.org/18/items/cu31924003065244/cu31924003065244.pdf
For a succinct review see Root, H.H. (June 17, 1909). British Bee Journal, Bee-Keepers’ Record and Adviser 39(1408):232.

viSimpson, J. (1958). The problem of swarming in beekeeping practice. Bee World 39(8):193-202. https://doi.org/10.1080/0005772X.1958.11095063

viiButler, C.G. (1954). The method and importance of the recognition by a colony of honeybees (A. mellifera) of the presence of its queen. Transactions of the Royal Entomological Society of London 105(2):11-29.

viiiMöbus, B. (1998). Brood rearing in the winter cluster, Part I. American Bee Journal 138(7):511-514.
Möbus, B. (1998). Rethinking our ideas about the winter cluster: Part II. American Bee Journal 138(8):587-591.
Parts I and II can be found at http://poly-hive.co.uk/recourses/mobus-bernard-his-workonswarming-and-wintering/brood-rearing-in-the-winter-cluster/
Möbus, B. (1998). Damp, condensation and ventilation, Part III: The sink, damp and condensation Part I repeated and Part III can be found at http://polyhive.co.uk/recourses/mobus-bernard-his-work-on-swarming-andwintering/ dampcondensation-and-ventilation/
The three parts were originally published in the July, August and September 1998 editions of the American Bee Journal.

ixAlexander, E.W. (1907a). A plurality of queens without perforated zinc: How the queens are introduced: The advantages of the plural-queen system. Gleanings in Bee Culture 35(17):1136-1138. https://babel.hathitrust.org/cgi/pt?id=umn.31951d00953180r;view=1up;seq=660
Alexander, E.W. (1907b). The plural queen system: All queens but one disappear at the end of the season. Gleanings in Bee Culture 35(17):1496. https://babel.hathitrust.org/cgi/pt?id=hvd.32044106180268;view=1up;seq=562
Alexander, E.W. (1907c). Queen rearing: Some questions answered concerning the age of drones: The two-queen system and other matters. Gleanings in Bee Culture 35(10):694. https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t7fr05w6z&view=1up&seq=708
Alexander, E.W. (1908). Gleanings in Bee Culture 36(18):1135. The plural two queen system: Is it profitable for bee-keepers to practice it? https://babel.hathitrust.org/cgi/pt?id=uc1.b3458203;view=1up;seq=357

xMacdonald, D.M. (1907). American and colonial papers: Extracts and comments. British Bee Journal and Bee-Keepers’ Adviser 35(1323):437-438.
Avery, G.W. (1907). The quiet season: Some retrospective bee notes. British Bee Journal and Bee-Keepers’ Adviser 35(1323):443-444. https://ia600201.us.archive.org/18/items/britishbeejourna1907lond/britishbeejourna1907lond.pdf

xiHand, J.E. (1907a). A season’s work with sectional hives: Swarm control and comb-honey production. Gleanings in Bee Culture 35(10):712-713. https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t7fr05w6z&view=1up&seq=726
Hand, J.E. (1907b). A season’s work with sectional hives: Swarm control and comb-honey production: How to find queens without handling frames: The two-queen system of honey production. Gleanings in Bee Culture 35(15):843-847. https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t7fr05w6z&view=1up&seq=857
Hand, J.E. (1907c). The plural queen system: No problem to introduce a number of queens to bees, but difficult to introduce them to each other. Gleanings in Bee Culture 35(21):1385-1386. https://babel.hathitrust.org/cgi/pt?id=umn.31951d00953180r;view=1up;seq=813
Bailey, W.M. (1907). The two queen system: Distances bees can fly: Caucasians short lived. Gleanings in Bee Culture 35(22):1447. https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t7fr05w6z&view=1up&seq=1463
Chambers, J.E. (1907a). Two or more laying queens in one hive: The plan not a success in the hands of the average bee-keeper. Gleanings in Bee Culture 35(17):1146-1147. https://babel.hathitrust.org/cgi/pt?id=umn.31951d00953180r;view=1up;seq=670;skin=mobile
Chambers, J.E. (1907b). Double queen colonies: More than two queens in one hive not a success, and these two must be kept separate. Gleanings in Bee Culture 35(17):1582.
Titoff, A. (1907). The plural queen system; Some doubts expressed as to the benefits derived from such a plan. Gleanings in Bee Culture 35(20):1328-1329. https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t7fr05w6z&view=1up&seq=1342
Wright, A.J. (1907). The Alexander plan of building up weak colonies, and a modification of it: Two queens in a hive as a method of preventing swarming. Gleanings in Bee Culture 35(21):1886. https://babel.hathitrust.org/cgi/pt?id=umn.31951d00953180r;view=1up;seq=814
Anon. Conversations with Doolittle (1908). Plurality of queens: When profitable and when not. Gleanings in Bee Culture 36(21):1307-1308. https://babel.hathitrust.org/cgi/pt?id=uc1.b4243668;view=1up;seq=1293 and https://babel.hathitrust.org/cgi/pt?id=uma.ark:/13960/t84j0sg9w;view=1up;seq=1325
Hand, J.E. (1908). The plural-queen system: Why is it more practicable with a division board-chamber than with an ordinary full-depth hive. Bee World 36(1):35-36. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=49
Whitney, W.M.M. (1908). The plural-system: A protest against the plan: Time and money should be spent on breeding better queens than in striving to make a lot of poor queens live together. Bee World 36(1):36-37. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=50
W.K.M. (1908). The Wells system. Gleanings in Bee Culture 36(3):203. https://babel.hathitrust.org/cgi/pt?id=hvd.32044106180250&view=1up&seq=117
Hand, J.E. (1908). The two-queen system: This plan makes it possible to keep the brood chamber packed with brood during the flow; Forcing honey into the supers: Wintering two queens in one hive not desirable. Bee World 36(3):155-156. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=167
Bussy, E. (1908). The plural queen system: A series of interesting experiments: Clipping the queen’s stings so they can’t kill each other: Do the bees take a hand in royal combat? Bee World 36(3):156-157. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=168
Gray, J. (1908). The plural-queen system. How an English expert looks at the question: The advantages and disadvantages. Bee World 36(3):157. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=169
Hand, J.E. (1908). The dual and plural queen systems: Conditions under which the may be used: A review of the whole question. Bee World 36(8):507-508. https://babel.hathitrust.org/cgi/pt?id=uc1.b3458202;view=1up;seq=517

xiiRoot (1910). loc. cit.

xiiiEds BBJ (Dec. 17, 1908). Death of Mr E.W. Alexander. British Bee Journal, Bee-Keepers’ Record and Adviser 36(1382):505.

xivPhillips, E.F. (1915). Beekeeping: A discussion of the life of the honeybee and of the production of honey., 520pp. The Rural Science Series, Bailey, L.H. (Ed.) The MacMillan Company London: MacMillan & Co., Ltd. https://victoriancollections.net.au/media/collectors/51d110e42162ef12e06aa06b/items/5341f5212162ef0a845d5a79/item-media/5341f6e42162ef0a845d648d/original.pdf

xvButler, C.G. (1954). The method and importance of the recognition by a colony of honeybees (A. mellifera) of the presence of its queen. Transactions of the Royal Entomological Society of London 105(2):11-29.

xviFarrar, C,L. (1953). Two-queen colony management. Bee World 34 (10):189-194. https://doi.org/10.1080/0005772X.1953.11094821

xviiSpoja, J. (1953). Observations on the operation of multiqueen colonies. Bee World 34(10):195-200. https://doi.org/10.1080/0005772X.1953.11094822

xviiiKovtun, F.N. (1949). How to make and use multiple queen colonies. Pchelovodstovo 26(9):29-30. Cited by Spoja (1953) and by Latif et al (1960) loc. cit.
Kovtun, F.N. (1950). Letter to the editorial office (multiple-queen colonies). Pchelovodstovo 27(2):112. Cited by Spoja (1953) loc.cit.

xixHall, C.A. (1908). Introducing queens: A modification of the Alexander method. Gleanings in Bee Culture 35(24):1592. https://babel.hathitrust.org/cgi/pt?id=umn.31951d00953180r;view=1up;seq=930

xxEditorial, Notices, &c. Franco-British Congress of Beekeepers (1908). British Bee Journal, Bee-Keepers’ Record and Adviser 36(1363):311-313.

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