Jerzy Woyke (Figure 1) was born in Malenin, Poland in 1926. He gained his MS degree from the Forestry and Agriculture Faculty of the University in Poznan in 1950. He gained a second MS degree from ...
The free living giant bee Apis dorsata migrates seasonally, but the biology of migrating swarms is unknown. A recent publication suggests, that arriving giant honey bees use wax specks from a previous season to recognize the nest sites. To verify this, I investigated the behavior of individual worker bees arriving to the nesting site. I video recorded 13 arriving swarms. I found that at first arrived the scout bees and next the swarm bees. Those bees arrived to two types of nesting places. One was to places where there were no wax specks were present, and the second was where wax specks from a previous season were present. However, the arriving bees were not interested in wax specks from previous seasons, suggesting that wax specks from previous nests are not necessary to attract arriving swarms and in places where wax specks are present, the arriving bees are not interested in them. An active nest of A. dorsata consists of a comb with brood and honey, and a protective curtain covering the comb by 3–6 layers of bees. The bees of the curtain do not sit on the comb. Numerous top worker bees create special hanging wax constructions namely the wax specks. The top bees hang below, holding the wax specks with their first pair of legs. The bees of the curtain hanging below hold the bees hanging from the wax specks. Thus, the bees create the wax specks as hanging constructions for bees of the protective curtain.
A detailed description of the behavior of eight arriving swarms of the giant honey bee Apis dorsata is presented. The scout bees from swarms select the proper settlement site for arriving swarms. It seems that the scouts examine the solidity (stability) of the support and accept a cleared not rotten area between the traces and remnants of former combs from previous season. Aggression between different scouts appears probably to be due to competition between scouts from different swarms for settlement places. The scouts run on the selected area. Possibly they show their potental properity. The number of scouts increases, but just before arrival of the swarm, the number of scouts decreases considerably. The arriving bees of the swarm do not congreate at once in a cluster, but they compose several curtains. The edges of the curtains are in a form of zigzag, and they appear as a kind of garlands. When the garlands start to unite, a structure similar to human face appears. Sometimes several small faces are visible. After short time the face(s) disappear(s) and only small round depressions of 3.5 cm diameter remain on the surface of the cluster. Next the depressons disappear, and the surface of the cluster becomes smooth. It takes about 1: 00 h from the time the first scouts are noticed until the final shape of the cluster is formed. This the first description of curtains, garlands and faces occurring during the settlement process of A. dorsata swarms to nesting places.
This paper emphasizes the topics concerning honey bee (Apis mellifera L.) mating biology, which have not been described in the recently published book of Koeniger et al. (2014). At the beginning of natural mating, the drone becomes paralyzed. However, the muscles in the abdomen continuously contract shrinking the abdomen till mating has ended and the pair have separated. It is not the queen that ends the nuptial flight. The termination of the nuptial flight is determined by the drone, which fails to remove the mating sign of the previous drone from the sting chamber of the queen. The mating sign originates from two or more drones. The queen also does not determine the age at which she starts oviposition. It is the last drone, which tried to mate, but failed to remove the mating sign of the predecessor, that determines the age that the queen starts oviposition. The book of Koeniger et al. (2014), together with this paper, present the current knowledge of the mating biology of honey bees.
Over a 37-year period, we observed 1011 active bee nests and abandoned combs of Apis dorsata and Apis laboriosa in Nepal, India, the Philippines, and Bhutan. This article focuses on the reasons for the different shapes of the nests. We discovered that differing ambient conditions were the reasons for the following three shapes of symmetrical nests: vertical semi-ellipse, semicircle, and the horizontal semi-ellipse. We noted that asymmetrical nests were constructed when there was lack of space to extend the comb equally in both external directions. An asymmetrical nest also appeared when remnants of a previous comb remained on one edge of the nest. Convex nests were constructed to avoid excess sun exposure. Concave nests appeared as a result of low temperatures during the night and part of the day (Nepal). An L-shape nest was constructed when there was lack of space available to extend the nest in a straight direction. The shape of the nests also determines the way the combs fall.
The giant honey bees Apis dorsata and Apis laboriosa construct nests of different shapes and sizes. A method was required to compare the shapes and sizes of these nests. We developed a nest shape index (NSI). It presents the ratio of the dimension of the upper horizontal attachment of the nest base (B), to the vertical dimension of the nest (V); NSI = B/V. We examined the NSI of A. dorsata nests. The results varied; NSI = 0.6–4.7. The shape of nests under a sloped support is characterized by the inclination index (II). This index presents the ratio of the vertical dimension of the nest (V) to the dimension perpendicular to the sloped base (P). The inclination index of the nests observed by us, varied; II = 1.2–3.0. The mean shape of A. laboriosa was NSI = 0.96. The difference between the NSI of A. dorsata and A. laboriosa is due to environmental conditions. These indexes make it possible to compare the shapes of different sized nests when real nests were measured or from measurement of nests in photographs. Knowing the indexes, one may assume in which environmental conditions the nests were constructed, and may draw and imagine the shapes of nests without ever having seen them. A nest in which the NSI = B/V = 2 is a semi-circle, a nest in which the B/V < 2 is a vertical semi-ellipse, and when B/V > 2 the nest is a horizontal semi-ellipse.
SUMMARYWild eye colour can be distinguished from brick, chartreuse and buff mutants: from 4 days before pupation by the presence of a pink rim to the compound eyes, and on the first day after pupation by the presence of pink dots in the compound eyes and by the colour of the ocelli. Brick mutants of both sexes are distinguishable on the third day after pupation, chartreuse on the fourth day in workers and the fifth day in drones: buff mutants start to colour on the sixth day in both sexes. The ocelli of wild bees are almost the same colour as the compound eyes, whereas in the mutant phenotypes they are colourless or white-grey.Eye colour is thus a good genetic marker in early development stages. On the other hand cordovan body colour can be distinguished from the wild type only one day before emergence from the cell, and is thus not useful.
SummaryThe morphology of the genitalia, and the mating behaviour of Apis cerana indica from Peshawar, Pakistan, are described. The morphology of the vagina of the A. cerana queen, in particular the open val vefold, makes instrumental insemination easier than in A. mellifera. In the drone, both the quantity of semen and the concentration of spermatozoa in it are lower than in A. mellifera. In the seminal vesicles about 1·5 million spermatozoa were found per drone, and in the ejaculated fluid about 1·0 million per drone.One queen, dissected immediately after a second mating flight, had a quantity of semen in the oviducts corresponding to that from 14 drones, so on both flights together this queen may have mated with some 30 drones. A mating sign was observed on newly mated queens, but it lacked the chitinized plates found in A. mellifera.It is concluded that, to ensure full insemination, A. cerana queens must mate with more drones than is necessary for A. mellifera queens. Evolutionary trends in the genus A...
The aim of this study was to develop an effective method of overwintering reserve honey bee queens in two-storey mini-plus mating nuclei and in 3-comb nuclei (frames 36 x 26 cm, Wielkopolski hive). The assay was performed during three wintering seasons (2005 - 2008) parallel at two centers in Poland: the Division of Apiculture at the University of Life Sciences (SGGW) in Warsaw, and the Apiculture Division at the University of Warmia and Mazury (UWM) in Olsztyn.The results showed that 59% of queens overwintered in mini-plus nuclei and 77% in 3-comb nuclei. Among queens in mini-plus nuclei 63% overwintered in bee yard and only 55% in cellar. Within queens in 3-comb nuclei, 62% overwintered in Olsztyn and 91% in Warsaw. The highest survival rate of 93% was observed in Warsaw during the first season. Due to low survival rate, it is not recommended to overwinter the queens in miniplus nuclei.
In bee colonies without open brood, e.g., after swarming, there is no need for royal jelly, and nurse bees thus do not produce it. According to many beekeepers, adding combs with open brood restarts the production of royal jelly by nurse bees, and the virgin queens then are better fed and start earlier oviposition. The purpose of this study was to investigate whether the presence of open brood and the strength of the colonies affect the onset of oviposition by queen bees. Open brood in colonies with virgins before and during mating flights did not accelerate the initiation of oviposition by the queens. In addition, no differences were identified in starting oviposition by queens in strong colonies of more than 30,000 worker bees, or in weak colonies with up to 1,000 workers. Overall, the results showed that neither open brood in the nests, nor the strength of the colonies affects the onset of oviposition by queen bees.
The viability of spermatozoa is a crucial parameter to appreciate semen quality and insemination potential of males both in natural mating and instrumental insemination. Here, we conducted a step-by-step investigation to address the questions why and at which step(s) the viability loss is occurring in spermatozoa of honeybees during natural mating and preparation for instrumental insemination. We detected the viability of spermatozoa in semen samples obtained from seminal vesicles and partly and fully everted endophalli of drones and in ejaculates collected into syringe tips, as well as the viability of spermatozoa in lateral oviducts of queens returning from the mating flight. A great diminish of spermatozoa viability (~10 %) was found in lateral oviducts of queens returning from mating flight (88.7 %) in comparison to viable spermatozoa in intact seminal vesicles of drones (98.1 %). Our results demonstrated that the decrease in spermatozoa viability occurs during the second stage of eversion of endophallus (viability loss, 3.3 %), and during injection of semen into the lateral oviducts of queens (viability loss, 6.1 %). The acting factor decreasing the viability of spermatozoa was the increased pressure occurring during the process of natural and instrumental insemination.
The performance of bee colonies greatly depends on the quality of the queens. The current research was conducted at the apiary of the Faculty of Agriculture, Zanjan University, in Zanjan, Iran. Together, 24 rearing colonies were assigned to 4 grafting larvae age groups: 1-day-old, 2-day-old, 3-day-old, and emergency queens. Two feeding groups, fed and not fed, were created. The effects of reared queens on biological characteristics and performance of honeybee colonies (Apis mellifera meda) headed by those queens were measured. Age of grafted larvae significantly influenced the results. The performance ratios of the most efficient colonies headed by queens reared from 1-day-old larvae compared with the least-efficient queens reared from 3-day-old larvae were 118% in brood production, 140% in bee population, and 154% in honey production. However, the age of grafted larvae did not affect colony defense behavior. Supplemental feeding of rearing colonies increased brood production to 111%, bee population to 116%, and honey production to 115%. A combination of the effect of age of larvae and supplemental feeding resulted in twice as much honey (12 kg) produced by colonies with queens reared from 1-day-old larvae in fed rearing colonies compared to those with queens raised from 3-day-old larvae in unfed rearing colonies.
Honey bee queens are highly polyandrous and mate in flight. Instrumental insemination is an essential tool that provides complete control of honey bee mating for research and breeding purposes. The technique requires specialized equipment to anesthetize and immobilize the queen and to collect and deliver semen from the drones. Semen is harvested from mature drones by hand eversion of the endophallus and collected into a syringe. The queen is placed in a chamber and anesthetized during the procedure of insertion of semen into the oviducts. Queens are introduced into colonies and their performance can equal to that of naturally mated queens, given proper technique and care.
Swarming and Migration ofApis dorsataandApis laboriosaHoney Bees in India, Nepal and BhutanThe migratory open air nestingApis dorsataandApis laboriosahoneybees migrate at least twice a year. DNA genotyping showed that the same swarms return to their natal nesting sites. We examined 23 nesting sites in Nepal, India and Bhutan, on which 587 colonies ofA. dorsataandA. laboriosanested. The results showed that the frequency of the periodic mass flights (PMF) performed by the colonies is a good indicator of the status of current colony performance. During the swarming period, both,A. dorsataandA. laboriosaissue several swarms. In some colonies, so many bees swarmed out, that those remaining in the maternal colonies did not cover the combs. After the rest of the brood emerged, all the bees of such colonies abscond during the swarming period. Thus, absconding appeared in results of total out swarming. The swarms do not migrate directly to the seasonal alternative nesting sites, but establish new colonies in the areas around. After environmental conditions deteriorate, all the bees with their queens abscond and migrate to alternate seasonal nesting sites. The next season, the swarms do not return to their original reproductive natal sites, but to those sites they occupied the previous season lately, where from they absconded.
Apis laboriosa and Apis dorsata are closely related migratory open-air nesting bees which nest at mountain cliffs. It is not known why they select certain cliffs over others. We examined 16 nesting cliffs, at which 258 A. laboriosa colonies nested. The studies were conducted in Nepal in 1998 and 1999 and in Bhutan in 2008. Most cliffs are of a grey to black colour, but all A. laboriosa bees selected as nesting sites light-coloured unweathered cliffs. The light cliffs appear as a result of water erosion, and the unweathered cleared surface assures strong fixation of the combs to the rock. This way the nest will not tear away from the cliff overhangs. Near some nests, a white area is present on the vertical wall of the cliff. This is the surface cleared by bees, which previously nested here. It is possible to spot actual or potential nesting cliffs of these bees from far away.
The research was conducted at the apiary of the Faculty of Agriculture, Zanjan University in Zanjan, Iran. Queens were reared in 24 Apis mellifera meda honey bee rearing colonies. The colonies were assigned to 4 grafting larvae age groups; 1 day old larvae, 2 day old larvae, and 3 day old larvae, and the last group reared emergency queen cells. The groups were divided into the 2 feeding groups: fed additionally and no fed. The effects of the age of the grafted larvae and the effects of supplemental feeding on 9 morphological characteristics of queens were measured. The results showed that the age of the larvae significantly affected the morphological characteristics of reared queens, and thus, their quality. Queens reared from 1 day old larvae were of the highest quality. These queens were significantly heavier (158.83 mg) and had significantly larger spermatheca (0.99 mm(3)) than queens reared from larvae 2 and 3 days old. Queens from emergency queen cells were of lower quality than queens reared from 1 day old larvae. However, queens from emergency queen cells were of higher quality than queens reared from 3 day old larvae. The supplemental feeding significantly increased most morphological characteristics of the reared queens. The different ages of the larvae did not significantly affect the wing length nor did supplemental feeding affect the wing length.
The hygienic behaviour of eight A. florea honey bee colonies was investigated in Thailand. Sealed brood was deep frozen or killed using entomological pins. The pins were 0.2 or 0.7 mm in diameter. The removal of killed brood by the bees was checked in intervals ranged from 24 to 60 h after it had been presented to the colonies. Almost eighty nine (89%) of the freeze-killed brood was removed by the bees within 24 h after being introduced into the combs, while 95% of the pin-killed brood was removed within 48 h after introduction. Thus, the A. florae colonies could be recognized as hygienic honey bees.
A study was carried out to investigate the effect of the original strengths of honey bee colonies supered in different ways, on the production of honey by Apis mellifera bees in the Terai region of Nepal. Bee colonies of three different original strengths, in which the bees covered 5, 10 and 20 combs, were supered in three different methods. The results showed that honey production was highly correlated to the number of worker brood cells in the colonies (r = 0.96, p = 0.003). Colonies of 5 comb initial strength (CIS), as farmers' practices in Nepal, produced the lowest amount of honey (30.1 kg per annum). Bees in colonies of 10 CIS with a deep super, produced twice as much honey (62.2 kg), and colonies of 20 CIS with deep supers produced even significantly more honey (74.5 kg). However, the relationship between the financial values of the produced honey to the cost of its production was the highest - 1.52 : 1 for colonies of 10 CIS with a deep super. Therefore, this bee colony management is recommended to the beekeepers in the Terai region and lower hills of Nepal. This finding has global application.
Summary Queens return from successful mating flights with the mating sign, which consists of chitinized plates of drone endophallus filled with mucus. The orange membranes covering both sides of the mating sign do not touch the hairy rim of the bursa copulatrix. The thin thread at the end of the mating sign is pushed forward on the surface of the sign in most queens. This makes the end of the sign look blunt. Simulated stages of natural mating showed that it is impossible for the orange membranes covering the cornua of the endophallus to become stuck to the mating sign of the same drone. Mating signs with additional pairs of orange membranes were found. In some queens, the semen or additional second mating signs were found at the end of the sign protruding from queen's sting chambers. Those three additional parts originated from drones, which attempted to mate, but failed to remove the mating sign of the predecessor. The thin thread at the end of the mating sign was pushed to the surface of the sign, because the next drone attempted to mate, but failed to remove the sign of its predecessor. The last drone, which failed to mate, left the orange membranes of his endophallic cornua at the mating sign of his predecessor. Thus, the mating sign originates from two drones. The chitinized plates and the mucus originate from one drone, but the orange membranes originate from the last drone/s, which attempted to mate, but failed to remove the sign of his predecessor.
Changes in light intensity effect flight intensity of honey bees. The ash cloud of the volcano Eyjafjallajokull covered the sky over Poland from 17 till 20 April 2010. We investigated whether the volcano cloud affected the flight intensity of foraging worker bees. We determined the flight intensity of bees during the two days of 17 and 20 April when the sky was covered by the cloud, and during the two days of 28 and 29 April when the cloud disappeared. We did seven counts on the number of bees returning to each of the 10 colonies studied. Each count lasted 5 min. The counts were done on the hour, from 10:00 to 16:00. The frequency distribution of the number of flights/5 min performed by bees differed significantly between different colonies, as well as between different days. High significant correlation was found between the number of combs covered by bees in different colonies and the flight activity. No significant correlation was found between the number of brood combs in the colonies and the flight activity. The daily mean number of flights/5 min performed by bees, differed significantly between different colonies, as well as between different days. The number of flights differed significantly on each of the four days. The flight activity decreased by 9%, during the first two days when the volcanic ash cloud covered the sky.