Bee behaviour module 6

Question 1

1 Define Polyethism

Answer 1

Functional specialization by caste (morphology) or by age in an insect colony leading to division of labour

Question 2

1. List the conditions that
   cause variations in polethestic duties

No No Super Ancestry Weather Disturbance

No no super AWD

Answer 2

• Shortage of pollen / stores
• Nosema
• Supercedure/swarming
• Genetic variation
• Cold / wet weather / winter
• Disturbance – eg hive manipulation

Question 3

Nas 9ODA 9HDA(holds swarm)

Name four pheromones

Answer 3

Nasonov, enables workers to recognise the queen, mark locations and attract drones to queens during mating flights. also used for orientation, particularly at hive entrance
Queen Mandibular pheromone, basis of queen substance 9-oxodec-2-enoic acid (9-ODA) which when distributed throughout colony prevents swarming and workers laying.
9-hydroxydec-2-enoic acid (9-HDA), attracts drones at long distanceand helps hold a swarm together.
Worker Mandibular gland, creates alarm pheromone, used by guard bees to ward off intruders (2-heptanone)
Alarm pheromone from sting gland, (isopentyl acetate) attracts other bees to sting

Question 4

1. List glandular development by age and production

Answer 4

• 3 - hypopharyngeal gland - brood food and royal jelly
• 3 - mandibular gland - brood food
• 12 - wax gland - wax flakes
• 16 - hypopharyngeal gland shrinks - produces invertase and glucose oxidase,
• 12+ mandibular gland - alarm pheromone (2-heptanone)
• 18 - sting gland (Isopentyl acetate)
• Nasonov increases with age; 28 = max;

Question 5

1. List worker duties between emergence in April until death

Answer 5

• 1-3 Cleaning cells, gorging on pollen
• 3-15 feeds larvae and tends queen
• 16-20 receives nectar ripens nectar into honey, packs pollen
• 10-18 makes wax, builds comb,
• ventilates, evaporates, controls temperature (homeostasis)
• 12-25 days guard duty, orientation flights
• 3-6 weeks foraging

Question 6

1. List worker duties between her emergence in OCTOBER until her death

Answer 6

• No income / forage
• Drastically less brood to feed and almost no hive duties
• External temp low, metabolic rate low, so food consumption low
• Focus is on heat conservation in cluster at 20-30C
• Keeping the colony warm, either through creating a denser cluster and/or shivering her dorsoventral muscles to generate metabolic heat
• Around Jan/Feb, as the queen starts laying again, raises temp 33-36C.
• Gradually pick up her duties, limited by the weather & availability of forage

Question 7

1. Orientation - describe the theory behind

‘move a colony less than 3 feet or more than 3 miles’

Answer 7

• Young bees take orientation flights noting local landmarks - bushes
• Successive flights get longer (half a mile)
• By foraging age, it knows local half mile really well
• Foraging bees will follow old routes home automatically
• Move hive -3’ / cut long grass / move landmarks: homecoming bees disorientated for a few hours until they adjust
• Move 3’+ and they return to old site and ‘wait’ for hive to appear
• Move -3 miles, forager may come across old flight paths and revert
• Move 3+ miles, = unlikely to cross old paths so reversion less likely
• Memory lost after about 2 weeks

Question 8

2 Describe colony mating behaviour 1

Answer 8

• Workers bees do not mate
• Only queens and drones, and they do so on the wing

Question 9

2 Drone mating - maturation

Answer 9

• Emerges after 24 days
• from colonies with 6000+ bees in April
• 14 days to mature while sperm finishes migrating to seminal vesicles
• Congregate on edges of frames feeding on honey
• Orientation flights followed by flights to DCA during warmest part of day

Question 10

2 Drone mating

1 attraction

Answer 10

• Attracted to Q by
  
  • Pher: Mandib (9-ODA) - upto 50m away
  • Pher: Renner Baumann (tergite) - 30cm
  • Sight: sees open bursa copulatrix at 1m
• Comet of D forms with strongest getting to her first

Question 11

2 Drone mating - 2 act

Answer 11

• Clasps Q from behind with all legs
• Bends abdomen, everts his endophallus into bursa copulatrix with violent contraction
• Q cannot release it.
• Becomes ‘paralysed’, loses hold and swings backwards
• Carried along by the queen as semen is ejaculated into the median and lateral oviducts
• Endophallus ruptures as D falls away, drops to the ground and dies
• The bulb remains in bursa copulatrix - emits UV light - attracts more D
• Mucus from the drone’s mucus glands coagulates and forms a seal “mating sign”
• Next drone removes mating sign

Question 12

02 mating

How many drones does a queen typically mate with

Answer 12

• 12-15

Question 13

2 Q mating - 0 maturation

Answer 13

• Emerges after 16 days and eliminates rivals in cells/fights
• 1-4 remains in col feeding herself honey - no RJ and no court
• Exoskeleton hardens; weather permitting takes orientation flights
• Exocrine glands develop esp mandibular: 9-ODA for D attraction
  
  • at 5 days, enough to attract D. Increases over 10 days
• After 5 days she is ready to take mating flights to DCA
• Workers ignore her at first
• When mature, ++ aggresive as she ages without mating (stale prevention)
• After 21 she cannot mate -> drone layer
• If due to inbreeding she produces diploid males, workers eat eggs - mechanism designed to limit drone numbers.

Question 14

2 Q mating - act

Answer 14

• Mating flights av 30 minutes.
• Between 12-4pm ideally at 20C +.
• Av 2-3km to DCA. At DCA attracts a comet of D.
• Q cannot release D once copulation starts.
• Mated on wing by several D, each removing the mating sign of the last
• Returns to the hive where workers remove last mating sign
• 2-3 mating flights (up to 20D) until her spermatheca is full of sperm.
• Forces the sperm from lateral and median oviducts to her vagina
• Sperm enters spermatheca via spermathecal duct: chemotaxis migration - rest lost but all drones represented
• Workers attend her, feed her royal jelly and form a court
• 3-4 days later: starts laying, maybe haphazardly before settling into gd pattern
• She never leaves the hive again, except to swarm.

Question 15

2. Drone - Describe a DCA 10

Answer 15

Definition: A discrete aerial site where drones fly anticipating the arrival of virgin queens.

1. Airspace 15-25m above ground where D congregate and fly around indep of Q(s)
2. 100m from apiary
3. Open/hilly ground sheltered from wind if poss
4. Magnetic attraction? - magnetite in trophocytes in gut
5. Mandib gland attracts Drones and Q
6. Area 200 x 100m
7. Attracts 12 to 10k drones from 5-6km radius (av 1km)
8. Minimises inbreeding
9. Mating height 10-40m av (can be less) inversely propotional to wind speed

Question 16

03 Egg laying - 2 rate

What drives the egg laying rate?

Answer 16

• Rate of egg-laying driven by how much Queen is fed
• Thus an artificial flow (feeding mimics a nectar flow) can accelerate egg laying in spring
  
  • Swarm prep - feeding drops off to prepare the queen for flying
  • Nectar flow end
  • Dearth – eg June gap, August gap
  • Winter as nest temperature falls below 33-36C to 20-30C

Question 17

03 egg laying - 3 brood pattern

Describe a good brood pattern

Answer 17

• Roughly spherical nest to make it easiler to keep warm in cluster
• Densely clustered concentric circles
• Placed in groups at the centre of the frame, but not nec of brood box
• Frames on edges of next tend to have less brood than central frames
• One egg per cell, placed upright and centred at the bottom of a cell
• Drone brood grouped on bottoms/sides of worker brood
• Some empty cells possibly to facilitate warming
• W shape of empty cells over wires
• Uses front legs to assess whether a cell sis W/D. Amputated leg tips -> haphazard laying

Question 18

3 Egg laying - Describe a typical season’s laying pattern

Answer 18

• Jan/Feb egg laying, but may be earlier if the weather warm
• Spring - queen’s laying rate governed by how much food is available.
• Drone eggs laid in early April in preparation of swarming (May/June)
• More forage (especially pollen) = more eggs
• Up to 1500 per day by late May
• June gap - may slow down. Some strains continue in dearth (Mediterranean)
• Autumn, younger queen may continue laying later into year than older queens, maintaining egg laying after a nectar flow has ended
• Ivy forage Sep-Oct stimulates egg laying, dep on weather and forage
• Late autumn egg laying drops right back
• November/December, likely to be off-lay in colder areas of the UK

Question 19

3 egg laying - 4

Why is a good brood pattern essential?

Answer 19

• In cold, the cluster can maintain the temperatures in the brood nest
  
  • can’t if brood is spread out all over the place
• More efficient for nurse bees to work areas of brood of the same age
• Workers store pollen in an arc around the brood nest so nurse bees don’t have far to go to reprovision, with honey above the nest area

Question 20

3. Egg laying - why does a queen inspect cells before laying?

Answer 20

• To see if it is clean enough – she is very picky
• To see if it already contains an egg/larva
• To check the width – worker /drone cell

Question 21

4. Season var -

What is the impact of a swarm in Jun on honey harvest

Answer 21

1. Depends on strain of bee, forage and weather
2. The harvest will be impacted adversely because, effectively, at least a month of foraging has been lost.
3. 60% of bees will have ghone with the old queen
4. Of those that remain, some will have to come off foraging duties and revert to being nurse bees
5. Meanwhile the queen will only start laying, at best, 20 days after emerging.
   
   1. Q=8 days to emerge, 4 days to mature, 4 days to mate (assuming perfect weather), 4 days to start laying, 21 days for new bees to start emerging.
6. However, it is likely that the bees will be able to store enough to survive the winter as they have the whole of July with the blackberries to feed on.

Question 22

4. Seasonal variation - what is the impact of the month bees swarm

Answer 22

• A swarm in may is worth a load of hay.
  
  • All ok for remaining bees
  • but may lack drones to mate new Q so Q may fail next spring from lack of sperm
• A swarm in June is worth a silver spoon
  
  • Temperatures more stable and drones mature.
  • Plus plenty fo time for new hive to establish create stores for winter.
• A swarm in July is not worth a fly
  
  • Too late in the year that the swarm lay down enough honey for winter.
• A swarm in August is worth a bale of sawdust
• A swarm in September is a swarm to remember

Question 23

4. Seasonal var

Answer 23

• Title: Av pop cycle over a typical year dep strain, forage, climate, weather
• Lowest adult point end Feb as winter bees die off
• Brood = adults twice that year
• Brood > adults Feb-April - critical period: brood risks being chilled as not enough adults to incubate
• Adults peak in June three weeks after brood peaks (when main flow has started and max foraging force req)
• Pop decreases rapidly as forager bees die off then slows w winter bees
• Pop max 40k-60k dep on fecunidity and strain of Q
• Pop builds in spring with flow (little stored)
• At max pop, they store large amounts for winter in a short time (less brood to care for)
• Reduced pop allows adequate reserves for winter
• Allow for local variations eg peaky graph

Question 24

4. Seasonal var pop 2

Answer 24

• The graph represents the amount of brood in a colony that issues a prime swarm at the end of May.
• x axis is the months of the year Jan to Dec
• Y axis is the amount of brood in 10,000 gradients.
• A Queen started to lay.
• B prime swarm issued.
• D New queen started to lay.
• F queen stopped laying.
• Shaded area. The bees emerging in this area will be winter bees
• Examining the colony at this point one should see:
  
  • No eggs or larvae but some brood.
  • Several swarm queen cells on the bottom and side edges of the comb.
  • Plenty of workers going about their business foraging and bringing in nectar.
• If the colony is inspected at point C: there may be a virgin queen in the colony and she may be on a mating flight. Opening up the colony may disrupt her return to the colony.
• The risked is reduced by having patience. Wait until you see pollen being taken into the colony, a sure sign that the queen has started to lay.

Question 25

4. Seasonal var pop 3

Answer 25

• I= first QC sealed and issue of the swarm
• 2= new queen hatches
• 3= new queen starts laying
• X= months from Man
• Y=000s of bees
• Pink= adult bees
• Blue= brood
• Dotted= how adult population would grown without a swarm

Question 26

05 colony defence mechanisms

Describe the many ways a colony defends itself from ‘attack’ - 10

Answer 26

1. Choice of nest - well hidden @3m above ground, 15cm2 entrance, 40l
2. Whole foraging force can defend, as well as bees 12 days+
3. Stings are ultimate defence because they die
4. Disease minimised by bees dying away from colony + necrophoresis
5. High brood temp minimises viruses and bacteria
6. Defectation outside hive or stored in winter
7. Propolis varnishs cells and embalms critters
8. Poisoned bees quickly evicted so stores uncontaminated and toxic pollen not used for brood.
9. Hygenic bees romove diseased brood fast
   
   1. AFB - resistant bees remove affected brood early to minimise infection
10. Some strains groom themselves better than others - removing varroa

Question 27

05 Queen substance

What messages does it convey

FRANCO 99

Answer 27

• F Foraging stimulator
• R Retinue stabilised
• A Attracts workers
• N Nasonov stimulator
• C QCell building inhibitor (9HDA)
• Ovary development inhibitor
• 9ODA attracts drones and swarm to queen
• 9HDA keeps swarm and colony stable

Question 28

5. Describe typical foraging behaviour

Answer 28

• Forage in daylight, good weather, windspeed 15mph
• Min temp 13˚C;
• Max temp 43˚C - then forage for water
• Flying 6-10’ above ground at max
• Forage up to 2.5 km from hive
• Pollen collection stim by brood pheromone

Question 29

5. Worker policing definition

Answer 29

• The phenomenon in Queen Right colony where workers eat eggs laid by other worker bees and show aggression towards the laying workers

Question 30

5. Worker policing - 4 benefits

What are the benefits of worker policing (12)

Answer 30

*Perpetuation of the Queen’s unique genes and characteristics
*Alignment of genetic interests so a totally cooperative group
*Reduces reproduction by nest mates with developed ovaries so that
*Almost all bees reared are the offspring of the queen
*Worker eggs in worker cells produce useless, stunted drones and use resources that cd have been used for the Queen’s offspring so energy not wasted raising them
*Workers are more related to the queen’s sons than the worker’s sons
So policing helps rear more closely related Drones, who will be stunted and are a waste of colony resources
Allows unbroken concentric circles of QE, so feeding more efficient
Nothing is wasted because worker police eat the eggs
Aggression acts as a deterrent to other potential LW

Question 31

5. Worker policing

Describe behaviour preventing laying workers and how it works 3

Answer 31

• Queen substance transmitted around the hive from court to workers via trophillaxis and presence of brood pheromones lead to WP worker policing,
• which involves workers eating any eggs laid by laying workers, usually within 2 hours,
• and showing aggression to laying workers

Question 32

5. Worker policing - under what circs would eggs be allowed to develop?

Answer 32

• No queen for 21 days, so no queen pheromones
• No brood, so no brood pheromones
• Unable to produce a queen cell - hopelessly queenless

Question 33

6 Communication -

Describe the behaviour of a forager on forage? 5

Answer 33

1. On encountering the forage, she will reference her entry spot to the area.
2. She may have to learn how to get pollen (anthers) and reach extra floral nectaries (eg field bean)
3. She will crawl or fly from flower to flower collecting pollen and/or nectar
4. She will stay constant to the species of flower
5. She will return to this spot to start her return journey when she has finished foraging
6. She will account for the movement of the sun while she does this

Question 34

06 sensors and stimuli x 11

Answer 34

1. S Trichodea - touch
2. S Basoconica - Taste
3. S placodea - smell
4. S Coeloconica CO2, RH, T
5. S Campanifomia - Stress/Strain
6. S Scolopophera - vibration
7. Trophocytes in abdo - magnetic charge
8. Hairs behind head - vertical
9. Hairs in Petiole - gravity
10. Compound eyes and ocelli - sight
11. Also Atmospheric pressure (Q&D for mating height)

Question 35

6. Communic - List pheramones and their messages

Answer 35

• Fertile Q - Queen substance( mandibular, Koschanvikov, Renner Bauman, Arnhart and Dufour)
  
  • Inhibit QCells and Laying Workers
  • Stim comb building, cell cleaning, brood rearing, forage and storage
  • Attracts court retinue and feeding
• Immature Q
  
  • Inhibit LW
  • Attract drones
• Workers
  
  • Guiding - Nasonov attracts,
  • Guarding - alarm pher 2-heptanone and isopentyl acetate
• Pher in comb
  
  • Attracts scouts in each of new nest
• Worker brood (brood phero)
  
  • Stim pollen foraging
  • inhibt LW ovary development
  • at +6000 encourages drone production
  • Feed me right stuff
  • id mother and caste/sex
  • Drone Brood
    
    • Kairomone for varroa
• Drone
  
  • Attracts Q&D to DCA

Question 36

6. Communic - Buzz Run

Answer 36

• warns bees about to swarm to warm up their flight muscles
• bees can’t fly if muscles are under 35˚C
• in preparation immediately before leaving the hive .
• Involves running through a small group of lethargic bees buzzing her wings, dashing over immobile bees, bulldozing between bees, turning this way and that.

Question 37

6. Communic - Key communication methods

Answer 37

1. Eyes - sight dark
2. Touch - antennal contact
3. Taste scent and touch: Trophillaxis
4. Scent: floral+ pheromones
5. Vibration
6. Dances

Question 38

6. Communic - Dance names and their meaning 8

Answer 38

1. Waggle dance - precise directions to food source 100m+ away
2. Round dance - food source somewhere nearby within 25m - seeley says adaptation of waggle.
3. Sickle dance 25-100m
4. Dorsoventral abdominal vibrating dance - DVAV - go down to dance floor and observe dances
5. Tremble dance - go and unload foragers
6. Piping dance - warm up your fligh muscles
7. Buzz run dance - prepare to leave nest/bivouac
8. Grooming dance - stamps feet and sways from side to side: groom me please.

Question 39

6. Communic - DVAV

Answer 39

• In the hive not on the dance floor
• info: nectar flow: umemp foragers: go to the dance floor to observe dances.
• The dancer approaches a bee from the front
• Mounts on its shoulders waggling its abdomen up and down.
• The other bee stands still until released.

Question 40

6. Communic - Foraging and dancing

What is the connection between foraging and dancing (6)

Answer 40

1. Back at the hive, a nectar source is advertised to unemployed foragers
2. On the dance floor using the round and the waggle dance.
3. The waggle dance gives precise directions based on a bearing on the sun’s cucrrent position
4. Samples by trophilliaxis and antennal touch during this convey scent and taste of teh forrage.
5. When HB are slow to unload - 40secs+ foragers dance a tremble dance away from dance floor to recuit unemployed nectar processors
6. When HB fast to unload - less than 20secs, foragers dance a DVAV away from dance floor to send processors down to dance floor

Question 41

6. Communic - grooming dance

Answer 41

• stamping feet and swaying from side to side
• elicits a grooming response.

Question 42

6. Communic - What are the key pheromones names for Q D B

Answer 42

• Queen
  
  • Mandibular
    
    • 9ODA (9-oxodec-2-enoic acid) - prevents swarming and laying workers, holds swarm together
    • 9 HDA - attracts drones at long distance
  • Renner Baumann QS and drones
  • Arnhart - footprint
  • Dufour - Eggs?
  • Koschevnikov -QS
    *Worker
    *Mandibular - alarm pheromone, used by guard bees to ward off intruders (2-heptanone)
    *Alarm pheromone from sting gland, (isopentyl acetate) attracts other bees to sting
• Drone
  
  • Mandibular - attracts Q & D to DCA
• Brood

Question 43

6. Communic - name worker pheromones

Answer 43

• Nasinov - come in, geraniol
• Arnhart - footprint - I was here
• Alarm scent
  
  • 2 heptanone - in foragers
  • Isopentyl acetate from 12 days

Question 44

6. Communic -

how does a forager recognise food source following a dance? 4

Answer 44

• The forager will fly out on the bearing given by the dancer
• She will recognise the scent of the pollen, nectar and flower from anntennal contact during tophillaxis when she she begged a sample on the dance floor
• She will recognise the taste of the nectar from the sample received by trophillaxis on the dance floor
• She will recognise the forage from the scent of the pollen, nectar and flower, as transmitted by the dancer during trophillaxis on the dance floor

Question 45

Communic - round dance

Answer 45

• Forage within 100m of hive
• On the dance floor.
• Looped circle one way and then the other.
• Share samples by Trophillaxis
• Discover its source by themselves.
• Danger of robbing - eg wet supers.

Question 46

6. Communic - Tremble dance

Answer 46

• In the hive away from the dance floor
• if unloading takes more than 40 seconds
• recruits nectar processors
• The dancer walks SLOWLY about on 4 legs,
• forelegs, also trembling, raised as if begging
• while its body trembles left and right, back and forth, side to side
• PIVOTING FROM HEAD

(WOTH 165)

Question 47

6. Communic - trophillaxis define

Answer 47

• The exchange of nectar or honey from one bee to another with antennal contact
• Note young bees obtain all food from trophillaxis from older bees
• Mainly older bees and foragers who offer

Question 48

6. Communic -

how does trophillaxis increase water collection in dearth

Answer 48

• The optimal sugar:water concentration in the honey sac is 50:50 and bees strive to maintain this through out their lives.
• Nectar = 30-90% water, usually meets the bees’ water requirements.
• In a dearth, bees eat honey stores = 20% water.
• Nurse bees produce brood food, which is 70-80% water.
• Nurse bees feed on honey, which may normally be diluted with nectar.
• As bees share this by trophillaxis, the sugar concentration of the honey sac will rise above 50%.
• Water foragers are motivated to forage for water, recruiting more water foragers with dances.
• With no nectar income, nurse bees will take large quantities of water from water receivers by trophillaxis.
• The faster receivers unload water foragers, the faster foragers return for more water.
• Unloading <60secs = get more; >60secs slow down >180 secs stop
  *

Question 49

6. Communic - Waggle dance

Answer 49

• Directions >100m from hive
• Dance floor if unloading less that 20 secs
• figure 8, waggling its abdomen size to side and buzz wings
• as travels the straight line of the 8
• then circles alternately left and right to repeat this step
• The angle to vertical (sun’s current position), gives the bearing to take from the sun on leaving hive.
• Trichodea gravity sensors between head and thorax
• The length of the waggle, and pips indicate distance to the forage
• Inter-ommatidial hairs on compound eyes - wind speed and distance
• Samples by trophillaxis
• Account for time
• Better ->longer dance -> more foragers

Question 50

6. Communic -

what is behaviour of nectar scout after she has collected some nectar

• exclude behaviour at hive

Answer 50

1. At the flower the bee swallows nectar down to honey sac
2. Adding invertase from her hypopharyngeal glands as she does so
3. This starts the breakdown of sucrose to glucose and fructose
4. She may eat some nectar to fuel her continued flight
5. Crawls/flies from flower to flower of the same species collecting nectar
6. Stays constant to this species, collect more nectar to ascertain forage quantity
7. This affects the duration of her dance back at the hive (excellent sources merit longer dances to recruit more foragers)
8. When she has a full load, about 35-45mg dep on distance, she flies back to her entry point at the forage
9. She navigates home using landmarks
10. Orientation of the sun, UV and polarised light helps navigation
11. Trophocytes in the bee’s abdomen contain molecules that help the bee to work out magnetic orientation

Question 51

7 Orientation

A worker honeybee has been given the direction and distance to a new forage source by observing a dancing bee. List the navigational methods used for determining direction and distance during the first flight of a new recruit to find the new forage source. (You do not need to describe the any dances) 8

Answer 51

1. The sensilla seta on the bee’s compound eyes allow it to estimate distance
2. Distance was conveyed by clicks inaudible the human ear during the dances, and the number of waggle sections per 15 seconds
3. The direction to the forage corresponds to the angle between the vertical point on the comb (which represents the sun’s current position) and the waggling part of the dance.
4. Hairs between head and thorax allow it to sense gravity and hence the vertical.
5. If the waggle was at 45˚ to the vertical, the bee exits the hive and flies along a bearing of 45˚ to the sun’s current position.
6. The bee can detect UV and polarised light coming from different directions and determine the precise angle between them.
7. As it approaches the forage, it will hone in on scent and taste of the flowers/pollen and nectar as well as any arnhart pheromones that remain from trips made by other bees, including the scout.
8. The bee can allow for the sun’s movement (15˚/hour) of the sun automatically

Question 52

7 Orientation

On arrival at the flowers, what additional cues confirm to the bee that this is the correct forage source? 3

Answer 52

1. During the dance the bee will have been given samples of the taste and smell of the forage by trophillaxis
2. She compares this with the taste and the scents she has been given by the dancer.
3. She will also be able to scent the arnhart pheromone remaining on the flowers visited by the dancing bee. This remains for 4 hours at 23˚C.

Question 53

V(SdSpUV) MSTELE(15)M(2)D(SS)W(OJ)

7 Orientation

Describe the mechanisms by which a foraging honey bee finds her way back to the colony entrance and how this is affected by experience. 30

Answer 53

1. Visual signals –
   
   1. sun direction
   2. sun polarisation and UV
   3. landmarks
2. the earth’s magnetic fields
3. smell
4. taste
5. The first time a bee returns from forage, she will have experienced the flight out to it, so already she has experience.
6. She also has powerful experience of the hive’s locality, thanks to her early orientation flights, which grew progressively wider as she matured towards becoming a forager
7. Landmarks are her principal guide after she has first found the forage. These include physical objects bushes, trees, roads, as well as colour and patterns and nectary guides
8. For example, if a bee is lifted from a forage source, placed in a dark container and moved to another spot where the hive was still visible but not the forage, and then released, after a moment of disorientated the bee will devise a new flight path back to the hive based on landmarks. (gould ) (Yate 6.21.3 )
9. Note that bees become disoriented if landmarks change – each cutting long grass around a hive will cause bees to be disoriented for several hours.
10. At the forage, the bee always arrives and departs from the same entry point to the forage
11. During her foraging trip on the crop, she somehow orientates and manages to compute, thanks to the sun’s UV and polarisation, where her entry point is at all times
12. allowing for the movement of the sun by 15˚ per hour.
13. She retains the memory of the forage, flight paths and hive location – this lasts for two weeks.
14. She will fly on the reciprocal and original bearing, while accounting for the movement of the sun
15. Scents and tastes along the way, including pheromones and hive and colony scents will guide her
16. She follows magnetic clues thanks to magnetite in trophocytes in her abdomen
17. She may account for the direction of the wind and the outward and reciprocal bearing on this (it won’t change much during one short foraging trip).
18. She assesses distance using the sensilla seta on her compound eyes.
19. She assesses wind direction using the organ of Johnson in her antennae
20. As she gets nearer the hive (half a mile), these landmarks will become very familiar from her initial orientation flights as a house bee and she will ‘fly on automatic pilot’
21. The automatic pilot aspect is so strong that, if the hive is moved more than 3’ and less than 3 miles, she will not be able to find it.
22. This is because she will automatically revert to old flight paths
23. Some say the bees will follow an odour plume out and back (Yate 6.21.1). This theory has not been proven. Obviously this will also depend on the direction of the wind.

Question 54

07 Orientation

What other aids may she use on her return to the hive and on subsequent flights to the same forage source? 6

Answer 54

1. The memory of its location and how to get there and back
2. The reciprocal and original bearing, while accounting for the movement of the sun
3. Scents and tastes along the way, including pheromones and hive/colony scents
4. Magnetic clues thanks to magnetite in trophocytes in her abdomen
5. Landmarks including colour and patterns and nectary guides
6. Sensilla on her eyes to assess distance

Question 55

7. Orientation

Briefly describe the behaviour of honeybees in relation to recognising the location of their hive. 4

Answer 55

1. Young bees go on orientation flights to learn the colour, shape and orientation of their hive in relation to immediate surroundings/landmarks
2. And the precise position of the hive in relation to nearby landmarks - everything within ½ mile by the time they start foraging
3. They recognise the nasonov pheromone at the hive entrance, and the hive and colony odour
4. They will recognise the position of the hive in relation to UV and polarised light - older, more experienced foragers can find their way home if moved from one location to another inside a dark container.

Question 56

08 Guard

During a moderate nectar flow, how would a guard bee react to a fully laden worker drifting into the wrong hive? 2

Answer 56

• In a moderate nectar flow, it is likely give it a desolultory challenge, scented with its antenna for 1-3 seconds. The incoming bee may simply get past the guards by acting confidently.
• Or if the laden worker acts submissively, curling its abdo and legs in and offering a bribe to get in it may let it in but follow it into the hive and observe for a few seconds.

Question 57

08 Guard

Recognise a robber bee? 2

Answer 57

1. Robbers have a characteristic zigzag flight as they approach the hive, because they are
   
   1. not familiar with the hive/entrance,
   2. weighing up the colony’s defenses
   3. alerting other robbers to the location of the colony being robbed
2. Habitual robbers become smooth, shiny and black, possibly as a result of the occupational hazards of fighting other bees (Winston 115-116)

Question 58

08 guard

What is hive odour and what factors affect it? 3

Answer 58

1. All honey bees have an odour which is colony specific.
2. This seems to come from the unique mix of nectars that are in the individual hive and are absorbed by the waxy surface of the bee
3. and from the common crop contents of all the inhabitants of the colony brought about by trophillaxis
4. as well as various in the mix of pheromones.

Question 59

08 guard

Seasonal impact on guarding 4

Answer 59

1. Colonies are more defensive during late summer and early autumn when they have large stores
2. In the early part of the year they are more relaxed about strange bees and even beekeepers.
3. Small colonies are less likely to attack than large ones.
4. In winter bees in the centre of the cluster are warm and need very little time to warm up their flight muscles to fly to the attack - eg when the beekeeper does an oxalic acid treatment
5. Some bees are genetically predispossed to be more defensive than others
   
   1. Eg African Bees
   2. Eg second generations of imported queens

Question 60

08 guard

How do you recognise a guard bee?

Answer 60

It’s stance

1. Standing on back and middle legs
2. Front legs raised
3. Antennae facing forward
4. Wings spread
5. Mandibles open

Pre-stinging behaviours include threat postures, buzzing, burrowing into predator, biting, pulling hairs

Question 61

MVDAPCO2

08 defence

Stimuli for stinging 6

Answer 61

1. Rapid movement
2. Vibration
3. Dark colours
4. Animal scent
5. Beekeeper’s perfume, perspiration
6. Bees sense CO2 for location best place to sting

Question 62

08 guard

Recognise a bee behaving submissively

Answer 62

1. The bee flies in straight and alights on the landing board
2. After being challenged, it may curl up legs, curl in abdomen
3. She may offer food
4. She may pull tongue through front feet
5. Guards will either remove such a bee without harming her or allow her to enter but follow her into the hive and watch her for a while

Question 63

08 guarding

describe guard behaviour at entrance in dearth (ignore stance)

Answer 63

1. Guards challenge every bee entering the site for 1-3 seconds by antennal contact
2. If it is a bee drifting in, it is likely to have flown in on a direct path, have a load of forage and to blithely ‘stroll’ in with confidence.
3. If challenged, they are likely to offer the guard food and beg their way in.
4. If it is an intruder intent on robbing, it is likely to have had a zigziag flight path in
5. This alerts the guards who may maul it, clamping on a leg wing, they might bite or pull hairs,
6. This will mark the intruder with 2-heptanone – and attract other bees from the hive to maul it and try to drive it off
7. The guard bee will curl her abdomen under enabling her to sting the intruder. Extruding sting releases sting scent from their sting scent gland.
8. Isolentyl acetate inhibits foraging and recruits more guards from the foragers.
9. Another sting scent pheromone - z-11-eicosen-1-ol boosts the effect of IA prolongs its effectiveness and attracts defenders to the area of the intruder.
10. IA elicits stinging in the same spot.
11. Bees as young as 12 days after emerging can be recruited to guard duty as thier sting gland has matured.

Question 64

08 Guard

how do bees defend against other pests trying to enter the hive

Answer 64

1. Acarine. Bees have no defense against acarine, which live in the trachea of affected bees, who develop K wing, and tremble on grass stalks outside the hive.
2. Varroa. Bees have no defense against varroa and tend to coexist with them, although some strains on honey bee groom themselves free of varroa better than others. A bad infestation will result in bald brood, reduced longevity, increased susceptibility to disease (eg deformed wing virus) and problems with orientation. In very bad cases, bees may abscond.
3. Wax moth. Bees tend to coexist with wax moth, although they will sometimes remove the larva.
4. Wasps and European hornets are hard to sting has they have a hard exoskeleton, so attacks are met with grappling and workers gang up and drive them off. Wasp attacks may be met by shimmering behaviour where workers shake violently from side to side, which often dissuades the attackers.
5. Asian Hornet. European bees have not yet adapted to the Asian Hornet and have no defenses.
6. Ants. Workers at the entrances turn their backs on ants and fan their wings and kick their rear legs which usually prevents ants from entering the nest.
7. Small hive beetle. Bees have no defense against SHB because they have very hard exoskeletons and tend to scurry very fast. They will try to confine them to the edges of frames, although the SHB has adaptive behavior and ‘begs’ for food. The bees end up feeding them by trophillaxis. When the colony is disturbed (eg during a manipulation), the beetles will escape confinement and lay hundred of eggs in cells. In a strong colony, bees will try to remove the larva.
8. Mice. Bees can coexist with mice, although the bees may kill and mummify in propolis and the cluster will move away from the mouse. If it is early enough in the year to start again, bees may abscond.

Question 65

08 Guard

What would the colony reaction be to an attack by wasps? 5

Answer 65

1. Wasps hover near nest attacking adults and removing thorax and carrying it back to their nest.
2. Wasps that gain entry eat brood/honey.
3. They have a characteristic zigzag flight as they approach the hive
4. Workers will shimmer in the entrance to try to put them off - (shake violently from side to side)
5. Guards immediately grapple wasps that land and workers gang up to maul and bite them and try to drive them off, curling their abdoment round to sting them.
6. marking them with 2-heptanone, so that if they escape into the hive they will be challenged by other workers.
7. Extruding the sting releases the sting scent pheromone isopentyl acetate which recuits more defenders

Question 66

09 nectar and water collection

How do honeybees mark a water source? 1

Answer 66

With the nasonov pheromone, although the arnhart pheromone will also linger for 4 hours at 23˚C

Question 67

09 Nectar and water collection - nectar to honey

Discuss the process of nectar collection and how it is converted into honey and stored within the nest. (description of bee dances is not required). 22

Answer 67

1. The bee arrives at a flower as a result of scouting/following ‘directions’ given in a waggle dance
2. In the dance, the bee will have been given samples of the nectar by trophillaxis
3. At the same time she will have picked up the scent of the pollen, flower, and nectar from the antennal contact during trophillaxis
4. She will navigate to the flower and be attracted by the scents she recognises, including arnhart pheromone which will linger on flowers for 4 hours at 23˚C; 4 days at 5˚C
5. Nectary guides and the scent of nectar will guide her to the nectaries, although she may have to learn how to reach these eg by nibbling a hole in the side of the field bean flower
6. She will extend her proboscis and suck up the nectar though its food canal.
7. As this passes over her hypopharyngeal plate, she adds invertase from her hypopharyngeal glands
8. This start to hydrolyse sucrose to fructose and glucose.
9. C₂₄H₂₂O₁₁ +H₂O + sucrase ¾> C₆H₁₂O₆ + C₆H₁₂O₆
10. She also adds glucose oxidase with converts glucose to gluconic acid and the antibacterial hydrogen peroxide, which will later help preserve the nectar when it has been converted to honey.
11. She navigates back to the hive by returning to the forage entry point, taking a reciprocal bearing from the sun that she came out on, allowing for the passage of time.
12. She will also navigate by landmarks and familiar odours – eg other flower scents in the area, as well as colony and hive scent and nasonov pheromones from the hive entrance.
13. Back at the hive, she unloads to a nectar receiver at the edge of the brood nest, who moves away from the brood nest to start the process of converting the nectar to honey
14. The house bee first ripens the honey: this involves regurgitating a droplet of nectar onto the end of her proboscis and swallowing it down about 100 times over 20 minutes.
15. She may add additional sucrase during this process.
16. This reduces the water content of the nectar by about 15%
17. She then spreads the nectar out in cells to evaporate further.
18. Other house bees fan the honey – replacing moist air inside the hive with dry air from outside the hive
19. This reduces the water content to about 20%
20. The house bees then pack the honey into cells and seal them with wax leaving a miniscule air gap.

Question 68

09 Nectar collection

Nectar collection - what is involved 8

Answer 68

• A bee either discovers a patch or her own, or has followed the directions given in a waggle dance to find it.
• During the dance, she will have received samples of the nectar by trophillaxis giving her the taste and smell of the nectar. Antennal contact during trophillaxis will have transmitted the scent of the pollen and flower.
• The bee arrives at the foraging patch and marks her entrance – she will arrive and leave from the same point every time
• Bee lands and uses nectary guides and nectar scent to find the flower’s nectaries
• She may have to learn how to reach nectaries - eg at field bean, will learn to copy bumble bee behaviour to access nectary from side or back by nibbling at the flower
• She will suck up the honey using her proboscis into her honeysac
• While collecting nectar at the flower, she adds sucrase from her hypopharyngeal glands. This mixture is swallowed to honey sac.
• Sucrase hydrolyses the sucrose (disaccharide) into glucose and fructose (monosaccharaides) which can be absorbed through the bee’s gut
  C₁₂H₂₂0₁₁ +H₂O C₆H₁₂0₆ + C₆H₁₂0₆

Question 69

09 Water and nectar

When is water collection noticeable

Answer 69

1. Late winter, early spring to disolve/dilute stores
2. Spring when nurses are feeding brood and relying on Honey stores for food
3. Hot weather when cooling required

Question 70

09 water and nectar collection

Give a description of the bee behaviour involved in nectar collection

Answer 70

1. A bee either discovers a patch or her own, or has followed the directions given in a waggle dance to find it.
2. During the dance, she will have received samples of the nectar by trophillaxis giving her the taste and smell of the nectar. Antennal contact during trophillaxis will have transmitted the scent of the pollen and flower.
3. The bee arrive at the foraging patch and marks her entrance – she will arrive and leave from the same point every time
4. Bee lands and uses nectary guides and nectar scent to find the flower’s nectaries
5. She may have to learn how to reach nectaries - eg at field bean, will learn to copy bumble bee behaviour to access nectary from side or back by nibbling at the flower
6. She will extend her proboscis and suck up the honey through the food canal into her honeysac
7. As this flows over her hypopharyngeal plate, sucrase is added from her hypopharyngeal glands.
8. Sucrase hydrolyses the sucrose (disaccharide) into glucose and fructose (monosaccharaides) which can be absorbed through the bee’s gut
   C12H22O11 + H2O= C6H12O6 + C6H12O6
   Sucrose + Water = Glucose + Fructose
9. She navigates back to the hive using a reciprocal bearing on the sun from her journey out, landmarks and scents
10. At the hive she will unload the nectar to a nectar receiver on the edge of the brood nest
11. She will unload the pollen into a cell on the outside of the brood nest.

Question 71

09 Water and Nectar collection

Honeydew 6

Answer 71

1. Honeydew from plants/hemiptera
2. Trees inc PLACEBOS HC
   
   1. Prunus Lime Ash Chestnut Elm Beech Oak Sycamore Hawthorn Conifer
3. Hemiptera feed on phloem and excrete Honey Dew
4. If it dries, it is called manna
5. Attacts bees when not other forage available
6. If dry, they wet it with saliva

Question 72

09 Nectar and water collection

How is water used in the colony? 5

Answer 72

1. To dissolve granulated honey
2. To dissolved honey stores
3. To cool through evaporation
4. To hydrate themselves - they have liquid faeces
5. To hydrate nurse bees who produce brood food which is 70-80% water

Question 73

09 water and nectar collection

How is water collected and transported back to the hive? 10

Answer 73

1. Collection varies according to demand - it is not stored in the hive
2. Site often marked with nasonov
3. They take up water in about 1 minute (av load 25mg)
4. 67% of field trips completed in 3 minutes; 90% in 10 minutes, water usually close by
5. Average 100 trips a day
6. They may collect water from a nearby puddle/pond
7. On cool/wet days, they may also sip water from dewy grass stems outside hive
8. They suck up water using their proboscis
9. They swallow this to their honey sac in which they transport it back to the hive.
10. Unloading <60secs -> more water; >60+ secs slow down; >180+ secs stop collection

Question 74

09 Water and nectar collection

Describe water collection in terms of preferred sites and regularity of visits. 4

Answer 74

1. 67% of field trips completed in 3 minutes; 90% in 10 minutes, so water is usually close by
2. They may collect water from a nearby puddle/pond
3. On cool/wet days, they may also sip water from dewy grass stems outside hive
4. Nasonov marking

Question 75

10 Nectar and water collection

Describe the interrelationship between nectar, honey and water in the colony in summer and winter

Answer 75

1. Bees can only metabolise sugars when the sugar:water ratio in their honey stomach is 50:50
2. They will strive to maintain this concentration throughout their lives
3. During summer, the nectar flow (usually 30-90% water)
4. This usually satisfies the bee’s requirement for water
5. In winter, or dearth, there is no nectar flow, so bees rely on honey stores, which are around 20% water.
6. Therefore, foragers must collect water in order to metabolise honey.

Question 76

11 Hydrolysis - define

Answer 76

1. Breaking down a compound with water to make other compounds
2. C₁₂H₂₂0₁₁ +H₂O C₆H₁₂0₆ + ₆H₁₂0₆
3. Sucrose + Water + invertase = Glucose and frucose

Question 77

11 nectar processing

What does nectar contain?

Answer 77

• 20-90% water, rest sugar S/F/G
  
  • Primrose 5% sugar
  • OSR 50% sugar
  • Horsechestnut 70% sugar
• MOLEPAVA
  
  • Mineral ash
  • Organic acides
  • Lipis
  • Enzymes
  • Proteins
  • Amino Acides
  • Vitamins
  • Aromatic compounds

Question 78

11. Ripening - define

Answer 78

1. Ripeing is the manipulation
2. by the house bee
3. whereby she swallows and regurgitates nectar + sucrase mixure
4. 100 times
5. over 20 minutes
6. to reduce the water content by 15%

Question 79

SWFMGOHP

11. Why does a bee make honey 6

Answer 79

1. Honey represents the colony’s stores for dearth, times of bad weather and winter
2. Nectar contains 20-70% water plus various combinations of fructose, sucrose and glucose, among other things including amino acids, organic acids and aromatic compounds.
3. If stored in this form, is would ferment/grow mould
4. So bees convert it into honey, which has 20% moisture content or less
5. Glucose oxidase is added from hypophryngeal glands
6. Converts glucose to gluconic acid and hydrogen peroxide. HP is antibacterial another layer of preservation

Question 80

11. Arriving at the flower to sucking up nectar 5

Answer 80

1. The bee lands and uses nectary guides and nectar scent to find the flower’s nectaries
2. (This scent was transmitted from the dancer to the prospective forage during trophillaxis during the waggle tail dance)
3. She might have to learn how to reach the nectaries
4. eg at field bean, will learn to copy bumble bee behaviour to access nectary from side or back by nibbling at the flower
5. She will suck up the honey using her proboscis into her honeysac
6. Passes over HPG adds sucrase

Question 81

11

How do honey bees convert nectar into honey?

Include a brief description of the physical chemical changes from the flower to the capped cell

Answer 81

1. While collecting nectar at the flower, the bee adds sucrase from its hypopharyngeal glands
2. This mixture is swallowed to the bee’s honey sac.
3. Sucrase hydrolyses the S (disaccharide) into G+F (monosaccharides) which can be absorbed through the bee’s gut
   C₁₂H₂₂0₁₁ +H₂O C₆H₁₂0₆ + ₆H₁₂0₆
   OR starts breakdown of S with H2O to G&F (HYDROLYSIS)
4. The forager transfers its load to house bees usually on the edge of the brood nest
5. House bees moves away from the broodnest,
6. They manipulate the nectar (called RIPENING), swallowing an regurgitating it back down to their honey sacs about 100 times over about 20 minutes
7. Adding more sucrase from its hypopharyngeal glands
8. This ripening reduced water content of the nectar by about 15%.
9. The house bee spreads the ripened nectar to dry on the surface of empty cells.
10. Fanning by other house bees removes the moist air from the hive and replaces it with dry air from outside reducing the water content to 18-20%.
11. The honey is moved and partially filled cells are filled and capped with wax seal;
12. Glucose oxidase, also added from the bee’s hypopharyngeal glands, converts glucose into gluconic acid and hydrogen peroxide. (the HP is antibacterial)

Question 82

11 Honey processing

Name three enzymes added to nectar

Answer 82

1. Sucrase
2. Gucose oxidase
3. Diastase

Question 83

12 pollen - bee bread

Why is pollen turned into bee bread and how is this achieved? 6

Answer 83

1. It would otherwise grow mould
2. Pollen is ‘glued’ to forager leg when front legs are wetted with nectar as it grooms the pollen into its corbiculae at the flower
3. Pollen is dropped off in cells by foragers
4. House bees mix it with more honey or nectar, which includes enzymes and bacteria and pack it into cells
5. These inhibit germination and growth of moulds
6. And give rise to lactic acid fermenation. The pollen becomes darker and sweeter and is known as bee bread.
7. Although only 50% of nutritional value of fresh pollen
8. Stored combs with pollen attracts pollen mite

Question 84

Faithful hairy survivalists mark nonstop daylight waggledances

12 pollen

What makes bees good pollinators?

]

Answer 84

Faithful hairy survivalists mark nonstop daylight waggledances

1. Faithful to one type of flower per visit
2. Hairy – bees covered in 3million hairs – many plumose – gap between hairs is 45μm = width of dandelion pollen grain (BBKA NEWS 10/17), which sweep up 15000 pollen grains/flower
   
   1. Pollination incidental to collection.
3. Can be moved en masse to pollinate crops
4. Survive en masse over winter and emerge in spring in large numbers to collect pollen
5. Mark forage with arnhart – so a beacon for new foragers – lingers 4 hours at 23˚C and 4 days at 5˚C
6. Nonstop foraging in daylight except in bad/cold weather
7. Forage in daylight so pesticide/herbicide spraying possible at night
8. Waggle dances communicate sources to other bees
9. Sun: Are able to use the sun, UV and oplarised light to navigate to source

Question 85

12 Pollen - ways it is collected 2

Answer 85

1. May bite filaments to dislodge pollen from ripe anthers
2. In trying to reach nectaries at the back of the flower the bee will brush against ripe anthers and collect pollen on plumose hairs before pushing it to the corbucula

Question 86

12 Describe Pollen collection in detail 20

Answer 86

1. Collection stim by QS and BP
2. Two types of bee collection:
   
   1. as a by-product of nectar collecting
   2. purposely collecting pollen.
3. Approximately 20% of foraging bees purposely collect pollen (approximately 1/3 of workers are foragers, 2/3 of a colony are house bees).
4. Arriving at the flower, by-producters get covered in pollen
5. Purposefiul collecters may bite filaments of the stamen to dislodge pollen from the ripe anthers.
6. Pollen lands on her body and catches in her plumose hairs
7. She flies/walks from flower to flower of the same species, incidentally cross pollinating from the pollen on her body as she goes
8. As she leaves a flower, she will groom the pollen into her corbiculae on her hind legs.
9. She moistens her legs with a bit of nectar, and then grooms the pollen from her head and first thoracic segment
10. The middle legs collect the pollen from the first legs and the rest of the thorax, particularly the ventral side
11. The hind legs collect the pollen from the middle legs and clean the abdomen.
12. When enough pollen is collected on the surfaces of her basitarsi, she will rake the pollen into the auricle on the opposite leg where it is held in place by the hairs of the rastellum
13. The retaining hairs of the auricle stop it falling out.
14. The teeth on the auricle grip it.
15. She then closes the tarsus against the tibia
16. This squeezes the pollen up and outwards in a patty blob on to the concave external surface of her tibia where the pollen is held in place by the hairs on the corbiculae.
17. Repeated several times over 3-18 minutes to build up a load of up to 8mg/leg.
18. A full load is approx 100 flowers
19. She will unload the pollen into a cell on the edge of the brood nest
20. Each journey takes 3-20 minutes in daylight hours in reasonable weather - low wind, temperatures above 13˚C

Question 87

12 Pollen external causes of regulation

What factors OUTSIDE the hive regulate the amount of pollen collected? 2

Answer 87

1. weather
2. un/availability of forage

Question 88

12 Pollen feedback mechanism - in brief

Outline the feedback mechanism involved. 6

Answer 88

1. Inhibitory cue from the protein-rich secretion from hypopharyngeal glands which is fed to foragers by trophilliaxis
2. The larger the P. reserves, the greater P. consumption by nurse bees
3. the more protein is fed to foragers by trophillaxis, the more foragers are inhibited from further P. collection.
4. If the nurses need to feed more larvae, there is less for foragers who are hungry for protein and increase the number of foragers and the per capita rate of collection
5. The reserve act as a buffer - rate of collecion is more variable that demand.
6. So the increased demand is responded to before the reserves in the nest fall too far

Question 89

12 Pollen internal causes of regulation

What factors within the hive regulate the amount of pollen collected? 2

Answer 89

1. presence of brood pheromone
2. number of cells prepared by house bees for pollen (Yates)

Question 90

12 Pollen

Two pheromones that stimulate pollen collection 2

Answer 90

1. Queen substance
2. Brood pheromone from open brood

Question 91

12 pollen reserves - detail

Describe how a honeybee colony regulates its pollen reserves?

Answer 91

Seeley (WOTH)

1. Pollen supply varies more day-to-day than demand -> buffer required of about kg -> colony must regulate supplies
2. Adjusts collecting rate in relation to reserve thro
   
   1. changes in number of pollen foragers
   2. per capita collection rate
3. Foragers fed no sensory feedback on quantity of stores
4. Feedback is indirect and involves non-foragers
5. Critical bees in this are the nurse bees, who are principle pollen users
6. They provide excitory feedback when there is little pollen in the hive
   
   1. Perhaps by preparing cells for pollen storage so unloading is quicker
7. Only evidence at present is inhibitory feedback from nurses when there is abundant pollen in the hive
8. Evidence suggests it is inhibitory cue from the protein-rich secretion from HG which is fed to foragers
9. Study required to assess key assumptions
   
   1. That the amount of secretion varies in relation to size of reserve
   2. That the amount received influences foragers collecting activity

Question 92

12 Pollen

Describe unloading2

Answer 92

• She grasps cell edge with front legs,
• arches abdo so hind legs dangle over cell
• Middle legs push off load into cell

Question 93

12 Pollen uses x 10

Answer 93

1. Protein
2. Nurse bees +P = BF + RJ d
   
   1. BF&RJ are a proteinous-rich fluid that enables brood/queen cells to grown and develop
   2. The more the Q is fed RJ, the more eggs she lays per day
3. Nurse bees feed P to older worker larvae
4. Capping brood
5. Reactivate or de-atrophy HPG when more nurses needed (eg after a swarm)
6. Gland development
7. Metabolic ‘Repairs’ eg to forager muscles
8. Stored - About 1kg is, pickled with honey to make bee bread to
9. Winter bees - to enlarge their HPG + fat bodies -> longevity
10. Lack of pollen -> stress, which can trigger chalk brood.

Question 94

12 Pollen

variations in who collect N and P

Collection duration

No. trips/day

Answer 94

• 60% of foragers collect nectar
• 25% collect Pollen
• 17% collect both
• Pollen is quicker (10 mins) CF nectar (30-80 mins)
• So will travel longer distances for P
• 10-15 trips/day

Question 95

CHAPP

13 propolis

sources

Answer 95

1. resinous substance obtained by HB from trees and wounds in woody plants.
2. CHAPP
3. Cherry,
4. Horse chestnut,
5. Alder,
6. Poplar
7. Pines

Question 96

13 Propolis

How is propolis unloaded?

Answer 96

1. Unloaded by a house bee
2. At the point of use
3. Takes about an hour to bite and pull off small pieces and
4. fix in place with mandibles

Question 97

13 Propolis

In what ways is propolis used by the honeybee colony?

Answer 97

1. eCAVES
2. Embalm
3. Cracks - filled to keep out wind and rain.
4. Antibacterial - preserves brood food and minimises infect
5. Varnish cells
6. Entrance narrowed
7. Smooth rough surfaces

Question 98

13. Propolis

Define and properties

Answer 98

• resinous substance obtained by HB from trees and wounds in woody plants.
• Antibacterial properties.
• Waterproof: disolves in meths, alcohol and ether
• Cold: hard and brittle when cold
• Warm: v sticky when warm - only collected in warm
• Orange to dark brown
• Chractristic smell: clean antiseptic

Question 99

13. propolis

How is propolis collected by honeybees? 2

Answer 99

1. In WARM weather, bite of small pieces and kneed with mandibles till pliable
2. Pass it to their forelegs
3. Transfer to corbiculae on same side by middle legs and patted into position with middle leg

Question 100

13. Propolis

uses by man

Answer 100

• Cosmetics
• Health food
• Tincture
• Varnish
• Wound dressing

Question 101

14 Swarm + supersedure

how a beekeeper can tell the difference between them.

Answer 101

1. Some say there tend to be fewer queen cells in a superseding colony, but with a medium number of QCs it is impossible to tell.
2. It is almost impossible to tell if a colony is superseding, swarming or even plans to destroy the cells and do neither.
3. A bee keeper can assess the amount of space remaining (if little, a swarm trigger) – colonies with a large amount of space are more likely to be superseding.
4. The age of the queen (old queens are a swarm trigger), unless she is in a captured swarm, in which case she is likely to be superseded because she was old when she swarmed and, having established the swarm, the bees may not consider her good enough to get through the winter.
5. Prior to a swarm, bees will be quiescent at the bottom of the combs and may have hypertrophied wax glands
6. Prior to a swarm, there will be a huge number of capped brood and very few eggs

Question 102

14 Swarm colonisation

From departure to the arriving at the new site

Answer 102

1. The queen’s mandibular gland (9-oxydec 2 enoic acid) attracts the bees in the swarm and holds it together,
2. while workers fan their nasonov glands to call in stragglers.
3. Scouts continue their search for a new site and dance locations of possible sites on the surface of the swarm using the waggle tail dance,
4. which gives a precise angle to the vertical (representing the sun in it’s current position) to fly out on, and the length of the waggle and pips and squeaks indicate how far away it is.
5. Scouts inspect the site inside and out taking about an hour for a good site, carefully walking the interior space and even flying it.
6. They are looking for a site off the ground, about 20-80 litrs in size (European bees), with a small, south-facing entrance of about 15cm² at least 300 m from the old colony.
7. When the scouts reach a quorum on a site, they stop doing waggle dances and start doing buzz run dances to get the bees to warm up their flight muscles in preparation for lift off from the temp site.
8. They take off in a cloud of bees hanging over the temp site for about 30 seconds then move off slowing in direction of the new site.
9. The scouts set off in a direct line to the new site, the Queens (9ODA) attracting the swarm bees.The first 30m fly at about 1km/hour and then gradually accelerate so that by 150m they have reached their top speed of 8km/hour
10. Only 5% of the bees know the destination. However, ignorant bees align to bees that seem to have purposeful flight.
11. The scouts ‘guide’ the swarm by Streaking through the colony from behind in the direction of the new site and then allowing the swarm to fly past it, and repeating this process until the reach the site.
12. At 90m before the site, they rapidly slow sown and swarm about 5m from goal
13. Over next 2-3 mins scouts appear at the entrance fanning nasonov
14. When they reach the site, the queen will enter the site and the bees will follow in after her.
15. Workers will collect on the outside and fan their nasonov glands to call in the stragglers

Question 103

14 swarming

What happens in the mother colony after the swarm has issued. 16

Answer 103

1. 40% of adult workers remain
2. Colony contains unsealed and sealed QCs + worker brood + stores
3. Workers continue to forage or may revert to feed brood. They will do this by consuming pollen to reactivate their hypopharyngeal glands.
4. Workers continue to seal QCs and rear worker brood so colony rebounds quickly
5. Worker brood is high during this period (42%) with most lost at egg and young larval stage - not enough adults to incubate/feed them.
6. Queen emerges
7. This usually supresses the emergnce of other queens, probably by DVAV on mature QCs
8. Emerged Q announces presence with pheramones and piping (pressing thorax against come and vibrating wing muscles)
9. While VQ pipes, rest of workers freeze in place
10. When there is a piping Q, workers will not chew away wax from capped ends of QCs preventing them emerging
11. Sometimes two queens emerge at once and may tolerate each other for a few hours/days before fighting.
12. Emerged Q’s attempt to kill sealed queends by cutting holes in cells and stinging occupants.
13. Workers ignore Q and she matures and feeds herself - no RJ/Court
14. Casts may emerge with virgin queens - some can contain up to three queens
15. Workers do not gorge fully and casts contain fewer younger bees issues in casts
16. No of casts correlated with amount of brood and numbe of adults int he colony at the time of swarming.
17. Colonies produce more casts in good years and areas with longer growing seasons.

Question 104

14 Swarming behaviour.

overview of behaviour 6

Answer 104

1. Starve queen so she can fly
2. Departing bees gorge and festoon at the bottom of lower combs
3. Leave after hour’s pipiing and then buzz run dance.
4. 60% leave; 40% remain with brood and stores
5. Castes with virgin queens may emerge 8 days later
6. Weather may delay departure for weather

Question 105

14 Swarming

Dances involved in swarming 4

Answer 105

1. Piping dance - get ready - warm up flight muscles 1 hour in advance
2. Buzz Run - let’s go - prepare to depart imminently
3. DVAV (dorsoventral abdominal vibrating dance) on queen to keep her moving
4. Waggle dance to indicate possible nest sites

Question 106

14 swarming

Describe swarm preparations

laying eggs to departing 22

And when might a beekeeper step in

Answer 106

1. Prep starts 2-4 weeeks before the first swram issues
2. Bees make cups; many are torn down. Number increases prior to queen rearing
3. QR starts when Q lays in cups, tho bees sometimes remove eggs/larva
4. The eggs hatch and the larva are fed royal jelly and long peanut-shpped cells build around them.
5. The beekeeper can do an artificial swarm at this point.
6. Around 15-25 is typical for a swarm
7. Q PUPAE inhibit more QC production
8. Timing - ideally in time of adult pop growth and plenth of brood to compensate for loss of swramer - QR starts when worker brood at it s peak
   
   1. QC destruction may ensure that colonies don’t swarm untilthere are sufficient adults and brood to populate 2 colonies.
9. The queen is feed more frequently and lays more eggs until the week before swarming.
10. Then the workers feed her less so she lays fewer eggs and her bdoment slijms down so she can fly
11. Workers jostle, shake, push and bit queen to keep her moving and slim her further
12. Queen examines QC frequently
13. Receivers are reluctant to accept loads from foragers
14. Scouts stop looking for new forage and start looking for a new home.
15. ALL Bees start gorging on honey about 10 days before swarming to ensure they have enough reserves which the swarm finally issues - all bees carry around 36mg of honey on honey stomach CF 10mg in non swarming colonies.
16. Their wax glands may become hypertrophied
17. They become v quiet prior to swarming and festoon onthe bottom of combs
18. Capping the first queen cell is the trigger to depart
19. The scouts dance the buzz run dance – running at and over lethargic groups of bees and queen buzzing their wings. This is the signal for the swarmers to warm up their flight muscles.
20. Queen is also bitten and chased
21. A torrent of workers pours out, hopefully with queen and takes to the air.
22. they settles nearby to check the queen is with them, if not, the return and start the issuing process again.

Question 107

What component of QS attracts bees to the cluster

What component of QS stablises the swarm

Answer 107

• 9ODA
• 9HDA

Question 108

Bite Prob Legs/mouth Hair Ab

How bees collect pollen

Answer 108

• Open flowers, bee bites the anthers with mandibles and uses forelegs to pull them towards her body
• Tubular flowers, workers insert proboscis into corolla searching for nectar and pollen is collected incidentally adhering to mouthparts and legs
• Closed flowers, bee forces petals apart with forelegs and gathers pollen on the mouthparts and legs
• Spike or catkin flowers, bees run along the spikes or catkins shaking off pollen onto her body hairs
• Presentation flowers, pressing abdomen against inflorescence, causing a pollen mass to be pushed out of the flowers

Question 109

14 swarming

What triggers swarming 6

When might a Beekeeper take action? 3

Answer 109

1. Lack of space to build more comb + reduced QS
2. 9ODA and 9HDA and Arnhart together inhibit queen rearing
3. The queen might be old. Level of QS production halves every year
4. Lots of workers - more workers req more QS
   
   1. Naturally swarming/superceding Queens produce less QS
5. Immature queen PUPAE inhibit further queen rearing
6. Overcrowding may prevent Q from moving along the bottom of combs and depositing Arnhart
7. Beekeeper can identify that there is a likelihood of the hive becoming congested and add supers accordingly.
8. If sees quiescent bees can do artifical swarm.
9. If sees QCs can do artifical swarm

Question 110

14 swarming

triggers short

Describe the conditions and time of year that are most likely to lead to swarming 5

Answer 110

1. Spring and early summer
2. As colony builds up rapidly with congestion in the hive
3. An old queen with reduced queen substance
4. No space to build new comb
5. Genetic predisposition to swarm (eg Carnolian bees if overcrowded)

Question 111

15 supercedure advantages

How can supersedure be of advantage to a honeybee colony? 3

Answer 111

1. Sick or damaged queen is replaced with minimal impact on the colony
2. The colony retains all the bees
3. Stores are not depleted, especially late in the year

Question 112

15 supersedure definition and types 3

Answer 112

1. Requeening of a honey bee colony a daughter queen without the colony swarming and without human intervention
2. Perfect: old queen present in the colony until the new one is mated and starts laying
3. Imperfect: old Q disposed of before new one emerges

Question 113

15 Supersedure

triggers short 5

Describe the conditions and time of year that are most likely to lead to supersedure

Answer 113

SPEND

1. Low levels of queen substance
2. Physical injury
3. Not enough eggs being laid for seasonal build up
4. Nosema
5. Too many drones in the hive
6. Tends to be later in the season – bees will want to replace a queen if they fear will not get them through the winter.

Question 114

16 choosing a nest site

How do bee prepare a new nest site 5

Answer 114

1. As soon as bees arrive they will remove any loose particles from the top down, cleaning the new nest site
2. They will seal unnecessary openings with propolis
3. Some bees will arrive with hypertrophied wax glands.
4. Others bees will hang in festoons for 24 hours while they metabolise the sugars they have brought with them,
5. Raising the temperature to 35˚C at which point bees can produce wax in quantity
6. They then build comb.

Question 115

16 new home assess

How do scout bees determine the internal capacity of a possible new home? 3

Answer 115

1. They walk and fly around it to gauge the volume, as well as inspect it from outside
2. They will inspect the volume during around 25 journeys around the inside measuring capacity by walking - principle means
3. Also sight. They need illunmination of 0.5 lux to do this.

Question 116

16 new home

Briefly list the factors that would encourage a swarm to occupy a cavity. 7

Answer 116

1. European honey bees prefer big sites (20-80litres)
2. (African bees prefer small sites (10litres)
3. A small entrance around 15cm²
4. At the bottom of the cavity
5. 3m meters above ground level
6. South facing
7. At least 300m from old colony
8. Ideally with comb from previous occupation
9. Ideally dry

Question 117

16. new home

How might the swarm utilise propolis to overcome deficiencies in the cavity? 3

Answer 117

1. Narrow the entrance
2. Fill in extra holes to stop draughts
3. Fill extra holes to stop water ingress
4. Line the cavity and smooth rough patches

Question 118

17 comb building

​​What would this comb look like? 12

Answer 118

1. Regular back-to-back array of hexagonal cells arranged in parallel series, each comb a precise distance (6-9mm) from its neighbours
2. Comb is build vertically from top of the cavity – bees sense gravity (hairs)
3. Comb shape tends to be in a catenary curve (the shape of bees festooning)
4. Hexagonal shape - may be the result of visco-elastic flow (Karihaloo 2013) or bees may construct hexagonal cells
5. Cells are set at 13˚ angle above the horizontal to stop honey/nectar poring out
6. Cells are build on two sides of the septum, off-set by 50% for greater rigidity
7. The brood cells are 11mm deep
8. Honey cells can be up to 16mm deep
9. Worker cells are 5 per inch
10. Drone cells are unusually at the sides or bottom and 4 per inch
11. Queen cells are elongaged conical cells hanging from comb edges
12. Two bee spaces between combs to facilitate back to back working – the spacing between combs is around 35mm.
13. Left to their own devices, bees build comb to suit the shape and fill dimensions of the cavity, filling it
14. Several bees contribute to one cell and several cells under construction simult. A bee may smooth wax or add to it
15. Cell walls 0.073mm ish
16. Ikg wax requires 8kg honey to make and supports 22kg honey
17. Workers use antennae to determin thickness and smoothness of cells walls.
18. Some species may try to orientate their combs in a NE-SW direction (about 50˚)

Question 119

17 comb building

List the factors that initiate 5

and inhibit comb building 2

Answer 119

1. During a big nectar flow
2. When combs are virtually full (only 20% of cells empty. WOTH 191)
3. It is dark
4. Brood is expanding
5. The queen is present
6. Not involved in swarm preparation
7. Not preparing for winter.

Question 120

17 comb building

Describe how bees construct new comb. 10

Answer 120

1. Bees must have had a pollen rich diet at some point in order to produce wax
2. Made from wax produced by four pairs of wax glands on the underside of the abdomen of middle aged bees (around 16-18 days old)
3. The bees gorge on honey (8:1 honey: wax), festoon while metabolizing the sugars and raise the temperature to 35˚C, at which temperature the wax will be more pliable
4. They remove the wax scales from the wax pockets with their middle leg and pass it to their mandibles with their front legs (workers with damaged legs cannot build or manipulate wax.)
5. The scale is chewed with the mandibles with secretions from the mandibular glands.
6. The scale is then deposited on the upper inside surface of the cavity to form a small ridge 2-4mm high
7. The comb is drawn down vertically - the bee has sensilla trichodea that sense gravity between the head and thorax and in the petiole.
8. Depressions are moulded into the sides and extra wax is deposited to form the cell walls, which are drawn at an angle of 13˚ from the horizontal to prevent honey dripping out
9. The bees may form a grid of hexagonal cells. Or they may make circular cells and the visco elastic flow of wax as it cools pulls it into a naturally occuring hexagonal shape. (Karihaloo 2013)
10. They don’t appear to need their antennae to build cells - workers with amputated antenna can still build perfect cells accroding to Winston.
11. The bees drawm out cells on both sies of the septum, with the cells on one side offset by 50% from the other for extra strength
12. The shape of the comb is due to the shape of bees handing in festoons to forma catenary curve
13. They start in different places and join up individual pieces as they come together, which accounts for thicker walls in some places.

Question 121

17 comb building

How do honeybees use wax comb? 6

Answer 121

1. To raise brood
2. To store pollen around the brood
3. To store honey above the brood
4. To hang nectar in while they reduce the water content
5. They may keep some empty cells in brood to facilitate warming – bees crawl inside
6. To create a space for living - evern foragers may spend up to 70% of their time hanging about idly on comb.
7. They build larger cells to the side/bottom of the comb for drones
8. They build peanut shaped cells when queen rearing

Question 122

18 summer winter

1. Describe the behaviour of an individual bee in a winter cluster. 4

Answer 122

1. In the outer shell where she will have her head into the cluster and her wings spread or in the inner festoon with the queen
2. She may crawl into empty cell to make colony denser - hypoxia 15% oxygen and 6% CO2 so ends up in suspended animation with v low metab
3. In cluster, as the temperatures drops she will move closer to her fellow workers to contract cluster and reduce surface:volume ratio
4. At ambient 0 and -5˚C she will shiver her dorsoventral muscles to generate metabolic heat
5. From time to time, on warmer days, the cluster will loosen so that bees can move through the hive and feed
6. From time to time on warmer days, she will go out on flights to void her rectum

Question 123

18 summer winter

How does normal colony regulate temperature in the brood area as the ambient temperature rises from 18˚C to over 36˚C. 8

Answer 123

1. The brood needs to be kept at a temperature of 30-35˚C – temperatures above 36˚C are harmful to brood

As internal temperature of the hive rises the bees have several escalating options:

1. Dehydrating nectar has a cooling effect on the hive.
2. Adults in the hive will disperse
3. Older workers (strong wing muscles) line up in chains facing the same way to ventilate the hive - fanning at the entrance and inside the hive to circulate the air.
4. One set of workers at the entrance will face inwards, fanning hot air out; another faces the other way on the other side to fan cooler air into the hive.
5. Water collection goes up as bees use water to cool the hive – water receivers ‘paint’ the water in puddles on capped cells or hang droplets in cells
6. Bees may tongue-lash over brood cells - repeated extend and contract their proboscis, pressing a droplet of water from their mouths into a thin film which can evaporate quickly
7. Bees will evacuate the hive cluster in a ‘beard’ outside the hive to reduce heat generated by their metabolism and by providing more room for ventilation and water evaporation.

Question 124

18 summer winter

How can a beekeeper help regulate high temperatures?

Answer 124

1. Providing sufficient space for bees to disperse in summer, or removing supers in winter so that they have a smaller space to keep warm in winter

In summer

1. Providing shade at noon or in the afternoon
2. Ensuring the crown board feed holes are open the roof ventilators are clear
3. Insulating the hive (to keep it cooler in summer and warmer in winter)
4. Trimming undergrowth to ensure good air flow around hive
5. Staggering supers and off-setting crown boards and roofs and raising the roof to facilitate air currents through the hive in extreme hea

Question 125

18 summer winter

What is the composition of a stable winter cluster? 13

Answer 125

1. The cluster starts for form below the stores at ambient 18˚C
2. Round ball run through with combs with top in contact with stores.
3. The outer shell is 1-3” deep - workers have their heads in and on the outside have their wings spread
4. Below an ambient temperature of 14˚C, the cluster has an a compact outer shell of quiet bees and an inner core of bees more loosely festooned around queen
5. with channels for ventilation
6. Some bees enter empty cells.
7. Bees in centr may suffer for hypoxia as cluster contracts, entering state of suspended animation from v low O2 (15%) and v high CO2 (6%) greatly reducing metabolic rate, so saving energy.
8. Core temperature 20-30˚C
9. 33-35˚C when queen is laying
10. The cluster contracts to conserve heat.
11. At 0˚C (Davies) to -5˚C (Winston) cluster stops contracting and the bees will generate metabolic heat by shiver their dorsoventral muscles.
12. The minimum temperature within the cluster is 13˚C which maintains the temperature of the outer shell at 8˚C
13. At 7.2˚C bees on the outside will fall off

Question 126

18 winter summer

How do bees prep for winter 12

Answer 126

1. During summer: forage and collect harvest (1)
2. Store food (2)
   
   1. Dehydrate nectar
   2. Ripen honey
   3. Seal with wax
3. Hoof out drones in Sept/Oct (1) to save stores
4. Dramatic reduction in population as queen goes off lay (1)
5. Autumn: gorge on pollen and and build up: (3)
   
   1. Hypopharnygeal glands
   2. Fat bodies
6. Propilise gaps in hive (1)
7. Poikilothermic – so take up temperature of surroundings (1)
8. Create homeostasis to keep nest temperature constant regardless of external conditions. (2)
   
   1. Start to cluster at 18C below food store
   2. NB Below cluster temperature of 8C bees cannot cling to cluster.
9. Outer shell is 1-3” deep (1)
10. Inner core is a festoon with queen at the centre – 20C (1)
11. Bees move slowly over frame consuming stores (1)
12. So shiver using indirect flight muscles (1)

Question 127

19 Drone layers

What are the causes of drone laying queen?

Answer 127

• Shortage of sperm - due to inadequate mating
• Shortage of sperm - due to age
• Disabilty - physically unable to fertilise eggs
• Genetic fault

Question 128

What are the signs of drone laying queen?

Answer 128

• Queen present
• Worker cells with drone cappings
• Stunted drones
• Good brood pattern
• During the season, Q produces small areas of drone brood with patches of drones
• As season progresses, beekeeper sees an increase in drone brood with SOME worker brood

Question 129

Describe treatment of a drone laying queen

Answer 129

• Remove drone layer and requeen
• Remove drone layer and unite.

Question 130

19 Laying workers

What are the causes of a laying worker? 4

Answer 130

• Queenless 21 days +
• No QS - so ovaries develop
• No brood pheramones - so ovaries develop
• No fertilised eggs to raise an emergency Q

Question 131

19 Describe signs of a laying worker

Answer 131

• Drones in worker cells
• Stunted drones
• Scattered laying pattern
• Workers lay more than one egg in a cell
• Colony endeavours to charge Q cells
• Once workers have begun laying eggs, colonies become more aggressive and there is fighting between workers

Question 132

What is the treatment for a laying worker

Answer 132

• Old bees so not much use to anyone
• Requeening hard as usually kill new Q
• If united, will kill Q in united colony
• Smoke, immediately shake out hive at bottom of the garden and allow to find own new homes.

Question 133

19 Laying worker

The sequence of events in a colony of a European strain of honeybee when a colony becomes queenless the beekeeper does not intervene. 9+1

Answer 133

1. Bees know they are queenless within 30 minutes.
2. With no queen and eventually no brood pheromones to inhibit their development, both ovaries and mandibular glands of workers enlarge
3. After 21 days after the queen has been lost, workers begin laying eggs.
4. Workers lay multiple unfertilised eggs haphazardly all over the place, producing only drones which are stunted because they are laid in worker cells, which are too small for proper development.
5. Once workers have begun laying eggs, colonies become more aggressive and there is fighting between workers
6. Sometimes one laying worker will develop a higher level of queenliness and be treated as a false queen who is attended by a retinue of workers who will groom her and lick her
7. This may be because her Dufour gland is producing more queen-like secretions.
8. Because there is no queen substance, workers stop foraging properly, and because there is almost no brood to feed, workers’ hypopharyngeal glands plump up, so they have greater longevity, although they will probably not survive the winter.
9. A colony with laying workers will generally not accept a new queen, and will kill an incumbent queen if united. The colony is therefore doomed.
10. Ultimately colonies die.

Question 134

19 Laying workers

The circumstances and likely causes that can lead to a colony becoming hopelessly queenless. 4

Answer 134

1. A new queen has emerged and fails to get mated within 20 days or gets killed during mating flight
2. The beekeeper is too assiduous in removing queen cells when doing an artificial swarm, possibly leaving non-viable queen cells and no eggs/2-day larva to raise an emergency queen.
3. After requeening when the colony rejects the new queen
4. If a colony becomes queenless, it may produce laying workers while the keeper is waiting for a new queen to arrive and reject the new queen.
5. Disease (eg Nosema) with Black Queen Cell Virus subsequently affecting the queen cells.

Question 135

20 disease behaviour

Sacbrood

Answer 135

1. No outward symptoms in adult bees
2. Symptoms in brood – can’t shed last skin during pupation
3. Hygienic behaviour -> leads to removal of infected larva
4. Nurse bees become infected by ingesting virus during removal or by feeding on contaminated food sources
5. Virus is then spread to larvae during nursing
6. Infected adults do not nurse larva for long
7. Become foragers earlier but rarely collect pollen, which they would contaminate

Question 136

KITESH

20 disease behaviour

CBPV

Answer 136

• KITESH
• K wings so can’t fly and crawl on grass stalks
• Isolation of infected bees
• Trembling
• Ejected on return by guards
• Smoking doesn’t send them down
• Hairless and shiny

Question 137

20 disease behaviour

nosema and its impact on behaviour and function of bees 11

Answer 137

1. No apparent symptoms but longevity halves in spring and summer bees
2. Affected bees can’t absorb protein
3. Hypopharangeal glands do not develop fully so they are unable to feed brood
4. So becomes a guard or forager earlier than normal
5. Internally, epithelial cells producing enzymes of ventriculus affected or may cease
6. In winter there is less protein stored in fat bodies so longevity affected
7. In winter, rectal contents may increase due to increase in water
8. This may be voided in the hive dring winter/bad weather - dysentery - which can fouls hive and cause further contamination through the colony as the bees clean the hive.
9. May infect queen (unusual), who stops laying
10. May cause supercedure
11. Queen pupa may suffer from Black Queen Virus so colony can become queenless
12. Can lead to reduction of adult bees until point of colony collapse

Question 138

20 Disease behaviour

Chalk brood

Answer 138

• No symptoms
• Hygienic behaviour - uncap and clean out affected brood
• May abscond in bad infestation - don’t see them for dust

Question 139

LRSL by ELSAB

20 disease behaviour

General diseases - give over view

Answer 139

• Pathogens affect
  
  • Life cycle
  • Reduce numbers
  • Strength
  • Longevity
• E Sometimes sick bees ejected or refused reentry
• L Lack of fit young bees reduces abilty to forage
• S Queen may be superceded (nosema)
• A Colonies may abscond (chalk brood sac brood)
• May prevent proper colony build up in spring (nosema)

Question 140

20 disease behaviour

Mice 3

Answer 140

• Problem in autumn winter because they enter an occupied hive and make a nest
• Rest of the year they would be stung - vibration and scent
• Bees Co-exist because bees remain in their cluster but cluster will move away from mouse nest
• May abscond

Question 141

20 disease behaviour

AFB

Answer 141

• may uncap sealed brood but unable to remove scales from cells
• workers attempt to clean out cells and become contaminated with spores
• queen doesn’t lay in dirty cells - pepperpot brood patterm

Question 142

20 Pest behaviour

Acarine

Answer 142

• No defensive mechanism
• Affected bees
  
  • tremble
  • K wing
  • Crawl on stalks

Question 143

20 Pest behaviour

What are the 14 general pests?

Answer 143

1. Acarine no defence
2. Varroa - no defence - grooming
3. Tropilalapse - notifiable - no defence
4. small hive beetle - notifiable - no defence
5. Asian Hornet - notifiable - no defence
6. Wasps defensive behaviour at entrance and inside hive
7. hornets - defensive behaviour at entrance and inside hive
8. Ants - turn back and kick up hind legs
9. Wax moth - sting and coexist
10. Mice - sting/enbalm/coexist
11. Woodpeckers no defence in winter
12. Badgers make bees defensive
13. Cattle can cause defensive behaviour - fence off
14. Vandals - sting

Question 144

20 Pest behaviour

Moths

Answer 144

• Not a predetor but a parasite
• Co-exist although they may treat incoming moths as introducers and maul and sting it
• May cause bald brood, destroy comb and brood
• Bees will remove larva
• Reduce life span of bees
• Moth faeces affects pupation and results in bees with v short bodies

Question 145

20 Pest behaviour

Varroa

Answer 145

• No defensive mechanisms
• Some strains groom themselves better than others
• V coexisting in pupating cells - drone=kairomone
• Bad infestation:
  
  • B bald brood
  • A Absconding
  • D DWV Deformed wing virus
  • W Weaker, deformed adults
  • O orientation impacted

Question 146

20 Pest behaviour

Small hive beetle

Answer 146

• Few defences so SHB can reproduce easily
• Strong colonies can remove larva
• Cannot expel adults (exoskelton too hard and beetles scurry about)
• Confine adults to edges of frames
• However, they feed SHB in response to begging behaviour

Question 147

Describe the worker and brood population throughout the year, assuming the colony does not swarm (7)

Answer 147

Winter to Spring
* Colony population will be winter bees. Minimum 5,000, typically 10,000 to 15,000.
* Little brood. Queen laying but only as much as stores and temperature allows.
* Bees in cluster loosening on warmer days to allow cleansing flights and collection of water.

Spring
* Queen’s laying rate increases as days lengthen but proportional to availability of nectar and pollen.
* Winter bees will be dying. For a few weeks there will be more brood than worker bees.
* As brood emerges, worker numbers increase and forage becomes available queen laying can increase to 1,500 eggs per day.

Early Summer
* With a good flow the hive could contain 40-50,000 bees.
* A dry spell could cause a break in forage with the queen going off lay. Reserve stores will be reduced.

Late Summer.
* Days shorten. Queen reduces laying. Good flow of nectar.
* Less brood to tend so more nectar is converted to honey as winter stores are built up.

Summer into Autumn
* Queen reduces laying further.
* Brood will develop into winter bees gorging pollen to build up fat bodies.
* Summer bees slowly die off.
* As temperature drops, days shorten and forage more or less ceases, egg laying may cease.
* Colony goes into cluster.

Question 148

Give reasons for the seasonal variations in the populations of drones and workers (8)

Answer 148

Drones
* Main purpose is to mate with a virgin queen.
* No drones in the colony over winter.
* March/April drone brood starts to appear.
* Colony will have mature drones before any swarm is issued. At least a 36 day lead time is required.
* Mature drones leave the colony every afternoon for Drone Congregation Areas as they seek to mate with virgin queens.
* Once a colony has swarmed and has a new mated queen that the colony is happy with then drones will be progressively barred from re-entering the hive and/or evicted.
* If the queen is suspect and supersedure is a possibility drones may have a stay of execution. But ultimately they will be evicted.

Workers
* Main role for workers in winter is to ensure the hive remains at an acceptable temperature, humidity and ventilated depending on external conditions and the amount of brood being produced.
* As days lengthen winter bees start to forage and have to tend increasing amount of brood. They begin to die off.
* Brood now exceed the number of workers for a few weeks until the new brood emerges.
* If there is congestion in the brood nest area, lots of young bees, a good flow of forage then the colony could swarm. Up to 50% of the workers could leave with the old queen. If there is a cast swarm a further 20% of the workers could leave.
* Swarm or no swarm, after the summer solstice the main effort is to build up stores so that the colony survives the winter months.

Question 149

Describe briefly the behaviour trait known as “worker policing”. (4)

Answer 149

In a queen right colony workers check the comb and eat worker-laid eggs and show aggression towards laying workers.
If a colony is queen less this behaviour ceases.
Queens are thought to mark their eggs with an extract from the Dufours gland situated in the sting chamber, which enables the normal workers to identify eggs laid by the queen.

Question 150

How is worker policing of benefit to a queen right colony? (11)

Answer 150

Worker laid eggs are a good source of protein, so a food supplement for the worker police.
In an emergency normal workers will ignore this signal and move eggs within the colony, e.g. into queen cups and supersedure cells.

In all colonies it is estimated that between 0.01% and 0.1% of workers lay eggs. This could account for up to 7% of all male eggs laid.
Worker laid eggs produce viable drones but they are smaller.
A colony has to support its drones so there is no advantage in having more drones than necessary. Worker policing helps control the numbers.

The colony benefits because queen laid drones are closer relatives than worker laid drones.
From the queen’s perspective, sons are 0.5 compared with grandsons that are 0.25.
From the police worker’s perspective, brothers are 0.25 while nephews are 0.15.

By contrast in a hopelessly queen less colony all the eggs laid by workers will be male. Laying worker numbers increase to between 5-24%. One laying worker can become dominant and will be treated like a queen. As the normal worker force diminishes so the colony slow dwindles and dies out.

Question 151

Name 5 different dances used by honey bees (5)

Answer 151

Choose from:
* Round
* Waggle
* Transition
* Dorsoventral abdominal vibrating dance (DVAV)
* Trembling
* Jostling run
* Spasmodic
* Buzzing run and shaking dance.

Question 152

Briefly describe two of the dances used in foraging (10)

Answer 152

Round
* Indicates forage within 100m of hive
* the bees recruited are left to discover the direction of the forage.
* To assist in the “discovery” the dancer will exchange
nectar with awaiting workers before and after the dance so that they can employ the odour of the
forage to orientate.
* A worker will only be inclined to do a round dance if the sugar content is high in, plus higher sugar content is also demonstrated by more vigorous dances (more
pronounced abdominal vibrations).

Wagtail
* Figure of eight waggle dance recruits foragers to nectar and pollen sources
* The dancer will dance in a straight line whilst shaking the body vigorously from side to side and buzzing, at the end of the straight section
* the worker will loop in one direction back to the start of the line, repeat the waggle and loop the other direction.
* During the dance the dancer will stop and exchange nectar with nearby workers, sometimes this is initiated by the dancer other times by the dance followers “begging” by means of a squeaking sound.
* Direction of the of the forage is indicated by the angle of the straight part of the dance to the vertical plane of the comb, this deviation is the angle in relation to the sun. The dance is normally in the vertical plane however if the dance is on the horizontal plane the direction of the straight line is the direction of the forage.
* The distance to the forage is indicated by the length of the straight line, the time of waggle and buzz, and the
total duration of the dance
* forage further away means a slower dance and slower the waggles. Distance is not a measure of physical length, rather energy required so that workers know how much food is required in their crop to complete the forage.

Question 153

List the behavioural events, which occur in a queen’s adult life between her emergence from the cell and the laying of good worker pattern. (15)

Answer 153

1. Initially the queen is left to care for herself while her glands, flight muscles, ovaries and spermatheca develop.
2. On emergence she will patrol the comb looking for other virgin queens.
3. She will toot in an attempt to identify mature queens yet to emerge. These mature queens will answer with a quack.
4. The emerged virgin queen will kill the queens in cells and if necessary fight with emerged queens.
5. After about three days the workers become increasingly aggressive towards the virgin queen. They will chase her round the comb and bite her.
6. By Day 5 she should be ready to mate. She will take a series of short orientation flights. Then, if the weather is suitable, she will set off on a series of mating flights.
7. She locates and enters a drone congregation area where she may mate with between 15 and 20 drones though not always in a single flight.
8. Back in the colony her mating sign will be removed by the workers, who will now give her their full attention. They will form a court of evolving workers who will feed and groom her.
9. She needs a couple of days while up to 5 million sperm are transferred from her oviducts to her spermatheca. Excess sperm will be discarded.
10. She will now move to the centre of a comb where cells have been polished ready for the queen to lay.
    She will measure the size of the cell with her front legs, withdraws, turns and lays an egg in the bottom of the cell. Initially she may lay more than one egg per cell but she will soon be laying normally.
11. She will lay from the centre outwards in concentric circles so producing a good pattern of worker comb.

Question 154

List nine tasks which may be performed by a worker after she emerges from a cell. These should be in chronological order along with approximate age. (9)

Answer 154

Task/Days of development
1 Cell cleaning 1-5
2 Capping Brood 2-8
3 Tending young brood 4-6
4 Tending old brood 7-12
5 Tending queen 7-15
6 Receiving nectar 10-18
7 Receiving pollen 10-16
8 Packing pollen 10-16
9 Comb building 12-20
10 Ventilating 13-18
11 Guarding 12-25
12 Foraging 21 - death
Note: series 3 to 9 are interchangeable depending on the needs of the colony

Question 155

No No Super PAWD

What conditions can cause variation in tasks? (6)

No No Super Pests Ancestry Weather Disturbance

Answer 155

Weather
Demand for food nectar and/or pollen.
Availability of food nectar and pollen
Swarming
Supersedure
Queenless colony
Demand for water
Time of year
Pests and pathogens.

Question 156

What is piping and how is it used within the colony? (11)

Answer 156

• Toots and quacks last several seconds and are not continuous.
• Worker piping is a single pulse of sound lasting less than a second.

1. Queen emits a series of pulsed high pitched sounds by pressing the thorax against the comb and operating her wing beating mechanism without beating her wings.
2. After a swarm has issued, the first queen to emerge announces her presence by tooting and releasing pheromone(s).
3. Mature queens, still confined in queen cells, answer with another distinct piping sound known as ‘quacking’.
4. Tooting and quacking is broadcast in the bee nest as vibrations on the comb.
5. Queens pipe only in the context of reproduction.
6. Tooting delays mature queens emerging for some hours. This delay allows house bees to inspect queen cells containing mature queens and sealing up any holes they may have made as they seek to emerge. This action can delay emergence by days.
7. Queens have also been known to pipe before going on a mating flight. This may protect her from being roughly handled by workers as the workers freeze when the queen pipes.

Question 157

Pipers are excited - let’s go in 1hr

How does the acoustic behaviour of honey bees impact on swarming? (4)

Answer 157

Mated queens sometimes pipe before swarming.
Worker piping differs from the queen as it occurs when a worker mounts and presses down on a fellow worker with her wings folded.
It occurs in swarms as they prepare to lift off and fly to their new home.
Pipers are excited bees that scramble through the swarm cluster, mount a fellow worker, pipe and move on. This is done by the scout bees and alerts the non-scouts to warm up their flight muscles ready for flight.
It starts about an hour before the swarm lifts off and reaches a climax as the moment arises.

There is another slightly different signal that inhibits actively dancing scout bees and so delays lift off.

Question 158

What senses do honey bees use to navigate while foraging (3)

Answer 158

• Visual
• Olfactory/smell
• Magnetic

Question 159

UV waves

How do bees orientate in cloudy sky when the sun is not visible. (2)

Answer 159

• Bees then rely on UV light waves rather than those of the visible spectrum.
• UV waves penetrate cloud, providing the cloud is not too dense.
• Foragers use the waves to locate the compass direction of the sun.

Question 160

Describe a nectar forager’s behaviour from when she watches a waggle dance to when she has unloaded the nectar. (10)

Answer 160

• The waggle dance will have given the forager the distance and direction to the nectar source. She may also have exchanged a drop of nectar to acquire the scent of the nectar.
• Having arrived in the area, she will identify the source by smell and will memorise the striation of the petals for future reference.
• Places her proboscis in the corolla and pumps the nectar up into her mouth.
• It may require her to visit up to 1,000 flowers to fill her honey crop. A load would be 30 to 50mg. Much depends on the flower, time of day, weather -wet or dry and the distance from the hive.
• With a high yielding patch she will travel a short distance between flowers, while in a low yielding area she will change direction more often and travel further.
• Uses landmarks to return to the hive.
• Met near the entrance by house bees. Transfers her load by trophallaxis to one or more house bees.
• In a good flow she will visit the same crop at the same time of day.
• She will enter and leave the source via the same place on each visit.
• For preference she will visit crops that are closer to the hive than others but the best value crops will be visited regardless of distance.
• It is a fine balance between the load carried and the distance travelled. A forager has a finite number of forager miles before she is worn out and dies.

Question 161

Describe the characteristic ‘stance’ of a guard bee ‘on patrol’ at the entrance of a hive. (2)

Answer 161

Standing on four legs; mandibles apart; antennae forward: wings open.

Question 162

Describe how guard bees at the entrance recognise a nest mate. (2)

Answer 162

Each colony has a distinct odour. The guard will use her antennae to check the nest mate as it enters the hive. If it is a match then entry is granted.

Question 163

Having recognised a potential robber or predator, describe the aggressive behaviour of the guard bee. (3)

Answer 163

• Rises onto her back legs, mandibles open and releases the pheromone 2-heptanone.
• If no heed is taken, the guard will fight and try to sting the intruder while releasing stronger pheromone. This is designed to summon reinforcements.
• Robbers don’t submit, they either run or fly away. If caught and stung then the first sting contains isopentyl acetate which guides others to sting in the same spot.

Question 164

List three stimuli likely to cause guard bees to sting an intruder. (3)

Answer 164

• Vibration.
• Scent.
• Movement.

Question 165

Worker bees may inadvertently drift to the entrance of an adjacent hive. Describe the behaviour of a young ‘intruder’ returning from an orientation flight. (2)

Answer 165

Becomes submissive. Curls up with legs and and abdomen trucked in.

Question 166

Worker bees may inadvertently drift to the entrance of an adjacent hive. Describe the behaviour of a nectar or pollen laden ‘Intruder’.

Answer 166

If she’s confident the guard will let her pass but may follow her in - just in case.

Question 167

Give the processes involved in converting nectar into honey. (5)

Answer 167

• Forager passes the nectar to house bee(s) by trophallaxis near the hive entrance.
• House bees place honey in cells in the honey storage area.
• Storage area bees take drops of honey and lash it on their tongues so exposing it to the heat of the hive (34.5℃). This happens many times and reduces the water content by up to 15%.
• Further evaporation is done by the constant circulation of air in the hive by bees fanning their wings inside the hive.
• Honey capped when water content less than 20%.

Question 168

Describe water collection in terms of preferred sites and regularity of visits. (3)

Answer 168

• Close to the colony.
• Stagnant water with organic matter preferred.
• 10 foraging trips per bee per day is the norm but up to 100 have been observed. Turn round time less than 10 minutes.

Question 169

Describe the interrelationship between nectar, honey and water, in the colony, in summer and winter. (5)

Answer 169

• In summer the water content of nectar coming into the hive varies between 30% and 70%. When it is being passed around the hive or stored there is usually sufficient ready to consume to keep the bees’ energy levels up. Water is also readily accessible if required.
• In winter it is too cold to collect water but bees expel water through their trachea which condenses on hive surfaces and can be recycled.
  Some stores are left uncapped and the hygroscopic nature of honey absorbs surface water increasing water content so that it is available as stores for consumption.

Question 170

Briefly describe how nectar is collected and transported to the hive. (2)

Answer 170

• Nectar is sucked via the proboscis.
• Glossa opens at the top into the ciborium. Ciborium muscles expand so liquid is drawn up into the mouthparts.
• Liquid passes to the pharynx where enzymes are added then moves along the oesophagus to the honey crop where it is stored.
• Passed to a house bee by trophallaxis just inside the hive entrance.

Question 171

How is nectar processed to provide a food store for later use? (8)

Answer 171

Two processes:

1. Evaporation of water to reduce the content to 17%-18%.
   Water is evaporated by the actions of the house bees.
   A drop of water is placed on the bee’s tongue, lashed in the air within the hive and swallowed. This is repeated many times resulting in the water content being reduced by 15%.
   Drops of honey are then placed into the top edge of an empty cell or an unsealed cell with nectar already in place.
   Bees create a current of air through the nest which helps further reduce the water content.
2. Chemical changes through the addition of enzymes.
   The enzymes invertase and glucose oxidase are produced in the bee’s hypophyngeal glands.
   Invertase breaks down the sucrose (a disaccharide) into glucose and sucrose (monosaccharides).
   C12 H22 O11 + H2O = C6 H12 O6 = C6 H12 O6

Glucose oxidase breaks down the glucose into Gluconic acid and hydrogen peroxide, which destroys bacteria. C6H12O7 + H2O2

Nectar is now ready to store at less than 20%.

Question 172

ECAVES

To what use is propolis put in the colony? (5)

Answer 172

• Filling cracks
• Reducing entrances
• Smoothing surfaces.
• Varnish cells before use.
• Embalm large intruders that cannot be evicted (Mice).
• Used with wax to strengthen comb where it joins the wall.

Question 173

Outline four different ways foragers collect pollen from the anthers of flowers. (4)

Answer 173

Open flowers.
* Bee bites the anther with her mandibles and uses her forelegs to pull them toward her body.

Tubular flowers.
* Bee inserts proboscis into the corolla searching for nectar. Pollen is collected inadvertently gathering on the hairs as they brush past the anthers.

Closed flowers.
* Bee forces petals apart with her forelegs and gathers pollen on her forelegs and mouthparts.

Spike or Catkin flower.
* Bee runs along the spikes shaking off pollen on to her body hairs where they stay owing to electrostatic effect.

Question 174

Why is pollen turned into bee bread and how is this achieved? (4)

Answer 174

Pollen is a vital source of protein and amino acids required to keep bees healthy. Some needs to be stored for future use when fresh pollen is not available. The process ensures that stored pollen stays moist and usable.

1. Forager delivers pollen to the cell. Backs in and uses one back leg to scrape pollen off the other back leg and then repeats.
2. The forager will have moistened her load with some regurgitated nectar that contains enzymes.
3. House bees pack the pollen down with their head and front feet adding further secretions from hypopharyngeal glands. Also contains lactic acid that starts a fermentation process that breaks down the outer shell of the pollen, which aids subsequent digestion.
4. Finally a thin layer of nectar is added to keep the bee bread free of pathogens.

Question 175

State two uses of pollen by bees (2)

Answer 175

• Main source of protein
• Required to develop their glands and other internal structures on emergence.
• Key ingredient of food for feeding worker and drone larvae in the latter stages.
• Mixing with wax to make brood cappings.
• Needed by workers in autumn to build up their fat bodies so that they can survive through winter.

Question 176

What factors within a hive regulate the amount of pollen collected? (2)

Answer 176

• Amount of larvae to be fed.
• Numbers of bees emerging and needing pollen in order to develop.
• Time of year and general state of the colony.

Question 177

Outline the feedback mechanism involved in the amount of pollen collected. (3)

Answer 177

• Proportion of bees foraging for pollen is determined by the amount of brood in the colony.
• Smell of the brood and contact with nurse bees tending the brood are each partly responsible.
• Access to the brood is most important. Larvae are more important than pupae.
• If brood is inaccessible then pollen collection decreases with bees foraging for nectar instead.
• Queenless colonies, with or without brood, result in increased nectar intake and less pollen.

Question 178

Name five typical sources of propolis. (5)

Answer 178

1. Horse chestnut,
2. alder,
3. poplar,
4. prunus
5. conifers.

Question 179

How is propolis collected by honey bees? (2)

Answer 179

• Foragers use their mandibles to scrape or pull resin from plant.
• Load is passed to the forelegs then transferred to the middle leg.
• It is then pressed into the pollen basket of the hind leg on the same side, i.e. different from pollen.

Question 180

How is propolis unloaded in the hive? (3)

Answer 180

1.Forager goes to where the propolis is required.
2.House bees nibble propolis from the forager.
3.House bees then spread it into place.

Question 181

Uses of pollen

Answer 181

• Consumed by nurse bees to produce brood food and royal jelly
• Fed to worker larvae
• Mixing with wax to make brood cell cappings porous
• Consumption by workers to re-activate their hypopharyngeal glands in order to feed
  larvae
• Consumed by workers in autumn to build up fat bodies and hypopharyngeal glands to
  extend life in order to over winter
• Consumed by workers for the orderly development of their glands, particularly the wax glands

Question 182

How much propolis is required
normally by a colony per annum? (1)

Answer 182

100g.

Question 183

Outline the swarming related activities that take place in the colony a few days before a swarm leaves. (10

Answer 183

• Worker brood rearing at its peak causes colony to become congested as worker population grows rapidly.
• Prompts the start of queen rearing. Queen cups constructed and may increase in number.
• Queen lays in queen cups.
• Worker bees start feeding some with royal jelly. Queen rearing begun.
• 8 days until swarm could issue.
• Up to now queen has been fed more frequently and laid more eggs. Over next few days she will be shaken by workers causing her to keep moving round the hive, laying fewer eggs and losing up to 25% of her body weight so that she is fit to fly.
• Not knowing when conditions will be suitable to swarm, the workers start feeding up on nectar and pollen. This may start up to 10 days before the swarm issues.
• Scout bees start the search for a new nest about three days before swarm leaves.
• Workers quieten down. Clusters seen festooned on the bottom of comb.
• Few hours before the swarm, scout bees start piping and shaking the workers, warning them to start warming up their flight muscles.
• Shaking signal stops and scouts and their recruits run back and forth in waves, buzzing to excite other workers.
• Queen chased, bitten and pulled along with excited workers.

Swarm issues.

Question 184

Describe the behaviour of a colony if it is to supersede the queen. (6)

Answer 184

• Assuming perfect supersedure when the old queen remains in the hive and continues to lay.
• Colony carries on more or less as normal. Brood production may slow if the queen is failing but it continues.
• No loss of honey production.
• Queen cells drawn out from worker cells and fewer in number, less than six.
• Queen cells may be away from the main brood area. All laid up at the same time. (Often wonder if the queen lays these eggs or the workers select and move.)
• Workers continue to look after the queen, tend brood etc with no actions towards preparing to swarm.
• Once new queen has emerged, mated, started laying and the workers are happy with her, in due course the old queen will quietly disappear.

Question 185

Describe T Seeley’s experiments on Appledore Island. Include how a swarm behaves when selecting a new home. (14)

Answer 185

Appledore Island chosen as no native bees and no large trees suitable for accommodating a swarm.
The experiments built on earlier work undertaken by Lindauer in Germany.

Experiment 1. To investigate the way way honey bees investigate a prospective nest.

• Empty light proof box placed in a hut with access to the outside. One side of the box was red glass. The inside of the box was gridded.
• Seeley and had team released a ‘swarm’.
• Every visit was recorded. How long inspections took: average 37 minutes.
• A mix of short visits inside the hive and out plus some rest periods then back to the parent colony. Time inside the hive was spent walking and flying.

Experiment 2 was to decide how bees choose a cavity of the desired volume and entrance size.
* Bees presented with a box with a movable inner wall so that volume could be adjusted to 15l or 40l. Entrance
15cm2 to 60cm2. Preferred option was 40l volume and the smaller entrance.
* Also discovered that light levels made no difference, the key was how far the bees walked inside the hive.

Lindauer had discovered that the scout bees dance on the outside of the swarm cluster to advertise the new site each has discovered. They recruit more scouts who may or may not endorse a site. Eventually a majority opt for one site but how does this occur. Seeley’s next experiments using more sophisticated equipment - video, set out to answer the question.
* Using video Seeley et al set about filming the scout bees’ dances throughout a swarm’s choice of its new home. They filmed over two and bit days then studiously studied the video noting how many bees were involved, how many dances they performed for each potential nest site until the swarm took off.
* These experiments were followed up with more work on Appledore to refine the findings such as the preferred time of day, how scouts either gave up or switched allegiance. Also to discover how the swarm remained cohesive as it moved to the new nest site.

Question 186

What is supersedure? (1)

Answer 186

The replacement of a resident queen by her daughters without swarming (thus ensuring the long term survival of the colony).

Question 187

State the difference between perfect and imperfect supersedure. (2)

Answer 187

• Perfect supersedure is when the old queen remains active in the colony and in lay until the new queen is mated and laying to the colony’s satisfaction.
• Imperfect supersedure is when the queen disappears for what ever reason and there is no overlap.

Question 188

ORCHIB

What can cause a colony to supersede? (6)

Answer 188

• Older queen producing less queen substance.
• Reduced rate of lay of fertile eggs.
• Colonies in large brood chambers are more likely to supersede than swarm.
• High drone population.
• Injured queen
• Queen runs out of sperm or becomes Barren.

Question 189

List the benefits of supersedure. (6)

Answer 189

• Colony gets a new queen.
• With perfect supersedure there is no break in egg laying so colony maintains it’s strength, produces stores and has better chance of overwintering.
• No need to relocate with all the danger that entails.
• Honey production maintained.
• With imperfect supersedure there may be a brood break that could help with varroa control.
• Non swarmy colony good for breeding.

Question 190

State the conditions within a colony and give the specific external stimuli that can lead to swarming. (9)

Answer 190

Internally
* Colony size around 12,000 bees stimulates queen rearing with swarm issuing when 20,000 bees.
* Increased brood production and a high number of nurse bees caring for the brood leads to congestion in the brood nest.
* Queen’s movement is restricted and her substance is not spread around the hive. Bees are concerned that queen is failing and start building queen cups.
* Winter bees have died off so the average age of the workers is 8 to 12 days, they are ready to build wax and tend the brood in the new nest.
* Need to be mature drones in the colony.

Externally
* Flow of both pollen and nectar so swarm can gorge prior to leaving and have food in abundance on arrival before new nest is established and ready to take stores.
* Flow also important for parent colony that will have far fewer bees
* Weather is warm and calm.

Question 191

Describe the changes in the colony in the weeks and days prior to swarming. (7)

Answer 191

• Drones produced to emerge up to a month before swarm issues.
• Queen rearing initiated when 12,000 bees. Queen cups constructed.
• Queen cells laid up 8 days before swarm can issue (in most cases).
• Queen is chased around. She loses 25% of her bodyweight so that she is fit to fly.
• Bees gorge up on honey and pollen.
• House bees stop accepting nectar from foragers.
• Scout bees start looking for new nest site.
• Scout bees start shaking house bees to start preparing them to fly. They build up their body temperature.
• First queen cell is capped.
• Wave of bees running through the hive increases excitement immediately prior to swarming.
• Swarm issues.

Question 192

Why is a good food supply required to support a swarm? (3)

Answer 192

• Pollen is needed so that wax glands are fully developed and ready to produce/ build comb on arrival in new site.
• Nectar is needed to provide energy for the flight and support the bees for three days on arrival in new site.
• Queen needs to be fed and built up again ready to start laying.

Question 193

Name three behaviours that may be observed in the colony just before the swarm leaves the hive. (3)

Answer 193

• Bees feeding up then festooning and resting at the bottom of the comb.
• Shake dance by the scouts and others as they gee up the house bees ready to fly.
• Scouts and some recruits rushing through the hive to stimulate action.
• Queen being nipped by house bees.

Question 194

40, 1 south, d, 15

List five qualities preferred by a swarm for the site of a new nest. (5)

Answer 194

1. 40l volume
2. 1 to 5 m above the ground.
3. South facing entrance.
4. Dry not drafty.
5. Entrance at bottom of cavity no more that 15cm2.

Question 195

How do scout bees tell the swarm where the new nest is and guide bees to it? (3)

Answer 195

• Scout bees are all agreed as to the preferred site following waggle dances on the outside of the clustered swarm.
• Scouts fly backwards and forwards through the swarm in flight keeping the swarm together.
• Nasanov pheromone emitted for other bees to follow.
• First scouts to arrive at the new site will fan the Nasanov pheromone so bees know they have arrived.

Question 196

BNFDFV

How do honey bees use wax comb? (6)

Answer 196

• To develop Brood - eggs, larvae and pupae. Also queen cells.
• To process Nectar.
• To store honey and pollen (Food).
• Platform for Dancing.
• Transmission of queen substance (Footprint).
• Queen piping Vibrations transmitted by comb as are worker dorsoventral oscillations.

Question 197

Need Flow Age Rate Swarm

What factors affect comb building? (4)

Answer 197

• Must be a need as processing wax requires much energy. Need for brood rearing, nectar ripening, pollen and honey storage.
• Needs to be a flow as lots of nectar and pollen needed, viz 6-8kg of honey to produce 1kg of wax.
• Age of the bees 10-20 days as wax producers.
• Queen’s laying rate will also be governed by availability of stores.
• Nest building after a swarm.

Question 198

Describe how bees construct new comb. (10)

Answer 198

• Need nest temperature of around 35℃.
• Need bees aged between 10 and 20 days old provided with ample stores to produce wax.
• Bees hang together in chains (festoon) and exude tiny transparent scales of wax from their wax glands.
• These are picked up by the pollen brush on the inside of the hind legs, transferred to the front legs, moistened with saliva and kneaded by front legs to make pliable.
• Wax is added to work of other bees and becomes part of the comb.
• Starts in several areas at the top so a ridge of wax is formed. This is drawn out and down.
• Bees have hairs on the plates where the head meets the thorax.
• These hairs are connected to nerves so when the position of the bee relative to gravity changes, head and thorax change position, the hairs move, nerves pick up the signal allowing the bee to orientate to its position in space.
• The bee knows when it is horizontal, vertical or at an angle allowing it to shape the cells and build the comb.
  *

Question 199

Describe the structure of new comb. (10)

Answer 199

• Hexagonal cells offset from the cell on the reverse side.
• 0.07mm thick
• At an angle of 13° to the horizontal.
• Worker cell 5.2 to 5.4mm wide.
• Drone 6.2 to 6.4 mm wide.
• Angle between cell wall is 120°.
• Composed entirely of beeswax.
• Propolis used to strengthen the comb bases.
• Distance between comb is 0.95cm.
• White in colour when fresh

Question 200

Name one factor which stimulates a queen to increase her laying rate.

Answer 200

Increased day length / increased income

Question 201

What is the average weight of pollen load carried by a honeybee?

Answer 201

8mg x 2 / 16mg

Question 202

What term is used for the mechanism of controlling the environment within the hive?

Answer 202

Homeostasis

Question 203

Describe the difference between drifting and robbing bees. 4

Answer 203

Robbing
* zig-zag flight pattern across the entrance.
Drifting
* enters confidently with load of nectar and/or pollen,
* becomes submissive when inspected by the guard bee curling her legs and tucking in the abdomen,
* may release a sample of nectar.

Question 204

List the duties that a worker honeybee undertakes between her emergence in October until her death? 8

Answer 204

1. Little or no brood to rear and no comb to build.
2. Large amounts protein in their fat bodies due to consumption of pollen that isn’t used to produce brood food or wax (life of bee is directly proportional to amount of pollen consumed).
3. Need to survive until following spring so they can feed new brood in spring ~24weeks.
4. Cluster tightens when ambient temp drops below 14°C to maintain internal temp of
   cluster~20-30°C (provided no brood present).
5. Vibrate wing muscles to keep temperature constant.
6. Dilute honey with water from condensation or collected on mild days, not usually able to forage outdoors
7. Uncapping honey to allow hydroscopic absorption of water
8. After ~Dec 21st (shortest day) queen starts laying and winter bees use their stored protein to feed brood.
9. Break in weather 10°C+ take cleansing flights.
10. Early spring on warm days forage mainly for pollen to feed brood with.
11. Start dying off early March – danger time for colony as brood exceeds adult bees.

Question 205

What form of learning relates to orientation flights?

Answer 205

Latent or observatory.

Question 206

A worker honeybee has been given the direction and distance to a new forage source by
observing a dancing bee. List the navigational methods used for determining direction and
distance during the first flight of a new recruit to find the new forage source. (You do not need
to describe any dances) 6 Marks

Answer 206

1. primary navigation tool is the sun.
2. able to compensate for the sun’s movement with passing time.
3. Use polarized light if sun not visible
4. need at least 10% of blue sky for accurate use of polarized light
5. aware of magnetic fields.
6. calculated by amount of energy expended during dance (von Frisch)
7. or using optic flow speed and time, (how quickly image of environment moves across their eyes).

Question 207

What are the factors that allow for efficient use of comb? 5

Answer 207

1. Combs must hang vertically,
2. combs must be parallel to each other,
3. there must be the correct bee space between each comb,
4. the cells need to be the correct size i.e. worker or drone.
5. hexagonal shape efficient
6. Angle of cell retains contents
7. Cells offset on each side for strength

Question 208

What are the triggers for a colony to start preparations for swarming? 5

Answer 208

1. Lack of pheromones –QMP & QTP,
2. hereditary factors,
3. age and condition of the queen,
4. average age of workers, swarming more likely when the average age is low,
5. external conditions – good nectar flow and supply of pollen,
6. rain may also make the colony more congested as the foragers cannot get out.

Question 209

Explain briefly the behaviour of a colony of honeybees from the construction of swarm
cells to the emergence of the first virgin queen. 10

Answer 209

1. Larvae in QCs are fed royal jelly for five days before being capped.
2. Foragers become scouts and investigate new nest sites.
3. Workers reduce the food fed to the queen in order to reduce her weight so that she can fly, this also reduces her egg-laying rate.
4. The queen is kept moving by workers performing the dorso-ventral abdominal vibration dance on her and biting her legs.
5. The workers load up with honey ready to swarm
6. Buzzing runs to prepare to swarm
7. The prime swarm leaves when the first cell is sealed.
8. Workers keep the queen cells warm
9. Workers nibble round the end of the cell to thin
10. Other virgins are kept within cells
11. Virgin queen emerges and pipes

Question 210

Probos-corolla cibarium sucks nectar enzym from hypo oesop to crop poll

A honeybee has landed on a flower. Describe how the nectar is collected and
transported back to the hive. Include changes that may happen to the nectar. 9

Answer 210

1. Her proboscis is inserted into the corolla of the flower
2. and the cibarium is expanded drawing nectar into her mouth.
3. Enzymes are added to the nectar from the hypopharyngeal
4. Glucose oxidase, diastase, and invertase
5. The nectar passes through the oesophagus to the crop (or honey stomach)
6. Pollen is filtered out by the proventriculus
7. which allows a proportion of the nectar to provide fuel for the return flight
8. it is stored in the crop by the forager until she returns to the hive.
9. It takes 30-80 minutes to collect a load of an average of 40 mg.

Question 211

What happens on her return to the hive with a full load of nectar up to the point the
honey cell is sealed? 18

Answer 211

1. Guard bees allowed into hive onto comb
2. If the source of nectar is good and nectar is required, the bee will dance round, sickle or waggle depending on the distance the forage is away from the hive, she may give a sample of nectar during the dance
3. she may have to wait for house bees to come to unload her nectar depending on the
   requirements of the colony at the time,
4. she unloads nectar to between 1 and 3 house bees, by exuding a droplet on her proboscis.
5. The bees touch each other’s antennae during this process (antennate).
6. Once unloaded the forager cleans antennae and wipes her proboscis through her front feet, before flying-off again.
7. The receiver house bee repeatedly exposes a drop of nectar on its partly folded proboscis and exposes it to air circulating in the hive - this could take up to 20 minutes
8. Enzymes continue to be added
9. An increase in air flow through fanning bees aid evaporation.
10. The drop of nectar is then place at the top of a cell where it hangs continuing the evaporation process - this occurs at around 40% water.
11. Once the nectar reaches 20% water (or less) and the cell is filled
12. the cell is sealed with wax to prevent the honey taking up moisture

Question 212

Give the definition and characteristics of a DCA and what attracts the drones. (5)

Answer 212

DCA definition. A discrete aerial site where drones fly anticipating the arrival of virgin queens.

Characteristics.
* 5 to 40 above ground.
* 100 x 200m in area.
* Often bounded by specific geographic features.
* Preferably over flat land but sheltered from the prevailing wind.
* Good light is important.

How drones find a DCA year on year with no previous knowledge is not known. Theories include:

• Topographical features assessed by sight and other senses.
• Distance from the apiary - usually more than 100m.
• Magnetic effects - drones develop large quantities of magnetite in their abdomens, this could be effected by the earth’s magnetic field.

[RAO thought. Drones pick up the virgin queen’s pheromones when she enters the DCA. Successful drones mount the queen and then drop to the ground dead after mating. At the end of the season there could be lots of dead drones in a particular DCA. Colin Butler demonstrated that queen substance can survive the elements for years. While they are on their early orientation flights, is it possible that drones pick up on these latent pheromones?]

Question 213

Describe the mating behaviour of a queen including the age for mating and the role of workers and drones. (10)

Answer 213

• A queen takes about 5 days to mature after emerging from her cell.
• As a virgin she is ignored by the workers at first and has to feed herself.
• As her glands develop and pheromones are emitted, the workers start to pester her and encourage her to go on mating flights.
• Once mated it takes a few days for the sperm to pass from her oviducts into her spermatheca to enable her to start laying.
• Workers may cluster at the entrance and fan their Nasanov glands to help the queen return to her home hive.
• On arriving back from mating the workers will clear her mating sign, form a retinue who will feed, clean and lick her thus distributing her pheromone round the hive.

The queen will have been on up to six mating flights, visiting DCAs that are well away from the hive. Queens fly further to DCAs than drones.

She will mate with up to 20 drones overall preferring to fly on calm, sunny days between 2 and 4pm, though sometimes earlier.

Drones leave the hive earlier, travel to nearby DCAs so that they can stay in the DCA for longer.
When a queen arrives they form a cone behind her.

The fastest approaching from behind and below, attracted by 9-oxodec-2-enoic acid (9ODA) emitted from her mandibular glands, rises above her and settles on her abdomen gripping her with all six legs.

The queen opens her sting chamber, The drone inserts his endophallus and is paralysed. He flips backwards causing ejaculation to occur. The drone falls away leaving the mucus mating sign. The drone dies.

The queen may mate several times on each flight until her oviducts are full.

Mating must take place within three to four weeks of the queen emerging. After that she is unable to mate and becomes a drone laying queen.

Question 214

Give three reasons why a queen might inspect a cell before laying in it (3)

Answer 214

• Is it clean and polished ready to accept an egg?
• What size is it? 5.2mm-5.5mm for a fertilised worker egg, 6.2-6.5mm for a drone egg.
• Is there an egg in there already?

Question 215

Outline the queen’s egg laying behaviour throughout the year with respect to the changing circumstances both within and outside the hive. (8)

Answer 215

• Once mated the queen needs a couple of days for the sperm to move from her oviducts into her spermatheca. She will then start to lay.
• The rate of lay is dependent on the supply of nectar and pollen and the temperature in the brood area (needs to be about 35℃).
• The queen walks over the comb. She measures the size of the cell with her front legs. Backs out, reverses in and places a single egg in the base of the cell. (She may place a couple of eggs in a cell in the early days as she gets the hang of things.)
• For the standard worker cell (5.2mm to 5.5mm) she will lay a fertilised egg. She achieves this by using her valve fold to press against the entrance of the spermathecal duct. A sperm enters the egg.
• For the bigger (6.2 to 6.5mm) cells, usually found on the edge of the brood area, she lays an unfertilised egg that will develop as a drone.
• As the days shorten and winter approaches, food will become scarce and her laying rate will slow and may stop altogether in a really cold spell. Similarly in a period of dearth in the summer she may also go off lay.
• As the days lengthen and we move from winter into spring, her rate of lay will increase peaking in May-June.
• If the colony becomes overcrowded in the swarming period, the queen will be starved so that she loses weight and is fit to fly with the swarm. Egg laying will reduce accordingly only to pick up once the swarm finds a new home and comb is built ready for egg laying to begin.

Question 216

What is a pheromone (2)

Answer 216

A chemical produced by one honey bee which has an effect on the physiology, behaviour or development of another honey bee. (Celia Davis)

Question 217

How does brood pheromone affect behaviour within the colony. (3)

Answer 217

• Brood pheromone suppresses ovarian activity in worker honey bees including when a colony is queen less.
• Enables nurse bees to determine the sex, caste and age of a larva so that it can be fed accordingly.
• Informs the colony when more food is required to feed increasing amount of brood and vice versa.
• Informs nurse bees when to cap a cell.
• Informs nurse bees if the larva is diseased so that it can be removed.

Question 218

Describe how four named worker pheromones affect behaviour within the colony. (8)

Answer 218

When disturbed a worker bee will raise her abdomen and release alarm pheromones by opening her sting chamber and protruding her sting.
It is thought to be produced by two masses of glandular cells lying against the quadrate plates of the sting mechanism. It attracts other bees to support the bee initiating the alarm.

• Mandibular pheromone 2 heptanone repels robbing bees at the hive entrance.
• Nasanov pheromone is used when marking the entrance to a nest. May be witnessed to help a queen to return to her colony after mating.
• A laying worker that becomes a false queen produces 9 ODA which inhibits the ovary development in other workers.
• Trail pheromone. In addition to marking food sources a returning forager will leave a trail pheromone at the entrance to her colony, which is an indicator to other foragers from that colony.

Question 219

When and how does drone pheromone affect behaviour. (2)

Answer 219

• Drone mandibular gland secretions have a role in attracting other drones to DCAs.
• A cuticular pheromone on the drone is used to identify a drone seeking entry to a nest.

Question 220

Name seven brood and adult bee diseases and give their specific effects on bee behaviour.

Answer 220

AFB
* May uncap sealed brood, but cannot remove scale once it is formed.
* Workers attempt to clean out cells and become contaminated with spores
* Queen does not lay in infected cells leading to pepper pot brood.

EFB
* Workers clean out infected larvae.
* Infected larvae demand more food by moving in the cell and are fed more or removed

Chalk brood
Cells are uncapped and chalk brood mummies are removed and left on the hive floor or in front of the hive.

Sacbrood virus
* Workers can uncap and remove the larvae, doing this infects the adult worker bee and although there is no apparent sign of disease
* its life is shortened and therefore it stops feeding larvae sooner
* it becomes a forager earlier than usual,
* few infected adults collect pollen.

Nosema
* The colony dwindles,
* queen may be superseded having stopped laying,
* foragers become disorientated.
* May stop feeding brood which dies.

Chronic bee
paralysis virus

Type 1
* Bees are unable to fly and crawl up grass stems outside the hive with trembling wings and bodies.
* Bees cluster on top bars

Type 2
* Bees nibble the hairs of affected bees and
force them out of the hive and prevent them returning.

Dysentery
Bees defecate in the colony

Question 221

Describe how the population of honeybees and brood in a healthy colony varies during
the year without swarming? 4

Answer 221

1. Jan- March adult numbers greater than brood which is increasing in size
2. April -June brood population overtakes that of adults peaking in June
3. Adult population peaks in July/August
4. Adult population reduces towards year end and brood practically peters out

Question 222

How might the population change with a severe infection of chalkbrood, from the spring
onwards and what would be the impact on the colony and honey harvest? 6

Answer 222

1. Fewer larvae would develop into adults
2. Fewer foragers and less nectar collected.
3. More time would be spent by house bees clearing out infected pupae and infecting the youngest larvae.
4. Eventually, the colony will dwindle out.
5. There will be reduced honey crop for the beekeeper to collect.

Question 223

Forage loads collected

Answer 223

• Nectar 30-50mg
• Water 25mg average
• Pollen 16mg (8mg x2)
• Propolis unknown
• Nectar collection is more energy efficient
  o 8:1 gain pollen
  o 10:1 gain nectar (40% nectar load)

Question 224

Other Bees/Wasps

Answer 224

• Eusocial bee - Buff tailed bumblebee (Bombus terrestris)
• Solitary bee - Ivy bee (Colletes hederae)
• Eusocial wasp - German wasp (Vespula germanica)
• Solitary wasp -European tube wasp (Ancistrocerus gazella)

Question 225

Other Bees and Wasps: Three characteristics of eusociality:

Answer 225

• Individuals of the same species co-operate in caring for the young.
• There is a reproductive division of labour with
  more or less sterile individuals working on
  behalf of the reproductives.
• There is an overlap of at least two generations in
  which the offspring contribute to colony labour

Question 226

Other bees and wasps: Numbers in UK

Answer 226

Bees
* One species of Honey bee
* 24 species of Bumblebee - 18 social and six “cuckoo”
* Approx 225 species of solitary bee.

Wasps
* Six species of social wasps: three Dolichovespula, three
Vespula and two Vespa (if you count Asian Hornet).
* 7,000 species of solitary wasps

Question 227

Eusocial Bee (Bombus Terrestris) - Lifecycle

Answer 227

1. After mating, new queen hibernates for winter and wakes in the spring
2. Queen builds the nest in preparation for egg laying and hatching of larvae
3. Eggs hatch, young worker bees emerge from cocoons
4. Colony developes
5. Founder queen dies, nest developes future males and queens
6. New queens and drones emerge from cocoons and leave the colony to mate

Question 228

Eusocial Bee (Bombus Terrestris) - Interaction with Honeybee

Answer 228

• Nests underground
• Some competition for nectar but bumblebees tend to have longer tongues.
• Far fewer bees per nest
• May pick up diseases from flowers

Question 229

Solitary bee (Collates hederae) - facts

Answer 229

• Last solitary bee to emerge each year. Males late August. Females in mid September to peak with ivy blossom in October (Changing?)
• Males outnumber females
• Nest soil is tamped and smoothed; soil microbes are applied to the walls, polymerises to form tough film.
• Single burrow up to 50cm deep with side branches. Single cell at the endwith the egg laid on a ball of pollen and nectar.
• Egg laid. Close up. Job done.
• Egg hatches. Larva feeds and pupates. Adult emerges next year
• Interaction with Honey Bee: negligible, perhaps competition for food.

Question 230

Common wasp - Vespula vulgaris
German wasp - Vespula germanica

Answer 230

• Queens emerge from hibernation in mid-March
• Workers are present from May to mid November
• Autumn queens leave the nest in September and October
• Males are on the wing from mid- August to mid-November.

Nest vary in size, location and shape:

Germanica nest grey/white. Vulgaris from old wood are brown

Question 231

Vespula germanica

Life Cycle

Answer 231

• Queen searches for and chooses nest site.
• Queen collects and masticates wood into pulp
• Creates cells and lays eggs
• First batch of workers emerge and take over
  nurse and foraging duties
• Adults are omnivores. Take flies, bugs, aphids,
  caterpillars etc.
• Feed chewed up remains to the larvae in the nest
• Larvae secrete sugary substances to feed adults
• As nest dwindles, adults look elsewhere for sugary
  foodstuffs.

Question 232

Eusocial Wasps
Interaction with Honey Bee

Answer 232

Feeds on some shared plants for nectar.

• Can rob and destroy weak honey bee colonies in late
  summer, early autumn

Question 233

Solitary Wasp

Ancistrocerus gazella
Small-notched mason wasp
European tube wasp

Answer 233

• Collects up to 20 caterpillars per nest.
• Lays egg. Seals cell with mud.
• Egg hatches after a few days.
• Larva eats prey in one or two weeks.
• Fully fed larva may be still for a couple of days.
• Voids waste and spins cocoon
• • In winter pre pupal stage lasts all winter.
• When no winter diapause, pre pupal stage lasts a week
  followed by pupation.
• Adult appears two weeks later.
• After emergence the adult moistens the mud partition to
  facilitate an exit.
• On the wing July - August