Photo: Ben Oldroyd
“Bees are in trouble”
“Bees are in trouble”
Why?
Colony loss
Colony loss
R. Upshaw vimeo.ocm
Winter losses
Colony loss
R. Upshaw vimeo.ocm
Winter losses
bees get too cold due to lack of food
or colony becomes too small to keep wa...
Colony loss
Colony collapse events
no or few adult bees, no dead bees
Colony loss
Colony collapse events
capped brood (pupae) and food remain
no or few adult bees, no dead bees
Colony loss
Colony collapse events
950
1905
~1909
1975
1999
But why should we care?
But why should we care?
Honey?
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
But why should we care?
Honey? POLLINATION!
Alternatives?
Alternatives?
Alternatives?
Alternatives?
Alternatives?
Alternatives?
Alternatives?
Varroa mite
Nosema
Causes of colony loss?
Varroa mite
Nosema
Causes of colony loss?
but not in Australia! (yet)
Viral diseases
Nosema
Causes of colony loss?
Varroa
Nosema
Causes of colony loss?
Causes of colony loss?
Climate change
Causes of colony loss?
Causes of colony loss?
Causes of colony loss?
Causes of colony loss?
Causes of colony loss?
Causes of colony loss?
Causes of colony loss?
What is the mechanism
of hive loss?
Hypothesis:
forager loss
drives hive loss
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
Few forag...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
Age
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
...
Honeybee demographics
eclosion
death
Age-based polyethism
Social inhibition:
Many foragers Hive bees stay in the hive
Few ...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
€
dF
dt
= R(H,F) − mF
recruitment death
E(H+F)...
Colony collapse demographic model
Hive bees:
Foragers:
€
dH
dt
= E(H + F) − HR(H,F)
eclosion
€
dF
dt
= R(H,F) − mF
recruit...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
€
dF
dt
= R(H,F) − mF
recruitment death
E(H+F)...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
€
dF
dt
= HR(H,F) − mF
recruitment death
E(H+F...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
recruitment death
E(H+F) increasing, saturatin...
€
E(H + F) = E(N) =
LN
N + w
maximum laying rate of queen
controls saturation rate
Khoury et al, 2011, PLoS ONE 6: e18491
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
recruitment death
E(H+F) increasing, saturatin...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
recruitment death
E(H+F) increasing, saturatin...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
recruitment death
E(H+F) increasing, saturatin...
Colony collapse demographic model
Hive bees:
Foragers:
eclosion recruitment
recruitment death
E(H+F) increasing, saturatin...
Colony collapse demographic model
Phase plane: m = 0.1: steady state solution
Khoury et al, 2011, PLoS ONE 6: e18491
Colony collapse demographic model
Phase plane: m = 0.3: colony death
Khoury et al, 2011, PLoS ONE 6: e18491
Colony collapse demographic model
Time course: m = 0.35: colony collapse
Colony collapse demographic model
Bifurcation diagram
death rate m
foragerpopulationatsteadystate
Khoury et al, 2011, PLoS...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
high death rate younger foragers
age at death
age at commencement of foraging
high death rate younger foragers
high death rate younger foragers
younger foragers
are less competent
higher
death rates
high death rate younger foragers
younger foragers
are less competent
higher
death rates
standard model
including young for...
But, but, but,……
But, but, but,……
What about food?
But, but, but,……
What about food?
and brood?
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
death
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate l...
eclosion
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate low soc...
eclosion
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate low soc...
eclosion
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate low soc...
eclosion
recruitment
Colony collapse demographic model
* Forager death rate drives population size
High death rate low soc...
stored
food
uncapped
brood
hive bees
foragers
Hive bees, foragers, food and brood
hive bees
foragers
collection
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers
collection
consumption
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers
Hive bees, foragers, food and brood
brood production depends on food
uncapped
brood
stored
food
hive bees
foragers
eclosion
Hive bees, foragers, food and brood
and on hive bees
brood production depends on food
uncapped...
hive bees
foragers
Hive bees, foragers, food and brood
eclosion
pupationuncapped
brood
stored
food
hive bees
foragers
Hive bees, foragers, food and brood
eclosion
delay
uncapped
brood
stored
food
hive bees
foragers
Hive bees, foragers, food and brood
social inhibition depends
on food
uncapped
brood
stored
food
hive bees
foragers
recruitment
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers
recruitment
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers
recruitment
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers
recruitment
Hive bees, foragers, food and brood
uncapped
brood
stored
food
hive bees
foragers forager death
Hive bees, foragers, food and brood
uncapped
brood
stored
food
Hive bees, foragers, food and brood
time in days
food
foragers
hive bees
brood
m = 0.1
Hive bees, foragers, food and brood
time in days
food
foragers
hive bees
brood
m = 0.3
Hive bees, foragers, food and brood
time in days
foragers
hive bees
brood
food
m = 0.42
Hive bees, foragers, food and brood
time in days
foragers
food
m = 0.5
hive bees
brood
food
brood
hive bees
foragers
where
Hive bees, foragers, food and brood
food
brood
hive bees
foragers
where
Hive bees, foragers, food and brood
food
brood
hive bees
foragers
where
Hive bees, foragers, food and brood
food
brood
hive bees
foragers
where
Hive bees, foragers, food and brood
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
increasing food
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
increasing food
limited
food
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
increasing food
limited
food
hive d...
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
increasing food
limited
food
hive d...
Steady state populations
brood
food
hive bees
foragers
analytic numerical
death rate m
m = 0.1 m = 0.3 m = 0.42 m = 0.5
Steady state populations
brood
food
hive bees
foragers
analytic numerical
m = 0.1 m = 0.3 m = 0.42 m = 0.5
analytic numeri...
Steady state populations
analytic numerical
food
foragers
brood
hive bees
food collection rate c
Steady state populations
analytic numerical
food
foragers
brood
hive bees
food collection rate c
Steady state populations
analytic numerical
food
foragers
brood
hive bees
food collection rate c
hive
dies
Steady state populations
analytic numerical
food
foragers
brood
hive bees
food collection rate c
hive
dies
limited
by food
Steady state populations
analytic numerical
food
foragers
brood
hive bees
food collection rate c
hive
dies
limited
by food...
Agreement with data
Adult bees
Khoury et al 2013 PLOS ONE
Agreement with data
Adult bees
Khoury et al 2013 PLOS ONE
input
Agreement with data
Adult bees
Khoury et al 2013 PLOS ONE
Agreement with data
Brood cells
Khoury et al 2013 PLOS ONE
Agreement with data
Food
Khoury et al 2013 PLOS ONE
Agreement with data
Khoury et al 2013 PLOS ONE
Food
Agreement with data
Khoury et al 2013 PLOS ONE
Food
small colonies
do worse than
predicted
Population changes with increased forager death ratew
food
brood
hive bees
foragers
m = 0.1 m = 0.42
strong
hive
Population changes with increased forager death ratew
food
brood
hive bees
foragers
m = 0.1 m = 0.42
perceptible change in...
Population changes with increased forager death ratew
food
brood
hive bees
foragers
m = 0.1 m = 0.42
perceptible change in...
Population changes with increased forager death ratew
m = 0.3 m = 0.42
no perceptible changes
food
hive bees
brood
forager...
So, does forager loss
drive hive loss?
So, does forager loss
drive hive loss?
Yes. It can
What else does this
model suggest?
What else does this
model suggest?
*Food supplies can remain even after hive
loss through forager death
What else does this
model suggest?
*Brood is relatively unaffected by
forager loss
*Food supplies can remain even after hi...
What else does this
model suggest?
*Brood is relatively unaffected by
forager loss
*Food supplies can remain even after hi...
How can we protect honey bees?
(and sustain our food supply)
How can we protect honey bees?
(and sustain our food supply)
How can we protect honey bees?
(and sustain our food supply)
* Pay attention to general hive health.
It is crucial!
* Pay attention to general hive health.
It is crucial!
* Take care with insecticide use
How can we protect honey bees?
(an...
* Pay attention to general hive health.
It is crucial!
* Take care with insecticide use
* Monitor adult bee populations in...
* Pay attention to general hive health.
It is crucial!
* Take care with insecticide use
* Maintain quarantine against varr...
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Lela Dowling, Bee Culture Magazine
Bees are in Trouble - Mary Myerscough
Bees are in Trouble - Mary Myerscough
Bees are in Trouble - Mary Myerscough
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Bees are in Trouble - Mary Myerscough

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Talk presented at Mathematics of Planet Earth Biosecurity meeting (A Planet at Risk) - Mary Myerscough

Abstract. Honey bees are vital to the production of many foods which need to be pollinated by insects. Yet in many parts of the world honey bee colonies are in decline. A crucial contributor to hive well-being is the health, productivity and longevity of its foragers. When forager numbers are depleted due to stressors in the colony (such as disease or malnutrition) or in the environment (such as pesticides) there is a significant effect, not only on the amount of food (nectar and pollen) that can be collected but also on the colony's capacity to raise brood (eggs, larvae and pupae) to produce new adult bees to replace lost or aged bees. We use a set of differential equation models to explore the effect on the hive of high forager death rates. In particular we examine what happens when bees become foragers at a comparatively young age and how brood rearing, food collection and colony survival is impacted by forager loss.

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Bees are in Trouble - Mary Myerscough

  1. 1. Photo: Ben Oldroyd
  2. 2. “Bees are in trouble”
  3. 3. “Bees are in trouble” Why?
  4. 4. Colony loss
  5. 5. Colony loss R. Upshaw vimeo.ocm Winter losses
  6. 6. Colony loss R. Upshaw vimeo.ocm Winter losses bees get too cold due to lack of food or colony becomes too small to keep warm
  7. 7. Colony loss Colony collapse events
  8. 8. no or few adult bees, no dead bees Colony loss Colony collapse events
  9. 9. capped brood (pupae) and food remain no or few adult bees, no dead bees Colony loss Colony collapse events
  10. 10. 950
  11. 11. 1905
  12. 12. ~1909
  13. 13. 1975
  14. 14. 1999
  15. 15. But why should we care?
  16. 16. But why should we care? Honey?
  17. 17. But why should we care? Honey? POLLINATION!
  18. 18. But why should we care? Honey? POLLINATION!
  19. 19. But why should we care? Honey? POLLINATION!
  20. 20. But why should we care? Honey? POLLINATION!
  21. 21. But why should we care? Honey? POLLINATION!
  22. 22. But why should we care? Honey? POLLINATION!
  23. 23. But why should we care? Honey? POLLINATION!
  24. 24. But why should we care? Honey? POLLINATION!
  25. 25. Alternatives?
  26. 26. Alternatives?
  27. 27. Alternatives?
  28. 28. Alternatives?
  29. 29. Alternatives?
  30. 30. Alternatives?
  31. 31. Alternatives?
  32. 32. Varroa mite Nosema Causes of colony loss?
  33. 33. Varroa mite Nosema Causes of colony loss? but not in Australia! (yet)
  34. 34. Viral diseases Nosema Causes of colony loss?
  35. 35. Varroa Nosema Causes of colony loss?
  36. 36. Causes of colony loss?
  37. 37. Climate change Causes of colony loss?
  38. 38. Causes of colony loss?
  39. 39. Causes of colony loss?
  40. 40. Causes of colony loss?
  41. 41. Causes of colony loss?
  42. 42. Causes of colony loss?
  43. 43. Causes of colony loss?
  44. 44. What is the mechanism of hive loss?
  45. 45. Hypothesis: forager loss drives hive loss
  46. 46. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment
  47. 47. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment
  48. 48. Honeybee demographics Age death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers brood
  49. 49. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers brood hive bee
  50. 50. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  51. 51. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  52. 52. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  53. 53. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  54. 54. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  55. 55. Honeybee demographics Age eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager
  56. 56. Honeybee demographics eclosion death Age-based polyethism Social inhibition: Many foragers Hive bees stay in the hive Few foragers Hive bees become foragers recruitment brood hive bee forager Bigger colonies raise more brood
  57. 57. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment € dF dt = R(H,F) − mF recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation social inhibition € dH dt = E(H + F) − HR(H,F) Khoury et al, 2011, PLoS ONE 6: e18491
  58. 58. Colony collapse demographic model Hive bees: Foragers: € dH dt = E(H + F) − HR(H,F) eclosion € dF dt = R(H,F) − mF recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation social inhibition Khoury et al, 2011, PLoS ONE 6: e18491
  59. 59. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment € dF dt = R(H,F) − mF recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation social inhibition € dH dt = E(H + F) − HR(H,F) Khoury et al, 2011, PLoS ONE 6: e18491
  60. 60. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment € dF dt = HR(H,F) − mF recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation social inhibition € dH dt = E(H + F) − HR(H,F) Khoury et al, 2011, PLoS ONE 6: e18491
  61. 61. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H social inhibition € dH dt = E(H + F) − HR(H,F) € dF dt = HR(H,F) − mF Khoury et al, 2011, PLoS ONE 6: e18491
  62. 62. € E(H + F) = E(N) = LN N + w maximum laying rate of queen controls saturation rate Khoury et al, 2011, PLoS ONE 6: e18491
  63. 63. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H € dH dt = E(H + F) − HR(H,F) € dF dt = HR(H,F) − mF Khoury et al, 2011, PLoS ONE 6: e18491
  64. 64. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation € dH dt = E(H + F) − HR(H,F) € dF dt = HR(H,F) − mF Khoury et al, 2011, PLoS ONE 6: e18491
  65. 65. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H fastest maturation social inhibition € dH dt = E(H + F) − HR(H,F) € dF dt = HR(H,F) − mF Khoury et al, 2011, PLoS ONE 6: e18491
  66. 66. Colony collapse demographic model Hive bees: Foragers: eclosion recruitment recruitment death E(H+F) increasing, saturating function; R(H,F) = α −σ F F + H proportion of foragers in colony € dH dt = E(H + F) − HR(H,F) € dF dt = HR(H,F) − mF Khoury et al, 2011, PLoS ONE 6: e18491
  67. 67. Colony collapse demographic model Phase plane: m = 0.1: steady state solution Khoury et al, 2011, PLoS ONE 6: e18491
  68. 68. Colony collapse demographic model Phase plane: m = 0.3: colony death Khoury et al, 2011, PLoS ONE 6: e18491
  69. 69. Colony collapse demographic model Time course: m = 0.35: colony collapse
  70. 70. Colony collapse demographic model Bifurcation diagram death rate m foragerpopulationatsteadystate Khoury et al, 2011, PLoS ONE 6: e18491
  71. 71. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes more adult bees raise more brood
  72. 72. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes more adult bees raise more brood
  73. 73. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes more adult bees raise more brood
  74. 74. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes more adult bees raise more brood
  75. 75. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes more adult bees raise more brood
  76. 76. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * BUT it gets worse….. more adult bees raise more brood
  77. 77. high death rate younger foragers age at death age at commencement of foraging
  78. 78. high death rate younger foragers
  79. 79. high death rate younger foragers younger foragers are less competent higher death rates
  80. 80. high death rate younger foragers younger foragers are less competent higher death rates standard model including young forager incompetence
  81. 81. But, but, but,……
  82. 82. But, but, but,…… What about food?
  83. 83. But, but, but,…… What about food? and brood?
  84. 84. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes
  85. 85. eclosion death recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes
  86. 86. eclosion recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes brood hive bees foragers eggs larvae pupae (capped brood) 3 days 6 days 12 days
  87. 87. eclosion recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes brood hive bees foragers eggs larvae pupae (capped brood) 12 days food important
  88. 88. eclosion recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes brood hive bees foragers eggs larvae pupae (capped brood) 12 days consumption and cannibalism
  89. 89. eclosion recruitment Colony collapse demographic model * Forager death rate drives population size High death rate low social inhibition loss of hive bees loss of brood * Colony collapse disorder may have many causes brood hive bees foragers eggs larvae pupae (capped brood) 12 days consumption and cannibalism cannibalism
  90. 90. stored food uncapped brood hive bees foragers Hive bees, foragers, food and brood
  91. 91. hive bees foragers collection Hive bees, foragers, food and brood uncapped brood stored food
  92. 92. hive bees foragers collection consumption Hive bees, foragers, food and brood uncapped brood stored food
  93. 93. hive bees foragers Hive bees, foragers, food and brood brood production depends on food uncapped brood stored food
  94. 94. hive bees foragers eclosion Hive bees, foragers, food and brood and on hive bees brood production depends on food uncapped brood stored food
  95. 95. hive bees foragers Hive bees, foragers, food and brood eclosion pupationuncapped brood stored food
  96. 96. hive bees foragers Hive bees, foragers, food and brood eclosion delay uncapped brood stored food
  97. 97. hive bees foragers Hive bees, foragers, food and brood social inhibition depends on food uncapped brood stored food
  98. 98. hive bees foragers recruitment Hive bees, foragers, food and brood uncapped brood stored food
  99. 99. hive bees foragers recruitment Hive bees, foragers, food and brood uncapped brood stored food
  100. 100. hive bees foragers recruitment Hive bees, foragers, food and brood uncapped brood stored food
  101. 101. hive bees foragers recruitment Hive bees, foragers, food and brood uncapped brood stored food
  102. 102. hive bees foragers forager death Hive bees, foragers, food and brood uncapped brood stored food
  103. 103. Hive bees, foragers, food and brood time in days food foragers hive bees brood m = 0.1
  104. 104. Hive bees, foragers, food and brood time in days food foragers hive bees brood m = 0.3
  105. 105. Hive bees, foragers, food and brood time in days foragers hive bees brood food m = 0.42
  106. 106. Hive bees, foragers, food and brood time in days foragers food m = 0.5 hive bees brood
  107. 107. food brood hive bees foragers where Hive bees, foragers, food and brood
  108. 108. food brood hive bees foragers where Hive bees, foragers, food and brood
  109. 109. food brood hive bees foragers where Hive bees, foragers, food and brood
  110. 110. food brood hive bees foragers where Hive bees, foragers, food and brood
  111. 111. Steady state populations brood food hive bees foragers analytic numerical death rate m
  112. 112. Steady state populations brood food hive bees foragers analytic numerical death rate m
  113. 113. Steady state populations brood food hive bees foragers analytic numerical death rate m increasing food
  114. 114. Steady state populations brood food hive bees foragers analytic numerical death rate m increasing food limited food
  115. 115. Steady state populations brood food hive bees foragers analytic numerical death rate m increasing food limited food hive dies
  116. 116. Steady state populations brood food hive bees foragers analytic numerical death rate m increasing food limited food hive dies food remains
  117. 117. Steady state populations brood food hive bees foragers analytic numerical death rate m m = 0.1 m = 0.3 m = 0.42 m = 0.5
  118. 118. Steady state populations brood food hive bees foragers analytic numerical m = 0.1 m = 0.3 m = 0.42 m = 0.5 analytic numerical
  119. 119. Steady state populations analytic numerical food foragers brood hive bees food collection rate c
  120. 120. Steady state populations analytic numerical food foragers brood hive bees food collection rate c
  121. 121. Steady state populations analytic numerical food foragers brood hive bees food collection rate c hive dies
  122. 122. Steady state populations analytic numerical food foragers brood hive bees food collection rate c hive dies limited by food
  123. 123. Steady state populations analytic numerical food foragers brood hive bees food collection rate c hive dies limited by food limited by brood survival
  124. 124. Agreement with data Adult bees Khoury et al 2013 PLOS ONE
  125. 125. Agreement with data Adult bees Khoury et al 2013 PLOS ONE input
  126. 126. Agreement with data Adult bees Khoury et al 2013 PLOS ONE
  127. 127. Agreement with data Brood cells Khoury et al 2013 PLOS ONE
  128. 128. Agreement with data Food Khoury et al 2013 PLOS ONE
  129. 129. Agreement with data Khoury et al 2013 PLOS ONE Food
  130. 130. Agreement with data Khoury et al 2013 PLOS ONE Food small colonies do worse than predicted
  131. 131. Population changes with increased forager death ratew food brood hive bees foragers m = 0.1 m = 0.42 strong hive
  132. 132. Population changes with increased forager death ratew food brood hive bees foragers m = 0.1 m = 0.42 perceptible change in food storage pattern and adult bee numbers strong hive
  133. 133. Population changes with increased forager death ratew food brood hive bees foragers m = 0.1 m = 0.42 perceptible change in food storage pattern and adult bee numbers no perceptible change in brood numbers strong hive
  134. 134. Population changes with increased forager death ratew m = 0.3 m = 0.42 no perceptible changes food hive bees brood foragers weak but viable hive but hive is doomed
  135. 135. So, does forager loss drive hive loss?
  136. 136. So, does forager loss drive hive loss? Yes. It can
  137. 137. What else does this model suggest?
  138. 138. What else does this model suggest? *Food supplies can remain even after hive loss through forager death
  139. 139. What else does this model suggest? *Brood is relatively unaffected by forager loss *Food supplies can remain even after hive loss through forager death
  140. 140. What else does this model suggest? *Brood is relatively unaffected by forager loss *Food supplies can remain even after hive loss through forager death *Number of adult bees may be the best indicator of colony well-being
  141. 141. How can we protect honey bees? (and sustain our food supply)
  142. 142. How can we protect honey bees? (and sustain our food supply)
  143. 143. How can we protect honey bees? (and sustain our food supply) * Pay attention to general hive health. It is crucial!
  144. 144. * Pay attention to general hive health. It is crucial! * Take care with insecticide use How can we protect honey bees? (and sustain our food supply)
  145. 145. * Pay attention to general hive health. It is crucial! * Take care with insecticide use * Monitor adult bee populations in hives How can we protect honey bees? (and sustain our food supply)
  146. 146. * Pay attention to general hive health. It is crucial! * Take care with insecticide use * Maintain quarantine against varroa * Monitor adult bee populations in hives How can we protect honey bees? (and sustain our food supply)
  147. 147. Lela Dowling, Bee Culture Magazine
  148. 148. Lela Dowling, Bee Culture Magazine
  149. 149. Lela Dowling, Bee Culture Magazine
  150. 150. Lela Dowling, Bee Culture Magazine
  151. 151. Lela Dowling, Bee Culture Magazine
  152. 152. Lela Dowling, Bee Culture Magazine
  153. 153. Lela Dowling, Bee Culture Magazine
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