private study only. The thesis may not be reproduced elsewhere without the permission of the Author.
~-lesting Biology of } ~bu~ r1.1c!er;1r.us P:i.b.
t hr r cquire~cn~~ for t~e dc~re~ of
· ·,ster of ·:ci erice in ~cu1 o~v ::1.t
i f-·u
• V •
Nel son Pc··1eroy
1977
·~o the bumbl ebees,
wt ose cooperation ~ade the ~roject poscihle.
ABSTRACT
1~st nat ural nest s of 3ombus ruderatus were found in underground locations. Under~round domici Jes were hi hly attractive t o queens of n.. £_~~~£_at~~ and
n.
t e:::_restr is, theset wo s:recies occuT).Vinr; 9y; of 45 domi.ciles. A design :.1od- i f ied to simplify in-pect i.on and colony removal vielded 62~ occupation by R. ruderat us only in 13 domici les. These percentages compare favourably with overseas results (various
ann. s u _ r-,3 SS t 1ose obtained previously
i n i .. Jew Zealanci.
Colonies were t ransferred from do~iciles to observation hives for study of their deveJopment . The observation
~ives were ~e~isne~ to provide the colonies ' needs for warmth , ,en t i l.'l t ion, sc: n i t at ion, and ~rooc-c omb sur"):)ort . The t ot al number of he8~ r roduccrl ner col o~y in both natural sit es anci observ·1~~ion. hi..1:8s .::rnrrer. fror:' ?3 t o 750 (mean =
420).
T·)_g production ·in
:2 ·
ru~ern.!~~ post - incipient colonies is r evulated by the quantitv of fresh cocoons , one eg~ clump being oade per five (ap_ r ox. ) substrate cocoons . Egg clumps consist of 1 - 5 (mean= 1 .5) eg~ cells , each cell containing 10 - 19 (mean = 14.3) eggs. Egg mortalit y, especially in multi-celled clump~ appeared to be high but could not be quantified.Larval rejection caused the loss of 13% to 36% of the larval populations of five colonies in observation hives.
The role of larval r~jection ap eared to be the control of adult size. It was very prevalent during colony decline when food short ages coincided with queen production.
3. ruderatus differs from most other "pocket makers "
in having a clear size distinction between workers and queens . Larvae which becGme queens consumed at l east twice as much pollen as most worker larvae. In most
colonies there was an interim period of male-only rear ing between worker and queen production. Colonies varied in their time of initiating male reduction, earlier male production being associated with less worker production and lower overall colony productivity. It is sug9:;ested that future research s~ould atte~ t to elucidat e the causes of male production :i.n }omb~ colonie"' .
V
ACKNOWLEDGEMENTS
I wi sh to thank my supervisors , Nr . L. Gurr , and Dr . R.II. Nilnes , for their help and encourageoent.
I have lad constructive discussions with the staff
~embers and _ellow students of the Department of Botany and ZoolOGY at ~assey University, and also with Drs. ~.P.
Viecfarlane and I3. J. Donovan of the DSIH at Li ncoln. I have benefited froJ stimulating correspondence with Dr. R.C.
Plowr i~ht of the Jniversity of ~oronto.
l':r . J.::. rr.:..1rtlcy, f'·Jano.c;er of the 1.-Jri;:;htson-Nh.1\ .Seed Centre at Christchurch, ~2de fu_ds available to ~ssist with travel and othe~ expenses.
f1.r . r:. F,"~rne·~'ill,: cnvc c1.:=:;s-' st;:;nce 1t1ith r:mny tcc:.1nical proble;.JS. [·, __ . \·.'.? . '!1t10r.1,.:, did the r.:otrilwork for tl:e obser- vo.tion hive~.;. ~.1~. i,. 1 .
:r .
:·ichnd.dt desicr:ed o.nd 1 ui lt t heelcc ,ronic circu~t:···. 1.::-;,..istci·::ce hith pi1otor.:;raphic equip:.1ent
w.:;s p ovide::i by;::-·. -L ':i-:itt . l.r. G.ll. \·!al ton cave ne pollen, tro.pped :'~~on l:is b8ehives.
rnhe field ':10::--k \·Ja,, done o:; t i1e property owned by t he late Mr . .
I. e .
[3arro,.v, and I t hank his family. hy father r.1anae;ed t his farm, and Inn srateful t o him for civing me a free run of the faro , its buildings and machinery. My mother happily accepted whole rooms of the house being taken over forbumblebees.
Eric, one of my brothers, generously loaned a motor
vehicle. Mr . R.G. Powlesland also helped with transport, and :i,.n other ways, including t he fin~ing of a Bombus ruderatus nest .
The monochrome photocraphs were processed by the Massey University Central Phot ographic Unit.
vii
Mrs. J.R. Parry typed the manuscr ipt.
Translations were done by Miss I.M. Verry, Mrs. L.I. Thompson , Dr. E.O. Campbell and Dr. A.J .A. Vieregg; and also by Ms. M. Kalin and ~;r . B. Pendrel, of the Universi ty of Toronto.
Rachel, my wife , helped in many ways, sharin~ both the dif f icult and t he happy ti~es.
CONTENTS
ABSTRACT
AC:CNO\·/LEDGEf'.,.3NT,.)
LIST OF TABLE:J LI3T OF FIGURES LI3T OF PLAT~~3
CEAP'T~~rt 1 II'T'J':JODUC t IOH
2.1.1
2 .1. 2 2.1.2.1 2.1.2.2 2.1.2.3 2.2 -2.2.1
2.2.2
2.2.3
2.2.LJ..
2.2.5
2 . 3
2.3.1 2.3 .2 2
-3-3
2.3.4
':.' ild Colonic s
Colo .ies from ?ield DoLliciles T~c 1974-75 season
mLoe 1 1075~ -~/ () ·~ sen~on
Trnnsfer of colonies to the laboratory
T c:r,porD ture Ventilo.tion 3ho.pe
Drainar~e
Tne Exit Hole DATA COLLECTION Brood Records Larval Rejection Foraging Data
Ad~lt male and queen food consumption
iv vi xi
xii
1
6
6 6 8 '10
12 16
17 17 23
25 25
26
32 32 34 35 37
2.3.5
2.4 2.,-1-.1 2.4-.2 2.4._,2_
CHAP'J:E:.1
3 3 . 1
3.1.1 -:;; 1 c..' )
,(
. .
3.2
3.2. 2.1 3.2. 2.2
3.2. 2.4
3
• 2 • < 7,3.2.4
3 . 3
3. 3. 1
3 . 3 . 1 . 1 3 . 3 . 1 . 2
3. 3. 2
3.3.2.1
3.3.2.2
3. 3 . 3
Destructive samplin3 of whole colonies
DA'11A ANALYSI S
5rood Development Larval Rejection Foras) ,·int n· HccordJ - s
RESUL'1'3 AND DISCUSSION
Natural ~est 8ites Do~icilc Occun~tions
Pi.1yr-;ic;,l onro::-;::.0~tion of t he clur:ip
~ode of nut~ition Lo.rval crowth Pupo.e
/,dul t G
i3HOOD ViO~tTALITY
En-,,. L H l f-:• ort:=.>... l i· t y s Hatchinc failure Oophagy
Larval Mortality Larva- eating Larval rejection
Post- spinning mortality
ix
Page 38
39 39 40 42
43 43
L!-3
46
so
50
53
5861
68 72 76
78 78
78 78
81 81 81
85
3.
4 .13.4.2 3.4.3
3.L~.4
COLONY DEV:SLOPNENT
Er;r, Product ion Brood Gr.o\·Jth Larval J~e,"jection
CHAPTF:R 4 GEI\TERAL DISCUSSION AND CONCLUSIONS 4. '1
4.2
ECOLOGY AND MANAGEMENT OF COI,ONIES COLONY DEVELOPMENT
J\PP2JDIX 1 Nest r:oterial Preferences
!.PE~::,DIX 3 Ventilo.tion
APPENDIX 4 Formula of Essi~s Aphid Fluid
APPENDIX 5 Results from the 1976 - 77 season
REFERENCES
Par;e
89 89
103 109
112
114
114 117
133
135 138
139
145
3.2
3.3
3. 4.
LI0T OF TABLES
Natural Nest Sites of Bo~bus ruderatus
Domicile Occupation L:esul t s
Relctive Dryness of Brick and Concrete Domicile:-:. !~a:rly in tile 1975-76 Season
nrood Eortal ity Occurrinc between Late
l ifth r~~ta~ (Sil k 3pinninc) and Adult St a~es
Hurbc::::- of Cocoons ~;'ound in Compl et ed B. ~uderatus olonies
xi
44
47
47
87
90
2.1 2.2
2.4
2 . 5 3 .1
3 . 2
3 . 7 3 . 8 3. 9
LIST OF FIGURES
Under5round domicile Brick surface do~icile
Licht wei'~h t-conc:::-et c surface domicile Nest-cha~ber of observation hive
Vestibules of observation hive
Percentn~cs of brood clumps formed on cocoons of v rious aces
Nurn e:r of conti:_;1.1011:; ec;c; cells per brood clur:1:p
!\.t_;e ol' broo(i c lu:nrs 1.-:hcn pollen-pocket f or~ed
J:reque~cy dist~ihution of larval head cop~ulc din~aters
Diagrams of developing brood clump
Relation between larval clump area and the biomass of larvae inside
Larval e;rowth rates
Frequency distribution of cocoon diameters
(A) Relation between cocoon "volume" and quantity of pollen consumed during larval sta~e. (B) Relation between cocoon diameters and dimensions of adult females emerging from them.
1,3
14- 22
27 52
55
56 57
59
69
71
73
74
3.10
3.11
3.12
Relation between cocoon diameter and pupal durntion
Food consumpt ion by newly emerged queens and males
Daily pollen int ake to colony versus t he quantity of l arvae present. The number of l arvae rejected that day and followinB niGht also shown
Relation ~ctween heed capsul e diameter and
dry weir)1t ror rejected and non-rejected lo.rV08
xiii Page
75
77
84
86
3.14.1 Spatio-tc~poral relntions of the brood clumps 91
or
colon;y 13.14.2 dpntio-tenporal relations of t he brood clumps 92 of colony 2
3.11.J-.3 Spatio-temporal
lD colon;y 3
3.14.4 Spatio-temporal in colony 4
3.14.5 Spatio-temporal in colony 5
3.15.1 Seasonal chane;es 3.15.2 Seasonal changes
3.15.3 Seasonal changes
relations of
relations of
relations of
in colony 1 in colony 2
in colony 3
the brood
t he brood
the brood
clumps
clumps
clumps
93
94
'95
97 98 .99
1
3 .1 5 .4
Seasonal chances in colony4 3 .1 5 . 5
Seasonal eh n~es in colony5
4.1 Flow diagram of the main factors affecting the development of B. ruderatus colonies.
100 101 118
LIST OF PLATES
Fronti spiece Thirty day old ~ombus ruderatus colony
1
2
3
4-
5
7
9 10 11
12
13
14-
15
The laboratory, view 1 The laboratory , view 2
Apparatus for automat ic photography of f oracin~ bumbl e~ee0
Fi fty dny old Dombus ruderatus col ony in observation hive
Poundress queen lo.,yinc; ecr-;s .Zc;c cell bcinc closed
A rupidly-crowinc clu~p of queen larvae EcG clu~p uroken open
Worker :eedinc lar va
Worker carryinc r ejected larva
Colony fi ve about twent y days since its inception
Colony f ive, about fort y days old Colony five , about sixty days old Colony five , about eighty days old
Incipient brood c·lump
xv
ii
30
30
36
51
54 54
62 62
65 65 106
106 107
~07 144
