SU M MARY. Fofty colonies of Apis mellifera macedonica ... (Nolan, 1925; Braun, 1942; Woyke, '1984), but only a few included queens 3- or 4-years old in their ...
01995IBRA
Journal of Apicultural Research 34(1):9-14 (1995)
Brood and honey production of honey bee colonies requeened at vanous frequencies MaRtn Kosranetou-Dervttnrutoou' ; t; Aruonels Txnaswoutou'' Dtnittrntos Tsettos' ; Korustaurtnos BLnoENopoutoso
'National Agricultural Research Foundation, Agricultural Research Station of Halkidiki, N Moudania 632 00, Greece 'Laboratory of Apiculture-Sericu lture, School of Agriculture, Aristotle University of Thessaloniki, Greece oNational
Agricultural Research Foundation, Cereal Institute, 570 01 Thermi, Thessaloniki, Greece
(Received 28 SePtember 1993, accepted subject to revision 30 December 1993, accepted for publication 9 November 1994)
"Author for correspondence
SU M MARY Fofty colonies of Apis mellifera macedonica were established in Greece with naturally-mated sister queens, two frames of sealed brood and 2 kg of bees, and requeened either every year (A), every two years (B), every three years (C), or left to replace their queens through supersedure (D). All colonies were managed in the same way during the seven-year experiment. Brood area was significantly higher in groups A and B than in C or D, except for the first two years, During the first three years there were no significant differences in honey production between any of the
groups, but group D subsequently produced significantly less honey than all other groups. There were generally no significant differences in
each year's honey production between colonies in groups A, B or C, suggesting that requeening every two or three years is adequate.
Keywords: queen honey bees, Apis mellifera macedonica, requeening, age, brood rearing, honey production, Greece
Kostarelou-Damianidou; Thrasyvoulou; Tselios; Bladenopoulos
10
INTRODUCTION It is
commonly accepted that the age of a queen honey
bee determines her performance, and bee colonies that are not requeened annually seldom produce good honey crops. Howeve[ few objective data exist to sup-
full-depth hive bodies, and four frames for those with one. Protein supplements were not provided since bees collected and stored plenty of pollen during spnng. All colonies were treated alike. They were all opened
port this contention. Most studies of the effects of
on the same day and were examined every three
older oueens have been based on comoarisons
weeks for queen replacement, brood diseases, avarr-
between colonies headed by queens 1- to 2-years old (Nolan, 1925; Braun, 1942; Woyke, '1984), but only a few included queens 3- or 4-years old in their experiments (Szabo & Lefkovitch, 1989; Diehnelt, 1986; Cavojsky, 1 982). There have been no studies comparing supersedure and younger queens. More surprisingly, the data do not fully support the conventional
wisdom on the poor performance of colonies with aged queens. Szabo and Lefkovitch (1989) did not show differences in honey or brood production among colonies headed by 1-,2- or3-year-old queens. Similarly, Cavojsky (1982) found no differences in 4-year-
old and younger queens. The alternative view, that sometimes young queens are poor while some older
queens lay well for several seasons in succession (Cale et al., 1978), is not supporled by enough research data.
We conducted a study on the performance of 40 colonies that were requeened at various frequencies for seven consecutive years, and recorded their brood and honey production in order to determine the effects of queen replacement frequency on brood and honey
ability of space for brood rearing and food storage. Combs with foundation were given to colonies which needed more space. During April and May all colonies were checked for swarm cells every week. Swarming was prevented by the removal of any occupied queen cell during the swarming season, but despite these efforts two colonies swarmed. The swarms were captured and rejoined with their parental colonies. Strengthening and removal of brood and population were not undedaken. All colonies, healthy or diseased, were treated at the
same time. Varroa was controlled with malathion (1985-1989) and fluvalinate (1989-1991) during autumn of each year. Chalkbrood was prevented by the destruction of affected combs, by good ventilation of hives, and by often removing the mummies from the floorboard. No chemotherapeutic agent was used against nosema disease, tracheal mite disease, American or European foulbrood, since these diseases were never found in the exoerimental colonies.
oroduction.
The area of brood was measured monthly from May to November every year by means of a clear acrylic plastic (Plexiglas) grid(4x4 cm), estimating that 4.28 cells
MATERIALS AND METHODS
occupy 1 cm'of comb surface. The amount of honey extracted from each colony was calculated annually from the difference in comb weioht before and after
This investigation was conducted at the Agriculture Research Station of Halkidiki. Greece, between June 1985 and May 1992. Forly colonies of Aprs mellifera macedonica were used. They all received newly emerged and naturally-mated sister queens. Each colony began with two frames of sealed brood and 2 kg of bees (about 20 000 bees per colony). Colonies were grouped in four treatments of 10 each, designated as A, B, C and D. Colonies of group A were requeened every year (1 985-1 991 ), group B every second year (1 985, 1 987, 1 989, 1 991 ), group C every third
year (1985, 1988, 1991), and group D were left to
requeen themselves (supersedure). All requeening was done in May. Numbered plastic discs were glued on to the thoraces of the queens to ensure proper identification and to determine if the queen was replaced by the bees of the colony.
Experimental colonies were located in the same apiary but were simultaneously moved to three or four areas in each year for food. In seasons when the quantity of honey was insufficient for wintering, colonies were fed with heavy syrup made of two pads sugar to one of water (2 : 1). Each was given enough syrup so that they stored adequate amounts of carbohydrate for winter:
eight frames of sealed food for colonies with two
extraction.
Statistical analysis was performed using separate ANOVA analyses for each year. Differences among treatments were assessed by Duncan's Multiple Range Test. Correlation coefficients between brood production and honey yield were calculated for each group ano year.
RESULTS Brood production in consecutive years Our experimental period began in June and ended in May of the following year, with the exception of the seventh year which lasted from June to November. Five colonies of group D became weak and died in 1 988-1 989, two in 1 989-1 990 and one in 1 990-1 991 . The other groups did not lose any colony throughout the experiment. Averages and statistical analysis were calculated using the surviving colonies for group D.
Colonies had the highest amount of brood in May (table 1). The process of requeening that was carried out after the May measurement affected brood production in June. Colonies regained the losses in July, and extended their brood rearing until August when
Kostarelou-Damianidou; Thrasyvoulou; Tselios; Bladenopoulos
12
the brood area began to decrease. The brood area was least in November and February. During October of 1986, 1987, and 1989 brood area increased in all groups, due to the flowering of the shrubs Erlca yertrcillata and E. carnea. During the first two years, no significant differences were obseryed among groups in total brood production (table 1). The differences began from the third year
when groups A and B produced significantly more brood than groups C and D. From then, the superiority
of younger queens was apparent in their greater amount of brood production. No significant differences were found among colonies with queens 1- or 2-years old in any one year in either environmentally favourable (1989-1990) adverse (1990-1991) years.
or
Colonies with 3-year-old queens or supersedure
Honey production in consecutive years During the first three years of the experiment no signif-
icant differences in total honey production were observed between the four groups (table 2). Group D produced about half of the honey produced by the other groups in 1 988, and every year after 1 989 group D continued to produce significantly less honey than the other groups. Dead colonies of D group were eliminated from the study.
During 1989 colonies of group C with 2-year-old queens gave the highest amount of honey, approximately 24 times more than that produced by colonies of group D in 1991 . With the exception of 1989, no sig-
nificant differences were found among colonies headed by 1 -, 2- or 3-year-old queens.
queens produced significantly less brood than groups A or B in all comparable years (table
1):35.5% less during autumn (September-November), 35.8% less during February, and 46.2% less during spring (March-May).
Thus, colonies with younger queens were more populous and had more bees for the winter cluster. This larger working force resulted in an earlier production of brood in February and a greater increase in spring.
Goefficient of variability and correlations Colonies were more or less homogeneous during the first three years but after that the coefficient of variation (CV) of all groups increased significantly (table 3). The unfavourable conditions of the fourth year seemed
to affect the brood rearing among groups differently,
TABLE 2. Honey production (kg) of colonies requeened at different frequencies (i t s.e.), Colonies in each group were requeened as follows: A, annually; B, every 2 years; C, every 3 years; D, left to natural supersedure.
Year
Group A
Group B
985 1 986 1987 1 988 1 989 1 990
27.4 + 1.6 a
27.2 r1.5 a 33.1 + 1.4 a
991
57.9+4.6a
31.4 t 0.9 a 20.2 + 1.3 a 51.8 t 7.0 a 21.6+1.7 a 57.0 + 4.4 a
Total
236.6
242.5
250.6
120.1
Means'
33.8 a
34.6a
35.8 a
17.1 b
1
1
28.6 +2.3 a 32.6 x 1.3 a 21.O +2.6 a 49.5t5.7 a 19.4 x 1.4 a
' Means with different letters are significantly different from each other
Group C 27.7 +1.3a 31.8 + 1.1 a 23.8 t 1.3 a 21.8 + 1.6 a 73.2 x.10b 17.4 t2.2 a
54.2x.5.1a
Group D
t
28.5 1.1 a 32.6 + 1.2 a 23.5 r 1.4 a
11.5+1.3b 14.3 6.7
t 10.1 c t 0.3 b
elrATh
(P < 0.05)according to Duncan's Multiple Range Test
Comparisons were made within years
TABLE 3. Coefficient of variation (CV) of brood production of colonies requeened at different
frequencies. Colonies in each group were requeened as follows: A, annuallyl B, every 2 years; C, every 3 years; D, left to natural supersedure Group A
Years 985-1 986-1 1 987-1 1 988-1 1 989-1 1 990-1 1 1
986 987 988 989 990 991
19.5 17.9 15.5
Group B 20.1 17.1
18.0
27.1
17.2 19.6
18.7 19.6
Group G
/.c
Group D
to.o
16.6 17.9
20.6
21.7
|
zc.l
48.1
27.5 36.5
53.9 34.7
Requeening frequency and brood and honey production
11
TABLE 1. Brood area in cm' (x * s.e.) of colonies reqeened at different frequencies (1 cm'= 4.28 cells). Colonies in each group were requeened as follows: A, annually; B, every 2 years; C, every 3 years; D, left to natural supersedure, The ages of queens in each group are shown.
Month
198S1986 Group A 1-yr-old
Jun Jul Aug Sep Oct Nov Feb Mar Apr May
752t1og 645t195 1 517 r2og 1 072!168
1
1986-1987
B Group C 1-yr-old 1-yr-old
Group
771
Month
!114
1473+171
078r188
1 365 x.172
1 081
1
1
Group C
Group D
2-yr-old
2-yr-old
2-yr-old
513r92
754 + 1O4 1 570
165
Group B
-yr-old
1
x122
22OO x.221
1465+186
1 573
r131
925
t
190
1 757
t25O
2057 1249
1349t113
r137
1
446t156
1175 1 363a132 1 757
595 a 90
603
1
572+154
1 928
*182
1 275 +
t74
577
118
t65
t
t69
427 +86
5Og + 72
487
t36
1 187
r
115
142+26
149 + 21
141
r22
259
t
43
172 +26
160
t22
165
t25
'128
13
123 + 21
125
r25
533
523 x 46
478
t24 t64
126
692+91
668
t
560 + 49
497 +76
492
r.86
1 409+168 2597t'191
198&1987 Group A -yr-old
Jun 602t82 Jul 2245t216 Aug 1 568t 119 Sep 977 t 138 Oct 1 292+105 Nov 282+32 Feb 154t30 Mar 731 t60 Apr 1 373t 139 May 24721197 Means'
-yr-old
Group A
146+23 168+30
441
1
763
1
570+142 1 5O4 r'177
1 653
1
1 171
t
65
107t180
2 586
Means' 1044a
Month
r118
r
Group D
198$1990 1-yr-old
Jun 1 093t105 Jul 2381 *24 Aug 2264t80 Sep 629 t 109 Oct 27841287 Nov 430 t 85 Feb 509 + 70 Mar 2459t296 Apr 3346t343 May 31 7 +251 Means'1909a
t
145
1115t134
50
i
'1 438 137 2 493 +220
t133
1 195
187 x.34
1 081
t
t117
2084 x.258
174
1
t127 r24
059t94
2 198
!
184
1043a
976 a
1138a
Group D
Group A
Group B
S-yr-old
1-yr-old
2-yr-old
998 a
994 a
1 225
178
166t30
1
054r86
2 102 + 176 1052 a
1988-1988
B Group C 1-yr-old 3-yr-old
Group
1
680i135 t311
846
t
134
1 '111 + 333
963 r.247
2 029
1 965
t
248
1 454
x.135
1
1 352 +327
1 245
!217
858 1322
97
410t161 686+149
573t159
592
t
352 + 167
141
328 + 55
251 + 90
277
!30
+24
83t54
66t17
102r30 514 r147
608t156
x173
1
518t67
1 282
t145
1 270
i59
878 x.1O4
BB0
r
1 296 *.92
997r149
981
t
154
t25
797
r35
275
150t29 517t108
187 75
342
1 157
+43
t
r
131
80
152t56
50+8
18
r50
430 + 76
685
t
206
r
1 277
r
183
843 x.121
1 036 x.232
1 590
t227
1 021
2 696
t
504
2214 r2OG
2067t470
3 624
!604
3115t498
952 b
Group D
917 x.70
t
1 5O7
1
superseded
1
1 198
132
C 1-yr-old
Group
496r155
621 +71
2067 t2O9
a
Group A
1 154
!147 2289 t248 2504*196
1 297
184
1072 ab
952 b
1142 a
Group D
Group A
superseded
1-yr-old
998t1105 1370+1539 1 114+1 117
1
86
152
2992
t466
1001 b
t
598
64
40t14
tzt !zo
1342+442
t
314r189 352 * 185 896
t792
555 a
1990-1991
B Group C 1-yr-old 2-yr-old
Group
971
r
90
2322 +241 2 109 +267 710
2802
r
128
t315
1 829
t
339
2 672 x.306
2374 581
r398
t
138
2 416 r.343
515+99
419t116
526t82
522
t
129
2334t313
2294
!
421
3523 t47O 3 200
t
1901 a
300
1
t
'106
715 L72
t51
420 x.298
474
388t1 676
270 x41
187 1
008+85
1 482
t192
106+40
112 x 42
21O
109r912
528
371+1 156
2 941 a 561
1
2643 t481
225611
1869a
1032 b
147
r 40 r70
1 413 a 105
2925 1262 912 a
B Group D Group C 2-yr-old 3-yr-old superseded 1 382t108 1157t301 '1 1041310 1 723 x171 9791305 1 210+589 718 t102 436 r 133 635 t 773 507 r 56 299t102 320t235 291 r34 2OBt43 192+280 130t30 166166 160t0 144 t125 203 t 51 185 t 64 536+106 423176 555 t 191 1 434r109 987 1129 1 077 x.437 2758 t2O5 1 683 t 154 1 461 t945 Group
968 a
652 b
685 c
'Means with different letters are significantly different from each other (P < 0.05) according to Duncan's Multiple Range Test Comparisons were made separately each year
Requeening frequency and brood and honey production
IJ
TABLE 4. Correlation coefficients ftl between brood area and honey production for each group and year.
Group A
Years 1
0.44
1
985 986 1 987 1 988 1 989 1 990
0.35
1
Group G
-0.46
Group D
-U.UC
0.32
0.73*
o.32
u.4b
0.58
0.'13
0.71.
0.46 0.04
o.37
0.02
0.77*
0.84*
0.76* 0.89*
0.20
0.30
0.45
0.75*
o.21
-o.20
-u.52
991
Group B
. Significant at P 0.05
..SignificantatP0
01
contributing to a higher CV. This hypothesis is supporled by the data of 1989-1990, which shows the best
1988-1989 produced less brood than colonies that were requeened every year (group A) or every two
honey production during the experimental period. In
years (group B), but the differences were not significant. lt seems that the young queens in group C were
that year the CV of groups A and B dropped, while that of C and D remained high. The high CV of group D can be explained by the variability that existed among the fecundity of mother queens and those of supersedure daughters. The CVfor months, across years and groups, ranged from 6.1 % to 80.1 %o.For all groups, October, November and February were the months with ihe highest CV (52%,31%o and29%o respectively) and June, July and May the months with the lowest (20%, 21%o and 19o/o respectively). lt seemed that some queens continued laying eggs late into the autumn while others staded their brood production earlier in the following spring.
Factors other than the age of the queen, the food resources or the environmental conditions could contribute to that variation. Table 4 indicates that the correlation coefficients (r) are
highly variable, often being significant, weak or negative. The rvalues were not consistent in either good or bad years or in colonies with queens of either one, two or more years of age.
DISCUSSION Previous studies comparing colonies headed by '1 year-old and 2-year-old queens provided contradictory results. Nolan (1925), Braun (1942), and Woyke (1984) found that 1-year-old queens produced more brood and higher amounts of honey than colonies with 2-year-old queens. Szabo & Lefkovitch (1989) found no significant differences either in brood or honey production between such colonies. Brunnich (1922) and Diehnelt (1986) found higher rates of brood rearing in colonies headed by 2-year-old queens than in those with 1-year-old queens. We found no significant differences among 1-year- and 2-year-old queens in any combination of comparisons in 1986-1987, 1988-1989, 1989-1990 and 1990-1991 (table 1). The queens that replaced the 3-year-old ones of group C in
affected by the smaller colony population in their inherited colonies and consequently produced less brood, but subsequently managed to compensate and overcome ihe loss of the previous year. The decline of brood production in colonies headed by older queens was apparent only during their third year, when they produced significantly less brood than 1-
year-old (1987-1988, 1990-1991),
or
2-year-old
990-1 991) queens. Our results with older queens agree with those of Diehnelt (1986), but are contradictory to those of Szabo & Lefkovitch (1989) who found no significant differences among colonies headed by 1-, 2- or 3-year-old queens. The differences are (1
restricted to brood production, since we found that colonies with 3-year-old queens produced similar amounts of honey to colonies with 1- or 2-year-old queens (table 3). This result is not unexpected, since the correlation of honey production and brood area is variable and not always positive (table 4).
ln all comparisons and combinations, colonies with supersedure queens gave significantly less honey and brood than all the other groups. During the last years of the experiment (1 988-1 989, 1 989-1 990, 1 990-1 991 ) they produced 34.4%,20.9Vo and21 .4Vo of thetotal brood and 24.5o/o,36.2%o and 5% of the honey production of the other groups, respectively. Fudhermore, eight out of 10 colonies with supersedure queens were lost over the seven years of experiment.
The positive correlations indicate that honey yield increases with increasing amounts of brood. The correlation is variable since honey production depends upon factors such as the length of adult life, the individual productivity (Woyke, 1984), the time of the year (Szabo & Lefkovitch, 1989), and environmental factors (Bar-Cohen et a1.,1978; Louveaux, 1967; Genc & Aksoy, 1993) in addition to brood area. Negative correlations are attributed to the interaction between brood and environmental factors on honey production.
Kostarelou-Damianidou; Thrasyvoulou; Tselios; Bladenopoulos
14
Our correlations were estimated by the relationships
between the annual amount of brood and weight of honey that each group produced during the seven years of the experiment. The known differences in r values between the favourable and the adverse years (Bar-Cohen et a|.,1978; Genc & Aksoy, 1993) contributed further to variation of our coefficients. The
range of r values reported by several other authors (Woyke, 1984; Moeller, 1958; Soller & Bar-Cohen, 1967) concurs with our research.
The overall results of this research indicate that colonies with queens one or two years of age do not differ significantly in either brood or honey production. Colonies with queens three years of age produce less brood than colonies with queens aged one or two years, but not significantly less honey. Colonies that were left to supersede their queens produced significantly less brood, less honey and had difficulty in surviving adverse conditions. Requeening can be effective every two or three years. More research is needed with different races of bees, under different climatic conditions with different food resources to evaluate further this impor.tant aspect of bee husbandry.
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(1
942) One-year and two-year-old queens. A merican Bee
Journal 82i356-357 (1 922) The influence of the age of the queen on the honey crop. 8ee World 4(1):6-7.
BRUNNICH, K
CALE, G H; BANKER, R; POWERS, J (1978) Management for honey production. /n Dadant & Sons (ed) The hive and the honey bee Dadant & Sons; Hamilton, lllinois, USA; pp 355-412 CAVOJSKZ V (1 982) [Comparative pedormance of queen honey bees of various ages I Pol' nohospoddrstvo 28(8)i 7 37 -7 42.
1202/85 DIEHNELT, W (1 986) A queen's best age: when to requeen.Gleanings in Bee Culture 114(3): 143.
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NOLAN, W J (1925\ The brood-rearing cycle of the honeybee. US Department Agriculture; Technical Bulletin No. 1349; 56 pp SOLLER, M; BAR-COHEN, R (1967) Some observations on the heri-
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SZABO, T l; LEFKOVITCH, L P (1 989) Effect of brood production and population size on honey production of honeybee colonies in Alberta, Canada. Apidologie 20(21: 157-163. 527 /9o
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