SummaryHere we cover a wide range of methods currently in use and recommended in modern queen rearing, selection and breeding. The recommendations are meant to equally serve as standards for both scientific and practical beekeeping purposes. The basic conditions and different management techniques for queen rearing are described, including recommendations for suitable technical equipment. As the success of breeding programmes strongly depends on the selective mating of queens, a subchapter is dedicated to the management and quality control of mating stations. Recommendations for the handling and quality control of queens complete the queen rearing section. The improvement of colony traits usually depends on a comparative testing of colonies. Standardized recommendations for the organization of performance tests and the measurement of the most common selection characters are presented. Statistical methods and data preconditions for the estimation of breeding values which integrate pedigree and performance data from as many colonies as possible are described as the most efficient selection method for large populations. Alternative breeding programmes for small populations or certain scientific questions are briefly mentioned, including also an overview of the young and fast developing field of molecular selection tools. Because the subject of queen rearing and selection is too large to be covered within this paper, plenty of references are given to facilitate comprehensive studies. Métodos estándar para la cría y selección de reinas de Apis mellifera ResumenSe describe una amplia gama de métodos actualmente en uso y recomendables sobre la cría actual de reinas, su selección y cruzamiento. Las recomendaciones tienen el propósito de servir de igual forma como estándares para fines apícolas tanto científicos como prácticos. Se describen las condiciones básicas y las diferentes técnicas de manejo para la cría de reinas, incluyendo recomendaciones para el equipo técnico adecuado. Dado que el éxito de los programas de mejora depende en gran medida el apareamiento selectivo de reinas, se dedica un subcapítulo a la gestión y control de calidad de las estaciones de apareamiento. Las recomendaciones para el manejo y control de calidad de las reinas completan la sección de cría de reinas. La mejora de las características de colonias por lo general, depende de ensayos comparativos entre colonias. Se presentan recomendaciones normalizadas para la organización de pruebas de rendimiento y la medición de los caracteres de selección más comunes. Aquellos métodos estadísticos y condiciones previas de datos para la estimación de valores de cruzamiento que integren los datos genealógicos y de rendimiento de tantas colonias como sea posible, se describen como los métodos de

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SUMMARYThe ectoparasitic mite Varroa destructor and honey bee pathogenic viruses have been implicated in the recent demise of honey bee colonies. Several studies have shown that the combination of V. destructor and deformed wing virus (DWV) poses an especially serious threat to honey bee health. Mites transmitting virulent forms of DWV may cause fatal DWV infections in the developing bee, while pupae parasitised by mites not inducing or activating overt DWV infections may develop normally. Adult bees respond to brood diseases by removing affected brood. This hygienic behaviour is an essential part of the beesʼ immune response repertoire and is also shown towards mite-parasitised brood. However, it is still unclear whether the bees react towards the mite in the brood cell or rather towards the damage done to the brood. We hypothesised that the extent of mite-associated damage rather than the mere presence of parasitising mites triggers hygienic behaviour. Hygienic behaviour assays performed with mites differing in their potential to transmit overt DWV infections revealed that brood parasitised by ʻvirulentʼ mites (i.e. mites with a high potential to induce fatal DWV infections in parasitised pupae) were removed significantly more often than brood parasitised by ʻless virulentʼ mites (i.e. mites with a very low potential to induce overt DWV infections) or non-parasitised brood. Chemical analyses of brood odour profiles suggested that the bees recognise severely affected brood by olfactory cues. Our results suggest that bees show selective, damage-dependent hygienic behaviour, which may be an economic way for colonies to cope with mite infestation. Supplementary material available online at

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The hygienic behaviour of the honey bees is considered to be a potential characteristic associated with resistance to Varroa destructor n.sp. In this study the heritability of the hygienic behaviour of Apis mellifera L. bees was estimated on the basis of the mother±daughter regression. Data were obtained from measurements of the bees' hygienic behaviour towards V. destructor-infested cells and towards pin-killed sealed brood. The heritability for the hygienic behaviour towards V. destructorinfested brood cells was h 2 0.18 (+ 0.27) and h 2 0.36 (+ 0.30) for the hygienic behaviour towards dead brood cells. The repeatability was likewise higher for the pin-killed brood assay (W 0.46) compared with the assay using living mites-infested brood cells (W 0.24). The genetic correlation between the behavioural responses to either the mite-infested or pin-killed brood cells was calculated to be r g 0.61 (+ 0.51) and the phenotypic correlation to be r p 0.11 (p 0.28, n 100). Since hygienic colonies demonstrate resistance to brood diseases such as American foulbrood and chalkbrood, it may be worthwhile to intensify the expression of the hygienic behaviour through selective breeding and thus strengthen these potential characteristics associated with resistance to V. destructor in honey bee stock. ZusammenfassungHeritabilita È t des Varroa-spezi®schen Hygieneverhaltens der Honigbienen (Hymenoptera:Apidae) Dem Hygieneverhalten der Honigbienen wird als potentieller Varroa-Toleranzfaktor besondere Beachtung geschenkt. In dieser Untersuchung wurde ± auf der Basis der Mutter-Tochter-Regression ± die Heritabilita Èt des Hygieneverhaltens von Apis mellifera L. ermittelt. Als Datengrundlage dienten Quanti®zierungen des Hygieneverhaltens der Bienen gegenu È ber mit Varroa destructor n.sp. in®zierter und gegenu È ber toter (`genadelte') gedeckelter Bienenbrut. Der daraus ermittelte Heritabilita Ètswert lag fu È r das Hygieneverhalten gegenu È ber mit Varroamilben in®zierter Brut bei h 2 0.18 ( + 0.27) und bei h 2 0.36 (+ 0.30) fu È r das Hygieneverhalten gegenu È ber toter Brut. Auch die Wiederholbarkeit war a Èhnlich ho È her bei dem Hygieneverhalten gegenu È ber toter Brut (W 0.46) im Vergleich zu W 0.24 ermittelt am Hygieneverhalten der Bienen gegenu È ber experimentell mit lebenden Milben in®zierten Brutzellen. Die genetische Korrelation zwischen diesen Verhaltensreaktionen wurde als r g 0.61 (+ 0.51) errechnet und die pha Ènotypische Korrelation als r p 0.11 (p 0.28, n 100). Da hygienischen Bienenvo È lkern eine erho È hte Widerstandsfa Èhigkeit gegenu È ber Amerikanischer Faulbrut und der Kalkbrut zugesprochen wird, erscheint es lohnenswert das Hygieneverhalten durch Selektion zu fo È rdern, um so auch diesen potentiellen Varroatoleranz-Parameter bei den Honigbienen zu sta Èrken. U.S.