title: Reproductive competition in the context of a superorganism

authors: Carta, Epernay

abstract: Population density is a fundamental ecological factor that shapes the intensity of competition within a species. Increased density often intensifies competition for resources and mates, driving the evolution of diverse adaptive responses in behaviour, morphology and physiology. A key area where density-mediated competition manifests is in sperm competition. This occurs when sperm from multiple males compete to fertilise the ova of a female. Species facing intense competition exhibit a range of adaptations, including adjustments in sperm production, ejaculate characteristics, and mating strategies. Elevated population density is a major factor that can exacerbate sperm competition, further amplifying these evolutionary pressures. Within this context, this thesis investigates the effects of apiary density on reproductive strategies and drone physiology in the honey bee (Apis mellifera), a highly social insect that exhibits a complex interplay between individual and colony- level responses. Apiary density, characterised by the number of colonies within a defined area, was manipulated across three treatments: low (8 colonies), medium (60-68 colonies), and high (120 colonies). Drone brood production and comb allocation (both drone and worker) were measured monthly to assess colony-level investment in reproduction versus worker production. Total sperm count and the proportion of viable sperm were assessed in drones collected from each density treatment to evaluate sperm competition dynamics. Heat Shock Protein 70 (HSP70) levels, a biomarker of cellular stress, were quantified in drones from each treatment using an enzyme-linked immunosorbent assay. Colonies in high-density apiaries exhibited significantly increased drone brood production and higher total sperm counts compared to those in low-density apiaries. However, worker comb production was significantly reduced in high-density apiaries, suggesting a trade-off between resource allocation towards reproduction and colony maintenance. Sperm viability varied across densities and seasons, with low-density treatments having a higher proportion of viable sperm in early summer and medium-density treatments showing higher viability in late summer. This suggests that while high-density may favour increased sperm quantity, lower densities might facilitate greater investment in sperm quality. Apiary density significantly predicted HSP70 concentration in drones, with drones from high-density apiaries displaying significantly elevated levels, indicating increased physiological stress. These findings suggest that colonies in high-density environments experience intensified sperm competition, responding by increasing drone and sperm production. However, the observed variation in sperm viability and reduced worker comb production implies that this augmented reproductive investment may compromise overall colony function. The elevated HSP70 levels further corroborate that drones in high-density apiaries are subjected to greater physiological stress. Crucially, this study provides a unique perspective by demonstrating that apiary density, a factor often controlled in beekeeping, acts as a significant driver of reproductive strategies and physiological stress at both the colony and individual drone level within a superorganism. This highlights the profound impact of a managed environmental parameter on the evolutionary dynamics of a highly social species, offering critical insights for sustainable apicultural practices.

text: Reproductive competition in the context of a superorganism Population density is a fundamental ecological factor that shapes the intensity of competition within a species. Increased density often intensifies competition for resources and mates, driving the evolution of diverse adaptive responses in behaviour, morphology and physiology. A key area where density-mediated competition manifests is in sperm competition. This occurs when sperm from multiple males compete to fertilise the ova of a female. Species facing intense competition exhibit a range of adaptations, including adjustments in sperm production, ejaculate characteristics, and mating strategies. Elevated population density is a major factor that can exacerbate sperm competition, further amplifying these evolutionary pressures. Within this context, this thesis investigates the effects of apiary density on reproductive strategies and drone physiology in the honey bee (Apis mellifera), a highly social insect that exhibits a complex interplay between individual and colony- level responses. Apiary density, characterised by the number of colonies within a defined area, was manipulated across three treatments: low (8 colonies), medium (60-68 colonies), and high (120 colonies). Drone brood production and comb allocation (both drone and worker) were measured monthly to assess colony-level investment in reproduction versus worker production. Total sperm count and the proportion of viable sperm were assessed in drones collected from each density treatment to evaluate sperm competition dynamics. Heat Shock Protein 70 (HSP70) levels, a biomarker of cellular stress, were quantified in drones from each treatment using an enzyme-linked immunosorbent assay. Colonies in high-density apiaries exhibited significantly increased drone brood production and higher total sperm counts compared to those in low-density apiaries. However, worker comb production was significantly reduced in high-density apiaries, suggesting a trade-off between resource allocation towards reproduction and colony maintenance. Sperm viability varied across densities and seasons, with low-density treatments having a higher proportion of viable sperm in early summer and medium-density treatments showing higher viability in late summer. This suggests that while high-density may favour increased sperm quantity, lower densities might facilitate greater investment in sperm quality. Apiary density significantly predicted HSP70 concentration in drones, with drones from high-density apiaries displaying significantly elevated levels, indicating increased physiological stress. These findings suggest that colonies in high-density environments experience intensified sperm competition, responding by increasing drone and sperm production. However, the observed variation in sperm viability and reduced worker comb production implies that this augmented reproductive investment may compromise overall colony function. The elevated HSP70 levels further corroborate that drones in high-density apiaries are subjected to greater physiological stress. Crucially, this study provides a unique perspective by demonstrating that apiary density, a factor often controlled in beekeeping, acts as a significant driver of reproductive strategies and physiological stress at both the colony and individual drone level within a superorganism. This highlights the profound impact of a managed environmental parameter on the evolutionary dynamics of a highly social species, offering critical insights for sustainable apicultural practices.

year: 2025

title: The Common European Framework of Reference for Languages: Should Pacific and Pacific Rim countries get on board the bandwagon?

authors: Crombie, Winifred

subjects: Europe

abstract: For many bureaucrats, the Common European Framework of Reference for Languages (CEFR) has an almost irresistible appeal. All that users have to do, apparently, is read its approximately 200 pages (plus appendices) and they will “find ... all [they] need to describe [their] objectives, methods and products” (Council of Europe (CoE), p. xii). Having done that, they will be in a position to “overcome the barriers to communication . . . arising from . . . different educational systems” and “facilitate . . . mobility through . . . mutual recognition of qualifications” (p. 1). The CEFR has some very influential advocates, both within and outside of Europe. Investment in it is extensive in some areas of the world. It emerges out of an organisation whose ethical credentials are beyond question, one that has been responsible for some interesting innovations in the area of language education. In such a context, it is not surprising that many countries outside of Europe, including Pacific and Pacific Rim countries, are developing an interest in it. On the basis of a close reading of the CEFR, this article concludes that it has considerably less to offer than it claims and, therefore, advises language educators in Pacific and Pacific Rim countries, particularly those involved in the teaching of indigenous languages, to be extremely cautious about adopting the approaches it recommends too readily.

text: The Common European Framework of Reference for Languages: Should Pacific and Pacific Rim countries get on board the bandwagon? For many bureaucrats, the Common European Framework of Reference for Languages (CEFR) has an almost irresistible appeal. All that users have to do, apparently, is read its approximately 200 pages (plus appendices) and they will “find ... all [they] need to describe [their] objectives, methods and products” (Council of Europe (CoE), p. xii). Having done that, they will be in a position to “overcome the barriers to communication . . . arising from . . . different educational systems” and “facilitate . . . mobility through . . . mutual recognition of qualifications” (p. 1). The CEFR has some very influential advocates, both within and outside of Europe. Investment in it is extensive in some areas of the world. It emerges out of an organisation whose ethical credentials are beyond question, one that has been responsible for some interesting innovations in the area of language education. In such a context, it is not surprising that many countries outside of Europe, including Pacific and Pacific Rim countries, are developing an interest in it. On the basis of a close reading of the CEFR, this article concludes that it has considerably less to offer than it claims and, therefore, advises language educators in Pacific and Pacific Rim countries, particularly those involved in the teaching of indigenous languages, to be extremely cautious about adopting the approaches it recommends too readily.

year: 2010-02

title: Delivering the Maori-language newspapers on the Internet

authors: Apperley, Mark

subjects: computer science

abstract: Although any collection of historical newspapers provides a particularly rich and valuable record of events and social and political commentary, the content tends to be difficult to access and extremely time-consuming to browse or search. The advent of digital libraries has meant that for electronically stored text, full-text searching is now a tool readily available for researchers, or indeed anyone wishing to have asscess to specific information in text. Text in this form can be readily distributed via CD-ROM or the Internet, with a significant impact on accessibility over traditional microfiche or hard-copy distribution. For the majority of text being generated de nouveau, availability in electronic form is standard, and hence the increasing use of full-text search facilities. However, for legacy text available only in printed form, the provision of these electronic search tools is dependent on the prior electronic capture of digital facsimile images of the printed text, followed by the conversion of these images to electronic text through the process of optical character recognition (OCR). This article describes a project undertaken at the University of Waikato over the period 1999 to 2001 to produce a full-text searchable version of the Niupepa or Maori- language newspaper collection for delivery over the Internet.

text: Delivering the Maori-language newspapers on the Internet Although any collection of historical newspapers provides a particularly rich and valuable record of events and social and political commentary, the content tends to be difficult to access and extremely time-consuming to browse or search. The advent of digital libraries has meant that for electronically stored text, full-text searching is now a tool readily available for researchers, or indeed anyone wishing to have asscess to specific information in text. Text in this form can be readily distributed via CD-ROM or the Internet, with a significant impact on accessibility over traditional microfiche or hard-copy distribution. For the majority of text being generated de nouveau, availability in electronic form is standard, and hence the increasing use of full-text search facilities. However, for legacy text available only in printed form, the provision of these electronic search tools is dependent on the prior electronic capture of digital facsimile images of the printed text, followed by the conversion of these images to electronic text through the process of optical character recognition (OCR). This article describes a project undertaken at the University of Waikato over the period 1999 to 2001 to produce a full-text searchable version of the Niupepa or Maori- language newspaper collection for delivery over the Internet.

year: 2002