Spring Phenology of Butterflies : The role of seasonal variation in life-cycle regulation

Animals and plants in temperate regions must adapt their life cycle to pronounced seasonal variation. The research effort that has gone into studying these cyclical life history events, or phenological traits, has increased greatly in recent decades. As phenological traits are often correlated to temperature, they are relevant to study in terms of understanding the effect of short term environmental variation as well as long term climate change. Because of this, changes in phenology are the most obvious and among the most commonly reported responses to climate change. Moreover, phenological traits are important for fitness as they determine the biotic and abiotic environment an individual encounters. Fine-tuning of phenology allows for synchronisation at a local scale to mates, food resources and appropriate weather conditions. On a between-population scale, variation in phenology may reflect regional variation in climate. Such differences can not only give insights to life cycle adaptation, but also to how populations may respond to environmental change through time. This applies both on an ecological scale through phenotypic plasticity as well as an evolutionary scale through genetic adaptation. In this thesis I have used statistical and experimental methods to investigate both the larger geographical patterns as well as mechanisms of fine-tuning of phenology of several butterfly species. The main focus, however, is on the orange tip butterfly, Anthocharis cardamines, in Sweden and the United Kingdom. I show a contrasting effect of spring temperature and winter condition on spring phenology for three out of the five studied butterfly species. For A. cardamines there are population differences in traits responding to these environmental factors between and within Sweden and the UK that suggest adaptation to local environmental conditions. All populations show a strong negative plastic relationship between spring temperature and spring phenology, while the opposite is true for winter cold duration. Spring phenology is shifted earlier with increasing cold duration. The environmental variables show correlations, for example, during a warm year a short winter delays phenology while a warm spring speeds phenology up. Correlations between the environmental variables also occur through space, as the locations that have long winters also have cold springs. The combined effects of these two environmental variables cause a complex geographical pattern of phenology across the UK and Sweden. When predicting phenology with future climate change or interpreting larger geographical patterns one must therefore have a good enough understanding of how the phenology is controlled and take the relevant environmental factors in to account. In terms of the effect of phenological change, it should be discussed with regards to change in life cycle timing among interacting species. For example, the phenology of the host plants is important for A. cardamines fitness, and it is also the main determining factor for oviposition. In summary, this thesis shows that the broad geographical pattern of phenology of the butterflies is formed by counteracting environmental variables, but that there also are significant population differences that enable fine-tuning of phenology according to the seasonal progression and variation at the local scale.

Anxiety disorders and depression and risk for drug use disorders across the life course: Sex and ethnic contrasts

The present study examined the linkage between mental (i.e., anxiety disorders and depression) and drug use disorders in a multi-ethnic (i.e., 25% Euro-American, 38% Hispanic/Latino, 33% African American, 4% other) sample of adults (N = 1638, age 18–93 years old). Risk for drug use disorders was examined, while attending to methodological issues of prior research including (1) psychiatric comorbidity, (2) variations in risk associated with sex, ethnicity, and age, and (3) temporal order between mental and drug use disorders. ^ Participants were assessed using the Composite International Diagnostic Interview (CIDI; World Health Organization, 1990). A life history calendar (Freedman et al., 1988) was used to aid the ordering of onsets of all disorders assessed. ^ Preliminary analysis indicated anxiety disorders and depression were significant predictors of drug use disorders, but after controlling for comorbidity and temporal order, anxiety disorders and depression were no longer predictive of drug use disorders. Findings are discussed in terms of their usefulness for prevention and treatment of drug use disorders. ^

Effect of ocean warming and acidification on the early life stages of subtropical Acropora spicifera

This study investigated the impacts of acidified seawater (pCO2 900 µatm) and elevated water temperature (+3 °C) on the early life history stages of Acropora spicifera from the subtropical Houtman Abrolhos Islands (28°S) in Western Australia. Settlement rates were unaffected by high temperature (27 °C, 250 µatm), high pCO2 (24 °C, 900 µatm), or a combination of both high temperature and high pCO2 treatments (27 °C, 900 µatm). There were also no significant differences in rates of post-settlement survival after 4 weeks of exposure between any of the treatments, with survival ranging from 60 to 70 % regardless of treatment. Similarly, calcification, as determined by the skeletal weight of recruits, was unaffected by an increase in water temperature under both ambient and high pCO2 conditions. In contrast, high pCO2 significantly reduced early skeletal development, with mean skeletal weight in the high pCO2 and combined treatments reduced by 60 and 48 %, respectively, compared to control weights. Elevated temperature appeared to have a partially mitigative effect on calcification under high pCO2; however, this effect was not significant. Our results show that rates of settlement, post-settlement survival, and calcification in subtropical corals are relatively resilient to increases in temperature. This is in marked contrast to the sensitivity to temperature reported for the majority of tropical larvae and recruits in the literature. The subtropical corals in this study appear able to withstand an increase in temperature of 3 °C above ambient, indicating that they may have a wider thermal tolerance range and may not be adversely affected by initial increases in water temperature from subtropical 24 to 27 °C. However, the reduction in skeletal weight with high pCO2 indicates that early skeletal formation will be highly vulnerable to the changes in ocean pCO2 expected to occur over the twenty-first century, with implications for their longer-term growth and resilience.

Biological and Physical Factors Affecting the Natural History and Evolution of Encapsulated Development

The evolution of reproductive strategies involves a complex calculus of costs and benefits to both parents and offspring. Many marine animals produce embryos packaged in tough egg capsules or gelatinous egg masses attached to benthic surfaces. While these egg structures can protect against environmental stresses, the packaging is energetically costly for parents to produce. In this series of studies, I examined a variety of ecological factors affecting the evolution of benthic development as a life history strategy. I used marine gastropods as my model system because they are incredibly diverse and abundant worldwide, and they exhibit a variety of reproductive and developmental strategies.

The first study examines predation on benthic egg masses. I investigated: 1) behavioral mechanisms of predation when embryos are targeted (rather than the whole egg mass); 2) the specific role of gelatinous matrix in predation. I hypothesized that gelatinous matrix does not facilitate predation. One study system was the sea slug Olea hansineensis, an obligate egg mass predator, feeding on the sea slug Haminoea vesicula. Olea fed intensely and efficiently on individual Haminoea embryos inside egg masses but showed no response to live embryos removed from gel, suggesting that gelatinous matrix enables predation. This may be due to mechanical support of the feeding predator by the matrix. However, Haminoea egg masses outnumber Olea by two orders of magnitude in the field, and each egg mass can contain many tens of thousands of embryos, so predation pressure on individuals is likely not strong. The second system involved the snail Nassarius vibex, a non-obligate egg mass predator, feeding on the polychaete worm Clymenella mucosa. Gel neither inhibits nor promotes embryo predation for Nassarius, but because it cannot target individual embryos inside an egg mass, its feeding is slow and inefficient, and feeding rates in the field are quite low. However, snails that compete with Nassarius for scavenged food have not been seen to eat egg masses in the field, leaving Nassarius free to exploit the resource. Overall, egg mass predation in these two systems likely benefits the predators much more than it negatively affects the prey. Thus, selection for environmentally protective aspects of egg mass production may be much stronger than selection for defense against predation.

In the second study, I examined desiccation resistance in intertidal egg masses made by Haminoea vesicula, which preferentially attaches its flat, ribbon-shaped egg masses to submerged substrata. Egg masses occasionally detach and become stranded on exposed sand at low tide. Unlike adults, the encased embryos cannot avoid desiccation by selectively moving about the habitat, and the egg mass shape has high surface-area-to-volume ratio that should make it prone to drying out. Thus, I hypothesized that the embryos would not survive stranding. I tested this by deploying individual egg masses of two age classes on exposed sand bars for the duration of low tide. After rehydration, embryos midway through development showed higher rates of survival than newly-laid embryos, though for both stages survival rates over 25% were frequently observed. Laboratory desiccation trials showed that >75% survival is possible in an egg mass that has lost 65% of its water weight, and some survival (<25%) was observed even after 83% water weight lost. Although many surviving embryos in both experiments showed damage, these data demonstrate that egg mass stranding is not necessarily fatal to embryos. They may be able to survive a far greater range of conditions than they normally encounter, compensating for their lack of ability to move. Also, desiccation tolerance of embryos may reduce pressure on parents to find optimal laying substrata.

The third study takes a big-picture approach to investigating the evolution of different developmental strategies in cone snails, the largest genus of marine invertebrates. Cone snail species hatch out of their capsules as either swimming larvae or non-dispersing forms, and their developmental mode has direct consequences for biogeographic patterns. Variability in life history strategies among taxa may be influenced by biological, environmental, or phylogenetic factors, or a combination of these. While most prior research has examined these factors singularly, my aim was to investigate the effects of a host of intrinsic, extrinsic, and historical factors on two fundamental aspects of life history: egg size and egg number. I used phylogenetic generalized least-squares regression models to examine relationships between these two egg traits and a variety of hypothesized intrinsic and extrinsic variables. Adult shell morphology and spatial variability in productivity and salinity across a species geographic range had the strongest effects on egg diameter and number of eggs per capsule. Phylogeny had no significant influence. Developmental mode in Conus appears to be influenced mostly by species-level adaptations and niche specificity rather than phylogenetic conservatism. Patterns of egg size and egg number appear to reflect energetic tradeoffs with body size and specific morphologies as well as adaptations to variable environments. Overall, this series of studies highlights the importance of organism-scale biotic and abiotic interactions in evolutionary patterns.

Age and Life Course Location as Interpretive Resources for Decisions Regarding Disclosure of HIV to Parents and Children: Findings from the HIV and Later Life Study

Studies of disclosure among older people living with HIV (PLWH) are uninformed by critical social-gerontological approaches that can help us to appreciate how older PLWH see and treat age as relevant to disclosure of their HIV status. These approaches include an ethnomethodologically-informed social constructionism that explores how ‘the’ life course (a cultural framework depicting individuals’ movement through predictable developmental stages from birth to death) is used as an interpretive resource for determining self and others’ characteristics, capacities, and social circumstances: a process Rosenfeld and Gallagher (2002) termed ‘lifecoursing’. Applying this approach to our analysis of 74 life-history interviews and three focus groups with older (aged 50+) people living with HIV in the United Kingdom, we uncover the central role that lifecoursing plays in participants’ decision-making surrounding disclosure of their HIV to their children and/or older parents. Analysis of participants’ accounts uncovered four criteria for disclosure: the relevance of their HIV to the other, the other’s knowledge about HIV, the likelihood of the disclosure causing the other emotional distress, and the other’s ability to keep the disclosed confidential. To determine if these criteria were met in relation to specific children and/or elders, participants engaged in lifecoursing, evaluating the other’s knowledge of HIV, and capacity to appropriately manage the disclosure, by reference to their age. The use of assumptions about age and life-course location in decision-making regarding disclosure of HIV reflects a more nuanced engagement with age in the disclosure decision-making process than has been captured by previous research into HIV disclosure, including on the part of people aging with HIV.

Learning to construct our identities over the life course: a study with lesbian, gay, bisexual and transgender adults in Scotland

To date, adult educational research has had a limited focus on lesbian, gay, bisexual and transgendered (LGBT) adults and the learning processes in which they engage across the life course. Adopting a biographical and life history methodology, this study aimed to critically explore the potentially distinctive nature and impact of how, when and where LGBT adults learn to construct their identities over their lives. In-depth, semi-structured interviews, dialogue and discussion with LGBT individuals and groups provided rich narratives that reflect shifting, diverse and multiple ways of identifying and living as LGBT. Participants engage in learning in unique ways that play a significant role in the construction and expression of such identities, that in turn influence how, when and where learning happens. Framed largely by complex heteronormative forces, learning can have a negative, distortive impact that deeply troubles any balanced, positive sense of being LGBT, leading to self- censoring, alienation and in some cases, hopelessness. However, learning is also more positively experiential, critically reflective, inventive and queer in nature. This can transform how participants understand their sexual identities and the lifewide spaces in which they learn, engendering agency and resilience. Intersectional perspectives reveal learning that participants struggle with, but can reconcile the disjuncture between evolving LGBT and other myriad identities as parents, Christians, teachers, nurses, academics, activists and retirees. The study’s main contributions lie in three areas. A focus on LGBT experience can contribute to the creation of new opportunities to develop intergenerational learning processes. The study also extends the possibilities for greater criticality in older adult education theory, research and practice, based on the continued, rich learning in which participants engage post-work and in later life. Combined with this, there is scope to further explore the nature of ‘life-deep learning’ for other societal groups, brought by combined religious, moral, ideological and social learning that guides action, beliefs, values, and expression of identity. The LGBT adults in this study demonstrate engagement in distinct forms of life-deep learning to navigate social and moral opprobrium. From this they gain hope, self-respect, empathy with others, and deeper self-knowledge.

Effects of flow regulation in the life-cycles of a mediterranean cyprinid species, the Iberian chub (Squalius carolitertii Doadrio, 1987)

Streamflow is considered a driver of inter and intra‐specific life‐history differences among freshwater fish. Therefore, dams and related flow regulation, can have deleterious impacts on their life‐cycles. The main objective of this study is to assess the effects of flow regulation on the growth and reproduction of a non‐migratory fish species. During one year, samples were collected from two populations of Iberian chub, inhabiting rivers with non‐regulated and regulated flow regimes. Flow regulation for water derivation promoted changes in chub’s condition, duration of gonad maturation and spawning, fecundity and oocyte size. However, this non‐migratory species was less responsive to streamflow regulation than a migratory species analysed. Findings from this study are important to understand changes imposed by regulated rivers on fish and can be used as guidelines for flow requirements implementations; RESUMO: O caudal é um dos fatores responsáveis pelo funcionamento dos ciclos de vida das espécies piscícolas dulciaquícolas. As barragens, e a regularização de caudal associada, podem ter impactes nos ciclos de vida destas espécies. O objetivo deste estudo prende‐se com a avaliação dos efeitos da regularização de caudal no crescimento e reprodução de uma espécie piscícola não‐migradora. A análise de amostras recolhidas em populações de escalo do Norte provenientes de dois rios de caudal regularizado e não regularizado, identificaram impactes significativos a nível da condição corporal, da maturação das gónadas e desova, da fecundidade e da dimensão dos oócitos. Esta espécie não‐migradora parece ser menos responsiva à artificialização do caudal que uma espécie migradora previamente analisada. Estes resultados permitem compreender as alterações impostas pela regularização do caudal e podem ser usados em programas de reabilitação fluvial.

The effects of habitat fragmentation on the demography and population genetic structure of Uromys Caudimaculatus

Habitat fragmentation can have an impact on a wide variety of biological processes including abundance, life history strategies, mating system, inbreeding and genetic diversity levels of individual species. Although fragmented populations have received much attention, ecological and genetic responses of species to fragmentation have still not been fully resolved. The current study investigated the ecological factors that may influence the demographic and genetic structure of the giant white-tailed rat (Uromys caudimaculatus) within fragmented tropical rainforests. It is the first study to examine relationships between food resources, vegetation attributes and Uromys demography in a quantitative manner. Giant white-tailed rat densities were strongly correlated with specific suites of food resources rather than forest structure or other factors linked to fragmentation (i.e. fragment size). Several demographic parameters including the density of resident adults and juvenile recruitment showed similar patterns. Although data were limited, high quality food resources appear to initiate breeding in female Uromys. Where data were sufficient, influx of juveniles was significantly related to the density of high quality food resources that had fallen in the previous three months. Thus, availability of high quality food resources appear to be more important than either vegetation structure or fragment size in influencing giant white-tailed rat demography. These results support the suggestion that a species’ response to fragmentation can be related to their specific habitat requirements and can vary in response to local ecological conditions. In contrast to demographic data, genetic data revealed a significant negative effect of habitat fragmentation on genetic diversity and effective population size in U. caudimaculatus. All three fragments showed lower levels of allelic richness, number of private alleles and expected heterozygosity compared with the unfragmented continuous rainforest site. Populations at all sites were significantly differentiated, suggesting restricted among population gene flow. The combined effects of reduced genetic diversity, lower effective population size and restricted gene flow suggest that long-term viability of small fragmented populations may be at risk, unless effective management is employed in the future. A diverse range of genetic reproductive behaviours and sex-biased dispersal patterns were evident within U. caudimaculatus populations. Genetic paternity analyses revealed that the major mating system in U. caudimaculatus appeared to be polygyny at sites P1, P3 and C1. Evidence of genetic monogamy, however, was also found in the three fragmented sites, and was the dominant mating system in the remaining low density, small fragment (P2). High variability in reproductive skew and reproductive success was also found but was less pronounced when only resident Uromys were considered. Male body condition predicted which males sired offspring, however, neither body condition nor heterozygosity levels were accurate predictors of the number of offspring assigned to individual males or females. Genetic spatial autocorrelation analyses provided evidence for increased philopatry among females at site P1, but increased philopatry among males at site P3. This suggests that male-biased dispersal occurs at site P1 and female-biased dispersal at site P3, implying that in addition to mating systems, Uromys may also be able to adjust their dispersal behaviour to suit local ecological conditions. This study highlights the importance of examining the mechanisms that underlie population-level responses to habitat fragmentation using a combined ecological and genetic approach. The ecological data suggested that habitat quality (i.e. high quality food resources) rather than habitat quantity (i.e. fragment size) was relatively more important in influencing giant white-tailed rat demographics, at least for the populations studied here . Conversely, genetic data showed strong evidence that Uromys populations were affected adversely by habitat fragmentation and that management of isolated populations may be required for long-term viability of populations within isolated rainforest fragments.

Certainties and uncertainties : ethics and professional identities of early childhood educators

This study is an inquiry into the professional identity constructions of early childhood educators, where identity is conceptualised as social and contextual. Through a genealogical analysis of narratives of four Queensland early childhood teachers, the thesis renders as problematic universal and fixed notions of what it is to be an early childhood professional. The data are the four teachers’ professional life history narratives recounted through a series of conversational interviews with each participant. As they spoke about professionalism and ethics, these teachers struggled to locate themselves as professionals, as they drew on a number of dominant discourses available to them. These dominant discourses were located and mapped through analysis of the participants’ talk about relationships with parents, colleagues and authorities. Genealogical analysis enabled multiple readings of the ways in which the participants’ talk held together certainties and uncertainties, as they recounted their experiences and spoke of early childhood expertise, relational engagement and ethics. The thesis concludes with suggestions for ways to support early childhood teachers and pre-service teachers to both engage with and resist normative processes and expectations of professional identity construction. In so doing, multiple and contextual opportunities can be made available when it comes to being professional and ‘doing’ ethics. The thesis makes an argument for new possibilities for thinking and speaking professional identities that include both certainty and uncertainty, comfort and discomfort, and these seemingly oppositional terms are held together in tension, with an insistence that both are necessary and true. The use of provocations offers tools through which pre-service teachers, teachers and teacher educators can access new positions associated with certainties and uncertainties in professional identities. These new positions call for work that supports experiences of ‘de-comfort’ – that is, experiences that encourage early childhood educators to step away from the comfort zones that can become part of expertise, professional relationships and ethics embedded within normative representations of what it is to be an early childhood professional.

Towards a method of constructing self-efficacy narratives.

This paper outlines a method of constructing narratives about an individual’s self-efficacy. Self-efficacy is defined as “people’s judgments of their capabilities to organise and execute courses of action required to attain designated types of performances” (Bandura, 1986, p. 391), and as such represents a useful construct for thinking about personal agency. Social cognitive theory provides the theoretical framework for understanding the sources of self-efficacy, that is, the elements that contribute to a sense of self-efficacy. The narrative approach adopted offers an alternative to traditional, positivist psychology, characterised by a preoccupation with measuring psychological constructs (like self-efficacy) by means of questionnaires and scales. It is argued that these instruments yield scores which are somewhat removed from the lived experience of the person—respondent or subject—associated with the score. The method involves a cyclical and iterative process using qualitative interviews to collect data from participants – four mature aged university students. The method builds on a three-interview procedure designed for life history research (Dolbeare & Schuman, cited in Seidman, 1998). This is achieved by introducing reflective homework tasks, as well as written data generated by research participants, as they are guided in reflecting on those experiences (including behaviours, cognitions and emotions) that constitute a sense of self-efficacy, in narrative and by narrative. The method illustrates how narrative analysis is used “to produce stories as the outcome of the research” (Polkinghorne, 1995, p.15), with detail and depth contributing to an appreciation of the ‘lived experience’ of the participants. The method is highly collaborative, with narratives co-constructed by researcher and research participants. The research outcomes suggest an enhanced understanding of self-efficacy contributes to motivation, application of effort and persistence in overcoming difficulties. The paper concludes with an evaluation of the research process by the students who participated in the author’s doctoral study.

Cytochrome p450 isozyme protein verified in the skin of southern hemisphere humpback whales (megaptera novaeangliae) : implications for biochemical biomarker assessment

Large mysticete whales represent a unique challenge for chemical risk assessment. Few epidemiological investigations are possible due to the low incidence of adult stranding events. Similarly their often extreme life-history adaptations of prolonged migration and fasting challenge exposure assumptions. Molecular biomarkers offer the potential to complement information yielded through tissue chemical analysis, as well as providing evidence of a molecular response to chemical exposure. In this study we confirm the presence of cytochrome P450 reductase (CPR) and cytochrome P450 isoenzyme 1A1 (CYP1A1) in epidermal tissue of southern hemisphere humpback whales (Megaptera novaeangliae). The detection of CYP1A1 in the integument of the humpback whale affords the opportunity for further quantitative non-destructive investigations of enzyme activity as a function of chemical stress.

Nascent entrepreneurship and the developing individual : early entrepreneurial competence in adolescence and venture creation success during the career

What predicts a person's venture creation success over the course of the career, such as making progress in the venture creation process and multiple successful venture creations? Applying a life span approach of human development, this study examined the effect of early entrepreneurial competence in adolescence, which was gathered retrospectively by means of the Life History Calendar method. Human and social capitals during the founding process were investigated as mediators between adolescent competence and performance. Findings were derived from regression analyses on the basis of prospective and retrospective data from two independent samples (N = 88 nascent founders; N = 148 founders). We found that early entrepreneurial competence in adolescence had a positive effect on making progress in the venture creation process. Nascent founders' current human and social capital also had a direct effect, but it did not mediate the effect of early competences. Finally, the data revealed that early entrepreneurial competence in adolescence positively predicted habitual entrepreneurship (multiple successful venture creations) exhibited over a longer period of the individual career (specifically, 18 years). In line with the results from prospective longitudinal studies on early precursors of entrepreneurship, our findings underscore the long neglected importance of adolescent development in the explanation of entrepreneurial performance during the subsequent working life.

Ruling Class Men: Money, Sex, Power

What is it like to be a master of the universe? The authors have researched the desires and fears of the world's most powerful men. The Murdochs, Packers, Kennedys, Agnellis and other men like them, directly determine the fates of thousands and influence the future of the world like no other people. Described as 'sacred monsters' by one of their own, they are carefully created to be what they are and to enjoy shaping the world in their own likeness. To learn about these often reclusive men, the authors extended the life-history technique to interrogate autobiographies, diaries and biographies and have created a composite picture, a collective portrait, of tycoons over three generations. The book carefully explores the childhoods, schooling, work and play, sexual activities, marriages and deaths of the wealthiest men who have ever lived. It exposes the nature of ruling-class masculinity itself.

Mitochondrial genome deletions and minicircles are common in lice (Insecta: Phthiraptera)

Background The gene composition, gene order and structure of the mitochondrial genome are remarkably stable across bilaterian animals. Lice (Insecta: Phthiraptera) are a major exception to this genomic stability in that the canonical single chromosome with 37 genes found in almost all other bilaterians has been lost in multiple lineages in favour of multiple, minicircular chromosomes with less than 37 genes on each chromosome. Results Minicircular mt genomes are found in six of the ten louse species examined to date and three types of minicircles were identified: heteroplasmic minicircles which coexist with full sized mt genomes (type 1); multigene chromosomes with short, simple control regions, we infer that the genome consists of several such chromosomes (type 2); and multiple, single to three gene chromosomes with large, complex control regions (type 3). Mapping minicircle types onto a phylogenetic tree of lice fails to show a pattern of their occurrence consistent with an evolutionary series of minicircle types. Analysis of the nuclear-encoded, mitochondrially-targetted genes inferred from the body louse, Pediculus, suggests that the loss of mitochondrial single-stranded binding protein (mtSSB) may be responsible for the presence of minicircles in at least species with the most derived type 3 minicircles (Pediculus, Damalinia). Conclusions Minicircular mt genomes are common in lice and appear to have arisen multiple times within the group. Life history adaptive explanations which attribute minicircular mt genomes in lice to the adoption of blood-feeding in the Anoplura are not supported by this expanded data set as minicircles are found in multiple non-blood feeding louse groups but are not found in the blood-feeding genus Heterodoxus. In contrast, a mechanist explanation based on the loss of mtSSB suggests that minicircles may be selectively favoured due to the incapacity of the mt replisome to synthesize long replicative products without mtSSB and thus the loss of this gene lead to the formation of minicircles in lice.