A Bolt on Year in Sustainability Studies. Dr Colin RA Hewitt

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1 A Bolt on Year in Sustainability Studies Dr Colin RA Hewitt

2 Discovery-led and Discovery-enabling Learning Strategy Our commitments Transformative teaching and learning We will ensure that every student has an opportunity to engage in a professionalising and transforming experience in a work placement or an internship, as a volunteer, an ambassador or a mentor. Our priorities, Transformative teaching and learning will be delivered by: Building internationalisation and sustainability into the curriculum and student experience

3 Our commitments Offering a vibrant, successful academic portfolio We will develop a distinctively flexible, exciting curriculum, so that students have opportunities to balance disciplinary depth and interdisciplinary breadth School of Biological Sciences Department of Geography Centre for Interdisciplinary Science Environmental Sustainability Strategy To promote and raise awareness of teaching and learning that provides students and staff with relevant sustainability literacy.

4 What is it? A zero-credit year/module between the 2nd and 3 rd year of any degree A year coherent with the core programme Setting: Industrial Institute or university EAA through the Erasmus Exchange Scheme Any other location worldwide or in the UK Awarded on a pass/fail basis

5 With a Year in Sustainability Studies Aim To give students experience of laboratory or qualitative research in the context of sustainable solutions to a global problems Subject-specific learning outcomes: Explain the responsibilities of the individual and corporate interests within a global community Describe contributions that differing disciplines can provide in seeking sustainable solutions to global problems Demonstrate how integration of diverse disciplines offer insight and practical suggestions for collaborative actions towards sustainable development Reflect on the key findings of laboratory or qualitative research in the contest of sustainability

6 With a Year in Sustainability Studies Non-subject-specific learning outcomes: Demonstrate the capacity to work on a sustainability-related project for > 9 months in an industrial setting, research institute or external university research Test an hypothesis by appropriate experimental techniques Conduct experimental procedures and demonstrate good laboratory practice AND/OR conduct qualitative or quantitative research Analyse and present experimental data in writing Locate appropriate literature sources and interpret their findings in relation to other work in the subject Produce a well written and presented report

7 Three Phases Can be taken in an industrial, institute or university setting in the EAA through the Erasmus Exchange Scheme, at any other location worldwide or in the UK. Phase 1 Sustainable Futures online programme Taken early in Semester 1 Introduction to Sustainable Development One Earth Energy and Climate Feeding the World Are we doomed? (Yes, almost certainly) Must pass (40%) to be awarded with a year in sustainability award

8 Three Phases Phase 2 Experimental/analytical research project Field-based Laboratory-based Qualitative Clearly related to sustainability and core degree

9 Three Phases Phase 2 Indicative projects for Biological and Natural Sciences students Genetically engineered crops and animals Microbial bioremediation of pollutants Biochemistry: bioenergy and biophotovoltaics Biosensors and pollution monitoring Sustainable biotechnology Sustainable fisheries - molecular approaches to traceability Microbiome barcoding and traceability Molecular markers in sustainable forestry Conservation genetics

10 Three Phases Phase 2 Indicative projects for Human Geography students Social movements for sustainability Justice in sustainability Economic alternatives to development Degrowth New understandings of prosperity Diverse knowledges of environmental value Corporate social responsibility

11 Three Phases Phase 3 Final Assessment Research project report in the style of a scientific or critical review paper Commentary on the sustainability context of the research project Discussion of the environmental social and economic aspects of the subject and the potential impact of the research project Must pass (40%) to be awarded with a year in sustainability award

12 Programme Approval Business and Academic cases Both written Needs approval at UPMG and in both colleges Up for discussion CSE have proposed a standard format and support for student industrial placements This includes An employability module Issues of governance Learning logs Monitoring & support If approved, the CMBSP would hope to join that programme with adaptations as appropriate

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14 Why Study Tropical Birds? Most tropical passerines little or no EPF Why? Asynchronous or synchronous breeding As nesting synchrony increases, so does EPF Most tropical birds nest asynchronously Cercomacroides tyrannina (F) Cercomacroides tyrannina (M) Studies of temperate zone birds have traditionally provided data to model generalities Tropical birds are just species oddities. Temperate zone bias

15 Temperate zone bias Why are temperate zone species thought to be typical? Where is the greatest diversity of avian species? Should we reverse questions about avian behaviour and physiology? Why do temperate birds lay so many eggs? Why do 90% of temperate species only defend territories when breeding whilst 87% of tropical species defend territories all year round Biodiversity.org

16 Climate an abiotic selection factor Tropical/subtropical zone Temperate zone

17 Impact of abiotic factors on biotic factors Suits synchronous breeding Suits asynchronous breeding Simmon R and Allen J,

18 Breeding season synchrony Hooded Warbler (Wilsonia citrina) Mangrove Swallow (Tachycineta Albilinea) %Clutch initiations %Clutch initiations Rufous-collared sparrow (Zonotrichia capensis) White-fronted Bee-eater, (Merops bullokoides) %Clutch initiations J F M A M J J A S O N D %Nesting J F M A M J J A S O N D

19 Selection factors Temperate Major selection factor is abiotic - climate Synchronous breeding season of two to three months Tropical Major selection factors are biotic food availability and predation Biotic selection shapes biodiversity, behaviour and morphology to a greater extent than in temperate zones e.g., broadly.. Frugivorous species breed in dry season Insectivorous species breed in the wet season Asynchronous breeding season four to eight months

20 Habitat disruption Biotic factors heavily impacted by disruption to habitat 15 million ha of tropical forest is cut per year (Wales is 2 million ha) Evolutionary history is being lost Conservation Rondônia1975 Rondônia 2009

21 Selection factors, biodiversity and latitude Accipitridae (Hawks and Kites) Troglodytidae (Wrens) Tyrannidae (Tyrant Flycatchers) Formicariidae (Antbirds) Rupornis magnirostris Troglodytes aedon Tyrannus savana Number of genera in each family Formicarius colma % of all passerine species breed in tropical regions High levels of adaptation to be expected

22 Behavioural adaptations Breeding Territoriality Food abundance Not independent variables

23 Breeding Extra-Pair Fertilisations Individual socially monogamous birds commonly participate in extra-pair copulations (EPC) that lead to extra-pair offspring (EPO) EPF strategically similar with leks Males provide an opportunity for females to assess a wide range of males for selection of the best possible traits Increase chances of males encountering females Outcome is that all fertilisations are extra-pair

24 Extra-Pair Fertilisations Neither random nor opportunistic strategic Covert visits to neighbouring territories Female advertisement of fertility Intense example of sperm competition Reflected by testis size Stuchbury and Morton 1995 Behaviour

25 What s in it for males? Sire more offspring without cost of care EPF and uncertain paternity imply that a male s EPO will be spread across several neighbouring nests Could focus the safety and productivity on the neighbourhood rather using all resources on their own social nest? Is there a self-interest to cooperate with other males and redirect paternal investment from care provided at their own nest towards neighbourhood roles? Could sustain public good among unrelated males in large groups be a mechanism of the evolution of cooperation? (Note that this is different from cooperative breeding ) Risk that an individual, who would be better off exploiting the public good, leaves others to pay the cooperative investment

26 What s in it for females? Females actively seek EPFs likely to be beneficial but how?? Risk of social mate withdrawing parental care if share of paternity becomes too low Benefits of a cooperative neighbourhood may outweigh the risk of lost care from her social mate? Females may seek EPC to ensure fertilization of their eggs - consistent with a fertility-insurance hypothesis, obtain more nuptial gifts or recruit increased paternal care at her nest

27 Genetic-benefit hypotheses of extra-pair mating Genetic quality of EPO improved by EPF with males of higher quality/more compatible genetics than their social partner Predicts EPO genetically/phenotypically superior than within- pair offspring (WPO) Compare features of EPO and WPO from the same nest only father different EPO found to be more fit than WPO w.r.t growth rate, fledging mass, size, or condition, survival to fledging, immune function and recruitment to the local breeding population in subsequent years Non-genetic explanations? Maternal effects?

28 Testosterone fluctuation an EPF-related adaptation? Temperate zone socially monogamous species Testosterone high during territory establishment and pair formation Testis size increases dramatically before breeding Testosterone drops once males start feeding young, Temperate zone polygynous species Lower male parental care - persistent mate attraction across a season Testosterone remains high maintains attraction and EPF success Tropical zone species Low testosterone all year Small testes High testosterone not required for territoriality and mate attraction Is high testosterone in temperate zone species is an adaptation to compete for mates and EPF during synchronous nesting?

29 Proximate cues for breeding Proximate factors that stimulate endocrine pathways to gonadal development In temperate species Long-term cues - photoperiod governs hormonal (LH & T) and gonadal changes between reproductive inactivity/activity Short-term cues - temperature, food, nest site availability lead to reproductive events Month Leicester Hour In tropical species Twelve hr to 12 hr 17 min sufficient to stimulate gonadal growth - not accompanied by changes in LH & T. Humidity and water availability Panamá Hour Month Hau et al., (1998) Proc. R. Soc. Lond. B 265, 89-95

30 Why no EPF in tropical species? EPF is the norm in socially monogamous, temperate passerines Tropical species: Clay Coloured Thrush Turdus grayi extensive EPF Most tropical passerines Dusky Antbird Cercomacroides tyrannina little EPF Why? A role for synchronous or synchronous breeding Most tropical birds nest asynchronously Turdus grayi Cercomacroides tyrannina (F) Cercomacroides tyrannina (M)

31 Why no EPF in tropical species? As nesting synchrony increases, so does EPF How can EPF be of benefit if the genetic-benefit hypothesis is correct? Stuchbury and Morton 1995 Behaviour

32 Is there a genetic contribution to the superior performance of EPO? EPO are larger and heavier and more likely to fledge than within pair offspring (WPO) Study of relationship between paternity and laying order in Blue Tits, Cyanistes caeruleus, Large clutches EPO typically occur in over 40% of broods Clutches hatch asynchronously over several days - size and mortality hierarchies Maggrath (2009) Current Biology 19, p

33 Laying order and hatch asynchrony Survival probability Wing length Proportion of EPO Body mass Tarsus length Relative laying order Hatch time (hr) Appears that EPO hatch earlier than WPO EPF more common in first half of clutch (Why?) Is development of EPO advanced compared with WPO? Maggrath (2009) Current Biology 19, p Hatch time (hr)

34 Doubts about evidence for the geneticbenefit hypotheses of EPF EPO hatch after WPO in clutches with greater synchrony Difference between EPO hatch and WPO hatch (hr) Hatching asynchrony (hr) Maggrath (2009) Current Biology 19, p % Difference between EPO and WPO EPO hatch before WPO in clutches with greater asynchrony Survival Wing Tarsus Body mass Offspring trait

35 Doubts about evidence for the geneticbenefit hypotheses of EPF Add to doubts about the strength of evidence for the genetic-benefit hypotheses of EPF Why don t tropical birds engage in EPF? Is there a role for territories? a role for clutch size? a role for food availability?

36 Territories Temperate zone Often migrate to small territory Territory establishment by male - defence essential to prevent EPF Song coincides with territoriality, mate attraction and threat of EPF Tropical zone Year-round residents holding large, stable territories High adult longevity and pairing Very low territory turnover Few opportunities to choose mates Defence is for real estate not against EPF Song used to defend territory, not to attract mates (when paired) or prevent EPF Primed, however, to attract mates e.g. Cercomacroides tyrannina

37 Experimental removal of Cercomacroides tyrannina from territory Cercomacroides tyrannina (F) Males replaced (n = 9, circles) Females replaced (n = 5, squares) Cercomacroides tyrannina (M) Morton et al. Behavioural Ecology 2000;11:

38 Clutch size and latitude Lack s hypothesis (1947) suggested clutch size determined by food abundance during the breeding period Northern species have large clutches because photoperiod is longer than those in the tropics. Life-history theory - high seasonality leads to high adult mortality high investment in reproduction and large clutch sizes because survival to next breeding season is low Ashmole suggested that high adult mortality reduces population density and increases per-individual resource availability in the breeding season Allows temperate birds to nourish large clutches

39 Clutch size Fewest species breed in higher latitude, temperate zones

40 Clutch size and latitude Precocial vs altricial mobility of offspring less intensive parenting Latitude - captures global environmental variation Energy availability more seasonal and reduced at higher latitudes Clutch sizes smallest in species with most aseasonal environments clutch size increases linearly with temperature seasonality Clutch size

41 For you to consider.. Is there a link between clutch size, and EPF and is this impacted upon by abiotic or biotic selection factors? How might clutch size may be related to the need for EPF in temperate species? If you lay a dozen eggs, why might it be advantageous to engage in EPF? If you lay one egg why might you remain with your social mate and avoid EPF?

42 Temperate species breeding and food availability Lack s Food Availability Hypothesis 1954 Birds should breed when the most food is available for raising young Winter moth caterpillar (Operophtera brumata) Great tit (Parus major) Van Noordwijket al.,journal of Animal Ecology, Vol. 64, No. 4 (Jul., 1995), pp

43 Food availability hypothesis in a temperate species Parus major Highest energy demand for birds starts ~30 days after laying of the first egg (9 days of egg laying+ 14 days of incubation, 7 days of nestling growth) 30 days post-1 st egg must coincide with peak caterpillar availability Birds will be too late in warm years - caterpillars develop quickly Too early in cold years - caterpillar development - slow Once clutch is laid and incubation started, birds can do nothing to match development of eggs with caterpillars (May be able to mitigate effects before incubation)

44 Biotic selection in temperate zone abiotic impact of climate Wrong timing index = (caterpillar half-fall date) minus (1st egg date) minus (30) Correlates with mean temperature 30 days after laying when energy demand peaks Wrong timing not explained by the seasonal increase in temperature. Wrong timing index Birds laid late/caterpillars dropped early Birds laid early/caterpillars dropped late Compatible with birds being less flexible in laying than caterpillars in developing Reproductive success not measured Mean temperature 30 days after laying C Van Noordwijket al.,journal of Animal Ecology, Vol. 64, No. 4 (Jul., 1995), pp

45 Food availability and breeding inconsistencies? %Nesting D J F M A M J J A S O N Flower abundance Long-tailed Hermit (Phaethornis Superciliosus) %Breeding ? D J F M A M J J A S O N? Fruit abundance White-collared Manakin (Manacus candei) %Clutches ? D J F M A M J J A S O N Arthropod abundance Tropical House Wren (Troglodytes aedon)

46 Food availability and individual fitness La Selva, Costa Rica insects appear to be most plentiful at La Selva in February-April which is when Manakin breeding activity starts. %Breeding Arthropods D J F M A M J J A S O N Fruit abundance White-collared Manakin (Manacus candei) Fruit is low in protein, nestlings are dependent on high-protein insect food. Breeding therefore coincides with peak arthropod abundance, not adult food availability. Levey D (1988) Ecological Monographs, Vol. 58, a

47 Food availability and individual fitness Tropical species lay low numbers of eggs Vacant breeding territories are scarce %Clutches Dispersal D J F M A M J J A S O N Arthropod abundance Tropical House Wren (Troglodytes aedon) Reproductive success limited by survival and dispersal of fledglings rather than ability to feed a large brood of nestlings Fledglings remain on parents territory for several months Post-juvenile moult

48 Post-Juvenile moult A complete moult requires synthesis of one-quarter of total body protein in the form of feathers and other epidermal structures High demand on energy and nutrients, especially protein

49 Food availability, moult and dispersal Post-juvenile and adult moult High demand on energy and nutrients, especially protein %Nesting Moult Arthropod abundance Tropical House Wren (Troglodytes aedon) 0 D J F M A M J J A S O N Reproduction timed to facilitate post-breeding activities, not activities associated with nesting itself Young B, (1994) The Condor 96:

50 Reproductive success controlled by genetic or environmental factors? Seychelles Warbler (Acrocephalus sechellensis) Lays a single egg per nesting attempt Endemic of the Seychelles islands No predators of adults Very stable territories Cousin Cousine Aride Praslin Birds All pairs on Cousin studied for 2yr pre translocation Pairs translocated to Aride and Cousine Controls left on Cousin Food abundance Aride x Cousine x Cousin + 1x Seasonal pattern identical Komdeur JA (1996) J. Biol. Rhythms 11:

51 Reproductive success controlled by genetic or environmental factors? Territories with nesting attempts following translocation Aride 6.1x (Food 3.4x) Cousine 2.9x (Food 1.8x) Cousin 1x (Food 1x) All other variables shown not to be significant Cousin % territories with nesting attempts Aride Cousine Komdeur JA (1996) J. Biol. Rhythms 11:

52 Link between reproductive success and food abundance Seychelles Warblers (Acrocephalus sechellensis) Highest reproductive success when nests begun two months before peak food abundance (on Cousin) Differences [in] reproductive timing and success by warblers on the islands are due not to genetic differences but entirely to environmental conditions. Cousin Aride Clutches per nest Building attempt (%) Month Komdeur JA (1996) J. Biol. Rhythms 11: Month

53 Summary

54 Picture credits Michael Woodruff NW_Ecuador-8 Hector Bottai _Dusky_Antbird_%28male%29.JPG Greg Gilbert Garita%2C_Alajuela%2C_Costa_Rica-8.jpg Dario Sanches Calibas Reynaldo Dario Sanches Vale_do_Ribeira,_Registro,_Sao_Paulo,_Brazil-8_(1).jpg Dario Sanches TICO_(Zonotrichia_capensis_)_( ).jpg Janet and Phil Aat Bender Robert Gallardo Waldemar Reczydło Anne Tanne Patrick Yates Terjee Kolaas Magnus Manske