Energetics and Social Behavior Lecture FW 469569 Energy

Energetics and Social Behavior Lecture FW 469/569

Energy Flow through Individual Animals Boyd, Bowen, and Iverson (2010)

Energy Intake Boyd, Bowen, and Iverson (2010)

Energy Storage and Conversion Boyd, Bowen, and Iverson (2010)

Energy Use Boyd, Bowen, and Iverson (2010) Basal Metabolic Rate (BMR)

Basal Metabolic Rate (BMR) • Minimal rate of metabolism of a thermoregulating endotherm Thermoregulating: maintaining constant body temperature Endotherm: heat drawn from internal processes, not the environment Post-absorbtive: not digesting a meal Within Thermal Neutral Zone: at temperature that doesn’t require additional processes to thermoregulate • Resting: inactive but awake, not experiencing physiological stress • Mature: not growing, not reproducing • Units: Energy/time, e. g. , k. J/h, kcal/h • • • Complications: • Circadian and annual cycles in metabolic activity

BMR versus Body Mass in Terrestrial Mammals 300 Oxygen consumption (liter O 2 h-1) 250 200 150 100 50 0 0. 001 0. 1 1 10 Body mass (kg) 10000 After Schmidt-Nielsen 1997

Mass Specific BMR 8 Mass specific oxygen consumption (liter O 2 kg-1 h-1) 7 6 5 4 3 2 1 0 0. 001 0. 1 1 10 Body mass (kg) 10000 After Schmidt-Nielsen 1997

Allometry • The systematic change in metabolic rate with increasing body size • Y = a ∙ mb • Thermoregulatory costs strongly related to surface area • For a sphere: • Surface area: A = 4 π r 2 • Volume: V = 4/3 π r 3 • Surface area / Volume: A/V α r 2/3 so b = 0. 67 • Empirical estimates: b ≈ 0. 75

Allometric Relationship of Metabolic Rate to Body Mass Schmidt-Nielsen 1997

Allometry of BMR Across Seabird Taxa 10000 BMR (k. J d-1) 1000 Sphenisciformes Procellariiformes Pelicaniformes Charadriiformes 100 10 10 1000 Body mass (g) 100000 After Ellis and Gabrielsen 2002

Energy Expenditures Above Basal Rate • Thermoregulation • Modes of heat loss: • Radiation • Convection • Conduction • Water is 24 x more conductive than air • Evaporation • Useful to avoid overheating • Regulating factors: • Surface area • Insulation quality Michael L. Baird

Energy Expenditures Above Basal Rate • Locomotion Schmidt-Nielsen 1972

Energy Expenditures Above Basal Rate • Growth and Development • Energy content of new tissue • Biosynthesis cost (heat lost) • Heat Increment of Feeding • Heat loss associated with digestion • Other activities • Grooming, social interactions, breeding, foraging, molt, etc.

Daily Energy Expenditure (DEE) • Energy required under free-roaming conditions and normal behavior • Incorporates BMR and • • Activity Thermoregulation Feeding All other energy expenditures • Sometimes expressed as Field Metabolic Rate (FMR)

Allometry of FMR Across Seabird Taxa BMR (k. J d-1) 10000 Sphenisciformes Procellariiformes 1000 Pelicaniformes Charadriiformes 100 1000 Body mass (g) 100000 After Ellis and Gabrielsen 2002

Allometry of FMR • Flight Style • Environment All. About. Birds. com Birt-Friesen 1989

Residuals from Allometric Fit are Informative Field Metabolic Rate (k. J/day). 10000 1000 Double-crested Cormorant • Flapping flight • Wetable plumage • Large clutch size • Measured FMR > Prediction 100 1000 Body Mass (g) 100000

Using Energetics to Quantify Community Trophic Structure • Organismal energy flow models useful to estimate prey consumption by predator populations, colonies • With general data on multiple predators and prey, can evaluate energy flow through entire communities • Can feed into ecosystem management of marine systems Roby et al. 2003

Data Analysis Exercise: Quantify Energy Flow into Oregon Seabird Community • Use a bioenergetic model to estimate prey consumption by major piscivorous seabird species in Oregon’s marine environment • Use published data where available; expert opinion where not • Use conventionally-structured organismal bioenergetics model extended to population and/or species • Incorporate prey quality (energy density, size) Field et al. 2006

Community Energetics Lab Techniques • Macro-enabled Excel File: “FW 469 569 Prey Consumption Calculator. v 1. xlsm” • Embedded Visual Basic code to calculate prey consumption base on data entered on “Input Data” worksheet • Most data available on additional worksheets in file for 4 prominent Oregon seabird species • Run macro, results loaded onto “Output” worksheet • Vary inputs to assess effects of uncertainty

Oregon Coast Picivorous Seabird Community: • Sooty Shearwaters: summer migrants Jen Zamon, NOAA Glenn Bartley/VIREO

Oregon Coast Picivorous Seabird Community: • Common Murres: breeders and partial migrants Roy Lowe USFWS

Oregon Coast Picivorous Seabird Community: • Leach’s Storm Petrels: breeders and off-shore migrants Roy Lowe USFWS C. Schlawe USFWS

Oregon Coast Picivorous Seabird Community: • Brandt’s, Double-crested, and Pelagic Cormorants Roy Lowe USFWS All About Birds Jan Arendtsz/Flickr Creative Commons Rick & Nora Bowers/VIREO

Lab 2 Part 2 Community Energetics: Objectives • Describe how individual energetic requirements of marine species along with their unique foraging ecology (i. e. diet) combine to determine community-level energy flow. • Evaluate the importance of prey quality (i. e. energy density) in determining total biomass or numbers of prey consumed • Practice using physiological models to address applied questions in marine ecology. • GRAD STUDENTS: • Add one or more species (bird or mammal, but similar trophic level) to the community-level energetics analysis in this lab exercise (not Caspian terns) • Develop plausible input parameters through allometry and/or literature research

Lab 2 Part 1: MAMU Time-Energy Budgets • How far from the coast can MAMU nest? • What prey quality is needed to support nesting? • What range is possible? • Could MAMU increase how often they feed chicks? Brian Woodbridge

Why study social behavior? • Social structure of a population captures the nuances of individual differences in social behavior but also efficiently summarizes the actions of the individuals and their relationships with each other. (Whitehead 2008) • Social structure is a key determinant of population biology, influencing fitness, gene flow and spatial pattern and scale (Wilson 1975) • “Qualities of Sociality” group size, demographic distributions, cohesiveness, amount and pattern of connectedness in communication, permeability of movement between social groups, degree to which the population contains distinct social units, differentiation of roles, integration of behavior, information flow and fraction of time devoted to social behavior. (Wilson 1975) • Conservation framework: Individual search strategies can be costly, time consuming and risky. Social sources of information circumvent these costs by enabling individuals to exploit the knowledge of experienced conspecifics. (Schakner et al 2017)

Analyzing social behavior • Ethology – accurately describing and modeling social behavior. Bottom-up approach to animal behavior: linked to psychology, uses description, then examines immediate causation, development, function and/or evolution (Hinde 1982) • Behavioral ecology - (Darwinian evolutionary biology) function of behavior and how a particular behavioral pattern influences survival and reproductive success (Krebs and Davies 1991)

Sea Lion behavior patterns Behavior of males changes both seasonally and instantaneously • Adult males have a size related dominance pattern (with specific behavioral repertoire) outside the breeding season • Reproductive season adult males maintain dominance relations with juvenile males and do not use the behavioral repertoire reserved for territorial defense • Adult males are territorial only toward other adult males during the breeding season AND only on traditional sites

Coding behavior patterns and analyzing coding data • How often individuals interacted than would be expected by random combinations (Poission distribution) Boundary display Fighting behavior

Male sea lion behaviors found in adults are also found in pups Boundary display Open mouth submission Fighting behavior Gentry 1974

Social analysis • Histograms, association indices or principal coordinate analysis • Testing for preferred/avoided companions • Testing for hierarchy establishment • Network analysis

Association indices 9 female sperm whales Whales were associated if they dived within 5 minutes of each other Data (sometimes) becomes more useful if it can be divided into age/sex classes

Network analysis: Social network describes island associated communities Kaua’i/Ni’ihau Island of Hawai’i and Lana’i O’ahu Mahaffy et al 2013

Inter-island movements Kaua’i individuals encountered off Hawai’i 2 individuals from Kaua’i observed together off Hawai’i. Ni’ihau Kaua’i (Baird et al. 2008) The two did not associate with Hawai’i island individuals O’ahu Molokai Lana’i Maui Island of Hawai’i Albertson et al 2013

Combining association indices with dyads and other data collections Looking at group composition (relatedness) and how often groups associate • Rough-toothed dolphins – weak matrileality (groups not composed of same haplotypes), but stable social structure (same group members are associated more often than expected by chance) • Humpback whales - Cooperative foraging groups are not related, but are composed of the same individuals year after year

Are groups more matrilineal than expected by chance? Random associations (many) Haplotype frequencies in region equal to that found in associations of groups (a few) Strict maternal structure Number of mt. DNA haplotypes in a group (1)

Matrilineality in rough-toothed dolphin groups Kauai: 15 groups Hawaii: 11 groups Oahu: 4 groups Moorea: 14 groups Raiatea: 2 groups Mass strandings Eastern Tropical Pacific: 3 groups Atlantic: 3 groups Total sample size, n = 214 60% of groups were composed of more than one haplotype

Dyad combinations determine if groups are more matrilineal than expected by chance Sampled group size n=4 haplotypes =2 possible dyads=6 But only 2 dyad combinations are from the same haplotype Observed haplotype identity 2 out of 6 = 2/6 = 0. 33

Are groups more matrilineal than expected by chance? Repeat 1, 000 for each region, calculate proportion of same haplotype dyads Sample regional mt. DNA haplotypes frequencies Hawaii Mean expected 0. 10 0. 15 0. 20 0. 25 Proportion of same haplotype dyads 0. 30

Rough-toothed dolphins show weak matrilineality 46% of groups Weak were more matrilineal than expected by chance (many) (few) Number of mt. DNA haplotypes in groups (1)

Chimpanzees of the sea? v. Weakly matrilineal v. Stable social structure

Useful software for analyzing association data

Important to consider sampling when testing association data • How do you define a “group” (dolphins within a certain distance of each other) • What percentage of animals do you sample in a group? (start with instantaneous scan and then focal follows of several animals) • How do you define association? (sea lions were considered associated if they occupied the same dock or jetty area; whales were associated if they dived within 5 minutes of each other) • How do you define a behavior? (start with instantaneous scan and focal follow to evaluate which is most common and most useful)

Lab 2 Part 3 Social Behavior Objective - evaluate important behaviors in male sea lion social structure • In a table, provide codes for most common (2 -3) behaviors (label behavior, description, age class participation) • Graph frequency observed of each behavior • Describe how the graph can be used to evaluate important behaviors in male sea lion social structure • Grad students: Write a research question about California sea lions that could be answered using a social network analysis.

Questions? FW 469/569