Behavioural ecology Flashcards

Question

Bias sex ratio in collard flycatchers

Answer 1

Females heterogametic - WZ Can potentially control sex ratios of offspring Females mated to males with larger forehead patches (more attractive) have ratio biases towards sons Experimentally increase patch size- female has more sons

Answer 2

Double mating experiments 2nd male gets 80% parentage (displaces first males sperm from storage sacs) therefore mate guarding

Answer 3

Used genetic markers to look at sperm storage when females doubly mated with a different combos of nonsib and sib when both males sib- inbreeding depression when at least one nonsib, female bias in favour of choosing sperm of nonsib (less likely to store sib sperm)

Answer 4

females prefer males with longer eyespan selection lines in lab for 13 generations- select only for eyespan female preference coevolved with it clear genetic correlation

Answer 5

penis driven by lymphatic system, inflates inside of females and corkscrew shaped female reproductive tract also coiled spirals are in different direction to make it harder to males to inseminate increasing phallus length drives greater number of female coils and an increased number of pouches(dead ends) in reproductive tract Brennan et al., 2007

Answer 6

Kolm et al., 2012 males have long extensions to operculum with blob on the end to attract females blob shape reflects food preferences in females eg. forest pond, ant shaped eg. open pond, beetle shapes higher proportion of ants in diet, greater shape similarity of ants and lure females fed ants in lab direct more bites at male ant shaped lures than those fed other things potentially too costly for females to resist lures because could compromise foraging success

Answer 7

female preference for sword tails stronger in species that lack swords suggests that males evolved swords to exploit pre-existing female preference but as they evolved, female preference decreased

Answer 8

loudest shouts best at attracting females small males can't shout loud enough =satellites, sit near calling male and try and get female when she arrives usually get ~50% of matings therefore, should decide to parasitise neighbour if calling 2x as loud as you 90% of decisions as predicted by this

Answer 9

Onthophagus large males with horns small males look like females which phenotype depends flexible developmental programme- depends on size and how much food as larva horned males guard nests with female inside small males dig into nest in attempt to sneak matings switch point at ~5mm pronotum width laboratory experiments show that threshold size for switch can be changed via selection select smaller sized, decrease threshold and vice versa

Answer 10

2 male morphs- large forceps (better at fighting) and small forceps UK mainland, almost all small forceps UK islands, greater proportions with big forceps allometric sigmoidal relationship-switch at a certain body size island switch size is lower due to higher population density- greater payoff for fighting even at smaller size

Answer 11

2 strategies: mouth twisted left, or twisted right 2 morphs present in 7 different species ESS= 50:50 proportion, because each host has two sides

Answer 12

10-13% population left handed left-handed, very slightly lower life expectancy left handed people have slight advantage in fights because rarer opponent harder to defend against eg. higher proportion of boxing champions left handed than predicted by chance

Answer 13

Fighting males with big jaws dispersing males with wings If all males fighters, good to be disperser because could mate with all females that emerge from fig that didn't include fighter wasp & vice versa

Answer 14

Philomachus pugnax dark ruffs- territorial, attract females white ruffs- sneak matings at edges of others' territories female mimics (only recently discovered) Don't know equilibrium between because disperse, can't tell mating success

Answer 15

males: orange throats- guard many females yellow throats- sneaks, look like females blue throats- guard one female oranges to best when high blue yellow do best when high orange blue do best when high yellow no equilibrium in wild, cycle between differing proportions can't change tactics in this eg. because morphological specialisations

Answer 16

nest in woodpecker holes in trees, chicks jump out when hatch some females parasitise nest and lay eggs in someone else's experimental tracking of chicks- can tell whose eggs are whose because characteristic shape Tag chick feet just prior to hatching to track Reproductive success tracking suggests that represents strategy not a tactic, because equilibrium value of 17% of eggs are parasitic (no sig. diff. from 20%, as predicted by equilibrium modelling) But females don't stick to same role throughout life- there may be a genetic propensity for one over another- does it depend on other individuals in pop? Potentially LRS of parasite better- don't have to care or guard box

Answer 17

Prunella modularis have different mating systems: Polygyny, female RS= 3.8 per season, male= 7.6 Monogamy, female=5.0, male=5.0 Polyandry, female= 6.8, alpha male= 4.1, beta male= 2.7 Polyandry best for females, will solicit matings with any new males they encounter Monogamy better for females than polygyny, fights between females for full territory Polygyny best for males, try to stop females fighting, and in polyandry, alpha consistently trying to chase off beta in polyandry Often see polyandry in dense vegetation

Answer 18

females wants equal chances of paternity from her chicks- 50:50, increases amount of paternal care chicks receive Normally, female tries to increase mating with beta to reach ratio If alpha male experimentally removed for period of time, will mate more with him when he's returned Males can't distinguish own chicks, therefore use mating share as indirect cue of paternity If beta mates, he will care Even if experimentally reduced brood to 1 No paternity markers- chicks and mothers don't want them, because they would reduce amount of care given (genetic suppression by both chicks and mothers?)

Answer 19

Houston-Davies model- "sealed bids"- evo game where efforts fixed and optimal investment of each carer reached over evo time Individual's best effort in reaction to partners effort decreases with shallow gradient and partner puts in more effort 4 different outcomes: ESS- stable equilibrium between two *optimal* ESS- male sole carer ESS- female sole carer Unstable - overcompensation of one, depending on starting effort eg. starlings together, male provides 12 feeds per hour, female 13 weigh down female, she provides only 7, male provides 14 weigh down male, he provides 7, female provides 15 Incomplete compensation- are they responding to partner's reduced effort or indirectly via increase in chick begging?

Answer 20

broadcast extra begging calls from speaker at nest box ensuring that playback didn't affect actual chick begging increased calls = increased parental effort playback to one of the two, ensuring not heard by other partner, both increase effort (more feeds per hour) both chick begging, and what other partner is doing affects care

Answer 21

males have iridescent feathers 76% young sired by EPCs social monogamy, with previous offspring as helpers Extra males bring yellow petals to female when in fertile period Females go to territories of other males in early morning to copulate 5% of the males in a pop get 50% of matings 1 "super stud" has sired 450 out of 1800 chicks genetically profiled females with many helpers- more EPCs, they will care for young even if male detects

Answer 22

Tribonyx mortierii Dominant male allows subordinate bother to copulate- more offspring can be raised (9.6 compared to 6.6 with a pair) with three parents helping Whereas is subordinate alone LRS = 0 Because brothers, r=1/5, greater benefit to alpha male to keep brother because larger genetic profit from altruism than selfishness

Answer 23

Can raise 5.2 children with polyandry compared to 3.75 with monogamy Hamilton's rule, dominant brother should share wife if 3/4 N2 > N1 Here 3.9 > 3.75 BUT very close Under harsh conditions, may pay to have younger brother around, help increase farming productivity, enables more children, might offset paternity loss Is female in control- for her advantageous as long as N2>N1 Importance of culture eg. Buddhists permit polyandry, but Hindus and Muslims don't

Answer 24

Young stay at home and help parents raise future broods

Answer 25

Brood sizes v small (1 or 2) Live only on Cousin Island Pair of breeders raise 0.8 young to dependence each year, with a helper, 1.2 Stay because full habitat on island (ecological constraint) and indirect fitness gain Translocation of birds to nearby Aride Island within weeks, birds breed and pops expand, no helping As habitat begins to fill up, start to have helpers again DNA profiles show that 40% chicks are sired by EPCs and female helpers lay some eggs female helpers only care if their mother is present - she produces 85% of the chicks in a nest, therefore if she is present, helpers probably related to chicks (0.85x0.85=0.72) But only 60% of chicks are produced by primary male, so if just he is present, chances of being related to chicks is 0.6x0.6=0.36

Answer 26

Live in clans of 30 plus- breeding parents plus their sons and immigrant wives at a point in the season, split into pairs to breed magpie predators can destroy nests, if no time to rebuild, will go to help other nearby pairs (immigrant females won't help though, because no kin) 16/17 choices, sons chose close kin to help over unrelated individuals Recognise by signature calls of family (learnt from parents/carers) broadcast churr calls, approach kin calls, and reply aggressively to non-kin

Answer 27

Hetaerina Americana Grether & Grey (1996) male bright red spot at base of wing=sexual signal But makes easier for prey to detect and avoid it (conspicuous) Catch and manipulate females, though with red spots added had lower relative gain rate, even when controlled for age and size

Answer 28

Fleishman, 1992 Anolis auratus territorial males defend with head bobbing display, bright dulap pulses in and out visually striking and attracts other males' attentions to maximise detectability, bob at high frequency different to that of background tree/leaf movement

Answer 29

Mimics background movement eg. branch in wind, similar f to background, harder for prey to detect

Answer 30

2 species in different Southern US states 1 zone found together mate attraction by calling G. olivacea higher f than G. caroliensis in sympatric overlap zone, diff in f exaggerated to minimise confusion, G. caroliensis lowers call f

Answer 31

``` Neotropical suboscines (innate, not complex songs) Sympatric species tend to have greater character displacement in song that allopatric species selection against inappropriate mating ```

Answer 32

Different behaviours are as maximally distinct as possible- hostile vs affectionate, change in tail/ears/back/face etc, greatest contrast

Answer 33

Most species repeat the same song multiple times Three species (Short and long-billed marsh wren and rock wren) don't repeat consistently, switch rapidly between song types, larger repertoires These three live in habitats where there aren't many other bird species and not much noise- where species recognition poses few problems

Answer 34

Long tail, jumping display, courtship bower

Answer 35

Spontaneous assertion display grabs attention of receiver with high amplitude and frequency movements (little information, no pattern), before moving onto informative display Ord & Stamps (2008) robot programmed to have: -no display -alert component and typical display or -alert component and novel display recorded receiver responses: -in bright light and at close distance, no difference in time it takes for robot to get attention -in dim light, takes longer when there was no alert Plus, lizards in dimmer conditions and environments with more background noise, tend to increase rate of alert components

Answer 36

8 species in forests of Kashmir small and green vary in amounts of ornamentation (bright plumage patches) across species patches play role in male individual territory defence high patch size and number means larger territory size (number of patches and the total reflectance at the feather tip is lower in higher light areas, when controlling for phylogenetic effects)

Answer 37

Male colour patches have different levels of conspicuousness in different levels of light to predators and other guppies eg. prawns don't detect orange well, therefore guppies often orange when prawns around, but less orange if fish is main predator

Answer 38

Davies & Kilner , 1998 gape colour in passerine chicks varies with light at nest (red in reed buntings, yellow in robins) Darker nest= wider mouth flanges that are whiter and less densely coloured relative to mouth, but mouth didn't change colour (increasing contrast, more detectable)

Answer 39

males golden colour with black crown and back, display to olive females creates display court, clears away leaves, helps show off plumage by increasing contrast with forest floor Uy & Endler, 2004, colour and brightness contrast of golden patches greater against cleared court than adjacent leaf litter

Answer 40

look deep blue-black but reflect UV strongly, therefore brightly coloured to conspecifics (mammals can't see) evolved because of habitat- high altitude forested streams and gorges that receive high levels of skylight rich in UV wavelengths, but nest and forage on ground so pays to be cryptic to mammalian predators

Answer 41

female workers recruit others to food resource figure of 8 and waggles abdomen in centre on vertical wax plane angle of centre part of dance to vertical = angle between sun and direction of food source from hide duration and intensity of waggle informs on distance and resource value fully cooperative behaviour

Answer 42

pheromonal lure that mimics female sex attractant of moths, spiders catch and eat male moths that arrive

Answer 43

Maevia indemens 2 male morphs: dark bodied and striped have different courtship and display at different differences dark morphs raises itself and front legs up striped flattens self females don't seem to prefer either morph, approach first male to move

Answer 44

``` Uca beebei (Christy, 1988) in a number of species, males build vertical pillar next to burrow exploits bias for crabs to detect things that break line of horizon(hide from predators in) eyes have narrow vertical pseudopupils where ommatidia densely packed = superior visual activity in vertical plane ``` experimentally, vertical rectangular shapes more consistently elicit spontaneous approach than horizontal rectangles of equal area Christy et al., 2003 arena with male burrows, some pillars are removed female approaches pillars in mating context females also run towards pillars when scared by predator (sensory bias evolved in non mating context) this bias also seen in predator situation for species that don't use pillars (mating preference before trait evolution)

Answer 45

Rowe et al., 1996 offered rtificially coloured food, one colour made distasteful when pyrazine odour present, even at beginning of experiment, do better when learning to avoid brightly coloured than duller coloured biased against bright colours even when not distasteful

Answer 46

Neumani papillator Proctor, 1991&92 'courtship trembling' ambush predators, grab prey items when detect water vibrations males approach waiting females and vibrate water so she grabs him and sees potential mate females deprived of food more likely to orient to and clutch vibrating males

Answer 47

Stalhandske, 2002 Pisaura mirabilis male provides silk wrapped prey item to mate gift mimic female egg sac, which she has propensity to protect and carry around males take advantage of this instinct? Experiment: paint courtship gifts to be as white as egg sacs (normally darker than them), latency of female to grab it reduced the brighter it gets

Answer 48

Engystomops pustulosus Ryan & Rand, 1993 calls have chucks and whines individual males have own calls different related species have different versions of call- all have basic 'whine' with specific additions splice together different call elements of different species' calls females have preference for different elements different species take advantage of ancestral female preference (perhaps one broad ranging bias?)

Answer 49

provisioning of food gift eg. spermatothylax (elaboration of spermatophore that contains sperm), eating spermatothylax by female increases time the ampulla is attached to her and transferring sperm therefore bigger gift = more sperm suggested that preference for gifts evolved in common ancestor predating evo of gift giving-- not all species crickets have gifts but some appear to have preference experiment: artificially supply spermatophylax for specis who don't normally have when present, spend more time eating and more time before she removes sperm ampulla, and less sperm left it in after

Answer 50

Basolo 1990, 1995 investigating female preference for sword on males in related fish female choose males with longer tails in green swordtail attach sword to southern platyfish (closely related but no tails), female preferred therefore preference must have evolved first in common ancestor but new molecular data suggest with revised phylogeny that either evolved together and tail first (Mayer et al., 1994) Therefore unsure

Answer 51

Mill et al., 2002 males have red plumage, determined by carotenoids and therefore indicating information about foraging success and nutritional condition (most animals cannot make carotenoids) males in population vary in colour intensity, poor quality males have less colour mate choice experiment, females prefer most intense colours- conveys honest information, gives fitness benefits to female brighter red males moult earlier and grow feather more quickly, and have higher carotenoid concentration in gut

Answer 52

FitzGibbon & Fanshawe, 1988 vertical jumping as a pursuit deterrent predator approach => stotting rather than running straight away predators choose gazelle with worst stot more vigorous stotting = better condition and harder to run down (reliable info about stamina and endurance)

Answer 53

Magpie- Pica pica Redondo & Castro, 1992 high pitched calling and brightly coloured gape in some birds amount of begging in magpie = honest signal of nutritional need greater time since fed to satiation, greater gape area and rate of calling (higher to older chicks) parental visit and feeding rates both increase when more chicks added or playback of more chicks added honest because cost of begging But McCarty, 1996 "the energetic cost of begging is surprisingly low" But measured energetic costs and not costs to fitness- need to actually know how begging costs affect growth

Answer 54

Mappes et al, 1996 Production cost attract females by drumming abdomen on ground (tiring) manipulate intensity with which males drum by exposing them to many for few females encourage to drum more = lose weight more quickly and lower fraction survive, therefore clear fitness cost males of low weight and low natural drumming intensity don't survive long in high drumming treatment

Answer 55

Hirundo rustica Maintenance cost male streamers for mate choice but aerodynamic cost measured performance in a flight maze after tail manipulation tail has costs- typical streamers exceed aerodynamically optimal length by approx. 12mm

Answer 56

Chlamydera maculata Madden (2002) corridor of twigs with court infront females prefer solanum berries, therefore males collect many manipulate by adding berries = elicits more aggressive attacks and stealing from other males (can't maintain bower with too many berries) when offered free supply of berries, number of berries they had at start positively correlated with how many they take appears to assess social standing in relation to neighbours and modulates display accordingly

Answer 57

Sergio et al., (2011) territorial decorate nests w eg. plastic waste old and young individuals have few nest ornaments middle aged have the most and more likely to take free offered ornaments nest decoration correlates with trespassing on territory by 'floaters' manipulate nest by adding ornaments, individuals in prime of life can defend off increased attack rate but others cant deal with increased aggression

Answer 58

Utetheisa ornatrix Dussord et al., 1991 male sex attractants during mating, male transfers chemicals n seminal fluid (toxic alkaloids) too female, who incorporates them into eggs (protects) - female wants males that provide lots of chemicals pheromone similar chemical composition to described chemical- therefore = informative of quality of how much alkaloid male can provide selection for reliability (males also want to provide alkaloids to offspring- therefore amount released relative to overall amount available)

Answer 59

Strategic Increased extra pair paternity -> in nest with half sibs selected to be more selfish and misrepresent hunger louder begging for food

Answer 60

Garcia & Ramirez, 2005 terminal yellow band on male tails, looks like worm when moved elicits feeding response in females (exploitation of feeding bias in mating context) more elaborate band decreases feeding response but sexual response remains (because costly and therefore honest signal - males and females with yellow bands suffered more damage confined to area of band when kept in outdoor ponds) Therefore initial evolution was exploitative but has become honest

Answer 61

eg. in reed warbler nest as cuckoo grows, gape increases, but doesn't grow more than overall gape size of al other chicks in nest therefore compensated by begging more intensely for a given level of hunger (Kilner et al., 1999)- have no genetic interest in future RS of adoptive parents Parents attend to begging calls- cuckoo parasitisation sufficiently rare that still pays to attend chick begging

Answer 62

Adams & Caldwell (1990) continues to display threat displays to repel intruders entering burrow, even when just moulted and unable to fight effectively intruders do best to respect signal, because in most cases represents real threat

Answer 63

Stations where larger fish/ turtles come to have parasites cleaned from gills have local client fish with small territories and free ranging fish which have multiple options of cleaning stations they could visit cleaners would prefer to eat the mucus and scales of clients (Grutter and Bshary, 2003) but if take a bite= lose a client, bad for rest of cleaner group more likely to cheat and take a bite out of local fish, because they have no choice of cleaning station Cleaners often work in pairs, number of mechanisms to reduce cheating eg. partner switching, punishment plus threat of reciprocity by client 2 different, unfixed strategies of cleaners: - low interest in small clients and rarely cheat larger clients - small minority cheat large non-predatory clients with 6x higher frequency (Bshary, 2002) cheaters give tactile stimulation with their fins, lowers basal and acute cortisol levels in clients- use to build new relationships with clients, to reconcile after cheating and as a pre-conflict management strategy with predators Observation of a cleaning interaction without conflict attracts new larger clients, so tactile stimulation of smaller fish can falsely entice larger that are then bitten to obtain mucus Injection of fish with cortisol or blocking of their glucocorticoid receptors increases this 'cheater' behaviour (Soares et al., 2014)

Answer 64

semantic communication in nonhuman animals acoustically different alarm calls to different predators each call causes different, seemingly adaptive responses Monkeys on the ground respond to leopard alarms by running into trees, to eagle alarms by looking up, and to snake alarms by looking down Seyfarth et al., 1980 played recordings of alarm calls in absence of predators and observed responses alarm length, amplitude and alarmist's age/sex class had little effect on response quality But, calls similar sounding in aggressive context eg. overlap between female calls given to snakes, eagles and during aggression eg. male barks in leopard and aggressive contexts Price et al., 2015 semantic properties of alarm calls bear little resemblance to human words, but acoustic variation, possibly with additional contextual information, allows listeners to select appropriate responses

Answer 65

field observations of killer whales have documented the existence of group-differentiated vocal dialects that are often referred to as traditions or cultures and are hypothesized to be acquired non-genetically Abramson et al., 2018 subjects were able to make recognizable copies of all familiar and novel conspecific and human sounds tested and did so relatively quickly suggests vocal variants observed in natural populations of this species can be socially learned by imitation

Answer 66

All three ‘peaks’ of large brain size evolution in mammals (odontocetes (esp. dolphins and sperm whales), humans and elephants) shared a common selective environment: extreme mutual dependence based on external threats from predators or conspecific groups In this context, social competition, and consequently selection for greater cognitive abilities and large brain size, was intense.

Answer 67

Form of teaching (fulfils all requirements) Tandem leaders know food location, tandem followers naïve leader only continued the tandem run when frequently tapped on her legs and abdomen by followers antenna The tandem leader therefore modifies its behaviour in the presence of the follower Imposes cost on leader because can move 4 times faster without follower- tandems slowed by frequent pauses where follower loops round, probably in search of landmarks (speed of run higher in presence of conspicuous landmarks than when absent) Followers found food more quickly when tandem running than when foraging alone Bidirectional feedback between the leader and follower is evident from their patterns of acceleration and deceleration as a function of the strength of the stimuli they present to one another- when gap between them too large, leader slows and follower speeds up Lessons learned by followers transferred when they become leaders for others- therefore propagates time-saving knowledge between individuals Bidirectional feedback distinguishes it from broadcasting in other ant species (eg. pheromone trails)- effective in big groups teaching works better in smaller societies, where info is valuable and easily lost (big brain is not a prerequisite of teaching)

Answer 68

David Lack- add or remove eggs Parents provide less food per offspring to enlarged broods- reduces offspring mass and survival (survival to 1 year reduced in enlarged clutches) - Smith et al., 1989 Parents act at optimal rate of feeding in each treatment

Answer 69

Larvae raised either in absence or presence of parents Better off when parents present (Eggert et al., 1998) Comparison between biparental care and sum of male and female uniparental care treatments- offspring grew larger and were more likely to survive to adulthood in biparental (first empirical evidence of synergistic effects of biparental cooperation on offspring fitness) - Pilakouta et al., 2018

Answer 70

Nedelec et al., 2017 repeated motorboat-noise playback compared to ambient noise increased defensive acts and reduced feeding and offspring interaction of brood-guarding males noise didn't affect offspring growth, but reduced the likelihood of survival (6/19 nests with motorboat noise suffered complete nest mortality, whilst all with ambient treatment survived)

Answer 71

Females deposit tadpoles in separate pools eg. bromeliads Lay unfertilised trophic eggs for developing tadpoles to feed on- survive better and growth larger with more trophic eggs provided (Dugas et al., 2016) Parental care had costs to both sexes, either through reduced productivity of reduced parental longevity

Answer 72

Isle of Rhum Females that successfully reproduced one year experienced survival and fecundity costs in the next breeding season- but results were partially dependent on population size

Answer 73

Individuals whose clutches had been experimentally removed had reduced oxidative stress and increased weight compared to breeding birds But these costs of reproduction only accrued by individuals breeding in groups with few or no helpers

Answer 74

Froy et al., 2013 Long-term data set to relate measures of parental care to reproductive success (offspring survival and quality, parental age) - care for over half a year per chick Generally improved their parental performance early in life likely due to increasing experience, but then showed decline in performance, likely due to senescence Very last reproductive attempt, invested heavily and achieved high reproductive success

Answer 75

Hoffman et al. (2010) examined relationships between maternal age and activity, mother and infant body condition, interbirth intervals, measures of behavioural investment in offspring, and offspring survival and fitness older mothers had lower body mass indices and were less active, had longer interbirth intervals, and spent more time in contact with infants, but had infants of lower masses and survival rates- mostly consistent with senescence, but some results consistent with predictions from terminal investment hypothesis

Answer 76

Praying mantid- female cannibalises male during mating Brown & Barry (2016) - males and females fed differently radiolabelled crickets and allowed to mate- half pairs progressed to sexual cannibalism, other half stopped Assessed relative allocation of both male-derived somatic materials and ejaculate materials into the eggs and soma of the female- male somatic investment does contribute to production of offspring eggs and reproductive tissues of cannibalistic females contained significantly more male-derived amino acids than those of non-cannibalistic females, and there was an increase in the number of eggs produced subsequent to sexual cannibalism- therefore increases male material investment in offspring In absence of cannibalism, males still 25% of their radiolabelled amino acids via the ejaculate

Answer 77

gametogenesis in two endangered frog species susceptible to Bd fungal pathogen Increased when infected Corroboree frog - infected males have thicker germinal epithelium, and a larger proportion of spermatocytes Whistling tree frog - infected males had more spermatic cell bundles in total, and a larger proportion of spermatozoa bundles and infected females ovaries and oviducts were larger in infected animals, and there were more cells present within the ovaries because mounting immune response costly and can reduce investment in other life history traits eg. reproduction - terminal investment hypothesis predicts will increase reproductive effort when threatened by disease However, if infected animals are increasing reproductive efforts and producing more offspring before succumbing to disease, it is possible that population-level selection for disease resistance will be minimized

Answer 78

Species that live in harsher, northern habitats had lower annual survival and laid larger clutches than species in more benign southern habitats Northern more willing to risk lives at benefit of offspring Where residual reproductive value is low, parental investment into current reproduction should be high and vice versa

Answer 79

males sole carers, females lay and desert Males eat small clutches/eggs at earlier developmental stage - weigh up costs of caring for egg vs benefits Females benefit from males caring, but males eat eggs they don't sufficiently benefit from Females avoid conflict by choosing males that already care for a clutch still at a v early stage

Answer 80

Males sole carers Will eat if doesn't think caring will sufficiently benefit him females appear to test whether a male will provide care by laying a few eggs and returning later to see whether eaten before providing with rest of eggs (Manica, 2010)

Answer 81

Conflict between maternally and paternally inherited genes in the offspring (some genes, only maternal one expressed, others, only paternal) Paternal genes favour taking increased resources from mother whilst maternal genes counteract them eg. insulin-like growth factor IGF2 and its receptor IFG2r IGF2 paternal, increases growth, IGF2r maternal, decreases reception of IGF2

Answer 82

females deposit androgens in eggs which affects offspring begging behaviour and can manipulate males into providing more care males responded more to testosterone mediated begging than females (Noguera et al., 2013) manipulate testosterone in egg, increases begging 2.5x that of unmanipulated

Answer 83

Males can increase RS by attracting second mate when they encounter a large carcass Females don't gain from this, lowers her RS When male attempts to attract another female by lifting abdomen and emitting pheromones (sterzeln behaviour), mate will knock him over to prevent and bite and bully him to stop future attempts (Eggert & Sakaluk, 1995)

Answer 84

Kill current offspring of a female to get females that are still caring for young to enter oestrus again Less likely to see infanticide in seasonal breeders, females can't resume reproduction Coercion strategy to force partner's parental investment away from current and towards future offspring Paternity confusion viable counter strategy for females, reduced likelihood of infanticide (Lukas & Huchard, 2014)

Answer 85

Each parent can increase RS by leaving mate to care and remate Females lay an egg per day and copulates with chosen male each day But timing of desertion crucial- if male leaves too early, mate may not finish clutch or mate with other males but if they wait too long, face female deserting (and then have to choose whether to care or desert) If female leaves before completing clutch males more likely to desert as well, but too late more likely male deserts Each tries to deceive other eg. female covers eggs with nesting material, making male believe clutch at earlier stage and therefore confuse when he should leave

Answer 86

Females increase apparent size of clutch at reduced cost by producing dummy eggs that don't contain yolk and don't hatch- but cheaper to produce while potentially deceiving males into investing into clutches they might otherwise eat

Answer 87

Many religions incorporate rules to ensure paternity eg. the Dogon of Mali traditionally expect women to sleep in special hut when menstruating, advertising their approach of fertile phase to whole community (esp husbands family) Results in lower extra-marital paternity

Answer 88

Negotiate amount of mouth brooding- influenced by male's state males expend less investment when in worse condition Can't feed when brooding Eggs exchanged from female to male once, when she signals for him to take over But if starve male before, female cares more before he takes over (ignores her signals) Other way round doesn't happen- he chooses when to take brood But, even when starved he still always cares for longer overall

Answer 89

No bias for male or female to care more | larger fish more likely to desert because more attractive

Answer 90

Offspring suckling prevents mothers conceiving Both determine rate at which offspring gain access to nipples as offspring get older, rate of milk feeds reduced from ~15 per hour at 4 months to less than 5 at 12 months Result of decrease in offspring attempts and increase in mothers rejections (rises from 40 to 80%) Gomendio et al., 1991 Expect to be in conflict over timing of weaning But strongest determinant of the reduced rate due to reduced number of attempts- therefore maybe not at conflict? Actual fitness correlates of behavioural changes weren't determined

Answer 91

Females wean offspring after two years of lactation But some females give birth to pups in two consecutive years- suckle a yearling and newborn simultaneously Older offspring often aggressive towards younger Female aggressively interferes with elder's attempts to subdue younger- sometimes causing actual injury to elder (Trillmich & Wolf, 2008) Appears mothers actually use aggression towards yearlings to assess how they should optimally distribute investment (yearlings already survived a year so their interests should align more)- if yearlings persist despite aggression= sign of need is honest- in mother's best interests to feed yearling at new-born's expense Actual conflict between mother and newborn But also sibling conflict

Answer 92

Haplodiploid, monogamous, queens equally related to male and female reproductive offspring But worker daughters more closely related to females (young queens) r=0.75 (brothers r=0.25) Queen wants 1:1 ratio, workers want 3:1 Hymenopteran systems, queens lay eggs at 1:1 ratio, but actual sex ratios are closer to 3:1 - workers skew survival of eggs/larvae towards their best interests But in polygamous, workers become equally related to brothers and half sisters (r=0.25)

Answer 93

Species where some colonies produce predominantly male reproductives and others mostly female eg. wood ants- colonies tend to invest more into females when queen singly mated, into males when mating with multiple males Queen lays ratio 1:1 but colony bias it one way or the other

Answer 94

Janzen & Warner, 2009 Mother's fitness (clutch hatching success) maximised at a lower individual egg mass than offspring fitness (individual hatching success) in two out of three species Observed egg masses closer to mother's optimum in these 2, suggesting that offspring have very little scope in influencing egg investment

Answer 95

Life bearing animals balance offspring number and size Offspring provisioned continually may have more power compared to in egg-laying species Soay sheep- mothers benefit from giving birth to twins, which increases their LRS, but twin offspring have lower birth weight- reduces own survival and lifetime fitness (Wilson et al., 2005) But have no data on fat content of mother's milk, suckling frequency, maternal behavioural care - don't have all evidence of parental investment And no data how these affect parental future reproduction

Answer 96

Placental, live bearing fish (matrotrophy) most maternal investment after egg fertilised, embryos may be able to influence amount of investment they receive 2 populations that differ in offspring number vs offspring provisioning: Moore Lake(ML)- more offspring than Wakulla Springs (WS) In gestation phase, ML offspring gain less weight than WS- difference in offspring demand of resources Offspring quality (weight) greatest for WS x WS But female fecundity (offspring #) greatest for WS fathers x ML mothers (suggests offspring in control of offspring provisioning)

Answer 97

mother's influence offspring behaviour in nest by supplying different levels of hormones in the egg Cross foster birds, increased fecundity cost to mothers the greater the mismatch between the begging level of their own brood and cross fostered brood Suggests that in natural conditions, mother's provisioning behaviour and offspring begging levels coadapted (would be difficult to detect conflict here)

Answer 98

tadpoles can have different phenotypes- omnivores that feed predominantly on detritus or carnivores that feed on larger prey (incl. conspecifics) Morphs often facultative- can switch between depending on diet- by supplying certain hormones a carnivorous diet induces and maintains morphological and behavioural changes When large prey (and therefore the hormones) rare, revert to omnivorous Ability of tadpoles to discriminate between kin and non kin varies with degree to which they are cannibalistic- individuals from more carnivorous species less likely to eat kin (S. bombifrons obligate carnivores, S. multiplicate has the two morphs)

Answer 99

First offspring to hatch in each uterus consumes all younger siblings and all unfertilised eggs in uterus- therefore only two end up surviving Offspring seem not to differentiate between full and half sibs- devour all (Chapman et al., 2013)

Answer 100

Some species- gregarious larvae w multiple larvae on one host, others aggressive larvae- fight and kill each other until one remains Comparative study showed that both species are capable of fighting, but gregarious larvae have reduced mobility during early larval stages- therefore if either species encounters another larva, 50% chance of survival- gregarious sp just don't encounter each other Increases number of eggs that become adults per host- but tend to be smaller

Answer 101

siblings fight over which gets to remain in natal territory | Philopatric birds have higher survival than those forced to disperse (Ekman et al., 2002)

Answer 102

Individuals only gain considerable fitness once dominant breeders in their group Dominance typically established via aggressive interactions among potential sibs (Stiver et al., 2006)

Answer 103

When flanked by male mice on either side The testosterone diffusing from male embryos changes development of genetically female fetus towards more masculine traits More aggressive, shorter lives, lower RS

Answer 104

Submissive behaviour increased subordinate offspring's suckling success Subordinates did better as they got older- training effects?

Answer 105

Females typically lay 2 eggs- one hatches 4 days earlier than other When food levels insufficient to sustain both chicks, one dies as direct or indirect consequence of aggression from senior chick Aggression modulated by hunger of senior Prevent from eating for a few days- tape throat- aggression increased as mass decreased Suggests facultative changing of aggression, based on perceived costs of having sibling Being aggressed doesn't seem to affect juniors' behaviour into adulthood If conditions favourable and both survive- both show similar levels aggression as adults

Answer 106

Individuals appear to target aggression towards less related sibs and unrelated conspecifics C. floridinam- females lay up to 2 eggs into moth eggs- usually an unfertilised male egg and a fertilised female egg Eggs then clonally reproduce up to over 1000 embryos each- create a polymorula Some embryos from female egg develop into soldiers (elongate, huge mandibles)- roam caterpillar and kill other embryos Soldiers sterile and die before caterpillar reached fifth instar, when other embryos consume host and disperse Giron et al. (2004)- two artificial parasitism experiments: resource limitation (and thus increased competition) doesn't alter soldier behaviour - both situations soldiers avoided killing members of their own clone But didn't spare brothers, likely because relatedness = 0.25 But killed brothers less than unrelated Otsuki et al., 2019 Mass killing of male clone larvae suggests intersexual conflict between male and female larvae, but mass killing is adaptive to all the killed males as well as the female soldiers that have conducted the killing because the killing increases their indirect fitness by promoting the reproduction of their clone sibs Models indicate that optimal number of surviving males for both male and female larvae is very small but not zero - matches field data, optimal killing rate of approximately 94–98% of the males in a mixed brood Maternal control of the sex ratio during oviposition, which is well known in other hymenopterans, is impossible in this polyembryonic wasp, therefore mass kill is necessary to maximize the fitness of the female killers and male victims, which can be seen as an analogy of programmed cell death in multicellular organisms

Answer 107

Overall amount of begging by a litter influences the amount of food each litter mate receives- more begging, more food Thus overall supply not fixed, begging cooperative act Individual increase of in begging can cause weight loss

Answer 108

Most young raised alone, but sometimes two-chick broods (Bebbington et al., 2017) Being raised with sibling has apparent costs of offspring- fed at lower rate and have lower body mass, negative effects on survival esp for subordinate Birds that survive these adverse conditions had at least similar RS as adults- tendency for those w siblings to live and breed longer

Answer 109

Mammals- how litter size and nipple number affect competition between siblings and influence growth patterns In species with more offspring per nipple, prenatal growth higher in carnivores and insectivores Postnatal growth showed reverse effect- females in full control of milk supply

Answer 110

Bell et al., 2004 2 Californian populations of stickleback examined the developmental stability of single behaviours (activity in an unfamiliar environment, aggressive behaviour and boldness under predation risk) and correlations between these behaviours Single behaviours unstable through time but some correlations between behaviours remained stable eg. boldness and aggression in one pop but not other Correlation emerged in those surviving trout predator (selection favours) Certain correlations apparent at some developmental stages but not others Therefore behavioural syndromes don't necessarily limit behavioural plasticity & ecological and developmental circumstances might favour different suites of traits

Answer 111

When given choice between joining shoal of bold or shoal of shy fish, bold and shy fish both prefer bold shoal But when hungry, shy fish more likely to go to shy (no hunger difference in bold fish when hungry)

Answer 112

eg. sticklebacks Rands et al., 2003 State-dependent, dynamic game model of foraging by a pair- each chose whether it should rest or forage at a given moment (to maximise own survival) When advantage to foraging together, equilibrium behaviour of both becomes synchronised, which causes differences in the energetic reserves of the two individuals- causes them to take on different roles Individual with lowest reserves = pace-maker, determines when pair should forage according to own needs Group coordination needs no knowledge of the state of others- use own state If low resources, go forage If relatively high resources and partner forages, join, because (depending on resource abundance) will be advantageous to you (usually positive feedback) Only if really high resources don't need to forage

Answer 113

Wolf et al., 2007 Trade off between current and future reproduction => polymorphic pops where some individuals put more emphasis on future reproduction that others (may be related to resource levels) Individuals with high future expectations have much to lose will be more risk-averse than those with low expectations Causes different behavioural types, behavioural consistency and correlations of behaviours across contexts. Explains common finding that explorative behaviour and risk taking traits such as aggression and boldness are common characteristics of animal personality

Answer 114

Bolder (less time to reemerge from shell after scared) individuals have smaller spermatophores than shy Fewer resources, less to lose, less careful Consistent at 2 different locations

Answer 115

Johnstone & Manica, 2011 Leadership is solution for maintaining group coordination when members of a group differ in preferred course of action intrinsic leadership = lambda, determines probability of choosing own preference over following another individual cost, k, determines cost of following Repeated coordination games Benefits maximised when individual imposes its choice on a group and is followed selection leads to evolutionary branching and diversification in intrinsic leadership among the members of a population even in the absence of any variation in state changes in k don't affect lambda much unless extreme in groups of 2, multiple equilibria exist in groups of more than 2, 2 equilibria appear- leaders and followers evolve (dimorphism) Personality types maintained by frequency-dependent selection (leaders gain by imposing their preferences on followers, but fail to coordinate effectively when interacting with other leaders) Fraction of intrinsic leaders in group increases with decreasing conflict - low conflict, many intrinsic followers and coordination achieved by random convergence on one leaders option High conflict- most intrinsic leaders and coordination can break down

Answer 116

eg. changes in aggressiveness with metabolic capacity Wold & McNamara, 2012 Hawk-Dove game, where metabolism affects aggressiveness: When cost of high metabolism is high, all have low metabolism When costs aren't too high, coexistence of some individuals with high and some with low metabolism High aggression can give gains needed to afford high metabolism- but only if costs limited and chances of winning in game are high ie. likely to come up against low aggression individual =Mixed ESS Can lead to behavioural syndromes Therefore, coevolution of behavioural and physiological traits also gives rise to adaptive physiological differences that are systematically associated with behavioural differences

Answer 117

Rey et al., 2013 Consistent behavioural differences in WT= proactive and reactive (bold and shy) - persist over time and context relates to underlying differences in regulated gene networks and predicted protein–protein interactions - differences can be mapped to distinct regions of the brain variation in gene expression between individuals accounts for >9% of observed variation in the brain transcriptome

Answer 118

Rev et al., 2013 Focus on groups of genes that interact For proactive and reactive individuals, certain pathways have similar levels f upregulation eg. Notch signalling But others have different levels eg. DNA replication/DNA strand elongation These levels different in fore and hind brain Sub-networks specific to different brain parts are expressed differently in the 2 behavioural types (but what does this actually mean?)

Answer 119

specifies that closely related species or populations experiencing different ecological conditions should differ in a suite of metabolic, hormonal and immunity traits that have coevolved with the life-history particularities related to these conditions consistent individual differences also covary with life history and physiological differences at the within-population, interpopulation and interspecific levels (Reale et al., 2010) eg. in domestic dogs (which have undergone extensive artificial selection) life history relates to personality: obedient (or docile, shy) breeds live longer than disobedient (or bold) ones and that aggressive breeds have higher energy needs than unaggressive ones these correlations likely due to correlated responses to artificial selection on personality (Careau et al., 2010) eg. boldness predicts predation of pigeons by raptors data from 27 GPS tracked homing pigeons - very strong effects despite small sample size Larger and darker pigeons more likely to be predated, plus when shorter flight initiation distance, and lower escape reaction time

Answer 120

Kern et al., 2016 Associations between personality, morphology and locomotor performance Zebrafish caught in wild and then selectively bred to have proactive or reactive stress-coping style After 4-7 generations, morphology and locomotor performance differed between personality lines: bold zebrafish exhibited a larger caudal region and higher fast-start performance than fish in the shy line Stronger difference in females Correlated response of locomotor performance likely due to pleiotropy or physical gene linkage Correlated response of body shape may be due to linkage disequilibrium

Answer 121

Kern et al., 2016 Size-selective harvesting associated with commercial and recreational fishing practices has been shown to alter life history traits through a phenomenon known as fishing-induced evolution due to direct or indirect correlations between behaviour and life history Juvenile Bluegill Sunfish caught by using seine net or a lure (angled) tested for boldness, water column use and activity Fish caught by angling more timid and had fewer ectoparasites Therefore fishing techniques can affect population demographics -conservation implication

Answer 122

Grinsted et al., 2013 Indian cooperative spider Task differentiation in simulated prey capture attempts cross 10-day behavioural assays in the field, independent of developmental stage Participation in prey capture positively correlated with level of boldness but not aggression Body size positively correlated with being the first spider to emerge from the colony as a response to prey capture but not with being the first to attack Boldness and aggression are positively correlated, but neither correlate with body size or developmental stage Spider groups of all asocial or all social problematic, need a mix for highest RS within mixed colonies, social and asocial equally successful because of task specialisation asocial= aggressive- prey capture and group protection social=parental care, raining offspring having a high density of asocial even in a mixed colony is bad because aggressive to each other

Answer 123

Leading individuals get large foraging benefits (first to arrive at food)- dominant individuals tend to lead, social affiliation to leader mediates arrival order King et al., 2008 probability of a group being synchronized increased with the number of pregnant females, but decreased with the number of sexually swollen females in a group Synchrony also declined throughout the day Due to changes in activity budget of both sexes related to reproductive strategies & changing levels of satiation in individuals synchrony was highest in a ‘closed’ woodland habitat, and lowest in ‘open’ desert habitat due to habitat differences in food patch configuration and/or predation risk

Answer 124

Harcourt et al., 2009 2 chambered tank- one fish either side, each has safe area and open feeding area with opaque barrier, can establish which are bolder, which are shyer Experimental set up with transparent barrier and randomly paired fish of different boldness either side Markov chain model to infer the individual rules underlying their joint behaviour Each responded to each others movements- if one foraging, other more likely to forage, if one returns to cover other more likely to follow But bolder individuals displayed greater initiative and less responsive to partners (and more determination-out for longer), shyer individuals the opposite- followed faithfully Shyer individuals, as followers, also elicited greater leadership tendencies in partner (therefore positive social feedback) Nakayama et al., (2010)- success in recruiting a partner affects an individual's tendency to initiate the next trip, if successful, more likely to initiate subsequent trip and be less responsive to partner bold fish were unaffected by failures to initiate a joint trip, shy individuals were less likely to attempt another initiation after a failure

Answer 125

Nakayama et al., 2012 Measured tendency to initiate foraging trip in bold and shy individuals before and after satiating just one of them Before- bold = leader After, depending on which fed If fed shy, became less active but bold didn't change behaviour and continued to lead If fed bold, it initiated fewer trips, so shy began to initiate more more frequently In pairs that only differed a bit in temperament, observed role reversal

Answer 126

3 fish, 1 bold, 1 shy, 1 focal 2 treatments: focal= shy and focal= bold Shy focal fish: no fish out, tendency to leave low bold fish only out, tendency to follow shy fish only out, greater tendency to follow both fish out, greatest tendency to follow Pay more attention to those with similar personalities Bold focal fish: similar pattern by more noise, not as statistically clear

Answer 127

Harcourt et al., 2010 Pairs prefer to synchronise trips because reduces perceived predation risk fish trained to expect food at one of two alternate locations, paired fish with different expectations Compared to isolated individuals members of a pair showed increased tendency to alternate between foraging sites, taking it in turns to visit each one's preferred site Markov chain model suggests that fish respond to a partner that breaks the pattern of alternation by themselves reverting to less regular behaviour Here, pattern of alternation is actively monitored and maintained

Answer 128

Groups must be 5+ Jolles et al., 2017 Open arena, sociability predicts spatial positioning and leadership in social contexts, no effect of boldness more social fish, more time in centre and less in periphery less social more likely to lead level of sociability in group can affect its aggregation- if all sociable, would cause swarm because overly responsive to each other If add food patch of area of cover into arena, now there is a goal Both sociability and boldness predict foraging success bolder and less sociable fish get the most (more time away from group, can find more), more sociable, just follow others round and achieve less (boldness has less of an effect in more sociable individuals)

Answer 129

use waggle dance to indicate potential locations if 1 nest box available, quick decision made but if eg. 5, can take ~5 hours to reach decision Quorum sensing: Scouting Dancing (info passed from scouts to naïve) once reached a certain level... Piping (by queen- makes continuous noise) Leaving and head to new nest Process figures out which site preferred overall

Answer 130

Progressive Decision made once threshold reached Scouts start with tandem runs (lead naïve individual to nest) Follow up with transporting eggs and other individuals Then nest moves (phase shift) =shared decision but only part of pop involved in scouting (2/3 of pop don't recruit others) Decision based on number of ants in new nest System very stochastic

Answer 131

Benefits of consensus decisions: maintenance of group cohesion, enhancement of decision accuracy compared with lone individuals and improvement in decision speed Sumpter & Pratt, 2008 Choice between 2 model fish (1 more attractive than other) Given time, choose attractive one But if show very quickly, some will pick other one If more individuals in a group shown, more likely to follow attractive model An individual's probability of exhibiting a behaviour is a sharply nonlinear function of the number of other individuals already performing this behaviour Results match well with quorum-consensus model (based on the concordant theorem)

Answer 132

track movements with radio collars Decision to follow is dependent on number of initiators and whether they all initiated in same direction or not (more likely to follow if many in same direction) Compromise vs choice depends on angle between initiations - is low angle, compromise, if large, go with majority

Answer 133

uninformed individuals can prevent a few opinionated individuals from hijacking decisions- still have to reach a majority decision eg. release fish from holders- aim for blue or yellow target minority trained to go to incorrect target increasing number of uninformed prevents overruling, group goes to preferred despite strongly opinionated individuals

Answer 134

Aphaenogaster senilis correlation between exploratory activity and brood rescue behaviour Brood rescue predicts colony growth

Answer 135

Leaving cover and crossing dangerous area (when leaving and how quick to cross) to feed affected by social context and experience Groups cross faster, feel safer (larger group = faster) Groups w experience of group cross faster than individuals with experience Leaving cover on first social trial affected by personality, but effect disappears on second trial (no effect on crossing) Increasing experience with group, build up group behaviour, becomes more and more different from individual behaviours Crossing= experience trumps personality But boldness still predicts feeding success irrespective of experience consistent individual difference depend on context

Answer 136

primary symbiont of aphids not in other insect groups aphids do poorly without it

Answer 137

significant genetic variation in Daphnia clone susceptibility and Pasteuria infectivity strong reciprocal selective pressure because infected Daphnia sterile and parasites need host to transmit Pasteuria castrates host and redirects resources to growth- larger hosts can contain more parasite spores intermediate virulence optimal because too low= too slow, too high= can't get as many spores in host

Answer 138

Lycaenid caterpillars secrete honeydew that ants drink ants defend caterpillars against parasitic wasps co-adaptations

Answer 139

Prefer smaller lodgepole pine cones because easier to de-seed Driven increased cone width, mass and length where only crossbills present Causes increased crossbill beak size, those with bigger beaks better survival Unless squirrels also present- grater selection pressure on lodgepole cones because remove cone from tree and eat it from base (crossbills eat from opposite end) Therefore haven't driven larger cones when both present

Answer 140

New Zealand Freshwater snails infected by castrating trematode (Microphallus spp.) parasite (snails not definitive host, infective stage released into ducks when they eat snails) snails infected by eggs, larvae develop in snails- duck consumption needed to complete life cycle, therefore snails inaccessible to ducks (deep) may be less affected Experiment: expose shallow and deep snails to sympatric (coevolving w shallow snails) and allopatric (non-evolving) parasites- measure infection prevalence parasites only adapted to and able to infect shallow snails Parasites best at infecting the most common snail genotypes Host genetic background therefore changing over time Jokela et al., 2009 cycling of gene frequencies in snail pop appears to be driven by parasites- evolution of more resistant snails and loss of most vulnerable The Red Queen hypothesis postulates that high infection rates in the common asexual clones could periodically favour the genetically diverse sexual individuals and promote the short‐term coexistence of sexual and asexual populations In study of snails, within 7–10 years, the most common clones were almost completely replaced by initially rare clones in two different habitats, and sexuals persisted throughout Sexuals more common in shallows and asexuals more common in deep, supports Red Queen (King et al., 2011) Parasites that encounter a resistant host are often killed (strong reciprocal selection)- secondary infections (needed for parasite survival) only happen in <3% of cases

Answer 141

Pseudomonas bacteria and phi-2 phage Poullain et al., 2007 The ability of coevolved phage isolates to infect coevolved bacteria host varies across populations At the phage population level, coevolution led to the evolution of broader infectivity range, but without an associated decrease in phage growth rate relative to the ancestor, whereas phage evolution in the absence of bacterial evolution led to an increased growth rate but no increase in infectivity range Brockhurst et al., 2007 Time shift assay: bacteria from past, present and future (1 transfer difference) were exposed to phage taken from a single transfer bacteria evolved increased resistance to phage infection over time - both resistance and phage infectivity increased with time Paterson et al., 2010: coevolve Pseudomonas and bacteriophage, monitor phage molecule evolution pathogens evolve more rapidly in coevolution that when alone Future bacteria best at resisting contemporary parasites

Answer 142

Rough-skinned newts have powerful anti predator toxins (TTX)- 100x more powerful than necessary to kill Garter snakes= their major predator- evolved resistance to TTX (Brodie et al., 2002) snake resistance is predicted by newt toxicity some pops have drastically increased TTX resistance, others haven't (hotspots with 100% snake resistance, others with 0% resistance)- depends on arms race & importance of ecological effects in different geographic locations

Answer 143

trematodes can only infect snails with specific 'matching' genotype rare snail genotypes therefore should be less frequently infected than common

Answer 144

if infected, not reproducing if don't infect, don't pass on therefore v strong reciprocal evolution Decastecker et al., 2007 experiment: Resting eggs and bacterial spores fall to bottom of lakes- covered in lake sediments produces fossil record extract 'sediment core', contains isolated hosts and parasites from over a 4 year period Do past and future parasites match contemporary host? Contemporary parasites best at infecting (because coevolved to) Never see resistance or infectivity ever reach 100% Snail resistance to past and future parasites decreases with increasing resistance to contemporary

Answer 145

Inhabit post glacial lakes in British Columbia 2 morphs: Limnetic (shallow) Benthic (deep) some lakes have both, some have 1 solitary intermediate morph Measured fitness effects of competition between 'ancestral' intermediates and limnetics (biggest ecological competitor) from a lake w both (Schluter, 1994) Expect competition with limnetics, selection towards benthic form Tested for: stabilising selection without competition directional selection towards benthic form w competition coexistence- reciprocal selection for divergence in body size Competition promoted v rapid morphological diversification in just 3 months- strength of comp between mediated by phenotypic similarity

Answer 146

Richard Lenski's E. coli- 12 flasks of E. coli in glucose-limited environment, since 1988, transfer 1% into new flask every day, by about 50k generations, 70% of pops fitter see same genes mutating and phenotypic changes in different populations- lots of synonymous mutations, all occurring in same genes, suggesting natural selection was finding the same path and potentially the genes had good pleiotrophy at 30,000 generations, Cit+ evolved, bacteria booming- strain evolved mutations to metabolise citrate in medium

Answer 147

see 7 cospeciation events and 3 host switches (may be due to extinction)

Answer 148

intracellular endosymbionts synthesise essential amino acids, can't survive outside cells, genes completely interlinked in metabolic processes 100-250 My association co-phylogeny v well matched (Clark et al., 2000)

Answer 149

``` Aphids and Buchnera Psyllids and Carsonella Mealybugs and Tremblaya Sharpshooters and Baumania Whitefly and Portiera Cicadas and Sulcia and Hodgkinia Hoppers and Sulcia ```

Answer 150

Tsetse fly and Wigglesworthia Assassin bug and Rhodococcus Bed bug and Wolbachia

Answer 151

Fungus resistance - Hamiltonella Parasitoid resistance - Rickettsia, Ricketsiella, Spiroplasmia Heat tolerance - Serratia, Hamiltonella, Regiella Body colour - Ricketsiella Host plant adaptation - Regiella

Answer 152

Thought to infect over 60% of insect species mutualist in nematodea male embryo killer in blue moon butterfly (in some pops, female: male =100:1) mutualist in bed bugs- vitamin B provider (Hosokawa et al., 2010- development stunted when Wolbachia experimentally removed w antibiotics- eggs fail to develop- rescued by providing vitamin B)

Answer 153

causes stomach cancers and stomach ulcers | but protects against oesophageal cancer

Answer 154

Bt insecticide for lepidopteran pests- rapid death from punching holes in gut not from starving to death but microbiome entering blood and causing sepsis (treat w antibiotics prior to Bt- mortality down from 100 to ~10%) Broderick et al., 20066

Answer 155

response to mitochondria in wrong place eg. with severe cut or bruise and cells split open=> bloodstream Huge inflammatory response

Answer 156

Release of 24 wild rabbits, 1859- ate all vegetation fencing, poisoning, hunting all unsuccessful to control pops Pox virus that killed European rabbits identified- released in 1950, spread 2000km in 3 months 1900 - 3 million rabbits 1959 - 24 99% fatality rate pop reductions of 40-99.8% Comparison of original released pox strain w that in rabbits later- had evolved to different strains with lower virulence- had been to virulent to transmit (stabilising selection) through time, evolution of rabbit resistance compare museum and wild-caught samples from last 150 years (1800s-2013), same genetic adaptations in 3 different parts of the world (UK, mainland Europe, Australia) sequenced nearly 20,000 genes to pinpoint mutations that emerged since 1850s- many of the genes that have changes played a key role in rabbit immune system

Answer 157

wasps oviposit offspring into fig ovule, which develop, emerge and mate wasps pollinate figs nematode parasites infect female wasps and kill released from female cadaver and infect offspring (vertical) and others (horizontal) vertical only, if 1 female oviposits inside egg variation in strategy depends on fig wasp behaviour (eg. number wasps per fig) greater proportion of figs with 1 wasp = increased relative fecundity worms that infect wasps that produce single foundress broods have lower virulence (because not horizontal) ???

Answer 158

how do they avoid death from coevolving pathogens? eg. bdelloid rotifers- dessication to esape pathogens on dispersal (anhydrobiosis) If disperse 'dry' by wind, lose fungal pathogens During anhydrobiosis, genomes physically fragment and generate novelty- repair includes lots of environmental DNA- use this novel material in adaptation

Answer 159

Moritz et al., 2001 Part of the Red Queen hypothesis suggests that sexual reproduction is maintained in populations because of the need to continually create genotypes that confer resistance against rapidly evolving pathogens and parasites Parthenogenetic individuals of the prickly gecko, Heteronotia binoei, are much more prone to infection by mites than are their sexual relatives This is consistent across localities with different combinations of parthenogenetic genotypes and sexual chromosome races and occurs despite the unusually high genetic diversity of the parthenogenetic form

Answer 160

Slime moulds -depends on surface adhesion proteins (direct) Long tailed tits- vocal patterns (imprinted, indirect) Ground squirrels - olfactory cues w genetic component (armpit hypothesis- direct and indirect) Malarial parasites - ??? (know they do somehow, adjust their investment into male and female transmission stages according to the number of conspecifics sharing host)- maybe via host immune response?

Answer 161

mixed genotype infections cause lower RBC count and greater weight loss in mice- more virulent suggests immune system less able to control antigenic diversity mounting immune response against more than 1 parasite genotype more costly (Taylor et al., 1998) due to increased replication of transmission stages of Plasmodium- overwhelms immune system

Answer 162

Pathogenic bacteria Pseudomonas aeruginosa produces costly siderophores= iron-scavenging molecules, beneficial to growth Cooperation favoured by high relatedness of infecting bacteria and low competition between them- more related, increased siderophore production

Answer 163

Mouse system Quorum sensing favoured by high relatedness of bacteria in mouse Mouse mortality rate higher in high relatedness treatment Infections by QS bacteria grew to higher densities

Answer 164

experimental evolution of 5 different phages in different mixes and monoculture determined whether coevo took place using 13,500 time-shift assays (Betts et al., 2018) diverse parasite communities accelerated host evolution parasite from future more infective hosts evolved increased resistance diverse community therefore = increased community more diverse communities more likely to contain high selective force Bacteria evolved resistance by modifying cell surface making pathogen attachment harder In response, phages modified structures so can attach Increased parasite diversity drove shifts from Red Queen dynamics to arms race dynamics Characterised by a selective sweep of SNPs for generalist resistance mutations in genes of host bacterium cell surface lipopolysaccharide (bacteriophage receptor) - strategy suitable for variety of enemies, maintaining resistance to specific parasites slow and less effective drove faster molecular evolution in host pops and greater genetic divergence among pops therefore exposure to multiple parasites influences rate and type of host-parasite coevo parasites also evolve greater infectivity parasite communities = hotspots for antagonistic coevolution increased parasite diversity = more alleles reaching fixation (stronger arms race)

Answer 165

Mass de-worming programmes to treat hookworms exacerbated intensity of malaria by 2-3x study of 4000 people in Indonesia w coinfection competition for RBCs Also strain effect, P. vivax less competitive than P. falciparum therefore modify treatments accordingly- in areas where P. vivax common, should test for malaria before giving deworming treatment

Answer 166

Microbe mediated host defence drives evo of reduced pathogen virulence In C. elegans, infect w Staphylococcus aureus ("bad") and "good" gut bacteria (defensive mutualist) Enterococcus faecalis When coinfected, good defends against bad and use siderophores, causes bad to reduce siderophore production, evolve to be less harmful Adaptive mutualism evolves Staph kills ~50-100% after 24hrs Enterococcus kills ~1% after one week But together, intermediate virulence- worm survival from 52% to 82%, occurred quickly (can see changes by generation 5) Good could defend against different staph strains Found mutation in E. faecalis associated with superoxide production, harmful to staph Therefore coinfections drove C. elegans evo Coevolving bacteria with nematode= changes in in both worm and its gut bacteria- in presence of parasite, all three evolved together to increase worm survival host adapted by accommodating increased number of protective bacteria (more susceptible) If amount of host protection too low, also still slightly virulent so not worth having but if too high then other microbes may be wiped out, removing the need for protection ????

Answer 167

sampled 4 leaves from each of 8 trees looked at interior and exterior bacterial diversity (much higher within due to phage control) isolate bacteria from surface and inside, expose to phage Phage do much better on their local bacteria (p<0.0001) No difference in scale of interactions between leaves within same tree Time shift experiment to determine whether coevolution: 6 time points, 8 trees, 24 bacteria per tree: bacteria evolved increased resistance to phages of past than future phages do best on bacteria from same month or recent past Horse chestnut trees with lower microbiome bacterial diversity more likely to have canker, and related to disease severity shifting bacterial community composition associates with disease (not driven by any one in particular)

Answer 168

Stoll et al., 2010 have repeatedly evolved in several lineages with same function of containing and controlling bacterial symbionts- hide them from immune system and stop them spreading into other tissue but these beneficial symbionts can be harmful elsewhere in body

Answer 169

mucus is quite impenetrable phage 15x more likely to find bacteria host if stuck to mucus mucus universal to all animals; phage universal to all mucus (always more phage than bacteria) original immune system

Answer 170

3 different algae have 3 different predators, but if predators present, algae group together exposure to predator poo also induces grouping

Answer 171

106 symbioses- bacterial symbionts and host MCMC glmm comparative analysis and bacterial host phylogenies used both transmission mode and symbiont function correlate with host dependence - vertical transmission and nutritional symbionts = higher host dependence decrease in host fitness greatest for loss of nutrient provisioning, vertically transmitted removed negative correlation of host dependence and symbiont genome size in vertically but not horizontally transmitted- therefore function and pop structure important in driving irreversible dependence between hosts and symbionts

Answer 172

13 years underground as nymph, emerge, reproduce and die Symbiont has split into separate completely complementary species within cicada (now ~ 15-70)- only survive if all present reduced genome because in tight coevolution (black queen hypothesis) risky because now ocked in coevo dynamic

Answer 173

Symbiogenesis eg. 2 spp of wol+ fig wasps, Nasonia giraulti and longiconis Have different strains of Wolbachia Reproductive isolation curable with antibiotics- death by distorted hologenome normally, mating => cytoplasmic incompatibility due to clashing wolbachia strains Or could it just be because hybrids have faulty immune systems that leave them vulnerable to any bacteria? shouldn't be able to change a genotype with an antibiotic

Answer 174

ants defend from grazers in return for honeydew and protection BUT appears mutualists, but enzyme prevents ant digesting other sugars, enslaved

Answer 175

Inadvertent lures released by aphid microbes for predatory hoverflies Poliovirus attaches to microbiome bacteria to gain access to host cells Mouse mammary tumour virus attaches to microbiome to disguise from immune system

Answer 176

Coral bleaching Warming causes coral to expel algae, makes coral weaker Acidification decreases saturation state of aragonite, not as energetically favourable/available to build coral skeleton Get diseases eg. white pox, black band, pink line, red band

Answer 177

Gros Michel banana clone wiped out by 2 fungi | New Cavendish clone, but now a new worse fungus has developed- risk

Answer 178

influenza H1N1 from pigs and birds, <0.1% fatality rate HIV-1 from primates, 100% fatality rate SARS Corona virus from bats, 10% fatality rate more likely as species ranges change with climate change

Answer 179

Climate change = range changes As geographical range of a parasite increases, becomes maladaptive to host Parasite adaptation is inversely proportional to fraction of a host's range it occupies Loss of hotspots

Answer 180

harnessing coevolution to restore beneficial microbial gut community eg. for people on long-term antibiotic treatment/chemotherapy But is it a drug or organ donation? will affect its availability- how food and drug administration regulates procedure, how much it costs and who profits

Answer 181

infected by nematodes and Spiroplasma Spiroplasma provides resistance to nematodes by encoding a Ribsome Inactivating Protein which cleaves specific sequences of ribosomal RNA of nematode but not insect Symbiosis so beneficial that it has spread trough many pops - enhances fertility 10-fold compared to unprotected flies

Answer 182

Wolbachia increases Drosophila resistance to broad range of RNA viruses, but also manipulated host reproduction by inducing cytoplasmic incompatibility Transmitted vertically from infected female to offspring (males = deadend) When male infected, sperm modified so embryo dies early in development But in females, encodes a 2nd factor that rescues embryo Therefore Wolbachia infected females have reproductive advantage, bacterium can invade pops and be maintained

Answer 183

Wolbachia decreases ability of mosquitos to carry and transmit Dengue and Zika virus Large scale releases of Wolbachia infected mosquitos underway

Answer 184

Siderophores scavenge metals (free iron) from environment a 'public good' in microbial communities can lead to cheating in P. fluorescens But community species sorting selects for high producing species- so intermediate levels of cooperation favoured Heavy metal contamination global problem In nature, 94 samples across heavy metal gradient samples analysed for heavy metal content, pH, bacterial composition, proportion siderophore-producing bacteria per sample, bacterial density (16S RNA) positive correlation between heavy metal content and siderophore production cheating probably doesn't matter (but only a correlation) In lab, Hesse et al., 2018: experimental evolution of natural compost communities after 3&6 weeks, sampled 24 clones per microcosm for siderophore production Production higher in cooper contaminated microcosms no obvious effects of cheating ????

Answer 185

Manipulate natural pathogens to decrease virulence eg. C. diphtheriae causes dihtheria, there are multiple strains virulent strains produce toxin, avirulent don't toxin is costly to produce but helps it to obtain resources Vaccinations that target toxins means net cost of producing, therefore outcompeted by avirulent But danger they might increase virulence vaccines against Plasmodium slow parasite growth but don't stop it could cause it to evolve higher intrinsic replication rate in compensation, could make it more virulent to unvaccinated people

Answer 186

Blind shrimp digs burrow Goby guards and when predator comes warns shrimp with tactile communication (flick w tail), hide together don't know enough to call coevo

Answer 187

cooperative hunting in coral reefs grouper = fast moray eel can fit between rocks corner prey referential gesture of grouper with face to indicate where prey is to demonstrate that gesture is due to coevo need to show = an adaptation (that grouper does because of association and that eel responds because of association)

Answer 188

Mock, 1987 Siblicidal species sibling aggression was significantly lower in two-chick broods than 3-chick broods when larger broods, parents don’t interfere with sibling fights but their presence seems to subtly decrease aggression (fewer “sever” fights occurred when parents were present) No favouritism of food distribution to dying chick victims Interpretation difficult, could mean one of two things: Either evolutionary conflict exists and offspring are winning or there is no conflict If no conflict, parents create asymmetries between offspring (eg. via hatch asynchrony) then leaving sibling conflict to run its course without further intervention -- siblicide and other investment-skewing selfish chick behaviours may serve parental interests well (equal allocation of resources likely to not be parental optimum)

Answer 189

(Head et al., 2005) Acheta domesticus Mated females with attractive or unattractive males, fitness components combined to give relative estimates for number of grandchildren produced females mated to attractive males incur a substantial survival cost, however costs are cancelled out and may be outweighed by the benefits of having offspring with elevated fitness Mostly due to increasing attractiveness of sons This reveals the value of estimating the net fitness consequences of a mating strategy by including measures of offspring quality in estimates of fitness

Answer 190

Bee et al., 2000 Rana clamitans Males broadcast calls to conspecifics in order to defend their territories Smaller male frogs have been observed to lower the dominant frequency of their calls in response to the low frequency calls of large male frogs, in order to give the appearance of being larger than in reality and maintain territory dominance when played the calls of smaller frogs, the relative lowering of frequency is less extreme, demonstrating that this behaviour isn’t a standard response to territorial contests, and instead a dishonest signal they produce to manipulate the outcome of contests that are not in their favour

Answer 191

Rubenstein et al., 2012 Injured fish produce alarm signals that signal danger to nearby conspecifics, increasing their chances of responding and surviving In fish that shoal with close kin, such communication could have inclusive fitness benefits to the signaller

Answer 192

Halliday & Slater, 1983 black patches on a bird’s throat and breast signals its status. Experiments- pale birds’ breasts are dyed black = increase their success against subordinates, but dominant birds are able to detect them from their behaviour and are more aggressive towards them as a result therefore cost selecting against dishonesty

Answer 193

Halliday & Slater, 1983 females choose males to mate with dependent on the size of the nuptial gift they present Occasionally, males manipulate rival males into handing over their gifts by mimicking a sexually receptive female and subsequently use the gift as their own to acquire matings- but this only happens infrequently compared to encountering receptive female, less costly to be deceived occasionally

Answer 194

Hughes et al., 2008 most significant potential conflict in social insect colonies is over which individuals become reproductive queens rather than sterile workers individual larvae will maximize their fitness by becoming queens whereas their nestmates will generally maximize fitness by forcing larvae to become workers -- evolutionary constraints are thought to prevent cheating by removing genetic variation in caste propensity However, one-fifth of leaf-cutting ant patrilines cheat their nestmates by biasing their larval development toward becoming queens rather than workers Mechanisms = general tendency to become larger adult, and relating to worker-queen developmental switch “royal” genotypes are rare both in the population and within individual colonies, to avoid suppression by cooperative genotypes, the efficiency of which is frequency-dependent Therefore cheating can be widespread in even the most cooperative of societies

Answer 195

piRNAs, in mammalian cells mainly in spermatogenic cells in testes, but also ovarian somatic cells and neuron cells in invertebrates, as well as in many other mammalian somatic cells -- endogenous system for silencing the expression of selfish genetic elements such as retrotransposons and thus preventing the gene products of such sequences from interfering with germ cell formation

Answer 196

Pulliam et al., 1982 cooperation in vigilance increases overall group vigilance. Potential for ‘cheating’ and not being vigilant? But, remains as ESS with everyone sharing vigilance, but for selfish reasons- spotting predator first allows greater chance of escaping.

Answer 197

Yucca plant pollinated exclusively by yucca moth, which depends entirely on feeding on the plant for survival Yuccas also provide protection deep within flowers for moth to lay eggs

Answer 198

Steiner & Whitehead, 1990 South Africa, bees use forelegs to collect oil from Diascia floral spurs, pollinating them in the process Studies have shows that in several populations mean foreleg length in bees correlates with spur length of specific Diascia population visited

Answer 199

Between cuckoo trickery and host defences Host patterns evolve - signatures that are replicable, distinct and hard to forge eg. Spottiswoode & Stevens, 2012: egg polymorphisms are extreme in the African tawny-flanked prinia and its parasite, the cuckoo finch used models of avian visual perception to analyse the appearance of prinia and cuckoo finch eggs from the same location over 40 years Egg colours of both species have diversified over time, expanding into avian colour space as expected under negative frequency-dependent selection Host and parasite appear to be closely tracking one another’s evolution, since parasites showed closer colour mimicry of contemporaneous hosts eg. (Lahti & Lahti, 2002) Village weaverbird eggs very different between individuals, hypothesised due to cuckoo parasitism Found that (1) interindividual egg variability permits individuals to discriminate between own and foreign eggs by rejecting eggs in proportion to the difference in appearance from their own; (2) village weavers remember the appearance of their own eggs and do not require a discordancy within their clutch, nor even the presence of one of their own eggs, in order to distinguish a foreign egg as such; and (3) colour and speckling contain the signature information by which village weavers can distinguish their eggs from foreign ones; whereas shape and mass, being less reliable, do not Lahti, 2005 Introduction of village weaverbirds onto islands lacking cuckoo parasitism, eg. Mauritius one century ago Here, between-individual variation and within-clutch consistency in egg appearance have both decreased compared to mainland birds supports the hypothesis that egg appearance in the African village weaver has been maintained by natural selection as a counteradaptation to cuckoo brood parasitism

Answer 200

(Brown, 1998) Nest in colonies that serve as information centres in which unsuccessful individuals locate and then follow successful individuals to aerial insect food resources In larger colony size: -increased number swallows departing colony per hour (individuals did not wait as long to locate appropriate foraging associates in large colonies as in small colonies) -decreased waiting time between an individual's arrival at its nest and its departure on its next foraging trip -returned with food for their nestlings more often and brought more food per trip -nestling body mass at 10 d of age increased significantly with colony size -adult body mass late in the nesting season during the period of feeding of nestlings increased Experimental reduction of large Cliff Swallow colonies to the approximate size of small colonies suggested that nestling and adult body mass in these colonies did not vary with colony location

Answer 201

one reason swarms arise is because the presence of multiple moving prey in swarms causes confusion for attacking predators Using an evolutionary model of a predator–prey system, Olson et al., (2013) show that predator confusion provides a sufficient selection pressure to evolve swarming behaviour in prey Plus, evolutionary effect of predator confusion on prey could in turn exert pressure on the structure of the predator's visual field, favouring the frontally oriented, high-resolution visual systems commonly observed in predators that feed on swarming animals

Answer 202

hard/durable matrix that bacteria live in, produced by quorum sensing bacteria reliant on it bacteria in centre can e protected from eg. antibiotics but hard to get nutrients when in centre open K+ channels and release K+ to direct glutamate from outer bacteria into centre =cooperative mechanism if all bacteria were clones, could it be considered multicellularity?

Answer 203

Price et al., 2019 background matching = resembling the colour and pattern of the environment may be ineffective in complex habitats where matching one patch may lead to increased visibility in other patches disruptive coloration = disguises body outlines, may be effective against complex backgrounds tested camouflage strategies of shore crab in rock pools and mudflats (species = highly variable, capable of plastic changes in appearance, and lives in multiple environments) predator (bird and fish) vision modelling and image analysis to quantify background matching and disruption in crabs from rock pools and mudflats rock pool individuals had significantly higher edge disruption than mudflat crabs, whereas mudflat crabs more closely matched the substrate than rock pool crabs for colour, luminance, and pattern expression of camouflage strategies dependent on the visual environment

Answer 204

Ling et al., 2019 Current understanding of collective behaviour in nature is based largely on models that assume that identical agents obey the same interaction rules, but in reality interactions may be influenced by social relationships among group members show that social relationships transform local interactions and collective dynamics tracked individuals’ three-dimensional trajectories within flocks of jackdaws, a species that forms lifelong pair-bonds flocks contain internal sub-structure, with discrete pairs of individuals tied together by spring-like effective forces Within flocks, paired birds interacted with fewer neighbours than unpaired birds and flapped their wings more slowly, which may result in energy savings But, flocks with more paired birds had shorter correlation lengths, which is likely to inhibit efficient information transfer through the flock This reveal a critical tension between individual- and group-level benefits during collective behaviour in species with differentiated social relationships

Answer 205

Kalb et al., 2019 Study of great tits in the field presented with taxidermy mounts of predators - tawny owl (low risk), sparrowhawk (high risk) observed them vary the number of D elements and the interval between those elements; produced significantly longer D calls with more elements and longer intervals between elements when confronted with a sparrowhawk, as well as a greater number of D calls suggests that the basic D calls are varied depending on threat intensity

Answer 206

When a female is sexually promiscuous, the ejaculates of different males compete for the fertilization of her eggs; the more sperm a male inseminates into a female, the more likely he is to fertilize her eggs. Because sperm production is limited and costly, theory predicts that males will strategically allocate sperm (1) according to female promiscuity (2) saving some for copulations with new females, and (3) to females producing more and/or better offspring Pizzari et al., 2003 Male fowl show status-dependent sperm investment in females according to the level of female promiscuity; they progressively reduce sperm investment in a particular female but, on encountering a new female, instantaneously increase their sperm investment; and they preferentially allocate sperm to females with large sexual ornaments signalling superior maternal investment