REPRODUCTION Ovis canadensis MATING SYSTEMS SEXUAL SELECTION Oviparity Ancestral reproductive mode of amniotes Turtles & Crocodilians Large clutches of small eggs (relative to adult size) Sea Turtle – no maternal care Aligator – basic maternal care (nest guarding) Hatchlings are PRECOCIAL require no further maternal provisioning Ovoviviparity (egg retention) lizards and snakes Live birth of small well-developed young, Small number of eggs hatch internally (maternal protection) Horned lizard (Phrynosoma) Rattlesnakes (Crotalus) Protection of young-- frequency of ovoviviparity increases with increasing environmental stress Extra provisioning-- some degree of “viviparity” (internal transfer of nutrients from mother to young) Young are PRECOCIAL independent at birth requiring no further maternal provisioning Rattlesnakes (Crotalus) Oviparity in birds In general, birds produce relatively large eggs (with large yolk) Clutch size is highly variable Bald eagle 1-3 Blue-winged teal 4-16 American Robin 3-4 Gray partridge Active birds – small clutches More sedentary birds – larger clutches 10-20 Oviparity in birds Reproductive constraints imposed by flight Limitations on egg size & clutch size A solution : sequential laying AND sequential hatching Barn Owl Kiwi Flightless birds have larger relative egg mass egg 8x the mass of a similar-sized flying bird Oviparity in birds Relative development of young Altricial Precocial In birds, even relatively precocial young require continued parental investment Oviparity in monotremes Small clutch (1-2); eggs very small relative to maternal size. Prehatching incubation period is very short compared to birds Young are extremely Atricial at hatching Require extended postnatal provisioning Postnatal care is strictly maternal (protection & lactation) Life history of Mesozoic prototherians Conjecture based on comparison to living groups Morganucodon Ptilodus Oviparous – some possibly ovoviviparous Altricial young Extended post-hatching maternal care Lactation Marsupials “ovoviviparous” or “semi-viviparous” Retention of egg shell membranes Limited placentation Neonate extremely altricial, requiring extended postnatal provisioning Postnatal care is strictly maternal (protection and lactation) Little developmental flexibility – fixed short gestation and extended lactation Wide variation in litter size Placentals – evolution of complete viviparity Elaborate placentation Prolonged gestation period Considerable developmental flexibility – gestation and lactation periods are variable Placentals – viviparity Neonatal development is variable, from: Ailurops highly altricial-- Equus -- to highly precocial but ALWAYS requiring extended postnatal provisioning Postnatal provisioning is exclusively maternal (lactation) Lactation Pre-weaning provisioning by females Why not male lactation? Dyacopterus spadiceus Francis. et al; "Lactation in Male Fruit Bats," Nature, 367:691, 1994. What limits PATERNAL care in mammals? Monogamy is rare in mammals (only 3 to 5% of all species) Generally FACULTATIVE (involving mate guarding, or due to scarcity of potential mates) Not OBLIGATE (involving direct paternal investment) Dolichotis IN CONTRAST Monogamy is common in birds (90% of all species) Monogramy is OBLIGATE Often involves mating for life Why the difference? Campephilus Males can directly increase their fitness through paternal care Parental Care in Mammals LACTATION makes maternal care absolutely essential in ALL mammals (to ensure direct fitness) Females may also help female kin rear offspring (indirect fitness) Cooperative female kin (the foundation of social behavior) Paternal care is restricted even in species that are monogamous Reproductive constraints of flight in bats Pteropus Haliaeetus Lasiurus Constraints of flight are much greater in viviparous mammals compared to oviparous birds Females have an unavoidable and prolonged mass increase during pregnancy Altricial young require protection and nursing (are often are carried in flight) RESULT: Most bats have small litters of 1-2 young (in a few species up to 4 young) Bats have very low reproductive output compared to other mammals of similar size (e.g. rodents) BUT they are much longer-lived Sexual strategies The sexes have different fitness perspectives: Females – mate quality, resource availability Males – mating opportunities Result: Male-male competition for mating opportunities Female choice of mates (and often associated resources) Mating systems Polygyny Single male mating with multiple females Harem polygyny -- male “control” of female groups to achieve exclusive matings Polyandry Single female mating with multiple males, AND males take on rearing responsibilities (i.e., “role reversal”) uncommon in birds (2% of species) Impossible (?) in mammals due to lactation Promiscuity Both sexes mating with multiple partners Many (most?) mammal species are promiscuous Multiple mating, but NOT necessarily indiscriminate mating May reflect male-male competition (“sperm competition”) e.g. – “copulatory plugs” in rodents May involve active selection by females For multiple mates (i.e., advantages of multiple-sired litters) Sexual Selection Differential selection for traits in one sex (males) through the action of male-male competition and female choice Size dimorphism Sexual combat structures Display behavior and structures