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Main Points

1) Diet -- primitive , cecal fermentation, and fermentation -- ruminant and an adaptive radiation -- example: the global distribution of mammalian

2) Metabolism and allometries -- universal patterns in ecology -- example: allometries for population density in carnivores

Terms: , meristem, frugivory, coprophagy, fermentation, , succession, metabolic rate, allometry, energy-equivalence rule

Pre-reading: Wednesday 11 October = NA Monday 16 October = Emlen and Oring

1 Diet: Insectivores and Carnivores

• insectivory (including myrmecophagy) and carnivory -- simplest digestive tract -- no fermentation or (or cecum is very small) -- food is readily digested, because of high energy content, no cell walls, and low fiber

2 Diet: Herbivores • Plants as food: -- leaves are low in -- less concentrated -- protected by biotic, chemical, and mechanical defenses

chili w/chemical defense barrel cactus bullhorn acacia (capsaicin) w/mechanical defense w/mechanical (thorns) and (spines) biotic defense (ants)

3 Diet: Herbivores • Plants as mammal food -- cellulose in cell walls -- ontogeny of plants -- graminoid vs. forb meristems

4 Diet: Herbivores • primitive monogastric digestion -- simple alimentary canal (AKA digestive tract) -- often leads to frugivory or omnivory

collared

bearded

5 Diet: Herbivores • hindgut or cecal fermentation -- enlarged cecum houses fermentation -- short rate of passage, low cellulose utilization -- may lead to coprophagy in smaller species

mountain tapir

6 Diet: Herbivores • foregut or ruminant fermentation -- fermentation occurs in four-chambered -- long rate of passage, high cellulose utilization,

American

7 Discussion Q: Families and Cervidae have proliferated in the past 15,000 years, while rhinos, , and equids have declined in number. Come up with a hypothesis for why this has happened.

15,000

5 MYA

34 MYA

8 vicuna saiga

water

9 • succession = predictable changes through time in ecological communities

10 • succession = predictable changes through time in ecological communities

succession -- materials pass twice as fast through cecal fermenters

-- protein assimilation less efficient than

11 • Grazing succession -- smaller have higher mass-specific metabolic rates

-- smaller animals have higher relative energy demands, but lower absolute energy demands

-- for smaller animals, intake is less, but assimilation must be high

12 Metabolism and body size

• metabolic rate = energy consumption or expenditure, typically measured in kJ or kCal consumed per day

13 Metabolism and body size

• metabolic rate = energy consumption or expenditure, typically measured in kJ or kCal consumed per day

• allometry = change in trait or process with body size that is non-linear

14 Metabolism and body size

• allometry = change in trait or process with body size that is non-linear

1:1 line

Positive allometry Log size rostrum Log Rostrum size increases steeply (slope >1) with skull size

15 Log skull size Metabolism and body size

• allometry = change in trait or process with body size that is non-linear

Negative allometry 1:1 line Head size increases slowly (slope <1) with

body size Log Log head size

16 Log body size Metabolism and body size

• metabolic rate

• allometry

linear axes log10 axes

Kcal/kg/day Kcal/kg/day

Kcal/day Kcal/day

17 body mass (kg) body mass (kg) Metabolism and body size

• metabolic rate

• allometry total metabolism (kJ) = a(mass^0.75)

mass-specific metabolism (kJ) = a(mass^-0.25)

18

Metabolism and body size Metabolic rate rate (W) Metabolic

Slope = 0.75

body mass (kg) 19

Metabolism and body size

)

2 Density Density km / (inds Slope = -0.75

100 101 102 103 104 105 106 107 From Brown 1995 20 body mass (g) Metabolism and body size

• Energetic-equivalence rule = change in abundance with body size is offset by change in metabolic rate with body size, so that populations exhibit invariance in energy use

with respect to size.

etabolism of of etabolism

population m

body mass (g) 21 body mass (g) body mass (g) Metabolism and body size

K. Roy. 2017.

“How to be an elephant” 22 http://katherineroy.com/elephants/ Digestion and global diversity of

• Predictors of plant quantity: -- more rain = more plants -- higher soil fertility = more plants

23 Digestion and global diversity of ungulates

• Predictors of plant quality: -- more rain = lower N and P content -- higher soil fertility = higher N and P content

24 Digestion and global diversity of ungulates

• Plants are most abundant in wet, high-fertility environments

~20 kg plains ~200 kg buffalo ~800 kg

25 Digestion and global diversity of ungulates

• Plants are most abundant in wet, high-fertility environments

• Plants are most nutritious in dry, high-fertility environments klipspringer ~20 kg ~200 kg buffalo ~800 kg

26 Dicussion Q: in light of the fact that plant quantity is most important to large ungulates, and plant quality is most important to small ungulates, come up with a hypothesis for where diversity should be highest.

klipspringer ~20 kg plains zebra ~200 kg buffalo ~800 kg

27 Global diversity of ungulates

3000 kg 15 kg

Plant available moisture Plant available N

From Olff et al. 2002.

28 Global diversity of ungulates

3000 kg 15 kg

Plant available moisture Plant available N

From Olff et al. 2002.

29 Global diversity of ungulates

Plant available moisture

Plant available N

30 Global diversity of ungulates

• High correspondence between predicted and observed diversity of large herbivores, based on rain and soil fertility

• Suggests that plant-available nutrients and moisture maintain species diversity of ungulates

31