Preparation Unit 1 Basics of Domestic Animal Biology

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1 Evolution, Domestication, Breeding

Estimated number of animal species in different groups and number of livestock species recruited from those Species Group total domestic Mammals 4 500 35 Birds 10 000 8 Amphibians & Reptiles 12 000 - Fish 22 000 (2) Insects 960 000 2 Others 400 000 2

Livestock or domestic livestock: Definitions Animalskeptordealtinforuseorprofit(TheConciseOxfordEnglishDictionary) Cattle, horses, poultry, and similar animals kept for domestic use but not as pets, especially on a farm or ranch (Collins English Dictionary Complete and Unabridged Harper Collins) Domestic animals, such as cattle or horses, raised for home use or for profit, especially on a farm(the American Heritage Dictionary of the English Language) Farm animals, with the exception of poultry. In Western countries the category encompasses primarily cattle, sheep, pigs, goats, horses, donkeys, and mules; other animals (e.g., buffalo, oxen, or camels) may predominate in other areas. (Merriam-Webster Concise Encyclopaedia) In the insurance sector a difference is made between livestock and bloodstock. Bloodstock are horses. Livestock are the rest. In accountancy a difference is made between livestock and deadstock. Deadstock are tractors and machinery and seed andotherinputs-i.e.stockthatisdeadatthe endof the yearoratleastnotalive. Livestock are things that are alive and are valued as stock at the end of the year. (Philip Colfox, 2012)

Numbers and biomass of domestic animals and humans Source: FAO Stat 2013 and author s calculations Species Numbers (million) Average live weight (kg) Biomass (million t) Cattle 1 485 350 520 Buffalo 198 400 79 Sheep 1 169 38 44 Goats 996 35 35 Camelids 27 + 9 380 / 90 11 Equines 59 + 44 + 10 300 / 140 / 280 27 Pigs 966 95 92 Poultry 24 075 1.6 38 Total 846 Humans 6 800 55 374

Global Biomass [%] of different livestock species Data Source: FAO 2014 & Author s calculations

The process of Evolution Evolution is change in the heritable traits of biological populations over successive generations. Evolutionary processes give rise to diversity at every level of biological organisation, including the levels of species, individual organisms, and molecules. All of life on earth shares a common ancestor known as the last universal ancestor, which lived approximately 3.5 3.8 billion years ago. Repeated formation of new species (speciation), change within species (anagenesis), and loss of species (extinction) throughout the evolutionary history of life on Earth are demonstrated by shared sets of morphological and biochemical traits, including shared DNA sequences. These shared traits are more similar among species that share a more recent common ancestor, and can be used to reconstruct a biological "tree of life" based on evolutionary relationships (phylogenetics), using both existing species and fossils. The fossil record includes a progression from early biogenic graphite, to microbial mat fossils, to fossilized multicellular organisms. Existing patterns of biodiversity have been shaped both by speciation and by extinction. More than 99 percent of all species that ever lived on Earth are estimated to be extinct. Estimates of Earth's current species range from 10 to 14 million, of which about 1.2 million have been documented. Source: https://en.wikipedia.org/wiki/evolution

The evolution of Ungulates

Mutation and natural selection driving evolution Source: https://en.wikipedia.org/wiki/evolution

Homologous bones in the limbs of tetrapods Source: https://en.wikipedia.org/wiki/evolution

The Evolution of Domestic Grazing Systems Evolution of Ruminants Gregariousness Coevolved with Grasses Microorganisms Humans Domestication Rapid Expansion in Miocene Expansion in Ruminant Diversity Cellulose Breakdown Accounted for by the Savannah Hypothesis Lactose Tolerance Patterns Milk Consumption rising

The Savannah Hypothesis Renato Bender, Phillip V. Tobias, Nicole Bender (2012) The Savannah Hypotheses: Origin, Reception and Impact on Paleoanthropology. Hist. Phil. Life Sci., 34 (2012), 147-184. http://hornshire.com/aah/2012%20savannah%20hypotheses%20-%20bender%20et%20tobias.pdf

If you want to learn more about evolution, check this website http://evolution.berkeley.edu/evosite/evohome.html

What is a species? Species: The lowest taxonomic rank, and the most basic unit or category of biological classification; it is the major taxonomic subdivision of a genus or subgenus. Species as the basic category of biological classification are composed of related individuals that resemble one another closely and are able to interbreed and produce fertile off- spring. Subspecies: A uniform, genetically distinct population within a species, often geographically isolated from the main body of the species. A species can contain several subspecies.

Phylogenetics of domestic ungulates Phylogenetics is the study of phylogenesis, or the evolutionary history, development and relationships among groups of organisms (e.g. species, or populations). These are discovered mainly through molecular data matrices, based on nucleic acids sequences and protein structures. Thus, the result of phylogenetic studies are hypotheses about the evolutionary history of taxonomic groups or their phylogeny. The evolutionary branching process may be depicted as a phylogenetic tree, and the place of each of the various organisms on the tree is based on a hypothesis about the sequence in which evolutionary branching events occurred. Taxonomy is the classification, identification and naming of organisms. It is usually richly informed by phylogenetics, but remains a methodologically and logically distinct discipline. The degree to which taxonomies depend on phylogenies (or classification depends on evolutionary development) differs depending on the school of taxonomy: phenetics ignores phylogeny altogether, trying to represent the similarity between organisms instead; cladistics (phylogenetic systematics) tries to reproduce phylogeny in its classification without loss of information; evolutionary taxonomy tries to find a compromise between them in order to represent stages of evolution Source: https://en.wikipedia.org/wiki/phylogenetics

Example: Taxonomy of the domestic goat Eukaryota Animalia Chordata Mammalia Artiodactyla Bovidae/Caprinae Capra Capra hircus = Domestic Goat

The Artiodactyla Family Tree (even-toed ungulates) Source: http://www.ultimateungulate.com/index.html

The Position of Bovini and Caprini in the Ruminant Family Tree Ruminants

Bovi idae Pseudoryx Bos Bison Bubalus Syncherus The Bovini subfamily tree Saola - Pseudoryx nghetinhensis Aurochs - Bos primigenius Wild Yak - Bos mutus Gaur - Bos gaurus Banteng - Bos javanicus Kouprey - Bos sauveli American Bison - Bison bison European Bison (Wisent) - Bison bonasus Wild Water Buffalo - Bubalus arnee Tamaraw - Bubalus mindorensis Lowland Anoa - Bubalus depressicornis Mountain Anoa - Bubalus quartesi African Buffalo - Syncherus caffer

The Caprini subfamily tree Bovidae * * * * * Genus Capra Genus Ovis

The Perissodactyla Family Tree (odd-toed ungulates) http://www.ultimateungulate.com/index.html The Equid Genus Tree

The Family Suidae Genus Babyrousa Babirusa (3 species) Hylochoerus Forest Hog (1 species) Phacochoerus Warthogs (2 species) Porcula Pygmy Hog (1 species) Potamochoerus Bush Pigs (2 species) Sus True Pigs (9 species) Sus scrofa Eurasian wild pig (wild form of the domestic pig, Sus domesticus)

Range Map of Sus scrofa

The Camelid (Tylopodae) Family Tree Ar rtiodactyla Camelus dromedarius Camelus bactrianus (ferus?) Lama (3 species) Vicugna vicugna

The process of domestication Loose contacts between man and animal, taming of individuals, with free breeding Confinement to human environment; with breeding in captivity Selective breeding organised by man to obtain certain characteristics; occasional crossing with wild forms Economic considerations of man leading to planned development of breeds with desirable properties Wild ancestors persecuted or exterminated

Temporal sequence of species in domestication Mammals domesticated in the pre-agricultural phase: Dog, Reindeer, Sheep, Goat Mammals domesticated in the early agricultural phase (crop-robbers): Cattle, Buffalo, Gaur, Banteng, Yak, Pig Mammals domesticated for transport: Horse, Bactrian Camel, Dromedary, Donkey, (Elephant) Mammals domesticated for pest control: Cat, Mongoose, Ferret Latecomers: Rabbit, various recent domesticants Without temporal classification: Birds, Insects, Fishes, Molluscs

Number of domestic species in different body size classes compared to the number of all known animal species 1000000 100000 10000 Number of all species [log] Number of domestic species 25 All species Domestic species 20 15 1000 10 100 5 10 1 0.01-0.05 0.05-0.1 0.1-0.5 0.5-1 1-5 5-10 Body length [m] 0

Behavioural pre-adaptations to domestication Social Structure Favourable Dominance hierarchy Large gregarious groups Male social group affiliation Persistent groups Unfavourable Territoriality Family groups or solitary Males in separate groups Open memberships Food Preferences Generalist herbivores or omnivores Dietary specialist or carnivore Captive Breeding Aggressiveness ( within or between species) Captive Temperament Polygamous/Promiscuous mating Males dominant over females Males initiate Precocial young Non-aggressive Tameable/readily habituated Readily controlled Solicits attention Limited agility Small home range Wide environmental tolerance Non-shelter seeking Implosive herd reaction to threat Pair bonding prior to mating Females dominant Females initiate Altricial young Naturally aggressive Difficult to tame Difficult to control Avoids contact/attention Highly agile/difficult to contain Large home range Narrow environmental tolerance Shelter seeking Explosive herd reaction to threat

Domestication of Ungulates in the Fertile Crescent Fertile Crescent Modified after: Zeder, M.A. (2008)

Centres of domestication based on archaeological and molecular genetics information Pig Rabbit Cattle Sheep Goat Pig Bactrian Camel Reindeer Horse Yak Pig Swamp Buffalo Chicken Turkey Pig Bali Cattle Chicken Guinea pig Llama Alpaca Donkey Cattle? Dromedary Cattle Water Buffalo Redrawn after: FAO 2008

The Range of Bos primigenius subspecies Taurine Cattle ~10 500 yr BP Zebu Cattle ~ 8 500 yr BP

Genetic diversity through breeding A breed is a specific group of domestic animals or plants having homogeneous appearance, homogeneous behaviour, or other characteristics that distinguish it from other animals or plants of the same species and that were arrived at through selective breeding. Despite the centrality of the idea of "breeds" to animal husbandry, no scientifically accepted definition of the term exists. A breed is therefore not an objective or biologically verifiable classification but is instead a term of art amongst groups of breeders who share a consensus around what qualities make some members of a given species members of a nameable subset. The term is distinguished from landrace, which refers to a regional variety of domestic (and sometimes feral) animal through relatively uncontrolled breeding. When bred together, animals of the same breed pass on these predictable traits to their offspring, and this ability - known as "breeding true - is a requirement for a breed. Plant breeds are more commonly known as cultivars. The offspring produced as a result of breeding animals of one breed with other animals of another breed are known as crossbreeds or mixed breeds. Crosses between animal or plant variants above the level of breed/cultivar (species, subspecies, botanical variety, even different genera) are referred to as hybrids. http://en.wikipedia.org/wiki/breed

Dromedary and Bactrian Camel Different Species or just Breeds? Source: S. Legel (1990) Nutztiere der Tropen und Subtropen

Three patterns in which natural or man s selection may lead to differentiation into breeds

Main cattle breed groups Taurine Breeds Bos primigenius Sanga Breeds Zebu Breeds

Distribution of cattle breed groups No Cattle Taurine Cattle Zebu Cattle Sanga Cattle Bos/Bibos Species

Taurine Bos (Bibos) Zebu Sanga

Genetic diversity in domestic livestock shown by approximate number of breeds world wide Species No of breeds Buffalo 17 (+) Camels Dromedaries Bactrian 97 16 Cattle 850 (+/-) Chicken 600 (+) Sheep 380 (+/-) Goats 110 (+) Horse 1400 Pig 200 (+/-)

Bos primigenius extinct since 1627 AD

Breeding back the Aurochs - An example for the creation of a breed Heck cattle are a hardy breed of domestic cattle. These cattle are the result of an attempt to breed back the extinct aurochs from modern aurochs-derived cattle in the 1920s and 1930s. Controversy revolves around methodology and success of the program. There are considerable differences between Heck cattle and the aurochs. Furthermore, there are other cattle breeds which resemble their wild ancestors at least as much as Heck cattle. Taurus cattle are an advanced form of Heck cattle, an attempt of breeding back the aurochs, the wild ancestor of domestic cattle. They are the result of crossbreeding traditional Heck cattle with aurochs-like cattle mostly from Southern Europe to achieve a greater resemblance to the aurochs and larger body size. Taurus cattle herds are located in Germany, Denmark, Hungary and Latvia and are used in conservation of natural landscapes and biodiversity. Taurus cattle is a breed that is still being crossbred and selected.

Breeds used for phenotypical reconstruction of B. primigenius

Schematic representation of the establishment of a composite breed

Back bred Aurochs Heck Cattle

Taurus cattle displaying aurochs-like colour dimorphism Source: Wikimedia Commons https://en.wikipedia.org/wiki/heck_cattle

Literature Cis van Vuure: History, Morphology and Ecology of the Aurochs (Bos primigenius). 2002 < http://members.chello.nl/~t.vanvuure/oeros/uk/lutra.pdf > Maas, P.H.J. (2014). Aurochs - Bos primigenius. In: TSEW (2015). The Sixth Extinction Website. <http://www.petermaas.nl/extinct>. Faysal Bibi et al. (2009): The Fossil Record and Evolution of Bovidae: State of the Field; Paleontologia Electronica 12 (3), 2009, http://palaeo-electronica.org Mason, I.L. 1996. A World Dictionary of Livestock Breeds, Types and Varieties. Fourth Edition. C.A.B. International. 273 pp. Renato Bender, Phillip V. Tobias, Nicole Bender (2012). The Savannah Hypotheses: Origin, Reception and Impact on Paleoanthropology. Hist. Phil. Life Sci., 34 (2012). http://hornshire.com/aah/2012%20savannah%20hypotheses%20-%20bender%20et%20tobias.pdf T. Ryan Gregory (2009) Artificial Selection and Domestication: Modern Lessons from Darwin s Enduring Analogy. Evo Edu Outreach (2009) 2:5 27. DOI 10.1007/s12052-008-0114-z