Nigerian Dwarf Goat

A BRIEF HISTORY

The Nigerian Dwarf goat is one of two miniature goat breeds in this country originating in West Africa.There are several recognized breeds of small and dwarf goats in West Africa, and it is from these native African herds that the first "miniature" goats were imported into the United States. Although illegal imports are suspected earlier, the first documented imports arrived in this country in the 1930's and 1940's, for the benefit of several private and public zoos and preserves.
There were two distinct types of small goats imported, one being short, cobby, with short legs, and the other being more slender, with a longer body and proportionally longer legs and a more streamlined appearance. In the earlier years, in fact until about the 1960's and 1970's, little distinction was made between the cobbier type that became the African Pygmy goat, and the more dairy-type goat that became known as the Nigerian Dwarf.
Over the years, selective breeding has resulted in the two distinct breeds that are recognized today in the United States: the African Pygmy Goat and the Nigerian Dwarf.

NIGERIAN DWARF REGISTRIES

There are several registries that maintain herdbooks for the Nigerian Dwarf Goat in North America.
Nigerian Dwarf Goats were first registered by the International Dairy Goat Registry (IDGR) in 1982, followed by the American Goat Society (AGS) in 1984, and the Canadian Goat Society (CGS) in 1985. A privately-owned breed-specific registry called the Nigerian Dwarf Goat Association (NDGA) was formed in 1996.
AGS is an all-breed purebred dairy goat registry, and is the largest register of Nigerian Dwarf Goats, having over over 20,000 Nigerian Dwarfs in its herdbooks as of September 2002. AGS sanctions a large and ever- increasing number of shows across the nation every year, with "hot spots" being Texas, the Northeast, and the Northwest.

BREED STANDARD

The breed standard for the Nigerian Dwarf Goat, as set forth by the American Goat Society, reads in part as follows:

Parts of the body are in balanced proportion; an animal with a disproportionately large head, or one with a relatively large body on short legs is not acceptable. The nose is straight, though there may be a small break or stop at the level of the eyes. The ears are upright.
The coat is soft, with hair that is short to medium length.
Any color or combinations of colors is acceptable.
Breed disqualifications are:
Curly coat
Pendulous ears
Roman nose
bucks over 23.6"
does over 22.4"
evidence of myotonia (this is associated with fainting goats)
The above breed standards are rather brief and don't really describe adequately what one of these little goats actually looks like. Using the AGS scorecard for a basis, this is what a Nigerian Dwarf doe should like:
General appearance will be of femininity and refinement, paired with good condition and strength.
Head will be refined and fairly long, with either straight or slightly dished profile. Eyes are large and soft, ears are medium-length and well set on the head. Face is very animated and expressive. Neck is long and slim, and blends smoothly into tight shoulders with high, sharp withers. Body is long, deepening from heart girth to flank to allow plenty of room for internal organs and kids. Ribs are flat, spaced wide apart, and well-sprung. Topline is level, with only a slight slope from hips to tail. Legs are wide apart, straight, with short, strong pasterns and short feet with tight toes. Skin is soft and pliable, with fine hair.
The crowning glory of any dairy goat is her udder, and the Nigerian Dwarf is no exception. The "perfect" udder is capacious, attached very high and wide, with milkable teats, and a smoothly blended foreudder.

WHAT ARE THEY GOOD FOR?

Milk of Course!
A well-bred, well-fed Nigerian Dwarf doe can produce a surprising amount of sweet milk for her small size. The 2000 All-Time Breed Leader for milk production is WGF Millie *D, who earned that honor by milking 1,545 pounds in a 305-day lactation, with butterfat averaging 5.8%. The All-Time Breed Leader for Butterfat, Gay-Mor Reggae's Nestle Quick 2*D, milked almost as much with 1,499 pounds with an amazing 7.1% butterfat!
Although the records above are of course exceptional, a properly cared for doe should easily produce 1 to 2 quarts a day of high-butterfat (averaging about 6%) , high-protein milk. This makes the breed an excellent choice for the family who can't use the gallon or more a day that a larger dairy goat can provide. Since Nigerian Dwarfs are also year-round breeders, they can also have their place in a commercial dairy as an off-season source of very rich milk. Many breeders enjoy making cheese, fudge, soap and other goat milk products.

Companion Animals

The Nigerian Dwarf goat is extremely popular as a pet and companion animal for people and other animals. It's calm, even temperament and engaging personality combined with its convenient size and flashy colors make it a great companion for all, including children, the handicapped and the elderly. Even breeding bucks can be handled easily and are non-aggressive. Most breeders disbud kids soon after birth, so horns are never a problem. Dwarfs have been used as companion animals for horses, llamas, and even ostriches and emus.

The Thrill of Competition

The best way to show others what you are accomplishing in your breeding program is in the show ring. Classes are usually held for milking does, junior does, bucks and occasionally wethers and dry (not milking) does.
Shows are excellent learning experiences, as unlike with many other animal shows, goat show judges are required to orally give reasons for their placings of a class.
Many goat shows include showmanship classes for children and adults alike. The showmanship classes for the younger set are usually held as "teaching" classes so the children are given a good foundation in the skills of properly showing a goat. Even very small children can usually handle a gentle, well-trained miniature goat, and it can be an excellent introduction for them into showing livestock, and a good confidence builder.

HUSBANDRY AND HEALTH CARE

Nigerian Dwarfs are a hardy, naturally healthy breed. They can thrive in all climates, but do need shelter from extreme conditions and especially from rain, as they hate to get wet. For the beginning goat keeper with just a few goats, a spacious dog house can work just fine. A good quality grass or alfalfa hay and/or free choice pasture should be the mainstay of their diet, with milking does and growing kids being given grain in accordance to their needs.
A vaccination program is usually recommended. Commonly given are Clostridium Types C & D Toxoid ( known as C&D) and tetanus. Ideally, the pregnant doe is given shots 2 to 4 weeks before kidding, in order to boost her antibody levels to better protect her kids at birth. Kids are born with no immunities - their protection comes from the immunities in their dam's first milk, or colostrum. If the dam was vaccinated, then kids should be vaccinated at 6 weeks, and boostered 2 to 4 weeks later. Adult goats should be boostered annually. Other vaccinations may be commonly given in your area - ask your vet for details.
Some parts of the country are deficient in certain minerals, such as selenium, zinc and copper. Deficiencies in these can cause kidding problems, hoof problems, poor hair coat, depressed immune systems, and other problems. Talk to your vet and/or breeders in your area for more information.
Goats are susceptible to parasites, both internal and external. Depending on location and management practices, goats should be wormed as often as every other month. Safeguard and Panacur (fenbendazole) and Ivermectrin products are commonly used and usually effective.

REPRODUCTION

Nigerian Dwarfs are a precocious and prolific breed, reaching sexual maturity as early as 7 weeks of age. For this reason it is very important to have buck and doe kids separated no later than 2 months of age. Bucks can be used for breeding as soon as they are ready and able, but most breeders chose to wait until doelings are at least 7 months of age, and some wait quite a bit longer, depending on the maturity of the doe. Twins are most common, followed by singles and triplets. Quads, quints and even more have occurred. The does are easy kidders and good mothers, seldom needing assistance in either delivering or caring for their babies. Most breeders do prefer to be close at hand during delivery however, just in case. The kids average about 3 pounds at birth and are usually on their feet and nursing within just a few minutes. By the end of the first day they are leaping and jumping and playing tag. This is a good time to make friends with the impressionable youngsters, as some dam-raised babies can be stand-offish if not handled regularly.
Kids can be bottle-raised if the owner wishes, and most breeders wean between 8 and 12 weeks of age. Kids should be exposed to high-quality hay and grain from an early age, and will often begin nibbling alongside their dams by a week or two of age.

COLOR

Color is part of the fun of breeding Dwarf goats. The American Goat Society accepts all possible colors, patterns and combinations. Dwarfs can be dalmation-spotted, pinto-patterned, tri-colored, or just beautiful solid shades of jet black, chocolate brown, gold or white. One of the most attractive patterns is called buckskin: a dark color - usually black or chocolate - covers the head, neck and shoulders in a cape effect, with the remainder of the body being a lighter color and the face and legs sporting coordinating stripes! Although not significant in the show ring or the milk pail, blue eyes are increasingly found in the breed.
See the AGS Chart of Suggested Colors and Patterns for Nigerian Dwarfs.

Pygmy VS Nigerian Dwarf

Pygmy Goats and Nigerian Dwarf goats are very similar with regards to general height ( 17"-19" at the shoulders is desirable ) and rearing techniques. I find that their personalities are very similar. It really depends on how they are raised and heredity with an emphasis on heredity. . . . (example: the friendly mom's tend to have friendlier kids). There are several very important differences. The pygmy is ( and judged in the show ring as ) more of a miniature "meat goat". Some of the qualities viewed to be favorable are: a dished face, short wide muzzle, short muscled neck, short stocky legs, large and deep body becoming deeper as viewed from the side moving from the front legs to the rear legs. The 2 allowed marking patterns to their coats are the Caramel and Agouti. There are shades of both but they are very specific marking patterns. Colors are explained more in depth further down this page. Only a white 'belly band' either complete or partial is allowed. Other random white markings are considered 'miss-marks'. Many have allowable "frosting" to the muzzle and ears.
The Nigerian Dwarf, on the other hand, is more of a miniature "milk goat". Some of it's qualities include : a long slender neck, a thinner body build, slender-more deer-like legs, a longer body--still getting deeper from front to back when viewed from the side, and of course a large balanced udder. There are no restrictions as to color or breed markings.





Gray Agouti Pygmy

Brown Nigerian Dwarf

Breeds of Dairy Goats

There are six types of dairy goats that are recognized by the American Dairy Goat Association. They are Nubians, LaManchas, Alpines, Oberhaslis, Togenburgs, and Saanens.

It is quite certain that the goat was one of the first domesticated animals in Western Asia. He is thought to have descended from the Pasang or Grecian Ibex, a species of wild goat found in Asia Minor, Persia, and other nearby countries. Old Testament verse mentions the goat more than 150 times. Dairy goats are not nearly as numerous as dairy cattle in the Europe, but, for many other countries of the world, they are the leading milk producers because they are well adapted to limited areas and require less specialized feed, most of which can be produced by the small land owner. There are actually more people consuming dairy goat products in the world than those consuming the products from dairy cattle.


Alpine –The French-Alpine is a breed of goat that originated in the Alps. The goats of Alpine type that were brought to the United States from France where they had been selected for much greater uniformity, size, and production than was true of the goats that were taken from Switzerland to France. These are hardy, adaptable animals that thrive in any climate while maintaining good health and excellent production.





LaMancha - Is a type of dairy goat noted for its `lack` of or much reduced external ears. The La Mancha breed is medium in size, and is also noted for a generally calm, quiet and gentle temperament, high butterfat content to the milk and good production.The LaMancha goat was developed in the 1930`s in Oregon by Eula F. Frey when she crossed some short-eared goats of unknown background with her outstanding Swiss and Nubian Bucks. The La Mancha has excellent dairy temperament and is an all-around sturdy animal that can withstand a great deal of hardship and still produce. Through official testing this breed has established itself in milk production with high butterfat.



Anglo-Nubian - This breed originated in England as a cross between the Old English Milch Goat and the Zariby and Nubian bucks imported from India, Russia, and Egypt. They have been exported to most countries from England and are sometimes referred to as "nubians" instead of the proper title of "Anglo-Nubian"The Nubian breed leads the way for the dairy breeds in butterfat production: it produces on average, 5% or more butterfat content.



Oberhasli - Is a dairy goat from the eponymous district of the Canton of Berne (Switzerland). Oberhasli are reddish brown in color with a black dorsal stripe, legs, belly, and face. This breed is a good choice for the person who wants a dairy goat, appreciates its vivid coloration, wants something slightly out of the ordinary, and who wants a quiet and gentle breed.






Saanen - The Saanen breed is white in color and the largest of the dairy breeds. Does typically weigh 150 lb (68 kg) or more, with bucks weighing over 200 lb (91 kg). The Saanen breed also produces the most milk (as a breed- there will be good and poor individuals in any breed) and tends to have a lower butterfat content, about 2.5%-3%. The Sannen temperament is as a rule, calm and mild mannered.


Toggenburg - Is a breed of goat, named after the region in Switzerland where the breed originated, the Toggenburg valley. Toggenburgs are medium in size, moderate in production, and have relatively low butterfat content (2-3%) in their milk. The breed standard is brown with white legs, facial stripes, muzzle, ears, and tail patch. Wattles, small rudimentary nubs of skin located on each side of the neck, are often present in this breed.

More on dairy goats.....

Nubians have very long, floppy ears and they can be any color. They have a convex nose and are one of the larger breeds of goats. Their milk tends to be higher in protein and butter fat than other breeds. They tend to be a little bit more stubborn than other dairy goats and make a distinctive sound. Even Nubian kids sound like they are complaining.






LaManchas have ears that are so small that it looks like they don't have ears and they can also be any color. They have a straight nose and are a small breed. The LaMancha sound is typical of other goats. In our experience, they are more calm and gentle than other breeds. When you own a LaMancha, be prepared to answer the question What did you do to its ears?






Alpines can be almost any color except solid white and light brown with white markings (toggenburg color); their face should be dished or straight. They have erect ears and are a medium-large breed. They are popular with dairies due the amount of milk they produce.

Oberhaslis have very specific color standards. They are a bay color, known as Chamoise, with a black dorsal strip, udder, belly, and black below the knees. They should also have a nearly black head. Another aceptable color would be all black but this is only acceptable for does. They have erect ears and are a medium-small breed.

Toggenburgs also have very specific color requirements. They are light brown and have white ears and lower legs. The side of the tail and two stripes down the face must also be white. They have erect ears and have the smallest height requirments of all the breeds, but most of the toggenburgs I've seen are pretty big. They grow a shaggier coat than other dairy goat breeds. They also are popular with dairies. In our experience, they tend to be a little wilder and more high strung than other breeds.

Saanens are usually pure white. They usually have a large udder capacity and are popular with dairies due to the quantity of milk they produce.

Dairy Goats

Dairy Goat Parts

Goat Milk

Goat milk is used for human consumption. In fact, more people in the world drink goat milk than cow milk, although in the US the opposite is true. Goat milk is similar nutitionally to cow milk, but it contains smaller fat globules and as a consequence it is easier for some people to digest and it does not require homogenization.

Goat milk is also used to feed many other animals. Usually, they are bottle feed, but goats will fairly easily adopt lambs (with a bit of coaxing), and we've even heard of a goat who would jump up on bale of hay to allow a foal (baby horse) to nurse.

Pygmy Coat Colors

CARAMEL PATTERN: Body color any shade of white through brown. Required Markings: Muzzle, crown, eyes and ears accented in white; vertical stripes on center front of socks same color as body; hoofs, sides and rear of socks, dorsal stripe and martingale accented in tones distinctly darker than main body color; Face: Accented in tones distinctly darker than main body color, amount ranges from partial to complete according to age and gender. Optional Markings: white patches or bellybands anywhere on barrel between point of elbow and stifle joint. LlGHT CARAMEL: Pure white to cream with the required darker breed markings. MEDIUM CARAMEL: Apricot to light tan with the required darker breed markings. DARK CARAMEL: Deep tan to deep red with the required darker breed markings. BROWN CARAMEL: Brown to dark brown with the required darker breed markings clearly visible.
White caramels

AGOUTI PATTERN: All body colors are acceptable. The predominant coloration is formed by the intermingling of lighter and darker hairs producing an appearance ranging from highly grizzled to nearly solid. Required Markings: Muzzle, crown, eyes and ears accented in white; hoofs, socks, face, and forehead distinctly darker than main body color.
Optional Markings: White patches or bellybands anywhere on barrel between point of elbow and stifle joint; dorsal stripe darker than the main body color; varying degrees of grizzling on areas of the body, such as the chest, shoulders, mane, and tail.
LIGHT GRAY AGOUTI: Black and white hairs intermingled with slightly more white hairs to give a light gray color, making the topcoat appear to be silver/pewter in shade MEDIUM GRAY AGOUTI: Black and white hairs intermingled in equal amounts giving the topcoat a blacker appearance close to an equal amount of salt and pepper DARK GRAY AGOUTI: Darker gray appearance with fewer white hairs, but still a salt and pepper appearance that is pronounced BLACK AGOUTI: Black mainly with only occasional intermingled white hairs LIGHT BROWN AGOUTI: Brown and white hairs intermingled making the topcoat appear a light cinnamon and sugar shade.MEDIUM BROWN AGOUTI: Brown and white hairs intermingle in equal amounts giving the topcoat a browner cinnamon and sugar appearance. DARK BROWN AGOUTI: Brown hairs intermingled with fewer white hairs, but still a cinnamon and sugar appearance that is a pronounced darker brown BROWN AGOUTI: Brown mainly with only occasional intermingled white hairs, darker brown mask and solid stockings clearly visible BLACK: All body hairs black in color. Required Markings: Muzzle, crown, eyes and ears accented in white. Optional Markings: White patches or bellybands anywhere on barrel between point of elbow and stifle joint; tail accented in white. SOLID BLACK: All body hairs black in color. Required Markings: none. Optional Markings: none

light and medium brown agouti

The Qualities of Pygmy Goat Milk

Written by: Maxine Kinne
Hopefully, you will one day decide to take up the pleasurable task of milking. When you do, there are some fundamentals to understand about the quality of Pygmy goat milk and what makes it different than other types of goat milk.
Milk traditionally weighs 8.6 lbs per gallon. Pygmy goat milk only weighs 8 lbs per gallon, because it has a significantly higher butterfat content that the milk of most other breeds. (Nubians come the closest to the Pygmy's fat content.)
Why is high butterfat content a good quality in milk? For one thing, it helps the milk resist off flavors due to a doe's diet. Strong flavored plants may impart their flavor to milk, especially when they are consumed within a few hours of milking. Higher fat content also extends the shelf life of milk. With very good milking technique and milk handling, Pygmy milk can retain its excellent flavor for about two weeks.
Fat, of course, is what gives many foods a desirable taste because it is rich and sweet. When I dried off my does after two solid years of milking and bought 3.8% cow milk from the store, I had to spike it with whipping cream to combat the flat, watery taste! Many health-conscious people now avoid fat. But since Pygmy milk separates fairly readily, unlike the naturally homogenized nature of dairy goat milk, most of it can be skimmed off.
The NPGA membership brochure lists the butterfat percentage at 6% to 9%. My milkers ranged from 4.5% to 11.75%; and average was about 6%. Fat percentages are tied to the total quantity of milk a doe gives. Does give about the same amount of butterfat all the time. Less milk is produced in early and late lactation, but the butterfat percentage remains stable. Thus, its percentage of the total yield is higher when the doe gives less milk. Butterfat is given at the end of the milking process, so it is very important to milk each doe out completely.
A study comparing minerals in the milk of dairy goats and Pygmies (West African Dwarf Goats) in The Third International Conference on Goat Production and Disease, 1979, found that Pygmy milk tested 65% higher in calcium, 19% higher in phosphorus, 75% higher in potassium, 26% higher in iron, and 10% higher in copper. It was 21% lower in sodium, 13% lower in magnesium and 40% lower in chlorine. Alpine and Saanen results were averaged for the dairy milks. This study supports another one cited by Alice Hall. These percentages mean that Pygmy goat milk is higher in things that are good for you and lower in things that are not.
As a meat breed, the Pygmy is more heavily boned and grows rapidly. They need a good ration of calcium and phosphorus, so heavy-boned mothers have sufficient calcium availability to kids. High milk fat content serves as energy for for fast growth.
Some people who have bottle fed many different kinds of milk replacer complain about unthrifty, slow-growing, light-boned kids. This has not been true for the many kids I have raised on Pygmy milk or 3.8% cow milk from the grocery store. I believe that critical management factors, like under- and overfeeding milk replacer, internal parasites and general sanitation, deserves a greater amount of blame than their choice of replacer.

Why Milk the Pygmy Goat?

Jennifer gave 1/2 gal/day (4.4#)at the peak of her lactation, and earned a 1984 production award.
In the early 1980s, my dairy goat friends talked me into milking my Pygmies. I also wanted to help establish quantity and quality standards for Pygmies. Once we tasted their delicious milk - it tastes just like milk, only better - we were hooked. I also found the milking routine soothing and even hypnotic. I bought a milking stanchion that was a too small for a friend's dairy goats, then I needed a doe in milk. Janny was a 2-year-old whose twins had just sold, and we learned the milking routine together. A does that has nursed her kids doesn't want her udder messed with by anyone. However, if you handle the udder from a young age and develop a close bond with the doe, she may be more forgiving than Janny or Jennifer were the first year I milked them.
The best way to start milking is to take the kids as they are born and remove them from the doe's presence entirely. You milk her just after kidding, so she accepts you as her kid. You've got bottle babies to raise, but that creates a special bond, as you replace their natural mother.
Excess colostrum beyond the kids' requirements can be frozen for future use. Colostrum from older does is higher in antibodies than from young does because older does have been exposed to more diseases.
As a child, my husband tasted some goat milk gone bad and was unimpressed with my venture. I didn't give him a chance to turn it down. Since his first taste, he has been a staunch proponent of the delicate taste and rich quality of Pygmy goat milk. The milk is very rich and sweet due to its high butterfat content.
Milking was a great skill to add to my repertoire. It is excellent relaxation therapy that helps you develop an extra special bond of love and trust with milking does. Getting to the barn to milk twice daily enhances your management - you notice details and tend to keep up with things a little better.
It is very gratifying to produce a high-quality product. I once entered my Pygmy milk in a dairy goat club milk products contest and won first place in the fluid milk division. My prize, appropriately enough, was miniature tin of Bag Balm.
For two years I was on DHI test (Dairy Herd Improvement) to help establish some official records for pygmies. DHI testing shows that pygmies are useful in the milk parlor in addition to their other qualities.
DHI expects goats and cows to milk for a 305-day (10-month) lactation, although their records may be shorter if the animal is dried off for any reason. Official records do not extend beyond 305 days, but extra production is credited toward lifetime production. An impartial tester weighs and takes a sample of each doe's milk, then sends it to a lab for several tests, such as solid non-fat protein percentage, fat content and a California Mastitis Test result. Other optional tests are available. Each month the owner receives a new DHIA report and paperwork for the next test.
Of the DHI options available, SCC is a very useful indicator of udder health, milk quality and sanitation. Management, in other words. Somatic cells are leukocytes (white blood cells), and a beyond certain number in the milk signals mastitis. The two different types of SCC counters (Coulter and Fossomatic) give quite different results because the Fossomatic mistakenly counts extra-cellular debris as leukocytes.
To be on DHI, I had to join an association that recognizes production testing for Pygmies. I double-registered the milkers and the sires of the offspring with AGS. Perhaps NPGA will one day have a testing program.
Pygmy milk and fat quantity requirements are one-third of the dairy goat standards. To earn a milking award, a Pygmy under two years old must produce 500# of milk or close to 18# of butterfat. The requirement rises slightly each year until it peaks at 5 years old: 582# of milk or 20# of fat.
Both does I tested the first year earned star milking awards. Dolly, a first freshener gave an average of just over 2 gallons of milk per week for ten months. She also produced 50# of fat in that lactation, which is more than two-and-a-half times that expected of a mature doe. Jennifer did respectably well as a second freshener, giving an average of 1.8 gallons a week.
A general rule of thumb is that an average Pygmy doe should give about ½ gallon per day at the peak of her lactation. A gallon of average fat milk weighs 8.6#, but Pygmy milk weighs 8 lbs. because fat is lighter.
For a steady supply of milk throughout the year, two does can be bred at different times. Many people, though, prefer to have two months off during the does' dry period before their next kidding.
When a doe freshens (comes into milk at kidding), she doesn't produce as much as she will later because her kids don't need much yet. Production increases for two months or so and slowly declines. This lactation curve matches kid growth and decreasing reliance on milk. Kids are nutritionally self-sufficient at about 10 weeks old. In other words, most does produce milk at the rate the kids need it. Lactations persistence means that a good milker's production stays at its peak for a long time.
Milking takes dedication. It is very important to milk twice daily at 12-hour intervals to maintain production. When milk is in the udder for 18 hours, the milk producing cells (alveoli) begin to shut down due to pressure. Once this production is lost in a lactation, it cannot be regained. As we all know, goats definitely like their routine, so regular milking times keep them happy.
Man can be infected with a few diseases passed in the milk of infected animals. Tests are available to identify disease carriers. Chances are that if a doe had any of them, you wouldn't want her on your farm, much less to drink her milk. Diseases most often mentioned as zoonoses are brucellosis, tuberculosis and Q-fever, though there are others. They are uncommon, but you should be aware of them.
Anything you put into your milker via injection or orally (eating, drinking, drenching) will come out in her milk. Each drug approved for use in goats has a withholding time on the label. Your veterinarian can advise you about these and extra-label drugs. Withholding means throwing out the milk of treated does (and observing slaughter times) for a certain number of days, weeks or months following drug use. Such milk may usually be frozen and for the next batch of kids.
It is doubtful that you will milk enough Pygmy does to have excess for sale. Laws regarding fluid milk sales vary by state. In some states it is illegal to sell raw (unpasteurized) milk for human consumption without a Grade A dairy license. Legal liabilities extend to raw milk sales or sometimes even in serving it in your own home. Mishaps involving contaminated raw milk make the news from time to time, and those cases emphasize this point. You are legally liable if your goat milk is contaminated and makes anyone sick. Most states heavily regulate the sale of milk, with sale defined as "dispensing, giving, delivery, serving or any other supplying..." Look into your state laws before serving raw milk.
There are many good reasons to milk a Pygmy goat or two. It is very enjoyable. After putting so much time, effort and care into your goats, they have few opportunities as unique as this to give you in return.
Good Milking ProceduresMaxine Kinne
Dairy guidelines proven to maintain udder integrity and milk quality start with establishing a routine milking order. Does get used to a certain order and production suffers if even the smallest things change.
Milking Order
1. Older does
2. First fresheners
3. Does with high somatic cell counts (if known)
4. Sick does without active mastitis
5. Does with mastitis
Does with mastitis are always milked last to prevent spreading bacteria to the next udder. In my experience, mastitis is rare in pygmies because they have relatively good udder attachments and small teat orifices. Sanitation management is very important in avoiding mastitis.
Somatic cells fight bacteria inside the udder that cause mastitis. There are always a certain number of somatic cells in the milk, but there are very many during any degree of mastitis. Lab testing can detect somatic cells in milk. On the farm, the California Mastitis Test is very reliable and can be used when you milk.
Does milk more willingly if they are trained to the stanchion first. Get them used to the stanchion by feeding them there.
Wash and dry your hands before you are going to milk. It is unnecessary to wash the entire udder before milking, but washing and drying each teat helps prevent mastitis.Wash each teat with a separate paper towel and discard. Dry each teat with a separate paper towel and discard. Never reuse a towel. (Teats can also be sanitized by predipping and drying each one with a separate paper towel instead of washing.)
Udder preparation relaxes the doe so she will let her milk down. Milk let-down is due to the oxytocin response. When a doe is excited or upset, she releases adrenaline which cancels oxytocin, and it is impossible for her to give milk until the adrenaline is used up. When it is, it takes another few minutes of preparation to elicit another oxytocin response. Loud, unusual noise or actions evoke fright and adrenaline, so a calm routine is always required. Massage the udder for 30 seconds or so after washing to feel udder texture. Lumps, hot spots or injuries may be found at this time.
Examine the few streams of milk from each teat for abnormalities. The dark, perforated insert in a strip cup lets milk though while retaining clots or strings. The Pygmy doe usually gives a small stream of milk, and a clean, dark cloth or a dark-colored dish works well. Check the milk from each teat at each milking. The first few streams of milk contain the most bacteria, so it is a good idea to discard this anyway.
Try to completely empty the udder within five minutes after udder preparation - oxytocin response is the greatest during those five minutes. Grasp the base of each teat just below the udder floor with your thumb, index and middle fingers. Delicate tissue may be injured if you squeeze any part of the udder. Milk in the teat cistern is trapped when you squeeze the base of the teat with your thumb and index finger. Expel the milk using all 3 fingers, your middle finger following the index finger in gentle squeezing. Pulling on the teat or sliding your fingers down it may injure udder or teat tissue. Alternate the streams, left and right. Developing coordination takes time - don't give up. Your hands may tire and cramp, but it gets easier as you build up milking muscles and coordination. At the end of the milking, massage the udder to release the last of the milk and milk that out.
After milking, dip the entire length of each teat with a good teat dip to kill bacteria at the orifice and help close the orifices. Teat dipping is very important. The dairyman I heard at at a seminar said, "If I had only one cow, and she had only one teat, I would dip that teat after each and every milking!"
If you have to reconstitute a dip, it is important to make it the proper strength. Strong solutions can damage the skin, and it is ineffective if it is too weak. Follow the label directions. Ideally, fresh teat dip is used on each teat at each milking. Contaminated teat dip is no longer effective and may harbor bacteria. I use a small glass jar with a 1"-deep plastic lid. I snap off the lid, fill it about 3/4 full, coat each teat, and discard the used portion. Besides the quality of the dip, good coverages is the most important part of dipping. A drop of teat dip should be visible on the end of each teat. (Fight-Bac teat spray also works very well instead of dipping.) After dipping or spraying, keep the doe standing for 15 minutes or so, maybe with fresh hay in the manger. If she lays down, the teat dip won't work as well as it should.
Milk is a delicate product and should be filtered immediately after milking to remove hair and other contaminants. Then it should be cooled as quickly as possible. Milk filters are available at feed stores and catalog suppliers. Strain into a glass jar and refrigerate immediately.
Milking sounds complex, but it is very easy. Having the right equipment is a big help. Good habits in procedure and technique are important in the long run, both for your enjoyment of the milking process and maintaining the health of your does.
Equipment for Milking the Pygmy Doe
Having the right tools of the trade make milking more pleasurable and preserve milk quality and udder health. Milking equipment must be kept very clean between uses, and it is best to let it air dry after it is washed.

Equipment List
Milking stanchion (or facsimile)
Stainless steel container with smooth finish
Small water bucket for udder wash
Paper towels to wash and dry teatsStrip cup (or facsimile)
Teat dip
Milk filters & filtering equipment
Glass storage jars

Does are usually immobilized for milking. Clipping her collar to the wall can work, but a stanchion elevates her and makes make it more convenient for you. Reserve a separate place for milking, away from distractions that may bother either of you. You usually sit at the doe's side and face her rear, although some people prefer to bend over the doe to milk from behind. Sitting beside her gives you a lot of physical control. If she acts up, press your head and shoulder into her flank to keep her slightly off balance against the wall.
Warm water for washing teats can be hauled to the barn in a small bucket. In the winter I use a large bucket filled with 140o F water, and refill the water trough after milking. The goats really appreciate a nice, warm drink when it's cold. Water intake is important to milk production, so providing plenty of it helps maintain milk production. Water is the main component of milk.
A smooth-finish, stainless steel mixing bowl works very well for Pygmy goats. A one-gallon bowl fits under larger does, but a shorter bowl may be needed for shorter does. Slant-sided bowls are very unstable for milking. Bowls with vertical sides are hard to find, however a mirror-finish saucepan without the handle can work well. The height of the bowl should leave room for your hands and arms between it and the doe's undercarriage.
After spending the time and effort to milk, treat the milk right to preserve its quality. Warm milk is an excellent medium for bacterial growth, so it must be strained through a filter and refrigerated immediately. Milk filters are sold by the box. I have folded 61/2" filters into cone shapes to fit a funnel. I have also used a Busy Liz food strainer and 21/4" filters to fit (Caprine Supply catalog). Cheesecloth and other fabrics are too porous to for filtering milk.
Immediately after milking, pour the milk through the filter into a glass container and refrigerate. Glass jars are perfect, nonporous milk storage containers. Bacteria accumulate in the pores of plastic and make it smell bad. The small pores in plastic also makes it impossible to disinfect. If you have a milk jug manufacturer in your area, you may be able to buy gallon or half-gallon jugs and lids. These are made for single use, but I have bleached them between uses and got quite a lot more mileage out of them. At the time I used these, they cost about $0.16 each.
As soon as the milk is filtered and refrigerated, wash all the milking equipment and let it air dry upside-down in a dish drainer. Dish towels contaminate the equipment. The fussier you are about maintaining equipment in top shape, the better your milk will taste.

Color and pattern genetics in the Pygmy

Color variation is part of the fascination of raising Pygmy goats. These colors all result from gene interactions, and a look at the genes present in the breed can reveal the colors and patterns that are possible in Pygmy goats.
A few concepts of color are important to grasp before any discussion of color genetics can be meaningful. Some of these concern the mechanisms by which color is produced. Color is due to melanin deposits in the hair. Each goat has a genetic directive for the placement of certain colors of melanin in the hair. White areas on goats, or entirely white goats, result from this directive being superseded by genetic directives that prevent melanin from being placed in the hair. This occurs through a wide range of mechanisms; the details are less important than the general fact. The importance of this is that white areas need to be viewed as covering up colored areas, much like white paint. The tendency of most people is to see color as added to a white background, when in reality the opposite is the case.
Melanin is deposited in the pigment granules responsible for color; these come in two basic types - eumelanin and pheomelanin. Eumelanin is usually black, but sometimes brown, and it is the pigment responsible for black and brown areas on goats, or rarely for dusky blue color. Usually the color choices with this pigment are either/or; the goat can either form black or brown, but not both. Pheomelanin pigment is responsible for red, tan, or yellow pigment. That sounds easy enough, but if pheomelanin is very dark it can be close in shade to the brown eumelanin pigment. Pheomelanin usually has a redder cast to it, though, and the difference between the two types is generally fairly obvious. "Red" pigment can also be pale enough to be cream or nearly white, and this can cause confusion in some instances.
Dominant and Recessive Genes
Finally, we get to some real genetics. Genetics can be confusing and is not always intuitive. It is important to remember that genes occur in pairs. Each parent transmits only one of its pair to its offspring. This sounds easy , but it is critical to remember that each goat only has two genes at each locus, or "address." So, while a goat has four grandparents, it can only have two genes at a color locus from these four, which means that someone got left out. We know that the sire and dam each must contribute one, but grandparents get much iffier.
An example may help. If a buck has a caramel father and an agouti mother, he may well be agouti, not caramel. If a doe has a caramel father and a black mother, she may be black. If we cross the black doe to the agouti buck, it is impossible to get a caramel kid, even though two grandparents are caramel. What happened in this case is that the caramel gene simply did not make it through to the parental generation, so it is not present to be donated to the grandkids.
Genes are also said to be dominant or recessive. This only describes their interactions with one another in pairs and says nothing about their prevalence in a population. A dominant gene can be rare, and a recessive gene can be common. The trick here is that a dominant, when paired with a recessive, masks the recessive. So, an animal with a dominant trait can have two doses of the dominant gene, or one dose of the dominant and one dose of the recessive, and the differences cannot be told by only looking. A goat with a recessive trait, in contrast, must have two doses of the recessive gene. The real importance of this is that recessive traits can pop up out of nowhere, since they can be masked by dominant genes. A dominant trait cannot pop up out of nowhere, since it cannot be masked. A good example of this is in Angus cattle. Every now and again, red calves occur in black herds, as red is recessive to black. The opposite situation - black calves in red herds - never occurs, for the red recessive cannot hide black.
Basic Colors
Three basic colors occur in Pygmy goats, and two patterns can be superimposed over them. The basic colors are black, dark brown/mahogany, and medium brown. As solid colors, these are fairly rare in the breed, but they are the underlying base for the common caramel and agouti patterns. In the Pygmy, black is dominant to the recessive medium brown, so medium brown can pop out as a surprise. Dark brown is dominant to black, and this seems to be the more common type of brown in the Pygmy. This means is that it usually takes a dark brown (or dark brown + pattern) goat to produce a dark brown (or dark brown + pattern) kid. The recessive medium brown is an exception to this rule, and will occasionally pop out as a surprise from black to black matings.

Color Table

There are practical implications of these genetic mechanisms for goat breeders. Black is the usual color, and most goats have two doses of black. As a result, most black-to-black matings produce black offspring. Dark brown is dominant to black, so it is impossible to tell if a dark brown goat has one dose or two doses of the dark brown gene. If it has two doses, it can only produce dark brown kids, regardless of the mate. If it has one dose of dark brown and one dose of black, it will produce about 50% black kids and 50% dark brown kids following matings to black goats. If two brown goats each have one black and one dark brown gene, then the resulting kid crops should be about 75% dark brown and 25% black. This works because each gene is contributed at random to the kids.
Medium brown is more rare than dark brown, and it is recessive to black. Medium brown goats, therefore, must have two medium brown genes. If medium brown goats are mated to black goats, the results depend on the black parent. If the black has two black genes, all the kids will have one black and one medium brown gene, and they will be black. If medium brown goats are mated to black goats with a medium brown parent, it is certain that they have one dose of the medium brown gene and can contribute that to half of their kids. As a result, they can be mated to medium brown goats to produce 50% medium brown and 50% black kids. The fun begins from an understanding that the recessive colors can lurk undetected for generations. So, if a black goat has an unseen medium brown gene and is only mated to black goats without a medium brown gene, the result will always be black kids. If one mate carries the medium brown, though, a medium brown kid can occur as a surprise.
So far, the basic lesson is that you have to breed to a dark brown to get one. Medium brown can occur as a surprise, but the only consistent way to produce them is to mate medium brown to medium brown, or medium brown to a black goat with a medium brown parent.
Agouti Pattern
Basic colors are usually "dressed up" in the Pygmy, and so are fairly rare in their basic form. One pattern that can be superimposed over the basic color is the agouti pattern. This is the familiar mixture of white hairs into the base coat color. Since this is superimposed over the basic color, the result is six choices for color: black, dark brown, medium brown, black agouti, dark brown agouti, and medium brown agouti. The agouti pattern varies in intensity, so that shades within the pattern are variable with some lighter and others darker. The agouti pattern is dominant to its absence, so agoutis are expected to sometimes produce solids, but two solids mated together should never produce an agouti. In the few instances in which this happens a further investigation usually reveals that one parent is a very dark agouti.
PIC 1
PIC 2
The basic pattern with the agouti is that some agoutis have two doses of the agouti gene. These mated to solid colored goats (usually black, but maybe dark or medium brown) result in all agoutis (on a black background, dark brown background, or medium brown background as determined by the previous examples). If an agouti has one agouti gene and one solid gene (as the kids from the previous matings would have to have, since they have a solid parent), the result of matings to solid colored goats would be 50% agoutis and 50% solids. Solid-to-solid matings cannot produce agoutis, since the gene is dominant and expresses itself even if present as a single copy of the gene.
Caramel Pattern
Caramel is the second major pattern. In genetic parlance this is called badgerface after a similar sheep color pattern, but the name caramel is used in pygmies, so let's stick with it. The caramel pattern consists of a tan or cream body with dark belly, dark legs, dark marks on the face and a dark stripe down the back. The caramel pattern can be superimposed over the basic colors, and the result is caramels with black marks, dark brown marks, or medium brown marks. Since the main feature of caramels is body color, they are frequently referred to by body color rather than the color of the belly, legs, face markings, and back stripe. These dark areas reveal the underlying color of the goat, though, and their color is therefore important. Caramel is dominant to the solid colors and is really superimposed over them. As a result, two caramels mated together can sometimes produce agoutis and solids. Two solids mated together cannot produce caramels. Neither can two agoutis mated together, or an agouti and a solid mated together. So, the basic rule is that you need to mate to a caramel to produce one. This is the key concept of a dominant gene.
Table 1 should help in understanding that dominant genes cover up recessive genes. If this theory is true, then individual caramel goats mated to solid goats should produce kid crops of three different sorts:
* All caramel
* 50% caramel and 50% agouti, no solid
* 50% caramel and 50% solid, no agouti
Caramel goats can have one or two doses of the caramel gene. As a result, they can produce various offspring. One theory is that each goat can only have two choices of caramel, agouti, or solid. This means that the colors in Table 1 could have these genetic makeups:

Table 1 P

The key point is that no one goat should provide all three choices to a kid crop. The question is, has anyone had a caramel goat that when mated to solid mates produced all three patterns? I have not come across one, but such a goat would be important in proving or disproving the above theory.

Table 2 P

The depth of the caramel color is variable, from nearly white to nearly brown, with tans and reds in between. The shade of color is controlled in a more complicated fashion than the basic pattern, and there are many questions about this. Most of these differences seem to be due to multiple genes, and the best recommendation is simply to mate the preferred shade back to itself (red x red, tan x tan). Then keep your fingers crossed. I have had lousy luck in my own herd in predicting shade. I have a passion for rich gold and always seem to get white, cream, or dark red, even out of a magnificent gold buck.
Spotting
Some Pygmy goats are white spotted. These patterns have never been adequately studied. The belting pattern, which is variable in expression, is dominant. Its expression is so variable that it is a difficult pattern to predict with any certainty. I suspect that there are other spotting patterns besides belting and that these are independent genetically, but none of these has been documented as to genetic mode of inheritance.
Frosting
The typical frosting on the Pygmy ears and muzzles is due to a dominant gene. This is so common in the breed that I have never seen a Pygmy without frosting. The amount varies, but at least some is usually there.
All the Possibilities
With all of these components it is possible to make a variety of color.

Table 3 P

It can be seen from the table that there are basically nine colors within the Pygmy goat breed. All of these can occur because the genetic machinery exists within the breed to form these colors, with or without added white spotting. Except for the solid dark brown and solid medium brown, all are accepted within the NPGA Breed Standard. Even though solid browns are not, they are very much possible because the genetic machinery is there.
These nine combinations are complicated by the addition of white spotting in some goats, which results in eighteen basic types, one spotted and one nonspotted. The important basic concept is that each goat is the combination of different factors. These are under separate control, so the choices can be viewed as a cafeteria with various choices for courses. The choices are:

Table 4 P

We can only pick one color trait from each group. As an example, a caramel goat with medium brown trim and a spot on the side is A3, B2, C2, D2, E1. Similar logic can show the choices for all the other colors. * I have never seen this in pygmies, so it may not be a choice.
A few other patterns probably pop up occasionally in the Pygmy. This is expected because caramels can mask a number of other patterns. These are probably culled because they are not recognized in the standard. They are just as much a part of the breed as similar recessive colors are in other breeds of livestock. I have seen one "reverse caramel" Pygmy, with a black body and light belly and legs. The pattern would be easily masked and pop out unexpectedly on occasion. This pattern and others reflect the Pygmy's African origin, where color is unimportant and survival and production are.
It is interesting to see how the color factors that are present in the Pygmy breed can interact to give various final colors. Colors add even more interest to an already interesting breed, and all of them should be appreciated by devoted followers of the breed.
Dr. Sponenberg is a Professor of genetics and pathobiology at Virginia Polytechnic Institute and State University and has worked to conserve rare American breeds for thirty years. Internationally known for his work in color genetics, he has authored numerous books and publications. His experience includes that of both an academic and an animal breeder, allowing him to utilize practical as well as theoretical aspects of breed conservation.

More on Genetics of Goat Color

D. Phillip Sponenberg, DVM, PhD

Introduction
General Considerations of Pigmentation
Loci Controlling Color
Agouti
Extension
Brown
Moonspots
Patterns of White Spotting
Angora White
Spotting
Belt
Roan
Flowery
Goulet
Algarve
Barbari
Ticking
Introduction
Goat color variations have only recently been studied, in contrast to the very early work on sheep color variation. While homologies to sheep color are frequently present, goat color inheritance differs from sheep in a number of important regards. The study of goat color has usually been tied to the need for production of certain colors (usually white) in certain breeds of goats such as Angora or cashmere bearing goats, as well as many breeds which have breed-specific colors or patterns. To best understand color genetics, it is helpful to focus first on the general biology of goat color, and then to delve into the genetic details. A clear understanding of the basic genetic phenomena is essential for those interested in the production of certain specific colors. A complete historic review of the world literature dealing with the loci and alleles affecting color in goats has recently been completed by the Committee on Genetic Nomenclature of Sheep and Goats (COGNOSAG). COGNOSAG recommendations on nomenclature are used in this chapter, which will focus on the action and interaction of the various loci that control pigmentation in goatsm and the biological processes that are involved.

General Considerations of Goat Pigmentation
Color in mammals depends on the presence of melanins in skin and hair (Searle, 1968). Melanins reside in cellular organelles called melanosomes, which are produced in melanocytes. Melanosomes reside in the melanocyte cytoplasm, and are also deposited into epidermis and hair by a process of exocytosis which transfers the melanosomes from dendritic processes of the melanocytes into these other (generally epidermal) cells. Melanocytes are specialized cells that migrate from the neural crest during embryological development, and melanocytes have close embryological ties to cells of the neurologic system. Pigmentation of skin and hair depends on melanocytes being present, and on their relative level of melanogenic and exocytotic activity. The two main mechanisms by which pigmentation is reduced or absent are absence or relative inactivity of melanocytes. Decrease or elimination of pigment by either of these basic mechanisms can be regional, or can involve entire haircoat.
Melanins are large polymers that are formed of varying amounts of tyrosine and cysteine (Prota, 1992; Jackson, 1994). Melanins occur as two types: eumelanin and phaeomelanin. Eumelanin is generally black or chocolate brown, and is predominantly made of tyrosine. Phaeomelanin is generally reddish brown or yellowish tan, and has varying amounts of cysteine in its polymers. The two types of melanin are important visually and genetically, and are usually considered as two discrete classes of pigment. Some overlap occurs between the two, but this overlap is usually trivial to an understanding of the genetic control of color in goats. Overlap can also occur visually, with very dark phaeomelanin being similar to lighter forms of eumelanin, but usually the two classes are distinct (Sponenberg et al. 1988, Sumner et al., 1994). The differences between eumelanin and phaeomelanin have been demonstrated by electron
magnetic resonance (Vsesolodov et al. 1981) as well as by analysis of chemical degradation products (Sponenberg et al., 1988). Other techniques to demonstrate the difference between the two melanins include histologic evaluation of melanosomes (Alexieva et al. ,1985; Renieri et al., 1995), and the use of color charts (Sumner et al., 1994).
Melanins are formed from tyrosine and cysteine by a process of catalysis, and the main enzyme involved in this process is tyrosinase (Jackson,1994). Tyrosinase activity is essential for melanin synthesis. Two other proteins, tyrosinase-related protein 1 (TRP-1) and tyrosinase-related protein 2 (TRP-2), are also important for melanogenesis. The exact function of TRP -1 is uncertain, although it is well documented to have an important role in the synthesis of eumelanin, and has little or no role in the synthesis of phaeomelanin. TRP-2 has dopachrome tautomerase activity.
Melanocytes are capable of forming both eumelanin and phaeomelanin, although they usually only produce one or the other at any one time. The dedication of melanocytes for eumelanin production depends on the presence of alpha melanocyte stimulating hormone (aMSH) which is a pituitary hormone (Jackson, 1994). Melanocytes have surface receptors that bind this hormone. When aMSH binds to the surface receptors the result is a cascade of events that activates adenylate cyclase. This activation in turn stimulates the melanocyte to produce eumelanin. In the absence of this signal, which is dependent on aMSH as well as the surface receptors, melanocytes produce phaeomelanin. The switch between eumelanogenesis and phaeomelanogensis depends on the function of the receptors to aMSH.
The control of melanocyte function is intricate, and many loci have mutants which affect different components of the melanogenic control mechanisms (Jackson, 1994). Some loci affect cell differentiation or migration from the neural crest. Other loci affect the morphology of the melanocyte or its ability to deposit melanosomes in hair and epidermis. Other loci directly affect the enzymes and related proteins that are responsible for melanogenesis. A few loci have mutants that affect the interaction of aMSH with the target melanocytes. All of these loci interact to give the final color phenotype of goats.
White has been an important color in many breeds of goats, especially those for fiber production. The expression of whiteness in mammalian hair can come about through a variety of different genetic mechanisms (Searle, 1968). Some of these mechanisms appear to be widespread throughout domesticated goat populations, but this is no guarantee that other mechanisms are not important to some specific populations.
One general mechanism for whiteness in mammals is due to white spotting. White spotting in mammals occurs in regions where skin or hair follicles lack melanocytes, since these regions are incapable for forming melainins. Absence of melanocytes can occur from a variety of mechanisms, including melanoblasts failing to differentiate from the neural crest, or failing to migrate to skin regions, or failing to survive once migration has taken place (Searle, 1968). The variety of mechanisms responsible for white spotting are affected by several different loci, each of which controls a different aspect of the process of populating skin and hair follicles with melanocytes.
A second general mechanism for whiteness is achieved through dilution of pigment, which is
accomplished by decreased effectiveness of melanin production by melanocytes. Melanocytes are present, but are ineffective at forming melanins, placing them in the wool and hair fibres, or both. Their ineffectiveness at melanosome production or transfer results in unpigmented or weakly pigmented hair and skin even though melanocytes are present. The effects of both white spotting and dilution are important in goats, and separating these two concepts is important to understanding pigmentation in goats.
The wild type color in goats must be considered as the standard type if the genetics of color is to make sense. The wild ancestor of the domesticated goats is most likely the bezoar of Anatolia and the Middle East. The color of the various species of wild goats has subtle differences one from the other, but similarities are striking (Novak, 1991). Wild goats are generally tan bodied (phaeomelanin) with pale bellies (dilute phaeomelanin, rather than nonpigmented white). Black areas (eumelanin) are present as a dorsal stripe and striping patterns on the heads and legs. Some sexual dimorphism is present, with bucks darker than does due to more extensive eumelanic patterns on the head and shoulders. The eumelanic areas, especially on males, become more extensive with age, and mature dominant bucks are therefore darker than are younger ones. The interplay of eumelanic areas with phaeomelanic areas is critical to the
overall appearance of the color pattern, as is the presence of intense dark phaeomelanic areas along with dilute, pale phaeomelanic areas.

Factors Affecting Color of Goats
The Agouti Locus
One of the main determiners of goat color is the Agouti locus. This locus has many allelic variants, making it one of the more complex loci governing goat color. Variation at the Agouti locus is widespread among goat breeds, such that Agouti locus phenomena account for the majority of color variants in most goat breeds. The concept of multiple alleles is essential to understanding the Agouti locus, and was first documented in sheep for patterns closely related to those of goats. Roberts (1928) first noted that multiple patterns (his badgerface, reversed badgerface, black, and white) segregated as alleles at a single locus. Rendel (1957) first suggested the Agouti locus as the site for these patterns. The four patterns suggested by Roberts were confirmed as multiple alleles at a single locus by Brooker and Dolling
(1969a). Alleles documented by these researchers, as well as several newly described ones, were ascribed by Adalsteinsson (1970) to the Agouti locus, and it is this work which put the entire study of sheep color on a sound comparative basis since this work was widely used as a base by other researchers. Demonstrating the character of the Agouti locus patterns in sheep was the breakthrough that finally provided for more rapid advances in the understanding of the genetic control of sheep color, because it put the science on a sound foundation of comparative genetics. Following this, sheep phenomena could be more readily coupled to homologous phenomena of other species including goats.
The manifestations of the Agouti locus in goats are relatively complex, although studies in mice indicate that the locus itself is relatively simple in its biology (Jackson, 1994). Many different alleles occur within the Agouti locus of goats, however all of the caprine Agouti locus patterns are characterized by symmetrical arrangements of eumelanic and phaeomelanic areas. In goats the eumelanic areas are usually black, although can be brown when the genotype at other loci modifies black to brown.

Mechanism of Action
The Agouti locus is responsible for the formation of a protein that acts to nullify the action of aMSH on melanocytes (Jackson, 1994). In regions in which this protein is present the melanocytes fail to respond to aMSH, and therefore form phaeomelanin and not eumelanin. Melanocytes in regions lacking this protein have full capability for stimulation by aMSH, and therefore form eumelanin. (Jackson, 1994). In regions with a pulsatile formation of the agouti protein the result is banded hairs. These are important in mouse phenotypes, and while present in some goat patterns they are less important as a cause of color variation than they are in mice.
In mice the Agouti locus has been extensively studied, and implications of homology suggest that its biology is similar in other species including goats. The series of murine Agouti alleles is a consistent array as it progresses from the most recessive to the most dominant allele (Jackson, 1994). The most recessive murine Agouti allele allows for no agouti protein production in the skin, and therefore results in a completely eumelanic phenotype. The more recessive alleles, above the one coding for a completely eumelanic phenotype, allow for agouti protein in ventral body regions, resulting in these being phaeomelanic. Successive alleles then add body regions to those of lower alleles, such that phaeomelanic areas of the coat are added successively as the series of alleles progresses from more recessive to more dominant. In a few of the intermediate alleles some regions have pulsatile agouti protein production, resulting in the banded hairs (on the dorsum) typical of the wild pattern in mice. These hairs have a eumelanic tip and bas e, and a phaeomelanic middle.
The murine Agouti locus patterns exhibit no reversal of pigment type among the alleles, so that regions which have become phaeomelanic from the action of more recessive alleles retain this pigment type in all patterns determined by the more dominant alleles. In this regard the murine Agouti locus acts as though the various alleles were simply shifting melanin production in an orderly stepwise progression from one extreme to the other, adding phaeomelanic areas body region by body region. The more dominant alleles result in phenotypes that are entirely phaeomelanic, while the most recessive alleles have phenotypes that are entirely eumelanic. The stepwise progress of Agouti locus patterns gave rise to early hypotheses that the locus was a complex of small loci that each controlled pigment type in only a specific body region.
Recent studies have shown that the complex hypotheses are mistaken, and that the locus is indeed single and simple rather than an array of miniloci (Jackson, 1994).
The most dominant murine Agouti allele is lethal yellow (Jackson, 1994). This allele is lethal to
homozygotes, and the mechanism of lethality appears to be that the very high levels of expression of the agouti protein disrupt fetal differentiation and development. The heterozygotes for this allele also have somatic manifestations of it, such as obesity, reduced fertility, and increased incidences of certain neoplasms.
The action of Agouti locus alleles in goats is not as neat and orderly as is the murine series. It is fairly common for the intermediate agouti patterns in goats to have reversals of pigmentation type, so that areas that are eumelanic in one pattern are phaemelanic in another, and vice versa. This reversal of pigmentation type results in some caprine patterns that appear to be opposites of one another, in contrast to the murine situation in which the Agouti locs patterns are characterized by a stepwise increase in phaeomelanic areas. The reversal of pigmentation patterns in caprine phenotypes is most remarkable in the "blackbelly" and "black and tan" patterns, which are nearly perfect opposites of one another in pigment distribution.
The action of caprine Agouti alleles is to consistently express phaeomelanic areas on a eumelanic
background. The result of this is that the tan or pale areas are dominant to the dark areas. This is a key concept to understanding the function of the Agouti locus. Some mild degree of intermediate dominance is present in some animals of some breeds with certain combinations of Agouti alleles, but as a general rule it holds true that the phaeomelanic areas are consistently expressed, whether heterozygous or homozygous. Heterozygotes for intermediate alleles therefore express the phaeomelanic areas for each of the alleles, and this results in phenotypes with combinations of both phaeomelanic patterns. Some heterozygotes for intermediate alleles and the entirely eumelanic nonagouti allele are darker than homozygotes for the same intermediate alleles, although this is more thoroughly documented in sheep.
This incomplete dominance has been noted for the badgerface allele, and it has also been found that overlapping phenotypes are more characteristic of females than of males (Brooker and Dolling, 1969a).
Goat Agouti locus patterns are numerous, and several have highly variable expression so that this locus is much more confusing in goats than in sheep. The patterns group themselves rather well, though, and considering them in groups can help to understand them. In this discussion the patterns will be discussed as occurring on a reasonably unmodified genetic background, so that the patterns take on a tan and black character. “Tan” (or gold or red) refers to pheomelanic areas, and “black” refers to eumelanic areas. While this is not universally true depending on what variants are present at other loci, it does hold true for goats that are “wild type” at all other color loci and is therefore a useful point of departure for understanding this incredibly complex and variable locus.
Some of the distinctions between patterns are subtle, but are very repeatable and are indeed significant indicators of the presence of different Agouti alleles. Goats, in general contrast to sheep, have several Agouti patterns that are extremely variable. These can be confusing, as their dark and light extremes resemble others of the patterns and it is only possible to be certain of the identity of the pattern by recognizing the range of effects over groups of individuals.
Patterns Capable of Producing White
Several of the goat Agouti patterns produce white goats at the pale extreme of expression. These patterns are interesting, as nearly all of them have darker (more rufous or more eumelanic) expressions that are clearly not white, and also are distinct from one another. The convergence of these on a starkly white phenotype at the pale extreme can easily result in confusion as to which of the alleles is present. A allele white may well occur as a distinct allele which consistently produces white goats and never darker colors. This is difficult to ascertain with no residual doubt. The overall level of intensity of the patterns varies with breed, and in some breeds that are long selected for white the modifiers that could provide for darker variants are lacking.
The allele white or tan yields unshaded, uniform colors that range from a fully intensely pheomelanic red, through yellow or gold, and to white. The genetic character of the modifiers is uncertain, although most manifestations of this allele cluster around deep red, medium yellow, or stark white and this suggests that the modifiers are few in number and may well be a single locus. This allele is a component of the Boer goat and other uniformly red breeds.
The shaded red allele results in white goats at the pale extreme, and at darker extremes is red with slight eumelanic shading on the body. Darker goats also have pale facial stripes from above eyes to nose, frequently with a thin dark eumelanic border to the white facial stripe.
Likely as a variant of white or tan is a pattern that is tan with darker tan trim in a pattern resembling the blackbelly pattern. This pattern may indicate a separate allele, or may simply be a dark manifestation of white or tan. One reason for thinking that this is a separate allele is that many gold and red goats do not have the darker trim of this variant and the two types consistently occur in different families.
The black mask allele is another that produces white in the palest extreme, but darker extremes can be red or yellow. Most have dark eumelanic shading on the head, and in the darker extremes this can result in considerably blackish regions, although with the retention of pale stripes from above eyes to nose.
The sable allele produces goats vary from very dark with red pheomelanin and extensive black shading to stark white. This pattern is extremely variable, and the extent of eumelanin and the depth of pheomelanin vary separately so that this single allele can produce patterns that are very distinct from one another. Most sable goats have white legs, white belly, and then are shaded to be deeper tan as well as more black over the top. Some have a shaded black backstripe from poll to tail, as well as a black shoulder stripe from withers, along shoulder, to the point of the shoulder and then to the midline. Faces are usually shaded with black, save for pale facial stripes.
Tan patterns with black trim
The wild type color pattern is produced by the bezoar allele, which has only moderately variable
expression. This, as with several other alleles, produces kids that are fairly light colored that then darken with age. The darkening is much more pronounced in males than in females, and is one of the dominance signals in the original species. The bezoar pattern is basically tan with a paler belly, perineum, and inside of the legs. Black trim includes facial shading which leaves pale facial stripes from above eyes to nose. The ears are rimmed in black, and a black stripe goes from poll to tail along the back. A black stripe is also common along the bottom edge of the neck. A black shoulder stripe is present. Black is present on the front aspect of the front legs as a stripe beginning beneath the carpus, and flaring out around the fetlock joint. A similar black stripe is present on the rear limb but is not discontinuous at the hock.
The wild riedell allele is named after the herd of origin, and is similar to the bezoar pattern but is overall much more eumelanic. It is minimally variable, and does not produce variants that could be easily confused with bezoar.
The “wild kolodzie” pattern occurs in a herd of Tennessee Myotonic goats. It is one of the extremely variable patterns, and this is evidence that it is not bezoar nor wild riedell. Light versions are white, while dark ones are nearly black. The most distinctive pattern resembles the bezoar pattern, although is generally darker than bezoar. The tops of the ears have distinctive mottled interplay of tan and black areas. Pale facial stripes are routine, sometimes only as dots above eyes as in the black and tan pattern described below. Striping on the legs resembles the bezoar pattern, as does the pale belly and perineum of this pattern.
The badgerface allele might more clearly be called blackbelly , and is similar to a homologous sheep pattern. These goats are tan, with specific black trim. The belly is black, as are the lower legs below tarsus and carpus. The inner aspects of the upper legs is also black, as is the perineum. A black backstripe is present from tail to poll. Black shoulder stripes are present, especially on males but generally on females as well. Dark facial stripes usually go from above eye, through the eye socket, and down toward or reaching the nose. The chin and throat are black, and this can continue down the bottom border of the neck as well. This is the common pattern of the Oberhasli goat breed.
The serpentina allele is named after a Portuguese goat breed. This breed is selected to have very pale pheomelanin, and whether this is due to the Agouti allele or to modifiers at other loci is uncertain. This pattern resembles blackbelly, but is somewhat paler in that the backstripe is discontinuous so that the neck usually lacks the stripe, and the shoulder stripe is minimal or missing. A pale region is present in the black area below the anterior aspect of the carpus.
Also in this group of patterns is the caramel of pygmy goats. This pattern retains the black belly and inner legs of the other others. The black lower leg is disrupted so that the anterior aspect is pale and the posterior remains black. This is most dramatic on the front limbs. The backstripe is incomplete, remaining on the tail and over the shoulders, which also have the black shoulder stripe. The neck lacks the stripe, and the head is extensively marked with black so that in older males the entire facial region can be black up to the ears. The bottom aspect of the ear is black, top is tan. In addition to the paler variants of blackbelly is a pattern occurring in Alpine goats, tan sides, in which the black areas of the blackbelly pattern are much more extensive. The result is that the sides remain tan, but the black extends and merges on the neck and head so that these are nearly or entirely black.
Patterns with anterior/posterior divisions of black and tan.
A group of a few alleles produces dramatic divisions between the anterior and posterior portions of the goat. These patterns do not appear to have homologues in sheep.
The peacock allele derives its name from a Swiss breed with this pattern. Generally, the posterior portion of the body is black and the anterior is tan. The posterior belly is black, as well as innner legs, and lower legs with a distinctive residual pale area distal and lateral on the anterior aspect of the front leg beneath the cannon bone. The top of the tail is usually tan, as well, although the rear half of the body is generally black. This varies from nearly entirely black to fairly pale with only the dorsal regions fully black and the other regions mixed tan and black. A triangular tan area usually remains on the lateral aspect of the thigh.
The tops of the ears are tan, bottoms are black. The head is distinctively marked with black facial striped from above and through the eye to the nose, with a second lower stripe in an arc from the base of the ear, and down below the eye to the cheek. The least black of the expressions of this pattern could resemble black belly, although the leg markings, lack of complete backstripe, and facial pattern are unique to this pattern.
A near reverse of the peacock pattern is called san clemente after the feral goats from that island which commonly exhibit this pattern. This is one of the Agouti locus patterns that is highly variable in degree of blackness. The result is that the blackest variants can be easily confused with the patterns produced by the black and tan or eyebar alleles, while the least black variants could be confused with a shaded red. The middle range of the pattern is very distinctive, with a tan belly, inner legs, and perineum. The lower legs have anterior black striping, which is discontinuous at the carpus (leaving it tan) and flares out around the fetlocks. The tops of the ears are black, undersides are tan. The head is black, with tan stripes
from above the eye to the nose. The repartida allele is one of a few that is typical of the Brazilian Repartida breed. This is similar to the middle expression of the the san clemente pattern, although the leg striping is very distinctive in that the anterior lower legs are tan, and the posteriors are pale. This is the reverse of most other Agouti locus patterns.
Grey or nearly grey patterns
Patterns within this group are not intuitively Agouti patterns, because they generally lack the distinctive symmetrical interplay of tan and black areas. However, they do reside at the Agouti locus in sheep, and segregation data suggest that this is true of goats as well.
The grey allele results in a relatively uniform mixture of black and white hairs over the entire goat, with no patterning. The shade varies depending on the relative proportion of black and white hairs. The palest ones are nearly white, while the darkest ones are still obviously grey. This pattern is uniform throughout the Azul Portuguese breed. The pygmy agouti grey allele causes the “agouti” pattern of grey in the African Pygmy goat. The mixture of white and black hairs is usually uniform over the body, but the legs are distinctly dark.
The grey striped allele results in a pattern that resembles a combination of toggenburg and grey but as a single allele. This is evidenced by the production of solid black kids by such goats. This is one of few Agouti alleles in goats that has a phenotypic appearance of a combination of two other Agouti alleles. Such combination alleles are much more common in sheep.
Black patterns with tan trim.
A group of dark patterns resides at Agouti, and some of these are subtle enough to be confusing in some individuals. The differences between them are usually in the head and leg patterning.
The toggenburg allele results in a black goat with distinctive tan trim. The trim is generally quite pale, although it is possible to generate goats with darker trim by appropriate modifiers that darken phaeomelanin. The pale areas include legs below carpus and tarsus, and facial stripes from above eyes to nose. The muzzle is also pale, as is the perineum but not the belly which is black. Scrotum/udder is tan. The top of the tail is black, bottom is tan. The top of the ear is generally pale, although the darkest manifestations have a distinctive shaded black bar down the middle of the ear. Usually there is a tan dot low on the cheek near the angle of the jaw, and possible one anterior to this as well. In the Toggenburg breed this pattern is further modified by having brown eumelanin, and also by having consistently pale pheomelanin.
The black and tan allele produces a pattern has a black body and head, with tan under the ear, small spots above the eyes, and usually ventrally on the throatlatch, belly, inside of legs, and perineum. The legs have black stripes on the fronts, with the foreleg stripe discontinuous just below the carpus, and an extension around the fetlock. The rear black stripe is continuous with the black of the body. Males darken somewhat with age, so that the tan “eyebrows” tend to be small or absent. Very similar to black and tan is eyebar which is similar except in having more extensive tan. The facial tan is a complete stripe from above eyes to nose. Most eyebar animals have two small tan dots low on the cheek, one near the lips and one posterior to this. The throat region is tan. The angel pattern is named after a goat by that name. This pattern is black with tan trim, including undersides of ears. A facial stripe runs from above eyes to nose, and is generally wider above the eyes and quite narrow near the nose. On some animals it is incomplete towards the nose. The chin is tan. A tan patch is present on the angle of the jaw. The perineum is tan, although the belly and scrotum/udder are black, which is one of the main distinctions of this pattern. The legs are pale with black stripes down the fronts. These are continuous from body to lower leg, and do not flare out over the fetlocks. They usually extend to fetlock or hoof, and end as a sharp point on the fetlock.
The lateral stripe allele results in a black goat with a tan stripe along the boundary of belly and body. The legs are pale, but with posterior (rather than anterior) black stripes.
Nearly black patterns
A few patterns are very dark. One is mahogany which is black with tan regions (usually tan and black mixed) over the body and especially over the backs of the thighs.
The tan cheek pattern is black except for round patches of tan below the eyes. The backs of the thighs are also usually mixed tan and black. The allele no pattern results in a completely (or nearly completely) black goat. Patterns such as grey striped appear to be the result of combining two distinct Agouti locus patterns, and are the phenomenon that made attractive the hypothesis that the Agouti locus is a complex of miniloci each controlling some aspect of pattern or body region. The presence of certain repeatable components across several patterns (black stripes on legs, shoulder stripes, backstripes) also makes this hypothesis attractive While the exact organization of the caprine Agouti locus is uncertain, it is unlikely to be much different than that of the murine locus, which has been shown to be simple rather than complex (Jackson,
1994). The caprine Agouti alleles do have differences from those of the murine locus, though, especially in the relatively high number of patterns that have reversals of pigment type distribution. Another major difference of the caprine alleles and those of other species is the tendency for some of them to have incredibly variable expression from blackest to least black, and from palest to most rufous.
Pleiotrophic Effects
Pleiotrophic effects for Agouti alleles have not been documented in goats, but similar alleles in sheep do have such effects. The AWt allele has pleiotrophic effects, in keeping with its status as the top dominant allele at the Agouti locus. Ewes with the AWt allele are on average less fecund than those without it. In Icelandic sheep, in which this has been best investigated, sheep bearing the AWt allele are on average 15% less fecund than those lacking this allele (Adalsteinsson, 1975b). In addition, sheep bearing the AWt allele are more strongly seasonal in reproduction than those lacking it (Dyrmundsson and Adalsteinsson, 1980). Obviously the breed genotype other than that at the Agouti locus is equally important in the overall fecundity and seasonality of any sheep, but the effects of this single allele are dramatic in breeds in which variants at the Agouti locus are segregating. However, it is worth a passing thought that the preference for
red in Boer goats selected for size and meat conformation might be assisted by the pheomelanic Agouti allele that characterizes the breed. Ryder et al. (1974) reported in Soay Sheep that the homozygous nonagouti genotype is less viable than others (usually black and tan). Soays are feral, and whether this trend holds true for sheep in less demanding environments is not known.
THE EXTENSION LOCUS
mechanism of action
The Extension locus encodes one member of a group of seven aMSH receptor proteins (Jackson, 1994). The aMSH receptor, following activation by attachment of aMSH, results in increased cAMP levels and increased protein kinase activity, which in turn activates eumelanin formation. In the inactive state phaeomelanin is produced. The wild type Extension locus allele in most species allows for control of eumelanin and phaeomelanin production to be governed by the Agouti locus as determined by the regional distribution of the agouti protein which inhibits eumelanin formation. Many species have dominant alleles at the Extension locus which act to increase adenylate cyclase either spontaneously or in response to aMSH. The specific mechanisms are different for different alleles in the mouse, and involve either responsiveness to aMSH or autonomous adenylate cyclase production (Jackson, 1994). Either mechanism results in an entirely eumelanic phenotype, and these alleles are usually considered as "dominant black" regardless of the underlying mechanism used to achieve this phenotype.
Recessive alleles at the Extension locus consistently result in an inactive receptor that is unresponsive to stimulation by aMSH. The inactive receptor results in a completely phaeomelanic phenotype. In this regard the Extension locus acts opposite to the Agouti locus since more dominant phenotypes are completely eumelanic, and more recessive ones are completely phaeomelanic. Extension locus alleles tend to affect the entire animal, so that this locus does not produce patterns as are typical of the Agouti alleles, with interplay of the two pigment types. The Extension locus is usually responsible for completely eumelanic or completely phaeomelanic phenotypes, although exceptions do occur in some other species
(Searle, 1968).
alleles
The Extension locus has few variants in goats, and these occur in only a handful of breeds. In those few breeds this locus has important interactions with the Agouti locus in determining color. The wild type allele at the Extension locus allows the expression of the Agouti locus. It is by far the most common Extension allele in nearly all breeds of goats, most of which have obvious segregation at the Agouti locus to account for the majority of color variation.
The dominant allele at the Extension locus is called dominant black and is symbolized ED. This allele causes a uniformly eumelanic coat, which is black on an otherwise unmodified background. Dominant black in goats has been documented in Angora goats, and anecdotal evidence suggests that it might also be present in some African breeds. In contrast to its rarity in goats, a similar allele is fairly frequent in sheep breeds of a wide variety of geographic origins.
The similarity of the phenotypes resulting from dominant black at the Extension locus and nonagouti at the Agouti locus can be confusing, as these two cannot be easily distinguished. Recent seletion in favor of eumelanic phenotypes by some Angora goat breeders has resulted in the presence of both mechanisms for black in some populations. Visual inspection alone cannot distinguish between these two biological mechanisms that achieve similar phenotypes. The persistence of both mechanism can result in some strange segregations of color unless the observer is aware that both the dominant black of the
Extension locus and the recessive nonagouti of the Agouti locus can be present.
A single occurrence of a red Angora kid from two obviously striped Agouti locus pattern parents points to the possibility of a recessive allele at Extension that codes for uniformly phaeomelanic color. No segregation data are available to substantiate this allele any further.
The Albino Locus
The Albino locus codes for the tyrosinase enzyme that is essential for melanogenesis. Recessive alleles at this locus are responsible for the production of abnormal forms of tyrosinase that either have reduced or nearly completely absent activity (Jackson, 1994). As a result, melanocytes are present but are incapable of melanogenesis. The Albino locus has never been documented to have variation in goats, although two different variants occur in sheep (Adalsteinsson, 1977, 1978a; Rowlett and Fleet, 1993).
Expression and Modification of Phaeomelanin
Phaeomelanic colors are subject to various modifications, some of which are well understood and many others of which are not. Many of these modifications do appear to have a genetic basis. Phaeomelanic colors are modified by independent loci controlling color directly, and are also modified by the physical structure of the hair coat. It is difficult to separate the results of these two phenomena. Some evidence points to a fairly limited number of modifiers for phaeomelanin intensity in goats. In some families the expression of phaeomelanin clusters around deep red, medium gold, and cream to nearly white. The lack of all intermediates between these three classes supports the contention that control may be at a single or a few loci with incompletely dominant alleles. Proof of this hypothesis is lacking, though.
Expression and Modification of Eumelanin
Eumelanic areas on goats are subject to modification by mechanisms distinct from those affecting phaeomelanic colors. One fairly common modification is the replacement of black eumelanin with brown eumelanin.
TheBrown locus
mechanism of action
Alleles assigned to the Brown locus are a common source of brown color in goats. This locus controls tyrosinase-related protein 1 (TRP-1), which has important but poorly documented action within melanocytes (Jackson, 1994). The recessive brown allele reduces activity of TRP -1, and the result is that all regions that are eumelanic have the black form replaced by a brown form. Brown melanosomes are structurally different than black ones (Alexieva et al., 1985). In the brown phenotypes the catalase normally involved in melanogenesis fails to protect the eumelanin from the action of hydrogen peroxide which is produced during melanogenesis.
The murine Brown locus has a few dominant alleles in addition to the recessive ones. These usually have apical pigmentation and basal pallor on hair shafts, and similar effects have not been documented in goats. The banding on hairs that is caused by these murine alleles is due to the buildup of toxic metabolites within the melanocytes, which increase as the hair cycle progresses. The result is that the apical portions are pigmented since melanocytes are functional, but as the hair grows the melanocytes undergo toxicosis and pigmentation is therefore diminished as the base of the hair is produced.
alleles
At least three different alleles can result in brown eumelanin in goats. Whether these all reside at the Brown locus is undetermined, but they are all likely candidates for this locus. One allele is brown, and is recessive. This results in a medium brown color in areas where other loci allow eumelanin. Brown eumelanin can replace black eumelanin in any of the Agouti locus patterns, as well as in goats with the dominant black allele of the Extension locus. The brown pigmented version of the various Agouti locus patterns is usually more subtle than is the black one, because the contrast of the pale areas with the brown is less than that of pale areas with black.
More common among goats are dominant alleles leading to brown eumelanin. It is likely that two different alleles occur. Dark brown changes black eumelanin to dark brown. The kids are usually born barely “off black” and can easily be confused as having black eumelanin rather than brown. By the age of a few months, though, the difference is obvious and remains so throughout life. Light brown, in contrast, results in a very obvious change from black to brown eumelanin. This is the allele that is responsible for the color of the Toggenburg breed.
At least in sheep, the Bb allele has pleiotrophic effects in addition to its action on color. Ryder et al. (1974) found that Soay sheep homozygous for Bb were less viable than sheep not homozygous for this allele. Homozygous sheep were also smaller bodied than those of other genotypes. This effect was most noticed in AaAaBbBb sheep. This work concerned the feral Soay sheep, in which the brown allele is relatively common. A similar effect on nonferal sheep has not been documented.
Moonspots
Moonspots are a conspicuous if unusual phenomenon in goat color. These are round to oval spots of varying size, and are superimposed over any background color. They are not white, but are usually very pale. They tend to be more intensely pigmented in kids than they are in adults, and can easily be confused with white spots in adults. The exact genetic control of these is uncertain, although it is likely that they have at least some dominant characteristics. The number and size of moonspots is highly variable, and they can easily be overlooked in some goats. They are most common and dramatic in AngloNubians as well as in other African and Indian breeds, although they do occur as a rare variant in a wide range of goat breeds.
Loci Controlling White Spotting
Patterns of white spotting are added independently to any colored background of goats. Multiple spotting patterns occur. Very few of these have any importance to goat breeds worldwide, but several occur in a wide range of breeds.
The Angora White Locus
The Angora White locus derives its name from the Angora breed, in which it is a common cause of white coat. It may not be the only genetic mechanism for the highly desired white mohair fiber of this breed, but it is one of the more common and is therefore important. A dominant allele, angora white, results in a completely white phenotype. Some evidence for slight incompleteness of dominance has been found, such that some heterozygotes have colored (usually black) stripes in horns and hooves. The allele is epistatic to Agouti and Extension, which makes breeding for color in Angora goats especially challenging.
Piebald Spotting
In mice the Piebald (S) locus has been documented to act on the differentiation of melanocytes at the neural crest, as well as on their migration from the neural crest to the rest of the body (Jackson, 1994). The result is an array of white regions on otherwise pigmented mice. The specific color of the pigmented regions is governed by the other coat color loci, so that white spotting can be superimposed over any color or pattern.
Mice with extremely white manifestations of spotting are afflicted with aganglionic megacolon, which is a lethal condition that occurs relatively early in life. This condition results from failure of neurons to migrate from the neural crest to the colon, much as the white spotting results from failure of the migration of melanocytes from neural crest to skin. The alleles for spotting at the S locus are recessive in most species, and result in unpigmented regions of skin and hair.
No definitive research in goats has documented alleles at the S locus. Patterns resembling the spotting of other species (including the taxonomically close sheep) suggest that this locus does indeed have variants in goats. Especially likely as candidates are minor white marks on head, tail, and distal legs. In addition, some obviously spotted goats have round patches of color over ears and eyes, and on the body. These are especially likely to betray S locus patterns. The dominance or recessiveness of these is undetermined. Spotting from alleles at this locus may be important in yielding the final color of the Boer goat.
Belt
A fairly repeatable marking in goats is a white belt over the midsection of the body. The belt can vary from narrow and incomplete, to very large and including one or more legs and the rump or shoulder and neck. This segregates as a dominant gene, and is likely to be one of the components of the Boer pattern. The belted pattern is among the most common white spotting patterns in goats.
Roan
Roan goats result from a mixture of white hairs into any background color. The amount of roaning varies tremendously, and in minimal grades might not be readily noticed. In more pronounced manifestations the intermixture occurs over the body and neck, but generally spares the head and legs which remain the background color. No segregation studies have been done, but results from some families suggest that this is a dominant allele. A dominant mode of inheritance is consistent with its action in most other species.
Flowery
The flowery pattern is a breed characteristic of the “Florida Sevillana” breed, and derives its name from that breed. This pattern consists of small (up to about 1 cm) spots of white scattered over any background color. The pattern varies, and maximal grades are very pale while minimal grades are very dark. The white speckling tends to be most intense on the ventral midline, and on many flowery goats the result is a nearly white ventral midline that feathers out into speckled roan sides, and then to a nearly solid colored topline. Segregation in a few families suggest that a dominant gene is responsible for this pattern.
Goulet
Goulet spotting derives its name from the herd of Tennessee Fainting Goats in which the pattern was first documented. Minimal grades of spotting include white ears with a few residual small spots of color, and facial white that usually does leave the nose, lips, and eyes with pigment. A few white spots, usually about 1 cm, are scattered over the rear flanks and hips, and the tail is generally white. Medium grades of spotting include white ears, facial white sparing the eyes and nose, and then a ragged interplay of white and colored areas over the body. Maximal grades are nearly white, with residual ragged colored spots
over the body.
Algarve
The Algarve breed sports a ragged and irregular spotting pattern that resembles the spotting of the Goulet pattern except that it consistently has colored ears even in very white individuals.
Barbari
The pattern of the Barbari goat is somewhat elusive, and consists of small colored spots in areas that are white in patterns consistent with the belting or spotting patterns. These dark spots are present at birth, in contrast to the ticking pattern in which they grow in later. This appears to be a modification of any spotting pattern, and the genetics of this have not been documented although early results are consistent with a dominant gene.
Ticking
Small spots of pigment grow into the fleece or haircoat of some goats that are spotted by virtue of any of the spotting patterns. This is called ticking, and the small spots are not present at birth but grow into the white areas later. Ticking is dominant to its absence, and varies considerably in extent. The mechanism that allows melanocytes to populate regions that are embryologically devoid of them has yet to be elucidated. This gene is analogous to ticking in dogs, on which species it is extremely variable in extent and in size of the tick marks (Little, 1957).