The Land of Painted Caves: Review

Jean Auel began her “Children of Earth” series over 30 years ago, with Clan of the Cave Bear. The Valley of Horses, The Mammoth Hunters, The Plains of Passage and The Shelters of Stone followed. Her latest addition, The Land of Painted Caves, continues to follow Ayla and Jondalar, still having difficulty communicating, and this time includes a tour of the cave paintings of France.

All are long books – The Land of Painted Caves is 828 pages in paperback and the others are about the same length. All are well researched. I discovered the series 30 years ago, primarily because of my interest in the Pleistocene and human evolution, and most of this review will be from that perspective.

From a writer’s point of view, the most recent book is full of information dumps, and rather weak on plot. That hasn’t stopped it from being a best seller, but there were times when I had to force myself to pick it up. I did manage to find a number of usable Twitter quotes, which are being posted and their contexts will be explained on February 29.

A good part of the book is description of the cave art of France. Auel does include a map keyed both to what the Zelandonii of her book called the caves and what archeologists call them, but I wanted to see some pictures of the cave art, not just descriptions. I actually searched the web for images from the caves, but found very few even when I knew the name of the cave. Good general references are http://myrencounter.blogspot.com/2011/07/land-of-painted-caves.html and http://www.donsmaps.com/indexauelfans.html, but they have more photos of the locations of the caves than of the actual paintings. White Hollow, identified as Lascaux, does have some images of the art at http://www.huffingtonpost.com/2010/09/12/inside-lascaux-the-versai_n_712645.html.

One possible source, at least for drawings of the cave art, is The Nature of Paleolithic Art, by Dale Guthrie. Dale is an artist himself, and while he suggests that a good deal of the “art” in the caves was equivalent to graffiti found in mens rooms, his first interest in cave art was as guides to reconstructing extinct animals. This is a huge book, with hundreds if not thousands of drawings of Pleistocene art from all over Eurasia, but putting the drawings in this book together with what Ayla saw would be a major project.

Leaving the art, there has long been a controversy in archaeology as to whether modern humans and Neanderthals (what Ayla calls the Clan) ever interbred, or whether such interbreeding was even possible. The argument went back and forth during the time period over which Auel’s books were being written. DNA for a time was used to claim such interbreeding never occurred. Then, less than a year ago, DNA evidence made it quite clear that such interbreeding had in fact happened. The basis of Auel’s books was if anything ahead of the archaeology of the time.

In one point, however, she was clearly wrong, though there was no way she could have known it at the time she started the series..  Jondalar and Ayla are described as being blonde and having blue and gray eyes respectively. Recent gene sequencing has strongly suggested that all blue and gray-eyed people are descended from a single common ancestor who lived between 6,000 and 10,000 years ago, well after the setting of Auel’s books. There is at least some argument that blonde hair may have evolved after the ice ages. Still, I cannot help but wonder if it could be derived from that Neanderthal admixture. If fair coloring is an adaptation to getting vitamin D in a region with little sunlight, such as Europe, the Neanderthals lived in Europe long before the Cro-Magnons arrived.

Cinderella (DVD Review)

I’m not a big fan of modern records. My taste runs to Mahler, the classics, Andrea Bocelli, Sarah Brightman, and the occasional soundtrack. I can’t say that I ever consciously listened to a record by Whitney Houston. Nevertheless I feel I knew her a little through a DVD I’ve watched many times: Rogers and Hammerstein’s Cinderella, as co-produced by Whitney Houston and Walt Disney Enterprises.

I’ve loved the music since I first heard it, on the original CBS broadcast with Julie Andrews as Cinderella in 1957. I don’t know if there even were color televisions then. We certainly didn’t have one, and my DVD of the original broadcast is in black and white.

The Disney version has a somewhat more traditional ending than the original broadcast, but the music is just as glorious, if not more so. And color? This DVD fairly pops with color, from the fabric swathes during the food dance (I don’t know what else to call it) to the row of gaily-patterned stockings on the feet of the girls waiting to try on the glass slipper.

The cast is multi-ethnic to a somewhat startling degree. Whoopi Goldberg and Victor Garber as the king and queen gave me no problems, nor did Paolo Montalban as the prince. Brandi as Cinderella gave life to some of my favorite songs, and Bernadette Peters was a wonderfully wicked stepmother. But the stepsisters…

There is nothing wrong with the performances of Jenne Cox and Natalie Dusselle. Their interaction, as rivals united only in despising Cinderella, was excellent. But I can’t help looking at things with a geneticist’s eye, and for years I was bothered by the idea that the thin, pale-skinned redhead and the plump chocolate-skinned woman were full sisters, both the daughters of a very pale-skinned mother. Recessives are unlikely to be that well hidden.

Finally, I realized that there is absolutely nothing said about how many times the stepmother had been married! Three times, obviously, with her two daughters having different fathers, and the third marriage being to Cinderella’s father. Now I can relax, my inner geneticist satisfied, and simply enjoy the DVD.

Whitney Houston played the fairy godmother with the aid of some special effects from the Disney team, and she was magnificent in the role. According to the featurette on the DVD, she was a real-life fairy godmother to Brandi, too.

I suppose it’s a bit light and fluffy by the standards of today’s movies, but it’s still one I can watch over and over again. Go Cinderella – and some day I’ll do a comparison of some of the DVDs and retellings of this story.

Horse Color Genetics Index

Color genetics is a long-time hobby of mine, and I’ve written a number of posts about the genes that determine color and pattern in horses. I also have an extensive website on genetics and coat color in dogs.

Horse Color Genetics Index

The Basic Colors of Horses 10/31/10
Palomino Genetics 11/06/10
Pearl: a Palomino Complication 11/14/10
Dun: a Wild-Type Dilution Gene in Horses 11/21/10
The Agouti and Extension Loci in Horses 11/27/10
Champagne: Another Dilution Gene in Horses 12/5/10
Silver Dapple – Another Dilution Gene in Horses 12/12/10
Two Rare Dilutions and a Summary of Dilution Genes in Horses 12/19/10
The Genetics of White on Horses 12/27/10
The Grey Gene in Horses 1/3/11
The Roan Gene in Horses 1/9/11
Other Patterns of White Hair in Horses 1/17/11
White Body Markings on Horses 1/24/11
The Tobiano Gene in Horses 1/31/11
Sabino Spotting in Horses 2/6/11
The Frame Gene in Horses 2/14/11
The Splashed White Gene in Horses 2/28/11
Horse Colors: Manchado and Brindle 3/7/11
White Horses 3/20/11
The Leopard Gene in Horses 3/29/11
The Leopard Gene in Horses (Continued) 4/2/11
The Leopard Gene in Horses Part 3 4/10/11
The Leopard Gene in Horses 4 4/17/11
Horse Color Genetics: Darkening Genes 4/24/11
Horse Color Summary 1 5/1/11
Horse Color Genetics, Final Summary 5/8/11

Other Posts to do with Horses

The Horses of Homecoming 4/17/10
Obstacle Racing (Homecoming Glossary) 6/26/10
A Circus Horse with no Circus 7/11/10
Tourist Trap: What’s It About? 8/16/11

Three Approaches to Health Care

There are three ways of approaching less than optimal health: prevention, treatment and cure. All have their strong and weak points; all are political hot cakes at the moment.

I think most individuals and societies would agree that the best solution is to stay in good health. It is also a solution that is not always possible. It is, however, the approach that is responsible for the dramatic drop in childhood death in developed countries.

Prevention measures are generally lumped as public health. Clean drinking water, proper sewage disposal, nutritious food, clean air and immunizations all fall into this category. So do measures intended to reduce accidental injury or death (such as seat belts) and those encouraging a healthier life style. Prevention would also include such highly controversial measures as not passing on genes known to have a deleterious effect on health.

In general prevention measures are good for the population, but affect individual choice. They may also affect the bottom line of corporations with a great influence on public policy, such as the food industry.

Some of what I need for treatment of diabetes. I want a cure!

By treatment I refer to ongoing treatment — the pill (or shot) for everything or treatments such as dialysis which must be repeated regularly for the life of the patient. Certainly it is better to have a treatment than not! I am alive today thanks to insulin. But treatment is generally expensive and is often lifelong. Further, treatment of this sort almost always has undesirable side effects. In my case, insulin can produce hypoglycemia which can kill. It should be noted that some “cures,” such as organ transplants, may then produce a condition in which lifelong treatment (anti-rejection drugs) is needed.

A cure implies a return to normal health. In some cases (such as the common cold) a cure is mainly a matter of time, with supporting treatment to prevent secondary infections or ease symptoms. Some cancers are curable with surgery, especially if caught early. Broken bones or other traumatic injuries can often be cured, especially in the young. A number of conditions, however, have no cure. Research on cures is ongoing, but the profit from a true cure is usually not as great as from lifelong treatment. Given that most health research today is profit-driven, research on cures tends to take a back seat to research on treatments.

I’ll probably return to this in the future, looking at one of the three approaches at a time. For right now, how would you order the importance of the three approaches?

Fairbanks weather 7/18/11

Red lychnis

Sunrise this morning was at 4:02 am, and for a change sunset will also be today, at 11:49 this evening. Day length is down to 19 hours 47 minutes, and we’re losing more that 6 ½ hours of potential sunlight a day. Not daylight, really – it’s still light enough to drive all night, and the sun dips less than 4° below the horizon. Granted it’s only a little more than 45° above the horizon at most, but for practical purposes we still have 24 hours of daylight.

Salmon lychnis--this one's about eye level.

The lychnis (Maltese Cross) is blooming, and I’m beginning to wonder if I have a new hybrid variety. I have lots of red ones, all descended via volunteer seedlings from a single red I purchased years ago. More recently, I bought a packet of salmon seed (actually a very light salmon pink) and raised several plants from it. The salmon is a taller, earlier plant, with umbels of small flowers; the red is shorter, blooms a little later and has relatively large flowers. Both self seed freely.

The first large blooms this year were on plants that looked like the reds, and the blooms were the size of reds, but they were lighter in color. Could pollen from the salmons have fertilized some of the reds? At any rate, I like the new color.

Possible hybrid? Flowers on this and the red are almost 2" across; those on the salmon are less than 3/4".

Horse Color Genetics: Final Summary

Last week we reviewed the base color and dilution loci. Today we will do a final review of the interspersed white hair and white marking genes, along with the darkening genes. Although the blog series will end today, links will be put on my author site to all of the posts in the series.

There are two main loci responsible for interspersed white hair. These are Grey (born dark with white hairs becoming more numerous with age) and Roan (born roan with white hairs constant or decreasing with age.)

The Grey locus is the syntaxin-17 (STYX17) locus on equine chromosome 25. It causes an initial increase in melanocytes  followed by their depletion. There are two alleles at this locus: grey and wild-type, with gray being incompletely dominant. (Horses with two copies of the grey allele lighten faster than horses with one grey and one wild-type allele, are less likely to develop a fleabitten appearance, and are more likely to develop melanomas with age.) At this time the progression of graying (dark vs. light mane and tail) and the color of dark hair (usually black, but some individuals become rose grey, with the dark hair remaining red) are not known to be subject to genetic control. In any case the final result is a mostly white horse.

The Roan locus is close enough to the Extension locus that there is significant linkage. It is considered part of the KIT linkage group on equine chromosome 3. There are two alleles: roan (dominant) and wild-type. At one time possession of two roan alleles was thought to be lethal, but this has now been shown not to be true. Classic roan causes interspersed white hairs on the body, but the legs, mane and tail normally remain dark. The frosty pattern, in which the mane and tail are also affected, may be a variant of roan, but the genetic mechanism is at present unknown. Scars commonly lack white hair, causing dark corn marks.

Spotting loci are far more numerous, and some produce roaning as well as white areas.

Minor spotting genes may be responsible for white facial and leg markings. These genes are present in most breeds, and facial and leg white tend to increase in tandem. Animals with wide blazes and no white on the legs, or with high stockings and plain faces are very often minimally marked animals with one of the other spotting genes.

The Tobiano locus is closely associated with the KIT locus, and hence on equine chromosome 3. There are two known alleles, tobiano and wild-type, with tobiano being an incomplete dominant. Generally tobianos are crisply marked, with white crossing the topline. Legs are normally white and the face is plain or has minor markings. Minimal tobianos may have high stockings with plain faces; in the maximal pattern only the head may be colored. Roan or colored spots known as paw prints may occur in white areas on animals with two tobiano alleles. There is a dominant modifier which in the presence of both tobiano and cream produces what is called a calico pattern—the yellow of the buckskin or palomino is broken up, with some areas being red.

The Frame locus is on equine chromosome 17, and is at the locus that controls endothelin receptor b (EDNRB.) The alleles are frame and wild-type. The frame allele is lethal in double dose, producing the so-called lethal white foal syndrome, so all frame horses should have one frame and one wild-type allele. The minimal expression of frame is extensive white on the head with colored legs. The maximal extent may have color confined to the topline and legs. The fact that the frame allele still seems sometimes to come out of nowhere need further clarification—a masking gene may also exist.

The sabino pattern is a combination of spotting and roaning, and extremely variable in expression. It may also have more than one genetic explanation. The Sabino-1 locus is part of the KIT complex (equine chromosome 3) and has two alleles, sabino and wild-type. The sabino allele is incompletely dominant over wild-type, as horses with two sabino alleles generally have more white (even to being almost completely white) than horses with one sabino and one wild-type allele. There are other mutations near the KIT locus that cause white spotting, some of which appear to be lethal in double dose.

The Splashed White locus is yet another that seems to be near the KIT locus, though not at it. The locus probably has two alleles, splashed white and wild-type, with splashed white behaving as an incomplete dominant. The minimal effect of splashed white may not be detectable, or the horse may be more extensively marked with white legs, possibly white underbody and generally white on the head, sometimes to the extent that the whole head is white. Splashed white is also associated with deafness.

Manchado is a relatively rare type of spotting found in several breeds in Argentina, though that may be because of the Argentine fascination with coat color. Parts of the body, often including the top of the neck (and mane) are white, often with round colored spots. The genetic basis is unknown.

White with pink skin and dark eyes may be a separate gene, possibly lethal in horses with two white alleles. At the moment, this is somewhat up in the air.

The Leopard locus is the Transient Receptor Potential Cation Channel, Subfamily M, Member 1(TRPM1) locus. It has two alleles, leopard and wild-type, but an enormous array of patterns. Leopard is incompletely dominant over wild-type—horses with two leopard alleles generally have fewer leopard spots than those with one leopard and one wild-type gene, and have a high incidence of night-blindness.

Finally, darkening due to black hair in the coat may occur in at least three forms. Black hair may be scattered throughout the otherwise red parts of the coat, producing a sooty effect. Black tipping on otherwise red hairs appears to be associated with the agouti locus, and produces shaded effects where the back appears darker than the rest of the horse. Actual black striping of the coat, brindle, is rare but documented. Some types of roan, especially sabino, may produce a type of brindle with white stripes. The genetics are unclear in all of these cases.

Horse Color Summary 1

As a final summary of horse color genetics, let’s go over the loci, what they do, and the alleles at each locus. My primary reference is Sponenberg.

The Agouti locus is widespread in mammals, and is involved with whether and where an animal produces eumelanin (black) or phaeomelanin (red) pigment. The alleles known in horses, listed with the most dominant first, are Wild Bay (Wild-type), Bay, Seal Brown and black. Agouti is hypostatic to Extension, meaning that the effects of the agouti alleles can be seen only if the extension gene allows the animal to produce both eumelanin and phaeomelanin. Note that at this locus, the redder the color, the more dominant.

The Extension locus is the same as the melanocortin receptor one locus, or MC1R. Like agouti, it influences whether eumalin or phaeomelanin gets into the coat and occurs in most mammals. The alleles are dominant black (still not confirmed), wild-type, and chestnut. This locus may also have genetic control over the depth of black tipping. Only wild-type and tipping allow the agouti genes to show. In this series, more black is dominant over more red. Extension is epistatic to agouti.

Agouti and extension determine the base color of the horse—bay, brown, black or chestnut.

The various dilution genes generally affect phaeomelanin and eumelanin differently, coarse and fine hair differently, and not uncommonly are associated with patterns of dilution.

The Dun locus has two alleles. Wild-type is dun and is dominant over non-dun, but the wild type is rare in many breeds. When present, dun dilutes both black and red pigment on the body, but the degree of dilution varies a great deal. Head, legs, mane and tail are generally much less affected than is the central body, and dorsal stripes almost always occur. “Zebra stripe” markings often occur on legs and the shoulder region. The center of the mane may remain light.

The Cream locus is also known as the membrane-associated transporter protein (MATP) locus. It probably has three alleles: Wild-type, pearl, and cream. The dominance hierarchy here is complex. A horse with two wild-type alleles is normal color. A horse with one wild-type and one pearl allele looks normal color except for slightly lighter skin. A horse with two pearl alleles will have red lightened to gold and black lightened to beige. A horse with one cream allele and one wild-type allele will have red lightened to gold and black lightened only very slightly. A horse with one cream and one pearl allele will have red lightened to pale cream or ivory and black lightened to beige. Finally, a horse with two cream alleles will be a very pale color, as red lightens to cream and black to a slightly dirty white.

The Champagne locus is the SLC36A1 locus. It has two alleles: Champagne (dominant) and wild-type. Champagne dilutes red to gold and black to brown or tan. The mane and tail are generally diluted less than is the body.

The Silver Dapple locus is the pre-melanosomal protein 17 (PMEL17) locus. It has two alleles, silver (dominant) and wild-type. The silver allele dilutes black strongly but has little or no effect on red. The allele also produces very strong dilution in mane, tail and lower legs, at times producing horses that appear black with white manes and tails. Far commoner are horses with a blue to chocolate body, often heavily dappled, with distinctly lighter manes and tails. At one time common primarily in ponies.

The Mushroom locus has not yet been located. Two alleles are suspected, wild-type (dominant) and mushroom (recessive.) Mushroom horses resemble silver dapples, but lack dappling and have tested chestnut at the extension locus.

Arab dilution is another possible locus. This is believed to be a recessive allele with a strong lightening effect on black but little or no effect on red. Both Mushroom and Arab dilution are very rare.

I will summarize patterns of white, including grey and roan, next week.

Horse Color Genetics: Darkening Genes

Horse colors are due to the interaction of a large number of genes, many of which we’ve discussed. These may be divided into base color genes, diluting genes, genes that cause interspersed white hairs and marking genes. One type we have not discussed, because the genetics are not really understood, is interspersed black hairs.

I’d like to point out one thing that Sponenberg does not cover: there are two distinct types of darkening due to the presence of black. It takes a magnifying glass and a great deal of patience to tell the difference, but darkening can be due either to interspersed black hairs (called sooty and it can occur on any base color) or to hairs that are red/yellow at the base and black at the tip (producing a shaded appearance and I think occurring only on bay, wild bay and some seal brown horses.)

For the rest of this discussion I will assume the horse is of one of the base colors, but sooty and shading can occur with any dilution or marking genes, or together with roaning or grey. You just have to remember what the other genes do to red and black pigment, or if they have different effects on coarse and fine hair.

Liver chestnuts are often sooty. It takes careful examination to tell if a chestnut has interspersed black hair, but when I was examining them with a magnifying glass, this was true of every liver chestnut I examined. Even red chestnuts often have a few black hairs mixed into the coat and the mane and tail. Bays can also be sooty, but this may be confused by the presence of shading.

In order to understand black shading, it is necessary to go back and look at how the agouti locus affects mammals in general. The locus got its name from a middle and South American rodent, the agouti. This animal has fur in which the individual hairs are banded in black and yellow. As it happens this is very common in mammals, and a number have banded hair. Unless the hair is very coarse this is not obvious—wild gray mice and rabbits, for instance, really have hair banded in black and light yellow.

The banding may vary from multiple bands on a hair to hair with red/yellow/cream bases and black tips. The banding may also vary with type of hair, with some hairs (often the coarsest) being solid black and others (often the finest) being predominantly yellow.

Remember bay and wild bay are due to genes at the agouti locus. Most bay horses have at least some banded hair on the body, usually with a red base and black tip. This is easiest to see around the edges of the ears, and the banded hairs tend to be most numerous along the spine and spreading down to cover the hips, shoulders and upper barrel. It’s been a long time since I actually looked at individual hairs with a magnifying glass or under a microscope—I was doing this in the late 60’s and early 70’s. But as I recall, just about every bay or buckskin horse I looked at had at least a few black-tipped hairs. In some, the black tipping produced a shaded effect on the body of the horse.

A few horses look black or seal brown in summer coat and quite different in winter. I recall two of these. Duchess was a typical seal brown in the summer—black with tan shading on her flanks, muzzle and under her tail. In winter she looked dark bay. Careful examination of her winter coat showed red near the skin with deep black tips. In her short summer coat, apparently only the tips showed.

The other was even more striking. I first saw him in winter coat, and thought at first he was a blue roan. Careful examination of his coat showed yellow bases with black tips rather than interspersed white hair—a buckskin with deep black tipping. In summer, I knew he was the same horse only because the stable owner identified him—he was a typical seal brown.

This is an area that needs much more research. Unfortunately with the prominence of DNA in genetic research, researchers seem not to be paying as much attention to the distribution of pigment in the hair.

The Leopard Gene in Horses 4

Not all horses with the leopard gene have blankets of any size, and not all have spots. The gene can also produce two specific types of roaning, called frost and snowflake.

These roan patterns are quite separate from that produced by the roan gene, which becomes less prominent with age and leaves head, legs, mane and tail dark. The leopard gene produces horses which are normally colored or at most have a few white hairs over the rump at birth, but develop roaning (frost) or scattered white spots (snowflake) as they age. In contrast to grey, the pattern eventually stabilizes rather than producing a pure white horse.

In frost, the roaning tends to be most prominent over the hips. So-called varnish marks are common — areas where the bones are close to the surface, such as the hipbones and nasal bones, retain pigment while the rest of the coat is roaned. An aged varnish roan may be almost white except for these varnish marks.

Snowflakes are small white spots scattered randomly over the body, but often most numerous and prominent on the front part of the horse. They tend to become larger and more numerous with age, until in extreme cases the horse appears white with colored specks. This gives what is often called a speckled pattern, not to be confused with flea-bitten grey. Note that not all of the photos shown at the link are true snowflakes — the term is used very loosely.

Both types of roan may be combined with any of the blanket or spotting patterns, or may occur alone. Figure 8.140 in Sponenberg is a beautiful example of a combination of snowflake, varnish roan, blanket and leopard spotting all on the same horse. (Put Figure 8.140 on the search inside field.) Since the leopard gene can produce any of these effects, alone or in combination, breeding leopard-pattern horses can lead to some interesting results.

The remaining named horse in Tourist Trap, Amber’s mount Splash, is a bay varnish roan with a small spotted blanket, in color rather like the horse in Figure 8.141. He’s a gelding, about 14.1 hands – just enough smaller than the other four to have problems with fords. Roi has seen only solid colored horses on Central, and his first look at Splash gives this impression:

“Amber’s [horse], a little bay roan with curious dark lines on its nose, looked less exotic until it turned as she halted it.  Then it became apparent that it had a large white area, punctuated by dark bay spots, over its hips.”

I may summarize the equine color loci and alleles next week with links back to where they are mentioned, but I have covered most of the known color genes in horses. That doesn’t mean more won’t be found!

The Leopard Gene in Horses (continued)

The pattern most people first think of in Appaloosa horses is the one that gave the gene its name—leopard. This pattern gives a white horse with round or oval spots of base color. There may be shading of the genetic base color on the flanks, behind the elbows or on the head.

Genetically, a leopard must have at least one Pattern-1 allele in order to have most or all of the body white. In addition, it must have one leopard allele and one wild-type allele at the TRPM1 locus. Two leopard alleles will lead to a few-spot leopard, with only a few colored spots. Other factors leading to the leopard pattern undoubtedly exist, but are still unknown.

The mane and tail may be mixed in color if some of the mane and tail hair grow from colored spots. The spots may have roan edges, called haloes, which normally develop after birth. Blacks tend to have more and larger leopard spots than do chestnuts, with bay being intermediate. Also, horses with black mixed in the coat (sooty) will sometimes have the black and red colors form separate spots.

Three of the horses in Tourist Trap have leopard markings.

Token is the mare ridden by Flame. She is fairly tall—around 16 hands. She is a chestnut leopard, white with copper spots. Genetically, she is homozygous for the most recessive of the extension alleles, has two copies of the Pattern-1 allele and one of the leopard allele. She is wild-type at all dilution, pinto spotting, grey and roan loci. She could have genes for minor white marking on face or feet, but they cannot be seen.

Dusty is the gelding ridden by Timi, who would just as soon not be riding. He is the calmest and laziest of the group, and the easiest for a novice rider to handle. He is also the least responsive to leg pressure. Dusty is a buckskin leopard, around 15 hands tall. He has wild-type extension genes, bay alleles at the agouti locus, and one cream and one wild-type gene at the cream locus. His pattern-1 and leopard alleles are the same as Token’s. He has quite a lot of white in his mane and tail, so they are not noticeably sparse.

Penny is the guide and her horse, Freckles, is a bay leopard gelding. Freckles is a little keener than the horses assigned to Penny’s clients, but he’s a bit younger and the cross-country trip is part of his training. Freckles’s underlying bay color is a little sooty, so he has both red and black spots. Genetically he is the same as Dusty but with sooty and without the cream allele.

The other two horses have the leopard allele but are not leopards, and I’ll talk about them next time. Today I have a balky computer, so this will be a very short post. If anyone has photographs that match the horses described, I’d love to be able to use them.