Category / foal

by Equus ferus Wild Horse Photography

The Tobiano Paints

 THE TOBIANO PAINTS

Disclaimer: The mustang photographs on this blog post are presented without genetic testing; we do not know the actual chromosomal make-up of the mustangs. We rely solely upon the horse’s phenotype, or how they appear physically: coat colour, white markings, eye color, mane, and tail colour.

Mammalian Pigmentation: 

The colors found in mammalian hair, skin, irises, and some internal organs is produced by the pigment melanin. Melanin appears as colored granules in these pigmented cells and occurs in two forms, eumelanin, and phaeomelanin. Eumelanin is responsible for brown and black colour, and phaeomelanin is responsible for reds and yellows (Bailey & Brooks, 2013).
Black tobiano (muddy) (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain
The absence of melanin will appear as white in mammals. The lack of color typically associated with Paint Horses is caused by the inability of these cells to produce the base colors from birth. The color loss results in large patches of white against a base coat of any colour (bay, chestnut, roan, grey, dun, champagne, silver dapple, palomino, brown, black, etc). Additionally, the white colour has pink skin beneath. 

Assorted tobiano horses (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain

There are other forms of white colouration in equines; for example gray or roan. Grey horse color is caused by the failure of melanocytes over time, so the hair starts normally pigmented but loses the ability to maintain the pigment so the horse eventually turns white(Sponenberg, 2009). Roan horses are roan from birth although they are often not recognizable until after the foal coat has shed. Roan horses typically retain the base color on their head, legs, mane, and tail. The skin beneath grey and roan horses is dark and these colors are not actual colours, but rather modifiers that act upon a base coat (Gower, 2016). 

Black tobiano demonstrating the “shield” (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus Wild Horse Photography ™© Karen McLain 
A tobiano horse may also be roan, appaloosa, sabinos, overo, or any other pattern as the tobiano pattern is not a mutually exclusive coat patterns.
Bay Roan tobiano   (Great Desert Basin, Utah) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Roan tobiano foal, palomino tobiano mare  (Great Desert Basin, Utah) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Equine chromosomes
The genes for four white coat patterns: roan (RN), sabinos (SB1), dominant white (W), and tobianos (TO) are located on the KIT gene.The KIT gene is responsible for sending instructions through cells that allow the cell to make specific proteins. The KIT proteins are found on the cell membrane where another protein called a “stem cell factor” binds to the KIT protein. When bound together, they activate the KIT protein, which in turn, activates other proteins within the cell. These proteins serve a variety of functions in mammalian cells such as growth, development, migration, and production of certain cell types such as interstitial gastrointestinal cells and melanocytes (Haase, Jude, Brooks, & Leeb, 2008).
Equine Chromosome #3
The KIT gene is located on the fourth chromosome at the 12 position and located very close is the ECA3 gene. The ECA3, or the third equine chromosome is the location of the chromosomal mutation responsible for tobianos. Although the KIT gene remains normal in these horses, the third chromosome has an area in the gene that has flipped. Approximately one-third the length of the chromosome is an area that is an exact mirror image in tobianos horses. Because the chromosomal inversion is adjacent to the KIT gene, it affects the KIT protein synthesis, and the cells cannot produce melanocytes- so the horse has areas of white. The test for tobianos examines the chromosome and looks for a ‘break’ (telomeric or centromeric) at the positions 13 and 21 on the third chromosome– this serves as an indication they separated and inverted during replication- the horse is genetically a tobianos (Bailey & Brooks, 2009).
The gene for tobiano horses is autosomal dominant. This means to be tobiano, a foal must have at least one tobiano parent, but they also may have two tobiano parents. If one parent is a tobiano, it does not matter what colour or pattern the other parent appears; the foal will be tobiano. If a horse matches the criteria for a tobiano, it is likely the horse has the genetic background of a tobiano, although there may be some mixing of other patterns (Gower, 2016). The patterns are not inherited exactly, however, the proportion of white to colour is inherited. In other words a horse with a lot of white will have offspring with a lot of white but this depends upon the other parent. Interestingly, the study by Woolf (1990) discovered male horses and those with chestnut coats have more white than female or bay horses. The researcher also noted that the inheritance of white leg markings and facial markings is multifactorial; there are many genes involved in the appearance of white markings (Woolf, 1990).

History:

Bay tobiano mare  (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
The Tobiano paint pattern was named for General Tobías from Brazil. The General brought the paint horses to Argentina in the mid-1800’s. Before their arrival, tobiano horses were rare and had been grouped with other spotted-type horses. After General Tobías’ arrival, they were renamed after the general and placed into a unique paint coat classification(Kerson, 2015; Sponenberg, 2009)
Tobiano Characteristics:
Bay tobiano mare (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
The tobiano is defined by several coat characteristics. As with all horses, unless genetically tested, we evaluate the coat pattern by the phenotype or the horses’ physical appearance. As a general rule, tobiano horses have the following characteristics (there are always exceptions to these rules):

  1. White cross the spine somewhere between the ears and the tail (Gower, 2016; Sponenberg, 2009)
  2. The body white appears to travel down in a vertical fashion (Gower, 2016)
  3. The edges of the white areas tend to be crisp and well-defined (Sponenberg, 2009)
  4. Legs are white and the edge of the socks/stockings is irregular (Gower, 2016; Sponenberg, 2009)
  5. Most tobiano have dark eyes although some tobiano horses have blue eyes  (Sponenberg, 2009)
  6.  Most tobianos have white areas within the mane and tail, this gives the appearance of a bicoloured tail, a trait usually seen only in tobiano horses. (Sponenberg, 2009)
  7. The predominantly solid coloured heads of tobianos are generally conservatively marked: thin blazes, simple stars (Sponenberg, 2009)

 Some tobianos have very little white

Black tobiano (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Minimally marked black tobiano stallion -note the bicoloured tail (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Bay tobiano stallion (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Black tobiano stallion (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Bay tobiano stallion (McCullough Peaks, Wyoming) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Some tobiano horses have very little base colour but they tend retain normally coloured heads even when extensively white.
Minimally marked light bay chestnut tobiano (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Light bay tobiano (note the minimal blaze) (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Extensively white black marked tobiano (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain
Some tobiano horses have marks within the white areas and there may be some bleeding of colour between the base colour and the white areas. The smaller spots in the white areas are called ink spots, bear tracks, cat’s paws. The areas of darker colour encroaching on the white areas are referred to as halos. There may also be some roaning at the edge of colour and white. This is a historical link between these markings and homozygosity. No genetic link has been found, however anecdotally, homozygous horses often present with these marking whilst heterozygous generally do not show these markings. 

Cat’s paws & halo effect (San Wash Basin, Colorado) © Equus ferus- Wild Horse Photography ™ © Karen McLain

Bay tobiano with the ‘halo’ effect (Black Hills Wild Horse Sanctuary, South Dakota) © Equus ferus- Wild Horse Photography ™ © Karen McLain
References
Bailey, E., & Brooks, S. (2009). Method for screening for a tobiano coat color genotype  #USPatent 8101354 B2.
Bailey, E., & Brooks, S. (2013). Horse Genetics (2nd ed.). Boston, Massachusetts: CABI.
Gower, J. (2016). Horse Color Explained: A Breeder’s Perspective. Brattleboro, Vermont: Echo Point Books & Media, Inc.
Haase, B., Jude, R., Brooks, S. A., & Leeb, T. (2008). An equine chromosome 3 inversion is associated with the tobiano spotting pattern in German horse breeds. 
Animal Genetics, 39(3), 306-309. doi:10.1111/j.1365-2052.2008.01715.x
Kerson, N. (2015). What Color is that?  A quick guide to horse color identification: Nancy Kerson- Self Published
Sponenberg, D. (2009). Equine Color Genetics (3rd ed.). Ames, Iowa: Wiley-Blackwell.
Woolf, C. M. (1990). Multifactorial inheritance of common white markings in the Arabian horse. J Hered, 81(4), 250-256.

A special thanks to Nancy Kerson for her brilliant book “What Color is that? A quick guide to horse color identification” and to the Black Hills Wild Horse Sanctuary. A worth sanctuary for wild horses.

by Equus ferus Wild Horse Photography

The Mustang Management Contraceptive Primer

MUSTANG MANAGEMENT CONTRACEPTIVE PRIMER 

This is not a debate on which is the best method of controlling wild horse numbers. These are simply the facts. It is clear science is far from perfect but research and observation can serve to give us an idea, a general sense of something which can compel us to look for more answers and continue research, preferably as humanely and as compassionately as possible. This is also not a debate as to whether mustangs should be classified as a native species in North America, returned native species, indigenous or invasive. They are here, with limited resources, and they are our responsibility.

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Porcine Zona Pellucida (PZP):  The compound PZP, which is short for Porcina Zona Pellucida, is derived from sow ovaries. When the pigs are slaughtered for the meat industry, the excess tissue not used for the food industry is either discarded or utilized for non-consumption purposes. Some tissue used for research and others for the preparation of drugs. Heparin is a potent anticoagulant given to almost every patient who has had surgery followed by an overnight stay in the hospital, to prevent the formation of blood clots. Heparin is derived from pig intestines. Insulin, given to diabetics, was originally made from cow, pig, and even whale pancreases. Currently there are still some available that contain animal products, although there are genetically modified human insulins and insulin analogs that are not animal based (http://iddt.org/about/gm-vs-animal-insulin).


Pesticide Classification: The FDA classifies PZP as a pesticide simply because they do not have a category for contraception. Pesticides control the numbers of a populations be it insect or mammal, and because they can be quite destructive, pesticides are deemed as negative chemical compounds. PZPdoes not have any direct effect on any of the plants or animals other than the inoculated mares.

How it works: PZP works by stimulating the immune system of a mare to produce antibodies which migrate through the horse’s body to an oocyte (egg). When the mare ovulates, the antibodies immediately surround the egg, making it impenetrable to sperm. The egg cannot be fertilized and there is no foal. The reproductive behavior remains relatively normal, the mare goes into estrus and is covered by a stallion but there is no resulting offspring <!–[if supportFields]> ADDIN EN.CITE Barber200118(Barber, Lee, Steffens, Ard, & Fayrer-Hosken, 2001)181817Barber, M. R.Lee, S. M.Steffens, W. L.Ard, M.Fayrer-Hosken, R. A.Immunolocalization of zona pellucida antigens in the ovarian follicle of dogs, cats, horses and elephantsTheriogenologyTheriogenology1705-17175582001May0093-691XWOS:000168957100010<Go to ISI>://WOS:00016895710001010.1016/s0093-691x(01)00514-3<![endif]–>(Barber, Lee, Steffens, Ard, & Fayrer-Hosken, 2001)<!–[if supportFields]><![endif]–>.

Risks: PZP is not without risks. The currently long acting PZP-22 can last approximately 22 months. PZP is based on the immune system of the mares and this can cause variation in the efficacy and duration of the contraceptive effect. As in humans utilizing long term injectable contraception (Depo-Provera), the mare’s return to fertility is quite variable <!–[if supportFields]> ADDIN EN.CITE Kirkpatrick200924(Kirkpatrick et al., 2009)242417Kirkpatrick, J. F.Rowan, A.Lamberski, N.Wallace, R.Frank, K.Lyda, R.The practical side of immunocontraception: zona proteins and wildlifeJournal of Reproductive ImmunologyJournal of Reproductive Immunology151-157831-22009Dec0165-0378WOS:000273024800028<Go to ISI>://WOS:00027302480002810.1016/j.jri.2009.06.257<![endif]–>(Kirkpatrick et al., 2009)<!–[if supportFields]><![endif]–>. The reason PZP is not offered to humans is because the efficacy rate is not high enough.

“For contraceptive treatment to be an effective management tool, it usually needs to be reversible (Kirkpatrick & Turner 1991). A long term study of feral horses showed that PZP was reversible even when females were treated for several years (Kirkpatrick & Turner 2002). However some females appeared not to return to full fertility after long-term PZP treatment and similar side effects were seen with GNRH treatments in deer (e.g. Miller et al. 2000a). Consequently, most wildlife contraceptives are reversible, or have minimal impact after prolonged use.”  (Gray & Cameron, 2010).

Prolonged use has demonstrated that some mares will never return to fertility. Kirkpatrick, Liu, Turner, Naugle, and Keiper (1992)<!–[if supportFields]><![endif]–> found that three factors determine the return to normal reproductive function: the amount of PZP administered, the number of antibodies produced by the mare, and ovarian dysfunction. Earlier studies also demonstrated damage to ovaries although the PZP preparation was crude in the earlier stages of development <!–[if supportFields]> ADDIN EN.CITE Kirkpatrick199226(Kirkpatrick et al., 1992)262617Kirkpatrick, J. F.Liu, I. M. K.Turner, J. W.Naugle, R.Keiper, R.LONG-TERM EFFECTS OF PORCINE ZONAE-PELLUCIDAE IMMUNOCONTRACEPTION ON OVARIAN-FUNCTION IN FERAL HORSES (EQUUS-CABALLUS)Journal of Reproduction and FertilityJournal of Reproduction and Fertility437-4449421992Mar0022-4251WOS:A1992HR77000018<Go to ISI>://WOS:A1992HR77000018<![endif]–>(Kirkpatrick et al., 1992)<!–[if supportFields]><![endif]–>.

Abscesses at the injection sites have been reported but these are temporary, and heal without complications.

PZP and Tuberculosis: Finally, there were some rumours floating around social media that PZP can cause tuberculosis. Although this may sound like science-fiction or the nefarious work of people against keeping the horses wild, there is some truth. The original method of getting PZP into the animals involved piggy-backing the molecule on a carrier molecule or adjuvant. Adjuvants are not biologically active but their presence can trigger an immune response. It may result in a false positive antibody response for tuberculosis. The animal doesn’t have the disease, and many human vaccines work this way by stimulating the body to form antibodies to something not biologically active.  The original choice for the adjuvant was a mycobacterium- the mycobacteria family are known to cause tuberculosis and many other diseases. Because the PZP was attached to an inactive mycobacterium, in some animals it cause a false-positive tuberculosis antibody response. They changedthe adjuvant for the preparation of PZP so now there is no mycobacterium involved. 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GONACON: This method works by injecting mares with a synthetic (man-made) chemical compound called GonaCon. This compound acts against a hormone called gonadotropin releasing hormone (GnRH). In the reproductive cycle of mammals, GnRH is produced by the anterior pituitary gland. This gland controls reproduction by stimulating the release of the follicle stimulating hormone and the luteinizing hormone (and others). ConaCon works by stopping the production of GnRH and subsequently the luteinizing hormone and without LH, there is no ovulation, no corpus luteum, and therefore, no egg/offspring <!–[if supportFields]> ADDIN EN.CITE Speroff201230(Speroff & Fritz, 2012)30306Speroff, L.Fritz, M. Clinical Gynecologic Endocrinology and Infertility 8th2012New Yok Lippincott Williams & Wilkins<![endif]–>(Speroff & Fritz, 2012)<!–[if supportFields]><![endif]–>. Like PZP, GonaCon can result in prolonged infertility (Ransom, 2014). The tables below represent one study each- these data are only reflecting the results one study and may not have generalisability to the entire population. The general consensus amongst zoos and researchers is that PZP is 90% effective when administered correctly.

In a study by <!–[if supportFields]> ADDIN EN.CITE Ransom201429Jason I. Ransom et al. (2014)292917Ransom, Jason I.Powers, Jenny G.Garbe, Heidi M.Oehler Sr, Michael W.Nett, Terry M.Baker, Dan L.Behavior of feral horses in response to culling and GnRH immunocontraceptionApplied Animal Behaviour ScienceApplied Animal Behaviour Science81-92157Equus caballusFertility controlGonadotropin releasing hormoneSocial behaviorWild horseWildlife contraception20148//0168-1591http://www.sciencedirect.com/science/article/pii/S016815911400135Xhttp://dx.doi.org/10.1016/j.applanim.2014.05.002<![endif]–>Jason I. Ransom et al. (2014)<!–[if supportFields]><![endif]–>, the researchers found there were fewer behavioural differences in mares treated with GonaCon, compared to those treated with PZP. They modeled their GonaCon study after the PZP study and found fewer alterations in the wild mare’s behavior. GonaCon is still a recent addition to the world of wildlife contraception and has potential as a potential management tool for equids. It shows promise but the long-term data is still unavailable.  


GonaCon can be given to males because the GnRH stimulates testosterone production in males. However, studies of stags treated with GonaCon resulted in antler deformity and other negative consequences. “In conclusion, the GnRH vaccination in male rusa deer resulted in the increase in GnRH antibody titer, which negatively correlated with blood testosterone. The decrease in blood testosterone might be involved in the lower semen quality and poor antler development” (Phraluk, O. et al, 2015). There is potential for use in stallions but we need more research in this area.


Because GonaCon a works systemically, not targeting the reproductive tract as specifically as PZP, the potential for side effects increases. The closer to the intended target a medication or treatment is administered, the more effective, the lower the dose, and adverse drugs reaction are substantially decreased. The Global Library of Women’s Health states: “In non-reproductive tissues, GnRH has been reported to modulate neuronal migration, visual processing, digestive tract function, and immune T cell chemotaxis. Studies in endometrial, ovarian, and prostate tumor cell lines have implicated GnRH in mediating cell growth, angiogenesis, invasion, and metastasis.” (http://www.glowm.com/section_view/heading/Gonadotropin-releasing%2520Hormone%2520(GnRH)%2520and%2520the%2520GnRH%2520Receptor%2520(GnRHR)/item/284)
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Table 1. (Represents data from one study)
Infertility over three-years with equine contraception (Kilian et al., 2006).
Infertility after one year
Infertility after two years
Infertility after three years
Spay- Vac PZP
100%
80%
80%
GonaCon
94%
60%
53%
Copper IUD
80%
29%*
14%*
*The assumption is the IUD’s had been expelled in the mare who became pregnant

Table 2. (Represents data from another study)

Foaling rates at three horse management areas after PZP treatment  (Ransom, J.,et al, 2011).
Foaling Rates
Type of contraception
Treated
Untreated
Little Book Cliffs
6.6%
60.1%
PZP in liquid form requires annual boosters
McCullough
31.7%
75%
PZP in pelleted form- designed to last two years
Pryor Mountain
17.7%
62.8%
PZP in liquid form requires annual boosters
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Any time we begin to tamper with Mother Nature- it places us at risk. Treating mares with immunocontraception and/or Gonacon can have negative consequences on the familial structures of a highly social animal. Treating mares with these two compounds can result in mare giving birth to foal during time of limited resources. A study determined that there are differences in parturition times for mare treated compared with mares that were not treated. Foals born during more normal times for wild horses have a higher survival rate. <!–[if supportFields]> ADDIN EN.CITE Ransom201331(J. I. Ransom, Hobbs, & Bruemmer, 2013)313117Ransom, J. I.Hobbs, N. T.Bruemmer, J.Contraception can Lead to Trophic Asynchrony between Birth Pulse and ResourcesPlos OnePlos One812013Jan1932-6203WOS:000315211500043<Go to ISI>://WOS:000315211500043e5497210.1371/journal.pone.0054972<![endif]–>(J. I. Ransom, Hobbs, & Bruemmer, 2013)<!–[if supportFields]><![endif]–>.  

However, all negative consequences of injectable contraception pale in comparison to the disruption of the social structure during round-ups. Separation of mares from stallions and their offspring occurs during round-ups and culling. Family bands are broken up and the horses face the terrible loss of their freedom. A balance must be found and the benefits and the negative outcomes must be weighed. There is a chance a mare treated with the above methods, may never foal again- but that mare remains free. There is always the potential she may resume ovulation however, she is will not spend her life in a holding pen. A study by Turner, Liu, Flanagan, and Rutberg (2007) indicated one mare out of sixteen in the study, did not have a normal return to fertility. Is a chance of infertility worth the price of freedom?
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OVARECTOMY: This is removal of a mare’s ovaries. This method is irreversible, the mare will never breed again. Behaviorally, it reverts a mare to a non-breeding state. There is not a lot of data and/or research regarding this permanent method in the wild horse population which is why research proposals are being requested by government agencies The risks are the this surgical procedure are post-operative infection and possible abortion of a fetus if the mare is gravid at the time of the procedure.  Additionally there is the loss of that mare’s potential contribution to the genetic diversity of that herd management area.  This loss is not well defined and may vary based of the genetic variation in a herd management area. Mares should be monitored for three weeks post-operative (optimally), before being returned to the range to live out their lives. The upfront cost is high, but the end result is permanent. It has been proposed for mare over a certain age, selected by the currently proposed studies, as means to control the populations which would allow the mares to live free without foaling year after year (Speroff & Fritz, 2012, National Research Council, 2013).


CHEMICAL STERILIZATION/VASECTOMY/NEUTERING:  These methods, performed on stallions, are permanent means of limiting wild horse populations. In chemical sterilization, stallions have a solution injected into the testes which causes necrosis and eventual tissue death of the testes. It is painful and carries a high risk of infection but is very cost-effective (Zhanwei, 1989),  Orchiectomy (removal of testes: aka gelding/neutering) is another permanent method which removes the testes of a stallion. Like chemical sterilization, ‘gelding’ causes behavioral changes and the stallions become less aggressive, there is less fighting and they cannot reproduce. Vasectomy involves severing the vas deferens of a stallion so that the communication between the testes and the penis is removed. The sperm are not able to be passed through the urethra during copulation. Behavior remains the same because the testes are still present and producing testosterone. Infection rates are much lower in vasectomies, the procedure is less painful but it requires a delicate touch, it may not be 100% effective, and the horse must be anesthetized or sedated (Speroff & Fritz, 2012, National Research Council, 2013).

ROUND-UPS: This method involves gathering the horses by use of helicopters, men on horses, ATV, trucks, and bait trapping. The bait-trapping is the least likely to cause physical harm but none of these methods are without significant risks. Helicopter round-ups have the highest incidences of morbidity and mortality.  The horses are gathered, separated and either returned to the horse management area, or they are removed to a holding facility for potential adoption.

RESERVE DESIGN:  This method of mustang management has merit in a perfect world. The theory is to find a place for the horses to live. A place that has natural horse-predators (may prove to be difficult to find), natural barriers to prevent migration in/out of the area, and neighbours sympathetic to mustangs.  The area has to have sufficient food, water, shelter, and other necessities for survival year round. Funds for this preserve may be obtained through ‘eco-tourism’ and public education regarding the wild horses are included in this plan.  Reserve design is a wonderful idea but it does not have practical application. The land that would be needed is not available, at least not at this time, for the numbers necessary.  The monitoring of these herds and the management of these herds must be carried out as well and the necessary people to oversee each area. The goal of ‘reserve design’ is to be self-sufficient in which the wild horses achieve homoeostasis with regards to population growth. Because the predation is very low, and we cannot safely import predators, this will prove to be challenging. Incorporating natural predators to assist in controlling the wild horse population can have deleterious effects on the domestic population of horses and of livestock. Historically, once a predator begins to hunt within the domestic population, the end result usually has rather negative consequences for the predator.


SELF-REGULATION: This involves leaving the mustangs alone to establish population equilibrium without any interference. There is no scientific evidence that this methods works and history has shown us wild horses do not fare well with a hands-off approach. Unfortunately the resources the mustangs have within the management areas are limited and the horses will suffer in one way or another if left unmanaged. There are three factors that determine a population’s ability to grow. They are: available resources, predation, and disease. Many of the wild horses at Cold Creek starved to death or had to be humanely euthanized because resources became compromised.  The resources may be limited due to naturally occurring factors such as drought or fire, or they may face competition from livestock grazing on the same land. Regardless of the mechanism, reduced resources will cause competition, and result starvation or disease in the wild horse population. Occasionally a mountain lion will take a foal or an older/injured horse but cougars are not primary predators of the mustangs . Wolves generally do not live in the horse management areas with any regularity. Disease is a concern with any animal left in overcrowded situations. Chronic Wasting Disease is a disease that began in the mule deer population, and has spread to other cervids (herbivorous even-toed mammals in which the male carries antlers). The current deer populations are substantially larger than the available resources in the north-east United States, and this disease has now been identified in white-tailed deer of the Adirondack Mountains. Similar to Bovine Spongiform Encephalopathy (BSE), or Mad Cow Disease, this disease is affecting all cervids, not just the deer (Chronic Wasting Disease Alliance, 2016).  If the populations of wild horses were left alone, they would increase to the point of starvation and disease. It would only be a matter of time before a disease mutated in that population and spread to the domestic horse population with devastating consequences.

DECLARING THE MUSTANG “ENDANGERED”: This method is to first declare the mustang a ‘native wild species’ and then have it declared endangered- which may prove difficult. The mustang is the same species as domestic horses. They fall within the same genera: Equus and species: caballus. They have no genetic markers or any other characteristic that differentiate them from their domestic counterparts.  There has been proposed theories that they behave differently but this was not enough for the United States Fish and Wildlife to declare mustangs as a separate subspecies.  The wild horses are identical genetically, physiologically, and behaviorally to the horses in your paddock.  There are currently 40-50,000 presumably wild horses in captivity at BLM holding facilities. There are an estimated 20-40,000 living wild on horse management areas, and several thousand more domesticated mustangs living with people. These numbers alone are not sufficient for endangered species status, or even threatened species if USFW was willing to grant them subspecies status. 

Proponents of making mustangs endangered believe that once they achieve the endangered species status, the mustangs would be granted the ultimate protection. However, advocacy groups would no longer have a say in their conservation; they would be managed by Fish and Wildlife. The now ‘endangered’ mustangs may be moved to locations to protect their numbers and they may very well lose their freedom if they were to ever to gain protected status. Our descendants would not be able to see these ‘endangered mustangs’ living free; they would only see them in zoos and protected reserves.


The belief that mustangs are a separate species or a separate animal from domestic horses is the first hurdle to overcome with this method. However, that has proven to be impossible. They are not separate; they are the same species just as a miniature horse and a Clydesdale are the same species. They are horses that have returned to the wild and been wildly successful at surviving and reproducing.   

“Managing them to extinction” is a catch-phrase often used to describe the situation of the wild horses. Mustangs will never become extinct because they aren’t recognized as a separate sub-species of the modern horse. However, there is a very good chance our children and their descendants will never see a free roaming mustang, and that would be the greatest tragedy of all. Regardless of their origin, regardless of their heritage, the mustangs are our responsibility. We need science to save the mustangs, scientists with the necessary credentials and expertise in wildlife management, ecology, contraceptive experts, and equine ethology all working to establish the best and most humane method of managing wild horses. Each management area is unique and each management area needs its own method of achieving appropriate and healthy numbers. The phrase “managing them to eradication” is more accurate, less sensational rhetoric and implies the same message without any controversy regarding the species/subspecies status of the wild mustangs.  


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Meredith Hudes-Lowder is a Nurse Practitioner in Women’s Health and an expert on contraception. Additionally, she has a bachelor’s of biology with a concentration in environmental education.  She will graduate in May 2016 with a Doctorate of Nursing Practice. She runs the largest exclusively wild horse photography site on Facebook with Karen McLain- they have almost half a million fans. Bruce Lowder was consulted for this blog. He is a wildlife expert, naturalist, and worked for U.S. Fish and Game.
References:

Barber, M. R., Lee, S. M., Steffens, W. L., Ard, M., & Fayrer-Hosken, R. A. (2001). Immunolocalization of zona pellucida antigens in the ovarian follicle of dogs, cats, horses and elephants. Theriogenology, 55(8), 1705-1717. doi:10.1016/s0093-691x(01)00514-3
Gray, M. E., & Cameron, E. Z. (2010). Does contraceptive treatment in wildlife result in side effects? A review of quantitative and anecdotal evidence. Reproduction, 139(1), 45-55. doi:10.1530/rep-08-0456


Chronic Wasting Disease Alliance (2016). Webage URL http://www.cwd-info.org/index.php/fuseaction/about.main. Accessed 01/11/2016.

Killian, G., Diel, N., Miller, L., Rhyan, J, Thain, D. (2006). Long-Term Efficacy of Three Contraceptive Approaches for Population Control of Wild Horses. USDA National Wildlife Research Center – Staff Publications.

Kirkpatrick, J. F., Liu, I. M. K., Turner, J. W., Naugle, R., & Keiper, R. (1992). LONG-TERM EFFECTS OF PORCINE ZONAE-PELLUCIDAE IMMUNOCONTRACEPTION ON OVARIAN-FUNCTION IN FERAL HORSES (EQUUS-CABALLUS). Journal of Reproduction and Fertility, 94(2), 437-444.  Retrieved from ://WOS:A1992HR77000018
Kirkpatrick, J. F., Rowan, A., Lamberski, N., Wallace, R., Frank, K., & Lyda, R. (2009). The practical side of immunocontraception: zona proteins and wildlife. Journal of Reproductive Immunology, 83(1-2), 151-157. doi:10.1016/j.jri.2009.06.257
Lyda, R. O., Hall, J. R., & Kirkpatrick, J. F. (2005). A comparison of Freund’s Complete and Freund’s Modified Adjuvants used with a contraceptive vaccine in wild horses (Equus caballus). J Zoo Wildl Med, 36(4), 610-616. doi:10.1638/04104.1

National Research Council, (2013). Using Science to Improve the BLM Wild Horse and Burro Program: A Way Forward. PDF accessed 01/12/16: http://www.nap.edu/catalog/13511/using-science-to-improve-the-blm-wild-horse-and-burro-program.

Phraluk, Orasa; Wajjwalku, Worawidh; Siriaroonrat, Boripat; Booddee, Orawan; Thongtip, Nikorn. (2015). Effects of immunization against gonadotropin releasing hormone on reproductive functions in male rusa deer (Rusa timorensis). The Thai Journal of Veterinary Medicine 45.1   (Mar 2015): 1,3-10.
Ransom, J.,   Roelle, J.,  Cade, B., Coates-Markle, L., Kane, A., Ransom, Jason I., Roelle, James E., Cade, Brian S.,Coates-Markle, L., & Kane, A. (2011) Foaling Rates in Feral Horses Treated With the Immunocontraceptive Porcine Zona Pellucida. WILDLIFE SOCIETY BULLETIN.
Ransom, J. I., Hobbs, N. T., & Bruemmer, J. (2013). Contraception can Lead to Trophic Asynchrony between Birth Pulse and Resources. Plos One, 8(1). doi:10.1371/journal.pone.0054972
Ransom, J. I., Powers, J. G., Garbe, H. M., Oehler Sr, M. W., Nett, T. M., & Baker, D. L. (2014). Behavior of feral horses in response to culling and GnRH immunocontraception. Applied Animal Behaviour Science, 157, 81-92. doi:http://dx.doi.org/10.1016/j.applanim.2014.05.002
Speroff, L., & Fritz, M. (2012). Clinical Gynecologic Endocrinology and Infertility (8th ed.). New Yok: Lippincott Williams & Wilkins.
Turner, J. W., Liu, I. K. M., Flanagan, D. R., & Rutberg, A. T. (2007). Immunocontraception in wild horses: One inoculation provides two years of infertility. Journal of Wildlife Management, 71(2), 662-667. doi:10.2193/2005-779

Zhanwei, S. (1989). Chemical castration of horses and mules by injecting testis with iodine tincture. Journal of Liaoning Animal Husbandry and Veterinary Medicine.

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Other resources:

Barber, M. R., Lee, S. M., & Fayrer-Hosken, R. A. (1998). Staining patterns to the zona pellucida of the dog, cat, horse and elephant with porcine zona pellucida (pZP) antisera. Theriogenology, 49(1), 307-307. doi:10.1016/s0093-691x(98)90660-4
Barber, M. R., Lee, S. M., Steffens, W. L., Ard, M., & Fayrer-Hosken, R. A. (2001). 

Immunolocatiom of zona pellucida antigens in the ovarian follicle of dogs, cats, horses and elephants. Theriogenology, 55(8), 1705-1717. doi:10.1016/s0093-691x(01)00514-3


Berger, J. (1983). Induced abortion and social factors in wild horses. Nature, 303(5912), 59-61.  Retrieved from http://dx.doi.org/10.1038/303059a0

Bowling, A. T., & Touchberry, R. W. (1990). Parentage of Great Basin Feral Horses. The Journal of Wildlife Management, 54(3), 424-429. doi:10.2307/3809652

Duncan, P. (1982). Foal killing by stallions. Applied Animal Ethology, 8(6), 567-570. doi:http://dx.doi.org/10.1016/0304-3762(82)90221-8
Feh, C. (1990). Long-term paternity data in relation to different aspects of rank for camargue stallions, Equus caballus. Animal Behaviour, 40(5), 995-996. doi:http://dx.doi.org/10.1016/S0003-3472(05)81007-3

Feh, C., & Munkhtuya, B. (2008). Male infanticide and paternity analyses in a socially natural herd of Przewalski’s horses: sexual selection? Behav Processes, 78(3), 335-339. doi:10.1016/j.beproc.2007.12.009

Gray, M. E. (2009). An infanticide attempt by a free-roaming feral stallion (Equus caballus). Biol Lett, 5(1), 23-25. doi:10.1098/rsbl.2008.0571

Gray, M. E., & Cameron, E. Z. (2010). Does contraceptive treatment in wildlife result in side effects? A review of quantitative and anecdotal evidence. Reproduction, 139(1), 45-55. doi:10.1530/rep-08-0456
Gray, M. E., Thain, D. S., Cameron, E. Z., & Miller, L. A. (2010). Multi-year fertility reduction in free-roaming feral horses with single-injection immunocontraceptive formulations. Wildlife Research, 37(6), 475-481. doi:10.1071/wr09175
Jordana, J., Pares, P. M., & Sanchez, A. (1995). ANALYSIS OF GENETIC-RELATIONSHIPS IN HORSE BREEDS. Journal of Equine Veterinary Science, 15(7), 320-328. doi:10.1016/s0737-0806(06)81738-7
Kavar, T., & Dovc, P. (2008). Domestication of the horse: Genetic relationships between domestic and wild horses. Livestock Science, 116(1-3), 1-14. doi:10.1016/j.livsci.2008.03.002

Kirkpatrick, J. F., Liu, I. M. K., Turner, J. W., Naugle, R., & Keiper, R. (1992). LONG-TERM EFFECTS OF PORCINE ZONAE-PELLUCIDAE IMMUNOCONTRACEPTION ON OVARIAN-FUNCTION IN FERAL HORSES (EQUUS-CABALLUS). Journal of Reproduction and Fertility, 94(2), 437-444.  Retrieved from ://WOS:A1992HR77000018

Kirkpatrick, J. F., Rowan, A., Lamberski, N., Wallace, R., Frank, K., & Lyda, R. (2009). The practical side of immunocontraception: zona proteins and wildlife. Journal of Reproductive Immunology, 83(1-2), 151-157. doi:10.1016/j.jri.2009.06.257

Kirkpatrick, J. F., & Turner, A. (2002). Reversibility of action and safety during pregnancy of immunization against porcine zona pellucida in wild mares (Equus caballus). Reproduction (Cambridge, England) Supplement, 60, 197-202.  Retrieved from http://europepmc.org/abstract/MED/12220160

Linklater, W. L., Cameron, E. Z., Minot, E. O., & Stafford, K. J. (1999). Stallion harassment and the mating system of horses. Anim Behav, 58(2), 295-306. doi:10.1006/anbe.1999.1155

Linklater, W. L., Cameron, E. Z., Stafford, K. J., & Minot, E. O. (2013). Removal experiments indicate that subordinate stallions are not helpers. Behav Processes, 94, 1-4. doi:http://dx.doi.org/10.1016/j.beproc.2013.02.005

Lyda, R. O., Hall, J. R., & Kirkpatrick, J. F. (2005). A comparison of Freund’s Complete and Freund’s Modified Adjuvants used with a contraceptive vaccine in wild horses (Equus caballus). J Zoo Wildl Med, 36(4), 610-616. doi:10.1638/04104.1

Madosky, J. M., Rubenstein, D. I., Howard, J. J., & Stuska, S. (2010). The effects of immunocontraception on harem fidelity in a feral horse (Equus caballus) population. Applied Animal Behaviour Science, 128(1–4), 50-56. doi:http://dx.doi.org/10.1016/j.applanim.2010.09.013

Mask, T. A., Schoenecker, K. A., Kane, A. J., Ransom, J. I., & Bruemmer, J. E. (2015). Serum antibody immunoreactivity to equine zona protein after SpayVac vaccination. Theriogenology, 84(2), 261-267. doi:10.1016/j.theriogenology.2015.03.012

Massei, G., & Cowan, D. (2014). Fertility control to mitigate human-wildlife conflicts: a review. Wildlife Research, 41(1), 1-21. doi:10.1071/wr13141

Nuñez, C. M. V., Adelman, J. S., Mason, C., & Rubenstein, D. I. (2009). Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season. Applied Animal Behaviour Science, 117(1–2), 74-83. doi:http://dx.doi.org/10.1016/j.applanim.2008.12.001

Pluhacek, J., & Bartos, L. (2000). Male infanticide in captive plains zebra, Equus burchelli. Anim Behav, 59(4), 689-694. doi:10.1006/anbe.1999.1371

Ransom, J. I., Hobbs, N. T., & Bruemmer, J. (2013). Contraception can Lead to Trophic Asynchrony between Birth Pulse and Resources. Plos One, 8(1). doi:10.1371/journal.pone.0054972

Ransom, J. I., Powers, J. G., Garbe, H. M., Oehler Sr, M. W., Nett, T. M., & Baker, D. L. (2014). Behavior of feral horses in response to culling and GnRH immunocontraception. Applied Animal Behaviour Science, 157, 81-92. doi:http://dx.doi.org/10.1016/j.applanim.2014.05.002

Schulman, M. L., Botha, A. E., Muenscher, S. B., Annandale, C. H., Guthrie, A. J., & Bertschinger, H. J. (2013). Reversibility of the effects of GnRH-vaccination used to suppress reproductive function in mares. Equine Veterinary Journal, 45(1), 111-113. doi:10.1111/j.2042-3306.2012.00577.x

Speroff, L., & Fritz, M. (2012). Clinical Gynecologic Endocrinology and Infertility (8th ed.). New Yok: Lippincott Williams & Wilkins.

Turner, J. W., Liu, I. K. M., Flanagan, D. R., & Rutberg, A. T. (2007). Immunocontraception in wild horses: One inoculation provides two years of infertility. Journal of Wildlife Management, 71(2), 662-667. doi:10.2193/2005-779



by Equus ferus Wild Horse Photography

Lethal White

Fleck and her foal 2015, most likely sire Picasso

Equus ferus– Equine Coat Colour Genetic -LETHAL WHITE

One of the mares at the Sand Wash Basin was believed to have given birth to a foal with Lethal White Syndrome.  And although we cannot be absolutely sure the foal was a Lethal White foal, the behaviour strongly suggests it might have been. Lethal White Syndrome has been talked about in groups and on Facebook, hence this blog post. Without a necropsy (autopsy on an animal) we won’t know for certain but here is what we do know…

 
Similar to Hirschsprung’s disease in humans, Lethal White Syndrome affects the colon by making it non-functional and in horses it also affects pigment of the coat. The affected foals are born pure white with blue/grey eyes and occasionally a smudge or darker colour on the body or near the tail however, they die within 72 hours of birth. The colon in these foals is a dead-end and the foals cannot pass feces. They do not act normally and exhibit signs of distress.

 

Lethal white foal (Picasso x Mingo) photo credit Nancy Roberts 2011
 
 
Picasso and Mingo 2012 photo credit Karen McLain
 
The trait, which is inherited, is carried by the horses who also carry the paint trait frame overo. Frame Overo horses typically have jagged white markings along the center of the body. The back and belly may remain solid colored so the effect is a framed area of white. They may have white faces (apron or bald face) and they may have blue eyes although not always. Some horses may minimally express the trait and the only evidence of the frame overo paint trait is a little spot of white along the neck and an unusually shaped blaze. Some horses may also carry other paint traits such as tobiano and they horses are referred to as toveros. Without genetic testing, nothing is certain so we are basing our assumptions on what we have observed and the reproductive history of the individual mustangs.
Yahtzee (R) & Van Gogh (L)
Photo credit Meredith Hudes-Lowder of Equus ferus Wild Horse Photography
In order to produce a foal with Lethal White Syndrome, both parents must be overo. Not all overo horses carry the trait according to the American Paint Horse Association (http://www.apha.com/breed/geneticsarticles/lethal-whites-1) and not all blue-eyed white foals carry the Lethal White gene. Because  Lethal White Syndrome is autosomal recessive, it means when two horses that are overo and each carry the gene , there is a 25% chance the foal with be born with the syndrome. If a dam with the trait and a sire with the trait have three normal foals, it does not mean the fourth foal will carry the trait; the chances a foal will inherit the syndrome resets each gestation and remains one in four with each subsequent preganncy. 
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In the Punnett Square below (Horse drawings by Karen McLain)
Oo outside the square on top represent the SIRE
Along the left side, the Oo represents the DAM
Both are Overo represented Oo and the carry the trait

 

 

The RESULT:
One Solid foal (unaffected) -25%
Two Overo foals- CARRIERS of the Lethal White Trait- 50%
One Lethal White Foal 25%

 

One Solid Foal (OO) 25%
One Overo Foal (Oo) 25%
ONe Overo Foal (Oo) 25%
One Lethal White Foal (oo) 25%
 
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PUNNETT SQUARE REPRESENTING AUTOSOMAL RECESSIVE INHERITANCE

 
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The photos below are by Danielle M. Williams and they show Fleck’s foal. Fleck is frame overo and the father is believed to be Eagle, a minimally marked frame overo so it is quite possible the foal carries the Lethal White Syndrome.  Sometimes Lethal White Foals do have darker pigment on the muzzle but this foal is not hunched over in the typical posture of a horse in gastric distress however the witness/photographer Danielle did say the foal did not look well and laid down frequently. She said the foal was unable to stand for any length of time. This is consistent with Lethal White Syndrome and the foals with the disorder often roll from side to side. Another possibility is that the foal may have perished in the fight between the band stallion Eagle and Diego who took over part of the band. Stallion infanticide is unfortunately unavoidable and may be more common than previously thought. Regardless of the manner of death, it is heartbreaking to see a young life extinguished so soon.
 
Fleck and her foal
Photo by Danielle M. Williams
 
Fleck and her foal
Photo by Danielle M. Williams
 
 
Fleck and her foal
Photo by Danielle M. Williams
Please email Meredith with any questions regarding the genetic behind Lethal White Syndrome or horse colour genetic in general.
Equus ferus- Wild Horse Photography
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Dr. Meredith Hudes-Lowder DNP, WHNP-BC, MSN, BSN, RNC, BS Biology
Meredith received a Bachelor of Science Degree from Binghamton University with an emphasis in ethology and genetics. She received a Bachelor of Science in Nursing also from Binghamton and a Masters of Nursing in Perinatal/Women’s Health from Stony Brook University. She currently practices medicine as a Nurse Practitioner in Manhattan for Advantage Care Physicians. She is also enrolled in the Doctoral program at Stony Brook and graduated in 2016 with a Doctorate of Nursing Practice. Her doctoral thesis was a research study on cervical cancer screening intervals. She is a member of several professional organizations and was inducted into Sigma Theta Tau- the nursing honor society in 2007.