Chemical Composition of Lick Soils - Functions of Soil Ingestion by Four Ungulate Species

7/13/22, 11:01 AM Chemical Composition of Lick Soils: Functions of Soil Ingestion by Four Ungulate Species
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Chemical Composition of Lick Soils: Functions of Soil Ingestion by Four Ungulate Species
October 2006 · Journal of Mammalogy 87(5):878-888

DOI:10.1644/06-MAMM-A-055R1.1
Authors:
Jeremy B. Ayotte Katherine L. Parker Joselito M. Arocena Michael P. Gillingham
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Citations (85) References (100)
Abstract
Perspectives on the importance of natural salt licks to ungulates have been

broadened beyond the role of providing sodium. This study examined the Discover the world's
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chemical compositions of wet and dry licks in north-central British Columbia, and
defined the benefits of licks to elk (Cervus elaphus), moose (Alces alces),
20+ million
Stone's sheep (Ovis dalli stonei), and mountain goats (Oreamnos americanus). members
We analyzed soils for buffering compounds (carbonates) and estimated
available elements from extractions with solutions at a low, digestive-tract pH. 135+ million
Even though the 2 types of licks are visually distinct, with different publications
concentrations of chemical components, they serve similar functions as 700k+ research
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concentrated sources of sodium, carbonates, magnesium, and sulfate. Sodium projects
and sulfate concentrations were typically higher at both wet and dry licks than
control sites. Carbonates and magnesium were elevated in soils from dry licks
and magnesium also was high in inflow waters to wet licks. We estimated
elemental intake by ungulates from the composition of forage samples. Forages
used by all 4 ungulate species were too low in sodium to meet requirements.
Spring and summer forages contained high potassium levels. Licks in our study,
therefore, provide ungulates with supplemental sources of sodium that are
particularly beneficial to offset increasing demands during lactation and with
carbonates to help stabilize rumen pH after forage changes in spring.
Supplemental sources of magnesium may be actively sought by ungulates when
high levels of dietary potassium affect absorption. To assess the importance of
clay in soils ingested at licks, we determined the clay mineral types at licks and
compared the buffering capacity of clay-filled fecal material collected at licks
with fecal material collected away from licks. Further studies are needed to
define the roles of clay in improving forage palatability and digestibility.
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Citations (85) References (100)
... Sodium plays a significant role in pregnant and lactating animals especially on maintenance of acid
base balance. However, studies have revealed presence of higher levels of other essential minerals in
the licks thus refuting arguments that sodium is the only reason for geophagy (Ayotte, 2006; Houston,
2011;and Nderi et al., 2015). For example, Katherine et al. (2004) reported high levels of magnesium for
wet and dry licks, which is thought to supplement diets when there are high concentrations of potassium
in spring forage that reduces absorption of magnesium by ruminants, thus interfering with rumen
fermentation. ...
... Results in Table 4, showed that there was no significant difference on digestible nutrients intake of the
basal diet (CP, CF, EE, NFE, TDN) (P > 0.05) due to consumption of natural licks from different sources
and a commercial lick. Ingestion of natural licks has been reported in both domestic and wild animals,
and it is associated with mineral supplementation, detoxification of plant secondary metabolites and
alleviation of digestive disorders (Ayotte et al., 2006) . This study shows that there were no significant
differences on lick intake (P > 0.05) among goats, which supports results of a previous study on sheep
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in Ethiopia (Sisay etDownload
al., 2007).citation
... Copy link
... The milk production and quality from goats fed on licks from different sources is presented in (Ayotte
et al., 2006) . The highest milk yield that was found in Kang'au lick might be related to the relatively
higher dry matter intake that was associated with this lick (Table 3) which led to ingestion of more
nutrients that was translated into milk production. ...
EFFECTS OF NATURAL LICKS ON FEED INTAKE, NUTRIENT DIGESTIBILITY, MILK PRODUCTION AND
QUALITY IN KENYA ALPINE DAIRY GOAT RATION
Article Full-text available
Nov 2015
Onesmus Nderi
View Show abstract
... Common warthogs salt lick mostly in the morning and afternoon hours [20]. The greatest peaks in
visits of such sites occur from December to February [6, 16,25]. The peak periods visit lick site coincide
with plant phenology (variable timing of plant growth and forage preferences) [17,27]. ...
... The peak periods visit lick site coincide with plant phenology (variable timing of plant growth and
forage preferences) [17,27]. The visitation periods associated with the increased physiological demands
of growth, or weight regain, motivated by electrolyte loss related to stress of sudden changes in forage
chemistry of the animal [6, 16,26,29]. Common warthogs use wet ( Figure 1) and dry licks ( Figure 2)
[6,17,26]. Wet lick is linked with the change in forage chemistry and the loss of sodium from sweat or
urine is much more than other times of the year [26] and underground water springs [6]. ...
... The visitation periods associated with the increased physiological demands of growth, or weight
regain, motivated by electrolyte loss related to stress of sudden changes in forage chemistry of the
animal [6,16,26,29]. Common warthogs use wet ( Figure 1) and dry licks ( Figure 2) [6, 17,26]. Wet lick
is linked with the change in forage chemistry and the loss of sodium from sweat or urine is much more
than other times of the year [26] and underground water springs [6]. Dry licks usually occur along
streams or riverbeds because un weathered soluble elements are deposits above less firm layers, and
become exposed by erosion as dry licks [6]. ...
Chemical Composition of Lick Soils and Functions of Soil Ingested by Common Warthogs (
Phacochoerus africanus ) in Dabena Valley Forest, Western Ethiopia
Article
Jan 2021
Alemayehu Edossa · Afework Bekele · Habte Jebessa Debella
View Show abstract
... A salt lick may take the form of a mineral water spring (Chong et al., 2005;Panichev et al., 2013;Sim
et al., 2020), dry land (Ramachandran et al., 1995;Chong et al., 2005), muddy pool (Brightsmith et al.,
2008;Razali et al., 2020;Sim et al., 2020) or rocks rich in specific minerals (Panichev et al., 2013;Razali
et al., 2020;Sim et al., 2020). Salt licks are abundant in temperate and tropical forests (Chong et al.,
2005; Ayotte et al., 2006; Tobler et al., 2009;Lee et al., 2010;Link et al., 2011;Hon and Shibata,
2013;Panichev et al., 2017;Kroesen et al., 2020). Artificial salt licks come in two forms based on two
types of commercially available salt: bagged and blocked. ...
... Many wildlife species are known to visit salt licks to consume soil (geophagy) or to drink water
(Krishnamani and Mahaney, 2000;Matsubayashi et al., 2006;Bravo et al., 2008); they include ungulates
(Ayotte et al., 2006 (Ayotte et al., , 2008Tobler et al., 2009;Blake et al., 2011;Lameed and Adetola,
2012;Ampeng et al., 2016;Molina et al., 2013), primates (Krishnamani and Mahaney,
2000;Matsubayashi et al., 2006;Link et al., 2011;Ampeng et al., 2016;Gomez-Hoyos et al., 2017), bats
(Nelson et al., 2005;Bravo et al., 2008;Voigt et al., 2008;Ghanem et al., 2013), birds (Brightsmith and
Munoz-Najar, 2004;Lee et al., 2010;Dudley et al., 2012;King et al., 2016;Griffiths et al., 2020), and
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reptiles (Hui, 2004;Magintan Copy link
al., 2015). Wildlife frequently uses salt licks for nutritional
supplementation (Kreulen, 1985;Ramachandran et al., 1995;Chong et al., 2005;Matsubayashi et al.,
2006;Bravo et al., 2010;Lameed and Adetola, 2012) but their usage varies depending on species, sex,
diet (Atwood and Weeks, 2002;Ayotte et al., 2006;Blake et al., 2011) and stage of reproductive activity
(Ping et al., 2011;Ampeng et al., 2016) and is affected by biotic and abiotic factors (Link et al., 2011;King
et al., 2016;Kroesen et al., 2020;Griffiths et al., 2020). ...
... Many wildlife species are known to visit salt licks to consume soil (geophagy) or to drink water
(Krishnamani and Mahaney, 2000;Matsubayashi et al., 2006;Bravo et al., 2008); they include ungulates
(Ayotte et al., 2006(Ayotte et al., , 2008Tobler et al., 2009;Blake et al., 2011;Lameed and Adetola,
2012;Ampeng et al., 2016;Molina et al., 2013), primates (Krishnamani and Mahaney,
2000;Matsubayashi et al., 2006;Link et al., 2011;Ampeng et al., 2016;Gomez-Hoyos et al., 2017), bats
(Nelson et al., 2005;Bravo et al., 2008;Voigt et al., 2008;Ghanem et al., 2013), birds (Brightsmith and
Munoz-Najar, 2004;Lee et al., 2010;Dudley et al., 2012;King et al., 2016;Griffiths et al., 2020), and
reptiles (Hui, 2004;Magintan et al., 2015). Wildlife frequently uses salt licks for nutritional
supplementation (Kreulen, 1985;Ramachandran et al., 1995;Chong et al., 2005;Matsubayashi et al.,
2006;Bravo et al., 2010;Lameed and Adetola, 2012) but their usage varies depending on species, sex,
diet (Atwood and Weeks, 2002; Ayotte et al., 2006; Blake et al., 2011) and stage of reproductive activity
(Ping et al., 2011;Ampeng et al., 2016) and is affected by biotic and abiotic factors (Link et al., 2011;King
et al., 2016;Kroesen et al., 2020;Griffiths et al., 2020). Apart from their important role in supplying
mineral supplements to wildlife, salt licks also provide pharmacological treatment against toxins
(Diamond et al., 1999;Dominy et al., 2004;Hui, 2004), diarrhea (Knezevich, 1998;Mahaney et al.,
1999;Wakibara et al., 2001), and endoparasites (Knezevich, 1998;Krishnamani and Mahaney, 2000;Hui,
2004), as well as playing a role in gut pH regulation (Krishnamani and Mahaney, 2000;Hui, 2004;Ayotte
et al., 2006;Molina et al., 2013). ...
The use of salt licks by birds in Peninsular Malaysia

Jun 2022
Nor Bazilah Razali · Mohammad Saiful Mansor · Nor Adibah Ismail · Shukor Md. Nor
View Show abstract
... The chemical properties of spring forage reduce the digestive efficiency of the rumen and impair
absorption (Kreulen 1985). Forage digestibility is further compromised for species like moose (Alces
alces) and mountain goat (Oreamnos americanus) because they consume forages high in plant defense
compounds (Ayotte et al. 2006) . Lick soils provide the necessary elements to help stabilize the rumen,
as well as supplement demands of lactation and growth (Kreulen 1985, Ayotte et al. 2006. ...
... Forage digestibility is further compromised for species like moose (Alces alces) and mountain goat
(Oreamnos americanus) because they consume forages high in plant defense compounds (Ayotte et al.
2006). Lick soils provide the necessary elements to help stabilize the rumen, as well as supplement
demands of lactation and growth (Kreulen 1985 , Ayotte et al. 2006 . ...
... It appears that the seep (drinking) at rock face licks provide ungulates with mineral elements. Ayotte
et al. (2006) mention that inflow waters at wet licks are particularly high in magnesium. Magnesium may
be sought after by ungulates when high levels of dietary potassium (a result of consuming lush spring
vegetation) inhibit nutrient absorption (Jones and Hanson 1985, Heimer 1988, Ayotte et al. 2006. ...
Observational Description of Alpine Ungulate Use at Mineral Licks in Southwest Alberta, Canada
Conference Paper Full-text available
May 2014
Michael Jokinen · M S Verhage · Robert Anderson · Doug Manzer
View Show abstract

... 6(2): mn0056, (2020) Los salados naturales son elementos del paisaje presentes tanto en bosques
templados como tropicales, los cuales se caracterizan por poseer altas concentraciones de minerales y
arcillas (Ayotte et al. 2006; Powell et al. 2009;Molina et al. 2014). Esta propiedad hace que diversas
especies herbívoras acudan allí para consumir lodo rico en minerales, comportamiento que se conoce
como geofagia. ...
... Esta propiedad hace que diversas especies herbívoras acudan allí para consumir lodo rico en
minerales, comportamiento que se conoce como geofagia. La geofagia puede ser realizada por las
especies con el fin de suplementar su dieta con minerales (Montenegro 2004; Ayotte et al. 2006; Powell
et al. 2009;Bravo et al. 2012), controlar el pH estomacal o reducir la toxicidad de compuestos
secundarios (Klaus & Schmidg 1998;Gilardi et al. 1999). ...
... Otras características como la concentración de minerales, estructura de la vegetación o el tipo de
salado, pueden influir en la cantidad y la frecuencia de las especies registradas, ya que salados con
concentraciones de minerales bajas o salados inundables pueden usarse con menor frecuencia por
algunas especies (Montenegro 2004; Ayotte et al. 2006; Cabrera 2012;Blake et al. 2013;Molina et al.
2014). Adicionalmente, debido a la baja cantidad de registros no fue posible probar si existe efecto de
la distancia al salado sobre la frecuencia o gremio trófico de mamíferos terrestres registrados, como si
ha sido evidenciado en la estructura y la diversidad de plantas en la región (Molina et al. 2018), lo cual
deja una pregunta abierta para próximas investigaciones. ...
Registros de mamíferos terrestres en un salado natural y sus inmediaciones en el Resguardo Indígena

Nonuya Villazul, departamento del Amazonas - Colombia
May 2020
Gabriel P. Andrade-Ponce
View Show abstract
... (significant); ∆ Depth was measured from top of excavation downwards on the vertical face ( Figure
4). (24) . In Aberdares National Park, Kenya particle size composition of lick and non-lick soils indicated
no difference in silt, less sand in lick soils and about two times as much clay in lick soils as in non-lick
soils (7). ...
... In Aberdares National Park, Kenya particle size composition of lick and non-lick soils indicated no
difference in silt, less sand in lick soils and about two times as much clay in lick soils as in non-lick soils
(7). The results of LMNP agree with the earlier findings (7, 24) where salt lick soils had higher clay
content, less sand and similar levels of silt as compared to non-lick soils (top soil). ...
... Therefore sodium concentrations in salt lick soils were about 3-12 times greater than in non-salt lick
soils. The results were consistent with those reported by many other researchers
(7,14,15,16, 24, 27,28). No laboratory analysis of plants eaten by animals was done in LMNP but
analysis elsewhere (15,16,28,29) showed that sodium was consistently low in animal plant diets. ...
Physical and chemical characteristics of animal natural salt lick soils in Lake Mburo National Park,
Uganda
Jan 2012
Ludwig Siefert · Michael Ocaido
View Show abstract
... Animals often travel long distances outside their usual habitat to visit specific mineral licks (Rice
2010, Link et al. 2011. The benefits of geophagy are still uncertain, but may include 1) detoxification of
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secondary plant compounds; citation Copy link
alleviation of gastrointestinal stress; and 3) nutrient supplementation to
meet metabolic demands (Jones and Hanson 1985, Kreulen 1985 , Ayotte et al. 2006 , Slabach et al.
2015. ...
... Studies have documented different species of ungulates visiting the different types of mineral licks.
For example, moose Alces alces and elk Cervus elaphus preferring wet licks and Stone's sheep Ovis
dalli stonei and mountain goats Oreamnos americanus target dry licks (Ayotte et al. 2006) . ...
... Mountain goats are exposed to significant risks when using these sites, indicative of the importance
of these mineral resources. We predict that the soils consumed at the mineral licks will be high in
minerals mountain goats have been shown to seek at other mineral lick study areas such as sodium
(Na), calcium (Ca), phosphorus (P) and magnesium (Mg) (Kreulen 1985 , Ayotte et al. 2006 , Slabach et
al. 2015. ...
Patterns of decadal, seasonal and daily visitation to mineral licks, a critical resource hotspot for
mountain goats Oreamnos americanus in the Rocky Mountains
Nov 2020 · WILDLIFE BIOL
Laura Kroesen · David S Hik
View Show abstract
... Many mineral licks have elevated concentrations of sodium, which is thought to be a key attractant for
animals such as Moose (Fraser and Hristienko 1981;Fraser et al. 1982;Tankersley and Gasaway 1983)
and Elk (Lavelle et al. 2014). Several studies have concluded that in addition to sodium, other elements
in licks such as carbonates, magnesium, and iron may be attractants to animals (Cowan and Brink
1949;Kreulen 1985;Kennedy et al. 1995; Ayotte et al. 2006) . Visitation to mineral licks by ungulates for
sodium and other attractants is thought to be related to milk production (Ayotte et al. 2006), the
demands of antler growth (Atwood and Weeks 2002), and elements required for improved rumen
function (Ayotte et al. 2008). ...
... Several studies have concluded that in addition to sodium, other elements in licks such as
carbonates, magnesium, and iron may be attractants to animals (Cowan and Brink 1949;Kreulen
1985;Kennedy et al. 1995;Ayotte et al. 2006). Visitation to mineral licks by ungulates for sodium and
other attractants is thought to be related to milk production (Ayotte et al. 2006) , the demands of antler
growth (Atwood and Weeks 2002), and elements required for improved rumen function (Ayotte et al.
2008). ...
... Carbonates (Ayotte et al. 2006) and magnesium found in licks may act to combat dietary deficiencies
and as compounds necessary for herbivores to regulate high levels of dietary potassium in spring
compared to winter forages (Dormaar and Walker 1996;Shackleton 1999;Parker and Ayotte
2004;Jokinen et al. 2016). Rea et al. (2013a) found that iron concentrations were high at licks in
northern British Columbia (BC). ...
Year-round patterns of mineral lick use by Moose (Alces americanus), deer, and Elk (Cervus
canadensis) in north-central British Columbia
Article
Mar 2021 · Can Field Nat

Carolyn Brianna Brochez · Roy V Rea · Shannon Michael Crowley · Dexter Hodder
View Show abstract
... The chemical properties of spring forage reduce the digestive efficiency of the rumen and impairs
absorption (Kreulen 1985). Forage digestibility is further compromised for species such as moose (Alces
alces) and mountain goat (Oreamnos americanus) because they consume forage high in plant defence
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compounds (AyotteDownload
et al. 2006)citation
. Lick soils provide theCopy link elements to stabilize the rumen as
necessary
well as supplement chemical needs exacerbated by lactation and growth (Kreulen 1985;Ayotte et al.
2006). ...
... Forage digestibility is further compromised for species such as moose (Alces alces) and mountain
goat (Oreamnos americanus) because they consume forage high in plant defence compounds (Ayotte
et al. 2006). Lick soils provide the necessary elements to stabilize the rumen as well as supplement
chemical needs exacerbated by lactation and growth (Kreulen 1985; Ayotte et al. 2006) . ...
... In Ayotte's (2004) study, the magnesium concentration was one-quarter that of our average
concentration, and the sodium concentration was half of our average concentration. Ayotte et al. (2006)
mentioned inflow waters at wet licks are particularly high in magnesium. Magnesium Multiple species
visited all forest mineral licks. ...
Frequency and Timing of Use of Mineral Licks by Forest Ungulates in Southwest Alberta
Technical Report Full-text available
Jan 2016
Mike Verhage · Michael Jokinen · Robert Anderson · Doug Manzer
View Show abstract
... Regions where wild animals intake mineral-rich soil or water are called mineral licks (Atwood and
Weeks 2002). The requirement for essential mineral nutrients (e.g., sodium, calcium, and magnesium)
has been widely recognized as the main driver of wildlife's access to mineral licks (Ayotte et al. 2006; Li
et al. 2014). ...
... In the Anzihe Nature Reserve, a 6-year (2016-2021) wildlife monitoring program has recorded eights
large or medium-sized mammalian species visiting the mineral licks, including seven species of
ungulates (Sambar Deer, Rusa unicolor, Red List of China's vertebrates category version 3.1: Near
Threatened; Takin, Budorcas taxicolor, Vulnerable; Tufted Deer, Elaphodus cephalophus, Near
Threatened; Reeves' Muntjac, Muntiacus reevesi; Chinese Serow, Capricornis milneedwardsii,
Endangered; Wild Boar, Sus scrofa, Least Concern; Forest Musk Deer, Moschus berezovskii, Critically
Endangered) and one species of primates (Tibetan Macaque, Macaca thibetana, Vulnerable) (Photo
2.1). These animals visited the mineral licks to intake the soil and/or water, which are rich in one or more
elements of sodium, calcium, and magnesium (Ayotte et al. 2006) . Distinct species usually visited
different mineral licks or visited the same mineral lick at different times (Table 1), probably to reduce
interspecific competition. ...
... Actually, this phenomenon has also been found in previous studies on ungulates (Hebert and Cowan
1971;Klaus and Schmidg 1998). For mammals, replenishing sodium and magnesium by salt licking can
offset high potassium ingestion during the spring (Ayotte et al. 2006; Klein, Froelich, and Krief 2008). In
our study area, we did observe a higher frequency of diarrhea in Sambar deer and Takin in the spring
(Photo 2.2b), which may lead to their higher frequency of salt licking. ...
Ecological significance and risks of mineral licks to mammals in a nature reserve on the Eastern
Qinghai-Tibet Plateau
Mar 2022
Xingcheng He · Zhixin Wen · Yujin Wang · Jianghong Ran
View Show abstract
... Additionally, three pooled soil samples were taken from each mineral and control site (randomly
located ~3 meters from each mineral site) for comparison on June 16, 2011 (prior to mineral re-
augmentation), July 13, 2011 (~60 days prior to opening of archery hunting season), September 15,
2011 (at opening of archery season), November 12, 2011 (at opening of firearm season) February 28,
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2012 (one year later).Download
Samplescitation Copy link
were sealed in soil sampling bags and sent to Ward Laboratories, Inc.
for chemical analysis of Na, Ca, P and pH, as elevated soil pH at natural mineral licks provides
supplemental carbonates that could assist in stabilizing rumen pH as spring forage begins to change
(Ayotte et al. 2006) . Mean results of minerals (Na, Ca, and P) and pH were paired by mineral site (N=5)
for each sample period and analyzed for statistical significance with a paired T-test. ...
... Calcium levels did not change in treated soils, which may be a result of Na cations replacing Ca
cations as they are both base cations (Ray Ward, personal communication). Elevated pH has also been
reported in soils from natural licks (Ayotte et al. 2006 ). However, Kennedy et al. (1995) found lower soil
pH at a natural lick which was attributed to finer soil textures. ...
... The need for P has been linked to deficiencies that arise due to late gestation, lactation and antler
growth (Brown 1990, Hewitt, 2011. The supplemental carbonates found in soils with elevated pH may
assist in stabilizing rumen pH as forage changes in the spring (Ayotte et al. 2006) . The timing of these
physiological needs coincides with increase WTD use observed in our study. ...
Longevity of Mineral Supplements within the Soil and Associated Use by White-Tailed Deer.
Transactions of the Nebraska Academy of Sciences 35:61-67.
Dec 2015
Brian C. Peterson · Keith D. Koupal · Andrew K Schissel · Cody M Siegel
View Show abstract
... While the exact reason for the deliberate ingestion of mineralised water or soil has yet to be fully
ascertained, it is likely mineral licks serve multiple functions that may vary for different species, sexes,
or at different temporal scales [2,3]. Natural mineral licks provide essential nutrients that are lacking in
the diet [3][4][5][6][7] or buffering compounds against dietary toxins, diarrhoea, endoparasites or to
adjust gut pH [8][9][10] [11] . ...
... Artificial saltlicks are therefore attracting a similar suite of species at rates similar to those of natural
mineral licks, and may be seen as an alternative to natural licks, at least in terms of behavioural
activities. How artificial saltlicks compare to natural mineral licks in terms of their chemical properties or
their implied health benefits has yet to be ascertained, however, several studies have shown that natural
mineral licks provide a wide variety of macro and micro minerals and gut buffering properties beneficial
to wildlife health [1,4,8, 11] . ...
Mammal Diversity at Artificial Saltlicks in Malaysia: A Targeted Use

Chapter Full-text available
Jan 2021
Boyd K. Simpson · Noraini Nasaruddin · Carl Traeholt · Shukor Md. Nor
View
... A wide variety of species of terrestrial mammals have been discovered across the world, and then the
major visitors of mineral licks are generally the herbivorous ungulates. These non-volant mammal
species fail to obtain a sufficent amount of sodium from the forages, and then high consumption onto
potassium-rich forages further exacerbates sodium stress experienced by the given species in the wild
(Ayotte et al., 2006; and Matsubayashi et al., 2007b). Henceforth, these species show high
dependencies on mineral licks to obtain, not only sodium mineral, but also clay minerals and other
essential and trace elements in sufficient amounts (Sienne et al., 2014;and Molina et al., 2014). ...
... However, the influence of seasonal variation in ambient temperature onto mammalian visitation
pattern to the licks is verified as insignificant (p > 0.05) by past studies (e.g., Hebert and Cowan,
1971;and Tankersley and Gasaway, 1983). The seasonal variation in lick use patterns of terrestrial
mammals are influenced by the change in sodium requirements, instead of being impacted by the
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seasonal change inDownload citation
local ambient temperature (AyotteCopy
et al.,link
2006; Matsubayashi et al., 2007a;and
Ampeng et al., 2016). Therefore, this particular explanatory variable is seldom being examined at the
given temporal scale in this field of study. ...
RELATIONSHIP BETWEEN MINERAL LICKS AND TERRESTRIAL MAMMALS IN THE TROPICAL

RAINFOREST OF MALAYSIA: A PERSPECTIVE REVIEW
Chapter
Jun 2021
Lim Wing Shen · Andy Russel Mojiol
View Show abstract
... pinchaque) were also reported to drank water at salt licks, though no quantitative data was shown
(Lizcano and Cavelier 2000). A number of studies has addressed physiological reasons of salt-lick use
by herbivorous animals, such as mineral supplementation (Moe 1993;Clayton and MacDonald
1999; Ayotte et al. 2006; Matsubayashi et al. 2007, Molina et al. 2014, detoxification of plant secondary
compounds by clay particle ingestion (Klaus et al. 1998;Gilardi et al. 1999;Houston et al. 2001), and
alleviation of gastrointestinal problems (Bechtold 1996;Ayotte et al. 2006, Molina et al. 2014. Malayan
tapirs are herbivores as other Tapiridae species that consume only plants as their food. ...
... pinchaque) were also reported to drank water at salt licks, though no quantitative data was shown
(Lizcano and Cavelier 2000). A number of studies has addressed physiological reasons of salt-lick use
by herbivorous animals, such as mineral supplementation (Moe 1993;Clayton and MacDonald
1999;Ayotte et al. 2006;Matsubayashi et al. 2007, Molina et al. 2014, detoxification of plant secondary
compounds by clay particle ingestion (Klaus et al. 1998;Gilardi et al. 1999;Houston et al. 2001), and
alleviation of gastrointestinal problems (Bechtold 1996; Ayotte et al. 2006 , Molina et al. 2014. Malayan
tapirs are herbivores as other Tapiridae species that consume only plants as their food. ...
Salt-lick use by wild Malayan tapirs (Tapirus indicus): behavior and social interactions at salt licks
Oct 2021 · EUR J WILDLIFE RES
Yuko Tawa · Shahrul Anuar Mohd Sah · Shiro Kohshima
View Show abstract
... Ungulate habitat selection can also be influenced by the availability of minerals in plant material or in
the soil as licks (Grasman & Hellgren 1993;Mungall & Sheffield 1994; Ayotte et al. 2006; Treydte et al.
2009;Watter et al. 2019). Phosphorus, sodium, and calcium are vital nutrients for ungulates as they are
needed for bone mineralisation, which is important for antler growth in deer, as well as sub-cellular
processes and genetic coding (Belovsky 1978;Grasman & Hellgren 1993;Mungall & Sheffield
1994;Dryden 2016;Griffith et al. 2017). ...
... Phosphorus is critical for ungulates (Grasman & Hellgren 1993;Mungall & Sheffield 1994;Dryden
2016) and, in their native range, chital use natural salt licks that are high in phosphate (Schaller,
1967;Moe 1993). In mineralogically-deficient environments, ungulates will actively graze in patches of
vegetation where there is higher phosphorus content or salt licks containing higher phosphorus (Moe
1993;McNaughton 1990; Ayotte et al. 2006; Treydte et al. 2009). The Charters Towers region is situated
on part of the oldest rock formations in Australia, meaning that deer may be particularly sensitive to
phosphorus levels in this deficient environment (Gillman & Bell 1978). ...
Ecology of chital deer in North Queensland

Thesis Full-text available
May 2022
Catherine Kelly
View Show abstract
... While the exact reason for the deliberate ingestion of mineralized water or soil has yet to be fully
ascertained, it is likely mineral licks serve multiple functions that may vary for different species, sexes,
or at different temporal scales (Kreulen, 1985;Atwood and Weeks, 2002). Natural mineral licks provide
essential nutrients that are lacking in the diet (Weeks and Kirkpatrick, 1976;Jones and Hanson,
1985;Heimer, 1988;Ceacero et al., 2009;Flueck et al., 2012) or buffering compounds against dietary
toxins, diarrhea, endoparasites or to adjust gut pH (Mahaney et al., 1995;Knezevich, 1998;Krishnamani
and Mahaney, 2000; Ayotte et al., 2006) . ...
... Artificial saltlicks are therefore attracting a similar suite of species at rates similar to those of natural
mineral licks, and may be seen as an alternative to natural licks, at least in terms of behavioral activities.
How artificial saltlicks compare to natural mineral licks in terms of their chemical properties or their
implied health benefits has yet to be ascertained, however, several studies have shown that natural
mineral licks provide a wide variety of macro and micro minerals and gut buffering properties beneficial
to wildlife health (Jones and Hanson, 1985;Kreulen, 1985;Mahaney et al., 1995; Ayotte et al., 2006) . ...
Mammal Diversity at Artificial Saltlicks in Malaysia: A Targeted Use

Nov 2020
Boyd K. Simpson · Noraini Nasaruddin · Carl Traeholt · Shukor Md. Nor
View Show abstract
... There are only suppositions about the phenomenon of consuming of mineral substances at kudurs by
animals. Among those are the requirement of sodium carbonate and other carbonate salts and of
mineral sorbents which regulate the buffering of the environment in the gastrointestinal tract, which
allows the reduction of the intestinal disorders in the period of spring change of the nutritive ratio
(Kreulen, 1985;Klaus, Schmid, 1998; Ayotte et al., 2006) ; the requirement of supplementing the
organism with the deficiency of Fe (Mahaney et al., 1990) or symbiontic microorganisms (Krishnamani,
Mahaney, 2002); control of parasite and regulation of pH in the intestinal tract (Krishnamani, Mahaney,
2002); the excretion of the chemical toxins from the organism with the help of the mineral sorbents
(Gilardi et al., 1999;Houston et al., 2001;Pebsworth et al., 2013). The revealing of the true meanings of
this phenomenon will permit learning of the ropes in the links of the organism with the environment and
in the physiological mechanisms of animals' adaptation to the geochemical and other environmental
factors; finally, it may be useful for medicine. ...
Results of geological and geochemical investigations of Kaplanov kudurs in Sikhote-Alin, Russian Far
East
Article
Sep 2018
Panichev A.M · Ivan V. Seryodkin · O.Yu. Zaumyslova · Pavel Khoroshikh
View Show abstract
... There is some evidence that elephants also conduct geophagy to support detoxification of
unpalatable secondary compounds in their diet (Mwangi, Milewski & Wahungu, 2004;Chandrajith et al.,
2009). In other ungulate species, clay may decrease the harmful effects of secondary plant compounds
Request full-text Download
and intestinal infections (Klauscitation Copy
& Schmidg, 1998; Ayotte link2006) . Soil is never consumed randomly
et al.,
within an elephants' home range, but instead it is consumed from specific spatially circumscribed sites
(Klaus & Schmidg, 1998). ...
African savanna elephants ( Loxodonta africana ) as an example of a herbivore making movement
choices based on nutritional needs
Feb 2019
Fiona Sach · Ellen S Dierenfeld · Simon C Langley-Evans · Lisa Yon
View Show abstract
... Aquatic plants, however, are not the sole sodium resource exploited by moose. Many workers have
documented moose making use of "Licks," or "Natural Salt Licks," which are natural geologic deposits
containing a variety of mineral concentrations that include manganese and sodium (e.g., Chamberlin et
al., 1977;Risenhoover and Peterson, 1986;Ayotte et al., 2008;Lavelle et al., 2014; also see Ayotte et al.,
2006 for review on classification of licks). Sodium and chlorine, the composition of table salt, are
required elements for the normal growth and development of bones, muscles, and maintaining
homeostasis of the nervous system, while also assisting in the regulation of hematologic pH levels (e.g.,
Denton, 1982). ...
The Nasal Complex of a Semiaquatic Artiodactyl, the Moose ( Alces alces ): Is it a Good Evolutionary
Model for the Ancestors of Cetaceans?
Article
Nov 2018
Samuel Marquez · Anthony Pagano · Carrie Mongle · Jeffrey T Laitman
View Show abstract
... In KLNP, one of such locations supporting kobs is the salt-licks spots. Ayotte et al. (2006) informed
that areas where animals actively ingest soil have commonly been referred to as salt-licks. Measuring
activity levels in animals provides important information about behavioral ecology which gives relevant
information in their management and conservation (Kilgo et al.,1998;Gervasi et al., 2006). ...
KOBS (KOBUS KOB ERXLEBER, 1777) ACTIVITIES AT SALT-LICK SPOTS IN KAINJI LAKE NATIONAL
PARK, NIGERIA LES ACTIVITES DES KOBS (KOBUS KOB ERXLEBER, 1777) AUX AFFLEUREMENTS…
Article
Jan 2015
Ogunjobi Johnson Adedayo
View Show abstract
new paper on kobs
Data Full-text available
Jan 2017
View
... Many wild plant-eating species can be observed spending time at natural or artificial salt licks,
basically to integrate a low-sodium diet (Hebert and Cowan, 1971;Klaus and Schmidg, 1998; Ayotte et
al., 2006) . External dam walls, where salty minerals regularly exude from the concrete bricks, represent
a particular kind of salt lick. ...
Gradient limits and safety factor of Alpine ibex (Capra ibex) locomotion

Jan 2017
Carlo Massimo Biancardi · Alberto E Minetti
View Show abstract
Jan 2015 · TROP ANIM HEALTH PRO
View Show abstract
... Mineral licks generally provide bio-metals such as sodium, calcium, iron, phosphorus, zinc, and trace
elements required for the development of bones, muscles, growth and functioning of other systems in
livestock and wildlife (Hogan, 2010;Black, Mosquera, Guerra, Loiselle, Romo & Swing, 2011). Research
has shown other uses and nutritional benefits of mineral licks to include selenium (Se), cobalt (Co) and
molybdenum (Mo) (Ayotte, Parker, Arocena, & Gillingham, 2006; Mills, & Milewski, 2007). Imbalances in
mineral profiles in soils and forages in semi-arid and arid regions of tropical countries, is the main
challenge in achieving the maximum production and reproduction potentials in livestock (Khan et al.,
2008, Khan, Ahmad, Ashraf, Valeem & McDowell, 2009). ...
Development of Novel Mineral Licks (Toka) and the Livestock Industry Boom in North-eastern Nigeria
Article
Jan 2017
I. H. Kubkomawa
View
... For example, spatial variation in rainfall and forage P has been related to the aggregation and
migration of wildebeest (Connochaetes taurinus) and the selection of foraging areas by elephants
Request full-text
(Loxodonta africana)Download citation
in Africa (McNaughton Copy et
1988, Hopcraft linkal. 2010, Pretorius et al. 2011. Similarly,
sodium (Na) distributions on the landscape have been associated with ungulate movements in North
America and Europe (Jones and Hanson 1985, Staaland and White 2001 , Ayotte et al. 2006 , Slabach
et al. 2015. ...
Mineral constraints on arctic caribou (Rangifer tarandus): a spatial and phenological perspective
Apr 2018
Keith Oster · Perry Barboza · David D. Gustine · Rachel D. Shively
View Show abstract
... В зоне расширения ареала вида солонцевание [Свиридов, 1978] или до 110 мин [Майманакова,
2013]. По данным зарубежных авторов, продолжительность солонцевания достигает 164 мин
[Ayotte et al., 2006] . Сравнение средних статистических показателей разовой продолжительности
литофагии в нативной и инвазионной частях ареала благородного оленя по критерию
достоверности Стьюдента показало, что различие достоверно при доверительной вероятности
0.90-0.99. ...
сравнительная характеристика солонцевания благородного оленя

Apr 2019
И.М. Охлопков · V. V. Stepanova · A. V. Argunov
View Show abstract
... As for foreign literature on the moose geophagia, one can refer to the works by Canadian and
American scientists (Hebert & Cowan, 1971;Carbyn, 1975;Fraser et al., 1980;Fraser & Hristienko,
1981;Tankersley & Gasaway, 1983;Couturier & Barrette, 1988;Filus, 2002; Ayotte et al., 2006; Ayotte et
al., 2008;Rea et al., 2013;Lavelle et al., 2014), as well as Finnish researchers (Heikillä & Härkönen,
1998). ...
Time-study of moose (Alces alces L., 1758) geophagia activity in the Central Yakutia
Dec 2017
V. V. Stepanova · A. V. Argunov · R.A. Kirillin · I. M. Okhlopkov
View Show abstract
... Geophagy is the deliberate ingestion of earth-like materials such as soil, ant or termite mounds, nests
(ovenbirds), chalk, and clay (Pebsworth et al. 2019). It has been observed throughout the animal
kingdom, in various species of birds and mammals including human and nonhuman primates (Ayotte et
al. 2006; Brightsmith and Muñoz-Najar 2004;Klaus et al. 1998;Krishnamani and Mahaney
2000;Matsubayashi et al. 2007;Young et al. 2011). Its occurrence in such a wide variety of species
suggests that it confers an ecological advantage on individuals (Abrahams and Parsons
1996;Krishnamani and Mahaney 2000;Pebsworth et al. 2019). ...
Geophagy in Wild Yellow-Tailed Woolly Monkeys (Lagothrix flavicauda) Is Brief and Rare

Aug 2020 · INT J PRIMATOL
Vinciane Fack · Sam Shanee · Régine Vercauteren Drubbel · Martine Vercauteren
View Show abstract
... Along with sodium and calcium, a wide range of other soil elements can be supplemented by
geophagy, including chromium, cobalt, copper, iodine, iron, magnesium, manganese, molybdenum,
phosphorus, potassium, selenium, sodium, sulphur and zinc Diamond 1999;Downs 2006). This
acquisition of essential minerals is also the most common hypothesis proposed for the occurrence of
geophagy in mammals (Kreulen 1985;Holdø et al. 2002; Ayotte 2006; Young et al. 2011). For example,
during low rainfall periods when sodium levels in plants are low, African elephants (Loxodonta africana)
in Zimbabwe consume sodium rich soils (Holdo et al. 2002). ...
More than eating dirt: a review of avian geophagy

Article
May 2019 · AFR ZOOL
Colleen T Downs · Ian P Bredin · Peter D Wragg
View Show abstract
... According to the data of other sources, the limit of red deer geophagia is up to 40 min (Sviridov, 1978)
or up to 110 min (Maimanakova, 2013). According to the data of foreign authors, the duration of
geophagia reaches 164 min (Ayotte et al., 2006) . ...
Comparative Characteristics of the Geophagic Activity of Red Deer (Cervus elaphus L., 1758, Cervidae,
Artiodactyla) in Yakutia in the Native and Invasive Parts of Its Range
Apr 2019
V. V. Stepanova · A. V. Argunov · I. M. Okhlopkov
View Show abstract
... This finding is consistent with other studies and could be attributed, in a limited way, to increased soil
ingestion and spore inhalation during animal feeding when vegetation is depleted [16][17][18][19][20]
[21]. It is likely that animals also exhibit increased geophagia (consumption of soil) during the dry
seasons, particularly in areas with mineral deficiencies [22] . Additionally, there is a social mixing of
animals, both wildlife and domestic animals that invade national conservation areas during the dry
seasons in search of pasture and water. ...
Temporal and spatial distribution of anthrax outbreaks among Kenyan wildlife, 1999–2017
Jan 2019
John Gachohi · Francis Gakuya · Isaac Lekolool · M. Kariuki Njenga
View Show abstract
Request full-text
... Unlike carnivoresDownload citation from their prey, the
that gain sodium Copy link
intrinsically low sodium in plant tissue means
phytophagous species must seek this vital nutrient elsewhere (Dudley et al., 2012). As such, sodium
deprivation is often considered a key driver of natural lick visitation (Holdø et al., 2002;Powell et al.,
2009;Bravo et al., 2012), but other elements such as calcium and magnesium may also constitute
motivating factors (Ayott e et al., 2006; Matsubayashi et al., 2007b), especially in tropical environments
where soils (and therefore, plants) are depleted of major cations (Emmons & Stark, 1979;Vitousek &
Sanford, 1986). ...
Cambodian Journal of Natural History Volume 2016 Issue 1

Book Full-text available
Jun 2016
Jennifer C. Daltry · Nicholas John Souter · Saveng Ith · Neil Furey
View
... Unlike carnivores that gain sodium from their prey, the intrinsically low sodium in plant tissue means
phytophagous species must seek this vital nutrient elsewhere (Dudley et al., 2012). As such, sodium
deprivation is often considered a key driver of natural lick visitation (Holdø et al., 2002;Powell et al.,
2009;Bravo et al., 2012), but other elements such as calcium and magnesium may also constitute
motivating factors (Ayott e et al., 2006; Matsubayashi et al., 2007b), especially in tropical environments
where soils (and therefore, plants) are depleted of major cations (Emmons & Stark, 1979;Vitousek &
Sanford, 1986). ...
Cambodian Journal of Natural History, 2016(1), 1-76.

Book Full-text available
Jul 2016
Saveng Ith · Neil Furey · Jennifer C. Daltry · Nicholas John Souter
View Show abstract
... These are known variously as licks, salt licks, mineral licks, clay licks or, in Peru, as colpas. Many
reasons have been put forward to explain geophagy Schmid 1998, Mills andMilewski 2007), the most
common of which appear to be dietary mineral supplementation, especially sodium (Ayotte et al. 2006) ,
and the fact that soil serves as a buffer against dietary toxins , Klein et al. 2008). ...
Expedition report: Icons of the Amazon: jaguars, pumas, parrots and peccaries in Peru (November -
December 2009)
Research Full-text available
Jun 2010
Alan Lee · Marcelo Mazzolli · Emma Tatum-Hume · Matthias Hammer
View Show abstract
... In chickens, high soil ingestion has been shown to decrease diet digestibility 12 . Others have
mentioned the role of soil ingestion in pathogen exposure 13 , and its interest for a mineral supply or to
buffer the pH of the digestive system 14, 15 , particularly in wildlife. However, soil ingested by
herbivores would contribute to their exposure to environmental pollutants (especially organic pollutants
as chlordecone that have a high affinity with soil OM 4 ) and to the transfer to animal tissues. ...
Dynamics of soil ingestion by growing bulls during grazing on a high sward height in the French West
Indies
Oct 2020
Claire Collas · Maurice Mahieu · Pierre-Marie Badot · Stefan Jurjanz
View Show abstract
... Most large mammals in Amazonia visit mineral licks. Mineral licks are naturally occurring areas in the
forest where animals exhibit geophagical behavior to obtain essential minerals lacking in their diet [6]
or clays that relieve indigestion caused by plant-based alkaloids [7]. At undisturbed mineral licks, the
ungulates, paca, and ateline primates are the most common visitors [8,9]. ...
The socio-cultural significance of mineral licks to the Maijuna of the Peruvian Amazon: implications for
the sustainable management of hunting
Oct 2020
Michael P Gilmore · Brian Griffiths · Mark Bowler
View Show abstract
... Elephants need food every 12 hours with a dry weight of 1.5% of their body weight during dry season
and 1.9% in the wet season 8 . Elephants directly receive minerals such as phosphorus (P), calcium
(Ca), sodium (Na), and potassium (K) from food especially in mineral rich soils 9 , but some nutrients
such as sodium (Na), sulfur (S) and magnesium (Mg) cannot be obtained from most plants 10 . Both
Asian and African wild elephants (Loxodonta africana) must receive these minerals from others sources
such as soil, drinking water [11][12][13][14] or termite mounds in the case of African elephants 15 . ...
Characteristics of General Soils, Natural Saltlicks and Artificial Saltlicks for Wild Asian Elephants
(Elephas Maximus) Conservation in the Western Forests of Thailand
Preprint Full-text available
Apr 2021
Rattanawat Chaiyarat · Salisa Kanthachompoo · Nikorn Thongtip · Monthira Yuttitham
View Show abstract
... The findings from the study presented here cast the pivotal role of predators stabilising the fauna
ecosystem by influencing the population of prey communities in the salt lick area 4, 42 . Thus, the
conservation of natural salt lick is not only important for prey, but also has direct bearing to the longevity
and survival of the Malayan tigers. ...
Impact of natural salt lick on the home range of Panthera tigris at the Royal Belum Rainforest, Malaysia
May 2021
Bryan Andrew Lazarus · Azlan Che Amat · Muhammad Muzammil Abdul Halim Shah · Ahmad Hafandi
View Show abstract

... The significance of PAM and PAN in understanding population dynamics has been well studied for
many grazer populations (Ahrestani et al., 2011;Illius & O'Connor, 2000;Ludwig et al., 2008;Olff et al.,
2002;Prins & Van Langevelde, 2008;Treydte et al., 2009) but less so for pure browsers with exception of
a study for black rhinoceros (Diceros bicornis) (Okita-Ouma et al., 2020) and for moose (Alces alces)
(e.g. Andersen, 1991; Ayotte et al., 2006; Ellis, 2016;Kielland & Bryant, 1998;Murray et al., 2006) but not
in relation to their population density. The aim of the present study was to test the relevance of PAM and
PAN measures, alongside population density, on populations of a browser species. ...
Effect of high population density of eastern black rhinoceros, a mega‐browser, on the quality of its diet

May 2021
Benson Okita · Richard Pettifor · Marcus Clauss · Herbert H T Prins
View Show abstract
... Saltlicks are naturally occurring deposits that are rich in minerals mainly sodium, potassium, fluorine,
chlorine, calcium, magnesium, sodium and zinc (Tracy and McNaughton, 1995). These minerals play a
significant role in aiding many species with digestibility as well as in detoxification of plant secondary
compounds via clay adsorption (Ayotte et al., 2006) . Previous study on saltlicks in Deramakot, Sabah
reported that 78.4% of the species identified to be living in the forest reserve was recorded at the
saltlicks (Matsubayashi et al., 2006). ...
Topographical differences impacting wildlife dynamics at natural saltlicks in the Royal Belum
rainforest
Article
Dec 2019
Muhammad Muzammil · Azwan Hamdan · Bryan Andrew Lazarus · Ahmad Hafandi
View Show abstract
... Mammals reportedly utilize natural licks as a source of essential minerals, particularly sodium which
is not sufficiently available in feed plants (Ayotte et al. 2006) . Ishige et al. (2017) used the 12S rRNA
mitochondrial DNA fragments as a markers for identification of mammal from water sample at a natural
salt-licks in Deramakot, Sabah, Malaysia. ...
Application of Environmental DNA (eDNA) Metabarcoding Method to Identify Threatened Sulawesi

Mammal Based on 12S rRNA Gene
Article
Dec 2021
Bambang Suryobroto · Ahmad Abdul Jabbar · Puji Rianti
View Show abstract
... Salt-licks occur naturally in certain locations in the forest where mineral salts are found on the ground
surface (Lameed and Adetola, 2012). Ayotte et al. (2006) informed that areas where animals actively
ingest soil have commonly been referred to as salt licks. Wild animal's rationale for licking salt is as a
result of mineral deficiency inside the plant species consumed by them (Ayeni, 1975). ...
COMPOSITION OF LARGE
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7/13/22, 11:01 AM Chemical Composition of Lick Soils: Functions of Soil Ingestion by Four Ungulate Species

October 2006 · Journal of Mammalogy 87(5):878-888

Jeremy B. Ayotte Katherine L. Parker Joselito M. Arocena Michael P. Gillingham

Citations (85) References (100)

Perspectives on the importance of natural salt licks to ungulates have been

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The use of salt licks by birds in Peninsular Malaysia

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Registros de mamíferos terrestres en un salado natural y sus inmediaciones en el Resguardo Indígena

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Mar 2021 · Can Field Nat

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Mammal Diversity at Artificial Saltlicks in Malaysia: A Targeted Use

RELATIONSHIP BETWEEN MINERAL LICKS AND TERRESTRIAL MAMMALS IN THE TROPICAL

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Ecology of chital deer in North Queensland

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Mammal Diversity at Artificial Saltlicks in Malaysia: A Targeted Use

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new paper on kobs

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сравнительная характеристика солонцевания благородного оленя

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More than eating dirt: a review of avian geophagy

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Cambodian Journal of Natural History Volume 2016 Issue 1

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Application of Environmental DNA (eDNA) Metabarcoding Method to Identify Threatened Sulawesi

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