Reese_considerations on the Potential Use of Cliffs

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ORIGINAL PAPER

Considerations on the potential use of cliffs and caves

by the extinct endemic late pleistocene hippopotami

and elephants of Cyprus

Eleftherios Hadjisterkotis &David S. Reese

Received: 22 January 2007 /Revised: 15 June 2007 /Accepted: 20 June 2007 / Published online: 27 July 2007# Springer-Verlag 2007

Abstract Most of the fossil mammal sites on Cyprus, as

well as on other Mediterranean islands, consist of largequantities of bones found in caves. Of 32 sites with

Phanourios minutus and 21 with Elephas cypriotes on

Cyprus, 19 were located in caves, two in rock-shelters, and

11 at open-air sites. Fifteen of them were littoral, four

coastal, and 13 inland. The purpose of this paper is to

examine possible reasons why Phanourios and Elephas

remains accumulated mainly in littoral and coastal caves.

Based on an analysis of the behavior exhibited by living

hippopotami and elephants, we assume that the extinct large

mammals of Cyprus entered these caves in search of fresh

water and to protect themselves from the heat in the cool

and moist cave environment. A further reason may have

been that these mammals entered the caves seeking mineral

licks to rectify possible mineral deficiencies, to bind

secondary plant compounds such as tannins, or to counter-

act acidosis. By entering caves, or even passing along

narrow paths through cliffs, they were at risk of becoming

trapped in natural traps, such as caves with their openings

facing upwards, sinkholes, and mire traps. There is no

evidence that Phanourios and Elephas remains were

accumulated by natural predators because on the island

there were no predators large enough to carry such large

mammals. The only exception are the remains in the

Akrotiri Aetokremnos rock shelter on the Akrotiri peninsu-

la, where there is evidence that the 218,459Phanouriosand330 Elephas remains were accumulated by the first human

settlers of Cyprus, about 10,000 years BP.

Keywords Fossils . Cyprus . Pygmy Hippopotami .

Pygmy elephants .Phanourios minutus .Elephas cypriotes

Introduction

The first evidence for the presence of humans on the island

of Cyprus comes from a rock shelter or small cave at

Akrotiri-Aetokremnos, where man-made artifacts were

found together with a large number of skeletal remains of

the extinct endemic pygmy hippopotamus (Phanourios

minutus) and pygmy elephant (Elephas cypriotes), as well

as many bird bones, reptile bones, edible marine mollusc

shells, lithics, shell and stone ornaments, etc. The site,

located on a steep sea cliff, dates to about 10,000 years ago

(Hadjisterkotis et al.2000; Simmons1991a,b,1996,1999,

2002; Simmons and Wigand1994).

The main argument that some researchers have given for

rejecting the association ofPhanouriosbones and the other

remains with humans at Aetokremnos is that most of the

fossil mammal sites on Cyprus, as well as on other

Mediterranean islands, consist of large quantities of bones

found in caves. Of 32 sites on Cyprus with Phanourios, of

which 21 also had Elephas, 19 were located in caves, two

probably in rock shelters, and 11 at open-air sites. Fifteen of

them were littoral, four coastal, and 13 inland (Reese 1995,

unpublished analyses). This was also observed at sites on

Crete (Lax 1996), Malta (Hunt and Schembri 1999), and

Sicily (Bonfiglio and Burgio 1992; Bonfiglio and Insacco

1992; Bonfiglio and Piperno 1996). Unlike at Aetokrem-

Eur J Wildl Res (2008) 54:122133

DOI 10.1007/s10344-007-0121-3

DO00121; No of Pages

Communicated by W. Lutz

E. Hadjisterkotis (*)

Ministry of the Interior,

1453 Nicosia, Cyprus

e-mail: [email protected]

D. S. Reese

Peabody Museum, Yale University,

P.O. Box 208118, New Haven, CT 06520-8118, USA


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nos, there is no evidence on these other islands that the

extinct endemic animals were associated with humans.

Reese (2001) provided a detailed study of the Aetok-

remnos fauna from the features and lowest stratum which,

in our opinion, leaves no doubt that the bones were

contemporary with the man-made artifacts. Hadjisterkotis

et al. (2000), in a paleoecological and paleoethological

analysis of the site and its remains, noted that perhaps anumber of species found in the rock-shelter entered for

nesting or hibernation or were brought there by the extinct

endemic genet (Genetta plesictoides). However, they noted

that the only way that the 218,459Phanouriosremains, 330

Elephasremains, 3,200 bird bones, and 73,000 marine shell

fragments could have gotten into the cave was if they were

brought in by humans. Most of the bones are broken and

many are burned.

Although numerous studies have now been published

that support the association between humans and Phanour-

ios at Aetokremnos (Simmons1991a,b,1996,1999,2001,

2002; Simmons and Reese 1993; Simmons and Wigand1994; Reese 1989, 1996, 2001; R eese et al. 1999;

Hadjisterkotis et al. 2000), a few researchers are still not

convinced of this association (noted in Reese 2001;

Ammerman and Noller 2005). Although Aetokremnos is

the only fossil site on the island with evidence on what

caused the accumulation of the above skeletal remains,

what brought the extinct endemic pygmy hippopotamus

and a pygmy elephant in the other 19 caves remains

unknown.

The purpose of the present paper is to examine the

possible reasons why Phanourios and Elephas remains

accumulated mainly in littoral and coastal caves. The main

hypothesis, which we examine here, is that these animals

entered caves, or used paths going through cliffs, for

several reasons and by doing so encountered a number of

natural traps. Consequently, this study also discusses

possible reasons why these animals would be attracted to

cliff caves and rocky beaches.

Materials and methods

Our analysis relies on the known behaviors of living

hippopotami and elephants, as well as on information from

archaeological and paleontological studies on related

pro boscidean s. The geological history of Cypr us is

reviewed to understand the formation of the littoral and

coastal caves of the island and some of the ecological

peculiarities relevant to its extinct large mammals. To

examine the hypothesis that animals had entered the caves

in search of water and for the cool environment, we review

the physiological needs of hippos and elephants as far as

water is concerned and the set of behaviors observed for

modern pachyderms to seek or struggle to get water, or to

protect their bodies from heat under drought conditions.

To investigate whether the extinct animals were attracted

to caves or to the coast in search of mineral licks or other

sources of minerals, the known mineral needs of living

elephants and the actions taken by them to satisfy such

needs are reviewed. To see if there were any mineral or

nutrient deficiencies in the vegetation of Cyprus thatwould force the extinct elephants and hippopotamus to

seek mineral licks or saline water, we investigate case

studies on the largest living wild herbivore on the island,

the Cyprus mouflon (Ovis gmelini ophion), and report

findings on domestic animals as well.

In the summer and fall of 1988, and in the winter, spring, and

summer of 1989, 85 samples of mouflon forage species were

collected and analyzed for nutrient content (Hadjisterkotis

1993). Selection for analyses was based on plants found in

the rumen and on visual observations of mouflon while

feeding in the wild (Hadjisterkotis 1993, 1996b). Plant

material collected mimicked plant parts selected by mou-flon. At least 100 g (fresh weight) of plant material was

harvested from a minimum of 25 individuals of each

species, with the exception of Mediterranean medlar

(Crataegus azarolus) fruit, which had a very bad crop year.

In this case, fruit was collected from only three trees.

Samples were kept for about 2 days at ambient temperature

until received at the Chemical Laboratory at the Ministry of

Agriculture in Nicosia. Dry matter was determined by

heating samples of a constant weight in a convection oven

at 60C for 24 h. Dried material was then ground to a

powder with a laboratory mill for further analytical

procedures. Zinc, copper, iron, molybdenum, manganese,

(ppm, mg/kg of dietary dry matter), and calcium (Ca

percentage of dry matter) were determined with a Perkin

Elmer atomic absorption spectrophotometer, after a 1-g

sample was wet digested with a mixture of 10 ml of

Nitric-perchloric acid. In the case of Ca, lanthanum oxide

(La2O310%) was added to the sample. Sodium, potassium,

and phosphorus (percentage of diet dry matter) were

determined by using a Corning-EEI flame photometer.

Nitrogen content was determined with the Kjeldahl method.

Crude protein content was estimated by multiplying the

nitrogen content by 6.25 (percentage of diet dry matter,

Horwitz 1960). Sulfur (percentage of dry matter) was

determined by using the dry ashingbarium sulfate method.

Due to technical difficulties, the microelements iodine, cobalt,

selenium, and fluorine were not examined. To determine if

there were any macromineral or micromineral deficiencies or

intolerable levels in the samples analyzed, the results were

compared with the values presented by the National Research

Council (1985, Tables 6 and 7).

To clarify whether the accumulated bone remains were

brought into the caves by predators, we review other known

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cases where proboscidean remains were carried in by humans

or other predators, and we list all the known predators found

on Cyprus from prehistoric times until today.

Results and discussion

The origin of Cyprus and its littoral caves, and the arrivalof large mammals

Cyprus begun forming between 92 and 85 million years

ago (time scale of Harland et al. 1982) with the formation

of the Troodos Massif, a fragment of uplifted oceanic crust

(Gass 1980; Mukasa and Ludden 1987; McCallum and

Robertson 1990; Robertson 1990). By the Late Miocene,

the Troodos Massif was a low-lying island, and the Kyrenia

Range, which had been deeply submerged, began to rise. A

severe compression and drastic uplift occurred in Cyprus in

the Pleistocene. The Troodos Massif, Kyrenia lineament,

and Mesaoria basin were uplifted together, forming a singlestructural unit (McCallum and Robertson1990). That uplift

raised Mt. Olympus to its present elevation of 1,951 m. In

addition to the regional tectonic uplift, sea level fluctuations

occurred throughout the Quaternary. These have contribut-

ed to the generation of the river terraces and raised beaches

of Cyprus and the formation of littoral caves from the

action of the waves on the rocks in combination with

underground flowing fresh water.

Since Cyprus is an oceanic island that was never

connected to the mainland, the only way large mammals

could arrive was by swimming (Hadjisterkotis and Masala

1995; Hadjisterkotis et al. 2000). When the first hippopot-

ami and elephants arrived from the mainland they most

likely had to face a completely different environment from

that which they were used to. The only rivers present on the

island today are torrential ones running during the winter

after strong rains. During the Late Pleistocene, it is

probable that hippos and elephants of the nearby mainland

inhabited the estuaries of the large rivers such as the Nile,

the Pyramos (modern Ceyhan river), or the Orontes, which

flow into the Mediterranean both to the north and to the

east of Cyprus. Hippopotamus amphibius first appear

during the Middle and Upper Pleistocene in coastal sites

in the southern Levant, such as Evron, Holon, Tabun,

Sefunim, and Kebara caves (Tchernov 1981, 1984; Faure

1986), and inhabited the Nile from at least the Pleistocene

(Reese 1985; Horwitz and Tchernov 1990) until the last

animal was shot in the Nile delta in the last century (Nowak

and Paradiso 1983). It also existed in Lebanese coastal

sites, such as Sidon, Naame, and Ras el-Kelb, and along the

river systems such as at Latamne (Faure 1986) since the

same period. The Orontes and Sinn rivers supported

hippopotamus populations up to the early Iron Age.

However, once hippos and elephants arrived on Cyprus,

in the hot summers, they were faced with a new, much drier

situation. This is evident from the adaptations that

Phanouriosunderwent. These hippos lost the characteristic

wide foot pad of the living large African hippopotamus

(H. amphibius) and developed a more-or-less pig-like foot

adapted to walking. The lowering of their eyes and nostrils

also indicates that they were adapted to a more terrestriallife than the common hippo (Houtekamer and Sondaar

1979; Spaan1996). These adaptations are an indication that

Phanourios probably spent less time in the water than its

ancestorH. amphibius.

The use of caves for water and for the cool environment

According to Haynes (1995), an adult elephant requires

great quantities of moisture/water each day to digest food

and regulate its body temperature. During dry seasons,

elephants will travel to water every day when possible,

although data from radiotelemetric studies show femalegroups regularly remaining without water for 48 h, even

when only 5 km from a water source. Male bands were

noted going 3 to 4 days without water, although never more

than 20 km from pans during that time. When there is no

water near feeding areas, elephants may travel long

distances to water every 2 to 3 days. If visits to water are

daily, elephants spend a relatively short time drinking,

spraying, and wallowing (less than an hour in daytime).

When visits are more widely spaced, the time spent at water

may be much longer. However, a recent report by Douglas-

Hamilton (2003) describes the elephants of the Gourma

region of the Malian Sahel. These animals eke out a living

at the northern extreme of the African elephants distribu-

tion, showing remarkable adaptations to their desert

environment and making long journeys. Based on data

from three radio-collared elephants, it was found that a

females home range exceeded 9,000 square miles. All

three followed the rains, eating and drinking their way from

water hole to water hole.

According to Haynes (1995), during the dry season,

elephants visit seeps to dig shallow wells 1 to 3 m deep to

the water level and wait for water to ooze into the opening.

This behavior was observed by early explorers such as

Chapman in 1862 (Tabler 1971), Davison (1930, 1930

1935), and others, and was seen mainly during the dry

season when pans dried up or were muddied by heavy use

(Davison 1930, 19301935). In very dry years, the water

oozes so slowly into the wells that elephants sometimes

stand at them waiting for hours to satisfy their thirst.

In the 1980s, according to aerial surveys, 1,0002,000

elephants were normally present near the seeps in Hwange

National Park in Zimbabwe during the dry seasons (Haynes

1995), and competition for access to the seeping water in

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wells became intense as the dry season progressed. In spite

of a succession of drought years, hundreds of elephants

continued to spend the dry season in the area of the seeps

each year.

Wells were often redug, cleaned out, and enlarged by

elephants, especially after traffic to the seeps and their use

as water sources increased dramatically in the middle of the

dry season (Haynes 1995). As the ground around wells istrampled bare, the sediments dry out, and wells often

collapse. If a carcass or skeleton lay within the depression

created by digging or trampling around a well opening, or

within the well itself, elephants would kick it or move it

with their trunks to get it out of the way. Elephants dug

through buried carcasses as well, after young animals had

died within wells and caused the holes to slump or collapse.

Weakened animals occasionally become mired in the

mud and muck of water-hole bottoms or banks (Murie

1934; Matthiesson and Porter1974; Shipman1975; Sinclair

1977) and such may have been the fate of Late Pleistocene

mammals (Haynes1995). Many of their bones would havebecome incorporated into the mud and sediments and might

become preserved, in contrast to the situation of bones left

lying on the ground surface during times of drought.

Hippopotami without water and mud baths, in a hot

environment, are known to become dehydrated and to have

difficulty surviving. According to Estes (1992), because

hippopotami have a unique skin (thin epidermis, no sweat

glands) which loses water at several times the rate of other

mammals, a hippo out of water in hot weather risks rapid

dehydration and overheating. In hippos, there are no sweat

glands either within the dermis or at the base of the few

hairs of the skin, where glands are almost always found in

most other mammals. There are, however, large subdermal

glands, made up of more than one type of cell, scattered

regularly over the body (Wright 1987). They are sparsely

distributed at a density in the common hippo of one per

square centimeter, but they make up for that by being

unusually large for a skin gland, with a weight of about 1 g

and a duct clearly visible to the naked eye. Hence, the

volume of secretion is quite high. These glands produce a

viscous fluid varying from colorless to pink and dark

brown. The color of the secretion gives a pinkish tinge to

the whole body.

Evaporation from the hippos skin was found to be very

high compared with that of other mammals and varied from

68 g m2 h1 when the skin was dry to 2,280 g m2 h1

when the skin was wet with secretion (Wright1987). The

latter value approaches the figures obtained by placing the

hygrometric capsule over filter paper soaked in a KCL

solution, suggesting that there is no control over water loss

from the skin. If a hippo is actually unable to control the

rate of water loss from its body, its ability to deal with heat

stress will be limited. It may be for this reason that the

hippo has evolved a semiaquatic life-style. The same is also

true of the air temperature at night, and this may be a

contributing reason for the hippos spending the night out of

water for feeding, often traveling quite long distances to

reach grazing areas (Eltringham 1999).

Olivier (1975) measured water loss from the skin of

living West African pygmy hippos (Choeropsis liberiensis).

The results revealed a much lower water loss comparedwith the larger common hippo. However, the temperature

was some 10C higher during the measurements of the

common hippo, and, as the water vapor pressure doubles

with each ten-degree rise, the water loss would also double.

If the values forChoeropsis are doubled, they come within

the range of the values for Hippopotamus, which seems

more reasonable and suggests that the physiology of the

skin as well as its anatomy is almost identical in the two

species.

The problems faced by hippos and elephants when they

lack water were demonstrated by Joubert and Joubert

(1988) in the film The Stolen River, and were also describedby Walker (1991) in the bookSavuti: the Vanishing River.

The Linyanti River brings water in to the Savuti Channel in

Botswana. Savuti was once a vast lake, then dried up for

about 80 years. The river began to flow again in 1957, but

in 1982, the channel stopped flowing for 7 years. Accord-

ing to Walker (1991),

maddened with thirst, the poor beasts continued to

flock to the well-remembered place, to find, not relief,

but foul mud; the stench from the rotting carcasses of

earlier arrivals; and death in its ugliest form...The

weaker individuals were immediately trapped, andtrampled down to rise no more, while the stronger

ones floundered desperately round and round, butting

one another in their agonies.

In the last water pool left, 100 hippos were trapped in the

mud (Joubert and Joubert1988). Most of them were killed

by the blazing sun. The surviving hippos left the mud to

seek the shade of trees. Some hippos lasted for several

seasons, surviving on the moisture of the early morning and

sheltering through the day, but by being away from the

protection of the aquatic environment, they died from

starvation or by being killed by lions and hyenas. The

elephants of the Savuti were able to dig into the ground at

the bottom of the empty canal, finding some water. The

limited supply of water forced the elephants to stop their

young from drinking, condemning them to death in order to

survive themselves.

Moss (1988) observed separation of cows with youngest

calves during a drought in Amboseli National Park in

Kenya. Old cows that are nursing may die during drought,

leaving their calves to die, mainly from predation. Haynes

(1995) reported seeing lactating cows without calves in tow,

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resulting from the calves death or abandonment by the

mother.

Similarly,Phanouriosfaced a dry and hot environment in

Cyprus and had to find ways to adapt. They had to search for

shady places that would offer the appropriate moisture for

their skin and drinking water. The best place for such

protection would be the littoral caves and the Mediterranean

Sea. Hippos are known to use the sea quite well; they havebeen known to swim from the East African mainland to the

island of Zanzibar, more than 35 km away. In addition, in the

Late Pleistocene/Early Holocene they reached the island of

Madagascar by overcoming the 450-km-wide Mozambique

Channel (Held 1989). Before Europeans introduced fire-

arms to southern Africa in the 1830s, hippos were reported

to swim in the sea as much as in rivers (Smith 1849).

It is likely that Phanourios made use of the sea for

cooling. However, even if they had the opportunity to use

the sea, they still needed water for drinking. So they had to

leave the sea to search for fresh drinking water and food.

This brought them in touch with the coastal and littoralcliffs and caves, and they probably soon discovered and

made use of the cool and moist environment of the caves.

Inside the caves, evaporation is reduced and any spring

water coming through the walls of the caves could create

water pools with drinking water. Many of these caves were

created from underground water dissolving the soft lime-

stone. However, entering such locations, they had to face

the possible natural traps left in the rocks by the action of

the sea waves or the underground water. The heat-stressed

hippos could enter the pools not knowing that getting out

was an impossible task. Elephants, when faced with a

steep-sided pool (contrary to the hippos) are reluctant to

enter. They lower their trunk and lift one of their hind legs

up, standing on only three legs in an attempt to lower their

body to reach the water. This might explain why so few

elephants were trapped in these caves.

Mineral utilization

A possible additional reason for entering caves may have been

the search for minerals. Animals occasionally eat small rocks

or soil impregnated with natural salts to rectify possible

mineral deficiencies, something known as geophagy and

pica behavior (Schaller 1977). Areas where ungulates

actively ingest soil have commonly been referred to as salt

licks (Ayotte et al. 2006). This name implies that licks serve

a universal role of supplementing diets with sodium.

Various reports exist showing that wild ungulates search

for salt (Stockstad 1950; Dalke et al. 1965; Geist 1971;

Schaller 1977; Weeks 1978; Watts 1979; Robbins 1993;

Hadjisterkotis 1997; Fraser and Reardon 1980; Holl et al.

1980; Ayotte et al.2006). Many studies have reported high

concentrations of sodium in lick material (Hebert and

Cowan 1971; Weeks and Kirkpatrick 1976; Reisenhoover

and Peterson 1986; Tracy and McNaughton 1995). How-

ever, many of the field observations of food preferences and

mineral lick use are difficult to interpret, as it is usually

unclear which element is being sought. Hadjisterkotis

(1993, 1997), from December 1985 until December 1988,

provided Cyprus mouflon with mineral blocks and table salt

in six stations spread inside Pafos Forest in Cyprus. Wildmouflon never licked any of the mineral blocks or the salt,

although there was evidence that they often bedded next to

them. It is likely that sometimes elements other than

sodium are sought in mineral licks (Chamberlin et al.

1977; Coates et al. 1990; Heard and Williams 1990;

Dormaar and Walker1996; Hadjisterkotis1997).

Knight et al. (1988) studied ungulates that visited lick

sites in the Kalahari and found (1) that browsers more often

than grazers visited licks, (2) that animals living in highly

seasonal (wet/dry) habitats countered springtime acidosis

by ingesting mineral soils during the time of abrupt dietary

shifts from dry forage to green sprouts, (3) that animalscorrected for the toxic compounds in plants, such as

tannins, by eating clay particles, and (4) that a high sodium

intake among herbivores in dry habitats was a regular

strategy that enabled them to survive dehydration by

protecting cell membranes from damage during osmosis.

In other parts of the world, deficiencies in macro- and trace

elements might be a result of digestive disorders associated

with spring forage change (Watts 1979; Ayotte et al. 2006).

The presence of many species of ungulates at mineral licks

in the spring [moose (Alces alces), Tankersley and Gasaway

1983; Ayotte et al.2006; elk (Cervus elaphus), Ayotte2004,

Ayotte et al. 2006; Williams 1962; Dalls sheep (Ovis dalli

dalli), Heimer1973, 1988; Tankersley 1984; bighorn sheep

(Ovis canadensis mexicana), Watts 1979; mountain goats

(Oreamnos americanus), Singer 1978; Ayotte et al. 2006;

and Stones sheep (Ovis dalli stonei), Ayotte et al. 2006]

coincides with the increased physiological demand of

lactation, growth, or weight regain, which can be aggravated

by electrolyte loss related to the stress of abrupt changes in

forage chemistry (Ayotte et al. 2006). One source of

physiological stress is the change from highly fibrous winter

diets to flourishing spring plant growth resulting from a

decrease in fiber and an increase in fermentable carbohy-

drates and proteins that alter rumen pH and weaken proper

microbial function. Ruminants exposed to sudden decrease

in dietary fiber produce less saliva, which is high in

bicarbonates (Kreulen1985). With less saliva, the buffering

capacity of the rumen is reduced (Church1975), which can

lead to a decrease in pH below the best possible conditions

for rumen microbes, and weight gain (Klaus and Schmid

1998; Kreulen1985).

Physiological stress can also occur when high levels

reduce the absorption and retention of other elements

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(Klaus and Schmid 1998; Kreulen 1985). Early-season

vegetation contains concentrations of potassium that elevate

the osmotic pressure of the digestive tract, interfere with

fecal water absorption, and lead to potentially unsafe

electrolyte loss (Weeks and Kirkpatrick 1976; Kreulen

1985; Klaus and Schmid1998).

According to Ayotte et al. (2006), supplemental sources

of carbonates during the transition to spring forage canimprove the buffering capacity of the rumen when saliva

secretion is compromised (Kreulen 1985). During spring

forage change, clay minerals can be associated with the

adsorption of potassium and short chain fatty acids to

cation exchange sites, adjusting and maintaining proper

osmotic balance and pH of the rumen, enhancing its

buffering capacity (Klaus and Schmid 1998; Kreulen

1985; Cairns-Smith and Hartman 1986). Some kinds of

absorptive clays are also able to bind secondary plant

compounds such as tannins (Johns and Duquette 1991) and

improve digestibility and feed conversion in domestic

animals (Smith1992).The search for salt and mineral licks to rectify possible

deficiencies has also been observed in living elephants.

However, almost no information is available on the

minimum levels of different nutritional substances required

for maintenance and production in wild elephants (Haynes

1995). There is good evidence that elephants benefit from

seeking out mineral soils to ingest, even at the expense of

optimized food intake. Moss (1988) has suggested that

African elephants living outside Amboseli National Park

traveled into the park to eat the salt found there.

Buss (1990) observed five elephants in Tanzania

(Ngorongoro Crater region) at a long, high trench that had

evidently been grubbed and gouged out over a long period

of time. Soil was gouged out with the tusks and transferred

by the trunk to the mouth. Some of the previously used

excavations that were too deep and inconvenient for easy

access had been deserted by the elephants. Chemical

analysis of the soil indicated relatively high manganese

content. Buss therefore suggested that elephants migrated to

the Ngorongoro Crater primarily for the manganese- and

cobalt-laden soil. Manganese is an essential element for

mammals, with Mn deficiency causing ataxia, abnormal

brain function, abnormal development of the skeleton, still

births, poor viability, etc. (Buss1990).

A recent study of three different South African elephant

populations suggests that reproduction rates may be

significantly depressed if the major food plants provide an

inadequate intake of phosphorus or if calcium intake is

excessive as compared with phosphorus (Koen et al. 1988).

A calciumphosphorus imbalance in association with

phosphorus deficiency may be a population-limiting factor

that can lead to longer interbirth intervals, delayed maturity,

and increasing age at first conception (Haynes 1995).

Elephants also need sodium in relatively large amounts

(Weir 1972), as well as iodine, cobalt, and selenium

(Milewski 2000; Milewski and Diamond 2000). According

to Hold et al. (2002), most studies on lick chemistry show

that lick soils consumed by African elephants have higher

Na+ concentrations than the surroundings soils (Ruggiero

and Fay1994; Stark1986; Weir1969), and it was assumed

that licks are generally used to supplement an inadequatedietary Na+ intake (Reisenhoover and Peterson 1986;

Weeks and Kirkpatric1976; Weir1972).

A need for extra sodium in the diet is met by visiting

mineral licks, where elephants excavate pits and even caves

with their trunks, seeking alternative foods, or by drinking

saline water (Weir 1972; Sukumar 2003, Fig. 5.7). The

Kalahari-sand region of Hwange National Park, Zimbabwe,

is particularly poor in soil nutrients including Na+

(Weir

1969; Hold et al. 2002). Plants in this habitat are,

therefore, presumably low in Na+ (McDowell 1985).

Elephants create and maintain depressions in patches of

Na+-rich soil in this area (Weir 1969), and positivecorrelation has been found between the number of

elephants visiting pans (clay-rich depressions that hold

rainwater during the dry season) and the Na+ content of

water in pans located in the Kalahari-sand region of

Hwange (Weir 1972). Hold et al. (2002) studied the use

of mineral licks by African elephants during the dry season

in a Kalahari-sand habitat in Hwange to investigate the role

of geophagy as a mechanism for supplementing low Na+

levels in browse and natural water supplies. According to

their findings, elephants in Hwange may have difficulty in

meeting their maintenance requirements of Na+ from forage

alone, but are probably able to obtain sufficient Na+ from

licks and some Na+-rich water sources to satisfy their

requirements of this mineral. In Kalahari-sand areas, licks

may be the primary source of Na+ supplementation in areas

without Na+-rich borehole water. At supplemented pans

with elevated water-Na+ concentrations, geophagy may not

be necessary as a means of Na+ intake. Hold et al. (2002)

concluded that Na+ appears to play an important role in

attracting elephants to licks and, thus, in affecting move-

ment and habitat-use patterns by this species.

There is evidence that not only modern elephants travel

long distances in search of minerals, but that mammoths

and mastodons did as well (Holman et al. 1988). These

authors suggested that proboscideans migrated from rela-

tively saltless areas in Ohio, Indiana, and Illinois to eat

sodium-rich earth in Michigan salt licks.

Holman et al. suggested that proboscideans are found in

such abundance at Michigan salt licks because they traveled

north into the periglacial forests seeking minerals that were

not available elsewhere. Other geologically similar areas

rich in salts also contain large accumulations of probosci-

dean bones, such as Big Bone Lick in Kentucky, USA.

Eur J Wildl Res (2008) 54:122133 127


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Based on the above evidence, the question arises

whether there are any mineral deficiencies on the island

of Cyprus which would have forced Phanourios and

E. cypriotes to search for salt licks, saline water and

minerals in cliffs and caves.

Since there are no wild elephants or hippos on Cyprus

today, the only way to investigate any mineral shortages in

soil or vegetation is by examining possible deficiencies ofthe living large mammals on the island. The only large wild

herbivore living on the island today is the Cyprus mouflon,

which inhabits Pafos forest (Hadjisterkotis1993,1996a,b,

1997, 1998, 1999, 2001, Hadjisterkotis and Bider 1997;

Hadjisterkotis and Van Haaften 1997; Hadjisterkotis and

Vakanas1997; Toumazos and Hadjisterkotis 1997; Serreri

et al.2000).

Fifteen elements have been demonstrated to be essential

for domestic sheep (National Research Council 1985).

Seven are major mineral constituents: sodium, chlorine,

calcium, phosphorus, magnesium, potassium, and sulfur.

The other eight are trace elements: iodine, iron, molybde-num, copper, cobalt, manganese, zinc, and selenium.

Hadjisterkotis (1993) examined the major forage species

selected by O. g. ophion for most of the above elements.

The study examined two different regions of the forest.

The first region was the center of the forest, which is an

undisturbed Mediterranean pine (Pinus brutia) and golden

oak (Quercus alnifolia) forest. The second region was an

area of mixed forest and agriculture, with many open

abandoned fields. The study concluded that O. g. ophion

during the late summer/early fall could have phosphorus,

protein, and sodium deficiencies (Hadjisterkotis 1993).

Although O. g. ophion are not expected to be subject to

dietary calcium deficiency, in some other parts of the

island, there is evidence that there might be such a

deficiency. In Mesaoria, cattle used to lick the plaster from

the walls, perhaps to rectify calcium deficiencies. The

sandy soils in certain villages of Mesaoria are deficient in

calcium.

During rumen analysis of O. g. ophion found dead in

their natural habitat, it was observed that a number of

individuals had eaten small stones of gabbro (Hadjisterkotis

1997). This type of stone has a higher phosphorus and

potassium content than diabase, and it breaks up much

more easily than the usual diabase. Therefore, based on the

above evidence, it was concluded that O. g. ophion ate

rocks and soil possibly to correct mineral deficiencies.

Mouflon might be deficient in sodium, particularly at the

edge of the forest during late summer and early fall

(Hadjisterkotis 1993, 1997). In Cyprus it was observed by

EH that domestic sheep foraging near the coast at Neta

village on the Karpas peninsula (under Turkish military

occupation since 1974) often visited rocky beaches to eat

salt from seawater pools. The water in these pools, naturally

collected during the winter storms, had evaporated, leaving

a layer of salt. According to Varnava, former Officer of the

Game and Fauna Service of the Ministry of the Interior,

sheep, goats, and donkeys licked salt from the rocky

beaches of Kormakitis in southwestern Kyrenia district

(personal communication to EH, 2002).

There is thus sufficient evidence that at least during

some seasons of the year the vegetation on Cyprus isdeficient in certain minerals and nutrients, particularly

phosphorus and sodium. Therefore, it is possible that the

extinct Cypriot mammals could have searched for these

minerals, as do wild and domestic herbivores today.

A further question to be discussed is whether hippos and

elephants enter caves to search for minerals. Redmond

(1982) reported that elephants go deep within a cave on

Mount Elgon in western Kenya to eat salt-laden rock. It is

possible that, in areas where trace metal concentration in

the topsoil are low due to leaching, and where these metals

are only available deep in the ground, they might be more

accessible through caves. However, by entering caves, orgoing through steep rocks on rocky beaches, these animals

are at risk of getting caught in natural traps.

Occasionally, mammoths entered caves to escape bad

weather, give birth, or search for water or salt. A striking

indication of this is the presence of quantities of mammoth

dung in dry caves in the American southwest (Mead et al.

1986).

Natural traps

Natural traps can be caves with their openings facing

upwards, sinkholes, tar pits, bog traps, mire traps, etc. Such

traps of modern elephants, but also of mammoth and

mastodon remains, are well known. According to Haynes

(1995), the main natural causes of mortality among modern

African elephants are accidental capture in natural traps

such as mud holes, drowning, or falling from steep slopes.

Most Late Pleistocene mastodon finds in North America

involve single skeletons or small numbers of individuals. A

few mass occurrences are known from artesian springs

(Saunders1977,1984). In South America, a mastodont has

been found in at least one mass site (Simpson and De Paula

Couto1957).

At Condover in Shropshire, England, four or five wooly

mammoths (Elephas primigenius), dating between 12,700

and 12,300 years BP, were found in a kettle hole in 1986

(Lister 1993; Lister and Bahn 1994). This is a feature

formed as a result of the melting of a large buried block of

ice left by a retreating glacier and the collapse of the

overlying sediments. The Condover kettle hole was 10 m

deep at its center, but at the end of the Pleistocene its sides

were not very steep. It is therefore most likely that the

mammoths entered it in search of food or drink and became

128 Eur J Wildl Res (2008) 54:122133


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mired in mud at the bottom. The Condover site probably

represents occasional deaths over several centuries.

Another spectacular accumulation discovered in 1914 in

downtown Los Angeles, CA, is the famous tarpits of

Rancho La Brea. From these tarpits, over 100 tons of fossils

were excavated, 1.5 million vertebrates, 2.5 million

invertebrates, often in densely concentrated masses. The

animals included 17 proboscideans [both mastodons andColumbian mammoths (Mammuthus columbi)]. Most of the

remains date between 40,000 and 10,000 years ago.

The most spectacular accumulation of large mammalian

fossils is perhaps the accumulation ofM. columbi remains

at a former sinkhole at Hot Springs, SD, USA (Agenbroad

and Mead 1994; Lister and Bahn 1994). About 50

individual mammoths were discovered up to 1994. It was

estimated that about 100 met their deaths in the sinkhole.

These mammoths died about 26,000 years ago, the trap

being open for 300 to 700 years before it finally filled up

with sediment. The individual mammoths are spread

throughout the sediment, indicating that they did not enterat the same time. Most of the animals were young males,

caught at their most adventurous age. Sinkholes in which

hippos and elephants got trapped were also found in the

Mediterranean region, particularly on Malta.

We assume that a pygmy elephant or a pygmy hippo

could quite easily get trapped in a natural trap. Carrington

(1962) describes how they used to trap elephants in Africa

with pitfalls, which he describes as the simplest and most

effective of all elephant hunting methods widely used in

Africa until recent times.

The supposed behavior of elephants when one of their

number falls into a pitfall was recorded by Pliny nearly two

thousand yeas ago. In his Natural History (book VIII,

chapter 8), he writes: In Africa they take them in pit-falls;

but as soon as an elephant gets into one, the others

immediately collect boughs of trees and pile up heaps of

earth, so as to form a mound, and then endeavor with all

their might to drag it out.

Is not certain how accurate the above description of

elephant behavior by Pliny actually is, but elephants are

known to assist other elephants of their group to free

themselves, particularly when they are trapped in muddy

pools. The pitfall is probably the oldest of all forms of

animal trap, and extinct proboscideans must have fallen

victim to it (Lister and Bahn1994, pp 122123).

Bunimovitz and Barkai (1996) suggested that the lowest

bone deposit at Aetokremnos is natural, not of anthropo-

genic origin. Hadjisterkotis et al. (2000) noted that the

piling of such a large number of hippos in such a small spot

in such steep cliffs usually takes place in the case of water

holes or sink holes with steep sides. Turner and Weaver

(1981, Fig. 7.3, p 108) describe one such death trapwater

hole in the Chocolate Mountains, CA, where the remains of

34 trapped bighorn sheep were found in a steep-sided

natural water tank in 1969. Bighorn were in peak

abundance in the Trap Cave deposits during the cold

phases of the Pleistocene. Two peaks, one at about

18,000 years ago and another 14,000 years ago, coincided,

respectively, with the glacial maximum and the severe

Older Dryas cold phase (Geist1999).

However, there is no geomorphic evidence for such atrap at Aetokremnos. The floor of the rock shelter did not

have any evidence of sedimentation and there are no

features on the cliffs or plateau above Aetokremnos to

account for a repeated natural jump site (Mandel and

Simmons 1997). The presence of one frog and an aquatic

snake could indicate the presence of water that attracted the

birds and mammals for drinking. However, the floor of the

rock shelter did not have any evidence of sedimentation or

other debris indicating the presence of a pond or running

water or a sink hole (Mandel and Simmons 1997, Mandel

1999).

Natural predators

Besides the possibility of a large mammal entering a cave

or a natural trap and dying there, complete animals

carcasses or parts of them can be brought into a cave by a

predator. According to Lister and Bahn (1994), mammoth

remains found in caves frequently were carried in by

humans or other predators. For example, juvenile mammoth

remains were accumulated in Friesenhahn Cave in Texas,

USA, by the extinct Scimitar cat (Homotherium), and in

Kents Cavern in southwestern England, by the Spotted

hyena (Crocuta crocuta). At Kents Cavern, a former hyena

den, thousands of hyena fossils have been excavated,

mingled with bones of juvenile woolly mammoths, many

of which bear the tooth marks of these scavengers. Neither

big cats nor hyenas could have hunted adult mammoths, but

they may have killed young animals.

Since on Cyprus there is no evidence of large predators

capable of killing pygmy hippos or elephants, the accumu-

lation of their bones cannot be attributed to predators or

scavengers. The only predator on Cyprus, besides man, at

the time of Aetokremnos was the extinct endemic genet

G. plesictoides(Steensma and Reese1999). Genets feed on

invertebrates, particularly insects and small rodents. Rep-

tiles, amphibians, birds, and other small mammals are also

taken, as well as wild fruits.

Based on the above reasoning, we therefore reject the

hypothesis that predators or natural traps were responsible

for the collection of hippos and elephants at Aetokremnos.

Conversely, we accept the conclusion of Hadjisterkotis et

al. (2000), Reese (1989, 1995, 1996, 2001; Reese et al.

1999), and Simmons (1991a, b, 1996, 1999, 2001, 2002)

that the bone accumulations are of anthropogenic origin and

Eur J Wildl Res (2008) 54:122133 129


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that the people of the Akrotiri peninsula were responsible

for the extinction of the two endemic mammal species in

that area.

However, it is possible that natural traps were the reason

for trapping these animals in caves elsewhere on the island,

attracted by the cave microclimate with possible availability

of water, or even concentrations of minerals. Shortage of

water under drought conditions conceivably drove them toseek water/moisture in caves or in hollow places in the

sides of cliffs and to look for protection from the blazing

sun during the long dry summers. Packed in these places to

avoid the burning Mediterranean sun and seeking the

moisture oozing from the cave walls, most possibly

suffered the torments of death from dehydration when

water sources gave out, or from thirst and hunger when

caught in natural traps.

The numerous sites on the island yielding large numbers of

trapped or otherwise killed mammals, together with the

consequences of the indiscriminate hunting of hippos and

elephants at Aetokremnos and the subsequent extinction ofthese species on Cyprus, record the final events of extinction.

They provide a complex but readable record of a process

whose outcome is clearly known. These deposits may not

record the death of the very last animal, but they are

evidence of major mortality processes that affected both

pygmy mammals on Cyprus.

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Documents

Reese_considerations on the Potential Use of Cliffs