Annales Societatis Geologorum Poloniae (2015), vol. 85: 507–514. doi: http://dx.doi.org/10.14241/asgp.2015.012
VER TE BRATE TRACKS IN LATE PLEIS TO CENE–EARLY
HO LO CENE (?) CAR BON ATE AEOLIANITES, PAPHOS, CY PRUS
Jesper MILN1,2, Georgios THEODOROU3, Da vid B. LOOPE4, Ioannis PANAYIDES5, Lars B. CLEMMENSEN6 &Maria GKIONI3
1
Geomuseum Faxe, stsj³llands Mu seum, stervej 2, DK-4640 Faxe, Den mark; e-mail: jesperm@oesm.dk
2
Nat u ral His tory Mu seum of Den mark,Uni ver sity of Co pen ha gen, ster Voldgade 5-7, DK-1350 Co pen ha gen K, Den mark
3
De part ment of His tor i cal Ge ol ogy and Palae on tol ogy, Ath ens Uni ver sity, Ath ens, Greece; e-mail: gtheodor@geol.uoa.gr
4
De part ment of Earth & At mo spheric Sci ences, Uni ver sity of Ne braska, Lin coln, NE 68588-0340, USA; e-mail: dloope1@unl.edu
5
Cy prus Geo log i cal Sur vey, 1415 Lefkosia, Cy prus; e-mail: ipanayides@spidernet.com.cy
6
De part ment of Geosciences and Nat u ral Re source Man age ment, Uni ver sity of Co pen ha gen, ster Voldgade 10, DK-1350 Co pen ha gen K, Den mark
Mil´n, J., Theodorou, G., Loope, D. B., Panayides, I., Clemmensen, L. B. & Gkioni, M., 2015. Ver te brate tracks in Late Pleis to cene–Early Ho lo cene (?) car bon ate aeolianites, Paphos, Cy prus. Annales Societatis Geologorum Poloniae, 85: 507–514.
Ab stract: In 2005, nu mer ous ver te brate tracks were dis cov ered in car bon ate aeolianites in and around the town of Paphos, in the south west ern part of Cy prus. The main track-bear ing ex po sure is lo cated in a pro tected ar chae o lo-g i cal site near the Alo-gia Solomoni Church in side the city of Paphos, where cross-sec tions throulo-gh tracks are abun dant in ver ti cal ex po sures of the aeolianite along Apostolou Pavlou Av e nue. Some ex po sures show as many as 10 tracks per m2 of ver ti cal ex po sure. Sev eral ad di tional tracks were found in the ex ten sive sub ter ra nean tomb com plex, the Tombs of the Kings, just out side Paphos. The ae olian de posit was formed when west erly to south-west erly winds drove fine- to me dium-grained cal car e ous sand on shore from the beach. This gen er ated low coas-tal dunes, rep re sented by 1–2-m-thick, cross-bed ded sets made up of grainflow and wind-rip ple strata, and sand sheets com posed en tirely of wind-rip ple strata. The sed i ment does not yet have an ab so lute date, but is con sid e-red to be of Late Pleis to cene to Early Ho lo cene age, as are many other coastal aeolianites in the Med i ter ra nean area. The Late Pleis to cene en demic fauna in Cy prus was lim ited to the dwarf hip po pot a mus Phanourios mi nor Desmarest, 1822, the dwarf el e phant Elephas cypriotes Bate, 1902, a small car ni vore Genetta plesictoides Bate, 1903, and (pos si bly) hu mans. The ex posed tracks are 5–15 cm in di am e ter, with a few tracks up to 23 cm in size. This range of size cor re lates well with the es ti mated foot size of dwarf hip po pot ami and dwarf el e phants. This low-di ver sity, en demic is land fauna pro vides a unique op por tu nity to cor re late tracks with trackmakers.
Key words: Cy prus, ae olian, foot prints, probocideans, hip po pot a mus, in su lar dwarf ism, track pres er va tion. Manu script re ceived 17 September 2014, ac cepted 22 January 2015
IN TRO DUC TION
Car bon ate aeolianites are known from a num ber of ar eas in the Med i ter ra nean re gion. Many of these Late Pleis to -cene and Ho lo cene aeolianites con tain ver te brate tracks. Im por tant sites com prise lo cal i ties on Mallorca, Sar dinia, and Rhodes. Well ex posed out crops of Mid dle and Late Pleistocene aeolianites have been known on Mallorca since the clas si cal works of Butzer and Cuerda (1962). Work by Clemmensen et al. (1997), Clemmensen et al. (2001), Fornós et al. (2002, 2009), Niel sen et al. (2004), has shown that the ae olian de pos its formed in a va ri ety of coastal-plain set tings, in clud ing clifffront and dis tal al lu vialfan en vi -ron ments; lo cally ex ten sive cliff-top ae olian sys tems were also de vel oped by dunes that mi grated far in land. Many of the aeolianites con tain tracks and trackways of the ru mi nant
goat Myotragus balearicus Bate, 1909 (Fornós et al., 2002). Tracks are seen on bed ding planes and as smallscale de for -ma tion struc tures in ver ti cal sec tions. Aeolianites also crop out abun dantly along the shores of west ern and north west -ern Sar dinia. These aeolianites are pri mar ily of Mid dle and Late Pleis to cene age (Andreucci et al., 2010ab; Pascuccci et
al., 2014), but rather loosely ce mented car bon ate aeolianites
of Ho lo cene age are pres ent lo cally (Andreucci et al., 2014). Most of the aolianites formed on coastal plains, backed by cliffs or low hills, or in front of val ley-heads. Rare tracks and trackways of “Praemegaceros” cazioti Depéret, 1897, (Artiodactyla, Cervidada) are seen (Fanelli et al., 2007). On Rhodes, trackbear ing aeolianites of Late Pleis to cene to Ho lo cene age were de scribed by Mil´n et al. (2007). The ae
olian de pos its of Rhodes form a lo cal sandramp ac cu mu la tion and con tain ichnological ev i dence of a di verse ver te -brate fauna, com pris ing foot prints made by me dium-sized endemic el e phants, ar tio dac tyls and cam els.
In 2005, nu mer ous tracks were dis cov ered by Theodo-rou (Pro ject UOA Re search ac count 7093 and 3370) in aeolianites in and around the town of Paphos in the south west -ern part of Cy prus (Theodorou et al., 2005). The main track- bear ing ex po sure is lo cated in the pro tected ar chae o log i cal site near the Agia Solomoni Church, in side the city of Paphos (Fig. 1), where crosssec tions through tracks are abun dant in the ver ti cal ex po sures of the aeolianite. Some ex po -sures show as many as 10 tracks per m2 in ex posed ver ti cal sec tions of the aeolianite. Sev eral ad di tional tracks were found in the ex ten sive sub ter ra nean tomb com plex, the Tombs of the Kings, just out side Paphos (Fig. 1). The aim of this study is to pres ent a pre lim i nary de scrip tion of the abun dant ver te brate tracks found in the aeolianites in the area in around Paphos, south west ern Cy prus and to link them to po ten tial trackmakers.
TRACK TER MI NOL OGY
Fos sil and subfossil an i mal tracks are most eas ily rec og nized, where they are ex posed on bed ding planes rep re -sent ing the orig i nal track ing sur face. But, as tracks are three-di men sional, struc tures that com monly de form the original sed i men tary fab ric to con sid er able depths be low the orig i nal track ing sur face, it is pos si ble to rec og nize tracks in ver ti cal sec tions or ran dom ero sional cuts through bed ding (Loope, 1986; Allen, 1989, 1997; Lea, 1996;
For-nós et al., 2002). The orig i nal sed i men tary sur face where the an i mal trod is termed the “track ing sur face” sensu Fornós et al. (2002), and the an i mal re spon si ble for the track is the trackmaker. The di rect im pres sion of the track-maker’s foot into the track ing sur face is termed the “true track” (Lockley, 1991). The force of the trackmaker’s foot not only de forms the track ing sur face, but is trans ferred ra di ally out ward into the sur round ing sed i ment, caus ing de -for ma tion of the subjacent lay ers as well (Allen, 1989, 1997). The de for ma tion struc tures in the lay ers subjacent to the true track at the track ing sur face are termed “un der-tracks” (Lockley, 1991). Undertracks pre serve less de tail than true tracks; they be come suc ces sively shal lower and wider and pre serve suc ces sively fewer an a tom i cal fea tures down ward (Man ning, 2004; Mil´n and Bromley, 2006, 2008; Jack son et al., 2010). The ra dial pres sure of the foot fur ther cre ates a “mar ginal ridge” of dis placed sed i ment around the tracks.
In track ing sub strates of a con sis tency that al low the trackmaker’s foot to pen e trate the sur face and sink down into the sub strate, the foot can cre ate ver ti cal or in clined walls from the bot tom of the true track to the track ing sur -face, termed trackwalls (Brown, 1999) or shafts (Allen, 1997). When the trackwalls are in clined, the track at the sur -face ap pears wider than the true track at the bot tom of the track and is termed the “over all track” (Brown, 1999). When tracks are emplaced in dry, loose sed i ments, the trackwalls col lapse on re moval of the foot, thus de stroy ing the shape of the true track; in ex treme cases, only a bowl-shaped de pres sion on the sed i ment sur face is left. If the track sub se quently is cov ered by sev eral thin ner lay ers of sed i ment, the lay ers will drape the con tours of the track and Fig. 1. Lo ca tion map. A. The Is land of Cy prus is lo cated in the east ern part of the Med i ter ra nean Sea, just south of Tur key. B. Map of Cy prus. Paphos is lo cated on the south west ern end of Cy prus. C. City map of Paphos with the two track-bear ing lo cal i ties in di cated.
can form a shallowing-up ward, stacked se quence of “ghost tracks” sensu Fornós et al. (2002). If ob served on hor i zon tal sur faces, these ghost tracks may be mis in ter preted as un der- tracks.
SEDIMENTOLOGY
The aeolianites of Paphos con sist of 1–2-m-thick cross- bed ded sets of both grainflow and windrip ple strata, dip -ping to wards the south-west (Fig. 2A, B), and sand sheets char ac ter ized by flat-bed ded, wind-rip ple strata. Rhizoliths larger than 1 cm in di am e ter are abun dant (Fig. 2C), in di cat -ing that woody plants par tially sta bi lized the dunes. The sed i ment con sists of fine- to me dium-grained cal car e ous sand, which was blown on shore from the beach, gen er at ing low coastal dunes and ae olian sand sheets.
Thin-sec tions of the aeolianite re veal meniscate cal cite ce ments (Fig. 3A) as well as abun dant micrite en ve lopes (Fig. 3B). The track-bear ing aeolianite does not yet have an ab so lute date, but is con sid ered to be of Late Pleis to cene to Early Ho lo cene age, as are many other coastal aeolianites in the Med i ter ra nean area (e.g., Fornós et al., 2009; Andreucci
et al., 2010b; Pascucci et al., 2014). This age is sup ported
by the fact that no Plio cene or Early Pleis to cene mam mals are known from Cy prus.
TRACK EX PO SURES
Agia Solomoni site
The main lo cal ity is within the city of Paphos, by the Agia Solomoni Church, and is a sub ter ra nean cem e tery, carved into the aeolianites. All the ob served tracks are found
VER TE BRATE TRACKS IN LATE PLEIS TO CENE–EARLY HO LO CENE, CYPRUS
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Fig. 2. Ae olian out crop at the Agia Solomoni ar chae o log i cal site in Paphos. A. Ex posed sec tion of the aeolianite, at the en trance to one of the sub ter ra nean burial cham bers. B. Rose di a gram of dune crossbed dip di rec tions show ing that the dunes were formed by sand, trans -ported on shore by south-west erly winds. C. Sec tion of the aeolianite with abun dant rhizoliths.
Fig. 3. Pho to mi cro graph of aeolianite. A. Quartz grains (Q) and di verse bioclasts, in clud ing red al gae (R), are vis i ble. Meniscate cal cite ce ment (ar rows) is con cen trated at points of grain con tact, and was pre cip i tated above the wa ter ta ble (within the vadose zone). Ce men ta tion postdates track for ma tion. B. Micrite en ve lope around a for mer ar agon ite grain (dashed line). The ar agon ite was dis solved af ter ae -olian trans port of the grain and its burial within the dune. Per co lat ing me te oric wa ter dis solved the ar agon ite, leav ing only its out line and the tubes within it that had al ready been coated by small cal cite crys tals. Ar agon ite is more sol u ble than cal cite and is com monly dis solved dur ing early fresh wa ter diagenesis.
ex posed in ver ti cal sec tions in the ae olian out crops and ap pear as flatbot tomed, bowlshaped de pres sions in the lam i -nated sand sheets. Some ex po sures show as many as 10 tracks per m2 of ver ti cal sec tion (Fig. 4A–C). At some ho ri
-zons, the tracks ap pear at evenly spaced in ter vals (Fig. 4A), and along other ho ri zons, tracks ap pear in pairs that partly over step each other (Fig. 4B). Hor i zon tal ex po sures of the track ing sur faces were not en coun tered at any lo cal ity. Fig. 4. Tracks at the main ex po sure at Agia Solomoni. A. Ex posed sec tion of the aeolianite show ing abun dant ho ri zons with tracks and rhizoliths. B. Sin gle tracks, show ing ho mog e nous in fill ing. C. Ho ri zon with two pairs of partly over lap ping tracks. D. Sev eral tracks ex -posed at the base of over hang ing bed.
How ever, an over hang ing part, show ing the nat u ral cast of a par tial track ing sur face, re vealed the con tours of a num ber of tracks, pre served in pos i tive epirelief be low the over hang (Fig. 4D). The ex posed track casts, how ever, are only pre served as bowlshaped casts and do not re veal any an a tom i -cal de tails of the trackmakers’ feet.
At the main ex po sure along Apostolou Pavlou Av e nue, a to tal of 87 tracks were mea sured, and their di men sions plot ted against their fre quency (Fig. 5). The ma jor ity of the tracks were be tween 4 and 16 cm in di am e ter, with sin gle tracks up to 23 cm in di am e ter.
The Tombs of the Kings
A sec ond lo cal ity with tracks was found 2 km north-west of Paphos har bour at the UNESCO World Her i tage Site, the Tombs of the Kings (Fig. 1). The Tombs of the Kings are an ex ten sive sub ter ra nean cem e tery dat ing back to 300 BC, where all the graves have been carved out of the rel a tively soft aeolianite. This has cre ated a large num ber of clean, well-ex posed ver ti cal sur faces in the aeolianite. The tracks are less abun dant here than at the main lo cal ity in Paphos, and only sin gle tracks were en coun tered, ex cept at one sur face in the atrium of tomb num ber 6 (Hadjisavvas, 2011), where two con sec u tive tracks were ob served in the same ho ri zon (Fig. 6).
DIS CUS SION
All the ob served tracks ap pear in the sandsheet de pos its and the in fill ing of the tracks is most com monly ho mog e -nous and structureless, which shows that the in fill ing of the tracks oc curred rap idly af ter the tracks were emplaced. On the ba sis of the anal y sis of the sed i ment and the pres ence of nu mer ous rhizoliths, the au thors in ter pret the tracks to have
been made on a lowre lief land scape com posed of partly veg e tated, small dunes and ae olian sand sheets. Al though dis so -lu tion of aragonitic skel e tal ma te rial led to rapid lithification by vadose cal cite, the tracks must have been emplaced in uncemented sand quite soon af ter its de po si tion.
The sizes of the tracks range from 4 cm to 23 cm in di -am e ter, with the ma jor ity be tween 5 and 15 cm in di -am e ter. How ever, when mea sur ing di am e ter in ver ti cal sec tions of tracks, the mea sure ment ob tained must be re garded as a min i mum di am e ter, as it of ten is im pos si ble to see if the sec -tion is through the mid dle of a track (the true di am e ter), or a tan gen tial sec tion (a smaller di am e ter).
Pos si ble trackmakers
The en demic Late Pleis to cene–Ho lo cene fauna of Cy prus, based on skel e tal re mains, is lim ited to very few spe -cies and a close cor re la tion be tween tracks and trackmakers should be pos si ble. Dwarf el e phants and dwarf hip pos have been doc u mented on Cy prus from Late Pleis to cene lay ers at Aetocremnos dur ing the ex ca va tions car ried out by A.
Sim-VER TE BRATE TRACKS IN LATE PLEIS TO CENE–EARLY HO LO CENE, CYPRUS
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Fig. 5 Graph show ing the fre quency of the tracks and their mea sured di am e ters.
Fig. 6. Tracks from the UNESCO World Her i tage Site, the Tombs of the Kings. A. Tracks found in the walls at the atrium of tomb num ber six, in di cated by white cir cle. B. Close-up of the two tracks, cir cled in A.
mons in the 1990s and at Agia Napa since 2001 by on go ing ex ca va tions by G. Theodorou. Both sites have brought to light an en demic fauna com posed of abun dant hip pos,
Pha-nourios mi nor Desmarest, 1822, and scarce el e phant bones
(Theodorou et al., 2007a).
Bate (1903, 1905) doc u mented the oc cur rence of a very small el e phant, Elephas cypriotes Bate 1902, the fos -sils of which have been col lected at nu mer ous sites all over Cy prus (Reese, 1995). How ever, the pos si bil ity of the oc -cur rence of a larger el e phant was dis cussed as early as 1929 by Vaufrey (1929). Later au thors also men tioned the pos si -bil ity of the ex is tence of a larger el e phant at Achna (Boekschoten and Sondaar, 1972). The ex is tence was first con -firmed in 2005, when a unique el e phant skull was ex ca vated by the Na tional Kapoditstrian Uni ver sity of Ath ens in col -lab o ra tion with the Geo log i cal Sur vey of Cy prus on the Xy- lofagou coast (Theodorou et al., 2005). The ma te rial from Xylofagou and Aetocremnos is com pa ra ble with the di men -sions of E. tiliensis Theodorou et al., 2007b, con firm ing the ex is tence of at least two Pleis to cene el e phant pop u la tions on Cy prus (Theodorou et al., 2007a). The Agia Napa el e phant re mains from the Phanourios mi nor site are too frag men tary to prove the ex is tence of the larger el e phant spe cies.
The rel a tive ages of the two pop u la tions are cur rently un known. It is clear, how ever, that the Aetocremnos find -ings prove the ex is tence of the larger el e phant spe cies at least 10,700 years ago. The Xylofagou skull (Theodorou et
al., 2005) was ex ca vated at a coastal lo cal ity from a
fossi-liferous layer over lain by lay ers of brack ish-wa ter sed i -ments de pos ited at least to a height of 8 metres above the mean sea level. This in di cates de po si tion within the cli ma tic max i mum of 60 – 70,000 years ago (Iliopoulos et al., 2011). The scarce el e phant re mains, ex ca vated at the hippo site at Agia Napa, have been dated at 13,500–11,000 years BP (Athanassiou et al., 2014). The avail able skel e tal ma te rial of the dwarf hippo, Phanourios mi nor Desmarest, 1822, sug -gests only one pop u la tion on Cy prus, as the size vari a tion falls within the nor mal dis tri bu tion for the ge nus.
A third sus pect for hav ing a role in the oc cur rence of trace fos sils in Cy prus is Homo sa pi ens. Up to now, there is no di rect ev i dence for a hu man pres ence on Cy prus dur ing the Late Pleis to cene. Dur ing the ex ca va tions that took place
at Agia Napa, on the east ern side of Cy prus, a well-preser-ved par tial skull and some ex trem i ties of an en demic genet,
Genetta plesictoides (Bate, 1903) were ex ca vated from the
un dis turbed Phanourios mi nor lay ers, dated at 13,500– 11,000 years (Theodorou et al., 2007b). Ge nets could not have mi grated by swim ming to Cy prus as el e phants or hip pos did. So, it is nec es sary to dis cuss the pos si bil ity that 1) ge nets fol lowed hu mans on their early trips well be fore the de po si -tion of the fossiliferous lay ers of Agia Napa, or 2) that they mi grated to the is land on nat u ral rafts. The pos si bil ity that hu -mans ar rived at Cy prus dur ing the last main ep i sode of low sea level can not be ex cluded, and Genetta plesictoides pos si -bly ac com pa nied them. The fact that the Agia Napa par tial skull is slightly mod i fied sug gests that Genetta plesictoides was on the is land for some con sid er able time be fore the spe- cimen was fos sil ized with the Phanourios mi nor re mains, that is, well be fore 13,500 y BP.
No data on the es ti mated foot sizes of ei ther Phanourios
mi nor or Elephas cypriotes are avail able, but on the ba sis of
skel e tal re con struc tions with an es ti mated amount of soft parts added, the au thors es ti mate the feet of Phanourios mi
-nor to have been 8–12 cm in di am e ter, and the feet of Elephas Cypriotes to be 20–25 cm in di am e ter. Feet of el e
-phants and hip po pot ami are sub-cir cu lar in out line with very short, blunt dig its, and as such, are likely to leave cir -cu lar im pres sions in the ae olian sands. Hu man feet on the con trary are elon gated, roughly three times lon ger than wide. There fore the di am e ter of a hu man track ob served in cross-sec tion can vary with a fac tor of three ac cord ing to the ori en ta tion of the sec tion ob served, con trary to the rounded feet of hip po pot ami and el e phants. When it is taken into con sid er ation that the ob served sec tions do not nec es sar ily rep re sent the max i mum di am e ter of the tracks, but also can rep re sent tan gen tial sec tions, the ob served sizes of tracks (Fig. 5), can con vinc ingly be ex plained by the avail able trackmak ers (Fig. 7).
Pre his toric, sea sonal vis its be tween Cy prus and the east ern Med i ter ra nean main land, as sisted by cli ma tic os cil -la tions and sea-level changes, could ex p-lain the de pos its at Akrotiri-Aetokremnos (~11000–10000 BC). The old est, un til now, per ma nent set tle ment on the is land seems to have been at Parreklishia Shillourokambos, around 8000 BC – Fig. 7. Sil hou ettes of the pos si ble trackmakers from the lim ited en demic fauna of Cy prus. All drawn to the same scale. A. Dwarf hip po -pot a mus, Phanourios mi nor. B. Dwarf el e phant, Elephas cypriotes, with the pos si ble larger spe cies in di cated in lighter grey. C. Hu man, Homo sa pi ens. D. Genet, Genetta plesictoides
the so-called PPNB (Pre-Pot tery Neo lithic B) pe riod. The round houses and the deep pits that pos si bly served as wells are the main char ac ter is tics of that pe riod. They are also found at Kastros and a lit tle bit later in the 6th mil len nium BC, and also in the Aceramic phase, at Khirokitia. The in -ter ac tion and ex change be tween Cy prus and the nearby main land cre ated a mixed-farm ing sub sis tence econ omy, based on the in tro duc tion of wild an i mals such as pigs, sheep, goat and cat tle. Later on, this was fol lowed by the in -tro duc tion of do mes ti cated plants and an i mals and, in the mid-third mil len nium BC, of met al lurgy. Clearly the Late Pleistocene–Early Ho lo cene age of the aeolinites per mits dis cus sion of the pos si ble role of hu mans in the for ma tion of some of the Paphos tracks (Bar-Yosef, 2001; Guilaiane and Briois, 2001; Hetherinton and Reid, 2010; Ferentinos et al., 2012; Knapp, 2013).
To sum up, the pos si ble trackmakers of the Paphos tracks are: 1) two pop u la tions (one small- and one large-bod ied) of el e phants; 2) dwarf hip po pot ami; and 3) pos si bly hu mans. No tracks small enough to have been emplaced by ge nets have been dis cov ered so far. Ab so lute dates for the Cyp riot en demic mam mals and hu mans are still highly in ad e quate. This is mainly true for the time of ar rival of these spe cies. Better known is the time of ex tinc tion of the larger el e phants from Aetocremnos and the ex tinc tion of dwarf hip pos, el e -phants and Genetta plesictoides from Aetocremnos and Agia Napa. The sizes of the Agia Napa el e phants must be dis -cussed in the fu ture if new ma te rial should come to light. In ad di tion, ab so lute dates for the aeolianites of Paphos are still miss ing.
Sites of fu ture in ter est
Since their ini tial finds of tracks in the Paphos aeolia-nites (Mil´n et al., 2009) and the Kattavia aeolianite of Rho- des (Mil´n et al., 2007), the au thors started a de tailed search for fur ther lo cal i ties with pos si ble track-bear ing aeolinites in the nu mer ous Greek is lands of the east ern Med i ter ra nean. So far, pos si ble tracks have been dis cov ered in aeolia-nites on the Greek is land, Elaphonisi Peloponnese, and a great num ber of tracks are found in the Zakynthos (Zante) Is land aeolianites. At both sites, the sizes of the tracks sug -gest that the tracks were made by Qua ter nary el e phants that in hab ited the is lands, un til they be came ex tinct dur ing the Late Pleis to cene. So far, no el e phant skel e tal re mains have been doc u mented from the Zante and Elaphonisi is lands. Other ex tremely scarce, small tracks have been lo cated in the Phalasarna area of south ern Crete in very young aeolianites.
Some of the nu mer ous tracks found at the Is land of Zante (Zakynthos) are large, but to date no el e phant skel e tal re mains have been found. How ever, Up per Palaeo lithic stone tools have been found in the aeolinites or at the over -ly ing palaeosol, in di cat ing that hu mans could have been the pro duc ers of those tracks (Kourtessi-Philippakis and Sorel, 1996). Very scarce re mains of nor malsized el e phants, in -clud ing a mo lar, have re cently been col lected from the sea bed at Kefallonia, in shal low wa ter. This site is cur rently un -der study by Theodorou, but up to now there were no more ad di tional dis cov er ies on the sea bed.
CON CLU SION
The coastal aeolianites, ex posed in sub ter ra nean grave com plexes within and around Paphos, southwest ern Cy -prus, were formed by south-west erly winds, blow ing fine-to me dium-grained cal car e ous sands on shore. The aeoliani-tes are pre sumed to be of Late Pleis to cene–early Ho lo cene age, and con tain abun dant ver te brate tracks, ex posed in ver -ti cal sec -tion.
The lim ited en demic fauna of Cy prus, con sists only of dwarf hip po pot ami, dwarf el e phants and ge nets. No tracks small enough to have been emplaced by ge nets were ob -served, in di cat ing that all of the tracks be long to hip pos and el e phants, with the pos si bil ity that some might have been made by early hu man set tlers.
Ac knowl edge ments
We would like to ded i cate this pa per to Rich ard G. Bromley, whose life long de vo tion to ichnology has been an end less source of in spi ra tion for us all. We thank the re view ers Joan J. Fornós and Spencer G. Lucas for their pos i tive and con struc tive com ments on the manu script.
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