ZEITSCHRIFT FUR ANGEWANDTE ZOOLOGIE

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1 Sonderdruck aus: ZEITSCHRIFT FUR ANGEWANDTE ZOOLOGIE GERMAN JOURNAL FOR APPLIED ZOOLOGY In Verbindung mit Kart Becker, Hannover; Ralf Garms 1 Hamburg; Karl G6,Bwald; Wurzburg; Waldemar Madel, Ingelheim; Lothar Renwrantz, Hamburg; Harald Rosenthal, Hamburg herausgegeben van PETER-DIEDRICH HANSEN Berlin 78. Jahrgang Zweites Heft 1991 DUNCKER & HUMBLOT / BERLIN

2 Distribution and bionomics of the Nile Soft-Shelled Turtle, Trionyx triunguis, in the Eastern Mediterranean By MAx KASPAREK and RAGNAR KINZELBACH Institute for Zoology, Technical University of Darmstadt (With 7 figures) Key w 0 r d s: Trionyx-Middle East-Zoogeography-Conservation 1. Introduction In the. original scientific description of "Testudo triunguis", FORSKAL (1775) noted that the species was rare in the Nile: "in Nilo rarior". Some 75 years later, it was found to occur in the countries of the Levant by RUSSEGGER (184}), and this wider distribution then became common knowledge (BOETT GER 1879, STRAUCH 1890, LORTET 1887). It was almost a century before the scientific community realised that its modern distribution extends as far north as Southwest Anatolia (BA~O(JLU 1973, BA~O(JLU & BARAN 1977, HATHAWAY 1972). The present contribution continues the collection of data on the Nile Soft Shelled Turtle Trionyx triunguis in the Middle East which began with HAAS (1951), ATATUR (1979) and KINZELBACH (1986). This seems to be justified for the following reasons: (a) Despite its wide range, Trionyx triunguis is threatened either by direct interference by man or by the increasing invasion of its habitats. This is especially true for the Asiatic part of its distribution area. In order to establish a protection strategy, data on distribution and population size are most urgently required. (b) A good deal of new information has become available, partly culled from literature sources not included in the compilations mentioned above, and partly from our own recent field activities in Turkey, Syria, Israel and Egypt.

3 138 Max Kasparek and Ragnar Kinzelbach 2. List of recent sites in the Middle East The sites are listed in groups according to the natural regions, from northwest to south-east (Fig. 1) Ko y c e giz - D aly a n we ti a nd c ompl ex Although the occurrence here of the Nile Soft-Shelled Turtle has become known only recently (BA~OGLU1973), this is now one of the best-known sites in Turkey. In additionto the published finds (ATATOR 1979, GRAMENTZ 1990, KINZELBACH 1986), numerous observations were made by various observers especially in The main occurrence is in the Dalyan River, which connects Lake Koycegiz with the Dalyan estuary. As many as 12 different individuals were observed each day and the localities include e. g. the thermal springs upstream of Dalyan town, including a nearby dead arm of the river, the river within Dalyan town and up and down-stream of it, the area of the fishery cooperative, the river branch near ancient Kaunos, and the outlet of the estuary to the sea. A few records are also from Siiliingiir Golii, which is a part of the Dalyan estuary. The observations there include a sighting of a mating pair on by tourists (cf. KINZELBACH & SCHEMEL 1987, where this observation was erroneously attributed to the Loggerhead Turtle, Caretta caretta). Within Koycegiz Golii, observations are available from the villages of Sultaniye and Horozlar on the western shore of the lake. The shores of the Dalyan River and the Dalyan estuary are generally muddy and covered by a dense vegetation (mainly the reed Phragmites australis), while those of Koycegiz G6hi are muddy, rocky or stony. A site which appears suitable for nesting is the sand dune which separates the estuary from the sea. In fact, f;l.esting was co\lfirmed there in 1987, when R. K. found two plundered nests on , and M. K. (together with K. CORBETT) independently found what appear to be the same nests on Both nests were close to each other (distance c.15 m) and were situated in a steep sand hill about 2.5 m from the water's edge. At exactly the same locality, M. K. again found two plundered nests in the following year, on On the same sand dune, ATATOR (1979) had earlier found a single Trionyx egg near the outlet of the estuary to the sea (Dalyanagzl), on However, no nest could be located on that occasion, So far, successful nesting in the K6ycegiz-Dalyan wetland complex has only been established by H. and M. BREIL. In September 1989 they saw a baby Soft-Shelled Turtle which had been caught by a local fisherman in the Dalyan River within Dalyan village. Its length was about 10 cm. An intensive search in the Dalyan River during the days that followed produced two

4 '-- i +\1.~ -,_ )r!... \ "- ('\-!2 ;<i. ~ so : g a. ~ s: ~ ",,, ~. -, ~, " g, :i 0 ~ f<0 ~. Fig. 1. Distribution of the Nile Soft-Shelled Turtle Trionyx triunguis and the Euphrates Soft-Shelled Turtle Ra/ems euphraticus, inthe Eastern Mediterranean. The numbers refer to chapter 2 of the text. Small dots: single records, large dots: populations, (///) distribution in the Nile basin in historical times. ~ W <0

5 l 140 Max Kasparek and Ragnar Kinzelbach Fig Sun bathing Soft-Shelled Turtles on the banks of Nahal Alexander, Israel. Photograph: M. KASPAREK. observations of juvenile Soft-Shelled Turtles, with estimated lengths of 10 14cm and 20 cm. The banks of an old river branch near the thermal springs above Dalyan village should be considered as a possible nesting site. The

6 Distribution and bionomics of Trionyx triunguis 141 Fig. 4. Habitat of the Soft-Shelled Turtle at Nahal Alexander, Israel. Photograph: M. KASPAREK. banks of the Dalyan River opposite the mouth of the dead arm of the river also provide suitable nesting habitats. The substrate is muddy and earthy, but as the zone between the cotton fields and lbe water's edge is easily heated because of its angle to the sun, nesting seems possible Dalaman river system The main sites are Kargm G6lii (= Kiikiirt G6lii), Kacag6F) and the channel which connects them both with the sea. ATATOR (1979) has called this areas "the richest in Trionyx of Anatolia", and he collected no fewer than 28 1 KocagOl, which is the larger of the two 'lakes, is sometimes also called Kargm GaIu.

7 142 Max Kasparek and Ragnar Kinzelbach specimens there in In 1988, the channel between Incebel Tepesi (Kargm G61ii) and the sea was checked several times between May and August (M. K, A. KlL19 and others). On each occasion, several adult individuals were observed, for example, observations of at least 10 different individuals on a 500 m long section of the channel on Furthermore, one dead individual was found on the bank of the channel and another on the sea shore close to where the channel opens into the sea. Local people reported its occurrence upstream in the channel, within the village of Kaplkargm (R. K, ), and in 1989 local people also confirmed it in Kocag61 (R. K, ). ATATOR (1979) found somenestsona sandysitetothesouth-eastofkargm G61ii: one nest on was some 15 m away from the water's edge. Two more nests were found on at distances of 6-7m and 10m from the splash-line. All three nests were 1-1.5m above the water-level. On , five tracks of the Soft-Shelled TuTtle were recorded in the fine-grained gravel on the left bank of the channel immediately before it opens into the sea (M. K et al.). Four of these had made attempts at laying a clutch of eggs (at distances of 7-32 m from the water), but it was not clear whether they had been successful. On later occasions in the same year, these tracks (the same?) were also recorded by other observers. In the lower course of the Dalaman Gayl, about 150 m upstream from the river mouth, R. K. observed tracks on leading from the river onto a sandy ridge which separates the river from the sea. A. KlLI9 found a dead specimen on the sea shore close to the river mouth on Other records are: One sighted in the channel, to the east of Dalaman Airport, joining Kagm~ay ACISU, on (R. K); track to a nest site on the sea shore 500 m east of Dalaman Airport on (R. K), a sandy site with rocks here and there (Fig. 5); carcass of a specimen killed by fishermen at the mouth of the Kargm~ayACISU on (R. K.). Several carcasses were found around the mouth of Sansu to the north of the Dalaman river mouth (Sangerme). They were swept down the river and carried onto the adjacent beach together with some 20 dead Mauremys caspica, on (R. K.). See Fig E~en Gayl (Patara beach) The only record is by A. KlLI9 et al. (unpubl.) who found a dead specimen on the beach near the mouth of E~en Gayl on

8 Distribution and bionomics of Trionyx triunguis 143. Fig. 5. Nesting habitat of the Nile Soft~Shelled Turtle in Turkey: Track to a nesting site on the Dalaman beach on Photograph: R. KINZELBACH. Fig. 6. The Soft-Shelled Turtle regularly leaves the water for sun bathing. Photograph: M. KASPAREK.

9 144 Max Kasparek and Ragnar Kinzelbach ' Fig. 7. Carcass of a Soft-Shell on the shore near Sangerme (Dalaman region) in Photograph: R. KINZELBACH Antalya area ATATOR (1979) found pleuralia on Lara beach to the east of the town on Belek area ATATOR (1979) collected a dead specimen and the femur of another one at the mouth of ACISU (= Belek Suyu) to the southeast of Belek (Serik region) on Its occurrence there was recently confirmed by R. K., who observed one individual on Manavgat River area In May 1965, SELLA (1981) found one specimen alive in a fishing net at "Side lagoon". The location was described as being 0.5 km from the shore and 12 km from any fresh water. The fishing net had been at a depth of 5 m. As there is no lagoon near Side, we suppose that the author was referring to

10 Distribution and bionomics of Tr1.onyx triunguis 145 the freshwater lake Titreyen GOlii and that the individual was found in the sea. At the same time, in May 1965, SELLA (Le.) also found the disarticulated skeleton of a Soft-Shelled Turtle on the beach near Side. ATATO'R (1979) received information from local fishermen on the occurrence of Soft-Shelled Turtles in the sea near Side and in the Manavgat River. During three visits in , no Soft-Shelled Turtles were observed in the Titreyen Giilii close to Manavgat River mouth (ATAT()R, pers. comm.). However, R. K. recently observed one individual on in the Titreyen Giilii and in a brackish water body east of the mouth of Manavgat Qayl Bozyazl area M. K. and M. Oz received a dead juvenile specimen from children at the mouth of Sansu on They also reported large individuals living in the same creek (cf. BARAN & KASPAREK 1989) Giiksu River Although ATAT()R (1979) received information on the occurrence of Soft Shelled Turtles from local fishermen, he could not confirm these statements with personal observations. RADDATZ found a dead specimen on the sea shore near Paradeniz Gii1ii on (cf. KiNZELBACH 1986), and M. K. found a disarticulated skeleton with a carapace length of 42 cm on the eastern side of the river mouth on (cf. BARAN & KASPAREK 1989). ATATO'R (1979) found plastron fragments at the mouth of a creek to the east of the delta area on (Tersakan Qayt). The exact location could not be identified Qukurova region In the lower section of the Berdan (= Tarsus) river, one large turtle which was observed by local people on (M. K. and A. KiL19) probably belonged to this species. BERK et a1. (1988) did not encounter the species when they sailed from the town of Tarsus to the mouth of the river on 12 and 13 May 1987, but they observed two adults in the sea near the river mouth on May 14 th. A disarticulated skeleton was present on Kazanh beach, which is some 10 km west of the river mouth, in August and September 1988, and another (?) on (M. K.). ATATO'R (1979) reported a dead Nile Soft Shelled Turtle in Mersin harbour on In the Seyhan river, its occurrence was first suggested by GRUVEL (1931) who mentioned a capture offshore, and more recently by BERK et a1. (1988) 10 Angewandte Zoologie 2/91

11 146 Max Kasparek and Ragnar Kinzelbach who observed adults in the sea near the river mouth on 14 and A dead specimen was found on the sea shore to the east of the river mouth on (WWF sea turtle team, cf. BARAN & KASPAREK 1989). PATTERSON collected a specimen in September 1965 on the sea shore near Karata (EISELT 1976). SELLA (1981) found a disarticulated skeleton on the shore near Akyatan Colii in June 1967'and ATAT()R (1979) skeletal fragments from two individuals near Karata on BERK et al. (1988) observed an adult in the sea to the south of Tuzla, not far from the mouth of a drainage channel on In 1988, a dead specimen was found on the beach to the east of Karata on 25.5., and a dead individual (different ones) on the sea shore to the south of Akyatan lagoon on and (WWF sea turtle team, cf. BARAN & KASPAREK 1989). The carcass of a Soft-Shelled Turtle was found near Agyatan ("Akyayan") Coliion (BERK et al. 1988) and (WWF sea turtle team, see BARAN & KASPAREK 1989). BERK et al. (l.c.) found a disarticulated skeleton without a head on the shore of a drainage channel to the west of Agyatan Colii, 3 km from the mouth, on While sailing on the lowest 20 km of the fast-flowing Ceyhan on , BERK et al. (l.c.) came across two single adults and a group of at least five adults near the village of Bebeli. Their body length was estimated at m. This group was sun bathing on sand banks and behaving socially, but was wary of man. Mating behaviour was also observed. Tracks were seen, on several other sand banks further downstream. In the bay of Iskenderun, SELLA (1981) reported a specimen in a fishing net at a depth of 10 m in May From the description of the locality (12 km from the shore, 25 km from fresh water), it seems that it must have been somewhere around Yumurtahk. In the Yumurtahk area, ATATtJR (1979) collected two individuals plus a skeleton which was found on the sea shore on Fishermen reported the regular occurrence of the species in the sea here in 1987 (M. K. and A. KLIG), and BERK et al. (l. c.) saw an adult in the sea near the shore on Fishermen of the <;ukurova are very familiar with Soft-Shelled Turtles which have the local name "Yahudi". After many interviews (A. KILn; and M. K.) it has become clear that they do not occur in the lagoons (but note the remarks by ZAHAVI 1968), but are regularly encountered in the sea Orontes (Asi Nehri): Lower section and Amik G61ti Its occurrence in this river has been known since the paper by RUSSEGGER (1843). HAAS (1951) reported "very large" specimens from Amik Coli! and

12 Distribution and bionomics of Trionyx triunguis 147 ATAT()R (1979) collected one individual at the mouth of the river near Samandagl on , and also found a disarticulated skeleton at Kii~iik Asi Kanah in the area of the former Amik Galii on Amik Galii, which once comprised some 28,400 hectares, was progressively drained in the 1960's (see e.g. KASPAREK, BlLGIN & AKIN 1989). The effect on the population of Soft-Shells is not known. However, the observation of mentioned above indicates that the species could survive in the network of drainage channels Orontes (Nahr al-asi): Ghab basin During our research on this river, we observed only twice an immature individual of about 20 cm in length in a ditch north of the small swamp near Ain Taqa in the Ghab Valley (Upper Orontes) on 25.3 and (KlN ZELBACH 1986, where an incorrect date is given). As the species-identification was not confirmed, it is always possible that these individuals belonged to Rafetus euphraticus Nahr al-kabir area PUNY was probably the first author to mention softshells from this site. After describing the methods of fishing sea turtles in the Red Sea, he turned abruptly to the Mediterranean (IX, 36): "In Phoenicio mari haud ulla difficultate capiuntur ultroque veniunt stato tempore anni in amnem Eleutherum effusa multitudine." The locality mentioned is not on the Red Sea, as some editors have suggested (naively interpreting "Phoenicio marii! as a synonym of the "Red Sea"),.but is undoubtedly in Phoenicia at the mouth of the Eleutheros, which a considerable body of evidence identifies as the modern Nahr al-kabir (South). PUNY is discussing a large turtle which is caught in the sea but which comes in large numbers into the river at a special season. It is clear that inforination referring to one of the sea turtles (Caretta caretta or Chelonia mydas) is confused with that on Trionyx triunguis. All three species may be caught in the sea and all come close to land at a special season, but only one goes into the river: the Soft-ShelL One adult individual was seen in the Damascusfish market on (R. K). It was caught offshore near Tartus, and a fisherman informed R. K that this species is regularly caught there. It probably comes from the nearby Nahr al-kabir (South) delta. This river, which today forms the frontier between the Lebanon and Syria, once had a number of outlets which covered a relatively wide area of wetlands. 10'

13 148 Max Kasparek and Ragnar Kinzelbach Nahr al-kalb (Beirut area) BOETTGER (1879) reported two specimens wbicb were collected by SCHRADER near Beirut in , and STRAUCH (1890) mentioned another specimen which had been collected at Beirut by STAUDINGER as early as All three specimens are labelled "Beirut", but probably refer to Nahr al-kalb. GRUVEL (1931) noted one specimen in the collection of the American University of Beirut which had been caught near the city, probably from Nahr al-kalb. HAAS (1951) mentioned its occurrence in that river, but it remains unclear whether he deduced tbis from the statements of BOETTGER and STRAUCH or whether he had new information Nahr al-litani The occurrence of the Nile Soft-Shelled Turtle was mentioned by HAAS (1951), although he did not give details or the source of this information Nahal Alexander and nearby coastal rivers There are specimens from the Kishon river near Haifa and the Yarkon river (Auja) near Tel Aviv in the collection of the Zoological Institute of the Hebrew University in Jerusalem. From Kfar Vitkiu, which lies between these localities, there is a batch of eggs in the museum (HAAs 1951). SELLA (1981) reported some records from Haifa bay in 1963 and 1972 and one record from Tel Aviv in According to YOM-Tov & MENDELSSOHN (1988), Nile Soft-Shelled Turtles lived in many of the west-flowing rivers of Israel and nested on their banks. However, they disappeared from most of them and are now more or less reduced to one coastal stream, Nahal Alexander (WERNER 1988), where reproduction takes place, Many of the eggs fail to hatch and some hatchlings have various anormalities, presumably due to poisonous and mutagenic effects of many pollutants. Currently, no young turtles live in the Alexander River; the population consists only of large to very large individuals, without apparent recruitment (YoM-Tov & MENDELSSOHN 1988). This was confirmed by M. K. in April 1990, when he observed at least 15 basking individuals on the banks of the river some 3 km above its mouth. None of the turtles was smaller than 50 cm. The water of the river was almost black and awful-smelling (see Fig. 4). The main pollutant seems to be a village whose inhabitants are specialised on Turkey-farming. All the organic refuse including dead Turkeys seems to be released to the river without purification. What is called a "feeding station" for Soft-Shells at Nahal Alexander

14 Distribution and bionomics of Trionyx triunguis 149 may be dead Turkeys, causing an unnatural high population density, eutrophication of the river and the above-mentioned loss of reproductivity. Through the efforts of MENDELSSOHN and the Nature Reserves Authority, the species is now protected in Israel and has been re-introduced to many suitable sites during the last 20 years, e. g. from a captivity population at Tel Aviv University Zoo. There are now about 200 specimens in Nahal Yarqon, Nahal Na'aman, Nahal Tanninim, Lake Kinneret, and some reservoirs in Northern Israel (ILANI 1983, ILANI & BOUSKILA 1982, ILANI & SHALMON 1984). Little is known on reproduction in these areas. The survival rate of the offspring is not known (ORTAL and WERNER, in litt.). A research programme includes the documentation of the development of the population and the registration of all individuals (ORTAL, in litt.) Lake Kinneret (= Lake Tiberias, Sea of Galilee) TRISTRAM (1888) emphasized that there is no evidence for the autochthonous occurrence of this species in Lake Kinneret. However, LORTET (1887) published some observations on Soft-Shelled Turtles at Lake Kinneret which suggest the former occurrence of an autochthonous population. SER RUYA (1978) and TCHERNOV (1988) cited two species of Trionyx from the Pleistocene Ubeidiya deposits 4 km SSW of Lake Kinneret. Soft-Shelled Turtles were released ilt Lake Kinneret in recent years for reintroduction. So, MENDELSSOHN released 2 individuals in 1965/66 (YoM-Tov & MENDELSSOHN 1988). The species is now reproducing at the lake (ILANI 1983). SERRUYA (1978) referred to an individual of Trionyx triunguis which was caught on Lake Kinneret by fishermen on It may have originated from the programme for the re-introduction of the species in Israel (see above). LORTET (1887), and subsequently also SIEBENROCK (1913), discussed the question of species identification without coming to any conclusion. A specific identification would be very welcome, since Lake Kinneret may originally have been colonised either by a population of Rafetus euphraticus or, more probably, by the Nile Soft-Shelled Turtle, invading through the Yizreel plain like other aquatic animals, or by both., Hula Nature Reserve MENDELSSOHN transferred 13 individuals from Nahal Alexander to Hula Nature Reserve in 1965/66. Soft-Shells were subsequently seen there on sev-

15 150 Max Kasparek and Ragnar Kinzelbach eral occasions, and judging by their size some of them reproduced there (YoM-Tov & MENDELSSOHN 1988) Bardawil lagoon SELLA (1981) reported a skeleton on the sea shore near Bardawillagoon in September The nearest fresh water is some 150 km away Nile delta and river Trionyx triunguis was originally widely distributed in the Nile. Almost without exception, all ancient Egyptian representations of turtles show the Nile Soft-Shelled Turtle, beginning with the protodynastic period and continuing, though becoming rarer, down to Graeco-Roman times (FISCHER 1968). From the 19 th dynasty on, the animal was interpreted as an embodiment of evil, hidden in the muddy darkness of the river (H. G. in: Lexikon der Agyptologie 1984). For this reason it was ritually persecuted. Nevertheless, the Soft-Shell was regularly eaten (BOESSNECK 1988). The species became rare later on, For the'19 th century, see e'.g. BOULENGER (1989). FLOWER (1933) receivedfouregyptianspecimens: oneeach from Kom Ombo (Aswan Prov.) in August 1902, from the Embaba district (Giza Prov.) in February 1908, from the Nile near Giza in January 1909, and from Ezbet Semada Saleh (2 miles north of Ibshawai in the al-fayum basin). There is no evidence of its occurrence in the Natron valley (FLOWER l.c.). Eventually, the species disappeared from the Egyptian part of the Nile. Rz6sKA (1976) considered that it had been extinct in Egypt since the 1930's. However, KINZEL BACH (1986) reported two recent observationsin 1985 near Luxorand Gizeh. After re-examining the sites 'in 1989 and questioning fishermen and officials, R. K. came to the conclusion that the record from near Gizeh referred to an escapee, perhaps even to one of the Trionyx species regularly imported as pets by animal traders from South Asia. The other record, from Luxor, is incontrovertible. Since the establishment of the Aswan High Dam and the creation of Lake Nasser in about 1979, this area has been re-settled by the Nile Soft-Shelled Turtle as well as by crocodiles and several birds associated with fresh water (cf. GOODMAN & MEININGER 1989). BAHA EL DIN & BAHA EL DIN (1990) found the shells of two soft-shells with fishermen at Abu Simbel on the western shore of lake Nasser. From this point, single individuals have obviously managed to migrate downstream to the Aswan region and, as already mentioned, to Luxor. A fisherman near Qena knew the animal from his father's time and from Lake Nasser. Asked why it was not found in Qena anymore, he emphasized that the softshell needs rocky

16 Distribution and bionomics of Trionyx triunguis 151 shores to rest on. This must be wrong, since the whole river below the Aswan cataract was previously colonised in spite of its natural lack of rocks. 3. Discussion 3.1. Nesting habitats In Africa, Trionyx uses sand banks and islands in rivers for egg deposition. To the best of our knowledge, habitats of this type are rare through most of the Asiatic part of its range; they are usually restricted to the river mouths. This may explain why the only reliable nesting sites recorded are from marine sand bows and dunes (Dalyan, Dalaman) which separate the sea from more or less distant brackish or fresh water bodies. On the other hand, young Soft-Shells have been ohserved in the Dalyan river which suggests that they also nest on Inland freshwater banks c curr enc e 0 ff s ho re The fossil Nile Soft-Shell or closely related species found in Neogene deposits of Northwest Anatolia, the Danube basin and parts of Central Europe lived in habitats which were always close to shallow, sometimes brackish marine basins. Present-day populations between the Nile estuary and Lake Koycegiz generally live in the lower sections and estuaries of rivers or in brackish former marine bays (e.g. K6ycegiz, Kocag61). Apart from the records discussed above from upper Orontes and Jordan, the Amik G6lii is the only site where the species lives at a distance of more than 30 km from the seashore. It often finds its way into the open sea. GRUVEL mentioned observations of the Soft-Shell off the Syrian coast as early as 1931, but this was considered to be somewhat exceptional behaviour. Since then, HATHAWAY (1972) and SELLA (1981) in particular added some more offshore records. EISELT (1976) quoted offshore catches from West Africa taken from VILLIERS (1958). So, there is no longer any reason to consider the occurrence of this species in the sea as accidental, caused only occasionally by floods. Our additional records provide confirmation of a more or less regular occurrence in the sea. But not every specimen found dead on the beaches, such as the carcasses mentioned above which were found along the splash-line of beaches near Dalaman or near Silifke, proves that the species spends much time at sea. Most of them were victims of pollution or persecution, and had been swept into the sea already dead.

17 152 Max Kaspal'ek and Ragnar Kinzelbach Furthermore, observations near Dalyan and Dalaman suggest that Soft Shells use marine beaches or the inland side of sand bars for egg laying when there are no suitable places at their freshwater habitats. The regular interchange between the sea and a river estuary among an extensive and undisturbed population may be reflected in PLINY'S short note on Eleutheros. There is another source from Antiquity which may suggest that Soft Shells even occurred in the highly saline offshore waters of the Red Sea. Two specimens are depicted in the temple of Hatschepsut ( B. C.) at Deir al-bahari, in scenes portraying Punt or at least the sea route to this southern country. They are in the company of many other aquatic animals which can undoubtedly be attributed to the Red Sea fauna. It is an open question whether the expedition to Punt encountered Soft-Shells in the open Red Sea or near river mouths somewhere far in the south, or whether the artist illustrated a report of large sea turtles by inserting a picture of the more familiar Soft-Shell. This ability to live in the sea or in brackish lagoons enabled Trionyx in the past to expand along the coastline of the Levantine Sea, and even to colonise lakes and shallow marine water-bodies in central and squth-eastern Europe in the upper Tertiary St a t us an d pro gno s e Surveys have suggested that the population throughout the Asiatic part of the range consists of less than 500 adults. In view of the increasing pressure to which it is being subjected, it is clear that by the end of this century the Nile Soft-Shell will have been exterminated in the Mediterranean. At the moment the population is enough for the species to survive. But the number of individuals will decrease rapidly because of the factors discussed below. It is still possible to prevent the extinction of this species in the region under consideration, but counterm~asuresmust start immediately. There are four areas where numbers reach a particularly high density: the Dalyan/Dalaman river systems in south-west Turkey, the <;ukurova delta region in southern Turkey, the mouth of the Nahr al-kebir in Syria, and Nahal Alexander in Israel. Nahal Alexander is the only site where a conservation-programme is underway Factors threatening Trionyx triunguis There are already many factors which are affecting the populations. Their impact will increase very rapidly as the demand for land intensifies, either by the growing resident human populations or by tourism.

18 Distribution and bionomics of Trionyx triunguis Accidents with nets, fishing lines, collisions with boats. - Human consumption. The species was eaten in Ancient Egypt. Being esteemed as a valuable gastronomic luxury for human consumption, the Nile Soft-Shelled Turtle has been hunted persistently for hundreds, or probably thousands, of years. Although this occasionally still occurs in Syria (see above), it is not thought to be a major threat to the species. - River pollution. Damage by direct poisoning, with effects on reproduction, eggs and young; and by indirect poisoning via the food chain. However, adult individuals do not seem to be very sensitive to polluted water, as the Dalaman River (SW Anatolia) and Alexander River (Israel) are extremely polluted by waste water from a paper mill and from a poultry farm, but Soft-Shells are still frequent there. - Persecution by fishermen. The Soft-Shelled Turtle is behated as pest species which is very aggressive and destroys fishing nets. - Changes in the waterbody by alterations in the amount and periodicity of the discharge. Changes in the riverbed, e. g. alteration of the vegetation (removal of submergent and emergent water plants, low-land vegetation on river shores), building activities (concrete quays, artificial subsirates), removing sand banks which are used.for sunbathing or nesting. - Disturbance by boat-traffic (e.g. Dalyan river), fishing, bathing, sitting around the water. - Scientific research. Soft-Shelled Turtles are still being killed for scientific research (ATATOR 1979). GRAMENTZ (1990) reported that he caught four Soft-Shells with fishing-hooks S tr ate g i es for th e pro t e et ion of Triony x triunguis Fishermen in Turkey fear the aggressive behaviour and the jaws of the "turtle with the nose" when it is trapped in their nets or hooked on their fishing lines. Regarded as a pest species, Soft-Shells are killed whenever fishermen come across them. In turtle-ridden Dalyan, horrific stories were deliberately spread in 1987, even by officials: for example, that a Soft-Shell killed a little girl by dragging her into the water or that a single Soft-Shell devours 10 kg of the best quality fish quality (Mugil spp.) each day, thus reducing the catches of the professional fishermen. Several strategies are available to prevent the extinction of this threatened species: (1). Education, information, enlisting public sympathy.

19 154 Max Kasparek and Ragnar Kinzelbach (2) Species protection through international conventions and national legisl~tion. It has to be given CITES status. At present it is regarded as endangered only in Ghana. It must also be included in the Berne Convention (Convention on the Conservation of European Wildlife and Natural Habitats, 1979). (3) Habitat protection. This is the most promising strategy, but must be backed-up by the other two just listed. This has been done for years in Israel, where a conservation programme is on the way to being accepted (ORTAL, in litt.). In Turkey, the establishment of Specially Protected Areas since 1988/., '89 may make it possible' to protect parts of the habitat, e. g. the Siiliingiir G61ii, as strictly protected genetic reserves. Habitat protection can be supported by direct measures, which have proved to be successful in Israel, e. g. building artificial sand banks for egg-laying and offering food Geographical segregation of Rafetus euphraticus and Trionyx triunguis The presence of two different species of Soft-Shelled Turtles in Southwest Asia was recognized as early as RUSSEGGER (1843). In Anatolia, the Mesopotamian catchment basin harbours Rafetus euphraticus (Daudin, 1802). West of it, in the basins of the Ceyhan and Lower Grontes, Trionyx triunguis (Forskal, 1775) is found. MEYLAN (1987) confirmed the clear separation of the Nile Soft-Shell from the Euphrates Soft-Shell, and the latter is now separated even at the generic level The line of separation is not known for certain in the countries of the Levant: it is not known which species inhabits the upper Orontes, nor is it known if Rafetus euphraticus existed in Lake Kinneret (Tiberias) at an earlier period. During the Pliocene and even the Pleistocene, parts of the Orontes and Jordan catchment basins were tributaries of the Euphrates river system (KINZELBACH 1987, KRupp 1987). So it may well be that in the Levantine rift valley Rafetus euphraticus was or still is autochthonous. The records from the Amik G6lii and Ghab should therefore be re-examined. The same applies to Lake Kinneret. However, the introduction of Trionyx triunguis may have resolved the situation there. The population in the coastal rivers of the southern Levant is autochthonous (HAAs 1951), and not just the result of occasional drift from the Nile as was suggested by FLOWER (1933). There are only two observations from the Grontes upstream of Amik G61ii. Unfortunately, the single specimen was neither caught nor photographed. So it cannot be decided whetherthe Ghab population, which in any case was highly relict, belongs to Trionyx triunguis or to the Mesopotamian Rafetus euphraticus. In its middle and upper sections the Grontes is known to belong

20 Distribution and bionomics of Trionyx triunguis 155 to the Mesopotamian faunal region, whereas its lower course is part of the Mediterranean faunal region (KINZELBACH 1986). SERRUYA (1978) named the Soft-Shell found in Lake Kinneret as Trionyx triunguis and considered it to be an early immigrant from Africa, like the crocodile. This view certainly has to be revised because the animals mentioned, together with the hippopotamus which also occurred in the countries of the Levant in historical times, are more likely to be relics of a former (Pikermian) distribution than relatively late immigrants from Africa. On the other hand, all the fossil Trionyx from the Pliocene and perhaps even from the Miocene inwest Anatolia and in Europe belong to T. triunguis or to close relatives (BACHMAYER 1966, TUNA 1988). The Nile Soft-Shell fits into a group of aquatic organisms which were widely distributed all around the Mediterranean Sea during the Upper Tertiary, including parts of the Danube Basin and of Central Europe. They disappeared north of the Alps due to the Pleistocene glaciations. Some of these "palaeomediterranean" animals managed to colonise the lost areas intermittently during the interglacials andlor quite recently, starting from refugia or taking advantage of mans' activities, e.g. species of the mollusc genera Corbicula, Dreissena, Ferrissia and the Crustacea Orchestia cavimana and Atyaephyra desmaresti. The Nile Soft-Shelled Turtle shares with most of these species a certain tolerance of brackish water. The localities of the Tertiary Trionyx in Europe also show close proximity to brackish inland waters or to shallow. coasts. This observation conforms with the behaviour of the recent Soft-Shells around the Eastern Mediterranean. Unlike most of the organisms mentioned above for comparison, Trionyx is extremely thermophilous. The geographic limit of this species, determined by temperature even nowadays, is located much further to the south. The distribution in Anatolia (Fig. 1) coincides with several isotherms, e.g. average number of days with frost below five, mean minimum temperature in January above 2.5 C, average temperature in November above 15 C and in March above 12.5'C (TANOGLU et al. 1961). For this reason the species was not able to expand again towards the north~west. The northwestern limit of its distribution in Anatolia is thus identical with that of the Loggerhead Turtle, Caretta caretta. A combination of habitat and climate was identified as limiting the distribution of that species (BARAN & KASPAREK 1989). Summary The distribution and status ofthe Nile Soft-ShelledTurtle Trionyx triunguis inthe Asiatic part of its range is described by means of a thorough literature review and

21 156 Max Kasparek ahd Ragnar Kinzelbach many new unpublished data. The size of the Asiatic population is thought not to exceed some 500 adults. Important sites are located in Turkey (Mugla and Adana districts), Syria (Nahr al-kabir), and Israel (Nahal Alexander). The species is being increasingly threatened. The reasons for this are listed, and proposals are made for its protection. Confirmation is given that the Nile Soft-Shell occurs. regularly in the sea, invading it for dispersal and using the beaches for nesting. Zusammenfassung Die Verbreitung und der Status der Afrikanischen Weichschildkrote Trionyx triun~ guis im asiatischen Teil ihres Areals wirdanhand einer Literaturtibersicht und zahlreichen neuen Angaben dargestellt. Der asiatische Gesamtbestand wird auf nicht iiber 500 Adulte geschatzt. Verbreitungsschwerpunkte liegen in der Tiirkei (Bezirke Mugla und Adana), in Syrien (Nahr al~kabir) und Israel (Nahal Alexander). Die Art ist zunehmend gefahrdet. Griinde dafiir werden angegeben, Vorschlage zum Schutz werden vorgestellt. Es wird bestatigt, dab die Afrikanische Weichschildkrote regelmiillig im Meer vorkommt. Es dient der Ausbreitung, und Strande werden zur Eiablage genutzt. Acknowledgements For data and discussion we are indebted to Doe;. Dr. M, ATAT'OR, H. and M, BREIL, A. KILIl;, L. LAURENT, R. ORTAL, and Prof. Dr, Y. WERNER, V. VAN DEN BERK provided a map with localities which were omitted from the publication by BERK et al (1988). Mr. A. C. PONT, London, kindly checked the English language of this paper. References ATATOR, M. K. (1979): Trionyx triunguis (Reptilia, Testudines) 'in morfoloji ve osteolojisi, Anadolu'daki biotop ve dagi.li~i iizerindeki ara~tirma1arve biyolojisine clair bazl gozlemler. - Ege 'Oniversitesi FenFakiiltesi Monografiler Serisi 18: 1-75, Izmir. ATATtrR, M. K, S. trctrnco (1987): A preliminary report on some peculiarities in the skull of the Euphrates Soft-Shelled Turtle, Trionyx euphraticu8. - Zoology in the Middle East, 1: 75-79, Heideiberg. BACHMAYER, F. (1966): Ein bemerkenswerter Schildkrotenfund aus dem Ober~Pannon der Schottergrube "Heidfeld" beim Flughafen Schwechat (Wien). - Ann. Naturhist. Mus. Wien 69: BAHAEL DIN, S. & M. (1990): The Abu Simbel experience. - Bull. Om. Soc. Middle East 25: 1-5, Sandy. BARAN I. & M. KASPAREK (1989): Marine Turtles Turkey: Status survey 1988 and recommendationsfor conservation and management. - Heidelberg, iv pp. M. (1973): A preliminary report on a specimen of Soft-shell Turtle from Southwestern Anatolia. - Ege tr.i.1iv. Fen Fak, Hmi Rap. Ser. no.172: BA~O(';.LU,

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