Salamandra infraimmaculata, Martens, 1885, Martens, 1885

in den Bosch, Herman A. J., 2021, Natural history notes on the Lebanese Fire salamander, Salamandra infraimmaculata (Salamandridae), Pod @ rcis 12 (1), pp. 4-15 : 3-10

DISTRIBUTION IN LEBANON

EISELT (1958) restricted the type locality "Syrien" by neotype designation to "Bscherreh (Becharreh) in nördlichen Libanon ", currently more commonly written Bcharrè. As the nation Lebanon is a fairly recent creation from 1943, and the type locality of S. infraimmaculata is Bcharrè , the form is presently more properly denoted as the ' Lebanese' Fire salamander, although at the time of the original description the area was denoted as Syrian and the form is consequently sometimes still referred to as the ' Syrian' Fire salamander. And indeed VON MARTENS (1885) just stated ' Syrien', and furthermore in my opinion provided a just as pitiable description copied from Ehrenberg (no citation given) “... bezeichnet als var. infraimmaculata … bei welchem die Rückenflecken eine unregelmässige Reihe auf der Mitte des Rückens bilden und alle Seitenflecken fehlen...” [described as var. infraimmaculata …. in which the dorsal spots form an irregular row on the middle of the back and all lateral spots are absent]. Remarkably enough, the doubtless originally nominate character of an immaculate black belly, is completely absent in the oldest descriptions (since 1885). I found no earlier citations. Added below is an excerpt of a Lebanese herpetological checklist I started to compile years ago (e.g. IN DEN BOSCH, 1998), but do note that this has not been updated since 2003 and is just meant to give a general impression of the local finds. Furthermore, a synonymy listing seems befitting here.

SYNONYMY

To avoid confusion by the various names encountered in literature over the years, the following synonymy is given, listed by date of publication, plus localities from literature as well as from museum records. The original citations can be found under the heading Literature.

Salamandra maculosa var. infraimmaculata VON MARTENS, 1885: 195 .

[“Loc.: Syrien. ...bei welchem die Rückenflecken eine unregelmässige Reihe auf der Mitte des Rückens bilden und alle Seitenflecken fehlen...”] The name infraimmaculata was proposed by Ehrenberg (his source unmentioned) but only published by VON MARTENS (1885) with the above mentioned – rather meagrely defined – line.

Salamandra maculosa Laur. - PERACCA, 1894: 18.

Salamandra maculosa - WOLTERSTORFF, 1905, 283.

Salamandra maculosa f. orientalis Wolt. - WOLTERSTORFF, 1932: 41.

Salamandra salamandra orientalis Wolterstorff, 1932 - MÜLLER & WETTSTEIN, 1933: 135.

Salamandra maculosa orientalis - WERNER, 1935: 227.

Salamandra salamandra orientalis Wolterstorff, 1932 - WERNER, 1939: 213.

Salamandra salamandra infraimmaculata - MERTENS, 1948: 187.

Salamandra salamandra infra-immaculata - FREYTAG, 1955: 72.

Salamandra salamandra infraimmaculata Martens - EISELT, 1958: 131.

Salamandra infraimmaculata - GASSER, 1978: 636.

Salamandra salamandra - WERNER & AVITAL, 1980: 192.

Salamandra salamandra infraimmaculata - DEGANI, 1986: 105.

Salamandra infraimmaculata infraimmaculata - JOGER & STEINFARTZ, 1995: 35.

Salamandra salamandra L. - HRAOUI-BLOQUET, 1997: 212.

Salamandra salamandra infraimmaculata Martens, 1885 - SABEH, 1997: 181.

LOCALITIES AND MATERIAL

Localities - Bcharré (1400 m) ( WOLTERSTORFF, 1932); Bcharré (1400 m) ( WERNER, 1935); surroundings of Bcharré, 1400-1600 m, 8 and 16vi1931 (two adults, one subadult) ( MÜLLER & WETTSTEIN, 1933); Cheba'a (in the most southern Lebanon) ( MÜLLER & WETTSTEIN, 1933); Mechmech ( Heinrich pers. comm. to Bischoff ); larvae, Haret Jandal , approx. 2 km N Bater, N33°37.574', E035°36.710', 900 m, (obs. In den Bosch, 6vi1996); Faraïya, (approx. 1325 m, pers. comm. locals to In den Bosch, 18v1998); Mt. Hermon (near Cheba'a?) ( PERACCA, 1894); Mt. Hermon ( WERNER & AVITAL, 1980).

Material - Aaqbé (3 km west of Rashaya) BM(NH) 1920.1.20.2373 ; Aammiq Swamp , J.R. Childress, 11v1969: CAS 159048 [not seen]; Broummana, M. Thompson: BM(NH) 1957.1.1.4; Machmouché (Jezzine), 900 m, Phares: BM(NH) 1937.6.1.1; Mt. Lebanon , from Zoological Museum Giza: BM(NH) 1927.8.12.11 . EISELT (1958) used material mentioned above under localities Bcharré (3 specimens NMW); Mt. Hermon (1 ZMT), plus Beirut (1 NMP) and Lebanon (1 NMW, 2 NMP, 2 HUJ). NMW 15501 he designated as neotype because of the loss of the holotype, and at the same restricted the terra typica to the surroundings of Bcharré .

Range - The range of nominate species is Israel, Lebanon, Syria, Hatay region, Turkey ( DEGANI, 1986; BARAN & ÖZ, 1994; JOGER & STEINFARTZ, 1995; BARAN & ATATÜR, 1998). The species reaches its southern distribution limit in the mountainous part of northern Israel, with the southernmost population living on Mt. Carmel near Haifa.

ECOLOGY

In e.g. DEGANI (1986) and WARBURG (1994) we find a lot of natural history data concerning northern Israel, DEGANI (1994) wrote on physiological adaptation to xeric habitats, and in SHARON et al. (1997) oogenesis and effect of xeric environments is described for Israeli populations close to Lebanon's southern border (Tel-Dan and Mt. Meron). In ecological respect publications on Israeli animals distinguish, probably correctly, between the populations since the most northern one is connected with a stream in which the larvae are deposited, the other two rely on pools and thus are much more dependent on rainfall. Ovulation probably takes place during spring and parturition occurs mainly between November and March when water is available and thus gestation takes less than a year. I found larvae of 1.5-3.0 cm in June in fairly slow flowing streams and adjacent small puddles, where they seemed to prefer depths of 20-30 cm, to a maximum of 50 cm, with a concentration of larvae found close to rocks and/or roots of plants growing on the banks. They live on all kinds of small invertebrates; I noticed many Gammarus . Water temperature was between 10-15°C. In general the species is active at night, with rain sometimes during the day. The animals copulate on land; the female deposits well-developed larvae in brooks.

SABEH (1997) collected the species in Lebanon between 560-1500 m (no localities mentioned). HRAOUI-BLOQUET (1997) gives 650-2000 m (again without localities). She also considers the animals to be viviparous; an obvious overstatement since larvae are of course deposited in the water (e.g. WARBURG et al., 1979), and not fully developed young salamanders appear. Nevertheless, births of fully developed juveniles are known for Salamandra salamandra populations in Spain and Portugal ( FREYTAG, 1955; FACHBACH, 1969; WOLTERSTORFF, 1928; LOURENÇO et al., 2019) and from Salamandra algira in Morocco (see DINIS & VELO-ANTÓN, 2017), but evidence of such a phenomenon in S. infraimmaculata is completely lacking. WERNER (1935) found a recently metamorphosed specimen near Bcharré in May.

ESTERBAUER (1992) reported for the Golan only larvae. Metamorphosed animals live in moist areas near streams. During the daytime these live concealed in holes, under stones, roots or stone heaps. They prey on rain worms and various insects.

REMARKS

MERTENS (1948) determined that Salamandra salamandra infraimmaculata Martens from Syria and Salamandra salamandra orientalis Wolterstorff from the Taurus mts. near Adana were identical and that infraimmaculata – a name used by Ehrenberg but only published with an extremely succinct description by von Martens – enjoyed priority. Based on serum protein patterns GASSER (1978) tentatively regarded the formerly as a subspecies of S. salamandra considered form a full species: Salamandra infraimmaculata .

The taxonomically interested are referred to a recent internet listing: https://amphibiansoftheworld.amnh.org/Amphib ia/ Caudata / Salamandridae / Salamandrinae /Sal amandra/Salamandra-infraimmaculata DEGANI (1986) grouped the Israeli salamanders with the Lebanese based on morphological evidence and plasma electrophoresis and regarded these as con-subspecific with S. s. infraimmaculata . VEITH (1994), however, confirmed the specific status of S. infraimmaculata . JOGER & STEINFARTZ (1995) stated that all Asian Salamandra should be included in the species S. infraimmaculata . For Lebanon, based on Israeli animals, they arrived at S. i. infraimmaculata . This seems somewhat crude, but they did sample two (of the three) populations in Israel of both the xeric and moist habitat represented in that country, and found no differences. In a more recent morphological paper concerning Israeli animals SHARON et al. (1997) do not seem to recognise the species status of the form and also attribute the taxon to the wrong author: Salamandra salamandra infraimmaculata Mertens. VEITH et al. (1998) hypothesised that the Iberian, Central and Eastern Mediterranean populations separated more or less simultaneously about 10 Mio. yrs BP.

The locality of PERACCA (1894) is deduced from the journey of Dr. Festa where Shuba is mentioned as base. WERNER (1939) reported "Nahr el Kebir bei Antiochia" as based on Wettstein (1928) where, however, the find is not described. It is referred to in MÜLLER & WETTSTEIN (1933) as "Nashr el Khebir bei Antiochia, Nordsyrien (coll. 1891, don. Steindachner)" and in Wolterstorff (1932) as "Nahr el Khebir bei Antiochia". This could lead to the assumption that the species is also present in the extreme northern part of Lebanon, since the Nahr el Kebir (or Nahr el Kabir) is the border river with Syria. EISELT (1966: 430) also noticed this and as a second alternative suggested the Nahr el Kebir which flows into the sea 10 km south of Hamidié (near Latakia). The latter seems most likely, since it is close to the present-day Hatay ( Turkey), formerly also known as Antiochia.

As maximum size in my vivaria I have measured a male of 115+ 90 mm and a female of 121+ 132 mm (head-body + tail), a total size of 205 mm and 55 g for a male, and a 253 mm and 115 g for a (possibly pregnant?) female. A pair from the initial batch, later donated to Sergé Bogaerts, measured 26 and 20.5 cm in March 2021. The sizes mentioned in the literature, 31.6 cm given by ESTERBAUER (1992) as reported to him, and by EISELT (1958) who deposited a 28 cm specimen (NHW 16148) in 1959 from Tarshiha, Israel, as well as a specimen from Mt. Carmel ( Israel), seem quite large compared to my captive specimens. WOLTERSTORFF (1905) already mentioned 30-32 cm for the Near-East and Syria.

It is remarkable that larvae said to have been obtained near Alexandria in the last century may actually have been caught in Lebanon by vacationing European residents of that Egyptian town who released these ( FLOWER, 1933).

One of the first reports containing ethologicalreproductive data on the Near Eastern form is DEGANI & WARBURG (1978), further WARBURG et al. (1979), and a summary of many findings at the southern limits of its distribution in DEGANI (1994). Larvae are said to be mostly produced in Nov-Dec, also Sept. rarely Aug., and in March-April, with 30-200 per female, dimensions 0.13- 0.40 g, 25-40 mm, delivered over 1-4 days per batch. The maximum amount of 200 seems quite a lot.

HOUSING AND FOOD

The wild-caught larvae were at first housed in a relatively small 40x25x 25 cm glass aquarium with some water plants (mainly Elodea ), cork bark and stones to avoid drowning. They were fed small invertebrates like Daphnia , Cyclops and whatever else was collected in my local Dutch ditches, small earthworms and even tiny mealworms, although the latter soon drowned and when non-moving were no longer of interest. A similar diet, but with larger prey, was fed to the (sub-) adults. With increasing size the animals were accommodated in a larger glass cage: 40x40x 100 cm fitted with a small plastic pond of 20x35x 6 cm. Water circulation was provided by an air stone connected to a small aquarium airpump.

In 1997-8 the animals were kept during the winter months in an unheated room with temperatures around 10°C. The salamanders stopped being active. With rising temperatures their normal night-active pattern resumed. In later years the terrarium was moved to another room with temperatures typically around 15°C in the colder season. In the warmer months, approximately May-August, temperatures outside rarely reach 30°C (usually 20-25°C in daytime). Indoors the low 20s are common in that period.

DEVELOPMENT

Shortly after returning to The Netherlands the larvae were measured: head-body lengths 24-27 mm, tails 18-20 mm. Already in the brook they showed only a slight yellowish coloration at the base of the legs (sometimes even absent). For older larvae see figure 5. Four specimens metamorphosed on July 17, 19, 26 and 28, 1996 and were then head-body 27-30 mm long with tails 23-27 mm, weights 0.93-1.08 g. Typically a few days before transforming into land-living salamanders they stopped feeding. The last larva did so November 9 and came on land November 14. That one measured 35+ 30 mm (head-body+tail) and weighed significantly more than the others at 2.24 g.

Their bellies were at first a dirty grey like in the older larvae (see fig. 7) but soon turned into dark grey. Before long it blackened as did the rest of the body. The dorsal and lateral yellow spots were at first tinged with a slightly orangey yellow ( fig. 6). Dorsally the spots were irregular, laterally much more rounded. Over the years the figuration of the spots barely changed. In contrast to the older S. s. salamandra larvae I know from localities in the southeastern part of the Netherlands, the Lebanese ones in this initial batch hardly showed the yellow spots near the base of the legs. Ventrally in just one metamorphosed specimen two tiny yellow specks were found posteriorly ( fig. 8), thus suggesting the scientific nomenclature is not unfounded. Commonly food was accepted after only a few days.

Sizes of initial metamorphosed S. infraimmaculata are given in table 1 . For further development see tables 2-4 .

In September 2001 one animal died and was donated to the ZFMK in Bonn (ZFMK 75922).

BEHAVIOUR

It was noteworthy to find that their initial walking was kind of leap-wise; a quick lateral swaying with a forward component, resulting in a at times very fast propulsion, at first even resembling hopping. This was no longer observed as they grew older. Also, somewhat unexpectedly, recently metamorphosed animals would lacertid-like run from a stone just above the waterline into the water when disturbed. I noticed shedded skins in the water a few (2-5) days after metamorphosis. This skin-shedding is commonly seen in the adults several times per year. It looks like that water is essential to the moulting process; these sheds were not seen on land, although the very thin and transparent cast-offs could be easily missed there.

In the warmer months the animals are mainly active at night, although during periods of very hot days, reaching almost 30°C, they seemed to enter brumation, which may last to October when feeding starts again. ANNUAL RHYTHM

Apart from the daily rhythm, mainly from dusk to dawn, an annual cycle is present and, in vivarium keeping, traceable because of droppings in the pool water and mealworms anew disappearing from a feeding bowl. Eating in captivity decreases around the middle of April and resumes around October. From the beginning of September-October renewed nightly activity can be deduced from finding some peat-soil in the water basin and the afresh disappearance of mealworms from the bowl. A drop of food-interest may occur in November. Commonly, mealworms in the feeding nap do not vanish anymore.

In May, under Dutch circumstances, the animals no longer appear to be active. When searching their terrarium in e.g. July, I found them under stones, fairly often several, up to four, together. In our wintertime, roughly from November to February when freezing temperatures are possible outside, the S. infraimmaculata become more active in January; the salamanders seem to sense the change in season even when indoors. For the Middle East, the above indicated periodicity is logical as reproduction seems to start around the beginning of the year (e.g. data of WARBURG, 2009 for Israel).

REPRODUCTION

The wild-caught larvae had apparently reached adulthood in 2003 as on March 9 in 2003 around 10 o'clock I found the first twenty larvae and one unfertilised whitish egg (diameter ± 8 mm) in the water. It had been a very cold and long winter; only now the ice on the ditches and pools started melting. None of the larvae (total lengths 22-29 mm) appeared viable, being rather skinny. Eight of the larvae were already dead or dying and showed a crooked tail ( fig. 9). One very transparent egg seemed on the face of it to contain a well-developed larva that soon wriggled free. According to the literature, a batch of 20- 40 eggs is common and is sometimes laid over a longer period, so I kept hoping for more. It turned out to be in vain. The few invertebrates collected in the still freezing waters outside could not tempt the larvae to feed. One after the other died, the last one on March 15. I did find another two eggs somewhat later, with non-moving embryos. After having been away for ten days, I found another five dead larvae in the water, and this pattern continued at intervals until the end of April. Uncertain if this was from one animal or two possible females.

Hoping that reproduction would be more successful elsewhere, I gave Sergé Bogaerts, who had ample vivarium experience with the genus, a couple in October 2005. He was surprised how much they had grown. Sergé later handed offspring to Max Sparreboom who also reported reproductive success. This aspect will be dealt with soon by BOGAERTS et al. (in prep.).