Introduction

The southeastern Mediterranean region is a crossroad of the Mediterranean, Irano-Turanian and East Sudanian savanna ecozones (ZOHARY, 1973; DANIN et al., 1975; POR, 1975; YOM-TOV & TCHERNOV, 1975; DANIN, 1992). The Jordan Valley, the northern part of the Great Rift Valley, which stretches along the coast of the Mediterranean Sea, is fringed by mountainous ridges from the west and the east, reaching their highest elevation of about 3,000 m on the Lebanon and Anti-Lebanon mountain ridges (Fig. 1a). The windward orientation of the ridges results in higher rainfall on the coastal plain thus providing a corridor for the Mediterranean biome to penetrate far south into the Arabian Peninsula. The Anti-Lebanon ridge borders both the Mediterranean and Irano-Turanian ecozones, while Sudanian faunistic and floristic elements penetrate the area through the Rift Valley (Fig. 1b).

The southernmost part of the Anti-Lebanon ridge (hereafter referred to as the Hermon) is situated in the northernmost part of Israel. Together with its foothills, it occupies less than 1% of the territory of the country, yet it possesses an unbelievable species richness in both flora and fauna of vertebrates and invertebrates, with high percentage of the species that occur within the borders of Israel being present solely or almost solely in the Hermon. The list for some taxons of animals and plants is given in Table 1. The highest percentage of unique species is recorded in Lepidoptera both for Rhopalocera - 32.7% (35: 107) and for Noctuidae - 19.3% (106: 548).

The vertical distribution of plants on the Hermon has been covered in detail (AUERBACH & SHMIDA, 1992; COHEN et al., 1981). In contrast, information on most insect groups is at best rudimentary. In this paper, we analyze the species composition, vertical distribution and chorotypical characteristics of the Noctuidae, the most abundant and species-rich family of the Lepidoptera worldwide as well as in the Mediterranean basin and on the Hermon (CHOI & MILLER, 2013).

Material and methods

Abbreviations used:

SMNH-TAU: Steinhardt Museum of Natural History, Tel Aviv University a.s.l.: Above sea level.

In seven years between 2002 and 2014 we operated six to 18 automatic light-traps, equipped with black-light ultra violet tubes (TL-D 18W/80), 220 V, in order to sample Noctuidae in the six main biomes identified on Mount Hermon (1-3 traps in each biome) along the main access road from Tel Dan National Park at 177 m a.s.l. (3314’37.03”N; 3538’45.85”E) to the Israel Cosmic Ray Center at 2050 m a.s.l. (3318’27.79”N; 3547’07.45”E). Each trap worked constantly for at least one complete year. A detailed description of the Israeli light trap network is given elsewhere (MÜLLER, KRAVCHENKO & SCHLEIN, 2005a; MÜLLER & KRAVCHENKO, 2005b; MÜLLER, 2006).

The distribution of the sampled biomes corresponds largely to the altitudinal zonation of the Hermon (AUERBACH & SHMIDA, 1992; COHEN et al., 1981; SHMIDA, 1977). It is comprised of four zones: 1) foothills with riverine forests and lush grassland (< 300 m a.s.l.); 2) evergreen Mediterranean maquis (300-1,300 m a.s.l.); 3) xeromontane open scrub with montane forest and montane steppe (1,300-1,900 m a.s.l.) and 4) alpine tragacanth vegetation (> 1,900 m a.s.l.).

The nomenclature, systematics, general distribution pattern, affiliation to chorotype, phenology and host plants of the Israeli noctuid species follow those used in previous studies (KRAVCHENKO et al., 2007a; KRAVCHENKO et al., 2007b). The collected voucher material is deposited in the Steinhardt Museum of Natural History, Tel Aviv University (SMNH-TAU).

We used Simpson’s similarity index to compare species composition and Fisher’s exact test was applied to examine the differences between the ratio of Mediterranean and Irano-Turanian species (MAGURRAN, 2004). Significance was taken at P < 0.05.

Results

SPECIES RICHNESS AND SPECIES COMPOSITION

From the collected material, 291 species in 125 genera were identified. Apart from the riverine forest where only 52 species were found, the number of species in each biome varied within a rather narrow range: 100 species were found in lush grassland, 121 in maquis, 109 in montane forest, 78 in montane steppe and 111 in the alpine tragacanth zone.

Only the following 18 polyphagous and migratory species with wide distribution ranges were found throughout the entire area: Agrotisipsilon (Hufn.), A. segetum (D. & Schiff.), Anarta trifolii (Hufn.), Cornutiplusia circumflexa(L.), Eublemma ostrina(Hb.), E. parva (Hb.), Helicoverpa armigera (Hb.), Heliothis peltigera ([D. & Schiff.]), Hypena lividalis(Hb), Leucanialoreyi (Dup.), Mythimnavitellina (Hb.), M. unipuncta (Hw.), Noctua pronuba (L.), Peridroma saucia (Hb.), Spodoptera cilium (Gn.), S. exigua (Hb.), S. littoralis (B.), Trichoplusiani (Hb.).

Other species showed a high level of discreteness in distribution. The majority were found only in one (128 species) or two (101 species) biomes. Preference for a certain biome is also pronounced at the genus level. All 14 species of the genus Dichagyris, all seven species of the genus Chersotis, 10 of the 12 species of the genus Hadena, and four of the five species of the genus Episema inhabit only the montane grassland and the tragacanth biomes of high altitudes, while species of the genera Acronicta and Lithophane are restricted to the riverine forest.

Excluding the 18 widely-distributed species, the similarity between the neighboring biomes, measured using Simpson’s index, was > 0.5 (greater than 50% of species common to both biomes). In all other cases, the index was much below this threshold. (Table 2).

CHOROTYPICAL DISTRIBUTION OF SPECIES

The majority (48%) of collected Noctuidae belong to the Mediterranean chorotype, followed by the Irano-Turanian (34%), Euro-Siberian (7%), Palaeotropic (6%) and Palaearctic (5%) types of distribution.

The Mediterranean species occur ubiquitously and comprise more than 50% of the total Noctuidae species in the maquis and montane forest, while in the montane grassland and the tragacanth they comprise only 27% (Fig. 2). In contrast to the Mediterranean species, the Irano- Turanian species do not exceed 6% of the total Noctuidae species below 1,900 m. However, in the montane steppes and tragacanth biomes they make up more than one-third of the species. About one- third of the discussed species in each of the biomes are polyphagous, ubiquitous, Palaearctic and Palaeotropic elements.

The highest numbers of Euro-Siberian species were collected in the riparian forest (eight species) and montane forest (six species), while the only two eremic species were found in maquis and lush grasslands: Autophila cerealis (Stgr.) and Condica viscosa (Frr.).

Twenty-one species are endemic and sub-endemic to the Levant. Five taxa were recently described from the Hermon and are known so far only from the type locality.

SPECIES RESTRICTED WITHIN ISRAEL TO THE HERMON

Although most of the species collected are widely distributed throughout the Mediterranean region of Israel, 106 of them were found only on the Hermon. These species occur only at medium and higher elevations with the highest species diversity in the tragacanth (20 species in the montane forest, 41 in the montane steppe, and 45 in the tragacanth) Fig. 3. For most of these taxa, the Hermon is the southernmost distribution border. Additionally, 11 Levantine endemics, sub-endemics, and five true Hermon endemics are found here.

The species were subdivided into those that are oromontane-inhabiting mountainous biomes throughout the entire area of their distribution (Table 3a) and non-oromontane which also occur in lowlands (Table 3b). The typical oromontane species are further subdivided (VARGA, 2003) into two major types: the European alpine species which inhabit the mesophilic mountains of Europe, and the xeromontane species which inhabit the dry mountains of Iran and Central Asia.

Of the 13 oromontane species inhabiting the steppe of the Hermon, nine are alpine and belong to the Mediterranean chorotype, and four are xeromontane and belong to the Irano-Turanian chorotype. The tragacanth biome is comprised of 43 oromontane species. They include 33 Irano-Turanian xeromontane species and ten alpine species that have a Mediterranean-type distribution.

In the montane forest, mainly arboreal Mediterranean (13 species) and Euro-Siberian (three species) are found at the southernmost border of their distribution (Table 2), only two species (A. pasiphae and O. gracilis) are Irano-Turanian. These species are predominantly monophagous on oaks (deciduous Quercus boissieriand .. libani and evergreen .. calliprinos) and Rosaceae (Cotoneaster nummularia, Crataegus monogyna and Prunusursina). The species assemblage of the montane steppe is more diverse, with 19 Mediterranean, 11 Palearctic, 10 Irano-Turanian and one European species.

Some of the Palaearctic and Euro-Siberian species (H. compta, H. munitalis, L. lusoria, M. confusa and A. anceps) that would be widely distributed in the center of their lowland ecozones are, on the Hermon, at their southern limit of distribution and are restricted to higher altitudes with temperate climate. M. confusa and A. ancepsare common agricultural pests in Europe, but at their limit of distribution on the Hermon they become rare.

There is also evidence that some of the species collected in this study were once more widely distributed. In historic collections of the SMNH-TAU, we found E. korsakovi, P. treitschkei, E. aquilina, E. robiginosa and E. cos specimens all collected from high elevations in the Judean desert at the beginning of the 20th century. D. melanura and D. singularis were even found in the mountains of the central Negev. Despite various studies, neither of these species have been recorded there recently (unpublished data of the authors).

A COMPARISON OF OROFAUNA OF THE LEBANON AND ANTI-LEBANON RIDGES

A series of publications on the noctuid fauna of Lebanon (CZERNY, 1932; ELLISON & WILTSHIRE, 1939; WILTSHIRE, 1940; TALHOUK, 1997) and our own data on Noctuidae of the Hermon, enable a comparison of the orofauna composition of the Lebanon and Anti-Lebanon ridges. Overall, 60 true montane species are found on the Hermon, 31 of which are also recorded from the Lebanon ridge (Table 3a). These species belong either to the Irano-Turanian or to Mediterranean chorotypes, but the ratio between the two chorotypes differs on each ridge. The Irano-Turanian species clearly dominate on the Hermon (41 Irano-Turanian species: 20 Mediterranean species), while on the Lebanon ridge the number of species of these two chorotypes is nearly equal (16 Irano-Turanian species: 19 Mediterranean species). These ratios differ significantly (Fisher’s exact test; P > 0.05).

Differences in the ratio of alpine and xeromontane species are caused by the lower number of xeromontane species on the Lebanon ridge, while the numbers of alpine species on both ridges are nearly equal. A total of 25 xeromontane species common on the Hermon are not found on the Lebanon ridge (Table 3a).

Discussion

All 106 species of Noctuidae restricted to the Hermon are found at the higher altitudes. Apart from 17 endemics, this group consists of species specific to mountain biomes of Europe and the Middle East. Some of the lowland Palaearctic and Euro-Siberian species exist at the southern limit of their distribution and only at high altitudes. The area most similar to the Hermon, with comparably high elevations and similar plant communities of thorn-cushion and dwarf-shrubs is found at least 600 km away in the Taurus Mountains of southern Turkey (PAROLLY, 2004; LATIF et al., 2006).

The Lebanon and Anti-Lebanon ridges, both reaching almost 3,000 m a.s.l., comprise the only location between southern Turkey and the Sinai Peninsula high enough to accommodate montane and alpine biomes. The connection between these biomes, and the similar xeromontane biomes of the Taurus Mountains to the north, was interrupted probably five to six thousand years ago, during the aridification of the Middle East (THOMPSON, 2000; VERHEYDEN et al., 2008). South of the Hermon, the closest places with similarly high altitudes, the Azir Mountains in Saudi Arabia and Santa Katherine Mountain on the Sinai Peninsula) support neither Mediterranean montane forest nor the Mediterranean tragacanth biomes on their higher altitudes (NATAN & WERNER, 1999). Therefore, for the Euro-Siberian, Mediterranean arboreal, and European oro-alpine species, the Lebanon and Anti- Lebanon ridges are the southernmost edge of their distribution. However, according to historical data, some species were found far southward along steppe tops of mountains, at least in the recent past (historical data collections in Tel Aviv).

The Lebanon ridge receives about 2,500 mm of annual precipitation, in contrast to the Anti- Lebanon ridge, which receives about one-fifth of that on its western slope (available on line at http: see in references). The eastern slope of the Anti-Lebanon ridge faces the Syrian Desert and as a result, the top of the ridge receives only 100-250 mm of annual precipitation resulting in xeromorphic vegetation (COHEN et al., 1981; SHMIDA, 1977). The highest part of the Anti-Lebanon ridge thus functions as the eastern limit for the Irano-Turanian xeromontane species.

Conclusions

Many invertebrate, Arthropoda, and plant species changed distribution depending on elevation in similar patterns, thus the cold climate resulted in an “elevational species richness pattern” (ROMDAL & GRYTNES, 2007). The pattern suggests that the highest species richness exists at mid-elevations and declines at higher elevations. This has been shown for different insects such as the Noctuidae in Azerbaijan (ALIEV, 1984), or macro-moths in Oregon, USA and South Korea (CHOI & AN, 2010). On the Hermon, this “hump-shaped” distribution was demonstrated for reptiles, birds (NATAN, WERNER, 1999), and vascular plants (WILSON & SHMIDA, 1984; LEVIN et al., 2007). However, the Noctuidae of the Hermon show no decline in the number of species toward the top. This is likely because this location constitutes a meeting point of the rich European alpine orofauna and xeromontane fauna, related to those of Iran and Central Asia.

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