Lakes Hula and Agmon: destruction and creation of wetland ecosystems in northern Israel

A portion of the former Lake Hula wetland (northern Israel) was re-flooded in spring 1994 and the physical, chemical and biological developments within the resulting new lake and wetland complex (Agmon) was followed closely by a multi-disciplinary scientific team. The first three years of study relating to Lake Agmon are reported in this issue of Wetlands Ecology & Management. We provide in this paper a general background on the Lake Hula Draining Project in the 1950s and the recent re-flooding and creation of the Agmon wetland.     

Repopulation and colonization by birds in the Agmon Wetland,Israel

Lake Hula and its surrounding wetland in northern Israel were drained in the late 1950s and the dried wetlands were transformed into a diverse agricultural region with a 0.3 km2 nature reserve. A portion of the extinct Hula wetland was re-flooded in April 1994 by constructing a small lake, Agmon, and 90 km of canals. The purposes of this study were (a) to document the re-population and colonization of the new Agmon wetland by birds after its flooding, (b) to evaluate bird species richness in this new wetland in comparison to that in a nearby mature Hula Nature Reserve, and, (c) to investigate if the species originally present in the Hula Valley before the drainage had been re-established. The new lake has succeeded in attracting a large variety of water birds to the newly formed habitats, especially ducks, egrets and herons, plovers, waders and snipes. The distribution of bird species among the different habitats was not random. The northern section of the lake, which is shallow and has an open shore, had the largest number of species during 1995 and 1996 and was preferred by plovers, waders and snipes. A large roosting and breeding colony of thousands of egrets and herons was formed in the dense cattails ( Typha domingensis) in the southern section of the lake. Unlike the nearby Hula Nature Reserve, the new wetland lacks a large seasonally flooded area and is less attractive for feeding and breeding plovers, waders and stilts. The new wetland also has very little muddy area without vegetation and attracts few of the species that regularly feed in this habitat in the reserve. The new wetland enlarges the area of only a few of the habitats found in the nearby reserve and therefore attracts fewer species than it might otherwise. Recommendations for management of the new wetland include increased diversity of habitats, restriction of human activities and optimization of conflicts that have arisen between the needs of the wetland and those of nearby agriculture.     

Phytoplankton-metaphyton seasonal dynamics in a newly-created subtropical wetland lake

The dynamics of the algal populations of Lake Agmon, a newly created shallow lake in the Hula Valley, Israel, were monitored following its filling in April 1994 through 1996. Additional limited field observations and measurements were taken throughout 1997. Following an initial establishment period, the dynamics of the algal populations showed a repetitive annual pattern comprised of three phases: I. a clear water phase in January–February, with low phytoplankton biomass and no metaphyton; II. a metaphyton dominance phase during March–June when mats of filamentous chlorophytes covered most of the lake's sediments while phytoplankton biomass remained low; and III. an intense phytoplankton bloom phase from June till December. The shifts from phase I to II and from phase II to III were gradual, resulting from interplay between phosphorus availability, the underwater light climate, temperature effects and zooplankton grazing pressure. The shift from phase III back to phase I was abrupt, due to winter flushing of Lake Agmon. The summer phytoplankton blooms intensified from 1994 to 1996 and shifted from chlorophyte dominance in 1994 and 1995 to cyanobacteria-dominance in 1996 and 1997. These observations, jointly with the nutrient chemistry of Lake Agmon, suggest intense eutrophication. Criteria based on phytoplankton taxonomy also indicate that Lake Agmon is eutrophic to hypertrophic. Due to the typical unstable nature of hypertrophic systems, careful management is essential to maintain the delicate ecological balance needed to ensure that the lake will fulfill its intended role as a center for eco-tourism.     

Biogeochemical evolution of a sulfur-iron rich aquatic system in a reflooded wetland environment (Lake Agmon,northern Israel)

Major biogeochemical processes in the newly created, shallow Lake Agmon (Hula Valley, northern Israel) were investigated from 1994 to 1996. Sediment cores, lake water and porewater were analyzed for chemical composition and spatial distribution. Sediment analyses revealed that Lake Agmon has two different sediment types: peat sediments in the northern and central parts, and marls in the southern part. The basic composition of the lake's water was controlled mainly by the mixing of two distinct water types (Jordan River and water drainage), and by evaporation. About 3/4 of the lake water originated from the Jordan Inlet, a quarter through the Z Canal Inlet (peat drainage) and a minor amount from groundwater seepage. Lake Agmon is unique among freshwater wetlands owing to its high SO ₄ ^2– content, which is ca. 1/3 that of sea water. This characteristic is ascribed to the dissolution of secondary gypsum, formed in the peat soils since the drainage of the historic Hula Marsh. Leaching gypsum from the shallow sediments during the first few months after flooding was followed by a later stage of constant diffusion and advection of SO ₄ ^2– from gypsum dissolution in deeper sediments. Gypsum dissolution in lake sediments contributed ca. half of the SO₄ ^2– and Ca²⁺ inputs to the lake. The concomitant increase of Ca²⁺ combined with alkalinity release due to organic matter decomposition in the sediments led to the precipitation of CaCO₃. This precipitation was enhanced by photosynthesis, particularly during summers, and consumed about a tenth of the Ca²⁺ and third of the alkalinity outputs from the lake. Iron-hydroxide was the main agent for microbial oxidation of organic matter, surpassing by far the role of sulfate, nitrate and manganese as electron acceptors. The produced Fe²⁺ was transported upward by diffusion and advection and oxidized to ferric iron at the sediment-water interface. There is evidence, however, that some sulfate was reduced, but most of the produced sulfide reacted with ferrous iron and accumulated in the sediments as FeS minerals. Therefore, despite high sulfate concentrations, the high iron availability restricted release of toxic sulfides into the water and thereby served to maintain reasonable water quality.     

Development of macrophytic vegetation in the Agmon Wetland of Israel by spontaneous colonization and reintroduction

The draining of the Lake Hula and swamps, northern Israel, during the late 1950s resulted in the loss of a very diverse and rare ecosystem. Oxidation of the peat soil resulted in ground surface subsidence, while heavy autumn winds have eroded the dry peat. Moreover, agriculture on the peat soils is restricted, because of a nitrate surplus. Predictions that the sinking would continue and that more areas would go out of agricultural production led authorities to re-flood a portion of the Hula Valley in 1994. The aim of the present study was to monitor the spontaneous establishment of vegetation in the re-flooded area, the Agmon wetlands, and to reestablish some of the major plant species lost from the valley when Lake Hula was drained. Within the first two years, 74 plant species colonized the wetland spontaneously. Five out of 11 species designated for reintroduction were successfully established. Cyperus papyrus and Cynodon dactylon demonstrated sustainable potential for lake-shore stabilization. Cyperus papyrus was reintroduced from seedlings and rapidly became the dominant riparian species, while Cynodon dactylon established spontaneously. Re-introduced Nymphaea alba clones were established only in enclosures protected from grazing by the semi-aquatic mammal Myocastor coypu. Nuphar lutea and Iris pseudacorus showed better resistance to grazing. These results demonstrate a high potential for successful re-establishment of much of the original Hula swamp macrophytic vegetation by either spontaneous colonization by extant species from the surrounding areas or by introduction of locally extinct species. As such, there is a good chance that the associated faunal components of the former Lake Hula and swamps that have returned to the region since the Hula rehabilitation project commenced will continue to flourish.     

Nutrient and plant dynamics in Lake Agmon Wetlands (Hula Valley,Israel): a review with emphasis on Typha domingensis (1994–1999)

Shallow lake Agmon is a newly created subtropical wetland in north-eastern Israel. The lake is part of the Hula Project aimed at slowing down deterioration processes of the peat soils, to establish infrastructure for ecotourism as an income for the land owners, and nutrient removal from Lake Kinneret inputs. An onset of benthic filamentous macro-green algae during late winter–spring season, followed by submerged macrophytes vegetation during spring–summer was documented. The phosphorus summer loads are mostly plant–mediated internal fluxes and nitrogen intensively removed from lake waters by sedimentation and denitrification. The summer phytoplankton, mostly colonial cyanobacteria, are P limited. During 1995 and early 1996, dense Typha domingensisstands were developed in the southern half of the Lake (chalk-marl bottom sediments). The P-limited Typhavegetation collapsed within less than half a year and reappeared in the south-eastern part of the lake where sediments were exposed and oxidized. It is hypothesized that phosphorus cycle is a strong dependant of macrophyte mediation, and P deficiency in the sediments predominantly affected Typhadecline and an increase of P availability later enabled the reappearance of the Typhastands.     

Temporal variation in fish community structure in a newly created wetland lake (Lake Agmon) in Israel

Lake Agmon was constructed in summer 1994 in the northern part of the drained Lake Hula swamp, as part of the Hula Reclamation Project. One of the main goals of the project was to attract aquatic and semi- aquatic birds to the area as a basis for planned eco-tourism and one of the main concerns of the project was the potential for increased proliferation of mosquitoes (including vector species of malarial parasites). There was a perceived need, therefore, to establish and maintain a suitable fish community. This present study documents the developments in the fish community in Lake Agmon between May 1995 and December 1996. The lake was initially stocked with two fish species, Tilapia zillii and Gambusia affinis, to control mosquitoes and to provide suitable small- bodied forage species for piscivorous birds. Subsequent stocking also included Oreochromis aureus, and Cyprinus carpio. Additional species were established by immigration from the Jordan River. Relatively large species ( e.g., Clarias gariepinus and C. carpio) flowed into the lake as eggs or juveniles, whereas the smaller species ( e.g., Hemigrammocapoeta nana, Pseudophoxinus kervillei and Aphanius mento) entered the lake as adults. In the first year, eight fish species were found in Lake Agmon: T. zillii, G. affinis, C. gariepinus, C. carpio, Capoeta damascina, P. kervillei, Acanthobrama lissneri and H. nana. Of these, T. zillii, G. affinis and H. nana were dominant, though the relative abundance of H. nana was very low in spring, and increased during summer. In the second year, O. aureus, A. mento and Anguilla anguilla also occurred in the lake. The density of P. kervillei increased, in comparison with the previous year. The relative proportions of juvenile and adult fish also changed in the second year. A high percentage of young T. zillii was found during the first spring, decreasing in the summer and autumn. The proportion of juveniles of both G. affinnis and H. nana was very low during the first year, but increased during the second. These results indicate that the fish community of Lake Agmon had not stabilized by the end of 1996 and that further changes in relative abundances, and species, age and sex compositions should be expected.     

Climatic and hydrological aspects of the Hula restoration project

Drying the Hula marshes (northern Israel) in the 1950s caused changes in sub-surface hydrology that severely compromised agricultural sustainability of the Hula Valley peatlands. The complex dynamics between climatic and hydrological regimes produced widely fluctuating elevations in the water table, leading to deterioration of the peat soils through enhanced aerobic decomposition, wind erosion, and underground fires. The year-round intensive and diverse cropping systems used in 1960s were gradually replaced in the 1980s by extensive winter forage crops, leaving the peat uncropped and exposed to increased deterioration in the dry season. The Hula Restoration Project was initiated in 1993 to moderate some of these after-effects of the draining of the Hula marshes. The project includes a 90-km network of water-level regulating supply and drainage canals and a 100-ha lake and a barrier to underground water flow from the lake to cultivated areas. We followed changes in underground water levels for the five years before and the three years after re-flooding of the valley in May 1994. Prior to re- flooding, water tables in the area fluctuated 2–4 m annually, depending on location and microrelief and there was a strong north to south flow of water. After re-flooding the lake initially had a strong influence on water table elevation in the adjacent areas, which later stabilized. The underground barrier shifted the predominant water flow westward toward a north-south canal near the edge of the valley and then southward along the canal. The restoration project has been successful up to now, there are few underground fires since re-flooding, and all agricultural lands can be cultivated al year long.     

Using forage crops to reduce nitrate accumulation in hula peat soils

Summary A two-year field experiment was made of growing Rhodes grass and alfalfa on peat soil of the Hula Valley, where high nitrate accumulation occurs under row crops or plantless soil and leads to the danger of contamination of Lake Kinneret. The aime of the investigation was to use these perennial high-nitrogen-consumer crops to prevent the nitrate accumulation in the peat soil. The results showed a drastic decrease in soil N-NO₃ under both species in two irrigation systems obtaining high dry matter yields with high plant nitrogen content without the use of any nitrogen fertilizer. Contribution from The Volcani Center, Agricultural Research Organization, Bet Dagan, Israel. 1977 series, no.263-E.     

Foraging,roosting, and nesting habitats of the avian fauna of the Agmon wetland,northern Israel

The foraging, nesting and roosting habitats of the avian fauna of a newly created Agmon wetland and surrounding cultivated peat land (5 km2) in the Hula Valley, northern Israel, were evaluated (January 1996–February 1997) to assess the value as a habitat and for wildlife tourism. We recorded 180 bird species (herons, dabbling ducks, kingfishers, waders, wagtails and raptors) in different habitats (the lake, shores, cattail and reed-bed stands, trees, temporary inundated areas). The most heavily used habitat for foraging, breeding, and roosting was a large cattail stand in the southern third of the lake. The foraging habitat and diet data of 97 avian species were determined. The most intensively used foraging habitats were cultivated fields, lake shore reed-beds, shallow canals and the cattail stand. Forty six species nested in the wetland during March-October, with 2,040 colonial and solitary nests mostly in the cattail stand, near-shore reed-beds and young trees. Roosts, also mostly in the cattail and reed-bed stands and in trees, were used by 55 species. Based on this survey of available habitats, we identified several key habitats that are either missing or require further development (e.g., temporarily inundated mud flats, reed-beds and bare islands). We also provide suggestions for improving the delicate balance between requirements of this developing wetland ecosystem and of the adjacent agricultural areas.     

Abstracts

Lake Agmon was created in the Hula Valley in Israel, in the northern part of the dried-up Hula Lake site, in summer, 1994. Samples of fish at ten stations were taken by electroshocker once a month, from January to December, 1996. The fish species whose food composition in the gut was examined in this study were: Tilapia zillii, Gambusia affinis, Clarias gariepinus, Cyprinus carpio, Pseudophoxinus kervillei, Acanthobrama lissneri, Hemigrammocapoeta nana and Oreochromis aureus. Food components were determined qualitatively and quantitatively. Assessment of the organism biomass in the gut contents was based on available data of phytoplankton and zooplankton. The food of G. affinis changed during the year, consisting high level of algae in winter and spring, and insects and crustaceans in summer. In T. zillii, a high percentage of algae were found in winter, and a high percentage of plants in summer. The food of H. nana consisted mainly of phytoplankton, but included high plants during most months; P. kervillei ingested mainly insects and crustaceans, with a high percentage of Daphnia spp.; and the diet of C. gariepinus comprised fish and invertebrates species, which varied with the seasons. C. carpio ate mainly invertebrates; O. aureus are vegetarian, and A. lissneri fed on both animals and plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

Present–absent: a chronicle of the dinoflagellate Peridinium gatunense from Lake Kinneret

A long-term record dating back to the 1960s indicates that Peridinium gatunense, an armored dinoflagellate, dominated the phytoplankton of Lake Kinneret (Sea of Galilee, Israel) until the mid-1990s, with a relatively stable spring bloom. However, since 1996, these blooms became irregular, failing to develop in 10 out of the past 16 years. During the later period, a significant correlation (R ² = 0.605, P = 0.013) was found between annual peak P. gatunense biomass and riverine inflow volume. In-lake surveys showed that patches of high P. gatunense densities were associated with water enriched with fresher inflowing Jordan River water. Supplementing laboratory cultures of P. gatunense with Hula Valley water stimulated its growth relative to un-enriched controls. A likely explanation to the recent irregular blooms of this dinoflagellate is a hydrological modification that was made in the catchment in the mid-1990s, preventing Hula Valley water from reaching Lake Kinneret in most years—except for exceptionally wet years. We propose that until the mid-1990s, the Jordan River water enriched Lake Kinneret with a growth factor (a microelement and/or organic compound) originating in the Hula Valley, which in recent years has arrived in sufficient quantities to support a bloom only in high-rainfall years.     

Seasonal Rotifer Dynamics in the Long-term (1969–2002) Record from Lake Kinneret (Israel)

Long-term (1969–2002) data record of biomass distribution of rotifers in Lake Kinneret is combined with previously published information on their metabolic activity and newly calculated population dynamics parameters to synthesize a model of their seasonal dynamics in Lake Kinneret. Nineteen rotifer species were recorded in routine samples collected in Lake Kinneret (Israel) in 7 offshore (deeper than 5 m), stations, at 12 discrete depths during 1969–2002. Organisms were sorted and counted (including external egg carrying females), biomass was measured and calculated for the entire lake stock (g_w.w m⁻²; mg l⁻¹). Rates of grazing, respiration and production were measured experimentally at three different temperature ranges. Results were extrapolated to the lake community for months with similar temperatures. Rotifera comprised 7% of total zooplankton biomass in Lake Kinneret whilst Cladocera and Copepoda 58 and 35% respectively. Rotifers were found to be more abundant during December–June and decline in summer months. Monthly (1969–2001) means indicated total grazing capacity of rotifers as 11%, respiration as 9% and production as 3.7% of the total zooplankton metabolic activity. Positive relations were indicated between rotifer and small bodied cladoceran numerical concentrations. Population growth models suggest that rotifers are not food limited in Lake Kinneret but that fish predation plays an important role in regulating abundance in spring-summer and fall.     

The impact of zooplankton status on the management of Lake Kinneret (Israel)

Monthly averages of standing stock wet biomass of zooplankton in Lake Kinneret (Israel) varied between 11 and 76 g m^–2 during 1969–1981, with the exception of two months. Averaged contributions of different groups were: Cladocera 58%, Copepoda 35% and Rotifera 7%. Total standing crop wet biomass is highest during January–June, averages varied between 35 and 50 g m^–2, and decreases during summer–fall (23–36 g m^–2). The winter biomass of Cladocera fluctuated between 22 and 35 g m^–2 and dropped to a range of 9–23 g m^–2 in summer, whereas copepod biomass varied very little around an average of 18 g (ww) m^–2 with the exception of low values from April to June. The stock biomass of Rotifera is relatively high during winter floods season (December-March) whilst in summer it is very low.Young stages of fish in Lake Kinneret feed mostly on zooplankton and zoobenthic forms. The most abundant fish in the Kinneret ecosystem, Mirogrex terraesanctae terraesanctae, also feed on zooplankton at the adult stage throughout the year, and herbivorous fish consume zooplankton during the summer when lake plankton resources are limited.The summer ecosystem of Lake Kinneret is characterised as a steady state type, in which the impact of the zooplankton-chain is of great importance. Increase of predation pressure on zooplankton by fish can disequilibrate the balanced trophic relations existing between nannoplankton production and zooplankton grazing capacity. Such a situation can lead to organics accumulation as nannoplankton blooms, resulting in water quality deterioration. Management options aimed at preventing collapse of zooplankton populations are discussed.     

Aphanizomenon ovalisporum (Forti) in Lake Kinneret, Israel

The filamentous cyanobacterium Aphanizomenon ovalisporum wasobserved for the first time in Lake Kinneret in August 1994and formed a prominent bloom from September through October.Aphanizomenon ovalisporum reappeared in diminished amounts inthe summer and fall of 1995. These events are the first recordof significant quantities of a potentially toxic nitrogen-fixingcyanobacterium in this lake. No definite provenance of inoculumhas been identified, although A.ovalisporum was also observedin a newly reflooded area (Lake Agmon) in the catchment. Unusuallyhigh water temperatures and low wind inputs were observed priorto and during the A.ovalisporum bloom period. These, togetherwith possibly enhanced availability of phosphorus or other growthfactors, may have contributed to the cyanobacterium growth in1994. Phosphorus limi tation, as indicated by high cellularalkaline phosphatase activity, the onset of stormy conditionsand a fall in water temperatures led to the demise of the 1994bloom. Although the A. ovalisporum bloom in 1994 had no seriousdirect impact on water quality, the continued presence of apotentially toxic cyanobacterium in Lake Kinneret, a major nationalwater supply source, is a cause for serious concern.     

Appearance and establishment of diazotrophic cyanobacteria in Lake Kinneret,Israel

1. We propose that the appearance and establishment of Nostocales (cyanobacteria) species of the genera Aphanizomenon and Cylindrospermopsis in the warm subtropical Lake Kinneret (Sea of Galilee, Israel) from 1994 was linked to changes in climate conditions and summer nitrogen (N) availability. 2. From 1994 to 2009, an increase in frequency of events of elevated water temperature (>29 °C) in summer, and to some extent a greater frequency of lower summer wind speed events, affected water turbulence and water column stratification, thus providing better physical conditions for the establishment of these populations. 3. In recent years, N‐depleted conditions in Lake Kinneret in early summer have promoted the development and domination of Nostocales that could gain an ecological advantage owing to their N₂‐fixing capability. 4. Nitrogen fixation rates coincided both with heterocyst abundance and with Nostocales biomass. The N supplied to the lake via nitrogen fixation ranged from negligible quantities when Nostocales represented only a minor component of the phytoplankton community to 123 tonnes when Cylindrospermopsis bloomed in 2005. This high N₂ fixation rate equals the average summer dissolved inorganic nitrogen load to the lake via the Jordan River.     

Holocene vegetation history of the southern Levant based on a pollen record from Lake Kinneret (Sea of Galilee), Israel

Lake Kinneret, also known as the Sea of Galilee and Lake Tiberias, is located in the northeast of Israel. At a lake level of 211 m b.s.l. (below mean sea level), the central basin is 43 m deep. The maximum length of the lake is 21 km (N–S) and its maximum width is 12 km (W–E). Lake Kinneret’s surface area is 166 km². A new 17.8 m long sedimentary core was drilled in 2010. Here, we present the entire palynological record from it, which covers the last?~?9,000 years. Special emphasis is given to the natural and human-influenced vegetation history of Galilee in comparison to that of the more southerly Dead Sea region. Significant signs of human impact are the first Olea (olive) increase during the beginning of the Chalcolithic period between 7,000 and 6,500 years ago, as well as the prominent Olea phase during the Hellenistic/Roman/Byzantine period between 2,300 and 1,500 years ago. Mediterranean macchia and bathas scrub vegetation, as known in the area today, has developed in the southern Levant under human impact since the last ca. 1,500 years.     

人为因素导致的盐度变化对Kinneret湖水细菌种群的影响

由于Ｋｉｎｎｅｒｅｔ湖周围的诸多盐泉被人为地导入其下游的约旦河,使得Ｋｉｎｎｅｒｅｔ湖水的盐度逐渐下降,这对作为主要工家业用水和饮用水资源的Ｋｉｎｎｅｒｅｔ湖来说是非常有价值的。然而,对湖水中的细菌来说则可能产生不同的影响。作者通过实验研究发有水盐度的变化导致细菌种群的改变,从而使我们观察到盐度与细菌之间的关系。这些结果表明,盐度变化对湖水水质保护对策和将来一段时间内湖水管理政策的制定都具有重要的     

Cyclopoid diversity in the basin of Lake Hula (Israel), after its partial reflooding

In a relatively small area of 60 km² which corresponds to the basin of the presently drained Lake Hula and its swamps (Upper Jordan Valley), 22 species of Cyclopoida are on record, with two more species reported in the pre-drainage times. The basin of Lake Hula is the southernmost enclave of permanent fresh waters in the Levant Province of Southwest Asia. Lacustrine and swamp environments existed in the area uninterruptedly for at least 700 000 years. A partial restoration project created a new lake. The basin is situated right in the middle of a main bird migration route. The fact that no less than 15 species were recorded from a set of small artificial and isolated experimental ponds seems to indicate that migrating birds and stocks of resting eggs are to account for this high diversity. Cyclops vicinus, Mesocyclops arcanus and Eucyclops serrulatus are the most common species. The success of the restoration is discussed.     

Feeding habits of larval Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret (Israel) II. Experimental study

Interactions between the larvae of Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret, Israel, and their zooplankton prey were studied experimentally. Prey species preference and size selectivities were measured. Larvae were hatched in the lab from eggs collected in the field, and fed different food items in various concentrations. The food items included lake zooplankton, algae, and commercial pellets. It was shown that small, first feeding larvae (7–8.5 mm SL) prefer small bodied zooplankters (< 180 µ). The effect of these food sources on larval growth was measured. It was found that larval Mirogrex grew at a higher rate when fed zooplankton prey sized from 63 µ–250 µ. Food items smaller than 63 µ, larger than 250 µ and Scenedesmus sp., produced less than optimal growth rates. The importance of Mirogrex feeding habits and their potential influence on the Kinneret ecosystem is considered.