Foraging, nesting, and roosting habitats of the avian fauna of the Agmonpdf by Hanna Kuzyo

169

Foraging, nesting, and roosting habitats of the avian fauna of the Agmon wetland, northern Israel S. Ashkenazi1 & Ch. Dimentman2 1 Ecology and Nature Conservation, P.O. Box 1057, Rosh Pinna 12100, Israel; 2 Department of Ecology, Systematic

and Evolution, The Hebrew University of Jerusalem, Jerusalem 91904, Israel Received 15 May 1997; accepted 23 June 1998

Key words: agriculture, breeding colony, cattail stands, diet composition, Hula Valley, management, peatland, re-creation project, restoration

Abstract 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.

Introduction Nearly 5 km2 in the Dead Sea Rift Valley (Afro-Syrian Rift Series), northern Israel (35◦430 E, 33◦ 030 N), was allocated in 1994 to water recreation, wildlife tourism and agriculture. The newly created Agmon wetland is located in the area of the former Lake Hula and swamps which was drained during the 1950s (Dimentman et al., 1992; Hambright and Zohary, 1998). The Agmon wetland consists of a variety of habitats, including a small shallow lake, a system of shallow canals, cattail and reed-beds, stands of trees, pasture land and cultivated fields (Table 1, Figure 1). The combination of the Agmon wetland, the Hula Nature Reserve (HNR), Enan Stream Nature Reserve, the commercial freshwater fish farms, local water reservoirs and cultivated fields all within a relatively small

area of the Hula Valley, creates a mosaic of diverse habitats for birds. The unique assemblage of waterbirds and raptors in this diverse area is very attractive for birdwatching. However, the close proximity of cultivated fields and fish farms requires a sensitive management program that benefits both the attracted bird populations and the local farmers. Suitable habitats for foraging, roosting and breeding determine the integrated value of a given area for supporting a year-round avian community. Each of these functions may require different habitats which might be used independently in different areas. However, the value of an area increases as it includes habitats for more than one function, reducing energy cost for daily movements among dispersed foraging, nesting and roosting habitats. In this study our goal was to assess the integrated importance of habitats available in the newly created wetland for its bird

Article: wetlhula14 Pips nr. 184302 (wetlkap:bio2fam) v.1.1 wethul14.tex; 7/02/1999; 21:01; p.1

170 Table 1. Agmon wetland habitats utilized by avian fauna (see Figure 1). Habitat

Habitat type

Description

1 2 3

Lake Agmon Cattails Typha domingensis Lake shores – riparian vegetation (a) and low grass (b)

4 5

Islands and islets Canals

6 7

Reed-beds of P. australis Bare – mudflats (a) and steep banks (b)

Periodic, mat-like floating vegetation

9 10 11

Temporary inundated areas Diverse pasture land Planted trees

12 13 14

Cultivated fields Roads – paved or dirt-roads Manmade – nest sites

Shallow lake, 0.5–1 m mean depth Permanently inundated in the southern lake Riparian: 12 km with Phragmites australis, Cyperus papyrus, T. domingensis and Scirpus litoralis; low grass: 2.5 km with Gramineae (Cynodon dactylon, Sorghum halepense or sown and harvested Triticum sp.) 2 islands planted (1994) with indigenous trees and 5 islets Transacting the area: < 0.5 m deep and 5–10 m wide – 80 km; boating canal: 1.5 m deep and 20 m wide – 5 km 5 sites, NE – 18.6 ha, NW – 5.8, E – 0.2, W – 5.9, S – 1.4 Temporally created in NW and SE along lake and island shores; banks > 1 m above water surface, 0.2 km Filamentous green algae, Potamogeton spp., Najas delilei and cattail Created after heavy rains and floods Fenced, with 20 reed-fringed shallow ponds, (grazed by donkeys) Large and old – 15 ha (Eucalyptus sp., Populus sp., Salix sp. and Tamarix sp.) and 10 ha of young planted (1994) indigenous trees Various crops Paved roads surrounding the lake (2.8 km), dirt-roads (23 km) Nesting boxes, floating platforms and concrete water passages

populations and to provide recommendations for management and long-term monitoring of indicator and vulnerable species for assessing changes in habitats.

Materials and methods The study was carried out during January 1996– February 1997 in the Agmon wetlands. We classified several habitats that seemed to be important for birds within the study area. Habitat areas were calculated with the aid of GIS from infra-red aerial photograph of the area taken in August 1996. Observations on foraging, nesting and roosting activities of the avian fauna were carried out in the Agmon wetland and adjacent areas (about 350 ha) at least once a week during daytime and once a month during the night. Observation, assisted by binoculars, telescope and a spotlight (at night), lasted 3–10 hours, and covered the entire day-time period within each month. The rest of the cultivated area was surveyed only once a month. We observed birds from a car, an observation tower (5 m high), or while walking. Bird presence and activity was recorded for each habi-

tat. Bird counts were made at the breeding colony, roost sites and in foraging observations. See Shy et al. (1998) for a more complete survey of bird populations in the wetland. Environmental conditions, plant phenology, presence of mammalian predators, seasonal changes in prey abundances and management activities (fish stocking, mowing, harvesting and water level fluctuations) were recorded. Foraging and diet We used a combination of direct observations, faeces, regurgitations and stomach content analysis for recording bird diets. For birds with partial digestion (e.g., European Coot, Moorhen, and some ducks), we collected fresh droppings from individuals exiting the water onto shore and preserved them in 70% ethanol, 4% formaldehyde or 1% Lugol’s solution. Food items were then analyzed under a binocular or compound microscope following sequential filtration with different mesh sizes. Food regurgitations of chicks of colonial waterbirds were collected from nests and their contents identified. Pellets were collected from two pairs of breeding Barn Owl nest boxes, every 4–20

wethul14.tex; 7/02/1999; 21:01; p.2

171

Figure 1. Map of the Agmon wetland showing habitats and main nesting sites for birds during January 1996â&#x20AC;&#x201C;February 1997. Refer to Table 1 for the legend and to Figure 2 for the area of habitats. The main nesting sites are indicated by a circle with an â&#x20AC;&#x153;xâ&#x20AC;? in it. The map was digitized from an aerial infra-red photograph taken during August 1996.

days. Occasionally we collected pellets from Kestrel nests. Bones in pellets were bleached (4% hydrogen peroxide) and identified using Mendelssohn and Yom-Tov (1987) guide plates. Individuals of Mallard, Northern Shoveler and European Coot were collected by shooting and their stomachs dissected and analyzed as were faeces. During direct observations of piscivorous species, we estimated fish lengths relative to bill length. The food items identified were recorded to the lowest taxonomic level possible. Breeding A species was recorded as breeding if a nest was found or adults were observed feeding chicks. Nests were marked with individual numbers on an aerial photograph. Dates of nest building, hatching and fledging were recorded. The number of colonial waterbird nests were estimated by combining counts from observa-

tion tower, counts of nests in defined areas and direct examination of nests. Roosting Roost sites were identified by direct observations on known roost sites and potential habitats. Roosting observation efforts in January 1996 were partial. An attempt was made to count all the birds that landed at the roost at twilight. After dark, it was difficult to count or distinguish between some species such as Little Egret and Cattle Egret. To minimize mistakes caused by low visibility, counts of roosting birds were made also after sunrise, when birds left the roost. Usually only one roost site was observed at a time. We therefore lacked information on the synchronous use of several roosts. We also recorded roost sites when not in use.

wethul14.tex; 7/02/1999; 21:01; p.3

172

Figure 2. Cumulative number of feeding (grey), breeding (black) and roosting (white) species using Agmon wetland habitats during January 1996â&#x20AC;&#x201C;February 1997 and habitat areas (ha). Refer to Figure 1 and Table 1 for habitat locations and descriptions.

Results The species and their status During the study period 180 species (and one subspecies) of birds were recorded in the Agmon wetland (Appendix 1). They consisted of residents (28.2%), summer breeders (23.7%), overwintering species (43.1%), migratory species (3.3%) and occasional vagrants (1.6%). Ten of these are species that ceased breeding after the Hula drainage in the late 1950s (Yom-Tov and Mendelssohn, 1988; Dimentman et al., 1992). Sixteen species observed in Agmon are endangered in either Israel or Europe or both and five are globally threatened (Ashkenazi and Hakham, 1987; Green, 1993; Tucker and Heath, 1994). Habitats and their utilization Several habitats were defined within the study area (Table 1, Figure 1). Integrated foraging, breeding, and roosting species richness in each habitat indicated that utilization of habitat for these functions is not correlated with habitat size (Figure 2). The mostused habitats for all the functions were cattail stands, reed-beds, riparian habitats and trees, although they were not the largest available habitats. The largest habitat area, cultivated fields, ranked second among most-used habitat and was used mainly for foraging.

Seasonally-inundated habitats and mudflats were important for ca. 20 species although they were available for very short periods during the year and were among the smallest-sized habitats. The relative absence of these shallow-water feeding habitat was demonstrated by 60 Avocets (Recurvirostra avosetta) which fed and roosted in the HNR during winter 1996â&#x20AC;&#x201C;1997, but did not use Agmon for any of the activities. Moreover, the Glossy Ibis uses similar feeding habitat bred and roosted in Agmon, but foraged only in the HNR. Most habitat areas were seasonally dynamic, thereby affecting their use. However, some catastrophic (human-induced and natural) changes in some habitats greatly affected habitat use patterns. For example, beginning in late 1996, the cattail stand began deteriorating such that by February 1997 few plants existed (see Kaplan et al., 1998). During this time, most colonial and solitary roosting and breeding species left the area, with the exception of Cattle and Little Egrets that remained faithful to the site throughout early 1997. Also mowing of reed-beds in 1997 greatly restricted roosting habitat for some species. In January 1996, ca. 70 Marsh Harriers roosted in the reed-beds, whereas in January 1997 the reed-beds were mowed and few Marsh Harriers were observed roosting in Agmon.

wethul14.tex; 7/02/1999; 21:01; p.4

173

Figure 3. Foraging-species density (number of species foraging per ha) in Agmon habitats during January 1996–February 1997. Refer to Figure 1 and Table 1 for habitat locations and descriptions.

Foraging and diet Cultivated fields were used for foraging by the largest number of species, though densely vegetated lake shores, shallow canals, the cattail stand, temporary inundated areas, the lake, low grass lake shores, reedbeds, and mats of floating vegetation were all important feeding habitats (Figure 2). Density of foraging species in different habitats (Figure 3) indicated that the most densely-used foraging habitats were the low grass lake shore, mudflats, vegetated lake shore, temporary inundated areas and mats of floating vegetation (i.e., the habitats smallest in area). Foraging habitats and diet data from direct observations (1,791), regurgitations (61), stomach contents (11), faeces (48) or pellets (257) of 97 species were recorded (Appendix 1, Appendix 2). The most diverse diet was found in Cattle Egret, which forages in cultivated fields and in shallow canal banks. Diverse and seasonally changing diet was detected also in species which fed in the lake, on banks grass and in cultivated fields such as European Coot, Moorhen and Mallard. Other species were very restricted in diet, such as Purple Coot which consumed mainly cattail and Kingfisher and Pied Kingfisher which consumed exclusively fish. Prey items recorded in diets of the largest number of avian species in the Agmon wetland (Appendix 2) were Chironomidae (in the diet of 36 species), Social Vole (Microtus socialis) (17), Hymenoptera (15), fish (19), Amphibia (11), Acrididae (11), terrestrial isopods, Porcellionidae (10), butterflies, Lepidoptera (9), dragonflies and damselflies, Odonata (8), and mole crickets, Gryllotalpidae (6).

Seven fish-eating heron species (Purple Heron, Black-crowned Night Heron, Little Egret, Squacco Heron, Grey Heron, Great White Egret and Little Bittern) fed in the area of the Agmon wetlands (Table 2). Fish mostly caught in canals, near the water inflow or outflow and in the southeastern shallow lake. In March, April, June, October and November 1996, the lake was stocked with small (20–40 g) fish, mostly tilapia (Tilapia zillii) and common carp (Cyprinus carpio) (Degani et al., 1998). Mass migration of the Great White Pelican through Israel occurs during April and October (Shmueli, 1996). The earliestarriving individuals of the wintering Great Cormorant were observed in October. Stocking in March did not attract any piscivorous birds. In April, about 80 Pelicans fed in the lake for about a week. In June, about 50 Black-crowned Night Herons foraged in Agmon for two days, but Pelicans roosting at the HNR were not attracted. Stocking in October attracted 300–1,500 Pelicans, dozens of herons and a few kingfishers and in November 300–500 Pelicans each time for a week. Analyses of pellets from two Barn Owl nests in the area indicated that 82% of the food items (n = 345) were Social Vole, 25% of which were young. Breeding Main habitats used for nesting by the largest number of species (Figure 2) were generally the smaller habitat areas (cattail stand, densely vegetated lake shores, trees, reed-beds and the pasture land. Large size habitats such as cultivated fields, canals or the lake were not favored by breeding species. The highest nest density was in steep banks, cattail stand, man-made nesting sites, mudflats and bare lake shores (Figure 5).

wethul14.tex; 7/02/1999; 21:01; p.5

174

Figure 4. Monthly patterns in breeding (solid bars) and roosting (open bars) species richness in the Agmon wetland during January 1996–February 1997.

Figure 5. Nest density (number of nests per ha) in Agmon wetland habitats during March–October 1996. Refer to Figure 1 and Table 1 for habitat locations and descriptions.

Among recorded species in the Agmon wetlands, 46 species (about 25.5%), nested in the study area. These consisted of 50% of the resident and summer breeders listed in Appendix 1. The rest were either rare breeders or species (such as raptors) with nesting habitat requirements which were absent in the area. The breeding season lasted eight months (end of March to end of October) with a peak (41 and 42 breeding species) in May and June, respectively (Figure 4). Altogether, 2,040 nests were recorded (1,670 in a mixed breeding colony and 370 solitary nests) (Appendix 1). Among the solitary nesters, Crested Lark, Spur-winged Lapwing, Moorhen and seven warblers were the most abundant breeders. Some of the species, such as Spur-winged Lapwing, Crested Lark, Moorhen, White-breasted Kingfisher, Pied Kingfisher,

Common Bulbul, Fan-tailed Warbler, Striped-backed Prinia, Reed Warbler, Clamorous Great Reed Warbler, Woodchat Shrike and Yellow Wagtail, had 2–3 successful broods within the breeding season. Moorhen had an unusually large breeding population in Agmon, compared with that in the HNR, possibly due to the abundance in Agmon of the submerged macrophyte Najas spp. (a major component of Moorhen diet). Seven species, Black-crowned Night Heron, Squacco Heron, Cattle Egret, Little Egret, Purple Heron, Glossy Ibis and Pygmy Cormorant, nested in a mixed-breeding colony on the cattail stand in the southern end of Lake Agmon. The solitary heron, Little Bittern, nested in the fringes of the colony. The breeding colony was first established (six species, 500 nests) in Lake Agmon in summer 1995 (Ashkenazi

wethul14.tex; 7/02/1999; 21:01; p.6

14 5 (35.7) 4.4 + 1.1 3–6 (5) 4 1

1 2

21 8 (38.1) 9.5 + 4.2 5–15 (8) 5 2 17 7 (41.2) 14.2 + 5.1 6–20 (5) 2 1 1 1 36 21 (58.3) 5.35 + 2.1 3–12 (14) 13 8

8 8 (100) 3.5 ± 0.5 3–4 (8) 4 4 No. of diet records No. of fish (% of diet) Fish length, cm (mean ± SD) Fish length range, cm (n) Unidentified fish Tilapia zillii Tilapia (red hybrid) Cyprinus carpio Gambusia affinis Clarias gariepinus Oncorhynchus mykiss

11 11 (100) 3 + 0.6 2–4 (7) 6 3

21 21 (100) 4.3 + 2.0 2–10 (13) 17 3

137 128 (93.4) 4.41 + 1.8 1–15 (122) 20 78 11 19

18 11 (61.1)

Purple Heron Black-crowned Night Heron Squacco Heron Great White Egret Little Egret Pied Kingfisher Little Bittern

Kingfisher

and T. Oron, unpublished data). In 1996 breeding activity started in mid March, but the colony was deserted three times due to motor boat activity. Successful breeding began only in April when dense algal mats prevented boating in the vicinity of the cattail stand. Desertion of the breeding colony was followed by desertion of the communal roost site on the cattail stand. Solitary ground-nesting birds were also affected from timing of human activity such as mowing and harvesting within the Agmon wetland. In 1996, two nests of Black-headed Yellow Wagtails, two nests of Spur-winged Lapwings, five nests of Bee-eaters and two nests of Crested Larks were destroyed during mowing, harvest or hay collection.

Parameter

Table 2. Fish species composition and size in the diets of piscivorous waterbirds at the Agmon wetland (January 1996–February 1997).

White-breasted Kingfisher

175

Roosting The most-used roost habitats were the cattail stand, reed-beds, and trees. Mudflats, temporary inundated areas and low grass lake shores were also favored habitats for roosting. Roads and irrigation systems were used for roosting by several species. Roost sites were used mainly in winter and during migration (spring and autumn). Most of roost sites were not in use during the summer (breeding season). However, the most used roost site, the cattail stand, was in use throughout the year by seven colonial nesting species, two overwintering herons (Grey Heron and Great White Egret) and several occasional roosting species such as swallows and wagtails. Fifty-five species were recorded roosting in the study area, with each species observed on 1–37 occasions and many of which were roosting throughout the year (Figure 4, Appendix 1). The number of species utilizing the Agmon wetland habitats at the same time for nesting and roosting varied between 9–45. The number of roosting species was higher in February 1997 (33) compared to February 1996 (15). Numbers of individuals of a species at a roost varied from 2 to 6,000 per night. The maximum number of roosting individuals per species per night was > 100 (30 species), > 1,000 (9 species), and > 5,000 (2 species). Roostjoining behavior varied in different species. Some arrived to a nearby pre-roost before sunset, e.g., Great White Egret and Stock Dove. Others joined the roost at dark, such as Mallard, Grey Heron and Marsh Harrier. Some arrived in large flocks, such as Cattle Egret, Little Egret, Glossy Ibis and Yellow Wagtail, while others joined the roost individually, e.g., Squacco and Grey Herons. In the communal roost site in the cattails, different species had a tendency to roost in a different

wethul14.tex; 7/02/1999; 21:01; p.7

176 section of the colony. This pattern was consistent until changes in species composition (departure of summer breeders such as Purple Heron and Squacco Heron) or changes in the habitat quality (deterioration of the cattail habitat or mowing of Common Reed stands) occurred.

Discussion Proper management of an area for avian species requires information on habitat specifications for the important functions of their life history (Kushlan, 1993; Erwin; 1996b, Hoffman et al., 1996). Although habitats for basic functions such as feeding, nesting and roosting may be independent and available in different places, there is an energetic advantage when habitats for several functions are proximal to each other (Hafner and Fasola, 1992; Hoffman et al., 1996; Rehfisch et al., 1996). Integration of species habitat requirements for foraging, breeding and roosting contributes to a better understanding of species resource requirements for planning a suitable management program on areas utilized on year-round basis. This is important particularly in management of a multi-purpose area, when attraction of populations to certain habitats (and avoiding damage to other habitats) is essential for minimizing land-use conflicts. We assume that the large number of species utilizing the area of Agmon resulted from the high diversity of habitats with their diverse, abundant food resources. However, the number of species utilizing these habitats was not related to habitat area. Rafe et al. (1985) found that both area and habitat diversity influence species richness and the number of species is better correlated with habitat diversity than with area. Presence of several congeneric species indicates a high suitability of the habitat to characteristics of the genus, and a richness and diversity which permit resource partitioning of closely related species. This was demonstrated by several waterbirds including 6 ducks (Anas spp.), 6 Tringa species and 3 crakes. Similarly, the Agmon wetland supports species having a special value in agricultural pest control, including 4 shrikes, 4 wagtails, 4 pipits, 5 falcons, 4 harriers and 4 eagles (Aquila spp.). Another correlate of well-established communities and diverse habitats is the complexity of the trophic levels in the ecosystem. The Agmon wetland system harbors a rich ensemble of species from herbivores such as Goose and Ruddy Shelduck,

to secondary or tertiary predators such as White-tailed Eagle. The value of the diverse habitats of Agmon for its bird assemblage was primarily for foraging (by highest number of species), but also for year-round roosting. Nesting habitats were utilized by the lowest number of species and for limited time during the year. However, nesting habitats for the endangered Marbled Teal, Black-headed Yellow Wagtail and Pratincole are available in Agmon. The deterioration of the most heavily used habitat for nesting and roosting, the cattail stand (Kaplan et al., 1998), in late 1996 will probably result in a decrease in breeding and roosting species richness. Although the site was very attractive for tourism, in terms of nature conservation, these colonial waterbirds have an alternative for both activities in the nearby HNR. However, habitats for the above mentioned endangered breeding species are rare or absent in the HNR and their maintenance in Agmon wetland may contribute to regional breeding species richness and global conservation of these species. Mudflats and temporary inundated habitats were among the smallest and short-lived (mostly 2â&#x20AC;&#x201C;7 days) habitats. However, when available they were utilized by waders and ducks both for feeding and roosting. Temporarily inundated habitats, usually created in winter, are attractive to breeding amphibians. Tadpoles were very attractive prey for several species, including herons, at the end of the winter. Shallow habitats are essential for waders which forage largely in water less then 18 cm deep (Rehfisch, 1994; Erwin, 1996b). Food and water conditions in such habitats are ideal for Mallards and Pintails which can swim but still dabble and tip up for food (Weller, 1990). We assume that enlargement of these habitats and increasing of their duration will increase the number of species and individuals which utilize them. Characteristics of diets of selected Agmon avian species A main feature that characterized the habitats and diets of many species in the Agmon wetland was the use of both wetland habitats and adjacent cultivated fields. The importance of cultivated field habitat for foraging emphasizes the need for an effective management program to avoid impact on crops while providing â&#x20AC;&#x153;biological pest control servicesâ&#x20AC;?. Cultivated fields provided rich and accessible insect and rodent fauna for species such as Cattle Egret, White Stork and several raptors. These fields had a large population of

wethul14.tex; 7/02/1999; 21:01; p.8

177 social voles with a long history of serious damage to crops, resulting in heavy use of rodenticides (YomTov and Mendelssohn, 1988). Mass consumption of social voles (one of the most important prey items for birds) in Agmon during the whole year may reduce vole damage to crops. Changes in food availability in the aquatic environment were reflected in the foraging activities and diets of European Coot, Moorhen and dabbling ducks. In January and early February 1996, there were no dabbling ducks in Lake Agmon while thousands of them fed in the HNR lake. Samples of zooplankton collected in both lakes indicated high winter densities of zooplankton in the HNR lake compared with in Lake Agmon. Similarly, an increase in the zooplankton densities in Lake Agmon during winter 1997 led to an increase in dabbling duck abundances. Fish in the Agmon wetlands have two distinctive groups which differ in availability: (1) The group of spontaneous colonizing indigenous fish, entered the main waterbody and the canals with the inflow water, in a continuous colonizing process; (2) Periodically stocked (from commercial freshwater ponds) fish fry, characterized by a few unpredictable pulses of large numbers at a time. Piscivorous species were observed eating fish in large groups only: (1) after stocking of the lake with fish; (2) during periods of low oxygen concentrations in the water in summer (mostly early in the morning); (3) after periods of low temperature (below 0 ◦ C for few hours (Tsipris and Meron, 1998); and (4) in situations when fish were trapped in shallow ponds following decreases in water level. In all four cases, fish densities were locally increased and the fish were more vulnerable to predation. The role of stocked fish for attracting large piscivorous birds for wildlife tourism is not clear. It was impossible to assess the impact of piscivorous birds due to lack of information on: (1) rates of natural fish immigration and emigration; (2) reproduction rates of fish within the lake; (3) the amount of fish removed by angling. Stocked fish were noticeably attractive only during the peak of the pelican migration (April and October), for a maximum of one week each. Fish fry eaten by the small herons in the lake were much smaller than the stocked fish size and were probably from naturally immigrated or reproduced fish. Creation of stable and attractive fish foraging habitat requires management of predictable and large fish populations. However, dense fish populations have a major impact on the distribution and abundance of other aquatic organisms and may cause changes to invertebrate fauna which affects

the non-piscivorous bird populations (Britton, 1982; Eriksson, 1987; Poysa et al., 1994). There is a need for further study for understanding the needed densities of fish and zooplankton in the lake for maintaining both piscivorous and zooplanktivorous birds for wildlife tourism. Nesting and roosting The richness of breeding species, the composition of the mixed-breeding colony and the long breeding season with multiple successful broods in some species, indicates that the Agmon wetland and surroundings are very productive habitats. Moreover, the establishment of a new heron, cormorant and ibis breeding colony at the Agmon wetland, two km from the traditional colony at the HNR, emphasizes the resource richness of the area for these species. All colonial breeding species in Agmon nested on the cattail stand. Most of solitary species nests were in the riparian habitats along the lake, the diverse pasture land, bare moderate or steep banks and trees. Nest densities in these small area habitats, in particular in the banks and riparian lake shores were the highest. Similarly, breeding bird densities in the United States are among the highest in riparian habitats (Carothers et al., 1974). Very little research has been devoted to understanding how roost availability contributes to longterm stability of avian populations (Draulans and Van Vessem, 1986; Rehfisch et al., 1996). There is limited information on the importance of factors determining roost-site selection such as energy limitations, predation, disease and weather conditions (Birkhead, 1973; Blanco, 1996; Rehfisch et al., 1996). Roost sites are particularly important for migrating or wintering populations like harriers, wagtails, swallows and various waders (Mitchell et al., 1988; Rehfisch et al., 1996). Roosting species richness was highest during the spring (March–April) and autumn (September– November). However, unusual amounts of flooded habitats in winter 1996–1997, in particular during February 1997, attracted a peak number of 33 roosting species. Flooded habitats were created on this occasion on the large islands, attracting foraging and roosting dabbling ducks, some rare waders and raptors. The number of roosting species during the breeding season was low due to dispersal of the solitary species for cryptic breeding activities. Eight of the colonial-roosting species (seven breeding species and the wintering Great White Egret) were faithful to a single roosting site (cattail stand). The only excep-

wethul14.tex; 7/02/1999; 21:01; p.9

178 tion was the Grey Heron which roosted alternately in two different habitats (cattail and trees). Glossy Ibis bred and roosted in 1996 only in Agmon (except for a single pair that nested in the HNR) although they bred and roosted in the HNR since the 1970s. We are uncertain whether there is an exchange between populations of some colonial roosting species with the HNR. However, breeding and roosting activities in the colony of the HNR were terminated in early August 1996 (T. Oron, pers. comm.). We assume that the increase in colonial roosting birds in Agmon from 6,000 in July to 10,000 in August was due to movement of the HNR populations to Agmon. Roosting populations decreased after the summer breeders migrated (August–September) and Black-crowned Night Herons shifted to night activity. The unusual double function of the cattail stand, the influx of the entire Hula Valley colonial waterbirds to the roost site in August, and the high site tenacity of the species, renders this habitat as critical for colonial waterbirds. Management implications The Agmon wetland area is a multi-purpose (recreation, wildlife tourism and agriculture) area that demands a very detailed, sensitive and dynamic management program to maintain the delicate balance among its different components. Management of the avian populations in this diverse area has to consider the available habitats in the entire Hula Valley, including the HNR and the agricultural areas. The management plans for the area should analyze allocation of habitats for species that are endangered, rare, or of conservation interest within the HNR and “tourism species” within the Agmon wetland. “Tourism species” are usually large (Pelican, Cormorants, Cranes), gregarious, easily observed (colonial waterbirds) or rare species (raptors, composed of 26 species in Agmon). However, endangered or rare species that feed, breed or roost in the newly created habitat despite the protected habitats of HNR, must receive special attention in management of the multi-purpose Agmon wetland. From a regional management perspective, and in terms of habitat use by the avian fauna, the Agmon wetland and the HNR are a part of a large mosaic of habitats. The avian fauna in the region use both sites either for foraging, nesting or roosting. Common Crane and Great Cormorant populations roosted only in the HNR, but foraged in surrounding cultivated fields and commercial fish ponds, respectively. Glossy Ibis foraged in the HNR, but nested and roosted only

in Agmon. Mallards use forage and roost sites either in the HNR or in Agmon in different periods. The movements between the sites for different activities, is facilitated by the Jordan River eastern and western canals which were the main flyways (connecting corridors) used in particular by cranes, herons and ducks after dark. Any disturbance in these flyways to and from roost sites should be avoided. Moreover, human activities near nest sites during mid March– September and near shorebird roost sites in March– May and September–November should be restricted. We suggest management guidelines used to avoid human disturbance to colonial breeding waterbirds in the US (Erwin, 1989, 1996a): 100–200 m buffer distance from wading birds nesting sites, 50–100 m from roosting shorebirds and at least 400 m from nesting cormorants. Shallow muddy foraging habitats were available only on a few occasions and in a very small area, when water level decreased. However, whenever created, mudflats immediately attracted a rich ensemble of foraging and roosting waders for a few days. The food availability of the large islands for most of the waterbird species could be improved by creating shallow impoundments to retain water more frequently and for longer periods. We recommend moderating slopes along the islands shores to create mudflats during the entire year, but in particular during the migrating seasons of wading birds (March–April and September–November). Most of the Lake Agmon shores were planned steeply, to preclude breeding of mosquitoes, mainly malaria-vector species (Anopheles spp.). However, mosquito predators such as the indigenous fish Aphanius mento, the exotic Mosquitofish (Gambusia affinis), amphibians and several invertebrates may eliminate the mosquito problem in such habitats. Neither of the two moderate-sized islands is accessible to the public and would provide protected habitat for species that require food characterized by temporary habitats and mudflats. Drying-out periods of certain parts of the mudflats (out of the migrating season) are recommended. This will create food resources characterizing “disturbed” habitats such as chironomid-dominated early-colonizer community (Rehfisch, 1994) which are important food sources during the breeding season (Cox and Kadlec, 1995). We suggest plowing the elevated parts of the large islands and dry pasture land in the end of March, to create nesting and roosting habitat for the endangered Pratincole. A series of small vegetated islets in the lake’s center are not used by birds. We suggest

wethul14.tex; 7/02/1999; 21:01; p.10

179 de-vegetating them by mowing followed by thermal treatment (black polyethylene) to create bare habitats for roosting and ground-breeding colonial species. We recommend to cover the surface of some of these islets with pebble, to create nesting habitats for terns and plovers that have very limited suitable habitat in the Hula Valley. Lake water levels should be managed very carefully to create rich seasonal food webs and available foraging and roosting habitats. It is suggested to mimic the natural water level fluctuations in the former Lake Hula: high water levels in winter, with a maximum in March, lowering water level gradually throughout the summer and increasing again beginning in November, after the fall migration. Increases in water level should be avoided during April–July to avoid flooding of ground nesting bird nests. Reed stands are among the most important habitats for breeding and roosting. We suggest that at least 6–8 ha of reed-beds should be protected from mowing to provide an area suitable for roosting harriers. This habitat may partially substitute for some of the functions provided by the deteriorated cattail. We advise vegetation be cut around the waterbody before the breeding season (mid-to-late March) and at the end of the nesting season (October). Also, it is recommended that harvested vegetation be removed from the fields within a week, to avoid ground nest destruction. The late mowing and silage making contributed in Europe to a decline in species such as Lapwing, Meadow Pipit and Yellow Wagtail (Anderson, 1995). Mowing of lake shores in winter is not recommended as it eliminates protected habitats and roosting sites for waterbirds. Also, it exposes the emergent vegetation to cold, stressful conditions. Some parts of this riparian habitat should not be mowed at all. On the other hand, mowing some areas (e.g., fallow agricultural fields surrounding the lake) in the fall will increase exposure of social voles to predation by migrating and wintering raptors, possibly decreasing the vole’s impact on winter crops.

Acknowledgments We thank E. Yas’ur for providing facilities and help with sampling habitats. G. Gississ, T. Oron, Y. Vaadia, O. Rabinovich and K. Merom provided information on presence of bird species. Y. Ayal, H. Bromley, I. Dor, D. Golani, M. Goren, G. Levi, H. Mienis, U. Pollingher, T. Shariv, E. Tchernov and Y. Yom-

Tov helped with taxonomic identifications. A. Kadmon helped with the GIS produced map. M. Erwin, Y. Yom-Tov and an anonymous reviewer provided valuable comments on the manuscript. The study was funded during January–December 1996 by Keren Kayemet L’Israel (JNF), Israel Land Development Authority and the Israel Ministry of Agriculture.

References Anderson, P. 1995. Ecological restoration and creation: a review. Biol. J. Linn. Soc. 56 (Suppl.): 187–211. Ashkenazi, S. and Hakham, E. 1987. Names of the vertebrates in Israel. Nature Conservation in Israel, Suppl. 1. Nature Reserves Authority, Jerusalem. Ashkenazi, S. and Yom-Tov, Y. 1997. The breeding biology of the black-crowned night-heron (Nycticorax nycticorax) and the little egret (Egretta garzetta) at the Huleh Nature Reserve, Israel. J. Zool. Lond. 242: 623–641. Birkhead, T.R. 1973. A winter roost of Grey Herons. British Birds 66: 147–156. Blanco, G. 1996. Population dynamics and communal roosting of White Storks foraging at a Spanish refuse dump. Colonial Waterbirds 19: 273–276. Britton, R.H. 1982. Managing the prey fauna. In: Scott, D.A. (ed.), Managing Wetlands and Their Birds. pp. 92–97. IWRB, Slimbridge, England. Carothers, S.W., Johnson, R.R. and Aitchison, S.W. 1974. Population structure and social organization in southwestern riparian birds. Amer. Zool. 14: 97–107. Cox, R.R. and Kadlec, J.A. 1995. Dynamics of potential waterfowl foods in Great Salt Lake marshes during summer. Wetlands 15: 1–8. Degani, G., Yehuda, Y., Jackson, J., and Gophen, M. 1998. Temporal variation in fish community structure in a newly created wetland lake (Lake Agmon) in Israel. Wetlands Ecol. Managmt. 6: 151–157. Dimentman, Ch., Bromley, H.J. and Por, F.D. 1992. Lake Hula: Reconstruction of the Fauna and Hydrobiology of a Lost Lake. The Israel Academy of Sciences and Humanities, Jerusalem. Draulans, D. and Van Vessem, J. 1986. Communal roosting in Grey Herons (Ardea cinerea) in Belgium. Colonial Waterbirds 9: 18– 24. Eriksson, M.O.G. 1987. Some effects of freshwater acidification on birds in Sweden. In: Diamond, A.W. and Filion, F.L. (eds.), The Value of Birds. pp. 183–190. International Council for Bird Preservation (ICBP) Tech. Pub. No. 6. Erwin, R.M. 1989. Responses to human intruders by birds nesting in colonies: experimental results and management guidelines. Colonial Waterbirds 12: 104–108. Erwin, R.M. 1996a. Dependence of waterbirds and shorebirds on shallow-water habitats in the mid-Atlantic coastal region: an ecological profile and management recommendations. Estuaries 19: 213–219. Erwin, R.M. 1996b. The relevance of the Mediterranean region to colonial waterbird conservation. Colonial Waterbirds 19 (Spec. Pub. 1): 1–11. Green, A.J. 1993. The status and conservation of the Marbled Teal Marmaronetta angustirostris. International Waterfowl and Wetlands Research Bureau Spec. Publ. 23, Slimbridge.

wethul14.tex; 7/02/1999; 21:01; p.11

180 Hafner, H. and Fasola, M. 1992. The relationship between feeding habitat and colonially nesting Ardeidae. In: Finlayson, C.M., Hollis, G.E. and Davis, T.J. (eds), Managing Mediterranean Wetlands and Their Birds. pp. 194–201. International Waterfowl and Wetlands Research Bureau Spec. Pub. 20, Slimbridge. Hambright, K.D. and Zohary, T. 1998. Lakes Hula and Agmon: destruction and creation of wetland ecosystems in northern Israel. Wetlands Ecol. Managmt. 6: 83–89. Hoffman, L., Hafner, H. and Salathe, T. 1996. The contribution of colonial waterbirds research to wetland conservation in the Mediterranean region. Colonial Waterbirds 19 (Spec. Pub. 1): 12–30. Kaplan, D., Oron, T. and Gutman, M. 1998. Development of macrophytic vegetation in the Agmon wetlands of Israel by spontaneous colonization and reintroduction. Wetlands Ecol. Managmt. 6: 143–150. Kushlan, J.A. 1993. Colonial waterbirds as bioindicators of environmental change. Colonial Waterbirds 16: 223–251. Mendelssohn, H. and Yom-Tov, Y. 1987. Appendix to volume 7: Mammals, skull and body measurements and plates. In: Alon, A. (ed.), Plants and Animals of the Land of Israel, An Illustrated Encyclopedia. pp. 111. Ministry of Defence, The Publishing House, The Society for Protection of Nature in Israel, Jerusalem. Mitchell, J.R., Moser, M.E. and Kirkby, J.S. 1988. Declines in midwinter counts of waders roosting on the Dee Estuary. Bird Study 35: 191–198. Poysa, H., Rask, M. and Nummi, R. 1994. Acidification and ecological interactions at higher trophic levels in small forest lakes: the perch and common goldeneye. Ann. Zool. Fenn. 31: 397–404. Rafe, R.W., Usher, M.B. and Jefferson, R.G. 1985. Birds on reserves: the influence of area and habitat on species richness. J. Appl. Ecol. 22: 327–335.

Rehfisch, M.M. 1994. Man-made lagoons and how their attractiveness to waders might be increased by manipulating the biomass of an insect benthos. J. Appl. Ecol. 31: 383–401. Rehfisch, M.M., Clark, N.A., Langston, R.H.W. and Greenwood, J.J.D. 1996. A guide to the provision of refuges for waders: an analysis of 30 years of ringing data from the Wash, England. J. Appl. Ecol. 33: 673–687. Shirihai, H. 1996. The Birds of Israel. Academic Press, London. Shmueli, M. 1996. Ecophysiology of the Great White Pelican (Pelecanus onocrotalus) during migration and wintering in Israel. M.Sc. Thesis, Israel Institute of Technology, Haifa (in Hebrew, with English summary). Shy, E., Beckerman, S., Oron, T. and Frankenberg, E. 1998. Repopulation and colonization by birds in the Agmon wetland, Israel. Wetlands Ecol. Managmt. 6: 159–167. Tsipris, J. and Meron, M. 1998. Climatic and hydrological aspects of the Hula restoration project. Wetlands Ecol. Managmt. 6: 91– 101. Tucker, G.M. and Heath, M.F. 1994. Birds in Europe: Their Conservation Status. Birdlife International (Birdlife Conservation Series No. 3) Cambridge, U.K. Weller, M.W. 1990. Waterfowl management techniques for wetland enhancement, restoration and creation useful in mitigation procedures. In: Kusler, J.A. and Kentula, M.E., (eds), Wetland Creation and Restoration, the Status of the Science. pp. 517–528. Island Press, Washington DC. Yom-Tov, Y. and Mendelssohn, H. 1988. Changes in the distribution and abundance of vertebrates in Israel during the 20th century. In: Yom-Tov, Y. and Tchernov, E. (eds), The Zoogeography of Israel. pp. 514–547. Dr. W. Junk, Dordrecht. Zahavi, A. 1957. The breeding birds of the Huleh swamp and lake (Northern Israel). Ibis 99: 600–607.

wethul14.tex; 7/02/1999; 21:01; p.12

Appendix 1. The conservation status and habitat utilization (January 1996–February 1997) of wetland bird species in the Agmon wetlands. Conservation status in Israel and Europe (Ashkenazi and Hakham, 1987; Tucker and Heath, 1994): () – Provisional, X – Extinct, E – Endangered, V – Vulnerable, R – Rare, D – Declining, I – Insufficiently known, S – Secure, T – Threaten undetermined, A – Remarkable increase, X-br – not breeding since the draining of Lake Hula (Zahavi, 1957; Dimentman et al., 1992), – global concern. Residence status – Partly based on Shirihai (1996): R – resident, R(nB) – non-breeding resident, W – overwinter, SB – summer breeder, rSB – rare summer breeder, S(nB) – non-breeding summer visitor, M – migrating, O – occasional visitor. (n) – number (nests, roosting or diet data), Nmax – max. roosting population, + < 7 diet records; nesting data: * – three eggs laid in Carrion Crow nest, ** – nesting potential (former breeder, habitat available). Scientific name

Common name

Status Israel

Status Europe

Residence

Nest No.

Nesting habitat

Diet data (n)

Feeding habitat

Roosting (n, Nmax )

Roosting habitat

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

Tachybaptus ruficollis Podiceps cristatus Podiceps grisegena Podiceps nigricollis Phalacrocorax carbo Phalacrocorax pygmeus Pelecanus onocrotalus Botaurus stellaris Ixobrychus minutus Nycticorax nycticorax Ardeola ralloides Bubulcus ibis Egretta garzetta Egretta alba Ardea cinerea Ardea purpurea Ciconia nigra Ciconia ciconia Plegadis falcinellus Platalea leucorodia Anser albifrons Anser anser Tadorna ferruginea Anas penelope Anas strepera Anas crecca Anas platyrhynchos Anas acuta Anas querquedula Anas clypeata Marmaronetta angustirostris

Little Grebe Great Crested Grebe Red-necked Grebe Black-necked Grebe Great Cormorant Pygmy Cormorant Great White Pelican Bittern Little Bittern Black-crowned Night-Heron Squacco Heron Cattle Egret Little Egret Great White Egret Grey Heron Purple Heron Black Stork White Stork Glossy Ibis Spoonbill White-Fronted Goose Grey-lag Goose Ruddy Shelduck European Wigeon Gadwall Teal Mallard Pintail Garganey Northern Shoveler Marbled Teal

V V, X-br R R S (X)1 , E S V E S V A S2 S S, X-br V S S V V R R R R R S3 S S3 S3 S E

S S S S S V R (V) (V) D V S S S S V R V D E S S V S V S S V V S E (#)

R+W W O W W R+SB W+M+R(nB) W+M SB R+SB SB R R+SB W W SB W+M rSB+M R+SB W W W W W W W R+SB W W W R

1 **

8, 96

1, 5

2 26

1, 5

25, 56 11, 1200 11, 2

2 2, 7a 6, 2

5, 2, 3a 2, 3a-b, 5, 12 5, 3a-b, 12, 2 12, 5, 3b 5, 3a, 2 12, 5, 2 12, 5, 2 12, 5

21, 300 13, 70 32, 5000 37, 414 33, 1088 27, 20 13, 21

2 2 2 2 2 2, 11 2

12 700 45 700 150

2, 3a, 5 2 2 2 2

** 40

8 17 35 113 94 18 + 21

+ 10

12 9

12, 200 25, 185

11, 14 2

2, 3a

7 + + 121

3b, 12, 2, 9 1, 9 2, 9, 1 12, 2, 1, 8, 9

7, 180 3, 29 6, 30 33, 3800 1, 10

3b, 13 1 2 2, 3b, 5 1

2, 3a

+ 26 +

9 1, 9 2, 9, 8, 5

2, 110 5, 6

2 2

181

wethul14.tex; 7/02/1999; 21:01; p.13

Code

182

Appendix 1. Continued.

wethul14.tex; 7/02/1999; 21:01; p.14

Code

Scientific name

Common name

Status Israel

Status Europe

Residence

32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

Aythya ferina Aythya nyroca Aythya fuligula Pernis apivorus Milvus migrans Haliaeetus albicilla Circaetus gallicus Circus aeruginosus Circus cyaneus Circus macrourus Circus pygargus Accipiter gentilis Buteo buteo Buteo rufinus Aquila pomarina Aquila clanga Aquila rapax Aquila heliaca Hieraaetus pennatus Pandion haliaetus Falco naumanni Falco tinnunculus Falco columbarius Falco subbuteo Falco peregrinus Alectoris chukar Francolinus francolinus Coturnix coturnix Rallus aquaticus Porzana porzana Porzana parva Porzana pusilla Gallinula chloropus Porphyrio porphyrio Fulica atra Grus grus

Pochard Ferruginous Duck Tufted Duck Honey Buzzard Black Kite White-tailed Eagle Short-toed Eagle Marsh Harrier Hen Harrier Pallid Harrier Montaguâ&#x20AC;&#x2122;s Harrier Goshawk Buzzard Long-legged Buzzard Lesser-spotted Eagle Spotted Eagle Tawny Eagle Imperial Eagle Booted Eagle Osprey Lesser Kestrel Kestrel Merlin Hobby Peregrine Chukar Black Francolin Common Quail Water Rail Spotted Crake Little Crake Baillonâ&#x20AC;&#x2122;s Crake Moorhen Purple Coot European Coot Common Crane

S3 S3 , X-br S3 S S E4 , X-br S V, X-br R R R R S V S S, X-br S R R R E S E I R S V V R R R R, X-br S R S S

S V (#) S S V R R S V E S S S (E) R E (#) E (#) R R (V)(#) D S S R V V V (S) S (S) R S R S V

W W W M rSB+W R(nB)+M SB+M W+M W+M W+M W+M W+M rSB+W+M R+W+M W+M W+M O W+M W+M W+M SB+W+M R W+M SB+M W+M R R SB+M rSB+W+M W W+M+S(nB) W+M+S(nB) R W+M rSB+R(nB)+W W

Nest No.

Nesting habitat

Diet data (n)

Feeding habitat

Roosting (n, Nmax )

Roosting habitat

3, 150

+ +

12, 3b 12, 3b

11, 70

11, 14

22 + + +

3a-b, 12, 2, 6 12, 3a-b, 6 12, 3a-b, 6 12, 3a-b, 6

15, 68 3, 5 2, 7

6 6 6

2, 2

3, 15

11, 300

2, 5, 8

21, 1200

2, 5, 8

14, 11

3b, 12

1 8

10 6, 10

13 12 16 570 18 164 +

2, 3a, 8 2, 3a, 8 2, 3a, 8 2, 3b, 8, 5 2, 3a, 8 2, 1, 3b, 8, 5 12

** 30

2, 3a, 5

Appendix 1. Continued. Scientific name

Common name

Status Israel

Status Europe

Residence

68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103

Rostratula benghalensis Himantopus himantopus Burhinus oeducnemus Glareola pratincola Charadrius dubius Charadrius hiaticula Charadrius alexandrinus Hoplopterus spinosus Vanellus vanellus Calidris minuta Philomachus pugnax Gallinago gallinago Gallinago media Limosa limosa Numenius arquata Tringa erythropus Tringa totanus Tringa stagnatilis Tringa nebularia Tringa ochropus Tringa glareola Actitis hypoleucos Larus ichthyaetus Larus minutus Larus ridibundus Larus cachinnans Sterna hirundo Sterna albifrons Chlidonias hybridus Chlidonias niger Chlidonias leucopterus Columba (livia X domestica) Columba oenas Streptopelia decaocto Streptopelia turtur Streptopelia senegalensis

Painted Snipe Black-winged Stilt Stone Curlew Pratincole Little Ringed Plover Ringed Plover Kentish Plover Spur-winged Lapwing Lapwing Little Stint Ruff Common Snipe Great Snipe Black-tailed Godwit Eurasian Curlew Spotted Redshank Redshank Marsh Sandpiper Greenshank Green Sandpiper Wood Sandpiper Common Sandpiper Great Black-headed Gull Little Gull Black-headed Gull Yellow-legged Gull Common Tern Little Tern Whiskered Tern Black Tern White-winged Black Tern Rock Dove X Domestic Dove Stock Dove Collared Dove Turtle Dove Laughing Dove

R S S E I S E S S S S R R S R S S S S S S S S R A S E I, X-br R, X-br S A R S S A

S V E (S) S D (E) (S) (S) (S) (S) (V) V D S D (S) S (S) D S S D S (S) S D D D S S S (S) D (S)

O R+M R+M SB+M SB+M+W W+M R+W+M R W+M W+M W+M W+M M W+M W+M W+M W+M W+M W+M W+M W+M W+M W+M W+M W+M W+M SB SB+W+M W+M M+S(nB) W+M R W+M R SB+M R

Nest No.

Nesting habitat

Diet data (n)

Feeding habitat

Roosting (n, Nmax )

Roosting habitat

7a, 9

3, 24

7a, 9

10, 12

air: 12, 1, 3a-b

3, 120

7a, 9

7a, 10, 12, 13

+ 35 +

7a 12, 3b, 13 12, 9

9, 300 4, 130

7a, 9 7a, 9

+ +

7a, 9 7a, 9

9, 30 5, 14

7a, 9 7a, 9

+ + +

5, 7a 3a, 5, 7a, 9, 8 1

+ + +

1 1, 12 1

1, 4

4, 3000 12, 1500

11 11

** **

4 4 1

11 11 14

+ + +

12 12, 13 12, 13

183

wethul14.tex; 7/02/1999; 21:01; p.15

Code

184

Appendix 1. Continued.

wethul14.tex; 7/02/1999; 21:01; p.16

Code

Scientific name

Common name

Status Israel

Status Europe

Residence

Nest No.

Nesting habitat

Diet data (n)

Feeding habitat

104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138

Clamator glandarius Tyto alba Otus scops Asio otus Asio flammeus Caprimulgus europaeus Apus apus Apus melba Apus affinis Halcyon smyrnensis Alcedo atthis Ceryle rudis Merops superciliosus Merops apiaster Coracias garrulus Upupa epops Galerida cristata Alauda arvensis Riparia riparia Hirundo rustica Hirundo daurica Delichon urbica Anthus campestris Anthus pratensis Anthus cervinus Anthus spinoletta Motacilla flava Motacilla flava feldegg Motacilla citreola Motacilla cinerea Motacilla alba Pycnonotus barbatus Erithacus rubecula Luscinia svevica Phoenicurus ochruros

Great Spotted Cuckoo Barn Owl Scops Owl Long-eared Owl Short-eared Owl Nightjar Swift Alpine Swift Little Swift White-breasted Kingfisher Kingfisher Pied Kingfisher Blue-cheeked Bee-eater Bee-eater Roller Hoopoe Crested Lark Skylark Sand Martin Swallow Red-rumped Swallow House Martin Tawny Pipit Meadow Pipit Red-throated Pipit Water Pipit Yellow Wagtail Black-headed Yellow Wagtail Yellow-headed Wagtail Grey Wagtail White Wagtail Common Bulbul Robin Bluethroat Black Redstart

V S V I R S I I S S S S I S V S S S S S S V I S S R S E R S S A S T S

S D (D) S (V) (D) S (S) (S) (S) D (S) (S) (D) (D) S (D) V D D S S V S (S) S S (S) (S) S S S S

SB+M R SB+W+M R+W+M SB+W+M W+M SB+W+M SB+W+M R R W+M R SB+M SB SB R+M R W W+M R+M SB+W SB+W SB+W W+M W+M W+M SB+W+M SB+M W+M W+M SB+W+M R W+M W+M SB+W+M

1 2

11* 14

+ 322

12, 11 12, 3b, 11

3b, 12

air: 12, 1

5, 3a, 2, 10, 11 5, 3a 5, 3a

14 11 21

air: 12, 3aâ&#x20AC;&#x201C;b

3b, 7a, 10, 12

4 8

7a, 3a 7a, 3a

+ 41 + 26 58 7 14 + + + + 46 44

12, 11 12, 3b, 13 12, 3b, 13 air: 2, 1, 12 air: 2, 1, 12 air: 2, 1, 12 air: 2, 1, 12 5, 9 12, 5, 9, 8 12, 5, 9 5, 9, 8 12, 5, 7a, 8, 9 12, 5, 7a, 8, 9

113 +

12, 5, 7a, 8, 9 11, 10

5, 8

Roosting (n, Nmax )

Roosting habitat

13, 150 14, 100 3, 1000 11, 6000 3, 1000 3, 1000

13, 3b 13, 3b 2, 6 2, 6 2, 6 2, 6

9, 300 6, 20

2, 6 2, 6

7, 200

2, 6

Appendix 1. Continued. Scientific name

Common name

Status Israel

Status Europe

Residence

139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173

Saxicola rubetra Saxicola torquata Oenanthe isabellina Oenanthe oenanthe Turdus merula Cettia cetti Cisticola juncidis Prinia gracilis Locustella luscinioides Acrocephalus melanopogon Acrocephalus schoenobaenus Acrocephalus palustris Acrocephalus scirpaceus Acrocephalus stentoreus Acrocephalus arundinaceus Hippolais pallida Sylvia melanocephala Sylvia curruca Sylvia atricapilla Phylloscopus collybita Phylloscopus trochilus Parus major Remiz pendulinus Lanius collurio Lanius minor Lanius excubitor Lanius senator Corvus monedula Corvus corone Sturnus vulgaris Passer domesticus Passer hispaniolensis Passer moabiticus Fringilla coelebs Serinus serinus

Whinchat Stonechat Isabelline Wheatear Wheatear Turdus Merula Cettiâ&#x20AC;&#x2122;s Warbler Fan-tailed Warbler Striped-backed Prinia Saviâ&#x20AC;&#x2122;s Warbler Moustached Warbler Sedge Warbler Marsh Warbler Reed Warbler Clamorous Great Reed Warbler Great Reed Warbler Hippolais pallida Sardinian Warbler Lesser Whitethroat Blackcap Chiffchaff Willow Warbler Great Tit Penduline Tit Red-backed Shrike Lesser Grey Shrike Great Grey Shrike Woodchat Shrike Jackdaw Carrion Crow Starling House Sparrow Spanish Sparrow Dead-sea Sparrow Chaffinch Serin

S S S S S S V S T V T I S R I S S S S S S S S S R S S S A A A S S S S

S (D) (S) S S S (S) (S) (S) (S) (S) S S (S) (V) S S S (S) S S (S) (D) (D) D V (S) S S S (S) (S) S S

M W R+W SB+W R R+M R+SB R SB+W+M R+M W+M M SB+W+M R SB+W+M SB+M R+W SB+W SB+W W+M M R W+M SB+M M R+M SB+M R+W R W R R+SB+M R W+M R+W+M

Nest No.

Nesting habitat

Diet data (n)

Feeding habitat

+ +

6, 3a, 10 6, 3a, 10

2 10 16

6, 3a 6, 3a, 10 6, 3a, 10

+ + +

6, 3a 6 6, 3a

6, 3a, 10, 2

6, 3a, 2

29 13 4

6, 3a, 10, 2 6, 2, 3a 6, 2, 3a

+ 14 +

6, 3a, 2 6, 3a, 2 6, 3a, 2

+ +

6, 3a 6, 3a

11, 10

1 6

11 11

14, 11

+ + + + + 38 +

11, 10 11, 10 12 12, 11 12 12, 11, 6 12, 11, 6

Roosting (n, Nmax )

Roosting habitat

1, 215

3, 300 4, 50 10, 320 5, 150

11 11 11 11

4, 40 4, 50

11 11

185

wethul14.tex; 7/02/1999; 21:01; p.17

Code

186 Appendix 2. Food items recorded in the diet of bird species at the Agmon wetlands (January 1996–February 1997). Bird species code from Appendix 1 (* – Predation attempt).

12, 5

12, 13

3a 1 SB+M R+W R+W R+W R+W SB+M W R+SB+W S (S) S V (S) S (S) V S S S R S I S Syrian Serin Greenfinch Goldfinch Linnet Rock Bunting Cretzschmar’s Bunting Reed bunting Corn Bunting Serinus syriacus Carduelis chloris Carduelis carduelis Carduelis cannabina Emberiza cia Emberiza caesia Emberiza schoeniclus Miliaria calandra 174 175 176 177 178 179 180 181

1 Extinct as resident since the early 1950’s (Yom-Tov and Mendelssohn, 1988), winters since 1982, breeds again since 1995 (Ashkenazi and Yom-Tov, 1997). 2 Decline in breeding populations in the Hula Valley since early 1990’s (Ashkenazi, unpublished reports 1991–1996). 3 Decline in wintering populations in the Hula Valley since early 1990’s (winter waterbird census 1990–1995, NRA unpublished reports). 4 Under reintroduction program at Hula Nature Reserve (NRA, unpublished reports).

11 3, 10

Diet data (n) Residence Status Europe Status Israel Common name Scientific name Code

Appendix 1. Continued.

Nest No.

Nesting habitat

Feeding habitat

Roosting (n, Nmax )

Roosting habitat

Food taxon

Bird species

Cyanophyta Microcystis sp. Lyngbia sp. Chlorophyta Filamentous green algae Rhizoclonium sp. Oedogonium sp. Spirogyra sp. Ulothrix sp. Schizomeris sp. Cladophora sp. Enteromorpha sp. Mougeotia sp. Bacillariophyta: Diatoms Angiospermae Fragments of plants Ceratophyllum demersum Najas delilei Potamogeton spp. Potamogeton nodosus Potamogeton berchtoldii Cyperus papyrus Typha domingensis Cynodon dactylon Cultivated crops Triticum sp. Alopecurus myosuroides Zea mays Medicago sativa Arachis hypogaea Helianthus annuus Ficus carica Morus nigra Unidentified seeds Arthropoda Crustacea: Cladocera Chydorus sphaericus Bosmina longirostris Daphnia spp. Crustacea: Cyclopoidea Crustacea: Ostracoda Crustacea: Isopoda Porcellionidae Arachnoidea Aranea: Lycosidae Aranea: Philodromidae Acari: Metastigmata Insecta

27, 64, 66 66 26, 27, 30, 31, 64, 66 27, 64, 66 64, 66 27, 64 66 66 66 66 66 64, 66 19, 24, 27, 30, 64, 66 64, 66 27, 31, 64, 66 31, 64, 66 64, 66 64 66 26, 27, 64, 65, 66 64, 66 27, 64, 66, 67, 120, 169, 170 64, 66 27, 75, 121, 168, 169 27 27, 58, 66, 92, 93, 120, 167 27, 120, 134 135, 162, 165 135, 169 27, 30, 58, 66, 120, 169, 181 12 66 30 25, 30, 66 66 66 12, 75, 76, 127, 128, 129, 130, 134, 159, 167 12 12 12 12 11, 12, 27, 53, 61–63, 75, 104, 113, 120, 130, 131, 134, 135, 152

wethul14.tex; 7/02/1999; 21:01; p.18

187 Appendix 2. Continued.

Appendix 2. Continued.

Food taxon

Bird species

Food taxon

Bird species

Ephemeroptera Odonata: Anisoptera Libellulidae Brachythemis leucosticta Crocothemis erythraea Odonata: Zygoptera Calopteryx syriaca Heteroptera Gerridae Dermaptera Dictioptera Mantodea Orthoptera Acrididae

62, 134, 152 131

Tabanidae Ephydridae Stratiomyidae Hymenoptera

19 27 27 1, 64, 69, 78, 89, 122–127, 129–131, 134 53, 164, 165 117 53, 122, 123, 124, 125

Acrida bicolor Calliptamus palestinensis Dociostaurus curvicerci Locusta migratoria (?) Oedipoda miniata Pezotettix judaica Pyrgomorphella granosa Tropidopola longicornis syriaca Tettigoniidae Festella festae Incertana incerta Platycleis sp. Tylopsis lilifolia Gryllidae Calliptamus palestinensis Gryllotalpidae Homoptera Aphidoidea Neuroptera Coleoptera Scarabaeidae Lepidoptera Vanessa cardui Unidentified caterpillar Diptera Chironomidae

Glyptotendipes sp. Psychodidae Muscidae Musca domestica Syrphidae Eristalis sp.

117, 120, 131, 152 131 130, 131, 134 120, 130, 131, 134, 151 130 64 12 12 12 11, 12, 75, 120, 131, 152 11, 12, 18, 75, 117, 120, 131, 140, 152, 162, 165 12 12 12 12 12 12 12 12 12, 120, 151 12 12 12 12 12 12 12, 53, 75, 113, 120, 165 27, 61, 62, 63, 64 130, 139 12, 53, 75, 113, 119, 120, 134, 148, 152, 165 53, 113, 165 12, 62, 66, 117, 134, 139, 181 12, 75, 120 12, 120 12, 64, 104, 130 12, 19, 27, 61–66, 71, 75, 78, 89, 122–131, 133, 134, 137, 140, 145, 146, 148, 151–153, 158, 159, 169 19 27 19

Vespa orientalis Apis sp. Formicidae Mollusca Gastropoda Melanoides tuberculata Melanopsis praemorsa Pisces Tilapia zillii Tilapia (red hybrid) Gambusia affinis Clarias gariepinus Cyprinus carpio Oncorhynchus mykiss Amphibia Unidentified adults Unidentified tadpoles Bufo viridis Rana ridibunda Reptilia Mauremys caspica Coluber jugularis asianus Aves Phalacrocorax carbo Bubulcus ibis Anas platyrhynchos Fulica atra Galerida cristata Hirundo rustica Motacilla alba Sturnus vulgaris Passer domesticus Mammalia Insectivora Crocidura sp. Crocidura leucodon Crocidura russula Rodentia Meriones tristrami Rattus sp. Mus musculus Microtus socialis Myocastor coypus

19, 64(?) 19 66 (fragments) 5–7, 9–11, 13–16, 39, 51, 66, 90, 92, 94, 113–115 10, 11, 13, 16, 66, 113, 114, 115 10, 13, 14, 51, 115 114 15, 16, 39 10, 13, 115 10 10, 11, 13, 16, 18 12, 13, 19, 88, 134, 145 10, 12, 16, 19 12, 13, 16, 113 113 18 37, 39, 105 39 37, 39 39, 49, unidentified Eagle 39, 49 36*, 39, 41, 56* 39 41* 105 105

53 53, 105 105 12, 53, 105 105 105, 113 12, 13, 105 10–16, 18, 36, 39, 40–42, 53, 105, 108, 164 Unidentified Eagle

wethul14.tex; 7/02/1999; 21:01; p.19

wethul14.tex; 7/02/1999; 21:01; p.20