foraging, nesting, and roosting habitats of the avian fauna of the agmonpdf

Wetlands Ecology and Management6: 169–187, 1998.© 1998Kluwer Academic Publishers. Printed in the Netherlands.

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

S. Ashkenazi1 & Ch. Dimentman21Ecology and Nature Conservation, P.O. Box 1057, Rosh Pinna 12100, Israel;2Department of Ecology, Systematicand 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 surroundingcultivated 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 cattailstand in the southern third of the lake. The foraging habitat and diet data of 97 avian species were determined. Themost intensively used foraging habitats were cultivated fields, lake shore reed-beds, shallow canals and the cattailstand. Forty six species nested in the wetland during March–October, with 2,040 colonial and solitary nests mostlyin the cattail stand, near-shore reed-beds and young trees. Roosts, also mostly in the cattail and reed-bed standsand in trees, were used by 55 species. Based on this survey of available habitats, we identified several key habitatsthat are either missing or require further development (e.g., temporarily inundated mud flats, reed-beds and bareislands). We also provide suggestions for improving the delicate balance between requirements of this developingwetland ecosystem and of the adjacent agricultural areas.

Introduction

Nearly 5 km2 in the Dead Sea Rift Valley (Afro-SyrianRift Series), northern Israel (35◦43′ E, 33◦03′ N),was allocated in 1994 to water recreation, wildlifetourism and agriculture. The newly created Agmonwetland is located in the area of the former LakeHula and swamps which was drained during the 1950s(Dimentman et al., 1992; Hambright and Zohary,1998).

The Agmon wetland consists of a variety of habi-tats, including a small shallow lake, a system ofshallow 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 Na-ture Reserve (HNR), Enan Stream Nature Reserve, thecommercial freshwater fish farms, local water reser-voirs and cultivated fields all within a relatively small

area of the Hula Valley, creates a mosaic of diversehabitats for birds. The unique assemblage of water-birds and raptors in this diverse area is very attractivefor birdwatching. However, the close proximity ofcultivated fields and fish farms requires a sensitivemanagement program that benefits both the attractedbird populations and the local farmers.

Suitable habitats for foraging, roosting and breed-ing determine the integrated value of a given area forsupporting a year-round avian community. Each ofthese functions may require different habitats whichmight be used independently in different areas. How-ever, the value of an area increases as it includeshabitats for more than one function, reducing energycost for daily movements among dispersed foraging,nesting and roosting habitats. In this study our goalwas to assess the integrated importance of habitatsavailable in the newly created wetland for its bird

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

Habitat Habitat type Description

1 Lake Agmon Shallow lake, 0.5–1 m mean depth

2 CattailsTypha domingensis Permanently inundated in the southern lake

3 Lake shores – riparian vegetation (a) Riparian: 12 km withPhragmites australis, Cyperus papyrus,

and low grass (b) T. domingensisandScirpus litoralis; low grass: 2.5 km with

Gramineae (Cynodon dactylon, Sorghum halepenseor sown and

harvestedTriticum sp.)

4 Islands and islets 2 islands planted (1994) with indigenous trees and 5 islets

5 Canals 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

6 Reed-beds ofP. australis 5 sites, NE – 18.6 ha, NW – 5.8, E – 0.2, W – 5.9, S – 1.4

7 Bare – mudflats (a) and steep banks (b) Temporally created in NW and SE along lake and island shores;

banks>1 m above water surface, 0.2 km

8 Periodic, mat-like floating vegetation Filamentous green algae,Potamogetonspp.,Najas delilei

and cattail

9 Temporary inundated areas Created after heavy rains and floods

10 Diverse pasture land Fenced, with 20 reed-fringed shallow ponds, (grazed by donkeys)

11 Planted trees Large and old – 15 ha (Eucalyptussp.,Populussp.,Salix

sp. andTamarixsp.) and 10 ha of young planted (1994)

indigenous trees

12 Cultivated fields Various crops

13 Roads – paved or dirt-roads Paved roads surrounding the lake (2.8 km), dirt-roads (23 km)

14 Manmade – nest sites Nesting boxes, floating platforms and concrete water passages

populations and to provide recommendations for man-agement and long-term monitoring of indicator andvulnerable species for assessing changes in habitats.

Materials and methods

The study was carried out during January 1996–February 1997 in the Agmon wetlands. We classifiedseveral habitats that seemed to be important for birdswithin the study area. Habitat areas were calculatedwith the aid of GIS from infra-red aerial photographof the area taken in August 1996.

Observations on foraging, nesting and roosting ac-tivities of the avian fauna were carried out in theAgmon wetland and adjacent areas (about 350 ha) atleast once a week during daytime and once a monthduring the night. Observation, assisted by binoculars,telescope and a spotlight (at night), lasted 3–10 hours,and covered the entire day-time period within eachmonth. The rest of the cultivated area was surveyedonly 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 populationsin the wetland. Environmental conditions, plant phe-nology, presence of mammalian predators, seasonalchanges in prey abundances and management activi-ties (fish stocking, mowing, harvesting and water levelfluctuations) were recorded.

Foraging and diet

We used a combination of direct observations, fae-ces, regurgitations and stomach content analysis forrecording bird diets. For birds with partial digestion(e.g., European Coot, Moorhen, and some ducks), wecollected fresh droppings from individuals exiting thewater onto shore and preserved them in 70% ethanol,4% formaldehyde or 1% Lugol’s solution. Food itemswere then analyzed under a binocular or compoundmicroscope following sequential filtration with dif-ferent mesh sizes. Food regurgitations of chicks ofcolonial waterbirds were collected from nests and theircontents identified. Pellets were collected from twopairs of breeding Barn Owl nest boxes, every 4–20

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Figure 1. Map of the Agmon wetland showing habitats and main nesting sites for birds during January 1996–February 1997. Refer to Table 1for the legend and to Figure 2 for the area of habitats. The main nesting sites are indicated by a circle with an “x” in it. The map was digitizedfrom an aerial infra-red photograph taken during August 1996.

days. Occasionally we collected pellets from Kestrelnests. Bones in pellets were bleached (4% hydro-gen peroxide) and identified using Mendelssohn andYom-Tov (1987) guide plates. Individuals of Mallard,Northern Shoveler and European Coot were collectedby shooting and their stomachs dissected and analyzedas were faeces. During direct observations of piscivo-rous species, we estimated fish lengths relative to billlength. The food items identified were recorded to thelowest taxonomic level possible.

Breeding

A species was recorded as breeding if a nest was foundor adults were observed feeding chicks. Nests weremarked with individual numbers on an aerial photo-graph. Dates of nest building, hatching and fledgingwere recorded. The number of colonial waterbird nestswere estimated by combining counts from observa-

tion tower, counts of nests in defined areas and directexamination of nests.

Roosting

Roost sites were identified by direct observations onknown roost sites and potential habitats. Roosting ob-servation efforts in January 1996 were partial. Anattempt was made to count all the birds that landedat the roost at twilight. After dark, it was difficultto count or distinguish between some species such asLittle Egret and Cattle Egret. To minimize mistakescaused by low visibility, counts of roosting birds weremade also after sunrise, when birds left the roost. Usu-ally only one roost site was observed at a time. Wetherefore lacked information on the synchronous useof several roosts. We also recorded roost sites whennot in use.

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Figure 2. Cumulative number of feeding (grey), breeding (black) and roosting (white) species using Agmon wetland habitats during January1996–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 sub-species) 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 va-grants (1.6%). Ten of these are species that ceasedbreeding after the Hula drainage in the late 1950s(Yom-Tov and Mendelssohn, 1988; Dimentman et al.,1992). Sixteen species observed in Agmon are endan-gered in either Israel or Europe or both and five areglobally 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 indi-cated that utilization of habitat for these functions isnot correlated with habitat size (Figure 2). The most-used habitats for all the functions were cattail stands,reed-beds, riparian habitats and trees, although theywere not the largest available habitats. The largesthabitat area, cultivated fields, ranked second amongmost-used habitat and was used mainly for foraging.

Seasonally-inundated habitats and mudflats were im-portant for ca. 20 species although they were availablefor very short periods during the year and were amongthe smallest-sized habitats. The relative absence ofthese shallow-water feeding habitat was demonstratedby 60 Avocets (Recurvirostra avosetta) which fed androosted in the HNR during winter 1996–1997, but didnot use Agmon for any of the activities. Moreover,the Glossy Ibis uses similar feeding habitat bred androosted 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 habi-tats greatly affected habitat use patterns. For example,beginning in late 1996, the cattail stand began deteri-orating such that by February 1997 few plants existed(see Kaplan et al., 1998). During this time, most colo-nial and solitary roosting and breeding species left thearea, with the exception of Cattle and Little Egrets thatremained faithful to the site throughout early 1997.Also mowing of reed-beds in 1997 greatly restrictedroosting habitat for some species. In January 1996, ca.70 Marsh Harriers roosted in the reed-beds, whereasin January 1997 the reed-beds were mowed and fewMarsh Harriers were observed roosting in Agmon.

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Figure 3. Foraging-species density (number of species foraging per ha) in Agmon habitats during January 1996–February 1997. Refer to Figure1 and Table 1 for habitat locations and descriptions.

Foraging and diet

Cultivated fields were used for foraging by the largestnumber of species, though densely vegetated lakeshores, shallow canals, the cattail stand, temporaryinundated areas, the lake, low grass lake shores, reed-beds, and mats of floating vegetation were all impor-tant feeding habitats (Figure 2). Density of foragingspecies in different habitats (Figure 3) indicated thatthe most densely-used foraging habitats were the lowgrass lake shore, mudflats, vegetated lake shore, tem-porary inundated areas and mats of floating vegetation(i.e., the habitats smallest in area).

Foraging habitats and diet data from direct obser-vations (1,791), regurgitations (61), stomach contents(11), faeces (48) or pellets (257) of 97 species wererecorded (Appendix 1, Appendix 2). The most di-verse diet was found in Cattle Egret, which foragesin cultivated fields and in shallow canal banks. Di-verse and seasonally changing diet was detected alsoin species which fed in the lake, on banks grass andin cultivated fields such as European Coot, Moorhenand Mallard. Other species were very restricted in diet,such as Purple Coot which consumed mainly cattailand Kingfisher and Pied Kingfisher which consumedexclusively fish.

Prey items recorded in diets of the largest numberof avian species in the Agmon wetland (Appendix 2)were Chironomidae (in the diet of 36 species), So-cial Vole (Microtus socialis) (17), Hymenoptera (15),fish (19), Amphibia (11), Acrididae (11), terrestrialisopods, Porcellionidae (10), butterflies, Lepidoptera(9), dragonflies and damselflies, Odonata (8), andmole crickets, Gryllotalpidae (6).

Seven fish-eating heron species (Purple Heron,Black-crowned Night Heron, Little Egret, SquaccoHeron, Grey Heron, Great White Egret and Little Bit-tern) fed in the area of the Agmon wetlands (Table 2).Fish mostly caught in canals, near the water inflow oroutflow and in the southeastern shallow lake.

In March, April, June, October and November1996, the lake was stocked with small (20–40 g) fish,mostly tilapia (Tilapia zillii ) and common carp (Cypri-nus carpio) (Degani et al., 1998). Mass migration ofthe Great White Pelican through Israel occurs duringApril and October (Shmueli, 1996). The earliest-arriving individuals of the wintering Great Cormorantwere observed in October. Stocking in March did notattract any piscivorous birds. In April, about 80 Peli-cans fed in the lake for about a week. In June, about50 Black-crowned Night Herons foraged in Agmonfor two days, but Pelicans roosting at the HNR werenot attracted. Stocking in October attracted 300–1,500Pelicans, dozens of herons and a few kingfishers andin November 300–500 Pelicans each time for a week.

Analyses of pellets from two Barn Owl nests inthe 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 numberof species (Figure 2) were generally the smaller habi-tat areas (cattail stand, densely vegetated lake shores,trees, reed-beds and the pasture land. Large size habi-tats such as cultivated fields, canals or the lake werenot favored by breeding species. The highest nestdensity was in steep banks, cattail stand, man-madenesting sites, mudflats and bare lake shores (Figure 5).

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Figure 4. Monthly patterns in breeding (solid bars) and roosting (open bars) species richness in the Agmon wetland during January1996–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 forhabitat 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 summerbreeders listed in Appendix 1. The rest were eitherrare breeders or species (such as raptors) with nest-ing habitat requirements which were absent in thearea. The breeding season lasted eight months (endof March to end of October) with a peak (41 and 42breeding species) in May and June, respectively (Fig-ure 4). Altogether, 2,040 nests were recorded (1,670in a mixed breeding colony and 370 solitary nests)(Appendix 1). Among the solitary nesters, CrestedLark, Spur-winged Lapwing, Moorhen and seven war-blers were the most abundant breeders. Some of thespecies, such as Spur-winged Lapwing, Crested Lark,Moorhen, White-breasted Kingfisher, Pied Kingfisher,

Common Bulbul, Fan-tailed Warbler, Striped-backedPrinia, Reed Warbler, Clamorous Great Reed Warbler,Woodchat Shrike and Yellow Wagtail, had 2–3 suc-cessful broods within the breeding season. Moorhenhad an unusually large breeding population in Ag-mon, compared with that in the HNR, possibly dueto the abundance in Agmon of the submerged macro-phyte Najas spp. (a major component of Moorhendiet). Seven species, Black-crowned Night Heron,Squacco Heron, Cattle Egret, Little Egret, PurpleHeron, Glossy Ibis and Pygmy Cormorant, nested ina mixed-breeding colony on the cattail stand in thesouthern end of Lake Agmon. The solitary heron, Lit-tle Bittern, nested in the fringes of the colony. Thebreeding colony was first established (six species, 500nests) in Lake Agmon in summer 1995 (Ashkenazi

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Table

2.F

ish

spec

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posi

tion

and

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1121

137

1836

1721

14

No.

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11(1

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21(1

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128

(93.

4)11

(61.

1)21

(58.

3)7

(41.

2)8

(38.

1)5

(35.

7)

Fis

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ngth

,cm

(mea

n±S

D)

3.5±

0.5

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0.6

4.3

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41+

1.8

5.35

+2.

114

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24.

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1.1

Fis

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ngth

rang

e,cm

(n)

3–4

(8)

2–4

(7)

2–10

(13)

1–15

(122

)3–

12(1

4)6–

20(5

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15(8

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6(5

)

Uni

dent

ified

fish

46

1720

1013

25

4

Tila

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33

788

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and T. Oron, unpublished data). In 1996 breedingactivity started in mid March, but the colony wasdeserted three times due to motor boat activity. Suc-cessful breeding began only in April when dense algalmats prevented boating in the vicinity of the cattailstand. Desertion of the breeding colony was followedby desertion of the communal roost site on the cattailstand. Solitary ground-nesting birds were also affectedfrom timing of human activity such as mowing andharvesting within the Agmon wetland. In 1996, twonests of Black-headed Yellow Wagtails, two nests ofSpur-winged Lapwings, five nests of Bee-eaters andtwo nests of Crested Larks were destroyed duringmowing, harvest or hay collection.

Roosting

The most-used roost habitats were the cattail stand,reed-beds, and trees. Mudflats, temporary inundatedareas and low grass lake shores were also favored habi-tats for roosting. Roads and irrigation systems wereused for roosting by several species. Roost sites wereused mainly in winter and during migration (springand autumn). Most of roost sites were not in use dur-ing the summer (breeding season). However, the mostused roost site, the cattail stand, was in use through-out the year by seven colonial nesting species, twooverwintering herons (Grey Heron and Great WhiteEgret) and several occasional roosting species such asswallows and wagtails.

Fifty-five species were recorded roosting in thestudy area, with each species observed on 1–37 occa-sions and many of which were roosting throughout theyear (Figure 4, Appendix 1). The number of speciesutilizing the Agmon wetland habitats at the same timefor nesting and roosting varied between 9–45. Thenumber of roosting species was higher in February1997 (33) compared to February 1996 (15). Numbersof individuals of a species at a roost varied from 2 to6,000 per night. The maximum number of roosting in-dividuals per species per night was> 100 (30 species),> 1,000 (9 species), and> 5,000 (2 species). Roost-joining behavior varied in different species. Somearrived to a nearby pre-roost before sunset, e.g., GreatWhite Egret and Stock Dove. Others joined the roost atdark, such as Mallard, Grey Heron and Marsh Harrier.Some arrived in large flocks, such as Cattle Egret, Lit-tle Egret, Glossy Ibis and Yellow Wagtail, while othersjoined the roost individually, e.g., Squacco and GreyHerons. In the communal roost site in the cattails, dif-ferent species had a tendency to roost in a different

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section of the colony. This pattern was consistent untilchanges in species composition (departure of summerbreeders such as Purple Heron and Squacco Heron)or changes in the habitat quality (deterioration of thecattail habitat or mowing of Common Reed stands)occurred.

Discussion

Proper management of an area for avian species re-quires information on habitat specifications for theimportant functions of their life history (Kushlan,1993; Erwin; 1996b, Hoffman et al., 1996). Althoughhabitats for basic functions such as feeding, nestingand roosting may be independent and available in dif-ferent places, there is an energetic advantage whenhabitats for several functions are proximal to eachother (Hafner and Fasola, 1992; Hoffman et al., 1996;Rehfisch et al., 1996). Integration of species habitatrequirements for foraging, breeding and roosting con-tributes to a better understanding of species resourcerequirements for planning a suitable management pro-gram on areas utilized on year-round basis. This is im-portant particularly in management of a multi-purposearea, when attraction of populations to certain habitats(and avoiding damage to other habitats) is essential forminimizing land-use conflicts.

We assume that the large number of species utiliz-ing the area of Agmon resulted from the high diversityof habitats with their diverse, abundant food resources.However, the number of species utilizing these habi-tats was not related to habitat area. Rafe et al. (1985)found that both area and habitat diversity influencespecies richness and the number of species is bet-ter correlated with habitat diversity than with area.Presence of several congeneric species indicates ahigh suitability of the habitat to characteristics of thegenus, and a richness and diversity which permit re-source partitioning of closely related species. This wasdemonstrated by several waterbirds including 6 ducks(Anasspp.), 6Tringa species and 3 crakes. Similarly,the Agmon wetland supports species having a specialvalue in agricultural pest control, including 4 shrikes,4 wagtails, 4 pipits, 5 falcons, 4 harriers and 4 eagles(Aquila spp.). Another correlate of well-establishedcommunities and diverse habitats is the complexityof the trophic levels in the ecosystem. The Agmonwetland system harbors a rich ensemble of speciesfrom herbivores such as Goose and Ruddy Shelduck,

to secondary or tertiary predators such as White-tailedEagle.

The value of the diverse habitats of Agmon for itsbird assemblage was primarily for foraging (by highestnumber of species), but also for year-round roosting.Nesting habitats were utilized by the lowest numberof species and for limited time during the year. How-ever, nesting habitats for the endangered Marbled Teal,Black-headed Yellow Wagtail and Pratincole are avail-able in Agmon. The deterioration of the most heavilyused habitat for nesting and roosting, the cattail stand(Kaplan et al., 1998), in late 1996 will probably resultin a decrease in breeding and roosting species richness.Although the site was very attractive for tourism, interms of nature conservation, these colonial waterbirdshave an alternative for both activities in the nearbyHNR. However, habitats for the above mentioned en-dangered breeding species are rare or absent in theHNR and their maintenance in Agmon wetland maycontribute to regional breeding species richness andglobal conservation of these species.

Mudflats and temporary inundated habitats wereamong the smallest and short-lived (mostly 2–7 days)habitats. However, when available they were utilizedby waders and ducks both for feeding and roosting.Temporarily inundated habitats, usually created inwinter, are attractive to breeding amphibians. Tadpoleswere very attractive prey for several species, includ-ing herons, at the end of the winter. Shallow habitatsare essential for waders which forage largely in waterless then 18 cm deep (Rehfisch, 1994; Erwin, 1996b).Food and water conditions in such habitats are idealfor Mallards and Pintails which can swim but still dab-ble and tip up for food (Weller, 1990). We assumethat enlargement of these habitats and increasing oftheir duration will increase the number of species andindividuals which utilize them.

Characteristics of diets of selected Agmon avianspecies

A main feature that characterized the habitats and dietsof many species in the Agmon wetland was the useof both wetland habitats and adjacent cultivated fields.The importance of cultivated field habitat for forag-ing emphasizes the need for an effective managementprogram to avoid impact on crops while providing“biological pest control services”. Cultivated fieldsprovided rich and accessible insect and rodent faunafor species such as Cattle Egret, White Stork and sev-eral raptors. These fields had a large population of

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social voles with a long history of serious damage tocrops, resulting in heavy use of rodenticides (Yom-Tov and Mendelssohn, 1988). Mass consumption ofsocial voles (one of the most important prey items forbirds) in Agmon during the whole year may reducevole damage to crops.

Changes in food availability in the aquatic envi-ronment were reflected in the foraging activities anddiets of European Coot, Moorhen and dabbling ducks.In January and early February 1996, there were nodabbling ducks in Lake Agmon while thousands ofthem fed in the HNR lake. Samples of zooplanktoncollected in both lakes indicated high winter densitiesof zooplankton in the HNR lake compared with inLake Agmon. Similarly, an increase in the zooplank-ton densities in Lake Agmon during winter 1997 ledto an increase in dabbling duck abundances.

Fish in the Agmon wetlands have two distinctivegroups which differ in availability: (1) The group ofspontaneous colonizing indigenous fish, entered themain waterbody and the canals with the inflow water,in a continuous colonizing process; (2) Periodicallystocked (from commercial freshwater ponds) fish fry,characterized by a few unpredictable pulses of largenumbers at a time. Piscivorous species were observedeating fish in large groups only: (1) after stocking ofthe lake with fish; (2) during periods of low oxygenconcentrations in the water in summer (mostly earlyin 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 shal-low ponds following decreases in water level. In allfour cases, fish densities were locally increased andthe fish were more vulnerable to predation. The roleof stocked fish for attracting large piscivorous birdsfor wildlife tourism is not clear. It was impossible toassess the impact of piscivorous birds due to lack ofinformation on: (1) rates of natural fish immigrationand emigration; (2) reproduction rates of fish withinthe lake; (3) the amount of fish removed by angling.Stocked fish were noticeably attractive only during thepeak of the pelican migration (April and October), fora maximum of one week each. Fish fry eaten by thesmall herons in the lake were much smaller than thestocked fish size and were probably from naturallyimmigrated or reproduced fish. Creation of stable andattractive fish foraging habitat requires management ofpredictable and large fish populations. However, densefish populations have a major impact on the distrib-ution and abundance of other aquatic organisms andmay cause changes to invertebrate fauna which affects

the non-piscivorous bird populations (Britton, 1982;Eriksson, 1987; Poysa et al., 1994). There is a need forfurther study for understanding the needed densities offish and zooplankton in the lake for maintaining bothpiscivorous and zooplanktivorous birds for wildlifetourism.

Nesting and roosting

The richness of breeding species, the composition ofthe mixed-breeding colony and the long breeding sea-son with multiple successful broods in some species,indicates that the Agmon wetland and surroundingsare very productive habitats. Moreover, the establish-ment of a new heron, cormorant and ibis breedingcolony at the Agmon wetland, two km from the tra-ditional colony at the HNR, emphasizes the resourcerichness of the area for these species. All colonialbreeding species in Agmon nested on the cattail stand.Most of solitary species nests were in the riparianhabitats along the lake, the diverse pasture land, baremoderate or steep banks and trees. Nest densities inthese small area habitats, in particular in the banksand riparian lake shores were the highest. Similarly,breeding bird densities in the United States are amongthe highest in riparian habitats (Carothers et al., 1974).

Very little research has been devoted to under-standing how roost availability contributes to long-term stability of avian populations (Draulans and VanVessem, 1986; Rehfisch et al., 1996). There is limitedinformation on the importance of factors determiningroost-site selection such as energy limitations, preda-tion, disease and weather conditions (Birkhead, 1973;Blanco, 1996; Rehfisch et al., 1996). Roost sitesare particularly important for migrating or winteringpopulations like harriers, wagtails, swallows and var-ious waders (Mitchell et al., 1988; Rehfisch et al.,1996). Roosting species richness was highest duringthe spring (March–April) and autumn (September–November). However, unusual amounts of floodedhabitats in winter 1996–1997, in particular during Feb-ruary 1997, attracted a peak number of 33 roostingspecies. Flooded habitats were created on this oc-casion on the large islands, attracting foraging androosting dabbling ducks, some rare waders and rap-tors. The number of roosting species during the breed-ing season was low due to dispersal of the solitaryspecies for cryptic breeding activities. Eight of thecolonial-roosting species (seven breeding species andthe wintering Great White Egret) were faithful to asingle roosting site (cattail stand). The only excep-

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tion was the Grey Heron which roosted alternately intwo different habitats (cattail and trees). Glossy Ibisbred and roosted in 1996 only in Agmon (except fora single pair that nested in the HNR) although theybred and roosted in the HNR since the 1970s. Weare uncertain whether there is an exchange betweenpopulations of some colonial roosting species withthe HNR. However, breeding and roosting activitiesin the colony of the HNR were terminated in earlyAugust 1996 (T. Oron, pers. comm.). We assume thatthe increase in colonial roosting birds in Agmon from6,000 in July to 10,000 in August was due to move-ment of the HNR populations to Agmon. Roostingpopulations decreased after the summer breeders mi-grated (August–September) and Black-crowned NightHerons shifted to night activity. The unusual doublefunction of the cattail stand, the influx of the entireHula Valley colonial waterbirds to the roost site in Au-gust, and the high site tenacity of the species, rendersthis habitat as critical for colonial waterbirds.

Management implications

The Agmon wetland area is a multi-purpose (recre-ation, wildlife tourism and agriculture) area that de-mands a very detailed, sensitive and dynamic manage-ment program to maintain the delicate balance amongits different components. Management of the avianpopulations in this diverse area has to consider theavailable habitats in the entire Hula Valley, includingthe HNR and the agricultural areas. The managementplans for the area should analyze allocation of habitatsfor species that are endangered, rare, or of conserva-tion interest within the HNR and “tourism species”within the Agmon wetland. “Tourism species” are usu-ally large (Pelican, Cormorants, Cranes), gregarious,easily observed (colonial waterbirds) or rare species(raptors, composed of 26 species in Agmon). How-ever, endangered or rare species that feed, breed orroost in the newly created habitat despite the pro-tected habitats of HNR, must receive special attentionin management of the multi-purpose Agmon wetland.

From a regional management perspective, and interms of habitat use by the avian fauna, the Agmonwetland and the HNR are a part of a large mosaicof habitats. The avian fauna in the region use bothsites either for foraging, nesting or roosting. Com-mon Crane and Great Cormorant populations roostedonly in the HNR, but foraged in surrounding cultivatedfields and commercial fish ponds, respectively. GlossyIbis foraged in the HNR, but nested and roosted only

in Agmon. Mallards use forage and roost sites eitherin the HNR or in Agmon in different periods. Themovements between the sites for different activities,is facilitated by the Jordan River eastern and west-ern canals which were the main flyways (connectingcorridors) used in particular by cranes, herons andducks after dark. Any disturbance in these flywaysto 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 hu-man disturbance to colonial breeding waterbirds inthe US (Erwin, 1989, 1996a): 100–200 m buffer dis-tance from wading birds nesting sites, 50–100 m fromroosting shorebirds and at least 400 m from nestingcormorants.

Shallow muddy foraging habitats were availableonly on a few occasions and in a very small area,when water level decreased. However, whenever cre-ated, mudflats immediately attracted a rich ensembleof foraging and roosting waders for a few days. Thefood availability of the large islands for most of thewaterbird species could be improved by creating shal-low impoundments to retain water more frequentlyand for longer periods. We recommend moderatingslopes along the islands shores to create mudflatsduring the entire year, but in particular during themigrating seasons of wading birds (March–April andSeptember–November). Most of the Lake Agmonshores were planned steeply, to preclude breeding ofmosquitoes, mainly malaria-vector species (Anophelesspp.). However, mosquito predators such as the indige-nous fishAphanius mento, the exotic Mosquitofish(Gambusia affinis), amphibians and several inverte-brates may eliminate the mosquito problem in suchhabitats. Neither of the two moderate-sized islandsis accessible to the public and would provide pro-tected habitat for species that require food character-ized by temporary habitats and mudflats. Drying-outperiods of certain parts of the mudflats (out of themigrating season) are recommended. This will cre-ate food resources characterizing “disturbed” habitatssuch as chironomid-dominated early-colonizer com-munity (Rehfisch, 1994) which are important foodsources during the breeding season (Cox and Kadlec,1995). We suggest plowing the elevated parts of thelarge islands and dry pasture land in the end of March,to create nesting and roosting habitat for the endan-gered Pratincole. A series of small vegetated islets inthe lake’s center are not used by birds. We suggest

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de-vegetating them by mowing followed by thermaltreatment (black polyethylene) to create bare habitatsfor roosting and ground-breeding colonial species. Werecommend to cover the surface of some of these isletswith pebble, to create nesting habitats for terns andplovers that have very limited suitable habitat in theHula Valley.

Lake water levels should be managed very care-fully to create rich seasonal food webs and availableforaging and roosting habitats. It is suggested to mimicthe natural water level fluctuations in the former LakeHula: high water levels in winter, with a maximumin March, lowering water level gradually throughoutthe summer and increasing again beginning in Novem-ber, after the fall migration. Increases in water levelshould be avoided during April–July to avoid floodingof ground nesting bird nests.

Reed stands are among the most important habi-tats for breeding and roosting. We suggest that at least6–8 ha of reed-beds should be protected from mow-ing to provide an area suitable for roosting harriers.This habitat may partially substitute for some of thefunctions provided by the deteriorated cattail. We ad-vise vegetation be cut around the waterbody beforethe breeding season (mid-to-late March) and at theend of the nesting season (October). Also, it is rec-ommended that harvested vegetation be removed fromthe fields within a week, to avoid ground nest destruc-tion. The late mowing and silage making contributedin Europe to a decline in species such as Lapwing,Meadow Pipit and Yellow Wagtail (Anderson, 1995).Mowing of lake shores in winter is not recommendedas it eliminates protected habitats and roosting sites forwaterbirds. Also, it exposes the emergent vegetation tocold, stressful conditions. Some parts of this riparianhabitat should not be mowed at all. On the other hand,mowing some areas (e.g., fallow agricultural fieldssurrounding the lake) in the fall will increase exposureof social voles to predation by migrating and winter-ing raptors, possibly decreasing the vole’s impact onwinter crops.

Acknowledgments

We thank E. Yas’ur for providing facilities and helpwith sampling habitats. G. Gississ, T. Oron, Y. Vaadia,O. Rabinovich and K. Merom provided informationon 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. Kad-mon helped with the GIS produced map. M. Erwin,Y. Yom-Tov and an anonymous reviewer providedvaluable comments on the manuscript. The studywas funded during January–December 1996 by KerenKayemet L’Israel (JNF), Israel Land DevelopmentAuthority and the Israel Ministry of Agriculture.

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Appendix 1.The conservation status and habitat utilization (January 1996–February 1997) of wetland bird species in the Agmon wetlands. Conservation status inIsrael 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), – globalconcern. 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).

Code Scientific name Common name Status Status Residence Nest Nesting Diet Feeding Roosting Roosting

Israel Europe No. habitat data (n) habitat (n, Nmax ) habitat

1 Tachybaptus ruficollis Little Grebe V S R+W 1 2 + 1 8, 96 2

2 Podiceps cristatus Great Crested Grebe V, X-br S W **

3 Podiceps grisegena Red-necked Grebe R S O

4 Podiceps nigricollis Black-necked Grebe R S W

5 Phalacrocorax carbo Great Cormorant S S W 16 1, 5

6 Phalacrocorax pygmeus Pygmy Cormorant (X)1, E V R+SB 10 2 25, 56 2

7 Pelecanus onocrotalus Great White Pelican S R W+M+R(nB) 26 1, 5 11, 1200 2, 7a

8 Botaurus stellaris Bittern V (V) W+M 11, 2 6, 2

9 Ixobrychus minutus Little Bittern E (V) SB 12 2, 3a, 5 8 5, 2, 3a

10 Nycticorax nycticorax Black-crowned Night-Heron S D R+SB 700 2 17 2, 3a-b, 5, 12 21, 300 2

11 Ardeola ralloides Squacco Heron V V SB 45 2 35 5, 3a-b, 12, 2 13, 70 2

12 Bubulcus ibis Cattle Egret A S R 700 2 113 12, 5, 3b 32, 5000 2

13 Egretta garzetta Little Egret S2 S R+SB 150 2 94 5, 3a, 2 37, 414 2

14 Egretta alba Great White Egret S S W 18 12, 5, 2 33, 1088 2

15 Ardea cinerea Grey Heron S, X-br S W ** + 12, 5, 2 27, 20 2, 11

16 Ardea purpurea Purple Heron V V SB 40 2 21 12, 5 13, 21 2

17 Ciconia nigra Black Stork S R W+M

18 Ciconia ciconia White Stork S V rSB+M + 12 12, 200 11, 14

19 Plegadis falcinellus Glossy Ibis V D R+SB 22 2 10 9 25, 185 2

20 Platalea leucorodia Spoonbill V E W

21 Anser albifrons White-Fronted Goose R S W

22 Anser anser Grey-lag Goose R S W

23 Tadorna ferruginea Ruddy Shelduck R V W

24 Anas penelope European Wigeon R S W 7 3b, 12, 2, 9 7, 180 3b, 13

25 Anas strepera Gadwall R V W + 1, 9 3, 29 1

26 Anas crecca Teal S3 S W + 2, 9, 1 6, 30 2

27 Anas platyrhynchos Mallard S S R+SB 10 2, 3a 121 12, 2, 1, 8, 9 33, 3800 2, 3b, 5

28 Anas acuta Pintail S3 V W 1, 10 1

29 Anas querquedula Garganey S3 V W + 9

30 Anas clypeata Northern Shoveler S S W 26 1, 9 2, 110 2

31 Marmaronetta angustirostris Marbled Teal E E (#) R 4 2, 3a + 2, 9, 8, 5 5, 6 2

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Appendix 1.Continued.



32 Aythya ferina Pochard S3 S W 3, 150 2

33 Aythya nyroca Ferruginous Duck S3, X-br V (#) W **

34 Aythya fuligula Tufted Duck S3 S W

35 Pernis apivorus Honey Buzzard S S M

36 Milvus migrans Black Kite S V rSB+W + 12, 3b 11, 70 11, 14

37 Haliaeetus albicilla White-tailed Eagle E4, X-br R R(nB)+M + 12, 3b

38 Circaetus gallicus Short-toed Eagle S R SB+M

39 Circus aeruginosus Marsh Harrier V, X-br S W+M ** 22 3a-b, 12, 2, 6 15, 68 6

40 Circus cyaneus Hen Harrier R V W+M + 12, 3a-b, 6 3, 5 6

41 Circus macrourus Pallid Harrier R E W+M + 12, 3a-b, 6 2, 7 6

42 Circus pygargus Montagu’s Harrier R S W+M + 12, 3a-b, 6

43 Accipiter gentilis Goshawk R S W+M

44 Buteo buteo Buzzard S S rSB+W+M

45 Buteo rufinus Long-legged Buzzard V (E) R+W+M

46 Aquila pomarina Lesser-spotted Eagle S R W+M

47 Aquila clanga Spotted Eagle S, X-br E (#) W+M

48 Aquila rapax Tawny Eagle S - O

49 Aquila heliaca Imperial Eagle R E (#) W+M 2, 2 11

50 Hieraaetus pennatus Booted Eagle R R W+M

51 Pandion haliaetus Osprey R R W+M + 1

52 Falco naumanni Lesser Kestrel E (V)(#) SB+W+M

53 Falco tinnunculus Kestrel S D R 2 14, 11 38 3b, 12

54 Falco columbarius Merlin E S W+M

55 Falco subbuteo Hobby I S SB+M

56 Falco peregrinus Peregrine R R W+M

57 Alectoris chukar Chukar S V R 1 10 3, 15 14

58 Francolinus francolinus Black Francolin V V R 8 6, 10 + 6

59 Coturnix coturnix Common Quail V V SB+M

60 Rallus aquaticus Water Rail R (S) rSB+W+M

61 Porzana porzana Spotted Crake R S W 13 2, 3a, 8

62 Porzana parva Little Crake R (S) W+M+S(nB) 12 2, 3a, 8

63 Porzana pusilla Baillon’s Crake R, X-br R W+M+S(nB) ** 16 2, 3a, 8

64 Gallinula chloropus Moorhen S S R 30 2, 3a, 5 570 2, 3b, 8, 5 11, 300 2, 5, 8

65 Porphyrio porphyrio Purple Coot R R W+M 18 2, 3a, 8

66 Fulica atra European Coot S S rSB+R(nB)+W 164 2, 1, 3b, 8, 5 21, 1200 2, 5, 8

67 Grus grus Common Crane S V W + 12

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68 Rostratula benghalensis Painted Snipe R - O

69 Himantopus himantopus Black-winged Stilt S S R+M 1 7a, 9 + 7a, 9 3, 24 7a, 9

70 Burhinus oeducnemus Stone Curlew S V R+M

71 Glareola pratincola Pratincole E E SB+M 3 10, 12 + air: 12, 1, 3a-b 3, 120 7a, 9

72 Charadrius dubius Little Ringed Plover I (S) SB+M+W

73 Charadrius hiaticula Ringed Plover S S W+M

74 Charadrius alexandrinus Kentish Plover E D R+W+M + 7a

75 Hoplopterus spinosus Spur-winged Lapwing S (E) R 48 7a, 10, 12, 13 35 12, 3b, 13 9, 300 7a, 9

76 Vanellus vanellus Lapwing S (S) W+M + 12, 9 4, 130 7a, 9

77 Calidris minuta Little Stint S (S) W+M

78 Philomachus pugnax Ruff S (S) W+M + 7a, 9 9, 30 7a, 9

79 Gallinago gallinago Common Snipe R (S) W+M + 7a, 9 5, 14 7a, 9

80 Gallinago media Great Snipe R (V) M

81 Limosa limosa Black-tailed Godwit S V W+M

82 Numenius arquata Eurasian Curlew R D W+M

83 Tringa erythropus Spotted Redshank S S W+M

84 Tringa totanus Redshank S D W+M

85 Tringa stagnatilis Marsh Sandpiper S (S) W+M

86 Tringa nebularia Greenshank S S W+M

87 Tringa ochropus Green Sandpiper S (S) W+M

88 Tringa glareola Wood Sandpiper S D W+M + 5, 7a

89 Actitis hypoleucos Common Sandpiper S S W+M + 3a, 5, 7a, 9, 8

90 Larus ichthyaetus Great Black-headed Gull S S W+M + 1

91 Larus minutus Little Gull R D W+M

92 Larus ridibundus Black-headed Gull A S W+M + 1

93 Larus cachinnans Yellow-legged Gull - (S) W+M + 1, 12

94 Sterna hirundo Common Tern S S SB 1 14 + 1

95 Sterna albifrons Little Tern E D SB+W+M

96 Chlidonias hybridus Whiskered Tern I, X-br D W+M **

97 Chlidonias niger Black Tern R, X-br D M+S(nB) ** 1, 4 2

98 Chlidonias leucopterus White-winged Black Tern S S W+M

99 Columba (livia X domestica) Rock Dove X Domestic Dove A S R

100 Columba oenas Stock Dove R S W+M + 12 4, 3000 11

101 Streptopelia decaocto Collared Dove S (S) R 4 11 + 12, 13 12, 1500 11

102 Streptopelia turtur Turtle Dove S D SB+M 4 11 + 12, 13

103 Streptopelia senegalensis Laughing Dove A (S) R 1 14

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104 Clamator glandarius Great Spotted Cuckoo V S SB+M 1 11* + 12, 11

105 Tyto alba Barn Owl S D R 2 14 322 12, 3b, 11

106 Otus scops Scops Owl V (D) SB+W+M

107 Asio otus Long-eared Owl I S R+W+M

108 Asio flammeus Short-eared Owl R (V) SB+W+M + 3b, 12

109 Caprimulgus europaeus Nightjar S (D) W+M

110 Apus apus Swift I S SB+W+M + air: 12, 1

111 Apus melba Alpine Swift I (S) SB+W+M

112 Apus affinis Little Swift S (S) R

113 Halcyon smyrnensis White-breasted Kingfisher S (S) R 7 7b 14 5, 3a, 2, 10, 11

114 Alcedo atthis Kingfisher S D W+M 11 5, 3a

115 Ceryle rudis Pied Kingfisher S (S) R 5 7b 21 5, 3a

116 Merops superciliosus Blue-cheeked Bee-eater I (S) SB+M

117 Merops apiaster Bee-eater S (D) SB 8 7b + air: 12, 3a–b

118 Coracias garrulus Roller V (D) SB

119 Upupa epops Hoopoe S S R+M + 12, 11

120 Galerida cristata Crested Lark S (D) R 90 3b, 7a, 10, 12 41 12, 3b, 13 13, 150 13, 3b

121 Alauda arvensis Skylark S V W + 12, 3b, 13 14, 100 13, 3b

122 Riparia riparia Sand Martin S D W+M 26 air: 2, 1, 12 3, 1000 2, 6

123 Hirundo rustica Swallow S D R+M 58 air: 2, 1, 12 11, 6000 2, 6

124 Hirundo daurica Red-rumped Swallow S S SB+W 3 14 7 air: 2, 1, 12 3, 1000 2, 6

125 Delichon urbica House Martin V S SB+W 14 air: 2, 1, 12 3, 1000 2, 6

126 Anthus campestris Tawny Pipit I V SB+W + 5, 9

127 Anthus pratensis Meadow Pipit S S W+M + 12, 5, 9, 8

128 Anthus cervinus Red-throated Pipit S (S) W+M + 12, 5, 9

129 Anthus spinoletta Water Pipit R S W+M + 5, 9, 8

130 Motacilla flava Yellow Wagtail S S SB+W+M 4 7a, 3a 46 12, 5, 7a, 8, 9 9, 300 2, 6

131 Motacilla flava feldegg Black-headed Yellow Wagtail E - SB+M 8 7a, 3a 44 12, 5, 7a, 8, 9 6, 20 2, 6

132 Motacilla citreola Yellow-headed Wagtail R (S) W+M

133 Motacilla cinerea Grey Wagtail S (S) W+M

134 Motacilla alba White Wagtail S S SB+W+M 113 12, 5, 7a, 8, 9 7, 200 2, 6

135 Pycnonotus barbatus Common Bulbul A - R 3 11 + 11, 10

136 Erithacus rubecula Robin S S W+M

137 Luscinia svevica Bluethroat T S W+M + 5, 8

138 Phoenicurus ochruros Black Redstart S S SB+W+M

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HannaWinner

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18

5




139 Saxicola rubetra Whinchat S S M + 6, 3a, 10

140 Saxicola torquata Stonechat S (D) W + 6, 3a, 10

141 Oenanthe isabellina Isabelline Wheatear S (S) R+W 1, 215 13

142 Oenanthe oenanthe Wheatear S S SB+W

143 Turdus merula Turdus Merula S S R

144 Cettia cetti Cetti’s Warbler S S R+M 2 6, 3a + 6, 3a

145 Cisticola juncidis Fan-tailed Warbler V (S) R+SB 10 6, 3a, 10 + 6

146 Prinia gracilis Striped-backed Prinia S (S) R 16 6, 3a, 10 + 6, 3a

147 Locustella luscinioides Savi’s Warbler T (S) SB+W+M

148 Acrocephalus melanopogon Moustached Warbler V (S) R+M 7 6, 3a, 10, 2 + 6, 3a, 2

149 Acrocephalus schoenobaenusSedge Warbler T (S) W+M

150 Acrocephalus palustris Marsh Warbler I S M

151 Acrocephalus scirpaceus Reed Warbler S S SB+W+M 29 6, 3a, 10, 2 + 6, 3a, 2

152 Acrocephalus stentoreus Clamorous Great Reed Warbler R - R 13 6, 2, 3a 14 6, 3a, 2

153 Acrocephalus arundinaceus Great Reed Warbler I (S) SB+W+M 4 6, 2, 3a + 6, 3a, 2

154 Hippolais pallida Hippolais pallida S (V) SB+M

155 Sylvia melanocephala Sardinian Warbler S S R+W

156 Sylvia curruca Lesser Whitethroat S S SB+W

157 Sylvia atricapilla Blackcap S S SB+W

158 Phylloscopus collybita Chiffchaff S (S) W+M + 6, 3a

159 Phylloscopus trochilus Willow Warbler S S M + 6, 3a

160 Parus major Great Tit S S R

161 Remiz pendulinus Penduline Tit S (S) W+M

162 Lanius collurio Red-backed Shrike S (D) SB+M 1 11 + 11, 10

163 Lanius minor Lesser Grey Shrike R (D) M

164 Lanius excubitor Great Grey Shrike S D R+M 1 11 + 11, 10

165 Lanius senator Woodchat Shrike S V SB+M 6 11 + 11, 10

166 Corvus monedula Jackdaw S (S) R+W + 12 3, 300 11

167 Corvus corone Carrion Crow A S R 4 11 + 12, 11 4, 50 11

168 Sturnus vulgaris Starling A S W + 12 10, 320 11

169 Passer domesticus House Sparrow A S R 18 14, 11 38 12, 11, 6 5, 150 11

170 Passer hispaniolensis Spanish Sparrow S (S) R+SB+M + 12, 11, 6

171 Passer moabiticus Dead-sea Sparrow S (S) R 1 11

172 Fringilla coelebs Chaffinch S S W+M 4, 40 11

173 Serinus serinus Serin S S R+W+M 4, 50 11

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186

Appendix

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ontin

ued.

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cien

tific

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onna

me

Sta

tus

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ting

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g

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urop

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ta(n

)ha

bita

t(n

,N max)

habi

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174

Serinus

syriacu

sS

yria

nS

erin

V-

SB

+M

3,10

11

175

Car

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isch

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Gre

enfin

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176

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177

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uelis

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178

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ber

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cia

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179

Em

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180

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ing

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W

181

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aria

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Cor

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(S)

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+W

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eth

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rly19

50’s

(Yom

-Tov

and

Men

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,198

8),w

inte

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nce

1982

,bre

eds

agai

nsi

nce

1995

(Ash

kena

zian

dY

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ov,1

997

).2D

eclin

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bree

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the

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port

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996)

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ley

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ter

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1990

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ublis

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.

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

Food taxon Bird species

Cyanophyta

Microcystissp. 27, 64, 66

Lyngbiasp. 66

Chlorophyta

Filamentous green algae 26, 27, 30, 31, 64, 66

Rhizocloniumsp. 27, 64, 66

Oedogoniumsp. 64, 66

Spirogyrasp. 27, 64

Ulothrix sp. 66

Schizomerissp. 66

Cladophorasp. 66

Enteromorphasp. 66

Mougeotiasp. 66

Bacillariophyta: Diatoms 64, 66

Angiospermae

Fragments of plants 19, 24, 27, 30, 64, 66

Ceratophyllum demersum 64, 66

Najas delilei 27, 31, 64, 66

Potamogetonspp. 31, 64, 66

Potamogeton nodosus 64, 66

Potamogeton berchtoldii 64

Cyperus papyrus 66

Typha domingensis 26, 27, 64, 65, 66

Cynodon dactylon 64, 66

Cultivated crops

Triticum sp. 27, 64, 66, 67, 120, 169, 170

Alopecurus myosuroides 64, 66

Zea mays 27, 75, 121, 168, 169

Medicago sativa 27

Arachis hypogaea 27, 58, 66, 92, 93, 120, 167

Helianthus annuus 27, 120, 134

Ficus carica 135, 162, 165

Morus nigra 135, 169

Unidentified seeds 27, 30, 58, 66, 120, 169, 181

Arthropoda 12

Crustacea: Cladocera

Chydorus sphaericus 66

Bosmina longirostris 30

Daphniaspp. 25, 30, 66

Crustacea: Cyclopoidea 66

Crustacea: Ostracoda 66

Crustacea: Isopoda

Porcellionidae 12, 75, 76, 127, 128, 129, 130,

134, 159, 167

Arachnoidea 12

Aranea: Lycosidae 12

Aranea: Philodromidae 12

Acari: Metastigmata 12

Insecta 11, 12, 27, 53, 61–63, 75, 104,

113, 120, 130, 131, 134, 135, 152

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Ephemeroptera 62, 134, 152

Odonata: Anisoptera 131

Libellulidae

Brachythemis leucosticta 117, 120, 131, 152

Crocothemis erythraea 131

Odonata:Zygoptera 130, 131, 134

Calopteryx syriaca 120, 130, 131, 134, 151

Heteroptera 130

Gerridae 64

Dermaptera 12

Dictioptera 12

Mantodea 12

Orthoptera 11, 12, 75, 120, 131, 152

Acrididae 11, 12, 18, 75, 117, 120, 131,

140, 152, 162, 165

Acrida bicolor 12

Calliptamus palestinensis 12

Dociostaurus curvicerci 12

Locusta migratoria(?) 12

Oedipoda miniata 12

Pezotettix judaica 12

Pyrgomorphella granosa 12

Tropidopola longicornis

syriaca 12

Tettigoniidae 12, 120, 151

Festella festae 12

Incertana incerta 12

Platycleissp. 12

Tylopsis lilifolia 12

Gryllidae 12

Calliptamus palestinensis 12

Gryllotalpidae 12, 53, 75, 113, 120, 165

Homoptera

Aphidoidea 27, 61, 62, 63, 64

Neuroptera 130, 139

Coleoptera 12, 53, 75, 113, 119, 120, 134,

148, 152, 165

Scarabaeidae 53, 113, 165

Lepidoptera 12, 62, 66, 117, 134, 139, 181

Vanessa cardui 12, 75, 120

Unidentified caterpillar 12, 120

Diptera 12, 64, 104, 130

Chironomidae 12, 19, 27, 61–66, 71, 75, 78,

89, 122–131, 133, 134, 137, 140,

145, 146, 148, 151–153, 158,

159, 169

Glyptotendipessp. 19

Psychodidae 27

Muscidae

Musca domestica 19

Syrphidae

Eristalis sp. 19



Tabanidae 19

Ephydridae 27

Stratiomyidae 27

Hymenoptera 1, 64, 69, 78, 89, 122–127,

129–131, 134

Vespa orientalis 53, 164, 165

Apissp. 117

Formicidae 53, 122, 123, 124, 125

Mollusca

Gastropoda 19, 64(?)

Melanoides tuberculata 19

Melanopsis praemorsa 66 (fragments)

Pisces 5–7, 9–11, 13–16, 39, 51, 66, 90, 92,

94, 113–115

Tilapia zillii 10, 11, 13, 16, 66, 113, 114, 115

Tilapia (red hybrid) 10, 13, 14, 51, 115

Gambusia affinis 114

Clarias gariepinus 15, 16, 39

Cyprinus carpio 10, 13, 115

Oncorhynchus mykiss 10

Amphibia

Unidentified adults 10, 11, 13, 16, 18

Unidentified tadpoles 12, 13, 19, 88, 134, 145

Bufo viridis 10, 12, 16, 19

Rana ridibunda 12, 13, 16, 113

Reptilia

Mauremys caspica 113

Coluber jugularis asianus 18

Aves 37, 39, 105

Phalacrocorax carbo 39

Bubulcus ibis 37, 39

Anas platyrhynchos 39, 49, unidentified Eagle

Fulica atra 39, 49

Galerida cristata 36*, 39, 41, 56*

Hirundo rustica 39

Motacilla alba 41*

Sturnus vulgaris 105

Passer domesticus 105

Mammalia

Insectivora

Crocidurasp. 53

Crocidura leucodon 53, 105

Crocidura russula 105

Rodentia 12, 53, 105

Meriones tristrami 105

Rattussp. 105, 113

Mus musculus 12, 13, 105

Microtus socialis 10–16, 18, 36, 39, 40–42, 53, 105,

108, 164

Myocastor coypus Unidentified Eagle

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foraging, nesting, and roosting habitats of the avian fauna of the agmonpdf

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