SOME STUDIES ON BIO-ECOLOGY OF PHEASANT TAILED JACANA, HYDROPHASIANUS CHIRURGUS IN PUNJAB, PAKISTAN
ZAHID IQBAL KHAN 008-GCU-Ph.D-Z-2005
Department of Zoology Faculty of Sciences
GOVERNMENT COLLEGE, UNIVERSITY
Lahore, Pakistan. 2009
SOME STUDIES ON BIO-ECOLOGY OF PHEASANT TAILED JACANA, HYDROPHASIANUS CHIRURGUS IN PUNJAB, PAKISTAN
by
ZAHID IQBAL KHAN 008-GCU-Ph.D-Z-2005
A thesis submitted in partial fulfillment of the requirements for the degree of
DOCTOR OF PHILOSOPHY in
ZOOLOGY
Department of Zoology Faculty of Sciences
GOVERNMENT COLLEGE, UNIVERSITY
Lahore, Pakistan.
2009
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In the Name of Allah, The Most Compassionate, The Most Merciful
This dissertation is dedicated to the memories of my Beloved Parents
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DECLARATION
I, Mr. Zahid Iqbal Khan Roll No 008-GCU-Ph.D-Z-2005 student of department of
Zoology in the subject of Zoology session 2004-2009, hereby declares that the matter printed in the thesis tilted Some studies on Bio-Ecology of Pheasant Tailed Jacana, Hydrophasianus chirurgus in Punjab, Pakistan is my own work and has not been printed, published and submitted as research work, thesis or publication in any form in any university, Research institution etc in
Pakistan or abroad.
Dated: ______Zahid Iqbal Khan
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RESEARCH COMPLETION CERTIFICATE
Certified that the research work contented in this thesis titled “Bio-Ecology of Pheasant Tailed Jacana, Hydrophasianus chirurgus in Punjab, Pakistan has been carried out and completed by Mr. Zahid Iqbal Khan Roll No 008-GCU-Ph.D-Z-2005 under my supervision
Supervisor:______Prof. Dr. Muhammad Sharif Mughal
Co-Supervisor:______Prof. Dr. Afsar Mian
Chairperson :______Prof. Dr. Muhammad Sharif Mughal Department of Zoology GC University, Lahore
Controller of Examination :______GC University, Lahore
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LIST OF CONTENTS
TITLE Page No. List of table List of figures Acknowledgment Abstract Chapter 1: INTRODUCTION 1 1.1. RATIONALE OF STUDY 1 1.2. PHEASANT-TAILED JACANA 3 1.3. OBJECTIVES 7 Chapter 2: REVIEW OF LITERATURE 9 2.1. POPULATION DYNAMICS 9 2.1.1. Population distribution 9 2.2.2 Population size 9 2.2.3. Population Fluctuation 10 2.2.4. Sex ratio 11 2.2.5. Migration 11 2.2. HABITAT 12 2.2.1. General 12 2.2.2. Physical Factors 13 2.3. FOOD AND FEEDING 14 2.4. BREEDING 16 Courtship 16 Egg Laying 17 Nesting 17 Habitat 18 Parental care 18 Territoriality 19 2.5. GENERAL BEHAVIOUR 20 2.6. POPULATION STRESS 21 Chapter 3: MATERIAL AND METHODS 23 3.1 STUDY AREA 23 3.2 POPULATION BIOLOGY 29 3.3. HABITAT 30 3.3.1. Sampling 30 3.3.2. Habitat Analysis 30 3.4.Food 32 3.5. Breeding 33
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TITLE Page No. Chapter 4: RESULTS 35 4.1.POPULATION BIOLOGY 35 4.1.1.Distribution and Estimates 35 4.1.2.Population Fluctuation 35 4.1.2.1. Annual fluctuation 35 4.1.2.2. Seasonal fluctuation 38 4.1.3 Sex structure 42 4.1.4.Age Structure 43 4.2.HABITAT 44 4.2.1. Physical factors: 44 4.3. Food and Feeding 51 4.3.1 General food consumption 51 4.3.2. Feeding behavior 53 4.4. Breeding 55 4.4.1 General Behavior 58 4.4.2 Defensive Response 58 4.4.3 Nest Site Selections 58 4.4.4 Agonistic Behavior 59 4.4.5 Parental Care 59 Chapter 5: DISCUSSION 60 5.1. POPULATION STUDIES 60 5.2. HABITAT 63 5.3. FOOD 66 5.4. BREEDING 67 5.5. MANAGEMENT 68 6. LITERATURE CITED 69 7. ANNEXURE ANNEXURE 1, Mean ±standard error of mean of different physico-climatic 81 parameters of water bodies during different calendar months between April 2003 to March 2007 at Marala (Punjab, Pakistan).
ANNEXURE 2, Mean standard error of physico-climatic parameters of 82 water during April 2003 to March 2007 at Head Qadirabad.
ANNEXURE 3, Mean standard error of physico-climatic parameters of water during April 2003 to March 2007 at Head Balloki. 83 ANNEXURE 4. Mean standard error of physico-climatic parameters of 84 water during April 2003 to March 2007 at site 2 (Head Sulemanki)
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LIST OF TABLES TITLE Page No.
Table 3.1. Mean maximum (Max) and minimum (Min) temperature and rainfall 25 (mm) recorded at four study sites during the last ten years. Table 4.1 Population density of Pheasant-tailed Jacana in different water 36 bodies during different years 2004 – 2006 Table 4.2 Monthly Variation of Pheasant-tailed Jacana Population at study 39 sites in years 2004 – 2006 Table 4.3 Sex Ratio in as exhibited by pool 2004 – 2006 population of 42 Pheasant-tailed Jacana at four study sites Table .4.4 Juvenile adult ratio for total Pheasant-tailed Jacana at four sites 44 Table 4.5 The rooted vegetation population structure at study area. 48
Table 4.6: The natural organization of rooted vegetation observed at different 49 study sites. Table 4.7: Vegetation types on the basis of dominant plant species with their 50 own composition at each site. Table 4.8 Feeding preference of Pheasant-tailed Jacana observed at study 52 sites during summer months. Table. 4.9 Feeding behavior of different age groups of Pheasant-tailed 53 Jacana observed at study sites. Table 4.10 Food intake to body weight ratio at different life stages of 54 Pheasant-tailed Jacana in Punjab (Pakistan). Table 4.11 Time consumed for feeding and resting during early and late 55 breeding seasons by Pheasant-tailed Jacana in Punjab (Pakistan). Table 4.12 Overall Breeding period of Pheasant Tailed Jacana during 2004 to 56 2007. Table 4.13 The overall breeding activity budget of Pheasant tailed jacana 57 marked peak activity observed at four study sites.
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LIST OF FIGURES
TITLE Page No.
Figure 1.1 & 1.2: Map showing Location of the Punjab and rivers of Punjab 24 (Pakistan)
Figure 2.1. Map showing the location of site 1 (Head Marala) 27
Figure 2.2. Map showing the location of site 2 (Head Qadirabad) 28
Figure 2.3. Map showing the location of site 3 (Head Balloki) 28
Figure 2.4. Map showing the location of site 4 (Head Sulemanki) 29 Fig 3.1 Population size of Pheasant-tailed Jacana at different localities 40 (A=Marala, B=Qadirabad, C=Balloki, D=Suleimanki) during different calendar months of different years (2004-2006)
Fig 4.1 (a) Air temperature (C) recorded during different calender months 45 at different study sites during 2004-2007.
Fig 4.2. Water temperature (oC) recorded during different calendar months 46 at different study site during 2004-2007. Fig 4.3. pH of the surface water recorded during different calendar months 47 at different study site during 2004-2007.
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LIST OF PICTURES TITLE Page No.
A male pheasant tailed jacana 85
A female pheasant tailed jacana 85
Female pheasant tailed jacana in trapanut habitat 85 Male pheasant tailed jacana with chick 85 A male pheasant tailed jacana in Eichhornia habitat 85 A Female pheasant tailed jacana near bank 85
A male pheasant tailed jacana with chick 86
An alert male pheasant tailed jacana to protect chick 86
A clutch of 4 Eggs of pheasant tailed jacana 86
A male pheasant tailed jacana at incubation 86
A one day old chick of pheasant tailed jacana 86
Two days old chick of pheasant tailed jacana 86
Four days old chicks of pheasant tailed jacana 87
Six days old chicks of pheasant tailed jacana 87
A chick jumping into water to avoid danger 87
A chick with complete motionless during danger 87
A chick with fully drowned body except nostrils 87
A chick in disturb habitat 87
Pheasant tailed jacana (juvenile) 88
Pheasant tailed jacana (Feeding) 88
Pheasant tailed jacana (Sub- adult) 88
Pheasant tailed jacana (Food) 88
A chick watching a coming danger 88
A chick fully camouflage during danger 88
A clutch of four eggs in Echhornia leaves 89
An alert male pheasant tailed jacana approaching danger 89
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Habitat destructions by human beings. 89
Habitat destructions for growing trapanut 89
Habitat destructions by human beings. 89
Habitat destructions by human beings. 89
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ABSTRACT
Pheasant Tailed Jacana, Hydrophasianus chirurgus is only beautiful bird maintain its presence in abandoned wetlands of Punjab during summer season. This study was designed to study the population status and its future potentials to estimate the threats and to propose the measures for its conservation if required. Literature suggested sites were selected and schedule of visits month and year wise was developed and sites were visited accordingly, data gathered was analyzed statistically. The population distribution appears, between 298 and 347 highest (83-96) at Marala and lowest (21-57) at Balloki, while density found highest (3/km2) and lowest (0.3/ km2), being the migratory bird March was observed as start of arrival and October was observed as completion of departure from the study areas. Female to male ratio appears 1:0.75 while juvenile to adult ratio observed 1:0.35 are sign of population stability. Physical factors of the habitat such as temperature, pH and phytosociological studies of the habitat distribution and frequency of rooted vegetation species were recorded to understand their association with the species. This shows 32 species distributed in 4 vegetation types within the habitats lies in study areas. The studies on biology of this bird were focused on food and feeding, food consumption which revealed that 56.3% of its food is consists of plant matter and 27.3% on animal matter. Breeding biology studies suggests the clutch size remains between 1-4 eggs, and most of the fertile females laid 2 consecutive clutches to hand over two different males for hatching and chick rearing, incubation calculated of 25-29 days hatchability found around 80%. Defensive responses, egg laying and chick care were observed to understand general behavior of the species. The results of this study suggested that major threats to Pheasant Tailed Jacana are the commercial activities within its habitat and immediate adjacent to them, which could be regulated for its sustainability.
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INTRODUCTION
1.1. RATIONALE OF STUDY
Pheasant-tailed Jacana (Hydrophasianus chirurgus), also known as the Lotus Bird or Lily
Trotter, is endemic to the South Asia, with its range of distribution extending from
Pakistan, north Kashmir, India, Nepal, Sri Lanka, Burma to the mainland China through
Myanmar, Taiwan, Java and Philippines (Baker, 1929; Oliver, 1962; Grizmek, 1972;
Woodcook, 1980; Lal, 2004). The species has a wide distribution in large bodies of the
Indus valley in Pakistan (Hasan, 2001), where it is especially frequent in swamps of Nara area, Hadero Lake and Ghouspur and the rice growing areas of the Sindh, i.e., Hyderabad,
Larkana and Thatta, and the Punjab, i.e., Sheikhupura, Gujranwala and Sialkot. The scattered individuals have, however been spotted up to an altitude of some 2,300 m above sea line in Murree Hills, Azad Kashmir, Besham, Kohistan, Gilgit and Panji in the northwest and Band Khushdil Khan in southwest (Balochistan). This jacana species has a resident status in the Sindh but is a regular summer visitor in the Punjab (Roberts, 1991;
Mirza, 2007). The species appears in good numbers in the water bodies created by barraging of the rivers and impoundments associated with the head-works and barrages, including, Qadirabad, Marala, Suleimanki, Balloki, Rasool, Taunsa and Chashma, appearing along the rivers Satluj, Ravi and Chenab, along with small dams appearing at smaller rivers or hill torrents (Hassan, 2001).
The Pheasant-tailed Jacana, with its dark feathers and a yellow nape on the back, black hand around neck, arms, white wing tips walking/ elegantly on floating leaves, present and fascinating look. The bird species adds beauty to the wetlands of the Indus plains, attracting attention of any visitor to such wetland. It is also a part of the food chain of the wetland ecosystem and has important role in the maintenance of the wetland ecosystem. The species has adapted to very specific habitat conditions, i.e., slow moving or almost stagnant, permanent freshwater water bodies, shallow enough to hold rooted aquatic vegetation having floating broad leaves. In most parts jacana is found on artificial water bodies, which are often located very close to human habitation and are directly influenced by the human activities. The Pheasant-tailed Jacana is presently not under a serious hunting pressure from the local human population and hence the changes in the population of this species are largely being controlled by the habitat changes. Thus the species, with a narrow ecological amplitude has potentials of working as an effective ecological indicator for the changes occurring in the wetland habitat (Odum, 1971;
Ormerod and Tyler, 1993).
The shallow wetlands, adapted as habitat by the Pheasant-tailed Jacana, are generally extremely productive, and constitute a rich source of food for man, his livestock, and also for breeding, wintering and migrating birds. The aquatic ecosystem of such water bodies is usually efficient in bioremediation of the pollution and ensures the supply of water for the direct use of man and his livestock and supports the irrigation, either directly or indirectly by ground water recharging. Recent environmental changes, caused by increasing human population and increased human activities under changing human demands, the conditions of the wetlands are generally deteriorating, which is liable to affect the human life. Under such a situation, the species, like, Pheasant-tailed Jacana, having potentials of working as good ecological indicators attain an important role, so that the changes in the wetland are perceived well in time and the impact of the remedial safeguards is monitored (Bhushan et al., 1993).
The Pheasant-tailed Jacana, having a value as an efficient ecological indicator and adding charm to a wetland, is thus an important species in the biodiversity of the area, demanding its conservation. Population studies are not available to suggest the changing 2 pattern in the population of the species in the area, yet conservationists and wildlife enthusiasts generally feel an eminent decline in the population of this species during the recent past and is sometimes attributed an endangered status in the Punjab (Pakistan), and the species has been included in 3rd Schedule under the Provincial Wildlife Act 1974. The general biology of the species with regard to Pakistan is known from compilation of different casual reports and through reports appearing on the species from different parts of its distribution range (Roberts, 1991). This demands that base line studies on the biology of the species may be instituted so that the biological potentials of the species are understood and the factors affecting its population level are determined. Such studies can be useful in developing conservation strategy for the species, ensuring the pleasant look of our wetlands, which will also help in future prediction on adverse changes being experienced by the wetlands under the cover of floating vegetation and remaining unnoticed by human observer. The present research has been instituted for the study of the biology of the Pheasant-tailed Jacana (Hydrophasianus chirurgus) with reference to its population summering in different water bodies located in the Punjab (Pakistan).
1.2. PHEASANT-TAILED JACANA
The jacanas [locally known as Piho, Pihuya (Hindi), Gund Katya, Gair Kov (Kashmir),
Suidal Sakdal, Miwa, Dalkukra, Bhepi, Jal Mangoo, Jal Mayur, Jal Kokra, Chitra Billai
(Belgali, Bihari), Lobodak (Santhah), Rani didaogophila, Little White Water Princess
(Cachar), Yem Paraba, Cockfowl (Manipuor), Pan Kanbula, Ballal Deru, Vilginava
(Sinhala), Miwa, Monal Pura (Tamil), Tamara Kazhi (Malayalam)] are included in the bird order Charadriiformes and the family Jacanidae. The order Charadriiformes is characterized by having elongated toes, but hind toe is shorter;
1st and 4th primaries attenuated, the first into a barb-less shaft terminating in a spatulated web, the fourth prolonged into an attenuated point; wings furnished with a strong sharp 3 spur at the carpel joint and bent; and both sexes assuming a nuptial plumage and are alike, but females longer (Ali and Ripley, 1987). This is one of the large avian orders and exhibits wide diversity in morphology, behaviour, and life histories (Sibley and. Ahlquist,
1990).
The order Charadriiformes includes three suborders, viz., Charadrii, Lari and Alcae (Fain and Houde, 2007). The suborder Alcae is characterized by the short-winged birds, and is represented by a single family, viz., Alcidae (auks). Suborder Lari includes the gulls
(Family: Laridae), terns (Family: Sternidae), skimmers (Family: Rynchopidae), jaegers, and skuas (Family: Stercorapiidae). Charadrii is a relatively large suborder includes 216 species which are usually divided into 11 families, i.e., Burhinidae (the thick-knees),
Charadriidae (plovers), Chionidae (sheathbills), Dromadidae (crab plovers); Glareolidae
(coursers and pratincoles), Haematopidae (oyster-catchers), Recurvirostridae (stilts and avocets), Rostratulidae (painted snipes), Scolopacidae (phalaropes, snipes and sandpipers), Thinocoridae (seedsnipes), and Jacanidae (jacanas).
The body sizes of the birds included in the family Jacanidae range between 15 to 58 cm, and the females are usually larger than the males. All the species of the family are with elongated toes and claws and are distributed in tropical freshwater wetlands having floating aquatic vegetation. Most species in this family have polyandrous mating system
(Betts, 1991). The classification of the family Jacanidae (Wetmore, 1965; Slikas, 1997;
Butchart, 2000) suggests that the family is represented by six genera, viz., Microparra,
Irediparra, Hydrophasianus, Metopidius, Actophilornis and Jacana, and eight species.
Four of these genera, i.e., Hydrophasianus, Metopidius, Microparra and Irediparra, are monotypic, while the two others, i.e., Actophilornis (A. africanus, in Africa; and A.
4 albinucha, in Medagascar) and Jacana (J. jacana in South America, and J. spinosa in
Central America) are represented by two species each (Slikas, 1997).
The phylogenetic relationship between different genera and species of the family has remained in controversy. Depending upon analysis of skeletal and muscular characters,
Strauch (1978) divided the genera of family into two groups, Hydrophasianus and Jacana falling in the one group, and Metopidius, Actophilornis, Irediparra, and Microparra in the second. Chu (1995), using cladistic approach, indicated that the origin of Jacanas was monophyletic. Sibley and Ahlquist (1990), DNA hybridization technique, supported the view of Chu (loc cit) and suggested that Actophilornis was much closer to Irediparra than to Jacana. Whitingham et al (2002) compared the sequences of mitochondrial cytochrome-b and ND5 genes in a phylogenetic analysis of six species of jacanas and suggested that members of the family Jacanidea comprised of two clades. Amongst four genera grouped in one clade, Irediparra and Microparra were closer to one another than
Metopidius and Actophilornis. Similarly Jacana jacana and Jicana spinosa presented a higher level of similarities than that with the genus Hydrophasianus, grouped in the other clade.
General descriptions of the Pheasant-tailed Jacana (Hydrophasianus chirurgus) have appeared in different publications, and the present one for this bird species has been developed from the accounts appearing in Oliver (1962), Roberts (1991), Grizmek
(1972), Hassan (2001), and Mirza (2007). Pheasant-tailed Jacana is a medium-sized bird, adults weighting 126 - 231 gm. Both the sexes of this Jacana species are similar to one another in their general morphology, yet females (450 - 54 cm in length) are bigger than the males (44 - 47 cm in length). The adults have a more colourful and longer plumage during the breeding season than that during the non-breeding season. With the breeding
5 plumage (May - October) the body length measures 64 – 79.5 cm (very prominent tail adding 20 – 32.5 cm into the body length), while it is 28 - 31 cm during the non-breeding season. The Pheasant-tailed Jacana shares the characteristic presence of long toes and claws (up to 10.2 cm), and a frontal shield with the other jacana species. The long legs claws and tertials enable the birds to walk and balance their body weight in floating vegetation without sinking (Ramachndra, 1993). The metacarpal spur at the bend in their wing can also be used as defensive organ against predators. The species can fly with its efficient wings (18.2 - 24.2 cm long), except during short molting periods. In the flying posture the neck extends forward and long legs appear trailing behind. During breeding season the species has a white face and neck with contrasting yellow nape, supplemented with conspicuous white wings with a dark chocolate-brown breast and belly, with long, down-curling central, jet-black tail feathers. The mantle and rump are dull brown and the brownish-white tertials are elongated, curving upward. In adults the first primary feather tips became speculated distally with an almost barbless quill or rachis. The rear part of the crown has dividing line down the side of neck, was black. The iris is dark brown and slender, slightly down curved bill (2.5 – 2.9 cm long) was bluish. During the non- breeding season, long tail vanishes, and mantle becomes sandy buff, the breast turns white leaving with a narrow black breast band and white supercelium, while whole of the crown and hind neck bears a dark brown plumage. The black line bordering the brownish- yellow neck patch becomes broader and extended to eye, and the bill is grey green.
Chicks have smaller forehead shield as compared with the adults, while young ones bear brown plumage with white under parts.
The adults of jacanas are less vocal, but in breeding season and while guarding the chicks their call may be compared with cat-like mewing, which changes into strident. Males and
6 females had same calls but males have a louder call and vocalize more often than the females.
1.3. OBJECTIVES
The review of literature suggest that no in depth field study is available on the biology of
Pheasant-tailed Jacana, especially with especially from Pakistan. The biology of this species is known from certain casual reports and observations recorded (Roberts, 1991).
Therefore a thorough study was required to understand the biology of this jacana species.
The present studies have been planned on the belief that Pheasant-tailed Jacana
(Hydrophasianus chirurgus), having its own genetic potentials, has the ability to survive under some specific set of ecological conditions, which are managed by the species different populations of the species under the conditions of Punjab, Pakistan. The underlying hypothesis of the present study is that the population structure of the Pheasant- tailed Jacana in different localities is controlled by different environmental factors of the specific locality, the specific objectives of the studies are to:
i. Study the population dynamics of Jacana in different localities during different
seasons.
ii. Study the biology of Jacana with reference to habitat, feeding and breeding
biology.
iii. Associate different population and biological parameters with different
physico-biotic factors.
iv. Acertain present and predict future possible stresses on survival of viable.
Jacana population and its habitat.
The study will help in evaluating the biological requirements of the population of Jacana
7 under the specific environmental conditions of Punjab (Pakistan). This will also provide base line information on the present status of the species, which can be utilized to visualize the future possible trends and the impact of future possible changes in the environmental conditions on its population in the wake of rapid climatic change scenario.
8 REVIEW OF LITERATURE
2.1. POPULATION DYNAMICS
2.1.1. Population Distribution
There is little information available on the presence of the Pheasant-tailed Jacana in some specific water bodies of Pakistan. General remarks indicate that this bird species was rare in southern Punjab, and was not seen in the Northern Punjab and the Sindh. The presence of the species was recorded from Gilgit (Baker et al., 1929), Murree Hills, Besham,
Kohistan and Indus valley (Roberts, 1991). Hasan (2001) recorded the presence of the
Pheasant-tailed Jacana in large bodies of water in the Indus valley.
Mirza (2007) believed that the Pheasant-tailed Jacana were present in the swamp of Nara area and the Indus river, while the birds of the spcies were also seen on the road side ponds of Kala Khatai (District Sheikhupura, Punjab). None of these reports describe the distribution of this Jacana species in different water bodies of Pakistan.
2.2.2 Population Size
Specific studies on population size of Pheasant-tailed Jacana are not available from
Pakistan. Humitirou and Sinkiti (1940), while discussing the distribution of this bird species in northern, central, and southern Taiwan, believed that the total population in the area, though varied with season and area, yet it was always less then 100 individuals.
Butchart (1998) conducted a three year study on dynamics of population of the Bronze- winged Jacana (Metopidius indicus) in a freshwater lake of Vembuar and suggested that the bird population was at its highest level during breeding season and the size of the population varied in relation to season and plant cover, but ranged between 41 to 62 individuals.
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Li and Mundkur (2004) have summarized the data on the winter census of the water birds and reported the sighting of 32 Pheasant-tailed Jacana during 1997, none in 1998, 452 in
1999, 477 in 2000 and 410 in 2001 at different water bodies of Pakistan. They however, did not mention the break down population, as recorded from different water bodies.
Azam et al. (2008), in his mid winter counts, could spot only 4 Pheasant-tailed Jacana at
Chashma Lake, Punjab (Pakistan), and could not spot any at Shahpur Dam, Jannah
Barrage, Nammal Lake, Jhalar Lake, Ucchali Lake, Khabbaki Lake and Kallar Kahar
Lake. No sighting has been reported from other wetlands of the Punjab (Pakistan).
2.2.3. Population Fluctuation
Birch (1957) proposed that population fluctuation in majority of the animal species is controlled by a number of environmental factors, such as, climate (air temperature, water temperature, topography, humidity) and population density. Roberts (1991) believed that the Pheasant-tailed Jacana in Pakistan were resident in the Sindh, where these were present mainly at Hadero Lake (southern Sindh) or Ghouspur (Jacobabad district) and used wetlands of Punjab as breeding grounds during summer. Mirza (2007) recorded some breeding birds as north as, up to Kashmir and Gilgit. No information is available on fluctuation in the population levels at any of the water body of the Punjab or Pakistan.
Deignan (1931) reported abundance of the breeding population of Pheasant-tailed Jacana in the Payao and Chieng Mai provinces in northern Thailand though the levels of non- breading population remained unrecorded. Tistr (1981) conducted a study on the population of this jacana species in Songkhla Lake in southern Thailand and suggested that population of jacanas fluctuated during different seasons of the year, and the
10 population of the birds was higher between January and March and low around
September and October.
2.2.4. Sex ratio
The available literature suggests that the jacanas frequently present a slightly tilted sex ratio in favour of males, though no reports directly concern Pheasant-tailed Jacana.
Butchart (1998) reported that, in the Bronze-winged Jacana (Metopidius indicus), population under suitable habitat conditions, the number of males has always been greater than the females. A sex of 1.9 male per female has been suggested for adult population of this jacana species. Jennie and Collier (1972) monitored the population of American
Jacana (Jacana spinosa) for 4 years and reported that the number of the females (average
15) were always less than the males (average 34), yielding a sex ratio of 2.3
(males/female). Stephen et al. (2004) found that male to female ratio ranged from 1.43:1 to 2.22:1 for the population of Wattled Jacana (J. Jacana). The female of the jacana are larger in size and hence probably faces a higher predation pressure or a higher cost of living in female results in the selective elimination of the females.
2.2.5. Migration
Though a detailed study on migration or movements of the Pheasant-tailed Jacana in
Pakistan are not available, yet observations available in literature suggest population movements from south to north and vice versa during spring and autumn, respectively.
The species has been recorded in the southern parts of Pakistan, i.e., Sindh, as resident, but could be recorded as breeder in northern parts of the country, i.e., Punjab, some times extending in to Murree Hills, Kashmir and as for as, Gilgit (Hasan, 2001). Kazmerczak et al. (2006) reported some local movements in the resident population of this species during drought in unfavourable conditions. Tistr (1981), the population of pheasant tailed
11 Jacana in Songkhla Lake (South Thailand) suggested that migrated locally with the season for breeding purposes. Though well defined population estimates are not available for the breeding and non-breeding populations, yet the distribution ranges presented
(Roberts, 1991) tend to suggest that the species shows population movements between season and the population spreads over a wider range during breeding periods. Such movements does not strictly fall under ecological migration, demanding geographically isolated breeding and non-breeding grounds (Odum, 1971).
2.2. HABITAT
2.2.1. General
Specific studies on the habitat requirements of the Pheasant-tailed Jacana are not available from Pakistan. The information regarding habitat of the species are derived from the general remarks appearing in the distribution notes recording general conditions of the habitual of the located bird.
Grizmek (1972), depending upon such records, suggests that the Pheasant-tailed Jacana is found in marshes, ponds and lakes having floating aquatic vegetation, though the species preferred emergent vegetation during winter. Butchart (1998) and Kazmerczak et al.
(2006) remarked that this species preferred shallow, freshwater wetlands that have floating and emergent vegetation, particularly the Lily pads, where they could feed and nest. Rao (1991) suggests that the species preferred the places where there was an abundant supply of water, though they also need thick mats of floating aquatic vegetation for shelter, nesting material and food supplies. He emphasized that every aquatic animal species has very specific habitat requirements, including water quality and biological diversity, which needs to be fully understood, for the species conservation and survival.
12 Roberts (1991) has given a rather detailed note on the habitat of the Pheasant-tailed
Jacana with reference to Pakistan. He suggested that the species in Pakistan is found to inhabit permanent swamps and lakes with good vegetation cover of floating and emergent plants in summer. However, during winter the species preferred to move to open water bodies with submerged growth of water weeds, including Hydrilla verticillata and
Vallisnaria spiralis. In summer the birds of the species preferred areas with rice fields and hence is dispersed in rice growing tracts of the Sindh (Hyderabad, Larakana and
Thatta) and the Punjab (Gujranwala, Sheikhupura and Sialkot). Hasan (2001) suggested that the Pheasant-tailed Jacana was found in all large bodies of water in the Indus valley, while Mirza (2007) suggested that the habitat of Pheasant-tailed Jacana is the swamps appearing along the river Indus and its tributaries in the Nara area. He, however, recorded that during winter the bird species was observed on the roadside ponds having thick mat of floating vegetation in Kalakhatai area (District: Sheikhupura).
2.2.2. Physical Factors
Beecher (1942) remarked that a correlation exists between physical characteristics of wetlands vegetation and the size of the population of different aquatic birds. Edward
(1962), while studying the factors regulating animal population, proposed a closer link between population density of the bird species and its food in controlling the fluctuation in animal populations. Haramis et al. (1986) emphasized that suitable habitat provides the food in the ecosystem, which is required to maintain a favourable energy flow to meet the requirement of the birds. Paulus (1988) was of the view that energy relationships between a bird and its habitat, as reflected by its body condition in relation to other individuals, adjusted through the time activity budget of the bird. Conversely, Sampath and
Krishnamurthy (1990) observed that bird population parameters, such as, species richness, relative density, and diversity of birds were the indicators of habitat quality. 13 Ormerod and Tyler (1993) strongly emphasized that aquatic birds are ecological indicators of water quality and subtle changes in hydrology or in physical or chemical limnological characteristics could exert profound effect on the habitat of aquatic bird communities.
Only direct study available on the habitat determinants of the Pheasant-tailed Jacana revealed that the water level played an important role over the invasive plant species,
Eichhornia crassipes, Salvinia molesta, Imperata cylindrical and Mikania Sp. which ultimately resulted in a decrease in the available foraging habitat for Jacana (Mahaulpatha et al., 2007).
2.3. FOOD AND FEEDING
Direct studies on the food and feeding of the Pheasant-tailed Jacana are not available.
Casual observations on this bird species have, however, appeared which give some indications of its food or feeding behaviour. Hayman et al. (1986) suggested that
Pheasant-tailed Jacana fed upon insects, invertebrates and seeds.
Woodcock (1980) while describing food and feeding behaviour of the Pheasant-tailed
Jacana, suggested that they were common in wetlands covered by weeds and fed upon aquatic plants, insects and larvae of invertebrates. Butchart (1998) and Lal (2004) believed that the Pheasant-tailed Jacana in India prefer to forage in weed carpeted stretches of water with emergent and floating vegetation, like water chestnut (Trapa bispinosa), where the birds could feed and nest. The bird preferred to move slowly across the floating water weeds, and probably picked insects, insect larvae, spawns of mollusks, and amphibians. The bird was thought to place a heavy reliance upon freshwater snails for its food. Butchart (loc cit.) recorded that during the breeding season birds of this
14 species feed on small aquatic invertebrates, though small seeds and other plant matter is also consumed. Lal (loc cit.) reported that the Pheasant-tailed Jacana was not particularly shy from humans and the birds continued the forage actively throughout the day, without caring for the presence/ movement of man around the water body. In another report,
Oliver (1962) suggested that Pheasant-tailed Jacana walk on vegetation surface for eating insects, small mollusks and they also like to feed occasionally upon minnow. The species also preferred seeds of water plant for their feeding.
Roberts (1991), while compiling the available information on the food and feeding behaviour of Pheasant-tailed Jacana with special reference to the area under Pakistan remarked that the birds of this species inhabit larger portions of wetlands where they prefer to forage across floating water weeds or lilly pads. He believed that the species prefers to consume insects, larvae, spawns of mollusks and amphibian. The species also likes to eat freshwater snails, small bivalves, mollusks, while some portion of their diet consists of seeds and succulents parts of water plants.
Hayman et al. (1986) reported that the jacanas preferred to feed on the invertebrate present on the floating vegetation or even on the surface of water. Grizmek (1972) has developed a note on the general biology of different animal species of the world and remarked that major source of food of all the different species of jacana is insects, associated with the aquatic habitat. Osbourne and Bourne (1977) recorded that the
Jacanas are found to inhabit freshwater marshes. They believed that the food of jacanas generally comprised of four types of seeds and five kinds of insects. Seeds comprised about 20% of the total diet by volume, while invertebrates made up some 80% of the average diet, which mainly included beetles and Clandera spp., apart from Lissorhoptrus spp., grasshoppers, shore flies, and crickets. Jenni (1974) suggested that the jacanas feed
15 on insects, picked up from the surface of floating vegetation, though they more frequently feed on ovule of water lilies.
2.4. BREEDING
Grizmek (1972) believed that jacanas, as a group, are polyandrous, and a female can mate with one to four males and defend their territories. These territories can be the size of a football ground. The defending behaviour begins with spreading of feathers with spur directed forward. Butchart et al. (1999) reported that female in polyandrous harems gave more copulation to the male. The jacanas generally lay four eggs at a time, and chicks hatch after 22 -28 days (Thongaree et al., 1995).
Courtship
Thong-aree et al. (1995), based upon their studies 1-Central Thailand reported that in
Pheasant-tailed Jacana courtship begins during the rainy season with some degree of pairing up of the individuals of two sexes. They believed that a female could pair with upto four males to successfully produce four clutches of eggs in one breeding seasons, when the food was abundant. The female fly acrobatically around the selected male and foraged close to him. The females also made loud calls to attract the male or some times the female tempted the male to fly up. After pairing, male and female flash their wings at each other before mating Chen et al., (2008)
Egg Laying
Thongaree et al. (1995) reported that in Pheasant-tailed Jacana egg laying was terminated after the female had laid the last egg of the clutch, even if eggs were removed. When the female had completed the laying of one full clutch, the male started incubating these eggs.
At the same time the female abandons the nest and pairs with another male. Eggs of 16 Pheasant-tailed Jacanas were pyriform in shape with one end rounded and the other rather pointed. The texture of the eggshell was smooth and shiny. The color of newly laid eggs varied from olive to dark green. Shell colour turned reddish or darkbrown when hatching approached. The thickness of the egg shell ranged from 0.35 to 0.40 mm.
Chen et al. (2008) suggested that during breeding season the males Pheasant-tailed Jacana spent more time in breeding activities while foraging time decreased. In contrast, females consumed more time in foraging while time spent on breeding activities remained low.
They also showed significant difference in foraging time between sexes during the early breeding stage. It was reasonable to understand that female Pheasant-tailed Jacanas spent more time in foraging than males, during the early breeding stage, owing not to be involved in incubation.
Nesting:
Chen et al. (2008) reported that female chooses a site within the territory to build a nest while males were responsible for building the nest. Subramanian et al. (2000) reported that the nest of Pheasant-tailed Jacana looked like a floating raft on surface of water without any depth for holding the eggs. Geetkia (1992) reported that the major part of the nesting in Pheasant-tailed Jacana occurred during June and July (22 out of 32 nests) and composed of floating structure of grasses and weeds with eggs just floating or half immersed in the water. The nests were loosely built on rafts of floating vegetations and took the form of a pile of solid vegetation with a shallow depression in the centre. The average diameter and depth of two nests was 13.7 cm and 3.4 cm.
Habitat:
Geetkia (1992) suggested that Pheasant-tailed Jacana selects a nesting site along the edge of the reservoir where aquatic plants are denser, consisting of water lily leaves or other 17 plant material appearing on top of a mat of floating vegetation. The dominant plant species around such sites include, Nelumbo nucifera, Nymphaea pubescens, Salvinia cucullata, Potomogeton malianus and Trapa quadrispinosa. Water depth at the nest sites varied from 0.3 to 2.0m.
Parental Care:
Wiktander et al. (2000), Thongaree et al. (1995), and Stephen et al. (2004) studied the parental investment of polyandrous breeding system and suggested that males showed greater parental investment than females. They reported that males took the most parental duties and females scarcely provide parental care, including the incubation (24 – 26 days, later being more frequent) and caring of the chicks from hatching until fledging (50 – 60 days). The male started incubating shortly after the female had laid the first egg. After the female laid an egg she walked away a few meters from the nest, when male came to attend the eggs. The male did not sit on the eggs all the time but remained close by while picking up food, but kept on turning the eggs at frequent intervals. Males incubated the eggs for the major part of the night and during the hotter part of the day, probably saving these from overheating or chilling. On disturbances males build the new nests, and eggs translocation was observed.
Territoriality:
Both the sexes of the Pheasant-tailed Jacana exhibit strong intra and inter-specific territorial defense around their nests. They announced their territories by loud calls.
During the initial period of incubation of eggs (first 2 weeks) the female is responsible for defending the nest (Thongaree et al., 1995). Stephen et al. (2004) suggested that females are heavier than males, showing dominant behaviour and developed more secondary
18 sexual characters including flashy facial ornamentation and wing spur. The territory defense was commonly shared between both the sexes. The competition for territory was so high that the females compete to get a chance of mating with more males. Larger and heavier females succeed to get more mates than smaller ones, and hence the body size has a role in inter-sexual competition for territories and mates.
The Jacanas as a group are polyandrous birds, and a female has been reported to mate with 1-6 (with an average of 1.8). The pairbond in jacanas showed flexibility, because when a female mates with a male to provide a clutch, they are chased by others before receiving clutch (Mace, 2000). Butchart et al. (1999) suggested that males defend their individual territories, while the female territory was defended by all of her mating males.
They believed that the size of the territory was not influenced by the availability of resources but was largely decided by the number of the males present in the female herm.
Thongaree et al. (1995) reported that a female in Pheasant-tailed Jacana matted with 1-4 males. Their report suggested that the nest was built by female, while male incubates the clutch of four eggs. The incubation period was around 25.3 days, and they observed that hatching success was 71% remaining/around. Chen et al. (2008) studied the polyandrous breeding system in Pheasant-tailed Jacana and found that most of the parental duties was performed by male only and during incubation, female took more time to feed whereas, male had to perform incubation duty and perform parental care by protecting the eggs from high temperature and ultraviolet rays especially at noon. They observed that male of
Pheasant-tailed-Jacana showed infantacidal behavior by threw away the eggs of previous males and accept the paternity of only two eggs. Previously, Chen et al. (2008) found that the interval between copulation of female with different males was shorter than 10 days.
It was suggested that sperm remained viable in the reproductive tract of female for at least 19 4 to 5 days and it was reported that sperm competition among the males was very intense so the paternity of the first two eggs was doubtful. So they suggested that pheasant tailed jacana male showed infantacidal behavior as a strategy to avoid cuckoldry when the paternity of eggs was doubtful.
Numerous predators prey on young jacanas, which include the purple gallinule, snakes, otters, and turtles and predation was reported to claim 20% of the chicks (Thongaree et al., 1995).
2.5. GENERAL BEHAVIOUR
The Pheasant-tailed Jacana is gregarious in nature during winter (Kazmerczak et al.,
2006), when these are very vocal and are concentrated in some part of the wetland (Lal,
2004). Kazmerczak et al. (2006) recorded the generalized call of the species, sounding like, kyoo, kyoo, kyoo, meoph meoph, k k k k nasal, tewn, tewn, tewn. Lal (2004), however, believed that the summer and winter calls are different in the species. The winter call sounds like meeoph, meeoph and then queeear but in summers the birds emit a territorial call, sounding like cat, mew, mew, meeoph often increased to ooph, ooph and alternated with quicker, shriller, meyu, meyu, meyu, and meyu. The birds of the species can swim very well and can dive during danger.
Pheasant-tailed Jacana uses elongated toes to walk upon floating vegetation like lotus, water hyacinth, trapanut and pistia Scot and corbonell 1986. They spent most of the time of day in sun, preening or foraging, during non-breeding period. Scott (1989) They also suggested that chicks just after hatching can swim and dive in water and used this ability to escape from predators. The adult pheasant tailed Jacana can camouflage very well in
20 aquatic vegetation even though they were bright in color during breeding season
(Grizmek, 1972).
2.6. POPULATION STRESS
Rao (1991) believed that currently none of the jacana species is threatened. However, pesticide, floods, and predation remain as serious threats to the survival of these bird species, effecting egg laying capacity, nesting, safer hatching of eggs and survival of the fledglings. He suggested that habitat destruction/allowed alteration remains the major threat to the future survival of the Pheasant-tailed Jacana populations. At places anthropogenic wetlands, managed for rice cultivation, are the most favourite habitat of the
Pheasant-tailed Jacana, which involve complex food web interactions between various biotic components of the rice ecosystem (Catling and Islam, 1999).
Bambaradeniya et al. (1998) expressed their concern about a fast disappearance of wetlands throughout the globe, which can indirectly affect all the species associated with such wetlands, including Pheasant-tailed Jacana. Scott and Carbonell (1986) believed that major threats to wetlands in each country include drainage of lakes and marshes, diversion of rivers, changes in patterns of flooding, and pollution.
21 MATERIAL AND METHODS
3.1 STUDY AREA
The Punjab (27.56 – 34.00NL, 69.36 – 75.22E), the north-eastern province of Pakistan, spread over an area 205,344 km2. The eastern boundaries of the province touch India, while Kashmir falls in the north, the Khayber Pakhtun Khwa (KPK) and the Balochistan in the west, and the Sindh province touches the southern boundaries of the province. The western extremities of the Himalayas extend in to the northwestern limits of the province, while the rest of the province is occupied by the Indus Plains, which represents a gradual natural north-south slope. Under the natural slopes and the mountains, five major rivers, i.e., Sutlej, Ravi, Chenab, Jhelum and Indus, emerge from different places in the north- warldly located Himalayas and traverse the province, joining one-another at different spots and ultimately into Indus river (Figure 1.1).
Climatically, the province is a sub-tropical country, having winter (November - January), spring (February - April), summer (May - June), Monsoon (July - August), and autumn
(September - October). The lowest temperatures are experienced in January (12C to
15C) and the highest in June (46C to 51C). The average temperatures are normally lower in the northern latitudes, and higher in the southern parts. The precipitate ranges between 15 mm and 316 mm in different parts of the study area. The area receives the rain through summer and winter monsoons. The eastward summer monsoons (July-
August) are heavier and more persistent. The splash summer rains cause the water ways to swell up, sometimes causing floods. The summer rains are heavier and more frequent in north-eastern parts and gradually decreasing as one moves towards western and southern parts (Table 3.1), so that
22 N
W E China Gilgit Baltistan n a S t r is P e F Azad Kashmir iv n R a W s h N u er g nd Riv f I um A Punjab hel J òMarala er iv Qadirò Abad R ia ab er d en iv Baluchistan n h R I C vi
Iran Baluki a R Sindh ò
Arabian Sea Sulemanki iver ò lej R Sut
ad jn an P Punjab Boundries ò Study sites Rivers 0 60 120 180 240 300 360 Kilometers
Figure 1.1 and Figure 1.2: Location map of the Punjab (Pakistan) and rivers of Punjab.
23
Table 3.1. Mean maximum (Max) and minimum (Min) temperature (C), and annual rainfall (mm) recorded at four study sites during 1996 – 2005. Source: Courtsy Pakistan Meterological Department
Marala Qadirabad Balloki Sulemanki
Rainfall Temperature Rainfall Temperature Rainfall Temperature Rainfall Temperature
(mm) Max Min (mm) Max Min (mm) Max Min (mm) Max Min
January 55.3 18.4 5.3 11.5 19.4 4.1 14 19 4 17.5 19.8 5.2
February 40.2 21.4 8.0 20.1 21.9 7.1 13 22 7 8.2 22.9 8.6
March 44.6 26.8 12.9 25.7 26.7 12.3 22 27 12 20.8 28.3 13.5
April 13.5 33.2 18.4 16.9 33.5 18 14 33 18 4.7 34.8 19
May 13.2 39.0 23.8 16.1 38.4 22.7 14 39 23 8.3 40.1 24.1
June 50.8 40.6 26.7 27.9 40.5 26.9 24 40 27 19.2 41.4 27.7
July 316.1 35.5 26.1 115 37.1 27 109 37 27 92.1 38.3 28
August 259.0 33.8 25.3 89.8 36.1 26.6 91 36 26 74.7 37.1 27.1
September 148.8 34.3 23.6 28.6 35.7 23.7 33 36 24 35 36.6 24.3
October 32.0 32.6 17.8 3.8 33 17 7 33 17 0.3 34.4 18
November 12.4 26.8 10.4 3.0 27.2 10.1 4 27 10 4.6 28.3 10.8
December 24.3 20.3 5.9 8.6 21.4 5.1 9 21 5 10.6 22.2 6.2
24 minimum rains are received in the southwestern sides of the province. The rainfall pattern suggests a cyclic change in the average annual rainfall, with 4-5 years of heavier rains, followed by equal number of the years with low rainfall. Floods have been regular feature, which use to support a series of wetlands around the rivers.
During the last century, a network of canals has been developed to support irrigation. The canals system of the Punjab is considered to be the Twentieth Century Engineering
Wonders. The rivers are connected with link canals. System of large earthen dams, reservoirs, barrages and headwork’s have been developed at process to requlate water into the canal system, creating man-made lakes of different sizes. Twelve head works and barrages are located at different rivers of the Punjab (Figure 1.2). Such a system has replaced the flood supported wetland with more reliable water bodies appearing in the form of such reservoirs. The water level in these reservoirs fluctuates with the pattern of rainfall, and hence sizes of these reservoirs vary during different parts of the year. The reservoirs characteristically have the main body of deeper water, which are associated with shallow ponds appearing towards the periphery of the main water body. Rooted vegetation, both floating and emergent, appears in these shallow water ponds, supporting different species of waders. The main birds of such areas include, Purple Gallemule
(Porphyrio porphyrio), White Breasted Waterhen (Amaurornis bicolor), Indian Moorhen
(Gallinula chloropus), Pond Heron (Ardeola grayii), Little Egrets (Egretta garzetta),
Grey Heron (Ardea cinerea) Purple Heron (Ardea purpurea), Pheasant-tailed Jacana
(Hydrophasianus chirurgus), Black Crowned Night Heron (Nycticorax nycticorax), Little bittern (Inobrychus sinensis), Red Watteled Lapwing (Vanellus indicus) (Whisler, 1986;
Bhushan et al., 1993; Ali et al.,1989; Stewart and Blin, 1976).
25 The present study on the biology of Pheasant-tailed Jacana concentrated on four lakes, head works.
1. Head Marala: Located in north-eastern parts of the Punjab (32.4o NL, 74.31 o E) at an altitude of 250 m, close to the town at River Chenab. (Figure 2.1). Apart from the main lake, ponds of shallow water (0.2 – 5.0 m deep) appear on the two sides (Figure 2.1), having an area of some 1,620 ha.
Figure 2.1. Map showing the location of site 1 (Head Marala)
2. Head Qadirabad: Located (32.19 o NL, 73.39 o E) at an altitude of 225 m near to
Gujranwala, Down stream of Head Marala on the River Chenab (Figure 2.2), having pond area of some 2,850ha. The depth of water in the ponds varied between 0.2 and 5.0 m.
26
Figure 2.2: Map showing the location of site 2 (Head Qadirabad)
3. Head Balloki: Constructed on the River Ravi (31.13 o NL, 73.52 o E) at an altitude of
362 m, some 40 km down stream of Lahore (Figure 2.3), with a total pond area of some
1,620 ha. The main supply of water is from the River Chenab, diverted through B.R.B link canal. The depth of water in pond area varies between 0.1 - 3.0 m.
Figure 2.3: Map showing the location of site 3 (Head Balloki)
27 4. Head Sulemanki: The head is located (29.49 o NL, 72.33o E) at an altitude of 177m in the south-eastern parts of the Punjab on the river Sutlej (Figure 2.4). The headwork develops a lake receiving main supply of water diverted from Chenab and Ravi rivers, as the Sutlej is almost dry when it enters Pakistan from India. The pond area during summers remains around 1,620 ha, where the depth of water remaining between 3 to 6.0 m.
Figure 2.4: Map showing the location of site 4 (Head Sulemanki)
3.2 POPULATION BIOLOGY
The four selected localities, having the major population of the Pheasant-tailed Jacana, i.e., Head Marala, Head Qaderabad, Head Balloki and Head Sulemanki, were visited on same dates of the different calendar months, between April 2004 and March 2007. The favourable jacana habitat patches at each locality were divided into four study sites. A fixed place at each of the study site was selected, from where the total habitat was visible.
It was regularly used for population census of Pheasant-tailed Jacana, with the help of binoculars (Olympus 8×40 mm) and field spotting scope (Olympus 15-60×60 mm). Care
28 was taken to avoid double counts. Each of the bird noted was recorded according to its gender and age.
The area under the favourable Pheasant-tailed Jacana habitat at each of the sampling site was calculated using GPS positioning (MAGELLAN SporTrak CQlor) supported by GIS
(ArcView GIS 3.3).
The census counts of the population were regarded as the population of the species exploiting the habitat at some specific time. For deriving the comparative population parameters of each site the population density of the bird numbers was calculated by dividing the population size with the estimated habitat area of each site and for each locality. The sex ratio (female to male ratio) was calculated by dividing the number of the adult females by the adult males. The adult to juvenile ratio was calculated by dividing the number of juveniles by the number of the adults, and females. The population data collected from different sites was pooled to derive different population parameters for different localities or the total population.
3.3. HABITAT
3.3.1. Sampling
The four study sites were visited during each calendar month between April 2004 and
March 2007 and the observations were recorded from two different locations of each locality. Prevailing weather conditions, and other relevant information were maintained for each study site.
3.3.2. Habitat Analysis
Physical factors:
29 Temperature: Temperature of surface (10 cm under surface of water) water and air (10 cm above surface of water) was recorded directly at each site, with the help of thermometer
(C mercury).
pH: The pH of surface water was measured directly with the help of field pH meter
(Lateron 0pH-206 meter), after standardizing of the pH meter with pH tablets, BDH chemicals, pool England.
Vegetation Analysis:
Estimation on the frequency of the different species of the rooted plant present in ponds of four study sites of each of locality, i.e., Marala, Qadirabad, Balloki and
Sulemanki, were made by using visual estimation and general approximation following
Brown- Blanquit method (Causton, 1988). The representative specimen of each plant species was collected in the field and identified at the Herbarium, Botany Department,
Government College University, Lahore, Nasir and Ali (1972) and Majid (1986) for different groups. The individually collected estimates were suitably pooled for general estimation.
30 3.4. FOOD
A total of 43 stomachs, along with their contents of Pheasant-tailed Jacana were obtained from the freshly killed birds from four study sites during 2004 – 2007 period.
Each stomach was opened in the field and its contents were flushed with distilled water and placed in a Petri dish containing distilled water. The identifiable parts of stomach content were visually separated and identified directly up to the lowest possible taxonomic level, using available identification keys (algae: Bold and Wynne, 1987; animals: Willoughby, 1976; Melleiby, 1986; Tonapi, 1986; Huet 1986, higher plants).
The number of different food items were counted and expressed as relative proportion of the total content.
Total number of fragments of species Relative frequency (%) = x 100 Total number of fragments analyzed
Means and standard errors of mean of different food items were calculated by using normal statistical methods (Mather, 1964, Sokal and Rohlf, 1969). Constancy of appearance of each food item was determined using plant ecology techniques (Mueller et al., 1974)
Frequency (%) = Number of samples showing species x 100 Total number of sampling point
Each bird was observed in the field for a period of 60 seconds, during different field tours, while picking up the food and recorded as field notes. During this period the number of the total feeding attempts and the successful ones were recorded directly as observed in the field. The successful attempts finally terminating in the final engulf of the
31 food picked was also recorded through field observations and engulf rate was calculated by dividing the successful engulfs by the total successful picks. The distance traveled for feeding from the nest was recorded as the best possible approximation.
Percentage of time consumed in feeding and resting during early and late breeding stages by every male, female and nestling was recorded to know the feed preference during breeding season.
The weight of the stomach contents (recorded to the nearest gram) was regarded as the food intake. The weight of each birds killed by hunters was also recorded to the nearest gram. Food intake to body weight was then calculated by dividing the food intake by the relevant body weight.
3.5. BREEDING
The established nests of the Pheasant-tailed Jacana were spotted through a search of the area with the help of the local hunters and staff of Wildlife Department, at different water bodies, during summers between 2004 to 2007. The nests were marked with appropriate tags, attached with the vegetation and were regularly visited for observations from a suitable location without disturbing the bird. A selected number of breeding females were also marked for observation through the territorial location and other visible morphological marks. The marked nests and the selected females were followed through daily early morning observations upon laying of the clutch (number of the eggs laid in a complete clutch), number of the days of incubation and number of the hatched eggs. A total of more than 250 nests were initially selected for studies. Some of the selected eggs could not be followed with reliability and were excluded from the final analysis.
Information on all the breeding parameters could be collected on 180 nests (Marala=36),
Qadirabad=67, Balloki=29, Suleimanki=48) and 52 females (Marala=13, Qadirabad-19,
32 Balloki=8, Suleimanki=12). The observation collected from different nests/ breeding females were pooled to develop generalizations for different localities, ignoring the individual ponds or the year.
33 4. RESULTS
4.1. POPULATION BIOLOGY
4.1.1. Distribution and Estimates
The data collected on the population of the adult Pheasant-tailed Jacana found on four study sites in the Punjab.(Table 4.1) suggest that during June (mid summer) the over all population ranged between 298 and 347 during different years. There were relatively higher numbers of the birds found at Marala (83 - 96), Qadirabad (95 – 106) and
Suleimanki (88 - 89) as compared to those present at Balloki (21 – 57). The density of the bird population was very high at Marala (14.4 – 16.7 birds/ km2), as compared to other three localities, i.e., Suleimanki (3.0 -3.1 birds/ km2), Qadirabad (2.0 – 2.3 birds/ km2), and Balloki (0.3 -0.8 birds/ km2).
4.1.2. Population Fluctuation
4.1.2.1. Annual fluctuation
The data obtained for three consecutive years (i.e., 2004, 2005, 2006) suggest that the population of this species though showed slight variability at years, with a relatively lower overall population size during 2005 (n=298), as compared to 2004 (n=323) and a slight rise in population size during 2006 (n=347). However, the difference between the years was not significant (chi-square, DF = 2, 3.72, not significant at 0.05 level). The trend of a non significant difference between the years was exhibited by the individual set of populations spotted at sub-localities and the sampling zones, as exhibited by the values of the individual and the heterogeneity chi-square (Table 4.1).
34 Table: 4.1 Population density of Pheasant-tailed Jacana in different water bodies during different years 2004 – 2006 2004 2005 2006 Area Chi Locality Density Density Density (km2) No No No square (per km) (per km) (per km)
Head Marala 5.75 92 16.0 83 14.4 96 16.7
0.8 32 40.0 32 40.0 33 41.3 0.02
3.51 216.0 23 6.6 23 6.6 0.12
0.69 17 24.6 17 24.6 18 26.1 0.04
0.75 22 29.3 21 28.0 22 29.3 0.03
Head Qadirabad 46.9 95 2.0 106 2.3 105 2.2
6.9 344.9 32 4.6 34 4.9 0.08
19.6 26 1.3 28 1.4 28 1.4 0.10
7.5 172.3 24 3.2 21 2.8 1.19
12.9 18 1.4 22 1.7 22 1.7 0.52
Head Balloki 68.09 48 0.7 21 0.3 57 0.8
8.32 26 3.1 8 1.0 26 3.1 10.80
13.57 70.5 5 0.4 7 0.5 0.42
28 80.3 4 0.1 12 0.4 4.00
19 70.4 4 0.2 12 0.6 4.26
Head Sulemanki 28.95 88 3.0 88 3.0 89 3.1
1.93 20 10.4 20 10.4 21 10.9 0.03
12.84 282.2 28 2.2 27 2.1 0.02
0.86 22 25.6 22 25.6 22 25.6 0.00
13.32 181.4 18 1.4 19 1.4 0.04
Total 150.5 323 2.1 298 1.98 347 2.4
Pooled Chisquare 21.49
Total Chisquare 2.37
Heterogeneity
Chisquare 19.12
35 2003 2004 2005 zone 1 32 32 33 zone 2 21 23 23 zone 3 17 17 18 zone 4 22 21 22 zone 1 34 32 34 zone 2 26 28 28 zone 3 17 24 21 zone 4 18 22 22 zone 1 26 8 26 zone 2 7 5 7 zone 3 8 4 12 zone 4 7 4 12 zone 1 20 20 21 zone 2 28 28 27 zone 3 22 22 22 zone 4 18 18 19
Table Analyzed Data 1
Repeated Measures ANOVA P value 0.0932 P value summary Ns Are means signif. different? (P < 0.05) No Number of groups 3 F 2.57 R squared 0.1463
Was the pairing significantly effective? R squared 0.8957 F 20.11 P value P<0.0001 P value summary *** Is there significant matching? (P < 0.05) Yes
ANOVA Table SS df MS Treatment (between columns) 48.38 2 24.19 Individual (between rows) 2839 15 189.2 Residual (random) 282.3 30 9.41 Total 3169 47
36 4.1.2.2. Seasonal fluctuation
Table 4.2 presents the data on the number of the birds of this species documented at different localities during study period. The table suggests that there was no population of this species on any of the locality under present study between November and March. The appreciable population suddenly appeared between the March and April census and suddenly disappeared between October and November census from all study sites.
The population of this bird species exhibited a reasonably consistent pattern of population fluctuation between different summer months (Figure 3.1). The April population exhibited a gradual rise between May and June, with a decline occurring between July and August. The population level showed a sudden rise during September, which was followed by a decline during October. There have been some fluctuations observed from this basic trend at isolated occasions, as for example no September rise in the population of Balloki during 2004 and 2006 and a very significant rise observed in September population at Marala and Qadirabad during year 2005.
.
37 Table 4.2: Monthly Variation of Pheasant-tailed Jacana Population at study sights in years 2004 to 2006
2004 2005 2006 Months
Marala Q/abad Balloki Sulemanki Total Marala Q/abad Balloki Sulemanki Total Marala Q/abad Balloki Sulemanki Total
April 53 60 25 54 192 63 62 30 54 209 58 63 28 54 203
May 72 78 37 67 254 78 86 42 67 273 75 82 40 67 264
June 92 85 48 88 313 95 106 21 88 310 97 103 35 88 323
July 68 77 38 74 257 87 88 24 74 273 80 84 31 74 269
August 78 83 30 84 275 87 84 18 84 273 84 86 25 84 279
September 85 85 31 83 284 86 89 26 83 284 87 89 27 83 286
October 56 52 34 57 199 53 46 25 49 173 57 55 24 55 191
38
100
90
80
70 2004 60 2005 50 2006 40
30
20
10
0 April May June July August September October A Months
120
100 2004 80 2005 60 2006
40
20
B 0 April May June July August September October Months
39 50
45
40
35 2004
30 2005 25 2006 20
15
10
5
0 April May June July August September October C Months
90
80
70 2004 60 2005 50
40 2006
30
20
10
0 April May June July August September October D Months
Figure 3.1: Population size of Pheasant-tailed Jacana at different localities (A=Marala, B=Qadirabad, C=Balloki, D=Suleimanki) during different calendar months of different years (2004-2006)
40 4.1.3 Sex Structure
The female to male sex ratio of 0.75 calculated for pooled population of Pheasant-tailed Jacana suggest that the males had a significant preponderance over the females (chi-square= 39.45,
DF= 1, highly significant at 0.05 level) (Table 4.3). This trend has been equally reflected in the populations of different localities, where the chi-square values remained significant. A very low value of heterogeneity chi-square (0.40, DF= 4, P>0.05) also suggest that this was a persistent trend in summer visiting birds.
The trend of preponderance of the males in the summering population has been equally reflected in all the monthly based samples, except for the May sample at Marala (females: male ratio of 1.05), while in May (female : male ratio of 0.97) and October samples, from Balloki
(ratio = 0.92), the frequencies of the two sexes were close to equal.
Table 4.3: Sex ratio in as exhibited by pool 2004-2006 population Pheasant-tailed Jacana at four study sites
Sex Ratio Locality Male Female (Female/Male)
Marala 298 224 0.75
Qadirabad 317 235 0.74
Balloki 182 144 0.63
Sulemanki 284 205 0.72
41 4.1.3. Age Structure
The overall population of the Pheasant-tailed Jacana presented a juvenile to adult ratio of 0.35
(juvenile to female ratio of 0.82) (Table 4.4). These ratios were maintained in the populations summering in all the four localities, with slight variations, the highest in Suleimanki (juvenile : adult = 0.41, juvenile : female 0.98), followed by Qadirabad (0.36, 0.84), Marala (0.34, 0.79) and the lowest at Balloki (0.27, 0.61).
Looking at juvenile to adult or juvenile to female ratio in the pooled seasonal samples (Table
4.4) suggest that there were no juveniles with the adults during April- June period, resulting in a juvenile : adult ratio of zero. The juveniles suddenly appeared between June and July census and continued to appear in varying ratios in all months there after (between July and October).
The juvenile to adult ratio gradually declined in the subsequent monthly samples at Marala,
Qadirabad, and Sulemanki, though it registered an increase in September October census data conducted at Balloki, where a very high juvenile to female ratio of 2.02 was recorded.
The major lot of the juveniles moved with the adults between October and November census.
Some small populations comprising of only the juveniles persisted at Balloki and Sulemanki during November and December even when the adult birds had moved from the area.
However, no juvenile was recorded at more northwardly located Marala and Qadirabad between October and March.
42 Table. 4.4 Juvenile adult ratio for total Pheasant-tailed Jacana at four sites
Total Juvenile / Juvenile / Locality Male Female Adults Female Ratio Adult Ratio
Marala 298 224 522 0.79 0.34
Qadirabad 317 235 552 0.84 0.36
Balloki 182 144 326 0.61 0.27
Sulemanki 284 205 489 0.98 0.41
4.2. HABITAT
Population of the Pheasant-tailed Jacana was recorded from the shallow ponds or parts of the water bodies having rooted vegetation. No population of this bird species was recorded from the areas having deeper water or associated dry terrestrial tracts.
4.2.1. Physical factors:
Temperature: The changes recorded in the ambient temperature of study sites have been presented in Figure 4.1 (detailed data appear as Annexure 1-4). The available data suggest that the air temperature at four localities, under present study remained in reasonable proximity to one another and exhibited a similar pattern of fluctuations. The temperature during the summer period of fluctuated between 16oC in April and 35oC June, (except at Sulemanki, where it rose to 40oC). The air temperatures were below 17C for the periods when there was no population of this Jacana species on these water bodies. The mean air temperature at different study sites was not significantly different at 0.05% level.
43 Changes in the air temerature during different calender months at different study sites
45 40 35 30 25 20 15 10 5 0 APR MAY JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR
Marala Qadirabad Balloki Sulemanki
Figure 4.1: (a) Air temperature (C) recorded during different calender months at different study sites during 2004-2007.
The water temperature at different water bodies followed a general pattern of the ambient temperature (Figure 4.2., Annexure 1 – 4). The surface water temperature was not significantly different from the air temperature at 0.05% level of significance, and it followed the same pattern, remaining between 20C in April and 40C in June. The water temperature remained below 15C during winter, when there was no population of Jacana.
44 Changes in the water temerature during different calender months at different study sites
45 40 35 30 25 20 15 10 5 0 APR MAY JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR Marala Qadirabad Balloki Sulemanki
Fig. 4.2: Water temperature (oC) recorded during different calendar months at different study
site during 2004-2007.
pH: The pH of the surface water in different ponds remained close to one another (Figure 4.3;
detailed summery in Annexure 1 – 4), and these values were not significantly different from
one another at 0.05 level of significance. The pH for the major part remained slightly basic
throughout the jacana breeding season, except for Balloki, where it was neutral or slightly
acidic during April and May. The pH for the major part remained between 7 and 8. No
prominent fluctuations in the pH of the surface water were recorded, and pH remained within
same limits even during the months when there was no population of Jacana in these water
bodies.
45 Changes in the pH during different calender months at different study sites
10 9 8 7 6 5 4 3 2 1 0 APR MAY JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR Marala Qadirabad Balloki Sulemanki
Fig. 4.3: pH of the surface water recorded during different calendar months at different study site during 2004-2007.
Rooted Vegetation: A total of 32 species of rooted plant species have been identified from the four water bodies, holding appreciable populations of the Pheasant-tailed Jacana. These species are distributed at different depth of water or in the immediate vicinity of such water bodies.
Each pond had a different composition of vegetation composition, depending upon its physico- biotic conditions (Table 4.5). The species having floating were represented by four spies, i.e.,
Nelumbo micifera, N. alba, Nymphaea lotus and Trapa bispinosa, which were associated with five other species, i.e., Eichhornia crasipes, Hydrilla verticillata, Vallisnariria spiralis, Psitia stratiotes and Chara sp., which provided a thick floating mat in the water. Every pond had one or more of these species, appearing in different combinations. The species, like, Arundo donax,
Typha angustata, Panicum paludorsum and Phragmites karka were present in shallow waters in the peripheral area of different ponds in high constancies. All other species were present in the areas surrounding the ponds, yet were associated with the water bodies.
46 Table 4.5: The rooted vegetation population structure at study area.
Mean Area/km2 Mean Vegetative type Population density/km2
Typha angustata – Arundo donax 222 8.0 27.8±2.24 Nelumbo nucifera – Panicum 173 7.5 23.0±2.12 paludosum Chara spp -Hydrilla verticillata 139 6.9 20.1±1.92 Pistia stratiotes - Arundo donax 166 8.832 18.79±2.16 Nelumbo nucifera -Arundo donax 157 19.38 8.15±0.72 Typha angustata - Nelumbo nucifera 164 35.1 4.6±0.60 Arundo donax –Nelumbo nucifera 240 69.0 3.47±0.71 Nelumbo nucifera - Typha augustata 163 75.00 2.17±0.08 Eishornia crasipes – Nelumbo 219 128.40 1.71±0.06 nucifera Pistia striotes – Eishornia crasipes 148 129.85 1.14±0.06 Nymphaea lotus - Nelumbo nucifera 148 133. 1.11±0.05 Trapa bispinosa Panicum paludosum 197 196.0 1.00±0.16 Nelumbo nucifera - Cyperus 157 83.20 0.89±0.07 diformes Eishornia crasipes - Nelumbo 100 135.7 0.75±0.01 nucifera Arundo donax –Typha angustata 60 190.0 0.36±0.01 Nelumbo nucifera - Cyperus 82 280.0 0.35±0.01 difformis
47
Table 4.6: The natural organization of rooted vegetation observed at different study sites. Marala Qadirabad Balloki
Species P1 P2 P3 P4 P1 P2 P3 P4 P1 P2 P3 P4 P1 P2 P3 P4
Ceratophylum + - - + + + - - - + - + + - + - demersum Eichhornia - + + - - + - + - +++ ++ - + +++ + + + crasipes Hydrilla + ++ +++ - + + - + +++ - + - + + + + + verticillata Ipomea equatica - + - - + - + + - + - - - + - - Lamina minor - + - - + - + + - + + + - - + + Azolla filiculoides - + - - + - + + - - - - - + - + Najas graminae + - + - - + - - - - + + + - + + Nelumbo nucifera ++ +++ + ++ +++ +++ +++ +++ + - +++ - +++ +++ + +++ + + + + + Nymphaea lotus - + + - - + + ++ +++ +++ ++++ - - - - - + Psitia stratiotes + - - +++ - + + + - + - - + + - ++ + Potoamogeton - + - + - - + + ++ + - + + - + -- perfoliatus Scirpus maritimus + + - - - + - + + ++ - + + - - + Trapa bispinosa ------+++ - - - - + Trewia polycarpa - + - - - + - + - + - + + + + - Typha elephantina ++ + - - + + - - - + + - + + - - Typha lantifolia + ++ - + + - + + + + + + + - + - Utricularia - - + - + - - + - - - + + - + + vulgaris Vallisnaria + + ++ - + - - + - - + - + + - - spiralis Panicum paludosum - +++ - + + + - - - - - +++ ++ - - - Chara spp - - +++ - + + - - + + - - + + + - + Dicanthium + + + + - + - + + - + + + + - + annulatum Nympaea alba - + + - - - - ++ ++ + ++++ - - - - + Saccharum + ------+ - + + + - + + + spontaneum Saccharum munja + - - - - + - - + ++ - - - - ++ + Oryza sativa + + - + - + - - - - + + - - - + Ipomoea sp. - ++ + - + - - - - + - + - - + + Cyperus difformis + ++ - + + - + + ++ - ++ - +++ + ++ +++ Phragmites karka + ++ - ++ + + + - + +++ + - - - + + Arundo donax +++ - + +++ +++ ++ +++ - + + ++ - - +++ +++ + + + Typha aungustata +++ + + - ++ +++ + +++ + + - + + + +++ ++ + + Trifolium sp. - - + + + ------+ Tritium indicum - - + + + ------+ + - Sulemanki
48
Table 4.7: Vegetation types on the basis of dominant plant species with their own composition at each site.
MARALA
P2: Nelumbo nucifera – Panicum paludosum P1: Typha angustata - Arundo donax
P3: Chara spp -Hydrilla verticillata P4: Pistia stratiotes - Arundo donax
QADIRABAD
P2: Typha angustata - Nelumbo nucifera P1: Nelumbo nucifera -Arundo donax
P3: Arundo donax –Nelumbo nucifera P4: Nelumbo nucifera - Typha angustata
BALLOKI
P2: Pistia striotes – Eishornia crasipes P1: Eishornia crasipes – Nelumbo nucifera
P3: Nymphaea lotus - Nelumbo nucifera P4: Trapa bispinosa Panicum paludosum
SULEMANKI
P2: Eishornia crasipes - Nelumbo nucifera P1: Nelumbo nucifera - Cyperus diformes
P3: Arundo donax –Typha angustata P4: Nelumbo nucifera – Cyperus difformis
(P indicates pond at different study sites)
49
4.3. FOOD AND FEEDING
4.3.1. Food:
The summary of the data on the feeding preferences of Pheasant-tailed Jacana has been presented in Table 4.8. The stomach contents analysis suggests that 39.8% (49.3% of identified part) of the food of the adult birds of this species is contributed by the animal matter, while the rest of 41.0% (50.7% of identified parts) come from the plant origin. Thus the animal and plant contribute almost equally as food of the Pheasant-tailed Jacana breeding in Punjab, Pakistan.
No stomach contents of chicks were included in the sample and hence the present results have
no bearing on the diet of the chicks.
Out of the animal part of the food of the species, larvae of Chaoborus sp. was the priority food
constituting (12.6%) of the stomach contents, followed by the carpenter ant (Camponotus sp.),
which was the second largest contributor (9.3%) in the diet. Small mollusks contributed 5.4%
and Donacia piscatrix (4.8%) was the least consumed.
Out of the plant matter, seeds of different water plants provided 7.9% of the food, while blades
of the marsh grasses (Spartina sp.) formed some 4.7% of the food. The anthers of the
unidentified aquatic rooted angiosperms were also picked up by this bird species and
contributed an average of 3.6% in its summer diet and floating filamentous algae provided
2.9% in the diet.
50 Table 4.8: Feeding preference of Pheasant-tailed Jacana observed at study sites during summer months.
Food Species Food Availability Preference Stomach Direct (%) index contents (%) Sighting (contents/ availability)
Animal 39.8 27.3 1.49
Chaoborus sp
(larvae) 12.6 - 3.2 3.93
Carpenter ant 9.3 + 10.2 0.91
Mollusks 5.4 + 9.3 0.58
Donacia piscatrix 4.8 - 4.6 1.04
Unidentified animal matter 7.7
Plant Matter 41.0 56.3 0.73
Seeds 7.9 - 12.4 0.63
Blades of marsh grass 4.7 - 9.3 0.51
Anthers
(unidentified) 3.6 + 9.7 0.37
Algae (filaments) 2.9 - 6.2 0.46
Unidentified plant matter 21.9 - 18.7 - unidentified 19.2 - - -
+ Presence, - Absent
51
4.3.2. Feeding behavior
The observation carried out on different age groups (Table 4.9) suggest that the chicks went for the maximum number of picks (29 attempts / minute), followed by adult birds (27 attempts / minute), juveniles (23 attempts / minute) while the minimum number of attempts were made by sub-adults (21 attempts / minute).
The sub-adults traveled the longest distance in search of food (17m/ minute), followed by juveniles (15 m / minute) and adults (12 m/ minute). The chicks moved the minimum and traveled only 8 m/ minute (Table 4.9).
The successful engulf rate of the picked food was the highest in adults (70.37%), followed by juveniles (65.22%) and Sub-adults (61.90%), which chicks (58.62%) had a comparatively low successful engulf rate (Table 4.9)
Table 4.9: Feeding behavior of different age groups of Pheasant-tailed Jacana observed at study sites.
Chicks Juvenile Sub-Adult Adults
Time observed (sec) 60 60 60 60
Attempts for feeding (#) 29 23 21 27
Distance traveled for feeding (m) 8 15 17 12
Engulf rate (%) 58.62 65.22 61.90 70.37
52
The food consumed per unit body weight was the higher in juveniles (19% of the body
weight), chicks (27%), and sub-adults (16), as compared with the adult birds (12%) (Table
4.10).
Table 4.10: Food intake to body weight ratio at different life stages of Pheasant-tailed Jacana in Punjab (Pakistan).
Chick Juvenile Sub-Adult Adult
Body weight (average, gm) 18 47 78 160.32
Food intake (average, gm) 3.24 8.93 12.48 19.24
Food intake / body weight (%) 27 19 16 12
The relative time spent in feeding activities during early breeding and late breeding periods slightly varied with the sex and age (Table 4.11). The females spent almost equal proportion of the time in feeding during early (42%) and late (41%) part of the breeding activities. However, male birds spent relatively a higher proportion of time in feeding during late breeding periods
(53%) as compared to that of during their early breeding period (38%). Juveniles spent a higher proportion of the time in feeding (49%) activity as compared to the adult males and females.
53 Table 4.11: Time consumed for feeding and resting during early and late breeding seasons by Pheasant-tailed Jacana in Punjab (Pakistan).
Early Breeding Late Breeding
Feeding Resting Feeding Resting
Male 38 22 53 8
Female 42 18 41 23
Juvenile 49 38 - -
4.4 BREEDING
The data collected on the breeding parameters of the Pheasant-tailed Jacana population summering between 2004 and 2006 at four water bodies under the present study has been presented in Table 4.12
The table suggests that females laid different number of the clutches during a breeding season, which ranged between 1 and 4 with the average remaining around 2 (n=180). The average number of the clutches laid at Marala (n=13, 2.00 ± 0.31), Qadirabad (n=19, 2.00 ± 0.29),
Balloki (n=08, 1.96 ± 0.41) and Sulemanki (n=12, 1.92 ± 0.37). These were not significant at
0.05% level.
The number of eggs laid per clutch varied between 2 and 4 (n=180), with a clutch observed having 5 eggs at maximum. The average number of the eggs produced per clutch was the
54 highest at Balloki (4.06 ±0.39), followed by Marala (3.71 ± 0.36), and Qadirabad (3.47 ± 0.40),
while the smallest size of the clutch was observed at Sulemanki (3.36 ± 0.28). The average
number of clutch produced at different localities was not significant different (p > 0.05), except
between Balloki and Sulemanki, where it was slightly significant (p < 0.05).
The field observations suggested that the incubation was totally attended by the males,. The
successful incubation lasted for 25 – 29 days (n=180). The average incubation period was
significantly higher (p < 0.05) at Balloki (28.0.2 ± 0.046 days), while the other three localities,
i.e., Qadirabad (27.14 ± 0.30 days), Sulemanki (26.97 ± 0.23 days), and Marala (26.87 days) were not significantly different from one another.
The average number of the eggs hatching per clutch was the highest at Marala
(3.00 ± 0.22), followed by Qadirabad (2.87 ± 0.29), Sulemanki (2.81 ± 0.46) and the lowest number of the eggs hatched per clutch was at Balloki (2.76 ± 0.41). The different localities were not significantly different for the number of the eggs hatched per clutch. The successful hatchability of the eggs was the highest at Sulemanki (83.7%), followed by Qadirabad
(82.7%), and Marala (81.8%), and the minimum hatchability was recorded for Balloki (68.0%).
Table: 4.12: Over all breeding period of Pheasant Tailed Jacana during 2004 to 2007. N Clutches Hatchery in year Eggs success Hatchability Incubation Mortality (%) (#) (#) (#) (days) (#)
Marala 36 2.00±0.31 3.71±0.36 81.8 3.00±0.22 26.86±0.26 1.00±0.31
Qadirabad 67 2.00±0.29 3.47±0.4 82.7 2.87±0.29 27.14±0.3 1.00±0.28
Balloki 29 1.96±0.41 4.06±0.39 68.0 2.76±0.41 28.02±0.36 2.00±0.48
55 Sulemanki 48 1.92±0.37 3.36±0.28 83.7 2.81±0.46 26.97±0.23 1.00±0.32
Depending upon the dates recorded for different breeding activities, the overall chronology of the breeding period of the species was developed.
56 4.4.1 General Behavior
The field observations, as and when recorded, on behavior exhibited by Pheasant tailed Jacana
are summarized as:
4.4.2 Defensive response
The bird developed an alert posture when a buffalo passed through the area at a distance of
some 50 m. Such a posture lasted for 40 second. The concern of the bird gradually increased
when the introducing animal approached a distance of some 25 m, and the normal posture was
retained as the intruder as the intruder distanced at 50 m or more away from the bird.
A three days old chick was observed responding indifferently to an approaching boat
when the boat was at a distance of 30 meters, the chick jumped into the water and camouflaged
himself by keeping most of his body submerged with nostrils exposed from the surface. The
chick even did not attempt an insect, which remained at distance of some four inches, in order
to keep himself motionless at camouflaged.
4.4.3 Nest site selections
The available data on the distribution of 180 nests suggest that 47% of the nesting occurred on
Nymphae lotus plus Eichhornia crassips leaves, and 23 of the nesting was recorded on Trapa
nut and Eichhornia leaves, 7% of the nesting appeared on Pistia and Eichhornia leaves, while
the other type of the vegetation supported only a smaller proportion of the nest.
The eggs were placed in all the nests with the narrow ends touching the nest and the broad end remaining away. The eggs were placed in a straight line, and incubated under the belly of the
male, both wings keeping tight. The distance between the two clutches laid by same female
57 remained around 50 m and the second clutch was laid after an interval of 21± 1.6 days (n =7, ranges(16 to 27 days).
4.4.4 Agonistic behavior
During incubation the breeding pair remained at a maximum distance of 20 m from nest, and with in a distance of 10 m under normal conditions.
During study crow (Corvus splendens), Marsh Harrier (Circus aeruginosus), and kite (Milvus migran) were observed attempting to rob on eggs and chicks. No successful robbing attempt was however recorded .However, mobbing behaviour with crows was very commonly noticed during course of study.
In August and September the leaves of Nyphae lotus and Trapa bispinosa change color from green to brownish which provided a better camouflage for the chicks.
4.4.5 Parental care
A male with four chicks were observed reacting to the intruder to a distance of some 12 m.
The male adopted an alert posture, produced alarm call to the chick, tried to gether the chick trying to bring. He could manage to get one of the chicks close to him, but was able to duly alert others. The chicks kept relaxed posture keeping their eyelid closed and depend on their parents call. The calls of the poreal pauses for longer duration they open their eyes.
During the rain, the male parent was seen protecting the chicks, keeping them under his wings, and the group remained motionless.
58 5. DISCUSSION
5.1. POPULATION STUDIES
The Pheasant-tailed Jacana is a wader and known to be associated with the relatively shallow
water bodies having a good growth of the rooted plants with surface floating vegetative parts
(Roberts, 1991). The bird species has been reported from the flooded fields with rice
cultivation (Mirza, 2007). With the development of the barrages and dams, the man-made lakes
have been created and the flood plains have contracted. Under the circumstances the major
population of the waders and waterbirds has shifted to such lakes. The initial reconnaissance of
the potential water bodies suggested that the major part of the population of the Pheasant-tailed
Jacana was present on the ponds associated with the Marala, Qadirabad, Balloki and
Sulemanki head-works. The lakes associated with the more westward located head-
works/barrages did not support an appreciable population of this species, which probably did
not exceed 50. The major part of the present study was thus concentrated on ponds appearing
on these four water bodies. No recent study is available on the summer counts of the bird taxa
associated with water bodies of Pakisan or the Punjab. The winter counts (Azam et al, 2008)
reported spotting of 4 birds of Pheasant-tailed Jacana at Chashma Barrage. Further studies are
eminently required to know the present distribution/status of this species in water bodies of the
Punjab/ Pakistan.
The present study suggests that a minimum of 300 – 400 birds (around 150 – 200 pairs) of
(Pheasant-tailed Jacana regularly summers in different water bodies. There are indications to
suggest that another 100-150 birds do summer at scattered location of the Punjab, as small
population. With the present information in hand a population of 400-550 Pheasant-tailed
Jacana is presently exploiting the water bodies of the Punjab during summers. No previous
estimates are available on the population of this jacana species summering in Punjab or
59 Pakistan. Casual distribution notes have, however, appeared at different times suggesting a
wide distribution of the birds of this species in the Indian subcontinent (Baker et al., 1929;
Oliver, 1962; Grizmek 1972; Woodcock, 1980; Rao, 1981; Roberts, 1991; Hasan, 2001; Lal,
2004; Mirza, 2007). The present one is the first report directly attempting a population study
on the summering population of this species in the Punjab. The population census on the water
birds carried out in winter suggested a population of 32 (1977), 0 (1998), 452 (1999), 477
(2000), 410 (2001), 334 (2002), 76 (2003) and 4 (2004) (David and Mundkar, 2004). Pheasant-
tailed jacana is predominantly summer visitor to Pakistan (Ali and Reply, 1987) and these
estimates to the population wintering in the Southern part of Pakistan, i.e. the province of
Sindh. The population of this bird species has been reported to remain below 100 in Taiwan
(Humiititrou and Tateisi, 1940). Similar small population (41 – 62 birds) of the Bronze-winged
Jacana (Metopidius indicus) (41-62) was observed at Vembanur Lake (Butchart et al, 1998).
The Jacana, as a group, requires very specific habitat, therefore the favourable habitat patches
for the species are limited, and hence a very large populations are not expected due to habitat
shrinkage.
The major part of the population of the Pheasant-tailed jacana, summering/ breeding in the
Punjab (Pakistan) comes from the southern latitudes, mainly from the province of the Sindh.
The population of the species appears to concentrate in the water bodies falling under the
Sindh province where it is believed to be residents (Roberts, 1991). It appears that population wintering in the Sindh scatters over a wider area during summers for the breeding purposes. A detailed country wide census during summer and winter is required for want of estimates on total wintering and breeding pairs in Pakistan. However the population and its breeding habitat in Punjab province needs exceeding conservation measures.
60 Not many studies are available on the annual or seasonal fluctuations in the population of any jacana species. In his general remarks, Deignan (1931) suggested an abundance of Pheasant- tailed Jacana in northern Thailand during summer and a higher populations of the species at
Songkhla Lake (southern Thailand) during January – March and a lower populations in the
September-October periods. This may indirectly indicate monthly population fluctuations
observed under the present study. The pattern of breeding population fluctuation, as suggested
in present study, indicates an initial build up of population during March – April, which can be justified through a gradual movement and influx of populations to the northern latitudes. The population appears to move gradually in and becoming stable in July – August, which can be associated with the major part of the breeding activities. The subsequent rise till October can be attributed to the addition of the new recruits into the population. Thereafter, it is hard to find any bird in Punjab in November onward till end of spring.
The movements between the summering (breeding) and wintering (non-breeding) grounds in the Pheasant-tailed Jacana can not be regarded as strict migration, demanding complete geographic isolation of breeding and non-breeding grounds with well defined areas where population of passage birds of the species can be observed only during migratory movements
(Odum, 1971). These movements can thus be regarded as local movements/population adjustments. Definite reasons for such movements are not known. Further study is required to analyze the reasons of such south to north movement of this species in the Punjab. The
presence of relatively moderate temperature in relatively northern latitudes of the Punjab, can
be one of the reason for such movements of the population during summer. Requirement of a
relatively wider area for the breeding population of the species can be the other reason for such
movements, disturbance and alteration of the habitat can be another reason.
61 A non-significant difference between the population’s levels of Pheasant-tailed Jacarna during three study years, and non-significant value of heterogeneity chi-square suggest that the population remained stable over the years. This can be regarded as a positive sign for the future long time survival of the species in these breeding grounds and enhances the importance of these sites for conservation purposes.
The present results have suggested a significant preponderance of males in the population, with a male to female ratio of 1.33. This is characteristics of the jacana, where a male to female sex ratios of 1.9 (Butchart, 1998), 2.2 (Jennie and Collier, 1972) and 1.43 – 2.22 (Stephen et al., 2004) have been suggested previously for Bronze-winged Jacana (Metopidius indicus). No report is in hand on the Pheasant-tailed Jacana, but the sex ratio remains within the range suggested for the Bronze-winged Jacana. Females having a larger body size and brighter colours, are believed to share more of the predation pressure. This is a successful strategy, where the males, which are responsible for incubation and care of the chicks, are saved from facing the undue predation pressure. Whereas, in most of the cases, a higher mortality of the males has been frequently reported in species where the males are larger and more colourful and females attend the duties of rearing of the offspring (Roberts, 1991). Not much is known about the male! Female sex ratio and predation pressure in wintering populations of this jacana species.
The juvenile to female ratio (0.82, juvenile to adult ratio of 0.35), suggests a successful breeding population at study sites. No previous comparable study is available on jacana, but an annual increase of over 30% population is a healthy sign and reflects negligible predation pressure and suggests a strong homeostasis in breeding habitat and population. On the onset of late breeding season the juveniles get into the dispersal phenomena with a few prefer to stay in their breeding haunts.
62
5.2. HABITAT
The analysis of the present distribution of the Pheasant-tailed Jacana largely go in conformity with the general observations appearing in literature that this bird species prefers the ponds having rooted vegetation with surface floating stems/leaves (Grizmek, 1972; Roa, 1991.
Roberts, 1991; Butchart, 1998. Hasan, 2001; Kazmerczak et al., 2006; Mirza, 2007). Changes with the pattern of flood appearing in the rivers to support paddy cultivation and development of established ponds associated with the man-made lakes around the barrages, it appears that the bird has gradually moved to these ponds for breeding, rather then remaining in frequently disturbed paddy fields.
There are reports available suggesting a correlation between physical characteristics of wetlands vegetation and the size of the population of different aquatic birds (Beecher, 1942;
Edward, 1962). A number of reports available suggests a close link with water quality in different wetlands habitat which influence the level of primary productivity ultimately influencing the tropic structures and total biomass throughout the aquatic food web.
(Hutchinson,1957;Lund, 1964. Greenberg, 1964; Summerfelt 1971; Latif, 1983; Baqai and
Rehana 1973; Wetzel; 1975; Chughtai, 1979; Vasishat and Jindal, 1980; Mandel,1980;
Benzie, 1984; Khatri, 1985. Haramis et al., 1986; Saleem 1986; Ali et al., 1988; Kushlan,
1989; Breininger and Smith, 1990; Sampath and Krishnamurthy, 1990; Chaudhry 1991, 1992;
Khan 1992; Velasquez, 1992; Ormered and Tyler, 1993; Weller,1994; Tamisier and Grillas
1994; Mary and Varghese, 1994; Kumar,1995; Ntiamoa and Baidu, 1998; Waran et al 2004;
Christina et al,. 2006; Emmanuel and Onyenmma, 2007; Mahaulpatha et al 2007; of the freshwater bodies in different parts of the work, but specific study on the jacana of Pheasant- tailed Jacana habitat is available. The present study concerns with the direct analysis of the
63 physico-climatic and biotic factors of the ponds holding appreciable populations of the
Pheasant-tailed Jacana. These analyses suggest that the temperature of air/water ranges between 16 and 35 oC. The pH remains slightly basic, while all other physico-chemical
parameters remained within the limits of a freshwater water body. There was non-significant
difference between different water bodies, with some degree of monthly variations. As the
variations between all such parameters were not significant, therefore association of such
factors with the population levels of the Pheasant-tailed Jacana could not be attempted. It appears that these factors constantly remain within certain limits, which are fully adapted by the species and hence are frequently not working as limiting factors to control jacana populations.
The variation in the overall populations of phytoplanktons (algae) or zooplanktons protozoa, rotifers, cladocerans, copepods) was not different between different ponds holding this species of the birds, and there was not much marked monthly variation in their populations during the summers, the breeding period of Pheasant-tailed Jacana in the area. The importance of the planktons in controlling the population of this species could not be determined under the present study. No previous study is available suggested a correlation of such factors with jacana populations. The present study did not attempt a microanalysis of species distribution or fluctuation in the water bodies, which could had provided some useful association with the jacana population levels.
The analysis of the rooted vegetation suggests that there were a total of 23 species of the rooted plants (angiosperm and fern) present along the periphery of the ponds catering jacana population. These species were present in different combinations and provided cover as will as foods were not directly important in controlling the jacana population. One or of the four species of angiosperm rooted plants with their surfacing floating leaves (Nelumbo micifera, N.
64 alba, Nymphaea lotus and Trapa bisoinosa) and five species of the plants species floating just under the surface water (Eishhornia crasipes, Hydrilla verticilata, Vallisnariria spiralis, Psitia stratiotes, Chara spp.) were present in the water having jacana populations. These species are mainly required for the breeding activities and nests are placed on the surface of the leaves of these species, where they are protected from the predators. No quantitative study was attempted on the relative distribution which could be used in the development of association with the jacana population levels. The species, like, Eishornia crassipes, Salvinia molesta,
Imperata cylindrical and Mikania sp. were indirectly associated with the presence of the jacana species (Malaulpatha et al., 2007).
The jacana appears to have adopted to survive in the open body of water rather than finding camouflage in the denser tall rooted vegetation present in the periphery of the pond. The general field observations suggest that the bird perfectly matches with the environment, and with its bright contrasting colour, it matches with the background vegetation giving a look of a flower emerging from the surface of the plant. A still bird hardly visible to a raptor, and remains obscured. The diving habit of the adult bird and chicks at the time of emergency is an additional defense of the bird.
5.3. FOOD
In the breeding season, the adult Pheasant-tailed Jacana places almost equal reliance on the animal and plant matter for meeting the requirements of its food. No previous report on the quantitative feeding preference of this species is available. The general remarks have suggested that this bird species depends upon insects and invertebrates, seeds (Hayman et al., 1986), larvae of invertebrate (Woodcock, 1980) and water chestnut (Butchart, 1998; Lal, 2004). The present information, however, concerns with the breeding population of this Jacana species and
65 has no bearing upon the winter diet of the animals. The animal species generally consumes more of the animal diet during the breeding period which are protein rich and helps meeting the higher energy needs of the breeding birds and the chicks (Ali and Ripley, 1987). The present results have no bearing on the different on diet consumed by chicks, juveniles and sub- adult, and also on the possible variation in the preference of food by the two sexes. Further studies in these aspects will be required for understanding the food requirements of this species of bird.
Roberts (1991), reported that the species preferred to consume insect larvae, and spawns of mollusks, amphibian, bivalve mollusks, and seeds and succulent plants. The present report partly confirms the observations of the Roberts (loc cit.), though it suggests that breeding jacanas place a higher reliance on larvae of Chaoborus sp. larvae, followed by Camponotus sp., while small mollusks and Donacia piscatrrix were consumed in relatively lower frequencies. Seeds of aquatic plants, leaves of marsh grasses were more frequently consumed, while anthers of the plants and floating algae also contributed an appreciable proportion.
The selection of the food by a species of animal is a compromise between the dietary requirements of the animal/animals, the adaptation of the species to efficiently pick upon the food item and the availability of such food items in nature (Odum, 1971). The present report concerns with a specific area and for a specific season and hence may be taken with caution when exproling these results for other areas. Variation can be expected between years, caused by changed availability.
The chicks going for a higher frequency of picks, followed by adults, juveniles and subadults and conversely longer distances traveled by the subadults, followed by juveniles, adults and chicks is new information in feeding behavior of this species. Definite explanation for such
66 behavior is hard to be developed with the information in hand. Further studies are required to
confirm such feeding behaviour.
The present results suggest that the juveniles, chicks and subadults picked up more food in
comparison with their body mass, as compared with the adults. This appears understandable as the energy requirement of the growing birds is higher as compared to that of the adults.
The variation in the proportion of the time spent by females and males on feeding and breeding can be explained on the requirement of the two sexes. The female, being responsible for laying of eggs and defending the territory, requires equally amount of time spent on the food and breeding during early and late breeding periods. The male, however, is responsible for incubation and rearing of chicks and hence to divert more time towards incubation. The depict of energy caused in males during incubation forces then to spend more time on feeding in later parts of the breeding season to keep a balance in their energy budget. Similar time budgeting has been previously reported for this species by Chen et al. (2008-a) based upon their studies on breeding population of Taiwan.
5.4. BREEDING
The present information on the breeding potentials suggests that of the species has a normal breeding behaviour as indicaterd for the species in different reports (Thongaree et al., 1995;
Grizmek, 1972; Thongaree et al., 1995 Butchert et al., 1999). The present results suggest that females lay an average of two clutches, with a range of 1 – 4, and is basically polyandrous. The clutch size ranged between 2 and 4, with mean clutch size remaining between 3-5. One of the exceptional female laid 5 eggs in a clutch. The incubation period ranges between 25 to 29 days.
The hatching success is around 75%, which is an indicator of healthy population buildup.
5.5. MANAGEMENT
67 The information collected during the present study suggests that the population of the
Pheasant-tailed Jacana breeding in the limits of the Punjab, Pakistan, is maintaining a stable
population. The sex ratio is slightly in favour of the males, which is adjusted through its
breeding behavior and polyandrous nature, where male are responsible for incubation and
rearing of chicks. A good juvenile to adult ratio is indicating of a good breeding potential of
the population and a normal recruitment. The species has sufficiently adjusted to a variety of
habitat conditions. Habitats provide ample food resources to cater the demands of the breeding
adult populations and juveniles. All these factors are sufficient to suggest that under the present
conditions the Pheasant-tailed Jacana is a least concerned bird species in the area and there is
no immediate threat to the survival of this species in the area.
The development of man-made lakes and associated ponds have ensured the availability of the
favourable habitat conditions. Such ponds provide a safer refuge for the bird, as compared with
the neighboring rice fields, or temporary ponds, appearing close to human settlements. The
jacana population is probably safer than it was in past, when habitat fluctuations and pollution
was a likely threat for the species survival.
Under the existing circumstances, the Pheasant-tailed Jacana does not require any human
intervention for the continued survival of the species in this part of the globe. However, constant monitoring of the population status and its viable breeding habitat conditions need to
be maintained and the wintering grounds needs to be studied for further analysis.
68
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79
ANNEXURES
80 ANNEXURE 1: Mean ±standard error of mean of different physico-climatic parameters
of water bodies during different calendar months between April 2003 to March 2007 at
Marala (Punjab, Pakistan).
Temperature (0C) pH
Air Water
April 19.97±0.71 17.27±0.32 7.90±0.23
May 32.67±0.65 27.83±1.12 7.43±0.09
June 36.87±0.84 33.77±1.10 7.13±0.46
July 31.63±0.59 27.73±0.62 7.80±0.29
August 31.33±0.68 25.80±1.21 8.03±0.64
September 28.87±0.35 23.63±1.16 8.37±0.18
October 26.3±0.50 21.3±0.31 7.3±0.29
November 20.87±0.76 17.7±0.64 7.63±0.26
December 17.73±0.69 15.33±0.43 7.5±0.17
January 10.7±0.53 8.2±0.47 8.0±0.34
February 12.57±0.70 9.27±0.84 7.30±0.12
March 17.83±0.64 13.70±0.53 7.90±0.23
81 ANNEXURE 2. Mean standard error of physico-climatic parameters of water during April 2003 to March 2007 at Head Qadirabad.
Air Water pH
temperature temperature
April 20.3±0.72 17.63±0.8 7.73±0.2
May 32±0.351 27.1±0.94 7.67±0.12
June 39.67±0.66 34.13±0.54 7.67±0.39
July 34.23±0.80 29.47±0.38 8.07±0.34
August 31.17±0.64 27.93±0.38 8.17±0.18
September 29±0.71 25.27±0.26 7.67±0.12
October 6.2±0.75 21.87±0.42 8.47±0.22
November 21.37±0.61 18.70±0.64 7.43±0.07
December 18.57±0.66 15.83±0.78 7.67±0.33
January 12.03±0.96 9.37±0.66 8.53±0.20
February 13.9±0.72 10.3±0.31 7.47±0.22
March 17.57±0.47 15.97±1.25 8.13±0.35
82 ANNEXURE 3. Mean standard error of physico-climatic parameters of water during April 2003 to March 2007 at Head Balloki.
Air Water pH
temperature temperature
April 21.37±1.04 18.00±0.96 7.20±0.17
May 32.63±1.21 28.00±2.33 6.80±0.21
June 40.13±0.83 34.13±0.84 7.17±0.38
July 34.53±0.66 30.40±0.61 8.47±0.22
August 32.17±0.68 27.33±0.49 7.70±0.40
September 29.53±0.38 24.97±0.55 7.87±0.54
October 25.67±0.97 23.50±0.74 8.13±0.32
November 22.53±0.64 19.60±1.27 7.83±0.71
December 19.43±0.82 17.77±1.03 7.80±0.21
January 13.03±0.82 9.70±0.46 8.63±0.49
February 15.53±0.95 11.90±0.87 8.23±0.58
March 18.47±0.83 15.23±0.98 7.70±0.47
83 ANNEXURE 4. Mean standard error of physico-climatic parameters of water during April 2003 to March 2007 at site 2 (Head Sulemanki)
Air Water pH
temperature temperature
April 18.97±0.87 16.3±0.95 7.67±0.32
May 32.83±2.32 29.17±1.98 7.30±0.12
June 41.37±1.96 39.2±5.30 7.10±0.44
July 35.40±1.19 30.93±1.43 7.83±0.55
August 32.33±0.64 27.80±1.32 7.47±0.20
September 28.67±0.91 23.77±0.98 8.07±0.19
October 24.67±0.94 20.20±0.96 8.00±0.26
November 21.30±0.21 17.50±0.90 7.93±0.38
December 19.83±0.93 15.83±1.03 7.57±0.24
January 13.13±0.38 9.97±0.55 8.67±0.37
February 15.30±0.72 10.93±0.45 6.97±0.39
March 18.57±0.41 15.27±1.06 8.57±0.8
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