let o fl ^ilasiopli^ · brothers jirif %han, 94usadiq (rafiq, musashir shamim, (dr. owais yaqoot...
TRANSCRIPT
r PRECISION OF HARD STRUCTURES US ESTIMATE AGE AND GROWTH OF SL
FRESHWATER TELEOST
DISSERTATION
SUBMITTED IN PARTIAL FULFILMENT OF THE R FOk THE AWARD OF THE DEGREE OF
le t ol f ilasiopli '•f-.
'•H^ ^^>^ 3n f-1
\ ^
SABAH ^jf'
UnderIhe Supervisior
DR. MOHAMMAD AFZAL KHAN
FE SCIEN MUNIVEh
£.\f i ^
3 0 JAN 20i5
DS4408
'SaM/f
Phone< External: 2700920/21-3430 Internal : 3430, 3431
DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY
ALIGARH - 202 002 INDIA
Sections: 1. ENTOMOLOGY , er^FISHERY SCIENCE & AQUACULTURE 3. GENETICS 4. NEMATOLOGY i P - /^U 1 ^ 5. PARASITOLOGY Dated. \9. ftH". )...T^.,
D. No /ZD
CERTIFICATE
This is to certify that the dissertation entitled "Precision of hard structures used to
estimate age and growth of selected freshwater teleost" has been completed under/ :" >; f . I ' . , , •
my supervision by Ms. Sabah. The work is original and independently pursued by the
candidate. It embodies some interesting observations contributing to the existing ' >i,
knowledge on the subject.
I permit the candidate to submit the dissertation work for the award of M. Phil.
Degree in Zoology.
riiv ^^'^ (Dr. Mohammad Afzal Khan)
Assistant Professor
Jic^owCecfgement
I cypress my sincere gratitude to J^CmigHty Mlafi who Ras given me strength to complete
tfiis worH^successfufCy and produce in its present form.
It is my priviCege and gives me immense pleasure to express sound perception of
gratitude to my Supervisor <Dr. Mohammad^fzal%jian for fiis continuous encouragement,
vafuaSCe and constructive suggestions. 'Wor^ng under his guidance is a matter of great
privifege.
I extend my thanlis and gratitude to (prof. Iifan ^hmad, Charrmait of the
(Department ofZoofogy, Jl.M.'V, for providing aCC necessary faciCities reiatedto my work^
I wouldatso [ike to thanh^<PTof. IqSaC^afvez, <Prof. Mu^tar_^hmad%han and
(Dr. SaCtanat (Parveenfor their vafuaBfe advice.
Special thanlis are also due for my seniors (Dr. Jiagma, !Mr %aish 'Miyan, iM.s.
Shahista 'Kfian, coffeagues Safman Kfian and_Afaq [Hazir Shah and to my friends Sumu\d
JAhad, J4rjumend Shaheen, !Kida %i^ani, Saima hameed, SharSa %ausar, Sayma Samreen.
SyedSuraya, (pyiiqaui TaSasum, ^iaffia 'Mustahj-m, Dania j^hmed, (Riishda Sharaf, 'Taihd
Saeed, Tarzana Imtiaz, Vzm.a Vsmain, Jlh. Latif 'Wain, !M. (Farced, Sajad (Bhat and
(prince Tariquefor their unconditional and constant help.
I am extremely proud to achjiowledge the contriSution of my family especially Dr.
Ahdul (p^sliidSheilfi, Kilal 9dali({^ Javaid Malil{, (RuSina 9dalih^ P^uhi'Malih^ C'handi
Mushtaq and (Dr. 'Mushtaq Wani and, sisters Shiraza Wani, Jdaadi !Mir, 9Audasir (Rgfiq,
(Dr <Enas Mushtaq, Jdafsa Maliki (Rayeesa Mali^i Madhiya 'Malih^ and yfuha 'Malif^
Brothers Jirif %han, 94usadiq (Rafiq, MuSashir Shamim, Yaqoot (Bhat, (Dr. Owais
Mushtaq, JLthar Malill and (Riyaza 'Wani Last, hut not the least, I am privileged to
express my deepest thanlis to the most precious gift of god, my loving parents and
grandparents for Being a source of unseen. But not unheard strength that installed courage
in me to complete this worh^
(SaBah)
CONTENTS
Particulars Page no.
INTRODUCTION 1-8
A SHORT DESCRIPTION OF FISHES 9-14
MATERIALS AND METHODS 15-20
RESULTS 21-25
DISCUSSION 26-33
SUMMARY 34
REFERENCES 35-53
APPENDIX
LIST OF TABLES
No. Title
1 Comparison of percent agreement (PA), average percent error (APE) and
coefficient of variation (CV) between the age readings of two independent readers
in Schizopyge cunnfrons, Schizopyge niger and Schizothorax esocinus.
2 Comparison of percent agreement (PA), average percent error (APE) and
coefficient of variation (CV) among ages assigned between structures iii
Schizopyge cunifrons, Schizopyge niger and Schizothorax esocinus.
3 Comparison of Pearson's correlation coefficient values in Schizopyge cuni/rons.
Schizopyge niger and Schizothorax esocinus.
4 Comparison of mean values of age estimates from different bony parts in
Schizopyge ciirvifrons, Schizopyge niger and Schizothorax esocinus.
5 von Bertalanffy growth parameters along with growth perfomiance index ( o ) in
Schizopyge curvif'rons, Schizopyge niger and Schizothoixix esocinus.
6 Estimated parameters of length-weight relationship (LWR) for Schizopvgc
curvifrons, Schizopyge niger and Schizothorax esocinus.
7 Length-length relationships (LLRs) between total length (TL), fork length (PL)
and standard length (SL) for Schizopyge curvifrons, Schizopyge niger and
Schizothorax esocinus.
LIST OF FIGURES
No. Title
1 Age bias plots for age comparisons between readers for scales, opercular
bones, otoliths, vertebrae and cleithra in Schizopyge cunifrons.
2 Age bias plots for age comparisons between readers for scales, opercular
bones, otoliths, vertebrae and cleithra in Schizopyge nigcr.
^ Age bias plots for age comparisons between readers for scales, opercular
bones, otoliths, vertebrae and cleithra in Schizothorax esocinus.
4 Age bias graphs for age comparisons between structures in Schizopyge
curvifrons.
5 Age bias graphs for age comparisons between structures in Schizopyge nigcr.
6 Age bias graphs for age comparisons between structures in Schizothorax
esocinus.
7 von Bertalanffy growth curves for Schizopyge curvifrons, Schizopyge niger
and Schizothorax esocinus showing back-calculated length at estimated age.
INTRODUCTION
Fishes are considered to be an excellent source of high quality protein, other essential
nutrients and minerals that are often difficult to obtain from many other sources. As
per FAO reports (2012), fish is the most valuable agricultural commodity traded
intemafionally with annual sales of nearly US$217.5 billion in the year 2010 and
increasing each year. Fisheries and aquaculture provided livelihoods and income for
an estimated 54.8 million people engaged in the primary sector of fish production in
2010, of which an estimated 7 million were occasional fishers and fish fanners. Asia
accounts for more than 87 percent of the world total followed by Africa (more than 7
percent), and Latin America and the Caribbean (3.6 percent). Capture fisheries and
aquaculture supplied the world with about 148 million tons offish in 2010. In 2009.
fish accounted for 16.6 percent of the global population's intake of animal protein and
6.5 percent of all protein consumed. Inland fisheries are a vital component in the
livelihoods of people in many parts of the world, in both developing and developed
countries. Total global inland capture fisheries production has increased dramatically
since the mid-2000s with reported (as submitted by countries) and estimated (by FAO
in cases of non-reporting countries) total production at 11.2 million tonnes in 2010, an
increase of 30 percent since 2004. Growth in the global fish calch from inland waters
is wholly attributable to Asian countries. With the remarkable increases reported for
2010 production by India, China and Myanmar, Asia's share is approaching 70
percent of global production (FAO, 2012).
The growth in fish catch from inland waters is also accompanied by drastic
decline in the number of several commercially important fish species from natural
environment. Inland waters are considered as being overfished in many parts of the
world, and human pressure and changes in the environmental condifions have
seriously degraded important bodies of freshwater (FAO, 2012). Freshwater fishes in
India are under threat for several reasons, but primarily due to unsustainable and
unethical fishing practices. Though there are only a few species of fishes that are in
trade, the state of all fi-eshwater fishes in India is in danger because of wrongful
methods of fishing. Other common threats that affect freshwater fish populations are
habitat loss due to dredging of lakes and rivers, filling, altering river courses, dams,
irrigation canals, and other reasons. Of the three species selected for the study, two
1
species are listed Vulnerable {Schizopyge curvifrons and Schizopyge niger) and one as
Lower risk-near threatened (Schizothorax esocinus) (Molur and Walker, 1998).
Studies on biological parameters of threatened fishes are highly significant for
management and conservation of populations in natural water bodies (Sarkar et al.,
2009; Hossain et al., 2012; Khan et al., 2012b).
India is one of the richest countries in terms of freshwater resources. India has
vast freshwater resources in the form of ponds, tanks, lakes, reservoirs, rivers, bheels,
canals, swamps, marshes, etc. Ponds and tanks account for about 2.25 million ha.
Lakes comprise 26 million ha, while reservoirs (major and medium) have spread over
an area of 3.0 million ha.'Rivers have a total length of more than 27,000 km. Canals
and channels and their network run up to 112,000 km. Bheels (water bodies fonned
due to the serpentine course of a flooded river), marshes and swamps constitute 7.9
million ha (Basavaraja, 2007). The water resources of Kashmir can be broadly divided
into lotic and lentic systems. The fomier comprises of the river Jhelum and various
streams which directly or indirectly join it, while the latter includes lakes, ponds,
wetlands and other similar aquatic habitats. River Jhelum flows from south to north
west up to Wular, from where it takes south westerly direction. The total length of the
river from the Veerinag up to Uri is about 239 km. There are a number of mountain-
and valley-lakes in Kashmir. Some of these are Dal, Anchar, Manasbal, Gangabal,
Wular lakes. The variety of ton-ential hill streams, rivers and lakes support good fish
diversity. These aquatic resources have been meeting the requirement of fishes for the
local people over the time immemorial. Over the years various changes have taken
place in the species composition, distribution and availability of fishes in the water
bodies of this region. The streams and lakes of the valley harbour a number of
indigenous fishes belonging to genus Schizothorax, Schizopyge, and other carps that
serve as an important food item for human consumption.
Studies on fish biology particularly morphometry, length-weight relationship,
condition factor, reproduction, food and feeding habit, etc. are important not only to
provide additional information to the existing knowledge but also in the utility of the
knowledge in increasing the technological efficiencies of the fishery entrepreneurs for
evolving judicious fisheries and aquaculture management. Age and growth analyses
of fish species are used to deteimine longevity, mortality, productivity, yield and
population dynamics, which in turn are essential for responsible management
(Holden, 1972; Hale and Lowe, 2008). Age estimation in fishes have been undertaken
using different hard structures such as, scales (Singh and Shamia, 1995; Andreu-Soler
et al., 2003; Aydin et al , 2003; Dua and Kumar, 2006; Karatas et al, 2007; Zhang
and Takita, 2007; Sarkar et al., 2008; Ozcan and Bahk, 2009; Ujjania, 2012), fin rays
and spines (Ezenwa and Ikusemiju, 1981; Morrow Jr. et al., 1998; Stevenson and
Secor, 1999; Kano, 2000; Sun et al., 2002; Mills and Chalanchuk, 2004; Penha et al,
2004; Whiteman et al., 2004; Metcalf and Swearer, 2005; Santamaria et al., 2009;
Tribuzio et al., 2010), otoliths (Doray et al., 2004; Monteiro et al., 2006; Qiu and
Chen, 2009; Ilkyaz et al, 2010; Ma et al., 2010; Jia and Chen, 2011; Huo et al., 2012).
opercular bones (Bardach, 1955; Qasim and Bhatt, 1964; Jellyman, 1980; Nargis.
2006; Gomez-Marquez et al, 2008), vertebrae (Guinn and Hallberg, 1990; Polat et al.
2001; Coelho and Erzini, 2002; Licandeo et al, 2006; Hale and Lowe, 2008; Kume et
al, 2008), cleithra (Govind and Gopal, 1966; Harrison and Hadley, 1979; Casselman,
1990) and some other bony parts. Scales have been widely used for ageing because
they are collected, prepared and read easily (Gursoy et al, 2005). However, several
researchers have reported that scales can provide unreliable estimates of fish age
(DeVries and Frie 1996; Maceina and Sammons, 2006), which has often been
attributed to reabsorption and deposition of false annuh due to stress and food
limitation and also because clarity of annuli is affected with increase in fish age (Khan
et al, 2011a) and have, therefore forced fishery scientists to use other calcified
stiTictures (Hammers and Miranda, 1991; Zoubi, et al, 2010). It is now well
established that a suitable structure for age estimation \'aries by species and
geographic location (Koch et al, 2009), thus the evaluation of the precision and
accuracy of bony structures should be studied (Polat et al, 2001). Precision is defined
as the reproducibility of repeated measurements on a given structure or the
repeatability of individual readings or age estimates, whether or not those
measurements (age readings) are accurate. It is not unusual for inaccurate age
readings to be highly reproducible (in other words, precisely wrong) or to show no
relationship between accuracy and precision (Campana et al, 1990; Campana and
Moksness, 1991; Campana, 1995). Therefore, precision cannot be used as a proxy for
accuracy. Nevertheless, a measure of precision is a valuable means of assessing the
relative ease of determining the age of a particular structure, of assessing the
reproducibility of an individual's age determinafions, or of comparing the skill level
of one ager relative to that of others (Campana, 2001). However, higher precision of
age estimates provides no insight into the biases or accuracy of age estimates among
or between structures. The application of a validation method is required for accurate
age estimations (Beamish and McFarlane, 1983), but the method of counting annuli in
hard structures and measuring the level of precision between structures and between
researchers is the best way to corroborate age estimations if a method of validation
cannot be established (Kimura et al., 2006; Gumus et al., 2010). Counting growth
increments (annuli) on otoliths, scales or other hard structures has been the most
common method for ageing fish; however, age estimation is often accompanied by
several sources of error that can have significant effects on many population
parameter estimates (Beamish and McFarlane, 1983) e.g., age underestimation may
result in optimistic estimates of growth and mortality rates, leading to the serious
overexploitation of the population and its eventual collapse (Campana, 2001).
Comparison of age estimates from various bony structures has been reported in a
number of fishes to identify the most suitable structure for a fish population (Gocer
and Ekingen, 2005; Sylvester and Berry, 2006; Jackson et al., 2007; Phelps et al.,
2007; Khan and Khan, 2009; Khan et al, 201 la, b; Ma et al, 2011).
However, there are no published reports available on precision of different
structures for ageing fishes selected for the present study {Schizopyge curvifrons,
Schizopyge niger and Schizothorax esocinus).
Age and growth are always used together in phraseology, but each terni has its
own distinct meaning, which, as cited by Goldman (2004), was eloquently stated by
DeVries and Erie (1996): "Age refers to some quantitative description of the length of
time that an organism has lived, whereas growth is the change in body or body part
size between two points in time, and growth rate is a measure of change in some
metric of fish size as a fimction of fime". Growth is one of the most important life
history processes influencing the fish population dynamics (Zhan, 1995). For a given
fish species, reliable estimation of growth is critical in developing sustainable
fisheries stock assessment and management (King, 1995; Gang et al, 2008). The main
objective of age and growth studies in fish is to estimate their mean size at each age
class and determine their growth parameters (Mather et al, 1995; Santamaria et al,
2009). Estimates of growth drive size- and age-structured stock assessment models
(Quinn and Deriso, 1999), scale yield calculations (Beverton and Holt, 1957; Gulland,
1983; Haddon, 2001) and are related to life history traits such as natural mortality and
4
age or length at maturity (Chamov, 1993; Jensen, 1998). Besides pure application, age
and growth studies are also important in describing the basic biology and ecology of
fishes (Weatherley and Gill, 1987; Cailliet and Goldman, 2004; Cope and Punt,
2007). Growth rate estimated trom length-at-age data are considered to assess fish
stocks with relatively higher accuracy than the method based on length- frequency
data. Fish growth data can be easily fitted with an appropriate mathematical fiinction
to generalize the growth process, so as to predict the growth trend and compare the
growth patterns among stocks or species (Rao, 1958; Gang et al., 2008). Several
mathematical models can be used for modelling fish growth, however, the von
Bertalanffy growth function (VBGF; von Bertalanffy, 1938) is by far the most studied
and most widely used of all length-age models in fisheries, and its parameters are
particularly useful in describing general fish growth (Chen et al., 1992; Quinn and
Deriso, 1999), deriving fisheries reference points (Clark, 1991; Williams and
Shertzer, 2003) and estimating life history parameters (Beverton and Holt, 1959;
Beverton, 1992; Cope and Punt, 2007). Since the incepfion of the VBGF, there has
been a steady evolufion in its use and estimation. Beverton and Holt (1957)
popularized the current form of the VBGF by changing the original three-parameter
fonnulation to include to rather than Lo. Several estimation techniques, ranging from
linear fitting (Walford, 1946; Stamatopoulos and Caddy, 1989) to nonlinear least
squares (Tomlinson and Abramson, 1961; Marquardt, 1963), iterative fitting (Rafail,
1973), and likelihood (Kimura, 1980) methods have subsequently been proposed
(Cope and Punt, 2007).
The growth rate of fish is an essential component of models used in stock
assessment offish populations; small variations in growth rates can have a significant
impact on modelling outcomes for populafion analysis (Megalofonou, 2000;
Santamaria et al., 2009).
Age and growth have been studied in a number of fishes from India such as
Cirrhinus mrigala (Ham.) from the river Ganga (Jhingran, 1959); Ophicephalus
punctatus Bloch from ponds in Aligarh (Qasim and Bhatt, 1966); Labeo rohita
(Ham.) from a pond (Moat) and rivers Ganga and Yamuna (Khan and Siddiqui,
1973); three Indian major carps from Hirakud reservoir (Mathew and Zacharia,
1982); Tachysurus dussu,nieri (Val.) along the Dakshina Kannada Coast
(Vasudevappa and James, 1988); Indian major Carp {Catla catla Ham. 1822) from
northern India (Johal and Tandon, 1992) and from selected water bodies of southern
Rajasthan (Ujjania, 2012); Johnieops sina (Pisces/Percifonnes) from Bombay
waters (Chakraborty, 1994); threadfm bream Nemipterus japonicas (Bloch) off
Bombay (Chakraborty, 1995); Channa marulius from Harike wetland (A Ramsar
site), Punjab (Dua and Kumar, 2006); Turbo brunneus occurring in Tuticorin coastal
waters (Ramesh et al., 2009); Sillago sihama (Forsskal) from Zuari estuary, Goa
(Shamsan and Ansari, 2010); silverbellies along Kerala coast (Abraham et al., 2011)
and Thunnus albacares (Bonnaterre, 1788) (Rohit et al., 2012). There have been few
studies on the age and growth of fishes belonging to sub-family Schizothoracinae viz.
Schizothorax richardsonii from the Garhwal hills, India (Singh and Shamia, 1995),
Schizopygopsis yoimghiisbandi younghiisbandi (Chen et al., 2009), Ptychobarbiis
dipogon (Regan, 1905) (Li and Chen, 2009), Schizothorax waltoni (Qiu and Chen,
2009); Schizothorax o'connori (Yao et al, 2009; Ma et al., 2010), and
Oxygymnocypris stewartii (Jia and Chen, 2011; Huo et al., 2012) in the Yarlung
Tsangpo river, Tibet. Despite the limited distribution of 5. curvifrons, S. niger and S.
esocinus and their importance to fishing, little is known about its biology and ecology.
Sunder and Subla (1984) demonstrated age and growth of Schizothorax cunnfrons by
back-calculation.
Fisheries management and research often requires the use of biometric
relationships in order to transfomi data collected in the field into appropriate indices
(Anderson and Gutreuter, 1983; Ecoutin and Albaret, 2003; Mendes et al., 2004). One
of the most extensively used in any analysis of fishery data is the length- weight
relationship (LWR). Furthennore, Length-weight relationship allows; i) estimation of
average weight of the fish of a given length group (Beyer, 1987); ii) conversion of
growth-in-length equations to growth-in-weight in order to estimate stock biomass
from limited sample size (Tarkan et al., 2009); iii) interspecific and inter populafional
morphometric comparison of fish species (Rajkumar et al., 2006); iv) study of the
ontogenic allometric changes in fish growth (Teixeira-de Mello et al., 2006) and v)
possible effects from parasites (Teixeira-de Mello and Eguren, 2008; Teixeira-de
Mello et al., 2011). Besides this, LWR can also be used in setting yield equations for
estimating the number of fish landed and comparing the population in space and time
(Beverton and Holt, 1957) and in frophic studies (Gonzalez-Gandara et al. 2003;
Sivashanthini, 2008). LWR parameters may change in relation to sex, body size,
6
temporal and / or spatial factors but variations of population structure, food
availability and fishing pressure can also result in different weight-length
relationships for the very same species (Giacalone et al., 2010). Knowledge on LWR
and length- length relationships (LLR) is useful in fish stock and population
assessments (Ricker, 1968). LLRs are important for comparative growth studies
(Moutopoulos and Stergiou, 2002; Hossain et al., 2006). Size conversions (e.g.,
calculated TL from SL) find applied value in fisheries for understanding several
aspects of population dynamics (Froese and Pauly, 2005; Ruiz-Campos et al., 2006).
Length and weight measurements in combination with age data can give
infonnation on the stock composition, age at maturity, life span, mortality,
growth and production (Beyer, 1987; Bolger and Connoly, 1989; King, 1996a, b; Diaz
et. al., 2000; Fafioye and Oluajo, 2005).
Condition factor is a quantitative parameter of the state of well-being of the
fish that will detemiine present and future population success by its influence on
growth, reproduction and survival (Hossain et al., 2006). Fulton's condition factor (K)
is widely used in fisheries and fish biology studies to describe a two-dimensional
weight-length relationship by converting it into a single statistic with the intention
describing the "condition" of that individual fish (Amason et al., 2009). The condition
of a fish reflects recent physical and biological circumstances, and fluctuates by
interaction among feeding conditions, parasitic infections and physiological factors
(Le Cren, 1951; Hossain et al., 2006). Condition factor is also a useful index for
monitoring feeding intensity, age, and growth rates in fish (Oni et al., 1983; Kumolu-
.Johnson and Ndimele, 2010).
LWR have been extensively studied in a number of fish species such as
Cyprinus carpio communis and Ctenopharyngodon idella from Himachal Pradesh
(Dhanze and Dhanze, 1997); Horabagrus brachysoma (Gunther) from Kerala (Kumar
et al., 1999; Anvar et al., 2008); Channa punctata (Bloch, 1793) from Western Ghats
rivers of Tamil Nadu (Haniffa et al., 2006); Rasbora daniconius (Hamilton-
Buchanan) from Sharavathi reservoir, Kamataka (Kumar et al., 2006); Puntius
filamentosus from Chalakudy river, Kerala (Prasad and Ali, 2007); Chiiala chitala
(Hamilton 1822) from the river Ganga basin, India (Sarkar et al., 2009); Tor putitora
(Hamilton, 1822) from the Ladhiya river, Uttarakhand (Pafiyal et al., 2010); fourteen
freshwater fish species from the Betwa (Yamuna river tributary) and Gomti (Ganga
7
river tributary) rivers (Sani et al., 2010); nine freshwater teleosts belonging to families
Clariidae, Channidae, Cyprinidae, Siluridae, Sisoridae and Bagridae collected from
river Ganga (Khan et al., 2011c); fourteen species belonging lo families Cyprinidae,
Bagridae, Siluridae and Pangasidae from Cauvery river at Hogenakal in south India
(Muralidharan et al., 2011) and Channa marulius and Heteropneustes fossilis form the
river Ganga (Khan et al., 2012a). LWR and LLR have been studied in Labeo bata,
Channa punctata, Ompok pabda and Mastacembelus armatus from river Ganga
(Khan et al, 2012b). Saha et al. (2009) studied LWR and condition factor in Thenus
orientalis (Lund, 1793) along East Coast of India. Gupta et al. (2011) studied the
LWR, LLR and condition factor in Ompok pabda (Hamilton 1822) (Silurifomies:
Siluridae) from the river Gomti, a tributary of the river Ganga. However, studies on
length-weight relationships for fresh water fish resources of Kashmir are scarce. Some
of the contributions available are: LWR and condition factor oi' Schizopyge curvifrons
from river Jhelum (Mir et al., 2012), Schizopyge esocinus from Kashmir waters
(Bhagat and Sunder, 1984; Dar et al., 2012), Scizopyge nlger from Dal lake (Shafi and
Yousuf, 2012); LWR of 5. plagiostomus, S. esocinus and S. labiatus from river Lidder
(Bhat et al. , 2010) and LWR and LLR for two species oi Schizopyge (S. curvifrons
and S. niger) and three species of Schizothorax {S. esocinus, S. labiatus and S.
plagiostomus) from the Kashmir valley of India (Khan and Sabah, 2013; fomis part of
this dissertation).
A critical appraisal of the available literature, as discussed above, warranted
the need to generate the basic biological information on the selected fish species from
Kashmir valley. In an attempt to respond to such a need, the present study was
undertaken with the following objectives:
1. to evaluate and compare age estimates, and reader precision between
different structures (i.e. scales, opercular bones, otoliths, vertebrae and
cleithra) for age determination of Schizopyge curvifrons, Schizopyge niger
and Schizothorax esocinus.
2. to fit the length-at-age data to the von Bertalanffy growth model, and
3. to investigate the length-weight, length-length relationship and condition
factor for all the three fish species.
/ i
^ short description
ofjlshes
Schizopyge curvifrons (Heckel, 1838)
Kingdom Animalia
Phylum Chordata
Subphylum Vertebrata
Superclass Pisces
Class Osteichthyes
Subclass Actinopterygii
Subdivision Teleostei
Order Cypriniformes
Family Cyprinidae
Subfamily Schizothoracinae
Genus Schizopyge
Species curvifrons
Schizopyge curvifrons (Heckel, 1838)
Common names
Sattar snowtrout English
S attar Kashmiri
Salient features
Body is elongate, ftisiform with a short, blunt and slightly prognathous upper jaw. S.
curvifrons differs from all other Kashmir valley Schizothoracinae fishes in having
more gill rakers (21-28) and in the thin lips, without enlarged lateral or median flaps
or the fleshy appearance. Scales are very small; lateral line with 95 to 121 scales.
Scales on anal sheath slightly larger than body scales (Kullander et al., 1999).
Colour: light brownish, belly silvery.
Geographical distribution
Asia: Afghanistan, Pakistan, India (Menon, 1999) and China (Walker and Yang,
1999). Reported from Iran (Coad, 1995), Uzbekistan (Kamilov and Urchinov, 1995),
Kazakhstan and Kyrgyzstan (Berg, 1964) (FishBase, 2012).
Fishery information
S. curvifrons is a good prized indigenous herbivorous cold freshwater teleost of
Kashmir Valley (Mir et al., 2012). This large-sized species attains a maximum length
of 40 cm (TL) and maximum weight of 1.25 kg (Talwar and Jhingran, 1991). This
species inhabits river, lakes and swamps. Mature adults undertake spawning migration
to incoming streams and breeding takes place amidst gravel and sandy beds (Raina
and Petr, 1999). Spawning period is extremely protracted, from May until the
beginning of August. Large specimens spawn earlier than the small ones (FishBase,
2012).
Schizopyge niger (Meckel, 1838)
Kingdom Animalia
Phylum Chordata
Subphylum Vertebrata
Superclass Pisces
Class Osteichthyes
Subclass Actinopterygii
Subdivision Teleostei
Order Cypriniformes
Family Cyprinidae
Subfamily Schizothoracinae
Genus Schizopyge
Species niger
Schizopyge niger (Heckel, 1838)
Common names
Alghad snowtrout English
Ale gaad Kashmiri
Salient features
Body is elongate, fiasiform, with a short, blunt and slightly prognathous upper jaw.
Lips thick but not expanded into wide folds. Barbels two pairs (maxillary and
mandibular). A series of enlarged scales are present along the anal-fin base. Scales are
very small, 92 to 100 in the lateral line (Kullander et al., 1999).
Colour
The fish is much darker in colour than other species of the genus.
Geographical distribution
Asia: Kashmir Valley in India. Inhabits lakes and adjoining channels (Talwar and
Jhingran, 1991).
Fishery information
This species attains a maximum length of 27.0 cm (SL) (Kullander, 1999) and
maximum weight of 2.7 kg (Talwar and .Thingran, 1991). S. niger doesn't normally
occur in running water and is chiefly found in lakes (Kullander et al., 1999). It feeds
on detritus. Breeding grounds are found in shallow parts of lakes, particularly on the
roots of willow trees (Raina and Petr, 1999; FishBase, 2012). S. niger being a truly
lacustrine fish does not show any spawning migration (Shafi and Yousuf, 2012).
1 -^
ScHizothorax esocinus Meckel, 1838
Kingdom Animalia
Phylum Chordata
Subphylum Vertebrata
Superclass Pisces
Class Osteichthyes
Subclass Actinopterygii
Subdivision Teleostei
Order Cypriniformes
Family Cyprinidae
Subfamily Schizothoracinae
Genus Schizothorax
Species esocinus
Schizotliorax esocinus (Heckel, 1838)
Common names
Chirruh snowtrout English
Chirruh Kashmiri
Salient features
S. esocinus differs from all other Kashmir valley Schizothoracinae in lower gill raker
number (8-15) and in the much longer jaws, without enlarged lips or tuberculate pads.
Body is elongate, fusifomi with a long snout, only slightly prognathous upper jaw or
jaws equal. A series of enlarged scales along the anal-fm base. Scales are very small,
96-108 in the lateral line (Kullander et al., 1999)
Colour
Colour pattern is distinctive with silvery colour and numerous dark, small irregular
spots on back and flanks of body. Fins silvery-grey, with similar dark spots, more
numerous at their bases.
Geographical distribution
India: Indus river and its tributaries in Ladakh and Kashmir valley; and Afghanistan
(Talwar and Jhingran, 1991).
Fishery information
This species attains a maximum standard length of about 20 cm (Talwar and Jhingran,
1991). This species occurs in mountain streams, rivers and lakes (Menon, 1999).
Inhabits sandy and gravel-bottomed rivers (Shrestha, 1990). Herbivore, feeding on
bottom detritus. Mature adults undertake spawning migration to incoming streams
where they breed amidst gravel and sandy beds (Raina and Petr, 1999). Fry always
occur in quiet parts of the streams or in the side branches of the main streams
(FishBase, 2012).
14
MATERIALS AND METHODS
1. Fish sampling and data collection
A total of 173 specimens of S. curvifrons (Total length = 18.2-42.9 cm), 126
specimens of 5. niger (Total length = 18.7-37.3 cm) and 199 specimens of 5. esocinus
(Total length = 12.4-61.1 cm) were collected from the Jhelum river and Dal lake of
the Kashmir valley, India from June 2011 to June 2012. The following parameters
were recorded for each individual:
1.1. Total length (TL): Distance in a straight line between the anterior most part
of the body (snout or premaxilla, whichever is making the anterior most extremity of
the body) to the tip of the tail. In case the lobes of the caudal fin differ in length, the
length of larger lobe was taken.
1.2. Fork length (FL): Distance from the anterior most part of the body to the
anterior limit of the median notch or the bifurcation of the caudal fm.
1.3. Standard length (SL): Distance from the tip of the snout or premaxilla to the
base of the caudal fm (hypural joint), where a groove fonns usually when the tail
bends from side to side.
1.4. Weight (W): Total weight of each fish (including gut and gonads) was taken.
Identification of fishes was done following Day (1878) and Kullander et al. (1999).
Length and weight measurements were recorded to the nearest 0.1 cm and 0.1 gm,
respectively.
2. Age reading techniques
Scales, opercular bones, otoliths, vertebrae and cleithra were examined to compare
their age estimates as per standard protocols (Khan and Khan, 2009; Khan et al..
2011 a)
15
2.1. Collection and preparation of scales
A minimum of 10 scales were removed with forceps from under the anterior part of
the dorsal fin. Scales were cleaned by first removing the extraneous matter and
mucous by washing them in tap water and then rubbing in between the finger tips.
They were then mounted between two glass slides and studied with the help of
compound microscope (Tandon and Johal, 1996).
2.2. Collection and preparation of opercular bones
The opercular bones were removed and dipped in boiling water for few minutes to
remove extraneous tissue. A bristled brush was used to remove tissue that boiling
water did not loosen. Cleaned opercular bones were dried at room temperature and
examined under transmitted fluorescent light with naked eye (Phelps et al., 2007;
Khan and Khan, 2009).
2.3. Collection and preparation of otoliths
Otoliths (sagittae) were removed with a pair of fine forceps, rinsed with distilled
water and stored dry in labeled envelopes. Otoliths were read whole by immersion in
glycerol and examined under microscope using reflected light. Otoliths with unclear
annual rings were ground with sandpaper to make the annuli more distinct for age
reading (Tandon and Johal, 1996; Khan and Khan, 2009).
2.4. Collection and preparation of vertebrae
Vertebrae (4" to 10*) were removed and placed in boiling water for 10-15 min to
clear the attached muscles. A bristled brush was used to remove tissues that boiling
water did not loosen. Vertebrae were examined under dissecting microscope using
reflected light (Phelps et al., 2007; Khan and Khan, 2009)
16
2.5. Collection and preparation of cleithra
Cleithra were removed from fresh specimens and the muscles were separated by
dipping them in boiling water for 5 minutes. The cleaned and dried cleithra were
examined under transmitted fluorescent light with a dissecting microscope (Euchner,
1988).
2.6. Measures of precision
Each structure was examined for age estimation independently by two readers. Age
assessments of all the fish samples were done in random order and without prior
infonnation on fish length, weight or date of collection. Such data was utilized to
calculate the precision of age estimation between the two readers. However,
consensus data (from the two readers for the specific structures) was utilized to
compare the age estimates between structures and also to evaluate the statistical
significance among the mean age estimates from different stmctures. A consensus
between readers was required in cases where structures exhibited disagreement in age
assignments by the readers (Khan and Khan, 2009). In S. ciinifrons and S. niger each
alternative structure (scales, opercular bones, vertebrae and cleithra) was paired with
the otoliths (showing high PA and low APE and CV values) and in case of S. csociniis
each alternative structure was paired with vertebrae (showing high PA and low APE
and CV values) to further interpret precision.
Age readings were tested for bias and precision. Age bias plots were constructed to
identify trends and sources of bias in discrepancies between age estimates among
successive readings. Age estimates were compared by calculating the average percent
error (APE), coefficient of variation (CV) and percent agreement (PA) between the
readers and between the pairs of ageing structures (Campana et al., 1995). APE was
derived using the formula presented by Beamish and Foumier (1981):
R I I
APEj = 100% X - ) ' ' " i = l J
Where x,;y is the rth age determination of theyth fish, Xj is the mean estimate of theyth
fish, and R is the number of times each fish is aged. The resulting value represents the
APEj oftheyYhfish.
17
CV, expressed as the ratio of standard deviation over mean, was computed following
Chang (1982):
ivR ^^'i ^y \U=1 R _ l
CVj = 100%x
Where CVj is the age precision estimate for thejth fish. Low values for CV and APE
indicate high levels of precision.
PA between structures was also calculated to further interpret precision and was
estimated as the proportion of each age on which both readers agreed.
3. Growth analysis
3.1. Estimation of growth parameters
To evaluate variability in growth, observed length-at-age data based on otoliths in S.
cun'ifrons and S. niger and vertebrae in S. esocimis were fitted to the von Bertalanffy
gi-owth function (Ricker, 1975), by non-linear least squares regression. The VBGF is
represented as:
Lt = Loo(l-e-k[t~to])
Where L == total length (cm) offish at age t;
Loo = asymptotic mean length;
K = rate constant that deteiTnines the rate at which Lt approaches 1^;
t = time or age of the fish and
tn = the hypothetical age at which the fish had zero length.
VBGF equation was also computed using one of the most common and non-lethal
ageing structure, the scales, in order to compare the VBGF parameters to that
obtained using the precise ageing structure.
The concept of growth performance index introduced by Pauly and Munro (1984)
makes it possible to directly compare the growth performance of different populations
of the same species, based on von Bertalanffy growth parameters. The performance
Index (0) is calculated as:
18
0 = logioK + 21ogioLoo
The performance index was estimated using the growth parameters Loo and K
obtained in this study.
3.2. Length-weight relationship
Length-weight relationships were determined by logarithmic transformation of the
linear regression equation:
log W = log a + b log SL
Where:
W = weight of the fish (g);
SL = standard length (cm);
a = intercept and
b = slope of the regression curve or growth coefficient
The degree of association between the variables was computed by the detemiination
coefficient, r". Additionally, 95% confidence limits (CL) of a and b were estimated.
3.3. Length-length relationship
Length-length relationships viz. total length vs standard length, standard length vs
fork length and fork length vs total length were calculated by linear regressions
(Hossain, 2010).
3.4. Condition factor
Condition factor is a quantitative parameter of the state of well-being of the fish that
will detennine present and future population success by its influence on growth,
reproduction and survival. The Fulton's condition factor (K) was calculated according
to the equation:
K=(W/SL^)x 100
Where:
W = weight of fish in grams
SL = standard length of fish in centimeters
A plump fish will show a larger ratio than a thin or lean fish of same length.
19
4. Data analysis
For each fish species, the data obtained on age estimation from different hard
anatomical parts were subjected to Pearson's correlation analysis between the
structures in order to establish the relationship among readings of different structures
of the same fish species. Percent agreement was calculated using the "Templates for
calculating ageing precision" by Sutherland (2006). Mean age readings (consensus
data) obtained from various bony structures were subjected to one-way analysis of
variance (ANOVA) followed by Duncan's multiple range test (DMRT) (Gomez and
Gomez, 1984) in order to explain whether the readings from different bony structures
of the same species showed significant differences among themselves (Khan and
Khan, 2009). Growth Parameters were estimated using non-linear regression methods,
as implemented in a Microsoft Excel based application developed by Cope and Punt
(2007). Length-weight and length-length relationship of the fish was examined by
simple linear regression analysis.
All statistical analyses were done using MS-Excel and SPSS (version 12.0).
20
^
Results
1. Age estimation
The precision of age estimates from different readers as well as different structures
showed variations within and among species. In Schizopyge curvifrons and
Schizopyge niger, highest mean age values were obtained from otoliths and lowest
from cleithra while in Schizothorax esocinus, vertebrae exhibited highest values of
mean age estimates while cleithra exhibited the lowest values. The relative precision
of age detemiination between readers and between structures were estimated using the
percent agreement (PA), the average percent error (APE) and the average coefficient
of variation (CV) as presented in Table 1 and 2.
1.1. Schizopyge curvifrons
In S. curvifrons, the specimens ranged in age from 1 to 6 years. Percent agreement
(PA) between the two independent readers was highest for the otoliths (95.4%)
followed by opercular bones (93.6%), vertebrae (91.3%), scales (86.1%) and cleithra
(82.7%) (Table 1). Between structures, highest PA was found between otoliths aiul
opercular bones (88.4%), followed by vertebrae (82.1%), scales (74.0%) and cleithra
(68.8%) (Table 2). In S. curvifrons, otoliths and opercular bones showed lov\est
values of APE and CV as compared to other structures substantiating high precision
in detecting annuli. There was no ageing bias between readers for otoliths while little
ageing bias was found between age estimates from scales and cleithra as shown by the
age bias graph (Fig. 1). Age bias graphs between structures revealed no age bias when
otoliths were compared to opercular bones and vertebrae whereas underestimation of
age was found with scales and cleithra for older fish species (Fig. 4).
The correlation coefficient between readers was higher for otoliths (0.977) and
opercular bones (0.970) as compared to other structures (Table 3). Similarly,
correlation coefficient between age estimates from different bony parts of S.
curvifrons showed that each ageing structure was correlated significantly (P<0.01)
with the other. Correlation coefficient between otoliths and opercular bones indicated
a near perfect concordance of age estimates from the two structures (0.937). Mean
values of age estimates from different structures, when compared using ANOVA
21
followed by DMRT, showed highest (P < 0.01) values for age readings from otoliths
followed by opercular bones, vertebrae, scales and cleithra (Table 4). Age readings
from otoliths, opercular bones and vertebrae did not show any significant (P < 0.01)
differences.
1.2. Schizopyge niger
In S. niger, PA of ages between readers was higher for otoliths (94.6%) than scales
(92.3%), vertebrae (84.6%), opercular bones (82.3%) and cleithra (80.0%) (Table 1).
When otoliths age estimates were compared with other structures, highest PA was
found between otoliths and scales (86.9%) followed by vertebrae (83.8%), opercular
bones (78.5%) and cleithra (73.8%) (Table 2). Otoliths and scales showed lowest
values of APE and CV as compared to other structures. No age bias was present
between readers for otoliths, scales and vertebrae while little ageing bias was found in
opercular bones and cleithra (Fig. 2). Between structures, little age bias was present
when otoliths were compared to scales, vertebrae and opercular bones whereas
cleithra showed underestimation of age (Fig. 5).
Correlation coefficient between readers was higher for otoliths (0.978) and
scales (0.970) as compared to other structures (Table 3). Correlation coefficient
between age estimates from different bony parts of S. niger showed that each part was
significantly (P < 0.01) correlated with the other. Otoliths exhibited highest values of
correlation with the scales (0.946) followed by vertebrae (0.924), opercular bones
(0.913) and cleithra (0.896). Mean values of age estimates from different structures
shovv'ed that age estimates obtained from otoliths were significantly (P < 0.05) higher
than that from cleithra but comparable (P < 0.05) to the values from scales, vertebrae
and opercular bones (Table 4).
1.3. Schizothorax esocinus
Between readers agreement for vertebrae was higher (96.0%) than opercular bones
(95.0%), scales (93.5%), otoliths (90.5%) and cleithra (87.4%) (Table 1). When age
estimates from vertebrae were compared with other structures, highest PA was found
between vertebrae and opercular bones (87.9%) followed by scales (86.4%)), otoliths
22
(80.9%) and cleithra (73.9%) (Table 2). Vertebrae, opercular bones and scales showed
low values of APE and CV. No indication of reader bias was noted for any structure
(Fig. 3). Between structures, little bias in age estimation was noted between vertebrae
and cleithra (Fig. 6) with cleithra showing underestimation of age in older fish.
The correlation coefficient between readers was also higher for vertebrae
(0.994) as compared to other structures (Table 3). In S. esocinus, correlation
coefficient between age estimates from different bony structures showed that each
part was correlated significantly (P < 0.01) with the other; highest values were found
between vertebrae and scales (0.974) followed by opercular bones (0.972), otoliths
(0.960) and cleithra (0.945). Mean values of age estimates from different structures
showed that values obtained from cleithra were significantly (P < 0.05) lower than the
readings obtained from all other structures except otoliths (Table 4). Also, the age
readings from vertebrae, opercular bones, scales and otoliths were comparable
(P<0.05) with each other.
2. Growth
2.1. Growth parameters
The von Bertalanffy growth parameters as well as growth perfonnance index (0) m
the selected fish species are presented in Table 5.
S. curx'ifrons
Following growth equation emerged using otoliths as the ageing structure:
Lt = 49.8 (1 - e-o-263(t+o.34>)
iS*. niger
Von Bertalanffy growth curve was fitted to the total length at age data using otoliths.
Following growth equafion was obtained:
Lt = 44.8 (1 - e-0-255(t+i.42)^
S. esocinus
The VBGF parameters obtained using vertebrae have been presented in the following
equation:
23
Lt = 66.6 (1 - e-o-27^ (t+o.34))
The von Bertalanffy growth curve is shown in Figure 7. Greater predicted
lengths for older specimens were estimated from scale-aged than otolith-aged fish.
The von Bertalanffy growth parameters constructed with scale data had a higher
asymptotic length and a lower growth coefficient than those found in the model based
on otoliths in S. curvifrons and S. niger and vertebrae in 5*. esocinus. Growth
performance index was found to be highest for S. esocinus (3.09) followed by S.
curvifrons (2.81) and S. niger (2.71), respectively.
2.2. Length-weight relationship
Length-weight data are presented in Table 6. The length-weight relationships were
calculated and represented in the form of following equations:
S. curvifrons Log W = -1.401 -|- 2.69 Log SL (n = 136: r = 0.992)
S. niger Log W = -1.304 + 2.66 Log SL (n = 173: r '= 0.982)
S. esocinus Log W = -1.925 + 3.08 Log SL (n = 163: r^- 0.995)
2.3. Length-length relationship
Estimated parameters of the LLRs are given in Table 7. The LLRs were calculated
and represented in the form of following equations:
.S". curvifrons TL = 2.39 -f- 1.09SL r = 0.998
FL = 1.68 + 1.04SL r = 0.999
TL = 0.63 + 1.05FL r^= 0.999
S. niger TL = 2.23 + l . l lSL ? = 0.993
FL= 1.38 + 1.06SL r^= 0.997
TL = 0.75-I-1.05FL r^= 0.997
S. esocinus TL = 0.93 + 1.15SL r^= 0.998
FL = 0.45-t-1.08SL r^= 0.997
24
TL = 0.48 + 1.06FL r^= 0.998
2.4. Condition factor
Condition factor (K) was found to be highest for S. niger (mean 0.49) and lowest for
S. curvifrons (mean 0.49).
S. curvifrons K = 0.49 (0.18-1.20)
S. niger K = 1.74 (1.24-2.01)
S.esociniis K = 1.51 (1.24-1.85)
25
o ^
- * - » <+H
o in OJO
c T 3 « D t-H
<U t * 03
0) ^ - * - < c OJ (U ^
"S - O —
> O
r ^
O
<A
a > i^^
^ t+H c o u o to
'o H f 8 U-I V <D o O -s; o •c
T d
OS -5 U
Co
< 03
i^ <ii o •5i^
S c 5« u C> o S V-. o (D . N
a ~5 (U o 5 0 Co 03 ^ t ^ "
U s; > c3
o
^ /^^ S
V < S
V CLi, 1^ u
, ^ j w c b€ <u ;^ B
G C3 Co
c (U • ' " '
o uo i-H ^ (U O
T3 03
o V^
o ZTi ' O
•§ CIH CL,
s o
!U s o
T3
o C
y-i
0)
^ CJ
H
O 0 0 o 0 0 l / ^ o tr-m ^ ro -+ c^ '-^ o 2: ?.
O N •st-
Csl rq r-o -H
<N 0 0 ON
( N OO (>1 ON V3 O N '—^ O iy~) m
O —' —' (N r^
ON •00 00
00 o
ON
O
m O N
O N O O oo
NO r ^
ON ON
NO
OO oo o 00
NO IT) O N
O
S
CO
C O
o ^ O
03
t; >
at CO ;-( W x: rt - * -» f ) <L)
GO r "1
NO
.bo
CO
o 00
ID
t: >
c/3 U
o 03 i-
o
O N ON
§ •5 o
o ~S o
-S cH
C3
t; >
en (U d o
03
O )-c
D
O
r^ IT) ^ o
( N ( N CM i /^
NO
1 0 oo NO
o . oo
00
— ^- — r^, m
t o i n r<-) O ON O^ oo
C/2 CO _ C
1 1 c/D rN
03
o
3 c
2
!U <L>
^ - * - » a> X^ 1 3 0)
a M m M cd
czi <D bO a bO P! O B c3
— > u • — ^
c o -^ KJ
'^ a >
-*—( o
• 4 — *
S (D
O • t - i 4-1
o o s
'o a o g t o
— X j-1 13
•\ ^ < o
o o , N
r^ '^ OJ r; o c«
s 03 w ; - i bo > 1 ^ N
< 2 H Co —
- 4 — ' t o C s: !U o a (D £ Q) s CO o
- 4 — »
bx) o gs o Cl,
^ 5 1) N
a •3 t+- l u o Co o C! o • -c/D V2
133 - 4 — '
a o
2 u
4->
f S
0) B-H
A « H
to
i
en
o
3 o
o I
m 0 0 (N
r~~
O ro CO
^^22 0 ^ en '^ i n r
00
!/-> t ^ o Cf-.
^ ( N Tl- ^ >o ^ O ^
^ ro ^ r-~ Os
IT ) o r 1 00
m LT)
^ r-^ o oo o 0 0 "O 0 0 o ^ ON o-oo oo CO
CO 0 0
m oo
oo m t^ 0 0 CO o
CO
> o on I
en
I (N
tu
en
i ^ 1 5 2 O = O
o O
.bo
en
G (U o
en ca X) (U X ) u rrl <1> ca O • d n
<u () en > \-t
o -^
<u O H o
o o ^ 5 O
ON 0\
•S o
o -«: o
.N
X en en CO
CO x; :3 O
en U
n 7) u CO
1 1
U H VH CO CO
n _o VH w 1 11 X X
CO
-s > > > t (U
>
en
S CO en
t+H O ; - ( tu
X
c
"^ o
cd
^ ^ ^ 0 0 s—* 'w ^ (N CN r "S -4—*
ON ON ON 1—H
O o O N
I j 1
o o O O 1 GO 0 0
o 0 0
d
I j 1
O N
d
O
C3
CD
>
S-H a\ '^ ' t ) - 4 -*—* ^ iy~> '^ ON X> '^ w O N O N ON (U O O O O
> r
l o ^ o O
> > r
o ON
CM O ;
1 N O
O N
t / 1
. V5
'o O
d d t: >
d
:5 r--0 0
OO (A CO
"o o> ON ON c <U
(U O o o O :3 o <U
(U
O 4 - '
o B
03
o
;-• 03
C C/2 3 o
r-- rr^ 3 o
r\l CD r-
3 o r<-) - 3
o t^
U S lU ON ON u . C^ - 4 — ' o
1
d d o
d
CQ o CQ 1
C3 o
o OO OO 'r^ -cd r- CM 0 0 O X) cd
t : > ON O N 6 <u 3 o
o O O 6 & o
t /3
13 o p
OO NO
NO
>
CD
13 o (/) I
^ r-
<u r-~ o r- C/3 0 0 O N & ON (N r- OO
C/3
t H
13 ON ON ON - 4 — » d d d c/ O o "o
o
'>3
S
>
O s •S o V _s; p.
y •^ D
on S to C/5 5
1 ) g '3 U tu s H
Q Q H ^ ^ CL, ^ &-w 1 00 1
• ^
O Co •<
^ ^
C/3
3 •5 o ^ , V X J
(N ON X I
(N ra ^
K ON ON 0 \ CN ON (N P T (^ t ^ "sl- r-
0 0 OO >r) 0 0 ^ <N (N <N ( N (N
o . N
C/j
a; .^0
en 0 0 0 0 X
' ih O
0 0 ( N OS •^
W) rn rn ON
(N r\i (N <>i '-' O •«
c3 _2 13 > c C3 <U
S
to
p
^ ' ^ X . p . X — ^ r* oo ^ m NO -—. lu r-- r<-i ri (N r r-uo r- o NO NO ON 1—1 (N r—( 00
1^ r-i m
<u
r<~, cn (N
CO c
c o
CQ
00
13 a
o
o
a, O
O
as ; - ( -8 v. >
S3
-4—*
u
V
e C
13
o
IT) o d A OH
>> C3 O
tX)
f—
g o o
-G O C3
o )-.
3 on u, 03
c • >
CO
13 >
u .bo X ^ IT) VO
O
S
o
Co
X u
o !=:
o
bO
p S feh K
U > O
fc t i l
K Ct
« ^ tl o (i> •^ CQ Cl N
c • ^
u u > CO
S)
o
oo
O
(N
in CM
o
O N O
00
CNl
o
IT)
o CO
^ (U •T3
, , x) CT t-< + ^ o o f )
a j X V-.
C3
J o
1^ C P3 . i i
t4-.
O
o I - 4 — '
00 p
o O
1/3
13 o 00
Cvj
o O
l O
o
(D CO
C3 a> CS O t: o
00 >
00
C/3
O
>
to
i . •5 w .b<)
s ^ li) ly t>0 H K a § o
.N •< -5 o o . N Co -5 O
Co
yCi
H
t l
H-1
u ID O N
wo
to-c
OS
-a
GO
o
c
'o C/5
0^
oo o^
I
(N 00 i n
o o
I 00 O
ON ro O
t ^
m
«n
NO
3
OJ
O H
to
o
C/3
(N
ON
O
oo (N m in '^ IN
NO
ON O
<N O
* O N
O
NO
.bo
o o O N
o
O N
in m
(N
<N
-a o C3 (N
x: P 00 o
^
00
in ON ON
r-
oo ON
(N
00 O
o o
I oo o o
o o
00
o
o
NO
^ 1 3
OH
H
NO in ON
o
(N
o
o in
NO i n mi
oo
o o
VO
in
i n
o
oo
NO
q oo
CM P
o T f ,—1
"O oo o C o , CNI c ^ - ^ r .
. 4 1 F—H 03 '-^ c3 bO <U •a T3 t; -C C
OQ =5 "S 00 ^
DQ
i n 00 oo
NO oo
i n
1 ^
a; i n cs
2 S
o
Q
to 3
O to
H
o
o
o
8
o
_i U u a o
Cl.
o
C
X)
g
13 O
NO
H
' ^ to
O T3
"So ^ (U ^
3 H
0 0 O N ON m r t^ 00 I ^ 0 0 ON ON O N O N O N O N ON ON ON
^ ^ i - ON O N ON ON O N ON ON ON O N
O O O d d d d d d
ON ^ i r i ^ NO i n i r> oo NO
x> p p q ^. ^ p p p
O N 0 0 r<^ m 00 IT) r<-) i n OO rt d ^ NO (N r^ r~~ ON • ^
oj ^ d (N ^ d d d d
J J J KJ - 1 h J _i J K J
G C/) GO U H 0 0 GO tu GO 00 u-_o r i3 ^ ^ Xi - O ^ ^ X) - O
"S + + + + + + + + + 3 a 03 rt cU « K) C3 C3 ct)
cr II II II II 1! 11 1 II tl tiJ J J J KJ J J _) -J -J
1— p
NO
H £_. c_ H
2 n-1
s
to
to
NO
CD
'o <u ?s a 0)
-5
o
o
Co
o
'3
o o
GO
£0
,o
J tin
"So
o
H
(U
"EH
a
o 1-1
s =3
o
u
>
ON
0 0 L. u •a
r-- C9 Ol k.
^ > 1
. f i
i r i T3 01
'd-
K C l it
Si
(N
l J3PB3J Xq pajBiuisa 3§v
O C N 0 C t ^ ^ > O ^ r ^ ( N — O
2 j3pB3j y{q pajBiuisa agy
o
V ^
X! in -8
- f a -
CD ^ <
J J3PB3J Xq pajBUijsa aSy
o
c
1 \ ° \
- ON
- oo "" u
- ^ ^
- ^ 2 e Ol
- <N »f
- O
3 c 5 0 N O o r - ^ i r ) ^ r n < N ^ ^ C
I japcaj Aq pajeuiisa aSy
- ON
- oo "" u
- ^ ^
- ^ 2 e Ol
- <N »f
- O
3 o O N o o r ~ - ^ i / ^ ^ m ( N - - ' 0
3 J3PB3J Kq pajBUitsa aSy
01
3 6D
O
\
- ON
u
" ^ S « « »- \ a> - \ u 4> N» - <N 6f
^ V <;
O C ^ O C t ^ ^ ^ O x J - r ^ f N ^ ^ C :> T —
Z Japeaj Aq pajBiuijsa aSy
3 aapeaj /(q pajEuiijsa aSy
' • ' \ •
t
JS
"3 U
\ \
\
oo u
• a OS
E
^j3pB3j Xq pajBiuijsa 3§v
o
O \
• 0 \
- oo i . CI
at u
- ^ >v X i
- ^ s - <-> 1
CI
• <
c 1 t i 1 1 - 1 1 1 1
Z J3pB3j iCq pa^eiupsa aSy 3
o \
o
\
- ON
- CO i,
•T3
01 I .
05 \ X 4J \
!3 H H - •r "S
O \ -*—' JQ H N ^
« \ • • 4
-- :« - r^ (y s \ " \», fcl)
- "^i • <
o X c O N O o O v c i O ' ^ r n t N - H C IS
Z aapBaj jCq pa^Buipsa aBy
o
- OO ^
«= \ h VC ; ^
i \ JQ - >0 - o
- ^ 5 ' \ s « \ " S •x - (~n oS
3 \ 01
Hi \
01
a. \ , 0 •x^ • o • o
o O N O O t ^ ^ i r ) T t r ^ ( N ^ ^ O
Z J3PB3J Xq pajBUiisa 3§v
<u
u
- 1 ^ u it
TS - VO 53
U
- </-) £1
T3 ^ » - h ^
« - r i s E «
0> r-l OJ
£ii
^_ <
3 j s p e a j Xq pajBiuisa aSy
3 en
\
\ 01
\ \
\ 4J \
2 \
• o o>
+ -CS
S 01 V
1 \ c 5 0 N 0 0 t ^ ' O i r ) ^ r 0 ( N ' - H C
^ T; J3PB3J Xq p3)Buiis3 s S y
• - ,
-
"o
0
i
\
1
0^
vO Xi T3
v Ml
O 0 N 0 0 r - ^ L r i ^ r - i ( N ^ - O
Z J3PB3J Xq p3:>BUIlS9 3§Y
m i
O O ^ o o ^ ^ \ C ^ o ^ m ^ N l
BJIIJI3I3
W
V
y: JZ
o O
0 ( j \ O C t ^ ^ i O ' ^ r O ( N
sauoq JBinojado
!••< "o
o
n \ I ; ^ \ 1 1 r-
o c 7 N o o r ^ ^ D i o r ) - m ( N ' - ^ o
S31B3S
.bo
t-*M
ON
in
O
_ O
f o
- ^
^
.A
o O
o c f \ o o r - ^ i o ^ m r N ^ - ' 0
B j q j i a o
00
C (U
"S
X)
O
•n a, S o u (U W) CO
&
1/1
2 IS QJ
<:
3 6E
O
o
(N
O O N O O C ^ V C > i n ^ r < ~ ) ( N ^ ^ O
S3[B3S o O N o o r ^ v o t r i ' ^ r n t N ' — o
sanoq JBjnojado
o
- o
\ - 00
- t>
U
u
t*; r^;
:•« - (N
n
O C ^ C 3 0 r - - v O > / 1 ^ r - , f N ^ ^ O
BJIHJPO
L O
in 3
Figure 7. von Bertalanffy growth curves for Schizopyge curvifrons, Schizopyge niger and Schizothorax esocinus showing back-calculated length at estimated age.
45
40
35 ^—V
s n u • ^ ^
J3 ?S W)
f> 20
« 15 o H 10
5
0
5. curvifrons
otoliths
- • - - scales
III I V
Age (years)
VI
40
35
? 3 0 u
---25
I 20 at
^ 10
5
0
5. niger
- •— otoliths
- • - - scales
III IV
Age (years)
VI
70
60 • S. esocinus
?50 i ^
£ 40 •
| 3 0 J
1 20 r
10 '\
0
1 II III IV V
Ages (years)
_ „, , . . ^ , . . .
- • — vertebrae
- • - - scales
VI
DISCUSSION
When assessing methods for age determination in fishes, it is important to consider
both accuracy and precision. Imprecise age estimates suggest variabiHty in ageing
criteria or the presence of unclear annuH that are difficuh to distinguish and count,
while inaccurate age estimates bias population parameters such as growth and
mortality (Quinn and Deriso, 1999). Despite the call for more statistically robust and
consistent analysis of ageing data in recent years (Beamish and Foumier, 1981;
Chang, 1982; Campana et al., 1994; Jackson et a!., 2007), many studies report
population data (i.e. length-at-age) without mention of the precision of their age
estimates. In the first study on precision of age estimates in S. cnrvifrons, S. niger and
S. esocinus inhabiting Kashmir valley, it v/as observed that the structures giving
precise age estimates varied among the selected fish species.
Otoliths
Agreement between age estimates from otoliths and other hard structures varied
among species as well as between readers (Table 1 and Table 2). In general, we found
that otoliths yielded the most precise age estimates for S. cunnfrons and S. niger with
age agreement between readers to be 95.4% and 94.6%. Studies have consistently
shown higher precision in ages assigned with otoliths than with scales. This is in
accordance to a mounting body of evidence that the scale method of age estimation
for fishes belonging to subfamily Schizothoracinae may be unreliable (Zhao et al..
1975; Li et al., 2009; Chen et al., 2009; Ma et al, 2011). Otoliths were reported to
provide the most reliable age estimation, while the annuli on vertebrae and opercular
bones were not very clear in Gymnocypris selineuoensis (Chen et al., 2002a, b; Ma et
al., 2011). Otoliths continue to grow and form annuli even as body growth slows and
asymptotic length is reached, and annuli reabsorption does not appear to occur during
periods of food limitation or stress (DeVries and Frie, 1996; Maceina and Sammons,
2006). Otoliths were reported to be the best structure for estimating yellow perch age
based on high reader agreement and low coefficient of variafion (CV) (Niewinski and
Ferreri, 1999). Gumus et al. (2007) determined age and growth of Scardinius
erythrophthalmus (Linnaeus, 1758) using five hard structures (scales, vertebrae,
opercular bones, lagenar and utricular otoliths) and reported otoliths as most reliable
26 . . . . . . ,
ageing structure. Isermann et al. (2003) demonstrated that whole-view otoliths were a
more time efficient method for ageing walleyes than scales or dorsal spines.
Boxrucker (1986) and Kruse et al. (1993) also suggested that ageing whole-view
otoliths was less time-consuming. Ages estimated from scales and spines were less
precise than from otoliths and scale ages showed underestimation in older specimens
of largemouth bass, smallmouth bass, yellow perch and brown bullheads collected
from upper Hudson river located in north-eastern USA (Maceina and Sammons,
2006).
Vertebrae
Amongst all age structures, vertebrae were found to be the most suitable ageing
structure in S. esocinus, with agreement of age estimates between readers to be
96.0%. The findings were similar to those of Ma et al. (2011) who reported that age
estimates of S. o 'connori from vertebrae and otoliths matched closely while opercular
bones appeared to underestimate age but for older fish, the counts diverged and
otoliths consistently provided higher age estimates. Filmalter et al. (2009) compared
otoliths and vertebrae as potential hard structures for ageing South African yellow fin
tuna, Thunnus albacares and found that growth increment counts from whole otoliths,
sectioned otoliths and vertebrae were not significantly different (t-test, p > 0.05).
Khan et al. (2011b) compared age estimates from otoliths, vertebrae, and pectoral
spines in African sharptooth catfish, Clarias gariepinus (Burchell) and found that
mean age estimates from otoliths were comparable (P > 0.05) to the values obtained
from vertebrae. Guinn and Hallberg (1990) reported that vertebrae and otoliths gave
similar age esfimates in Burbot, Lota lota (Linnaeus). Vertebrae were the most
reliable bony structure for ageing shad {Alosa pontica Eichwald, 1838) inhabiting the
Black Sea, as it had the highest agreement and the lowest ageing error (Yilmaz and
Polat, 2002). The findings of Liu et al. (2009) indicate that the vertebrae are suitable
calcified structure for age determination of the sharptail mola.
Scales
In S. niger, when otoliths age estimates were compared with other ageing structures
(i.e., scales, opercular bones, vertebrae and cleithra), highest PA (86.9%) and lowest
27
APE (3.61%) and CV (5.10%) values were reported between otoliths and scales. In
most of the cyprinids, age has been estimated from scales (Kamilov, 1984; Phelps et
al., 2007). In addition to having clear and sharp annuli, scales also have the
advantages such as easy collection, preparation and being non-destructive to the fish
(DeVries and Frie, 1996). Because scales are commonly used to age fish, numerous
studies have assessed the accuracy and precision of scale age (e.g., Erickson, 1983;
Boxrucker, 1986; Kruse et al., 1993; Long and Fisher, 2001). Amongst all age
structures, scales were found to be the most suitable ageing structure in L. rohita and
C. marulius (Khan and Khan, 2009). Precision of ages detennined from scales and
otoliths were similar in black crappies collected from South Dakota waters (Kruse et
al., 1993). Several researchers have reported that scales can provide unreliable
estimates of fish age (Boxrucker, 1986; Hammers and Miranda, 1991; Khan and
Khan, 2009). Scale ages were on an average 9 years less than the ages estimated from
sectioned otoliths in Morone saxatilis (Linnaeus) older than 20 years, but scales were
reported to esfimate age adequately up to the age of 12 years (Secor et al., 1995). In
some scienfific reports, the use of scales had been criticized mainly because of the
frequent underesfimation of the ages in older fish (Beamish and McFarlane, 1987).
Imprecise age enumeration from scales has been attributed to reabsorption and
deposition of false annuli due to stress and food limitation, and annuli becoming
obscure because scale growth tends to cease as fish grow older (Beamish and
McFarlane, 1987; DeVries and Frie, 1996; Maceina and Sammons, 2006).
Opercular bones
Esfimates from otoliths in S. curvifrons were compared with other ageing structures
(i.e., scales, opercular bones, vertebrae and cleithra) and were found to match closely
with opercular bones with highest PA (88.4%) and lowest APE (1.75%) and CV
(2.47%) values. Age bias graphs between structures (Figure 4) indicated that age
estimates obtained from otoliths and opercular bones were in a good agreement. Khan
and Khan (2009) suggested that annuli in opercular bones provided the precise age
estimation in Catla catla. Opercular bones were reported to be superior to scales for
the age estimafion of common carp (McConnell, 1952). The determination of age and
growth of fish from opercular bones is well established in a number of fishes and have
more reliable age estimates than scales, vertebrae, spines or other hard parts in Esox
28
lucius (Linnaeus) (Frost and Kipling, 1959). In S. niger and S. esocinus annuli on
opercular bones showed mean age estimates comparable to those from all other
structures except cleithra while in S. curvifrons mean age estimates from opercular
bones were comparable to all other structures except scales and cleithra.
Cleithra
Cleithra are commonly used to age fishes (e.g., Casselman and Crossman, 1986;
Sharp and Bernard, 1988; DeVries and Frie, 1996; Quist et. al., 2007); however, few
studies have been conducted to evaluate the precision among age estimates using
cleithra. Govind and Gopal (1966) found valid growth rings in the cleithral bone of
Silonia childrenii. Laine et al. (1991) noted that scales are not as well-suited for use in
age determination of pike as compared to cleithra. Brennan and Cailliet (1989)
evaluated a variety of calcified age structures (pectoral fin rays, opercles, clavicles,
cleithra, medial nuchals, and dorsal scutes) used to age white sturgeon Acipenser
transmontanus and found that ages estimated from these structures did not vary
significantly. However, in the present study, mean values of age estimates from
cleithra was found to be significantly different to those from all other structures
except scales in S. curvifrons, opercular bones in S. niger and otoliths in S esocinus.
Nuevo et al. (2004) found otoliths and cleithra to be unsuitable structures for age
deteraiination of bighead carp from the Mississippi River. Quist et al. (2007)
evaluated between reader precision and agreement of otoliths age estimates from
scales, fin rays, cleithra, and opercular bones and found that exact agreement between
readers was highest for otoliths and fin rays and lowest for opercular bones, cleithra,
and scales.
Growth parameters
Estimates of the size of fish in relation to age (time) are used for estimating certain
growth parameters. Each of these parameters describes the inherent characteristics of
growth of the species. The von Bertalanffy growth function is commonly used in
describing fish growth (Gallucci and Quinn, 1979; Misra, 1980; Chen et al, 1992).
This function has only three parameters, making it statistically robust compared to
models with more parameters (Booth, 1997). The VBGF parameters are commonly
29
used in mortality estimates and fisheries stock assessment modelling (Dominguez et
al., 2006). The von Bertalanffy equation follows the assumption that fish grows
towards some theoretical maximum length, and the growth rate declines as the fish
reaches its ultimate length (Beverton and Holt, 1957).
The growth coefficient (k) is a useful index for estimating the potential
vulnerability of stocks to excessive exploitation and for comparing life history
strategies (Pratt and Casey, 1990; Musick, 1999). Branstetter (1987) categorized the k
values as 0.05-0.10/year for slow growth species, 0.10-0.20/year for species with
moderate growth, and 0.20-0.50/year for rapid growth. In the present study, the value
of K was in the range of 0.20-0.30/year. A high value of k indicates a high metabolic
rate and such fishes mature at an early age or at a size which is large in relation to
their asymptotic length, LQO (Qasim, 1973).
The Loo of S. niger (44.8 cm) was larger than the maximum obser\'ed size of
37.3 cm which was likely due to the less number of large specimens. Growth model
estimates are greatly affected by the lack of very young or old individuals (Cailliet
and Goldman, 2004; Ma et al., 2010). In S. ciirvifrons (N=-173) and S. esociniis
(N=199) due to relatively high sample size and rather good agreement of Lx with the
observed maximum length, values appear to be more reliable and clearly underline the
trend of higher growth coefficients in smaller sized species. This in coiToboration
with the findings of Frisk et al. (2001).
In general, growth parameters need to be checked for quality and validity
(Karlou-Riga and Sinis, 1997), while a negative value close to zero for to is a good
indicator of the reliability of the determined ages (Kerstan, 1985; Gang et al., 2008).
The estimate of to in the present study ranged from -0.30 to -1.45, minimum for 5.
esocimis (-1.42) and maximum for 5'. curvifrons (-0.34).
Growth performance values were all in the same range, as should be expected
for related species, but still there was variation between species. In S. esocinus, which
also had the highest k value, growth performance was found to be highest (3.09)
fohowed by S. curvifrons (2.81) and S. niger (2.71).
The growth parameters may vary regionally or methodologically; it also
depends on differences in size of the largest individual sampled, species, sex and age.
VBGF parameters were generated using otoliths and scales in both S. curxnfrons (PA
30
74.0%) and S. niger (PA 86.9%) in order to compare the results from the two studies.
Scales were selected because they are often the most common structure for age
estimation on account of their advantages discussed elsewhere. The length-at-age data
derived from otoliths and scales were significantly different in S. curvifrons (t-test for
paired comparison; t = 3.159, df = 5, p < 0.05) but no significant difference was found
in S. niger (t = 2.588, df = 4, p > 0.05). In S. esocinus, VBGF parameters were
generated using vertebrae and scales (PA 86.4%) in order to compare the results from
the two studies. The mean total length derived from vertebrae and scales were
compared but no significant difference was found (t-test for paired comparison; t =
2.562, df = 5, p > 0.05). Estimates of asyinptotic length derived from scales were
higher for both species together with a moderate growth coefficient that shows that
their size increased at similar ages. However, the method of age detennination from
scales could be still applicable to analysis of growth for these species, because of
similar growth performance indices. These results suggest that management actions
based on estimates of growth would not be significantly influenced by the type of
structure used.
Length-weight relationship (LWR) and length-length relationship (LLR)
No infonnation on LWRs and LLRs of the selected fish species was available in
FishBase (Froese and Pauly, 2011). When the b value in LWR was equal to or did not
show statistically significant deviation from 3, the growth was isometric, whereas the
positive or negative allometric growth occurred when the b value deviated
significantly from 3: positive if b > 3 and negative if b < 3 (Ricker, 1975). The LWRs
suggested isometric growth in S. esocinus (3.08) which was in agreement to the
reports of Bhagat and Sunder, 1984 (3.0034), Bhat et al., 2010 (3.0034) and Dar et al.,
2012 (2.7148 for males and 2.8618 for females) respectively. The negative allometric
growth for the remaining species implies that the allocation of energy is more towards
axial growth rather than to biomass. In S. curvifrons, the allometric coefficient b of
the LWR was reported to be negatively allometric (b < 3) throughout the year except
March, July and October where the growth was isometric (b=3) (Mir et al., 2012).
Shafi and Yousuf (2012) reported the value of b for S. niger as 3.07 for males, while
in females it was 2.77 and in case of pooled data its value was 3.07. The value of b
reported by Shafi and Yousuf (2012) for S. niger is different from the present study
31
which can possibly be due to several factors such as the habitat, number of specimens
examined and length ranges and length types used. As per the data available in
FishBase (2011), we reported the new maximum standard length for S. esocinus (50.8
cm) and S. niger (32.6 cm). As suggested by Petrakis and Stergiou (1995), use of
LWRs should be strictly limited to the observed length ranges applied in the
estimation of the linear regression parameters. All LLRs were highly significant (P <
0.001), with determination coefficients (r^) > 0.98.
Condition factor (K)
The most useful tools used for evaluation of fish populations, are the mathematical
equations or parameters intended to estimate fish condition, using the relation
between total length and weight (Le Cren, 1951, Bolger and Connolly, 1989,
Ritterbusch-Nauwerck, 1995). The condition factor (K) is an index reflecting
interaction between biotic and abiotic factors in the phvsiological condition of the
fishes. It shows the well-being of the population during various life cycle stages
(Angelescu et al., 1958; Gupta et al., 2011). The condition of 5'. curvifrons and 5.
esocinus from Jhelum river and 5. niger from Dal lake of Kashmir valley was
evaluated using the Fulton"s condition factor. The values obtained tilted the range of
values most commonly reported for these species. In S. curvifrons, the condition
factor was calculated month-wise and it ranged from 1.0-1.95 (Mir et al., 2012). Shafi
and Yousuf (2012) reported Fulton's condition factor for S. niger in the range 0.996-
12.4 (minimum of 0.996 in November and maximum of 1.24 in March). Sex-wise
analysis of 'K„' values in S. esocinus revealed that the mean 'K,,' value was 0.96 in
females and 0.91 in males (Dar et al., 2012). Slight differences in the mean values
between the results obtained and the published data could be the result of different
seasons at capture and the fact that the influence of fish length on the condifion factor
was not taken into consideration in the analyses. According to Le Cren (1951), 'K,,'
greater than 1 indicated good general condition of fish. Pandey & Sharma (1997)
studied the condition of four exofic carps and only the common carp, Cyprinus carpio
communis was found to have value above 1 ( 1.0109). Pandey & Sharma, 1998
reported high 'K ' values for Labeo rohita (1.0129) and C. catla (1.0007) and low
values for Cirrhinus mrigala (0.9967).
32
It may be concluded from the present study that otoUths were the most precise
ageing structure in S. curvifrons and S. niger while vertebrae showed most clear and
sharp rings in S. esocinus. The estimated ages were 1-6 years for S. curvifrons and S.
esocinus and 1-5 years for S. niger. The von Bertalanffy growth model was obtained
as Lt = 49.8 ( l - e-0'263(t+o.34^ -j ^ curvifrons, Lt = 44.8 (1 - e-o-255(t+i.42)^ i^ ^
niger and Lt = 66.6 (1 - e"°-^^^ (t+0.34) -^^ esocinus. The von Bertalanffy growth
parameters were significantly different between otoliths and scales in S. curvifrons but
did not differ significantly between otoliths and scales in S. niger and between
vertebrae and scales in 5. esocinus. The study of LWR in selected fish species showed
an almost isometric pattern of growth in S. esocinus while in S. curvifrons and S.
niger the value of b was found to be negatively allometric. Estimated parameters of
the LLRs were highly significant (P < 0.001), with detemiination coefficients (r^) >
0.98. K in the selected fish showed values within the range 0.18-1.20 in S. curvifrons,
1.24-2.01 in S. niger and 1.24-1.85 in S. esocinus.
33
Q^ummaty
SUMMARY
The present study was undertaken with a view to evaluate and compare age estimates
from different ageing structures (scales, opercular bones, otoliths, vertebrae and
cleithra); to fit the length-at-age data to the von Bertalanffy growth model; and to
investigate the length-weight, length-length relationships and condition factor in
Schizopyge curvifrons, Schizopyge niger and Shizothorax esocinus. Standard
procedures were followed to prepare and study the ageing structures. The age and
total length data were used for estimating the parameters of von Bertalanffy growth
equation. Body measurements were taken as per the standard procedures to study
length-weight, length-length relationships and condition factor. In S. curvifrons and S.
niger, percent agreement between-readers was highest for otoliths i.e. 95.4% and
94.6%, respectively and in S. esocinus, percent agreement was highest for vertebrae
(96.0%). When otolith ages were compared with other alternative structures viz.,
scales, opercular bones, vertebrae and cleithra highest percent agreement was found
between otoliths and opercular bones (88.4%) in S. cur\nfrons and between otoliths
and scales (86.9%) in S. niger. In S. esocinus, highest agi'eement was found between
vertebrae and opercular bones (87.9%). Otoliths exhibited highest values (P < 0.05) of
mean age estimates in S. cunifrons and S. niger while in S. esocinus highest \alues of
mean age estimates were found for vertebrae. Correlation coefficient of age
estimation from different bony parts of S. cur\nfrons, S. niger and S. esocinus showed
that age estimates from all the age structures were significantly (P < 0.01) coixelated
with each other. The von Bertalanffy model growth parameters were estimated as L^
- 49.8, K = 0.263, to = -0.34 for S. curvifrons; Loo= 44.8, K = 0.255, to = -1.42 for S.
niger and Loo = 66.6, K = 0.278, to = -0.34 for S. esocinus. The length-weight
relationship equations were: Log W = -1.40 -I- 2.69 Log SL (S. curvifrons); Log W =
-1.30 -I- 2.66 LogSL (S. niger) and Log W = -1.92 -I- 3.08 Log SL (5. esocinus).
Results for length-length relationships for selected fish species indicated that the
values between total length and standard length; fork length and total length, and;
standard length and fork length were highly correlated (r > 0.9). The mean condition
factor of S. curvifrons, S. niger and S. esocinus was 0.49, 1.74 and 1.51, respectively.
34
C^^ermces
REFERENCES
Abraham, K. J.; Murty, V. S. R.; Joshi, K. K., 2011: Age and growth studies in
silverbellies along Kerala coast. J. Mar. Biol. Ass. India. 53(2), 172-177.
Anderson, R.; Gutreuter, S., 1983: Length, weight and associated structural indices.
In: Fisheries Techniques. Nielsen, L. and Johnson, D. (Eds). American
Fisheries Society, Bethesda, MD, pp. 283-300.
Andreu-Soler, A.; Oliva-Patema, F. J.; Femandez-Delgado, C.; Torralva, M., 2003:
Age and growth of the sand smelt, Atherina boyeri (Risso 1810), in the Mar
Menor coastal lagoon (SE Iberian Peninsula). J. Appl. Ichthyol. 19, 202-208.
Angelescu, V.; Gneri F. S.; Nam, A., 1958: La del Mar Argentine hake (biology and
taxonomy) (The hake of the Argentino Sea: biology and taxonomy). Seer.
Mar. Serv. Hydrogels. Nav. Pubhc, HI004: 1-224.
Anvar, A. P. H.; Prasad, G.; Balasubramanyam, N. K.; Chandran, L. R.; Raghavan, R.
P., 2008: Weight-length relation of an Asian catfish, Horabagrus brachysoma
(Gunther, 1864), (Siluriformes: Horabagridae) from rivers of the western
ghats, Kerala, India. Acta Ichtyol. Pise. 38(1), 41-44.
Amason, T.; Bjomsson, B.; Steinarsson, A., 2009: Allometric growth and condition
factor of Atlantic cod (Gadiis morhiia) fed to satiation: effects of temperature
and body weight. J. Appl. Ichthyol. 25, 401-406.
Aydin, R.; Calta, M.; Sen, D., 2003: Age and growth of Capoeta tnitia (Pisces:
Cyprinidae) from Keban Dam lake, Turkey. Arch. Pol. Fish. 11(2), 237-243.
Bardach, J. E., 1955: The opercular bone of the yellov/ perch, Perca flavescens, as a
tool for age and growth studies. Copeia 2, 107-109.
Basavaraja, N., 2007: Freshwater fish seed resources in India, pp. 267-327. In: M.G.
Bondad-Reantaso (ed.). Assessment of fi-eshwater fish seed resources for
sustainable aquaculture. FAO Fisheries Technical Paper. No. 501. Rome,
FAO. 2007. 628p.
Beamish, R. J.; Foumier, D. A., 1981: A method for comparing the precision of a set
of age determinafions. Can. J. Fish. Aquat. Sci. 38, 982-983.
Beamish, R. J.; McFarlane, G. A., 1983: The forgotten requirements for age
validation in fisheries biology. T. Am. Fish. Soc. 112, 735-743.
35
Beamish, R. J.; McFarlane, G. A., 1987: Current trends in age determination
methodology. In: Age and Growth of Fish [Eds] Summerfelt, R. C. and Hall,
G. E., Iowa State University Press, Ames, Iowa: 15-42.
Berg, L. S., 1964: Freshwater fishes of the U.S.S.R. and adjacent countries, volume
2, 4th edition. Israel Program for Scientific Translations Ltd, Jerusalem.
(Russian version published 1949).
Beverton, R. J. H., 1992: Patterns of reproductive strategy parameters in some marine
teleost fishes. J. Fish. Biol. 41(Suppl. B), 137-160.
Beverton, R. J. H.; Holt, S. J., 1957: On the dynamics of exploited fish populations.
Chapman and Hall, London, UK.
Beverton, R. J. H.; Holt, S. J., 1959: A review of the Hfe-spans and mortality rates of
fish in nature, and their relationship on growth and other physiological
characteristics. In: Ciba Foundation Colloquia on Ageing. Vol. 5. The lifespan
of animals. J. & A. Churchill Ltd., London, UK.
Beyer, J. E., 1987: On length-weight relationship. Part 1. Corresponding the m.ean
weight of a given length class. Fishbyte 5(1), 11 - 13.
Bhagat, M. J.; Sunder, S., 1984: Some biological aspects of Schizothoraicthys
esocinus Heckel, from Kashmir waters with a note on its utility in culture.
J. Inland Fish. Soc. India 16,42-47.
Bhat, F. A.; Yousuf, A. R.; Balkhi, M. H.; Mahdi, M. D.; Shah, F. A., 2010: Length-
weight relationship and morphometric characteristics of Schizothorax spp. in
the River Lidder of Kashmir. Indian J. Fish. 57,11)-16.
Bolger, T.; Connolly, L., 1989: Ihe selecfion of suitable indexes for the measurement
and analysis offish condition. J. Fish Biol. 34, 171-172.
Booth, A. J., 1997: On the life history of the lesser gurnard {Scorpaeniformes
Triglidae) inhabiting the Agulhas Bank, South Afiica. J. Fish Biol. 51, 1155-
1173.
Boxrucker, J., 1986: A comparison of the otolith and scale methods for aging white
crappie in Oklahoma. N. Am. J. Fish. Manage. 6, 122-128.
Branstetter, S., 1987: Age and growth estimates for blacktip, Carcharhinus limbatus
and spinner, C. Brevipinna sharks from the northwestern Gulf of Mexico.
Copeia 4, 964-974.
Brennan, J. S.; Cailliet, G. M., 1989: Comparative age-deteniiination techniques for
white sturgeon in California. T. Am. Fish. Soc. 118, 296-310.
36
Cailliet, G. M.; Goldman, K. J., 2004: Age determination and validation in
Chondrichthyan fishes. In: Carrier, J., Musick, J. A., Heithaus, M. (Eds), The
Biology of Sharks and their Relatives. CRC Press, New York, pp. 399-447.
Campana, S. E.; Annand, M. C ; McMillan, J. I., 1995: Graphical and statistical
methods for determining the consistency of age determinations. T. Am. Fish.
Soc. 124, 131-138.
Campana, S. E.; Fowler, A. J.; Jones, C. M., 1994: Otolith elemental fingerprinting
for stock identification of Atlantic cod {Gadus morhua) using laser-ablation
ICPMS. Can. J. Fish. Aquat. Sci. 51, 1942-1950.
Campana, S. E.; Moksness, E., 1991: Accuracy and precision of age and hatch date
estimates from otolith microstructure examination. ICES J. Mar. Sci. 48, 303-
316.
Campana, S. E.; Zwanenburg, K. C. T.; Smith, J. N., 1990: "'"iV^^Ra determination
of longevity in redfish. Can. J. Fish. Aquat. Sci. 47, 163-165.
Campana, S., 1995: Expert age determination of 4VW and 4X haddock otoliths by
national and international laboratories. DFO Atl. Fish. Res. Doc. 95/120.
Campana, S.E., 2001: Accuracy, precision and quality control in age detennination.
including a review of the use and abuse of age validation methods. J. Fish
Biol. 59(2), 197-242.
Casselman, J. M., 1990: Growth and relative size of calcified structures of fish. T.
Am. Fish. Soc. 119,673-688.
Casselman, J. M.; Grossman, E. J., 1986: Size, age and gi-owth of trophy muskellunge
and muskellunge-northern pike hybrids - The Cleithruin Project, 1979-1983.
In: Hall G. E.(ed.), Managing muskies - a treatise on the biology and
propagation of muskellunge in North America, American Fisheries Society
Special PubUcation, 15, Bethesda, 93-110.
Chakraborty, S. K., 1994: Growth and mortality estimates of a sciaenid Johnieops
sina (Pisces/ Perciformes) from Bombay waters. Indian J. Mar. Sci. 23, 244-
246.
Chakraborty, S. K., 1995: Growth, mortality and yield per recruit of threadfin bream
Nemipterus japonicas (Bioch) off Bombay. Indian j . Mar. Sci. 24, 107-109.
Chang, W. Y. B., 1982: A statistical method for evaluating the reproducibility of age
determination. Can. J. Fish. Aquat. Sci. 39, 1208-1210.
37
Chamov, E. L., 1993: Life history invariants:some explorations of symmetry in
evolutionary ecology. Oxford University Press, Oxford, UK.
Chen, F.; Chen, Y. F.; He, D. K., 2009: Age and growth of Schizopygopsis
younghusbandi younghusbandi in the Yarlung Zangbo River in Tibet, China.
Environ. Biol. Fish. 86,155-162.
Chen, Y. F.; He, D. K.; Chen, Y. Y., 2002a: Age discrimination of Selincuo
schizothoracini {Gymnocypris selincuoensi) in Selincuo Lake, Tibetan Plateau.
Acta Zoolog Sin 48, 527-533, in Chinese, with English abstract.
Chen, Y. F.; He, D. K.; Duan, Z. H., 2002b: Annuli characters of Selincuo
schizothoracini {Gymnocypris selincuoensi) in Selincuo Lake, Tibet. Acta
Zoolog Sin 4^, 384-392, in Chinese, with English abstract.
Chen, Y.; Jackson D. A.; Harvey H. H., 1992: A comparison of von Bertalanffy and
polynomial functions in the modeling fish growth data. Can. J. Fish. Aquat.
Sci. 49, 1228-1235.
Clark, W. G., 1991: Groundfish exploitation rates based on life history parameters.
Can. J. Fish. Aquat. Sci. 48, 734-750.
Coad, B. W., 1995: Freshwater fishes of Iran. Acta Sci. Nat. Acad. Sci. Brno. 29(1),
1-64.
Coelho, R.; Erzini, K., 2002: Age and growth of the undulate ray. Raja undulata, in
the Algarve (southern Portugal). J. Mar. Biol. Ass. U.K. 82, 987-990.
Cope, J. M.; Punt, A. E., 2007: Admitting aging error when fitting growth curves: an
example using the von Bertanlanffy growth function with random effects. Can.
J. Fish. Aquat. Sci. 64, 205-218.
Dar, S. A.; Najar, A. M.; Balkhi, M. H.; Rather, M. A.; Sharma, R., 2012: Length
weight relationship and relative condition factor of Schizopyge esocinus
(Heckel, 1838) form Jhelum River, Kashmir. Int. J. Aqua. Sci. 3(1), 29-36.
Day, F., 1878: The Fishes of India; being a natural history of the fishes known to
inhabit the seas and fresh waters of India, Burma and Ceylon, Vol. 1.
published by Bernard Qualitch, 15 Piccadilly London, London: pp. 529-533.
DeVries, D. R.; Erie, R. V., 1996: Determination of age and growth. Pages 483-512
In: Murphy, B. R. and Willis, D. W. (Eds). Fisheries techniques, 2nd edition.
American Fisheries Society, Bethesda, Maryland.
38
Dhaiize, R.; Dhanze, J. R., 1997: Biology of scale carp and grass carp. 1. Length-
weight relationship and growth performance under the agroclimatic zone 1 of
Himachal Pradesh. Indian J. fish. 44(3), 255-263.
Diaz, L. S.; Roa, A.; Garcia, C. B.; Acero, A.; Navas, G., 2000: Length-weight
relationships of demersal fishes fi-om the upper confinental slope off
Colombia. The ICLARM Quarterly 23(3), 23-25.
Dominguez, S. R.; Pajuelo J. G.; Lorenzo J. M., 2006: Age and growth of the
sharpsnout seabream Diplodus puntazzo (Cetti, 1777) inhabiting the Canarian
archipelago, estimated by reading otoliths and by back-calculation. Fish. Res.
81: 142-148.
Doray, M.; Stequert, B.; Taquet, M., 2004: Age and growth of blackfin tuna {Thunmis
atlanticus) caught under moored fish aggregating devices, around Martinique
island. Aquat. Living Resour. 17, 13-18.
Dua, A.; Kumar, K., 2006: Age and growth patterns mChanna marulius from Harike
wetland (a Ramsar site), Punjab, India. J. Environ. Biol. 27(2), 377-380.
Ecoufin, J. M.; Albaret, J. J., 2C03: Length-weight relafionship of 52 fish species from
West African estuaries and lagoons. Cybium 27, 3-9.
Enckson, C. M., 1983: Age detemiinafion of Manitoban walleyes usmg otoliths.
dorsal spines, and scales. N. Am. J. Fish. Manage. 3, 176-18 L
Euchner, R. B., 1988: Collection, preparation and use of northern pike {Esox hicitis)
cleithra for age detennination. Ministry of Environment and Parks.
Recreafional Fisheries Branch, Fort St. John, B.C. Rep. No. PCE 20.
Ezenwa, B. I. O.; Ikusemiju, K., 1981: Age and growth determinations in the catfish,
Chtysichthys nigrodigitatus (Lacepede) by use of the dorsal spine. Fish Biol.
19,345-351.
Fafioye, O. O.; Oluajo, O. A., 2005: Length- weight relafionships of five fish species
in Epe lagoon, Nigeria. Aft-. J. Biotechnol. 4(7), 749-751.
FAO, 2012: The state of world fisheries and aquaculture. FAO Fisheries and
Aquaculture Department, Rome.
Filmalter, J. D.; Weyl, O. L. F.; Sauer, W., 2009: Otoliths and vertebrae as potential
hard structures for ageing south African yellowfin tuna Thunnus albacores.
Afr. J. Mar. Sci. 31(2), 271-276.
39
Frisk, M. G.; Miller, T. J.; Fogarty, M. J., 2001: Estimation and analysis of biological
parameters in elasmobranch fishes: a comparative life history study. Can. J.
Fish. Aquat. Sci. 58, 969-981.
Froese, R.; Pauly, D. (Eds), 2005: FishBase. World Wide Web electronic publication.
Available at: http://www.fishbase.org; accessed on 20 September 2005.
Froese, R.; Pauly, D. (Eds), 2011: FishBase. World Wide Web electronic publication.
www.fishbase.org: accessed on August 2011.
Froese, R.; Pauly, D. (Eds), 2012: FishBase. World Wide Web electronic publication.
www.fishbase.org: accessed on August 2012.
Frost, W. E.; Kipling, C, 1959: The detennination of age and growth of pike, Esox
hiccius (Linnaeus) from scales and opercular bones. J. Anim. Ecol. 23, 314-
341.
Gallucci, V. F.; Quinn II, T. J., 1979: Reparameterizing, fitting, and testing a simple
growth model. T. Am. Fish. Soc. 108, 14-25.
Gang, H.; Bo, F.; Huosheng, L.; Junfeng, Z., 2008: Age and gi'owth characteristics of
crimson sea bream Paragyrops edita Tanaka in Beibu gulf J. Ocean Univ.
China 7(4), 457-465.
Giacalone, V. M.; D'Aiina, G.; Badalamenti, F.; Pipitone, C, 2010: Weight-length
relationships and condition factor trends for thirty-eight fish species in trawled
and untrawled areas off the coast of northern Sicily (central Mediterranean
Sea). J. Appl. Ichthyol. 1-4.
Gocer, M.; Ekingen, G., 2005: Comparison of various bony structures for age
determinafion of Liza ramada (Risso) population from the Mersin bay. E.U. J.
Fish. Aquat. Sci. 22, 211-213.
Goldman, K. J., 2004: Age and growth of elasmobranch fishes. In: Musick, J. A. And
Bonfil, R. (Eds) Elasmobranch Fisheries Management techniques. Asia Pacific
Economic Cooperation, Singapore, pp. 97-132.
Gomez, K. A.; Gomez, A. A., 1984: Statisfical procedures for agricultural research.
Singapore: John Wiley and Sons. p. 680.
Gomez-Marquez, J. L.; Pena-Mendoza, B.; Salgado-Ugarte, I. H.; Arredondo-
Figueroa, J. L., 2008: Age and growth of the tilapia, Oreochromis niloticus
(Perciformes: Cichlidae) from a tropical shallow lake in Mexico. Rev. Biol.
Trop. 56(2), 875-884.
40
Gonzalez-Gandara, C; Perez-Diaz E.; Santos-Rodriguez, L.; Arias-Gonzalez, J. E.,
2003: Length-weight relationships of coral reef fishes from the Alacran reef,
Yucatan, Mexico. NAGA, ICLARM Quarteriy 26(1), 14-16.
Govind, B. V.; Gopal Y. S., 1966: Cleithrum bone as an aid in the age and growth
studies ofSilonia childrenii (Sykes). Sci. Cult. 32(3), 156-158.
Guinn, D. A.; Hallberg, J. E., 1990: Precision of estimated ages of burbot using
vertebrae and otoliths. 1990. Alaska Department of Fish and Game. Fishery
Data Series No. 90-17, Anchorage.
Gulland, J. A., 1983: Fish stock assessment: a manual of basic methods. John Wiley
& Sons, Chichester, UK.
Gumus, A.; Bostanci, D.; Yilmaz, S.; Polat, N., 2007: Age determination of
Scardinius erythrophthalmiis (Cyprinidae) inhabiting Bafra Fish Lakes
(Samsun, Turkey) based on otolith readings and marginal increment analysis.
Cybium 31(1), 59-66.
Gumus, A.; Sahinoz, E.; Dogu, Z.; Polat, N., 2010: Age and growth of the
Mesopotamian spiny eel, Mastacembelus mastacembelus (Banks & Soiender,
1794). from southern Anatolia. Turk. J. Zool. 34, 399-407.
Gupta, B. K.; Sarkar, U. K.; Bhardwaj, S. K.; Pal, A., 2011: Condition factor, length-
weight, length-length relationships of an endangered fish Ompok pabda
(Hamilton 1822) (Siluriformes: Siluridae) from the River Gomti. a tributary of
the River Ganga, India. J. Appl. Ichthyol. 27, 962-964.
Gursoy, C; Tarkan, A. S.; Gaygusuz, O.; Acipinar, H., 2005: A comparison of ageing
techniques to improve precision of age estimation from fish scales. E.U. J.
Fish. Aquat. Sci. 22(3-4), 423-425.
Haddon, M., 2001: Modelling and quantitative methods in fisheries. Chapman and
Hall, London, UK.
Hale, L. F.; Lowe, C. G., 2008: Age and growth of the sfingray Urobatis halleri at
Seal beach, California. J. Fis Biol. 73, 510-523.
Hammers, B. E.; Miranda, L. E., 1991: Comparison of methods for esfimating age,
growth, and related populafion characteristics of white crappies. N, Am. J.
Fish. Manage. 11, 492-498.
Haniffa, M. A.; Nagaraj'an, M.; Gopalakrishnan, A., 2006: Length-weight
relafionships of Channa punctata (Bloch, 1793) from Western Ghats rivers of
Tamil Nadu. J. Appl. Ichthyol. 22, 308-309.
41
Harrison, E. J.; Hadley W. F., 1979: A comparison of the use of cleithra to the use of
scales for age and growth studies. T. Am. Fish. Soc. 108, 452-456.
Holden, M. J., 1972: Are long-term sustainable fisheries for elasmobranchs possible?
J. cons. - Cons. int. explor. mer. 164, 360-367.
Hossain, M. D., 2010: Length-weight, length-length relationships and condition
factors of three schibid catfishes from the Padma River, northwestern
Bangladesh. Asian Fish. Sci. 23, 329-339.
Hossain, M. Y.; Ahmed, Z. F.; Leunda, P. M.; Jasmnne, S.; Oscoz, J.; Miranda, R.;
Ohtomi, J., 2006: Condition, length-weight and length-length relationships of
the Asian striped catfish Mystus vittatus (Bloch, 1794) (Silurifonnes:
Bagridae) in the Mathabhanga River, southwestern Bangladesh. J. App.
Ichthyol. 22, 304-307.
Hossain, M. Y.; Rahman, M. M.; Fulanda, B.; Jewel, M. A. S.; Ahamed, F.; Ohtomi,
J., 2012: Length-weight relationships of five threatened fish species from the
Jamuna (Brahmaputra river tributary) river, northern Bangladesh. J. Appl.
Ichthyol. 28, 275-277.
HUG, B.; Xie, C. X.; Ma, B. S.; Yang, X. F.; Huang, H. P., 2012: Age and growth of
Oxygymnocypris stewartii (Cyprinidae: Schizothoracinae) in the Yarlung
Tsangpo river, Tibet, China. Zool. Studies 51(2), 185-194.
Ilkyaz, A. T.; Metin, G.; Soykan, O.; Kinacigil, H. T., 2010: Age, growth and sexual
development of solenette, Buglossidium luteum (Risso, 1810), in the central
Aegean sea. J. Appl. Ichthyol. 26,436-440.
Isen-nann, D. A.; Meerbeek, J. R.; Scholten, G. D.; Willis, D. W., 2003: Evaluation of
three different structures used for walleye age estimation with emphasis on
removal and processing fimes. N. Am. J. Fish. Manage. 23, 625-631.
Jackson, N. D.; Garvey, J. E.; Colombo, R. E., 2007: Comparing aging precision of
calcified structures in shovelnose sturgeon. J. App. Ichthyol. 23, 525-528.
Jellyman, D. J., 1980: Age, growth, and reproduction of perch, Perca fluviatilis L., in
lakepounui. New Zeal. J. Mar. Freshw. Res. 14(4), 391-400.
Jensen, A. L., 1998: Simulation of reladons among fish life history parameters with a
bioenergetics-based population model. Can. J. Fish. Aquat. Sci. 55(2), 353-
357.
Jhingran, V. G., 1959: Studies on the age and growth of Cirrhinus mrigala (Ham.)
from the river Gang. Proc. Nat. Inst. Sci. India 25B(3), 107-137.
42
Jia, Y. T.; Chen, Y. F., 2011: Age structure and growth characteristics of the endemic
fish Oxygymnocypris stewartii (Cypriniformes: Schizothoracinae) in the
Yarlung Tsangno river, Tibet. Zool. Studies 50(1), 69-75.
Johal, M. S.; Tandon, K. K., 1992: Age and growth of the carp Catla catla (Hamilton,
1822) from the northern India. Fish. Res. 14, 83-90.
Kamilov, B. G., 1984: Morphology of growth structures in silver carp
Hypophthalmichthys molitrix, in relation to estimation of age and growth rate.
J. Ichthyol. 6, 1003-1013.
Kamilov, G.; Urchinov, Z. U., 1995: Fish and fisheries in Uzbekistan under the
impact of irrigated agriculture, p. 10-41 In: T. Petr (ed.) Inland fisheries under
the impact of imgated agriculture;Central Asia. FAO Fisheries Circular No.
894.
Kano, Y., 2000: Age and growth of the Ajime-loach, Niwaella delicata, in the Yura
River, Kyoto, Japan. Ichthyol. Res. 47(2), 183-186.
Karatas, M.; Cicek, E.; Basusta, A.; Basusta, N., 2007: Age, growth and mortality of
common carp (Cyprimts carpio Linneaus, 1758). J. Appl. Biol. Sci. 1(3), 81-
85.
Karlou-Riga, C; Sinis A., 1997: Age and growth of horse mackerel, Trachurus
trachurus (L.), in the Gulf of Saronikos (Greece). Fish. Res. 22, 157-171.
Kerstan, M., 1985: Age, growth, maturity and mortality estimates of horse mackerel
{Tachurus trachurus) from the waters of Great Britain and Ireland in 1984.
Arch. Fischwiss. 36(1/2), 115-154.
Khan, M. A.; Khan, S., 2009: Comparison of age estimates from scale, opercular
bone, otolith, vertebrae and dorsal fin ray in Labeo rohita (Hamilton), Catla
catla (Hamilton) and Channa marulius (Hamilton). Fish. Res. 100, 255-259.
Khan, M. A.; Khan, S.; Miyan, K., 2011a: Precision of aging structures for Indian
major carp, Cirrhinus mrigala.from the River Ganga. J. Freshwat. Ecol.
26(2), 231-239.
Khan, M. A.; Khan, S.; Miyan, K., 2012a: Length-weight relationship of giant
snakehead, Channa marulius and stinging catfish, Heteropneustes fossilis
fi-om the river Ganga, India. J. Appl. Ichthyol. 28, 154-155.
Khan, M. A.; Khan, S.; Miyan, K., 2012b: Studies on length-weight and length-length
relationships of four freshwater fishes collected from River Ganga. J. Fish.
Aqaut. Sci. 7(6), 481-484.
43
Khan, M. A.; Sabah, 2013: Length-weight and length-length relationships for five fish
species from Kashmir valley. J. Appl. Ichthyol. 29, 283-284.
Khan, R. A.; Siddiqui, A. Q., 1973: Studies on the age and growth of rohu, Labeo
rohita (Ham.) from a pond (Moat) and rivers Ganga and Yamuna. Proc. Indian
natn. Sci. Acad. 39, 582-597.
Khan, S.; Khan, M. A.; Miyan, K., 201 lb: Comparison of age estimates from otoliths,
vertebrae, and pectoral spines in African sharptooth catfish, Clarias
gariepinus (Burchell). Eston. J. Ecol. 60(3), 183-193.
Khan, S.; Khan, M. A.; Miyan, K.; Mubark, M., 2011c: Length-weight relationships
for nine freshwater teleosts collected from River Ganga, India. Int. J. Zool.
Res. 7(6), 401-405.
Kimura D. K.; Kastelle, C. R.; Goetz, B. J.; Gburski, C. M.; Buslov, A. V., 2006:
Corroborating the ages of walleye pollock {Theragra chalcogramma). Mar.
Freshwater Res. 57, 323-332.
Kimura, D. K., 1980: Likelihood estimates for the von Bertalanffy growth curve. Fish.
Bull. 77, 765-776.
King, M., 1995: Fisheries Biology, Assessment and Management. Reprinted edition.
Fishing News Books, Oxford, UK, 1-341.
King, R. P., 1996a: Length-weight relationships of Nigeria Freshwater fishes. Naga
ICLARIMQ 19(3), 49 -52 .
King, R. P., 1996b: Length-weight relationship of Nigerian Coastal water fishes.
Fishbyte 19(4), 53 - 58.
Koch, J. D.; Quist, M. C; Hansen, K. A., 2009: Precision of hard structures used to
esfimate age of bowfin in the upper Mississippi river. N. Am. J. Fish.
Manage. 29, 506-511.
Kruse, C. G.; Guy, C. S.; Wilhs, D. W., 1993: Comparison of otolith and scale age
characteristics for black crappies collected from South Dakota waters. N. Am.
J. Fish. Manage. 13, 856-858.
Kullander, S. O.; Fang, F.; Delling, B.; Ahlander, E. 1999: The fishes of the Kashmir
Valley. In: River Jhelum, Kashmir Valley, impacts on the aquatic
environment. Nyman, L. (Ed), pp. 99-163.
Kumar, K. H.; Kiran, B. R.; Purushotham, R.; Puttaiah, E. T.; Manjappa, S., 2006:
Length-weight relationship of Cyprinid fish, Rasbora daniconius (Hamilton-
44
Buchanan) from Sharavathi reservoir, Kamataka. Zoos' Print Journal 21(1),
2140-2141.
Kumar, S. G.; Mercy, T. V. A.; John, K. C , 1999: Length weight relationship in the
catfish Hombagrus brachysoma (Gunther). Indian J. Fish. 46, 191-193.
Kume, G.; Furumitsu, K.; Yamaguchi, A., 2008: Age, growth and age at sexual
maturity of fan ray Platyrhina sinensis (Batoidea: Platyrhinidae) in Ariake
bay, Japan. Fish. Sci. 74, 736-742.
Kumolu-Johnson, C. A.; Ndimele, P. E., 2010: Length-weight relationships and
condition factors of twenty-one fish species in Ologe lagoon, Lagos, Nigeria.
Asian J. Agric. Sci. 2(4), 174-179.
Laine. A. O.; Momot, W. T.; Ryan, P., 1991: Accuracy of using scales and cleithra for
agmg northern pike from an oligotrophic Ontario Lake. N. Am. J. Fish.
Manage. 11,220-225.
Le Cren, E. D., 1951: The length-weight relationship and seasonal cycle in gonad
weight and condition in the perch {Perca fluviatilis). J. Anim. Ecol. 20(2),
201-219.
Li, X. Q.; Chen, Y. F.; He, D. K.; Chen, F., 2009: Otolith characteristics and age
determination of an endemic Ptychobarbus dipogon (Regan, 1905)
(Cypnnidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet. Environ.
Biol. Fish. 86,53-61.
Li, X.; Chen, Y., 2009: Age structure, growth and mortality estimates of an endemic
Ptychobarbus dipogon (Regan, 1905) (Cyprinidae: Schizothoracinae) in the
Lhasa river, Tibet. Environ. Biol. Fish 86, 97-105.
Licandeo, R. R.; Lamilla, J. G.; Rubilar, P. G.; Vegas, R. M., 2006: Age, growth, and
sexual maturity of the yellownose skate Dipturus chilensis in the south-eastern
Pacific. J. Fish Biol. 68,488-506.
Liu, K.; Lee, M.; Joung, S.; Chang, Y., 2009: Age and growth estimates of the
sharptail mola, Masturus lanceolatus, in waters of eastern Taiwan. Fish. Res.
95, 154-160.
Long, J. M.; Fisher, W. L., 2001: Precision and bias of largemouth, smallmouth, and
spotted bass ages estimated fi'om scales, whole otoliths, and sectioned otoliths.
N. Am. J. Fish. Manage. 21, 636-645.
45
Ma, B. S.; Xie, C. X.; Huo, B.; Yang, X. F.; Huang, H. P., 2010: Age and growth of a
long-lived fish Schizothorax o'connori in the Yarlung Tsangpo river, Tibet.
Zool. Studies 49(6), 749-759.
Ma, B.; Xie, C; Huo, B.; Yang, X.; Li, P., 2011: Age validation and comparison of
otolith, vertebrae and opercular bone for estimating age of Schizothorax
o 'connori in the Yarlung Tsangpo river, Tibet. Env. Biol. Fish. 90, 159-169.
Maceina, M. J.; Sammons, S. M., 2006: An evaluation of different structures to age
freshwater fish from a northeastern US river. Fisheries Manag. Ecol. 13, 237-
242.
Marquardt, D. W., 1963: An algorithm for least-squares estimation of nonHnear
parameters. J. Soc. Indust. Appl. Math. 11(2), 431-441.
Mather, F. J., Ill; Mason, J. M.; Jones, A. C, 1995; Historical document: life history
and fisheries of Atlantic bluefin tuna. NOAA Tec. Mem. NMFS-SEFSC 370,
165 pp.
Mathew, K.; Zacharia, P. U., 1982: On the age and growth of three Indian major carps
from Hirakud reservoir. Bull. Dept. Mar. Sci. Univ. Cochin. XIII, 81-95.
McConnell, W. J., 1952: The opercular bone as an indicator of age and growth of the
carp, Cyprinus carpio Linnaeus. Trans. Am. Fish. Soc. 81, 138-149.
MegaJofonou, P., 2000: Age and growth of Mediten-anean albacore. J. Fish Biol. 57,
700-715.
Mendes, B.; Fonseca P.; Campos, A., 2004: Weight -length relationships for 46 fish
species of the Portuguese west coast. J. Appl. Ichthyol. 20, 355-361.
Menon, A. G. K., 1999: Check list - fresh water fishes of India. Rec. Zool. Surv.
India, Misc. Publ., Occas. Pap. No. 175, 366 p.
Metcalf, S. J.; Swearer, S. E., 2005: Non-destructive ageing in Notolabrus tetricus
using dorsal spines with an emphasis on the benefits for protected, endangered
and fished species. J. Fish Biol. 66, 1740-1747.
Mills, K. H.; Chalanchuk, S. M., 2004: The fin-ray method of aging lake whitefish.
Ann. Zool. Fennici 41, 215-223.
Mir, J. I.; Shabir, R.; Mir, F. A., 2012: Length-weight relationship and condition
factor of Schizopyge curvifrons (Heckel, 1838) from River Jhelum, Kashmir,
India. Worid J. Fish Mar. Sci. 4(3), 325-329.
Misra, R. K., 1980: Stafistical comparison of several growth curves of the von
Bertalaffy type. Can. J. Fish. Aquat. Sci. 37, 920-926.
46
Molur, S.; Walker, S., 1998: Report of the workshop conservation assessment and
management plan for freshwater fishes of India. Zoo Outreach Organization,
Conservation Breeding Specialist Group, (CBSG), Coimbatore, India, 156 p.
Monteiro, P.; Rentes, L.; Coelho, R.; Correia, C; Goncalves, J. M. S.; Lino, P. G.;
Ribeiro, J.; Erzini, K., 2006: Age and growth, mortality, reproduction and
relative yield per recruit of the bogue, Boops boops Linne, 1758 (Sparidae),
from the Algarve (south of Portugal) longline fishery. J. Appl. Ichthyol. 22,
345-352.
Morrow, Jr. J. V.; Kirk, J. P.; Killgore, J.; George, S. G., 1998: Age, growth, and
mortality of shovelnose sturgeon in the lower Mississippi river. N. Am. J.
Fish. Manage. 18, 725-730.
Moutopoulos, D. K., Stergiou, K.I., 2002: Length-weight and length-length
relationships offish species from Aegean Sea (Greece). J. App. Ichthyol. 18,
200-203.
Muralidharan. M.; Arunachalam, M.; Raja, M., 2011: Length-weight relationships for
fish species from Cauvery river at Hogenakal in South India. J. Appl. Ichthyol.
27, 968-969.
Musick, J. A., 1999: Criteria to define extinction risk in marine fishes. Fisheries 24.
6-14.
Nargis. A., 2006: Determination of age and growth of Catla catla (Hamilton) from
opercular bones. J. Bio-sci. 14, 143-145.
Niewinski, B. C ; Ferreri, C. P.; 1999: A comparison of three structures for estimating
the age of yellow perch. N. Am. J. Fish. Manage. 19, 872-877.
Nuevo, M., Sheehan, R. J.; Heidinger, R, C, 2004: Accuracy and precision of age
determination techniques for Mississippi River bigliead carp
Hypophthalmichthys nobilis (Richardson 1845) using pectoral spines and
scales. Arch. Hydrobiol. 160,45-56.
Oni, S. K.; Olayemi, J. Y.; Adegboye, J. D., 1983: Comparative physiology of three
ecologically distinct fresh water fishes, Alestes nurse (Ruppell), Synodontis
schall (Bloch), S. Schneider and Tilapia zilli (Gervais). J. Fish Biol. 22, 105-
109.
Ozcan, G.; Balik, S., 2009: Age and growth of Bassan barbell, Barbus pectoralis
(Actinopterygii: Cypriniformes: Cyprinidae), under conditions of a dam
reservoir. ACTA Ichthyologica et Piscatoria 39(1), 27-32.
47
Pandey, A. C; Sharma, M. K., 1997: A preliminary study on the relative condition
factor of exotic carps cultivated in sodic soil pond. Indian J. Fish. 44(2), 221-
223.
Pandey, A. C; Sharma, M. K., 1998: A Preliminary Study on the Relative condition
factor of exotic Carps Cultivated on Sodic Soil Pond Conditions in U.P, India.
Indian J. Fish. 45(2), 207-210.
Patiyal, R. S.; Sharma, R. C; Punia, P.; Goswami, M.; Lakra, W. S., 2010: Length-
weight relationship of Tor putitora (Hamilton, 1822) from the Ladhiya river,
Uttarakhand, India. J. Appl. Ichthyol. 26, 472-473.
Pauly, D.; Munro, J. L, 1984: Once more on the comparison of growth in fish and
invertebrates. ICLARM Fishbyte 2, 21.
Penha, J. M. F.; Mateus, L. A. F.; Barbieri, G., 2004: Age and growth of the porthole
shovelnose catfish {Hemisorubim platyrhynchos) in the Pantanal. Braz. J. Biol.
64(4), 833-840
Petrakis, G.; Stergiou, K. I., 1995: Weight-length relationships for 33 fish species in
Greek waters. Fish. Res. 21, 465-469.
Phelps, Q. E.; Edwards K. R.; Willis, D. W., 2007: Precision of five structures for
estimating age of common carp. N. Am. J. Fish. Manage. 27, 103-105.
Polat, N.; Bostanci, D.; Yiimaz, S., 2001: Comparable age determination in different
bony structures of Pleuronectes flesus lusciis Pallas, 1811 inhabiting the Balck
sea. Turk. J. Zool. 25, 441-446.
Prasad, G.; Ali, P. H. A., 2007: Length-weight relationship of a Cyprinid fish Puntiiis
filamentosus from Chalakudy river, Kerala. Zoos" Print Journal 22(3), 2637-
2638.
Pratt, H. L. J.; Casey, J. G., 1990: Shark reproductive strategies as a hmiting factor in
directed fisheries, with a review of Holden's method of estimating growth
parameters. In: Pratt, H. L. J., Gruber, S. H., Taniuchi, T. S. (Eds.),
Elasmobranchs as Living Resources: Advances in the Biology, Ecology,
Systemafics, and the Status of the Fisheries NOAA Technical Report NMFS
90, pp. 97-109.
Qasim, S. Z., 1973: Some implicafions of the problem of age and growth in marine
fishes fi-om the Indian waters. Indian J. Fish. 40(2), 351-371.
Qasim, S. Z.; Bhatt, V. G., 1964: Occurrence of the growth in the opercular bones of
the fi-eshwater murrels Ophicephaluspunctatus (Bloch). Curr. Sci. 33, 19-20.
48
Qasim, S. Z.; Bhatt, V. G., 1966: The growth of the freshwater murrel, Ophicephalus
punctatus Bloch. Hydrobiol. 27(3-4), 289-316.
Qiu, H.; Chen, Y. F., 2009: Age and growth of Schizothorax waltoni in the Yarlung
Tsangpo river in Tibet, China. Ichthyol. Res. 56, 260-265.
Quinn, T. J.; Deriso, R. B., 1999: Quantitative fish dynamics. Oxford University
Press, New York.
Quist, M. C; Jackson, Z. J.; Bower, M. R.; Hubert, W. A., 2007: Precision of hard
structures used to estimate age of riverine Catostomids and Cyprinids in the
Upper Colorado River Basin. N. Am. J. Fish. Manage. 27, 643-649.
Rafail, S. Z., 1973: A simple and precise method for fitting a von Bertalanffy growth
curve. Mar. Biol. 19, 354-358.
Raina, H. S.; Petr, T., 1999: Coldwater fish and fisheries in the Indian Himalayas:
lakes and reservoirs, pp. 64-88. In T. Petr (ed.) Fish and fisheries at higher
aUitudes:Asia. FAO Fish. Tech. Pap. No. 385. FAO, Rome. 304 p.
Rajkumar, M.; Antony, P. J.; Trilles, J. P., 2006: Length- weight relationship of asian
seabass (Lates calcarifer Bloch, 1790) from Pichavaram mangrove waters.
South East coast of India. Asian Fish. Sci. 19, 177-183.
Ramesh,, R.; Ravichandran, S.; Rameshkumar, G., 2009: Analysis of age and growth
rate of Turbo brunnens. World J. Dairy & Food Sciences 4(1), 56-64.
Rao, C. R., 1958: Some statistical methods for comparison of growth curves.
Biometrics 14, 1-17.
Ricker, W. E., 1968: Methods for Assessment of Fish Production in Freshwaters.
Blackwell Scientific Publications, Oxford, 313 p.
Ricker, W. E., 1975: Computation and interpretation of biological statistics offish
populations. J. Fish. Res. Board Can. 191, 382 p.
Ritterbusch-Nauwerck, B., 1995: Condition or corpulence, fitness or fatness: a
discussion of terms. Arch. Hydrobiol. 46, 109-112.
Rohit, P.; Rao, G. S.; Rammohan, K., 2012: Age, growth and population structure of
the yellowfin tuna Thunnus albacores (Bonnaterre, 1788) exploited along the
east coast of India. Indian J. Fish. 59(1), 1-6.
Ruiz-Campos, G.; Gonzalez Acosta, A. F.; De La Cruz Aguero, J., 2006: Length-
weight and length-length relationships for some continental fishes of
northwestern Baja California, Mexico. J. App. Ichthyol. 22, 314-315.
49
Saha, S. N.; Vijayanand, P.; Rajagopal, S., 2009: Length-weight relationship and
relative condition factor in Thenus orientalis (Lund, 1793) along east coast of
India. Curr. Res. J. Biol. Sci. 1(2), 11-14.
Sani, R.; Gupta, B. K.; Sarkar, U. K.; Pandey, A.; Dubey, V. K.; Lakra, W. S., 2010:
Length-weight relationships of 14 Indian freshwater fish species from the
Betwa (Yamuna River tributary) and Gomti (Ganga River tributary) rivers. J.
Appl. Ichthyol. 26,456-459.
Santamaria, N.; Bello, G.; Corriero, A.; Deflorio, M.; Vassallo-Agius, R.; Bok, T.; De
Metrio, G., 2009: Age and growth of Atlantic bluefin tuna, Thunnus thynmis
(Osteichthyes: Thunnidae), in the Mediterranean sea. J. Appl. Ichthyol. 25, 38-
45.
Sarkar, U. K.; Deepak, P. K.; Negi, R. S., 2009: Length-weight relationship of clown
knifefish Chitala chitala (Hamilton 1822) from the River Ganga basin, India.
J. Appl. Ichthyol. 25, 232-233.
Sarkar, U. K.; Negi, R. S.; Deepak, P. K.; Lakra, W. S.; Paul, S. K., 2008: Biological
parameters of the endangered fish Chitala chitala (Osteoglossiformes:
Notopteridae) from some Indian rivers. Fish. Res. 90, 170-177.
Secor, D. H.; Trice, T. M.; Homick, H. T., 1995: Validation of otolith-based ageing
and a comparison of otolith and scale-based ageing in mark-recaptured
Chesapeake Bay striped bass, Morone saxatilis. Fish. Bull. 93, 186-190.
Shafi, S.; Yousuf, A. R., 2012: Length-weight relationship and condition factor of
Schizothorax niger (Heckel, 1838) Misra from Dal lake, Kashmir. Int. J. Sci.
Res. Pub. 2(3), 1-3.
Shamsan, E. F.; Ansari, Z. A., 2010: Study of age and growth of Indian sand whiting,
Sillago sihama (Forsskal) from Zuari estuary, Goa. Indian J. Mar. Sci. 39(1),
68-73.
Sharp, D.; Bernard, D. R., 1988: Precision of estimated ages of lake trout from five
calcified structures. N. Am. J. Fish. Manage. 8, 367-372.
Shrestha, T. K., 1990: Resource ecology of the Himalayan waters. Curriculum
Development Centre, Tribhuvan University, Kathmandu, Nepal. 645 p.
Singh, D.; Sharma, R. C, 1995: Age and growth of a Himalayan teleost Schizothorax
richardsnii (Gray) from the Garhwal hills (India). Fish. Res. 24, 321-329.
50
Sivashanthini, K., 2008: Length-weight relationships and condition of Gerreids
(Pisces: Gerreidae) from the Parangipettai waters (SE coast of India). Asian
Fish. Sci. 21,405-419.
Stamatopoulos, C; Caddy, J. F., 1989: Estimation of von Bertalanffy growth
parameters: a versatile linear regression approach. J. Cons. Int. Explor. Mer.
45, 200-208.
Stevenson, J. T.; Secor, D. H., 1999: Age determination and growth of Hudson river
atlantic sturgeon, Acipenser oxyhnchus. Fish. Bull. 97, 153-166.
Sun, C. L.; Wang, S. P.; Yeh, S. Z., 2002: Age and growth of the swordfish {Xiphias
gladius L.) in the waters around Taiwan detennined from anal-fin rays. Fish,
Bull. 100, 822-835.
Sunder, S.; Subla, B. A., 1984: Occurrence of growth rings on the hard parts of
Schizothorax curvifrons Heckel. Curr. Sci. 53(16), 860-862.
Sutherland, S. J., 2006: Templates for calculating ageing precision, http:
//www.nefsc.noaa.gov/fbi/age-prec/) (accessed 2008-11-16).
Sylvester, R. M.; Deny, C. R., 2006: Comparison of white sucker age estimates from
scales, pectoral fin rays, and otoliths. N. Am. J. Fish. Manage. 26, 24-31.
Talwar, P. K.; Jhingran, A. G., 1991: Inland fishes of India and adjacent countries, vol
1. A.A. Balkema, Rotterdam. 541 p.
Tandon, K. K.; Johal, M. S., 1996: Age and Growth in Indian Freshwater Fishes.
Delhi: Narendra Publishing House.
Tarkan, A. S.; Ozulug, iM.; Gaygusuz, O.; Gaygusuz, C. G.; Sac, G., 2009: Length-
weight relafionships of six freshwater fishes from small streams flowing into
Lake Sapanca, NW Turkey. J. App. Ichthyol. 25, 230-231.
Teixeira-de Mello, F.; Eguren, G., 2008: Prevalence and intensity of black-spot
disease in fish community from Canada del Dragon stream (Montevideo.
Uruguay). Limnefica 27, 251-258.
Teixeira-de Mello, F.; Gonzalez-Bergonzoni, 1.; Viana, F.; Saizar, C , 2011: Length-
weight relafionships of 26 fish species from the middle secfion of the Negro
River (Tacuarembo-Durazno, Uruguay). J. Appl. Ichthyol. 27, 1413-1415.
Teixeira-de Mello, F.; Iglesias, C; Borthagaray, A. I.; Mazzeo, N.; Vilches, J.; Larrea,
D.; Ballabio, R., 2006: Onthogenic allometric coefficient changes. Implicances
of diet shift and morphometric attributes in Hoplias malabaricus (Bloch)
(Characiforme, Erythrinidae). J. Fish Biol. 69, 1770-1778.
51
Tomlinson, P. K.; Abramson, N. J., 1961: Fitting a von Bertalanffy growth curve by
least squares. California Department of Fish and Game, Sacramento, Calif
Fish. Bull. No. 116.
Tribuzio, C. A.; Kruse, G. H.; Fujioka, J. T., 2010: Age and growth of spiny dogfish
{Squalus acanthias) in the Gulf of Alaska: analysis of alternate growth
models. Fish. Bull. 108, 119-135.
Ujjania, N. C, 2012: Comparative age and growth of Indian major carp {Catla catla
ham. 1822) in selected water bodies of Southern Rajasthan, India. Res J.
Recent Sci. 1, 17-22.
Vasudevappa, C; James, P. S. B. R., 1988: Age and growth of the marine catfish
Tachysunis diissumieri (Val.) along the Dakshina Kannada coast. In: M.
Mohan Joseph (Ed.) The first Indian Fisheries Forum, Proceedings. Asian
Fisheries society, Indian branch, Mangalore. pp. 225-228.
Von Bertalanffy, L., 1938: A quanfitafive theory of organic growth (Inquires on
growth laws II). Hum. Biol. 10,181-213.
Walford, L., 1946: A new graphical method of describing the growth of animals. Biol.
Bull. 90, 141-147.
Walker, K. F.; Yang, H. Z., 1999: Fish and fisheries in western China. FAO Fish.
Tech. Pap. 385, 237-278.
Weatherley, A. H.; Gill, H. S., 1987: The biology offish growth. Academic press,
London, UK.
Whiteman, K. W.; Travnichek, V. H.; Wildhaber, M. L.; Delonay, A.; Papoulias, D.;
Tillett, D., 2004: Age estimafion for shovelnose sturgeon: A cautionary note
based on annulus formation in pectoral fin rays. N. Am. J. Fish. Manage. 24,
731-734.
Williams, E. H.; Shertzer, K. W., 2003: Implications of hfe-history invariants for
biological reference points used in fishery management. Can. J. Fish. Aquat.
Sci. 60(6), 710-720.
Yao, J. L.; Chen, Y. F.; Chen, F.; He, D. H., 2009: Age and growth of an endemic
Tibetan fish, Schizothorax o 'connori, in the Yarlung Tsangpo river. J. Freshw.
Ecol. 24(2), 343-345.
Yilmaz, S.; Polat, N., 2002: Age determination of Shad (Alosa pontica Eichwald,
1838) Inhabiting the Black Sea. Turk. J. Zool. 26, 393 - 398.
52
Zhan, B. Y., 1995: Fisheries Assessment. Agriculture Press of China, Beijing,18-51
(in Chinese).
Zhang, J.; Takita, T., 2007: Age and growth of Ilisha elongate (Teleostei:
Pristigasteridae) in Ariake sound, Japan: comparison among populations in
western north Pacific ocean. Fish. Sci. 73, 971-978.
Zhao, L. H.; Wang, S. H.; Zhao, T. Q., 1975: The age and growth of Gymnocypris
przewalskii przewalskii (Kessler). In: Institute of Biology, Qinghai Province
(Eds) The Fish Fauna of Qinghai Lake Region and Biology of Gymnocypris
przewalskii przewalskii (Kessler). Science Press, Beijing, 37-45 (in Chinese).
Zoubi, A.; Lamrini, A.; Berraho, A.; Hamouda, A., 2010: A comparison of the
precision of red mullet (Mullidae) age detennination by the use of different
bony structures at the strait of Gibraltar region. Rapp. Comm. Int. Mer Medit.
39, 707.
b^'4^Cg
' ^ ^ <
53
•J^' ^
J. Appl. Ichthyol. 29 (2013), 283-284 © 2012 Blackwell Veriag GmbH ISSN0175-S659
Received: February 5, 2012 Accepted: April 4, 2012
doi: 10.1111/j.1439-0426.2012.02061 ,x
Technical contribution
Length-weight and length-length relationships for five fish species from Kashmir VaUey By Mohammad Afzal Khan and Sabah
Department of Zoology. Section of Fishery Science and Aquaculture, Aligarh Muslim University, Aligarh, India
Summary Length-weight (LWR) and length-length relationships (LLR) are presented for two species of Schizopyge [S. curvi-frons (Meckel, 1838) and S. niger (Heckel, 1838)] and three species of Schizothorax [S. esocinus (Heckel, 1838), S. labiatus (McCleUand, 1842) and S. plagiostomus (Heckel, 1838)] from the Kashmir valley o( India. A total of TiS specimens were sampled and measured from June to December 2011. No information regarding LWRs and LLRs of these species was available in FishBase.
IntrodnctioD
Knowledge on length-weight (LWR) and length-length relationships (LLR) is useful in fish slock and popxUation assessments (Ricker, 1968; Kara and Bayhan, 2008). Size conversions (e.g. calculated TL from Sh) are needed for comparison of species. The present study was undertaken with the objective to estimate the length-weight and length-length relationships for five fish species collected from the Jhelum River and Dal Lake of the Kashmir valley in India.
Materials and mediods
Data on length and weight of S. curvifrons, S. esocinus, S. labiatm, and S. plagiostomus from the Jhelum River and S. niger from Dal lake (Kashmir valley, India) were collected from June to December 2011. Total length (TL), standard
length (SL) and fork length (FL) to the nearest O.i cm and weight (W) to the nearest 0.1 g were recorded for each individual. Identification of fishes was done foffowing Day (1878) and Knllander et al. (1999).
Length-weights were determined by logarithmic transformation of the Unear regression equation; log W = log a + b log SL, where W is the weight of the fish (g), SL is the standard length (cm), a is the intercept and b the slope of the regression curve (Ruiz-Campos et al., 2010). The degree of association between the variables was computed by the determination coeffcient, r^ (Golzarianpour et al., 2011). LLRs viz. TL vs SL, SL vs FL and FL vs TL were calculated by linear regressions (Hossain, 2010). All statistical analyses were done using SPSS version 16.0 (IBM Corporation, Chicago, IL).
Results and discussion
Length-weight statistics for two species of Schizopyge and three species of Schizothorax are presented in Table I. The negative allometric growth for three species imphes that the allocation of energy is more toward axial growth rather than to isometric growth. This may be due to a number of biological factors that are known to influence LWR (Gonzalez-Acosta et a/., 2004; Ruiz-Campos et a/., 2006); however, these factors were not accounted for in the present study. Estimated parameters of the LLRs are given in Table 2. As expected, the TL was roughly 10% longer than SL.
Table 1 Estimated parameters of LWR for two species of Schizopyge and three species of Schizothorax: in bold, new maximum length records not found to date in electronic data bank FisbBase.oig
Length (cm)
range
Length Scientific name Study N Min Max a 95% CL b 95% CL ,2 type
Schizopyge Present study 136 14.5 37.0 0.039* 0.028-0.056 2.69 2.582-2.798 0.992 SL curvifrons
Schizopyge niger Present study 173 14.7 32.6 0.049* 0.026-0.093 2.66 2.453-2.864 0.982 SL Schizothorax Present study 163 10.3 50.8 O.OU* 0.008-0.016 3.08 2.98-3.17 0.995 SL esocinus Bhagat and Sunder,
1984; 187 U.O 55.6 -~5.10 — 3.02 - 0.956 TL
Bhat et al., 2010 70 5.7 42.0 -^5.16 - 3.00 - - TL Schizothorax Present study 142 17.6 38.7 0.044* 0.019-0.993 2.64 2.39-2.89 0.972 SL
labiatus Bhatet al., 2010; 40 9.2 25.5 -5.25 - 3.09 - - TL Schizothorax Present study 121 18.5 41.5 0.022' 0.011-0.042 2.86 2.66-3.05 0.982 SL plagiostomus Bhatet al., 2010 136 9.6 52.0 -4.96 - 2.95 - - TL
N, total number of samples; a, intercept; b, slope; CL, Coafsdeace hmits; r^. Coefficient of determination. *Anti-log a.
IJ.S. Copyright Qearance Cealre Code Statement. 0 n 5 - 8 6 5 9 / 2 O l 3 / 2 9 O l - 2 8 3 $ 1 5 . 0 0 / 0
284 M. Afzal Khan and Sabah
Table 2 LLRs between total length (TL), fork length (FL) and standard length (SL) for two species of Schizopyge andthree species of Schizo-thorax
Species N Equation a b r"
Schizopyge 136 TL=a + bSL 2.39 1.09 0.998 cunrifrons FL=a + bSL 1.68 1.04 0.999
TL=a + bPL 0.63 1.05 0.999 Schizopyge 173 TL=a + bSL 2.23 1.11 0.993
niger FL=a + bSL 1.38 1.06 0.997 TL=a + bFL 0.75 1.05 0.997
Schizothorax 163 TL=a + bSL 0.93 1.15 0.998 esocinus FL=a + bSL 0.45 1.08 0.997
TL=a + bFL 0.48 1.06 0.998 Schizothorax 142 TL=a + bSL 1.53 1.11 0.998
labiatus FL=a + bSL 0.82 1.061 0.996 TL=a + bFL 0.80 1.05 0.997
Schizothorax 121 TL=a + bSL 2.20 1.08 0.996 plagiostomus FL=a + bSL 0.64 1.06 0.998
TL=a + bFL 1.53 1.02 0.999
N, total number of samples; TL, total length; FL, fork length; SL, standard length; r^, coefficient of determination.
No information on LWRs and LLRs of the selected fish species was available in FishBase (Froese and Pauly, 2011). However, Bhat et al. (2010) reported values of b for S. esocinus (3.0034), S. labiatus (3.0997) and S. plagiostomus (2.9467) in the Lidder River of Kashmir. Bhagat and Stmder (1984) reported the value of b parameter for S. esocinus as 3.0034. The value of b reported by Bhat et al. (2010) for S. labiatus is different from the present study, which is possibly due to several factors such as the habitat, number of specimens examined and length ranges and length types used. As per the data available in FishBase (2011), reported herein is the new maicimnm standard length for S. esocinus (50.8 cm), S. labiatus (38.7 cm), S. plagiostomus (41.5 cm) and Schizopyge niger (32.6 cm). However, Bhat et al. (2010) reported a higher maximum length (TL = 52 cm) for S. plagiostomus. As suggested by Petrakis and Stergiou (1995), use of LWRs should be strictly limited to the observed length ranges applied in the estimation of the linear regression parameters.
References
Bhagat, M. J.; Sunder, S., 1984: Some biological aspects of Schizo-thoraicthys esocinus Meckel, from Kashmir waters with a note on its utility in culture. J. Inland Fish. Soc. India 16, 42-47.
Bhat, F. A.; Yousuf, A. R.; Balkhi, M. H.; Mahdi, M. D.; Shah, F. A., 2010: Length-weight relationship and morphometric characteristics of Schizothorax spp. in the River Lidder of Kashmir. Indian J. Fish. 57, 73-76.
Day, F., 1878: The Fishes of India; being a natural history of the fishes known to inhabit the seas and fresh waters of India. Burma and Ceylon, Vol. 1. pubUshed by Bernard QuaUtch, 15 Piccadilly London, London: pp. 529-533.
Froese, R.; Pauly, D. (Eds), 2011; FishBase. World Wide Web electronic publication. Available at: http://www.fishbase.org. Version 3, (accessed on 20 March 2012).
Golzarianpour, K.; Abdoli, A.; Kiabi, B. H., 2011: Length-weight relationships for nine nemacheilian loaches (Teleostei: Nemache-iUdae) from Iran. J. Appl. Ichthyol. 27, 1411-1412.
Gonzalez-Acosta, A. F.; De La Cruz-Aguero, G.; De La Cruz-Agu-ero, J., 2004: Length-weight relationships of fish species caught in a mangrove swamp in the Gulf of Cahfomia (Mexico). J. Appl. Ichthyol. 20, 154-155.
Hossain, M. Y. 2010: Length-weight, length-length relationships and condition factors of three schibid catfishes from the Padma River, northwestern Bangladesh. Asian Fish. Sci. 23, 329-339
Kara, A.; Bayhan, B., 2008: Length-weight and length-length relationships of the bogue Boops hoops (Linneaus, 1758) in Izmir Bay (Aegean Sea of Turkey). Belg. J. Zool. 138, 154-157.
Kullander, S. O.; Fang, F.; Delling, B.; Ahlander, E, 1999: The fishes of the Kashmir Valley. In: Nyman, L. (Ed). River Jhelum. Kashmir Valley, impacts on the aquatic environment. Swedmar. Goteborg. pp. 99-163.
Petrakis, G.; Stergiou, K. I., 1995: Weight-length relationships for . 3 fish species in Greek waters. Fish. Res. 21, 465-469.
Ricker, W. E. 1968: Methods for Assessment of Fish Production in Freshwaters. Blackwell Scientific Pubhcations, Oxford, pp. 313.
Ruiz-Campos, G.; Gonzalez-Acosta, A. F.; De La Cruz-Aguero, J.. 2006: Length weight and length-length relationships for some continental fishes of northwestern Baja California, Mexico. J. Appl. Ichthyol. 22, 314-315.
Ruiz-Campos, G.; Ramirez-Valdez, A.; Gonzalez-Guzman, S.; Gonzalez-Acosta, A. F.; Acosta Zamorano, D., 2010: Length-weight and Length-length relationships for nine rocky tidal pool fishes along the Pacific coast of the Baja California Peninsula, Mexico. J. Appl. Ichthyol. 26, 118-119.
Author's address: Mohammad Afzal Khan, Section of Fishery Science & Aquaculture, Department of Zoolog>', Ali-garh Muslim University, Aligarh - 202 002, India. E-mail: [email protected]
Acknowledgements
The authors are thankful to the Chainnan, Department of Zoology, Aligarh Muslim University, Aligarh, India for providing the necessary facilities for the study.
1 Using five calcified structures to describe age and growth of three fish species from
2 Kashmir Valley
4 Sabah and Mohammad Afzal Khan*
5 * Corresponding author.
6 E-mail address: khanmatzal(@yahoo.com. (M. Afzal Khan)
7 Abstract
8 Schizothoracinae are the most important food fish of Kashmir valley. Age determination
9 methods and growth rates are poorly described for these species. The present study estimated
10 the precision of age estimates between different structures (scales, opercular bones, otoliths.
11 vertebrae, cleithra) by calculating the percentage of agreement (PA), average percentage of
12 error (APE), and coefficient of variation (CV). Maximum ages of 6 years were observed for
13 Schizopyge curvifrom and Schizopyge niger and 5 years for Schizolhorax esociniis.
14 respectively. In S. curvifrons and S. niger, percent agreement between-readers was highest for
15 otoliths i.e. 95.4% and 94.6% respectively and in S. esocimis. percent agreement was highest
16 for vertebrae (96.0%). The von Bertalanffy growth parameters were as follows: Loo = 49.8
17 cm, K = 0.263 year"' and to =-0.34 year for S. curvifrons (based upon otoliths). Loo= 44.8 cm.
18 K = 0.255 year"' and to = -1.42 year for S. niger (based upon otoliths), and Loo = 66.6 cm. K =
19 0.278 year" and to = -0.34 year for S. esocinus (based upon vertebrae).
20 Keywords
21 Precision. Growth. Schizopyge curvifrons. Schizopyge niger, Schizolhorax esocinus.
22 Communicated for publication in the journal, Ichthyological research (Springer
23 publication).