Fishery v4n1p54 Fa PDF

Journal of Survey in Fisheries Sciences 4(1) 54-93 2017
A Review of Fish Taxonomy Conventions and Species

Identification Techniques
Keat-Chuan Ng C.1; Aun-Chuan Ooi P.1; Wong W.L.1; Khoo G.1*
Received: December 2016 Accepted: March 2017
Abstract
Taxonomists, ecologists, geneticists or researchers from other biological fields who
wish to adopt fish as a constituent of their studies often become discouraged when they
find that ichthyology is a complex subject. In fish-based studies, the failure to recognize
fishes as distinct biological units can lead to wrong diagnosis. Hence, this review paper
attempts to clarify and discuss the latest schools of thought pertaining to fish taxonomy
and the techniques for species identification. It is hoped that the contents and
illustrations in this paper will assist researchers in laying a good foundation to inform
their studies.
Keywords: Fish, Morphology, Molecular, Taxonomy, Species identification
1-Faculty of Science, Universiti Tunku Abdul Rahman, Jalan Universiti, Bandar Barat,
31900 Kampar, Perak, Malaysia.
*Corresponding author's Email: gideonkhoo@utar.edu.my
55 Keat-Chuan Ng et al., A Review of Fish Taxonomy Conventions and Species Identification …
Introduction interbreed in natural conditions. In the

The term “fish” is usually a convenient “evolutionary species” concept, a species
description for a group of poikilothermic is a representative of a lineage having its
(cold-blooded) aquatic vertebrates under own evolutionary affinity and historical
the Chordata phylum that breathe with destiny. As for the “phylogenetic species”
gills (Nelson, 2006). Scientifically, the concept, species is viewed as a
collective term of “fish” primarily refers monophyletic set of organism with
to Agnatha (jawless fishes), common ancestors. In practice, each of
Chondrichthyes (sharks and rays), these major concepts is prone to some
Sarcopterygii (lobe-finned fishes), and level of subjectivity. Regardless of
Actinopterygii (ray-finned fishes). concept, wildlife scientists, especially
Actinopterygians, the bony or ray-finned ichthyologists, typically identify and
fishes, are without a doubt the majority of name fishes by either by their consistency
fishes found in freshwaters. in morphological, and more recently,
Actinopterygians have lepidotrichia which molecular characteristics.
are characterized by fins of membranous While most researchers are concerned
webs held together by bony spines, or with fishes as a food source and their
rays. This niche character differentiates work involves enriching the body of
Actinopterygians from Sarcopterygians aquaculture knowledge, there are some
which possess lepidotrichia that are who are interested in their diversity,
fleshy. distribution patterns, ecology and
Although in early 20th century, the functional physiology. Recently, there has
ichthyologist Regan (1910) defined a fish also been an overwhelming interest in the
species as a product of interrelated molecular constitution of fishes (Wong et
communities with common morphological al., 2011; Pereira et al., 2013; Rakshit et
features (today this is termed as al., 2015 Quraishia et al., 2015) and their
“morphospecies”), it should be noted that function as biological indicators to
the species classification concept differs monitor waterbody pollution (Fonge et
among scientists. While authorities such al., 2011; Khodadoust et al., 2013;
as Nelson (2006) and Mayr (1942) accept Authman et al., 2015). Correspondingly,
the “biological species” concept, others the interest in fish has expanded
like Simpson (1951) promotes the exponentially, and the ichthyology
“evolutionary species” concept. Then discipline is often sought to contribute too
there is Cracraft (1983) who prefers to many other fields of studies (Padilla and
adopt the “phylogenetic” or “cladistic” Williams, 2004; Lauder et al., 2007; Feist
species concept. and Longshaw, 2008; Rudkowska et al.,
The oldest and most widely practised 2010). Generally, species is the basic unit
“biological species” concept postulates in these studies and sound taxonomy is a
that species are part of a group prerequisite to prevent confusion and
composition that breed or can potentially misinterpretation.
Journal of Survey in Fisheries Sciences 4(1) 2017 56
However, researchers who wish to rules, and yet set out to conduct fish-
adopt fish as a major or minor component based studies and publish papers. Since
of their studies will be discouraged when they lack field experiences, many are
they discover that ichthyology is not an unaware of distinctive fish characteristics
easy subject, more so fish taxonomy. such as phenotypic polymorphisms,
Also, it is a common knowledge that there sexual dimorphism and behavioural
is already an acute shortage of fish divergence due to the regional speciation
taxonomists and finding a fish taxonomist process (Waugh, 2007). Failing to
to assist in a study is often difficult. Such recognize such exceptions can lead to
is the same case with many other taxa. wrong species diagnosis. Some
Since fish diversity can be high, researchers restrict themselves to
especially in tropical countries, a laboratories, ornamental fish and
taxonomist is typically overwhelmed with aquaculture farms, and their specimens
constant scientific name revisions, field may look very different from the wild
collection expeditions and managing a type which the scientific name was
museum. Moreover, taxonomy cannot be derived from. Ideally, it is advisable to
commercialized and it sees very little treat a selective bred variety raised in
funding in many countries. Thus, the artificial conditions differently.
discipline rarely attracts students or can Otherwise, publication with nomenclature
sustain career taxonomists in the errors will be perpetuated through citation
universities. Such a problem has caused and cause a chaotic situation.
taxonomic errors in published papers and In lieu of concerns highlighted, this
the condition is now widely known as the review attempts to provide a concise
“taxonomy impediment” (Wheeler et al., introduction to this complex but important
2004; de Carvalho et al., 2007). It is often field. To understand fish as a biological
said that the discipline of taxonomy unit, the readers should also have some
would be extinct earlier than most background in zoology. Our objective is
endangered species. not to polarize techniques for fish
On hindsight, the advancement of identification because we are convinced
molecular, computerized and statistical that each technique has its merits. Their
techniques have led many to believe that shortcomings will, however, be discussed
“species” can be easily characterized by to assist researchers in making sound
nucleotide sequences, software and judgments and decisions.
mathematical calculations, and the This paper reviews and discusses
knowledge in taxonomy is no longer taxonomic concerns of freshwater species
needed. Ebach and Holdrege (2005) warn that belong to the Actinopterygii class,
us that there are a growing number of and we assume that readers have some
researchers who have never wet their feet familiarity with ray-finned fish anatomy
in the rivers to observe species or build and the common species. Given that a
competency in applying nomenclature picture is worth a thousand words, we
believe that taxonomy can be easier to clustering species that looked similar
master if the reader is provided with because they believed that these species
detailed scientific illustrations and colour share a common biological lineage. In the
images. This review is by no means mid-18th century, the evolution hypothesis
comprehensive but we hope it will set a was a new and strange concept.
good foundation for those who wish to Eventually this was proven to be factual
make a competent start in species-level in the advent of modern molecular and
research. genetic assessment technologies. In
general, today, there is a consensus
Why are there do many species? among scientists that all species on Earth
Teleost species form the largest category are interlinked with a hierarchical and
among vertebrate animals and their evolutionary tree with millions of
numbers have reached more than 32,500 branches. Each branch itself is a
valid marine and freshwater species under representation of the natural history of a
515 families (Nelson, 2006). In the global particular species and its pedigree. But
context, there are roughly 11,952 of why did the branches have to extend and
freshwater species (Helfman et al., 2009). multiply in the first place? What causes
Ricklefs (2004) suggests that competition the divergence?
and mutualism among species have an When gene flow among populations is
effect on species abundance while Wright interrupted by natural (e.g., geographic
et al. (2003) speculate that species barriers created by earthquakes) or
diversity is the result of productivity from artificial means (e.g., man-made barriers
available “energy” in a particular region. in aquaculture farms and dams), two types
If a region lacks energy (e.g., low food of speciation will occur, namely allopatric
availability in the desert), species richness speciation and sympatric speciation
is typically low. However, in the case of (Butlin et al., 2016) (Fig. 2). Allopatric
fish, MacArthur’s (1969) hypothesis speciation occurs when a population is
seems to be the most probable as he separated by a barrier and such isolation
suggests that species diversity is prevents the two or more sub-populations
accelerated if there are more areas to host from mating. Given time, the lack of gene
suitable habitats. The hypothesis flow among the sub-populations will
corresponds with the reality today as large cause biological incompatibility and
freshwater regions seem to demonstrate divergence would be triggered (Mayr,
more genera and species, for example the 1959; Turelli et al., 2001; Singh 2012).
Neotropical region (705 genera, 4,035
species) compared to the Australian
region (94 genera, 261 species) (Leveque
et al., 2008) (Fig. 1).
Early taxonomists, namely Linnaeus
and Darwin, had started cataloguing and
Figure 1: Major freshwater fish Eco regions encompassing 1,054 rivers as classified by the Fish-
SPRICH database (Source: Brosse et al., 2012).
Figure 2: Modes of speciation.
In sympatric speciation, biologists branch would still maintain a certain

highlight that ecological shift and degree of morphological similarity (Butlin
resource competition are the key drivers et al., 2016).
(Mayr, 1947; Dieckmann and Doebeli,
1999; Bolnick and Fitzpatrick, 2007; Taxonomy and systematics
Mallet et al., 2009). They reasoned that The word taxonomy originated for Greek
new species may arise within a population word taxis, meaning arrangement, and
from biological reproductive barriers nomos, meaning law. The science of
between mutants that are better adapted biological taxonomy is responsible for
and parent populations. Nonetheless, discovering, describing, classifying,
despite further speciation in both space naming and treating each species as the
and time, and regardless of allopatric or basic unit. Species are given names in
sympatric speciation, species within a accordance to the protocol set by
Linnaeus’ binomial nomenclature system studies require considerable identification

(Enghoff, 2009). Systematics on the other skills, experiences and familiarity with
hand is the science of distinguishing local species. Nonetheless, this may not
orderliness and classification of a taxon be as complex as it sounds because each
into hierarchical series to emphasize their species is normally distinctive in
interrelationships (Mayr, 1942; Guerra- appearance and has a certain “look” (Fig.
Gracia et al., 2008). The word 3).
systematics stems from Greek word At a higher level and broader
systema. Nelson (2006) tells us that a perspective, systematists are experts who
systematist seeks the broadest outlook to examine historical discrepancies,
resolve family, relative, order or grouping ambiguities, errors and variant names of
orderliness. Taxonomy and systematics species, genus and family. For example,
are not entirely different schools of there is the problem with regards to
thought, but rather overlapping fields whether Eleotridae or Eleotrididae should
(Wilson, 1985; Lincoln et al., 1998). be used for the members of sleeper fishes
Kapoor (1998) and Wägele (2005) also (Robins, 1991). Thankfully, species
highlight that the term “systematics” is classification and naming consistency is
often used synonymously with taxonomy. slowly being resolved and currently
It must be clarified that this review adopts governed by the International
Nelson’s (2006) argument that biological Commission on Zoological Nomenclature
classification is based on systematic (ICZN). However, fish taxonomic
studies and taxonomy is part of problems cannot be resolved quickly as
systematics. desired because ICZN also has to address
A proper taxonomy is a first-hand and issues affecting other taxa. For the fish
exhaustive undertaking. Whether one taxon, appreciatively, there are dedicated
adopts the “biological species” concept ichthyologists who take it upon
(Mayr, 1942; Nelson, 2006), themselves to diligently keep track of the
“evolutionary species” concept (Simpson, latest development and progressively
1951) or the “phylogenetic species” publish the most updated information.
concept (Cracraft, 1983) as explained Every species belongs to a genus
earlier, all identification processes start (plural: genera), every genus to a family,
which examining morphotypes at the every family to an order, every order to a
earliest stage of discovery. Specimens are class, every class to a phylum (plural:
physically scrutinized from small phyla) and finally all phyla are placed
microscopic configuration of scales to under an overarching kingdom (Fig. 4).
large membrane patterns of the caudal fin. Each phylum is regarded as a
Each physical variance, no matter how representation of a large grouping of
small, on a fish body is useful species that shares a common ancestor in
information. Naturally, all fish-based evolution.
Figure 3: Species from each family have a similar morphological profile to provide identification
clues (Source: Adapted from Rainboth, 1996).
Figure 4: Taxonomic ranking and naming convention.
Ray-finned fish which is reviewed in this ranking. According to ICZN regulation,

paper belongs to the Actinopterygii class the family-group name must always end
under the Chordata phylum in the with the “idae” suffix (e.g., Cyprinidae)
Animalia kingdom. Such is the ranking and the subfamily-group name must end
and principle of taxonomy practice used with the “inae” suffix (e.g., Cyprininae).
to organize all biological units. At time of At a higher level, the order-group name
writing, some experts have arrived at should end with the “iformes” suffix (e.g.,
some agreement and the “Family-group Cypriniformes) and the suborder-group
Names of Recent Fishes” published by name must end with the “oidea” suffix
Van der Laan et al. (2014) seems to be the (e.g., Cyprinioidea). The family names are
most updated for family naming. always capitalized but never italicized.
In fish taxonomy, there are established
conventions for expressing taxonomic
How species are named and why its scientific name Trichogaster leeri is a
In Latin, species means “a kind, unique name and there is no chance that
appearance and quality”. Depending on the name may be mistaken with other
locality, people have various vernacular species. Ideally, a scientific name may
and common names for a fish species. For even tell something about the species' key
example, the common names Pearl characteristics (Fig. 5), its habit,
Gourami, Diamond Gourami and Mosaic discoverer and perhaps the location where
Gourami all refer to the same species. it was first found.
Therefore, the use of common names can
be confusing and misleading. However,
Figure 5: The species name Leptobarbus rubripinna is derived from Greek word “leptós” which
means thin or slender, and the Latin word “barbus” meaning barbel. Rubripinna
originates from the Latin words “ruber” and “pinna” which mean red and fin,
respectively.
Another interesting example is the brittani) was named after a prominent

fighting fish species, Betta persephone, ichthyologist Maurice Kottelat, and a
which is named after Persephone the species that is named Anguilla borneensis
daughter of Zeus in Greek mythology. tells us that it was first described from
Also known as the princess of darkness specimens found in Borneo island. Those
because she is said to rule the underworld, familiar with Latin or Greek, the
Persephone was the perfect epithet for B. traditional language used in scientific
persephone which typically inhabits the names, would also be able to tell that the
black water in peat-swamp habitats of catfish Clarias leiacanthus should have
Southeast Asia. Another fighting fish pectoral spines with smooth anterior edge;
species the Betta gladiator need no in Greek, “leios” means smooth and
further explanation as to why the epithet “akathos” means thorn. The Bihunichthys
was given. monopteroides was named after a popular
The Kottelatia genus (with only one food in Southeast Asia; “bihun” is a local
recognized cyprinid species, Kottelatia name for rice noodle and “ichtys” means
fish in Greek. Sure enough, it is a very hosts an isolated Channa gachua

small and thin spineless eel that resembles population …). This is to ensure that the
rice noodles. From examples mentioned, a species name is outstanding and can be
scientific name can be communicative and easily singled out during reading. When
gives species stable and universal handwritten, species name should be
designations for easy retrieval. underlined for the same reason. The first
The scientific naming that we practice part may be used alone but the second
today would not be possible without the part is never used by itself.
foundation laid by Carl Linnaeus (1707- A species name must be written in full
1778). Often confused by the many the first time is it expressed in a
dialectal names during his specimen manuscript (e.g., Cyprinus hyperdorsalis)
collection work in various countries, and thereafter the first part, or genus
Linnaeus was convinced that species name, can be abbreviated with initial
names must be standardized. In 1735, he capital letter (e.g., C. hyperdorsalis) on
published a small pamphlet titled Systema the condition that there is no
Naturae (The System of Nature) to misconstruing with other genera (i.e.,
introduce his new system of giving and bearing in mind “C. can also mean
organizing species names. Linnaeus also Channa or Clarias if these genera appear
decided that species names should be on the same paper). In cases where an
given in Latin or Greek in two parts. Thus abbreviated initial capital letter may cause
the binomial nomenclature system was confusion, two letters may be used (e.g.,
conceived. Although the Systema Ch. for Channa or Cl. for Clarias). There
Naturae was meant for naming plants, it is no absolute rule and the objective is to
soon gained popularity due to its avoid misinterpretation.
practicality and it was quickly adopted by Additionally, “sp.” (Singular) or “spp.”
taxonomists of various taxa. (Plural) may be used to represent
The rule of binomial nomenclature “species” to indicate partially identified
dictates that the first part of species name species with the genus known. For
comprises of its genus (noun) and it is example, when a specimen is recorded as
always capitalized (e.g., Cyprinus). The Cyprinus sp. it denotes that a specimen of
second part is used to describe the the Cyprinus genus cannot be identified to
species’ attribute or epithet (adjective) species level, possibly due to it being a
and it is never capitalized (e.g., Cyprinus small juvenile which makes positive
hyperdorsalis; “hyper” meaning “over” in identification difficult. For species with
Greek and “dorsalis” meaning the back problematic identification, it may also be
part of the body in Latin). The first and recorded with the term “cf.” added
second part is always italicized. However, between the scientific names. It is simply
if the neighbouring texts are italicized, a short term for “confer” or “compare
then the first and second name would be with”. For example, when a species is
non-italicized (e.g., in mesohabitat that referred as Rasbora cf. elegans, it implies
the species it most likely belongs to but when submitting a manuscript for peer
the designation is still marred by review, it would be wise to countercheck
unresolved taxonomic issues or more the journal's submission instructions on
work is needed to be completely sure. how to quote the target species.
Alternatively, the term “aff.” is sometimes
added between the scientific name when a Species identification at the
specimen cannot be matched to any morphological level
species known to science, or it is a new Since humans learnt how to hunt fish,
species that is yet to be named. The term species were identified based on some
is a short form of “affinis” in Latin which simple anatomical features. Observation
means “akin to”. The insertion of “aff.” is of specimen anatomy and differentiating
to associate the possibility of a new fish species based on their morphological
species with the closest species (e.g., features is the most practical, rapid and
Rasbora aff. elegans). low cost method. Besides experienced
It would be advisable to describe the local fisherman and fish mongers, people
genus, species, author and year in full who live by the river or wetland would
when a species name is expressed as part learn to identify fishes at a young age.
of a manuscript title, or the first time it is This is due to knowledge and memory
written in the manuscript. The author(s)’ acquired from long-term observation or
name(s) who first coined the binomial through oral tradition maintained by
name and year of publication should be elders. Such traditional knowledge has
included (e.g., Cyprinus hyperdorsalis been interweaved into modern
Nguyen, 1991) because it is a good ichthyology by many researchers
practice to ensure the manuscript author is (Calamia, 1999; Drew, 2005; Stacey et
explicitly referring to a species described al., 2008; MacLean et al., 2009; Ferreira
by a particular taxonomist in a particular et al., 2014), and the term for it is
year. The author’s name is typically “traditional ecological knowledge” (TEK)
enclosed in parentheses when the genus is (Berkes et al., 2000). For those keen in
no longer the original one used. taxonomy, however, there is a need to
Alternatively, the author’s name is adopt a more precise and sound approach.
enclosed within parenthesis when a Fish identification is the most
species has been transferred from the important component in any fish-based
original genus in which it was first study and acquiring reasonable
described; e.g. Cyprinus melanes (Mai, competency always begins with the study
1978). This puts the author and the reader of fish anatomy. Fundamentally, fish
in a safer position because literature that species have to “go with the flow”. It is
adopts fish as a subject can be flawed by apparent that the anatomy of fish is
synonyms and obsolete species’ names. It shaped by the physical properties of
should be noted that the author citation is water, the essential liquefied medium in
treated differently in various taxa. Also, which the fish adopts as habitat. Water is
dense but holds relatively small amounts and it forms the main body that lies
of oxygen which affects fish respiratory between the head and caudal (Fig. 6). In
function. Water also absorbs light in most species, the trunk is narrowed down
higher intensity than air which affects fish at an area called the caudal peduncle
visual capacity. Water can flow fast or not where it is connected to the caudal fin,
at all, and this affects how fishes which is a prominent feature on the body
manoeuvre in water. Such a complex and of a fish.
fluid environment calls for special Body flexure attained by contractions
adaptations to live in and fishes have of the myomeres and thrust from fins is
evolved precisely to fit in. responsible for fish propulsion. Fin shapes
In general, most biological adaptations and sizes vary tremendously and they are
in fish occur in the body, mouth, fins, skin used for stabilizing, reversing, stopping,
coloration and reproductive traits. The descending, ascending and manoeuvring.
body of a bony ray-finned fish comprises In morphometric (Figs. 7 and 8) and
3 sections; the head, trunk and caudal or meristic identification of fishes, the
tail. The head is a region from the mouth positions of fins, numbers and types of
tip to the posterior edge of the gill cover. ray or spine composition are useful (Figs.
The trunk contains the abdominal cavity 9, 10 and 11).
Figure 6: An example of key morphological features of a species from the Cyprinidae family
(Source: Adapted from Rainboth, 1996).
Figure 7: Common morphometric data collected for fish identification (Source: Adapted from
Rainboth, 1996).
Figure 8: Common measurements between key points to construct patterns that quantify
morphometric variance between species (Source: Adapted from Strauss and Bookstein,
1982).
Figure 9: A close-up view of fin rays which epitomises fish under the Actinopterygii class.
Figure 10: An example of fin positions and ray structures of a fish with a contiguous dorsal fin.
Figure 11: A specimen from the Gastromyzon genus with specialized suctorial pectoral fins and
fused pelvic fins for adhering to substratum rocks in riffles.
There are two fundamental types of fin “ribbon-fins” (Curet et al., 2011), and
rays, namely the true spinous rays and what makes them so special is that the
soft rays that form the framework for fins. entire stretch of the anal fin is actuated by
Spinous rays are stiff and typically muscles along the body length. In eels, a
unbranched, and they are located in a ribbon-fin may be present but the caudal
single fin’s anterior part while soft rays fin is almost absent and they rely mostly
consist of longitudinal supports and are on snake-like rectilinear locomotion for
typically branched. Even so, there are swimming.
exceptions. Some species like the Most bony fishes have homocercal
Silurichthys indragiriensis and Wallago caudal fins that can be forked, rounded,
attu propel themselves forward or truncated, and other symmetrical and non-
backward by wavelike flexure of long symmetrical forms (Fig. 12). Generally,
anal fins. Ichthyologists call these fins fishes with tapered body and fins are
proficient in high-speed propulsion in fast As expected, scale counts are crucial for
flowing waters while fish with rounded fish identification (Fig. 14). Some genera
body and fins are associated with low- such as Clarias, Mystus and Ompok are
speed movement in slow waters. without scales, but they are no less
Mouth form and position also vary vulnerable. The skin of scaled and scale-
greatly among fishes (Fig. 13). In most less fishes also secretes mucus to function
fishes, the mouth is located terminally at as a sealant or protection against
anterior tip of the head (e.g., Pristolepis infection, and to reduce hydraulic friction
spp. and Oreochromis spp.), but in some while swimming.
others it may be inferior (beneath the Evidently, some of the most interesting
snout, e.g., Pangio spp.) or superior features of fishes are their pigmentation
(upturned, e.g., Belodontichthys dinema) patterns (Fig. 15) which can be used as
depending on their feeding habits. critical references for taxonomic
Generally, terminal mouths belong to identification purposes. Body colouration
species that prefer to bite or seize their is indeed an interesting subject in the
prey while those with inferior mouths are context of fish as a biological indicator.
bottom feeders. Insectivorous fishes such Like all animals, fishes cannot synthesize
as the archer fish (Toxotes spp.) typically pigments. They have to ingest colourant
have superior mouths and they feed on pigment like carotenoids from the food
insects that fall on the water surface. they consume within their habitats, and
Some freshwater species like the river these can include fruits, insects and
pipefish of the genus Doryichthys have a phytoplankton (Grether, 2000). This can
tubular mouth to suck food from crevices. reflect their foraging tendencies, habitat
Most fishes have protective scales health and especially the presence of
which vary greatly in size, structure, insect diversity as a rich source of
squamation (scale coverage), pigments. Many studies have shown that
arrangement, sequence and colouration. fish mating behaviour is affected by
Scientifically, scales of teleost fishes can pigment availability from feeding habitats
be classified into the placoid, cosmoid, and this, in turn, affects mate choice and
elasmoid and ganoid categories (Sudo et population abundance (Endler, 1980;
al., 2002). Teleost scales possess Basolo, 1990; Houde, 1997; Blount et al.,
outstanding hydrodynamic properties and 2003).
provide a resistant layer to protect fishes
from injury (Bruet et al., 2008). Scaled
skin is also known to play a critical role in
supporting fish locomotion by
synchronizing wave propagation (Long et
al., 1996), and by accumulating potential
energy like tendons (Hebrank and
Hebrank, 1986) for swimming efficiency.
Figure 12: Common caudal fin shapes (Source: Adapted from Rainboth, 1996).
Figure 13: Common mouth types.
Figure 14: Common meristic data collected by taxonomists (Source: Adapted from Rainboth, 1996
and Ambak, 2012).
Figure 15: Common terms used to describe body markings and patterns on a representative
teleost.
The fundamental colour constituent in or fully (Slavík et al., 2016). Individuals

fishes is the dermal chromatophore unit affected by albinism are typically pale,
(pigment cell) which consists of the: 1) light pink, white or yellow (Lechner and
melanophore which contains melanin Ladich, 2010). Their eyes are generally
(browns, blacks, and greys), 2) pink because blood is seen through the
xanthophore and erythrophore which colourless retina (Van Grouw, 2006).
harbour carotenoids (yellows, orange and Conversely, leucism is a condition
red), and 3) iridophore, or sometimes characterized by the reduction or absence
termed as iridocyte, which naturally of most, if not all, of the pigment cell
reflects external light source and provides types. Leucoptic individuals are generally
fishes their iridescence (Hawkes, 1974; pale, white or yellow but they exhibit
Fujii et al., 1989; Metz et al., 2006). normal retinal pigmentation (Quigley and
Chromatophores can naturally contract or Wallace, 2013). Melanism is the exact
expand to induce or change colours to opposite of albinism where excessive
blend into the aquatic environment for development of dark-coloured pigment
camouflaging purposes. In some cases, a melanin occurs and the affected individual
combination of chromatophore units can is typically dark brown or black (Regan,
produce interesting colours. For example, 1961). It is a form of polymorphism that
in the Siamese fighting fish (Betta is widespread in fishes, and it may be
splendens), a wide variety of bodily genetically determined and/or influenced
colours can be exhibited by the by the environment (Price et al., 2006).
permutation of iridophores, For example, melanin polymorphism has
melanophores, erythrophores and been report in Gambusia holbrooki,
xanthophores (Khoo et al., 2012; Khoo et Siphateles bicolor mohavensis and
al., 2014). Amphilophus labiatus (Horth, 2002;
It must be cautioned that identifying Henkanaththegedara and Stockwell, 2011;
species by their body colouration and Sowersby et al., 2014). When an
patterns alone is not entirely robust. In individual exhibits abnormal deep yellow,
certain cases, individuals in a natural orange or reddish pigmentation on the
population may be affected by genetically body, this is attributed to excessive
inherited conditions that cause them to development of the xanthophores and
exhibit albinism, leucistic, melanistic and erythrophores (Khoo et al., 2012).
xanthic abnormalities. Albino individuals Xanthic variety of the Cyprinus carpio
such as Silurus glanis, Astyanax and Carasszus auratus with red, orange
mexicanus, Hydrolagus colliei and and yellow pigmentation were first
Genidens planifrons have been reported reported and domesticated in China and
(Dingerkus et al., 1991; Jeffery, 2006; Japan as ornamental species (Kajishima,
Reum et al., 2008; Leal et al, 2013) and 1977; Balon, 1995). Wild individuals
these individuals are unable to synthesize from species such as Cyprinodon
tyrosine and melatonin hormones partially bifasciatus have also been found to be
xanthic (Carson 2011). On the other hand, those of terrestrial vertebrates. In fact,
an axanthic individual that lacks they are endowed with more sensitive and
xanthophores may be black, white, and/or complex sensory organs to detect
blue (Lewand et al., 2013). For example, vibration and sound. Fishes possess the
Parichy (2006) reported that axanthic cephalic-lateralis system (Nelson, 1972)
individuals have been found in species that comprises a series of neuromast
from the Danio genus. sensory cells that run across the outer
Being fully aquatic, fishes have vastly layer of the head and body (Fig. 16).
different sensory systems compared to
Figure 16: An illustration of the fish cephalic-lateralis system consisting of neuromast sensors that
run across canals on the head and body (Source: Adapted from Nelson, 1972 and Iwata
and Jeon, 1995).
The neuromast contains fine hair cells schooling synchronization (Partridge and
oriented in a manner to detect the Pitcher, 1979). The patterns and
direction of vibration. Vibration signals configuration of cephalic-lateralis system
are then transmitted through special on fish bodies are sometimes used for fish
canals and pits containing endolymphic identification such as those from the
fluid to amplify the signals and thereafter Kryptoglanis, Pseudorasbora and
pass on to the brain. The location of the Caecieleotri genus (Moncey, 2012;
lateralis system in fish can reflect their Kawase and Hosoya, 2015; Walsh and
feeding habits. Bottom feeders tend to Chakrabarty, 2015).
have the lateralis system on top of the In the case of cryptic species when the
body to detect vibration from predators species are too difficult to be
that are lurking above. Those which feed differentiated because they show small
on the water surface normally have the external anatomical and morphological
system along the ventral margin of the deviations between species in the same
body to track the presence of predators genus, ichthyologists have relied on the
below. Studies have also shown that the internal organs to characterise them
lateralis system is responsible for obstacle (McCune, 1981). Where necessary, gill
avoidance, triggering startle response and rakers from the first gill arch on the left
side of the body are also counted for archive and share Fourier spectrum (FFT),
meristic characterization such as species wavelet analysis (WT) and curvature scale
from the Coregonus, Garra and Labeo space analysis (CSS) data of otoliths
genus (Amundsen et al., 2004; Ayoade et (Lombarte et al., 2006). At time of this
al., 2004; Krupp and Budd, 2009). writing, the AFORO archive contains
Correspondingly, in some species, the 4,672 high resolution images of 1,441
pharyngeal teeth found on the fifth gill species from 221 families for reference.
arch are counted from left to right to assist
in meristic identification of species such Species Identification at the Molecular
as those from the Moxostoma, Cycleptus, Level
Danio and Epalzeorhynchos genus Morphological characterization is not
(Eastman 1977; Pasco-Viel et al., 2010). entirely robust. As mentioned earlier, the
All teleost fishes have an inner ear as concept of “species” is still subjected to
an auditory system. Instead of bony debate and not all taxonomists assign
ossicles, fishes have three pairs of meaningful categories to organisms by
calcareous “ear stones”, or scientifically morphotype. In many cases when
termed as otoliths (Greek for “ear certainty cannot be attained, researchers
stones”). The lapillus (“little stone” in use terms like “subspecies”, “strain” or
Latin) functions in maintaining body “variant”. These are highly subjective and
balance and orientation, and the other confusing. In fish especially,
two, namely asteriscus (“little star” in morphological plasticity between
Latin) and sagittal (“arrow” in Latin), individuals of the same species is
peruse acoustic reception. Otoliths are inherent. For example, body colour tones
enclosed in a membranous sac together and polymorphism exhibited by
with sensory hair cells or ciliary bundles. individuals of the same species may vary
Detection of mechanical signals will considerably depending on the diet
occur when there are dynamic interactions regime, habitat and season such as species
between otoliths and cilia (Assis, 2003; from the Betta, Poecilia and Danio genus
Campana, 2005). Since Koken (1884) (Khoo et al., 1997; Price et al., 2008).
reported that fish species can be And this is where molecular analysis has
characterised by having otoliths of become a viable alternative.
different shapes and sizes, otolith Each organism is characterized by a
morphometry is fast becoming a trend to unique set of biological attributes that
identify fishes and some species such as enhance its fitness to survive in a niche
Netuma bilineata, Nuchequula nuchalis, environment. Correspondingly, to adjust
Coilia dussumieri and Garra rufa (Chen to any changes in the environment, an
et al., 2011; Thuy et al., 2015; Salimi et organism is naturally subjected to genetic
al., 2016; Yedier et al., 2016). In 2006, drift, mutation or variation
the AFORO online database was launched (polymorphism) as a mechanism to adapt.
(http://www.cmima.csic.es/aforo/) to Such natural phenomenon provides
markers at molecular level to detect order or sequence in each gene is unique

individual or species uniqueness (Sanger to every species. Since Razin and Rottem
et al., 1977). All molecular methods (1967) have successfully employed
depend on DNA marker or protein protein analysis for characterising
sequence analysis and comparison to microorganism species, modern advances
determine molecular divergence over have adapted the concept of analysing A-
evolutionary time based on the null G-C-T bases in various DNA genetic
hypothesis of molecular evolution, or markers. In fish identification, the
better known as the “neutral theory” common DNA analyses applied are length
(Kimura, 1968). Essentially, all methods polymorphism (RFLP), amplified
assume that individuals from the same fragment length polymorphism (AFLP),
species have specific DNA (or protein) random amplified polymorphic DNA
sequences that vary, to a certain extent, (RAPD), microsatellites or simple
from individuals that belong to other sequence repeat (SSR), single nucleotide
species. Nonetheless, the variance, or polymorphism (SNP) and expressed
“signature” of speciation (Sbordoni, sequence tag (EST) markers (O’Reilly
2010), is dependent on time and space and and Wright, 1995; Khoo et al., 2003;
subjected to biological productivity of Sampaio et al., 2003; Teletchea, 2009;
individuals, dispersal pattern and natural Chauhan and Rajiv, 2010; Khoo et al.,
genetic drift. Therefore, genetic 2011; Kress et al., 2015).
variability also occurs among individuals Fish genome typically contains
of the same species. This means roughly a billion nucleotide pairs (Stepien
establishing a credible locus, or the and Kocher, 1997), and analysing all of
position of a gene (Khoo et al., 2011) them would be too daunting and the
from a phenotype is a prerequisite as results would cause an information
master reference for comparison purposes. overload. Since Herbert et al. (2003)
DNA markers can be classified in two discovered a technique to amplify the
different types, namely type I which are mitochondrial cytochrome c oxidase
markers associated with genes with subunit 1 (COI) gene. There is now a
known function and type II which are consensus to analyse a 648-base pair (bp)
markers are with anonymous genomic region of COI to rapidly identify a fish
segments (Chauhan and Rajiv, 2010). For species (Cawthorn et al., 2012). Such an
example, allozyme markers are classified approach is now known as DNA
as type I protein markers and barcoding and it has gained widespread
microsatellites and other neutral markers acceptance as a fast, cost effective and
are considered type II (Hinsinger et al., standardised technique. So far, the Fish
2015). The DNA in all organisms is a Barcode of Life Initiative has barcoded
composition of four chemical bases – 7,882 fish species, which is only
adenine (A), guanine (G), cystosine (C), approximately 25% of the estimated
and thymine (T) (Avise, 1994). Their
31,220 species present globally (Jinbo et metabarcoding" to refer to their ability to

al., 2011). automatically identify multiple species
It should be stressed that identified from a single bulk sample (Taberlet et al.,
species through DNA barcode is only as 2012). Because NGS technologies can
good as the voucher specimen's DNA extract massive numbers of reads, the
barcode made available and archived in results increase the chances of finding and
platform such as GenBank. Hence, if a annotating matches (Hemmer-Hansen et
DNA barcode is inaccurate or al., 2014). So far, technological platforms
compromised due to issues such as that are able to produce and analyse
carryover DNA contamination, gigabases of DNA sequence include
incomplete genotyping of loci throughout Illumina, Roche, AB SOLiD, 454 GS
the genome, low/wrong quality DNA FLX, Ion Torrent, HeliScope, Starlight
material, human errors, biases and some and PacBio (Rothberg and Leamon, 2008;
other critical risks (Bridge et al., 2003; Pandey et al., 2008; Davey and Blaxter,
Forster, 2003; Moritz and Cicero, 2011; Hemmer-Hansen et al., 2014). They
2004; Smith and Burgoyne, 2004; Ebach differ from each other based on amount of
and Holdrege, 2005; Meyer and Paulay, sequence information generation,
2005; Will et al., 2005), the identification chemistry protocol for sequencing and the
process would be erroneous. Ross et al. length of sequence read (Mehinto et al.,
(2008) proposed that reference sequences 2012).
of at least five or more voucher specimens
from various geographical sites should be Reference Type and Traceability
acquired. Unfortunately, this also means The hallmark of the taxonomy discipline
that DNA analysis is not a one minute is its persistency in collecting, preserving
procedure. To conduct it properly, the and managing specimens as essential
process of acquiring exemplar genotypes physical references for species-level
is tedious and costly because some research. Although ecologists,
species are found in large regions and conservationists, aquaculturists or
various countries. researchers who adopt fish as a part of
In the recent years, DNA sequencing their studies need not master the specifics
techniques have made substantial progress in specimen management, it is useful to
and they are able to analyse 30 to 1,500 briefly understand the ICZN zoological
nucleotides (nt) for hundreds of thousands code and terms as a precaution against
to millions of DNA molecules in a single negligence when tracing the correct
process within a complex or degraded binomial nomenclature and description of
DNA source (Davey and Blaxter, 2011; fish.
Mehinto et al., 2012). These are classified Fundamentally, in taxonomy,
as next-generation sequencing (NGS) a “type” is a specimen of a distinct
technologies (Tillmar et al., 2013) and species which is used as master reference
sometimes termed as "DNA (Krell and Wheeler, 2014). Each species
or subspecies that is scientifically named around the world for the type series (a
by the original author (discoverer) is range of specimens showing variation in
traceable to a name-bearing specimen the species). These comparable specimens
which was first found and kept in a are termed as “paratypes” and they have
particular museum. A new species cannot no name-bearing role.
be described and registered formally In situations when the researcher fails
without depositing a single physical to establish a holotype, the existing two or
whole fish known as a “holotype” in the more specimens collected and deposited
museum (Clemann et al., 2014; Kumar in the museums can be used to describe
and Hassan, 2015). As sexual dimorphism and name a species. These deposited
occurs in many fish species, it is good specimens are known as “syntypes”
practice to deposit an opposite sex although such practice is now rarely used
specimen of the holotype and it is in contemporary taxonomy. A master
assigned as an “allotype” (Jorge et al., reference specimen selected from
2014). Also, polymorphisms may occur in syntypes is termed as “lectotype” and
certain fishes and a holotype is not when a lectotype is finally established, all
expected to be a typical representative other syntypes shall be, by default,
(Hulsey, 2005), although in an ideal case reassigned as “paralectotypes”.
it should be. To mitigate this, Correspondingly, any duplicate specimen
morphotypes that show morphological of the lectotype is called an
variants of a species can be established, “isolectotype”. Finally, specimens that
and molecular analysis is usually carried have been erroneously described, named
out to determine whether the variations or labelled are annotated as “non-types”.
are due to polymorphism or if it is a new All terminologies pertaining to types
species (Simonov, 2008). mentioned in this section are elaborated in
A researcher may also designate a detail by ICZN at the website
duplicate specimen of the holotype and http://www.iczn.org/iczn/index.jsp.
such a specimen is termed as an The responsibility rests on the
“isotype”. When a better specimen is taxonomist to maintain the specimens and
eventually deposited, the holotype is not make them accessible for current and
superseded. According to the ICZN code, future studies (Krell and Wheeler, 2014;
such a specimen which provides better Rocha et al., 2014). To consolidate a large
clarity is known as an “epitype” and it quantity of voucher specimen database in
may be deposited when the holotype is a museum, the taxonomist is also
evidently imprecise. In the case when a expected to occasionally publish
holotype is lost or damaged, a “neotype” monographs that contain summarised
specimen may also be deposited as information of all species in a group to
replacement. While there can only be one update the scientific community (Grinnell,
holotype, taxonomists can continue to 1910; Pyke et al., 2010; Kottelat, 2013).
deposit specimens in any museums
In the advent of technological In principle, a scientific finding must be

advancement, non-destructive methods reproducible. When an author declares
such as high-resolution photography, and publishes the description of a new
computerised tomography (CT) scan and species, a specimen (i.e., haplotype) must
radiography methods that can record the be deposited permanently in the museum
physical characteristics of specimens, the so that the author's claim can be critically
work of a taxonomist has expanded over appraised and reappraised by others
the years. There are already attempts to (Rocha et al., 2014). The specimen can be
digitalize specimens using these refuted or disputed by other ichthyologists
technologies. For example, Berquist et al. to allow for revisions. Similarly, when a
(2012) have progressively scanned researcher conducts a fish inventory
specimens with the magnetic resonance investigation that results in a scientific or
imaging (MRI) technology and created an journal report, it is a good practice
online digital archive called Digital Fish (although not compulsory) to collect
Library (DFL, voucher specimens for future verification
http://www.digitalfishlibrary.org) to share and to promote transparency. This
high-resolution and high-contrast visual approach also provides physical records
data. The Biovisualization Center at the that allow researchers to scrutinize the
University of Washington has also just inter- and intraspecific variation of
initiated a program that applies the CT species collected at different periods of
scanning technology and the captured 3D time and localities.
morphology data of fish specimens is Specimens are typically acquired by
shared through the Open Science various capturing methods as discussed
Framework website widely elsewhere by literature such as by
(https://osf.io/ecmz4/wiki/Fishes/). Such handnets, traps, seine nets, electrofishing
innovations have the potential to replace and even by buying directly from
pale and degraded voucher specimens that fishermen or the local fresh markets. Once
have been preserved for a long period of obtained, they must be immediately
time in alcohol. Morphological and labelled with 1) serial number, 2) species
meristic database compilation can be name, 3) name of collector, 4) location, 5)
instantly peer-reviewed and shared online date of collection, 6) GPS coordinates,
with any amateurs or experts located and 7) some description of the specimens’
anywhere around the world. This is an colour and body markings in fresh should
opportunity for scientists to engage more be recorded (Fischer, 2013; Motomura
with the society and garner support from and Ishikawa, 2013). This may be
the public and policymakers for fish supported by high resolution photographs
conservation. of the whole fish and specific parts with
distinctive characters (Figs. 17, 18 and
Specimen Collection and Preservation 19).
Figure 17: Fresh specimen is photographed facing left together with a ruler to indicate scale.
Figure 18: A live specimen is best photographed with high-resolution photography to record subtle
details and colouration.
Figure 19: Noticeable variations of the same species should be photographed for species-level
studies. For example, in the case of Tor tambra (above), individuals in the same
population may display differing forms of mental lobe and barbels (Roberts, 1999).
Specimens should be cleaned and can (2006) suggest that in the absence of
either be frozen or immediately fixed in refrigeration during field work, specimens
formalin or alcohol before being fixed in 10% buffered formalin or 95%
transferred to depositories. alcohol should be analyzed with DNA
In fish taxonomy, the left side of the processes within one week to produce the
fish body is examined. Therefore if best results.
muscle tissue is required for molecular In the museums and depositories,
analysis, it should be dissected from the taxonomists are cautious when fixing and
right side of the body. In small fishes storing specimens (Fig. 20) since any flaw
where muscle tissues are difficult to in the process will devalue their academic
excise, the right pelvic fin may be significance. Specimens are usually fixed
collected as an alternative (Shiozawa et in 10% formalin solution and large sized
al., 1992). Typically, 1.0 cm3 tissue from specimens are incised on the right side to
the fresh muscle contains enough genomic enable better absorption of formalin
material for molecular study and it should (Schander and Halanych, 2003; Garrigos
be stored in 95% alcohol and then et al., 2013). For the same purpose,
refrigerated at -20°C (Motomura and formalin may also be injected into the
Ishikawa, 2013). Preferably, it should be abdomen by using a syringe (Motomura
free from fat and blood which may and Ishikawa, 2013).
hamper the DNA purification process
(Wong et al., 2012). Chakraborty et al.
Figure 20: Fixed specimens are usually stored in glass jars with heads pointing downwards. Note
that specimens become pale and lose their colouration quality, which is why
photography of live specimens is crucial as part of the data collection process.
Taxonomic Key 9. Field notes – description of sympatric

After fixing, labelling and cataloguing a and syntonic species found in the same
specimen, a competent taxonomist would habitat when the specimen is collected.
carry out the standard morphometrics, 10. Remarks or comments – description of
meristics and sometimes molecular precautions as a measure against
analyses as mentioned earlier. As a misidentification and any other useful
minimum, the taxonomist would information for effective identification.
subsequently generate a report with the If a potential new species is
following information (Fischer, 2013) for encountered, the specimen and full-
each species; colour photographs are to be sent to an
1. Species name, author and year. authority of the genus for further
2. Material examined – description of type investigation. This shall be recorded in
material and voucher specimen this section.
examined. A competent taxonomist would update
3. Diagnosis – description of the the field “taxonomic key” for the
specimen’s key markers or corresponding genus once a new species
morphological features that has been identified. It is a conventional
differentiates it from nearest congeners tool meant for quick and practical
from the same watershed and other identification based on the major
watersheds. Diagrams may be included morphological characters (Fig. 21) of a
for clarity. species (Fischer, 2013). A typical
4. Description – description of major taxonomic key is organized in a series
morphological and meristic data. diagnostic characteristic of a species that
5. Pigmentation in life – description of lead the user to the correct name of a
colour, marking and patterns on the given specimen. A taxonomic key is
body and fins of a fresh specimen. called a dichotomous key (in Latin
6. Colour in formalin or alcohol – “dichotomous” means in two parts)
description of colour, markings and because only two marker options are
patterns on the body and fins of the offered in each step (Fig. 22). The
fixed specimen. markers provided may be quantitative
7. Distribution – description of location (e.g., scale count) or qualitative (e.g.,
where the specimen and the species body colour). Once created, the key can
can be typically found according to be continuously improved by taxonomists
published literature. based on feedback from users.
8. Etymology – description of the
Greek/Latin word or rationale behind
the scientific name (binomial
nomenclature) assigned to the species.
Figure 21: The key characteristics of a genus are typically described at the start of the taxonomic
key. For example, the Glyptothorax genus’s key characteristic is the presence of a
unique thoracic adhesive apparatus (arrow) that can only be observed in the ventral
view.
Figure 22: An example of a dichotomous key produced by Silfvergrip (2009, p.25) for the
identification of freshwater eels from the Anguillidae family.
Integrative taxonomy family due to species and subspecies from

There are species within the same genus the family have sympatric distribution and
that possess small variations that are not the morphological characters among them
easy to discriminate. This is compounded are hard to distinguish (Castle and
by the unstable and convoluted taxonomic Williamson, 1974; Aoyama et al., 2000;
history of some species which can Arai and Wong 2016). For example,
experience numerous revisions of species Sugeha and Suharti (2009) found
names because taxonomists may not share difficulty in distinguishing Anguilla
the same assumptions and approaches. bicolor bicolor and Anguilla bicolor
Correspondingly, it is fair to expect that pacifica based on morphological
the science of taxonomy and systematics attributes alone. However, they were able
is constantly in flux and revisions may be to make a distinction between the species
frequent. by their adult sizes. Generally, A. bicolor
A classic example is the case of bicolor is longer and heavier than A.
Neolissochilus spp. and Tor spp. found in bicolor pacifica. They also found that A.
Southeast Asia. These genera are typically bicolor bicolor tend to occur in Sumatra
prone to trophic polymorphism and and Java region and A. bicolor pacifica is
display conspicuous oral morphology found in Sulawasi and New Guinea region
variation (Roberts and Khaironizam, of Indonesia. As such, they have proposed
2008) that have confused even the most that the two subspecies can be
experienced ichthyologists. For example, distinguished by the geographic approach.
in early 20th Century, members of the Subsequently, Teng et al. (2009) resolved
Neolissochilus genus were placed in the the issue by conducting a phylogenic
Barbus genus (Boulenger, 1893; Duncker, study and validated the notion that
1904). Later they were reassigned Anguilla species and subspecies can
respectively to Labeobarbus, cluster regionally (Fig. 23).
Crossochilus, Puntius and Acrossocheilus The case of Anguilla genus is a classic
(Weber and de Beaufort 1916; Ahl 1933; example how the combination of
Fowler 1934; Herre and Myers 1937; morphological and molecular approaches,
Smith 1945) before being finalized as generally termed as integrative taxonomy
Neolissochilus, a new genus created by (Goulding and Dayrat, 2016), are crucial
Rainboth (1996). in gaining and expanding the
In cases when the taxonomy of a understanding of taxonomy and diversity
particular genus has stabilized, in fish. This demonstrates that the
misidentification can still be common broadest range of methods should be
because small variations can be difficult utilized to answer and solve taxonomic
to be distinguished. For example, in the concerns.
past decades, there were numerous reports
that highlighted misidentification of the
freshwater eels from the Anguillidae
Figure 23: The morphological phylogenetic tree of the genus Anguilla showing correlation species
occurrence in various regions (Source: Teng et al., 2009).
Acknowledgements between two species of freshwater eels,

We wish to extend our deepest gratitude Anguilla celebesensis and A. interioris.
to all past and present taxonomists around Ichthyological Research, 47, 157-161.
the world whose life time work and Arai, T. and Wong L.L., 2016.
publications have provided substance for Validation of the occurrence of the
this manuscript. tropical eels, Anguilla bengalensis
bengalensis and A. bicolor bicolor at
References Langkawi Island in Peninsular
Ahl, E., 1933. Ueber eine kleine Malaysia, Malaysia. Tropical
Fischsammlung aus dem Toba-See in Ecology, 57(1), 23-31.
Sumatra. Sitzungsberichte der Assis, C.A., 2003. The lagenar otoliths of
Gesellschaft Naturforschender teleosts: Their morphology and its
Freunde, 514–516. application in species identification,
Amundsen, P., Bøhn, T. and Våga, phylogeny and systematics. Journal of
G.H., 2004. Gill raker morphology and Fish Biology, 62(6), 1268–1295.
feeding ecology of two sympatric Authman, M.M.N., Zaki, M.S., Khallaf,
morphs of European whitefish E.A. and Abbas, H.H., 2015. Use of
(Coregonus lavaretus). Annales fish as bio-indicator of the effects of
Zoolologici Fennici, 41, 291–300. heavy metals pollution. Journal of
Aoyama, J., Watanabe, S., Ishikawa, S., Aquatic Research and Development, 6,
Nishida, M. and Tsukamoto, K., 328.
2000. Are morphological characters
distinctive enough to discriminate
Avise, J.C., 1994. Molecular markers, Review of Ecology, Evolution, and

natural history, and evolution. New Systematics, 38, 459–487.
York: Chapman and Hall. Boulenger, G.A., 1893. List of the fishes
Ayoade, A.A., Sowunmi, A.A. and collected by Mr E.W. Oates in
Nwachukw, H.I., 2004. Gill southern Shan State and presented by
asymmetry in Labeo ogunensis from him to the British Museum. The Annals
Ogun River, Southwest Nigeria. and Magazine of Natural History,
Revista de Biologia Tropical, 5(1), 12(6), 198–203.
171-175. Bridge, P.D., Roberts, P.J., Spooner,
Balon, E.K., 1995. Origin and B.M. and Panchal, G., 2003. On the
domestication of the wild carp, unreliability of published DNA
Cyprinus carpio: From sequences. New Phytologist, 160, 43–
Romangourmets to the swimming 48.
flowers. Aquaculture, 129, 3-48. Britton, J.R. and Davies, G.D., 2006.
Basolo, A.L., 1990. Female preference First record of the white catfish
predates the evolution of the sword in Ameiurus catus in Great Britain.
swordtail fish. Science, 250, 808–810. Journal of Fish Biology, 69, 1236–
Berkes, F., Coldings, J. and Folke, C., 1238.
2000. Rediscovery of Traditional Bruet, B., Song, J., Boyce, M. and
Ecological Knowledge as Adaptive Ortiz, C., 2008. Materials design
Management. Ecological Applications, principles of ancient fish armour.
10, 1251-1262. Nature Materials, 7, 748-756.
Berquist, R.M., Gledhill, K.M., Butlin, R.K., Galindo, J. and Grahame,
Peterson, M.W., Doan, A.H., Baxter, J.W., 2016. Sympatric, parapatric or
G.T., Yopak, K.E., Kang, N., allopatric: the most important way to
Walker, H.J., Hastings, P.A. and classify speciation? Philosophical
Frank, L.R., 2012. The Digital Fish Transactions of the Royal Society B,
Library: Using MRI to Digitize, 363, 2997–3007.
Database, and Document the Calamia, M.A., 1999. A methodology for
Morphological Diversity of Fish. PLoS incorporating traditional ecological
ONE, 7(4), e34499. knowledgewith geographic information
Blount, J.D., Metcalfe, N.B., Birkhead, systems for marine resource
T.R. and Surai, P.F., 2003. management in the Pacific.
Carotenoid modulation of immune http://www.spc.int/Coastfish/publicatio
function and sexual attractiveness in ns/bulletins/traditional-
zebra finches. Science, 300, 125–127. management/212-traditional-
Bolnick, D.I. and Fitzpatrick, B.M., information-bulletin-10.html
2007. Sympatric Speciation: Models Campana, S.E., 2005. Otolith science
and Empirical Evidence. Annual entering the 21st century. Marine and
Freshwater Research, 56, 485–495.
Carson, E.W., 2011. Low but Stable vertebrate voucher specimens, with
Frequency of Xanthic Phenotypes in a guidelines for ethical collection.
Population of the Twoline Pupfish, Memoirs of Museum Victoria, 72, 141-
Cyprinodon bifasciatus. The American 151.
Midland Naturalist, 166(2), 462-466. Cracraft, J., 1983. Speciation and its
Castle, P.H.J. and Williamson, G.R., ontology: The empirical consequences
1974. On the validity of the freshwater of alternative species concepts for
eel species Anguilla ancestralis from understanding patterns and processes
Celebes. Copeia, 2, 569-570. of differentiation. Speciation and its
Cawthorn, D.M., Steinman, H.A. and Consequences, 28–59.
Witthuhn, R.C., 2012. DNA Curet, O.M., Patankar, N.A., Lauder,
barcoding reveals a high incidence of G.V. and MacIver, M.A., 2011.
fish species misrepresentation and Mechanical properties of a bio-inspired
substitution on the South African robotic knifefish with an undulatory
market. Food Research International, propulsor. Bioinspiration and
46, 30–40. Biomimetics, 6(2), 1-9.
Chakraborty, A., Sakai, M. and Davey, J.W. and Blaxter, M.L., 2011.
Iwatsuki, Y., 2006. Museum fish RADSeq: Next-generation population
specimens and molecular taxonomy: A genetics. Briefings in Functional
comparative study on DNA extraction Genomics, 9(5), 416-423.
protocols and preservation de Carvalho, M.R., Bockmann,
techniques. Journal of Applied F.A., Amorim, D.S., de Vivo, M., de
Ichthyology, 22(2), 160-166. Toledo-Piza, M., Menezes, N.A., de
Chauhan, T. and Rajiv, K., 2010. Figueiredo, J.L., Castro, R.M., Gill,
Molecular markers and their A.C., McEachran, J.D., Compagno,
applications in fisheries and L.J., Schelly, R.C., Britz,
aquaculture. Advances in Bioscience R., Lundberg, J.G., Vari, R.P.
and Biotechnology, 1, 281-291. and Nelson G., 2007. Revisiting the
Chen, W., Al-Husaini, M., Beech, M., taxonomic impediment. Science, 307,
Al-Enezi, K., Rajab, S. and Husain, 353.
H., 2011. Discriminant analysis as a Dieckmann, U. and Doebeli, M., 1999.
tool to identify catfish (Ariidae) On the origin of species by sympatric
species of the excavated archaeological speciation. Nature, 400, 354-357.
otoliths. Environmental Biology of Dingerkus, G., Seret, B. and Guilbert,
Fishes, 90, 287–299. E., 1991. The first albino Wels, Silurus
Clemann, N., Rowe, K.M.C., Rowe, glanis Linnaeus, 1758, from France,
K.C., Raadik, T., Gomon, M., with a review of albinism in catfishes
Menkhorst, P., Sumner, J., Bray, D., (Teleostei: Siluriformes). Cybium, 15,
Norman, M. and Melville, J., 2014. 185–188.
Value and impacts of collecting
Drew, J.A., 2005. Use of Traditional Fischer, J., 2013. Fish identification tools
Ecological Knowledge in Marine for biodiversity and fisheries
Conservation. Conservation Biology, assessments: Review and guidance for
19, 1286-1293. decision-makers. FAO Fisheries and
Duncker, G., 1904. Die fische de Aquaculture Technical Paper No. 585.
Malayichen Halbinsel. Mitteilungen Rome: FAO.
aus dem Naturhistorischen Museum in Fonge, B.A., Tening, A.S., Egbe, A.E.,
Hamburg, 21, 133–207. Awo, E.M., Focho, D.A., Oben, P.M.,
Eastman, J.T., 1977. The Pharyngeal Asongwe, G.A. and Zoneziwoh,
Bones and Teeth of Catostomid Fishes. R.M., 2011. Fish (Arius heudelotii
American Midland Naturalist, 97(1), Valenciennes, 1840) as bioindicator of
68-88. heavy metals in Douala Estuary of
Ebach, M.C. and Holdrege, C., 2005. Cameroon. African Journal of
DNA barcoding is no substitute for Biotechnology, 10(73), 16581-16588.
taxonomy. Nature, 434. Forster, P., 2003. To err is human.
Endler, J.A., 1980. Natural selection on Annals of Human Genetics, 67, 2-4.
color patterns in Poecilia reticulata. Fowler, H.W., 1934. Zoological results
Evolution, 34, 76–91. of the third de Schauensee Siamese
Enghoff, H., 2009. What is taxonomy? – Expedition, Part I – Fishes.
An overview with myriapodological Proceedings of the Academy of Natural
examples. Soil Organism, 81(3), 441– History of Philadelphia, 86, 67-163.
451. Fujii, R., Kasuwaka, H., Miyaji, K. and
Feeley, K.J. and Silman, M.R., 2011. Oshima, N., 1989. Mechanism of skin
Keep collecting: accurate species coloration and its changes in the blue-
distribution modelling requires more green damselfish. Zoological Science,
collections than previously thought. 6, 477-486.
Diversity and Distributions, 2011, 1-9. Garrigos, Y.E., Hugueny, B., Koerner,
Feist, S.W. and Longshaw, M., 2008. K., Ibañez, C., Bonillo, C., Pruvost,
Histopathology of fish parasite P., Causse, R., Cruaud, C. and
infections– Importance for populations. Gaubert, P., 2013. Non-invasive
Journal of Fish Biology, 73, 2143– ancient DNA protocol for for fluid-
2160. preserved specimens and phylogenetic
Ferreira, H.M., Reuss-Strenzel, G.M., systematics of the genus Orestias
Alves, J.A. and Schiavett, A., 2014. (Teleostei: Cyprinodontidae). Zootaxa,
Local ecological knowledge of the 3640 (3), 373-394.
artisanal fishers on Epinephelus itajara Godfray, H.C.J., 2002. Challenges for
(Lichtenstein, 1822) (Teleostei: taxonomy. Nature, 417, 17-19.
Epinephelidae) on Ilhéus coast – Bahia Goulding, T.C. and Dayrat, B., 2016.
State, Brazil. Journal of Ethnobiology Integrative taxonomy: Ten years of
and Ethnomedicine, 10(51), 1-15. practice and looking into the future.
Archives of Zoological Museum of Henkanaththegedara, S.M. and

Lomonosov Moscow State University, Stockwell, C.A., 2011. Melanism In
54, 116-133. Endangered Mohave Tui Chub
Grether, G.F., 2000. Carotenoid Siphateles Bicolor Mohavensis Snyder
limitation and mate preference 1918 (Cypriniformes: Cyprinidae).
evolution: a test of the indicator Western North American Naturalist,
hypothesis in guppies (Poecilia 71(1), 127-130.
reticulata). Evolution, 54, 1712-1724. Herre, A.W. and Myers. G.S., 1937. A
Grinnell, J., 1910. The methods and uses contribution to the ichthyology of the
of a research museum. Popular Science Malay Peninsula. Bulletin of the
Monthly, 77, 163-169. Raffles Museum, 13, 5-75.
Guerra-Gracia, J.M., Espinosa, F. and Hinsinger, D.D., Debruyne, R.,
Garcia-Gomez, J.C., 2008. Trends in Thomas, M., Denys, G.P.J.,
Taxonomy today: an overview about Mennesson, M., Utge, J. and Dettai,
the main topics in Taxonomy. A., 2015. Fishing for barcodes in the
Zoologica Baetica, 19, 15-49. Torrent: from COI to complete
Hawkes, J.W., 1974. Structure of fish mitogenomes on NGS platforms. DNA
skin. Cell and Tissue Research, 149, Barcodes, 3, 170–186.
147-172. Horth, L., 2002. Melanic body colour
Hebert, P.D.N., Ratnasingham, S. and and aggressive mating behaviour are
de Waard, J.R., 2003. Barcoding correlated traits in male mosquitofish
animal life: Cytochrome c oxidase (Gambusia holbrooki). Proceedings of
subunit 1 divergences among closely the Royal Society of London Series B,
related species. Proceedings of the 270, 1033-1040.
Royal Society B, 270, 96-99. Houde, A.E., 1997. Sex, color, and mate
Hebrank, M.R. and Hebrank, J.H., choice in guppies. Princeton: Princeton
1986. The mechanics of fish skin – University Press.
lack of an external tendon role in 2 Hulsey, C.D., Hendrickson, D.A. and
teleosts. Biological Bulletin, 171(1), García de León, F.J., 2005. Trophic
236-247. morphology, feeding performance and
Helfman, G.S., Collette, B.B., Facey, prey use in the polymorphic fish
D.E. and Bowen, B.W., 2009. The Herichthys minckleyi. Evolutionary
diversity of fishes: Biology, evolution Ecology Research, 7, 1–22.
and ecology. Oxford: Wiley-Blackwell Iwata, A. and Jeon, S.R., 1995.
Publishing. Descriptions of Cephalic Canals of
Hemmer-Hansen, J., Therkildsen, N.O. Lophiogobius ocellicauda. Japanese
and Pujolar, J.M., 2014. Population Journal of Ichthyology, 42, 329-333.
Genomics of Marine Fishes: Next- Jeffery, W.R., 2006. Regressive
Generation Prospects and Challenges. evolution of pigmentation in the
Biological Bulletin, 227, 117-132.
cavefish Astyanax. Israel Journal Assignment of RAPD Markers to

of Ecology & Evolution, 52, 405-422. Multipoint Linkage Groups. Marine
Jinbo, U., Kato, T. and Ito, M., 2011. Biotechnology, 5, 279-293.
Current progress in DNA barcoding Khoo, G., Lim, T.M. and Phang,
and future implications for V.P.E., 2011. A Review of PCR-Based
entomology. Entomological Science, DNA Fingerprinting Using Arbitrary
14(2), 107-124. Primers in Tropical Ornamental Fishes
Jorge, F., Perera, A., Rocas, V., of South East Asia. Journal of
Carretero, M.A., Harris, D.J. and Advanced Medical Research, 1, 71-93.
Poulin, R., 2014. Evolution of Khoo, G., Lim, T.M. and Phang,
alternative male morphotypes in V.P.E., 2012. Ultrastructure of
oxyurid nematodes: A case of erythrophores and xanthophores of the
convergence? Journal of Evolutionary Siamese Fighting Fish, Betta
Biology, 27, 1631-1643. splendens. The Israeli Journal of
Kajishima, T., 1977. Genetic And Aquaculture, 64, 1-7.
Developmental Analysis Of Some New Khoo, G., Lim, T.M. and Phang,
Color Mutants In The Goldfish, V.P.E., 2014. Cellular basis of metallic
Carassius Auratus. Genetics, 86, 161- iridescence in the Siamese fighting
174. fish, Betta splendens. The Israeli
Kapoor, V.C., 1998. Principles and Journal of Aquaculture, 66, 1-10.
practices of animal taxonomy. Science Khoo, G., Loh, E.Y.F, Lim, T.M. and
Publishers. Phang, V.P.E., 1997. Genetic
Kawase, S. and Hosoya, K., 2015. variation in different varieties of
Pseudorasbora pugnax, a new species Siamese fighting fish using isoelectric
of minnow from Japan, and focusing of sarcoplasmic proteins.
redescription of P. pumila (Teleostei: Aquaculture International, 5, 537-549.
Cyprinidae). Ichthyology Exploration Kimura, M., 1968. Evolutionary rate at
of Freshwaters, 25(4), 289-298. the molecular level. Nature, 624-626.
Khodadoust, D., Ismail, A., Zulkifli, Koken, E., 1884. Über Fisch-Otolithen,
S.Z. and Tayefeh, F.H., 2013. Short insbesondere uber diejenigen der
Time Effect of Cadmium on Juveniles norddeutschen Oligocän-
and Adults of Java Medaka (Oryzias Ablagerungen. Zeitschrift der
javanicus) Fish as a Bioindicator for deutschen Geologischen Gesellschaft,
Ecotoxicological. Life Science Journal, 36, 500-565.
10(1), 1857-1861. Kottelat, M., 2013. The Fishes of the
Khoo, G., Lim, M.H., Suresh, H., Gan, Inland Waters of Southeast Asia: A
D.K.Y., Lim, K.F., Chen, F., Chan, Catalogue and Core Bibliography of
W.K., Lim, T.M. and Phang, V.P.E., the Fishes Known to Occur in
2003. Genetic Linkage Maps of the Freshwaters, Mangroves and Estuaries.
Guppy (Poecilia reticulata):
The Raffles Bulletin of Zoology, 27, 1- 2008. Global diversity of fish (Pisces)
663. in freshwater. Hydrobiologia, 595,
Krell, F.T. and Wheeler, Q.D., 2014. 545-567.
Specimen collection: Plan for the Lewand, K.O., Hyde, J.R., Buonaccorsi,
future. Science, 344, 815-816. V.P. and Lea, R.N., 2013. Orange
Kress, J.W., Garcia-Robledo C., coloration in a black-and-yellow
Uriarte M., and Erickson, D.L., rockfish (Sebastes chrysomelas) from
2015. DNA barcodes for ecology, central California. California Fish and
evolution, and conservation. Trends in Game, 99(4), 237-239.
Ecology and Evolution, 30, 25-35. Lincoln, R., Boxshall, G. and Clark,
Krupp, F. and Budd, K., 2009. A new P.A., 1998. A dictionary of Ecology,
species of the genus Garra (Teleostei: Evolution and Systematics.
Cyprinidae) from Oman. International Cambridge: Cambridge University
Journal of Ichthyology, 15(2), 117- Press.
120. Lombarte, A., Chic, Ò., Parisi-
Kumar. V. and Hassan, M.A., 2015. Baradad, V., Olivella, R., Piera, J.
Methods and Procedures of Sampling, and García-Ladona, E., 2006. A web-
Preservation and Identification for Fish based environment from shape analysis
Taxonomy Studies. World Journal of of fish otoliths. The AFORO database.
Fish and Marine Sciences, 7(2), 105- Scientia Marina, 70, 147-152.
108. Long, J.H., Hale, M.E., McHenry, M.J.
Lauder, G.V., Anderson, E.J., and Westneat, M.W., 1996. Functions
Tangorra, J. and Madden, P.G.A., of fish skin: flexural stiffness and
2007. Fish biorobotics: kinematics and steady swimming of Longnose gar
hydrodynamics of self-propulsion. The Lepisosteus osseus. Journal of
Journal of Experimental Biology, 210, Experimental Biology, 199(10), 2139-
2767-2780. 2151.
Leal, M.E., Schulz, U.H., Albornoz, MacArthur, R.H., 1969. Patterns of
P.L., Machado, R., Ott, P.H., 2013. communities in the tropics. Biological
First record of partial albinism in two Journal of the Linnean Society, 1, 19-
catfish species of genidens 30.
(Siluriformes: Ariidae) in an MacLean, M., Breeze, H. and Doherty,
estuary of Southern Brazil. P., 2009. Using Fish Harvesters’ Local
Brazilian Archives of Biology and Ecological Knowledge (LEK) in
Technology, 56, 237- 240. Support of Identifying Ecologically
Lechner, W. and Ladich, F., 2010. How and Biologically Significant Areas
do albino fish hear? Journal of (EBSAs) on the Offshore Eastern
Zoology, 283, 186-192. Scotian Shelf. Oceans and Habitat
Leveque, C., Oberdorff, T., Paugy, D., Report 2009. Nova Scotia: Fisheries
Stiassny, M.L.J. and Tedesco, P.A., and Oceans Canada.
Mallet, J., Meyer, A., Nosil, P. and Motomura, H. and Ishikawa, S., 2013.
Feder, J.L., 2009. Space, sympatry Fish collection building and procedures
and speciation. Journal of manual. The Kagoshima University
Evolutionary Biology, 22, 2332-2341. Museum, Kagoshima and the Research
Mayr, E., 1942. Principles of Systematic Institute for Humanity andature,
Zoology. New York: McGraw-Hill. Kyoto.
Mayr, E., 1947. Ecological factors in http://www.museum.kagoshima-
speciation. Evolution, 1, 263-288. u.ac.jp/staff/motomura/CollectionMan
Mayr, E., 1959. Isolation as an ual_lowres.pdf
evolutionary factor. Proceedings of the Nelson, G.J., 1972. Cepahlic sensory
American Philosophical Society, 103, canals, pitlines, and the classification
221-230. of esocoid fishes, with notes on
McCune, A.R., 1981. Quantitative galaxiids and other teleosts. American
description of body form in fishes: Museum Novitates, 2492, 1- 49.
Implication for species level taxonomy Nelson, J.S., 2006. Fishes of the World,
and ecological influences. Copeia, 4, 4th Edition. New Jersey: John Wiley
897-901. and Sons Inc.
Mehinto, A.C., Martyniuk, C.J., Spade, O’Reilly, P. and Wright, J.M., 1995.
D.J. and Denslow, N.C., 2012. The evolving technology of DNA
Applications of next-generation fingerprinting and its application to
sequencing in fish ecotoxicogenomics. fisheries and aquaculture. Journal
Frontiers in Genetics, 3, 1-10. of Fish Biology, 47, 29–55.
Metz, J.R., Peters, J.M.J and Flik, G., Padilla, D.K. and Williams, S.L., 2004.
2006. Molecular biology and Beyond ballast water: aquarium and
physiology of the melanocortin system ornamental trades as sources of
in fish: A review. General and invasive species in aquatic ecosystems.
Comparative Endocrinology, 148, 150- Frontiers in Ecology and the
162. Environment, 2(3), 131-138.
Meyer, C.P. and Paulay, G., 2005. DNA Pandey, V., Nutter, R.C. and Prediger,
Barcoding: Error Rates Based on E., 2008. Applied biosystems SOLID
Comprehensive Sampling. PLoS system: ligation-based sequencing, In:
Biology, 3(12). Next Generation Genome Sequencing:
Moncey, V., 2012. Occurrence, Towards Personalized Medicine.
distribution and troglomorphisms of Weinheim: Wiley-VCH.
subterranean fishes of peninsular India. Parichy, D.M., 2006. Evolution of Danio
Current Science, 102(7), 1028-1034. pigment pattern development.
Moritz, C. and Cicero, C., 2004. DNA Heredity, 1–11
barcoding: promise and pitfalls. PLoS Partridge, B.L. and Pitcher, T.J., 1979.
Biology, 2. The Sensory Basis of Fish Schools:
Relative Roles of Lateral Line and
Vision. Journal of Rakshit, A., Paul, A., Bhattacharjee, S.,

Comparative Physiology, 135, 315- Banik, T., Saran, R., Mandal, B.,
325. Poddar, D. and Gangopadhyay, K.,
Pasco-Viel, E., Charles, C., Chevret, P., 2015. Cytogenetic and molecular
Semon, M., Tafforeau, P., Viriot, L. profiling of spotted snake head fish
and Laudet, V., 2010. Evolutionary Channa punctatus (Bloch, 1793) from
Trends of the Pharyngeal Dentition in three districts (Nadia, Hooghly and
Cypriniformes (Actinopterygii: north 24 Parganas) of west
Ostariophysi). PLoS ONE, 5(6). Bengal, India. International Journal of
Pereira, L.H.G., Hanner, R., Foresti, F. Fisheries and Aquatic Studies,
and Oliveira, C., 2013. Can DNA 3(1), 312-319.
barcoding accurately discriminate Razin, S. and Rottem, S., 1967.
megadiverse Neotropical freshwater Identification of Mycoplasma and
fish fauna? BMC Genetics, 14(20), 1- other microorganisms by
14. polyacrylamide-gel electrophoresis of
Price, A.C., Cameron, J., Weadick, cell proteins. Journal of Bacteriology,
Shim, J. and Rodd, F.H., 2006. 94, 1807-1810.
Pigments, Patterns, and Fish Behavior. Regan, C.T., 1910. The Asiatic fishes of
Zebrafish, 5(4), 297-307. the family Anabantidae. Proceedings
Pyke, G.H. and Ehrlich, P.R., 2010. of the Zoological Society of London,
Biological collections and ecological / 767-787.
environmental research: A review, Regan, J.D. 1961. Melanism in the
some observations and a look to the poecillid fish, Gambusia affinis (Baird
future. Biological Reviews, 85(2), 247- and Girard). American Midland
266. Naturalist, 65, 139–143.
Quigley, D.T.G. and Wallace, E., 2013. Reum, J.C., Paulsen, C.E., Pietsch,
Note first record of leucism in hake T.W. and Parker-Stetter, A.L., 2008.
Merluccius merluccius (L.). Bulletin of First Record of An Albino
the European Association of Fish Chimaeriform Fish, Hydrolagus
Pathologists, 33(3), 97-98. Colliei. Northwestern Naturalist, 89,
Quraishia, S.F., Panneerchelvam, S., 60– 62.
Zainuddin, Z. and Rashid, N.H.A., Ricklefs, R.E., 2004. A comprehensive
2015. Molecular Characterization of framework for global patterns in
Malaysian Marine Fish Species using biodiversity. Ecology Letters, 7, 1-15.
Partial Sequence of Mitochondrial Roberts, T.R. and Khaironizam, M.Z.,
DNA 12S and 16S rRNA Markers. 2008. Trophic Polymorphism in the
Sains Malaysiana, 44(8), 1119- Malaysia Fish Neolissochilus Soroides
1123. and other Old World Barbs (Teleostei,
Rainboth, J., 1996. Fishes of Cambodian Cyprinidae). Natural History Bulletin
Mekong. Rome: FAO. of the Siam Society, 56(1), 25-53.
Roberts, T.R., 1999. Fishes of the Candida albicans strains. Journal of

Cyprinid Genus Tor in The Nam Clinical Microbiology, 41, 552-557.
Theun Watershed (Mekong Basin) Of Sanger, F., Nicklen, S. and Coulson
Laos, With Description Of A New A.R., 1977. DNA sequencing with
Species. Raffles Bulletin of Zoology, chain-terminating inhibitors.
47(1), 225-236. Proceedings of the National Academy
Robins, C.R., Bailey, R.M., Bond, C.E., of Sciences, 74(12), 5463-5467.
Brooker, J.R., Lachner, E.A., Lea, Sbordoni, V., 2010. Strength and
R.N. and Scott, W.B., 1991. Common Limitations of DNA Barcode under the
and scientific names of fishes from the Multidimensional Species Perspective.
United States and Canada, 5th edition. Tools for Identifying Biodiversity:
American Fisheries Society Special Progress and Problems, 275-280.
Publication No. 20. Schander, C. and Halanych, K., 2003.
Rocha, L.A. et al. (120 co-authors), DNA, PCR and formalinized animal
2014. Specimen collection: An tissue - A short review and protocols.
essential tool. Science, 344, 814-815. Organisms Diversity &
Ross, H.A., Murugan, S. and Li, Evolution, 3(3), 195-205.
W.L.S., 2008. Testing the Reliability Shiozawa, D.K., Kudo, J., Evans, R.P.,
of Genetic Methods of Species Woodward, S.R. and Williams, R.N.,
Identification via Simulation. 1992. DNA extraction from preserved
Systematic Biology, 57(2), 216–230. trout tissues. Great Basin Naturalist,
Rothberg, J. and Leamon, J., 2008. The 5(1), 29-34.
development and impact of 454 Silfvergrip, A.M.C., 2009. CITES
sequencing. Nature Identification Guide to the Freshwater
Biotechnology, 26(10), 1117-1124. eels (Anguillidae). Swedish Museum
Rudkowska, I., Marcotte, B., Pilon, G., of Natural History, Swedish
Lavigne, C., Marette, A. and Vohl, Environmental Protection Agency.
M.C., 2010. Physiological Genomics, Simonov, V.M., 2008. An approach for
40(3), 189-194. genetic fish groups identification using
Salimi, N., Loh, K.H., Dhillon, S.K. and adaptability and characteristics of
Chong, V.C.. 2016. Fully-automated morphotype. Bulgarian Journal of
identification of fish species based on Agricultural Science, 14(2), 145-149.
otolith contour: using short-time Simpson, G.G., 1951. The species
Fourier transform and discriminant concept. Evolution, 5, 285-298.
analysis (STFT-DA). Peer J, 1-15. Singh, B.N, 2012. Concepts of species
Sampaio, P., Gusmao, L., Alves, C., and modes of speciation. Current
Pina-Vaz, C., Amorim, A. and Pais, Science, 103(7), 784-790.
C., 2003. Highly polymorphic Slavík, O., Horký, P. and
microsatellite for identification of Wackermannová, M., 2016. How
does agonistic behaviour differ in
albino and pigmented fish? Peer J, 4, Mechanical Systems Machine Elements

e1937. and Manufacturing, 45(4), 1100-
Smith, H.M., 1945. The freshwater fishes 1105.
of Siam or Thailand. Bulletin of the Sugeha, H. and Suharti, S., 2009.
United States Natural Museum, 188, 1- Discrimination and Distribution of
622. Two Tropical Short-Finned Eels
Smith, L.M. and Burgoyne, L.A., 2004. (Anguilla bicolor bicolor and Anguilla
Collecting, archiving and processing bicolor pacifica) in the
DNA from wildlife samples using IndonesianWaters. Publications of the
FTA(R) databasing paper. BMC Seto Marine Biological Laboratory, 9,
Ecology, 2004, 4(4), 1-11. 1-14.
Sowersby, W., Lehtonen, T.K. and Taberlet P., Coissac E., Pompanon F.,
Wong, B.B.M., 2014. Background Brochmann C., Willerslev E., 2012.
matching ability and the maintenance Towards next-generation biodiversity
of a colour polymorphism in the red assessment using DNA metabarcoding.
devil cichlid. Journal of Evolutionary Molecular Ecology, 21, 2045-2050.
Biology, 1-8. Teletchea, F., 2009. Molecular
Stacey, N., Karam, J., Dwyer, D., identification methods of fish species:
Speed, C. and Meekan, M., 2008. Reassessment and possible
Assessing Traditional Ecological applications. Reviews in Fish
Knowledge of Whale Sharks Biology and Fisheries, 19, 265–293.
(Rhincodon typus) in eastern Teng, H.Y., Lin, Y.S. and Tzeng, C.S.,
Indonesia: A pilot study with fishing 2009. A new anguilla species and a
communities in Nusa Tenggara Timur. reanalysis of the phylogeny of
Canberra: Charles Darwin University. freshwater eels. Zoological Studies,
Stepien, C. and Kocher, T.D., 1997. 48(6), 808-822.
Molecular systematics of fish: Chapter Termvidchakorn, A. and Hortle, K.G.,
1, Molecules and morphology in 2013. A guide to larvae and juveniles
studies of fish evolution. Academic of some common fish species from the
Press. Mekong River Basin. MRC Technical
https://www.utoledo.edu/nsm/lec/pdfs/ Paper No. 38. Phnom Penh: Mekong
moschp1.pdf River Commission.
Strauss, R.E. and Bookstein, F.L., 1982. Thuy, T.T., Hau, T.D., Nguyen, T.T.D.
The Truss: Body Form Reconstructions and Thanh, T.T., 2015. Diversity of
In Morphometrics. Systematic Zoology, otolith morphology in Nuchequula
31(2), 113-135. nuchalis (Temminck and Schlegel,
Sudo, S., Tsuyuki, K., Ito, Y. and 1845) larvae and juvenilles collected
Ikohagi, T., 2002. A study on the in the Tien Yen Estuary, Northern
surface shape of fish scales. JSME Vietnam. Tropical Natural
International Journal Series C – History, 15(1), 69-79.
Tillmar, A.O., Dell'Amico, B., barcoding and the need for integrative
Welander J, and Holmlund, G., taxonomy. Systematic Biology, 54(5),
2013. A universal method for species 844-851.
identification of mammals utilizing Wilson, E.O., 1985. Time to revive
next generation sequencing for the systematics. Science, 230.
analysis of DNA mixtures. PLoS ONE, Wong, L.L., Peatman, E., Lu, J.,
8(12), e83761. Kucuktas, H., He, S., Zhou, C.J.,
Turelli, M., Barton, N.H. and Coyne, J. Nanakorn, U. and Liu, Z.J., 2011.
A., 2001. Theory and speciation. DNA barcoding of catfish: Species
Trends in Ecology and Evolution, authentication and phylogenetic
16(7), 330-343. assessment. PLoS ONE, 6(3), e17812.
Van der Laan, R., Eschmeyer, W.N. Wong, P.B.Y, Wiley, E.O., Johnson,
and Fricke, R., 2014. Family-group W.E., Ryder, O.A., Brien, S.J.O,
names of Recent fishes. Zootaxa, Haussler, D., Koepfli, K.P., Houck,
3882(2), 1-230. M.L., Perelman, P., Mastromonaco,
Van Grouw, H., 2006. Not every white G., Bentley, A.C., Venkatesh, B.,
bird is an albino: Sense and nonsense Zhang, Y.P., Murphy, R.W. and
about colour aberrations in birds. G10KCOS, 2012. Tissue sampling
Dutch Birding, 28, 79–89. methods methods and standards for
Wägele, J.W., 2005. Foundations of vertebrate genomics. GigaScience, 1,
phylogenetic systematics. Verlag Dr. 8.
Friedrich Pfeil. München: Germany. Wright, S., Gray, R.D. and Gardner,
Walsh, S.J. and Chakrabarty, P., 2015. R.C., 2003. Energy and the rate of
A new genus and species of blind evolution: inferences from plant rDNA
sleeper (Teleostei: Eleotridae) from substitution rates in the western
Oaxaca, Mexico: First Obligate Cave Pacific. Evolution, 57, 2893-2898.
Gobiiform in the Western Hemisphere. Yedier, S., Kontaş, S., Bostanc, D. and
Copeia, 104(2), 506-517. Polat, N., 2016. Otolith and scale
Waugh, J., 2007. DNA barcoding in morphologies of doctor fish (Garra
animal species: Progress, potential and rufa) inhabiting Kangal Balıklı Çermik
pitfalls. BioEssays, 29, 188-197. thermal spring (Sivas, Turkey). Iranian
Weber, M. and de Beaufort, L.F., 1916. Journal of Fisheries Sciences, 15(4),
The fishes of the Indo-Australian 1593-1608.
Archipelago. Leiden: E.J. Brill.
Wheeler, Q.D., Raven, P.H. and
Wilson, E.O. 2004. Taxonomy:
Impediment or expedient? Science, 303
(5656), 285.
Will, K.W., Mishler, B.D. and Wheeler,
Q.D., 2005. The perils of DNA

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Journal of Survey in Fisheries Sciences 4(1) 54-93 2017

A Review of Fish Taxonomy Conventions and Species

Keat-Chuan Ng C.1; Aun-Chuan Ooi P.1; Wong W.L.1; Khoo G.1*

Received: December 2016 Accepted: March 2017

Keywords: Fish, Morphology, Molecular, Taxonomy, Species identification

Introduction interbreed in natural conditions. In the

Figure 2: Modes of speciation.

In sympatric speciation, biologists branch would still maintain a certain

Linnaeus’ binomial nomenclature system studies require considerable identification

Figure 4: Taxonomic ranking and naming convention.

Ray-finned fish which is reviewed in this ranking. According to ICZN regulation,

Another interesting example is the brittani) was named after a prominent

fish in Greek. Sure enough, it is a very hosts an isolated Channa gachua

Figure 13: Common mouth types.

The fundamental colour constituent in or fully (Slavík et al., 2016). Individuals

markers at molecular level to detect order or sequence in each gene is unique

31,220 species present globally (Jinbo et metabarcoding" to refer to their ability to

In the advent of technological In principle, a scientific finding must be

Taxonomic Key 9. Field notes – description of sympatric

Integrative taxonomy family due to species and subspecies from

Acknowledgements between two species of freshwater eels,

Avise, J.C., 1994. Molecular markers, Review of Ecology, Evolution, and

Archives of Zoological Museum of Henkanaththegedara, S.M. and

cavefish Astyanax. Israel Journal Assignment of RAPD Markers to

Vision. Journal of Rakshit, A., Paul, A., Bhattacharjee, S.,

Roberts, T.R., 1999. Fishes of the Candida albicans strains. Journal of

albino and pigmented fish? Peer J, 4, Mechanical Systems Machine Elements

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