A draft guide to learning and teaching ichthyology

using the FishBase information system[1]


Daniel Pauly[2]

Rainer Froese[3]


Maria Lourdes Palomaresc



This guide provides a structure and case study material for a computer-based course in ichthyology for upper undergraduate and graduates students in biology or environmental science.

The key resource made accessible through this guide is FishBase, a large database on the biology of fish, available on CD-ROM (for the Windows operating system) and on the Internet (

Following brief introductions to ichthyology and to FishBase, and to the use of the latter to teach the former, the key aspects of ichthyology are presented in five chapters covering Evolution and classification; Morphology and biodiversity; Reproduction; Physiology; and Fishes as part of ecosystems.

For each of these chapters, one or several ‘Exercises’ are presented describing how the relevant topics are covered in FishBase and describing how to access that information. ‘Tasks for the Student’ are provided, along with Internet links to relevant sources other than FishBase.

It is anticipated that this guide will improve as our experience with FishBase as a teaching tool improves. Thus, a final chapter describes how users (both students and teachers) may contribute to the frequent updates that are anticipated for this guide, and to completing the coverage by FishBase of the biology of fishes.


2.      Introduction

2.1.   What is ichthyology?

2.2.   What is FishBase?

2.3.   Why use one to teach the other?

3.      Evolution and Classification

3.1.   Phylogeny and classification

3.2.   Darwin and natural selection

3.3.   The species concept

3.3.1.      What’s in a name?

3.3.2.      Subspecies vs. populations

3.3.3.      Within-species diversity

3.3.4.      Common names

3.3.5.      Exercise 1

4.      Morphology and Biodiversity

4.1.   Diversity of Indo-Pacific shore fishes

4.2.   Diversity of shapes and sizes

4.2.1.      Exercise 2

4.3.   Diversity of brain sizes

4.3.1.      Exercise 3

4.4.   Diversity of growth and mortality

4.4.1.      Exercise 4

4.5.   Diversity of habitats: inferences from occurrence records

4.5.1.      Exercise 5

4.6.   Diversity of colors and sexual selection

4.6.1.      Exercise 6

4.7.   Diversity of food and feeding habits

4.7.1.      Table 1

4.7.2.      Exercise 7

5.      Reproduction

5.1.   The reproductive load concept

5.2.   Small eggs and no worries

5.2.1.      Exercise 8

5.3.   Large eggs and parental investment

5.3.1.      Exercise 9

5.4.   Variation on the basic themes

5.4.1.      Exercise 10

6.      Physiology

6.1.   Metabolism, gills and size

6.1.1.      Table 2

6.1.2.      Exercise 11

6.2.   Food consumption

6.2.1.      Exercise 12

6.3.   Estimating food consumption from empirical models

6.3.1.      Exercise 13

7.      Fish as Part of Exploited Ecosystems

7.1.   Food webs and trophic levels

7.2.   Trophic levels and sizes of fish

7.3.   Formal description of food webs

7.3.1.      Exercise 14

8.      Contributing to FishBase

9.      Acknowledgements

10.  References

11.  Appendices

11.1.                    Appendix A: Ichthyology resources on the net

11.2.                    Appendix B: fish-related web resources for UBC students

2.     Introduction

2.1.                      What is ichthyology?

Ichthyology, commonly defined as “the study of fish” or “that branch of zoology dealing with fish” has a long documented history, dating thousands of years back to the ancient Egyptians, Indians, Chinese, Greeks and Romans (Cuvier 1828).

This long, sustained interest in fish is due to their double role as highly speciose denizens of a fascinating, yet alien world, and as human food. It has generated, over the centuries, highly heterogeneous information—mainly taxonomic, but also referring to zoogeography, behavior, food, predators, environmental tolerances, etc.

This huge amount of information, embodied in a widely scattered literature, has gradually forced ichthyologists to specialize, and thus accounts on fish are now either global, but highly specialized (e.g. Eschmeyer’s Catalog of fishes (1998) or Pietsch and Grobecker’s Frogfishes of the world (1987) to name two outstanding representatives), or local and deep (e.g. Fryer and Iles’ Cichlid Fishes of the Great Lakes of Africa (1972) or Groot and Margolis’ Pacific Salmon Life Histories (1991).

Thus, with a few exceptions such as the massive Diversity of fishes (Helfman et al. 1997), texts are lacking which bring together, on a global basis, all aspects of ichthyology, such that they can be used for a specialized course, and/or independent learning.

2.2.                      What is FishBase?

FishBase is an information system available in the form of CD-ROMS and on-line, at, covering all fishes of the world in a fashion that is both global and deep. FishBase 99, whose accompanying book is available both in English and French, covers over 23,000 species of fish, i.e. most of the 25,000 extant species, and addresses the needs of a vast array of potential users, ranging from fisheries managers to biology teachers. The features of FishBase that enable it to meet such a wide range of needs reside in its architecture, which makes extensive use of modern relational database techniques.

Other features of FishBase are:

·        all information on a given species in the database is accessible through a unique scientific or common name;

·        the wide use of multiple choice field structures standardized qualitative information;

·        numeric fields record quantitative information that has been previously standardized;

·        numerous cross-relationships between data tables enable previously unknown relationships to be discovered; and

·        complementary databases provided by colleagues and linked to FishBase proper, contribute to making the combined package the most comprehensive data source of its kind.

2.3.                      Why use one to teach the other?

For teachers of aquatic biology, or of specialized ichthyology courses, the uses of FishBase will range from practical solutions to theoretical issues:

·        FishBase is directly useable as data source (i.e., as an electronic encyclopedia on fish), thus complementing classical sources of information on fish, e.g., the Zoological Record or Aquatic Science and Fisheries Abstracts, and helping overcome the lack of scientific literature, especially in developing countries;

·        the many pictures in FishBase can be used, just as those in taxonomic books, to provide students with a visual impression of the morphological and color diversity of fish, and/or of specific features of various groups;

·        students will be able to assess the state of knowledge on various groups of fish, and thus obtain some guidance in identifying worthwhile projects; and

·        the synoptical view that FishBase produces by assembling and structuring all available information on one species will help students to obtain material for study (see above) and, perhaps more importantly, to develop a sense of how scattered bits of knowledge can be used to ‘reconstruct’ species, and to show how these fit into their environments, thus encouraging a ‘holistic view’, as now required for most of what we do in the biological sciences.

Thus, a series of lectures on ichthyology may be conceived, based on the following elements:

·        show FishBase pictures through an introductory lecture, to highlight the diversity and colorfulness of fish and similarity of external morphology in related groups (this hopefully would serve to generate interest in the course as a whole, and introduce fish classification);

·        compare the early classification schemes in Cuvier (1828) with a recent one, e.g., that in the Catalog of fishes (Eschmeyer 1998), ‘hosted’ by FishBase and largely identical with the widely used classification in Nelson (1994);

·        introduce the species concept and its requirements (a formal description with figures, a binomen, a holotype, a type locality, etc.) and implications (synonymies, sister species, etc.), using FishBase as source of examples, and its Glossary for definition of terms;

·        define the characteristics (meristics, morphometrics) through which fish species are usually defined and hence identified, and compare identification through keys with computer-based identification using the appropriate FishBase routine (see ‘Quick Identification’);

·        show how museum and other occurrence records, as included in FishBase, can be used to define distribution ranges and habitats, which can then be used for ecological inferences;

·        show how the latitudinal ranges of fish species can be used to test various hypotheses, e.g., on the relationship between fish biodiversity and shelf area (for marine species) or land area (for freshwater species);

·        define and illustrate various life history strategies, and analyze their frequency distribution throughout the world. Show, e.g., that salmon-type anadromy is extremely rare in subtropical or tropical species (it is well documented only in hilsa, Tenualosa ilisha, ranging from Iraq to Myanmar). Show how students can identify the relative frequencies of different strategies and draw inferences from these;

·        let each student select a species, print out the relevant FishBase synopsis and complement it based on a literature review (and send the result to the FishBase Team); and

·        show or let students derive quantitative relationships between different expressions of fish physiology (e.g., respiration, growth) and temperature (and hence latitude) and identify modifying factors (salinity, gill size, food type, etc.).

In the context of higher education, FishBase may also serve as background for Bachelor’s or Master’s theses wherein an area of ichthyology not presently or suitably covered by the tables in the latest version of FishBase would be ‘broken up’ into choice, numeric and text fields, entered and then analyzed on a comparative basis[4].

3.     Evolution and Classification

3.1.                      Phylogeny and Classification

There are different ways in which objects can be classified and the human mind is very good at generating criteria for classification. This is why the following list, assembled by the Argentinean author Jorge Luis Borges, and purportedly extracted from an ancient Chinese encyclopedia (Lakoff 1987), strikes us as funny:

“…it is written that animals are divided into:

·        those that belong to the Emperor;

·        embalmed ones;

·        those that are trained;

·        suckling pigs;

·        mermaids;

·        fabulous ones;

·        stray dogs;

·        those that are included in this classification;

·        those that tremble as if they were mad;

·        innumerable ones;

·        those drawn with a very fine camel’s hair brush;

·        others;

·        those that have just broken a flower vase;

·        those that resemble flies from a distance.”

The two major criteria that are used to classify things (neither met by Borges’ list), are utility or affinity:

·        Utility generates classifications whose objects are easy to find. An example of such a classification would be a dictionary, whose entries are arranged alphabetically;

·        Affinity, on the other hand generates classification wherein adjacent objects s are straightforward to compare (because adjacent entries share important features).

In the European middle ages, animal books (‘Bestiarum’) were usually ordered alphabetically. However, such ordering eventually struck people as odd, especially as people realized, in the course of long debates on ‘universals’ (on whether names are ‘natural’ attributes of things, or not), that names are arbitrary labels.

Thus, authors gradually began seeking for natural classifications, wherein organisms are ordered by affinities, these affinities being initially conceived as reflective of the general rules which god used when creating these organisms.

The work of Linnaeus, whose Systema Naturae, the tenth edition of which in 1758 still marks the beginning of zoological nomenclature, is an example of such attempts to identify the underlying affinities among plants and animals. The resulting ‘natural’ classifications have started to make sense, however, only since Darwin, in The Origin of Species (1859), provided a rationale for affinities, that is, shared ancestry. Darwin not only provided a basis for the affinities between organisms, however. He also provided a mechanism by which new species and higher taxa emerged out of common ancestors. This mechanism he called natural selection.

3.2.                      Darwin and Natural selection

Natural selection is the core of Charles Darwin’s work and is best defined in his own terms: “many of every species are destroyed either in egg or [young or mature (the former state the more common)]. In the course of thousand generations infinitesimally small differences must inevitably tell; when unusually cold winter, or hot or dry summer comes, then out of the whole body of individuals of any species, if there be the smallest differences in their structure, habits, instincts [senses], health, etc., <it> will on an average tell; as conditions change a rather larger proportion will be preserved: so if the chief check to increase falls on seeds or eggs, so will, in the course of 1,000 generations, or ten thousand, those seeds (like one with down to fly) which fly furthest and get scattered most ultimately rear most plants, and such small differences tend to be hereditary like shades of expression in human countenance. (Darwin 1842)

Natural selection, thus, consists of three elements:

·        organisms usually produce far more progeny than their habitat can accommodate;

·        each member of the progeny differs in some inheritable attributes or properties;

·        there is a tendency for those progeny with attributes or properties that are more suitable for the habitat in question to suffer a lower rate of mortality and to reproduce better than their siblings.

These three features jointly cause animals and plants to try to track fluctuation of the environment. In this process, and in conjunction with other mechanisms such as the ‘founder effect’ and the effect of neutral selection, isolated populations can become so different from a mother species that they will not be able to mate if the barrier that once separated them disappears.

3.3.                      The species concept

Species are “groups of actually (or potentially) interbreeding natural populations which are reproductively isolated from other such groups” (Mayr 1942, p. 120).

3.3.1.      What’s in a name?

Since species are the basic rank of biological nomenclature, naming species is very important and we now follow for this a model proposed by Linnaeus, (see above), wherein the species is defined by a so-called binomen consisting of a unique genus name, always starting with a capital letter, and a species epithet , which is never capitalized; both are written in italics font. With regard to the capitalization rule, simply recall that the binomen is the short version of an earlier mode of description wherein a whole paragraph was used to describe, and thereby define, a species. The binomen, thus, was the start of a sentence.

An important addition to a species name is the name of the author who first described that species and the date of that description; as in, for example, the Linnaean species Salmo trutta Linnaeus, 1758. At times you will encounter a species, e.g. Oncorhynchus mykiss, with an author’s name and date in brackets, e.g. (Walbaum, 1792). In this case, it means that the species whose epithet is mykiss was originally described as a memeber of another genus, in this case Salmo, and due to better understanding of its relationships with other trouts, was subsequently moved into the genus Oncorhynchus which it is now a member.

Another rule important to animal species names are that the genus part of the name must be unique to the animal kingdom. From the year 2000 on, it must also be unique among all organisms. Thus, when a generic name is coined, the author must verify that this name has never been used by any other zoologist, and, from 2000 on, by any botanist, bacteriologist, etc. The apparently daunting task is not impossible, however, as global catalogues of organism names are now being created; the most important of these is the Species 2000 catalogue (see

3.3.2.      Subspecies vs. populations

Given the mechanism of natural selection, every fish population can be conceived as being a potential new species. All one needs to imagine is that populations become isolated from others long enough for their members to lose the ability to mate with those of other populations. However, as long as some members of each population continue to mate with members of other populations of the same species, a mating barrier will not emerge (only a small gene flow is required to prevent the emergence of a mating barrier). Thus populations, though it might be easy to define them in terms of attributes such as number of scales or spines or body proportions, should not be given full taxonomic status because (contrary to species) they usually do not maintain themselves over a long period. Not having taxonomic status also means they should not have formal names, such as the trinomen that are still frequently used today, e.g. Oreochromis niloticus niloticus. The third part of the trinomen refers to a subspecies, which is, in fact, a population, or, to use a term much used in earlier times, a ‘race’.

3.3.3.      Within-species diversity

Species differ as to the extent of their diversity. Some species consist of a single population of a few individuals — these are often endangered species. Others have wide ranges and a rich structure of populations – the situation which tempted authors to define subspecies as populations at opposite ends of a geographical range often differ in several characters. However, its is usually not objectively defined within-species diversity which has motivated authors to define subspecies, but national or local research traditions, and the resources available for taxonomy. Thus, Berg (1965) established numerous subspecies and even lower taxa for the fishes of adjacent lakes and rivers of the former Soviet Union, while subspecies are rarely proposed by taxonomists working on the many coral reef species of the Indo-Pacific, although their distribution spans thousands of kilometers, and detailed studies may justify this (at least if one believes in subspecies).

3.3.4.      Common names

The common names of fish are what most people know about most fish. Thus, capturing the common names of fish in various languages captures most of what people who speak these languages know about fish. For this reason, FishBase includes over 90,000 names of fish in over 100 languages, ranging from widespread languages such as English or Spanish, to languages spoken by few speakers, such as Haida in Haida Gwaii, British Columbia. Anthropologists, notably Berlin (1965), have established that essentially all ethnic groups in the world spontaneously differentiate a similar number (about 500) of ‘kinds’ of organisms, the kinds roughly corresponding to genera, with important species being named, as well as some of their life history stages.

The sounds in fish names also generate interesting patterns. Thus, small fishes (i.e., fishes with small values of Lmax) tend to have names containing high pitch sound such as ‘i’ or ‘ee’, while large fish tend to have names with lower pitch sounds, such as ‘a’, or ‘aa’ (Berlin 1992; Palomares et al. 1999).

3.3.5.      Exercise 1

Task for the Student:
  • Look at the scientific names of ten species whose author name is in brackets and identify for each the original name and several synonyms. List and define the different kinds of synonyms.
  • Identify a language with at least 50 different names in FishBase. Relate the number of species with i/ee sounds in their names against the maximum length reported for those species, i.e., test the occurrence of a sound-size association for fish in the language in question. [Tips: use the Information by country/island search to get a list of species and their common names; sort by language; get maximum size information from the Species Summary page and see item (5) of on how to export data to a spreadsheet (Excel format) for further analysis.]

Classification related topics covered in FishBase:


FishBase:  To search for terms included in the FishBase online glossary, go to and use the Glossary search by either typing in a term or browsing the index provided. Note that here, and for nearly all other terms in the glossary, you can click on the hypertext link to the Encyclopedia Britannica online.
Darwin, Charles: See Box 10 in The Expeditions Table
Species concept: Eschmeyer's Genera of Fishes; Eschmeyer's Species of Fishes
Go to, use the Scientific name search either by typing in the genus and species names or by browsing the provided index and select the Summary button. In the Species Summary page, click on the Synonyms link, e.g., Oncorhynchus mykiss
Subspecies:  The STOCKS Table
Go to and search for Oreochromis niloticus niloticus
Population:  The STOCKS Table
Threatened species: See Status field in The STOCKS Table
Go to, use the Information by country/island search, type in the country of interest and select the Threatened button.
Common names: See Fig. 6 in The COMMON NAMES Table
Go to and use the Common name search by typing in the name or by browsing the provided index. If a list of species is returned, click on the species of interest to access the Species Summary page. Then click on the Common names link, e.g., click on the Haida name ‘Skaagwun
Max. length (Lmax): The POPCHAR Table
Go to to search for a species as described above. Once in the Species Summary page, click on the Max. age & size link to obtain a list of maximum lengths, e.g., Salmo trutta
See also Key Facts example for Growth and Life span in Exercise 4 .



4.     Morphology and Biodiversity

The diversity of fish is larger than for any other vertebrate group. Not only are there more species of fish (25,000) than of all other vertebrates together, but also the range of body shapes and sizes of fish is larger than for mammals, birds or reptiles. Consequently, the range of habitat occupied is larger as well.

4.1.                     Diversity of Indo-Pacific shore fishes

The triangle formed by Indonesia, the Philippines and New Guinea, collectively referred to as the ‘East Indies’, form the center of marine fish biodiversity in the Indo-Pacific, with about 2,800 species naturally occurring there. These numbers drop with distance from this center to about 500 species in Hawaii and 120 species in the Easter Islands. The number of endemic species, i.e., fishes that do not occur outside a given area, increases with distance from the center, supporting one hypothesis that species evolved in the outer region and accumulated in the center. Another hypothesis holds that species evolved in the rich and stable habitats of the East Indies and were carried to the periphery by currents. Randall gives 5 explanations for fish biodiversity in the Indo-Pacific:

·        Sea surface temperatures in the East Indies were more stable during the glacial periods and thus extinction rates were lower than in the periphery;

·        Shelf area in the East Indies is much longer than that of the periphery, again making extinctions less likely;

·        Dispersal of shore fishes to remote islands occurs during the planktonic larval phase which lasts from several days to several weeks. However, the larval phase of many species is not long enough for long stretches of open ocean water, thus restricting their distribution;

·        Existing current patterns support dispersal of fish larvae from the area as well as convergence of larvae of species that have evolved in the periphery towards the East Indies;

·        During the last 700,000 years, there have been at least 3 ice age events that reduced the water level in the East Indies and separated populations long enough to become different species.

4.1.1.      Exercise 2

Task for the Student:
  • FishBase provides maps with species numbers by FAO area, country and ecosystem. It also provides maps with indications of how many species have been collected with in a cell of a grid system. Discuss pros and cons of these approaches in mapping species diversity. [Tip: see biodiversity maps at].
  • Use Randall’s 5 explanations to discuss the pros and cons of the ‘dispersal from center versus the ‘immigration from periphery’ hypothesis.

Biodiversity related topics in FishBase:


Distribution: The FAOAREAS Table; The COUNTRIES Table; The COUNTREF Table; The OCCURRENCES Table
Plot occurrence records, families by FAO area, species by FAO area, species by climate, etc. using the Biodiversity Maps routines in


4.2.                     Diversity of shapes and sizes

The shapes of fish are also extremely diverse, and include – besides the torpedo shape perceived as ‘typical’ for fishes and termed ‘fusiform’– shapes ranging from the serpentine (in the Anguilliformes and other orders) to the avian (in ‘flying fishes’), with Latimera chalumnae sporting limbs resembling, but not being used as, those of land-based tetrapods.

Shape and other morphological features are the key characteristics used to date for classifying fishes, and hence understanding their classification requires a basic overview of the basic shapes of fishes, as can be obtained from the outline drawings included, for each of the existing 500 fish families.

Size is the most important attribute of individual organisms; it determines what can be their food, and the extent to which they can be the prey of other organisms. Size also determines how much food an animal requires to eat, how fast it can swim, and to a large extent, where it can live.

The maximum size of fish can range from one centimeter in Philippine gobies, e.g., Pandaka pygmea to 13-15 meters in the Whale shark, Rhincodon typus. This diversity of size allowed widely different environments to be colonized, ranging from temporary puddles to the central gyres of the open ocean. However, colonizing these environments required other adaptations, involving growth and mortality rates, and their various correlates, discussed below.

4.3.                     Diversity of brain sizes

The brain size per body weight of adult animals is related to the sensory and behavioral capabilities of the respective species. For example, fishes with well-developed electrosensing capabilities are known to have large brains. The brain is the organ with the highest energy and oxygen demand, and thus, fishes as well as other animals have evolved brain sizes that are neither too small nor too large respective to the niches they occupy in nature.

4.3.1.      Exercise 3

Task for the Student:
  • Based on your general knowledge about the fish and their habitat, rank the following groups according to their brain size: coral reef fish, deep sea fish, herrings, sharks, coelacanths. Explain in 1-2 sentences why you ranked each group as such.
  • For the groups listed above, find typical examples, look at their brain size compared to other fishes, and use these data to prove or disprove your hypothesis about their respective groups. [Tip: fish common names often contain parts of group names].

Brain size related topics covered in FishBase:


Brains: See Box 27 and Fig. 42 in The BRAINS Table.
In the Species Summary page, click on the Brains link to obtain brain weight measurement data. Click on the Relative brain weight graph link to obtain a plot of encephalization coefficient (brain weight) vs. body weight.
To make a list of species with brain weight measurements, use the Information by topic search in, click on the Brains option.


4.4.                      Diversity of growth and mortality

In spite of this wide diversity of fish sizes, clear patterns do emerge: tropical fish tend to be smaller and faster-growing than their cold-water counterparts and their natural mortality tends to be higher. This is due to high temperature elevating the metabolic rates of tropical fish relative to their cold-water counterparts.

Correspondingly, the natural mortalities experienced by fish, which are a function of their sizes, range from values which exterminate an entire cohort in a few months, e.g., the round herring, to 50 and more years in the lake sturgeon and 150 years in the orange roughy. These enormous differences in life span allow fish to respond differently to habitat variations. Small, short-lived fish track such variations, for example, when growing up in temporary puddles and laying desiccation-proof eggs before they dry up, thus being able to live through dry periods or produce a successful cohort every 1-3 years or so (as may happen in such long-lived fish as cod).

4.4.1.      Exercise 4

Task for the Student:
  •   Identify 2 families, one tropical, one temperate, whose representatives share similar maximum sizes, and compare the distribution of their growth parameters on an auximetric grid.
  • Compare natural mortality (M) estimates for 10 species of tropical fish, ranging between 50 and 100 centimeters maximum length, with 10 species of fish with similar sizes from cold waters and test for a temperature effect. [Tip: temperature and M values maybe found in the Key Facts page.]

Size, growth and mortality related topics covered in FishBase:


Auximetric grid: Go to, use the Information by Family search, select a family, click on the Graphs option, select Auximetric graph, click on View graph.
Morphology: The MORPHOLOGY Table
Biodiversity: and The OCCURRENCES Table
Shapes (Fam. picts.): Go to, use the Information by Family search by choosing the Family of interest from the drop-down list and select the Family information button. In the Families page, click on the Pictures link to view the outline drawing representative of the Family.
Shapes (swim. mode): See Figs. 45 and 46 in The SWIMMING and SPEED Tables
[Note: Information on swimming modes is currently available only in the CD-ROM version. However, some biological information are available in the Species Summary page under the Biology field and in the Key Facts page. Swimming mode can also be inferred from the aspect ratio or the shape of the caudal fin. To make a list of species with such information, use the Information by topic search in, click on the Swim. type option and jot down the scientific name(s) of the species of interest. Then open the Species Summary and Key Facts page for that species or look at its Picture.]
Sizes (lengths): See Fig. 5 and Box 5 in The SPECIES Table; Fig. 13 and Box 13 in The POPCHAR Table; Fig. 29 in The ECOLOGY Table; Max. size field in the Species Summary page; click on the Max. age & size link for further information, e.g., in cod or select the Key Facts button in the Scientific name search (see Exercise 1 above) to display size related parameters, as in the example for, Oncorhynchus mykiss
Growth: See POPULATION DYNAMICS and link to The POPGROWTH Table and discussion on Auximetric Analyses
Download “Color versions of the graphs contained in Pauly D. 1998. Tropical fishes: patterns and propensities. J. Fish Biol. (53(A): 1-17” from
In the Species Summary page, click on the Growth link to obtain a list of growth and mortality parameters for different populations of a species, e.g., for Gadus morhua, then click on the Auximetric graph link to view a plot of growth coefficients vs. body lengths, e.g., for the cod.
Life span: See Fig. 16 in The POPGROWTH Table; Fig. 23 and Box 16 in section on Natural Mortality; Sizes (lengths) above; Life span field, e.g., for the rainbow trout.

4.5.                      Diversity of habitats: inferences from occurrence records

Fish inhabit more diverse habitats than any other group of vertebrates, ranging from Himalayan or Andean brooks at 4000 meters to abyssal depth at 10 kilometers, spanning an extremely high range of pressures. The range of temperatures that can be tolerated is also very large, from minus 2oC as tolerated by the Antarctic fish, Pagothenia borchgrevinki (which sport anti-freeze substances in their blood; see Eastman and Devries 1985); to up to 40o C for Oreochromis alcalicus, which lives at the edge of a hot spring in Lake Nakuru in Kenya. (This does not consider the temperature tolerance of deep-sea vent fishes, which have not yet been studied in detail).

Because fish occur only in habitats which they can tolerate, and tend to be abundant in those habitats to which they are best adapted, occurrence records kept by museums can be used to reconstruct the habitat preferences of fishes whose ecology is otherwise unknown. Such records have been named bioquads because they refer to biodiversity and consist of four key elements: (a) the name of the organism; (b) the place where it was caught; (c) the source or person who sampled or identified it; and (d) the date. FishBase makes wide use of bioquads for documenting the distribution of fish and this can be emulated by ichthyology students who may assemble bioquads from FishBase and other sources, notably the Internet. (see Appendix A for sources of bioquads).

4.5.1.      Exercise 5

Task for the Student:
  • Select a species in FishBase and print a point map as well as the point information. See whether you can find additional points in ichthyological museum collections (see Appendix A). Identify problematic records. Infer from the habitat, for which records exist, the ecological requirements of that species. [Note: links to point maps are available from the Species Summary page. Point information details are shown by clicking on a point (or dot) in a map.]

Distribution and occurrence related topics covered in FishBase:


Biodiversity: and see Biodiversity maps in
Environmental Info.: Under The SPECIES  Table, see Fig. 5 and Box 5 in Environmental Information
See Biology, Environment and Climate zone fields in the Species Summary page, e.g., for Pagothenia borchgrevinki.
Habitat and feeding: See Fig. 28 and Box 19 in The ECOLOGY Table
See Main food, Trophic level and Food consumption fields in Key facts page, e.g., for Oncorhynchus mykiss; clck on Diet link to obtain detailed information of food items.
To make a list of species with habitat and feeding information, use the Information by topic search in and click on the Diet option.
Occurrence: The OCCURRENCES Table and see also The INTRODUCTIONS Table
To make a list of species with introductions information, use the Information by topic search in and click on the Introductions option. See also Biodiversity Maps and plot, e.g., occurrence records by museum, familes by FAO area, species by climate zone.

4.6.                      Diversity of color and sexual selection

Fish are beautiful; they have beautiful colors and fascinating body shapes, one of the reasons why people keep them in aquaria. Color patterns in fish have been long misunderstood. Pre-Darwinian authors thought that god had given fish such marvelous colors so that predators would find it easier to see and catch them. We know, since Darwin, that such coloring, if it serves any function at all, must benefit directly the ones who sport it and not their predators as is obvious in the many color patterns that camouflage the owner, or confuse predators, by, e.g., displaying large eyes in the wrong places. Darwin also proposed a reason why non-camouflaging striking coloring should exist, and that is sexual selection.

Essentially, the males entice the females to choose them by displaying nicer colors than other males; they compete in terms of their ‘beauty’, this being related to good genes (remember Darwin did not know of genes and that part of his theory was very hard on him). Recently, the Zahavi’s have complemented Darwin’s version of sexual selection through a new concept, the handicap principle, which takes into account that the colors and other adornments which males use to entice females to choose them are costly to produce (Zahavi and Zahavi 1997). Hence, the color and other adornments represent a handicap and the males capable of displaying these attributes thus must have really good genes for life-supporting traits. We may call this ‘truth in advertisement.’

The idea is that sporting highly symmetrical patterns, as, for example, in Pomacanthus imperator, implies that the fish in question had a harmonious development since development problems, due to genetic problems, parasites or disease (also indicative of ‘bad genes’) would always lead to asymmetries. Also, for colors that do not necessarily camouflage the fish, sporting them indicates that the fish in question has been able to evade predators. Some fish imitate the color patterns of other species to fool prey or predators (mimicry).

4.6.1.      Exercise 6

Task for the Student:
  • Read chapter XII, Secondary sexual characters of fishes, amphibians and reptiles, in Charles Darwin’s Descent of Man, vol.2. Give a one-page summary of the argument and re-express the main line of Darwin’s argument using fish other than the one in that chapter.
  • Give examples from FishBase for species that use color patterns for a) camouflage, b) predator confusion, c) sexual selection.
  • Give one example of mimicry in fishes. Explain the benefits gained. [Tip: common names of such species often contain the word ‘mimic’].  

Morphology related topics covered in FishBase:


Morphology See links to information on MORPHOLOGY AND PHYSIOLOGY
Reproduction See links to information on REPRODUCTION and spawning
To make a list of species with morphology and reproduction information, use the Information by topic search in, and select the appropriate topic button. [Note: Information on morphology is currently available only in the CD-ROM version. However, some biological information are available in the Species Summary page under the Biology field and in the Key Facts page. To make a list of species with morphology information, use the Information by topic search in, click on the Morphology option and jot down the scientific name(s) of the species of interest. Then open the Species Summary and Key Facts page for that species or look at its Picture.]

4.7.                      Diversity of food and feeding habits

Given the diversity of their sizes and habitats it is obvious that fish should also have a wide diversity of food and feeding habits. Thus fish range from feeding on microscopic phyto- and zooplankton to engulfing entire adult fishes, such as is done by Whale sharks or gulpers, respectively. Attempts to link fish to their ecosystems have led to a huge literature on their food and feeding habits. Unfortunately, some of this is useless because it is reported in the wrong units, i.e., frequency of occurrence of certain items in a number of stomachs sampled. Still, there are enough studies in which the proper units have been used (percent contribution in weight, energy or volume to total stomach contents) for a clear idea to emerge of what fish generally eat in their typical habitat. Given knowledge of the average trophic level of their diet items, the trophic level of fish whose stomach content has been studied can thus be computed, which allows evaluation of the position the consumers occupy in the food web.

4.7.1.      Table 1

Hierarchy of food items, simplified from the FishBase table used to compute trophic levels (TL) from diet composition data. Therein, the TL of a consumer is 1 + (mean TL of the prey items).

Food I Food II   Food IIIa   TL  
Detritus detritus debris; carcasses 1.0
Plants phytoplankton blue-green algae; dinoflagellates; diatoms; green algae; other phytoplankton   1.0
  other plants benthic algae/weeds; periphyton; terrestrial plants 1.0
zoobenthos sponges/tunicates sponges; ascidians 1.0
  cnidarians hard corals and other polyps 1.0
  worms Polychaetes; other annelids; non-annelids 1.0
  mollusks chitons; bivalves; gastropods; octopi;, other mollusks 1.0
  benthic crustaceans ostracods;; isopods; amphipods; other small forms 1.0
    shrimps; lobsters; crabs stomatopod; other large forms  
  Insects Insects 1.0
  Echinoderms sea stars/brittle stars; sea urchins; sea cucumbers; etc 1.0
  other benthic inverts Other benthic invertebrates  
zooplankton jellyfish/hydroids jellyfish/hydroids 1.0
  planktonic crustaceans copepods; cladocerans; mysids; euphausiids; etc.   1.0
  other planktonic inverts n.a./other planktonic invertebrates 1.0
  finfish fish larvae 1.0
nekton Cephalopods squids/cuttlefish 1.0
  Finfish Bony fish and small sharks or rays 1.0
others Herps Salamanders/newts; toads/frogs; turtles and other reptiles 1.0
  Birds sea and shore birds  
  Mammals Small cetaceans and pinnipeds  

a)       in FishBase, these food items have distinct trophic levels (and associated standard errors), not presented here.

4.7.2.      Exercise 7

Task for the Student:
  • Find published studies on the diet composition of three different species of fish: one mainly herbivore; one omnivore, and one typical carnivore.
  • Compute their trophic levels using the classification of diet items and trophic level in Table 1 .

Food and feeding habits related topics covered in FishBase:


Trophic Ecology: See Box 18 and links to information on diet composition, food items, predators, daily ration and food consumption in TROPHIC ECOLOGY.
See Box 20 in The ECOLOGY Table
See Box 21 in The FOOD ITEMS Table
See Figs. 33-34 and Boxes 22-23 in The PREDATORS Table
See also Box 12. Mean size of fish in fisheries catches
[Note: information contained in the Ecology table is available only in the CD-ROM version of FishBase. However, some ecological information are available in the Species Summary page under the Biology and Environment fields and in the Key Facts page. To make a list of species with such information, use the Information by topic search in, select the Ecology button and jot down the scientific name(s) of the species of interest. Then open the Species Summary and Key Facts page for that species.]

5.     Reproduction

5.1.                      The reproductive load concept

Fish usually reproduce when they have reached about half of the maximum size they are likely to reach (Lmax). The size at which maturity is first reached is called Lm and the fraction Lm/Lmax, called reproductive load, tends to be higher in small than in large fish. Thus, a goby with Lmax=10 cm will have a value of Lm = 7 cm, while in a Basking shark with Lmax » 10 m, Lm will be about 4 m. Given that fish of different sizes have different growth rates, their different Lm values imply very different ages at first maturity (tm).

5.2.                      Small eggs and no worries

Fish differ from most other vertebrates in that for most species, parental care is very limited or non-existent. The typical bony fish produces a large number of small eggs which hatch and become part of the phytoplankton, and which must beware of their parent (or their congeners) if these are zooplankton feeders.

The high fecundity of bony fish has led many to believe that they can be exploited very strongly, i.e., that there will always be some recruits even if the parental stock is much reduced. This is called the ‘million egg fallacy’ and it has caused untold damage to fisheries, especially cod fisheries. Still, it is useful to know the relationship between numbers of eggs spawned and the weight of the mothers.

5.2.1.      Exercise 8

Task for the Student:
  • Identify in FishBase, a given species with a number of estimates of eggs in the ovaries of females of different sizes, and establish a fecundity/length relationship. Using that relationship and a length-weight relationship, calculate the relative fecundity (eggs/body-weight) of a female at 95% of its maximum size and compare that with the relative fecundity of a female at first maturity, i.e., near 60% of Lmax.
  • Relate the finding (above) to the exponent of the maturity/length relationship.

Reproductive load related topics covered in FishBase:


Reproductive load: See Figs. 36-37 and Box 25 The MATURITY Table; see item (h) under Fields in The POPGROWTH Table; see also Fecundity related fields in The SPAWNING Table
Use the Key facts (from the Key facts link in the Species Summary page or from the Scientific name search selecting the Key facts button) and/or Species Summary pages to obtain more information on reproduction.
In the Key Facts page, click on the Growth & mortality data link and in the resulting list, click the Reproductive load graph link to view a plot of Lm/L vs. L. This graph is also available in the Species Summary, through the Growth link.
Egg sizes: ICHTHYOPLANKTON and see Egg diameter field in The EGGS Table; refer to Exercise 9 .

5.3.                      Large eggs and parental investment

There are many fish which give birth to live young or which construct nests for their eggs, or which practice buccal incubation, e.g., in the Nile tilapia (see also Fish Quiz link in Exercise 7). Some other fish, notably the cartilaginous sharks and rays, give birth to fully-formed pups or produce very large eggs from which fully-formed young are hatched.

5.3.1.      Exercise 9

Task for the Student:
  • Write a one-page essay on why most fish broadcast their eggs and exert no parental care, given the fact that parental care reduces the mortality of the young and is practiced by numerous successful groups.

Reproductive strategies (spawning behavior) related topics covered in FishBase:


Reproduction: From click the Fish Quiz button. Play the Biodiversity quiz and test your knowledge of ‘guarding’ by fishes.
See discussions under REPRODUCTION and see Reproductive guild field in The REPRODUCTION Table
In the Species summary page, click on Reproduction and/or Spawning.
To list species with reproduction and spawning information in FishBase, use the Information by topic search in and click on the Reproduction or Spawning options. Click on a species to obtain more detailed information on these topics.
Egg size: See Egg diameter field in The EGGS Table; see also discussions in ICHTHYOPLANKTON
In the Species summary page, click on Eggs then Meristic characters.
To list species with egg and larvae information in FishBase, use the Information by topic search in and click on the Eggs, Egg dev’t. or Larvae options. Click on a species then click on Meristic characteristics to obtain information on egg size.

5.4.                      Variations on the basic theme

As noted by Darwin, fish are extremely labile in their sex determination, i.e., there are lots of fish which change sex, at least, far more than in other vertebrate classes (e.g., wrasses, parrot fishes, groupers). These are called hermaphrodites. In some fishes the different life (and sex) stages differ so much in color and/or form that they were originally described as different species. Fish also give us neat examples of parasitic males, and other aberrant (?) behaviors.

5.4.1.      Exercise 10

Task for the Student:
  • Give one example of a hermaphroditic species where subsequent development phases look very different.
  • Write a one-page essay about the different forms of hermaphroditism that exist and their distribution among fish families, and latitudinally.
  • Write a one-page essay on the group(s) in which parasitic males occur and give possible reasons for their preponderance among these groups.

Reproductive strategies (sex change) related topics covered in FishBase:


Hermaphroditism: See discussion on Mode field and pay particular attention to Fig. 35 and Box 24 in The REPRODUCTION Table
See Biology field in Species summary page for the mangrove rivulus and click on the Reproduction link for more information.
To list species with information on reproduction in FishBase, use the Information by topic search in and click on the Reproduction option.

6.     Physiology

The basic building blocks of fish bodies are proteins. Proteins have structure at several levels. The primary structure is determined by a sequence of the component amino acids, themselves with a structure determined by their sequence of atoms of carbon, hydrogen, etc. The secondary structure of most protein is a primary coil, similar to a braid.  A third-level structure can emerge when the braids fold onto themselves, with various loops weakly connected by hydrogen bonds. It is this tertiary structure which determines the external shape of a protein, e.g. of an enzyme and hence how it will lock into ‘receptors’, often other molecules on the surface of cells.

6.1.                      Metabolism, gills and size

Thermal noise is ubiquitous above absolute zero (0 Kelvin) and one of its effects is to destroy the tertiary structure of protein, thus rendering it ineffective. As a result, animals must break down such denatured molecules into their constituent parts and re-synthesize them. This is the reason why it costs energy to maintain a living body, even when it ‘does’ nothing, nor grows. In mammals and birds, which maintain more or less constant internal body temperatures, enzyme systems are geared such that the rate of synthesis matches a certain level of thermal noise, i.e. that which occurs at 37 to 38 degrees temperature. In fish, which except for large scombroids and some large sharks, cannot maintain a constant body temperature, different external temperatures thus imply different levels of thermal noise and hence rates of protein denaturation. Thus, metabolic rate must vary with temperature and it does so essentially in function of the need to re-synthesize protein.

However, it must be understood that the oxygen consumed by a fish is not its oxygen demand but the oxygen supplied to it via its gills, i.e., the fish would use more oxygen if it could get it. Hence, the amount of oxygen consumed by a fish is an imperfect measure of its real ‘need’ for oxygen. Gill size grows in proportion to a power of body weight that is less than one, i.e., the bigger fish of a given species become, the smaller the gill area per body weight becomes. Hence, big fish, given a certain level of activity, will tend to run out of oxygen faster than small fish of the same species, other things being equal.

6.1.1.      Table 2

Ten species in FishBase with growth parameters, at least one length-weight relationship and three records each of gill area and oxygen consumption per unit body weight.

Common name

Scientific name

Sea trout

Salmo trutta trutta


Carassius auratus auratus

Blackfin icefish

Chaenocephalus aceratus


Platichthys flesus


Stizostedion vitreum


Cirrhinus mrigala

Skipjack tuna

Katsuwonus pelamis


Tinca tinca

Common carp

Cyprinus carpio carpio


Rutilus rutilus

6.1.2.      Exercise 11

Task for the Student:
  • Choose a species from Table 2. Estimate for that species the exponent of a log-log relationship between gill area and body weight, and between oxygen consumption and body weight, and plug into this equation the value for the maximum size reported for that fish in a given habitat. [Tip: maximum lengths by locality are found using the Max. size & age link in the Species Summary page.]
  • Compute the gill area per unit weight and oxygen consumption per unit weight at which the fish stops growing. [Tip: Lmax and L¥ may have something to do with this.]

Metabolism related topics covered in FishBase:


Gill area and size: See Figs. 47-48 in  The GILL AREA Table; The OXYGEN Table; Fig. 51 and Boxes 27-28 in The GENETICS Table
To list species with gill area information in FishBase, use the Information by topic search in and click on the Gill area option. Click on the species of interest then click on the Gill area vs body weight graph link.

6.2.                      Food consumption

Like other heterotrophic organisms, fish need food to survive and grow. Within ecosystems, trophic (feeding) relationships and energy flows largely define the function of various species. There are two ways of presenting species-specific consumption:

·        At the individual level, i.e., as the consumption of a particular food type by a fish of a certain size, in the form of a daily ration (Rd); or

·        At the population level, i.e., as the consumption (Q) by an age-structured population of weight (B), in the form of population-weighted consumption per unit biomass (Q/B).

There are a number of methods that can be used to estimate the daily ration of fish: studying the changes in stomach content in the course of a day, direct observation of captive fish, etc. One of these techniques is to infer ration from daily oxygen consumption, which is justified since the oxygen consumed is ultimately combined with the food consumed to generate ATP (adenosine triphosphate, the substance used to fuel internal metabolism). This is illustrated through an example for red piranha, Pygocentrus nattereri, adapted from Pauly (1994):

Data were analyzed using a multiple (log) linear regression which yielded, for prediction of the metabolic rate (C, in mg02 · h-1) in small Pygocentrus nattereri, the model

C = 0.387 · W0.539· O21.13,                                                                                            … 1)

where W is the live weight of the fish in g, and O2 is the oxygen content of the water, in mg 1-1. The overall fit is good (R = 0.950); the standard errors of the exponents are 0.163 and 0.205, respectively, for 4 degrees of freedom. Given the small range of weights considered here, the relatively large standard errors about the estimates, and the low number of degrees of freedom, it would not be appropriate to assume that the slope linking O2 consumption and body weight is, in P. nattereri, significantly different from that proposed by Winberg (1960) for most fishes larger than guppies, i.e., 0.7 - 0.8. This implies that the equation above can be used only for a small range of weights, here 20 to 160 g.

For a 100 g fish in water with 6 mg O21-1, the equation above predicts an O2 consumption of 35 mg·h-1, i.e., 841 mgO2 ·day-1. An estimate of daily energy consumption (Q) can be obtained from this using the approach of Wakeman et al. (1979), wherein

Rd = (
DW + RESP)/0.75,                                                                                           … 2)

where Rd is the ration, i.e., daily energy consumption in kcal,
DW the energy content of the (daily) growth increment, and RESP is the oxygen consumption.

The first derivative (i.e. growth rate) of the von Bertalanffy equation in terms of wet weight is

dw/dt = 3KW ((W
¥/W)1/b-1)                                                                                       … 3).

This, solved for W
¥ = 756 g, K = 0.893/365 = 0.00245 day-1, and b = 3, gives for a 100 g fish a daily growth increment of 0.706 g, corresponding to 0.706 kcal if the calorific value of fish wet weight is set equal to unity (Brett & Blackburn 1978). The available information on body composition of red piranha flesh (Junk 1976, in Smith 1979) is 8.2 % fat, 15.0 % protein, and 4.4 % ash, not very different from values reported from other fishes (Bykov 1983). Thus, if an oxycaloric equivalent of 0.00325 kcal·mg-1 O2 is assumed, as in other fishes (Elliot and Davidson 1975), the above estimate of 841 mg O2 day-1 becomes 2.733 kcal day-1. Thus,

Rd = 0.706 + 2.733/0.75                                                                                             … 4)

or 4.585 kcal day-1 for a 100 g piranha. Food conversion efficiency (K1 = (dw/dt)/ Rd ; Ivlev 1966) would then be K1 = 0.154.

6.2.1.      Exercise 12

Task for the Student:
  • Compute for species in Table 2, the gill area per unit weight and oxygen consumption used only for maintenance. [Tip: fish cease growing when they approach W¥ and conversion between total and fork length can be done from a picture.]

Ration related topics covered in FishBase:


Daily ration: The RATION Table and see links to food consumption in TROPHIC ECOLOGY
In the Species Summary page, click on the Ration link for more information.
To list species with daily ration information in FishBase, use the Information by topic search in and click on the Ration option.


6.3.                      Estimating food consumption from empirical models

The method outlined above to deal with the ration of fishes lead to point estimates, pertaining to a single size or age (group). A fish population consists, however, of different size (age) groups, with small sizes and ages being far more abundant than large sizes and ages. Thus, drawing inferences from one (or several) ration estimate(s) pertaining to a given size (range) of fish, to a population containing a multitude of size groups, requires a knowledge of the size (age) structure of the population. An approach for performing this inference is given in FishBase.

A large number of such inferences, from ration to population weighted food consumption estimates (Q/B), have been performed in recent years, notably Palomares and Pauly (1998). These estimates of Q/B can be used in the context of empirical models to predict Q/B from other, easy-to-estimate parameters. One such equation is

log Q/B = 7.964 – 0.204logW¥ – 1.965T’ + 0.083A + 0.532h + 0.398d                     … 5)

where Q/B is the food consumption, W¥ is the asymptotic weight in grams, T’= 1000/(°C+273.15), A is the aspect ratio of the caudal fin = h2/s, h=1 and d=0 for herbivores, h=0 and d=1 for detritivores, and h=0 and d=0 for carnivores.

Here, one key input is the aspect ratio of the caudal fin defined as in Figure 1 . Fish with tails with high aspect ratio consume more food than fish with low aspect ratio tails, other things being equal. Needless to say, equation (5) above cannot be used for fish (e.g. eels) which do not use their caudal fin as their main propulsive organ. Other approaches can be used in such cases.

6.3.1.      Figure 1


6.3.2.      Exercise 13

Task for the Student:
  • Identify through FishBase, pictures of 3 species of fish covering a wide range of caudal fin aspect ratio: one with an aspect ratio of around 1; one with an aspect ratio of around 3-4, and one with an aspect ratio of above 7. [Tip: copy a square grid on a transparency and count the number of square units or cells occupied by the caudal fin to estimate the fin area.]
  • Use the aspect ratio, the body size, and the temperature of the habitat to infer Q/B given equation (5) above, and: (a) a herbivorous diet; or (b) a carnivorous diet. [Tip: the equation is also implemented in the Key facts page.]

Food consumption related topics covered in FishBase:


Food consumption:
To get a list of species with food consumption information, use the Information by topic search in and select the Food consumption button.
Or look for a particular species using the Scientific name search and select the Key Facts or Species Summary, click on the Food consumption link to get more information on food consumption.

7.     Fish as part of exploited ecosystems

7.1.                      Food webs and trophic levels

Fish populations do not live by themselves. Rather, they are embedded in ecosystems where they perform their roles as consumers and prey of other organisms, including larger fishes.

7.2.                      Trophic levels and sizes of fish

The role of fishes within ecosystems is largely a function of their size: small fish are more likely to have a vast array of predators than very large ones. On the other hand, various anatomical and physiological adaptations may lead to dietary specialization, enabling different fish species to function as herbivores, with a trophic level of 2.0, or as carnivores, with trophic levels typically ranging from 3.0 to about 4.5.

Moreover, trophic levels change during ontogeny of fishes. Larvae, which usually feed on herbivorous zooplankton (TL= 2.0) consequently have a trophic level of about 3.0. Subsequent growth enables the juveniles to consume larger, predatory zooplankton and small fishes or benthic invertebrates; this leads to an increase in trophic level, often culminating in values around 4.5 in purely piscivorous, large fishes.

7.3.                      Formal description of food webs

For formal descriptions of the role of fish in ecosystems and their responses to changes in fishing, and other changes, see the Ecopath modeling tool at

There is a strong link between Ecopath and FishBase, i.e., FishBase has a special routine to assemble and print out information on the fish of a given area or ecosystem, such that ecosystem models can be straightforwardly constructed.

7.3.1.      Exercise 14

Task for the Student:
  • Use the diet composition data previously standardized (see section 2.5 Diversity of food and feeding habits, above) and calculate the trophic level implied by the diet, given the prey trophic levels (Table 1) of FishBase.
  • Assemble diet composition studies for different sizes of the same species of fish, preferably in the same population, and show trophic level changes with ontogeny.
  • Identify a marine ecosystem that interests you and run the FishBase routine which extracts information on the fish of that and similar (or adjacent) ecosystems. Use this information to draw a food web incorporating the diet information on major fish and invertebrates in that ecosystem.

Food web related topics covered in FishBase:


Food webs: see Box 18 in; see also Box 20 in
Ecopath parameters: Use the Information by topic search in and select the Ecopath parameters button to get a list of species ordered by habitat type and size. The list indicates for which of these species Ecopath related parameters, i.e., growth parameters, Q/B, diet and predator information, are available. It is also possible to output the list as an Excel file as described in Exercise 1 .
Trophic levels & catches: In the Species Summary page, click on link to FAO stats for information on mean trophic levels and catches. [Note: FAO catches is only available in the CD-ROM version. To list species with available catch data in the FishBase CD-ROM, use the Information by topic search and click the FAO catches option. Or search the FAO Fishery Statistics online database at]
Diets: In the Species Summary page, click on links to Diet and Predators. [Note: links to Diet and Food consumption are also available in the Key facts page.]
To list species with available diet data in FishBase, use the Information by topic search and click on the Diet option.

8.     Contributing to FishBase

The FishBase project is a large, international, non-profit venture which started in 1989 and whose latest product, FishBase 99 (Froese and Pauly 1999) covers all the fish in the world - at least in terms of nomenclature. In terms of biology and ecology the coverage is, however, rather spotty and it is paradoxically in the well-studied temperate areas that the coverage is most incomplete, at least relative to the available literature. The reason for this is that FishBase was funded by the European Commission to cover countries in Africa, the Caribbean and the Pacific (‘ACP’) that are associated with the European Union.

For FishBase to realize its potential as the integrated, computerized system of fish most useful to the global ichthyological community, it requires input from users, including students. Thus you are encouraged to contribute to FishBase, notably by sending reprints or photocopies of material used for your analyses, as well as other information which you think should be incorporated (with complete sources!). You are also welcome to submit photos or slides whose originals would be returned to you after they have been scanned. See ‘How to become a collaborator and why’ for details on the manners in which such contributions are acknowledged; also note that you retain all rights to any submitted photo).

9.     Acknowledgements

We wish to thank Ms. Donna Shanley, for creating a first workable draft out of a jumble of text notes, references and Internet URLs. Without her dedication and skill, the making of this guide would have continued to be postponed forever.

10.             References

Berg, L.S. 1965. Freshwater fishes of the U.S.S.R. and adjacent countries. In 3 Vol., 4th edition. Israel Program for Scientific Translations Ltd., Jerusalem. [Original Russian version published in 1949].

Berlin, B. 1992. Ethnobiological classification. Princeton Univ. Press, New Jersey. 335 p.

Bykov, V.P. 1983. Marine fishes: chemical composition and processing properties. Amerind Publishing Co., New Delhi. 322 p.

Brett, J.R. & J.M. Blackburn. 1978. Metabolic rate and energy expenditure of the spiny dogfish, Squalus acanthias. J. Fish. Res. Board Can. 35: 816-821.

Cuvier, G. 1828. Historical portrait of the progress of ichthyology, from its origin to our own time. Translated by A.J. Simpson and edited by T.W. Pietsch (1995). The Johns Hopkins University Press, Baltimore. 366 p.

Darwin, C. 1842. The Essay of 1842 p. 1-53 In: F. Darwin (ed.). The Foundation of The Origins of Species: two essays written in 1842 and 1844. Cambridge, 1909. [Vol. 10 of ‘The Works of Charles Darwin’. Pickering & Chatto, London, 199 p.]

Eastman, J.T. and A.L. Devries. 1985. Adaptations for cryopelagic life in the Antarctic notothenioid fish Pagothenia borchgrevinki. Polar Biol. 4:45-52.

Elliott, J.M. and W. Davidson. 1975. Energy equivalents of oxygen consumption in animal energetics. Oecologia 19: 195-201.

Eschmeyer, W.N. (Editor). 1998. Catalog of fishes. California Academy of Sciences, San Francisco. 3 vols. 2905 p.

Fryer, G. and T.D. Iles. 1972. Cichlid Fishes of the Great Lakes of Africa. Oliver & Boyd, Edinburg, UK.

Froese, R. and D. Pauly (Editors). N. Bailly and M.L.D. Palomares (Translators). 1999. FishBase 99: Concepts, structure, et sources des données. ICLARM, Manila, Philippines. 324 p.

Groot, G. and L. Margolis. Editors. 1991. Pacific Salmon Life Histories. University of British Columbia, Vancouver, Canada. 564 p.

G.S. Helfman, B.B. Collette and D.E. Facey. 1997. The diversity of fishes. Blackwell Science, MA. 512 p.

Ivlev, V.S. 1966. The biological productivity of waters. [translated by W.E. Ricker]. J. Fish. Res. Board Can. 23: 1717-1759.

Junk, W.J. 1976. Biologia de água doce e pesca interior, p. 105. In: Relatorio Anual de INPA, Instituto Nacional de Pesquisas da Amazonia, Manaus.

Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae secundum Classes, Ordinus, Genera, Species cum Characteribus, Differentiis Synonymis, Locis. 10th ed., Vol. 1. Holmiae Salvii. 824 p.

Lakoff, G. 1987. Women, fire and dangerous things: what categories reveal about the mind. University of Chicago Press, Chicago, 614 p.

Linnaeus, C. 1758. Systema naturae per regna tria naturae secundum classes, ordinus, genera, species cum characteribus, differentiis synonymis, locis. 10th ed. Vol. 1. Holmiae Salvii. 824 p.

Mayr, E. 1942. Systematics and origin of species. Columbia University Press, New York. 334 p.

Nelson, J.S. 1994. Fishes of the world. 3rd edition. John Wiley and Sons, New York. 600 p.

Palomares, M.L.D. and D. Pauly. 1998. Predicting food consumption of fish populations as functions of mortality, food type, morphometrics, temperature and salinity. Mar. Freshw. Res. 49:447-453.

Palomares, M.L.D., C.V. Garilao and D. Pauly. 1999. On the biological information content of common names: a quantitative case study of Philippine fishes, p. 861-866. In: B. Séret & H,-Y. Sire (eds.) Proc. 5th Indo-Pac. Fish Conf., Nouméa, 1997.

Pauly, D. 1994. Quantitative analysis of published data on the growth, metabolism, food consumption, and related features of the red-bellied piranha, Serrasalmus nattereri (Characidae). Environ. Biol. Fish. 41:423-437.

Pietsch, T.W. and D.B. Grobecker. 1987. Frogfishes of the world. Stanford University Press, Stanford, California. 420 p.

Randall, J.E. 1998. Zoogeography of shore fishes of the Indo-Pacific region. Zoological Studies 37(4):227-268.

Smith, N. 1979. A pesca no rio Amazonas. Instituto Nacional de Pesquisa da Amazonia, Manaus.

Wakeman, J.M., C.R. Arnold, D.E. Wohlschlag and S.C. Rabalais. 1979. Oxygen consumption, energy expenditure and growth of the red snapper (Lutjanus campecheanus). Trans. Amer. Fish. Soc. 108: 288-292.

Walbaum, J. J. 1792. Petri Artedi Sueci Genera piscium. In quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Redactis speciebus 242 ad genera 52. Ichthyologiae, pars iii. Artedi Piscium 1-723.

Winberg, G.G. 1960. Rate of metabolism and food requirements of fishes. Fish. Res. Board Can. Transl. Ser. (194). 239 pp.

Zahavi, A. and A. Zahavi. 1997. The handicap principle: a missing piece of Darwin's puzzle.  Oxford University Press. 304 p.


11.              Appendix A: Web sites of institutions/organizations with ichthyological collections and/or associated links

This site includes a page with links to Museums and Collections world-wide: (click on museums)

Albany Museum - (Freshwater Ichthyology):

American Museum of Natural History: (no link to Department of Herpetology & Ichthyology, however link to Center for Biodiversity Conservation available at

Aquatic animals mailing lists maintained by individuals from various institutions

Australian Museum – Ichthyology:

BBCWS - Biodiversity and Biological Collections Web Server:

Bernice P. Bishop Museum - Ichthyology Department:

Burke Museum, University of Washington, Seattle

California Academy of Sciences - Ichthyology Department:

Canadian Museum of Nature - Research and Collections - Recherche et collections: or

Carnegie Museum of Natural History, Pittsburgh:

Chicago Academy of Sciences Nature Museum:

Coleccion Nacional de Peces (Mexico):

FishGopher - The FishGopher Project:

Florida Museum of Natural History - Ichthyology Page:

Grice Marine Laboratory, University of Charleston, SC:

Humboldt State University Fish Collection:

Illinois Natural History Survey - Fish Collection:

Instituto Nacional de Pesquisas da Amazônia - Neodat page:

James Ford Bell Museum of Natural History - Fish Collection: (database still under construction)

J. L. B. Smith Institute of Ichthyology:

Massachusetts Museum of Natural History - Fish collection:

Milwaukee Public Museum - Vertebrate Zoology Section:

Museu de Ciências e Tecnologia PUCRS - Laboratório de Ictiologia:

Museu Nacional, Universidade Federal do Rio de Janeiro - Setor de Ictiologia:

Muséum d'histoire naturelle de la Ville de Genève - Catalogue des types:

Muséum National D'Histoire Naturelle, Paris – Search:

Museum of Comparative Zoology, Harvard University - Fish Page:

Museum of Southwestern Biology, University of New Mexico - Division of Fishes

National Museum of Natural History, Smithsonian Institution - Division of Fishes

Natural History Museum (London) – collections:

Natural History Museum of Los Angeles County: (click on Ichthyology; note that collection database still not available)

Naturhistorisches Museum der Burgergemeinde Bern - (Vertebrate animals - collections): (under E.A. Göldi collection, click on Gopher, then on Search Pisces collections; note that this is a downloaded file and may take more than 5 minutes)

Naturhistoriska Riksmuseet (Swedish Museum of Natural History) – Collections, Ichthyology Section, Department of Vertebrate Zoology:

Nova Scotia Museum (click Backstage, then Collections, note that data computerization is incomplete and the collection is not yet searchable)

NEODAT - Neotropical Biodiversity Database:

Oklahoma Museum of Natural History - Collections and Research: (Division of Ichthyology is not on line)

Ohio State University’s Museum of Biological Diversity: (internal link to Museum of Zoology currently not available)

Philadelphia Academy of Natural Sciences:

Provincial Museum of Alberta - Ichthyology Program – Holdings:

Royal British Columbia Museum - Ichthyology and Herpetology Collection:

Royal Ontario Museum - Centre for Biodiversity and Conservation Biology:

Santa Barbara Museum of Natural History:

Scripps Institute of Oceanography - Marine Vertebrates Search:

Staatliches Museum fur Naturkunde, Stuttgart, Germany:

Texas Natural History Collection (Austin) - Ichthyology Division:

The Field Museum of Natural History, Chicago, Illinois - Fish collection:

Tulane University Museum of Natural History – Fishes:

University of Alabama Ichthyological Collections:

University of Alaska Museum, Fairbanks - Aquatics collection:

University of Alberta Laboratory for Vertebrate Paleontology:

University of Alberta Museum of Zoology – Collections:

University of Arizona Zoological Collections - Fish Collection:

University of Arkansas Museum, Fayetteville:

University of California Museum of Paleontology - Vertebrate Collection:

University of Georgia Museum of Natural History: (Icthyology collection not online).

University of Kansas Natural History Museum - Division of Ichthyology: (click on Collections)

University of Michigan Museum of Zoology - Division of Fishes:

University of Nebraska State Museum - Division of Zoology

University of Washington Fish Collection

Virginia Institute of Marine Science - Ichthyology Collection

Western Australia Museum - Aquatic Zoology

Yale University Peabody Museum of Natural History - Ichthyology Collection

Zoological Museum, University of Copenhagen - Ichthyological Section

Zoologisches Museum, Universität Hamburg - Ichthyological Collection

12.             Appendix B:

University of Alberta Ichthyology Web Resources:

The University of British Columbia Library:

Aquatic Sciences and Fisheries Abstracts:


[1] Draft of December 1999. To be tested in the Fish 445 class, January-March, 2000, UBC, Vancouver;

[2] Fisheries Centre, 2204 Main Mall, University of British Columbia, Vancouver, B.C., Canada, V6T 1Z4 e-mail: ;

[3] FishBase Project, 3rd floor Collaborator’s Center, IRRI, Los Baños, Philippines; e-mail:,

[4] The FishBase project leader (Rainer Froese; e-mail: would appreciate being informed of plans for such projects, which may lead to new information or new tables being added to future versions of FishBase (see also Contributing to FishBase below).


Last modified by Eli, 17.01.05   (