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This review deals with annual production by the fresh-water animal groups: fishes, zoobenthos, and zoo-plankton. Included are the concept and terminologyof productivity, methods for estimating annual pro-duction, the production/biomass ratio, and levels ofproduction by the three groups from the world litera-ture. [The SC!5 indicates that this paper has been citedin over 180 publications.]

Animal Production in Fresh Waters

Thomas F. WatersDepartment of Fisheries & Wildlife

University of MinnesotaSt. Paul, MN 55108

November 7, 1988

In 1974, Professor Amyan Macfadyen, editorof Advances in Ecological Research, wrote tomeand suggested a review of secondary pro.duction measurement. I immediately agreedbecause, at that date, the subject of freshwaterproduction had been researched with some in-tensity for the past decade, and the first majoraccumulation of data had been done. The timeseemed particularly opportune to criticallysummarize methods, which had reached apoint of soundness and broad applicability,and to collect into one place most of theworlds production data. Another objectivewas to clarify the concept and terminology ofproduction, at that time in a bit of a mess inthe literature. I felt it would be helpful to alsoconvert the melange of units in the literatureinto a single expression for each of the threemajor groups: kg/ha/yr (wet) for fish, kg/ha/yr(dry) for zoobenthos, and g/m/yr (dry) for zoo-plankton.

The definitive beginning of aquatic produc-tion literature is usually credited to the paperof P. Boysen-jensent on a marine benthos.

But the first great stimulation, conceptuallyand mathematically sound, was that presentedindependently by W.E. Ricker2 and K.R.AlIen3what we now call the instantaneousgrowth rate (ICR) method for estimating fishproduction. Even so, a long lag occurred be-tween the mid-1940s and important further ad-vances; H.B.N. 1-lynes,4 in reviewing streamecology literature through 1966, consideredproduction research to be in its infancy. Byabout the mid-1960s, however, the Interna-tional Biological Program (lBP), emphasizingproductivity, had given great impetus to re-search on the subject, and a few landmark pa-pers had also appeared by that time. Further-more, a methodological breakthrough for ben-thos production was presented by Hynes5that now, after further development, we termthe size-frequency method.

By the mid-i 970s, then, a viable, decade-long literature had accumulated. Thus, I thinkthe reason for the frequent citation is that myreview served as a linchpin for further work,appropriately located in time. Subsequent au-thors could refer to the definition of produc-tion without giving a complicated explanationof their own; the methods were given in asomewhat recipe-book style and could bereadily referenced; and the literature data,summarized in tabular form with commonunits, could be easily examined.

No comparable review has been subsequent-ly published, although A.C. Benke reviewedin depth the concept and methods for benthos,and R.H.K. Mann and T. Penczak7 summa-rized production by river fishes. The revisedIBP handbook on secondary productivity infresh waters deals with a number of relatedsubjects.

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During the most recent decade, a large num-ber of additional estimates have been pub-lished, particularly on fishes and stream ben-thos. Methods used are basically the same buthave been fine-tuned in some critical respects.Methods used today most commonly appearto be the ICR method for fish and the size-fre-quency method for benthos.

This Weeks Citation Classic FEBR~JARY13,1989Waters I F. Secondary production in inland waters.

Advan.Ecol. Res. 10:91-164, 1977.[Department of Entomology, Fisheries and Wildlife, University of Minnesota. St. Paul, MNI

1. Boysen-JensenP. Valuation of the LimOord. I. Studies on Inc fish-food in the Limljord 1909-1917, its quantity, variationand annual production. Rep. Dan. BioL Sm. 26:344. 1919. (Cited 30 times since 1945.)

2. Picker WE. Production and utilization of fish populations. EcoL Monogr. 16:373-9!. 1946. (Cited 14t asses.)3. Allen K R. Some aspects of the production and cropping of fresh waters. Trans. Roy. Soc. N. Z. 77:222-8. 1949.

(Cited 20 times.)4. Hynes H B N. The ecology of running waters. Liverpool, England: Liverpool University Press. (1970) 1982. 440 p.

(Cited 745 times.)5. . The invertebrate faunaof a Welsh mountain stream. Arch. Hydrobtol. 57:344-88. 1961. (Cited 220 times.)

1See also: Hynes H B N. Citation Classic. (BarrettS T. comp.( Contemporaryclassicsin plant, animal, andenvironntental sciences. Pttiladelphia: IS! Press. 1986. p. 164.1

6. Benke A C. Secondary production of aquatic insects. (Resh V H & Rosenberg 0 M. eds.( The ecology of aquatic insects.New York: Praeger, 1984. p. 289-322. (Cited 20 tintes.(

7. Mann R H K & Penczak T. Fislt production in rivers: a review. Polish Arch. 1-fydrobiol. 33:233-47, 986.8. DownIng J A & RIgler F H, eds. A ntanaai on methodsfor the assessment of secondaryprodacfisify infresh wafers.

Oxford. England: Blaclraell Scientific. 984. 501 p.

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