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Chapter 4 • Zooplankton

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Chapter 4 • Zooplankton CHAPTER 4 • ZOOPLANKTON The animals making up the zooplankton are taxonomically and structurally diverse. They range in size from microscopic, unicellular organisms to jellyfish several metres in diameter (refer to Figure 1.2). Although all zooplankton are capable of movement, by definition none are capable of making their way against a current. By definition also, all zooplankton — indeed all animals and some micro-organisms — are heterotrophic. That is, they require organic substrates (as opposed to inorganic ones) as sources of chemical energy in order to synthesize body materials. Unlike plants, which carry out autotrophic production by utilizing solar energy to reduce carbon dioxide, animals obtain carbon and other essential chemicals by ingesting organic materials. Animal species differ in how their energy is obtained: some species are herbivores which consume plants; others are carnivores which are capable of eating only other animals; and some species are predominantly detritivores which consume dead organic material. Many animals, however, are omnivores with mixed diets of plant and animal material. Different types of zooplankton often are placed in categories which describe their diets. In addition to size categories and positions in food chains, zooplankton can be subdivided into classifications based on habitat (oceanic vs. neritic species; see Section 1.2) and taxonomy. They also form two categories depending upon the length of residency in the pelagic environment; holoplankton (or permanent plankton) spend their entire life cycles in the water column, whereas meroplankton are temporary residents of the plankton community. The meroplankton includes fish eggs and fish larvae (the adults are nektonic), as well as the swimming larval stages of many benthic invertebrates such as clams, snails, barnacles, and starfish. The more common types of holoplankton and meroplankton are described below in Sections 4.2 and 4.3, respectively. 4.1 COLLECTION METHODS Zooplankton larger than 200 /xm (refer to Figure 1.2) traditionally have been collected by towing relatively fine-mesh nets through the water column. Plankton nets vary in size, shape, and mesh size (Figure 4.1), but all are designed to capture drifting or relatively slow-moving animals that are retained by the mesh. The simplest nets are conical in shape, with the wide mouth opening attached to a metal ring and the narrow tapered end fastened to a collecting jar known as the cod end. This type of net can vary in length and diameter and in mesh size, and it can be towed vertically, horizontally, or obliquely through the desired sampling depths. Such a net will filter water and collect animals during an entire towing period. More sophisticated nets are equipped to be opened and closed at selected depth intervals, and a series Copyright © 1997. Elsevier Science & Technology. All rights reserved. & Technology. © 1997. Elsevier Science Copyright of such nets may be attached to a single frame to allow sampling of different discrete depths during a single towing operation. Analyses of the collected samples permit a more detailed picture of the vertical distribution of zooplankton within a particular area. Because many zooplankton migrate Figure 4.1 A 'Bongo' zooplankton sampler, consisting of duplicate plankton nets, being vertically during each 24-hour period, the time of sampling is also critical in Parsons, T. retrieveR., & Lalli,d C.o n(1997). boar Biologicald a researc oceanographyh vessel .: An introduction. Retrievedtrackin fromg http://ebookcentral.proquest.comthese changes in depth distribution. Created from ucsd on 2020-05-15 19:29:38. 75 The selection of a particular net depends upon the type of organisms desired and the characteristics of the water being sampled. For example, a fine-mesh net (with a mesh opening of ca. 100-200 /xm) obviously must be used to collect small mesozooplankton. However, the same net is not suitable for sampling larger, relatively fast-swimming zooplankton, like fish larvae, because it clogs quickly and must be towed slowly to avoid tearing. In deep water, where zooplankton tend to be larger and are less abundant, it is common to use a very large coarse-mesh net. All nets can be equipped with a flowmeter that estimates the total volume of water filtered during a tow; this permits a quantitative representation of the zooplankton collected. The newest towed samplers, such as the Batfish shown in Figure 4.2, simultaneously measure salinity, temperature, depth, and chlorophyll a concentration while counting and sizing zooplankton that pass through an optical sensor. Such devices can be set to undulate over a set depth range, thus making it possible to obtain samples at several depths. No single sampler is capable of capturing all zooplankton within its path. Zooplankton smaller than 200 /xm (nano- and microplankton) cannot be satisfactorily sampled in nets; instead, a known volume of water is collected in sampling bottles or by pumps from defined depths and the smallest zooplankton are concentrated and removed by filtration, centrifuging, or settling and sedimentation. Both planktonic protozoans and phytoplankton can be counted in the concentrated water samples. Figure 4.2 A towed Batfish plankton sampler which simultaneously estimates phytoplankton and zooplankton abundance while recording environmental parameters. F, fluorometer for detecting chlorophyll a; L, light sensor; OPC, optical zooplankton counter; PI, intake for zooplankton sampling; SB, stabilizer; STD, salinity-temperature-depth sensor; T, towing arm. Copyright © 1997. Elsevier Science & Technology. All rights reserved. & Technology. © 1997. Elsevier Science Copyright Parsons, T. R., & Lalli, C. (1997). Biological oceanography : An introduction. Retrieved from http://ebookcentral.proquest.com Created from ucsd on 2020-05-15 19:29:38. 76 Further, it is now recognized that some zooplankton are capable of avoiding towed samplers, which they detect either visually or by sensing turbulence created in advance of the moving gear. In addition, certain gelatinous plankton are so fragile that they are impossible to collect intact in nets; others disintegrate rapidly in the preservatives that are routinely used to store collections. Crustaceans usually comprise the majority of zooplankton in net collections because most are too small to avoid capture, and because their hard exoskeletons protect them from damage and distortion in nets and preservatives. Thus the numbers of Crustacea relative to other types of zooplankton may be overestimated, and net-collected plankton may not provide a true representation of the natural plankton community in many areas. The numerical dominance and biomass contribution of crustaceans, especially copepods, needs to be reassessed in many localities. Zooplankton can also be observed directly in the field, either by scuba diving down to depths of about 30 m (Figure 4.3) or, in deeper waters, by using manned submersibles or ROVs (remotely operated vehicles) which are tethered to ships and coupled with underwater video cameras of high resolution. These techniques have resulted in discoveries of new species, particularly of fragile forms; in an awareness of the problem of underestimating numbers and biomass of these animals from net collections; and in new behavioural observations of many species. Bioacoustic methods, Figure 4.3 A scuba diver counting the number developed from the use of sonar to locate fish schools, are also being apphed of zooplanl<ton (larvacea) contained within a to locate and estimate densities of larger zooplankton, such as euphausiids, frame of known dimensions. which form dense aggregations. 4.2 HOLOPLANKTON: SYSTEMATICS AND BIOLOGY There are approximately 5000 described species of holoplanktonic zooplankton (excluding protozoans) representing many different taxonomic groups of invertebrates (Table 4.1). Those groups that are commonly found in the sea and that form significant fractions of the plankton community are described below. In addition to providing descriptive anatomical accounts, particular attention is given to describing the food and feeding mechanisms of each group as these are important in the discussions of food webs and energy transfer which follow. The smallest of the zooplankton are certain unicellular protists (Table 4.1). Included are many species of dinoflagellates that are partly or wholly heterotrophic (see Section 3.1.2 for a discussion of autotrophic species). These heterotrophic dinoflagellates feed on bacteria, diatoms, other flagellates, and ciliate protozoans that are either drawn to the predator by flagella-generated water currents, or that are trapped in cytoplasmic extensions of the dinoflagellate. Some species are only capable of functioning as heterotrophs; others that contain chloroplasts may also function as autotrophs part of the time. The best known heterotrophic dinoflagellate is Noctiluca scintillans (Figure 4.4), which has the form of a gelatinous sphere 1 mm or more in diameter. Noctiluca often occurs in Copyright © 1997. Elsevier Science & Technology. All rights reserved. & Technology. © 1997. Elsevier Science Copyright dense swarms near coasts, and it feeds on small zooplankton (including fish eggs) as well as on diatoms and other phytoplankton. A taxonomically diverse group of flagellated protists, commonly called zooflagellates, includes all those species that are colourless and strictly Parsons, T. R., & Lalli, C. (1997). Biological oceanography : An introduction. Retrievedheterotrophic
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