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Fauna of New Zealand 68 - Landcare Research

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28<br />

larva. Another pattern is for larvae to form a row with larvae<br />

side-by-side (Chance & Craig 1986; Ciborowski & Craig<br />

1989; Eymann 1991). This is known to increase the velocity<br />

<strong>of</strong> water between the larvae, and thence through the fans,<br />

would increase particle flux (Ciborowski & Craig 1989),<br />

and was suggested as a strategy for use when food levels<br />

were low. Although not illustrated in this work, that is a<br />

common pattern for larvae <strong>of</strong> Austrosimulium species, such<br />

as A. unicorne and A. bicorne, which occur at low density<br />

under stones (DAC & TKC pers. obs.). Both these species<br />

occur at high altitude in pristine waters, where concentrations<br />

<strong>of</strong> particulate matter appear to be low.<br />

Colbo & Moorhouse (1979) noted that at high population<br />

levels larvae <strong>of</strong> A. bancr<strong>of</strong>ti and A. pestilens formed<br />

clumps on stones for the former and on vegetation for the<br />

latter, echoing earlier comments on those two species by<br />

Mackerras & Mackerras (1948). All other species formed<br />

spaced distribution. In general, microdistribution <strong>of</strong> A.<br />

bancr<strong>of</strong>ti larvae depended on age <strong>of</strong> larva, water velocity,<br />

and nature <strong>of</strong> the substrate. Horne et al. (1992) showed that<br />

larvae <strong>of</strong> A. furiosum were normally found in velocities<br />

<strong>of</strong> 0.2–0.3 m/s, whereas Simulium ornatipes Skuse in the<br />

same habitat in velocities <strong>of</strong> 0.9–1.3 m/s. Austrosimulium<br />

furiosum (Skuse) larvae were particular about the choice<br />

<strong>of</strong> microhabitat and these velocity preferences were taken<br />

as evidence for microhabitat partitioning — this difference<br />

is important when benthic sampling to ensure accurate<br />

results are obtained. For <strong>New</strong> <strong>Zealand</strong> Austrosimulium,<br />

even for dense populations, such as occur at times with A.<br />

tillyardianum, larvae are always spaced.<br />

Where velocities could be determined during this study,<br />

larvae were rarely taken at velocities below 0.3 m/sec or<br />

above 0.8 m/sec for stone-loving species (A. tillyardianum,<br />

A. multicorne) and there was a sharp cut-<strong>of</strong>f below and<br />

above these limits (see Appendix 1). The upper limit appears<br />

to relate to shear stress <strong>of</strong> the water on the substrate<br />

since that was where even a thin layer <strong>of</strong> periphyton was<br />

scoured <strong>of</strong>f the substrate. This can be seen in Fig. 483<br />

(Kowhai River, Kaikoura) where larvae were only taken<br />

from the narrow dark band <strong>of</strong> stones along the edge — those<br />

with a thin layer <strong>of</strong> periphyton. Higher velocities were<br />

recorded for species on vegetation, but it was not possible<br />

to determine the exact velocity immediately proximate to<br />

the larvae. Nothing is known regarding velocity tolerances<br />

for species (e.g., A. unicorne, A. bicorne) living under<br />

perched stones.<br />

Jowett (2000) gave the mean depth for larvae <strong>of</strong> an<br />

Austrosimulium sp. in a small stream as 0.08 m and the<br />

mean velocity as 0.18 m/s, the latter low by our observations.<br />

He also quantified that larvae in small streams prefer<br />

higher velocity than that <strong>of</strong> the mean velocity, and we agree<br />

fully. This phenomenon allows rapid assessment <strong>of</strong> a locality<br />

for the likely presence <strong>of</strong> simuliid larvae — substrates<br />

Craig, Craig & Crosby (2012): Simuliidae (Insecta: Diptera)<br />

in higher velocity are examined first.<br />

Quinn & Hickey (1990a, b) in a <strong>New</strong> <strong>Zealand</strong>-wide<br />

survey <strong>of</strong> 88 rivers showed that maximum density <strong>of</strong><br />

unidentified Austrosimulium larvae was to be found on<br />

substrates with sizes up to large cobble (128–256 mm<br />

diameter). A rationale for that was the increased complexity<br />

and provision <strong>of</strong> a three-dimensional habitat; aeration<br />

is also improved. In an earlier study Pridmore & Roper<br />

(1985) examined macroinvertebrates in runs and riffles <strong>of</strong><br />

three North Island streams. They showed statistically that<br />

unidentified Austrosimulium spp. were significantly more<br />

abundant in riffles than runs in the Rangitukia Stream.<br />

Although in farmland, the stream substrate was andersite<br />

and basalt cobble usually favoured by A. tillyardianum,<br />

and also within its distributional range.<br />

Boothroyd & Dickie (1991), in an examination <strong>of</strong><br />

colonisation <strong>of</strong> artificial substrates by aquatic invertebrates,<br />

showed that while chironomids dominated the drift<br />

and substrates, A. australense could at times comprise<br />

up to 49% <strong>of</strong> that fauna. Drifting by simuliid larvae is a<br />

well known phenomenon (Crosskey 1990) and the initial<br />

dominance by simuliids on fresh substrates is in keeping<br />

with other studies. Similarly, Death (1996, 2000) dealt<br />

with colonisation <strong>of</strong> aquatic invertebrates after substrate<br />

disturbance. Austrosimulium larvae colonised readily, but<br />

were eventually replaced by more slowly colonising species.<br />

This is in full agreement with Harding et al. (2000)<br />

and Suren (2000), with Austrosimulium dominating in<br />

streams after disturbances by forestry, and their occurrence<br />

in urban milieu. In his study on A. tillyardianum Crosby<br />

(1970) constructed a stream deviation having a substrate<br />

<strong>of</strong> small stones, and found that larvae colonised the new<br />

substrate in 3–4 days.<br />

Collier (1995) examined some 29 lowland waterways<br />

in Northland. While an unidentified Austrosimulium occurred<br />

at 19 <strong>of</strong> those sites, it was not dominant enough to<br />

determine habitat requirements. Collier et al. (1998) also<br />

examined physical parameters in relation to macroinvertebrate<br />

fauna in 20 lowland Waikato streams. Austrosimulium<br />

larvae occurred at all sites. In keeping with most other such<br />

studies, larvae were found in faster water, and a range <strong>of</strong><br />

physico-chemical parameters, such as water depth, dissolved<br />

oxygen, and temperature accounted for the greater<br />

part <strong>of</strong> larval densities on macrophytes. As usual for most<br />

such studies, the simuliid species was not identified, but<br />

was assuredly A. australense.<br />

Simuliid larvae are well known to form large concentrations<br />

at lake outlets, an explanation for which has been<br />

the availability <strong>of</strong> planktonic food material (Crosskey 1990,<br />

Adler et al. 2004). As part <strong>of</strong> a series <strong>of</strong> studies on lake<br />

outlets, Harding (1992) examined the physico-chemical<br />

parameters and invertebrate fauna <strong>of</strong> three lakes in Westland.<br />

There were marked differences between the inlet

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