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Mechanical disruption of seagrass in the digestive tract of the dugong

Mechanical disruption of seagrass in the digestive tract of the dugong

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<strong>Mechanical</strong> <strong>disruption</strong> <strong>of</strong> <strong>seagrass</strong> by <strong>the</strong> <strong>dugong</strong><br />

J. M. Lanyon and G. D. Sanson<br />

distributions, especially if <strong>the</strong>y are skewed. The change <strong>in</strong><br />

median particle size between 0 and 500 gr<strong>in</strong>ds is <strong>in</strong>dicative <strong>of</strong><br />

<strong>the</strong> overall effects <strong>of</strong> gr<strong>in</strong>d<strong>in</strong>g for each plant species. These<br />

values (=‘breakability <strong>in</strong>dex’) were compared.<br />

Results<br />

Size <strong>of</strong> mouthparts<br />

The occlusal surface area <strong>of</strong> cheek teeth ranged from 24 mm 2<br />

<strong>in</strong> a <strong>dugong</strong> aged o0.5 years to 485 mm 2 <strong>in</strong> an adult <strong>dugong</strong><br />

aged 21 years, and was significantly regressed aga<strong>in</strong>st skull<br />

length and age (F=32.12; d.f.=1, 51; Po0.0001; adj<br />

R 2 =0.36; n=52; Fig. 2). However, <strong>the</strong> coefficient <strong>of</strong> variation<br />

(CV) <strong>in</strong> occlusal surface area was high (CV=35.05),<br />

even when only adults were compared.<br />

The surface area <strong>of</strong> <strong>the</strong> lower horny pad ranged from<br />

901 mm 2 <strong>in</strong> <strong>dugong</strong>s aged 0.5 years to 6280 mm 2 <strong>in</strong> <strong>the</strong><br />

largest adults, with a mean <strong>of</strong> 4933 109 for adult <strong>dugong</strong>s.<br />

Horny pad surface area was highly significantly regressed<br />

aga<strong>in</strong>st body size (F=433.28; d.f.=1, 53; Po0.0001; adj<br />

R 2 =0.91; n=54) and followed an exponential growth<br />

curve aga<strong>in</strong>st age (Fig. 2). The CV for <strong>the</strong> horny pad surface<br />

area <strong>of</strong> adult <strong>dugong</strong>s (CV=12.73) was considerably lower<br />

than that for <strong>the</strong> dental occlusal surface area <strong>of</strong> adults. The<br />

absolute surface area <strong>of</strong> <strong>the</strong> horny pads was up to n<strong>in</strong>e times<br />

greater than <strong>the</strong> occlusal surface area <strong>of</strong> <strong>the</strong> cheek teeth.<br />

Digesta analysis<br />

Diet composition<br />

Twenty-seven <strong>of</strong> <strong>the</strong> 29 <strong>dugong</strong>s had fed on more than one<br />

<strong>seagrass</strong> genus and some on up to four or five. Seagrass<br />

genera recorded most frequently <strong>in</strong>cluded Halophila, Halodule,<br />

Cymodocea, Thalassia and Zostera. One <strong>dugong</strong> had<br />

<strong>in</strong>gested some Enhalus. The most frequently <strong>in</strong>gested genera<br />

Surface area (mm )<br />

7000<br />

6000<br />

5000<br />

4000<br />

3000<br />

2000<br />

1000<br />

0<br />

SA teeth<br />

Sa horny pad<br />

Satotal<br />

Log. (SA teeth)<br />

Log. (Sa horny pad)<br />

Log. (Satotal)<br />

y = 1102.9 Ln(x) + 1793.1<br />

R = 0.9309<br />

y = 1044.7 Ln(x) + 1674<br />

R = 0.9182<br />

y = 58.382 Ln(x) + 118.46<br />

R = 0.3648<br />

a<br />

0 5 10 15 20 25 30 35 40 45 50<br />

Age (years)<br />

Figure 2 Surface area (mm 2 ) <strong>of</strong> (a) functional cheek teeth, (b) lower<br />

horny pad and (c) total functional mouthparts (cheek teeth plus lower<br />

horny pad; all log-transformed) plotted aga<strong>in</strong>st age (years). L<strong>in</strong>es <strong>of</strong><br />

best fit <strong>in</strong>cluded.<br />

c<br />

b<br />

recorded were Halophila and Halodule. Two <strong>dugong</strong>s had<br />

fed on only one <strong>seagrass</strong> genus, Halodule. Ten per cent <strong>of</strong><br />

<strong>dugong</strong>s had identifiable algal fragments <strong>in</strong> <strong>the</strong>ir digesta.<br />

However, because digesta could only be identified from <strong>the</strong><br />

largest particle size classes, <strong>seagrass</strong>es that were more amenable<br />

to breakdown dur<strong>in</strong>g mastication (see below) would<br />

have been underestimated. Consequently, composition <strong>of</strong><br />

diet could not be exam<strong>in</strong>ed as a variable affect<strong>in</strong>g particle<br />

size breakdown.<br />

Stomach contents<br />

In general, <strong>the</strong> stomach contents were well macerated.<br />

However, <strong>the</strong>re was considerable variation <strong>in</strong> particle size<br />

distribution curves for <strong>the</strong> stomach contents <strong>of</strong> different<br />

<strong>dugong</strong>s. Figure 3 shows some representative stomach<br />

particle size distributions <strong>of</strong> <strong>dugong</strong>s <strong>of</strong> different ages.<br />

Median particle size ranged between 302.8 and 3454.3 mm,<br />

with a mean <strong>of</strong> 1241.8 141.98 mm (n=29; Table 1). The<br />

CV <strong>in</strong> median particle size (for stomachs pooled) was high<br />

at 62.6.<br />

The multiple regression model, with median particle size<br />

as <strong>the</strong> dependent variable, and total surface areas <strong>of</strong> <strong>the</strong><br />

cheek teeth and <strong>the</strong> horny pad as <strong>in</strong>dependent variables,<br />

accounted for only 1% <strong>of</strong> <strong>the</strong> variation, R 2 =0.01, with<br />

none <strong>of</strong> <strong>the</strong> factors significantly regressed (P40.05). This<br />

suggests that particle size distributions <strong>in</strong> <strong>the</strong> stomachs <strong>of</strong><br />

<strong>dugong</strong>s are not simply a function <strong>of</strong> <strong>the</strong> masticatory surface<br />

areas.<br />

Whole guts<br />

Particle size frequency histograms for each <strong>of</strong> five gut<br />

regions <strong>of</strong> three <strong>dugong</strong>s (for which <strong>the</strong> contents <strong>of</strong> each<br />

gut region were available) are shown <strong>in</strong> Fig. 4. Table 2<br />

compares <strong>the</strong> range, mean SE and CV for median particle<br />

size <strong>of</strong> each <strong>of</strong> five gut regions. Median particle size<br />

decreased progressively down <strong>the</strong> gut with a reduction<br />

<strong>in</strong> particle size distribution occurr<strong>in</strong>g at each gut region<br />

(Fig. 5), <strong>in</strong>dicat<strong>in</strong>g significant post-oral particle size reduction.<br />

There was variation between specimens, both <strong>in</strong><br />

particle size distributions with<strong>in</strong> each gut region and differences<br />

<strong>in</strong> distributions between regions (Fig. 4).<br />

There was consistent particle reduction down each gut<br />

despite <strong>the</strong> fact that diet composition probably altered along<br />

each gut. Species composition <strong>of</strong> digesta was based on<br />

stomach digesta analysis only, because it was usually impossible<br />

to identify <strong>seagrass</strong> to generic level past <strong>the</strong><br />

stomach, due to fur<strong>the</strong>r particle reduction. However <strong>in</strong> <strong>the</strong><br />

case <strong>of</strong> two <strong>of</strong> <strong>the</strong> three <strong>dugong</strong>s, Z. capricorni was identifiable<br />

<strong>in</strong> <strong>the</strong> h<strong>in</strong>dgut.<br />

The amount <strong>of</strong> particle size reduction that was attributable<br />

to mastication by <strong>the</strong> mouthparts could not be quantified.<br />

However, based on <strong>the</strong> degree <strong>of</strong> mastication <strong>of</strong> digesta<br />

<strong>in</strong> <strong>the</strong> cardiac region <strong>of</strong> <strong>the</strong> stomach (i.e. all particles were<br />

less than 8 mm <strong>in</strong> maximum dimension), this was generally<br />

considerable. By exam<strong>in</strong><strong>in</strong>g <strong>the</strong> changes <strong>in</strong> median particle<br />

280<br />

Journal <strong>of</strong> Zoology 270 (2006) 277–289 c 2006 The Authors. Journal compilation c 2006 The Zoological Society <strong>of</strong> London

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