We examined clinging ability, subdigital pad area and body mass in 14 pad-bearing lizard species h m three families to test three predictions: (1) clinging ability and pad area should be tightly correlated among species; (2) pad area and clinging ability should scale similarly to body mass among 14 species; and (3) functional similarity in clinging capabilities should exist among species despite differences in body mass. Our results c o n f i i two predictions; clinging ability is tightly correlated with pad area, even when the effects of body size are removed, and the lizards examined are approximately functionally similar in their clinging capabilities. Nevertheless, despite the tight correlation between pad area and clinging ability, pad area scales with body mass by a lower slope than clinging ability. Overall, these results indicate that although pad area is a strong determinant of clinging ability, other factors enable these lizards to maintain functional similarity.
Sea urchin skeletons are strengthened by flexible collagenous ligaments that bind together rigid calcite plates at sutures. Whole skeletons without ligaments (removed by bleaching) broke at lower apically applied forces than did intact, fresh skeletons. In addition, in three-point bending tests on excised plate combinations, sutural ligaments strengthened sutures but not plates. The degree of sutural strengthening by ligaments depended on sutural position; in tensile tests, ambital and adapical sutures were strengthened more than adoral sutures. Adapical sutures, which grow fastest, were also the loosest, suggesting that strengthening by ligaments is associated with growth. In fed, growing urchins, sutures overall were looser than in unfed urchins. Looseness was demonstrated visually and by vibration analysis: bleached skeletons of unfed urchins rang at characteristic frequencies, indicating that sound traveled across tightly fitting sutures; skeletons of fed urchins damped vibrations, indicating loss of vibrational energy across looser sutures. Furthermore, bleached skeletons of fed urchins broke at lower apically applied forces than bleached skeletons of unfed urchins, indicating that the sutures of fed urchins had been held together relatively loosely by sutural ligaments. Thus, the apparently rigid dome-like skeleton of urchins sometimes transforms into a flexible, jointed membrane as sutures loosen and become flexible during growth.
The feeding mechanism of Mellita quinquiesperforata (Leske) has been examined in detail. This sand dollar is a deposit feeder, ingesting particles mostly in the range of 100-250 µm. The particles are picked out of the substrate individually by specialized long barrel-tipped podia, which form a narrow palisade surrounding the geniculate spine fields on the oral surface. Selected food items are passed to short barrel-tipped podia, thence from podium to podium until they reach the food grooves where they are finally aggregated into mucus cords. The cords are passed to the mouth by the activity of food groove podia. At the peristome, the cord is passed between the circumoral spines by large food groove podia and steered into the mouth by five pairs of buccal podia. The lantern is powerfully muscled and has hardened teeth which crush diatoms and fracture many sand grains. For this reason, there is an apparent accumulation of fine particles (<50 µm) in the gut. Analysis of size frequencies of the material in the mucus cords and substrate indicates that no selection of fine particles occurs and, in fact, that they are virtually absent from the native sediment. An account of spine and podial morphology and distribution is included with descriptions and measurements of surface ciliary currents. It is shown that the formerly accepted sieve hypothesis of feeding cannot be entirely rejected on theoretical grounds. However, during feeding there was no evidence of the operation of any of the elements of the supposed sieve mechanism. Furthermore, the ciliary currents are not fast enough to account for the movement of most ingested material. Patterns of ciliary flow on the oral surface are not simply centripetal, but are much more complex than previously supposed.
Used singly, the fluorochrome tags tetracycline and calcein have yielded important insights into sea urchin biology, especially regarding growth. We present a new method of tagging using sequential fluorochrome markers, as well as a more precise method of quantifying growth. Such polyfluorochromes enable repeated markings that allow measurement of multiple growth points and unique identification of individuals or groups. We marked sea urchins, Strongylocentrotus droebachiensis, with four fluorochromes: alizarin complexone, calcein, calcein blue, and tetracycline. All fluorochromes marked both by injection and immersion. We examined the clarity of the mark produced with low, metabolically scaled doses, and higher doses similar to those that have been previously used. We tested the effect of fluorochromes on survival, growth, jaw size, and gonad size by marking a size range (3.9-44.3 mm in diameter) of urchins with either one or all four fluorochromes. We quantified growth using a nominal diameter, that is a fitted constant, times the cube root of weight, which increased the precision of measurements by a factor of six relative to measured diameter. Growth rate was a decreasing function of diameter except for a growth lag in the smallest urchins. Growth rate data for all sizes were fitted using: gamma distributions; Tanaka functions; and, for larger sizes, straight lines (von Bertalanffy model). All treatments produced clear marks, with higher doses producing more reliably clear marks. Tetracycline marking did not affect growth; other treatments produced only transient slowing of growth in the marking month. Growth rate, survival, gonad production, and jaw weight did not differ between control and treatments during the following 6 months. Thus, polyfluorochromes produce reliable marks that do not significantly affect growth or gonad production.
Clams of the species Donax variabilis migrate shoreward during rising tides and seaward during falling tides. These clams spend most of the time in the sand, emerging several times per tidal cycle to ride waves. Migration is not merely a passive result of waves eroding clams out of the sand; rather clams actively jump out of the sand and ride specific waves. Such active migration is experimentally demonstrated during a falling tide by comparing the motion of dead and live clams; live clams emerge from the sand and move seaward even when dead ones do not. As low tide approaches, live clams become progressively less active. They cease migrating for 2 hours around low tide and resume jumping to migrate shoreward after the tide has turned. During the rising tide, far from being passive, the clams jump out to ride only the largest 20% of waves. Specifically, they choose swash that have the largest excursion, i.e., those swash that move furthest on the beach.
Indirect predator-induced effects on growth, morphology and reproduction have been extensively studied in marine invertebrates but usually without consideration of size-specific effects and not at all in post-metamorphic echinoids. Urchins are an unusually good system, in which, to study size effects because individuals of various ages within one species span four orders of magnitude in weight while retaining a nearly isometric morphology. We tracked growth of urchins, Strongylocentrotus droebachiensis (0.013–161.385 g), in the presence or absence of waterborne cues from predatory Jonah crabs, Cancer borealis. We ran experiments at ambient temperatures, once for 4 weeks during summer and again, with a second set of urchins, for 22 weeks over winter. We used a scaled, cube-root transformation of weight for measuring size more precisely and for equalizing variance across sizes. Growth rate of the smallest urchins (summer: <17 mm diameter; winter: <7 mm diameter) decreased by 40–42% in response to crab cues. In contrast, growth rate of larger urchins was unaffected in the summer and increased in response to crab scent by 7% in the winter. At the end of the 22-week experiment, additional gonadal and skeletal variables were measured. Cue-exposed urchins developed heavier, thicker skeletons and smaller gonads, but no differences in spine length or jaw size. The differences depended on urchin size, suggesting that there are size-specific shifts in gonadal and somatic investment in urchins.
We measured coelomic pressure in sea urchins to determine whether it was high enough to support a pneu hypothesis of growth. In Strongylocentrotus purpuratus the pressure was found to fluctuate rhythmically about a mean of -8 Pa, and was negative for 70% of the time. This is at variance with the theoretically required positive pressures of the pneu hypothesis. Furthermore, there were no sustained significant differences between the pressure patterns of fed and starved urchins, presumed to be growing and not growing, respectively. The rhythmical fluctuations in pressure were caused by movements of the lantern which changed the curvature and tension of the peristomial membrane. We developed a mathematical and morphological model relating lantern movements, membrane tension, and pressure, that correctly predicts the magnitude of the fluctuations. Pressures predicted by the model depend also on coelomic volume changes. In Lytechinus variegatus simultaneous retraction of the podia, which causes expansion of the ampullae, resulted in an 8.8 Pa increase in coelomic pressure, relative to the pressure during simultaneous podial protraction.
Externally visible, growing calcitic structures can be marked using fluorochromes. Such marks are useful for field recapture studies in ecology, evolution, and aquaculture as well as for studies on mechanisms of growth and development. We marked 2-month-old sea urchins (Strongylocentrotus droebachiensis) with the fluorochromes calcein, calcein blue, and tetracycline by batch-marking via immersion. Neither growth nor survival was affected by marking. Marks were externally visible on the skeletal plates, demipyramids, and spines of 100% of the marked sea urchins up to 63 d post-marking. After 342 d, marks were still externally visible on 100% of calcein-marked, 98% of calcein blue-marked, and 22% of tetracycline-marked sea urchins. Marks were brightest on calcein-marked and faintest on tetracycline-marked sea urchins, in correspondence to the fluorochrome dose. Growth marks in the aboral oculogenital ring, followed for 333 d, showed that genital plate growth in the hoop direction was greatest adjacent to the anal suture and that both the oculogenital ring and periproct grew less than isometrically. Internal marks (not externally visible) were subsequently seen on 99% of the demipyramids at 342 d post-marking. Such fluorochrome marks on demipyramids have previously been used to measure aboral and oral-end demipyramid growth to allometrically calibrate diametrical growth rates of field sea urchins. We found that although aboral demipyramid marks were always clear, 13% of marks on the oral end were obscured. However, we show that measuring only aboral end growth is sufficient for allometric calibration. In a separate experiment, multiple marks of the above three fluorochromes plus alizarin complexone, administered by injection to larger (12.9-37.1 mm diameter) sea urchins, persisted internally for at least 2 years. Multi-color, internally and externally visible, persistent marks will enhance experimental designs in laboratory, field, and common garden experiments.
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