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Two sniffing strategies in palinurid lobsters

J. A. Goldman* and S. N. Patek{dagger}

Biology Department, Duke University, Durham, NC 27708, USA
* Present address: American Institute of Biological Sciences, 1444 Eye Street, NW, Suite 200, Washington, DC 20005, USA



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Fig. 1. The antennules and flagella of Palinurus elephas. (A) An anterior view of P. elephas (carapace length 138 mm) showing the location of the antennae (ant) and antennules. The antennules are composed of antennular peduncles with three segments (p1, p2 and p3) and antennular flagella (af). Scale bar, 26 mm. (B) The antennular flagella with the medial flagellum (mf) towards the top of the figure and the lateral flagellum (lf) below. Note the discrete aesthetasc tuft (at; length 8 mm) on the lateral flagellum. Scale bar, 1.7 mm. (C) An orthogonal view of the diffuse array of aesthetasc tips with the guard hairs removed.

 


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Fig. 2. Illustration of the video analysis technique used for tracking antennule movement. (A) Video images of a flicking right antennule show the motion of the lateral flagellum (lf) during the closing phase; the medial flagellum (mf) remains stationary. The location of the antennule edge was determined by superimposing a line tangent (tl) to the motion of the arc swept by the lateral flagellum. To clearly show the lateral flagellum's motion, this illustration presents images and tangent lines at the start (tl0) and end (tl1) of a 22 ms time period (images were actually sampled every 2 ms). Scale bar, 4.9 mm. (B) The pixel intensity profile from the tangent line (tl) was used to locate the lateral flagellum's edge and to calculate position and speed during a flick. For illustration purposes, we show a small number of pixels with an intensity profile including only white (background) or dark stippling (flagellum) rather than the hundreds of pixels and continuous shades of gray actually used in the analyses. Times tl0 and tl1 correspond to the tangent lines denoted as tl0 and tl1 in panel A. Arrows indicate the moving edge of the lateral flagellum.

 


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Fig. 3. Scanning electron micrographs of the setae found on the lateral and medial flagella of Palinurus elephas. The lateral flagellum (A-D) has an aesthetasc tuft, which is flicked through the water. The medial flagellum (E-F) remains stationary during the flick. Distal is towards the left of the page in images A-E. (A) The lateral region of the aesthetasc tuft shows the aesthetascs (ae), which form paired rows on each annulus (an) of the flagellum. A single guard hair (g) per annulus flanks both medial and lateral edges of the aesthetasc tuft. Paired simple companion hairs (sc) only are located on the lateral side of the aesthetasc tuft. Scale bar, 0.5 mm. (B) Paired plumose companion hairs (pc) are found only on the medial region of the aesthetasc tuft. Scale bar, 0.5 mm. (C) The ventro-lateral side of the lateral flagellum shows the positions of setae along the aesthetasc tuft. The asymmetric setae (as) are found on the lateral side of the aesthetasc tuft just proximal to the guard hairs (g). Scale bar, 0.5 mm. (D) The lateral region of the aesthetasc tuft is shown with guard hairs and companion hairs removed so that the paired arrangement of aesthetasc rows per annulus is visible. Scale bar, 0.5 mm. (E) Long plumose setae extend along the medial flagellum. Scale bar, 0.5 mm. (F) Each plumose hair on the medial flagellum has an array of setules extending along its axis. Scale bar, 0.01 mm.

 


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Fig. 4. A cross-sectional view of the antennule flagella and setae during a flick. The lateral flagellum is rapidly flicked ventrally during the closing phase (towards the bottom of the figure) and opened more slowly. Aesthetascs are aligned such that fluid flows between rows. The medial flagellum is not actively moved and its plumose setae remain slightly below the lateral flagellum during the flicks. The location of the medial flagellum's simple setae shifts medio-laterally along the length of the flagellum.

 


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Fig. 5. Schematics of relative antennule lengths mapped onto a palinurid phylogeny. Medial flagellum is towards the top of the figure. The antennule lengths were divided by body size to show proportional differences across taxa. The phylogeny is modified from a morphological phylogeny (S. N. Patek and T. H. Oakley, submitted). The phylogenetic relationships of the Palinuridae have proved difficult to resolve, possibly due to fast radiations over ancient time scales (Patek, 2001Go); morphological and molecular rDNA sequence data have not consistently resolved genus-level relationships (Baisre, 1994Go; Patek, 2001Go; S. N. Patek and T. H. Oakley, submitted). Homarus americanus (Nephropidae), Scyllarus arctus (Scyllaridae) and Parribacus antarcticus (Scyllaridae) are included for outgroup comparisons.

 


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Fig. 6. Comparisons of lateral flagellum length (A) and peduncle length (B) for a given body size (carapace length) across lobster species. Circles represent individuals from the Panulirus genus. Squares represent members of the Palinuridae other than Panulirus. Triangles represent members of the Scyllaridae and Nephropidae.

 


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Fig. 7. Kinematic chronology of four representative flicks produced by a Palinurus elephas individual. (A) The lateral flagellum is shown at the starting position (arbitrarily set as zero) in which it is opened at a maximum distance from the medial flagellum. Then, the flagellum rapidly closes (red bracket) and slowly opens (blue bracket) four times. (B) The closing phase (red bracket, negative speed values) is faster and of shorter duration than the opening phase (blue bracket, positive speed values) of each flick.

 


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Fig. 8. Sampled water area ({Delta}) varies dramatically with small increments in lateral flagellum length. Panulirus species (circles) sample orders of magnitude larger water area per one degree of a closing phase than do other palinurid species (squares) and nephropid and scyllarid lobsters (triangles).

 

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