5. Ventilation

Slots and openings in the cabinet are provided for ventilation. To ensure reliable operation of the product and to protect it from overheating, these openings must not be blocked or covered. Do not block the openings by placing the product on a bed, sofa, rug or other similar surface. Do not place the product in a built-in installation such as a bookcase or rack unless proper ventilation is provided or the manufacturers instructions have been adhered to.
IMPORTANT PRODUCT SAFETY INSTRUCTIONS
Electrical energy can perform many useful functions. But improper use can result in potential electrical shock or fire hazards. This product has been engineered and manufactured to assure your personal safety. In order not to defeat the built-in safeguards, observe the following basic rules for its installation, use and servicing.
6. Wall or Ceiling Mounting
The product should be mounted to a wall or ceiling only as recommended by the manufacturer.

ATTENTION:

Follow and obey all warnings and instructions marked on your product and its operating instructions. For your safety, please read all the safety and operating instructions before you operate this product and keep this manual for future reference.
ANTENNA INSTALLATION INSTRUCTIONS
1. Outdoor Antenna Grounding
If an outside antenna or cable system is connected to the product, be sure the antenna or cable system is grounded so as to provide some protection against voltage surges and built-up static charges. Article 810 of the National Electrical Code, ANSI/NFPA 70, provides information with regard to proper grounding of the mast and supporting structure, grounding of the lead-in wire to an antenna discharge unit, size of grounding conductors, location of antenna discharge unit, connection to grounding electrodes, and requirements for the grounding electrode.

INSTALLATION

1. Grounding or Polarization
(A) Your product may be equipped with a polarized alternating-current line plug (a plug having one blade wider than the other). This plug will fit into the power outlet only one way. This is a safety feature. If you are unable to insert the plug fully into the outlet, try reversing the plug. If the plug should still fail to fit, contact your electrician to replace your obsolete outlet. Do not defeat the safety purpose of the polarized plug. (B) Your product may be equipped with a 3-wire grounding-type plug, a plug having a third (grounding) pin. This plug will only fit into a grounding-type power outlet. This is a safety feature. If you are unable to insert the plug into the outlet, contact your electrician to replace your obsolete outlet. Do not defeat the safety purpose of the grounding-type plug.

LIGHT OFF/AUTO/ON Switch

DANGER
The video light can become extremely hot. Do not touch it either while in operation or soon after turning it off, otherwise serious injury may result. Do not place the camcorder into the carrying case immediately after using the video light, since it remains extremely hot for some time. When operating, keep a distance of about 30 cm (11-13/16") between the video light and people or objects. Do not use near flammable or explosive materials. It is recommended that you consult your nearest JVC dealer for replacing the video light.
RECORDING Advanced Features

Fade/Wipe Effects

EFFECT Button
These effects let you make pro-style scene transitions. Fade- or wipe-in works at recording start, and fade- or wipe-out works at recording end or when you enter RecordStandby mode.
Set the Power Switch to PRO. Press EFFECT repeatedly until the desired modes name and indication appear. They are displayed for approx. 2 seconds, then the name disappears so that only the indication remains. The effect is reserved.
Press the Recording Start/Stop Button to activate fadein/out or wipe-in/out. To cancel a fade or wipe, press EFFECT repeatedly until OFF appears. OFF is displayed for approx. 2 seconds and the fade/wipe standby mode is canceled.
Fades in/out to a black screen.

MOSAIC (Fader)

Gradually turns/returns the picture into/from a mosaic pattern.

SHUTTER (Wipe)

A black screen moves in from the top and bottom, closing over the image like a shutter, or a new image pushes open the black screen vertically from the center.

SLIDE (Wipe)

A black screen moves in from the left to gradually cover the image, or a new image moves in from right to left.

DOOR (Wipe)

Wipes in as the two halves of a black screen open to the left and right, revealing the scene, or wipes out and the black screen reappears from left and right to cover the scene.
NOTES: Pressing and holding the Recording Start/Stop Button allows you to vary the length for the image during fade in/out or wipe in/out. The screen becomes slightly reddish when the Fade/Wipe is used with Sepia ( pg. 22). With the Electronic fog filter mode ( pg. 22) engaged, the image fades in/out to a white screen.

CW:CORNER (Wipe)

Wipes in on a black screen from the upper right to the lower left corner, or wipes out from lower left to upper right, leaving a black screen.

WW:WINDOW (Wipe)

The next scene gradually wipes in from the center of the screen toward the corners, covering the previous scene.
RECORDING Advanced Features (cont.)
Program AE With Special Effects

P.AE Button

Power Switch
Set the Power Switch to PRO. Press P.AE repeatedly until the desired modes name and indication appear. They are displayed for approx. 2 seconds, then the name disappears so that only the indication remains. The mode is activated. To cancel the effect, press P.AE repeatedly until OFF appears. OFF is displayed for approx. 2 seconds.
NOTES: Only one mode can be engaged at a time. The screen becomes slightly reddish when the Fade/ Wipe ( pg. 21) is used in the Sepia mode. The screen becomes slightly dark in the High Speed Shutter mode. Use in well-lit situations. In the High Speed Shutter or Sports modes, picture color may be adversely affected if the subject is lit by alternating discharge-type light sources such as flourescent or mercury-vapor lights.
The scene being shot is recorded in sepia-tinted (reddish-brown) monochrome, giving the effect of an older movie. Use together with Wide ( pg. 24) for the authentic look of a classic Hollywood movie.

TWILIGHT

Dusk, twilight scenery, fireworks, etc., look more natural and dramatic. The following happens when Twilight mode is selected: Auto gain control is turned off. White Balance is set to :FINE (day mode), but can also be manually changed to another mode ( pg. 30). Auto Focus becomes available only in the range of 10 m (32 ft.) to infinity. To focus when the subjectto-camera distance is less than 10 m (32 ft.), use manual focusing ( pg. 29).

SPORTS

High shutter speed clearly captures fast-moving action.

ND:ND EFFECT

A black mist darkens the picture, as when an ND filter is used. Helps to counter the effects of glare on the subject.

S1/2000

Captures faster action than Sports mode.

FG:FOG

Makes the picture look misty white, as when an external fog filter is attached to the lens. Softens the image and gives it a fantasy look.

NEGA POSI

The colors of a picture are reversed.

NEGA POSI mode

Using Menus For Detailed Adjustment
This camcorder is equipped with an easy-to-use, on-screen menu system that simplifies many of the more detailed camcorder settings.

NORMAL

OFF TITLE DATE/TIME DISP.
Allows you to superimpose one of eight preset titles ( pg. 26). Makes the date/time settings appear in the camcorder or on a connected monitor ( pg. 26).
Menu Screen Available Using The MENU Button
This Menu Screen cannot be accessed while recording. D. ZOOM ON Allows you to use the Digital Zoom. By digitally processing and magnifying images, zooming is possible from 16X (the optical zoom limit), to a maximum of 400X digital magnification. Digital Zoom is not available. Only optical zoom (maximum 16x magnification) can function. When set to OFF during Digital Zoom, zoom magnification changes to 16X. Usually the distance to a subject where the lens is in focus depends on the zoom magnification. Unless there is a distance of more than 1 m (3.25 ft.) to the subject, the lens is out of focus at the maximum telephoto setting. When set to ON, you can shoot a subject as large as possible at a distance of approx. 60 cm (23-5/8"). Depending on the zoom position, the lens may go out of focus.

TELE MACRO

OFF ON

: Factory-preset

EN Menu Screen Available Using The MENU Button (cont.)
S-VHS* (S-VHS ET*) TAPE LENGTH REC TIME INT. TIME TITLE LANG. DATE/TIME JLIP ID NO. DEMO MODE ON OFF Records in S-VHS on a VHS or S-VHS cassette ( pg. 13). Records in VHS on a VHS or S-VHS cassette ( pg. 13).
Allows you to set the tape length depending on the tape used ( pg. 12). Refer to Animation and Time-Lapse ( pg. 27). Refer to Time-Lapse ( pg. 27). Allows you to select the language (ENGLISH, FRENCH, SPANISH or PORTUGUESE) of Instant Titles ( pg. 26). Allows you to set the current date and time ( pg. 11). This number is necessary when connecting the camcorder to a device such as a computer using the J terminal (JLIP). The numbers range from 01 to 99. Factory setting is 06. ON Demonstrates certain functions such as Fade/Wipe, etc. When DEMO MODE is set to ON and the Menu Screen is closed, the demonstration starts. Operating the Power Zoom Lever during the demonstration stops the demonstration temporarily. If the Power Zoom Lever is not moved for more than 1 minute after that, the demonstration will resume. NOTE: When a tape whose Erase Protection tab is in the position that allows recording is loaded in the camcorder, demonstration is not available. Automatic demonstration will not take place.

*S-VHS ET is displayed when a VHS cassette is loaded, and S-VHS is displayed when an S-VHS cassette is loaded ( pg. 13). If a cassette is not loaded, S-VHS will be displayed in the Menu Screen. S-VHS ET is preset to OFF.

Date/Time Insert

Allows you to display the date and time in the camcorder or on a connected color monitor, as well as to record them manually or automatically. You should have already performed the Date/ Time Setting procedure ( pg. 11).
Set the Power Switch to PRO. Press the Select Wheel to display the Menu Screen. Rotate the Select Wheel to move the highlight bar to DATE/TIME DISP. while in Record-Standby, then press it to display the Date/Time DISP. Menu. Rotate the Select Wheel to move the highlight bar to the desired mode, then press it. The desired mode is activated. The Menu Screen reappears with the highlight bar on RETURN. Press the Select Wheel to close the Menu Screen.
Select Wheel Power Switch

DEC 2 5. 9 9

AUTO DATE DEC 25. 99 (DATE) AM 10 : 25 : 00 (TIME) DEC 25. 99 AM 10 : 25 : 00 (DATE&TIME) No indication (OFF)
NOTES: DISPLAY The selected display can be recorded. If you do not want to record the display, select the OFF mode before shooting. AUTO DATE Your camcorder automatically records the date for about 5 seconds after recording is initiated in the following situations: After changing the date. After loading a cassette. After Auto Date Record mode is selected by rotating the Select Wheel. In this mode, the date is replaced after 5 seconds with AUTO DATE but this is not recorded.

Instant Titles

The camcorder has eight preset titles in memory. You can superimpose one of them over the video image. Instant Titles can be displayed not only in English but also in French, Spanish and Portuguese. Change the setting in TITLE LANG. in the Menu Screen. ( pg. 23, 25).
T I TLE OF F HA P P Y B I R T HDA Y OUR V ACA T I ON ME RR Y CHR I S TMA S A S P E C I A L DA Y HA P P Y HO L I DA Y S OUR N EW BAB Y WE DD I NG DA Y CONGRA T U L A T I ON S EX I T
Set the Power Switch to PRO. Press the Select Wheel to display the Menu Screen. Rotate the Select Wheel to move the highlight bar to TITLE while in Record-Standby, then press it to display the TITLE Menu. Rotate the Select Wheel to move the highlight bar to the desired mode, then press it. The desired mode is activated. To make the title indication disappear, select OFF. The Menu Screen reappears with the highlight bar on RETURN. Press the Select Wheel to close the Menu Screen.

LCD Monitor Brightness Adjustment: Rotate the Select Wheel (BRIGHT) towards + to brighten, or towards to darken. The Bright Level Indicator is displayed under the Date/ Time Display when you play back images recorded in Wide Mode ( pg. 24). Speaker Volume Control: Slide the Power Zoom Lever (VOL.) towards + to turn up the volume, or towards to turn down the volume. The Speaker Volume Indicator is displayed under the Date/Time Display when you play back images recorded in Wide Mode ( pg. 24). Still Playback: Pauses during playback. 1) Press 4/6 during playback. 2) To resume normal playback, press 4/6 again. Noise bars will appear and the picture will become monochrome during Still Playback. Shuttle Search: Allows high-speed search in either direction. 1) Press 3 for forward or 2 for reverse search during playback. 2) To resume normal playback, press 4/6. During playback, press and hold 2 or 3. The search continues as long as you hold the button. Once you release it, normal playback resumes. Noise bars appear and pictures may become monochrome or darken during Shuttle Search. This is normal.

P.STABILIZER Button

PLAYBACK Features
Tracking: Eliminates noise bars that appear on-screen
during playback. Factory-preset is Auto Tracking. To activate Manual Tracking: 1) Press P.AE and EFFECT simultaneously. MT appears. 2) Press P.AE or EFFECT as many times as necessary until the noise bars disappear. To return to Auto Tracking, press P.AE and EFFECT simultaneously. AT blinks. When Auto Tracking finishes, the indication disappears. When noise bars appear during playback, the camcorder enters the Auto Tracking mode and AT is displayed. Manual Tracking may not work with tapes recorded on other VCRs or camcorders.

COUNTER R/M Button

TBC (Time Base Corrector): Removes jitter from
fluctuating video signals to deliver a stable picture even with old tapes. Factory-preset: TBC is engaged. To activate/release the TBC mode, press P.STABILIZER during playback for more than 1 second. When the TBC mode is activated, TBC is displayed. The TBC indicator turns green (gray with a camcorder equipped with a black/white viewfinder) while TBC is working. TBC does not work during still playback and shuttle search. The TBC indicator turns white while TBC is not operative. It may take a few seconds before TBC actually starts working. The picture may be distorted if the TBC mode is activated or deactivated at the edit-in/-out points or when normal playback resumes after still playback or shuttle search. If the playback picture is distorted when TBC is set to on, turn off TBC.
Counter Memory Function: Makes it easier to
locate a specific tape segment. 1) Press COUNTER R/M and hold for more than 1 second. The counter resets to 0:00:00. 2) Press COUNTER R/M for less than 1 second. M appears. 3) After recording or playback, press 5, then 2. The tape automatically stops at or close to 0:00:00. 4) Press 4/6 to start playback. The counter memory functions in the Fast-Forward and Rewind modes. To disable the Counter Memory Function, press COUNTER R/M for less than 1 second so that M disappears.

To AUDIO IN

* When connecting the cables, open this cover.

Basic Connections

Tape Dubbing
Make sure all units are turned off. Connect the camcorder to a TV or VCR as shown in the illustration ( pg. 34). If using a VCR. go to step 3. If not. go to step 4. Connect the VCR output to the TV input, referring to your VCRs instruction manual. Turn on the camcorder, the VCR and the TV. Set the VCR to its AUX input mode, and set the TV to its VIDEO mode.
Following the illustration on pg. 34, connect the camcorder and the VCR. Set the camcorders Power Switch to PLAY, turn on the VCRs power, and insert the appropriate cassettes in the camcorder and the VCR. Engage the VCRs AUX and Record-Pause modes. Engage the camcorders Play mode to find a spot just before the edit-in point. Once it is reached, press 4/6 on the camcorder. Press 4/6 on the camcorder and engage the VCRs Record mode. Engage the VCRs Record-Pause mode and press 4/6 on the camcorder. Repeat steps 4 through 6 for additional editing, then stop the VCR and camcorder when finished.
NOTES: It is recommended to use the AC Power Adapter/ Charger as the power supply instead of the battery pack ( pg. 9). If your VCR has an S-Video input connector, connect the camcorder and the VCR using an optional S-Video cable. This can improve the dubbed picture quality. To monitor the picture and sound from the camcorder without inserting a tape, set the camcorders Power Switch to CAMERA (AUTO or PRO.), then set your TV to the appropriate input mode. Make sure you adjust the TV sound volume to its minimum level to avoid a sudden burst of sound when the camcorder is turned on. If you have a TV or speakers that are not specially shielded, do not place the speakers adjacent to the TV as interference will occur in the camcorder playback picture.

RM-V715U (provided)

The Full-Function Remote Control Unit can operate this camcorder from a distance as well as the basic operations (Playback, Stop, Pause, Fast-Forward and Rewind) of your VCR.

Installing The Batteries

The remote control uses two AAA (R03) size batteries. See General Battery Precautions ( pg. 46).
Open the battery compartment cover as illustrated. Insert two AAA (R03) size batteries in the correct direction. Replace the battery compartment cover.
Transmitted beam effective area

Remote sensor

When using the remote control, be sure to point it at the remote sensor. The illustration shows the approximate transmitted beam effective area for indoor use. The transmitted beam may not be effective or may cause incorrect operation outdoors or when the remote sensor is directly exposed to sunlight or powerful lighting.

Functions Buttons

With the camcorders Power Switch set to CAMERA (AUTO or PRO.). Transmits the beam signal. Zoom in/out ( pg. 18) Retake (rewind), Quick Review ( pg. 19) Animation, Time-Lapse ( pg. 37) Playback start ( pg. 31) Rewind, Reverse Shuttle Search ( pg. 31) Stop ( pg. 31) Insert Editing ( pg. 38) With the camcorders Power Switch set to PLAY.

1 Infrared beam

transmitting window
2 ZOOM (T/W) Buttons 3 PLAY Button 4 REW Button 5 STOP Button 6 INSERT Button 7 REC TIME Button

ANIM. Button

8 START/STOP Button 9 FF Button 0 PAUSE Button ! A.DUB Button @ INT. TIME Button

SELF TIMER Button*

Functions the same as the Recording Start/Stop Button on the camcorder. Retake (forward) ( pg. 19) Time-Lapse ( pg. 37) Fast-Forward, Forward Shuttle Search ( pg. 31) Pause ( pg. 31) Audio dubbing ( pg. 39)
* This function is unavailable with this camcorder.
FEATURE: Animation and Time-Lapse

Interval time indicator

1M I N 1 / 2S
The remote control lets you set/release Interval Time and Recording Time without using the Menu Screen.

Recording time indicator

Animation 1) Set Instead of performing steps 1 and 2 on pg. 27 (Animation), press REC TIME on the remote control. Each time REC TIME is pressed, the Recording Time indicator appears, changing in the following order: 1/4S, 1/2S, 1S, 5S and no indication (off). 2) Release Instead of using the Menu in step 5 on pg. 27, press REC TIME on the remote control until the Recording Time indicator disappears. Time-Lapse 1) Set Instead of performing steps 1 through 3 on pg. 27 (Time-Lapse), press INT. TIME and REC TIME on the remote control. Each time INT. TIME is pressed, the Interval Time indicator appears, changing in the following order: 15S, 30S, 1MIN, 5MIN and no indication (off). Each time REC TIME is pressed, the Recording Time indicator appears, changing in the following order: 1/4S, 1/2S, 1S, 5S and no indication (off). 2) Release Instead of using the Menu in step 5 on pg. 27, press INT. TIME and REC TIME on the remote control until the indicators disappear. NOTE: Also refer to page 27.

START/STOP

REC. TIME INT. TIME
USING THE REMOTE CONTROL UNIT (cont.)

Insert Editing

You can record a new scene onto a previously recorded tape, replacing a section of the original recording with minimal picture distortion at the in- and out-points. The original audio remains unchanged. NOTE: Use the RM-V715U remote control unit to perform this procedure.
Power Switch COUNTER R/M Button
Set the Power Switch to PLAY. Play back the tape, locate the Edit-Out point and press PAUSE. Press and hold COUNTER R/M on the camcorder for more than 1 second to reset the tape counter, then press again for less than 1 second so that M appears. Press REW to go a little beyond the beginning of the scene you want to replace, press PLAY to view the recorded tape, and press PAUSE exactly at the Edit-In point where the new scene should start. Press and hold INSERT, and without releasing INSERT, press and release PAUSE. The Insert Editing indicator appears and the Insert Editing mode is engaged. When the scene the camcorder is aimed at appears on the screen, go to the next step. To begin Insert Editing, press START/STOP. When the counter reaches 0:00:00, editing stops automatically and the camcorder enters the Insert-Pause mode. To end Insert Editing, press STOP.

Insert editing indicator

M 0 : : 3 4

Tape Counter

PLAY REW STOP INSERT

START/STOP PAUSE

NOTES: During Insert Editing, the original audio will be heard from the speaker. After step 5, Retake ( pg. 19) can be performed. The tape may stop slightly before or after the designated end point, and noise bars may appear. Neither indicates a defect in the unit. Only the video signal is recorded in the Insert Editing mode.

Audio Dubbing

Audio dubbing indicator

M 0 : : 2 0

During Audio Dubbing
You can record a new soundtrack on a prerecorded tape (normal audio only). The sound is recorded from the builtin microphone. Perform steps 1, 2 and 3 of the Insert Editing procedure ( pg. 38) before continuing. NOTE: Use the RM-V715U remote control unit to perform the following procedures.

Microphone

Press REW to go a little beyond the beginning of the scene onto which you want to dub the new audio, press PLAY to view the recorded tape, then press PAUSE at exactly the point where dubbing should start. Press and hold A. DUB, and without releasing A. DUB, press and release PAUSE. The Audio DubStandby mode is engaged. To begin Audio Dubbing, press PLAY. When the counter reaches 0:00:00, Audio Dubbing stops automatically and the camcorder enters the Audio Dub-Standby mode. To end Audio Dubbing, press STOP.

PLAY REW STOP

START/STOP PAUSE A.DUB
NOTES: Do not press FF or REW during Audio Dub-Standby, or the edit points will not be accurate. If the microphone is too close to the TV, or if the TVs volume is too high, whistling or howling may occur. NOTES (for Insert Editing and Audio Dubbing): Insert Editing and Audio Dubbing may not work correctly if the tape contains blank segments, or if the recording speed was changed during the original recording. To edit/dub onto a tape, make sure the Erase Protection tab is in the position that allows recording. If not, slide the tab. Some cassettes have removable tabs. If the tab has been removed, cover the hole with adhesive tape. Do not press STOP during Insert Editing or Audio Dubbing, or the edit points will not be accurate. During Insert Editing and Audio Dubbing, when InsertPause or Audio Dub-Standby is engaged, the counter may go slightly past 0:00:00. Recording, however, stops at exactly 0:00:00.

If, after following the steps in the chart below, the problem still exists, please consult your JVC dealer. The camcorder is a microcomputer-controlled device. External noise and interference (from a TV, a radio, etc.) might prevent it from functioning properly. In such cases, first disconnect its power supply unit (battery pack, AC Power Adapter/Battery Charger, etc.) and clock battery; and then re-connect it and proceed as usual from the beginning.

SYMPTOM

POSSIBLE CAUSE(S)
The battery pack has not been attached correctly ( pg. 8). The battery pack is not charged ( pg. 8). The power supply has not been correctly connected ( pg. 9). The battery pack has completely discharged. Remove the cassette and disconnect the power source, then after a few minutes, try turning the power back on. If it still does not come on, consult your nearest JVC dealer.

No power is supplied.

The power suddenly goes off and does not come back on by itself.

RECORDING

Recording cannot be performed. Make sure the Erase Protection tab is in the position that allows recording. If not, slide the tab. Some cassettes have removable tabs. If the tab has been removed, cover the hole with adhesive tape ( pg. 10). The camcorder Power Switch has not been set to CAMERA (AUTO or PRO.) ( pg. 16). Before recording in the Animation mode at the very beginning of a tape, set the camcorder to the Recording mode for about 5 seconds, so that the tape runs smoothly. Using the Fade-in function at this point is a good way to begin an animated program ( pg. 27, 37).
Recording does not start. Animation or Time-Lapse is not available.
The tape is running, but there is no playback picture. Playback picture is blurred or interrupted. The counter indication is blurred during Still playback. The TV has not been set to its VIDEO mode or channel. If A/V connection is used, the TVs VIDEO/TV Switch has not been set to VIDEO. The video heads are dirty or worn out. Consult your nearest JVC dealer for head cleaning or replacement. This is normal.

TAPE TRANSPORT

The tape stops during fast-forward or rewind. Rewinding or fast-forwarding cannot be performed. The Counter Memory Function has been activated ( pg. 32). The tape is already fully wound on one reel or the other.
Press DISPLAY for longer than 1 second to make the indications appear ( pg. 17). The battery pack is running low. If the LCD monitor is not open over 60 degrees, the EJECT Switch does not function. The Power Switch is set to AUTO. Certain combinations of modes or effects are not possible to use. This sometimes occurs when the contrast between the background and the object is great. It is not a defect of the camcorder. Sunlight is directly entering the lens. This is not a defect of the camcorder. The light used to illuminate the LCD monitor causes it to be hot. Close the LCD monitor to turn it off or set the Power Switch to OFF, and let the unit cool down. The LCD monitor and the viewfinder are made with highprecision technology. However, black spots or bright spots of light (red, green or blue) may appear constantly on the LCD monitor or in the viewfinder. These spots are not recorded on the tape. This is not due to any defect of the unit. (Effective dots: more than 99.99 %.) This may occur when the surface or the edge of the LCD monitor is pressed. Wipe them gently with a soft cloth. Gently wipe in a semicircular motion, as wiping strongly spreads the stain and it wont come out easily. This is normal while recording in the EP mode. Some noise may be present while recording in the SP mode ( pg. 12). Turn the camcorders power off, then disconnect the power source. After a few minutes, try turning the power on again. If the error number still appears in the viewfinder, consult your nearest JVC dealer. The diopter needs to be adjusted ( pg. 15). The scene being shot is adversely affecting the operation of Auto Focus ( pg. 29). If you remove the power source from the camcorder while the power is on, all settings and selections are erased. Make sure to turn the camcorders power off before disconnecting the power source. If you have not performed Date/Time Setting ( pg. 11), the date and time are not displayed/recorded correctly. Be sure that Date/Time setting is carried out. The Lens Cover Warning indicator may blink when the camcorder is used in dark areas, regardless of the position of the lens cover.

Plug adapter

General Battery Precautions

INFORMATION

This device complies with Part 15 of FCC Rules. Operation is subject to the following two conditions: (1) This device may not cause harmful interference, and (2) this device must accept any interference received, including interference that may cause undesired operation. Change or modifications not approved by the party responsible for compliance could void the users authority to operate the equipment. This equipment has been tested and found to comply with the limits for a Class B digital device, pursuant to Part 15 of the FCC Rules. These limits are designed to provide reasonable protection against harmful interference in a residential installation. This equipment generates, uses, and can radiate radio frequency energy and, if not installed and used in accordance with the instructions, may cause harmful interference to radio communications. However, there is no guarantee that interference will not occur in a particular installation. If this equipment does cause harmful interference to radio or television reception, which can be determined by turning the equipment off and on, the user is encouraged to try to correct the interference by one or more of the following measures: Reorient or relocate the receiving antenna. Increase the separation between the equipment and receiver. Connect the equipment into an outlet on a circuit different from that to which the receiver is connected. Consult the dealer or an experienced radio/TV technician for help. If the remote control or cassette adapter is not functioning even if it is being operated correctly, the batteries are exhausted. Replace them with fresh ones. Use only the following batteries: Remote control.. AAA (R03) size x 2 (RM-V715U) Cassette adapter.. AA (R6) size x 1 (C-P6U or C-P7U) Please make note of the following rules for battery use. When misused, the batteries can leak or explode. 1. When replacing batteries, refer to page 36 for the remote control, or read the cassette adapter (C-P6U/C-P7U) instructions. 2. Do not use any different size of batteries from those specified. 3. Be sure to install batteries in the correct direction. 4. Do not use rechargeable batteries. 5. Do not expose the batteries to excessive heat as they can leak or explode. 6. Do not dispose of the batteries in a fire. 7. Remove the batteries from the unit if it is to be stored for an extended period to avoid battery leakage which can cause malfunctions.

Battery Packs

The battery packs are nickel-cadmium or nickel metal-hydride batteries. Before using the supplied battery pack or an optional battery pack, be sure to read the following cautions: 1. To avoid hazard. do not burn. Terminals. do not short-circuit the terminals. do not modify or disassemble. use only specified chargers. 2. To prevent damage and prolong service life. do not subject to unnecessary shock. avoid repeated charging without fully discharging. charge in an environment where temperatures are within the tolerances shown in the chart below. This is a chemical reaction type batterycooler temperatures impede chemical reaction, while warmer temperatures can prevent complete charging. store in a cool, dry place. Extended exposure to high temperatures will increase natural discharge and shorten service life. avoid prolonged uncharged storage. remove from charger or powered unit when not in use, as some machines use current even when switched off. NOTES: It is normal for the battery pack to be warm after charging, or after use. Temperature Range Specifications Charging. 10C to 35C (50F to 95F) Operation. 0C to 40C (32F to 104F) Storage. 10C to 30C (14F to 86F) Recharging time is based on room temperature of 20C (68F.) The lower the temperature, the longer recharging takes.

Cassettes

To properly use and store your cassettes, be sure to read the following cautions: 1. During use. make sure the cassette bears the VHS-C mark. be aware that recording onto prerecorded tapes automatically erases the previously recorded video and audio signals. make sure the cassette is positioned properly when inserting. do not load and unload the cassette repeatedly without allowing the tape to run at all. This slackens the tape and can result in damage. do not open the front tape cover. This exposes the tape to fingerprints and dust. 2. Store cassettes. away from heaters or other heat sources. out of direct sunlight. where they wont be subject to unnecessary shock or vibration. where they wont be exposed to strong magnetic fields (such as those generated by motors, transformers or magnets). vertically, in their original cases.

CAUTIONS (cont.)

LCD Monitor
1. To prevent damage to the LCD monitor, DO NOT. push it strongly or apply any shocks. place the camcorder with the LCD monitor on the bottom. 2. To prolong service life. avoid rubbing it with coarse cloth. 3. Be aware of the following phenomena for LCD monitor use. These are not malfunctions: While using the camcorder, the surface around the LCD monitor and/or the back of the LCD monitor may heat up. If you leave power on for a long time, the surface around the LCD monitor becomes hot.

Main Unit

1. For safety, DO NOT. open the camcorders chassis. disassemble or modify the unit. short-circuit the terminals of the battery pack. Keep it away from metallic objects when not in use. allow inflammables, water or metallic objects to enter the unit. remove the battery pack or disconnect the power supply while the power is on. leave the battery pack attached when the camcorder is not in use. 2. Avoid using the unit. in places subject to excessive humidity or dust. in places subject to soot or steam such as near a cooking stove. in places subject to excessive shock or vibration. near a television set. near appliances generating strong magnetic or electric fields (speakers, broadcasting antennas, etc.). in places subject to extremely high (over 40C or 104F) or extremely low (under 0C or 32F) temperatures. 3. DO NOT leave the unit. in places of over 50C (122F). in places where humidity is extremely low. (below 35%) or extremely high (above 80%). in direct sunlight. in a closed car in summer. near a heater. 4. To protect the unit, DO NOT. allow it to become wet. drop the unit or strike it against hard objects. subject it to shock or excessive vibration during transportation. keep the lens directed at extremely bright objects for long periods. direct the eyepiece of the viewfinder at the sun. carry it by holding the viewfinder or the LCD monitor. Be sure to hold the main unit with both hands or use the grip. swing it excessively when using the shoulder strap.

WARRANTY (Only in U.S.A.)

LIMITED WARRANTY

CONSUMER VIDEO 1-90
JVC COMPANY OF AMERICA warrants this product and all parts thereof, except as set forth below ONLY TO THE ORIGINAL PURCHASER AT RETAIL to be FREE FROM DEFECTIVE MATERIALS AND WORKMANSHIP from the date of original retail purchase for the period as shown below. (The Warranty Period) PARTS

90 DAYS

THIS LIMITED WARRANTY IS VALID ONLY IN THE FIFTY (50) UNITED STATES, THE DISTRICT OF COLUMBIA AND IN THE COMMONWEALTH OF PUERTO RICO. WHAT WE WILL DO: If this product is found to be defective, JVC will repair or replace defective parts at no charge to the original owner. Such repair and replacement services shall be rendered by JVC during normal business hours at JVC authorized service centers. Parts used for replacement are warranted only for the remainder of the Warranty Period. All products and parts thereof may be brought to a JVC authorized service center on a carry-in basis except for Television sets having a screen size 25 inches and above which are covered on an in-home basis. WHAT YOU MUST DO FOR WARRANTY SERVICE: Return your product to a JVC authorized service center with a copy of your bill of sale. For your nearest JVC authorized service center, please call toll free: (800) 252-5722. If service is not available locally, box the product carefully, preferably in the original carton, and ship, insured, with a copy of your bill of sale plus a letter of explanation of the problem to the nearest JVC Factory Service Center, the name and location of which will be given to you by the toll-free number. If you have any questions concerning your JVC Product, please contact our Customer Relations Department. WHAT IS NOT COVERED: This limited warranty provided by JVC does not cover: 1. Products which have been subject to abuse, accident, alteration, modification, tampering, negligence, misuse, faulty installation, lack of reasonable care, or if repaired or serviced by anyone other than a service facility authorized by JVC to render such service, or if affixed to any attachment not provided with the products, or if the model or serial number has been altered, tampered with, defaced or removed; 2. Initial installation and installation and removal for repair; 3. Operational adjustments covered in the Owners Manual, normal maintenance, video and audio head cleaning; 4. Damage that occurs in shipment, due to act of God, and cosmetic damage; 5. Signal reception problems and failures due to line power surge; 6. Video Pick-up Tubes/CCD Image Sensor, Cartridge, Stylus (Needle) are covered for 90 days from the date of purchase; 7. Accessories; 8. Batteries (except that Rechargeable Batteries are covered for 90 days from the date of purchase);from the date of purchase); There are no other express warranties except as listed above. THE DURATION OF ANY IMPLIED WARRANTIES INCLUDING THE IMPLIED WARRANTY OF MERCHANTABILITY, IS LIMITED TO THE DURATION OF THE EXPRESS WARRANTY HEREIN. JVC SHALL NOT BE LIABLE FOR THE LOSS OF USE OF THE PRODUCT, INCONVENIENCE, LOSS OR ANY OTHER DAMAGES, WHETHER DIRECT, INCIDENTAL OR CONSEQUENTIAL (INCLUDING, WITHOUT LIMITATION, DAMAGE TO TAPES, RECORDS OR DISCS) RESULTING FROM THE USE OF THIS PRODUCT, OR ARISING OUT OF ANY BREACH OF THIS WARRANTY. ALL EXPRESS AND IMPLIED WARRANTIES, INCLUDING THE WARRANTIES OF MERCHANTABILITY AND FITNESS FOR PARTICULAR PURPOSE, ARE LIMITED TO THE WARRANTY PERIOD SET FORTH ABOVE. Some states do not allow the exclusion of incidental or consequential damages or limitations on how long an implied warranty lasts, so these limitations or exclusions may not apply to you. This warranty gives you specific legal rights and you may also have other rights which vary from state to state. JVC COMPANY OF AMERICA DIVISION OF JVC AMERICAS CORP. 1700 Valley Road Wayne, N.J. 07470

The Journal of Experimental Biology 206, 2749-The Company of Biologists Ltd doi:10.1242/jeb.00496
Swimming performance studies on the eastern Pacic bonito Sarda chiliensis, a close relative of the tunas (family Scombridae)

II. Kinematics

Hawkins J. Dowis1, Chugey A. Sepulveda2, Jeffrey B. Graham2 and Kathryn A. Dickson1,*
1Department of Biological Science, California State University Fullerton, Fullerton, CA 92834-6850, USA and 2Center for Marine Biotechnology and Biomedicine and Marine Biology Research Division, Scripps Institution of
Oceanography, University of California San Diego, La Jolla, CA 92093-0204, USA
*Author for correspondence (e-mail:kdickson@fullerton.edu)
Accepted 7 May 2003 Summary length. The lateral displacement and bending angle of The swimming kinematics of the eastern Pacic bonito each intervertebral joint during a complete tailbeat cycle Sarda chiliensis at a range of sustained speeds were analyzed to test the hypothesis that the bonitos swimming were determined for the bonito at a swimming speed mode differs from the thunniform locomotor mode of of 90cms1. The pattern of mean maximum lateral tunas. Eight bonito (fork length FL 47.52.1cm, mass displacement (zmax) and mean maximum bending angle 1.250.15kg) (mean S.D.) swam at speeds of (max) along the body in the bonito differed from that of 50130cms1 at 182C in the same temperatureboth chub mackerel Scomber japonicus and kawakawa controlled water tunnel that was used in previous studies tuna Euthynnus affinis; zmax was highest in the bonito. of tunas. Kinematics variables, quantied from 60Hz This study veries that S. chiliensis is a carangiform video recordings and analyzed using a computerized, twoswimmer and supports the hypothesis that the thunniform dimensional motion analysis system, were compared with locomotor mode is a derived tuna characteristic associated published data for similar sized tunas at comparable with changes in this groups myotomal architecture. The speeds. Bonito tailbeat frequency, tailbeat amplitude and nding that yaw and zmax were greater in the bonito than stride length all increased signicantly with speed. Neither in both mackerels and tunas suggests that swimming yaw (6.00.6%FL) nor propulsive wavelength (12065% kinematics in the bonito is not intermediate between that sh total length) varied with speed, and there were no of tunas and mackerels, as would be predicted on the basis mass or body-length effects on the kinematics variables of morphological characteristics. for the size range of bonitos used. Relative to similar sized yellown (Thunnus albacares) and skipjack (Katsuwonus pelamis) tunas at similar speeds, the bonito has a lower Key words: locomotion, swimming, kinematics, Scombridae, eastern Pacic bonito, Sarda chiliensis, thunniform, carangiform, tuna. tailbeat frequency, a higher yaw and a greater stride
Introduction Tunas (family Scombridae, tribe Thunnini) are the only teleost shes known to conserve metabolically derived heat to maintain the temperature of the slow-twitch myotomal locomotor muscle (red muscle, RM) elevated above ambient water temperature (Carey et al., 1971; Graham, 1973; Block, 1991). In addition, tuna RM is in a more anterior and medial position (closer to the vertebral column) than it is in other teleosts (Kishinouye, 1923; Graham et al., 1983; Ellerby et al., 2000; Graham and Dickson, 2000). Because this will reduce conductive heat loss from the RM across the body surface, the internalization of RM was hypothesized to have been a precursor to the evolution of regional endothermy in the tunas (Block et al., 1993; Block and Finnerty, 1994; Westneat and Wainwright, 2001). Swimming in tunas has been classied as thunniform locomotion, characterized by minimal lateral undulation of most of the body and thrust generation by rapid oscillations of the high-aspect-ratio caudal n (Fierstine and Walters, 1968; Lighthill, 1970; Webb, 1975; Lindsey, 1978). Many morphological specializations of tunas are associated with thunniform swimming. These include the anteriormedial RM, streamlined body shape, elongated myotomes, RMtendon skeleton connections, and narrow-necking of the caudal peduncle (Fierstine and Walters, 1968; Lighthill, 1969, 1970; Webb, 1975; Magnuson, 1978; Ellerby et al., 2000; Graham and Dickson, 2000; Westneat and Wainwright, 2001). It has been proposed that the RM position in tunas evolved to enhance swimming performance by affecting the mechanical

2750 H. J. Dowis and others
transfer of muscle contractile force to the backbone and caudal propeller (Westneat et al., 1993; Ellerby et al., 2000; Graham and Dickson, 2000). The transition to thunniform locomotion was hypothesized to have occurred prior to the evolution of endothermy, in response to changing oceanographic conditions (Graham and Dickson, 2000). Testing and distinguishing among these hypotheses requires knowledge of the tunas sister groups and mapping morphological characteristics onto a scombrid phylogeny (Block and Finnerty, 1994; Graham and Dickson, 2000). The Scombridae is composed of a monotypic subgroup (the buttery mackerel Gasterochisma melampus) and four tribes: Scombrini (mackerels), Scomberomorini (Spanish mackerels), Sardini (bonitos) and Thunnini (tunas) (Collette, 1978; Collette et al., 2001). According to phylogenies based on both morphological and gene-sequence data, the 15 species of tunas form a derived, monophyletic clade, and their closest relatives are the bonitos (Collette, 1978; Block et al., 1993; Finnerty and Block, 1995; Carpenter et al., 1995; Graham and Dickson, 2000; Collette et al., 2001). Because of this sister-taxon relationship, examination of the bonitos is essential for determining the sequence of character state changes that led to the specializations of the tunas. This study quanties swimming kinematics in the eastern Pacic bonito Sarda chiliensis so that the trait of thunniform locomotion can be mapped precisely onto the scombrid phylogeny. Studies of scombrid swimming kinematics have focused primarily on mackerels and tunas (Gray, 1933; Fierstine and Walters, 1968; Magnuson, 1970; Videler and Hess, 1984; Dewar and Graham, 1994b; Shadwick et al., 1998; Knower et al., 1999; Gibb et al., 1999; Donley and Dickson, 2000; Nauen and Lauder, 2000; Dickson et al., 2002). Donley and Dickson (2000) distinguished the kinematics of juvenile chub mackerel Scomber japonicus and kawakawa tuna Euthynnus affinis, and emphasized the importance of comparing similar-sized sh at comparable speeds. They found that, at the same speeds, the tuna swam with higher tailbeat frequencies, lower tailbeat amplitudes, lower stride lengths and less lateral displacement along most of the body, than did the chub mackerel. Some kinematics data have been reported for bonitos, but comparisons with tunas and mackerels led to conicting conclusions. The relationship between tailbeat frequency and speed for a 16cm total length (TL) Atlantic bonito Sarda sarda derived from data in Pyatetskiy (1970) was similar to that of similar sized (14.8 and 16.7cmTL) chub mackerel, but lower than that of a 16.2cm TL kawakawa tuna (Donley, 1999; Donley and Dickson, 2000). In contrast, Altringham and Block (1997) reported similar tailbeat frequencies in S. chiliensis and yellown tuna Thunnus albacares swimming in a large, cylindrical tank, but swimming speed and sh size, which both affect tailbeat frequency, varied interspecically. Block (personal observation, cited in Block and Finnerty, 1994) indicated that the Atlantic bonito swims in a more stiffbodied (tuna-like) fashion than mackerels do. The most comprehensive study to date of bonito swimming (Ellerby et al., 2000) used sonomicrometry and electromyography to measure RM strain and activity patterns, which have been correlated with swimming mode (Wardle et al., 1995; Altringham and Ellerby, 1999; Knower et al., 1999; Altringham and Shadwick, 2001), at several positions along the body in S. chiliensis (6071cm fork length, FL). Muscle activity patterns in the bonito were found to be more similar to those of tunas than to those of mackerels. On the other hand, based on the extent of maximum lateral displacement of ve points along the dorsal midline measured from videotapes of bonito swimming steadily in a large, open tank, Ellerby et al. (2000) concluded that the bonito swims in the carangiform mode like the mackerel. They found lateral displacement of the bonito to be similar to that of the Atlantic mackerel (Scomber scombrus, 3034cmTL) from Videler and Hess (1984) and greater than that of a 44cmFL yellown tuna derived from Dewar and Graham (1994b). From the existing data, it is not possible to determine unequivocally if the swimming mode of bonitos is more similar to that of mackerels or tunas, or if it is intermediate between the two. Therefore, thunniform locomotion cannot be mapped onto the scombrid phylogeny at a specic position to determine if that trait evolved before or after the divergence of the tunas and bonitos. Furthermore, because most kinematic variables vary with swimming speed or sh size, it is important that interspecic comparisons are made at the same speeds in similar sized individuals. Thus, the objective of the present study was to quantify the swimming kinematics of the eastern Pacic bonito at a range of controlled speeds, and then to compare the bonito to tunas of similar size that have been swum at similar speeds in the same respirometer (Dewar and Graham, 1994b; Knower, 1998; Knower et al., 1999) and to intervertebral lateral displacement and bending angle data for chub mackerel and kawakawa tuna (Donley and Dickson, 2000). Materials and methods Swimming kinematics This study utilized the same bonito Sarda chiliensis Cuvier as in the companion energetics study (Sepulveda et al., 2003), which describes the procedures for sh collection, handling and metabolic measurements in a large, temperaturecontrolled, swimming tunnel respirometer. After oxygen consumption rates were measured at a series of swimming speeds (experiments lasting up to 16h), the sh was allowed to swim steadily at a low speed for at least 15min before video analysis was initiated. The dorsal view of the sh was then videotaped at 60Hz, using a JVC super VHS camcorder (model GR-SXM520; Frys Electronics, Anaheim, CA, USA) mounted directly over the respirometer working section, for 215min at each speed, from 50cms1 to 130cms1 in increments of 10cms1. All sh did not swim at all of the speeds. During videotaping, respirometer water temperature was maintained at 182C, the temperature at which the bonitos were acclimated in the laboratory prior to experimentation.

Bonito swimming kinematics 2751
A Motus 3.2 motion analysis system (Peak Performance Technologies, Inc., Englewood, CO, USA) was used to analyze videotaped segments that met the following criteria: (i) the sh was positioned in the middle of the chamber, away from the walls and bottom, (ii) the sh was swimming steadily through at least seven complete tailbeat cycles, and did not move forward or backward in the chamber, and (iii) both the head and tail of the sh were in the eld of view. Two points, the tip of the upper lobe of the tail and the tip of the snout, were followed through time by digitizing sequential video frames for 714 complete tail beats at each swimming speed. Using the measured total length of the sh, a scaling factor was calculated for each video segment so that pixels could be converted to centimeters. Using methods described in Donley and Dickson (2000), kinematic variables were quantied for each bonito at each test speed. Tailbeat frequency (in Hz) was calculated by following the tip of the tail through time and dividing the number of consecutive tail beats by the amount of time, in seconds, that it took to complete those tail beats. Tailbeat amplitude (cm) and yaw (cm) were determined by measuring the distance between the lateral-most positions of the tip of the tail and of the tip of the snout, respectively, during a complete tailbeat cycle (the excursion of the tail from one side of the body to the other and back again). Mean tailbeat amplitude and yaw values were computed at each speed for each sh. Stride length, the distance (cm) that the sh moves forward in each tailbeat cycle, was calculated by dividing swimming speed by tailbeat frequency. Relative values (as %FL) of tailbeat amplitude, yaw and stride length were also determined. The propulsive wavelength (the length of the wave of undulation that travels down the body of the sh from snout to tail tip during swimming) was obtained by dividing propulsive wave velocity by the corresponding tailbeat frequency. Propulsive wave velocity was determined from the amount of time (s) between the peaks in lateral displacement at the tip of the snout and the tip of the tail; then the TL (cm) of each individual (the distance between the two points) was divided by the mean progression time in order to obtain the propulsive wave velocity in cms1 at each speed. Propulsive wavelength was measured in both cm and as a percentage of TL (%TL). Based on the known size of the eld of view that was videotaped and the resolution of 400 horizontal lines for super VHS, the spatial resolution of the measurements was 0.1250.183cm, which is equivalent to ranges of 0.250.41%FL and 0.230.38%TL. Depending on tailbeat frequency, there were 1840 video elds per tail beat. The lateral displacement and bending angle of each intervertebral joint during the tailbeat cycle were determined using the techniques of Jayne and Lauder (1995) and Donley and Dickson (2000). With the Peak Performance Motus system, 32 points approximately equally spaced around the dorsal outline of each individual were digitized in consecutive frames for one complete tailbeat cycle at a swimming speed of 90cms1 (relative speeds of 1.631.85TLs1), a speed at which all bonito swam. The points were converted into complete curves using a cubic spline function, and a midline was calculated for each frame (Jayne and Lauder, 1995). Lateral view X-rays were taken of each individual, and the lengths of the skull, each vertebra, and the hypural plate were measured with digital calipers. Each midline was divided into segments representing the measured skeletal elements, and the position in each frame of each intervertebral joint and of the snout and tail tip were calculated. Using a Microsoft Excel macro written by Jayne and Lauder (1995), the lateral displacement (z, using the terminology of Jayne and Lauder, 1995) and angle of exion () of each intervertebral joint throughout the tailbeat cycle were calculated. Then, maximum lateral displacement (zmax) and maximum bending angle (max) during the tail beat were determined for each joint along the body for each bonito. Mean zmax and max values for each intervertebral joint for the eight bonito studied were compared with data for all intervertebral joints in the chub mackerel and kawakawa tuna studied by Donley and Dickson (2000) at speeds of 75100cms1. For these comparisons, the maximum lateral displacement of each joint, measured in cm, was converted to relative sh length (%TL), and the position along the body of each intervertebral joint was expressed as %TL. The zmax at 0%TL is one-half of yaw, and zmax at 100%TL is one-half of tailbeat amplitude, as dened above. Statistical analysis The bonito kinematic variables were assessed for signicant effects of swimming speed (cms1) and sh size (mass in g and length in cm) and for signicant interactions between these factors. Minitab (version 13.1) was used to create the data le, calculate interaction terms and test for normality. SAS (version 8.2) was used to perform repeated-measures multiple regression analyses on tailbeat frequency (in Hz), tailbeat amplitude, yaw and stride length (in both cm and %FL), propulsive wavelength (in cm and %TL), and on zmax (in %TL) and max (in degrees). The initial statistical models for tailbeat frequency, tailbeat amplitude, yaw, stride length and propulsive wavelength included the main effects of speed, mass and FL, as well as all possible interaction terms. The models for zmax and max included mass, sh total length, position along the body, and all possible interactions, but did not include swimming speed because only one speed was used. For each variable, the full model was subjected to a backward stepwise reduction to t the best model to the data; each nonsignicant term was dropped until a nal model that included all signicant terms was obtained. We then determined if tailbeat frequency (in Hz), tailbeat amplitude, yaw and stride length (all expressed as %FL), and propulsive wavelength (in %TL) in the bonito differed signicantly from published data for two species of tuna (Thunnus albacares and skipjack Katsuwonus pelamis). Because there was limited access to the raw data for the tunas, two-sample t-tests (Dixon and Massey, 1969) were used to detect signicant interspecic differences in mean values or in the slopes and y-intercepts of linear regressions reported in the literature.

2752 H. J. Dowis and others
We tested for signicant interspecic differences in mean maximum lateral displacement of the body midline and mean maximum exion angles, at all intervertebral joints along the body, between the bonito and the chub mackerel and kawakawa tuna from Donley and Dickson (2000). Repeatedmeasures multiple regression analyses were run in SAS to determine if there were any signicant effects of position along the body (%TL), species, or position species on zmax and max. The position term was squared in order to incorporate curvature into the equation for a more accurate model of the data, but the coefficients of the squared terms were not interpreted. Signicant interspecic differences are indicated as signicant terms in the nal regression model, after a backward stepwise reduction process was completed. A signicance level of P=0.05 was used in all statistical analyses. Results Swimming kinematics of the eastern Pacic bonito A total of eight bonito were analyzed, FL=47.52.1cm, range 45.050.5cmFL, mass 1.250.15kg (means S.D.). When the effects of sh size were accounted for, tailbeat frequency increased signicantly with speed (P<0.0001) in the bonito (Fig.1). When the effects of speed were accounted for, there was no signicant effect of sh length or mass on tailbeat frequency. The range of tailbeat frequencies was 1.53.2Hz. Tailbeat amplitude, yaw and stride length were assessed for size and speed effects using the absolute values in cm, as well as relative values (%FL). When the effects of sh size were accounted for, both tailbeat amplitude and stride length increased signicantly with swimming speed (Figs2 and 3), but yaw did not vary signicantly with speed. When the effects of speed were accounted for, there were no signicant effects of FL or mass on tailbeat amplitude, yaw or stride length. When these three kinematics variables were expressed as %FL, no signicant size effects were detected. Tailbeat amplitude ranged from 16 to 24%FL, and the range of stride length was 6291%FL. The yaw for the bonito ranged from 5.2 to 6.9%FL (6.00.6%FL, mean S.D.). Propulsive wavelength (in cm and in %TL) did not vary signicantly with sh size or with swimming speed (P=0.065) in the bonito. Thus, a mean for each individual and a grand mean for all eight sh were calculated. The propulsive wavelength was 110129%TL (1206%TL, mean S.D.). Both mean maximum intervertebral lateral displacement (zmax) and mean maximum intervertebral bending angles (max) varied signicantly with position along the body (P<0.0001) in the bonito (Figs4 and 5). Minimum mean zmax occurred at 30%TL (the joint between vertebrae 11 and 12) and maximum zmax occurred at the tail tip. Minimum mean max occurred at 18%TL (the joint between the rst and second vertebrae) and maximum mean max was at the joint between the last vertebra and the hypural plate (92%TL). There were no signicant effects of sh mass or length on bonito zmax or max.

Tailbeat frequency (Hz)

100 Speed (cm s1) 120 140
Fig.1. Relationship between tailbeat frequency and swimming speed for Sarda chiliensis (each symbol denotes one individual) compared with (A) 42cm and 53cm yellown tuna Thunnus albacares (open triangles) (Dewar and Graham, 1994b) and (B) 4044cm yellown (open triangles) and 3841cm skipjack tuna Katsuwonus pelamis (solid triangles) (Knower et al., 1999). Best-t regression equation (regression coefficients S.D.) for the bonito data: tailbeat frequency = 0.0170.002 speed + 0.750.15 (N=8). Broken lines are 95% condence intervals of the regressions.
Interspecic comparisons Signicant effects of sh size were not detected for any of the kinematics variables measured in the present study, most likely due to the small size range of the bonito studied. Therefore, interspecic comparisons were made using the mean values of yaw, propulsive wavelength, and zmax and max at different positions along the body, and the regressions of the other kinematics variables versus swimming speed for the bonito. In these comparisons, we have assumed that temperature differences among the studies compared (with reported temperatures ranging from 18C to 28C) do not contribute signicantly to differences in tailbeat frequency, tailbeat amplitude, yaw and stride length. This assumption is based on a number of studies that have found little to no effect of temperature on these kinematics variables when sh are
Bonito swimming kinematics 2753
Mean maximum lateral displacement (%TL) Speed (cm s1) Tailbeat amplitude (%FL) Position along the body (%TL) 100
Fig.2. Tailbeat amplitude as a function of swimming speed for Sarda chiliensis (each symbol denotes one individual) and two sizes of yellown tuna, Thunnus albacares (inverted solid triangle for 42cm and upright solid triangle for 48cm) from Dewar and Graham (1994b). The solid line is the best-t regression equation (regression coefficients S.D.) for the bonito: tailbeat amplitude = 0.060.01 speed + 14.00.95 (N=8). Dotted lines represent 95% condence intervals of the regression. FL, fork length.
Fig.4. Maximum lateral displacement (means S.D.) at each intervertebral joint as a function of relative position along the body for Sarda chiliensis (solid triangles), kawakawa tuna Euthynnus affinis (open circles) and chub mackerel Scomber japonicus (open squares). Tuna and mackerel data are from Donley and Dickson (2000).

90 Stride length (%FL) 100 Speed (cm s1) 60 Position along body (%TL) Mean maximum intervertebral angle (degrees) 0
Fig.3. Stride length versus swimming speed for Sarda chiliensis (each symbol denotes one individual) and two species of tuna, yellown Thunnus albacares (upright solid triangles) and skipjack Katsuwonus pelamis (inverted solid triangles). Tuna data represented by solid lines are from Dewar and Graham (1994b) and by broken lines from Knower (1998). Best-t regression equation (regression coefficients S.D.) for the bonito: stride length = 0.320.08 speed + 51.156.98 (N=8). Dotted lines are the 95% condence intervals of this regression.
Fig.5. Maximum bending angle (means S.D.) at each intervertebral joint as a function of relative position along the body for Sarda chiliensis (solid triangles), kawakawa tuna Euthynnus affinis (open circles) and chub mackerel Scomber japonicus (open squares). Tuna and mackerel data are from Donley and Dickson (2000).
acclimated to the measurement temperature and comparisons are made at a given speed (for a review, see Dickson et al., 2002). Temperature does affect swimming performance in shes, primarily through changes in water viscosity that signicantly impact swimming at low Reynolds numbers
(Fuiman and Batty, 1997; Johnson et al., 1998) and by affecting muscle power output and patterns of muscle ber recruitment, leading to higher maximum sustainable speeds at higher temperatures (e.g. Rome and Swank, 1992; Altringham and Block, 1997; Rome et al., 2000). Tailbeat frequency increased signicantly with speed in both yellown and skipjack tunas (Dewar and Graham, 1994b; Knower 1998; Knower et al., 1999), as it did in the bonito (Fig.1). The slopes of the tailbeat frequency versus speed
2754 H. J. Dowis and others
Table1. Comparative data for yaw and propulsive wavelength (as a percentage of sh length) in scombrid shes
Yaw Species %FL %TL PWL %FL 8996 %TL PWL range %FL %TL Fish length (cm) FL 421.6 482.3841 15.125.5 (21.42.7) 45.050.16.027.1 TL Speed (cms1) 479.8 923.Speed range (body lengths s1) FLs1 ~1.1 ~1.9 1.12.7 1.53.7 1.65.0 TLs1 References 3
Tunas Yellown tuna 4.30.38 Thunnus albacares Yellown tuna 2.80.15 Thunnus albacares Yellown tuna Thunnus albacares Skipjack tuna Katsuwonus pelamis Kawakawa tuna 3.51.2 Euthynnus affinis Bonito Eastern Pacic bonito Sarda chiliensis Eastern Pacic bonito Sarda chiliensis Mackerels Atlantic mackerel Scomber scombrus Chub mackerel Scomber japonicus Chub mackerel Scomber japonicus 6.00.6 5.60.6 5.01

1202 1136

118134

110129

49.055.0

1.02.9

1.02.7

3.03.6 3.81.5 2.20.1062

84105 14.023.4 (20.33.4) 15.626.3 (20.94.0)

3034 14.825.3

30-105 1.45.9 1.44.8

3.911.2

Values are means S.D. or range (N.B. PWL values are means S.E.M.). FL, fork length; TL, total length; PWL, propulsive wavelength. 1Dewar and Graham (1994b); 2Knower (1998); 3Donley and Dickson (2000); 4This study; 5Ellerby et al. (2000); 6Videler and Hess (1984); 7Dickson et al. (2002).
relationships did not differ signicantly (P>0.25) between the bonito and two groups of yellown tuna (FL 421.6cm and 533.0cm, means S.D.) (data from Dewar and Graham, 1994b) or between the bonito and 4044cmFL yellown tuna and 3841cmFL skipjack tuna (data from Knower et al., 1999). Because the slopes did not differ, comparisons were made between the y-intercepts of these lines. The intercepts of the tailbeat frequency versus speed relationships for the 42cm and 53cm yellown tuna groups did not differ signicantly from that of the bonito (P>0.25) (Fig.1A), but the intercepts for the 4044cm yellown and 3841cm skipjack tuna were signicantly higher than for the bonito (P<0.0005) (Fig.1B). Because the original data from Knower et al. (1999) were provided to us, 95% condence intervals for the tailbeat frequency versus speed relationships were calculated and plotted (Fig.1B). The lack of overlap of the 95% condence intervals over the range of speeds studied suggests that the bonito swim at a given speed with signicantly lower tailbeat frequencies than do similar sized yellown and skipjack tunas. The only published data for tailbeat amplitudes and yaw at known speeds in tunas that are of comparable size to the bonito in the present study are from yellown tuna (Dewar and Graham, 1994b). Tailbeat amplitude did not vary signicantly with speed in the yellown tuna, and the values (mean S.D.)
of Dewar and Graham (1994b) are compared with the bonito data plotted in Fig.2. It appears that, at similar speeds, the tailbeat amplitude of the bonito does not differ signicantly from that of one group of yellown (FL 421.6cm, swimming at a speed of 402.8cms1; means S.D.) but is higher than that of the larger yellown (482.2cmFL, swimming at 1006.5cms1). The yaw for the bonito (6.00.6%FL, mean S.D.) is signicantly greater (P<0.001) than the values for both the 42cm (4.30.38%FL) and 48cm (2.80.15%FL) yellown tuna groups (Table1). A high yaw value (5%TL) was also observed for the eastern Pacic bonito by Ellerby et al. (2000). Stride length increased signicantly with speed in the yellown and skipjack tunas (Dewar and Graham, 1994b; Knower 1998), as it did in the bonito (Fig.3). The slopes of the stride length versus speed relationships did not differ signicantly (P>0.25) between the bonito and the two groups of yellown tuna from Dewar and Graham (1994b), or between the bonito and the yellown and skipjack tunas from Knower et al. (1999). The y-intercepts of the four tuna stride length versus speed regressions were signicantly lower than that of the bonito (P<0.001). Thus, at a given speed, stride length is greater in the bonito than it is in similar-sized tunas (Fig.3). The range of values for relative propulsive wavelength (as

Bonito swimming kinematics 2755
%FL) in the bonito (Table1) overlapped with data for yellown tuna from Dewar and Graham (1994b) (1249%FL and 12317%FL for 42cmFL and 48cmFL groups, respectively; means S.E.M.), but were higher than values reported by Knower (1998) (103%FL in 4044cmFL yellown tuna and 97%FL in 3841cmFL skipjack tuna). The patterns of mean maximum intervertebral lateral displacement (zmax) and mean maximum bending angle (max) at all intervertebral joints in the bonito were compared with data from juvenile kawakawa tuna and chub mackerel (Donley and Dickson, 2000), the only scombrid species that have been analyzed in this manner. Overall, mean zmax was signicantly higher in the bonito than in both the tuna (P<0.0001) and the mackerel (P=0.032). Lateral displacement at the snout and posterior to 40%TL is greater in the bonito than in the other two species (Fig.4). The pattern of zmax along the body differed between the bonito and the kawakawa tuna, as indicated by a signicant tuna position interaction (P=0.0025), but did not vary signicantly between the bonito and the chub mackerel. The minimum mean zmax occurred at 38%TL (between vertebrae 10 and 11) in the chub mackerel and at 41%TL (between vertebrae 15 and 16) in the kawakawa tuna, compared with 30%TL in the bonito; maximum mean zmax occurred at the tip of the tail in all three species. Mean maximum bending angles in the bonito differed signicantly from those in both the kawakawa tuna (P=0.0016) and the chub mackerel (P=0.032), as did the pattern of max versus position along the body, as indicated by signicant species position interaction terms (P<0.0001). The mackerel had higher bending angles than the bonito in the anterior third of the body, and the tuna had lower bending angles than the bonito at approximately 6575%TL (Fig.5). The position of minimum mean max was 40%TL (between vertebrae 11 and 12) in the chub mackerel and 18%TL (between vertebrae 1 and 2) in both the kawakawa tuna and the bonito; maximum mean max occurred at the intervertebral joint anterior to the hypural plate in all three species (at 87, 90 and 92%TL in the mackerel, tuna and bonito, respectively). Discussion The objective of this study was to characterize the swimming mode of the eastern Pacic bonito under controlled conditions (i.e. at a range of steady swimming speeds) and to compare specic kinematics variables between bonitos, tunas and mackerels. This study was designed to determine if the bonito utilizes a mode of locomotion similar to the thunniform mode used by tunas, in which minimal lateral displacement of the body occurs during steady swimming. Our analyses demonstrate that the swimming kinematics of the eastern Pacic bonito are signicantly different from those of yellown and skipjack tunas. Relative to comparably sized tunas swimming at similar speeds, the bonito swims with a lower tailbeat frequency, greater yaw, higher stride length and a greater degree of lateral displacement along the body. These ndings conrm the tentative conclusion from comparison of tailbeat frequency versus speed relationships (Donley, 1999; Donley and Dickson, 2000) and the conclusion of Ellerby et al. (2000), based on midline lateral displacement data, that the bonitos use the carangiform locomotor mode, and supports the hypothesis that thunniform locomotion is an autapomorphy of the tunas, associated with the anterior and medial RM position. Because the mackerels that have been studied are all less than 35cm in length, we cannot compare tailbeat frequency, tailbeat amplitude or stride length data for similar sized mackerels and bonitos swimming at comparable speeds. However, interspecic comparisons of the variables that are apparently independent of sh size, yaw and zmax expressed as a percentage of sh length, show that the bonito swims with signicantly more lateral displacement along most of the body, including the snout, than do juvenile chub mackerel. Thus, swimming kinematics in the bonito may not be intermediate between that of tunas and mackerels, as would be predicted on the basis of morphological characteristics. Swimming kinematics variables Although we concluded that tailbeat frequency at a given speed is signicantly lower in the bonito than it is in similar sized tunas, based on comparison with data for yellown and skipjack tunas from Knower et al. (1999), the relationships between tailbeat frequency and speed did not differ signicantly between the bonito and the yellown tuna from Dewar and Graham (1994b). There was much greater variability in the Dewar and Graham (1994b) data than in that of Knower et al. (1999), which may be due to the methods that were used to record tailbeat frequency. Dewar and Graham (1994b) used visual observations and a stopwatch to determine the time required for a sh to complete 20 tail beats while swimming steadily, whereas Knower et al. (1999) used frameby-frame analysis of video footage to calculate tailbeat frequency. The high variability may also be a consequence of the inclusion of lower swimming speeds by Dewar and Graham (1994b). If a sh swims at a speed that is below the minimum speed required for hydrostatic equilibrium (Magnuson, 1978), it may use sporadic swimming motions to maintain position, which can lead to high variability in kinematics data. Consequently, we believe that the values measured in Knower et al. (1999) are a more accurate representation of tuna tailbeat frequency at steady, sustainable speeds. However, it should be noted that the tunas used in Knower et al. (1999) all were smaller in FL than the bonito in the present study and thus would be expected to use higher tailbeat frequencies at a given speed than larger individuals. Because no size effects were observed in either study, it is not possible to extrapolate the data sets to a common sh size. When we compared the largest tuna (44cmFL yellown) from Knower et al. (1999) and our smallest bonito (45cmFL), tailbeat frequency was higher at a given speed in the tuna than in the bonito. Because tailbeat frequencies were lower in the bonito, we expected that tailbeat amplitudes would be higher in the bonito compared to tunas. The bonito did swim at a given speed with a higher tailbeat amplitude than the 48cm yellown, but there

2756 H. J. Dowis and others
was no difference between the bonito and 42cm yellown (Fig.2). In addition, the maximum lateral displacement of the tip of the tail (one-half of the tailbeat amplitude) was signicantly greater in the bonito than in the juvenile tuna of Donley and Dickson (2000) when expressed as %TL (Fig.4). Thus, the limited tailbeat amplitude data that are available provide some support for a difference in swimming mode between the bonito and tuna. Stride length in the bonito was higher than the values reported for tunas by both Dewar and Graham (1994b) and Knower et al. (1999), indicating that the bonito moves farther with each tailbeat. Altringham and Block (1997) also noted greater stride lengths in free-swimming bonito (4247cmTL) compared with larger yellown tuna (5881cmTL). These data further support the difference in swimming mode between tunas and the bonito. Yaw, the result of anterior recoil forces generated by oscillation of the tail, is minimized in scombrid shes by narrow necking of the caudal n, a large muscle mass and a high body depth (Lighthill, 1969; Lindsey, 1978; Webb, 1978, 1998). Magnuson (1978) found that maximum body thickness (the average of maximum height and maximum width) for seven tuna species ranged between 20.8 and 23.5%FL, but was only 18.4%FL for Sarda chiliensis and 16.0%FL for Scomber scombrus. Decreased yaw has been used to distinguish thunniform locomotion from other swimming modes (Fierstine and Walters, 1968; Dewar and Graham, 1994b; Ellerby et al., 2000). In the present study, the bonito had signicantly higher yaw than did similar sized yellown tuna at similar speeds, which supports the hypothesis that tunas utilize a different swimming mode than do bonitos. Because yaw (as a percentage of sh length) apparently does not vary signicantly with sh size, we examined yaw values from a number of other scombrid shes (Table1) and also found yaw in the bonito to exceed that in kawakawa tuna and Atlantic and chub mackerels. This interspecic difference is reected in the midline lateral displacement at the tip of the snout (one-half of yaw) (Fig.4). Thus, although morphological characteristics indicate that yaw in the bonito would be intermediate between that in mackerels and tunas, yaw was highest in the bonito. Propulsive wavelength values (as %FL or %TL) for tunas, mackerels and the eastern Pacic bonito overlap (Table1), and no consistent pattern was detected. Propulsive wavelength has been used previously to categorize swimming mode, and should be greater for thunniform than for carangiform swimmers (Lindsey, 1978). However, Donley and Dickson (2000) found that propulsive wavelength as a percentage of body length was greater in the chub mackerel than in the kawakawa tuna, and varied with sh size. Although propulsive wavelength in scombrids is not known to vary with swimming speed, it does vary with temperature (Dewar and Graham, 1994b; Donley and Dickson, 2000; Dickson et al., 2002). Studies with other sh species indicate that propulsive wavelength varies with axial position (Blight, 1977) and within a given species (Long and Nipper, 1996) suggest that this variable should not be used as a criterion for distinguishing sh swimming modes (see Long and Nipper, 1996; Donley and Dickson, 2000). The intervertebral exion angles were higher in the chub mackerel than in the bonito and in the kawakawa tuna (Fig.5). These angles reect intervertebral lateral displacement, the number of vertebrae and vertebral exibility. The larger angles in the mackerel can be attributed primarily to the smaller number of vertebrae in the chub mackerel (31) relative to the kawakawa (39) and the bonito (44); when there are fewer intervertebral joints, larger angles are required as the body midline is displaced laterally a given distance. In all three species, low max values were found for the intervertebral joints just anterior to the hypural plate, and max values at this position were highest in the chub mackerel because the mackerel has one relatively large vertebra in this position, whereas the tuna and bonito have two or three much shorter vertebrae (Collette, 1978). The interspecic differences in vertebral number may also contribute to differences in yaw and in the pattern of zmax along the body. Videler (1985) suggested that a greater number of vertebrae would lead to a greater degree of lateral exibility. This may explain why the bonito swims with greater lateral displacement than do the kawakawa tuna and chub mackerel (Fig.4). However, if vertebral number was the only factor involved, lateral displacement would be lowest in the chub mackerel, which has the fewest vertebrae of the three species, but lateral displacement is lowest in the tuna (Fig.4). The lower zmax and max values observed in the tuna may result from specializations for axial stiffness and/or the anteriormedial RM position. Relative to other scombrids, tunas have enlarged neural and hemal spines, larger zygapophyses that link adjacent vertebrae, more epipleural ribs and more extensive branching of tendons as they insert onto the backbone within the horizontal septum, and bony caudal keels which are thought to stiffen the caudal region (Kishinouye, 1923; Fierstine and Walters, 1968; Collette, 1978; Hebrank, 1982; Westneat et al., 1993). Tunas also have a well developed vertical septum containing collagen bers in a crossed-ber array, and some tuna species possess bony projections (lattices; Kishinouye, 1923) that extend between their hemal spines that may stiffen the skeleton (Westneat and Wainwright, 2001). Furthermore, in the tunas Euthynnus, Katsuwonus and Thunnus, the rst vertebra is partially or fully sutured to the skull (Collette, 1978). All of these characteristics may reduce axial exibility, but their contribution to differences in swimming kinematics remains to be determined empirically. Differences in swimming mode between mackerels, bonitos and tunas may also be related to the position of the RM, its pattern of activation, and how muscle contractile force is transferred to the skeleton to produce swimming movements. In bonitos and mackerels, the lateral RM is rmly attached to the skin and is connected to the backbone via posterior oblique tendons (POTs) within the horizontal septum that insert onto the backbone at higher angles than they do in tunas (Westneat et al., 1993; Graham and Dickson, 2000; Westneat and

Bonito swimming kinematics 2757
Wainwright, 2001). Contraction of RM therefore results in localized bending in the chub mackerel (Shadwick et al., 1998) and most likely also in the eastern Pacic bonito (Ellerby et al., 2000; Altringham and Shadwick, 2001). In the tunas, little of the RM is rmly attached to the skin and the POTs are longer and insert onto the backbone at a lower angle (Westneat et al., 1993; Graham and Dickson, 2000). Tuna RM transfers contractile force further caudally, allowing RM contraction in tunas to be uncoupled from local bending (Knower et al., 1999; Shadwick et al., 1999; Altringham and Shadwick, 2001). Because of the POT morphology, muscle contractile force is also transferred caudally with a higher velocity ratio, but a lower mechanical advantage, in tunas compared with mackerels and bonitos (Westneat et al., 1993; Graham and Dickson, 2000), and this is reected in the higher tailbeat frequencies and lower tailbeat amplitudes in tunas. Future studies are needed to test how differences in vertebral number, structures that affect axial stiffness, total muscle mass, RM position and connective tissue linkages between the locomotor muscle, skin and skeleton affect scombrid swimming kinematics. Conclusions The results of this kinematics study support the hypothesis that thunniform locomotion is a derived characteristic of the endothermic tunas associated with the anterior, medial position of the RM. The traits of anteriormedial RM, thunniform locomotion and endothermy all map onto the scombrid phylogeny after the divergence of the bonitos and tunas, and we cannot determine if the anteriormedial RM evolved initially for a less exible swimming mode and secondarily as a way to conserve metabolically derived heat. However, when combined with the swimming energetics data for the bonito (Sepulveda et al., 2003), we have shown that an increase in energetic efficiency is apparently not associated with thunniform locomotion. Sepulveda et al. (2003) found that the net cost of transport during sustained swimming was similar in the eastern Pacic bonito and the yellown tuna studied by Dewar and Graham (1994a), but that total metabolic costs were higher in the tuna due to a higher standard metabolic rate. This corresponds with the results of similar size-matched comparisons of juvenile chub mackerel and kawakawa tuna (Donley and Dickson, 2000; Sepulveda and Dickson, 2000; Korsmeyer and Dewar, 2001). Thus, there is no evidence that increased swimming efficiency was the selective advantage leading to the evolution of the thunniform locomotor mode. It may be that the advantages of endothermy, not swimming efficiency, led to the evolution of the anteriormedial RM in tunas, because heat loss from RM across the body surface would be reduced. If so, thunniform locomotion may simply be a consequence of changes in the biomechanical linkages of the locomotor muscle with the backbone and caudal propeller necessitated by this RM position. The next step in trying to determine at what point thunniform locomotion evolved within the family Scombridae will require making comparisons among the 15 tuna species. Although it is assumed that all of the tunas use thunniform locomotion, swimming in many tuna species has not been studied. Efforts should be directed at describing the swimming mode of the most basal tuna, Allothunnus fallai. Although this species does not possess all of the circulatory specializations for endothermy that are found in the other tunas, its RM is located in an anterior, medial position and a small central heat exchanger is present (Graham and Dickson, 2000). Characterizing the swimming kinematics of this species and determining if it is able to elevate RM temperature will establish whether the anteriormedial RM evolved prior to the evolution of thunniform locomotion or prior to the evolution of endothermy. This research was funded by NSF grant #IBN-9973916, the California State University (CSU) Fullerton Departmental Associations Council, an intramural grant from the CSU State Special Fund for Research, Scholarship and Creative Activity, the Scripps Institution of Oceanography Directors Office, and the Birch Aquarium at Scripps. Modications of the respirometer system were funded by NSF grants IBN-9607699 and IBN-0077502. Bonito were collected under California Department of Fish and Game scientic collecting permits. All experimental protocols were approved by the University of California San Diego and California State University Fullerton Institutional Animal Care and Use Committees. We thank D. Bernal, C. Chan, J. Donley and H. Lee for assistance with sh collection and maintenance and with the experiments, G. Noffal for use of the Peak Motus system, G. Lauder for providing the Excel programs used in the body bending analyses, J. Donley for her generous assistance with those programs and with data analysis, and K. Messer for invaluable statistical assistance. We are indebted to T. Knower for supplying tuna data and H.J. Walker for assistance with the sh X-rays. We also thank M. Horn, S. Murray, J. Videler and an anonymous reviewer who provided useful comments on drafts of the manuscript. References

Altringham, J. D. and Block, B. A. (1997). Why do tuna maintain elevated slow muscle temperatures? Power output of muscle isolated from endothermic and ectothermic sh. J. Exp. Biol. 200, 2617-2627. Altringham, J. D. and Ellerby, D. J. (1999). Fish swimming: Patterns in muscle function. J. Exp. Biol. 202, 3397-3403. Altringham, J. D. and Shadwick, R. E. (2001). Swimming and muscle function. In Tuna Physiology, Ecology, and Evolution, Fish Physiology, vol. 19 (ed. B. A. Block and E. D. Stevens), pp. 314-341. New York: Academic Press. Blight, A. R. (1977). The muscular control of vertebrate swimming movements. Biol. Rev. 52, 181-218. Block, B. A. (1991). Endothermy in sh: thermogenesis, ecology, and evolution. In Biochemistry and Molecular Biology of Fishes, vol. 1 (ed. P. W. Hochachka and T. P. Mommsen), pp. 269-311. New York: Elsevier. Block, B. A. and Finnerty, J. R. (1994). Endothermy in shes: A phylogenetic analysis of constraints, predispositions, and selection pressures. Environ. Biol. Fish. 40, 283-302. Block, B. A., Finnerty, J. R., Stewart, A. F. R. and Kidd, J. (1993). Evolution of endothermy in sh: mapping physiological traits on a molecular phylogeny. Science 260, 210-214.
2758 H. J. Dowis and others
Carey, F. G., Teal, J. M., Kanwisher, J. W., Lawson, K. D. and Beckett, K. S. (1971). Warm bodied sh. Amer. Zool. 11, 137-145. Carpenter, K. E., Collette, B. B. and Russo, J. L. (1995). Unstable and stable classications of scombrid shes. Bull. Mar. Sci. 56, 379-405. Collette, B. B. (1978). Adaptations and systematics of the mackerels and tunas. In The Physiological Ecology of Tunas (ed. G. D. Sharp and A. E. Dizon), pp. 7-39. New York: Academic Press. Collette, B. B., Reeb, C. and Block, B. A. (2001). Systematics of the tunas. In Tuna Physiology, Ecology, and Evolution, Fish Physiology, vol. 19 (ed. B. A. Block and E. D. Stevens), pp. 5-30. New York: Academic Press. Dewar, H. and Graham, J. B. (1994a). Studies of tropical tuna swimming performance in a large water tunnel. I. Energetics. J. Exp. Biol. 192, 13-31. Dewar, H. and Graham, J. B. (1994b). Studies of tropical tuna swimming performance in a large water tunnel. III. Kinematics. J. Exp. Biol. 192, 4559. Dickson, K. A., Donley, J. M., Sepulveda, C. and Bhoopat, L. (2002). Effects of temperature on sustained swimming performance and swimming kinematics of the chub mackerel Scomber japonicus. J. Exp. Biol. 205, 969980. Dixon, W. J. and Massey, F. J. (1969). Introduction to Statistical Analysis. New York: McGraw-Hill Book Company. Donley J. M. (1999). Swimming kinematics of two scombrid shes. MS thesis, California State University, Fullerton, USA. Donley, J. M. and Dickson, K. A. (2000). Swimming kinematics of juvenile kawakawa tuna (Euthynnus affinis) and chub mackerel (Scomber japonicus). J. Exp. Biol. 203, 3103-3116. Ellerby, D. J., Altringham, J. D., Williams, T. and Block, B. A. (2000). Slow muscle function of Pacic bonito (Sarda chiliensis) during steady swimming. J. Exp. Biol. 203, 1-13. Fierstine, H. L. and Walters, V. (1968). Studies in locomotion and anatomy of scombroid shes. Mem. S. Calif. Acad. Sci. 6, 1-31. Finnerty, J. R. and Block, B. A. (1995). Evolution of cytochrome b in the Scombroidei (Teleostei): molecular insights into billsh (Istiophoridae and Xiphiidae) relationships. Fish. Bull. US 93, 78-96. Fuiman, L. A. and Batty, R. S. (1997). What a drag it is getting cold: Partitioning the physical and physiological effects of temperature on sh swimming. J. Exp. Biol. 200, 1745-1755. Gibb, A. C., Dickson, K. A. and Lauder, G. V. (1999). Tail kinematics of the chub mackerel, Scomber japonicus: testing the homocercal tail model of sh propulsion. J. Exp. Biol. 202, 2433-2447. Graham, J. B. (1973). Heat exchange in the black skipjack and the blood-gas relationship of warm-bodied shes. Proc. Natl. Acad. Sci. USA 70, 19641967. Graham, J. B., Koehrn, F. J. and Dickson, K. A. (1983). Distribution and relative proportions of red muscle in scombrid shes: consequences of body size and relationships to locomotion and endothermy. Can. J. Zool. 61, 2087-2096. Graham, J. B. and Dickson, K. A. (2000). The evolution of thunniform locomotion and heat conservation in scombrid shes: New insights based on the morphology of Allothunnus fallai. Zool. J. Linn. Soc. 129, 419-466. Gray, J. (1933). Studies in Animal Locomotion. I. The movements of sh with special reference to the eel. J. Exp. Biol. 10, 88-104. Hebrank, M. R. (1982). Mechanical properties of sh backbones in lateral bending and in tension. J. Biomech. 15, 85-89. Jayne, B. C. and Lauder, G. V. (1995). Speed effects on midline kinematics during steady undulatory swimming of largemouth bass, Micropterus salmoides. J. Exp. Biol. 198, 585-602. Johnson, T. P., Cullum, A. J. and Bennett, A. F. (1998). Partitioning the effects of temperature and kinematic viscosity on the C-start performance of adult shes. J. Exp. Biol. 201, 2045-2051. Kishinouye, K. (1923). Contributions to the comparative study of the socalled scombroid shes. Jour. Coll. Agric., Tokyo Imperial Univ. 8, 293475. Knower, T. (1998). Biomechanics of thunniform swimming: electromyography, kinematics, and caudal tendon function in the yellown tuna and the skipjack tuna. PhD dissertation, Scripps Institution of Oceanography, California, USA. Knower, T., Shadwick, R. E., Katz, S. L., Graham, J. B. and Wardle, C. S. (1999). Red muscle activation patterns in yellown (Thunnus albacares) and skipjack (Katsuwonus pelamis) tunas during steady swimming. J. Exp. Biol. 202, 2127-2138. Korsmeyer, K. E. and Dewar, H. (2001). Tuna metabolism and energetics. In Tuna Physiology, Ecology, and Evolution, Fish Physiology, vol. 19 (ed. B. A. Block and E. D. Stevens), pp. 36-71. New York: Academic Press. Lighthill, M. J. (1969). Hydromechanics of aquatic animal propulsion. Ann. Rev. Fluid Mech. 1, 413-446. Lighthill, M. J. (1970). Aquatic animal propulsion of high hydromechanical efficiency. J. Fluid Mech. 44, 265-301. Lindsey, C. C. (1978). Form, function, and locomotory habits in sh. In Fish Physiology, vol. 7 (ed. W. S. Hoar and D. J. Randall), pp. 1-13. New York: Academic Press. Long, J. H., Jr and Nipper, K. S. (1996). Body stiffness in undulating vertebrates. Amer. Zool. 36, 678-694. Magnuson, J. J. (1970). Hydrostatic equilibrium of Euthynnus affinis, a pelagic teleost without a swim bladder. Copeia 1970, 56-85. Magnuson, J. J. (1978). Locomotion by scombrid shes: Hydromechanics, morphology and behavior. In Fish Physiology, vol. 7 (ed. W. S. Hoar and D. J. Randall), pp. 240-315. New York: Academic Press. Nauen, J. C. and Lauder, G. V. (2000). Locomotion in scombrid shes: morphology and kinematics of the nlets of the chub mackerel Scomber japonicus. J. Exp. Biol. 203, 2247-2259. Pyatetskiy, V. Y. (1970). Kinematic swimming characteristics of some fast marine sh. In Hydrodynamic problems in Bionics, Bionika, vol. 4, Kiev (translated from Russian, JPRS 52605, pp. 12-23. Natl. Tech. Inf. Serv., Springeld, Virginia, 1971). Rome, L. C. and Swank, D. (1992). The inuence of temperature on power output of scup red muscle during cyclical length changes. J. Exp. Biol. 171, 261-281. Rome, L. C., Swank, D. M. and Coughlin, D. J. (2000). The inuence of temperature on power production during swimming. II. Mechanics of red muscle bres in vivo. J. Exp. Biol. 203, 333-345. Sepulveda, C. and Dickson, K. A. (2000). Maximum sustainable speeds and cost of swimming in juvenile kawakawa tuna (Euthynnus affinis) and chub mackerel (Scomber japonicus). J. Exp. Biol. 203, 3089-3101. Sepulveda, C., Dickson, K. A. and Graham, J. B. (2003) Swimming performance studies on the eastern Pacic bonito (Sarda chiliensis), a close relative of the tunas (family Scombridae). I. Energetics. J. Exp. Biol. 206, 2739-2748. Shadwick, R. E., Katz, S. L., Korsmeyer, K. E., Knower, T. and Covell, J. W. (1999). Muscle dynamics in skipjack tuna: timing of red muscle shortening in relation to activation and body curvature during steady swimming. J. Exp. Biol. 202, 2139-2150. Shadwick, R. E., Steffensen, J. F., Katz, S. L. and Knower, T. (1998). Muscle dynamics in sh during steady swimming. Amer. Zool. 38, 755-770. Videler, J. J. (1985). Fish swimming movements: A study of one element of behaviour. Neth. J. Zool. 35, 470-485. Videler, J. J. and Hess, F. (1984). Fast continuous swimming of two pelagic predators saithe (Pollachius virens) and mackerel (Scomber scombrus): a kinematics analysis. J. Exp. Biol. 109, 209-228. Wardle, C. S., Videler, J. J. and Altringham, J. D. (1995). Tuning into sh swimming waves: body form, swimming mode and muscle function. J. Exp. Biol. 198, 1629-1636. Webb, P. W. (1975). Hydrodynamics and energetics of sh propulsion. Bull. Fish. Res. Bd. Can. 190, 1-159. Webb, P. W. (1978). Hydrodynamics: nonscombroid sh. In Fish Physiology, vol. 7 (ed. W. S. Hoar and D. J. Randall), pp. 190-237. New York: Academic Press. Webb, P. W. (1998). Swimming. In The Physiology of Fishes, 2nd Edition (ed. D. H. Evans), pp. 3-24. Boca Raton: CRC Press. Westneat, M. W., Hoese W., Pell, C. A. and Wainwright, S. A. (1993). The horizontal septum: Mechanisms of force transfer in locomotion of scombrid shes (Scombridae, Perciformes). J. Morphol. 217, 183-204. Westneat, M. W. and Wainwright, S. A. (2001). Mechanical design for swimming: muscle, tendon, and bone. In Tuna Physiology, Ecology, and Evolution, Fish Physiology, vol. 19 (ed. B. A. Block and E. D. Stevens), pp. 272-308. New York: Academic Press.