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Facultative Cleaning by the Forcepsfish, Forcipiger Flavissimus (Chaetodontidae)

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Matthew Thomas Craig at NOAA National Marine Fisheries Service
Matthew Thomas Craig
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Abstract

On 7 June 2006 a specimen of Forcepsfish, Forcipiger flavissimus, was observed engaging in cleaning behavior of a Bigscale Soldierfish, Myripristis berndti (Holocentridae). This behavior is described here as novel for the species. Anecdotal reports of similar behavior indicate that this species engages in cleaning with several hosts including other fishes and sea turtles. Reports of this behavior at other localities and among a variety of butterflyfish species indicate that cleaning may be a more important foraging strategy than previously thought.
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Copeia, 2007(2), pp. 459–461
Facultative Cleaning by the Forcepsfish,
Forcipiger flavissimus (Chaetodontidae)
MATTHEW T. CRAIG
On 7 June 2006 a specimen of Forcepsfish, Forcipiger flavissimus, was observed
engaging in cleaning behavior of a Bigscale Soldierfish, Myripristis berndti (Holocen-
tridae). This behavior is described here as novel for the species. Anecdotal reports of
similar behavior indicate that this species engages in cleaning with several hosts
including other fishes and sea turtles. Reports of this behavior at other localities and
among a variety of butterflyfish species indicate that cleaning may be a more important
foraging strategy than previously thought.
C
LEANING by marine fishes is a commonly
observed behavior in reef systems. At least
131 species of fishes and crustaceans have been
reported to perform this activity (Coˆte´, 2000).
This behavior has been observed in a wide variety
of fish families, although it is usually associated
with Labridae, such as the common cleaner
wrasses of the genus Labroides. While some fishes
utilize this behavior as an exclusive means of
foraging (Coˆte´, 2000), others perform this task
when conditions favor this behavior. Such be-
havior is termed facultative cleaning and has
been observed in several families including the
Labridae, Gobiidae, and Chaetodontidae. On 7
June 2006 while collecting fishes at Johnston
Atoll (north-central Pacific), a specimen of
Forcepsfish, Forcipiger flavissimus (Chaetodonti-
dae), was observed cleaning a large Bigscale
Soldierfish, Myripristis berndti (Holocentridae).
The behavior was noted as atypical for the
species, and subsequent dives were conducted
to photograph and observe the interaction.
M
ATERIALS AND METHODS
The behavior was first observed at Johnston
Atoll (16u459N, 169u319W) in a large cave
approximately 4.0 m wide by 5.0 m high in
30 m water depth at approximately 0930 hrs.
The cave opened to the bottom of a steep drop-
off and was fully connected with the upper reef
flat by a hole in the ‘‘roof.’’ The light levels were
generally low, and the dominant species within
the cave was Myripristis berndti. Three subsequent
dives were made to observe the behavior, and
photographs were taken to portray the interac-
tion in greater detail. In total, the cleaning
behavior was observed over 10.5 hours by four
divers.
R
ESULTS
Bigscale Soldierfish exhibited typical daytime
shoaling behavior inside the large cave described
above. In the presence of divers, the soldierfish
did not immediately posture for cleaning, and
required some adjustment period before they
attempted to solicit cleaning from the Forceps-
fish. The soldierfish slowly approached the
cleaner, opened its mouth to full gape, depressed
its first dorsal fin, and extended its second dorsal,
pelvic, and pectoral fins. The pectoral fins were
used in a slow sculling motion to maintain a still
position. The Forcepsfish then approached the
soldierfish and inserted its protruding snout fully
into the soliderfish’s mouth and plucked items
from inside the mouth and from the gill rakers.
On more than one occasion, the Forcepsfish was
mobbed by several (3–8) soldierfish. Cleaning
sessions lasted approximately 5–25 s. Photo-
graphs of the behavior were taken although the
strobe consistently distracted the fish and caused
a cessation in the cleaning behavior (Fig. 1).
Cleaning behavior was observed at two sites
that were approximately 5 km apart. Interesting-
ly, while both M. berndti and F. flavissimus were
present in high numbers on shallow portions of
the reef (10–15 m), the cleaning behavior was
only seen at depth (approximately 30 m at both
sites) within the confines of caves where light
levels were low. Even more noteworthy is that
cleaning behavior occurred in the presence of
the Cleaner Wrasse Labroides phthirophagus.In
many instances, the soldierfish ignored the
#
2007 by the American Society of Ichthyologists and Herpetologists
cleaner wrasse, and preferentially postured for
the Forcepsfish.
D
ISCUSSION
The feeding mechanics of butterflyfishes with
elongate jaws have been described in detail
(Ferry-Graham et al., 2001). While the prevailing
hypothesis to explain this adaptation is that it
provides an advantage when foraging in reef
crevices and cracks and on the tube feet of sea
urchins, the current observation adds another
means by which a protruding snout may be
advantageous. While this is the first documented
case of Forcepsfish cleaning behavior, casual
observations indicate that this species interacts
with and cleans a variety of hosts such as
trumpetfishes (Aulostomidae), surgeonfishes
(Acanthuridae), and Green Sea Turtles (Chelonia
mydas; J. Hoover and J. Randall, pers. comm.).
Observations of the Forcepsfish cleaning Bigscale
Soldierfish on the north shore of O’ahu, Hawaii
indicate that this interaction is not unique to
Johnston Atoll (C. Meyer, pers. comm.).
Cleaning is listed as a common behavior for
relatively few chaetodontids (e.g., the Barberfish,
Johnrandallia nigrirostris; Hobson, 1969; Burgess,
1978; Thompson et al., 2000). However, cases
have been reported in which chaetodontids
function as cleaners, including the eastern
Pacific Scythe Butterflyfish, Prognathodes falcifer
(Lea and Richards, 2005), the central-western
Atlantic Banded Butterflyfish, Chaetodon striatus
(Sazima and Sazima, 2001), and others (van
Tassell et al., 1994; Coˆte´, 2000). In some species,
this behavior has only been observed in juveniles
(e.g., the Indo-Australian Chaetodon plebius, the
Indo-Pacific Heniochus diphrutes, and the Easter
Island endemic Chaetodon litus; Allen et al.,
1998) indicating an ontogenetic component to
this foraging strategy. Given the numerous rec-
ords of cleaning behavior at scattered locations
that involve several genera and species, it seems
that cleaning may be a much more important
part of chaetodontid foraging than previously
thought.
A
CKNOWLEDGMENTS
This observation was made while surveying
fishes as part of the Northwestern Hawaiian
Islands Marine National Monument coral reef
research partnership (MOA-2005-008/6882 and
MOA-2005-008/7189). B. Bowen, J. Hoover, S.
Karl, R. Kosaki, C. Meyer, J. Randall, L. Rocha, J.
Schultz, and J. Zamzow provided additional
observations in the field and reports of cleaning
behavior by the Forcepsfish. J. Hyde, S. Karl, R.
Lea, D. Pondella, II, and L. Rocha provided
useful comments on earlier versions of this
manuscript. I thank the officers and crew of the
NOAA Ship Hi’ialakai who provided support at
sea. This is SOEST contribution No. 6919 and
HIMB contribution No. 1244.
L
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ALLEN, G. R., R. STEENE, AND M. ALLEN. 1998. A Guide
to Angelfishes and Butterflyfishes. Odyssey Pub-
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B
URGESS, W. E. 1978. Butterflyfishes of the World.
T.F.H. Publications Inc., Ltd., Neptune City, New
Jersey.
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ERRY-GRAHAM, L., P. C. WAINWRIGHT,C.D.HULSEY,
AND D. R. BELLWOOD. 2001. Evolution and mechan-
ics of long jaws in butterflyfishes (Family Chaeto-
dontidae). Journal of Morphology 248:120–143.
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OBSON, E. S. 1969. Comments on certain recent
generalizations regarding cleaning symbiosis in
fishes. Pacific Science 23:35–39.
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EA, R. N., AND D. V. RICHARDS. 2005. The Scythe
Butterflyfish, Prognathodes falcifer (Pisces: Chaeto-
dontidae), observed as a facultative cleaner. Bulle-
Fig. 1. Photographs of Forcepsfish, Forcipiger flavissimus (Chaetodontidae), cleaning the Bigscale
Soldierfish, Myripristis berndti (Holocentridae), at Johnston Atoll.
460 COPEIA, 2007, NO. 2
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104:152–153.
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AZIMA, C., AND I. SAZIMA. 2001. Plankton-feeding
aggregation and occasional cleaning by adult
butterflyfish, Chaetodon striatus (Chaetodontidae),
in southwestern Atlantic. Cybium 25:145–151.
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HOMSON, D. A., L. T. FINDLEY, AND A. N. KERSTITCH.
2000. Reef Fishes of the Sea of Cortez: The Rocky
Shore Fishes of the Gulf of California, Revised
Edition. The University of Texas Press, Austin.
VAN TASSELL, J. L., A. BRITA, AND S. A. BORTONE. 1994.
Cleaning behavior among marine fishes and
invertebrates in the Canary Islands. Cybium
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HAWAII INSTITUTE OF MARINE BIOLOGY, P.O. 1346,
K
ANEOHE,HAWAII 96744. E-mail: mtcraig@
hawaii.edu. Submitted: 28 Sept. 2006. Accepted:
22 Jan. 2007. Section editor: J. Snodgrass.
CRAIG—FACULTATIVE CLEANING BY FORCEPSFISH 461
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Full-text available
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  • CYBIUM
Butterflyfishes of the genus Chaetodon (Chaetodontidae) are mostly bottom-feeders, and juveniles of some Pacific species are known to clean other fish. Herein we report on occasional cleaning by adult individuals of the Western Atlantic butterflyfish Chaetodon striatus, during their plankton-feeding aggregation. Four species of reef fish solicited cleaning to the butterflyfish amidst the aggrega-tion. RÉSUMÉ.-Rassemblement alimentaire pour la capture de plancton et nettoyage occasionnel par des Chaetodon striatus (Chaetodontidae) adultes, dans l'Atlantique occidental. La plupart des poissons-papillons du genre Chaetodon (Chaetodontidae) recherchent leur nour-riture sur le fond et les jeunes de quelques espèces du Pacifique sont connus comme nettoyeurs d'autres poissons. Nous avons observé un comportement de nettoyage occasionnel pratiqué par des Chaetodon striatus adultes, dans l'Atlantique occidental, quand ils se regroupent pour capturer des organismes planctoniques. Quatre espèces de poissons récifaux sont venus se faire nettoyer par le groupe de chaeto-dons.
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Evolution and Mechanics of Long Jaws in Butterflyfishes (Family Chaetodontidae)
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  • May 2001
  • J MORPHOL
We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger. J. Morphol. 248:120–143, 2001. © 2001 Wiley-Liss, Inc.
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Comments on Certain Recent Generalizations Regarding Cleaning Symbiosis in Fishes
Article
  • Jan 1969
Many fishes in the marine environment remove ectoparasites, diseased tissue, and other deleterious material from the bodies of other aquatic animals (Eibl-Eibesfeldt, 1955; Randall, 1958; Limbaugh, 1955; and others). In reporting on these "cleaners," Limbaugh (1961), in a posthumous report edited by Howard M. Feder, attempted some broad generalizations that contrast cleaning in tropical and temperate seas; these were later presented by Feder (1966) in his review of cleaning symbiosis in the marine environment. These generalizations are in need of critical review, as they are now becoming entrenched in the literature (e.g., Gotshall, 1967; Marshall, 1965; Ommanney, 1966).
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E-mail: mtcraig@ hawaii.edu. Submitted: 28 Sept
  • Feb 2006
  • 22
  • Hawaii Institute
  • P O Marine Biology
HAWAII INSTITUTE OF MARINE BIOLOGY, P.O. 1346, KANEOHE, HAWAII 96744. E-mail: mtcraig@ hawaii.edu. Submitted: 28 Sept. 2006. Accepted: 22 Jan. 2007. Section editor: J. Snodgrass.
A Guide to Angelfishes and Butterflyfishes
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ALLEN, G. R., R. STEENE, AND M. ALLEN. 1998. A Guide to Angelfishes and Butterflyfishes. Odyssey Publishing, Sydney.
Evolution and ecology of cleaning symbiosis in the sea
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COˆTÉCOˆCOˆTÉ, I. M. 2000. Evolution and ecology of cleaning symbiosis in the sea. Oceanography and Marine Biology Annual Review 38:311-355.