15 Sea Turtle Epibiosis
Michael G. Frick and Joseph B. Pfaller
In the marine environment, any exposed, undefended surface will eventually be colonized by marine
propagules (Wahl, 1989). Colonization of inanimate structures (e.g., dock pilings and boat hulls) is
called fouling, while colonization of other marine organisms is called epibiosis. Epibiosis results in
spatially close associations between two or more living organisms (Harder, 2009), in which a single
host (or basibiont) supports one or more typically opportunistic colonizers (or epibionts) (Wahl
and Mark, 1999). Epibiosis is the most common form of symbiosis in the marine environment and
15.1 Introduction ..........................................................................................................................399
15.2 Common Forms ....................................................................................................................401
15.2.1 Sessile Forms ............................................................................................................401
15.2.2 Sedentary Forms ....................................................................................................... 401
15.2.3 Motile Forms ............................................................................................................401
15.3 Communities and Community Dynamics ............................................................................ 402
15.3.1 Pelagic/Oceanic Communities ..................................................................................402
15.3.2 Benthic/Neritic Communities ...................................................................................402
15.3.3 Obligate Communities ..............................................................................................403
15.3.4 Community Distribution ...........................................................................................403
15.3.5 Community Succession ............................................................................................404
15.4 Ecological Interactions .........................................................................................................405
15.4.1 Effects on Epibionts ..................................................................................................405
15.4.2 Effects on Host Turtles .............................................................................................405
15.4.3 Ecological Inferences ...............................................................................................406
15.4.4 Ecological Implications ............................................................................................407
15.5 Conceptual Model of Epibiosis .............................................................................................408
15.5.1 Geographic Overlap ..................................................................................................408
15.5.2 Ecological Overlap ...................................................................................................408
15.5.3 Balance of Costs and Benets ..................................................................................409
15.6 Considerations ...................................................................................................................... 410
Acknowledgments .......................................................................................................................... 410
15.A Appendix A: Annotated Bibliography of Selected Sea Turtle Epibiont Studies
and Reports Listed by Geographic Region ........................................................................... 411
15.A.1 Caribbean–Western Atlantic..................................................................................... 411
15.A.2 Mediterranean–Eastern Atlantic .............................................................................. 411
15.A.3 Indo–West Pacic ..................................................................................................... 411
15.A.4 Eastern Pacic .......................................................................................................... 412
References ...................................................................................................................................... 412
400 The Biology of Sea Turtles, Volume III
may be classied into several types of associations (e.g., mutualism, commensalism, parasitism)
depending on the interactions between a host and its epibionts (Leung and Poulin, 2008).
Sea turtles often act as hosts to a wide variety of epibionts, most of which are unspecialized
organisms normally found associated with inanimate structures in the surrounding marine
environment (i.e., “free living”). These types of epibiotic associations are known as facultative
commensalisms (Wahl and Mark, 1999). That is, the host receives no direct benet from the epibiont
and the epibiont demonstrates little to no substrate specicity. For these associations to occur, the
various settlement cues that facultative commensal epibionts utilize when selecting substrata must
also be present on sea turtles (Zardus and Hadeld, 2004). Alternatively, there are several epibionts
that are found almost exclusively on sea turtles (Frick et al., 2011a). These associations are known
as obligate commensalisms, whereby the epibiont is dependent on the host turtle for survival, but the
welfare of the host turtle is not dependent on the presence or behavior of the epibiont. While some
obligate commensal epibionts are known to perform activities that might be considered benecial
to the host turtle, there are no examples of obligate mutualisms, in which both the host turtle and
the epibiont depend on each other for survival. Future studies, however, may identify such obligate
mutualisms. Most obligate (and facultative) commensal epibionts do not derive nutrients from the
tissue of the host turtle and are not parasitic; instead the host turtle simply provides a foraging
platform (Frick et al., 2002a). On the contrary, several sea turtle epibionts are known to derive
nutrients from the tissue of the host turtle and, therefore, represent associations known as parasitism
(Leung and Poulin, 2008). Parasitic epibionts of sea turtles are rare, but these associations may have
important consequences for the health of host turtles (Greenblatt et al., 2004).
Following a rich history of anecdotal reports dating back to Darwin (1851, 1854), the study of
epibiosis in sea turtles has received considerable attention in recent years. The vast majority of studies
describe the diversity of epibiota, and speculate on the possible causes and effects of these associations.
From these descriptive studies, we have learned a great deal with respect to the wonderful diversity of
epibiotic forms found associated with sea turtles (Appendix A). Fewer studies, however, approach sea
turtle epibiosis from the community perspective. These studies not only describe diversity of epibiota
but also consider the structuring of epibiotic communities and the complex suite of interactions
occurring on the turtle across space and time. Finally, even fewer studies attempt to quantify and
understand the ecological interactions between turtles and their epibiota. These studies have allowed
researchers to better understand the ecological and evolutionary implications of epibiosis, and to
decipher the valuable information that can be gleaned from studying sea turtle epibionts.
Despite the antiquity of some sea turtle epibiont observations, the study of sea turtle epibiosis remains
in a prolonged state of infancy when compared to the breadth of information that has recently and
quickly accrued on sea turtle migrations and home ranges (largely through the deployment of satellite
tags). Likewise, our understanding of sea turtle genetics and molecular phylogeny exceeds that of basic
facets of sea turtle ecology—including diet, foraging behavior, and epibiotic associations. Given the
documented declines of turtle populations in some areas, it has become imperative for scientists to
understand how sea turtles interact with the constituents of the habitats they occupy, be it while foraging
or through epibiosis. Such information allows scientists to view sea turtles within the context of a
complex and ecologically rich marine environment, and it aids in modeling the potential impacts that
certain natural and anthropogenic-driven events may have upon sea turtles and the habitats they utilize.
In this chapter, we begin by introducing many of the common forms of epibionts known to be
associated with sea turtles. Second, we describe several common epibiotic community types, and
discuss the spatial and temporal factors by which epibiotic communities are structured. Third, we
propose a number of costs and benets that may affect sea turtle–epibiont interactions and discuss
the ecological inferences and implications of sea turtle epibiosis. Lastly, we outline a conceptual
model of epibiosis with which researchers may apply to better understand the factors that affect
their particular epibiotic systems and more easily decipher the important biological information that
can be gleaned from studying epibiotic interactions.
401Sea Turtle Epibiosis
15. 2 COMMON FORMS
The diversity of epibionts known from sea turtles is exceptional. For example, loggerhead (Caretta
caretta) and hawksbill turtles (Eretmochelys imbricata) are known to host 200+ and 150+ epibiont
taxa, respectively. For this reason, we have not included an itemized list of epibionts from each
turtle species. Instead, we have included a list of references that include records of epibionts from
sea turtles separated by geographic region (Appendix A) and encourage investigators to examine
the studies cited in this chapter.
15. 2.1 SESSILE FORMS
Sessile forms attach directly to a substrate and do not move around freely. These forms are the most
common and conspicuous epibionts of sea turtles. Most sessile forms have motile, planktonic larvae
that recruit to suitable substrata, where they attach and transform into adults. For these organisms,
the carapace and skin of sea turtles must possess certain settlement cues that larvae recognize,
including water ow characteristics, chemical signals, and surface rugosity. Of the sessile forms
documented from sea turtles, the most noticeable are barnacles (Cirripedia). Barnacles attached
to the carapace of sea turtles are considered “pioneer” species that facilitate the colonization of
subsequent epibiota (see Section 15.3.5; Frick et al., 2002b). Some coronuloid barnacles embed
themselves in the skin and soft tissues of sea turtles (e.g., Chelolepas cheloniae). Through chemical
mediation, these barnacles become encased in connective tissue, which aids in strengthening the
shell of the barnacle while protecting the host tissue from further injury (Frick et al., 2011a). Other
sessile forms include algae, foraminiferans, poriferans, cnidarians (Hydrozoa and Anthozoa),
mollusks (Bivalvia), bryozoans, and tunicates. Many of these sessile forms are colonial and can
reproduce asexually. As a result, some colonies are known to grow quite large and overtake much
of the carapace of the host turtle. In such situations, aggregations of sessile forms provide additional
surface area for the recruitment of other sessile epibiota, and create numerous crevices and spaces
for the colonization of various motile epibionts (see later).
15. 2.2 SEDENTARY FORMS
Sedentary forms live a semi-sessile existence, in which motile individuals construct refugia or
tubes attached to a substrate. Sea turtles host a variety of sedentary forms, including polychaete
worms, amphipods, and tanaids (Frick et al., 1998, 2004b). Some sedentary forms create only
small (1–2 mm long) tubes to dwell in, while others, particularly sabellariid worms and Corophium
amphipods, will aggregate into dense communities—creating reef-like structures consisting of
hundreds of individual tubes bonded together. These “worm reefs” can become quite large (up to
10 cm high) and cover the entire carapace of the host turtle (Frick et al., 2004b). These complex
structures also provide suitable habitat for the colonization of small motile epibionts.
15. 2.3 MOTILE FORMS
Motile forms do not directly attach to a substrate and are capable of free movement throughout
their lives. These organisms may colonize sea turtles directly from the plankton (similar to sessile
forms) or secondarily colonize turtles after initially recruiting to their primary habitat. In the latter
case, colonization may occur when resting turtles contact pelagic or benthic substrata. Motile
forms reported as sea turtle epibionts include protozoans, sipunculid worms, platyhelminth worms,
annelid worms (hirudineans and polychaetes), mollusks (Polyplacophora and Gastropoda), dipterans
(ightless marine midges), decapods (Brachyura, Anomura, Caridea), copepods, ostracods,
peracarids (amphipods, isopods, and tanaids), echinoderms (Ophiuroidea and Echinoidea), and sh
402 The Biology of Sea Turtles, Volume III
(Genera Echeneis and Remora; “shark suckers”). Most motile forms are small and cryptic, and live
within the gaps and sinuses provided by aggregations of sessile and sedentary epibionts. Moreover,
the deposition of sediment between sessile aggregations provides habitat for small infaunal animals
that live in the trapped mud layer (e.g., polychaete worms, amphipods, and clams). For these reasons,
the presence of most motile forms is often dependent on the preceding colonization of other sessile
and sedentary epibiota. Two exceptions are Caprella amphipods, which cling tightly to the host
carapace via limbs with hooked dactyls, and Planes crabs, which hide in the inguinal notch between
the carapace and tail (Chace, 1951). Not surprisingly, these are two of the more common motile
epibionts of sea turtles around the world.
15. 3 COMMUNITIES AND COMMUNITY DYNAMICS
15. 3.1 PEL AGIC/OCEANIC COMMUNITIES
All extant sea turtles, except the atback turtle (Natator depressus), utilize pelagic and oceanic
habitats during juvenile life stages (Bolten, 2003) and some continue to use these habitats throughout
adulthood (e.g., Dermochelys coriacea and eastern Pacic Lepidochelys olivacea). Adult and
subadult loggerhead turtles (C. caretta) are considered mostly neritic, but some individuals make
occasional forays into the pelagic/oceanic environment (Frick et al., 2009; Reich et al., 2010). During
pelagic/oceanic life stages, sea turtles may host communities of pelagic organisms that are typically
found associated with drifting otsam (e.g., Sargassum) and jetsam. These organisms primarily
include pedunculate barnacles of the genera Lepas and Conchoderma, and grapsid crabs of the
genus Planes. Lepas spp. and Conchoderma spp. are ubiquitous throughout the world’s oceanic
environment and are known to colonize a variety of other nektonic hosts (e.g., Reisinger and Bester,
2010; Pfaller et al., 2012). Studies on Planes crabs from oceanic-stage sea turtles represent the most
detailed information on sea turtle–epibiont symbiosis to date (Davenport, 1994; Dellinger et al.,
1997; Frick et al., 2000a, 2003b, 2004a, 2006, 2011b; Pons et al., 2011). Other less frequent epibionts
of the pelagic/oceanic community may include pelagic sea slugs (Fiona pinnata), sea spiders
(Endeis spinosa), pelagic tunicates (Diplosoma gelatinosum), and crabs of the genera Portunus
and Plagusia (Frick et al., 2003a, 2011b; Loza and López-Juardo, 2004). The presence of pelagic/
oceanic epibionts on sea turtles outside these areas strongly suggests that these turtles have recently
migrated from the pelagic/oceanic environment, providing valuable insights into cryptic migratory
behaviors and habitat preferences of sea turtles.
15. 3. 2 BENTHIC/NERITIC COMMUNITIES
After early life stages in pelagic/oceanic areas, most cheloniid sea turtles transition to more
coastal and benthic habitats—presumably in search of food, and later for mates (Bjorndal, 1997).
In benthic/neritic habitats, sea turtles become exposed to intense colonization pressure by marine
propagules (larvae and spores) seeking to colonize submerged substrata and begin their benthic
existence. The skin and especially the carapace of sea turtles provide suitable substrata for a variety
of benthic/neritic organisms (Frick et al., 1998, 2000a; Schärer, 2001). As previously mentioned,
the recruitment of sessile and sedentary forms (e.g., barnacles, tubicolous worms, and tunicates)
facilitates the colonization of smaller motile forms (e.g., crabs, amphipods, mollusks, etc.), which
inhabit the gaps and crevices between sessile aggregations. After prolonged exposure to settlement
by local plants and animals in a given area, the epibiotic communities of sea turtles begin to
resemble the adjacent benthic environment. For this reason, the species composition of benthic/
neritic communities is largely dependent on the geographic region or habitat in which the host turtle
occupies (Frick et al., 1998; Schärer, 2001). Complex benthic/neritic communities are most evident
on nesting female turtles, which tend to remain relatively sedentary and localized during the nesting
period (Frick et al., 2000b).
403Sea Turtle Epibiosis
15. 3. 3 OBLIGATE COMMUNITIES
Obligate communities are composed almost entirely of organisms that are known exclusively as
epibionts of sea turtles and other motile marine organisms. That is, these communities are largely
independent of the habitat in which the turtle occupies (i.e., pelagic/oceanic vs. benthic/neritic). The
predominant epibiont of obligate communities is the coronuloid barnacle Chelonibia testudinaria.
This ubiquitous species is the most frequently reported epibiont of sea turtles and is also known
to colonize crabs, sirenians, and crocodilians (Newman and Ross, 1976; Zardus and Hadeld,
2004; Cupul-Magaña et al., 2011; Nifong and Frick, 2011). Chelonibia testudinaria occurs in
great numbers on some turtles and appears to function as a “pioneer” for the development of more
extensive and diverse epibiotic communities (Frick et al., 2002b; Rawson et al., 2003). Aggregations
of C. testudinaria provide refugia for other obligate epibionts, such as the ruby-eyed amphipod
(Podocerus chelonophilus) and the robust tanaid (Hexapleomera robusta). However, both species
will also cling directly to the skin and carapace of host turtles, and P. chelonophilus will also
aggregate around epidermal lesions and eat necrotic tissue from the wounds of host turtles (Moore,
1995). Other obligate epibionts of sea turtles include marine red alga (Polysiphonia carettia), which
is known only from cheloniid sea turtles (Senties et al., 1999), and several other species of coronuloid
barnacles that are wholly chelonophilic (Ross and Frick, 2011). While some individual turtles are
known to host strictly obligate communities (Frick et al., 2010a), most communities composed
primarily of obligate epibionts also contain some facultative forms.
15. 3. 4 COMMUNITY DISTRIBUTION
The spatial distribution of epibiont communities on host turtles may be inuenced by a complex
suite of factors, including recruitment dynamics, water ow patterns, differential disturbance among
body regions, and inter- and intraspecic interactions (Pfaller et al., 2006). In general, studies that
examine or anecdotally report on the distribution of sea turtle epibionts have found that epibiont
communities tend to aggregate on the carapace, as opposed to the skin or plastron (Gramentz,
1988; Fuller et al., 2010). Extra-carapacial epibionts mostly include barnacles, parasitic leeches,
and Planes crabs (Chace, 1951; Gramentz, 1988; Frick et al., 1998; Hayashi and Tsuji, 2008). Some
barnacles occur only along the plastral sutures (e.g., Stomatolepas transversa) (Young, 1991), while
others mostly occur along the leading edges of the front ippers (e.g., Stephanolepas muricata)
(Frick et al., 2011a). Limb movements, unfavorable water ow patterns, and the sloughing of skin
by the host turtle probably restrict the recruitment and development of extra-carapacial epibionts.
Nevertheless, information on the distributions of extra-carapacial epibionts is still lacking (Frick
et al., 2011a).
Most studies that examine the spatial distribution of epibiont communities on sea turtles have
focused on the carapace, where the densest and most diverse communities are found (Frick et al.,
1998). These studies indicate that epibiotic communities tend to be distributed in nonrandom patterns.
Most studies report a tendency for epibiont communities to cluster along the vertebral scutes and
across the posterior third of the carapace (Caine, 1986; Matsuura and Nakamura, 1993; Frick et al.,
1998; Pfaller et al., 2006). Such nonrandom distributions are thought to reect the preference of
lter-feeding epibionts (e.g., barnacles) for elevated ow rates along the vertebral scutes and the
favorable settlement conditions for other epibiota along the posterior of the carapace where ow rates
are reduced (Pfaller et al., 2006). Recruitment of “pioneer” species in these areas (e.g., Chelonibia
barnacles and Polysiphonia alga) will then facilitate the accumulation of more diverse epibiotic
communities (Gramentz, 1988; Frick et al., 2000b; Fuller et al., 2010). Additionally, the colonization
and persistence of epibionts on the anterior costal scutes may be reduced by contact from the front
ippers (Caine, 1986; Dodd, 1988) and/or removal during “self-grooming” (Schoeld et al., 2006;
Frick and McFall, 2007). Other studies show mostly random distributions among barnacle species
with some spatial structuring among different size classes of barnacles (Fuller et al., 2010).
404 The Biology of Sea Turtles, Volume III
Recently, Moriarty et al. (2008) conrmed that the obligate commensal barnacle, Chelonibia
testudinaria, is capable of substantial (but slow) post-settlement locomotion. Individual C.
testudinaria were shown to move across multiple scutes from areas of low water ow to areas with
better lter-feeding conditions. Such movements may be triggered by differential ow rates over the
carapace or/and the presence of conspecics that disrupt ow patterns. As previously mentioned,
Chelonibia spp. are important “pioneer” species for epibiotic communities (Frick et al., 2000) and
post-settlement locomotion will certainly affect the spatial distribution of epibiotic communities.
However, as the density of C. testudinaria and other epibiota increases, post-settlement locomotion
and survival will be reduced, and the overall distribution may become more reective of differences
in recruitment patterns (Pfaller et al., 2006).
Debilitated turtles will host epibionts, especially barnacles, over their entire external surface
area—including portions of the mouth regularly exposed to the outside environment. These
“barnacle bill” turtles will often suffer severe deformations as a result of barnacle colonization.
Current information indicates that such turtles are immunosuppressed or lethargic prior to barnacle
colonization and that limited mobility by the host likely facilitates rapid and prolic colonization
of barnacles (Deem et al., 2009). Nevertheless, because healthy turtles may also support massive
aggregations of epibionts over much of their bodies, it is difcult to judge the health of a turtle
simply by examining epibiont loads and percentage coverage (see Deem et al., 2009).
15. 3. 5 COMMUNITY SUCCESSION
Prior to the colonization of macroorganisms, all structures exposed to seawater initially undergo
a similar sequence of events (Wahl, 1989): (1) biochemical conditioning, whereby surfaces absorb
dissolved macromolecules; (2) bacterial colonization; and (3) unicellular eukaryote (e.g., yeasts,
protozoa, and diatoms) colonization. To our knowledge, these critical stages in the process of
epibiosis in sea turtles have never been explored.
The temporal succession of “macro”-epibiont communities on host turtles remains poorly
understood, as well. To date, there is one study that examines temporal succession of epibiont
communities from individual turtles over an extended period of time (Frick et al., 2002b). Using,
ipper-tagging data, photography, and in situ assessments, epibiont data were collected from the
carapaces of nesting loggerhead turtles (C. caretta) in Georgia, United States, over the course
of 3 months. General observations of community succession were similar to those reported for
neritic, epibenthic communities (Dean, 1981). Community succession is typically initiated when
hard, sessile forms like barnacles (C. testudinaria in Frick et al., 2002b) colonize a relatively
bare carapace. These “pioneers” facilitate the subsequent colonization of other epibiota by
increasing the surface area for colonization and changing water ow patterns (Pfaller et al.,
2006). Secondary colonizers include other sessile forms (e.g., hydrozoans and bryozoans) and
sedentary forms, which take refuge within the interstices of the barnacles (e.g., tanaids). The
accumulation of sediments among primary and secondary sessile forms then facilitates the
colonization of sessile tunicates and many small, motile forms. Tunicates and other secondary
sessile forms tend to overgrow and kill the barnacles beneath them. Tunicates (Molgula
manhattensis) appear to be the climax species of the carapace epibiont community on nesting
loggerheads in Georgia, United States. Aggregations of M. manhattensis occasionally cover the
entire carapace at the end of the season, providing innumerable gaps and crevices for a diverse
array of motile epibionts.
At or before reaching terminal succession, epibiont communities may be partially or
catastrophically disturbed by various biotic and abiotic factors. Turtles that accumulate benthic/
neritic communities may immigrate to different, less favorable habitats, causing the less tolerant
epibionts to die and slough off. In some cases, this may completely clear the carapace of epibiota.
Moreover, community succession may be disrupted when host turtles “groom” themselves
by actively rubbing against submerged structures to remove epibiota (Heithaus et al., 2002;
405Sea Turtle Epibiosis
Schoeld et al., 2006; Frick and McFall, 2007). Evidence of such behaviors is often present in the
form of longitudinal scratch marks on the carapace (Caine, 1986; Frick and McFall, 2007). Lastly,
predatory epibionts (e.g., Planes crabs and several gastropods) and sh may systematically clean/
remove certain epibionts (Davenport, 1994; Losey et al., 1994; Frick et al., 2000a, 2011b; Pfaller
et al., 2008; Sazima et al., 2010). These factors may lead to partial or complete turnover of the
epibiotic communities of sea turtles.
15. 4 ECOLOGICAL INTERACTIONS
15. 4.1 EFFECTS ON EPIBIONTS
Epibionts may benet from epibiosis through reduced competition and predation. These are major
factors affecting the ability of marine propagules to successfully colonize a substratum (Enderlein
and Wahl, 2004). Thus, when risk of predation is high or when settlement area is limited—whether
by high population densities (e.g., on benthic structures) or by low substrata availability (e.g., on
pelagic otsam)—epibiosis of sea turtles may be benecial for the survival of marine propagules
(Wahl, 1989; Pfaller et al., 2012). Some “burrowing” barnacles may avoid predation by encasing
themselves within the tissue of host turtles via chemical mediation (Frick et al., 2011b). Epibionts
may also benet from improved energetic positioning. Filter-feeding epibionts, such as barnacles,
may benet from favorable feeding currents on host turtles (Pfaller et al., 2006), while photosynthetic
epibionts, such as algae, may benet from increased oxygen and light availability (Shine et al.,
2010). Furthermore, epibionts may benet though range expansion and increased genetic mixing by
hitchhiking on migratory turtles (termed phoresis). Researchers have hypothesized that sea turtles
may act as long-distance dispersal vectors for benthic marine invertebrates (Schärer and Epler,
2007; Harding et al., 2011).
Epibiosis may be costly to epibionts when turtle behaviors cause physical disturbance and
unfavorable uctuations in physiological conditions (Wahl, 1989). Contact between turtles during
mating, or between turtles and submerged structures (e.g., rock or coral ledges), may physically
damage epibionts, especially those with fragile, erect body forms (e.g., leafy bryozoans and soft
corals). As previously mentioned, sea turtles are also known to actively remove epibionts by scraping
against submerged structures (Heithaus et al., 2002; Schoeld et al., 2006; Frick and McFall, 2007).
Moreover, epibionts that are sensitive to desiccation may die when turtles emerge to nest or bask at
the surface (Caine, 1986; Bjorndal, 2003). Similarly, epibionts that are sensitive to uctuations in
temperature, salinity, or pressure may not survive when turtles migrate and/or dive. Another cost
for certain epibionts might be reduced access to food resources and mates, which would ultimately
cause reduced longevity and reproductive capacity. These costs might favor epibionts capable of
asexual reproduction and dietary versatility.
15. 4. 2 EFFECTS ON HOST TURTLES
Epibiosis may be costly to host turtles when epibionts cause increased weight and drag. In
extreme cases, epibiotic loads have been reported that effectively double the mass and volume of
juvenile sea turtles (Bolten unpubl. data in Bjorndal, 2003). Epibionts attached to the carapace
may increase drag by disrupting the laminar ow over the carapace (Logan and Morreale, 1994)
and those embedded in the leading edge of the front ippers may increase drag while swimming
(Wyneken, 1997; Frick et al., 2011a). The energetic costs of hosting epibionts are likely greatest
when turtles undertake long-distance migrations and least when turtles remain relatively sedentary
(e.g., females during internesting periods). Because otherwise healthy turtles will often support
massive epibiont aggregations (Deem et al., 2009), turtles are apparently capable of overcoming
the costs associated with “epibiotic drag” and should not be judged as healthy or unhealthy
simply by examining epibiotic loads (see Deem et al., 2009). Furthermore, the aforementioned
406 The Biology of Sea Turtles, Volume III
“barnacle bill” turtles tend to accumulate their prolic barnacle loads after (not before) becoming
lethargic at the surface.
Epibiosis may also be costly to host turtles when certain epibionts detrimentally affect the health
of host turtles. A number of common epibionts of sea turtles (e.g., platyhelminth worms, annelid
worms and barnacles) are thought to be the cause of or related to infections of sea turtles (George,
1997; Alfaro, 2008). Tissue damage caused by burrowing epibionts may increase the vulnerability
of host turtles to pathogens (George, 1997). Some coronuloid barnacles (e.g., C. cheloniae, S.
muricata, and Cylindrolepas darwiniana) become embedded within hard and soft tissues of host
turtles causing deep-tissue wounds that can sometimes leave impressions on the underlying bone
(Hendrickson, 1958; Green, 1998; Frick and Zardus, 2010; Frick et al., 2010a). Platylepas decorata
have also been found imbedded in the beaks of host turtles causing severe beak deformation, which
may lead to reduced foraging capacity and death of the host turtle (see Green, 1998; Frick and
Zardus, 2010). Other non-barnacle forms may act as disease vectors of pathogens. Parasitic marine
turtle leeches (Ozobranchus sp.) not only consume host tissue but also are believed to act as disease
vectors for the dispersal of the bropapilloma-associated herpes virus found in latent tumors that
often cover, deform, and debilitate host turtles (Greenblatt et al., 2004). Commensal gastropods of
sea turtles may act as intermediate hosts for spirorchiid blood ukes (Frazier et al., 1985), which can
have devastating effects on host turtles (George, 1997).
Host turtles may benet from epibiosis through improved optical, chemical, or electrical
camouage. Predators may not recognize hosts as potential prey items if epibiotic communities
visually or chemically resemble the surrounding benthic communities (Rathbun, 1925; Fishlyn and
Phillips, 1980; Feifarek, 1987; Frazier et al., 1991). Moreover, dense epibiotic communities may
disrupt electric elds produced by hosts, allowing hosts to avoid predation by predators that utilize
electrolocation when searching for prey (e.g., sharks) (Ruxton, 2009). Hosts may also benet from
epibiosis through associational defense and cleaning. Epibionts with chemical or structural defenses
(e.g., toxins, sharp projections, or hard outer coverings) may deter predation on host turtles (Wahl
and Mark, 1999; Bjorndal, 2003). Predatory epibionts may provide a cleaning benet by consuming
other epibionts—some of which may be harmful—from the surface of host turtles (Davenport,
1994; Sazima et al., 2010).
15. 4. 3 ECOLOGICAL INFERENCES
Studies of epibiosis have helped elucidate cryptic life history attributes of sea turtles and informed
the implementation of conservation measures. While such studies will not and should not supplant
the use of tag-return data, satellite telemetry, stable-isotope analyses, or population genetics,
studying epibiosis can provide a time- and cost-effective alternative to elucidate the geographic
ranges, habitat preferences, and migratory corridors of sea turtles. Using primarily examples from
the well-studied epibiont community of loggerhead turtles in the northwestern Atlantic Ocean,
we illustrate the types of ecological inferences that can be gained by studying the epibionts of
Epibiont data have been used to elucidate the foraging locations of loggerhead turtles nesting
along the Atlantic coast of Florida, United States. These turtles occasionally host epibionts that are
geographically restricted to far southern Florida, the Bahamas, and the Caribbean (Caine, 1986;
Pfaller et al., 2008). Such associations suggest that these nesting turtles had recently migrated from
more southerly areas where their range overlapped with free-living populations of the epibionts.
Data from ipper-tag returns, satellite telemetry, and stable-isotope analyses have conrmed
that turtles nesting in Florida frequently utilize these more southerly, tropical waters during
nonbreeding seasons (Meylan, 1983; Foley et al., 2008; Pajuelo et al., 2012). Caine (1986) further
extrapolated these epibiont data to suggest the presence of two discrete nesting assemblages along
the southeastern United States, one to the north and one to the south of Daytona Beach, Florida
(approximately 29° N latitude). Several years later this hypothesis was rather precisely conrmed by
407Sea Turtle Epibiosis
molecular data (Bowen et al., 1993; Encalada et al., 1998) and now these two nesting assemblages
receive markedly different conservation status (Turtle Expert Working Group, 2009).
In another example from nesting loggerhead turtles in Florida, United States, Reich et al. (2010)
supplemented stable-isotope data with epibiont community data to suggest a bimodal foraging
strategy by female loggerheads prior to their arrival at breeding grounds. Because isotopic signatures
(depleted vs. enriched δ13C) can vary along multiple environmental continua, the incorporation of
epibiont data in this study provided additional support for an oceanic versus neritic dichotomy,
as opposed to dietary or latitudinal gradients. These results have important implications for role
of adult loggerhead turtles in the oceanic environment and the management policies that serve to
Epibiont data have also been used to assess the foraging migrations of juvenile and subadult
loggerhead turtles. Killingley and Lutcavage (1983) used duel isotopic proles (δ18O and δ13C) from
the shells of C. testudinaria to reconstruct the movements of subadult loggerheads between oceanic
habitats in the northwest Atlantic and estuarine habitats in the Chesapeake Bay (Maryland and
Virginia). Moreover, Limpus and Limpus (2003) used the presence of particular epibionts (Planes
sp. and S. muricata) and morphological features to identify which juvenile turtles caught in neritic
habitats in the southwest Pacic Ocean had recently recruited from the open ocean. In both studies,
epibiont data provided valuable insights in to cryptic host movements that otherwise would have
been very difcult to obtain.
Lastly, in another interesting application of epibiont data, Eckert and Eckert (1988) measured
the size distribution of epibiotic barnacles (Conchoderma virgatum) on nesting leatherback turtles
to extrapolate the time of arrival to the tropical nesting region. Because reproduction in these
barnacles is typically restricted to tropical regions, their colonization of turtles is limited to the
period when turtles also occupy tropical waters. Based on reproductive periodicity and established
growth rates of barnacles (Eckert and Eckert, 1987), the authors determined that turtles do not
arrive from temperate latitudes until just prior to nesting and orient directly toward their preferred
nesting beach (Eckert and Eckert, 1988). These data have provided important information on the
cryptic migratory behavior of leatherback turtles and have better informed the implementation of
15. 4. 4 ECO LOGICAL IMPLICATIONS
The ecological implications of sea turtle epibiosis remain one of the most poorly understood aspects
of this nascent eld. Aside from many of the direct effects of epibiosis on host turtles and epibionts
discussed earlier (Sections 15.4.1 and 15.4.2), sea turtle epibiosis may have other less obvious,
indirect effects on the marine communities and habitats that sea turtles inhabit.
Several authors have discussed the potential role of sea turtles as dispersal vectors for a
diverse array of marine invertebrates over broad geographic regions (Bjorndal and Jackson,
2003; Schärer and Epler, 2007; Harding et al., 2011; Lezama et al., 2012). Hitchhiking on highly
mobile hosts may facilitate genetic mixing and/or range expansion for epibionts capable of
reproducing on turtles or after arriving in distant locations (Rawson et al., 2003). These factors
may be particularly important for invertebrate taxa with limited dispersal capacities (Schärer
and Epler, 2007). Turtle-mediated genetic mixing may aid in maintaining the genetic diversity
and homogeneity of marine invertebrate populations (Rawson et al., 2003), but may also inhibit
biological diversication by impeding local adaptation or random divergence. Moreover, turtle-
mediated range expansion may promote biological diversication if newly established populations
subsequently remain isolated from their source populations, or disrupt ecosystem functioning
when invaders compete with or consume resident species.
A recent study has drawn attention to the potential for turtle-mediated introductions of
nonindigenous and potentially invasive species. Harding et al. (2011) report the rst records of the
nonindigenous veined rapa whelk (Rapana venosa) as an epibiont of loggerhead turtles in Virginia
408 The Biology of Sea Turtles, Volume III
and Georgia. R. venosa is a generalist shellsh predator native to Asia that has recently been
introduced in to the Chesapeake Bay (Harding and Mann, 1999). However, the size and stage of the
epibiotic individuals on turtles in Georgia indicate the presence of an extra-Chesapeake breeding
population of this invasive species. The authors suggest that turtle-mediated dispersal is currently
the only compelling explanation for the occurrence of R. venosa on turtles in Georgia. These
ndings have important implications for the future management of invasive marine invertebrates.
Sea turtles are known to modify the physical structure of their habitat in a number of ways
(Bjorndal and Jackson, 2003). Thus, another unexplored ecological implication of sea turtle
epibiosis might be the extent to which turtles modify hard-bottom habitats when actively
removing epibiota. This behavior involves turtles pushing their carapace against the underside of
rock ledges and vigorously scrapping against the rock to remove epibiota, particularly barnacles
(Frick and McFall, 2007). The rock ledges often erode during such behaviors, leaving behind
scours or arched ledges, which turtles may return to for subsequent “self-grooming.” The extent
to which these habitat modications affect the surrounding reef or hard-bottom communities
15. 5 CONCEPTUAL MODEL OF EPIBIOSIS
As we accumulate studies of epibiotic diversity in sea turtles, we have begun to formulate a
conceptual framework to better understand and learn from these epibiotic interactions. While
there have been several broad reviews on epibiosis (see Wahl, 1989; Wahl and Mark, 1999; Harder,
2009; Wahl, 2009), there has been no attempt to construct a conceptual framework to explain such
associations. The conceptual model of epibiosis depicted in Figure 15.1 outlines three hierarchical
factors inherent to epibiotic interactions: (1) geographic overlap (Figure 15.1A), (2) ecological
overlap (Figure 15.1B), and (3) the balance of costs and benets to hosts and epibionts that dictate
the likelihood of epibiosis once in close proximity (Figure 15.1C). Because the factors that affect
epibiotic interactions—as displayed in this conceptual model—are inherent to the biology of the
species involved, we can learn about the ecology and evolution of these species by studying epibiosis.
Such a conceptual framework will hopefully allow researchers to better understand the factors
that affect their particular epibiotic systems and more easily decipher the important biological
information that can be gleaned from studying epibiosis in sea turtles.
15. 5.1 GEOGRAPHIC OVERLAP
A necessary prerequisite for epibiosis is geographic overlap between the range of the host turtle and
the range of the epibiont (Figure 15.1A). Logically, without geographic overlap, epibiosis between a
host turtle and any potential epibiont would never occur. This is an obvious criterion for epibiosis.
However, because the host turtles are highly mobile, the occurrence of particular epibiont taxa
with more limited distributions can reveal information about cryptic host movements. Studies of
sea turtle epibionts have provided important information on the migratory behavior of loggerhead
and leatherback turtles (Caine, 1986; Eckert and Eckert, 1988), and subsequently informed the
implementation of conservation measures.
15. 5. 2 ECOLOGICAL OVERLAP
Where geographic ranges overlap, epibiosis will then depend on the spatial and temporal overlap
in ecology of the host turtles and potential epibionts (Figure 15.1B). Local geographic areas are
typically heterogeneous mosaics of different habitats, each characterized by different ecological
communities of plants and animals (e.g., saltmarshes, coral reefs, pelagic areas). The species
composition of local communities may also vary through time, especially for seasonal differences
409Sea Turtle Epibiosis
in recruitment of larval propagules. Host turtles may utilize many different habitats or may show
preferences for certain habitats during different behaviors (e.g., foraging, resting, and mating) or
life stages, or at different times of the year. In order for epibiosis to occur, the host turtles must
occupy the same habitat at the same time as free-living populations of potential epibionts. Thus, the
epibionts associated with a given host turtle should reect the assemblage of plants and animals that
occupy the habitats where the hosts spend time. For example, sea turtles that tend to inhabit benthic/
neritic habitats tend to host different epibionts than turtles that tend to inhabit pelagic/oceanic
habitats (Limpus and Limpus, 2003; Reich et al., 2010). Such information can be used to assess
interspecic and intraspecic differences in habitat use, which is critical for the implementation of
effective conservation strategies.
15. 5. 3 BALANCE OF COSTS AND BENEFITS
Once in close proximity, there is a complex balance of costs and benets for host turtles and
potential epibionts that ultimately determine the likelihood of epibiosis. Figure 15.1 displays a
2D likelihood surface in which each axis represents a continuum from high benet to high cost.
The various positions of different hosts and epibionts along these cost-benet axes depend on
the net cost or benet experienced during epibiosis. Because the relative costs and benets are
Likelihood of epibiosis
FIGURE 15.1 Conceptual model of epibiosis. (A,B) Venn diagrams showing the geographic and ecological
overlap between hosts and epibionts, respectively. (C) Graph showing the likelihood of epibiosis based on the
balance of cost and benets to hosts and epibionts (Bepibiont, benet to the epibiont; Bhost, benet to the host;
Chost, cost to the host; Cepibiont, cost to the epibiont).
410 The Biology of Sea Turtles, Volume III
different for different turtle–epibiont pairs, some associations are more likely and therefore more
frequent than others. Epibiotic interactions in which both species experience a net benet would
have a high likelihood of occurring and therefore would be more frequent (quadrant I). Such
mutually benecial associations would favor mechanisms for active attraction and may develop
into obligate associations over evolutionary time. On the other end of the continua, interactions
in which both species suffer high costs would have a low likelihood and would effectively never
occur (quadrant III). Interactions in which one species incurs a high cost while the other receives
minimal benet would also have a low likelihood (bottom left of quadrants II and IV), as the
former species would actively avoid such interactions and the latter would gain very little by
exploiting the former. Conversely, if one species receives a high benet at a high cost to the
other species (top left of quadrant II and bottom right of quadrant IV), then such associations
might exhibit patterns similar to that of parasitic interactions (top left of quadrant II only). Lastly,
interactions in which one species receives a high benet while the other incurs little or no cost
would have a higher likelihood and would be relatively frequent (top left of quadrants II and IV).
This last scenario characterizes many of the interactions between sea turtles and their epibiota,
and is typically referred to as commensalism (Leung and Poulin, 2008).
As previously mentioned, epibionts may benet from epibiosis through reduced spatial
competition and predation (Wahl, 1989; Enderlein and Wahl, 2004; Pfaller et al., 2012), improved
energetic positioning (Pfaller et al., 2006; Shine et al., 2010), and range expansion (Schärer and
Epler, 2007; Harding et al., 2011), while coping with costs associated with physical disturbance
(Wahl, 1989; Schoeld et al., 2006; Frick and McFall, 2007), transport to unfavorable physiological
environments (Caine, 1986; Bjorndal, 2003), and reduced access to food resources and mates.
Host turtles may benet from epibionts through optical, chemical, or electrical camouage
(Rathbun, 1925; Fishlyn and Phillips, 1980; Feifarek, 1987; Frazier et al., 1991; Ruxton, 2009)
and associational defense and cleaning (Davenport, 1994; Wahl and Mark, 1999; Bjorndal,
2003), while coping with costs associated with increased weight and drag (Logan and Morreale,
1994; Bjorndal, 2003), and tissue da mage and associated susceptibility to pathogens (George,
1997; Greenblatt et al., 2004). The balance of costs and benets to host turtles and epibionts will
ultimately determine the likelihood—and therefore the frequency—of epibiosis for most turtle–
15. 6 CONSIDERATIONS
Studies that seek to elucidate the relationships that exist between sea turtles and other marine
organisms require investigators to adopt an interdisciplinary approach to data collections and
analyses. Knowledge of the standard measurements and preservation methods employed by taxon
specialists is important to properly report and archive marine algae and invertebrate specimens
(Lazo-Wasem et al., 2011). A familiarity with the life histories and general biology of the marine
organisms that utilize the habitats occupied by sea turtles is essential for identifying situations that
bring sea turtles into contact with the marine organisms they consume and those that attach to
them. An understanding of the major systematic characters that dene the major family-groups of
local marine ora and fauna is helpful for identication, and to adequately ascertain evolutionary
relationships between sea turtles and other marine organisms.
We sincerely thank Rebecca Pfaller for valuable editorial and technical assistance. We thank
Kristina L. Williams of the Caretta Research Project, Karen Bjorndal, Alan Bolten, Peter Eliazar,
and our other colleagues at the Archie Carr Center for Sea Turtle Research for their encouragement,
advice, and support, and we greatly appreciate the helpful comments of anonymous reviewers that
improved an earlier draft of the present chapter.
411Sea Turtle Epibiosis
15. A APPENDIX A: ANNOTATED BIBLIOGRAPHY OF SELECTED SEA TURTLE
EPIBIONT STUDIES AND REPORTS LISTED BY GEOGRAPHIC REGION
15. A.1 CARIBBEAN–WESTERN ATLANTIC
Bacon, 1976 (Trinidad); Bugoni et al., 2001 (Rio Grande do Sul: Brazil); Cardenas-Palomo and
Maldonado-Gasca, 2005 (Yucatan: Mexico); Caine, 1986 (South Carolina, Florida); Farrapeira-
Assunção, 1991 (Brazil); Frazier et al., 1985 (Georgia, Florida); Frazier et al., 1991 (Georgia);
Frazier et al., 1992 (Georgia; Rio Grande do Sul: Brazil); Frick et al., 1998 (Georgia); Frick and Slay,
2000 (Georgia); Frick and Zardus, 2010 (Panama, Georgia, and Florida); Frick et al., 2000a, 2000b
(Georgia); Frick et al., 2002a,b (Georgia); Frick et al., 2003a (Jumby Bay: Antigua); Frick et al.,
2004b (Georgia); Frick et al., 2006 (Florida); Frick et al., 2010a (Nova Scotia, Georgia); Frick
et al., 2010b (Georgia and Florida); Gruvel, 1905 (Antilles Sea); Henry, 1954 (Florida, Texas);
Hunt, 1995 (Florida); Ives, 1891 (Yucatan: Mexico); Killingley and Lutcavage, 1983 (Virginia);
Lutcavage and Musick, 1985 (Virginia); Nilsson-Cantell, 1921 (Florida); Nilsson-Cantell, 1939
(Bay of Chacopata: Venezuela); Pereira et al, 2006 (Almofala: Brazil); Pilsbry, 1916 (Cape Frio:
Brazil; Delaware, Florida, New Jersey; Point Patuca: Honduras; West Indies); Plotkin, 1996
(Texas); Richards, 1930 (New Jersey); Rudloe et al., 1991 (Florida); Schwartz, 1960 (Maryland);
Walker, 1978 (North Carolina); Wass, 1963 (Virginia); Wells, 1966 (Florida); Weltner, 1897
(Florida; Cuba; Bahia: Brazil); Young, 1991 (Brazil); Zavodnik, 1997 (Rovinj: Croatia); Zullo and
Bleakney, 1966 (Massachusetts; Nova Scotia: Canada); Zullo and Lang, 1978 (South Carolina).
15. A.2 MEDITERRANEAN– EASTERN ATLANTIC
Badillo-Amador, 2007 (Mediterranean Sea); Barnard, 1924 (Table Bay: South Africa); Broch, 1924
(Baie du Levrier: Mauretania; Gambia); Broch, 1927 (Rabat: Morocco); Carriol and Vader, 2002
(Finmark: Norway); Caziot, 1921 (Nice: France); Chevereaux and de Guerne, 1893 (between Algeria
and Balaeres); Darwin, 1854 (Africa; Mediterranean Sea); Davenport, 1994 (Madeira); Frazier et al.,
1985 (Peloponnesus, Zakynthos Island: Greece); Frick et al., 2010a (Gabon: Africa); Gauld, 1957
(Accra: Ghana); Geldiay et al., 1995 (Koycegiz-Dalyankoy: Turkey); Gramentz, 1988 (Malta, Zacharo,
Zakynthos: Greece; Lampedusa: Italy); Gruvel, 1903 (Palermo: Italy; Alexandria: Egypt); Gruvel, 1931
(Gulf of Alexandrette); Haelters and Kerckhof, 1999 (DeHaan: Belgium); Haelters and Kerckhof, 2001
(Oostende: Belgium); Holothuis, 1952 (Ouddorp: the Netherlands); Holothuis, 1969 (Ameland Island: the
Netherlands); Kitsos et al., 2005 (Aegean Sea); Kolosvary, 1939 (Rovigno, d’Istria: Croatia); Kolosvary,
1943 (Alexandria: Egypt; Palermo, Sicily: Italy); Kolosvary, 1951 (Mediterranean Sea); Koukouras
and Matsa, 1998 (Aegean Sea; Levantine Basin); Lanfranco, 1979 (St. Julian’s: Malta); Lucas, 1968
(Mediterranean Sea); Margaritoulis, 1985 (Zakynthos: Greece); Nilsson-Cantell, 1921 (Bibundi:
Cameroon); Nilsson-Cantell, 1931 (Mediterranean Sea); O’Riordan, 1979 (Dingle: Ireland); O’Riordan
and Holmes, 1978 (Ventry Harbor: Ireland); Pilsbry, 1916 (Taranto: Italy; Cape of Good Hope: South
Africa); Quigley and Flannery, 1993 (Dingle Bay: Ireland); Relini, 1968 (Gulf of Trieste: Italy); Relini,
1969 (Adriatic Se a); Relini, 1980 (Adriat ic Sea); Sezgin et al., 2009 (Turkey); Smaldon and Lyster, 1976
(Skarvoy: Norway; Crail, Kirkcudbrightshire: Scotland; Cornwall: England); Stubbings, 1965 (Hann,
Saloum River: Senegal); Stubbings, 1967 (Goree, Hann: Senegal); Utinomi, 1959 (Banyuls-sur-Mer:
France); Zakhama-Sraieb et al., 2010 (Gulf of Gabès: Mediterranean Sea).
15. A.3 INDO–WEST PACIFIC
Annandale, 1906 (Rameswaram Island: India; Gulf of Manaar); Balazs, 1978 (Hawaii); Balazs,
1980 (Hawaii); Balazs et al., 1987 (Hawaii); Borradaile, 1903 (Minikoi Island: India); Broch,
1916 (Broome: Australia); Broch, 1931 (Gulf of Thailand; Nagasaki: Japan); Broch, 1947
(Ream: Cambodia; Indochina); Bustard, 1976 (Great Barrier Reef: Australia); Daniel, 1956
(Tuticorin, Drusadai Islands, Royapuram Coast, Madras Coast: India); Daniel, 1962 (Little Andaman
412 The Biology of Sea Turtles, Volume III
Island: India); Darwin, 1854 (Low Archipelago: French Polynesia; Australia); Dawydoff, 1952 (Pulo
Condore: Vietnam; Ream: Cambodia); Deraniyagala, 1939 (Bentota: Ceylon); Dobbs and Landry,
2004 (Great Barrier Reef: Australia); Fernando, 1978 (Porto Novo: India); Glazebrook and Campbell,
1990 (Torres Strait: Australia); Fischer, 1886 (Pulo Condor: Vietnam); Foster, 1978 (North Island: New
Zealand); Frazier, 1971 (Aldabra Atoll); Frazier et al., 1985 (Orissa: India; Tanzania: Africa); Frazier,
1989 (Dwarka Island: India); Frazier et al., 1992 (Orissa, Gujarat: India; Karachi, Pakistan);Gordon,
1970 (Hawaii); Gruvel, 1903 (Seychelles; Mallicolo: Vanuatu; Djibouti; Sandwich Island; Cochinchina:
Vietnam); Gruvel, 1907 (Andaman Islands: India); Gruvel, 1912 (Tuamotu Archipelago: French
Polynesia); Hayashi and Tsuji, 2008 (Okinawa: Japan); Hendrickson, 1958 (Johor, Sarawak: Malaysia);
Hiro, 1936 (Wakayama Prefecture, Aichi Prefecture: Japan); Hiro, 1937a (Baberudaobu Island: Palau);
Hiro, 1939 (Toyama Bay: Japan); Jones, 1990 (Australia); Jones et al., 1990 (Tasmania; Australia);
Jones et al., 2000 (summary of distribution); Kruger, 1911b (Sagami Bay: Japan); Kruger, 1912 (Timor
Sea); Lanchester, 1902 (Kota Bharu: Malaysia); Limpus et al., 1983a (Campbell Island: Australia);
Limpus et al., 1983b (Crab Island: Australia); Limpus et al., 2005 (Raine Island: Australia); Loop
et al., 1995 (Milman Island: Australia); Losey et al., 1994 (Hawaii); Matsuura and Nakamura, 1993
(Kagoshima Prefecture: Japan); McCann, 1969 (North Island: New Zealand); Monroe and Limpus,
1979 (Queensland: Australia); Mustaquim and Javed, 1993 (Sandspit Beach: Pakistan); Newman et al.,
1969 (Hawaii); Newman and Abbott, 1980 (California); Nilsson-Cantell, 1921 (Western Australia:
Australia); Nilsson-Cantell, 1930a (Enoe Island: Malaysia); Nilsson-Cantell, 1932 (Bentota: Sri Lanka);
Nilsson-Cantell, 1937 (Singapore); Nilsson-Cantell, 1938 (Maldives; Kilakarai, Andaman Islands,
River Hooghly, mouth of Ganges: India); Pillai, 1958 (Quilon: India); Pilsbry, 1916 (Hawaii; Caroline
Islands; Ana: Japan; Saigon: Vietnam); Pilsbry, 1927 (Hawaii); Ren, 1980 (Xisha Islands); Ren, 1987
(China); Ross, 1981 (Oman); Smaldon and Lyster, 1976 (Kuala Lumpur: Malaysia); Tachikawa, 1995
(Japan); Utinomi, 1949 (Hakata Bay: Japan); Utinomi, 1958 (Sagami Bay: Japan); Utinomi, 1966
(Amakusa: Japan); Utinomi, 1969 (Kharg: Iran); Utinomi, 1950 (Tanabe Bay: Japan); Utinomi, 1970
(Hakui, Cape Kyoga-misaki, Kamo, Nezugaseki, Sado Island: Japan); Wagh and Bal, 1974 (Bombay:
India); Weltner, 1897 (Massaua: New Guinea; Torres Strait); Weltner, 1910 (Ile Europa); Zann and
Harker, 1978 (Queensland: Australia); Zardus and Balazs, 2007 (Hawaii).
15. A.4 EASTERN PACIFIC
Angulo-Lozano et al., 2007 (Sinaloa: Mexico); Beaumont et al., 2007 (Galapagos Islands: Ecuador);
Brown and Brown, 1995 (Peru); Darwin, 1854 (Mexico; Galapagos Islands: Ecuador); Frazier et al.,
1985 (Galapagos Islands: Ecuador); Frazier et al., 1992 (Santa Rosa: Ecuador); Frick et al., 2011a,b
(Baja California: Mexico; Eastern Tropical Pacic; Galapagos: Ecuador); Green, 1998 (Galapagos
Islands: Ecuador); Henry, 1941 (La Paz: Mexico); Henry, 1960 (Gulf of California, Guaymas: Mexico);
Hernandez-Vasquez and Valadez-Gonzalez, 1998 (Jalisco: Mexico); Hubbs, 1977 (California);
Kolosvary, 1943 (San Jose: Guatemala); Lazo-Wasem et al., 2011 (Jalisco: Mexico); MacDonald,
1929 (Cocos Island: Costa Rica); Newman et al., 1969 (Baja California: Mexico; Eastern Pacic);
Pilsbry, 1916 (Baja California: Mexico; Galapagos Islands: Ecuador); Ross and Newman, 1967 (Baja
California: Mexico); Stinson, 1984 (California); Vivaldo et al., 2006 (Michoacan, Oaxaca: Mexico);
Weltner, 1897 (western Mexico; California; Valparaiso: Chile); Young and Ross, 2000 (Sonora:
Mexico); Zullo, 1986 (Galapagos Islands: Ecuador); Zullo, 1991 (Galapagos Islands: Ecuador).
Alfaro, A. 2008. Synopsis of infections in sea turtles caused by virus, bacteria and parasites: An ecological
review. In A.F. Rees, M.G. Frick, A. Panagoloulou and K. Williams, compilers. Proceedings of the 27th
Annual Symposium on Sea Turtle Biology and Conservation. NOAA Technical Memorandum NMFS-
SEFC-569, Miami, FL, p. 5.
Annandale, N. 1906. Report on the Cirripedia collected by Professor Herdman, at Ceylon, in 1902. Ceylon
Pearl Oyster Fisheries, Supplemental Report 31: 137–150.
413Sea Turtle Epibiosis
Bacon, P. 1976. Cirripedia of Trinidad. Studies on the Fauna of Curacao and Other Caribbean Islands
Badillo-Amador, F.J. 2007. Epizoítos y parásitos de la tortuga boba (Caretta caretta) en el Mediterráneo
Occidental. PhD thesis, Facultat de Ciencies Biologiques, Univesitat de Valencia, Valencia, Spain,
Balazs, G.H. 1978. A hawksbill turtle in Kaneohe Bay, Oahu. Elepaio 38: 128–129.
Balazs, G.H. 1980. Synopsis of the biological data on the green turtle in the Hawaiian Islands. NOAA Technical
Memorandum NMFS-7, pp. 1–141.
Balazs, G.H., R.G. Forsyth, and A.K.H. Kam. 1987. Preliminary assessment of habitat utilization by Hawaiian
green turtles in their resident foraging pastures. NOAA Technical Memorandum NMFSC-SWFC-71,
Barnard, K.H. 1924. Contributions to the crustacean fauna of South Africa. Annotates of the South African
Museum 20: 1–103.
Beaumont, E.S., P. Zárate, J.D. Zardus, P.H. Dutton, and J.H. Seminoff. 2007. Epibiont occurrence in
Galapagos green turtles (Chelonia mydas) at nesting and feeding grounds. In A.F. Rees, M.G. Frick,
A.Panagoloulou, and K. Williams, compilers. Proceedings of the 27th Annual Symposium on Sea Turtle
Biology and Conservation. NOAA Technical Memorandum NMFS-SEFC-569, Miami, FL, p. 8.
Bjorndal, K.A. 1997. Foraging ecology and nutrition of sea turtles. In P.L. Lutz and J.A. Musick, eds. The
Biology of Sea Turtles. CRC Press, Boca Raton, FL, pp. 199–231.
Bjorndal, K.A. 2003. Roles of loggerheads in marine ecosystems. In A.B. Bolten and B.E. Witherington, eds.
Biology and Conservation of the Loggerhead Sea Turtle. Smithsonian Institution Press, Washington, DC,
Bjorndal, K.A. and J.B.C. Jackson. 2003. Roles of sea turtles in marine ecosystems: Reconstructing the past.
In P.L. Lutz, J.A. Musick, and J. Wyneken, eds. The Biology of the Sea Turtles, Vol. II. CRC Press, Boca
Raton, FL, pp. 259–274.
Bolten, A.B. 2003. Variation in sea turtle life history patterns: Neritic vs. oceanic developmental stages. In P.L.
Lutz, J.A. Musick, and J. Wyneken, eds. The Biology of the Sea Turtles, Vol. II. CRC Press, Boca Raton,
FL, pp. 243–257.
Borradaile, L.A. 1903. VII. The Barnacles (Cirripedia). In S. Gardner, ed. The Fauna and Geography of the
Maldive and Laccadive Archipelagoes, Being the Account of the Work Carried on and of the Collections
Made by and Expedition during the Years 1899 and 1900, Vol. I. University Press, Cambridge, MA,
Bowen, B.W., J.C. Avise, J.I. Richardson, A.B. Meylan, D. Margaritoulis, and S.R. Hopkins-Murphy. 1993.
Population structure of loggerhead turtles (Caretta caretta) in the northwestern Atlantic Ocean and
Mediterranean Sea. Conservation Biology 7: 834–844.
Broch, H. 1916. Cirripedien. Results of Dr. E. Moberg’s Swedish scientic expeditions to Australia 1910–13.
Kungliga Svenska Vetenskaps Akademiens Handlingar 52: 1–16.
Broch, H. 1924. Cirripedia. Parasitologia Mauritanica, Materiaux pour la Faune Parasitologique en Mauritanie.
Arthropoda (2e Partie). Bulletin Comite d’Etudes Hist. Sci. l’Afrique Occidentale Francaise (October–
December, 1924): 1–21.
Broch, H. 1927. Studies on Moroccan cirripeds (Atlantic Coast). Bulletin de la Societe des Sciences Naturelles,
Maroc 6–7: 11–38.
Broch, H. 1931. Indomalayan Cirripedia. Papers from Dr. Th. Mortensen’s Pacic Expedition 1914–1916.
Videnskabelige Meddelelser Dansk Naturhistorisk Forening 91: 1–146.
Broch, H. 1947. Cirripedes from Indochinese shallow-waters. Avhandlinger Norske Videnskaps-akademi
Oslo I 7: 1–32.
Brown, C.A. and W.M. Brown. 1995. Status of sea turtles in the southeastern Pacic: Emphasis on Peru. In
K.A. Bjorndal, ed. Biology and Conservation of Sea Turtles, Revised edition. Smithsonian Institution
Press, Washington, DC, pp. 235–240.
Bugoni, L., L. Krause, A.O. de Almeida, and A.A. De Pádua Bueno. 2001. Commensal barnacles of sea turtles
in Brazil. Marine Turtle Newsletter 94: 7–9.
Bustard, H.R. 1976. Turtles of coral reefs and coral islands. In O.A. Jones and R. Endean, eds. Biology and
Geology of Coral Reefs, Vol. III (Biology 2). Academy Press, NY, pp. 343–368.
Caine, E.A. 1986. Carapace epibionts of nesting loggerhead turtles: Atlantic coast of U.S.A. Journal of
Experimental Marine Biology and Ecology 95: 15–26.
Cardenas-Palomo, N. and A. Maldonado-Gasca. 2005. Epibiontes de tortugas de carey juveniles Eretmochelys
imbricata en el Santuario de Tortugas Marinas de Rio Lagartos, Yucatan, Mexico. CICIMAR Oceanides
414 The Biology of Sea Turtles, Volume III
Carriol, R.P. and W. Vader. 2002. Occurrence of Stomatolepas elegans (Cirripedia: Balanomorpha) on a
leatherback turtle from Finnmark, northern Norway. Journal of the Marine Biological Association of the
United Kingdom 82: 1033–1034.
Caziot, E. 1921. Les Cirripedes de la mer de Nice. Bulletin Society Zoology, France 46: 51–54.
Chace, F.A. 1951. The oceanic crabs of the genera Planes and Pachygrapsus. Proceedings of the United States
National Museum 101: 65–103.
Chevereaux, E. and J. de Guerne. 1893. Crustaces et Cirripeds commensaux des Tortues marines de la
Mediterranee. Comptes Rendus des Seances de l’Academie des Sciences 116: 443–445.
Cupul-Magaña, F.G., A. Rubio-Delgado, A.H. Escobedo-Galván, and C. Reyes-Nuñez. 2011. First report
of marine barnacles Lepas anatifera and Chelonibia testudinaria as epibionts on American crocodile
(Crocodylus acutus). Herpetology Notes 4: 213–214.
Daniel, A. 1956. The Cirripedia of the Madras Coast. Bulletin of the Madras Government Museum 6: 1–40.
Daniel, A. 1962. A new species of platylepadid barnacle (Cirripedia: Crustacea) from the green turtle from
Little Andaman Island. Annals and Magazine of Natural History (Series 13) 5: 641–645.
Darwin, C. 1851. A Monograph on the Subclass Cirripedia, with Figures of All Species. The Lepadidae, or,
Pedunculated Cirripedes. Ray Society, London, U.K. 400pp.
Darwin, C. 1854. A Monograph on the Subclass Cirripedia, with Figures of All the Species. The Balanidae, the
Verrucidae, etc. Ray Society, London, U.K. 684pp.
Davenport, J. 1994. A cleaning association between the oceanic crab Planes minutus and the loggerhead sea
turtle Caretta caretta. Journal of the Marine Biological Society of the United Kingdom 74: 735–737.
Dawydoff, C. 1952. Contribution a l’etude des invertebrates de la faune marine benthique de l’Indochine.
Bulletin biologique de la France et de la Belgique 37(Suppl): 127–131.
Dean, T.A. 1981. Structural aspects of sessile invertebrates as organizing forces in an Estuarine fouling
community. Journal of Experimental Marine Biology and Ecology 53: 163–180.
Deem, S.L., T.M. Norton, M. Mitchell, A. Segars, A.R. Alleman, C. Cray, R.H. Poppenga, M. Dodd, and
W.B.Karesh. 2009. Comparison of blood values in foraging, nesting, and stranded loggerhead turtles
(Caretta caretta) along the coast of Georgia, USA. Journal of Wildlife Diseases 45: 41–56.
Dellinger, T., J. Davenport, and P. Witz. 1997. Comparisons of social structure of Columbus crabs living on
loggerhead sea turtles and inanimate otsam. Journal of the Marine Biological Association of the United
Kingdom 77: 185–194.
Deraniyagala, P.E.P. 1939. The tetrapod reptiles of Ceylon. Ceylon Journal of Science, Colombo Museum,
Natural History Series 1: 1–412.
Dobbs, K.A. and A.M. Landry, Jr. 2004. Commensals on nesting hawksbill turtles (Eretmochelys imbricata),
Milman Island, northern Great Barrier Reef, Australia. Memoirs of the Queensland Museum 49: 674.
Dodd, C.K. Jr. 1988. Synopsis of the biological data on the loggerhead sea turtle Caretta caretta (Linnaeus
1758). U.S. Fish and Wildlife Service Biological Report 88: 1–110.
Eckert, K.L. and S.A. Eckert. 1988. Pre-reproductive movements of leatherback sea turtles (Dermochelys
coriacea) nesting in the Caribbean. Copeia 1988: 400–406.
Encalada, S.E., K.A. Bjorndal, A.B. Bolten, J.C. Zurita, B. Schoeder, E. Possardt, C.J. Sears, and B.W. Bown. 1998.
Population structure of loggerhead turtle (Caretta caretta) nesting colonies in the Atlantic and Mediterranean
as inferred from mitochondrial DNA control region sequences. Marine Biology 130: 567–575.
Enderlein, P. and M. Wahl. 2004. Dominance of blue mussels versus consumer-mediated enhancement of
benthic diversity. Journal of Sea Research 51: 145–55.
Farrapeira-Assunção, C.M. 1991. Revisão do gênero Chelonibia Leach, 1817 na costa Brasileira (Crustacea,
Cirripedia). Abstracts of XVIII Congresso Brasileiro de Zoologia, Salvador, Brazil, 133pp.
Feifarek, B.P. 1987. Spines and epibionts as antipredator defenses in the thorny oyster Spondylus americanus
Hermann. Journal of Experimental Marine Biology and Ecology 105: 39–56.
Fernando, S.A. 1978. Studies on the biology of barnacles (Crustacea: Cirripedia) of Porto Novo region, South
India. PhD dissertation, Center of Advanced Study in Marine Biology, Annamalai University, Tamil
Nadu, India, 213pp.
Fischer, P. 1886. Description d’un nouveau genre de Cirripedes (Stephanolepas) parisite des tortues marines.
Actes de la Société Linnéenne de Bordeaux 40: 193–196.
Fishlyn, D.B. and D.W. Phillips. 1980. Chemical camouaging and behavioural defenses against predatory seastar
by three species of gastropods from the surfgrass Phyllospadix community. Biological Bulletin 158: 34–48.
Foley, A.M., B.A. Schroeder, and S.L. MacPherson. 2008. Post-nesting migrations and resident areas of Florida
loggerheads. In H. Kalb, A. Rohde, K. Gayheart, and K. Shanker, compilers. Proceedings of the 25th
Annual Symposium on Sea Turtle Biology and Conservation. NOAA Technical Memorandum NMFS-
SEFSC-582, Miami, FL, pp. 75–76.
415Sea Turtle Epibiosis
Foster, B.A. 1978. The marine fauna of New Zealand: Barnacles (Cirripedia: Thoracica). New Zealand
Oceanographic Institute Memorandum 69: 1–160.
Frazier, J. 1971. Observations on sea turtles at Aldabra Atoll. Philosophical Transactions of the Royal Society
of London, Series B 260: 373–410.
Frazier, J. 1989. Observations on stranded green turtles, Chelonia mydas, in the Gulf of Kutch. Journal of the
Bombay Natural History Society 86: 250–252.
Frazier, J.G., I. Goodbody, and C.A. Ruckdeschel. 1991. Epizoan communities on marine turtles. II. Tunicates.
Bulletin of Marine Science 48: 763–765.
Frazier, J.G., D. Margaritoulis, K. Muldoon, C.W. Potter, J. Rosewater, C. Ruckdeschel, and S. Salas. 1985.
Epizoan communities on marine turtles. I. Bivalve and gastropod mollusks. Marine Ecology 6: 127–140.
Frazier, J.G., J.E. Winston, and C.A. Ruckdeschel. 1992. Epizoan communities on marine turtles. III. Bryozoa.
Bulletin of Marine Science 51: 1–8.
Frick, M.G., K. Kopitsky, A.B. Bolten, K.A. Bjorndal, and H.R. Martins. 2011a. Sympatry in grapsoid crabs
(genera Planes and Plagusia) from olive ridley sea turtles (Lepidochelys olivacea), with descriptions of
crab diets and masticatory structures. Marine Biology 158: 1699–1708.
Frick, M.G., P.A. Mason, K.L. Williams, K. Andrews, and H. Gerstung. 2003a Epibionts of hawksbill turtles
in a Caribbean nesting ground: A potentially unique association with snapping shrimp (Crustacea:
Alpheidae). Marine Turtle Newsletter 99: 8–11.
Frick, M.G. and G. McFall. 2007. Self-grooming by loggerhead turtles in Georgia, USA. Marine Turtle
Newsletter 118: 15.
Frick, M.G., A. Ross, K.L. Williams, A.B. Bolten, K.A. Bjorndal, and H.R. Martins. 2003b. Epibiotic associates
of oceanic-stage loggerhead turtles from the southeastern North Atlantic. Marine Turtle Newsletter
Frick, M.G. and C.K. Slay. 2000. Caretta caretta (loggerhead sea turtle) epizoans. Herpetological Review
Frick, M.G., K.L. Williams, A.B. Bolten, K.A. Bjorndal, and H.R. Martins. 2004a. Diet and fecundity of
Columbus crabs, Planes minutus, associated with oceanic-stage loggerhead sea turtles, Caretta caretta,
and inanimate otsam. Journal of Crustacean Biology 24: 350–355.
Frick, M.G., K.L. Williams, A.B. Bolten, K.A. Bjorndal, and H.R. Martins. 2009. Foraging ecology of oceanic-
stage loggerhead turtles Caretta caretta. Endangered Species Research 9: 91–97.
Frick, M.G., K.L. Williams, M. Bresette, D.A. Singewald, and R.M. Herren. 2006. On the occurrence of Columbus
crabs (Planes minutus) from loggerhead turtles in Florida, USA. Marine Turtle Newsletter 114: 12–14.
Frick, M.G., K.L. Williams, E.J. Markestyn, J.B. Pfaller, and R.E. Frick. 2004b. New records and observations
of epibionts from loggerhead sea turtles. Southeastern Naturalist 3: 613–620.
Frick, M.G., K.L. Williams, and M. Robinson. 1998. Epibionts associated with nesting loggerhead sea turtles
(Caretta caretta) in Georgia. Herpetological Review 29: 211–214.
Frick, M.G., K.L. Williams, and D. Veljacic. 2000a. Additional evidence supporting a cleaning association
between epibiotic crabs and sea turtles: How will the harvest of Sargassum weed impact this relationship?
Marine Turtle Newsletter 90: 11–13.
Frick, M.G., K.L. Williams, and D.C. Veljacic. 2002a. New records of epibionts from loggerhead sea turtles
Caretta caretta (L.) Bulletin of Marine Science 70: 953–956.
Frick, M.G., K.L. Williams, D. Veljacic, J.A. Jackson, and S.E. Knight. 2002b. Epibiont community succession
on nesting loggerhead sea turtles, Caretta caretta, from Georgia, USA. In A. Mosier, A. Foley and
B. Brost, compilers. Proceedings of the 20th Annual Symposium on Sea Turtle Biology and Conservation.
NOAA Technical Memorandum, NMFS-SEFSC-447, Miami, FL, pp. 280–282.
Frick, M.G., K.L. Williams, D. Veljacic, L. Pierrard, J.A. Jackson, and S.E. Knight. 2000b. Newly documented
epibiont species from nesting loggerhead sea turtles (Caretta caretta) in Georgia. Marine Turtle
Newsletter 88: 103–108.
Frick, M.G. and J.D. Zardus. 2010. First authentic report of the turtle barnacle Cylindrolepas darwiniana since
its description in 1916. Journal of Crustacean Biology 30: 292–295.
Frick, M.G., J.D. Zardus, and E.A. Lazo-Wasem. 2010a. A new Stomatolepas barnacle species (Cirripedia:
Balanomorpha: Coronuloidea) from leatherback sea turtles. Bulletin of the Peabody Museum of Natural
History 51: 123–136.
Frick, M.G., J.D. Zardus, and E.A. Lazo-Wasem. 2010b. A new coronuloid barnacle subfamily, genus and
species from cheloniid sea turtles. Bulletin of the Peabody Museum of Natural History 51: 169–177.
Frick, M.G., J.D. Zardus, A. Ross, J. Senko, D. Montano-Valdez, M. Bucio-Pacheco, and I. Sosa-Cornejo. 2011b.
Novel records of the barnacle Stephanolepas muricata (Cirripe dia: Bala nomo rpha : Coronulo idea ); in clud -
ing a case for chemical mediation in turtle and whale barnacles. Journal of Natural History 45: 629–640.
416 The Biology of Sea Turtles, Volume III
Fuller, W.J., A.C. Broderick, R. Enever, P. Thorne, and B.J. Godley. 2010. Motile homes: A comparison of the
spatial distribution of epibiont communities on Mediterranean sea turtles. Journal of Natural History
Gauld, D.T. 1957. An annotated check-list of the Crustacea of the Gold Coast. I. Cirripedia. Journal of the West
African Science Association 3: 10–11.
Geldiay, R., T. Koray, and S. Balik. 1995. Status of sea turtle populations (Caretta caretta caretta and Chelonia
mydas) in the northern Mediterranean Sea, Turkey. In K.A. Bjorndal, ed. Biology and Conservation of
Sea Turtles, Revised edition. Smithsonian Institution Press, Washington, DC, pp. 425–437.
George, R.H. 1997. Health problems and disease of sea turtles. In P.L. Lutz and J.A. Musick, eds. The Biology
of the Sea Turtles. CRC Press, Boca Raton, FL, pp. 363–385.
Glazebrook, R.S. and R.S.F. Campbell. 1990. A survey of the diseases of marine turtles in northern Australia.
2. Oceanarium-reared and wild turtles. Diseases of Aquatic Organisms 9: 97–104.
Gordon, J.A. 1970. An annotated checklist of Hawaiian barnacles (Class Crustacea: Subclass Cirripedia)
with notes on their nomenclature, habitats and Hawaiian localities. Hawaii Institute of Marine Biology
Technical Report 19: 1–130.
Gramentz, D. 1988. Prevalent epibiont sites on Caretta caretta in the Mediterranean Sea. Naturalista Siciliano
Green, D. 1998. Epizoites of Galapagos green turtles. In R. Byles and Y. Fernandez, compilers. Proceedings of
the 16th Annual Symposium on Sea Turtle Biology and Conservation. NOAA-Technical Memorandum
NMFS-SEFSC–412, Miami, FL, p. 63.
Greenblatt, R.J., T.M. Work, G.H. Balazs, C.A. Sutton, R.N. Casey, and J.W. Casey. 2004. The Ozobranchus
leech is a candidate mechanical vector for the bropapilloma-associated turtle herpesvirus found latently
infecting skin tumors on Hawaiian green turtles (Chelonia mydas). Virology 1: 101–110.
Gruvel, A. 1903. Revision des Cirrhipedes appartenant a la collection du Museum d’Histoire Naturelle.
Nouvelles Archives du Museum D’Histoire Naturelle de Paris (Series 4), 5: 95–170.
Gruvel, A. 1905. Monographie des Cirrhipedes ou Theocostraces. Masson et Cie, Editeurs, Paris, 472pp.
Gruvel, A. 1907. Cirrhipedes opercules de le l’Indian Museum de Calcutta. Memories of the Asiatic Society of
Bengal 2: 1–10.
Gruvel, A. 1912. Mission Gruvel sur la cote occidentale d’Afriques (1909–1910) et collection du Museum
d’Histoire Naturelle, Les Cirrhipedes. Bulletin du Museum D’Histoire Naturelle, Series 1: 344–350.
Gruvel, A. 1931. Crustaces de Syrie. Les etat de Syrie, Paris, France, pp. 397–435.
Haelters, J. and F. Kerckhof. 1999. Een waarneming van de lederschildpad Dermochelys coriacea (Linnaeus,
1758), en de eerste waarneming van Stomatolepas dermochelys Monroe and Limpus, 1979 aan de
Belgische kust. De Strandvlo 19: 30–39.
Haelters, J. and F. Kerckhof. 2001. Opnieuw een klapmuts Cystophora cristata Erxleben, 1777 en een
lederschildpad Dermochelys coriacea (Linnaeus 1758) aan onze kust. De Strandvlo 21: 81–83.
Harder T. 2009. Marine epibiosis: Concepts, ecological consequences and host defence. Marine and Industrial
Biofouling 4: 219–31.
Harding, J.M. and R. Mann. 1999. Observations on the biology of the veined rapa whelk, Rapana venosa
(Valenciennes, 1846) in the Chesapeake Bay, USA. Journal of Shellsh Research 24: 9–18.
Harding, J.M., W.J. Walton, C.M. Trapani, M.G. Frick, and R. Mann. 2011. Sea turtles as potential dispersal
vectors for non-indigenous species: The veined rapa whelk as an epibiont of loggerhead sea turtles.
Southeastern Naturalist 10: 233–244.
Hayashi, R. and K. Tsuji. 2008. Spatial distribution of turtle barnacles on the green sea turtle, Chelonia mydas.
Ecological Research 2007 (On-line Journal of the Ecological Society of Japan), 5pp.
Heithaus, M.R., J.J. McLash, A. Frid, L.M. Dill, and G.J. Marshall. 2002. Novel insights into green turtle
behaviour using animal-borne video cameras. Journal of the Marine Biological Association of the
United Kingdom 82: 1049–1050.
Hendrickson, J.R. 1958. The green sea turtle, Chelonia mydas (Linn.), in Malaya and Sarawak. Proceedings of
the Zoological Society of London 130: 455–535.
Henry, D.P. 1941. Notes on some sessile barnacles from Lower California and the west coast of Mexico.
Proceedings of the New England Zoological Club 18: 99–107.
Henry, D.P. 1954. The barnacles of the Gulf of Mexico. Gulf of Mexico, its origin, waters and marine life. U.S.
Fish and Wildlife Service, 55, Fishery Bulletin 89: 443–446.
Henry, D.P. 1960. Thoracic Cirripedia of the Gulf of California. University of Washington Publication
Oceanography 4: 135–158.
Hernandez-Vasquez, S. and C. Valadez-Gonzalez. 1998. Observations on the epizoa found on the turtle
Lepidochelys olivacea at La Gloria, Jalisco, Mexico. Ciencias Marina 24: 119–125.
417Sea Turtle Epibiosis
Hiro, F. 1936. Occurrence of the cirriped Stomatolepas elegans on a loggerhead turtle found at Seto.
Annotationes Zoologicae Japonenses 15: 312–320.
Hiro, F. 1937a. Cirripeds of the Palao Islands. Palao Tropical Biological Station Studies 1: 37–72.
Hiro, F. 1939. Studies on the cirripedien fauna of Japan. V. Cirripeds of the northern part of Honshyu. Science
Reports of the Tohoku Imperial University (Series 4) 14: 201–218.
Holothuis, L.B. 1952. Enige interessante, met drijvende voorwerpen op de Nedelandse kust aangespoelde
zeepissebedden en zeepokken. De Levende Natuur 55: 72–77.
Holothuis, L.B. 1969. Enkele interessante Nederlandse Crustacea. Zoologische Bijdragen 2: 34–48.
Hubbs, C.L. 1977. First record of mating ridley turtles in California, with notes on commensals, characters, and
systematics. California Fish and Game 63: 263–267.
Hunt, T.L. 1995. Preliminary survey of commensals associated with Caretta caretta. In J.I. Richardson and
T.H. Richardson, compilers. Proceedings of the 12th Annual Workshop on Sea Turtle Biology and
Conservation. NOAA-Technical Memorandum NMFS-SEFSC-361, Miami, FL, pp. 204–207.
Ives, J.E. 1891. Crustacea from the northern coast of Yucatan, the harbor of Vera Cruz, the west coast of
Florida and the Bermuda Islands. Proceedings of the Academy of Natural Sciences of Philadelphia 43:
Jones, D.S. 1990. The shallow-water barnacles (Cirripedia: Lepadomorpha, Balanomorpha) of southern
Western Australia. In F.E. Wells, D.I. Walker, H. Kirkman and R. Lethbridge, compilers. Proceedings of
the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western
Australia. Western Australia Museum, Perth, Australia, pp. 333–437.
Jones, D.S., J.T. Anderson and D.T. Anderson. 1990. Checklist of the Australian Cirripedia. Technical Report
of the Australian Museum 3: 1–38.
Jones, D.S., M.A. Hewitt, and A. Sampey. 2000. A checklist of the cirripedia of the South China Sea. Rafes
Bulletin of Zoology 8(Suppl): 233–307.
Killingley, J.S. and M. Lutcavage. 1983. Loggerhead turtle movements reconstructed from 18O and 13C pro-
les from commensal barnacle shells. Estuarine Coastal Shelf Science 16: 345–349.
Kitsos, M., M. Christodoulou, C. Arvanitidis, M. Mavidis, I. Kirmitzoglou, and A. Koukouras. 2005.
Composition of the organismic assemblage associated with Caretta caretta. Journal of the Marine
Biological Association of the United Kingdom 2005: 257–261.
Kolosvary, G. 1939. Ueber Fundortsangaben adriatischer Balanen. Bollettino dei Musei di Zoologia ed
Anatomia Comparata della R. Università di Torino (Series III) 47: 37–41.
Kolosvary, G. 1943. Cirripedi thoracica in der Sammlung des ungarischen National-Museums. Annales
Historico-Naturales Musei Nationalis Hungarici 36: 67–120.
Kolosvary, G. 1951. Les balanids de la Mediterranee. Acta Biologica 2: 411–413.
Koukouras, A. and A. Matsa. 1998. The thoracican cirriped fauna of the Aegean Sea: New information, checklist
of the Mediterranean species, faunal comparisons. Senckenbergiana Maritima 28: 133–142.
Kruger, P. 1911b. Zur Cirripedien fauna Ostasiens. Zoologischer Anzeiger, Leipzig 38: 459–464.
Kruger, P. 1912. Uber ostrasiatische Rhizocephalen. Anhang: Uber eninige intersante Vertreter der Cirripedia
thoracica. Abhandlungen der Mathematisch-Physikalische Klasse der Königlich Bayerischen Akademie
der Wissenschaften, II Supplement Band 8: 1–16.
Lanchester, W.F. 1902. On the Crustacea collected during the “Skeat Expedition” to the Malay Peninsula.
Proceedings of the Zoological Society of London 2: 363–381.
Lazo-Wasem, E.A., T. Pinou, A. Peña de Niz, and A. Feuerstein. 2011. Epibionts associated with the nesting
marine turtles Lepidochelys olivacea and Chelonia mydas in Jalisco, Mexico: A review and eld guide.
Bulletin of the Peabody Museum of Natural History 52(2): 221–240.
Lanfranco, G. 1979. Stomatolepas elegans Costa (Crustacea, Cirripedia) on Dermochelys coriacea Linn., taken
in Maltese waters. Central Mediterranean Naturalist 1: 24.
Leung T.L.F. and Poulin R. 2008. Parasitism, commensalism, and mutualism: Exploring the many shades of
symbioses. Vie Milieu 58: 107−115.
Lezama, C., A. Carranza, A. Fallabrino, A. Estrades, and M. López-Mendilaharsu. 2012. Unintended back-
packers: Bio-fouling of the invasive gastropod Rapana venosa on the green turtle Chelonia mydas in the
Río de la Plata Estuary, Uruguay. Biological Invasions doi:10.10007/s10530-012-0307-9.
Limpus, C.J. and D.J. Limpus. 2003. Biology of the loggerhead turtle in western south Pacic Ocean foraging
areas, In A.B. Bolten and B.E. Witherington, eds. Biology and Conservation of the Loggerhead Sea
Turtle. Smithsonian Institution Press, Washington, DC, pp. 93–113.
Limpus, C.J., D.J. Limpus, M. Munchow, and P. Barnes. 2005. Queensland turtle conservation project: Raine
Island turtle study, 2004–2005. Queensland Government Conservation Technical and Data Report,
418 The Biology of Sea Turtles, Volume III
Limpus, C.J., J.D. Miller, V. Baker, and E. Mclachlan. 1983a. The hawksbill turtle, Eretmochelys imbri-
cata (L.), in north-eastern Australia: The Campbell Island rookery. Australian Wildlife Research 10:
Limpus, C.J., C.J. Parmenter, V. Baker, and A. Fleay. 1983b. The Crab Island sea turtle rookery in the north-
eastern Gulf of Carpentaria. Australian Wildlife Research 10: 173–184.
Linnaeus, C. 1758. Systema Naturae. Holmiae, Editio Decima, Reformata, Vol. 1, 824pp.
Logan, P and S.J. Morreale. 1994. Hydrodynamic drag characteristics of juvenile L. kempi, C. mydas, and C.
caretta. In B.A. Schroeder and B.E. Witherington, compilers. Proceedings of the 13th Annual Symposium
on Sea Turtle Biology and Conservation. NOAA-Technical Memorandum NMFS-SEFSC-341, Miami,
FL, pp. 248–252.
Loop, K.A., J.D. Miller, and C.J. Limpus. 1995. Nesting by the hawksbill turtle (Eretmochelys imbricata) on
Milman Island, Great Barrier Reef, Australia. Wildlife Research 22: 241–252.
Losey, G., G.H. Balazs, and L.A. Privitera. 1994. Cleaning symbiosis between the wrasse, Thalassoma dup-
erry, and the green turtle, Chelonia mydas. Copeia 1994: 684–690.
Loza, A.L. and L.F. Lopez-Juardo. 2004. Comparative study of the epibionts on the pelagic and mature
female loggerhead turtles on the Canary and Cape Verde Islands. In R.B. Mast, B.J. Hutchinson, and
A.H. Hutchinson, compliers. Proceedings of the 24th Annual Symposium on Sea Turtle Biology and
Conservation. NOAA Technical Memorandum NMFS-SEFSC-567, Miami, FL, p. 100.
Lucas, M. 1968. Les cirrhipedes l’Europe. Les Naturalistes Belges 49: 105–160.
MacDonald, R. 1929. A report of some cirripeds collected by the S.S. “Albatross” in the eastern Pacic during
1891 and 1904. Bulletin of the Museum of Comparative Zoology 69: 527–538.
Margaritoulis, D. 1985. Preliminary observations on the breeding behaviour and ecology of Caretta caretta
in Zakynthos, Greece. 2e Congrès international sur la zoogéographie et l’écologie de la Grèce et des
régions adjacentes, Athens, Greece, September 1981, 10, pp. 323–332.
Matsuura, I. and K. Nakamura. 1993. Attachment pattern of the turtle barnacle Chelonibia testudinaria on the
carapace of nesting loggerhead turtles Caretta caretta. Bulletin of the Japanese Society for the Science of
Fish (Nippon Suisan Gakkaishi) 59: 1803.
McCann, C. 1969. First southern hemisphere record of the platylepadine barnacle Stomatolepas elegans (Costa)
and notes on the host Dermochelys coriacea (Linne). New Zealand Journal of Marine and Freshwater
Research 3: 152–158.
Meylan, A.B. 1983. Marine turtles of the Leeward Islands, Lesser Antilles. Smithsonian Institution Atoll
Research Bulletin 278: 1–43.
Moore, P.G. 1995. Podocerus chelonophilus (Amphipoda: Podoceridae) associated with epidermal lesions of
the loggerhead turtle, Caretta caretta (Chelonia). Journal of the Marine Biological Association of the
United Kingdom 75: 253–255.
Monroe, R. and C.J. Limpus. 1979. Barnacles on turtles in Queensland waters with descriptions of three new
species. Memoirs of the Queensland Museum 19: 197–223.
Moriarty, J.E., J.A. Sachs, and K. Jones. 2008. Directional locomotion in a turtle barnacle, Chelonibia
testudinaria, on green turtles, Chelonia mydas. Marine Turtle Newsletter 119: 1–4.
Mustaquim, J. and M. Javed. 1993. Occurrence of Chelonibia testudinaria (Linnaeus) (Crustacea: Cirripedia)
in coastal waters of Pakistan. Pakistan Journal of Marine Science 2: 73–75.
Newman, W.A. and D.P. Abbott. 1980. Cirripedia: The barnacles. In R.H Morris, D.P. Abbott and E.C. Haderlie,
eds. Intertidal Invertebrates of California. Stanford University Press, Stanford, CA, pp. 504–535.
Newman, W.A., V.A. Zullo, and T.H. Withers. 1969. Cirripedia. Treatise on Invertebrate Paleontology, Part R,
Arthropoda 4: R206–R295.
Nifong, J.C. and M.G. Frick. 2011. First record of the American alligator (Alligator mississippiensis) as a host
to the sea turtle barnacle (Chelonibia testudinaria). Southeastern Naturalist 10: 557–560.
Nilsson-Cantell, C.A. 1921. Cirripedian-Studien. Zur kenntnis der Biologie, Anatomie und Systematik dieser
Gruppe. Zoologiska Bidrag från Uppsala 7: 75–395.
Nilsson-Cantell, C.A. 1930a. Diagnoses of some new cirripeds from the Netherlands Indies collected by the
expedition of His Royal Highness the Prince Leopold of Belgium in 1929. Bulletin de Musee Royal
d’Histoire Naturelle de Belgique 6: 1–2.
Nilsson-Cantell, C.A. 1931. Revision der Sammulung recenter Cirripedien des Naturhistorichen Museums in
Bael. Verhandlungen der Naturforschenden Gesellschaft in Basel 42: 103–137.
Nilsson-Cantell, C.A. 1932. The barnacles Stephanolepas and Chelonibia from the turtle Eretmochelys
imbricata. Ceylon Journal of Science, Section B (Spolia Zeylanica) 16: 257–264.
419Sea Turtle Epibiosis
Nilsson-Cantell, C.A. 1937. On a second collection of Indo-Malayan cirripeds from the Rafes Museum.
Bulletin of the Rafes Museum 13: 93–96.
Nilsson-Cantell, C.A. 1938. Cirripeds from the Indian Ocean in the collection of the Indian Museum, Calcutta.
Memoirs of the Indian Museum 13: 1–81.
Nilsson-Cantell, C.A. 1939. Recent and fossil balanids from the north coast of South America. Capita Zoologica
O’Riordan, C.E. 1979. Marine fauna notes from the National Museum of Ireland 6. Irish Naturalists Journal
O’Riordan, C.E. and J.M.C. Holmes. 1978. Marine fauna notes from the National Museum of Ireland.
5. Passengers on the North Atlantic currents. Irish Naturalists Journal 19: 152–153.
Pajuelo, M., K.A. Bjorndal, K.J. Reich, M.D. Arendt, and A.B. Bolten. 2012. Distribution of foraging habitats
of male loggerhead turtles (Caretta caretta) as revealed by stable isotopes and satellite telemetry. Marine
Biology 159: 1255–1267.
Pereira, S., E.H.S.M. Lima, L. Ernesto, H. Matthews, and A. Ventura. 2006. Epibionts associated with Chelonia
mydas from Northern Brazil. Marine Turtle Newsletter 111: 17–18.
Pfaller, J.B., K.A. Bjorndal, K.J. Reich, K.L. Williams, and M.G. Frick. 2006. Distribution patterns of epibionts
on the carapace of loggerhead turtles, Caretta caretta. Journal of the Marine Biological Association of
the United Kingdom Marine Biodiversity Records 1: e36.
Pfaller, J.B., M.G. Frick, F. Brischoux, C.M. Sheey III, and H.B. Lillywhite. 2012. Marine snake epibiosis:
A review and rst report of decapods associated with Pelamis platurus. Integrative and Comparative
Biology 52: 296–310.
Pfaller, J.B., M.G. Frick, K.J. Reich, K.L. Williams, and K.A. Bjorndal. 2008. Carapace epibionts of loggerhead
turtles (Caretta caretta) nesting at Canaveral National Seashore, Florida. Journal of Natural History.
Pillai, N.K. 1958. Development of Balanus amphitrite, with a note on the early larvae of Chelonibia testudinaria.
Bulletin of the Central Research Institute of Kerala, University of Kerala, Series C 6: 117–130.
Pilsbry, H.A. 1916. The sessile barnacles (Cirripedia) contained in the collections of the U.S. National Museum;
including a monograph of the American species. U.S. National Museum Bulletin 93: 1–366.
Pilsbry, H.A. 1927. Littoral barnacles of the Hawaiian Islands and Japan. Proceedings of the Academy of
Natural Sciences of Philadelphia 79: 305–317.
Plotkin, P.T. 1996. Occurrence and diet of juvenile loggerhead sea turtles, Caretta caretta, in the northwestern
Gulf of Mexico. Chelonian Conservation and Biology 2: 78–80.
Pons, M., A. Verdi, and A. Domingo. 2011. The pelagic crab Planes cyaneus (Dana, 1851) (Decapoda,
Brachyura, Grapsidae) in the southwestern Atlantic Ocean in association with loggerhead sea turtles
buoys. Crustaceana 84: 425–434.
Quigley, D.T. and K. Flannery. 1993. Southern marine fauna and ora from S.W. Ireland. Porcupine Newsletter
Rainbow, P.S. and G. Walker. 1977. The functional morphology of the alimentary tract of barnacles (Cirripedia:
Thoracica). Journal of Experimental Marine Biology and Ecology. 28: 183–206.
Ranzani, C. 1817–1818. Osservazioni su i Balanidi. Bologna, opuscoli Scientici I (1817): 195–202; II (1817):
269–276; III (1818): 63–93.
Rathbun, M.J. 1925. The spider crabs of America. Bulletin— United States National Museum 129: 1–598.
Rawson, P.D., R. Macnamee, M.G. Frick, and K.L. Williams. 2003. Phylogeography of the coronulid barnacle,
Chelonibia testudinaria, from loggerhead sea turtles, Caretta caretta. Molecular Ecology 12: 2697–2706.
Reich, K.J., K.A. Bjorndal, M.G. Frick, B.E. Witherington, C. Johnson, and A.B. Bolten. 2010. Polymodal
foraging in adult female loggerheads (Caretta caretta). Marine Biology 157: 651–663.
Reisinger, R.R. and M.N. Bester. 2010. Goose barnalces on seals and a penguin at Gough Island. African
Zoology 45: 129–132.
Relini, G. 1968. Segnalazione di du cirripedi nuovi per l’Adriatico. Bolletin de Societie du Adriatica Sciencia
Trieste 56: 218–225.
Relini, G. 1969. La distribuzione dei Cirripedi Toracice nei mari Italiani. Archaeologie Botanica Biogreoria
Italia 4, 45: 169–186.
Relini, G. 1980. Cirripedi toracici. Guide per il Riconoscimento delle Specie Animali delle Acque Lagunari e
Costiere Italiane 2: 1–122.
Ren, X. 1980. Turtle barnacles of the Xisha Islands, Guangdong Province, China. Studia Marina Sinica
420 The Biology of Sea Turtles, Volume III
Ren, X. 1987. Studies on Chinese Cirripedia (Crustacea) VIII. Supplementary Report. Studia Marina Sinica
Richards, H.G. 1930. Notes on the barnacles from Cape May County, New Jersey. Proceedings of the Academy
of Natural Sciences of the Philadelphia 83: 143–144.
Ross, J. 1981. Hawksbill turtle Eretmochelys imbricata in the Sultanate of Oman. Biological Conservation 19:
Ross, A. and M.G. Frick. 2011. Nomenclatural emendations of the family-group names Cylindrolepadinae,
Stomatolepadinae, Chelolepadinae, Cryptolepadinae, and Tubicinellinae of Ross & Frick, 2007—
Including current denitions of family-groups within the Coronuloidea (Cirripedia: Balanomorpha).
Zootaxa 3106: 60–66.
Ross, A. and W.A. Newman. 1967. Eocene Balanidae of Florida, including a new genus and species with a
unique plan of “turtle-barnacle” organization. American Museum Novitates 2288: 1–21.
Rudloe, J., A. Rudloe and L. Ogren. 1991. Occurrence of immature Kemp’s ridley turtles, Lepidochelys kempi,
in coastal waters of northwest Florida. Northwest Gulf Science 12: 49–53.
Ruxton, G.D. 2009. Non-visual crypsis: A review of the empirical evidence for camouage to sense other than
vision. Philosophical Transactions of the Royal Society B364: 540–557.
Sazima, C., A. Grossman, and I. Sazima. 2010. Turtle cleaners: Reef shes foraging on epibionts of sea turtles in the
tropical southwestern Atlantic, with a summary of this association type. Neotropical Ichthyology 8: 187–192.
Schärer, M.T. 2001. A survey of the epibiota of Eretmochelys imbricata (Testudines: Cheloniidae) of Mona
Island, Puerto Rico. Revista de Biologia Tropical 51: 87–89.
Schärer, M.T. and J.H. Epler. 2007. Long-range dispersal possibilities via sea turtle—A case study for Clunio
and Pontomyia (Diptera: Chironomidae) in Puerto Rico. Entomological News 118: 273–277.
Schoeld, G., K.A. Katselidis, P. Dimopoulos, J.D. Pantis, and G.C. Hays. 2006. Behaviour analysis of the
loggerhead sea turtle Caretta caretta from direct in-water observation. Endangered Species Research
Schwartz, F.J. 1960. The barnacle, Platylepas hexastylos, encrusting a green turtle, Chelonia mydas mydas,
from Chincoteague Bay, Maryland. Chesapeake Science 1: 116–117.
Senties, G.A., J. Espinoza-Avalos, and J.C. Zurita. 1999. Epizoic algae of nesting sea turtles Caretta caretta
(L.) and Chelonia mydas (L.) from the Mexican Caribbean. Bulletin of Marine Science 64: 185–188.
Sezgin, M., A.S. Ateş, T. Katağan, K. Bakir, and Ş. Yalçin Özkilek. 2009. Notes on amphipods Caprella
andreae Mayer, 1890 and Podocerus chelonophilus (Chevreux and Guerne, 1888) collected from the
loggerhead sea turtle, Caretta caretta, off the Mediterranean and the Aegean coasts of Turkey. Turkish
Journal of Zoology 33: 433–437.
Shine, R., F. Brischoux, and A.J. Pile. 2010. A seasnake’s colour affects its susceptibility to algal fouling.
Proceedings of the Royal Society B 277: 2459–64.
Smaldon, G. and I.H.J. Lyster. 1976. Stomatolepas elegans (Costa, 1840) (Cirripedia): New records and notes.
Crustaceana 30: 317–318.
Stinson, M.L. 1984. Biology of sea turtles in San Diego Bay, California, and in the northeastern Pacic. Master
thesis, San Diego State University, San Diego, CA, 578pp.
Stubbings, H.G. 1965. West African Cirripedia in the collections of the Institut Francais d’Afrique Noire,
Dakar, Senegal. Bulletin de l’Institut Français d’Afrique Noire, Series A 27: 876–907.
Stubbings, H.G. 1967. The cirriped fauna of tropical West Africa. Bulletin of the British Museum (Natural
History) Zoology 15: 229–319.
Tachikawa, H. 1995. Notes on three species of stalked barnacles found from a turtle barnacle on the carapace
of a green turtle, Chelonia mydas. Nanki Seibutu 37: 67–68.
Turtle Expert Working Group. 2009. An assessment of the loggerhead turtle population in the western northern
Atlantic Ocean. NOAA Technical Memorandum NMFS-SEFSC-575.
Utinomi, H. 1949. Studies on the cirripedian fauna of Japan. VI. Cirripeds from Kyusyu and Ryukuyu Islands.
Publications of the Seto Marine Biological Laboratory 1: 19–37.
Utinomi, H. 1950. Cirripeds commonly taken by dredging near Tanabe Bay (Record of collections dredged
from off Minabe Prov. Kii, IV). Nanki Seibutu 2: 60–65.
Utinomi, H. 1958. Studies on the cirripedian fauna of Japan. VII. Cirripeds from Sagami Bay. Publications of
the Seto Marine Biological Laboratory 4: 281–311.
Utinomi, H. 1959. Thoracic cirripeds from the environs of Banyuls. Vie et Milieu 10: 379–399.
Utinomi, H. 1966. Fauna and ora of the sea around the Amakusa Marine Biological Laboratory. Part VI.,
Cirriped Crustacea. Amakusa Marine Biological Laboratory, July 1966, pp. 1–11.
421Sea Turtle Epibiosis
Utinomi, H. 1969. Cirripedia of the Iranian Gulf. Videnskabelige Meddelelser Dansk Naturhistorisk Forening
Utinomi, H. 1970. Studies on the cirripedian fauna of Japan. IX. Distributional survey of thoracic cirripeds
in the southeastern part of the Japan Sea. Publications of the Seto Marine Biological Laboratory
Vivaldo, S.G., D.O. Sarabia, C.P. Salazar, A.G. Hernandez, and J.R. Lezama. 2006. Identication of parasites
and epibionts in the olive ridley turtle (Lepidochelys olivacea) that arrived to the beaches of Michoacan
and Oaxaca, Mexico. Veterinaria Mexico 37: 431–440.
Wagh, A.B. and D.V. Bal. 1974. Observations on the systematics of sessile barnacles from the west coast of
India-1. Journal of Bombay Natural History Society 71: 109–123.
Wahl, M. 1989. Marine epibiosis. I. Fouling and antifouling: Some basic aspects. Marine Ecology Progress
Series 58: 175–89.
Wahl, M. 2009. Epibiosis: Ecology, effects and defences. In M. Wahl, ed. Marine Hardbottom Communities.
Ecological Studies Series, Vol. 206. Springer, Berlin, Germany, pp. 61–72.
Wahl, M., O. Mark. 1999. The predominately facultative nature of epibiosis: Experimental and observational
evidence. Marine Ecology Progress Series 187: 59–66.
Walker, G. 1978. A cytological study of the cement apparatus of the barnacle, Chelonibia testudinaria Linnaeus,
an epizoite on turtles. Bulletin of Marine Science 28: 205–209.
Wass, M.L. 1963. Check list of the marine invertebrates of Virginia. Virginia Institute of Marine Science
Gloucester Point, Virginia, Spec. Sci. Rept. No. 24 (revised): 1–56.
Wells, H.W. 1966. Barnacles of the northeastern Gulf of Mexico. Quarterly Journal of the Florida Academy
of Science 29: 81–95.
Weltner, W. 1897. Verzeichnis der bisher beschriebenen recenten Cirripedienarten. Mit Angabe der im berliner
Museum vorhandenen Species und ihrer Fundorte. Archiv für Naturgeschicthe 1: 227–280.
Weltner, W. 1910. Cirripedien von Ostafrika. In: Reise in Ostafrika. V.A. Voeltzkow, Stuttgart, Germany,
Wyneken, J. 1997. Sea turtle locomotion: Mechanisms, behavior, and energetics. In P.L. Lutz and J.A. Musick,
editors. The Biology of Sea Turtles. CRC Press, Boca Raton, FL, pp. 165–198.
Young, P.S. 1999. Subclasse Cirripedia. In: L. Buckup and G. Bond-Buckup, eds. Os Crustáceos do Rio Grande
do Sul. Porto Alegre, Ed. Universidade/UFRGS, pp. 24–53.
Young, P.S. and A. Ross. 2000. Cirripedia. In: J.E.L. Bosquets, E.G. Soriano and N. Papavero, eds. Bioversidad,
Taxonomia y Biogeographia de Arthropodos de Mexico: Hacia una Sintesis de su Conocimiento. Vol. II.
Universidad Nacional Autonomia de Mexico, Mexico, pp. 213–237.
Zakhama-Sraieb, R., S. Karaa, M.N. Bradai, I. Jribi, and F. Char-Cheikhrouha. 2010. Amphipod epibionts
of the sea turtles Caretta caretta and Chelonia mydas from the Gulf of Gabès (central Mediterranean).
Journal of the Marine Biological Association of the United Kingdom Marine Biodiversity
Records 3: e38.
Zann, L.P. and B.M. Harker. 1978. Egg production of the barnacles Platylepas ophiophilus Lanchester,
Platylepas hexastylos (O. Fabricius), Octolasmis warwickii Gray and Lepas anatifera Linnaeus.
Crustaceana 35: 206–214.
Zardus, J.D. and G.H. Balazs. 2007. Two previously unreported barnacles commensal with the green sea turtle,
Chelonia mydas (Linnaeus, 1758), in Hawaii and a comparison of their attachment modes. Crustaceana
Zardus, J.D. and M.G. Hadeld. 2004. Larval development and complemental males in Chelonibia testudi-
naria, a barnacle commensal with sea turtles. Journal of Crustacean Biology 24: 409–421.
Zavodnik, D. 1997. Chthamalus montagui and Platylepas hexastylos two cirriped crustaceans new to the east-
ern Adriatic Sea. Natura Croatica (Croatian Nat. Hist. Mus.) 6: 113–118.
Zullo, V.A. 1986. Quaternary barnacles from the Galapagos Islands. Proceedings of the California Academy of
Sciences 44: 55–66.
Zullo, V.A. 1991. Zoogeography of the shallow water cirriped fauna of the Galapagos Islands and adjacent
regions in the tropical Eastern Pacic. In M.J. Jones, ed. Galapagos Marine Invertebrates. Taxonomy,
Biogeography and Evolution in Darwin’s Islands. Plenum Publishing Company, New York.
Zullo, V.A. and J.S. Bleakney. 1966. The cirriped Stomatolepas elegans (Costa) on leatherback turtles from
Nova Scotian waters. Canadian Field-Naturalist 80: 163–165.
Zullo, V.A. and W.H. Lang. 1978. Order Cirripedia. In: R.G. Zingmark, ed. Annotated Checklist of the Biota of
the Coastal Zone of South Carolina. University of South Carolina, Columbia, SC, pp. 158–160.
422 The Biology of Sea Turtles, Volume III
Allee, W.C., A.E. Emerson, O. Park, T. Park, and K.P. Schmidt. 1949. Principles of Animal Ecology. W.B.
Saunders Co., Pennsylvania, PA.
Allen, E.R. and W.T. Neill. 1952. Know your reptiles: The diamondback terrapin. Florida Wildlife 6: 42.
Angulo-Lozano, L., P.E. Nava-Duran, and M.G. Frick. 2007. Epibionts of olive ridley turtles (Lepidochelys
olivacea) nesting at Playa Ceuta, Sinaloa, Mexico. Marine Turtle Newsletter 118: 13–14.
Aradas, A. 1869. Desrizione di una nuova specie del genere Coronula. Atti della Accademia Gioenia di Scienze
Naturali in Catania 43: 215–224.
Arndt, R.G. 1975. The occurrence of barnacles and algae on the red-bellied turtle, Chrysemys r. rubiventris
(LeConte). Journal of Herpetology 9: 357–359.
Aurivillius, C.W.S. 1894. Studien uber cirripedien. Kongliga Svenska Vetenskaps-Akademien 26: 1–107.
Ayling, A.M. 1976. The strategy of orientation in the barnacle Balanus trigonus. Marine Biology 36: 335–342.
Aznar, F.J., J.A. Balbuena, and J.A. Raga. 1994. Are epizoites indicators of a western Mediterranean striped
dolphin die-off? Diseases of Aquatic Organisms 18: 159–163.
Baer, J.G. 1951. Ecology of Animal Parasites. University of Illinois Press, Urbana, IL.
Barnes, M. 1989. Egg production in cirripedes. Oceanography and Marine Biology Annotated Review
Bleakney, J.S. 1967. Food items in two loggerhead sea turtles, Caretta caretta caretta (L.) from Nova Scotia.
Canadian Field-Naturalist 81: 169–272.
Booth, J. and J.A. Peters. 1972. Behavioral studies on the green turtle (Chelonia mydas) in the sea. Animal
Behaviour 20: 808–812.
Bourget, E. 1977. Shell structure in sessile barnacles. Naturaliste Canadien 104: 281–323.
Briggs, K.T. and G.V. Morejohn. 1972. Barnacle orientation and water ow characteristics in California grey
whales. Journal of Zoology 167: 287–292.
Cake, E.W., Jr. 1983. Symbiotic associations involving the southern oyster drill Thais haemastoma oridana
(Conrad) and macrocrustaceans in Mississippi waters. Journal of Shellsh Research 3: 117–128.
Carr, A. 1952. Handbook of Turtles. Cornell University Press, New York.
Carr, A. 1964. Transoceanic migrations of the green turtle. Bioscience 14: 49–52.
Carr, A. 1965. The navigation of the green turtle. Scientic American. 12: 79–86.
Chen, Y. 1989. Cirripedia. In C. Wei and Y. Chen, eds. Fauna of Zhejiang: Crustacea. Zhejiang Science and
Technology Publishing House, Hangzhou, Zhejiang Province, China, pp. 38–73.
Crisp, D.J. 1960. Mobility of barnacles. Nature 188: 1208–1209.
Crisp, D.J. 1974. Factors inuencing the settlement of marine invertebrate larvae. In P.T. Grant and
A.M. Mackie, eds. Chemoreception in Marine Organisms. Academic Press, Inc., NY, pp. 177–265.
Crisp, D.J. 1983. Chelonibia patula (Ranzani), a pointer to the evolution of the complemental male. Marine
Biology Letters 4: 281–294.
Crisp, D.J. and J.D. Costlow, Jr. 1963. The tolerance of developing cirripede embryos to salinity and temperature.
Oikos 14: 22–34.
Crisp, D.J. and H.G. Stubbings. 1957. The orientation of barnacles to water currents. Journal of Animal Ecology
Dall, W.H. 1872. On the parasites of the cetaceans of the N.W. coast of America, with descriptions of new
forms. Proceedings of the California Academy of Sciences 4: 299–301.
deAlessandri, G. 1895. Contribuzione allo studio dei Cirripedi fossili d’Italia. Bollettino della Società
Geologica Italian 13: 234–314.
deAlessandri, G. 1906. Studi monograci sui cirripedi fossili d’Italia. Palaeon Italica 12: 207–324.
Eckert, K.L. and S.A. Eckert. 1987. Growth rate and reproductive condition of the barnacle Conchoderma
virgatum on gravid leatherback sea turtles in Caribbean waters. Journal of Crustacean Biology
Fabricius, O. 1790. Beschribung zweiter neuer Gattung Meereichein (Lepades) nebst der Islandischen
Kammuschel (Ostrea islandica) mit abbildungen. Schriften der Berlinischen Gesellschaft
Naturforschender Freunde 1: 101–111.
Fabricius, O. 1798. Tillaeg-til Conchylie-Slaegterne Lepas, Pholas, Mya, Solen. Skrivter av Naturhiftorie
Selskabet Kiobenhavn 4: 34–51.
Felix, F., B. Bearson, and J. Falconi. 2006. Epizoic barnacles removed from the skin of a humpback whale after
a period of intense surface activity. Marine Mammal Science 22: 979–984.
Fischer, P. 1884. Cirrhipedes de l’archipel de la Nouvelle-Caladonie. Bulletin de la Société Zoologique de
France 9: 355–360.
423Sea Turtle Epibiosis
Frazier, J. 1986. Epizoic barnacles on pleurodiran turtles: Is the relationship rare? Proceedings of the Biological
Society Washington 99: 472–477.
Frazier, J.G. and D. Margaritoulis. 1990. The occurrence of the barnacle, Chelonibia patula (Ranzani, 1818) on
an inanimate substratum (Cirripedia, Thoracica). Crustaceana 59: 213–218.
Frick, M.G. and A. Ross. 2002. Happenstance or design: An unusual association between a turtle, and octocoral
and a barnacle. Marine Turtle Newsletter 97: 10–11.
Gamez Vivaldo, S., D. Osorio Sarabia, C. Peneores Salazar, A. Garcia Hernandez, and J. Ramirez Lezama.
2006. Identicacion de parasitos y epibiontes de la tortuga golna (Lepidochelys olivacea) que arribo a
playas de Michoacan y Oaxaca, Mexico. Veterinaria Mexico 37: 431–440.
Geraci, J.R. and D.J. St. Aubin. 1987. Effects of parasites on marine mammals. International Journal of
Parasitology 17: 407–414.
Gittings, S.R., G.D. Denis, and H.W. Harry. 1986. Annotated guide to the barnacles of the northern Gulf of
Mexico. Texas A&M University Sea Grant College Program 86–402: 1–36.
Grant, C. 1956. Aberrant lamination in two hawksbill turtles. Herpetologica 12: 302.
Gray, J.E. 1825. A synopsis on the genera of Cirripedes arranged in natural families, with a description of some
new species. Annals of Philosophy 10: 97–107.
Greef, S.R. 1885. Ueber die Fauna der Guinea-Inseln S. Thome und Rolas. Sitzungsberichte der Gesellschaft
zur Beforderung der gesammten Naturwissenschaften zu Marburg 41–80.
Guess, R.C. 1982. Occurrence of a Pacic loggerhead turtle, Caretta caretta gigas Deraniyagala, in the waters
off Santa Cruz Island, California. California Fish and Game 68: 122–123.
Gutmann, W.F. 1960. Funktionelle morphologie van Balanus balanoides. Abhandlungen der Senckenbergischen
Naturforschenden Gesellschaft 500: 1–43.
Healy, J.M. and D.T. Anderson. 1990. Sperm ultrastructure in the Cirripedia and its phylogenetic signicance.
Records of the Australian Museum 42: 1–26.
Hiro, F. 1937b. Studies on the cirripedien fauna of Japan. II. Cirripeds found in the vicinity of the Seto Marine
Biological Laboratory. Memoirs of the College of Science, University of Kyoto (Series B) 12: 385–478.
Jackson, C.G., Jr. and A. Ross. 1971. The occurrence of barnacles on the alligator snapping turtle, Macroclemys
temmincki (Troost). Journal of Herpetology 5: 188–189.
Jackson, C.G., Jr. and A. Ross. 1972. Balanomorph barnacles on Chrysemys alabamensis. Quarterly Journal of
the Florida Academy of Science 35: 173–176.
Jackson, C.G., Jr. and A. Ross. 1975. Epizoic occurrence of a bryozoan, Electra crustulenta, on the turtle
Chrysemys alabamensis. Transactions of the American Microscopy Society 94: 135–136.
Jackson, C.G., Jr., A. Ross, and G.L. Kennedy. 1973. Epifaunal invertebrates of the ornate diamondback
terrapin, Malaclemys terrapin macrospilota. American Midland Naturalist 89: 495–497.
Johnson, T.W., Jr. and R.R. Bonner. 1960. Lagenidium callinectes Couch in barnacle ova. Journal of the Elisha
Mitchell Society 76: 147–149.
Kadovich, J. 1961. Relationship of some marine organisms of the northeast Pacic to water temperatures,
particularly during 1957 through 1959. California Fish and Game, Fisheries Bulletin 112: 1–62.
Karuppiah, S., A. Subramanian, and J.P. Obbard. 2004. The barnacle, Xenobalanus globicipitis (Cirripedia,
Coronulidae), attached to the bottle-nosed dolphin, Tursiops truncatus (Mammalia, Cetacea) on the
southeastern coast of India. Crustaceana 77: 879–882.
Kasuya, T. and D.W. Rice. 1970. Notes on the baleen plates and arrangement of parasitic barnacles of the gray
whale. Scientic Reports of the Whales Research Institute 22: 39–43.
Kato, M., K. Hayasaka, and T. Matsuda. 1960. Ecological studies on the morphological variation of a sessile
barnacle, Chthamalus challengeri. III. Variation of the shell shape and of the inner anatomical features
introduced by the population density. Bulletin of the Marine Biological Station of Asamushi, Tohoku
University 10: 19–25.
Key, M.M., Jr., J.W. Volpe, W.B. Jeffries, and H.K. Voris. 1997. Barnacle fouling of the blue crab Callinectes
sapidus at Beaufort, North Carolina. Journal of Crustacean Biology 17: 424–439.
Kim, I.H. and H.S. Kim. 1980. Systematic studies on the cirripeds (Crustacea) from Korea. 1. Balanomorph
barnacles (Cirripedia, Thoracica, Balanomorpha). Korean Journal of Zoology 23: 161–194.
Kitsos, M., M. Christodoulou, S. Kalpakis, M. Noidou, and A. Koukouras. 2003. Cirripedia Thoracica
associated with Caretta caretta (Linnaeus, 1758) in the northern Aegean Sea. Crustaceana 76: 403–409.
Klepal, W. 1987. A review of the comparative anatomy of the males in cirripedes. Oceanography and Marine
Biology: Annual Review 25: 285–351.
Kolosvary, G. 1942. Studien an Cirripedien. Zoologischer Anzeige 137, pp. 138–150.
Kolosvary, G. 1947. Der balaniden der Adria. Annales Historico-Naturales Musei Nationalis Hungarici
424 The Biology of Sea Turtles, Volume III
Kruger, P. 1911a. Beitrage zur Cirripedien fauna Ostasiens. Abhandlungen der Mathematisch-Physikalische
Klasse der Königlich Bayerischen Akademie der Wissenschaften, II Supplement-Band 6: 1–72.
Lamarck, J.B.A. de M. de. 1802. Mémoire sur la Tubicinelle. Annales du Museum National d’Histoire Naturelle
Lang, W.H. 1979. Larval development of shallow water barnacles of the Carolinas (Cirripedia: Thoracica) with
keys to naupliar stages. NOAA-Technical Report-NMFS Circular-421, pp. 1–39.
Leach, W.E. 1817. Distribution systematique de la class Cirripèdes. Journal de Physique de Chimie et d’Histoire
Naturelle 85: 67–69.
Leach, W.E. 1818. Cirripedes. In: Supplement to the fourth and fth editions of the Encyclopedia Britannica
Lezama, C., M. Lopez-Mendilaharsu, F. Scarabino, A. Estrades, and A. Fallabrino. 2006. Interaction between
the green sea turtle (Chelonia mydas) and an alien gastropod (Rapana venosa) in Uruguay. In M.G. Frick,
A. Panagopoulou, A.F. Rees and K. Williams, compilers. Proceedings of the 26th Annual Symposium on
Sea Turtle Biology and Conservation, Miami, FL, pp. 64–65. ISBN: 960-87926-1-4.
Linnaeus, C. 1767. Systema naturae per regna tria naturae—Edito duodecima, reformata. Holmiae 1: 533–1327.
Lutcavage, M.E. and J.A. Musick. 1985. Aspects of the biology of sea turtles in Virginia. Copeia 1985: 449–456.
Meischner, D. 2001. Seepocken auf einer Meeres-Schildkröte, ein ökologisches Idyll. Nature und Museum
Mignucci-Giannoni, A.A., C.A. Beck, R.A. Montoya-Ospina and E.H. Williams Jr. 1999. Parasites and
commensals of the West Indian manatee from Puerto Rico. Journal of the Helminthological Society of
Washington 66: 67–69.
Miranda, L. and R.A. Moreno. 2002. Epibionts from Lepidochelys olivacea (Eschscholtz, 1829) (Reptilia:
Testudinata: Cheloniidae) in the central south region of Chile. Revista de Biologia Marina y Oceanograa
Mokaday, O., Y. Loya, Y. Achituv, E. Geffen, D. Graur, S. Rozenblatt and I. Bricker. 1999. Speciation versus
phenotypic plasticity in coral inhabiting barnacles: Darwin’s observation in an ecological context.
Journal of Molecular Evolution 49: 367–375.
Monroe, R. 1981. Studies on the Coronulidae (Cirripedia): Shell morphology, growth, and function, and their
bearing on subfamily classication. Memoirs of the Queensland Museum 20: 237–251.
Mörch, O.A.L. 1852. Cephalophora Catalogus Conchyliorum (Cirripedia) 1: 65–68.
Newman, W.A. 1996. Sous-classe des Cirripèdes (Cirripedia Burmeister, 1834), Super-ordres des Thoraciques
et des Acrothoraciques (Thoracica Darwin, 1854—Acrothoracica Gruvel, 1905). Traité de Zoologie
Newman, W.A. and A. Ross. 1971. Antarctic Cirripedia. American Geophysical Union, Antarctic Research
Series 14: 1–257.
Newman, W.A. and A. Ross. 1976. Revision of the balanomorph barnacles; including a catalog of the species.
San Diego Society of Natural History 9: 1–108.
Newman, W.A. and A. Ross. 1977. A living Tesseropora (Cirripedia: Balanomorpha) from Bermuda and the
Azores: First records from the Atlantic since the Oligocene. Transactions of the San Diego Society of
Natural History 18: 207–216.
Newman, W.A. and A. Ross. 2001. Prospectus on larval cirriped setation formulae, revisited. Journal of
Crustacean Biology 21: 56–77.
Nilsson-Cantell, C.A. 1930b. Cirripedes. In: Resultates Scientiques du Voyage aux Indes Orientales
Neerlandaises de LL. AA. RR. le Prince et la Princesse Leopold de Belgique. Mémoires du Musee Royal
d’Histoire Naturelle de Belgique 3: 1–24.
Nilsson-Cantell, C.A. 1957. Thoracic cirripeds from Chile. Lunds Universitets Arsskrift N.F. (Series 2),
Nogata, Y. and K. Matsumura. 2006. Larval development and settlement of a whale barnacle. Biology Letters
Orams, M.B. and C. Schuetze. 1998. Seasonal and age/size-related occurrence of a barnacle (Xenobalanus
globicipitis) on bottlenose dolphins (Tursiops truncatus). Marine Mammal Science 14: 186–189.
Ortiz, M., R. Lalana, and C. Varela. 2004. Caso extremo de epibiosis de escaramujos (Cirripedia: Balanomorpha),
sobre una esquilla (Hoplocarida: Stomatopoda), en Cuba. Revista de Investigaciones Marinas 25: 75–76.
Pasternak, Z, A. Abelson, and Y. Achituv. 2002. Orientation of Chelonibia patula (Crustacea: Cirripedia) on the
carapace of its crab host as determined by the feeding mechanism of the adult barnacles. Journal of the
Marine Biological Association of the United Kingdom 82: 583–588.
Peterson, M.N.A. 1966. Calcite: Rates of dissolution in a vertical prole in the central Pacic. Science
425Sea Turtle Epibiosis
Pilsbry, H.A. 1910. Stomatolepas, a commensal barnacle in the throat of the loggerhead turtle. American
Naturalist 44: 304–306.
Pitombo, F.B. 2004. Phylogenetic analysis of the Balanidae (Cirripedia: Balanomorpha). Zoologica Scripta
Rees, E.I.S. and G. Walker. 1977. A record of the turtle barnacle Chelonobia [sic] testudinaria (L.) in the Irish
Sea. Porcupine Newsletter 5: 189.
Rice, D.W. and A.A. Wolman. 1971. The life history and ecology of the gray whale (Eschrichtius robustus).
American Society of Mammalogists, Special Publication 3: 1–142.
Ridgway, S.H., E. Linder, K.A. Mahoney, and W.A. Newman. 1997. Grey whale barnacles Cryptolepas
rhachinecti infest white whales, Delphinapterus leucas, housed in San Diego Bay. Bulletin of Marine
Science 61: 377–385.
Riedl, R. 1963. Fauna und Flora der Adria. Cirripedia only, pp. 10–15, pls. 1–2, pp. 18–19, 252–258, Figs.
83–84, 2 maps. Paul Parey Verlag. Hamburg.
Ross, A. 1963a. A new Pleistocene Platylepas from Florida. Quarterly Journal of the Florida Academy of
Science 26: 150–158.
Ross, A. 1963b. Chelonibia in the Neogene of Florida. Quarterly Journal of the Florida Academy of Science
Ross, A. 1964. Type locality of Platylepas wilsoni Ross. Quarterly Journal of the Florida Academy of Science
Ross, A. and W.K. Emerson. 1974. Wonders of Barnacles. Dodd, Mead and Co., New York.
Ross, A. and M.G. Frick. 2007. From Hendrickson (1958) to Monroe and Limpus (1979) and beyond: An
evaluation of the turtle barnacle Tubicinella cheloniae. Marine Turtle Newsletter 118: 2–5.
Ross, A. and W.A. Newman. 1995. A coral-eating barnacle, revisited (Cirripedia, Pyrgomatidae). Contributions
to Zoology 65: 129–175.
Ross, A. and W.A. Newman. 2000. Pyrgoma kuri Hoek, 1913: a case study in morphology and systematics of a
symbiotic coral barnacle (Cirripedia: Balanamorpha). Contributions to Zoology 68: 245–260.
Ryder, J.A. 1879. Strange habitat of a barnacle on a gar pike. American Naturalist 8: 453.
Samaras, W.F. and F.E. Durham. 1985. Feeding relationship of two species of epizoic amphipods and
the gray whale, Eschrichtius robustus. Bulletin of the Southern California Academy of Sciences
Scaravelli, D. 1998. Segnalazioni faunistiche. 29. Stomatolepas elegans (O.G. Costa 1838) (Crustacea
Thoracica Balanidae). Quaderno di Studi e Notizie di Storia Naturale Della Romagna 10: 78.
Scarff, J.E. 1986. Occurrence of the barnacles, Coronula diadema, C. reginae and Cetopirus complanatus
(Cirripedia) on right whales. Scientic Reports of the Whales Research Institute 37: 129–153.
Schmitt, W.L. 1965. Crustaceans. University of Michigan Press. Ann Arbor, MI.
Seigel, R.A. 1983. Occurrence and effects of barnacle infestation on diamondback terrapins (Malaclemys
terrapin). American Midland Naturalist 109: 34–39.
Shields, J.D. 1992. Parasites and symbionts of the crab Portunus pelagicus from Moreton Bay, eastern Australia.
Journal of Crustacean Biology 12: 94–100.
Southward, A.J. 1986. Class Cirripedia (barnacles). In W. Sterrer, ed. Marine Fauna and Flora of Bermuda. A
Systematic Guide to the Identication of Marine organisms. John Wiley & Sons, New York, pp. 299–305.
Southward, A.J., ed. 1987. Barnacle Biology. A.A. Balkema, Rotterdam, the Netherlands, i–xxii + 443pp.
Southward, A.J. 1998. New observations on barnacles (Crustacea: Cirripedia) of the Azores region. Arquipelago.
Life and Marine Sciences 16: 11–27.
Spears, T.L., L.G. Abele, and M.A. Applegate. 1994. A phylogenetic study of cirripeds and their relatives
(Crustacea: Thecostraca). Journal of Crustacean Biology 14: 641–656.
Steenstrup, J.J.S. 1851. Videnskabelige Meddelelser fra den Naturhist. Forening i Kjöbenhavn, for Aaret, 1851.
Table 3, Fig. 11–15.
Stunkard, H.W. 1955. Freedom, bondage and the welfare state. Science 121: 811–816.
Walker, L.W. 1949. Nursery of the gray whales. Natural History 58: 248–256.
Weltner, W. 1895. Die Cirripedien von Patagonien, Chile and Juan Fernandez. Archiv für Naturgeschicthe
Williams, K.L. and M.G. Frick. 2008. Tag returns from loggerhead turtles from Wassaw Island, GA.
Southeastern Naturalist 7: 165–172.
Williams, A.B. and H.J. Porter. 1964. An unusually large turtle barnacle (Chelonibia patula) on a blue crab
from Delaware Bay. Chesapeake Science 5: 150–153.
Withers, T.H. 1928. The cirriped Chelonibia caretta Spengler, in the Miocene of Zanzibar Protectorate. Annals
and Magazine of Natural History, Series 10, 2: 390–392.
426 The Biology of Sea Turtles, Volume III
Withers, T.H. 1929. The cirriped Chelonibia in the Miocene of Gironde, France and Vienna, Austria. Annals
and Magazine of Natural History, Series 10, 4: 566–572.
Withers, T.H. 1953. Catalogue of Fossil Cirripedia in the Department of Geology, Vol. 3, Tertiary. British
Museum (Natural History), 396 pp.
Yasuyuki, N. and K. Matsumura. 2006. Larval development and settlement of a whale barnacle. Biology Letters
Young, P.S. 1991. The superfamily Coronuloidea Leach (Cirripedia, Balanomorpha) from the Brazilian coast,
with redescription of Stomatolepas species. Crustaceana 61: 190–212.
Zangerl, R. 1948. The vertebrate fauna of the Selma Formation of Alabama. Part I. Fieldiana: Geology Memoirs
Zann, L.P. 1975. Biology of a Barnacle (Platylepas ophiophilus Lanchester) Symbiotic with Sea Snakes.
University Park Press, Baltimore, MA, pp. 267–286.
Zullo, V.A. 1963. A preliminary report on systematic and distribution of barnacles (Cirripedia) of the Cape Cod
region. Marine Biology Laboratory, Woods Hole, MA. Systematics Ecology Program, 33pp.
Zullo, V.A. 1969. Thoracic Cirripedia of the San Diego formation, San Diego County, California. Contributions
in Science Las Angeles County Museum 159: 1–25.
Zullo, V.A. 1979. Marine ora and fauna of the northeastern United States, Arthropoda: Cirripedia. NOAA
Technical Report NMFS Circular 425: 1–29.
Zullo, V.A. 1982. A new species of the turtle barnacle Chelonibia Leach, 1817, (Cirripedia, Thoracica) from the
Oligocene Mint Spring and Byram Formations of Mississippi. Mississippi Geology 2: 1–6.