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Keith Philippe
Undergraduate, University of Minnesota, April 2007
Marine Biology, Prof. Peter W. Sorensen
Existing literature indicate that certain species of whales are well known for harboring
body parts of their host have definite organization. These include species richness,
population density, and microhabitat selection. Lice community organizations result from
This article describes parasitic occurrence of lice (cyamid amphipods) on three species of
right whales: (i) The North Atlantic (Eubalaena glacialis); (ii) The Southern (Eubalaena
australias); and (iii) a poorly known remnant population, Eubalaena japonica. To help
accomplish its objective, this review will focus primarily on: (i) evolutionary trends of
right whales and lice; (ii) microhabitat selection of lice on right whales; and (iii)
Louse (pl. lice) are small creatures usually less than 5mm long. Their first instar nymphs
(a phase between two periods of molting in the development of an insect larva or other
invertebrate animal.) are less than 1mm long. Lice are dorsoventrally1 flattened and
1
axis joining the dorsal and ventral surfaces
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possess many short setae2. With little variation in color, lice are generally off-white with a
few having brownish or reddish hues. They generally remain near the surface of the skin
where they feed, copulate, and lay eggs. Most lice have large claws on their tarsi in which
Although various forms of social symbiosis exist between whales and lice, right whales
and their louse symbionts appear to share both commensalic (relationship in which one
party gains some benefit, whilst the other suffers no serious disadvantage) and phoretic
relationships.
Most cetaceans carry populations of benign ectoparasites called cyamids (whale lice),
isopods, lampreys and remoras (Kim, 1985). Cyamids spend all life stages 3 on whales,
feeding on the outer surface of their host’s epidermis. They roam widely on their hosts,
but because they have no free-swimming stage they can migrate only between whales
cyamid population is well defined and sharply bounded, consisting of many virtually
identical and constantly moving habitat islands (whales) that occasionally replace them
selves through a simple birth-and-death process (Seger, 2005). Right and grey whales
(Eschrichtius robustus) are unusual in that each carries large populations of three cyamid
2
a stiff hair like or bristle like structure, esp. in an invertebrate.
3
The life stages of louse include three phases: (i) the egg or nit; (ii) the nymph; and (iii) the adult louse.
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representatives that are not common among other cetaceans. Grey whale cyamids
(Cyamus ceti, Cyamus kessleri, Cyamus scammoni) are distantly related to right whale
cyamids. Right whale cyamids are traditionally referred to as Cyamus ovalis, Cyamus
gracilis and Cyamus erraticus. Every adult right whale carries populations of all three
cyamid species. On a typical adult, the callosities4 provide about 0.5 m2 of substrate for
C. ovalis which is by far the most abundant of the three species with about 5000 adults
per adult right whale, based on an estimated density of roughly one adult cyamid per cm2
The Cyamids of Right Whales occupy ecologically suitable regions of whale hosts. The
details on such selectivity are not fully understood. Cyamus ovalis ,for example, blankets
the callosities (raised patches of roughened skin on the head); and their white bodies
make the callosities far more pronounce against the black skin of the whale’s head and
back. Each right whale has distinct callositive patterns that can be identified in
photographs; owing to the visual contrast and sharp delineation of callosity boundaries
created by a living blanket of Cyamus ovalis. Cyamus gracilis occupies pits and grooves
between the elevated patches of callosity tissue. Adults of this species are smaller and
thinner than adult Cyamus ovalis. Cyamus. erraticus occupies smooth skin in the genital
and mammary slits; it is highly mobile and opportunistically colonizes wounds and other
areas of reduced water flow not occupied by the other two species; it also ‘blooms’ on the
heads of young calves, but these concentrations disappear within a few months (Seger,
2005).
4
a thickened and hardened part of whale skin; a callus.
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Morphological and behavioral adaptations of parasitic arthropods are strongly correlated
with mammalian skin type. The mammalian skin provides numerous microhabitats for
different parasitic arthropods to reside and obtain food (Kim,1985). There is much
speculation on what whale lice feed on when attached to their hosts. Some suggest
cyamids feed on shedding integument. Others indicate neither Anoplura nor Mallophage
Some cyamids occur on more than one cetacean species, but few cetaceans normally host
more than one species of cyamid at a given period. The slow-swimming right whales and
grey whale (Eschrichtius robustus) are unusual in that each carries large populations of
three cyamid species that do not occur regularly on any other cetaceans. The grey whale
cyamids (Cyamus ceti, Cyamus kessleri, Cyamus scammoni) are only distantly related to
each other and to the right whale cyamids, and vice versa (Seger, 2005).
researchers used nucleic material in cyamid and Eubalaena mitochondria. Their results
indicated that one subspecies of Right whale (North Atlantic) have long been isolated
from its southern counterparts. It was further suggested that the same species of louse
how a focal whale cyamid was able to migrate from the Northern Atlantic to the Southern
Oceans. One possibility included a right whale migrating across the equator thousands of
years ago allowing the lice to move from host to host. The challenges of this theory was
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the governing fact that the equator, being a major barrier to blubber-clad whales, can
Researchers believe that infestation of whale cyamid on host is mediated by: (i) social
behavior of whales; (ii) interactions between whale heterospecifics and conspecifics; and
(iii) delivery of calf. Melon-headed whales are highly social and usually travel in groups
of 500 to 1,000 individuals. Occasionally, groups as large as 2,000 individuals have been
observed (Wardle et al., 2000). Other scientists further believe that slow movements of
whales serve as ideal conditions for lice shifts and relocation. Near shore drifting may
also influence lice migration and subsequent habitation. Scheffer included a notable
example of how mobility and drift location affect commensal transmission. Cyamids are
host specific. Scientists indicate that this was expected considering the circumstances of
their life history. Young whale-lice are released from the female brood pouch as miniature
adults with no free-swimming stage(s). The adults have appendages that are adapted for
swimming. In other amphipods these appendages are modified, often greatly reduced.
The peraeopods or walking legs dig deeper into the whale skin as currents (or collectors)
tend to pull the cyamid body away from the host. The probability of cyamid transmission
from one host to another, apart from other restrictions of specificity, is mediated by
bodily contact. There is contact between adult and calf at birth and in suckling, between
adults during courtship and copulation; and accidental or occasional contact through
closeness in migration, in play and in attack upon one another. Of these, only the latter
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In their search to find out more about local interactions, large-scale movements and
population histories of Right whales, researches have used Right whale cyamids as
Polymorphism within and differentiating among populations; Gene tree estimates; and
In recent years, human exploitation of Right whale populations has reduced whale and
cyamid populations. While researchers have identified some cyamids of Right whales and
developed relationships between the two, no single hypothesis completely explains the
In conclusion, whales and cyamids have a long history of commensalisms and phoresy.
Not only are whale cyamids host specific, but, in the case of Cyamus ovalis and Cyamus
gracilis on right whales, it appears as if lice selectively occupy bodily regions on whales.
The most common methods of lice redistribution on right whales are through: (i) contact
between adult and calf at birth and in suckling; (ii) contact between adults during
courtship and copulation; (iii) sedated whale movements; and (iv) accidental contact
through closeness in migration, in play and in attack upon one another. The redistribution
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References
Bohannan, John. Listening to Lice. Science Now. (September, 2005), p1-2, 2p, 1bw.
http://search.ebscohost.com/login.aspx?direct=true&db=aph&AN=1834035
6&site=ehost-live
Marquardt, W.C.; Demaree, R.S.; and Robert B. Grieve. Mallophaga and Anoplura: The
Lice. Parasitology and Vector Biology. California, Harcourt Academic
Press. (month n/a, 2000). p.549.
Seger J.; Kaliszewska, Z.A.; Rowntree, V.A.; Barco, S.G.; Benegas, R.; Best, P.B.;
Brown, M.W.; Brownell Jr., R.L.; Carribero, A.; Harcourt, R.; Knowlton,
A.R.; Marshalltilas, K.; Patenaude, N.J.; Rivarola, M.; Schaeff, M.C.;
Sironi, M.; Smith, W.A.; and Tadusu K.Yamada. Blackwell Publishing, Ltd.
Population Histories of Right whales (Cetacea: Eubalaena) Inferred from
Mitochondrial sequence diversities and divergences of their whale lice
(Amphipoda: Cyamus). Molecular Ecology (June, 2005) 14, 3439–3456
Wardle, W.J.; Haney, A.T.; and Graham A.J. Worthy. New Host Record for the Whale
Louse (Isocyamus Delphinii, AMPHIPODA, Cyamidae). Crustaceana.
Koninklijke Brill NV, Leiden, (month n/a, 2000) 73 (5): pp.639-641
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