Journal of Invertebrate Pathology 106 (2011) 79–91

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Journal of Invertebrate Pathology

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Diseases of spiny lobsters: A review

J.D. Shields ⇑
Department of Environmental and Aquatic Animal Health, Virginia Institute of Marine Science, The College of William & Mary, Gloucester Point, VA 23062, USA

a r t i c l e i n f o a b s t r a c t

Keywords: Spiny lobsters have few reported pathogens, parasites and symbionts. However, they do have a diverse
Panulirus argus virus 1 fauna comprised of a pathogenic virus, several bacteria, protozoans, helminths and even symbiotic crus-
Palinurid taceans. A few idiopathic syndromes have also been reported, but these appear correlated with lobsters
Jasus
held in poor conditions. Fungal and bacterial pathogens present significant threats for rearing spiny lob-
Panulirus
Symbiont
sters in aquaculture settings, but only one pathogen, Panulirus argus virus 1, is thought to have damaged a
Pathogen fishery for a spiny lobster. No doubt others will emerge as lobsters are brought into aquaculture setting
Fishing and as fishing pressure intensifies with stocks become more susceptible to anthropogenic stressors.
Ó 2010 Elsevier Inc. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
2. Panulirus argus virus 1 (PaV1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
3. White spot syndrome virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
4. Gaffkemia – Aerococcus viridans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
5. Shell disease . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
6. Vibriosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
7. Fouling bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
8. Water molds and Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
9. Oomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
10. Fusarium solani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
11. Peritrich ciliates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
12. Microsporidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
13. Helminths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
14. Digenetic trematode infections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
15. Cestoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
16. Nemertea – Carcinomertes spp. and Pseudocarcinonemertes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
17. Miscellaneous metazoan symbionts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
18. Nicothoidae – parasitic copepods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
19. Amphipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
20. Fouling organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
21. Disease syndromes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
21.1. Turgid lobster syndrome. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
21.2. Pink lobster syndrome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
21.3. Mass mortalities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
22. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

⇑ Fax: +1 804 684 7186.

E-mail address: jeff@vims.edu

0022-2011/$ - see front matter Ó 2010 Elsevier Inc. All rights reserved.
doi:10.1016/j.jip.2010.09.015
80 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
1. Introduction
Spiny lobsters in the genera Jasus, Panulirus, and Palinurus sup-
port worldwide fisheries with annual landings of over 70,000 m
(Booth, 2006; Phillips and Melville-Smith, 2006). There are surpris-
ingly few diseases reported from these lobsters given the size and
value of their fisheries. However, recent attempts to culture lob-
sters have failed because of mortalities associated with potential
agents, most notably bacterial infections (Kittaka, 1997; Bourne
et al., 2004, 2006). That, plus the emergence of a pathogenic virus
in the Caribbean spiny lobster (Shields and Behringer, 2004), high-
light the potential damage that can be caused by unrecognized
pathogens. I review the microbial diseases, parasites, and symbi-
onts of the spiny lobsters, Panulirus spp., Palinurus spp., and Jasus
spp. For earlier reviews of the diseases of spiny lobsters see Evans
and Brock (1994) and Evans et al. (2000). For a broader perspective
on the diseases of clawed and spiny lobsters, see Shields et al.,
(2006). Several overviews and syntheses of crustacean diseases
are also available (Couch, 1983; Johnson, 1983; Overstreet, 1983;
Brock and Lightner, 1990; Sindermann, 1990; Meyers, 1990;
Shields and Overstreet, 2007), as well as those presented in this is-
sue of the Journal of Invertebrate Pathology.
2. Panulirus argus virus 1 (PaV1)
PaV1 is the first naturally occurring pathogenic virus found
infecting a lobster. As its name implies, it infects P. argus. The virus
has a tropism for mesodermal cells, namely certain hemocytes
(hyalinocytes and semi-granulocytes), soft connective tissue cells,
some hematopoietic tissues, and fixed phagocytes (Shields and
Behringer, 2004; Li et al., 2008). PaV1 is currently an unclassified
virus, but it shares several morphological features with the Herpes-
viridae and the Iridoviridae (Shields and Behringer, 2004). For
example, it is an unenveloped, icosahedral, DNA virus with a nucle-
ocapsid approximately 182 nm in size that develops within the nu-
clei of host cells. The Herpesviridae are icosahedral, but they are
enveloped and have a conserved size of approximately 125 lm.
The Iridoviridae are icosahedral and unenveloped, but they are
usually formed within the cytoplasm, whereas PaV1 is formed
within the nucleus.
Lobsters heavily infected with PaV1 are often lethargic or slow.
Their hemolymph does not clot and is milky in color, from cellular
debris and exudates (Shields and Behringer, 2004). PaV1 initially
infects the fixed phagocytes in the hepatopancreas (Li et al.,
2008). These cells line the hepatic arterioles and, as implied by
their name, they phagocytize non-self particles, including virions
of PaV1. The infected fixed phagocytes lyse, and the virus spreads
to spongy connective tissues as well as hemocytes. In late stages
of the infection, virtually all of the spongy connective tissue cells
become infected, with some apparently proliferating around the
hepatopancreas (Fig. 1). The tubules of the hepatopancreas atrophy
in late stages of infection (Li et al., 2008). Reserve inclusion cells in
the hepatopancreas and other tissues lose their glycogen reserves
and the host probably dies from metabolic wasting. Few lobsters
have concurrent bacterial infections (Shields and Behringer, 2004,
Shields, unpubl. data), which is surprising considering that the pri-
mary defensive cells of the host, the hyalinocytes and semi-
granulocytes, are targets for the virus.
Until recently, histology has been the primary tool for diagnos-
ing PaV1 infections in P. argus. Pathological alterations to the fixed
phagocytes and spongy connective tissues, with Cowdry-like inclu-
sions and enlarged nuclei, are the primary pathologies used in
diagnosis. Histopathological changes can be observed in the tissues
within 10–15 days after infection. Using a 110-bp piece of the viral
genome, a DNA probe and dot-blot assay were developed for use in
diagnostics (Li et al., 2006). A fluorescent-in situ-hybridization
(FISH) technique was used in conjunction with the probe to visual-
ize infected tissues. The probe was specific to the virus and sensi-
tive to 10 pg. Infected cells were visualized in many tissues. A
primer set for the polymerase chain reaction (PCR) has also been
developed for diagnosis (Montgomery-Fullerton et al., 2007). The
primer set amplified a 499-bp piece of viral DNA. It was specific
for the virus and sensitive to 1.2 fg. The primer set showed a high
acuity with histological findings. A primary cell-culture method
has been used to quantify the virus in infected hemolymph as a
TCID50 (Li and Shields, 2007). For cultures, naïve hemocytes, partic-
ularly hyalinocytes and semi-granulocytes, were separated using
density centrifugation, then cultured in modified Leibovitz med-
ium, prior to exposure to the virus. Virally infected cells were dem-
onstrated in cultures using an in situ hybridization technique (Li
and Shields, 2007). Uninfected cell cultures retained a high viabil-
ity through 15 or more days.
PaV1 has been shown to induce a remarkable behavior in
healthy, uninfected lobsters. Such lobsters have the ability to de-
tect and avoid diseased lobsters (Behringer et al., 2006). This is
the first example of avoidance of diseased individuals by healthy
animals other than in humans. Moreover, the avoidance behavior
occurs around the time infected lobsters become infectious! This
has important consequences for the denning behavior of P. argus,
because the animals are normally gregarious and the ability to
detect and avoid disease may serve to limit the spread of the
disease in the host population (Behringer et al., 2006). The nat-
ure of the avoidance is likely chemical detection as spiny lob-
sters have an exquisite sense of smell. PaV1 has been
transmitted to uninfected lobsters via inoculation, ingestion, con-
tact and through seawater (Butler et al., 2008). Contact and
water-borne transmission are apparently the primary modes of
transmission in nature. Lobsters inoculated with infected hemo-
lymph can develop acute infections and die within 30–90 days,
but some take much longer to show signs of infection. Lobsters
infected more naturally, via contact or water-borne transmission,
die within 200 days, with mortality highly correlated with smal-
ler sized animals. Infectivity declines significantly with lobster
size. In lobsters exposed via contact, 63% of animals <25 mm car-
apace length (CL) became infected, as opposed to 33% of those
30–40 mm CL, and 11% of the large juveniles (40–50 mm CL),
respectively (Butler et al., 2008). Transmission coefficients used
in mass-balance models were calculated for the different modes
of infection. Contact transmission among the smallest juveniles
had the highest transmission coefficient; that for water-borne
transmission was about five times lower. Thus, the virus appears
to be transmitted by direct contact among small juveniles and
over short distances through the water column.
In short-term mark-recapture studies, overtly diseased lob-
sters had a lower survival rate than animals without disease
(Behringer et al., 2008). In the laboratory, lightly infected lob-
sters showed no significant differences in movement compared
to uninfected animals, but as the disease progressed, infected
lobsters were much less active than healthy animals. The phys-
iological state of intermolt lobsters from the field was assessed
using hemolymph serum proteins as a general indicator of
health. Overtly diseased lobsters had significantly lower hemo-
lymph protein concentrations than presumably uninfected lob-
sters, by as much as 4–5 mg/dl protein levels (Behringer et al.,
2008). Interestingly, starvation did not increase the susceptibility
of animals to viral infection. Given that lightly infected animals
were still highly active, they were thought to facilitate dispersal
of the virus to new habitats.
The discovery of PaV1 coincided with a decline in the landings
of P. argus from the Florida Keys fishery (Shields and Behringer,
2004). Whether the virus caused this decline or not remains to
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91 81

Fig. 1. PaV1 infections in lobsters. (A) Heart of a juvenile lobster infected with PaV1. Note the hypertrophied, infected connective tissue cells lining the arterioles (arrows).
Bar = 100 lm. (B) Hepatopancreas from a heavily infected juvenile lobster. The hemal sinus (star) shows heavy infiltration by infected hemocytes and spongy connective
tissue cells. Bar = 50 lm. (C) Fixed phagocytes (arrow) around an arteriole (lumen marked with star) in the hepatopancreas of an uninfected lobster. Bar = 50 lm. (D) Fixed
phagocytes around an arteriole (lumen marked with star) in the hepatopancreas of an infected lobster. Note the hypertrophied cells with emarginated chromatin in infected
cells (arrows). Bar = 50 lm.

be determined, but there is no doubt that the agent is widespread
and pathogenic to lobsters in their early benthic juvenile stage.
Lobsters infected with PaV1 have been found in several places
around the Caribbean Sea, including the Florida Keys, US Virgin Is-
lands, Mexico, and Belize (Butler et al., 2008; Huchin-Mian et al.,
2008, 2009). PaV1 is widespread in the Florida Keys and Florida
Bay, particularly in the shallow juvenile nurseries. In field studies,
the mean prevalence was 7%, with local ‘‘hot spots” of up to 30%
(Shields and Behringer, 2004). The size predilection for the virus
stands out in the field studies because prevalence was 16% among
the smallest juveniles (<20 mm CL), whereas it was 5% among the
largest juveniles (>40 mm CL); in field surveys of adults, <1% of
adults were visibly infected. Off of Puerto Morelos and the Chin-
chorro Bank, Mexico, prevalence was as high as 10.9% in juvenile
lobsters, but there was no apparent change in host density with in-
creased prevalence of disease. (Lozano-Álvarez et al., 2008). The
differences in the nursery habitat between the Florida Keys and
the area around Puerto Morelos are large enough to warrant fur-
ther attention to density issues and how habitat restrictions poten-
tially alter the host-pathogen association.
PaV1 appears to be specific to P. argus (Butler et al., 2008). In
infection trials, three potential alternate hosts (the spotted lobster,
P. guttatus, the stone crab, Menippe mercenaria, and the channel
crab, Mithrax spinosissimus) were inoculated with hemolymph
from an infected Caribbean spiny lobster. After 80 days, none of
the three alternate hosts showed signs of disease using histology.
This is particularly interesting because the spotted lobster, P. gutt-
atus, is a close congener with P. argus (Ptacek et al., 2001), and its
lack of observable disease suggests that PaV1 is highly specific to P.
argus.
The recent emergence of PaV1 and its potential association with
a decline in the spiny lobster fishery in the Florida Keys is cause for
some concern. Several viruses have severely damaged the shrimp
aquaculture and fishery industries (Flegel, 1997; Lightner and
Redman, 1998); therefore, programs should be developed to mon-
itor for PaV1 in populations of the spiny lobster from around the
Caribbean Sea to establish baselines and to document its potential
effect on artisanal and regional fisheries. Infections have been
identified in two culture facilities in Florida (Shields, unpubl. data),
and we know that it occurs in several countries around the Carib-
bean Sea. Given that lobsters and lobster tails are shipped interna-
tionally (see Huchin-Mian et al., 2009), plus the fact that nascent
aquaculture ventures have been attempting to culture spiny lob-
sters from enzootic areas, lobsters should be screened for the virus
as well as other microbial infections to control for infections as
well as to prevent their potential spread to other areas (see
Table 1).
3. White spot syndrome virus
Spiny lobsters are not naturally infected with the white spot
syndrome virus (WSSV), but with epidemics of WSSV occurring in
native shrimp stocks, it seems inevitable that it will spread into
new host species. Indeed, at least 42 crustaceans, including three
species of Panulirus, can serve as experimental reservoir hosts for
WSSV (see Supamattaya et al., 1998; Rajendran et al., 1999; Mus-
thaq et al., 2006). In experimental infections, a DNA probe was used
to detect WSSV in the tissues of P. versicolour and P. penicillatus
(Chang et al., 1998). Lobsters survived for at least 70 days, with
the virus present in their tissues, but they were not examined his-
tologically for signs of disease. That is, the lobsters were infected
but were not obviously diseased. PCR has also been used with a
virus-specific primer set to detect WSSV in several species of lobster
(P. versicolour, P. penicillatus, P. ornatus and P. longipes) fed infected
82 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91

Table 1
Parasites and symbionts naturally occurring in spiny lobsters, including the tissues in which they are found.

Parasite Host Tissue Key reference

Virus
PaV1 P. argus Connective tissues, Shields and Behringer (2004)
hemocytes
Bacteria
Aerococcus viridans H. americanus, H. gammarus, P. Systemic Snieszko and Taylor (1947), Rabin (1965),
argus Stewart et al. (1966), Stewart et al. (1980), Bobes et al. (1988)
Several genera P. argus, P. ornatus External Porter et al. (2001), Bourne et al. (2006), Payne et al. (2007)
Vibrio harveyi J. edwardsi Systemic, phyllosoma Diggles et al. (2000)
V. alginolyticus Panulirus homarus Systemic Hameed (1994), Abraham et al. (1996)
Vibrio spp. P. argus, P. laevicauda Systemic Silva dos Fernandes Vieira et al. (1987)
Leucothrix mucor Several genera Carapace, eggs Nilson et al. (1975), Steenbergen and Schapiro (1976), Harper and Talbot
(1984)
Fungia
Atkinsiella panulirata P. japonicus Larva Kitancharoen and Hatai (1995)
Haliphthoros sp. P. argus,J. verreauxi Pueruli, small juveniles Fisher et al. (1975), Diggles (2001)
Fusarium solani P. cygnus Cuticle McAleer and Baxter (1983)
Didymaria palinuri P. elephas Gills Sordi (1958)
Ramularia branchiales
Protozoans
Microsporidia P. argus
Panulirus spp. Muscle? Muscle Bach and Beardsley (1976), Dennis and Munday (1994), Kiryu et al.
(2009)
Peritrich ciliates Jasus edwardsii Kittaka (1997), Handlinger et al. (1999), Bourne et al. (2004)
Helminths
Turbellaria Panulirus spp. On mouthparts Shields (unpubl. data)
Thulakiotrema genitale P. cygnus Gonad Deblock et al. (1991)
Cymatocarpus solearis P. argus Abdominal muscle Gómez del Prado-Rosas et al. (2003)
Tetraphyllid cestode Panulirus spp. Foregut Shields (unpubl. data)
Carcinonemertes P. interruptus Eggs Shields and Kuris (1990)
wickhami
Carcinonemertes P. cygnus Eggs Campbell et al., (1989)
australiensis
Carcinonemertes sp. J. edwardsii Eggs Frusher and Shields (unpubl. data)
Crustaceans
Choniomyzon panuliri Panulirus spp. Eggs Pillai (1962)
Paramphiascopsis sp. J. edwardsii, J. verreauxi Gills Booth (pers. comm.)
Parapleustes commensalis P. interruptus Eggs Shoemaker (1952)
Gitanopsis iseebi P. japonicus Branchial chamber Yamato (1993)
Octolasmis californiana P. interruptus Gills, carapace Newman (1960)
Octolasmis spp. P. polyphagus Gills, carapace Jeffries et al. (1982)
Trilasmis fissum P. japonicus, P. penicillatus Mouthparts Bowers (1968)
Balanomorphs P. argus Carapace Eldred (1962)
Paralepas spp. Spiny lobsters Carapace Jones (2003)
a
The Phylum Fungi no longer includes the fungus-like Phycometes and other classical forms that are now placed in a separate Kingdom, the Chromista.

shrimp (Wang et al., 1998). The virus was detectable in all of the
lobsters, but none developed disease. Lobsters (Panulirus homarus)
inoculated with WSSV-infected shrimp tissues died, presumably
from the infection (Rajendran et al., 1999; Musthaq et al., 2006),
but the question is unresolved because animals fed infected tissues
did not develop infections (Musthaq et al., 2006), and controls con-
sisting of uninfected tissue homogenates were not used. However,
in the latter study, all of the lobsters injected with the virus died
over 160 days.
It is surprising that no other viruses are known from lobsters.
Shrimp have at least 20 and the blue crab, Callinectes sapidus, has
at least eight pathogenic viruses (Lightner and Redman, 1998;
Shields and Overstreet, 2007). Several shrimp viruses have spread
as pandemics through the Americas and Asia with catastrophic re-
sults for shrimp aquaculture (Flegel, 1997; Lightner and Redman,
1998). Perhaps more viruses are known from shrimp because they
are intensively cultured; hence, there are more opportunities for
outbreaks under crowded conditions. Nonetheless, clawed and
spiny lobsters support large fishing industries in which diseased
animals would be observed or recognized. Perhaps the paucity of
viruses in lobsters is simply a reflection of the fact they are only
now being investigated for intensive aquaculture, and that most
of the focus on lobsters has been on their fisheries and not on
the more susceptible larval and juvenile stages.
4. Gaffkemia – Aerococcus viridans
Gaffkemia, or red-tail disease, is a serious disease of clawed lob-
sters, primarily Homarus americanus, but also H. gammarus. Out-
breaks usually occur in holding facilities; it is not common in
natural populations of lobsters (Stewart et al., 1966; Keith et al.,
1992; Lavallée et al., 2001). Aerococcus viridans, a Gram-positive
bacterium, is the causative agent of gaffkemia, and it is a common
soil microbe with many variants. The variant that is pathogenic to
lobsters is known as A. viridans var. homari. The pathogen can be
spread via shipment and holding of diseased clawed lobsters;
therefore, further risk assessment should be initiated in areas
where there could be likely introductions of contaminated water
or discarded lobsters.
Aerococcus viridans reportedly occurs naturally in P. argus from
Cuba. Bobes et al. (1988) isolated and cultured A. viridans from six
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
animals that presented signs of disease. The animals had reddish
discolored exoskeletons, lethargy, pink hemolymph and coagulop-
athies. The bacterium was susceptible to 10 of 13 antibiotics in cul-
ture (Bobes et al., 1988). While Koch’s postulates were not
attempted, this finding has further implications for the transporta-
tion of spiny lobsters. Aerococcus viridans var. homari has been iso-
lated from other crustaceans from Massachusetts (Rabin and
Hughes, 1968) and the Gulf of Mexico (Liuzzo et al., 1965). While
other crustaceans can be carry the bacterium, clawed lobsters are
apparently the only hosts in which it is highly pathogenic (Cornick
and Stewart, 1975).
The route of transmission of Aerococcus viridans is through dam-
age to the host exoskeleton, not through intact cuticle or ingestion
(Stewart et al., 1969). The bacterium is not part of the natural
microflora of lobsters (Stewart, 1975). It has been isolated from
lobster holding facilities; and the isolation of the agent usually
indicates the presence of diseased lobsters (Goggins and Hurst,
1960; Kellog et al., 1974; Stewart, 1980). Unfortunately, it can re-
main viable in the benthos for long periods where it can act as a
reservoir of infection (Stewart, 1984). Aerococcus viridans has also
been documented in muds adjacent to a holding facility for clawed
lobsters in Anaheim Bay, California, and was likely transported
there through diseased clawed lobsters (Kellog et al., 1974).
Gaffkemia can be experimentally induced in P. interruptus, but
the lethal dose is higher than that for H. americanus (Schapiro
et al., 1974; Steenbergen and Schapiro, 1974). That is, spiny lob-
sters show a high degree of resistance to experimental infections
(Schapiro et al., 1974; Schapiro and Steenbergen, 1974). Because
most studies of gaffkemia have been carried out in the clawed lob-
sters, there will be no further treatment of it here. For reviews of
gaffkemia, see Stewart (1980) and Brock and Lightner (1990).
5. Shell disease
Shell disease, or shell disease syndrome, is a progressive chitin-
olysis and necrosis (bioerosion) of the exoskeleton of crustaceans
(Rosen, 1970). Virtually all crustaceans are susceptible to shell dis-
ease, but it is most notable in large decapods. The syndrome first
appears as small pits or erosions in the cuticle and progresses to
large eroded lesions (Rosen, 1970; Getchell, 1989; Noga et al.,
1994). The bacterial form, which is referred to as classical, or enzo-
otic, shell disease, was first described from the American lobster
(Hess, 1937). Recently, an epizootic form of the syndrome has
emerged in American lobsters from Rhode Island (see Castro
et al. (2006), for review). This new form of the disease has not been
reported from palinurid lobsters. For comprehensive reviews of
shell disease in crabs and lobsters see Shields et al. (2006) and
Shields and Overstreet (2007).
Classical shell disease is caused by a number of chitinoclasitic,
Gram-negative bacteria. Vibrios (Vibrio vulnificus, V. parahaemoly-
ticus, V. alginolyticus) are the most common bacteria associated
with shell disease, but other species have also been isolated from
the lesions, including Shewanella spp. and Aeromonas hydrophila
(Geddes et al., 2003; Porter et al., 2001; Reuter et al., 1999). Both
V. alginolyticus and a V. harveyi-like bacterium were isolated from
the hemolymph and lesions occurring on P. homarus reared in
the laboratory (Abraham et al., 1996). Using the 16s and 23s rDNA
intergene regions, Porter et al., (2001) examined the bacterial fau-
na of lobsters with and without classical shell disease. DNA finger-
printing found no specific etiological agent associated with the
lesions, but bacteria in several genera were identified, including
Vibrio, Pseudoaltermomonas, Pseudomonas and Shewanella. The na-
tive bacterial flora was thought to be responsible for shell disease
lesions in spiny lobsters (Porter et al., 2001), and many of these are
indeed chitinoclastic.
83
Classical shell disease requires a portal of entry through the epi-
cuticle, and mechanical abrasion is the usual mode of transmission.
Shell disease starts as small pits or ‘‘burn” marks in the cuticle. The
lesions occur on the sternum or legs of American lobsters (Rosen,
1967; Johnson, 1983), but they are more commonly found on the
uropods of spiny lobsters (Porter et al., 2001; Geddes et al.,
2003). As the pits coalesce, they develop into large, brown or black,
irregularly shaped lesions (Rosen, 1967). The lesions can penetrate
through the cuticle into underlying tissues of the lobster, but they
rarely do so; instead they develop laterally with further erosion of
the cuticle (Overstreet, 1978; Johnson, 1983). The exposed cuticle
is weakened and brittle, and discolored from the deposition of mel-
anin (Johnson, 1983; Smolowitz et al., 1992). Melanization and cel-
lular infiltration occur as part of the host response to shell disease
in American lobsters (Smolowitz et al., 1992, Smolowitz et al.,
2005). Pseudomembrane formation can occur in particularly se-
vere cases of shell disease, particularly in epizootic shell disease
(Smolowitz et al., 2005). It is no doubt the same process in spiny
lobsters. Individuals usually overcome the disease by molting
(Rosen, 1967; Castro and Angell, 2000), but animals that molt less
frequently can presumably succumb to infection.
In spiny lobsters, P. argus, Panulirus cygnus and Jasus edwardsii,
another form of shell disease occurs. It is known as ‘‘tail fan necro-
sis”, an aptly named syndrome because the uropod and telson ap-
pear melanized and partly or fully eroded away (Porter et al., 2001;
Geddes et al., 2003). In clawed lobsters, shell disease lesions are
more frequently seen on the ventral side of the claws, tail and car-
apace (Estrella, 1991). Shell disease is normally found at low (<1%)
levels in healthy wild populations of crustaceans. Iversen and
Beardsley (1976) examined 9000 P. guttatus, and found only two
infected individuals. They cite several unpublished reports indicat-
ing a very low prevalence of shell disease in P. argus.
Classical shell disease is often associated with pollution
(Gopalan and Young, 1975; Young and Pearce, 1975; Couch,
1983; Morado et al., 1988; Gemperline et al., 1992; Weinstein
et al., 1992; Ziskowski et al., 1996), and spiny lobsters require rel-
atively pristine water quality parameters; therefore, shell disease
is not usually found at a high prevalence in palinurid lobsters.
However, clawed lobsters held in crowded pounds with poor water
quality can develop fatal shell disease (Taylor, 1948, in Sinder-
mann, 1989). Therefore, in culture systems, efforts should be direc-
ted at maintaining good water quality and reducing crowding
effects. Lobsters with moderate to severe forms of shell disease
are not esthetically pleasing and are not used in the ‘‘live food”
trade. Indeed, epizootic shell disease has had an economic impact
on the clawed lobster fishery off Rhode Island, because the cuticle
of affected animals is severely compromised making the animals
unsuitable for live trade (Castro et al., 2006).
Antibiotics have been used to treat shrimp and lobsters with
shell disease, but none are currently registered for use in crusta-
cean aquaculture in the USA. Some such as furanace may be used
in other countries. Juvenile lobsters have been successfully treated
with Malachite green (Fisher et al., 1976a,b, 1978). Shrimp have
been treated with Malachite green, formalin, or antibiotic baths
(penicillin–streptomycin, furanace, erythromycin, oxolinic acid)
(Tareen, 1982; Brock, 1983; El-Gamal et al., 1986). Contaminated
aquaria should be disinfected with bleach solutions, and it is a
good practice to do this prior to any laboratory culture. Animals
with severe lesions should be destroyed to prevent further spread
in impoundments.
6. Vibriosis
Vibriosis is an infection by any species of Vibrio. Vibrios are
ubiquitous in the marine environment, and several species cause
84 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
serious diseases to invertebrates, fishes and even humans. Lob-
sters, in turn, can be exposed to several pathogenic species of
Vibrio. Vibrio alginolyticus, V. harveyi, V. parahaemolyticus and
V. anguillarum are facultative, opportunistic pathogens that have
been implicated in disease in clawed and spiny lobsters (Bowser
et al., 1981; Brinkley et al., 1976; Jawahar et al., 1996). Vibrios
can also contaminate seafood products including lobsters (Wong
et al., 1999); and there are many cases of human food poisoning
from improper handling and preparation of marine crustaceans.
Lobsters should be properly cooked prior to eating or freezing,
and all surfaces used in the preparation of lobsters should be
cleaned and disinfected afterwards.
Vibrio infections can cause serious disease in spiny lobsters, but
few outbreaks have been reported. This is probably because spiny
lobsters are usually kept in short-term culture, but not long-term
culture where there would be more opportunity for disease under
crowded conditions. However, outbreaks are known to occur in lar-
val cultures. Indeed, vibriosis may be the single most important
impediment to successful larval culture (Bourne et al., 2004). Cul-
tures of phyllosoma larvae of Sagmariasus(=Jasus) verreauxi experi-
enced significant mortalities due to a luminescent strain of V.
harveyi in New Zealand (Diggles et al., 2000). Infected larvae were
opaque and had small red spots throughout the body. The hepato-
pancreas was atrophied and the hepatopancreatic tubules had
numerous bacterial plaques. Mortalities occurred over 4 weeks.
The bacterium is an opportunistic pathogen and attacked injured
larvae more than uninjured ones. Two drugs, sulfadimidine and tri-
methoprim, were found to enhance survival the lobster larvae
(Diggles et al., 2000).
The dynamics of bacterial flora have been studied in cultures of
larval spiny lobsters. Larval cultures of Panulirus ornatus had a high
microbial diversity, with representatives from many bacterial taxa
(Bourne et al., 2006; Payne et al., 2007). Denaturing gradient gel
electrophoresis (DGGE) was used to study diversity in larval cul-
tures (Bourne et al., 2004; Payne et al., 2007) as well as biofilms
in culture aquaria (Bourne et al., 2006). There were few changes
in the bacterial flora in relation to mortality. In fact, bacterial den-
sities decreased significantly over time and had no relationship
with mortality (Bourne et al., 2004). However, larval mortality
was high, reaching 100% in 13–24 days. Vibrios were visualized
and enumerated using a fluorescence in situ hybridization (FISH)
technique (Webster et al., 2006). Vibrios increased both on and
within phyllosoma larvae starting about seven to ten days after
hatching, and reached high levels 18 days after hatching. Two iso-
lates of V. harveyi were used in a challenge study and one strain
caused more mortality than the controls (Bourne et al., 2006). Sev-
eral species of Vibrio and other bacteria have been identified from
cultures of the phyllosoma of J. edwardsii (Handlinger et al., 1999).
An isolate of V. harveyi was obtained from the diseased gut of an
infected larva. More research is needed in this area because it is
not clear how larval nutrition, host defenses, temperature, culture
stress and disease agents interact in culture systems.
Vibrio alginolyticus, V. parahemolyticus and a V. anguillarum-
like species were isolated from P. argus and P. laevicauda from
commercial landings in Brazil. Prevalence varied from 2.8% in V.
anguillarum to as high as 45.7% in P. alginolyticus (Silva dos Fernan-
des Vieira et al., 1987). Unfortunately, the disease status of the lob-
sters was not reported. Vibrio alginolyticus and a V. harveyi-like
bacterium were isolated from shell disease lesions and hemolymph
of juvenile and adult P. homarus from India (Hameed, 1994). In
inoculation trials, the Vibrios became systemic and induced shell
disease-like lesions in larvae of Penaeus indicus and P. homarus
(Hameed, 1994). Several tissues showed histopathological altera-
tions. In a separate study by Abraham et al., (1996), mortality
reached 100% in P. homarus injected with 108 bacterial cells, but
there was no mortality in lobsters inoculated with 107 bacterial
cells. I have obtained similar results in inoculation trials with Vib-
rios in J. edwardsi (Shields, unpubl. data).
Vibrios are frequently isolated from the hemolymph of crusta-
ceans; and the fact is that crustacean hemolymph is not necessarily
sterile. In a debate in the 1970s, Bang (1970), Colwell et al., (1975)
and Tubiash et al., (1975) found a high prevalence of bacteria, par-
ticularly Vibrios in the hemolymph of apparently healthy blue
crabs. These were thought to have been acquired through abra-
sions or other wounds in the cuticle that occurred because of the
rough handling of crabs en route to markets (Johnson, 1976). In-
deed, crabs are handled roughly during capture and transport to
processing plants (Shields, pers. obs.). However, in a definitive
study, species of Vibrio were shown to occur in the hemolymph
and various other tissues of freshly caught, unstressed crabs (Davis
and Sizemore, 1982). Nonetheless, we rarely isolate bacteria from
the hemolymph of spiny lobsters. Thus, while it is possible that
spiny lobsters can carry background levels of septicemia, there
are few reports of such in adults, nor have I observed many bacte-
rial isolates in routine health examinations of P. argus and J. edwar-
dsii (Shields, unpubl. data).
7. Fouling bacteria
Filamentous bacteria are ubiquitous on the eggs of marine crus-
taceans, and spiny lobsters are no exception (Johnson et al., 1971;
Bland and Brock, 1973). Leucothrix mucor is a common filamentous
species that has been observed on or isolated from cultured larvae
of the American lobster (Nilson et al., 1975; Dale and Blom, 1988).
It and other fouling agents are common fouling organisms in larval
culture of spiny lobsters (e.g., Kittaka, 1997; Bourne et al., 2004,
2006; Payne et al., 2007). Leucothrix mucor is very common in sea-
water systems. It is a saprophyte of dead algae. I have observed it
on crustacean eggs and limbs, including amphipods, crabs and lob-
sters. Filamentous bacteria can be difficult to control because they
are so common and because they grow readily in the organically
enriched environments encountered in culture systems. Heavy
infections are thought to contribute to larval mortality. Control re-
quires careful site selection and system design, good water quality
parameters and strict attention to nutrition. Outbreaks of L. mucor
have been controlled with antibiotics (Steenbergen and Schapiro,
1976; Fisher et al., 1976a,b; Sadusky and Bullis, 1994).
8. Water molds and Fungi
The ‘‘lower fungi” have been reclassified as protists in the King-
dom Chromista (Cavalier-Smith, 1993). The Oomycota include the
Oomycetes and Phycomycetes after Margulis et al., (1990). They
are now referred to as water molds. The true Fungi include Deuter-
omycetes, Ascomycetes and Basidomycetes and a few other classes
of ‘‘higher” fungi.
9. Oomycetes
A few oomycetes are known from spiny lobsters. These typically
infect the eggs or larvae, but a few infect adult lobsters (e.g.,
Alderman, 1973). Atkinsiella panulirata is an oomycete described
from phyllosoma of P. japonicus (Kitancharoen and Hatai, 1995).
Haliphthoros mildfordensis has been implicated in mortalities in
aquaculture facilities for postlarval spiny and clawed lobsters (Fish-
er et al., 1975, 1978; Nilson et al., 1975 Diggles, 2001). An outbreak of
Haliphthoros sp. caused mortalities in a grow-out facility in New Zea-
land (Diggles, 2001). The disease infected the pueruli and juveniles
of J. edwardsii, which exhibited morbidity, including lethargy and
loss of appetite. Only lobsters less than 30 mm CL became infected.
Perhaps this was due to larger animals having a thicker or more resil-
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
ient cuticle. Antibiotic treatment using Malachite green and forma-
lin was successful in preventing the spread of disease. The outbreak
was associated with poor water quality (Diggles, 2001).
Melanization of infected tissues is a common defense against
Haliphthoros and other mold and fungal infections in crustaceans.
Therefore, black or brown spots are commonly seen in necropsies
involving mold infections. The quinones and melanin, which are
activated by the prophenoloxidase pathway, cause these black spots.
Prophenoloxidase occurs naturally in the cuticle and its activated
enzyme, phenoloxidase, is activated through a well known cellular
process involving hemocytes. Melanin is highly toxic to many micro-
bial pathogens (for review, see Jiravanichpaisal et al. (2006)).
Maintaining animals in conditions of excellent water quality is
the preferred method to control water mold outbreaks in cultures
of lobsters. Ozonated or dechlorinated seawater should be used in
cultures. Water changes to maintain low organic loads and low
ammonium levels is also critical. Furanace and other antimold
drugs, such as Malachite green, can be used to treat infected ani-
mals (Abrahams and Brown, 1977; Fisher et al., 1978), but these
drugs are not approved for use in aquaculture by the Food and
Drug Administration in the USA and in several other countries.
10. Fusarium solani
A deuteromycete fungus, F. solani, causes disease in cultured
crustaceans. This species and other unidentified species have
caused disease in shrimp (Lightner and Fontaine, 1975), spiny lob-
sters (McAleer and Baxter, 1983) and clawed lobsters (Alderman,
1981). Black lesions resembling shell disease occur on the cuticle
of infected hosts. An outbreak of F. solani was reported from P. cyg-
nus off Western Australia (McAleer and Baxter, 1983). The lesions
occurred on the abdomen, uropods, telson and pereopods. The fun-
gus was isolated from lobsters and from seawater samples in areas
with diseased lobsters. Environmental factors were thought to
have facilitated the outbreak, but they were not identified. Given
the location of the lesions around the uropods and telson, water
quality was likely an issue.
The pathology of Fusarium infections has been studied in
shrimp and clawed lobsters. The fungus penetrates through the
cuticle, which causes the formation of lesions (Lightner and Fon-
taine, 1975; Fisher et al., 1978). The lesions often become melan-
ized through the phenoloxidase pathway. Lesions containing
cellular aggregates, necrotic tissues, and fungal hyphae can be ob-
served in histology. The hyphae of Fusarium produce diagnostic mi-
cro- and macroconidia (Lightner and Fontaine, 1975).
Two other deuteromycetes, Didymaria palinuri and F. (Ramular-
ia) brachialis were isolated from moribund P. elephas and H. gamm-
arus from the Mediterranean Sea (Sordi, 1958), but no attempts
were made to infect healthy lobsters. It is not clear if the fungi
were primary pathogens or opportunists. Little data were reported
from these infected animals.
11. Peritrich ciliates
Peritrich and suctorian ciliates are often found as commensals
on the external surfaces and embryos of crustaceans. Heavy infes-
tations often indicate poor water quality or the presence of a dis-
ease. They are not usually pathogenic but heavy infections can
impose a respiratory debt to the host (Schuwerack et al., 2001).
They can be quite abundant in larval cultures of spiny lobsters
(e.g., Handlinger et al., 1999; Bourne et al., 2004). Their presence
on crustacean embryos generally indicates that a female host is
not feeding well or is not preening or otherwise taking care of her-
self. Several studies have found peritrichs in the genera Vorticella
and Zoothamnium, suctorian ciliates Acineta and Ephelota, several
85
cyanobacteria and diatoms in the clutches of clawed lobsters
(Dannevig, 1928, 1937; Nilson et al., 1975; Harper and Talbot,
1984; Dale and Blom, 1988).
Vorticella sp., Navicula sp., L. mucor and other organisms fouled the
larvae of J. edwardsii (Kittaka, 1997). Fouled phyllosoma larvae died,
but their cause of death was not documented. Interestingly, other foul-
ing agents, including Saprolegnia sp., hindered fouling by the ciliates.
Chilodonella-like protists and Epistylis spp. have also been reported as
fouling agents (Handlinger et al., 1999; Bourne et al., 2004). Wescodyne
and Malachite green have been used to control Vorticella infestations on
clawed lobsters (Boghen, 1982). Baths of formalin, oxytetracycline,
erythromycin and streptomycin were used to control fouling organ-
isms on P. ornatus (Bourne et al., 2004). Weak formalin solutions have
been used to effect pathogen control in many culture situations, includ-
ing ciliate infections in fishes.
12. Microsporidia
The Microsporidia is a phylum of small, intracellular and intra-
nuclear parasites. The phylum is named after the small, highly
refractory spore produced as the transmissive stage. All members
of the phylum have an extrusible polar tube that is used to inject
the sporoplasm into a new host cell. The Microsporidia were once
considered a phylum within the Protozoa, but they are now consid-
ered to be closely related to the true Fungi (for review, see Lee et al.
(2008)). Some species, such as Nosema apis, can survive in their
cadavers for very long periods.
Microsporidia are very rare in lobsters. A single infection was
reported from P. argus from Florida (Bach and Beardsley, 1976),
but no details were given. More recently, Kiryu et al., (2009) found
two lobsters with Ameson-like microsporidiosis from Florida. The
microsporidians are aptly named as those in the lobster were
1.4  1.0 lm in size (Kiryu et al., 2009). In Australia, at least one
species of Ameson sp. is known to infect spiny lobsters, P. cygnus
and P. ornatus (Dennis and Munday, 1994; Owens and Glazebrook,
1988). In both systems, infections occur in the muscles. They cause
a distinctive liquefactive and coagulative necrosis of the muscles,
with heavy infections causing the muscle fibers to turn creamy
white in color. A similar condition in crabs and shrimp is known
as ‘‘cotton” disease. The flesh of these animals when cooked has
a consistency like that of cotton; and it is very unappetizing
(Shields, pers. obs.). Very little is known about infections in lob-
sters. In blue crabs, Ameson michaelis infections are transmitted
via cannibalism (Overstreet, 1978), but their rarity in lobsters sug-
gests that other pathways are involved or that lobsters may be
accidental hosts. Microsporidia are known to damage prawn aqua-
culture (Flegel et al., 1992; Hudson et al., 2001) so there is a prec-
edent for their potential infection of lobster populations. However,
in field surveys from the Florida Keys that involved several hun-
dred animals, there was no obvious histological infection by any
microsporidian (Shields, unpubl. data). Given the live market for
lobsters and their shipment to international markets, the potential
transport of microspores in infected hosts should be investigated
as a risk factor, albeit many microsporidians show high host spec-
ificity, which could lower the risk to other hosts.
13. Helminths
Few platyhelminthes reportedly use lobsters as hosts. There are
no published reports of turbellarians on spiny lobsters, but I have
observed them on the mouthparts and gills of several species of
Panulirus, from the Great Barrier Reef (Shields, unpub. data). Tur-
bellaria have been reported from the egg clutches of crabs (Kuris
et al., 1991), and it would not be surprising to find them in the
clutches of other crustaceans, including lobsters. There are a few
86 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
reports of digenetic trematodes from palinurid lobsters, but there
are no published reports of cestodes from them other than my
anecdotal observations below.
14. Digenetic trematode infections
Lobsters serve as the second intermediate host to digenetic
trematodes. They bear the metacercaria, or the encysted stage.
The metacercarial cysts are usually quite small (<1 mm) and can
be detected using microscopy. As the name implies, the microphal-
lid trematodes are known for their small phallus. Several species
use crustaceans as intermediate hosts. While the metacercariae
of most trematodes are difficult to identify to the species level,
the microphallids develop sexual characteristics as metacercariae
making it possible to identify and describe them. Thulakiotrema
genitale is a microphallid found encysted as metacercaraie in the
gonads of Panulirus cygnus (Deblock et al., 1991). The prevalence
in Western Australia was quite high, ranging from 47% to 87%.
The vertebrate definitive host is unknown, but it is likely to be a
fish. Cymatocarpus solearis(=C. undulatus) is a brachycoeliid trema-
tode. Metacercariae infect the abdominal muscles of P. argus
(Gómez del Prado-Rosas et al., 2003). The cysts of C. solearis are
large enough to see by eye (1.5 mm in size). The parasite was rel-
atively abundant in lobsters, with a prevalence of 35.8% and a
mean intensity of 26 cysts per host. The loggerhead turtle Caretta
caretta serves as the definitive host, where it lives in the intestines
(Linton, 1910; Caballero, 1959). Light infections of metacercariae
probably do not result in significant pathology, albeit in other sys-
tems, particularly those involving the nervous systems of their
hosts, they can cause alterations to host behaviors.
15. Cestoda
Lobsters may serve as an intermediate host for cestodes, bear-
ing the metacestode, which is usually encysted in the connective
tissues. Metacestodes vary in size, some are microscopic whereas
others can be quite large. I have observed metacestodes of a tetra-
phyllidean cestode in the foreguts of Panulirus spp. and Scyllarides
sp. from the Great Barrier Reef (Shields, unpub. data). The tetra-
phyllidean cestodes use elasmobranchs as definite hosts. Tetra-
phyllidean metacestodes cannot be identified to species unless
fed to and recovered from an elasmobranch host. This can be diffi-
cult to do in the laboratory, but it is possible (see Sakanari and
Moser, 1989). There are no published records of metacestodes from
lobsters. This is intriguing because they no doubt serve as interme-
diate hosts for other parasites that use elasmobranchs, other fishes,
and turtles as definitive hosts.
16. Nemertea – Carcinomertes spp. and Pseudocarcinonemertes
Two species of Carcinonemertes are known from spiny lobsters.
They are egg predators and, at least on crabs, they can occur in epi-
demics where they damage reproduction on a population level
(Wickham, 1979, 1986; Hobbs and Botsford, 1989; Shields and
Kuris, 1988; Kuris et al., 1991). Carcinonemertes can be difficult to
detect on lobsters because they only infest the egg masses and only
during reproduction; that is, one must assess their presence when
lobsters carry eggs.
Panulirus interruptus from Southern California are infested by
Carcinonemertes wickhami (Shields and Kuris, 1990). It is a rela-
tively large carcinonemertid worm, 30–50 mm in length. Panulirus
cygnus from Western Australia are infested by C. australiensis
(Campbell et al., 1989). It is a relatively small worm for the genus
at 7 mm in length. Jasus edwardsii from Tasmania is also infested by
a carcinonemertid worm, but it has yet to be identified (Frusher
and Shields, unpubl. data). It is a larger, more robust species than
C. australiensis.
Both C. wickhami and C. australiensis are monostyliferous hoplo-
nemerteans; that is, they each bear a single stylet on the proboscis
armature. They are dioecious, and use internal fertilization for
reproduction. Other members of the Carcinonemertidae can be
hermaphroditic (Roe, 1986; Shields, 2001). The eggs are extruded
into a mucilaginous egg strand, which adheres to a seta on the ple-
opod within the clutch. The egg strands can also be used to indicate
the presence of the worms. The biology and ecology of these nem-
erteans is unknown. Comparative studies on the life history of
these species would be very interesting.
Clawed lobsters bear a nemertean that is unlike the carcinon-
emertids. It is known as Pseudocarcinonemertes homari and it prob-
ably belongs in the family Tetrastemmidae. Charmantier et al.,
(1991) used freshwater baths to rid clawed lobsters of infestations.
Freshwater killed worms after only 4 min. Freshwater baths and
Malachite green have been used to control infestations of C. errans
on Cancer magister (Wickham, 1988). The fact that carcinonemert-
ids can occur in epidemics on crab species indicates that these
worms have the potential to occur in outbreaks on lobsters. Care
should be taken in transporting ovigerous lobsters to new locations
to avoid the introduction of their worms and other parasites.
Broodstock should also be examined prior to establishing colonies
of lobsters.
17. Miscellaneous metazoan symbionts
As yet, spiny lobsters are not known to be infested with any
member of the Cycliophora. Symbion pandora and S. americanus
are the only described members of the phylum Cycliophora (Obst
et al., 2006). The phylum is highly specialized with both S. pandora
and S. americanus living on the mouthparts of the Norway lobster
and American lobster, respectively (Funch and Kristensen, 1995;
Obst et al., 2006). However, I have observed rotifers and rotifer-like
animals on the mouthparts and gills of spiny lobsters from the
Great Barrier Reef, and given what is now known about the Cyclio-
phora, these organisms may be of further scientific interest.
18. Nicothoidae – parasitic copepods
Nicothoid copepods are a family of highly specialized symbionts
that live on other crustaceans. Nicothoidae were once in the family
Choniostomatidae (see Boxshall and Lincoln, 1983). The copepods
live in the egg clutches or marsupia of their hosts. They are egg
predators and in some cases they are very similar to host eggs;
in fact, they are considered egg mimics (Hansen, 1897; Bowman
and Kornicker, 1967). Choniomyzon panuliri is a nicothoid copepod
that infests the clutches of Panulirus spp. (Pillai, 1962). Several spe-
cies of Panulirus from the Great Barrier Reef, Australia, have a sim-
ilar species in their egg clutches (Shields, unpubl. data). The larval
stages of species of Choniosphaera occur in the gill chambers of
non-ovigerous crab hosts, but this has not been examined for
Choniomyzon. The feeding habits of at least one species of Nicothoi-
dae can cause significant damage to the fecundity of individual
hosts (Shields and Wood, 1993).
Two other copepods have been reported from spiny lobsters.
The harpacticoid copepod Paramphiascopsis sp. is apparently sym-
biotic in the gills of male and female J. edwardsi and S. verreauxi
(Booth, personal communication). Members of this genus appar-
ently occur in symbiotic relationships with other invertebrates
(Soyer, 1973; Hicks, 1986). Another harpacticoid copepod occurs
in the gills of Panulirus spp. from the Great Barrier Reef, Australia.
It is similar to Sunaristes (Shields, personal observation).
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
19. Amphipods
Spiny lobsters often have amphipods living in close association
with them, either on their egg clutches or within their branchial
chambers. The relationship is likely commensal, but those found
on the egg clutches are probably predatory on the eggs of their
hosts. Parapleustes commensalis occurs on the pleopods and egg
clutches of P. interruptus and Cancer anthonyi from Southern
California (Shoemaker, 1952; Shields pers. observation) and Para-
lithodes californiensis (Wicksten, 1982). It ate crab eggs in experi-
mental systems (Shields, unpubl. data). Ischyroceros sp. infests
the egg clutches of Paralithodes camtschaticus (Kuris et al., 1991).
It was correlated with significant egg mortality on that host; hence,
the potential of amphipods as egg predators on lobsters. Gitanopsis
iseebi has been reported from the branchial chamber of Panulirus
japonicus (Yamato, 1993). Other members in the family of amphi-
olochid amphids are commensals; therefore, it is likely to be a
commensal. Isaea elmhirsti lives as a commensal on the mouthparts
of H. gammarus (McGrath, 1981, in Costello et al. (1990)). Amphi-
pods are highly motile and they can swim away from a host when
it is captured; therefore, some of these relationships may be hard
to study. Symbiotic amphipods warrant attention, particularly in
studies of fecundity or reproductive behaviors or when establish-
ing broodstock for nascent culture studies.
20. Fouling organisms
A plethora of organisms can foul the exoskeleton of lobsters and
other crustaceans. Some are opportunists that simply require a
hard surface. Others are obligate symbionts. Some are also useful
as indicators of the health of their host. The gooseneck barnacles,
Octolasmis spp., use a number of decapod crustaceans as hosts,
including panulirid and scyllarid lobstes (Jeffries et al., 1982).
These barnacles live on the exoskeleton and within the gill cham-
bers of their hosts. They are highly adapted to the life history pat-
terns of their hosts. One species, O. bullata, had a prevalence of 48%
on Panulirus polyphagus off Singapore (Jeffries et al., 1982). No
other species were found on the lobster, but other species were re-
corded from a number of different decapods from the area. Octolas-
mis angulata has been recorded from P. versicolor from Western
Australia (Jones, 2004). Octolasmis californiana was first reported
from the California spiny lobster, P. interruptus (Newman, 1960),
but it is known to occur on several decapods (Newman and Abbott,
1980). Another lepadomorph barnacle, Trilasmis fissum hawaiiensis
is specific to P. japonicus and P. penicillatus where it lives on the se-
tae of the mouthparts (Bowers, 1968). This relationship is quite
intriguing and indicative of the specialization that can occur in
barnacles.
There are few reports of balanomorph, or acorn barnacles, on
spiny lobsters. However, Chelonibia patula and a few other species
sometimes infest them. Chelonibia patula is a host generalist, found
on turtles, drift wood, decapods and other hard surfaces. It will set-
tle on virtually any hard surface. Species of Paralepas occur on
spiny lobsters from Australia (Jones, 2003). Several species of bal-
anomorph barnacles have been found on spiny lobsters P. argus
(Eldred, 1962). Heavy infestations of acorn barnacles may indicate
that the host lobster is not in good health and may be suffering an
underlying infection. Such hosts should receive priority for health
examinations for rare infections.
Bryozoans, hydroids, serpulorbid molluscs, bivalve molluscs
and many other animals can be found as fouling organisms on
the shells of spiny lobsters, but there are few records of their
occurrence. In horseshoe crabs and true crabs, the fouling com-
munity has been used to address ecological questions on succes-
sion (Key et al., 1996, 1997, 1999). I have seen fewer fouling
87
organisms on lobsters than on crabs, perhaps because many crab
species have terminal molts whereas lobsters often do not.
21. Disease syndromes
21.1. Turgid lobster syndrome
Turgid lobster syndrome is an unusual condition that occurs
in J. edwardsii from New Zealand (Diggles, 1999) and Panulirus
ornatus from Australia (Evans, pers. communication). The etiol-
ogy of the syndrome is unknown, but the condition is easy to
identify because the thin arthrodial membranes between the
tergites swell from fluid pressure, causing the animals to become
stiff and lifeless. Affected animals have difficulty moving so they
stop feeding and become lethargic. It occurs in short-term aqua-
culture facilities and is probably the result of poor water quality,
but that remains to be determined. Up to 50% of the affected
animals can die (Diggles, 1999). Animals can be bled with a syr-
inge to relieve fluid pressure, and those treated this way often
survive.
A similar condition was described in lobsters from Japan
(Wada et al., 1994). Lobsters were lethargic and had visible
swelling of the arthrodial membranes. Histology revealed an
inflammation of the myocardium as well as infiltration of hemo-
cytes into affected cardiac muscles. Interestingly, the changes to
the heart were visible as white nodules in the heart. The etiology
of the syndrome was not determined, but it was not a microbial
agent. Both wild and laboratory-held animals showed signs of
the syndrome.
21.2. Pink lobster syndrome
An unusual syndrome occurs in P. cygnus from Western Austra-
lia. The condition is known as pink lobster syndrome and the flesh
of affected animals becomes pink or orange (see Shields et al.,
2006). Affected lobsters usually die from the syndrome. They show
signs of lethargy and have a bitter flavor, making them unpalat-
able. Very little information is available on this syndrome, but it
appears to occur in holding facilities as well as in nature. This con-
dition should be further documented to determine if it is caused by
an infectious or non-infectious agent.
21.3. Mass mortalities
Several mass mortalities have been documented in spiny lob-
ster populations. An interesting mortality event was recorded from
a tagging study that was undertaken to estimate the size of the
population of P. ornatus at Yule Island, Papua New Guinea (Dennis
et al., 1992). The mortality of adult lobsters was high, 95%, during
the fishing period; however, only a third of the mortality was
attributed to fishing pressure, the rest was thought to be due to
physiological stress from reproduction and migration, but disease
agents were not examined. These events are different from those
that alter habitat, thereby forcing migrations such as observed
with hypoxia events. Lobster mortalities are relatively common
along the coast of South Africa (see Cockcroft, 2001). Hypoxia
caused by harmful algal blooms develops in deep waters and forces
lobsters, Jasus lallandii, to move into extremely shallow littoral
areas, which are subject to rapid tidal changes. Lobsters cannot es-
cape exposure to air, and when caught in the intertidal zone, they
die in mass strandings. Ecological disturbances to the Florida Keys
have altered the habitats of the juvenile lobsters causing them to
move into less impacted areas (Butler et al., 1995). The potential
for increased exposure to pathogens has not been studied in these
88 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
systems, but the emergence of PaV1 shortly after these events is
very intriguing.
22. Conclusions
Spiny lobsters have few reported parasites, diseases and symbi-
onts. Several species of spiny lobster have supported fisheries for
many years, so one would expect their diseases to be well docu-
mented in their fished populations. However, this is not the case.
This is primarily because fisheries focus on animals of legal size,
which are typically adults. In my experience, diseases are more
common and more serious in postlarval and early juvenile stages,
a segment of the population that is not sampled by the fishery. Fur-
ther, it is likely that new emerging diseases will become significant
issues in nascent aquaculture industries for spiny lobsters. In fact,
several microbial pathogens are known to be problematical to cul-
tures of larval and juvenile clawed lobsters (see Fisher et al.,
1976a,b, Nilson et al., 1975, 1976; Brock and Lightner, 1990). These
include bacterial (Vibrio spp., L. mucor), fungal (Lagenidium spp.)
and protozoal (peritrich ciliates) agents, all of which can be diffi-
cult to control in closed systems. No doubt more symbionts and
disease agents will emerge as fisheries become more intensively
managed and aquaculture production force animals into high den-
sity, stressful conditions.
The presence of emerging pathogens and disease outbreaks are
often signs that a population is under significant stress. Identifying
the stressors that ultimately lead to or contribute to outbreaks can
be very difficult. Such factors as degrading water quality, seasonal
hypoxia or increased temperatures have been linked to disease or
death in other systems; and they are likely to operate in popula-
tions of spiny lobsters. While such stressors have yet to be impli-
cated in disease in spiny lobster populations, several have been
implicated in mortalities of clawed lobsters. For example, a major
mortality occurred in Long Island Sound in 1999. Several factors
were thought to have contributed to the mortality, including tem-
perature stress, hypoxia, increased manganase concentrations,
pesticides, other pollutants, Neoparamoeba sp. or combinations of
these factors (Pearce and Balcom, 2005; Howell et al., 2005). It
can be very difficult to undertake multifactorial studies on multiple
stressors, but in the case of the 1999 mortality, experiments indi-
cated that lobsters cannot tolerate combinations of severe hypoxic
conditions, poor water quality and temperature stress (Dove et al.,
2004, 2005; Draxler et al., 2005a,b; Robohm et al., 2005). Thus, it is
important to understand both the proximate and ultimate causes
that give rise to emerging pathogens to determine their potential
threat to lobster fisheries.
At present, the virus PaV1 in P. argus represents the most sig-
nificant pathogen to any spiny lobster. It infects the smallest
juveniles and, therefore, would normally go unnoticed in popula-
tions. However, fishermen in the Caribbean spiny lobster fishery
use juveniles as social attractants (bait) in fished traps and
lethargic diseased lobsters would be discarded. Thus, fishermen
may spread diseased animals, or at least the virus, into new fish-
ing grounds (Shields and Behringer, 2004). While contact trans-
mission is the most efficient mode for the spread of PaV1,
water-borne transmission can also occur, albeit at a much lower
efficiency (Butler et al., 2008). However, healthy lobsters can de-
tect and avoid diseased animals, which makes contact transmis-
sion less likely to occur in nature (Behringer et al., 2006). The
ramifications of transmission to the spread of PaV1 is unclear,
but it is clear that the virus can threaten the valuable Caribbean
fisheries (Shields and Behringer, 2004). Recent emerging dis-
eases, such as PaV1 in P. argus and Hematodinium in the Norway
lobster (e.g. Field et al., 1992), highlight the fact that we know
very little about the pathogens in our lobster stocks.
Acknowledgments
This work was partially supported by NSF Grants OCE-0136894
and OCE BE-UF-0723662. This is contribution #3101 from the Vir-
ginia Institute of Marine Science.
References
Abraham, T.J., Rahman, M.K., Joseph, M.T.L., 1996. Bacterial disease in cultured spiny
lobster, Panulirus homarus (Linnaeus). J. Aquacult. Trop. 1, 187–192.
Abrahams, D., Brown, W.D., 1977. Evaluation of fungicides for Haliphthoros
milfordensis and their toxicity to juvenile European lobsters. Aquaculture 12,
31–40.
Alderman, D.J., 1973. Fungal infection of crawfish (Palinurus elephas) exoskeleton.
Trans. British Mycol. Soc. 61, 595–597.
Alderman, D.J., 1981. Fusarium solani causing an exoskeletal pathology in cultured
lobsters, Homarus vulgaris. Trans. British Mycol. Soc. 76, 25–27.
Bach, S.D., Beardsley, G.L., 1976. A disease of the Florida spiny lobster. Sea Frontiers
22, 52–53.
Bang, F.B., 1970. Disease mechanisms in crustacean and marine arthropods. In:
Snieszko, D.F. (Ed.), A Symposium on Diseases of Fish and Shellfish. Amer. Fish.
Soc. Spec. Publ. 5. Bethesda, Maryland, pp. 383–404.
Behringer Jr., D.C., Butler III, M.J., Shields, J.D., 2006. Avoidance of disease in social
lobsters. Nature 441, 421.
Behringer Jr., D.C., Butler, M.J., Shields, J.D., 2008. Effect of PaV1 Infection on
Caribbean Spiny Lobster (Panulirus argus) movement, condition, and survival.
Mar. Ecol. Prog. Ser. 359, 26–33.
Bland, J.A., Brock, T.D., 1973. The marine bacterium Leucothrix mucor as an algal
epiphyte. Mar. Biol. 23, 283–292.
Bobes, R., Diaz, J., Diaz, E., 1988. Aislamiento e identificacion de Aerococcus viridans
var. Homari en la langosta Panulirus argus con sintomas de septicemia. Rev.
Invest. Marin. 9, 97–103.
Boghen, A.D., 1982. Effects of Wescodyne and malachite green on parasitic ciliates
of juvenile American lobsters. Progr. Fish-Cultur. 44, 97–99.
Booth, J.D., 2006. Chapter 10 Jasus Species. In: Phillips, B.F. (Ed.), Lobsters: biology,
management, aquaculture and fisheries. Blackwell Scientific, UK, pp. 340–357.
Bourne, D.G., Høj, L., Webster, N.S., Swan, J., Hall, M.R., 2006. Biofilm development
within a larval rearing tank of the tropical rock lobster, Panulirus ornatus.
Aquaculture 260, 27–38.
Bourne, D.G., Young, N., Webster, N., Payne, M., Salmon, M., Demel, S., Hall, M., 2004.
Microbial community dynamics in a larval aquaculture system of the tropical
rock lobster, Panulirus ornatus. Aquaculture 242, 31–51.
Bowers, R.L., 1968. Observations on the orientation and feeding behavior of
barnacles associated with lobsters. J. Exp. Mar. Biol. Ecol. 2, 105–112.
Bowman, T.E., Kornicker, L.S., 1967. Two new crustaceans: the parasitic copepod
Sphaeronellopsis monothrix (Choniostomatidae) and its myodocopid ostracod
host Parasterope pollex (Cylindroleberidae) from the southern New England
coast. In: Proc. US Nat. Mus., vol. 123, pp. 1–28.
Bowser, P.R., Rosemark, R., Reiner, C.R., 1981. A preliminary report of vibriosis in
cultured American lobsters, Homarus americanus. J. Invertebr. Pathol. 37, 80–85.
Boxshall, G.A., Lincoln, R.J., 1983. Some new parasitic copepods
(Siphonostomatoida: Nicothoidae) from deep-sea asellote isopods. J. Nat. Hist.
17, 891–900.
Brinkley, A.W., Rommel, F.A., Huber, T.W., 1976. The isolation of Vibrio
parahaemolyticus and related vibrios from moribund aquarium lobsters. Can.
J. Microbiol. 22, 315–317.
Brock, J.A., Lightner, D.V., 1990. Diseases caused by microorganisms. In: Kinne, O.
(Ed.), Diseases of Marine Animals, Diseases of Crustacea. Biologische Anstalt
Helgoland, vol. III. Hamburg, Germany, pp. 245–349.
Brock, J.A., 1983. Diseases (infectious and noninfectious), metazoan parasites,
predators, and public health considerations in Macrobrachium culture and
fisheries. In: McVey, J.P. (Ed.), Handbook of Mariculture and Crustacean
Aquaculture. CRC Press, Boca Raton, Florida, pp. 329–370.
Butler, M.J., Behringer, D.C., Shields, J.D., 2008. Transmission of a pathogenic virus
(PaV1) and its effects on the survival of juvenile Caribbean spiny lobster,
Panulirus argus. Dis. Aquat. Org. 79, 173–182.
Butler IV, M.J., Hunt, J.H., Herrnkind, W.F., Childress, M.J., Bertelsen, R., Sharp, W.,
Matthews, T., Field, J.M., Marshall, H.G., 1995. Cascading disturbances in Florida
Bay, USA: cyanobacteria blooms, sponge mortality, and implications for juvenile
spiny lobsters Panulirus argus. Mar. Ecol. Prog. Ser. 129, 119–125.
Caballero, G., 1959. Tremátodes de las tortugas de México. VII. Descripción de un
tremátodo digéneo que parasita a tortugas marinas comestibles del Puerto de
Acapulco, Guerrero. Anales del Instituto de Biología, Universidad Nacional
Autónama de México 30, 159–166.
Campbell, A., Gibson, R., Evans, L.H., 1989. A new species of Carcinomertes
(Nemertea: Carcinonemertidae) ectohabitant on Panulirus cygnus (Crustacea:
Panuliridae) from Western Australia. Zool. J. Linn. Soc. 95, 257–268.
Castro, K.M., Angell, T.E., 2000. Prevalence and progression of shell disease in
American lobster, Homarus americanus, from Rhode Island waters and the
offshore canyons. J. Shellfish Res. 19, 691–700.
Castro, K.M., Factor, J.F., Angell, T., Landers Jr., D.F., 2006. The conceptual approach
to shell disease revisted. J. Crustac. Biol. 26, 646–660.
Cavalier-Smith, T., 1993. Kingdom Protozoa and its 18 phyla. Microbiol. Rev. 57,
953–994.
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
Chang, P.S., Chen, H.C., Wang, Y.C., 1998. Detection of white spot syndrome
associated baculovirus in experimentally infected wild shrimp, crab and
lobsters by in situ hybridization. Aquaculture 164, 233–242.
Charmantier, G., Charmantier-Daures, M., Waddy, S.L., Aiken, D.E., 1991. Salinity
tolerance and osmoregulation in the nemertean Pseudocarcinonemertes homari,
an egg predator of American lobster, Homarus americanus. Can. J. Fish. Aquat.
Sci. 48, 209–214.
Cockcroft, A.C., 2001. Jasus lalandi ‘walkouts’ or mass strandings in South Africa
during the 1990s: an overview. Mar. Freshw. Res. 52, 1085–1094.
Colwell, R.R., Wicks, T.C., Tubiash, H.S., 1975. A comparative study of the bacterial
flora of the hemolymph of Callinectes sapidus. Mar. Fish. Rev. 37, 29–33.
Cornick, J.W., Stewart, J.E., 1975. Red crab (Geryon quinquedens) and snow crab
(Chionoecetes opilio) resistance to infection by the lobster pathogen Aerococcus
viridans (var.) homari.. J. Fish. Res. Bd. Can 32, 702–706.
Costello, M.J., Holmes, J.M.C., McGrath, D., Myers, A.A., 1990. A review and catalogue
of amphipod Crustacea in Ireland. Irish Fish. Invest. Ser. B, No. 33 (1989), 70 pp.
Couch, J.A., 1983. Diseases caused by Protozoa. In: Provenzano, A.J., Jr. (Ed.), The
Biology of the Crustacea, Pathobiology, vol. 6. Academic Press, New York, pp.
79–111.
Dale, T., Blom, G., 1988. Epigrowth organisms on reared larvae of the European
lobster, Homarus gammarus. Fauna 40, 16–19.
Dannevig, A., 1928. Beretning for Flødevigens utklekningsanstalt 1926–1927.
Arsberetning Vedkommende Norges Fiskerier 150, 156.
Dannevig, A., 1937. Beretning for Flødevigens utklekningsanstalt 1936–1937.
Arsberetning Vedkommende Norges Fiskerier, pp. 1–6.
Davis, J.W., Sizemore, R.K., 1982. Incidence of Vibrio species associated with blue
crabs (Callinectes sapidus) collected from Galveston Bay. Texas. Appl. Environ.
Microbiol. 43, 1092–1097.
Deblock, S., Williams, A., Evans, L.H., 1991. Contribution a l’etude des
Microphallidae Travassos 1920 (Trematoda). Description de Thulakiotrema
genitale n. gen., n. sp., metacercaire parasite de langoustes australiennes. Bull.
Mus. Nat. Hist. Natur., Paris 12, 563–576.
Dennis, D.M., Munday, B.L., 1994. Microsporidiosis of palinurid lobsters from
Australian waters. Bull. Eur. Assoc. Fish Pathol. 14, 16–18.
Dennis, D.M., Pitcher, C.R., Prescott, J.H., Skewes, T.D., 1992. Severe mortality in a
breeding population of ornate rock lobster Panulirus ornatus (Fabricius) at Yule
Island, Papua New Guinea. J. Exp. Mar. Biol. Ecol. 162, 143–158.
Diggles, B.K., 1999. Diseases of spiny lobsters in New Zealand. In: Evans, L.H., Jones,
B.J., (Ed.), International Symposium on Lobster Health Management. Curtin
University, Adelaide, Australia, pp. 18–34.
Diggles, B.K., 2001. A mycosis of juvenile spiny rock lobster Jasus edwardsii (Hutton,
1875) caused by Haliphthoros sp., and possible methods of chemical control. J.
Fish Dis. 24, 99–110.
Diggles, B.K., Moss, G.A., Carson, J., Anderson, C., 2000. Luminous vibriosis in rock
lobster Jasus verreauxi (Decapoda: Panuliridae) phyllosoma larvae caused by
infection with Vibrio harveyi. Dis. Aquat. Org. 43, 127–137.
Dove, A.D.M., Allam, B., Powers, J.J., Sokolowski, M.S., 2005. A prolonged
thermal stress experiment on the American lobster. J. Shellfish Res. 24,
761–766.
Dove, A.D.M., LoBue, C., Bowser, P., Powell, M., 2004. Excretory calcinosis: a new
fatal disease of wild American lobsters Homarus americanus. Dis. Aquat. Org. 58,
215–221.
Draxler, A.F.J., Robohm, R.A., Wieczorek, D., Kapareiko, D., Pitchford, S., 2005a. Effect
of hatibat biogeochemicals on survival of lobsters (Homarus americanus). J.
Shellfish Res. 24, 821–824.
Draxler, A.F.J., Sherrell, R.M., Wieczorek, D., Lavigne, M.G., Paulson, A.J., 2005b.
Manganese concentration in lobster (Homarus americanus) gills as an index of
exposure to reducing conditions in eastern Long Island Sound. J. Shellfish Res.
24, 815–820.
Eldred, B., 1962. The attachment of the barnacle, Balanus amphitrite niveus Darwin,
and other fouling organisms to the rock shrimp, Sicyonia dorsalis Kingsley.
Crustaceana 3, 203–206.
El-Gamal, A.A., Alderman, D.J., Rodgers, C.J., Polglase, J.L., MacIntosh, O., 1986. A
scanning electron microscope study of oxolinic acid treatment of burn spot
lesions of Macrobrachium rosenbergii. Aquaculture 52, 157–171.
Estrella, B.T., 1991. Shell disease in American lobster (Homarus americanus, H. Milne
Edwards, 1837) from Massachusetts coastal waters with considerations for
standardizing sample. J. Shellfish Res. 10, 483–488.
Evans, L.H., Brock, J.A., 1994. Diseases of spiny lobster. In: Phillips, B.F., Cobb, J.S.,
Kittaka, J. (Eds.), Spiny Lobster Management. Blackwell Scientific Publications,
Oxford, pp. 461–471.
Evans, L.H., Jones, J.B., Brock, J.A., 2000. Diseases of spiny lobster. In: Phillips, B.F.,
Kittaka, J. (Eds.), Spiny Lobsters: Fisheries and Culture. Fishing News Books,
Blackwell Science, Oxford, pp. 586–600.
Field, R.H., Chapman, C.J., Taylor, A.C., Neil, D.M., Vickerman, K., 1992.
Infection of the Norway lobster Nephrops norvegicus by a Hematodinium-
like species of dinoflagellate on the west coast of Scotland. Dis. Aquat.
Org. 13, 1–15.
Fisher, W.S., Nilson, E.H., Shleser, R.A., 1975. Effect of the fungus Haliphthoros
milfordensis on the juvenile stages of the American lobster Homarus americanus.
J. Invertebr. Pathol. 26, 41–45.
Fisher, W.S., Nilson, E.H., Follett, L.F., Shleser, R.A., 1976a. Hatching and rearing
lobster larvae (Homarus americanus) in a disease situation. Aquaculture 7, 75–
80.
Fisher, W.S., Nilson, E.H., Steenbergen, J.F., Lightner, D.V., 1978. Microbial diseases of
cultured lobsters: a review. Aquaculture 14, 115–140.
89
Fisher, W.S., Rosemark, R., Nilson. E.H., 1976b. The susceptibility of cultured
American lobsters to a chitinolytic bacterium. In: Proc. World Maricult. Soc., vol.
7, pp. 511–520.
Flegel, T.W., Boonyaratpalin, S., Fegan, D.F., Guerin, M., Sriurairatana, S., 1992. High
mortality of black tiger prawns from cotton shrimp disease in Thailand. In:
Shariff, M., Subasinghe, R.P., Arthur, J.R. (Eds.), Disease in Asian aquaculture 1.
Fish Health Section. Asian Fisheries Society, Manila, pp. 181–197.
Flegel, T.W., 1997. Major viral diseases of the black tiger prawn (Penaeus monodon)
in Thailand. World J. Microbiol. Biotechnol. 13, 433–442.
Funch, P., Kristensen, R.M., 1995. Cycliophora is a new phylum with affinities to
Entoprocta and Ectoprocta. Nature 378, 711–714.
Geddes, M.C., Musgrove, R., Thomas, C.J., 2003. Investigation of tail fan necrosis in
live-held adult southern rock lobsters. Developments in Rock Lobster
Enhancement and Aquaculture III, April 2001, Adelaide.
Gemperline, P.J., Miller, K.H., West, T.L., Weinstein, J.E., Hamilton, J.C., Bray, J.T.,
1992. Principal component analysis, trace elements, and blue crab shell disease.
Anal. Chem. 64, 523a–532a.
Getchell, R.G., 1989. Bacterial shell disease in crustaceans: a review. J. Shellfish Res.
8, 1–6.
Goggins, P.L., Hurst Jr., J.W., 1960. Progress repot on lobster gaffkyaremia (red tail).
Maine Department of Sea Shore Fisheries 1-9. Mimeo. [Not seen by author.
Cited in Stewart, 1980.].
Gómez del Prado-Rosas, M.C., Álvarez-Cadena, J.N., Lamothe-Argumedo, R., Grano-
Maldonado, M.I., 2003. Cymatocarpus solearis a brachycoeliid metacercaria
parasitizing Panulirus argus (Crustacea: Decapoda) from the Mexican Caribbean
Sea. Anales del Instituto de Biologia, Universidad Nacional Autónoma de
Mexico, Serie Zoologia 74, 1–10.
Gopalan, U.K., Young, J.S., 1975. Incidence of shell disease in shrimp in the New York
Bight. Mar. Pollut. Bull. 6, 149–153.
Hameed, A.S.S., 1994. Experimental transmission and histopathology of brown spot
disease in shrimp (Penaeus indicus) and lobster (Panulirus homarus). J. Aquacult.
Tropics 9, 311–321.
Handlinger, J., Carson, J., Ritar, A.J., Crear, B.J., Taylor, D.P., Johnston, D., 1999.
Disease conditions of cultured phyllosoma larvae and juveniles of the southern
rock lobster (Jasus edwardsii, Decapoda; Palinuridae). In: Evans, L.H., Jones, J.B.
(Eds.), Proceedings of the International Symposium on Lobster Health
Management, Adelaide, September 1999. Curtin University of Technology,
Perth, Australia, pp. 75–87.
Hansen, H.J., 1897. The Choniostomatidae, a family of Copepoda, parasites on
Crustacea Malacostraca. Host & Son, Copenhagen. pp. 1–206.
Harper, R.E., Talbot, P., 1984. Analysis of the epibiotic bacteria of lobster (Homarus)
eggs and their influence on the loss of eggs from the pleopods. Aquaculture 36,
9–26.
Hess, E., 1937. A shell disease in lobsters (Homarus americanus) caused by
chitinivorous bacteria. J. Biol. Bd. Can. 3, 358–362.
Hicks, G.R.F., 1986. Revised keys to Paramphiascopsis Lang (Copepoda,
Harpacticoida, Diosaccidae) including a new species from deep water off New
Zealand. J. Nat. Hist. 202, 389–397.
Hobbs, R.C., Botsford, L.W., 1989. Dynamics of an age-structured prey with density-
and predation-dependent recruitment: the Dungeness crab and a nemertean
egg predator worm. Theor. Pop. Biol. 36, 1–22.
Howell, P., Benway, J., Giannini, C., McKown, K., Burgess, R., Hayden, J., 2005. Long-
term population trends in American lobster (Homarus americanus) and their
relation to temperature in Long Island Sound. J. Shellfish Res. 24, 849–858.
Huchin-Mian, J.P., Rodríguez-Canul, R., Arias-Bañuelos, E., Simá-Álvarez, R., Pérez-
Vega, J.A., Briones-Fourzán, P., Lozano-Álvarez, E., 2008. Presence of Panulirus
argus Virus 1 (PaV1) in juvenile spiny lobsters Panulirus argus from the
Caribbean coast of Mexico. Dis. Aquat. Org. 79, 153–156.
Huchin-Mian, J.P., Briones-Fourzán, P., Simá-Álvarez, R., Cruz-Quintana, Y., Pérez-
Vega, J.A., Lozano-Álvarez, E., Pascual-Jiménez, C., Rodríguez-Canul, R., 2009.
Detection of Panulirus argus Virus 1 (PaV1) in exported frozen tails of subadult-
adult Caribbean spiny lobsters Panulirus argus. Dis. Aquat. Org. 86, 159–162.
Hudson, D.A., Hudson, N.B., Pyecroft, S., 2001. Mortalities of Penaeus japonicus
prawns associated with microsporidean infection. Australian Vet. J. 79, 504–
505.
Iversen, E.S., Beardsley, G.L., 1976. Shell disease in crustaceans indigenous to South
Florida. Prog. Fish-Cultur. 38, 195–196.
Jawahar, A., Kaleemur, R., Leema, J., 1996. Bacteial disease in cultured spiny lobster,
Panulirus homarus (Linnaeus). J. Aquacult. Trop. 11, 187–192.
Jeffries, W.B., Voris, H.K., Yang, C.M., 1982. Diversity and distribution of the
pedunculate barnacle octolasmis in the seas adjacent to Singapore. J. Crustac.
Biol. 2, 562–569.
Jiravanichpaisal, P., Lee, B.L., Söderhäll, K., 2006. Cell-mediated immunity in
arthropods: hematopoiesis, coagulation. Melanization and Opsonization
Immunobiol. 211, 213–236.
Johnson, P.T., 1976. Bacterial infection in the blue crab, Callinectes sapidus: course of
infection and histopathology. J. Invertebr. Pathol. 28, 25–36.
Johnson, P.T., 1983. Diseases caused by viruses, rickettsiae, bacteria, and fungi. In:
Provenzano, A.J. (Ed.), The Biology of Crustacea, Pathology, vol. 6. Academic
Press, New York, pp. 1–78.
Johnson, P.W., Sieburth, J.M., Sastry, A., Arnold, C.R., Doty, M.S., 1971. Leucothrix
mucor infestation of benthic Crustacea, fish eggs, and tropical algae. Limnol.
Oceanogr. 16, 962–969.
Jones, D.S., 2003. The biogeography of Western Australian shallow-water barnacles.
In: Wells, F.E., Walker, D.I., Jones, D.S. (Eds.), The Marine Flora and Fauna of
Dampier, Western Australia. Western Australian Museum, Perth, pp. 479–496.
90 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
Jones, D.S., 2004. Barnacles (Cirripedia: Thoracica) of the Dampier Archipelago,
Western Australia. Rec. West. Austral. Mus. Suppl. 66, 121–157.
Keith, I.R., Paterson, W.D., Airdrie, D., Boston, L.D., 1992. Defense mechanisms of the
American lobster (Homarus americanus): vaccination provided protection
against gaffkemia infections in laboratory and field trials. Fish Shellfish
Immunol. 2, 109–119.
Kellog, S., Steenbergen, J.F., Schapiro, H.C., 1974. Isolation of Pediococcus homari,
etiological agent of gaffkemia in lobsters, from a California estuary. Aquaculture
3, 409–413.
Key Jr., M.M., Winston, J.E., Volpe, J.W., Jeffries, W.B., Voris, H.K., 1999. Bryozoan
fouling of the blue crab Callinectes sapidus at Beaufort, North Carolina. Bull. Mar.
Sci. 64, 513–533.
Key Jr., M.M., Volpe, J.W., Jeffries, W.B., Voris, H.K., 1997. Barnacle fouling of the blue
crab Callinectes sapidus at Beaufort, North Carolina. J. Crust. Biol. 17, 424–439.
Key Jr., M.M., Jeffries, W.B., Voris, H.K., Yang, C.M., 1996. Epizoic bryozoans,
horseshoe crabs and other mobile benthic substrates. Bull. Mar. Sci. 58, 368–
384.
Kiryu, Y., Behringer, D.C., Landsberg, J.H., Petty, B.D., 2009. Microsporidiosis in the
Caribbean spiny lobster Panulirus argus from southeast Florida, USA. Dis. Aquat.
Org. 84, 237–242.
Kitancharoen, N., Hatai, K., 1995. A marine oomycete Atkinsiella panulirata sp. nov.
from philozoma of spiny lobster, Panulirus japonicus. Mycoscience 36, 97–104.
Kittaka, J., 1997. Culture of larval spiny lobsters: a review of work done in northern
Japan. Mar. Freshw. Res. 48, 923–930.
Kuris, A.M., Blau, S.F., Paul, A.J., Shields, J.D., Wickham, D.E., 1991. Infestation by
brood symbionts and their impact on egg mortality in the red king crab,
Paralithodes camtschatica, in Alaska: Geographic and temporal variation. Can. J.
Fish. Aquat. Sci. 48, 559–568.
Lavallée, J., Hammell, K.L., Spangler, E.S., Cawthorn, R.J., 2001. Estimated prevalence
of Aerococcus viridans and Anophryoides haemophila in American lobsters
Homarus americanus freshly captured in the waters of Prince Edward Island.
Canada. Dis. Aquat. Org. 46, 231–236.
Lee, S.C., Corradi, N., Byrnes III, E.J., Torres-Martinez, S., Dietrich, F.S., Keeling, P.J.,
Heitman, J., 2008. Microsporidia evolved from ancestral sexual fungi. Current
Biol. 18, 1675–1679.
Li, C., Shields, J.D., 2007. Primary cell culture of hemocytes from the spiny lobster
and their susceptibility to Panulirus argus virus 1 (PaV1). J. Invertebr. Pathol. 94,
48–55.
Li, C., Shields, J.D., Ratzlaff, R.E., Butler, M.J., 2008. Pathology and hematology of the
Caribbean spiny lobster experimentally infected with Panulirus argus virus 1
(PaV1). Virus Res. 132, 104–113.
Li, C., Shields, J.D., Small, H.J., Reece, K.S., Hartwig, C.L., Cooper, R.A., Ratzlaff, R.E.,
2006. Diagnosis of Panulirus argus virus 1 (PaV1) in the Caribbean spiny lobster
using fluorescence in situ hybridization. Dis. Aquat. Org. 72, 185–192.
Lightner, D.V., Fontaine, C.T., 1975. A mycosis of the American lobster, Homarus
americanus, caused by Fusarium sp.. J. Invertebr. Pathol. 25, 239–245.
Lightner, D.V., Redman, R.M., 1998. Strategies for the control of viral diseases of
shrimp in the Americas. Fish Pathol. 33, 165–180.
Linton, E., 1910. Helminth fauna of the Dry Tortugas. II. Trematodes. Publ. Carnegie
Inst. Wash. 133, pp. 1–98.
Liuzzo, J.A., Novak, A.F., Ortego, J.R., 1965. Physiological changes induced by gamma
irradiation of bacteria from shrimp. J. Food Sci. 30, 710–713.
Lozano-Álvarez, E., Briones-Fourzán, P., Ramírez-Estévez, A., Placencia-Sánchez, D.,
Huchin-Mian, J.P., Rodríguez-Canul, R., 2008. Prevalence of Panulirus argus
Virus 1 (PaV1) and habitation patterns of healthy and diseased Caribbean spiny
lobsters in shelter-limited habitats. Dis. Aquat. Org. 80, 95–104.
Margulis, L., Corliss, J.O., Melkonian, M., Chapman, D.J., 1990. Handbook of
Protoctista. Jones and Bartlett Publishers, Boston. 914 pp.
McAleer, R., Baxter, M., 1983. Black shell disease of the western rock lobster caused
by Fusarium solani. In: Proc. 8th Cong. Internat. Soc. Human Anim. Mycol.,
Massey University, Palmerston North, New Zealand, 1982, pp. 378–382.
McGrath, D., 1981. Benthic macrofaunal studies in the Galway Bay area. Vol.II. The
benthic macrofauna of the Galway Bay area. Part 1. General introduction,
Arthropoda. Ph.D. Thesis, National University of Ireland, 225 pp. (Not seen by
author. Cited in Costello et al., 1990.).
Meyers, T.R., 1990. Diseases caused by protistans. In: Kinne, O. (Ed.), Diseases of
Crustacea. Biologische Anstalt Helgoland, vol. III. Hamburg, Germany, pp. 350–
368.
Montgomery-Fullerton, M.M., Cooper, R.A., Kauffman, K., Shields, J.D., Ratzlaff, R.E.,
2007. Detection of Panulirus argus virus 1 by PCR in Caribbean spiny lobsters.
Dis. Aquat. Org. 76, 1–6.
Morado, J.F., Sparks, A.K., O’Clair, C.E., 1988. Preliminary study of idiopathic lesions
in the Dungeness crab, Cancer magister, from Rowan Bay. Alaska. Mar. Environ.
Res. 26, 311–318.
Musthaq, S.S., Sudhakaran, R., Balasubramanian, G., Sahul Hameed, A.S., 2006.
Experimental transmission and tissue tropism of white spot syndrome virus
(WSSV) in two species of lobsters, Panulirus homarus and Panulirus ornatus. J.
Invertebr. Pathol. 93, 75–80.
Newman, W.A., 1960. Octolasmis californiana, spec. nov., a pedunculate barnacle
from the gills of the California spiny lobster. Veliger 3, 9–11.
Newman, W.A., Abbott, D.P., 1980. Cirripedia. In: Morris, R.H., Abbott, D.P., Haderlie,
E.C. (Eds.), Intertidal Invertebrates of California. Stanford University Press,
Stanford, California, pp. 504–535.
Nilson, E.H., Fisher, W.S., Shleser, R.A., 1975. Filamentous infestations observed on
eggs and larvae of cultured crustaceans. In: Proc. World Maricult. Soc., vol. 6, pp.
367–375.
Nilson, E.H., Fisher, W.S., Shleser, R.A., 1976. A new mycosis of larval lobster
(Homarus americanus). J. Invertebr. Pathol. 27, 177–183.
Noga, E.J., Engel, D.P., Arroll, T.W., McKenna, S., Davidian, M., 1994. Low serum
antibacterial activity coincides with increased prevalence of shell disease in
blue crabs Callinectes sapidus. Dis. Aquat. Org. 19, 121–128.
Obst, M., Funch, P., Kristensen, R.M., 2006. A new species of Cycliophora from the
mouthparts of the American lobster, Homarus americanus (Nephropidae,
Decapoda). Orgs. Divers. Evol. 6, 83–97.
Overstreet, R.M., 1978. Marine Maladies? Worms, Germs, and Other Symbionts
from the Northern Gulf of Mexico. Mississippi-Alabama Sea Grant Consortium,
Gulf Coast Research Laboratory, Ocean Springs, Mississippi. MASGP-78-021,
140 pp.
Overstreet, R.M., 1983. Metazoan symbionts of crustaceans. In: Provenzano, A.J., Jr.
(Ed.), The Biology of the Crustacea, Pathobiology, vol. 6. Academic Press, New
York, pp. 156–250.
Owens, L., Glazebrook, J.S., 1988. Microsporidiosis in prawns from northern
Australia. Austral. J. Mar. Freshw. Res. 39, 301–305.
Payne, M.S., Hall, M.R., Sly, L., Bourne, D.G., 2007. Microbial diversity within early-
stage cultured Panulirus ornatus phyllosomas. Appl. Environ. Microbiol. 73,
1940–1951.
Pearce, J., Balcom, N., 2005. The 1999 Long Island Sound lobster mortality event:
Findings of the Comprehensive Research Initiative. J. Shellfish Res. 24, 691–698.
Phillips, B.F., Melville-Smith, R., 2006. Chapter 11. Panulirus Species In: Phillips, B.F.
(Ed.), Lobsters: biology, management, aquaculture and fisheries. Blackwell
Scientific, UK. pp. 359–384.
Pillai, N.K., 1962. Choniomyzon gen. nov. (Copepoda: Choniostomatidae) associated
with Panulirus. J. Mar. Biol. Assoc., India 4, 95–99.
Porter, L., Butler, M., Reeves, R.H., 2001. Normal bacterial flora of the spiny lobster
Panulirus argus and its possible role in shell disease. Mar. Freshwater Res. 52,
1401–1405.
Ptacek, M.B., Sarver, S.K., Childress, M.J., Herrnkind, W.F., 2001. Molecular
phylogeny of the spiny lobster genus Panulirus (Decapoda: Palinuridae). Mar.
Freshw. Res. 52, 1037–1047.
Rabin, H., Hughes, J.T., 1968. Studies on host-parasite relationships in gaffkemia. J.
Invertebr. Pathol. 10, 335–344.
Rabin, H., 1965. Studies on Gaffkemia, a bacterial disease of the American lobster
Homarus americanus (Milne-Edwards). J. Invertebr. Pathol. 7, 391–397.
Rajendran, K.V., Vijayan, K.K., Santiago, T.C., Krol, R.M., 1999. Experimental host
range and histopathology of white spot syndrome virus (WSSV) infection in
shrimp, prawns, crabs, and lobsters from India. J. Fish Dis. 22, 183–191.
Reuter, R.E., Geddes, M.C., Evans, L.H., Bryars, S.R., 1999. Tail disease in southern
rock lobsters (Jasus edwardsii). In: Evans, L.H., Jones, B.J. (Eds.), International
Symposium on Lobster Health Management. Curtin University, Adelaide, pp.
88–91.
Robohm, R.A., Draxler, A.F.J., Wieczorek, D., Diane Kapareiko, D., Pitchford, S., 2005.
Effects of environmental stressors on disease susceptibility in American
lobsters: a controlled laboratory study. J. Shellfish Res. 24, 773–880.
Roe, P., 1986. Parthenogenesis in Carcinonemertes spp. (Nemertea: Hoplonemertea).
Biol. Bull. 171, 640–646.
Rosen, B., 1967. Shell disease of the blue crab, Callinectes sapidus. J. Invertebr. Pathol.
9, 348–353.
Rosen, B., 1970. Shell disease of aquatic crustaceans. In: Snieszko, S.F. (Ed.), A
Symposium on Diseases of Fishes and Shellfishes. Am. Fish. Soc. Spec. Publ. 5.
Washington, DC, pp. 409–415.
Sadusky, T.J., Bullis, R.A., 1994. Experimental disinfection of lobster eggs infected
with Leucothrix mucor. Biol. Bull. 187, 254–255.
Sakanari, J.A., Moser, M., 1989. Complete life cycle of the elasmobranch
Lacistorhynchus dollfusi Beveridge and Sakanari 1987 (Trypanorhyncha). J.
Parasitol. 75, 806–808.
Schapiro, H.C., Steenbergen, J.G., 1974. Active immunity to gaffkemia in lobsters. In:
Avault, J.W.J. (Ed.), Proceedings of the Fifth Annual Meeting World Mariculture
Society. Louisiana State University Press, Charleston, SC, pp. 145–147.
Schapiro, H.C., Mathewson, J.H., Steenbergen, J.F., Kellogg, S., Ingram, C.,
Nierengarten, G., Rabin, H., 1974. Gaffkemia in the California spiny lobster,
Panulirus interruptus: Infection and immunization. Aquaculture 3, 403–408.
Schuwerack, P.-M., Lewis, J.W., Jones, P.W., 2001. Pathological and physiological
changes in the South African freshwater crab Potamonautes warreni Calman
induced by microbial gill infections. J. Invertebr. Pathol. 77, 269–279.
Shields, J.D., Behringer Jr., D.C., 2004. A new pathogenic virus in the Caribbean spiny
lobster Panulirus argus from the Florida Keys. Dis. Aquat. Org. 59, 109–118.
Shields, J.D., Kuris, A.M., 1988. Temporal variation in abundance of the egg predator
Carcinonemertes epialti (Nemertea) and its effect on egg mortality of its host, the
shore crab, Hemigrapsus oregonensis. Hydrobiologia 156, 31–38.
Shields, J.D., Kuris, A.M., 1990. Carcinonemertes wickhami n. sp. (Nemertea), an egg
predator on the California lobster, Panulirus interruptus. Fish. Bull. 88, 279–287.
Shields, J.D., Overstreet, R.M., 2007. Parasites, symbionts, and diseases. In: Kennedy,
V., Cronin, L.E. (Eds.), The biology and management of the blue crab. University
of Maryland Sea Grant Press, pp. 299–417.
Shields, J.D., Wood, F.E.I., 1993. Impact of parasites on the reproduction and
fecundity of the blue sand crab Portunus pelagicus from Moreton Bay, Australia.
Mar. Ecol. Prog. Ser. 92, 159–170.
Shields, J.D., 2001. Ovicides julieae n. gen., n. sp. (Nemertea: Carcinonemertidae) on
xanthid crabs from the Great Barrier Reef, Australia. J. Crust. Biol. 21, 304–312.
Shields, J.D., Stephens, F.J., Jones, J.B., 2006. Chapter 5: Pathogens, parasites and
other symbionts. In: Phillips, B.F. (Ed.), Lobsters: biology, management,
aquaculture and fisheries. Blackwell Scientific, UK, pp. 146–204.
 J.D. Shields / Journal of Invertebrate Pathology 106 (2011) 79–91
Shoemaker, C.R., 1952. A new species of commensal amphipod from a spiny lobster.
Proc. US Nat. Mus. 102, 231–233.
Silva dos Fernandes Vieira, R.H., Cavalcante, D.S.P., Saker-Sampaio, S., 1987.
Algumas espécies do género Vibrio em Lagostas e camaróes. Arqu. Cienc. Mar.
26, 1–5.
Sindermann, C.J., 1989. The shell disease syndrome in marine crustaceans. NOAA
Technical Memorandum, National Marine Fisheries Service, NMFS-F/NEC-64. 43
pp.
Sindermann, C.J., 1990. Responses of shellfish to pathogens. In: Sinderman, C.J.,
(Ed.), Principal Diseases of Marine Fish and Shellfish, Diseases of Marine
Shellfish, 2nd ed., vol. 2. Academic Press, San Diego, pp. 247–300.
Smolowitz, R.M., Bullis, R.A., Abt, D.A., 1992. Pathologic cuticular changes of winter
impoundment shell disease preceding and during intermolt in the American
lobster, Homarus americanus. Biol. Bull. 183, 99–112.
Smolowitz, R., Chistoserdov, A.Y., Hsu, A., 2005. A description of the pathology of
epizootic shell disease in the American lobster, Homarus americanus, H. Milne
Edwards 1837. Shellfish Res. 24, 749–756.
Snieszko, S.F., Taylor, C.C., 1947. A bacterial disease of the lobster (Homarus
americanus). Science 105, 500.
Sordi, M., 1958. Micosi dei Crostuacei decapodi marini. Riv. Parassitol. 19, 131–137.
Soyer, J., 1973. Paramphiascopsis paromolae n.sp., Copépode Harpacticide récolt6 sur
les lamelles branchiales du Crustacè Dècapode Paromola cuvieri (Risso).
Crustaceana 24, 90–96.
Steenbergen, J.F., Schapiro, H.C., 1974. Gaffkaemia in California spiny lobsters. In:
Avault, J.W.J. (Ed.), Proceedings of the Fifth Annual Meeting of the World
Mariculture Society. Louisiana State University Press, Charleston, SC, pp. 139–
143.
Steenbergen, J.F., Schapiro, H.C., 1976. Filamentous bacterial infestations of lobster
and shrimp gills. Am. Zool. 15, 816.
Stewart, J.E., 1975. Gaffkemia, the fatal infection of lobsters (genus Homarus)
caused by Aerococcus viridans (var.) homari: a review. Mar. Fish. Rev. 37, 20–
24.
Stewart, J.E., 1980. Diseases. In: Cobb, J.S., Phillips, B.F. (Eds.), The Biology and
Management of Lobsters. Academic Press, New York, pp. 301–342.
Stewart, J.E., 1984. Lobster diseases. Helgoland. Meeresunter. 37, 243–254.
Stewart, J.E., Cornick, J.W., Spears, D.I., 1966. Incidence of Gaffkya homari in natural
lobster (Homarus americanus) populations of the Atlantic region of Canada. J.
Fish. Res. Bd. Can. 23, 1325–1330.
Stewart, J.E., Dockrill, A., Cornick, J.W., 1969. Effectiveness of the integument and
gastric fluid as barriers against transmission of Gaffkya homari to the lobster
Homarus americanus. J. Fish. Res. Bd. Can. 26, 1–14.
91
Supamattaya, K., Hoffmann, R.W., Boonyaratpalin, S., Kanchanaphum, P., 1998.
Experimental transmissions of white spot syndrome virus (WSSV) from black
tiger shrimp Penaeus mondon to the sand crab Portunus pelagicus, mud crab
Scylla serrata and krill Acetes sp. Dis. Aquat. Org. 32, 79–85.
Tareen, I.U., 1982. Control of diseases in the cultured population of penaeid shrimp,
Penaeus semisulcatus (de Haan). J. World Maricult. Soc. 13, 157–161.
Tubiash, H.S., Sizemore, R.K., Colwell, R.R., 1975. Bacterial flora of the hemolymph of
the blue crab, Callinectes sapidus: Most probable numbers. Appl. Microbiol. 29,
388–392.
Wada, S., Takayama, A., Hatai, K., Shima, Y., Fushimi, H., 1994. A pathological study
on cardiac disease found in spiny lobsters. Fish. Sci. 60, 129–131.
Wang, Y.-C., Lo, C.-F., Chang, P.-S., Kou, G.-H., 1998. Experimental infection of white
spot baculovirus in some cultured and wild decapods in Taiwan. Aquaculture
164, 221–231.
Webster, N.S., Bourne, D.G., Hall, M., 2006. Vibrionaceae infection in phyllosomas of
the tropical rock lobster Panulirus ornatus as detected by fluorescence in situ
hybridisation. Aquaculture 255, 173–178.
Weinstein, J.E., West, T.L., Bray, J.T., 1992. Shell disease and metal content of blue
crabs, Callinectes sapidus, from the Albemarle-Pamlico estuarine system, North
Carolina. Arch. Environ. Contam. Toxicol. 23, 355–362.
Wickham, D.E., 1979. Predation by Carcinonemertes errans on eggs of the Dungeness
crab, Cancer magister. Mar. Biol. 55, 45–53.
Wickham, D.E., 1986. Epizootic infestations by nemertean brood parasites on
commercially important crustaceans. Can. J. Fish. Aquat. Sci. 43, 2295–2302.
Wickham, D.E., 1988. Nemertean worm predation. In: Sindermann, C.J., Lightner,
D.V. (Eds.), Disease Diagnosis and Control in North American Marine
Aquaculture. Develop. Aquacult. Fish. Sci., vol. 17. Elsevier Scientific
Publishing Company, New York, pp. 226–229.
Wicksten, M.K., 1982. Crustaceans from baited traps and gill nets off Southern
California. Calif. Fish Game 68, 244–248.
Wong, H.C., Chen, M.C., Liu, S.H., Liu, D.P., 1999. Incidence of highly genetically
diversified Vibrio parahaemolyticus in seafood imported from Asian countries.
Inter. J. Food Microbiol. 52, 181–188.
Yamato, S., 1993. A new amphilochid species (Crustacea: Amphipoda:
Amphilochidae) from a spiny lobster. Publ. Seto Mar. Biol. Lab. 36, 99–106.
Young, J.S., Pearce, J.B., 1975. Shell disease in crab and lobster from New York Bight.
Mar. Pollut. Bull. 6, 101–105.
Ziskowski, J., Spallone, R., Kapareiko, D., Robohm, R., Calabrese, A., Pereira, J., 1996.
Shell disease in American lobster (Homarus americanus) in the offshore,
northwest-Atlantic region around the 106-mile sewage-sludge disposal site. J.
Mar. Environ. Eng. 3, 247–271.