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Behavior and Sensory Biology of Slipper Lobsters

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7
Behavior and Sensory Biology of Slipper Lobsters
Kari L. Lavalli, Ehud Spanier, and Frank Grasso
CONTENTS
7.1 Introduction .................................................................................................................................. 134
7.2 Larval (Naupliosomas and Phyllosomas) Behavior...................................................................... 135
7.3 Postlarval (Nisto) Behavior ..........................................................................................................141
7.4 Juvenile Behavior ......................................................................................................................... 142
7.5 Adult Behavior.............................................................................................................................. 142
7.5.1 Feeding Behavior............................................................................................................. 143
7.5.2 Sheltering Behavior and Substrate Preferences............................................................... 146
7.5.3 Mating Behavior .............................................................................................................. 148
7.5.4 Intra- and Interspecific Interactions.................................................................................149
7.5.5 Diel-Activity Patterns ...................................................................................................... 150
7.5.6 Predators and Antipredator Behavior ..............................................................................151
7.5.7 Movement Patterns .......................................................................................................... 152
7.5.7.1 Daily and Seasonal Horizontal Patterns.......................................................... 153
7.5.7.2 Swimming Behavior (Vertical Movements).................................................... 154
7.6 Sensory Biology of Scyllarids ...................................................................................................... 157
7.6.1 Chemoreception............................................................................................................... 158
7.6.2 Mechanoreception ........................................................................................................... 168
7.6.3 Vision............................................................................................................................... 169
7.6.4 Control Mechanisms and Sensory-Motor Integration .....................................................170
7.7 Conclusions .................................................................................................................................. 170
References.............................................................................................................................................. 171
Abstract
Scyllarid lobsters are a diverse group of animals (85 species) that possess varied early
life-history strategies and live in dissimilar habitats and depths — as such, they pos-
sess diverse behaviors, and no “stereotypical” behavioral descriptions are possible. For
species that specialize on bivalves, feeding behavior is probably the closest to being ste-
reotypical, with the sharp-pointed dactyls of the pereiopods acting to pry open bivalves
or remove the flesh from gastropods. Antennules, which are normally organs of dis-
tant chemoreception (olfaction) appear to play a contact chemoreception role (taste)
in the species of this family. Setal types on mouthparts and pereiopods are less diverse
than in the nephropids and palinurids, but are likely to represent bimodal chemo-
mechanosensors and mechanosensors. Substrate preferences are varied, depending on
final habitat of the adults. Some species (e.g., Ibacus and Thenus) are well adapted for
133
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134 The Biology and Fisheries of the Slipper Lobster
digging into the substrate and show an advanced digging behavior compared to others
that are adapted for sheltering in crevices or under living structures (corals, sponges)
and do so either solitarily or communally. Only a few species apparently undertake
migrations from adult grounds, while the majority appear to make nomadic move-
ments only within their adult habitats. Solitary species and social species seem to have
little agonism toward conspecifics, except over food items. Mating behavior has not
been observed; however, here again, there is diversity among species in spermatophore
placement (internal or external) and persistence. Scyllarides,Ibacus, and Thenus spe-
cies appear capable of burst-and-coast swimming to effect what are presumed to be
foraging movements within their habitats or to escape from predator threats. Neuro-
physiological control over such swimming has only been investigated in Ibacus species
and demonstrates loss of giant interneurons that are used by other decapod forms for
escape swimming. Little is known of the sensory inputs affecting scyllarid behaviors.
7.1 Introduction
Despite being recognized for thousands of years as a decapod lobster, slipper lobster behavior and sensory
capabilities, like so many other aspects of their biology, have not been well studied. It is argued that this is
due to their relative insignificance in commercial fisheries operations, even though they have been locally
consumed in many cultures for hundreds of years (see Holthuis 1991; Spanier & Lavalli, Chapter 18).
Only within the last two decades have researchers turned their eyes toward these lobsters and, of these,
only limited studies have focused on their basic biology. The few studies on behavior that currently
exist focus mainly on several species in three subfamilies: the Arctidinae — Scyllarides aequinoctialis
(Lund, 1793), Scyllarides astori Holthuis, 1960, Scyllarides latus Latreille, 1802, and Scyllarides nodifer
(Stimpson, 1866); the Ibacinae — usually Ibacus peronii Leach, 1815, but occasionally Ibacus chacei
Brown & Holthuis, 1998 and Ibacus ciliatus (Von Siebold, 1824); and the Theninae — both Thenus
orientalis (Lund, 1793) and Thenus indicus Leach, 1816.
In terms of sensory biology, numerous anatomical drawings exist in taxonomic descriptions of the
more than 80 species (e.g. see the works of Holthuis: Holthuis 1985, 1991, 1993a, 1993b, 2002; Brown
& Holthuis 1998); these drawings provide information regarding setal positions on pereiopods, but there
is little information on setal types, their functions, or the information they provide for the guidance of
behavior. Even in studies directly examining scyllarid feeding appendages (e.g., Suthers & Anderson
1981; Ito & Lucas 1990; Mikami et al. 1994; Johnston & Alexander 1999; Malcom 2003; Weisbaum
& Lavalli 2004), only a few have provided a detailed analysis of setation — fewer still have provided
information on the development of the setation pattern from larval to adult stages (Ito & Lucas 1990;
Mikami et al. 1994). In the related and well-studied nephropid lobster, Homarus americanus H. Milne
Edwards, 1837 setation developmental patterns, innervation, and responsiveness have been the focus of
numerous studies on mouthparts, pereiopods, and antennules (Factor 1978; Derby 1982; Derby & Atema
1982a, 1982b; Derby et al. 1984; Moore et al. 1991a, 1991b; Lavalli & Factor 1992, 1995; Gomez
& Atema 1996; Guenther & Atema 1998; Kozlowski et al. 2001), which have had important impacts in
every aspect of nephropid lobster biology and its large fishery. A similar situation exists in the well-studied
palinurid lobster, Panulirus argus (Latreille, 1804) where setal innervation and responsiveness have been a
major focus (Derby & Ache 1984; Daniel & Derby 1988; Derby 1989; Steullet et al. 2000a, 2000b, 2001,
2002; Cate & Derby 2001; Derby & Steullet 2001; Derby et al. 2001; Garm et al. 2004). Currently, only
several species of Scyllarides and Ibacus ciliatus have been examined specifically for purported sensory
structures on appendages typically used for feeding (pereiopods) and orientation (antennules) (Mikami
et al. 1994; Malcom 2003; Weisbaum & Lavalli 2004).
This lack of knowledge, not only of adult, but also of larval, postlarval, and juvenile behavior and sensory
capabilities is problematic because these species are increasingly being exploited as by-product in many
localities and as targeted fisheries in certain places in the world (see Chapters 11 to 18 for information on
scyllarid fisheries). Behavior and sensory input affecting behavior contributes to the survival, growth, and
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 135
ultimately, the reproductive success of individuals — thus, this information is vital to an understanding
of why certain individuals, populations, and species are more successful than others in adapting to their
environments and changes within those environments. Our lack of understanding of behavioral patterns of
all life history stages, as well as the basic biology of how (or if) they migrate, when they mature, how they
find mates and food, how and why they are distributed as larvae, postlarvae, juveniles, and adults, and the
kinds of natural challenges the larvae, postlarvae, juveniles, and adults face, can lead to overexploitation
and population depletion, as has already occurred in several regions (see, for example, DiNardo & Moffitt,
Chapter 12; Hearn et al., Chapter 14; Radhakrishnan et al., Chapter 15; Haddy et al., Chapter 17; and
Spanier & Lavalli, Chapter 18).
Because these animals are held readily in laboratory settings, there is potential for understanding
many behaviors and sensory abilities, as has been done for nephropid lobsters, Homarus americanus and
H. gammarus (Linnaeus, 1758), and for many of the palinurid lobster species. This chapter summarizes
the limited behavioral and sensory research on the few species that have been studied and aims to stimulate
in-depth research focused on these topics for more scyllarid species in both naturalistic laboratory settings
(sensu, Atema 1986; Jones 1988; Cowan & Atema 1990; Barshaw & Spanier 1994a) and in field settings
(sensu, Karnofsky et al. 1989a, 1989b; Wahle & Steneck 1991, 1992; Barshaw et al. 2003).
7.2 Larval (Naupliosomas and Phyllosomas) Behavior
The life history of scyllarids is similar to that of both nephropids and palinurids and can be divided
into a series of developmental phases that occupy different ecological niches (Table 7.1; see Figure 1.2
in Lavalli & Spanier, Chapter 1). Compared to information accumulated for such phases in nephropid
lobsters, especially Homarus americanus, and palinurid lobsters, especially Panulirus argus, we know
very little about the corresponding phases in scyllarids. Much of the lack of knowledge is due to lack of
success in sampling the different phases, particularly the larval, postlarval, and juvenile stages. This is
clearly an area where more research is needed.
Observations of larvae are made from sampling of wild specimens (via plankton tows) and laboratory
culture. In the laboratory, females have hatched their embryos during night or early morning hours
(Sims 1966; Robertson 1968). Scyllarids typically begin their pelagic lives as phyllosomal larvae that are
flattened, leaf-like, transparent, planktonic, zoeal forms with long appendages and long cephalic shields
(Phillips et al. 1981). Unlike the postlarvae, juveniles, or adults, the second antennae of early phyllosomal
instars are initially small compared to the first antennae (antennules) and grow to a length similar to the
antennules; they are not broad and flat (Robertson 1969a). However, some species of scyllarids hatch as a
naupliosoma (prelarva or prezoea), and remain in this form for a few minutes to several hours, depending
on species, before molting into the first-stage phyllosoma (Booth et al. 2005; Sekiguchi et al., Chapter 4).
The species that apparently hatch as this early form include: Scyllarides aequinoctialis (Robertson, 1968,
1969a), S. herklotsii (Herklots, 1851) (Crosnier 1972), S. latus (Martins 1985a), Ibacus alticrenatus
(Bate, 1888) (Lesser 1974), I. ciliatus (Harada, 1958, although the observed individuals may have been
I. novemdentatus Gibbes, 1850 instead, as per Holthuis, 1985). It is not clear if other scyllarids complete
this stage within the egg or if the naupliosoma stage has simply not been observed given its short existence
(Sekiguchi et al., Chapter 4). Some rearing studies indicate that certain species simply, however, do not
hatch as naupliosomas (e.g., I. peronii — Stewart et al., 1997).
Dispersal of phyllosomas varies among species and depends largely on whether the parental stock is
found within lagoons formed by coral island barrier reefs or in deeper waters (Baisre 1994; Johnson 1971a,
1971b; Yeung & McGowan 1991; Coutures 2000). Those hatched in coastal lagoons tend to remain there,
while those hatched in deeper waters gradually move shoreward, such that final-stage phyllosomas of some
scyllarids are found much closer to shore than is typical for palinurid phyllosomas (Booth et al. 2005;
Sekiguchi et al., Chapter 4). Scyllarides aequinoctialis,S. astori,S. herklotsii (Herklots, 1851), S. nodifer,
and S. squammosus (H. Milne Edward, 1837) all have oceanic distributions of their phyllosomas and are
presumed to be dispersed in a manner similar to that for palinurids, since few mid- to late-stage larvae are
found in inshore regions (Robertson 1969a; Johnson 1971b; Phillips et al. 1981; McWilliam & Phillips
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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136 The Biology and Fisheries of the Slipper Lobster
TABLE 7.1
Summary of Life-History Characteristics of Scyllaridae and Associations with Environment and Habitat
Naupliosoma Phyllosoma Nisto
Juvenile to
Subadult Adult
Location Near hatching site Oceanic, beyond the shelf break and over ocean
basins (Arctides guineensis,Scyllarides
aequinoctialis,S. astori,S. herklotsii,S. nodifer,
S. squammosus,Parribacus antarcticus,
Antipodarctus aoteanus)
Pelagic to benthic; when pelagic, tends to be
located over adult grounds (e.g., Scyllarides
astori) or, alternatively in some species,
swims towards such grounds from offshore
(except Ibacus chacei)
Benthic Benthic
Arctides — tropical to subtropical localities in
all oceans except the eastern Atlantic
Scyllarides — western, eastern, & central
Atlantic, Indo-Pacific, Mediterranean &
Oceanic and within and beyond the shelf break
(Ibacus peronii,Crenarctus bicuspidatus,
Eduarctus modestus,Galearctus timidus,
Scyllarus depressus)
Red Seas
Parribacus — tropical to subtropical waters in
Pacific, Atlantic, and Indian Oceans
Evibacus — Pacific Ocean (west coast of
Central America)
Ibacus — Indo-West Pacific
Coastal, over the continental shelf, as well as
within and beyond the shelf break (Evibacus
princeps,Ibacus ciliatus,Ibacus novemdentatus,
Ibacus peronii,Biarctus sordidus,Chelarctus
cultrifer,Crenarctus bicuspidatus,Eduarctus
martensii,Galearctus kitanoviriosus,Petrarctus
demani,P. rugosus,Scammarctus batei,Scyllarus
americanus,S. chacei,S. depressus)
Thenus — Indo-West Pacific
Acantharctus — one species each in Atlantic,
Pacific, & Indian Oceans
Remaining scyllarinid species — Indo-West
Pacific
Coastal (Thenus orientalis, some Scyllarus species)
Depth (m) Unknown Pelagic, but depth varies from surface waters to
deeper waters (some reports state that
phyllosomas have been found to depths of 2000
m, with heavy concentrations at 500–600 m)
Pelagic to benthic (burying during day in sand
but swimming in the water column at night:
Thenus orientalis,Eduarctus martensii)
Usually the same
as adults, at
least for species
that settle
closer inshore;
unknown for
species that
may settle
offshore
Arctides — 5–146
Scyllarides — 0–380
Parribacus — 0–20
Evibacus — 2–90
Ibacus — 20–750
Thenus — 8–100
Acantharctus — 20–57
Antarctus — 122–440
Antipodarctus <90
Bathyarctus — 26–800
Biarctus — 0–73
Chelarctus — 100–333
Crenarctus — 0–108
Eduarctus — 4–112
Galearctus — 30–390 or more
Gibbularctus — 12–57
Petrarctus — 5–282
Remiarctus — 18–260
Scammarctus — 152–531
Scyllarus — 0–329
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Behavior and Sensory Biology 137
Habitat Selection Unknown Choice of depth may be the result of negative
phototaxis and responses to depth of the mixed
layer
Varies considerably from spatially
complex reefs to sandy or muddy
habitats
Recruitment in to benthic habitat:
Thenus orientalis and T. indicus
primarily in summer, but can occur
throughout year
Usually the same as
adults, but may
differ in species that
settle offshore and
later return to adult
grounds
Arctides — rocky habitats
Scyllarides — rocky to shelly sand or mud
Parribacus — rocky to sandy
Evibacus — shelly sand to mud
Ibacus — shelly sand to mud on continental
shelves and slopes
Thenus — shelly sand to mud
Acantharctus — rubble, sand, shelly sand
Antarctus — unknown
Antipodarctus — rocky, and among sponges
and corals
Bathyarctus — rubble, shelly sand to mud
Biarctus — reefs, shelly sand to mud
Chelarctus — rock, mud
Crenarctus — rock, coral reefs, shelly sand
Eduarctus — rubble, sand, mud
Galearctus — reefs, corals, mud
Gibbularctus — shelly sand to mud
Scammarctus — shelly sand to mud
Scyllarus — highly variable, but includes
habitats with algal vegetation
Food Unknown Most likely soft foods, such as larvae, small
zooplankton, jellyfish
Nonfeeding at least in Eduarctus
martensii,Petarctus demani,Thenus
orientalis; if feeding, most likely it is
on small, soft foods
Unknown; presumed
to be similar to
adults based on
structure of
mouthparts and gut
Small benthic invertebrates: mollusks,
polychaetes, crustaceans.
Arctides,Scyllarides,Parribacus specialize on
bivalves (families Arcidae, Mytilidae,
Isognomonidae, Pinnidae), limpets, and sea
urchins (Tripneutes) while
Thenus prefers scallops, but also consumes
mollusks (including cephalopods and
gastropods), fish and fish eggs, crustaceans
(shrimp and barnacles), ostracods, polychaetes,
and detritus
Predators Unknown Despite being transparent to blend in with water
column, they are consumed by pelagic and coastal
fish, such as pilot fish, albacore, rudderfish,
sunfish (Ibacus alticrenatus); also found in
stomachs of zooplankton feeding fish and
barracuda (Scyllarus spp.)
Unknown; cryptically colored and
capable of pigment changes
throughout day (transparent while in
water column; pigmented while on
benthos)
Largely unknown;
each species tends
to have cryptic
coloration for their
preferred substrate.
Scyllarides: dusky
grouper, combers,
and rainbow wrasse
Each species has cryptic coloration for their
preferred substrate.
Scyllarides: queen triggerfish; spotted gully
shark; tiger shark; dusky, red, gag, and
goliath groupers
Scyllarus: scorpionfish, dusky flounder;
snakefish; high hat drum; clearnose skata
Unidentified: blackbar soldierfish, bigeye,
yellowtail snapper, great barracuda, gray
snapper; dog snapper; black margate
(Continued)
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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138 The Biology and Fisheries of the Slipper Lobster
TABLE 7.1
(Continued)
Naupliosoma Phyllosoma Nisto Juvenile to Sub-adult Adult
Movements Typically
nonswimming
because legs are
bound
Diel vertical migrations (surface waters at night;
deeper waters during day) — seen in Crenarctus
bicuspidatus,Scyllarus and Scyllarides species off
the Florida Keys, USA, and Chelarctus cultrifer
Forward swimming via vigorous
beating of pleopods; backward
swimming via abdominal flexion;
passive sinking
Some species may
exhibit horizontal
migratory
movements to adult
grounds (e.g.,
Ibacus chacei)
Nomadism: Ibacus,Thenus
Migratory movements: Ibacus chacei
Capable of walking long distances;
capable of alternating walking with
“burst-and-coast” swimming for
relatively long durations (up to 40 min
in Thenus spp.)
Growth one molt to first
phyllosoma stage
Number of molts is highly variable among
subfamilies: oceanic species tend to have more
molts (11–13), those found within and beyond the
shelf break to oceanic areas have an intermediate
number of molts (7–10), those found within and
beyond the shelf break to coastal regions are
highly variable regarding the number of molts
(4–12)
one molt to juvenile stage Largely unknown
Ibacus — likely
>5 molts before
attaining
physiological
maturity
(2 years); molt
increments =
20–35% premolt
size
Thenus — growth
rate rapid in first
year, slowing for
T. indicus in second
year after reaching
70 mm CL (size of
50% maturity is
52 mm CL)
Scyllarides
S. nodifer can attain
30 cm TL in 16–18
mo in laboratory;
S. latus — molt
increments =24%
Variable.
Scyllarides S. astori molts every
18–24 months; growth
increment =∼
6% of premolt size;
females attain sexual maturity between
21 and 23 cm TL (7–8 years); S. latus
molt increments =6%; S. squammosus
attains sexual maturity at 66–67 mm
CL or 47.6 mm TW
Ibacus — yearly molts; some individuals
apparently molt once every two years;
molt increments =11–15% premolt
size; females grow larger than males;
lifespan 15 years
Thenus — growth rate slows for
T. indicus in second year after attaining
sexual maturity and reaching 70 mm
CL; in T. orientalis, growth rate
remains high for several years until
reaching 120 mm CL (50% sexual
maturity is reached at 58 mm CL in
Australian waters and 107 mm TL in
Indian waters; smallest mature female
in Indian waters is 93.5 mm CL);
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 139
females grow larger, but more slowly than
males; in Indian waters, size at 50% maturity
is reached in 1.5 years; lifespan 4–5 years
Scyllarus S. arctus females are mature at
70 mm TL; S. pygmaeus females reach sexual
maturity at 23 mm TL; S. rugosus attains
sexual maturity at 17–25.2 mm CL
Citations See Sekiguchi et al.,
Chapter 4
Distribution: Gurney (1936), Prasad & Tampi
(1960), Johnson (1968, 1971a, 1971b),
Robertson (1968, 1969a, 1969b, 1971),
Crosnier (1972), Ritz & Thomas (1973),
Shojima (1973), Prasad et al. (1975), Ritz
(1977), Phillips et al. (1981), Atkinson &
Boustead (1982), McWilliam & Phillips
(1983), Barnett et al. (1984), Olvera Limas &
Ordonez Alcala (1988), Yeung & McGowan
(1991), Rothlisberg et al. (1994), Coutures
(2000), Inoue et al. (2001), Minami et al.
(2001), Webber & Booth (2001), Sekiguchi &
Inoue (2002), Higa & Shokita (2004)
Food: Shojima (1963), Sims & Brown (1968),
Mikami et al. (1994), Mikami & Takashima
(2000); see also Mikami & Kuballa, Chapter
5; Johnston, Chapter 6
Predators: Lyons (1970); Bailey & Habib
(1982)
Distribution: Barnett et al. (1986),
Jones (1988); Webber & Booth
(2001)
Food: Barnett et al. (1986); Mikami &
Greenwood (1997); Mikami &
Takashima (1993, 2000); Johnston,
Chapter 6; Jones, Chapter 16
Distribution: Stewart
& Kennelly (1998),
Haddy et al. (2005),
Sekiguchi et al.,
Chapter 4
Food: Johnston,
Chapter 6
Predation: Martins
(1985b), see
Webber & Booth,
Chapter 2
Growth: Rudloe
(1983), Stewart &
Kennelly (1998),
Haddy et al. (2005)
Distribution: Holthuis (1985, 1991, 2002),
Chan & Yu (1986, 1993), Brown & Holthuis
(1998), Spanier & Lavalli, (1998), Davie
(2002), Webber & Booth, Chapter 2
Food: Cau et al. (1978), Suthers & Anderson
(1981), Martins (1985b), Lau (1987, 1988),
Spanier (1987), Jones (1988), Johnston &
Yellowlees (1998), Martínez (2000); see also
Radhakrishnan et al., Chapter 15 and Jones,
Chapter 16
Predation: Lyons (1970), Martins (1985b),
Smale & Goosen (1999)
Movements: Jones (1988), Stewart & Kennelly
(1998), Haddy et al. (2005)
Growth: Kagwade & Kabli (1996a, 1996b),
Relini et al. (1999), Stewart & Kennelly
(2000), Kizhakudan et al. (2004),
Subramanian (2004), DeMartini et al. (2005),
Hearn (2006), Hearn et al., Chapter 14;
Radhakrishnan et al., Chapter 15; Jones,
Chapter 16; Bianchini & Ragonese, Chapter 9
TL =total length; CL =carapace length; TW =tail (abdomen) width.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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140 The Biology and Fisheries of the Slipper Lobster
1983; Yeung & McGowan 1991; Coutures 2000; see Table 1 in Booth et al. 2005). Likewise, larvae
of Arctides and Parribacus species, as well as several genera of scyllarinids (Antipodarctus aoteanus
(Powell, 1949), Crenarctus bicuspidatus (De Man, 1905), Galearctus timidus (Holthuis, 1960), and Scyl-
larus depressus (S.I. Smith, 1881)) have oceanic distributions that often place them in waters that are
deeper than the depth at which their corresponding adult forms are found (Johnson 1971b; Robertson
1971; Phillips et al. 1981; Webber & Booth 2001). In contrast, species of Evibacus,Ibacus,Biarctus,
Chelarctus,Eduarctus,Petarctus,Scammarctus,Thenus, and most Scyllarus are not widely dispersed from
their parental grounds. While swimming behavior of phyllosomas has not been assessed for most scyllarid
species, it may be better developed in species that are dispersed farther offshore (e.g., C. bicuspidatus)
than in species closer to shore (e.g., Thenus spp.), as determined by development and setation of the perei-
opods and their associated exopodites in various instars of phyllosomas, and the pleopods in the nisto
stage (Williamson 1969; Phillips et al. 1981; Minami et al. 2001; see Robertson 1971; Ito & Lucas 1990;
Marinovic et al. 1994; Mikami & Greenwood 1997 for descriptions and diagrams of these appendages in
various species and in different instars).
Coutures (2000) noted that, at least in S. squammosus, first-stage phyllosomas reach the surface rapidly
post-hatching and are positively phototropic. Likewise, Stewart et al. (1997) noted that first-stage phyllo-
somas of I. peronii were active swimmers that were strongly attracted to light. Some larvae are known to
undertake diel vertical migrations (e.g., C. bicuspidatus and Chelarctus cultrifer (Ortmann, 1897)), but
data are limited as to the extent of these migrations and the species-specific preferences for various depths
(Phillips et al. 1981; Minami et al. 2001; Booth et al. 2005; Sekiguchi et al., Chapter 4), as well as the
efficacy of their swimming behavior. It is likely that smaller instars with less developed pereiopods ver-
tically migrate less than later, larger instars with better developed pereiopods (Yeung & McGowan 1991).
Those species or instars that do exploit diel vertical migrations may use passive transport by occupying
vertical strata that move them in specific directions (Booth et al. 2005; Sekiguchi et al., Chapter 4).
Thus, if the larvae are positively phototropic, they may be transported back toward parental grounds or to
juvenile grounds from offshore areas by using surface drift to move inshore. Alternatively, if negatively
phototropic, larvae may use specific subsurface countercurrents or gyres to avoid displacement by surface
currents and to maintain their position over deeper waters (Johnson 1971b; Berry 1974; Sekiguchi 1986a,
1986b; Yeung & McGowan 1991; Lee et al. 1992, 1994; Fiedler & Spanier 1999; Inoue et al. 2000, 2001;
Sekiguchi & Inoue 2002). However, Fiedler & Spanier (1999) found that phyllosoma larvae of Scyllarus
arctus (Linneaus, 1758) in the Eastern Mediterranean were sampled considerable distances offshore and
in long-term, persistent or recurrent eddies. Data from physical oceanography studies imply an increased
probability that phyllosomas can be “trapped” in these eddies for relatively long periods of time, leading to
offshore drift (but see McWilliam & Phillips (1983) who argue that macrozooplanktonic forms, including
Crenarctus (formerly Scyllarus)bicuspidatus, are more abundant in surface waters than in subsurface
eddies). Such offshore drift within a mostly closed basin, such as the Mediterranean, would have advant-
ages — the larvae could possibly reach unexploited or mostly open coastline habitats where they would
have little competition for food and shelter with other decapods. But, the larvae also risk being “lost at
sea” and perishing if they do not reach a proper settlement habitat by the time they metamorphosed into
the nisto stage; this would be particularly problematic for species in large, open ocean basins. Fiedler &
Spanier (1999) suggested that longer planktonic lives and wide dispersal should be correlated with wider
niche breadth of juveniles and adults, and might be more important in species that live in environments
with limited resources, such as those found in the Mediterranean.
Some phyllosomas (various species of Ibacus,Scyllarides,Parribacus, and Scyllarus, as well as Thenus
orientalis,Petrarctus demani (Holthuis, 1946), and Eduarctus martensii (Pfeffer, 1881)) even travel
attached to the aboral surface of jellyfish medusae (Shojima 1963, 1973; Thomas 1963; Herrnkind et al.
1976; Barnett et al. 1986; Booth & Mathews 1994), which may affect their dispersal or allow them
to remain relatively near shore where such medusae are common (Booth et al. 2005; Sekiguchi et al.,
Chapter 4). A clearer understanding of the phototropic responses of the different instars of the phyllosomas,
their sensory abilities with regard to gravity reception and orientation, as well as more information on the
actual depth ranges of their vertical distribution, would further understanding of the recruitment processes
of these larval forms.
Phyllosomas appear to be raptorial predators, using their pereiopods to hold onto food items, which
are then shreaded by the maxillipeds and masticated by molar processes of the mandibles (Mikami &
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Behavior and Sensory Biology 141
Takashima 1994). Scyllarid phyllosomas lack an exopodite on the third maxilliped — a diagnostic feature
that clearly distinguishes these phyllosomas from all palinurid phyllosomas except Jasus spp. and which
suggests differences in feeding strategies or abilities (Booth et al. 2005). Mostly fleshy foods are inges-
ted corresponding to the masticatory abilities of the mouthparts and cardiac stomach; such food types
should be readily available in the water column (Mikami et al. 1994; Booth et al. 2005; Sekiguchi et al.,
Chapter 4). Some Scyllarus larvae have been observed holding hydromedusae, but it is not known if these
were subsequently ingested (Shojima 1963). Nematocysts have been found in the fecal matter of large,
unidentified phyllosomas (Sims & Brown 1968), which suggests that at least some species may use the
hydromedusae for food.
The developmental period for scyllarid larvae is far more variable than that for palinurids, and can last
from a few weeks to at least nine months (Booth et al. 2005). Nauplisomas are small (1 to 2 mm total length,
TL) and short-lived forms. While first-stage phyllosomas can be quite small (see Sekiguchi et al., Chapter 4
for size information on many species), the size of phyllosomas at final stage is highly variable among the
species, ranging from 10 to 80 mm TL (Booth et al. 2005; Sekiguchi et al., Chapter 4). Final-stage phyllo-
somas typically have shorter, broader second antennae and well-developed, feathery exopodites on their
pereiopods (Robertson 1971; Ritz & Thomas 1973). Table 7.1 summarizes the distribution, movement
patterns, food preferences, potential predators, and growth for larval forms.
7.3 Postlarval (Nisto) Behavior
The final-stage phyllosoma metamorphoses into the nisto, or postlarval (megalopa) stage, which, like
spiny lobster pueruli and clawed lobster postlarvae, recruits into the benthic environment. Nistos are
neither completely planktonic nor completely benthic — they are caught in plankton tows demonstrating
that they are pelagic at least part of the duration of this phase (Booth et al. 2005). Some scyllarid nistos are
excellent swimmers (using their abdominal pleopods — Robertson, 1968), while other species are poor
swimmers; some are also capable of executing tail flips (backward swimming) as a means of escape when
disturbed (Lyons 1970; Higa & Saisho 1983; Barnett et al. 1984, 1986; Ito & Lucas 1990). Webber &
Booth (2001) suggest that these swimming differences exist due to marked differences in the size of
pleopods among different species. However, this suggestion has not been adequately tested.
Like the phyllosoma, the nisto is initially completely transparent, which makes it cryptic in the water
column and, no doubt, helps it to avoid predation. In many species of scyllarids, the nisto appears to bury
into soft substrates during the day and swim actively at night; some species even change coloration daily
between these two habitats to remain cryptically colored in both (Barnett et al. 1986; Booth et al. 2005).
Nistos look like adult scyllarid lobsters — the pereiopod exopodites are lost in the metamorphosis, the
second antennae have become flattened, broad, and plate-like, and the abdomen varies in length to about
twice the length of the carapace (e.g., Scyllarus spp.) or about half of the length of the carapace (e.g.,
Ibacus peronii) with functional pleopods (swimmerets) that bear natatory (swimming) setae (Williamson
1969; Robertson 1971; Ritz & Thomas 1973). Some researchers noted the similarity of nistos to adult
Ibacus and Parribacus forms (rather than adult Scyllarides forms) and, as such, nistos were formerly
referred to as a “pseudibacus” (Chace 1966; Crosnier 1972; Holthuis 1993b). The duration of the nisto
phase lasts from seven to 24 days (see Table 1 in Booth et al., 2005). The nisto is typically nine to 13 mm
in carapace length (CL) (Michel 1968; Lyons 1970; Crosnier 1972), but can reach a size of 20 mm CL in
some species (Booth et al. 2005).
As with spiny lobster pueruli, the nisto appears to rely on energy reserves, rather than to actively feed
(Sekiguchi et al., Chapter 4; Mikami & Kuballa, Chapter 5). However, its proventriculus has features that
are transitional between the phyllosoma and the juvenile (Johnston, Chapter 6) and suggest that the ability
to process and sort ingested food particles is more advanced than it is in phyllosomas. While the phyllo-
soma lacks a cardio-pyloric valve that divides the anterior cardiac chamber from the posterior chamber,
the nisto has this feature. But, like the phyllosoma, the nisto lacks a gastric mill, suggesting that food, if
consumed, is similar in softness to that of the phyllosoma, and is primarily masticated by the mouthparts
prior to ingestion (Mikami & Takashima 1993; Johnston, Chapter 6).
Nistos are taken in plankton tows and occasionally are found on spiny lobster pueruli collectors. Better
methods need to be developed to sample this phase in both the pelagic and benthic realms. Additional
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142 The Biology and Fisheries of the Slipper Lobster
studies examining the substratum preferences and methods of substrate sampling could easily be conducted
in laboratory settings to better understand how the nisto makes the transition to juvenile or adult habitat
and to better elucidate the sensory modalities involved in this transition. Table 7.1 summarizes the little
information available about nisto distribution, habitat selection, movement patterns, predators, and growth
patterns.
7.4 Juvenile Behavior
In contrast to an ever-increasing body of knowledge of the juvenile life of clawed and spiny lobsters,
almost nothing is known about the habits of juvenile slipper lobsters. The nisto stage in most species
lasts approximately a half-month to a month. The size of the first juvenile, when known, is dependent on
species, but varies from about 10 to 60 mm CL. In the few species where the molt increment of juveniles
has been determined, juveniles appear to have greater growth increments than adults and more frequent
molting (Stewart & Kennelly 2000; Haddy et al. 2005). In some species (e.g., Ibacus spp., Thenus spp.,
Scyllarides nodifer), growth is rapid and sexual maturity is attained at a relatively young age (two to
three years) (Rudloe 1983; Stewart & Kennelly 2000; Courtney 2002; Haddy et al., Chapter 17); in other
species (e.g., S. astori), growth is slower with sexual maturity occurring at an age >6 years (Hearn 2006).
Large juveniles (subadults) of some species are sampled in pots and trawl nets as by-catch (e.g., Ibacus
spp. — Graham et al., 1993a, 1993b; Graham & Wood, 1997; Thenus spp. — Kizhakudan et al., 2004;
Radhakrishnan et al., Chapter 15), but apparently have not been retained for laboratory studies. In other
species, juveniles have not been sampled at all (e.g., S. latus — Spanier & Lavalli, 1998). A small
preserved male S. latus of 34.3 mm CL was recorded in the Museum of Zoology of the University of
Florence, “La Specola” — it had been collected by a scientific trawl in Italian waters possibly at a depth
>400 m. This record may suggest that, at least in this species, the nisto settles on the substrate in deep water
and the juveniles develop there. Similar suggestions have been made for other scyllarids. For example,
S. astori displays a narrow size range in fishery samples, with almost a complete absence of juveniles
(few individuals smaller than 20 cm TL). Hearn (2006; see also Hearn et al., Chapter 14) suggested that
the juveniles occupy a different spatial niche from adults and are far more cryptic than adults. Scyllarides
nodifer juveniles have not been sampled with adults in the month of April in the Gulf of Mexico, suggesting
that they may have different spring movement patterns than adults (Hardwick & Cline 1990). Likewise,
Ibacus peronii juveniles appear to live in a different habitat than adults and migrate shoreward from
offshore waters to recruit into adult grounds (Stewart & Kennelly 1997). Interestingly, swimming abilities
may differ between nistos and juveniles: in S. nodifer,Scyllarus americanus (S.I. Smith, 1869), S. chacei
Holthuis, 1960, and S. depressus, the relative size of the pleopods is reduced in the juvenile compared
to the nisto and remains reduced until the genital pores develop (Lyons 1970). Since the animals are
experiencing growth in both their carapace and abdomen at this time, but the pleopods initially are about
the same size as those found in the nisto, it is likely that, at least in these species, swimming abilities are
reduced (Lyons 1970). However, as the animals approach sexual maturity, the pleopods increase in size,
presumably for the purpose of oviposition in females.
Table 7.1 summarizes what is known about the distribution, food preferences, predators, and growth
patterns of the juvenile forms. As can be seen in this table, most of what is known about this phase is
speculative, based on assumptions made about adult distribution and biology. Thus, this phase of slipper
lobster life is in great need of further study. However, it is clear that in order to obtain sufficient numbers of
small individuals for such studies, specific sampling techniques must be developed that target the juveniles,
which may prove difficult if many of the species have juvenile development in deep, oceanic waters.
7.5 Adult Behavior
Adults are captured more frequently than other life stages, either by diving, trawling, or pots (see Spanier &
Lavalli, Chapter 18 for detailed information on fishing techniques) and, as a result, adults of several
species have been brought into the laboratory or in controlled field settings for the purposes of studying
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 143
growth (see Bianchini & Ragonese, Chapter 9), larval biology and aquaculture (see Mikami & Kuballa,
Chapter 5; Haddy et al., Chapter 17), and behavior (see Martins 1985b; Lau 1987, 1988; Spanier 1987;
Almog-Shtayer 1988; Jones 1988; Spanier et al. 1988, 1990, 1993; Spanier & Almog-Shtayer 1992;
Barshaw & Spanier 1994a, 1994b; Barshaw et al. 2003; Jones, Chapter 16). One result of this access
is that more is known about adult behavior than any other ontogenetic phase. Various species have also
been captured, tagged, and released to determine wild-growth patterns and movements (see Bianchini &
Ragonese, Chapter 9; DiNardo & Moffitt, Chapter 12; Hearn et al., Chapter 14; Jones, Chapter 16;
and Haddy et al., Chapter 17 for descriptions of such studies). Even so, these studies have tended to
focus only on a few species and, given the highly variable distribution and habitat preference among all
species of scyllarids, care should be taken before generalizing among species within a subfamily or across
subfamilies. This section identifies the species examined and focuses on the behaviors that have been
examined in the laboratory, namely feeding behaviors, shelter preferences and substrate selection, mating
behavior, intraspecific interactions, diel activity patterns, movement patterns, and antipredator behavior.
7.5.1 Feeding Behavior
Much of the work on external feeding mechanisms has focused on the Scyllarides spp. — Scyllarides
aequinoctialis,S. haanii (De Haan, 1841), S. nodifer,S. latus,S. squammosus,S. tridacnophaga Holthuis,
1967 (Holthuis 1968; Lau 1987, 1988; Malcom 2003; Malcom & Lavalli, personal observations), and
the two Thenus spp. — Thenus indicus and T. orientalis (Jones, 1988). One study has used Parribacus
antarcticus (Lund, 1793) and Arctides regalis Holthuis, 1963 (Lau 1988). It is important to characterize
the feeding behavior for specific genera within a subfamily. The descriptions that follow are based on
these studies and should not be overgeneralized for two established reasons. First, the various species
within the four subfamilies differ in food preferences. Some species appear to be invertebrate generalists
(Parribacus antarcticus), while others appear to be molluskan specialists (Scyllarides spp.). Secondly,
fundamental differences exist in the structure of the mouthparts of genera among different subfamilies:
the Arctidinae (Arctides and Scyllarides species) and Ibacinae (Evibacus,Ibacus, and Parribacus species)
possess multi-articulated flagella on their maxillipeds, whereas the Scyllarinae (Acantharctus,Antarctus,
Antipodarctus,Bathyarctus,Biarctus,Chelarctus,Crenarctus,Educartus,Galearctus,Gibbularctus,
Petrarctus,Remiarctus,Scammarctus, and Scyllarus species) and Theninae (Thenus species) lack such a
flagellum on their first and third maxillipeds and bear only a single-segmented flagellum on their second
maxilliped (Webber & Booth, Chapter 2). Ibacinae species have a mandibular palp of only two segments,
while Arctidinae have a three-segmented mandibular palp (Webber & Booth, Chapter 2). There is also a
fundamental difference that separates the Theninae from the other subfamilies — the fifth pereiopod of
the female is achelate in Thenus spp., but is chelate in genera within the other subfamilies (Webber &
Booth, Chapter 2; Jones, Chapter 16). Such differences can affect the use of appendages during the feeding
sequence.
Scyllarides spp. appear to have become specialized for feeding on bivalves (usually scallops, clams,
mussels, or oysters). Bivalves have existed since the pre-Cambrian era and have colonized much of the
world’s marine and aquatic environments. Given their specialization for feeding on bivalves, it seems
almost certain that the radiation of Scyllarides spp. into the central, eastern, and Western Atlantic Ocean,
as well as into the Indo-Pacific (east Asia-Australasia region, east Pacific, and western Indian Ocean)
(Webber & Booth, Chapter 2) followed the beds of bivalves around the world. Where clawed lobsters
crush bivalve shells with their claws and spiny lobsters use their mandibles to crack and chip away at bivalve
shells to access the meat, Scyllarides spp. have evolved an elegant feeding mechanism that involves using
the physics of the bivalve shell to their advantage, while, at the same time, overcoming the disadvantage
of the extremely effective adductor muscles that keep molluskan valves closed. In essence, they “shuck”
bivalves (Lau 1988; Spanier 1987), using tactile and olfactory senses, as well as a guided mechanical
advantage, to avoid the cost that a “brute” force mechanism would require. In contrast, clawed lobsters
use repetitive loading via their crusher claw to cause fracture lines in the rigid shell of bivalves (Moody &
Steneck 1993). This works because there is little organic matrix that would blunt the cracks created by
repetitive loading. Spiny lobsters lack claws but have, in their place, strong mandibles, which they use
to bite cracks into the valve edges of bivalves (Randall 1964; Lavalli, personal observations). Once a
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144 The Biology and Fisheries of the Slipper Lobster
sufficiently large hole is bitten into the edge, the lobster can dig molluskan flesh out of the shells with its
more-rounded pereiopod dactyl tips (Smale 1978). In the case of both claw-loading and biting of bivalves,
the process to crack the shell takes some time; probably, this time exceeds that needed by slipper lobsters to
shuck shells. Scyllarides spp., like spiny lobsters, lack claws, and apparently also lack strong mandibles;
thus, they resort to using their pereiopods — specifically the sharp, pointed, and chitin-reinforced dactyl
tips — to open bivalves (Spanier 1987; Lau 1988).
During the feeding sequence, Scyllarides spp. typically approach a bivalve with their antennules down
near the substrate. Upon encountering the mollusk, the lobster picks up the shell with the first two to three
pairs of pereiopods and repetitively probes the outer valves with its antennules, as though “smelling” and
assessing the shell for its possible value (Lau 1987; Malcom 2003). This behavior contrasts with that of
clawed and spiny lobsters that do not use their antennules for such probing activities, but instead use them
to distantly “chemo-orient” to the food source (Devine & Atema 1982; Zimmer-Faust & Spanier 1987;
Moore & Atema 1991; Moore et al. 1991a, 1991b; Beglane et al. 1997; Nevitt et al. 2000; Derby et al.
2001). In clawed and spiny lobsters, the setose pereiopods are used for initial assessment of food items
(Derby & Atema 1982b). However, after the initial assessment of the shell by the antennules, lobsters
(S. aequinoctialis and S. nodifer) grasp the bivalve and orient its shell rim properly for access by the
pereiopods. With the bivalve thus firmly held with either the first, third, and fourth pairs, or the second,
third, and fourth pairs of pereiopods, the lobster then uses the dactyl tips of either the second or first pair
of pereiopods to repetitively probe the edges of the valves (Malcom 2003). By such repetitive probing,
they eventually wedge the dactyl tips into the shell edge and then insert the tips further and further into
the shell — a process known as “wedging” (Lau 1987). Once one pair of pereiopod dactyls is inserted,
another pair — usually those of the second and the third pereiopods — is used to cut the mantle tissue
along the pallial line (line of attachment to the valve). Then the lobster uses a “scissoring” motion of the
first two pairs of pereiopods to increase the opening angle and to provide access to the adductor muscles
(Figure 7.1A and Figure 7.1B; Malcom 2003). The second or third pereiopod cuts the adductor muscles,
so that the valves open freely. With the valve rims separated, the meat is repetitively scraped from the
surface of the valves and passed directly to the third maxillipeds (Figure 7.1C and Figure 7.1D; Lau 1987;
Malcom 2003). These appendages are used to stretch the flesh and pass the strands back to the subsequent
five pairs of mouthparts for ingestion. Until the molluskan flesh is actually passed back to the third max-
illipeds, the antennules make repeated downward motions to probe inside the valves, to touch the flesh,
and to touch the shell as the pereiopods scrap the flesh from it (Malcom 2003).
(B)
(E)
(A)
(D)
(C)
(F)
FIGURE 7.1 The feeding process of scyllarid lobsters. (A) investigative behavior, using the antennules; (B) probing of the
shell by the first two sets of pereiopods; (C) insertion of the pereiopods into the shell; (D) wedging and subsequent cutting of
the adductor muscles; (E) opening of the shell and removal of the bivalve flesh; (F) scraping of the empty shell. (Video stills
by C. Malcom; used with permission.)
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Behavior and Sensory Biology 145
In S. squammosus this scheme is modified (Lau 1987): the lobster uses the dactyls of the first two pairs
of pereiopods to pry open shell lips; the dactyls of the third pair (or sometimes the second pair) are used to
sever the adductor muscle, and the fourth and fifth pair of pereiopods brace the bivalve against the lobster
and the lobster against the substrate. If the shell is cemented onto a substrate (e.g., oysters), then the lobster
inserts dactyls into the shell while rocking back and forth, presumably to exhaust the adductor muscle
with the repetitive pulls and pushes. For shells attached to substrates via byssal threads, the lobster simply
pulls the shells off and then wedges. If the initial attempt at wedging fails, S. squammosus may attempt
to chip the edge of the shell with its mandibles. Occasionally, lobsters will sample the shell with their
antennules and then simply hold the shell with their dactyls poised two to three cm above the opening and
will wait until the bivalve reopens, whereupon they will plunge the dactyls into the opening, wedging the
valves fully open and cutting the adductor muscles. Typically these different schemes range in duration
from 10 to >40 min (Lau 1987).
The opening of giant clams (Tridacna spp.) by S. tridacnophaga was observed by C. Lewinsohn and
reported in Holthuis (1968). The giant clam uses byssal threads to attach itself to a substrate. The lobster
manipulates the clam to expose its dorsal surface from whence the byssal thread attachments protrude and
then it plunges its dactyls into this exposed and vulnerable region, which causes the clam to gape. At this
point, the lobster turns the shell over and inserts its pereiopods to further wedge the valves open.
Additional observations by Lau (1987) on S. haanii show that this species carries out a scheme similar
to S. squammosus and has similarly shaped, blunt dactyls. In contrast, A. regalis and P. antarcticus have
more tapered, sharper dactyls that appeared, at least to Lau, to be less suited to wedging behaviors. His
observations indicate that these species can perform a type of wedging behavior to open bivalves, but
the time required to do so is longer than that for Scyllarides species. In addition, P. antarcticus appears
to consume a wider variety of prey species than S. squammosus and, while mollusks were still the most
important prey group, they tended to be better represented by gastropods and chitons than by bivalves (Lau
1987). Soft, fleshy, sea anemones represented the second largest proportion of the diet of P. antarcticus,
with annelids, sipunculids, echinoderms, and small arthropods also being present (Lau 1987).
Thenus spp. employ a tactic more similar to clawed lobsters to locate food — they nocturnally search
an area of soft sediment by repetitive probing of their first two pairs of pereiopods, while continuously
moving their antennules up and down and flicking the lateral filaments (Jones 1988). Food odors elicit
chemo-orientation behaviors and directed movements to the source. Contact with the food source results
in grasping and manipulation by the pereiopods (Jones 1988). Presumably they open bivalves in a similar
manner to Scyllarides,Arctides, and Parribacus species, but the exact sequence of leg and mouthpart
movements has not been described (see Jones, Chapter 16 for additional information). In the laborat-
ory, Thenus spp. readily take scallops (Amustium and Chlamys spp.), goatfish (Upeneus spp.), shrimp
(Metapenaeopsis and Trachypenaeus spp.), and lizardfish (Synodus,Saurida, and Tracinocephalus spp.),
but avoid urchins and cuttlefish (Jones 1988; see Table 7 in Jones, Chapter 16).
While the preferred food of Scyllarides spp. appears to be molluskan bivalves, these lobsters are also
known to take sea urchins, crustaceans, sponges, gastropods, barnacles, sea squirts, algae (Ulva spp.),
and fish flesh from gut content analyses. Different species may well prefer different food items, and
setting (laboratory vs. nature) may exert an effect. For example, wild S. latus appear to consume bivalves
(Venus verrucosa Linnaeus, 1758, Glycymeris pilosa (Linnaeus, 1767), Pinctada radiata (Leach, 1814),
Brachidontes semistriatus (Krauss, 1848), Spondylus spinsosus Schreibers, 1793, and limpets (Martins
1985b; Spanier 1987; Spanier, personal observations)). Laboratory tests with S. latus demonstrate that
they prefer soft flesh to crabs and bivalves to soft flesh or snails, while no preference was shown between
choices of oysters, clams, and limpets (Almog-Shtayer 1988). In the laboratory, S.squammosus prefers
Ostrea spp. and Isognomon spp., but will take limpets (Lau 1988), and gut-content analyses of wild-caught
S. squammosus indicate that gastropods are also taken. Wild S. astori prefer white sea urchins, Tripneustes
depressus (A. Agassiz, 1863) (Martínez 2000), but, nonetheless, display a varied diet of molluskan prey
(mostly mussels, pen shells, oysters, arks and cockles) as seen by stomach content analyses (Martínez,
2000).
Parribacus antarcticus gut contents include several species of chitons and gastropods, red algae, sedi-
ment (also seen in Thenus spp.), polychaetes, sabellids, nemertines, sipunculids, sea cucumbers, sea stars,
ostracods, copepods, amphipods, stomatopods, small anomurans, carideans, and brachyurans (Lau 1988).
Thenus spp. also seem to have a broad diet with gut contents of wild-caught animals showing a distinct
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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146 The Biology and Fisheries of the Slipper Lobster
preference for mollusks, followed by sediments, fishes, crustaceans, and polychaetes, with occasional
incidences of sipunculids, sponges, and formaniferans (Kabli 1989). Apparently, wild T. orientalis will
take cuttlefish (Sepia spp.) and squid (Lologio spp.), as well as arrow worms, fish eggs, and barnacles,
but these items may be taken more in scavenging activities, rather than as directly, sought-after, and
captured items (Kabli 1989; see Radhakrishnan et al., Chapter 15 for more information). Kabli’s (1989)
data conflict with laboratory findings of Jones (1988), in which Thenus spp. always avoided cephalopods.
Differences in preferred prey and methods of consumption may allow for niche separation in the
cases where multiple species within a subfamily or multiple genera among the subfamilies overlap in
distribution. They may also allow for niche separation in cases where slipper lobster species are sympatric
with palinurids and nephropids. It is clear from what already is known that certain species may be
generalists that maximize intake of a variety of invertebrates (and some vertebrates), while others may
be highly specialized to feed on a few molluskan species or echinoderm taxa (e.g. S. astori and urchins).
A better understanding of diet and feeding mechanics in the Ibacinae, Scyllarinae, and Theninae would
improve our ability to manage entire ecosystems, especially in cases where multiple taxa are targeted for
fisheries exploitation (e.g., in the Galápagos where spiny, slipper, and urchin fisheries exist) and where
there is an important trophic relationship among the targeted species.
7.5.2 Sheltering Behavior and Substrate Preferences
From species descriptions and various sampling programs/cruises, the general habitat and depth prefer-
ences of most adult scyllards is known (see Table 7.1 for a listing of such preferences and appropriate
references). However, few studies exist that examine the sheltering behavior and preferred physical prop-
erties of scyllarid shelters. Only Jones (1988), working on Thenus spp., and Spanier & Almog-Shtayer
(1992) and Barshaw & Spanier (1994a), working on Scyllarides latus, have conducted laboratory assess-
ments of sheltering behavior and preferred configurations of shelters. Only Spanier et al. (1988, 1990)
and Spanier & Almog-Shtayer (1992) have assessed sheltering behavior in wild settings, although Sharp
et al. (Chapter 11) describe habitat preferences and sheltering behavior of S. aequinoctialis,S. nodifer,
and Parribacus antarctus and Jones (1993) examined abundance and population structure of Thenus spp.
over different habitats and depths to determine if there was a species-specific distribution pattern. This
section summarizes these studies; however, generalizations to other genera based on these studies are
unwise given the broad range of habitats exploited by scyllarid species.
Microhabitat preferences have been determined for S. latus by providing lobsters with a variety of
shelter designs in artificial reefs made of used car tires (for a review of such studies, see Spanier &
Lavalli 1998). Scyllarides latus significantly preferred horizontally oriented dens to vertically oriented
dens where light levels were higher. They also preferred shelters with small, multiple openings, like those
between tires, over those with larger entrances (in the tires themselves) (Spanier et al. 1990; Spanier
& Almog-Shtayer 1992). When additional “back doors” were experimentally blocked, lobsters stopped
using the single-opening dens. These preferences are translated in the use of naturally constructed dens:
all natural dens had two openings and most had a top covering and side walls, as well as some kind of
a back structure (Spanier & Almog-Shtayer 1992). During daylight hours, light in these natural shelters
was 10 to 20 times less than that in the open-reef habitat. In laboratory choice tests, using opaque or
transparent plexiglass pipes for shelters, lobsters significantly preferred opaque to transparent shelters of
the same shape and size. They also preferred medium-sized shelter diameters (20 to 30 cm) that were
open on both ends (Spanier & Almog-Shtayer 1992). A similar preference was shown in the field in an
artificial tire reef where lobsters preferentially chose to live in medium-sized diameter shelters formed
between adjacent tires, rather than in the large, central hole of the tires themselves (Spanier et al. 1988;
Spanier & Almog-Shtayer 1992).
When held in captivity, even in large, naturalistic aquaria with biogenic rocks and sand, S. latus ceased
to show substrate preferences within two months, after initially spending significantly more time on
rocky substrates (Barshaw & Spanier 1994a). In contrast, Chessa et al. (1996) reported that in laboratory
experiments, lobsters preferred rough artificial substrates (plastic carpet) over smooth ones only after
having experience with each for some time. The rougher substrates allowed the lobsters to cling with
their nail-like dactyls, which protects their vulnerable ventral surface from predators (see Section 7.5.6
for additional details). Other than these two reports — one using natural structures and one using artificial
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 147
structures, substrate preferences have not been well studied in other Scyllarides spp. and can only be
inferred from areas where they are caught.
The adults of most species of Scyllarides are found on hard substrates, with some species also inhabiting
soft-bottom habitats or shifting between soft bottom and rocky habitats (see Table 7.1 and Webber & Booth,
Chapter 2). In dive surveys, S. aequinoctialis tended to be more common in high-relief, coral habitats with
ready-made shelters, while S. nodifer was found on both patch reefs with high-relief and unconsolidated
sediments (Sharp et al., Chapter 11). It is likely that Scyllarides species sampled both on hard and soft
substrates result when lobsters that usually shelter in hard substrates are collected in soft substrates during
their short- and long-term movements (see Section 7.5.7 for descriptions of these movements) — such
a suggestion was made by Ogren (1977) to explain the distribution of S. nodifer. Nevertheless, a few
species have been reported only on soft substrates (e.g., S. elisabethae (Ortmann, 1894)). Holthuis (1991)
states that S. elisabethae seems to dig into the mud; he also mentions that S. aequinoctialis buries in the
sand, although others report that this species is a reef dweller that shelters within coral-rock caves and
under coral heads (Moe 1991). Likewise, Hardwick & Cline (1990) have reported that because they were
caked in mud, S. nodifer in the northern Gulf of Mexico may bury in sediments during the daylight. It is
assumed that such digging into soft substrates is an antipredator adaptation, similar to that seen in other
lobster genera living long term or temporarily on soft substrates (e.g., Homarus americanus in offshore
habitats — Cooper & Uzmann, 1977, 1980).
When in shelters, S. aequinoctialis and S. nodifer rarely occupy the den floor, but instead hang from the
ceilings and walls. This is also common in S. latus that are occupying caves (Spanier, personal observa-
tions). Likewise, Parribacus antarcticus, which prefers similar high-relief habitats to those preferred by
S. aequinoctialis, also occupies the den wall, rather than the floor. Scyllarides aequinoctialis, in particular,
appears to have a high degree of den fidelity and the same individual can be found day after day occupying
the same den (Sharp et al., Chapter 11).
Thenus spp. live on soft-bottom substrates into which they bury during the day (Jones 1988; see Jones,
Chapter 16 for a complete description of their burial patterns and substrate use). However, there is a
differential spatial distribution of the two currently recognized valid species (that have yet to be fully
described in the literature; see Webber & Booth, Chapter 2 and Jones, Chapter 16), T. indicus and
T. orientalis. In the laboratory, T. indicus prefers more fine-grained sediments (mud/silt) and T. orientalis
prefers coarser particle sizes (sand) (Jones 1988); these preferences are carried over into wild distri-
butions of the two species which spatially separate not only by depth, but also by the coarseness of
the substrate particles (Jones 1993). Because the preferred substrate is soft and ready-made shelters
are not available, as they would be in rocky or coral habitats, Thenus spp. bury themselves into the
substrate, usually following nocturnal activity periods. Burial behavior involves backward movements
and repetitive extensions of the abdomen, followed by rapid abdominal flexions — these movements
cover the dorsal surface of the carapace and antennae with 1 cm of sediment, leaving only the anten-
nules and eyes exposed (Jones 1988; see Jones, Chapter 16). The entire sequence takes 2 min (Jones
1988). Antennular flicking occurs while buried, albeit at a low frequency, and a gill current is effected
by scaphognathite beating that draws water beneath the eyes and out at the base of the antennules
(Jones 1988). Partial burial is also observed, but this generally occurs during short resting sessions
throughout the nocturnal activity period. Contrary to expectations from the name “shovel-nosed” lob-
ster, the broad, flat, antennae are not used for burial purposes (Jones 1988), but in ovigerous females
of Scyllarus arctus, these antennae of Thenus spp. are apparently used for burial (see Pessani & Mura,
Chapter 13).
Ibacus spp. also are found on soft-bottom substrates and are presumed to bury into those sediments,
much in the same manner as Thenus spp. (see Haddy et al., Chapter 17 for more information). A recent
study (Faulkes 2006) of digging in I. peronii provided a description of the sequence of behaviors involved.
Normally pereiopods three to five are used in a typical alternating tripod gait (Wilson 1966; Johnston &
Yellowlees 1998). Digging is initiated when all pereiopods are inserted into the substrate and the abdomen
is fully flexed and pressed down such that the tailfan contacts the substrate. The abdomen is then extended
which causes the substrate (usually sand) to be pushed backward and results in the abdomen submerging.
The abdomen then flexes and reextends several times, and the pereiopods may be repositioned, by lifting
out the anterior to posterior legs in a metachronal wave and reinserting them in a more posterior position
than before. The sequence ends with a few tail flips that cover the exposed portions of the carapace
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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148 The Biology and Fisheries of the Slipper Lobster
completely with sand. Ibacus peronii is a slow digger, taking >4 min and an average of 17 cycles of
abdominal extension and flexion to completely submerge into the sand. As with Thenus spp., the broad,
flat antennae are not used in this behavior.
7.5.3 Mating Behavior
Because scyllarids copulate, spawn, and brood readily in the laboratory, some aspects of their reproductive
biology are known; however, less is known of the actual mating behavior or the rituals involved during
the mating process. It is likely that the permanence of the spermatophore differs among the genera; it is
also likely that fertilization is internal in some genera and external in others.
In Scyllarides latus, males produce white, gelatinous spermatophores, which they carry around on the
base of their fourth and fifth pereiopods (Almog-Shtayer 1988; Spanier, personal observations). These
are transferred externally to females. It is not clear whether females retain the spermatophores externally
and fertilize their eggs externally, or whether they somehow manipulate the spermatophore and store it
internally. In some Scyllarides species, females have been observed carrying spermatophores externally
6 to 10 days or less prior to egg extrusion (S. latus — Martins 1985b; Almog-Shtayer, 1988), while in others,
the lack of observable spermatophores prior to egg extrusion has led to a belief that the spermatophore is
stored internally and fertilization is internal (S. nodifer — Lyons 1970; and S. squammosus — DeMartini
et al. 2005). Females of most Scyllarides species can spawn multiple broods in a season due to short
brooding periods (but see Hearn et al., Chapter 14 for a contrasting view, in which S. astori broods once
annually), and these broods are usually carried during warm (spring and summer) months. Only in S. latus
have both eggs and spermatophores been observed simultaneously (Almog-Shtayer 1988); however, in
this species, multiple broods have not been observed, with only one peak spawning occurring in spring
and summer months of April to July (Spanier, personal observations).
Ibacus species deposit persistent, gelatinous spermatophores near the genital openings and it is thought
that egg extrusion and fertilization (external) occurs shortly after the spermatophore is deposited on the
female (Stewart & Kennelly 1997; Haddy et al. 2005). Ibacus chacei and I. peronii appear capable of
spawning at least twice per year; however, there is no evidence that I. alticrenatus and I. brucei Holthuis,
1977 have multiple broods, as females in these species possess inactive ovaries at various times during a
year (Haddy et al. 2005). Broods are usually spawned and carried in the colder months (autumn to spring)
with hatching in warmer months (spring through summer).
Parribacus antarcticus has been observed bearing spermatophores that were hard and black, somewhat
similar in form to those typically seen on most spiny lobsters species (reported in Lyons 1970). Apparently
they are capable of carrying both eggs and a spermatophore simultaneously, and can fertilize multiple
broods with the same spermatophore (Lyons 1970; Sharp et al., Chapter 11). They can also repetitively
mate and have been observed with a fresh spermatophore atop a spent one (Sharp et al., Chapter 11). It is
not clear what other species within the genus do with regard to spermatophores.
Thenus species do not deposit a persistent spermatophore, so egg extrusion typically occurs within
hours of mating. Even after 2000 h of remote video observations, no precopulatory courtship behavior has
ever been observed (Jones 1988). The manner of spermatophore transfer is unknown, although Kneipp
(1974) noted that spermatophores were deposited on both sides of the female’s sternum below the genital
openings after a rapid (few seconds) embracement of the ventral surfaces of the animals. Fertilization
is presumed to be external and to occur shortly after mating (Jones 1988; Kizhakudan et al. 2004; see
Radhakrishnan et al., Chapter 15 and Jones, Chapter 16 for more information), and in T. orientalis,
the spermatophoric mass appears to be lost 12 h after mating (Kizhakudan et al. 2004). Spawning in
T. orientalis apparently occurs between September and April (Kabli & Kagwade 1996), with the highest
frequency of ovigerous females occurring November to January (spring and summer). A second spawning
peak can occur in March, so it is likely that this species may be capable of spawning multiple broods
each year (Kabli & Kagwade 1996); however, in the Red Sea, only a single spawning period was noted
(Branford 1982).
Scyllarus species mating habits are not well known. In the laboratory, S. depressus was seen to hatch
larvae and three days later to have extruded another brood (Robertson 1971). No external spermatophore
was noted on the sternum of the female and fertilization was presumed to be internal. Lyons (1970)
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 149
sampled numerous S. depressus females in the Gulf of Mexico, but also never saw the presence of a
spermatophoric mass. However, in S. rugosus, spermatophoric masses were seen on intermolt females
after nocturnal mating and ovipositioning occurred several hours later (Kizhakudan et al. 2004).
7.5.4 Intra- and Interspecific Interactions
In those species for which diving censuses have been conducted, it is clear that slipper lobsters can
range from being solitary to highly gregarious. From observations on species that are gregarious (e.g.,
S. latus, Barshaw & Spanier, 1994a), there is intraspecific competition over food items and, generally, the
largest females are dominant over other lobsters (Barshaw & Spanier 1994a). The least aggressive of such
encounters is the “approach/retreat” sequence common to all lobsters (Atema & Cobb 1980) — one lobster
walks toward the other, which responds by walking away or otherwise avoiding the approaching lobster.
A more aggressive encounter involves use of the flattened second antennae and is called a “flip.” Here the
lobster jerks up its flattened second antennae under the opponent’s carapace, attempting to dislodge it. In
the most intense aggressive behavior, the attacking lobster grabs the anterior portion of the opponent and
holds on with the dactyls of the pereiopods. The opponent usually tail flips, as does the attacking lobster,
which causes the opponent to end up on its back. Often after this “face grab” maneuver, both lobsters are
holding onto the same food item, ventral side to ventral side, and they continue in this fashion until one
lobster finally relinquishes its hold on the food item (Barshaw & Spanier 1994a).
Only one observation of cannibalism was observed in S. latus where an individual was feeding on a
conspecific postmolt, and it was not clear if the cannibalizing individual killed the conspecific during
or after its molt, or if it was simply opportunistically feeding on an individual that died during ecydsis
(Spanier, personal observation).
A tendency for gregarious sheltering among S. latus was observed in a survey of natural dens where 95%
of lobsters cohabited with one or more conspecifics (Spanier & Almog-Shtayer 1992). This gregarious
behavior also was demonstrated by very large lobsters (>100 mm CL) in the field, which differs from
gregarious spiny lobster species where the tendency for cohabitation is greater in smaller and medium-
sized animals (Spanier & Zimmer-Faust 1988). Fishermen have reported aggregations of 50 to 60 lobsters
in the same shelter or shelter-providing structure (reported in Spanier & Almog-Shtayer, 1992).
Similar clustering behavior was observed among S. latus individuals in naturalistic habitats and with
artificial shelters in laboratory tanks. In the absence of a predator, freshly caught, laboratory-held lobsters
significantly preferred an opaque artificial shelter compared to a transparent shelter of the same shape
and size, and formed clusters within this opaque shelter. When they were supplied with no shelter but
with shade, the lobsters concentrated under the shade (Spanier & Almog-Shtayer 1992). When neither
shelter nor shade was supplied, they showed distinct gregarious behavior similar to the defensive “rosette”
observed in migrating Caribbean spiny lobsters under attack by a triggerfish (Herrnkind 1980; Kanciruk
1980; Herrnkind et al. 2001). However, field predation studies indicate this gregarious behavior does not
confer any advantage on exposed individuals within the group who are under attack by fish predators. They
suffer an equal amount of predation as do solitary animals exposed to the same fish predators and gain
only a small advantage of time, as predatory attack patterns are less focused when lobsters are grouped
(Lavalli & Spanier 2001).
Reports of gregarious behavior also exist for S. nodifer (Moe 1991), but these may be question-
able. Rudloe (1983) reported that individuals established their own residences in the laboratory and
that social interactions were limited to displacement of other individuals on mussel clumps. In diver sur-
veys, Sharp et al. (Chapter 11) report that S. nodifer was observed sheltering alone 100% of the time,
and S. aequinoctialis was observed sheltering alone 84 out of 86 instances, and with a second conspecific
only two out of 86 times. In the same survey, Parribacus antarcticus was observed sheltering twice —
once with a spiny lobster and once with a conspecific. In similar surveys of S. astori in the Galápagos
archipelago, divers found that these lobsters were primarily solitary (Hearn et al., Chapter 14). In a study
of the population biology of congeners in the Hawaiian archipelago, Morin & MacDonald (1984) reported
that S. haanii was a solitary species, whereas S. squammosus tended to occur in groups. It seems that, at
least in the genus Scyllarides, there is great variability in the sociality of the species, and this suggests
that other genera may be equally variable in their sociality.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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While Thenus spp. appear to be solitary, they are capable of being held together in tanks, but display
little to no interest in other individuals. Agonistic encounters only appear to arise over food items (as
is the case for S. latus, described earlier), and involve maneuvering for better mechanical advantage in
possession of the food item (via pereiopod movements and tail flipping) (Jones 1988; see Jones, Chapter 16
for more details). Mikami (1995) noted that the phyllosomal stages of Thenus spp. can be cannibalistic,
but only if insufficient food is present in the rearing chambers.
Obviously, more work is needed to understand the nature of associations between conspecifics when they
occur, as well as to determine their value to the individual lobster, and how widespread they are through
the various genera of scyllarids. It is possible that in those species that display communal sheltering or
sheltering with at least one conspecific, the “guide effect,” seen in juvenile Panulirus argus (Childress &
Herrnkind 1996, 2001), may play a role. In the guide effect, individuals are attracted to shelters that
harbor conspecifics, as chemo-orientation to the odor of the conspecific reduces the time necessary to find
an appropriate shelter. Collective denning was highly correlated with conspecific density and scarcity of
local shelters, rather than with lobster size, molt condition, shelter type, or predator density. Childress &
Herrnkind (1996, 2001) suggested that this guide effect benefited the shelter-seeking individual by reducing
the time of exposure on the substrate with its associated predator risk. Once sheltered with conspecifics
(particularly those with weapons, as is the case with spiny lobsters and their spinose, long antennae),
the risk of predation is reduced upon the individual via collective prey vigilance and defense, making
gregarious behavior a beneficial trait (e.g., P. argus: Zimmer-Faust & Spanier 1987; Spanier & Zimmer-
Faust 1988; Butler et al. 1997; Herrnkind et al. 2001; Jasus edwardsii (Hutton, 1875): Butler et al. 1999).
It is likely that collective denning may provide a similar benefit to scyllarids, at least in those species that
live in rocky habitats and for which large shelters are available.
7.5.5 Diel-Activity Patterns
Adult specimens of all scyllarids appear to be well camouflaged due to their flattened morphology and
various coloration patterns that blend into their preferred substrates (see Webber & Booth, Chapter 2
for an expanded discussion of distribution and morphological features). However, for those species that
live in shallow, brightly illuminated waters, this camouflage provides only limited concealment against
diurnal predators. Therefore, like most other lobsters, scyllarids, by and large, appear to be nocturnally
active. However, it is not clear whether deeper-water scyllarids display activity patterns different from
shallow-water species during daytime hours, given the reduction in light at depth. Again, very few studies
examining diel activity patterns exist for scyllarids— only Scyllarides and Thenus spp. have been examined
in laboratory settings (Jones 1988; Spanier & Almog-Shtayer 1992; Barshaw & Spanier 1994a).
Scyllarides spp. forage during the night and shelter during the day on the ceilings of caves, in crevices
in vertical rocky walls (e.g., Barr, 1968; Martínez et al., 2002 for S. astori; Spanier & Lavalli, 1998 for
S. latus), and in other natural dens, as well in artificial reefs in the field (including ship wrecks), or in
man-made shelters in the laboratory (S. nodifer: Rudloe, 1983; S. latus: Spanier et al., 1988, 1990; Spanier
& Almog-Shtayer, 1992; Spanier, 1994). However, in laboratory holding tanks, where predators are not
encountered for a long time, they tend to shift to diurnal activity (Spanier et al. 1988, 1990; Spanier &
Almog-Shtayer 1992), foraging for bivalves and even carrying bivalves back to their shelters to consume
at a later time (Spanier et al. 1988). In addition, when visibility is low during the day, as is common during
and following a storm, lobsters can be detected in the entrances of their shelters and exposed on substrates
(Spanier & Almog-Shtayer 1992; Spanier, personal observations). This latter behavior suggests, as has
been found for Thenus spp. (Jones 1988, see below), that increases in S. latus activity may be linked to
crepuscular periods of the diel cycle.
Based on trawl surveys, it appears that the shallow-water species, Ibacus chacei and I. peronii, also are
largely inactive during the day — catches per 60 min tow during the day averaged 1.8±2.0 (normalized)
and 4.9 ±4.4 individuals, respectively, vs. 37.8 ±28.4 (normalized) and 9.3 ±7.8 at night, respectively
(Graham et al. 1993a, 1993b, 1995, 1996). However, catch rates for I. alticrenatus and I. brucei, both
deep water species, are high during the day (see Haddy et al., Chapter 17 for more information). This
result suggests that diel patterns may vary depending on the preferred depth of a species.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 151
Jones (1988) has provided the most detailed analysis of diel activity patterns to date for any scyllarid
species (see Jones, Chapter 16 for details). Thenus indicus and T. orientalis are shallow-water species that
vary their activities around the diurnal phase, the nocturnal phase, and the crepuscular phases. During
daylight hours, they remain buried with only eyes and antennules exposed and the antennules maintain
a low-frequency flicking rate. During twilight (the first crepuscular phase), lobsters unbury and begin to
forage for food via nomadic movements on the substrate punctuated by brief bursts of swimming activity.
This activity drops off after about 4 h into the nocturnal phase, with lobsters remaining still on the substrate
or partially buried. The major activity here is grooming. At dawn (the second crepuscular phase), activity
again increases and consists primarily of foraging behavior. Lobsters then rebury completely into the
sediment and remain inactive during the day (Jones 1988).
7.5.6 Predators and Antipredator Behavior
By and large, predators of slipper lobsters are currently known only from gut-content analyses of sampled
fish or invertebrate species, but little is known of how various genera escape predation and how the predat-
ors subdue the prey. The general assumption is that the blunt tubercular (or smooth) surface of a flattened
carapace, along with the cryptic coloration patterns of each species provides for a highly cryptic lifestyle
that provides excellent concealment of individuals against their preferred habitat. Barshaw & Spanier
(1994b) noted that lobsters tethered in the open placed themselves alongside rocks, where their cryptic
coloration made them visually difficult to detect; alternatively, they tucked under whatever macroalgae
was in the vicinity in an attempt to hide. In addition, an activity pattern that directly avoids diurnal pred-
ators is assumed to reduce potential predator risk. Finally, in the event that an individual is detected and
attacked, the escape response — rapid, abdominal flexions, resulting in swimming movements — aid the
lobster in escaping to an area where it can reconceal itself. However, none of these assumptions has been
well studied and, given that some scyllarids may undergo migrations over terrains that may or may not
be as cryptic, it is clear that antipredator behaviors need to be more carefully examined.
Scyllarides latus represents the one species in which the response to predator attack has been well
studied (Spanier et al. 1988, 1991, 1993; Barshaw & Spanier 1994b; Lavalli & Spanier 2001; Barshaw
et al. 2003). This response consists of three strategies, two of which are typically executed in sequence:
(1) the “fortress strategy” in which the animal grasps the bottom and attempts to outlast its attacker’s
motivation to penetrate its hard shell (described in Barshaw et al., 2003); (2) the “swimming escape”
response (described in Barshaw & Spanier, 1994a, 1994b, and Barshaw et al., 2003); and (3) remaining
sheltered in dens (Spanier et al. 1988; Spanier & Almog-Shtayer 1992). Lacking claws (like Homarus spp.)
or long, spinose antennae (like spiny lobsters; see Zimmer-Faust & Spanier 1987; Spanier & Zimmer-
Faust 1988; Lozano-Alvarez & Spanier 1997; Herrnkind et al. 2001) with which to fend off swimming
predators, S.latus has developed a shell that is at least twice as thick and more durable to mechanical
insult than clawed or spiny lobsters that live in the same general region (Barshaw et al. 2003; Tarsitano
et al. 2006). They use their short, strong legs to grasp the substrate and resist being dislodged (Barshaw &
Spanier 1994a, 1994b). This clinging force can reach maximum magnitudes of 3 to 15 kg (8 to 29 times
the body weight of the lobster) and linearly correlates with lobster size (Spanier & Lavalli 1998). When
this “fortress defense” fails, they are exceptionally deft swimmers capable of evasive maneuvers like
barrel rolls (presumably using their flat, broad antennae like reciprocal aileron stabilizers on an airplane
wing) en route to a shelter (Spanier et al. 1991). Also they may suddenly change the direction of their
swimming, presumably to confuse the chasing predator, a tactic known as “protean” behavior. This is an
energetically costly response to a threat and is generally used as a last resort. Barshaw et al. (2003) argue
that S. latus has matched the energy invested by clawed lobsters in claws and spiny lobsters in antennae
by increasing only moderately the thickness of their shells and bettering their swimming-escape behavior.
If this strategy is shared among all genera within the family, it would appear to be highly successful, as
slipper lobsters are the most diverse group of lobsters with more than 85 species distributed worldwide
(Booth et al. 2005; Webber & Booth Chapter 2).
As demonstrated with S. latus, slipper lobsters also display a variety of shelter-related behaviors that
provide a third highly effective survival strategy (Barshaw & Spanier 1994a). By combining nocturnal
foraging with diurnal sheltering, as well as carrying food to their shelters for later consumption rather
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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152 The Biology and Fisheries of the Slipper Lobster
than remaining exposed while feeding, slipper lobsters may fully minimize their exposure to diurnal
predators. Open bivalves have been found in shelters occupied by lobsters, as well as in a 40 m diameter
surrounding the occupied shelters. When such empty shells are removed by divers, new ones appear
in a manner of days (Spanier & Almog-Shtayer 1992). Horizontally oriented shelters supply shade and
reduce visual detection by diurnal predators. Small shelter openings also supply shade but, in addition,
increase physical protection against large diurnal predators, especially fish with high body profiles, such
as the gray triggerfish. Clinging may enable the lobsters to survive an attack inside a den and even in
open areas. Multiple shelter openings enable escape through a “back door” if a predator is successful
in penetrating the den. They can then escape by using their fast tail-flip swimming capability (Spanier
& Almog-Shtayer 1992). The tendency for cohabitation with conspecifics may be adaptive because of
collective “prey vigilance” and defense or concealment among cohorts (“selfish herd” response or the
“dilution effect” sensu Hamilton, 1971). If all else fails, their thick carapace, designed to effectively blunt
cracks, may serve them in times of exposure to attacking predators.
The function of sheltering as a predator-avoidance mechanism against diurnal fish was tested in a series
of field-tethering experiments on S. latus by Barshaw & Spanier (1994b). Tagged lobsters were tethered
with monofilament line inside and outside an artificial reef. All predation events occurred only during
the day. Predation by the gray triggerfish, Balistes carolinensis Gmelin, 1789, a high-body-profile, large,
diurnal fish, was significantly less on lobsters tethered in the reefs compared to those tethered in open
areas. The lobsters tried to cling to the substrate, relying on their armature and lack of movement to protect
them. Because the lobster could not tail flip to a shelter, the fish was eventually able to turn the lobster
over and consume it by biting through its thinner, vulnerable, ventral surface.
There have been some observations (in natural habitats) of specimens of S. latus sharing dens with the
Mediterranean moray eel, Muraena helena Linnaeus, 1758, with no apparent predator–prey interactions
between the fish and the lobsters (Spanier & Almog-Shtayer 1992; Martins, personal communication).
Sharing shelters with a moray eel may have mutual benefits: the octopus, Octopus vulgaris Cuvier, 1797,
is prey for the moray eel and may be a predator of the lobster (at least it is in laboratory settings; Spanier,
personal observations). Thus, by cohabiting, the lobster may be protected by a moray which preys on
octopus, and the moray eel may take advantage of any octopus attracted to the shared den by the presence
of its prey — the lobster (Spanier & Almog-Shtayer 1992). However, this association needs to be further
studied to elucidate any such mutualistic interaction.
Besides the grey triggerfish, dusky groupers (Epinephelus guaza Linnaeus, 1758) have been reported
as predators of adult and juvenile S. latus and combers (Serranus spp.) and rainbow wrasse (Coris julis
Linnaeus, 1758) apparently prey on juvenile S. latus (Martins 1985b). The spotted gully shark, Triakis
megalopterus (Smith, 1839), has been reported to feed on S. elisabethae in South Africa (Smale & Goosen
1999), and tiger sharks, Galeocerdo cuvieri (Peron & LeSueur, 1822), red grouper, Epinephelus morio
(Valenciennes, 1828), and gag grouper, Mycteroperca microlepis (Goode & Bean, 1879), have been repor-
ted as predators of S. nodifer (Lyons, 1970). Queentriggerfish, Balistes vetula Linnaeus, 1758, and Goliath
groupers or jewfish, Epinephelus itajara (Lichtenstein, 1822), are the main predators of S. aequinoctialis
(Lyons, 1970). Scorpionfish, Scorpaena brasiliensis Cuvier, 1829, dusky flounder, Syacium papillosum
(Linnaeus, 1758), high hat drum, Equetus acuminatus (Bloch & Schneider, 1801), and clearnose skate,
Raja eglanteria Bosc, 1802, feed on various Scyllarus species (S. americanus,S. chacei,S. depressus)
(Lyons, 1970). Unidentified scyllarids have also been found in the stomachs of various zooplankton feed-
ing fish (blackbar soldierfish, Myripristis jacobus Cuvier, 1829; bigeye, Priacanthus arenatus Cuvier,
1829; and yellowtail snapper, Ocyurus chrysurus (Bloch, 1791)); great barracuda, Sphyraena barracuda
(Walbaum, 1792); gray snapper, Lutjanus griseus (Linnaeus, 1758); dog snapper, L. jocu (Bloch &
Schneider, 1801); and black margate, Anisotremus surinamensis (Bloch, 1791) (Lyons 1970).
7.5.7 Movement Patterns
Slipper lobsters demonstrate two modes of movement: (1) slow, benthic walking movements that may be
nomadic within a small home range or migratory from inshore, shallower waters to offshore, deep waters;
and (2) swimming movements that can be used for rapid escape or, as some have suggested, for vertical
movements (swimming) in the water column.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 153
7.5.7.1 Daily and Seasonal Horizontal Patterns
Tagging studies of only a few species (Scyllarides astori,S. latus, and Thenus spp.) have been used not
only to determine growth and maturity indices, but also to determine movement patterns. These studies
demonstrate that some species remain within local grounds year-round, while other species demonstrate
two annual patterns of movement: local nomadic movements made while inshore and migratory offshore
movements. Such studies are summarized below.
In a study of S. latus, 314 lobsters caught at an artificial tire-reef complex were tagged between the car-
apace and the abdomen, using numbered spaghetti tags (Spanier et al. 1988). Lobsters were also marked
by puncturing small holes in the telson and were released at their site of capture. In a later phase of this
study, they were also tagged between the third and fourth abdominal segments. Thirty-two percent were
recaptured at least once, 9% were recaptured twice, and 2.6% were recaptured thrice. All but 3% of the
recaptured tagged lobsters retained their spaghetti tags. These remaining 3% were identified by the holes
punctured in their telsons or the scars left by these holes after molting. However, a later laboratory study of
tag retention in these lobsters (Spanier & Barshaw 1993) found that only 40% of the animals retained tags
positioned between the carapace and abdomen after molting, indicating that many individuals may not have
been properly reidentified if captured. During the inshore lobster season (February to June in the south-
eastern Mediterranean), lobsters left their shelters at night to make short-term movements to forage and
bring back food (mostly bivalves). More than 71% of tagged lobsters were recaptured in the artificial reef
site repeatedly during the season. Time between repeat captures was one to 17 weeks (mean 29 days) and
this rate probably represents short-term movements for foraging or local nomadism (Spanier et al. 1988).
In contrast, only 7.2% of the tagged lobsters were recaptured in the same man-made shelter site after
more than half a year. Time between these captures ranged between 10 and 37 months (mean 338 days) and
may represent long-term movements or migration. Returning lobsters have to orient and locate the small
artificial reef site in the widespread continental shelf. Due to limited cooperation with local fishermen,
only 11 tagged lobsters were reported outside the artificial shelter site. Six were caught by divers about
300 to 800 m off the reef site. The rest were caught by fishermen in the late part of the summer two to
three months after tagging, 20 to 35 km north of the site and at depths >50 m (Spanier et al. 1988).
For several years, a seasonal survey of all lobsters was conducted in the earlier-mentioned artificial
tire-reef complex (Spanier et al. 1988, 1990). Lobsters appeared in the reefs in early winter (December
to early February), with their numbers peaking in the spring (March to May). From June onward, their
numbers decreased in the shallow part of the continental shelf and they disappeared from shallow water in
August/September until the beginning of the following winter. This seasonality correlates with water tem-
perature. Lobsters appear in the shallow part of the continental shelf (15 to 30 m depth) when water
temperatures are the lowest for the southeastern Mediterranean region (15 to 16C), and their numbers
decrease when water temperatures rise to 26 to 27C (Spanier et al. 1988). A similar trend is seen in the
yield of the commercial fisheries off the Mediterranean coast of Israel.
Several traps (see Figure 18.1 for an example of the type) set offshore, off-season, and at a depth of
48 m caught lobsters in October. Water temperature at these trap sites was 23.6C, while in the much
shallower artificial reef site at 18.5 m where no lobsters were detected, it was 27.7C. Also lobsters were
caught during the fall at depths greater than 50 m by a rough bottom trawl (Spanier et al. 1988). This
limited information suggests that slipper lobsters off the coast of Israel seasonally move to deeper and
more northerly waters (i.e., colder water). By migrating, the colder-water lobsters may avoid the high, and
perhaps unfavorable, summer and autumn temperatures in the shallow waters of the Levant basin of the
Mediterranean. Today these temperatures may rise as high as 31C, which may cause molting difficulties
or abnormalities (also reported by Spanò et al., 2003). Some lobsters kept in the laboratory with ambient
water supply in the fall died while molting or only incompletely molted. This happened after they were
exposed for over two months to water temperatures of 26C and higher (Spanier, personal observations).
Thus, one possible function of the seasonal shallow-to-deep migration may be to meet physiological and
behavioral requirements for molting. Molting in deeper habitats may also be a predator-avoidance strategy
during this vulnerable period. Berry (1971) pointed out that while most spiny lobsters are associated with
hard substrates that supply shelters, some deep water species appear to exhibit behavioral adaptations for
soft substrates, perhaps because of fewer predators in greater depths. Slipper lobsters could switch to soft
bottoms at greater depths for the same purpose.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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154 The Biology and Fisheries of the Slipper Lobster
Of 115 S. latus tagged by Bianchini et al. (2001) in 1995 to 1997, 29 individuals were later recovered up
to 70 weeks after tagging. One tagged female was caught by a trammel net, after being 1575 days at large,
at 5 km from the place of release. Contrary to the findings of Spanier et al. (1988), the recurrence of
tagged specimens during every period of the year, at least in Sicily, reduces the possibility of widespread
seasonal horizontal migrations, although some specimens might displace vertically. However, surface
temperatures in Sicilian waters range from 14 to 23C and differ considerably from those in Israeli waters
(15 to 31C); thus, movement pattern differences may vary in a temperature-dependent fashion.
In a mark-recapture program in the Galápagos Islands, Hearn et al. (see Chapter 14 for more details)
reported that of a total of 1926 S. astori tagged and released back into the wild, 116 (6 %) were recaptured
and reported by the local fishermen. No information was reported on distances between release and capture
locations; thus, it is unknown whether S. astori migrates to deeper water to molt, or simply behaves in a
more cryptic fashion during this vulnerable period (as reported for S. latus by Spanier et al., 1988).
Via repetitive diving surveys of the same dens, it appears that S. aequinoctialis displays den fidelity
(Sharp et al., Chapter 11), but it is not clear if they remain in a single locale over long periods of time or
if they migrate in a manner similar to S. latus. It is also not clear how they are able to orient back to the
same shelter after foraging movements.
Jones (1988) and Courtney et al. (2001) conducted a rather ambitious tag and release program involving
>13,000 lobsters with recapture rates ranging from 7 to 10%. From these studies, the following picture of
Thenus spp. movements emerges: lobsters are capable of moving 24 km from point of release, but do not
do so in any particular direction, as would be expected if these movements represented migration. Their
movement pattern is clearly nomadic, which, given their preference for spatially simple, sedimentary
substrates, is logical, as there is no shelter to repetitively locate. It also appears that these lobsters do not
migrate, as Thenus orientalis sexes are not segregated at any time during the year, indicating that females
do not leave the fishing grounds for either shallower or deeper waters (Kagwade & Kabli 1996a).
Likewise, in a tagging program involving nearly 3900 Ibacus peronii, with recapture rates of 14.3%,
the movement pattern also appeared to be nondirectional nomadism, with lobsters being recaptured close
to their point of release. One cannot generalize, however, about Ibacus spp. because 94 (13.1%) of 557
tagged I. chacei demonstrated a northward migration from released locations and covered approximately
0.15 to 0.71 km/day for a total of traveling time of 310 km in 655 days (Stewart & Kennelly 1998; see
Haddy et al., Chapter 17 for more information).
For those lobsters that engage in a migration involving a return to a specific location (e.g., S. latus),
individuals seem to be capable of shoreward homing movements to arrival back at to their preferred location
(e.g., artificial reef site for S. latus in Israeli waters). The advantage of this homing ability is obvious.
Natural rocky outcrops that supply shelters with the physical parameters preferred by lobsters constitute
a very small portion of the shallow continental shelf along many coasts. Thus, it may be advantageous
for lobsters to “recall” these preferred natural sites or dens (or artificial sites/dens that would be found
in artificial reefs and ship wrecks; see Figure 18.3 in Chapter 18 for an example) and to return to them
after short- as well as long-term movements. It seems that they just walk or walk and swim relatively long
distances to return. The mechanism by which they orient and locate such preferred habitats is completely
unknown. They may use some geomagnetic cues or magnetic maps, as has been reported for spiny lobsters
(Lohman 1984, 1985; Boles & Lohman 2003). Lohman (1984, 1985) found magnetic remenance in the
Caribbean spiny lobster, Panulirus argus, along with the ability to detect geomagnetic fields, while Boles
& Lohman (2003) demonstrated that spiny lobsters could orient homeward and navigate without any cues
from their outbound, displacement trips. Some species of slipper lobsters may have similar abilities.
7.5.7.2 Swimming Behavior (Vertical Movements)
In mechanical terms, tail-flip swimming in crustaceans constitutes locomotion in which a single “append-
age” — the abdomen — produces thrust by a combination of a rowing action and a final “squeeze”
force when the abdomen presses against the cephalothorax (Neil & Ansell 1995). Although the tail-flip
response is known in adults and juveniles of all three major taxonomic group of lobsters (e.g., Ritz &
Thomas 1973; Jones 1988; Newland & Neil 1990; Newland et al. 1992; Jacklyn & Ritz 1986; Jackson &
Macmillan 2000; Jeffs & Holland 2000), as well as in other crustaceans (e.g., crayfish, shrimp, squat
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 155
lobsters, stomatopods), it is best developed in slipper lobsters. Tail flipping is first developed in the nisto
phase, where it can vary among species in strength (Robertson 1968; Lyons 1970; Barnett et al. 1986;
Higa & Saisho 1983).
The hydrodynamics of swimming in slipper lobsters has been well studied in Scyllarides latus (Spanier
et al. 1991; Spanier & Weihs 1992, 2004), Ibacus peronii and I. alticrenatus (Jacklyn & Ritz 1986; Faulkes
2004), and Thenus orientalis (Jacklyn & Ritz 1986; Jones 1988). In only one of these species (Ibacus
peronii) has the neural circuitry controlling the tail flip been studied (Faulkes 2004). Faulkes (2004) argues
that the external morphology of the more flattened species of scyllarids, combined with their proclivity
for holding their abdomen in a flexed position, is likely to have resulted in the loss of giant interneurons
that are involved in the control circuitry of escape behavior. While the neuroanatomy in Ibacus peronii
supports this idea to the extent that these neurons appear absent and a similar argument has been made
for hermit crabs and galathean crabs (squat lobsters) (Faulkes & Paul 1997, 1998) this idea remains to
be more fully examined at the behavioral and neuroanatomical levels in the Ibacus spp., as well as in the
allied genera of Parribacus,Scyllarides, and Thenus.
Scyllarides latus uses a “burst-and-coast” type of swimming (see Weihs 1974) in response to a predator
(e.g., triggerfish) or harassment from divers. Large-amplitude movements of the tail propel the lobster
quickly backward, with periods of acceleration reaching top velocities of three body lengths per sec; these
movements are followed by periods of powerless gliding, decelerating to velocities of less than one body
length per sec (Figure 7.2). The force per tail beat ranges between 1.25 to more than 3.00 N and correlates
with body length because additional force is needed to move the greater mass of larger animals, rather
than to increase speed and acceleration. The intermittent fast-escape swimming is only of short duration
and does not appear to be used for foraging or long-range movements; instead it is an emergency response,
whereby the animal invests considerable energy resources to reduce its exposure time in the open area
until it can reach safety.
Spanier & Weihs (1992) suggested that the flattened second antennae of S. latus (mistakenly called
“shovels” or “flippers”) along with their movablejoints, play an important hydrodynamic role in controlling
the swimming movement. Essentially, they serve as stabilizers and rudders in “take off,” acceleration,
gliding, turning, and landing. Significant lift is created during backward tail flips, and articulation of
the flattened second antennae (a.k.a. “rudders”) alters the distributions of this lift so that pitching and
rolling movements are possible. Essentially the flattened antennae are articulated hydrofoils that move in
different horizontal and vertical planes; they can form continuous or separated surfaces via an overlapping
or spreading out of the component segments (Spanier 2004; Spanier & Weihs 2004). When spreading,
the flattened antennae can increase surface area by 56.2 to 79.5%. Exact movements and resultant forces
T
T
FIGURE 7.2 Drawing of a swimming sequence of the Mediterranean slipper lobster, Scyllarides latus, based on video
recordings; top view: above, side view: below (reprinted from Spanier, E., Weihs, D., & Almog-Shtayer, G. 1991. J. Exp.
Mar. Biol. Ecol. 145: 15–31, with permission from Elsevier).
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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156 The Biology and Fisheries of the Slipper Lobster
A
M
D
C.G.
T
50–60°
FIGURE 7.3 Forces and counterforces produced during take off in swimming behavior of Scyllarides latus. The abdominal
flexion results in a propulsive force (T) that produces a head-up (A) movement due to a misalignment in the animal’s center
of gravity (C.G.). To compensate, the lobster deflects the 2nd antennae 50 to 60upward from the body axis. This produces a
drag force (D) that results in a counterbalancing head-down movement (M). (Drawing based on videorecordings by R. Pollak;
used with permission.)
are described via videorecording analysis of take off, swimming, and turning. In take off, propulsive
abdominal flexion produces a head-up moment due to a misalignment in the lobster’s center of gravity. To
compensate for this force and its misalignment, the lobster deflects its flattened antennae 50 to 60upward
from the body axis (known as “rotation” in aircraft); this, in turn, produces a drag force that results in a
counterbalancing head-down movement (Figure 7.3).
This alteration in lift via the second antennae is also seen in Ibacus spp. and Thenus spp. (Jacklyn &
Ritz 1986). In contrast, spiny lobsters found in similar habitats and ranges (e.g., Jasus edwardsii and
J. novaehollandiae Holthuis, 1963), produce a negligible amount of lift during each tail flip and do not
possess antennae that were shaped or positioned properly to control any created lift (Jacklyn & Ritz
1986). As a result, spiny lobster tail flips are not efficient for continuous swimming or maneuvering
(Ritz & Jacklyn 1985; Jacklyn & Ritz 1986).
Neil & Ansell (1995) compared data of swimming performance for a number of decapod, mysid, and
euphausid crustaceans. Maximum velocity of body movement achieved during the tail flip was similar
across the adults in each group, and ranged from 10 to 300 mm body lengths, although this represented
a 30-fold difference in the velocities expressed as body lengths per second. Scyllarides latus was ranked
as the fastest of those tested, with a maximum velocity of close to 1 m/sec compared to 0.6 m/sec in the
clawed lobster Nephrops norvegicus (Linnaeus, 1758) (Newland et al. 1988). In a more recent analysis,
Spanier & Weihs (2004) identified the contribution of the tail as the propulsor, the legs as landing gear, and
the second antennae as control surfaces. They also examined secondary hydrodynamic effects of carapace
curvature and the longitudinal ridge associated with vortex production and control. A possible function
has been postulated for this ventrolateral curvature (keel) of the lobster carapace. It may be similar to the
scale armature of rigid-body boxfishes. In a detailed study of these fish, Bartol et al. (2002) found that the
ventral keels produce vortices that serve to stabilize motion, resulting in a smooth swimming trajectory.
This could be the case for slipper lobsters as well, and might explain why certain species of scyllarids have
a more pronounced keel than others. Furthermore, while in the coast portion of swimming, deflection of
the second antennae would increase drag; thus, the lobster returns its second antennae (and antennules)
to a position that is in line with the body, and shifts its legs forward in an effort to minimize drag. With
the advent of another abdominal flexion (to create acceleration), the lobster must repeat the deflection
of the second antennae, but does so at a smaller deflecting angle of 10 to 30. Turning is achieved by
differentially tilting and spreading the second antennae (Figure 7.4).
Ibacus peronii and I. alticrenatus have been examined both for the hydrodynamics of swimming as well
as for the neural circuitry controlling swimming. In their description of the tail-flip response, Jacklyn &
Ritz (1986) argued that because the abdomen was fully extended before being flexed, and because some
flexions occurred with only a partial extended abdomen, these flexions must be mediated by the lateral
giant (LG) interneurons. Typically the decapod startle response involves a circuit in which medial giant
(MG) interneurons and LG interneurons trigger a single, short-latency, powerful abdominal flexion; fast
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 157
FIGURE 7.4 An example of differential tilting and spreading of the 2nd antennae by Scyllarides latus to produce turning
movements during swimming. (Drawing based on videorecordings by R. Pollak; used with permission.)
flexor motor giant neurons (MoG) receive synaptic input from the LG and MG interneurons and control
the exact flexor response of the abdominal musculature (Wine & Krasne 1972; Mittenthal & Wine 1978).
The MG interneurons fire when stimuli come from an anterior direction and their firing results in a direct,
backward movement. In contrast, the LG interneurons fire when the stimulus is at the posterior end of the
animal and cause the abdomen to lift off the surface, dragging the animal into the water column (Wine &
Krasne 1972; Cooke & Macmillan 1985; Newland & Neil 1990). However, swimming by abdominal
extensions and flexions can be mediated by nongiant neurons (Reichert et al. 1981; Reichert & Wine
1983; Faulkes 2004).
Ibacus spp. do not respond to sudden tactile stimuli with tail flips — they simply flatten their eyestalks
into their eye cups; however, attempts to turn them over result in swimming via repeated tail flips (Faulkes
2004). Histological analysis of the abdominal ventral nerve cord showed that I. peronii lacked both MG
and LG interneurons, as well as the MoG neurons. This result is surprising because these neurons modulate
the fast-escape response in crayfishes, spiny lobsters, and clawed lobsters and have been the subject of
considerable research (Wine & Krasne 1982). In the place of these highly conserved control circuits,
I. peronii has fast flexor neurons and flexor inhibitor motor neurons that appear to mediate the tail-flip
response. Ibacus alticrenatus, although not as well studied as I. peronii, also apparently lacks the tail-
flip escape circuitry (Faulkes 2004). Faulkes (2004) argues that the adaptive value of the giant-mediated
escape tail-flip response should be inversely proportional to the amount of time the abdomen is flexed
and, because it is flexed normally in Ibacus spp., such a tail flip response would only produce minimal
movement and thus be rather ineffective as an escape mechanism. While the paucity of data currently do
not permit a thorough evaluation of this provocative idea, it is another example of the degree to which the
specializations of slipper lobsters offer valuable avenues for basic research.
Jones (1988) distinguished two types of swimming activity in Thenus species: locomotion or free
swimming and escape swimming, based on the presence/absence of an overt stimulus and the speed of
the swimming response (29 cm/sec for the first type; 1 m/sec for the latter). In free swimming, lift is
generated by the body shape (aerofoil) and by the downward thrust of the abdomen, while drag is reduced
by all pereiopods being extended anteriorly (Jacklyn & Ritz 1986). Height is controlled by the second
antennae and each flexion helps maintain the animal above the sediment (Jacklyn & Ritz 1986; Jones
1988). In contrast, escape swimming was always effected after a direct stimulus or threat was applied
and consisted of an abdominal flexion that was proportional to the magnitude of the stimulus (see Jones,
Chapter 16 for more information). Faulkes (2004) argues that these two separate responses need not be
the result of separate and distinct neural circuitry, but could simply represent two extremes of the nongiant
flexor motor neuron effected tail flips.
7.6 Sensory Biology of Scyllarids
The sensory world of the slipper lobster unfortunately has received little attention when compared to
that world for nephropid and palinurid lobsters, as well as other decapods such as crayfish and certain
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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158 The Biology and Fisheries of the Slipper Lobster
species of crabs. Extrapolating from morphological homologies with these other genera, it is likely that
chemical and mechanical stimuli will be of comparable central importance to the natural history of slipper
lobsters. However, the one likely exception is that vision may make a greater contribution to the umveldt
of many species within this family that live in shallow waters. Like other lobsters, slipper lobsters have
many appendages (cephalic, thoracic, abdominal) that appear to bear many different types of setae which
may or may not have a sensory function. Unlike other lobsters, many species of slipper lobsters have been
described as having a fine “pubescence” covering their cephalothorax that may provide those species with
additional sensory information. Few studies have actually examined the sensory function of appendages
or the setae borne upon such appendages; therefore, this section will describe what is known about the
different sensory modalities that are likely to be used by slipper lobsters. Also, where possible, this section
will describe potential appendages and setae that, by likely morphological homology, can reasonably be
expected to serve as primary receptors for sensory modalities typical in other lobsters. It is our hope that
by laying out potential functions for appendages and the setae borne upon them, future research will be
directed toward a systematic study of neurophysiological responses of those appendages and setae to a
variety of sensory stimuli. With this purpose in mind, Table 7.2 provides a summary of appendages found
in slipper lobsters for all life-history stages described, setae borne upon them when identified by various
researchers, purported sensory functions, and associated references.
7.6.1 Chemoreception
Like many other animals, lobsters use chemoreception to orient to and locate food and conspecifics.
Extensive research in the nephropid and palinurid families of lobsters, has distinguished “distance”
chemoreception (smell) and “contact” chemoreception (or taste) on morphological, neurophysiological
and behavioral grounds (Atema 1977, 1980; Schmidt & Ache 1992, Voigt & Atema 1994, Atema & Voigt
1995). In these lobsters, chemo- and mechanoreceptors on the antennules guide the animal to the vicinity
of odor sources; those on the distal ends of the walking legs are used in local probing searches of the
substrate, and those of the mouthparts evaluate the “palatability” of the objects lifted by the legs. Descrip-
tions of feeding behavior in various species of Scyllarides (Lau 1987; Malcom 2003) and Thenus (Jones
1988), and differences in morphology of the feeding appendages make the utility of these distinctions
questionable for slipper lobsters.
In nephropids and palinurids, the antennules are thought to be essential for distance chemoreception
and help the animal orient to a food, conspecific, or substrate source (Reeder & Ache 1980; Devine &
Atema 1982; Moore et al. 1991b; Boudreau et al. 1993; Basil & Atema 1994; Beglane et al. 1997; Koehl
et al. 2001; Goldman & Patek 2002; Koehl 2006). A key feature of antennule morphology subserving this
function is the ability to reach beyond the fluid-boundary layer surrounding the animal’s own body into
the potentially odor-bearing free stream of the ambient current. The long antennules of clawed and spiny
lobsters are well suited to this function, but the short antennules of slipper lobsters are not (Grasso & Basil
2002). Scyllarides aequinoctialis,S. notifer, and S.latus performed poorly compared to H. americanus
and P. argus when challenged to locate food sources under flow conditions identical to those under which
the latter easily tracked free-stream borne food odors over distances of two meters (Grasso, unpublished
observations). In slipper lobsters, antennules obviously play a role in distance chemoreception as part of
the detection process, but based on morphological considerations and behavioral observations, the sensing
strategy is clearly different.
Once the (clawed or spiny) lobster reaches the proximity of an odor source, the pereiopods take over
the decision-making and act as “near-field” and contact chemoreceptors. This function is mediated by
numerous chemo- and mechanosensory setae covering their broad, blunt dactyls. Such setae are largely
absent from the dactyls of slipper lobsters (discussed later) and, in their place, are epicuticular caps
tapering to sharp points — well adapted to the mechanical task of shucking bivalve shells, but not for
chemosensing. In spiny and clawed lobsters, the antennules have little to no participation in contact
chemoreception which the pereiopods are morphologically specialized to serve.
When searching for buried food items, it is unclear how slipper lobsters detect such items, as they
lack setal tufts on the tips of their pereiopod dactyl segments. Nonetheless, they are capable of digging
bivalves out from 3.5 cm of sediment (Almog-Shtayer 1988), and increase flicking rates of the antennules
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Behavior and Sensory Biology 159
TABLE 7.2
Possible Behavioral and Sensory Functions of Scyllarid Appendages
Appendage Possible Function References
Antennules (=1st antennae)
Lateral flagellum Olfaction (tracking odor plumes)
aesthetascs present in two rows per segment
Chemical food recognition (observed to touch surface and interior of
bivalve shells) or debris recognition
have toothbrush setae (similar to hooded setae in palinurids)
have asymmetric setae (similar to those seen in palinurids) that
initiate grooming in palinurids in response to specific chemicals
Mechanoreception (for current information)
simple setae present as “guard hairs” and form a water channel
have asymmetric setae — in palinurids, these are innervated by both
mechano- and chemosensory neurons
Odor-plume tracking not observed, but orientation to food
source observed by Jones (1988) and increased antennule
flicking observed by Jones (1988) & Lau (1987)
scyllarids: Weisbaum & Lavalli (2004)
Lau (1987, 1988); Jones (1988); Malcom (2003)
scyllarids: Weisbaum & Lavalli (2004)
palinurids: Cate & Derby (2002), Schmidt et al. (2003);
Schmidt & Derby (2005)
No neurophysiological studies done on scyllarids
scyllarids: Weisbaum & Lavalli (2004)
palinurids: Schmidt et al. (2003); Schmidt & Derby (2005)
nephropids: Guenther & Atema (1998)
Medial flagellum Unknown; chemoreceptive in nephropids Tierney et al. (1988)
Basal segment Unknown; location of statocyst in nephropids Cohen (1955, 1960)
Antennae (=2nd antennae)
Flagella Rudders and stabilizers (articulated hydrofoils) used while swimming
Flagellar segments have great range of movement
Mechanoreception (for water flow information)
Setae present on most edges; unknown types
Spanier & Weihs (1992, 2004); Spanier (2004)
See various figures in Holthuis (1985, 1991, 2002) for setal
arrangements
Bases Unknown; location of nephropore in nephropids Bushmann & Atema (1994)
Mandibles
Endopod Mastication of food; grasping, shearing, and ripping of food; in some species
can be used to chip edges of bivalve shells
Larvae: divided into three portions:
Upper portion =blunt, multipronged, canine-like process
Middle portion =sharp incisor teeth along inner, medial edge
Lower portion =flattened, tuberculate process
Adults: asymmetric; left mandible larger with calcified molar process
that extends over right mandible; both left and right mandibles have a
serrated, tooth-like, incisor process ventromedially
Lau (1987)
Mikami et al. (1994)
Suthers & Anderson (1981); Johnston & Alexander (1999)
(Continued)
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160 The Biology and Fisheries of the Slipper Lobster
TABLE 7.2
(Continued)
Appendage Possible Function References
Palp Unknown; presumably aids in directing food into mouth
Adult: noncalcified; bears densely arranged pappose setae along its
upper margins (setal barriers?)
Suthers & Anderson (1981); Johnston & Alexander (1999)
First maxillae
Endopod
[Biramous (distal
endite + proximal
endite)]
Unknown; presumably involved in food handling and grooming based on setal
types present
Larvae: bears “spines” and setae, the number of which varies with larval
instar and species; some species are described as having “masticatory”
spines (cuspidate setae?)
Adult: distal endite large and covered with simple, stout setae; proximal
endite small with two multidenticulate, cuneate setae projecting
terminally
Robertson (1968); Johnson (1971b); Barnett et al. (1986); Ito & Lucas
(1990); Mikami & Greenwood (1997); Webber & Booth (2001)
Johnston & Alexander (1999)
Exopod No information
Second maxillae
Endopod Unknown; presumably used to handle food; has potential
chemo-mechanosensory setae (simple) as well as setae thought to seal or act
as barrier (pappose)
Larvae: paddle-shaped lobe with several plumose setae that gradually
expands distally, loses the plumose setae, and becomes flatter with
anterior lobes
Adults: distal and proximal endites fused; inner margin of distal endite
covered with pappose setae; inner margin of proximal endite covered
with simple setae
Barnett et al. (1984); Phillips & McWilliam (1986); Ito & Lucas (1990);
Mikami & Greenwood (1997); Webber & Booth (2001)
Johnston & Alexander (1999)
Exopod
(Scaphognathite)
Gill bailer; drives gill current
Larvae: develops slowly as a posterior lobe
Adults: fringed with plumose setae; aboral surface covered with simple
setae; oral surface covered with tapered setae
Robertson (1968); Ito & Lucas (1990); Webber & Booth (2001)
Johnston & Alexander (1999)
First maxillipeds
Endopod Unknown; presumably used to handle food; has potential
chemo-mechanosensory setae (simple), mechanosensory setae (plumose);
and setae thought to act as a barrier (pappose)
Larvae: not present or simply a rudimentary bud in early instars of some
species
Adults: distal and proximal endites reduced and fused to exopod; lined
with simple, pappose, and plumose setae
Robertson (1968); Johnson (1970, 1971b); Ito & Lucas (1990)
Suthers & Anderson (1981); Johnston & Alexander (1999)
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Behavior and Sensory Biology 161
Exopod Unknown; fusion with endopod probably limits use as a current generating
structure
Adults: terminal flagellum in I. peronii, but not in T. orientalis; lined
with simple, pappose, and plumose setaeaSuthers & Anderson (1981); Johnston & Alexander (1999)
Second maxillipeds
Endopod
(5 segments)
Unknown; presumably used to handle food; has setae that are likely
chemo-mechanosensory
Larvae: spines and setae on distal segments only (dactyl, propus, carpus)
Adults: fused basis and coxa; reduced ischium; carpus and propus have
short, simple, stout setae on upper margins; dactyl bears simple, stout
setae (hooklike) along edges
Barnett et al. (1984); Webber & Booth (2001)
Suthers & Anderson (1981); Johnston & Alexander (1999)
Exopod Unknown; lacks a flagellum, so cannot generate or redirect currentsaSuthers & Anderson (1981); Johnston & Alexander (1999)
Third maxillipeds
Endopod
(5 segments)
Presumably used to recognize food, to tear food, to groom antennules; has
setae for these purposes and crista dentate for maceration of food
Larvae: setae distributed on all segments, but most numerous on
distal-most segments (dactyl, propus)
Adults: ischium has only small, blunt, crista dentate; merus and carpus
have numerous setae covering the oral surface; inner margin of merus
bears multidenticulate setae; dactyl has simple, stout setae covering
entire oral and portions of aboral surface
Barnett et al. (1984); Webber & Booth (2001)
Suthers & Anderson (1981); Johnston & Alexander (1999)
Exopod Unknown; may be involved in current generation in some species
Larvae: lacking in early larvae; bud present in later larval stages
Adults: reduced and lacking flagellum in T. orientalis; elongate with
flagellum in I. peroniia
Webber & Booth (2001)
Suthers & Anderson (1981); Johnston & Alexander (1999)
Pereiopods
First pereiopods Used to grasp food in larvae and to “wedge” open bivalves in adults; have setal
types that are likely chemo- and mechanosensory; in larval instars, exopods
are used as swimming organs and then lost in metamorphic molt to nisto
(recognizable as vestiges in some species)
Larvae: well developed in early instars with exopods bearing paired
natatory setae that increase in number in successive instars; curving
subexopodal spines present in latter instars — these become fine, short
setae; dactyl is claw-like
Adults: large and thick; right and left are symmetrical; dactyl bears
cuspidate and simple setae in rows on both oral and aboral surfaces;
propus, carpus, and ischium bear cuspidate setae in rows (longer on
aboral surface; shorter on oral surface); some species have conate setae
on merus, others have teasel-like or paintbrush setae
Robertson (1971); Lau (1987); Ito & Lucas (1990)
Robertson (1968, 1971); Johnson (1971b); Phillips & McWilliam
(1986); Ito & Lucas (1990)
Malcom (2003)
(Continued)
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162 The Biology and Fisheries of the Slipper Lobster
TABLE 7.2
(Continued)
Appendage Possible Function References
Second pereiopods Used to grasp food in larvae and to wedge open bivalves in adults; have setal
types that are likely chemo- and mechanosensory; in larval instars, exopods
are used as swimming organs and then lost in metamorphic molt to nisto
(recognizable as vestiges in some species)
Larvae: well developed in early instars with exopods bearing paired
natatory setae that increase in number in successive instars; curving
subexopodal spines present in latter instars — these become short, fine
setae; dactyl is claw-like
Adults: longest and slimmest leg; right and left are symmetrical; dactyl
bears cuspidate and simple setae in rows on both oral and aboral
surfaces; propus, carpus, and ischium bear cuspidate setae in rows
(longer on aboral surface; shorter on oral surface); some species have
conate setae on merus, others have teasel-like or paintbrush setae and
cuspidate setae
Robertson (1971); Lau (1987); Ito & Lucas (1990)
Robertson (1968, 1971); Johnson (1971b); Phillips & McWilliam
(1986); Ito & Lucas (1990)
Malcom (2003)
Third pereiopods Used in walking gait; probing of substrate, and digging sequence in adults; in
larval instars, probably used to grasp food; exopods are used as swimming
organs and then lost in metamorphic molt to nisto (recognizable as vestiges in
some species)
Larvae: in first instars of some species, less well developed than legs one
and two with only an exopodal bud; curving subexopodal spines present
in latter instars; dactyl is claw-like
Adults: shorter in length but wider; two tufts of cuspidate and simple
setal on dactyl, with rows of cuspidate and simple setae present on oral
and aboral surface; propus, carpus, merus, and ischium bear rows of
cuspidate setae that are longer on the aboral surface and shorter on the
oral surface; conate setae on portions of merus in some species,
cuspidate or paintbrush setae in other species
Robertson (1971); Jones (1988); Ito & Lucas (1990); Faulkes (2004)
Robertson (1968, 1971); Johnson (1971b); Phillips & McWilliam
(1986); Ito & Lucas (1990)
Malcom (2003)
Fourth pereiopods Used in walking gait; probing of substrate, and digging sequence; in larval
instars, probably used to grasp food; exopods are used as swimming organs
and then lost in metamorphic molt to nisto (recognizable as vestiges in some
species)
Larvae: rudimentary or absent in earliest instars of some species;
develops exopod with natatory setae; dactyl is claw-like
Adults: smaller than 3rd leg; two tufts of cuspidate and simple setal on
dactyl, with rows of cuspidate and simple setae present on oral and
aboral surface; propus, carpus, merus, and ischium bear rows of
cuspidate setae that are longer on the aboral surface and shorter on the
oral surface; conate setae on portions of merus in some species,
cuspidate or paintbrush setae in other species
Robertson (1971); Jones (1988); Ito & Lucas (1990); Faulkes (2004)
Robertson (1968); Johnson (1971b); Phillips & McWilliam (1986); Ito
& Lucas (1990)
Malcom (2003)
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Behavior and Sensory Biology 163
Fifth pereiopods Grooming of abdomen (and eggs in females); chemoreception(?); walking;
probing the substrate; exopods are used as swimming organs and then lost in
metamorphic molt to nisto (recognizable as vestiges in some species)
Larvae: rudimentary in earliest instars of some species with no trace of
an exopod; in later instars, leg may be present but is initially nonsetose
and then develops with exopod and natatory setae
Adults: sexual dimorphism presentb
Male: leg is achelate; two tufts of cuspidate and simple setal on
dactyl, with rows of cuspidate and simple setae present on oral and
aboral surface; propus, carpus, merus, and ischium bear rows of
cuspidate setae that are longer on the aboral surface and shorter on
the oral surface
Female: leg is chelate or subchelate and larger than that in males;
distal tip of dactyl and of propus articulate and bear a structure
(“brush pad”) set into a semicircular groove and made of long
cuspidate and simple setae in some species and of only long
cuspidate setae in others; this structure is flanked by simple setae at
the proximal end rows of cuspidate and simple setae present on oral
and aboral surface; propus, carpus, merus, and ischium bear rows of
cuspidate setae that are longer on the aboral surface and shorter on
the oral surface
Robertson (1971); Jones (1988); Ito & Lucas (1990); Faulkes (2004)
Robertson (1968); Johnson (1971b); Ritz & Thomas (1973); Berry
(1974); Ito & Lucas (1990)
Malcom (2003)
Pleopods Swimming current generation in forward swimming (nistos); bear plumose
setae along fringes
Robertson (1971)
Uropods Unknown; used to cover ventral abdomen in some species Faulkes (2004)
Telson Unknown; used to cover ventral abdomen in some species Faulkes (2004)
Information on appendages is based on a variety of scyllarid species: (1) antennules =Scyllarides species (S. aequinoctialis,S. latus,&S. nodifer); (2) second antennae =Scyllarides latus; (3) mouthparts,
larvae =a variety of species, see references for specifics; (4) mouthparts, adults =Ibacus peronii &Thenus orientalis; (5) pereiopods =Scyllarides species (S. aequinoctialis,S. latus,&S. nodifer).
Refer to Johnston, Chapter 6 for detailed descriptions and figures of mouthparts and Jones, Chapter 16 for detailed descriptions of feeding behavior and use of appendages in Thenus spp.
aExopods or flagella of exopods of the maxillipeds are missing in the Theninae and Scyllarinae, but present in the Arctidinae and Ibacinae.
bNot clear if sexual dimorphism exists for all species, because some species descriptions are based on males alone (e.g., Holthuis 1993; Tavares 1997).
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164 The Biology and Fisheries of the Slipper Lobster
while doing so, or when food items are deposited in their holding tanks (Jones 1988; Lavalli, personal
observations). Jones (1988) reports that Thenus spp. probe the substrate with their first two pairs of
pereiopods, while at the same time raising and lowering the antennules above the substrate surface as
described by Lau (1987). Given that epicuticular caps cover the distal portion of the dactyls, it is unclear
how the pereiopods detect buried food, and more observations are needed to elucidate food detection
mechanisms of scyllarids.
If feeding on bivalves, leg motions are suspended several times during the manipulation and initial prob-
ing phases while the lobster brings its distance chemoreceptor organs (antennules) beneath the carapace
and into contact with the bivalve (Malcom 2003). This is surprising for two reasons. First, the chemo-
sensory maxillipeds are better positioned to reach the shell. Second, as mentioned above, the tactile and
proprioceptive sense from the legs should be sufficient to provide feedback information during opening.
But, because slipper lobsters are unique in lacking distally placed dactyl setal tufts, they may not be able
to obtain sufficient sensory feedback without using the antennules.
The above makes clear that chemo-orientation in slipper lobsters deviates from the model that emerged
from extensive studies of food-searching behavior in clawed and spiny lobsters. Compared against this
model, three interlocking hypotheses are suggested: (1) periopod chemosensory ability has been replaced
by epicuticular caps suitable for shucking bivalves; (2) the antennules have, at least partially, filled this
sensory void and perform the functional role of the dactyls in clawed and spiny lobsters; (3) the distance
chemoreceptive function of the antennules has been reduced to accommodate the added role of contact
chemoreception. Despite these drastic modifications, the antennules continue to be used to examine
the odor source directly, in contact chemoreception, thereby seemingly blurring the distinction between
“smell” and “taste.” Given that the chemosensory areas of the brains of slipper lobsters do not differ
dramatically in architecture or volume from those of spiny or clawed lobsters (Sandeman et al. 1993;
Sandeman 1999), studies of chemoreception focused on these hypotheses are likely to put the smell and
taste distinction into context and reveal unexpected insights into the organization of the crustacean brain.
The annuli of the lateral flagella of the antennules are covered with a variety of different types of
setae (Figure 7.5H and Figure 7.5I); many of these have already been identified as chemoreceptors
and mechanoreceptors in clawed and spiny lobsters. The positioning of the different types of setae is
highly organized and appears not to differ significantly among the several Scyllarides species examined
by Weisbaum & Lavalli (2004): aesthetascs, asymmetric, modified simple, and hemiplumose setae
(Figure 7.5H) are only found in the tuft region between the distal and proximal ends of the lateral
flagellum. Simple setae are found on all regions of all annuli of the lateral flagellum, and toothbrush
setae (Figure 7.5I) are concentrated on the annuli of the base region or on the proximal annuli of the
tuft region (Weisbaum & Lavalli 2004). This positioning of the setae is ideal as they occur where stimuli
carried by currents, possibly generated by the exopodites of the mouthparts, can reach them and provide
an assessment of the shell and flesh within or where bottom currents can provide information about distant
sources. It is likely that aesthetascs on the slipper lobster antennule serve the same chemosensory function
that has been demonstrated for nephropid and palinurid lobsters (Spencer 1986; Derby & Atema 1988;
Michel et al. 1991; Ache & Zhainazarov 1995; Cate et al. 1999; Steullet et al. 2000a), that simple setae are
bimodal (chemo- and mechanosensory) as they are in palinurids (Cate & Derby 2001), that asymmetric
setae are mechanosensory as they are in blue crabs (Gleeson 1982) or bimodal (chemo- and mechano-
sensory) as they are in palinurids (Schmidt et al. 2003; Schmidt & Derby 2005), and that toothbrush setae
are bimodal chemo-mechanosensory homologues of hooded sensilla in palinurids (Cate & Derby 2002).
The asymmetric setae apparently are necessary to elicit antennular grooming in palinurids, but, thus far,
no studies on antennular grooming in scyllarids have been conducted.
The sensory-motor mechanisms that slipper lobsters use in shucking are unknown at this time. Visual
cues can be excluded because the shucking process takes place beneath the animal and is outside the field
of view of its dorsally placed eyes, except, possibly, in Thenus spp. in which the eyes are at the lateral-
most edges of the carapace. However, Jones (1988) tested the response of Thenus spp. to the introduction
of their favorite food item and found that visual cues alone did not elicit searching responses — only
cues with a chemical signal did. Touch, proprioceptive, and chemosensory modalities are the most likely
candidates and there is evidence to suggest that each is involved. The touch and proprioceptive senses that
could control this task are located in the pereiopods. For manipulation, the positions of each pereiopod
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Behavior and Sensory Biology 165
AB
CD
EFG
HI
Ms
Hp
sSc
TSc
FIGURE 7.5 Setal types found on scyllarid pereiopods and antennules. (A) dactyl tip of left 2nd pereiopod of Scyllarides
aequinoctialis; (B) simple setae set in groove on oral surface of left 5th pereiopod propus of S. nodifer; (C) cuspidate seta
of right oral surface of 3rd pereiopod propus of S. aequinoctialis; (D) teasel-like seta with setules from left 1st pereiopod,
aboral surface of S. aequinoctialis; (E) cuspidate and simple setae of female’s right 5th pereiopod dactyl, oral surface of
S. aequinoctialis; (F) conate setae on merus shield of 3rd right pereiopod, oral surface of S. nodifer; (G) miniature simple-
type setae covering the cuticular surface of the aboral surface of propus of S. aequinoctialis; (H) aesthetasc (arrow), modified
simple setae (Ms), and hemiplumose setae (Hp) found on the ventral surface of the antennular flagellum of Scyllarides species;
(I) toothbrush setae found on the dorsal surface of the antennular flagellum of Scyllarides species showing scale (Sc), setules
(S), and textured scaling on flagellar surface (TSc). Scale bars: A =200 µm; B =200 µm; C =200 µm; D =20 µm;
E=50 µm; F =250 µm; G =350 µm (box =20 µm); H =273 µm; I =20 µm. (Modified from Malcom, C. 2003.
Description of the setae on the pereiopods of the Mediterranean Slipper Lobster, Scyllarides latus, the Ridged Slipper Lobster,
S. nodifer, and the Spanish Slipper, S. aequinoctialis. M.Sc. thesis: Texas State University at San Marcos, San Marcos, Texas.
(A–F); Weisbaum & Lavalli 2004 (G,H); used with permission.)
segment, as well as each pereiopod in its entirety, informed by proprioception, are likely to signal the
size, orientation, and location of the bivalve shell through signaling of joint angles. Points of contact for
application of manipulative forces may be sensed through tactile sensors on the dactyl tips or through the
tension in muscle organs. The pereiopods of Scyllarides species do bear numerous tufts of setae (Malcom
2003), but most are placed on the more proximal segments of the propus, carpus, merus, and ischium,
rather than the distal dactyl (see Figure 7.5A, Figure 7.6A to Figure 7.6D). Those that are present on the
proximal edge of the dactyl segment are usually damaged, most likely due to the abrasive action they suffer
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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166 The Biology and Fisheries of the Slipper Lobster
S
C
P
D
I
M
S
I
D
M
P
C
C
P
MI
D
S
C
P
M
S
ID
H1, J1
H1, J1
J
J
I
J1
J1
J1
JI
J1
JI
H1, J1
JI
J1
J1
JI
JI
J1
J1
J1
H1, J1
r
r
r
(A) (B)
(C) (D)
r
J1J1
J1
r
rJ1
J1
J1
J1
J1
J1
J1
FIGURE 7.6 Outer, or aboral, views of walking legs of Scyllarides spp. (A) left 1st pereiopod; (B) left 2nd pereiopod;
(C) right 3rd pereiopod; (D) right 4th pereiopod. D — dactyl; P — propus; C — carpus; M — merus; I — ischium; S — meral
shield; J1— cuspidate setae; H1— simple setae, — stripped dactyl setal pit. (Modified from Malcom, C. 2003. Description
of the setae on the pereiopods of the Mediterranean Slipper Lobster, Scyllarides latus, the Ridged Slipper Lobster, S. nodifer,
and the Spanish Slipper, S. aequinoctialis. M.Sc. thesis: Texas State University at San Marcos, San Marcos, Texas; used with
permission.)
while probing and shucking. However, the setae may pick up information from the surrounding water as
the flesh of the bivalve is exposed to that water after valve opening. Setal types present consist mostly
of simple, cuspidate, teasel-like, conate, and miniature simple (Figure 7.5B to Figure 7.5G) (Malcom
2003). Simple setae have been demonstrated to have both chemo- and mechanoreceptive functions in
clawed (Derby 1982) and spiny lobsters (Cate & Derby 2001); cuspidate setae are implicated in grasping
food (Farmer 1974). However, a more recent study on the mouthparts of spiny lobsters (mandibular palp,
medial rim of the basis in the first maxilla and first maxilliped, propus and dactyl of the second maxilliped,
and dactyl of the third maxilliped), found that simple setae apparently respond to displacement of the setal
shaft and cuspidate setae are highly sensitive mechanoreceptors (Garm et al. 2004). It is not clear if such
setae on the pereiopods would serve similar functions, but it is likely given that three pairs of mouthparts
(the maxillipeds) are thoracic-derived appendages and, essentially, highly modified legs. Malcom (2003)
observed some trends while documenting the setal morphology of S. latus,S. nodifer and S. aequinoctialis.
Pereiopods grow progressively smaller toward the posterior end of the lobster’s body, with the fifth male
pereiopods being the smallest. Male and females are dimorphic with regard to the fifth pereiopods, with the
female having a subchelate structure on which sits a pad of stiff, short setae (a “brush pads”; Figure 7.7).
This structure no doubt is involved in the grooming of eggs. The propus, carpus, merus, and ischium
become more ridged along their lateral and medial edges, starting with the third pereiopods, which is
where the pereiopods start becoming smaller in length and width. The oral surfaces tend to have shorter
cuspidate setae, often much shorter than those found on the aboral surfaces. Lateral edges of the aboral
surface tend to have longer cuspidate setae; these become progressively shorter as one moves medially.
At the distal end of each segment (propus, carpus, merus, and ischium only), long cuspidate setae are
present. Miniature setae are widespread on the pereiopods and result in a sculpturing look on the shell
surface.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 167
C
P
M(A) (B)
I
dt
D
P
C
M
I
H1, J1
H1, J1
D
J
J
1J1
J1
J1
J1
J1
J1
J1
J1J1
J1
J1
J1
J1
J1
J1
FIGURE 7.7 Inner, or oral, views of the left 5th pair of walking legs of Scyllarides spp. (A) male; (B) female, showing false
chelae formed by dactyl and propus segments. D — dactyl; P — propus; C — carpus; M — merus; I — ischium; S — meral
shield; J1— cuspidate setae; H1— simple setae, dt — dactyl tuft. (Modified from Malcom, C. 2003. Description of the
setae on the pereiopods of the Mediterranean Slipper Lobster, Scyllarides latus, the Ridged Slipper Lobster, S. nodifer, and
the Spanish Slipper, S. aequinoctialis. M.Sc. thesis: Texas State University at San Marcos, San Marcos, Texas; used with
permission.)
From numerous descriptions and illustrations of phyllosomal stages and nistos, it is clear that all stages
bear setae on both pereiopods and antennules (see Sims 1966; Johnson 1970, 1971b, 1975; Crosnier
1972; Ritz & Thomas 1973; Berry 1974; Phillips & McWilliam 1986, 1989; Robertson 1968, 1971;
Martins 1985a; Ito & Lucas 1990; Sekiguchi 1990); however, only a few studies have actually described
those setae. Typically, phyllosomal pereiopods are described as bearing natatory setae on the exopods
(Robertson 1971). Martins (1985a) states that these setae bear fine setules in S. latus phyllosomas and
these are illustrated by Harada (1958, his Figure 1) for Ibacus ciliatus and by Johnson (1968, his Figure 1
and Figure 2) for Evibacus princeps S.I. Smith, 1869 — the setules would increase the surface area of the
setae, presumably making them more like paddles than rods, and thereby aid in swimming. Pereiopods
are also described as bearing spines, most likely for the purpose of repelling a predator attack, and hook-
like setae on their dactyls that probably grasp food. As for the antennules, it appears that the lateral and
medial filaments increase in size by increasing the number of segments present, and also increase in the
amount of setation borne upon them (Figures 45, 53, 59, 63, 66, 70 for Parribacus antarcticus in Johnson
1971b; Figures 77, 80, 86 for S. squammosus in Johnson 1971b; Figure 4 for S. astori and Figure 10 for
E. princeps in Johnson 1975; Figure 2 to Figure 9 for Eduarctus (formerly Scyllarus)martensii in Phillips
& McWilliam1986; Figures 1d, 2d, 3d, 4d, 5d, 6d, 7d, 8d, 9a for Petarctus (formerly Scyllarus)demani in
Ito & Lucas 1990). Robertson (1968; Figure 16) states that the number of aesthetasc tufts increases with
each successive phyllosomal stage in S. americanus and provides figures illustrating these changes, while
Robertson (1971; Figure 14 to Figure 23) shows a nice sequence for the development of the antennules
in the phyllosoma of S. depressus.
As with nephropids and palinurids, the six pairs of mouthparts bear numerous setae on their various
segments. Adult mouthpart structure has only been described in Ibacus peronii (Suthers & Anderson
1981) and Thenus orientalis (Johnston, 1994; Johnston & Alexander 1999), while phyllosomal mouthpart
structure has been investigated in I. ciliatus (Mikami et al. 1994) and Petarctus (formerly Scyllarus)
demani (Ito & Lucas 1990). Scanning electron microscopy has not been used for a complete analysis of
setal types and distribution; thus, only descriptions from specimens viewed under light microscopy are
available and many of these are incomplete.
Adult mouthparts tend to have a simple setation pattern with fusion of endites — this makes them
considerably less complex than the mouthparts of nephropid or palinurid lobsters. A detailed description
of the mouthparts is provided in Johnston, Chapter 6 and will not be repeated here. Instead, only the
setal types, where known, will be described. Mandibular palps are lined along their upper margins with
pappose setae (Suthers & Anderson 1981; Johnston & Alexander 1999). The first maxilla bears simple
setae on the outer (aboral) surface of the distal endite and bears two multidenticulate, cuneate setae on
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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168 The Biology and Fisheries of the Slipper Lobster
the proximal endite. The proximal endite of the second maxilla is covered with simple setae and the inner
margin of the distal endite is covered with pappose setae (Johnston & Alexander 1999). The margin of the
scaphognathite is fringed with plumose setae (Johnston & Alexander 1999), as it is for nephropid lobsters
(Lavalli & Factor 1992, 1995), which presumably aid in the production of the current by increasing the
surface area of the beating scaphognathite. The endite and exopod of the first maxilliped are fused and both
are lined with simple, pappose, and plumose setae. A terminal flagellum of the exopod is present in some
species, but not in others (see Webber & Booth, Chapter 2; Johnston, Chapter 6 for more information).
In the second maxilliped, short simple setae are found along the upper margins of the propus and carpus,
while longer, hook-like setae are found along the edges of the dactyl (Suthers & Anderson 1981; Johnston
& Alexander 1999). As in the case of the first maxillipeds, the third maxilliped exopod is lacking or
severely reduced in some species, but present in others; additionally the teeth of the ischium are small,
blunt, and reduced compared to those found in nephropid or palinurid lobsters. Multidenticulate setae are
borne upon the merus and simple setae cover the entire inner (oral) surface and part of the outer (aboral)
surface of the dactyl (Suthers & Anderson 1981; Johnston & Alexander 1999).
In the phyllosomal stages, the mouthparts generally increase in the number of setae (often called
spines) present on various surfaces. Little information on the specific types of setae are available, although
Robertson (1968) and Ito & Lucas (1990) state that the lobe constituting the second maxilla bears plumose
setae in the early stages on the proximal but not basal segment, but loses these in later stages as the endites
fully form and separate from the paddle-like scaphognathite.
Evidence from nephropid and palinurid lobsters suggests that simple setae are most likely to be bimodal
chemo- and mechanoreceptors (Derby 1982; Cate & Derby 2001), although those present on mouthparts
appear to be highly sensitive to bend (Garm et al. 2004) and may allow for fine-tuning of manipulation of
the food bolus as ingestion proceeds. Cuspidate setae seem to have multiple functions — from their robust
appearance, they appear to be well suited for stabbing, tearing, and shredding of food particles and may
serve a mechanosensory function (see Garm 2004; Garm et al. 2004 for a discussion) to allow feedback to
the lobster concerning the food-handling process. Plumose setae are not implicated as a sensory structure,
but are thought to be advantageous on appendage segments that create currents (see Lavalli & Factor 1992,
1995; Garm 2004 for a discussion of current generation). Alternatively, they may act as gaskets, sealing
the space between the edge of the scaphognathite and the branchial chamber (Farmer 1974; Factor 1978)
or as filters to prevent entry of particulate matter into the branchial chamber (Farmer 1974). The function
of pappose setae and multidenticulate setae has yet to be determined for any lobster species; however,
Farmer (1974) suggested that pappose setae might be used for grooming purposes, while Garm (2004)
suggests that they act as setal barriers. Setae described as stout spines or spinous processes are likely to be
used for the purpose of gripping particles or pieces of food (Farmer 1974) or may have a mechanosensory
purpose in the same manner that cuspidate setae do (Garm et al. 2004). The other setal types encountered
on the pereiopods, mouthparts, and antennules may be implicated in grooming, touch or hydrodynamic
reception, or fine-food handling.
7.6.2 Mechanoreception
The carapace and abdomen of slipper lobsters, like those of other decapod crustaceans, are covered with
stiff, presumably mechanosensory setae (Figure 7.8), and in some species, these setae are so numerous
and long that the species are said to be covered in a fine pubescence.InScyllarides species, this basic
pubescent pattern shows two distinct elaborations. First, the hairs are arranged in groups of 7 to 12 on
small islands of hard, protruding cuticle (about 500 islands are present on the carapace, depending on the
species). Second, this pattern continues out onto the surfaces of the flattened antennae to the extent that
the antennae, when extended, appear to form a continuous surface with the carapace. These structures
likely serve a hydrodynamic sensing role that provides important sensory feedback while swimming, or
they could be used to detect motion above the lobster while it is walking or still upon the substrate.
The setae present on the second antennae are likely mechanoreceptors used for the purpose of detecting
water currents or vortices as the animal is engaged in swimming activity. Since these appendages act
as rudders that alter the pitch and roll of the lobster’s body while engaged in tail flipping or coast-and-
burst swimming (see Section 7.5.7.2), the numerous setae are likely to represent an important feedback
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Behavior and Sensory Biology 169
A
BC
FIGURE 7.8 Setal tufts found covering the surface of the carapace of Scyllarides latus. (A) setal tufts of tubercles;
(B) arrangement of setae (likely cuspidate or simple setae) within carapace tufts; (C) single tuft, showing distribution of
setae around tubercle. (Photo by Lavalli, Spanier & Grasso.)
system to the lobster. It is possible that setae on the legs and the carapace may also provide important
sensory feedback during swimming. However, significantly more work is needed to elucidate the feedback
mechanisms that control the swimming response in these lobsters.
While not investigated in slipper lobsters, statocysts are present in other families (Cohen 1955) and are
used by individuals within those families to maintain balance. Slipper lobsters are known to initiate tail
flips in response to being flipped over (or even in response to the attempt to flip them over) (Barshaw &
Spanier 1994a; Faulkes 2004), so it is likely that their statocysts are well developed. It is also likely that
the statocyst is heavily relied upon during swimming, so that the second antennae can adjust the pitch of
the animal.
7.6.3 Vision
The eyes of slipper lobsters are stalked, which should provide a broader field of vision. For Thenus spp.,
this field of view is extended as the eyes are located on the lateral edges of the carapace. In all other
species, eyes are more intermediately placed and fit into cups or orbits, which presumably protect the
eyes from mechanical damage. Currently, no descriptions of the compound eyes, their ommatidia, or
the visual pigments exist. No studies have systematically examined the importance of visual signals in
various slipper lobster species. It is likely that vision plays some role in swimming, as blindfolded animals
are not as easily stimulated to initiate swimming when harassed. In addition, while swimming, the eyes
apparently are directed in such a way to provide backward vision (Spanier, personal observations). Finally,
when Thenus and Ibacus spp. bury, they do not cover up their eyes, but leave them exposed, along with the
antennules (Jones 1988; Faulkes 2006), a behavior that strongly suggests that vision may be important.
Clearly, this sensory modality needs to be better investigated.
Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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170 The Biology and Fisheries of the Slipper Lobster
7.6.4 Control Mechanisms and Sensory-Motor Integration
Studies of the advanced sensory-motor integration and precise control in shucking by slipper lobsters
promise insights into the functional organization of the bäuplan of the crustacean central nervous system.
Although no specific investigations of the central mechanisms that control feeding behavior in slipper
lobsters have been conducted, comparisons of what is known from well-studied clawed and spiny lobsters,
and freshwater crayfishes suggest what we will learn. For all three of the latter taxa, the location of the
eyes means that vision can play no roll in the positioning and manipulation of the bivalve during feeding
(although some role of vision is possible for clawed lobsters during crushing of the shell). However,
at least in Thenus spp., vision may play a role as the eyes are situated on the lateral margins of the
carapace (see Jones, Chapter 16 for more information). The precise positioning and manipulation of
the shell by slipper lobsters requires them to make finer sensory discriminations and to have more precise
motor control of dactyl tip force and positioning and tighter feedback between these sensory and motor
functions than in spiny or clawed lobsters consuming bivalves. Considerable evidence suggests that neutral
control programs for the legs are situated in the thoracic ganglia (Ayers & Davis 1977, 1978; Ayers &
Crisman 1993; Clarac et al. 1987; Cruse & Saavedra 1996; Jamon & Clarac 1995, 1997; Domenici et al.
1998, 1999). Elucidation of the neural mechanisms controlling the dexterous feeding behavior in slipper
lobsters will lead to a deeper understanding of sensory-motor control in crustaceans and arthropods in
general.
Observations of shucking behavior and comparative neuranatomy raise further questions about the
neural control of the behavior that can profitably, and uniquely, be explored with slipper lobsters. For
example, slipper lobsters are observed to suspend leg activity as the antennules are brought into contact
with the bivalve during initial assessment of the shell (Malcom 2003). Antennular chemo- and mechanore-
ceptors synapse in the cerebral ganglion in decapod crustaceans (Sandeman 1990; Sandeman et al. 1992).
Does the cerebral ganglion inhibit the motor programs controlling leg movement? How, if at all, does
the information collected by the antennules and processed in the cerebral ganglion affect the thoractic
ganglia? These are fundamental questions of nervous organization and function that apply through the
homologous arthropod bäuplan to lobsters (Fraser 1982; Sandeman et al. 1993).
7.7 Conclusions
Like so many other aspects of their basic biology, little is known of the behavior of the different scyllarid
species. Where studies exist, the focus has been on species within three subfamilies: Scyllarides,Ibacus,
and Thenus, most likely because these are relatively large lobsters that are by-product in other fisheries
or that are specifically targeted. While many Parribacus species are targeted by the aquarist trade, their
commercial importance is significantly less and subsequently little is known of their behavior. The picture
that emerges from the few studied species suggests that, with the possible exceptions of feeding behavior
on bivalves and swimming behavior, there is no “stereotypical” pattern of behavior that describes the
“general” scyllarid — this is because there is no general scyllarid. Bivalve feeding behavior and burst-
and-coast swimming behavior has only been investigated in a few species; thus, it is not clear if the
behavioral patterns seen thus far are common to all scyllarids. The 85 species that have been described
differ widely in their depth and substrate preferences, their early larval life-history strategy, their dietary
preferences, and their reproductive strategies.
Behavioral studies, in and of themselves, should not be difficult to undertake on these species. More
and more scyllarids are being successfully cultured in the laboratory from the naupliosoma or first-
stage phyllosoma. These successes should allow for detailed studies of depth preferences, phototactic
responses, responses to thermoclines, and swimming behavior in flow. If phyllosomas are reared to the
nisto stage, studies using techniques that have been employed for nephropids and palinurids should allow
the determination of how nistos sample the benthos, what causes them to choose specific substrates, and
how they avoid predators. Most adults and juveniles are readily held in the laboratory, and lessons learned
from nephropid and palinurid lobsters demonstrate that many “natural” behaviors can be examined if the
study animals are provided with sufficient space within a naturalistic setting. Videotaped analysis removes
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Behavior and Sensory Biology 171
the need for tedious, in-person observations, and allow for a more detailed analysis of both the behavior
studied and the order in which behavioral sequences are carried out.
While behavioral research on these lobsters is limited, studies on the sensory structures and their func-
tions are almost nonexistent. This is remarkable because of the unique behaviors observed in this family.
The dramatic change in the dactyls of the pereiopods with the concomitant loss of setae at the distal end
of the dactyl suggests that the mode of finding food buried within the benthos differs greatly in scyllarids
from that seen in nephropids and palinurids. Furthermore, the observations by a number of workers that
the antennules are used to apparently directly sample the food item (i.e., contact chemoreception) during
its manipulation, suggests that the sensory structures of the antennule may differ in their responsiveness
to various odor stimulants. Additional observations of distance orientation to food-odor plumes, some-
thing that is well studied and easily performed by nephropids and palinurids, suggest that fundamental
differences exist in the central nervous control of feeding behavior in scyllarids.
The flattened second antennae, which are very important in the swimming response of several species
thus far studied, must have dynamic hydroreceptors that provide feedback to the lobster during swimming,
so that it can effect turning maneuvers and change its body’s pitch angle. The numerous surface setae on
the carapace may also provide a feedback role here. Likewise, the statocyst of slipper lobsters is probably
extremely well developed to provide rapid feedback to the lobster while it engages in coast-and-burst
swimming movements. In addition, while vision has been discounted compared to chemoreception and
mechanoreception in other species, the swimming abilities of these lobsters and the fact that species that
bury do not cover over their eyes, suggests that it may be a more important and a better developed sense
in scyllarids.
As stated by Faulkes (2004), the loss of the giant interneuron that effect the escape tail-flip response of
scyllarids begs numerous questions concerning the evolution of decapod nervous systems and the response
of those nervous systems to deleted or altered components. It also provides a comparative approach because
some scyllarids spend much of their time with their abdomen mostly tucked under their body, while others
leave the abdomen extended. The nervous system of the former species could be compared to other
decapods which have similar postures and have lost the giant interneurons, while the latter species could
be compared to others that have not lost the interneurons.
Behavior is, ultimately, the result of the interplay of genes, physiology, and the environment. Sensory
input affects and finely tunes behavioral output. Together, the sensory biology and behavior of individuals
affects the success of the populations comprising the species and the ability of those populations to adapt
to their environments. A failure to understand these aspects of the animal’s biology can have profound
effects upon the ability of the species to survive and thrive under exploitation. As slipper lobsters are
becoming more and more the target of fisheries, it is critical that we begin to focus our attention upon
them, not only because they are exploited, but also because they are, in and of themselves, interesting
animals that can provide evolutionary insights into other decapod species and arthropods as a whole.
References
Ache, B.W. & Zhainazarov, A.B. 1995. Dual second-messenger pathways in olfactory transduction. Curr. Opin.
Neurobiol. 5: 461–466.
Almog-Shtayer, G. 1988. Behavioural-ecological aspects of Mediterranean slipper lobsters in the past and
of the slipper lobster Scyllarides latus in the present. M.A. thesis: University of Haifa, Israel.
165 pp.
Atema, J. 1977. Functional separation of smell and taste in fish and Crustacea. In: Le Magnen, J. & MacLeod, P.
(eds.), Olfaction and Taste, Vol. 6: pp. 165–174. London: Information Retrieval.
Atema, J. 1980. Smelling and tasting underwater. Oceanus 23: 4–18.
Atema, J. 1986. Review of sexual selection and chemical communication in the lobster, Homarus americanus.
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Atema, J. & Cobb, J.S. 1980. Social behavior. In: Cobb, J.S. & Phillips, B.F. (eds.), The Biology and Management
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Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/bu/detail.action?docID=283296.
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Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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Lavalli, Kari L., and Ehud Spanier. The Biology and Fisheries of the Slipper Lobster, edited by Kari L. Lavalli, and Ehud Spanier, CRC Press, 2007.
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... S. latus è una specie tipica degli ambienti rocciosi e delle praterie di Posidonia oceanica, dal comportamento gregario, che trascorre le ore diurne nascosta in anfratti e si alimenta durante la notte (Lavalli et al., 2007). Si trova prevalentemente tra i 2 e i 50 m di profondità, ma può spingersi a profondità molto maggiori (400 m) (Pessani & Mura, 2007). ...
... Si trova prevalentemente tra i 2 e i 50 m di profondità, ma può spingersi a profondità molto maggiori (400 m) (Pessani & Mura, 2007). Le magnose si nutrono di bivalvi e gasteropodi e svolgono l'attività riproduttiva una volta l'anno, durante i mesi estivi (Holthuis, 1991;Lavalli et al., 2007;Pessani & Mura, 2007). ...
... L'interesse commerciale per la specie, favorito anche dalle sue grandi dimensioni, ha condotto al sovrasfruttamento delle popolazioni in gran parte dell'areale e soprattutto nelle Azzorre e in Italia e (Bianchini & Ragonese, 2007;Pessani & Mura, 2007). Le tipologie di pesca più utilizzate per la cattura delle magnose sono la pesca con le reti da posta (tramagli), le nasse e la pesca subacquea (Holthuis, 1991;Spanier & Lavalli, 2007). ...
... S. latus è una specie tipica degli ambienti rocciosi e delle praterie di Posidonia oceanica, dal comportamento gregario, che trascorre le ore diurne nascosta in anfratti e si alimenta durante la notte (Lavalli et al., 2007). Si trova prevalentemente tra i 2 e i 50 m di profondità, ma può spingersi a profondità molto maggiori (400 m) (Pessani & Mura, 2007). ...
... Si trova prevalentemente tra i 2 e i 50 m di profondità, ma può spingersi a profondità molto maggiori (400 m) (Pessani & Mura, 2007). Le magnose si nutrono di bivalvi e gasteropodi e svolgono l'attività riproduttiva una volta l'anno, durante i mesi estivi (Holthuis, 1991;Lavalli et al., 2007;Pessani & Mura, 2007). ...
... L'interesse commerciale per la specie, favorito anche dalle sue grandi dimensioni, ha condotto al sovrasfruttamento delle popolazioni in gran parte dell'areale e soprattutto nelle Azzorre e in Italia e (Bianchini & Ragonese, 2007;Pessani & Mura, 2007). Le tipologie di pesca più utilizzate per la cattura delle magnose sono la pesca con le reti da posta (tramagli), le nasse e la pesca subacquea (Holthuis, 1991;Spanier & Lavalli, 2007). ...
... A third and major group of lobsters, the Scyllarids (slipper lobsters), also appear nocturnally active. However, the prevalence of endogenous rhythms in slipper lobsters remains elusive (see Lavalli et al., 2007). Mediterranean slipper lobsters (Scyllarides latus) exemplify a commercially and biologically important species throughout their range. ...
... After synchronizing their activity patterns to an imposed LD cycle, these lobsters continued to express the same activity pattern in constant conditions, maintaining a circadian rhythm for the duration of the trial (Figures 4 and 5). Few studies have examined diel activity patterns in Scyllarid lobsters despite their economic and ecological importance throughout their range (Lavalli et al., 2007). Jones (1988) has provided the only other detailed analysis of diel activity patterns for Scyllarid species (T. ...
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Although the natural history for Mediterranean slipper lobsters (Scyllarides latus) is well established, there exists a disproportionate lack of important biological and physiological data to verify many key traits, including to what extent endogenous rhythms modulate aspects of their behaviour. Although Scyllarids appear nocturnally active, few studies exist that quantify this tendency. Our overall objective was to test the hypothesis that adult slipper lobsters are nocturnal and to determine if their diel activity rhythms are under the influence of an endogenous circadian clock. In the laboratory, we exposed a total of 16 animals (CL avg ¼ 92.6 + 6.6 mm; CL, carapace length) to a 12 : 12 light : dark (LD) cycle for 7-10 d, followed by ***constant dark (DD) for 15-20 d. Activity was assessed using a combination of time-lapse video and accelerometers. Of a total of 16 lobsters, we analysed data from 15 (one mortality). All 15 lobsters were evaluated using video. Thirteen of these lobsters were also evaluated using acceler-ometers. All lobsters were more active during night-time than during daytime and synchronized their activity to the LD cycle, expressing a diel activity pattern (t ¼ 24.04 + 0.13 h). In DD, lobsters maintained a circadian rhythm with a t of 23.87 + 0.07 h. These findings may provide insight into the behaviour of these animals in their natural habitat and help explain their ability to anticipate dawn and dusk.
... The phyllosoma larval stage of lobsters feed primarily on marine invertebrate larvae, including small zooplankton and jellyfish (Jones, 2007). They are often observed attached to the aboral surface of jellyfish medusae, where they may ride and feed on the medusae until they metamorphose into the benthic nisto stage (Lavalli et al., 2007;Wakabayashi and Phillips, 2016). Although many lobster species are oceanic, phyllosomas are also abundant in coastal waters and shelf areas with depths of ~ 40 to 50 m (Barnett et al., 1984;Jones, 2007), where coastal acidification is a major environmental concern (Duarte et al., 2013;Wallace et al., 2014). ...
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Ocean acidification (OA) can alter the behaviour and physiology of marine fauna and impair their ability to interact with other species, including those in symbiotic and predatory relationships. Phyllosoma larvae of lobsters are symbionts to many invertebrates and often ride and feed on jellyfish, however OA may threaten interactions between phyllosomas and jellyfish. Here, we tested whether OA predicted for surface mid-shelf waters of Great Barrier Reef, Australia, under ∆ pH = −0.1 (pH ~7.9) and ∆pH = −0.3 (pH ~7.7) relative to the present pH (~8.0) (P) impaired the survival, moulting, respiration, and metabolite profiles of phyllosoma larvae of the slipper lobster Thenus australiensis, and the ability of phyllosomas to detect chemical cues of fresh jellyfish tissue. We discovered that OA was detrimental to survival of phyllosomas with only 20% survival under ∆pH = −0.3 compared to 49.2 and 45.3% in the P and ∆pH = −0.1 treatments, respectively. The numbers of phyllosomas that moulted in the P and ∆pH = −0.1 treatments were 40% and 34% higher, respectively, than those in the ∆pH = −0.3 treatment. Respiration rates varied between pH treatments, but were not consistent through time. Respiration rates in the ∆pH = −0.3 and ∆pH = −0.1 treatments were initially 40% and 22% higher, respectively, than in the P treatment on Day 2 and then rates varied to become 26% lower (∆pH = −0.3) and 17% (∆pH = −0.1) higher towards the end of the experiment. Larvae were attracted to jellyfish tissue in treatments P and ∆pH = −0.1 but avoided jellyfish at ∆pH = −0.3. Moreover, OA conditions under ∆pH = −0.1 and ∆pH = −0.3 levels reduced the relative abundances of 22 of the 34 metabolites detected in phyllosomas via Nuclear Magnetic Resonance (NMR) spectroscopy. Our study demonstrates that the physiology and ability to detect jellyfish tissue by phyllosomas of the lobster T. australiensis may be impaired under ∆pH = −0.3 relative to the present conditions, with potential negative consequences for adult populations of this commercially important species.
... The response of slipper lobsters to predator attack (e.g., by gregarious triggerfish) has been well studied [79,98,[102][103][104][105][106][107][108] and consists of three strategies, two of which are typically executed in sequence: (i) the "fortress strategy" in which the animal grasps the bottom and attempts to outlast its attacker's motivation to penetrate its hard shell (described in [107]); (ii) the "swimming escape" response (described in [102,[105][106][107]); and (iii) remaining sheltered in dens [79,103]. Lacking claws (like Homarus spp.) or long spinose antennae (like spiny lobsters; see [109][110][111][112]) with which to fend off swimming predators, slipper lobsters have developed a shell that is thicker and more durable to mechanical insult than clawed or spiny lobsters [107]. ...
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The light characteristics of an ecosystem drive evolutionary adaptations in visual traits, enhancing the diversity and abundance of species living there. The visual systems of crustaceans are highly diverse and often correspond to the optical properties of their preferred environments. Although habitat depth is known to greatly influence visual specialization in marine crustaceans, it remains unclear whether depth drives visual adaptions in nocturnal species. Slipper lobsters (Scyllaridae) are nocturnal benthic marine crustaceans distributed throughout a wide range of depths. In order to understand the visual adaptive capabilities of slipper lobsters inhabiting different depths, we characterized the eye structures of a shallow-water species ( Parribacus japonicas ), an intermediate-depth species ( Scyllarides squammosus ) and a deep-water species ( Ibacus novemdentatus ). Moreover, we measured by electroretinogram (ERG) the spectral sensitivities and temporal resolutions for each species using the following light stimuli: UV (λ max 386 nm), blue (λ max 462 nm), green (λ max 518 nm), yellow (λ max 590 nm), and red (λ max 632 nm). Our histological experiments show that all three species possess a typical superposition compound eye with square facets, and their ERG measurements revealed a single sensitivity peak for each species. Notably, peak spectral sensitivity corresponded to habitat depth, with the estimated peak for I. novemdentatus (493.0 ± 9.8 nm) being similar to that of S. squammosus (517.4 ± 2.1 nm), but lower than that of P. japonicus (537.5 ± 9.9 nm). Additionally, the absolute sensitivities at respective peak wavelengths for I. novemdentatus and P. japonicus were higher than that of S. squammosus . No differences were observed among the three species for maximum critical flicker fusion frequency (CFF max ) across light stimuli. However, P. japonicus had lower CFF max values than the other two species. These data suggest that all three nocturnal slipper lobsters are likely monochromatic and well adapted to dim light environments. Significantly, the deep-water slipper lobster displayed higher spectral sensitivities at shorter wavelengths than the shallow water species, but temporal resolution was not compromised.
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Following recent advances in hatchery technology and large-scale larval rearing of spiny and slipper lobsters, a greater understanding of key nutritional requirements is now imperative to facilitate feed development for juvenile culture. However, there is a lack of relevant information available for the slipper lobsters, particularly Thenus species. This study sought to evaluate the potential requirements for and effects of a fresh ingredient component in a formulated feed on the growth performance of hatchery-reared juvenile Thenus australiensis. This was assessed using six formulated experimental feeds incorporating fresh blue mussel flesh (BM; Mytilus galloprovincialis) in a geometric series (0, 1.6, 3.1, 6.3, 12.5 and 25.0% dry matter) administered continuously over 9 weeks. An additional dietary treatment of blue mussel half shells (BMHS) was included as a reference feed to establish a benchmark of growth potential. Survival and moult frequency were unaffected by feed treatment. Lobsters fed BMHS displayed a clustered or synchronised moulting pattern with two major moulting events lasting several days each during the experiment. In contrast, lobsters provided experimental feeds moulted continuously throughout the experiment. The BMHS produced approximately 2-fold greater growth compared to experimental feeds, while BM incorporated into experimental feeds had no beneficial effect on growth at any inclusion level. Growth mirrored feed intake where it was also found that feed intake on a dry matter basis was approximately 2-fold higher for BMHS compared to experimental feeds, while biological feed conversion ratios were similar for all feeds. Thus, dry matter intake appeared to be primarily responsible for the difference in growth rates achieved. Bulk chemical analysis revealed that BMHS-fed lobsters had significantly lower ash content coupled with higher gross energy, total lipid and polar lipid contents on a proportional basis compared to lobsters fed formulated experimental feeds. Lobster haemolymph Brix values were also significantly higher in the BMHS treatment lobsters. Higher Brix coupled with decreasing feed intake suggests these lobsters were on average at a more advanced stage within the moult cycle and offers a possible explanation for differences in body chemistry. In conclusion, while growth rates were lower in animals fed formulated feeds compared to benchmark growth performance, overall survival, moulting and growth performance revealed the potential of the basal experimental feed formulation as a reference for future nutrition research with this species without a requirement for inclusion of fresh BM. The research highlights that significant opportunities exist to improve slipper lobster growth performance through maximising feed intake.
Chapter
This chapter presents a comprehensive overview of the global distribution of lobsters (nephropid, palinurid and scyllarid) in all oceans. Lobsters are found in tropical, subtropical and temperate regions, from the intertidal to great depths. Many species of the genus Panulirus prefer rocky or coral reefs and some are found in sandy/muddy substrates. The tropical zone has the largest number of species (174), followed by the subtropics with 71 species and the temperate region with 16 species. Among the 63 species of palinurids, 39 are distributed in the tropics and 19 species in the subtropical zone with many species overlapping in their distribution. Under the family Nephropidae, 32 species are found in the tropical belt, 15 in the subtropical zone and 10 species in the temperate zone. Three species under two genera, Thymops and Thymopides are distributed in the southern Atlantic and Indian Ocean region (50oS). The Indo-West Pacific is the region with maximum diversity with the nephropids represented by 29 species, palinurids by 36 species and the scyllarids by 57 species. Among the total number of scyllarids, 63 species are distributed in the tropical zone, 26 in the subtropical and 3 species in the temperate region.
Chapter
Marine lobsters are high-valued crustaceans occupying a variety of habitats in tropical, subtropical and temperate oceans, from continental shelves and slopes to deep sea ridges, remote seamounts, lagoons and even estuaries. The highly diverse form and size are suited to their wide distribution in a range of shallow coastal and oceanic habitats. They are economically important, forming sustained profitable fisheries in many countries. In 2016, world capture production was 3,08,926 t, with an additional 2000 t from aquaculture. Apart from their economic importance, they are an important component of the marine ecosystems, playing pivotal roles as a prey and a predator. Lobster has been one of the most extensively studied group, and literature search has yielded more than 15,000 entries including research papers, technical reports and popular articles on taxonomy, biology, physiology, ecology, fisheries and aquaculture of clawed, spiny and slipper lobsters. The knowledge base created on individual species is critical in our understanding of the life history strategies and ecology of larvae, juveniles and adults. Advances in molecular techniques have been instrumental in resolving the conflicting hypotheses of evolutionary relationships among different taxa. Recent success in breeding and seed production has prompted researchers to develop commercially viable technologies for aquaculture of a few species, which is expected to relieve the mounting pressure on the natural resource. Lobster has been subjected to intensive exploitation due to high price in the international markets and except in a few countries where impeccable management strategies have ensured sustained production, the resource has been under heavy pressure due to poor enforcement of fishing and marketing regulations. The need to develop alternate management strategies, including co-management, is emphasised so that the resource could be conserved and their sustainability ensured. This volume attempts to present available information on biology, fisheries and aquaculture of lobsters with special focus on lobster research in India.
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The technique of underwater acoustic localization (UAL) is widely used to track submerged targets. In the present study, we consider UAL in the specific application of locating the Mediterranean slipper lobster, Scyllarides latus (S. latus), for the aim of continuous and long-term investigation of its movement patterns. Since visual inspections only offer a snapshot to the life of these nocturnal crustaceans, we used underwater acoustic tags and deployed the first testbed to track a community of S. latus through UAL. The testbed included 19 acoustically tagged S. latus, as well as a sensor network of four receivers that record the time-of-arrival of detected tag’s emissions. The testbed operated for six months, and the data collected were analyzed offline. In this paper, we describe the design details of our testbed. We discuss the considerations in choosing both the type of acoustic tags and the structure of the testbed. We also present our algorithm to time synchronize the receivers, and the localization procedure. Our preliminary results show that, for this long-term deployment, the widely used model of clock offset for the receivers is too simplified, and that time synchronization must consider also the clock’s skew. We also show that the tagged lobsters are mostly detected by fewer than three receivers, thereby making localization using traditional trilateration methods highly challenging. Finally, preliminary results for the localization of the tagged lobsters in the deployment area are presented.
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Full-text available
Phyllosoma larvae collected to date in Japanese and Taiwanese waters have been classified into two genera (Linuparus, Panulirus) of the Palinuridae, four genera (Ibacus, Parribacus, Scyllarides, Scyllarus) of the Scyllaridae, and one genus (Palinurellus) of the Synaxidae. However, phyllosoma larvae of three Scyllarus species (S. bicuspidatus, S. cultrifer, S. kitanoviriosus) are absolutely dominant among the larvae collected in the waters. Scyllarus larvae are abundant in coastal waters while those of Panulirus are often collected in offshore/oceanic waters. Based on previous and ongoing studies dealing with spatial distributions of phyllosoma larvae in Japanese and Taiwanese waters, it appears that phyllosoma and nisto larvae of the Scyllarus are retained within coastal waters north of the Kuroshio Current. On the other hand, the life history of the Panulirus (particularly P. japonicus) may be completed within the Kuroshio Subgyre: their phyllosoma larvae may be flushed out from coastal waters into the Kuroshio, then transported through the Counter Current south of the Kuroshio into the water east of Ryukyu Archipelago and Taiwan where they attain the subfinal/final phyllosoma or puerulus stages, once again entering the Kuroshio and dispersing into coastal waters.
Article
Twelve late-stage phyllosoma larvae of Chelarctus cultrifer (Ortmann, 1897) were collected in the offshore area northeast of Yonaguni Island on 23 November 1998, transported to the laboratory, and reared individually in circular acrylic chambers on a diet of fresh clam meat. Two final-stage phyllosomas metamorphosed into the nisto stage after 14 and 30 days, and these nistos then molted to the juvenile stage after 10 and 11 days, respectively. The morphology of the late-stage phyllosomas collected off Yonaguni Island, other phyllosomas from the Northwestern Pacific, and the nisto and juvenile stage exuvia of the reared larvae are described and illustrated in detail.