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CRUSTACEAN ISSUES ] 3<br />
%. m<br />
II<br />
<strong>Decapod</strong> <strong>Crustacean</strong> <strong>Phylogenetics</strong><br />
edited by<br />
Joel W. Martin, Keith A. Crandall, and Darryl L. Felder<br />
£\ CRC Press<br />
J Taylor & Francis Group
<strong>Decapod</strong> <strong>Crustacean</strong> <strong>Phylogenetics</strong><br />
Edited by<br />
Joel W. Martin<br />
<strong>Natural</strong> History Museum of L. A. County<br />
Los Angeles, California, U.S.A.<br />
KeithA.Crandall<br />
Brigham Young University<br />
Provo,Utah,U.S.A.<br />
Darryl L. Felder<br />
University of Louisiana<br />
Lafayette, Louisiana, U. S. A.<br />
CRC Press is an imprint of the<br />
Taylor & Francis Croup, an informa business
CRC Press<br />
Taylor & Francis Group<br />
6000 Broken Sound Parkway NW, Suite 300<br />
Boca Raton, Fl. 33487 2742<br />
Contents<br />
Preface<br />
JOEL W. MARTIN, KEITH A. CRANDALL & DARRYL L. FELDER<br />
I Overviews of <strong>Decapod</strong> Phylogeny<br />
On the Origin of <strong>Decapod</strong>a<br />
FREDERICK R. SCHRAM<br />
<strong>Decapod</strong> <strong>Phylogenetics</strong> and Molecular Evolution 15<br />
ALICIA TOON. MAEGAN FINLEY. JEFFREY STAPLES & KEITH A. CRANDALL<br />
Development, Genes, and <strong>Decapod</strong> Evolution 31<br />
GERHARD SCHOLTZ. ARKHAT ABZHANOV. FREDERIKR ALWES. CATERINA<br />
BIEFIS & JULIA PINT<br />
Mitochondrial DNA and <strong>Decapod</strong> Phylogenies: The Importance of 47<br />
Pseudogenes and Primer Optimization<br />
CHRISTOPH D. SCHUBART<br />
Phylogenetic Inference Using Molecular Data 67<br />
FERRAN PALERO & KEITH A. CRANDALL<br />
<strong>Decapod</strong> Phylogeny: What Can Protein-Coding Genes Tell Us? 89<br />
K.H. CHU, L.M. TSANG. K.Y. MA. T.Y. CHAN & P.K.L. NG<br />
Spermatozoal Morphology and Its Bearing on <strong>Decapod</strong> Phylogeny 101<br />
CHRISTOPHER TUDGE<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 121<br />
AKIRA ASAKURA<br />
A Shrimp's Eye View of Evolution: How Useful Are Visual Characters in 183<br />
<strong>Decapod</strong> <strong>Phylogenetics</strong>?<br />
MEGAN L. PORTER & THOMAS W. CRONIN<br />
<strong>Crustacean</strong> Parasites as Phylogenetic Indicators in <strong>Decapod</strong> Evolution 197<br />
CHRISTOPHER B. BOYKO & JASON D. WILLIAMS<br />
The Bearing of Larval Morphology on Brachyuran Phylogeny 221<br />
PAUL F. CLARK
vi Contents<br />
II Advances in Our Knowledge of Shrimp-Like <strong>Decapod</strong>s<br />
Evolution and Radiation of Shrimp-Like <strong>Decapod</strong>s: An Overview 245<br />
CHARLES H..I.M. ERANSEN & SAMMY DE GRAVE<br />
A Preliminary Phylogenelic Analysis of the Dendrobranchiata Based on 261<br />
Morphological Characters<br />
CAROLINA TAVARES. CRISTIANA SERE.IO & JOEL W. MARTIN<br />
Phvlogeny of the Infraorder Caridea Based on Mitochondrial and Nuclear 281<br />
Genes (Crustacea: <strong>Decapod</strong>a)<br />
HEATHER D. BRACKEN. SAMMY DE GRAVE & DARRYL L. FEEDER<br />
III Advances in Our Knowledge of the Thalassinidean<br />
and Lobster-Like Groups<br />
Molecular Phylogeny of the Thalassinidea Based on Nuclear and 309<br />
Mitochondrial Genes<br />
RAFAEL ROBLES. CHRISTOPHER C. TUDGE, PETER C. DWORSCHAK, GARY C.B.<br />
POORE & DARRYL L. FBLDER<br />
Molecular Phylogeny of the Family Callianassidae Based on Preliminary 327<br />
Analyses of Two Mitochondrial Genes<br />
DARRYL L. FELDER & RAFAEL ROBLES<br />
The Timing of the Diversification of the Freshwater Crayfishes 343<br />
JESSE BREINHOLT. MARCOS PEREZ-LOSADA & KEITH A. CRANDALL<br />
Phylogeny of Marine Clawed Lobster Families Nephropidae Dana. 1852. 357<br />
and Thaumastochelidae Bate. 1888, Based on Mitochondrial Genes<br />
DALE TSHUDY. RAFAEL ROBLES. TIN-YAM CHAN, KA CHAI HO. KA HOU CHU,<br />
SHANE T. AHYONG & DARRYL L. FELDER<br />
The Polychelidan Lobsters: Phylogeny and Systematics (Polychelida: 369<br />
Polychelidae)<br />
SHANE T. AHYONG<br />
IV Advances in Our Knowledge of the Anomttra<br />
Anomuran Phylogeny: New Insights from Molecular Data 399<br />
SHANE T. AHYONG, KAREEN E. SCHNABHL & ELIZABETH W. MAAS<br />
V Advances in Our Knowledge of the Brachyura<br />
Is the Brachyura Podotremata a Monophyletic Group? 417<br />
GERHARD SCHOLTZ & COLIN L. MCLAY
Contents vii<br />
Assessing the Contribution of Molecular and Larval Morphological 437<br />
Characters in a Combined Phylogenetic Analysis of the Supcrfamily<br />
Majoidea<br />
KRISTIN M. HUI.TGREN, GUILLERMO GUHRAO, HERNANDO RL. MARQUES &<br />
EHRRAN P. PALERO<br />
Molecular Genetic Re-Examination of Subfamilies and Polyphyly in the 457<br />
Family Pinnotheridae (Crustacea: <strong>Decapod</strong>a)<br />
EMMA PALACIOS-THEIL. JOSE A. CUESTA. ERNESTO CAMPOS & DARRYL L.<br />
FELDER<br />
Evolutionary Origin of the Gall Crabs (Family Cryptochiridae) Based on 475<br />
16S rDNA Sequence Data<br />
REGINA WETZER. JOEL W. MARTIN & SARAH L. BOYCE<br />
Systematics, Evolution, and Biogeography of Freshwater Crabs 491<br />
NEIL CUMBERLIDGE & PETER K.L. NG<br />
Phylogeny and Biogeography of Asian Freshwater Crabs of the Family 509<br />
Gecarcinucidae (Brachyura: Potamoidea)<br />
SEBASTIAN KLAUS. DIRK BRANDIS. PETER K.L. NG. DARREN C.J. YEO<br />
& CHRISTOPH D. SCHUBART<br />
A Proposal for a New Classification of Porlunoidea and Cancroidea 533<br />
(Brachyura: Heterotremata) Based on Two Independent Molecular<br />
Phylogenies<br />
CHRISTOPH D. SCHUBART & SILKE RRUSCHRL<br />
Molecular Phylogeny of Western Atlantic Representatives of the Genus 551<br />
Hexapanopeus (<strong>Decapod</strong>a: Brachyura: Panopeidae)<br />
BRENT P. THOMA. CHRISTOPH D. SCHUBART & DARRYL L. FELDER<br />
Molecular Phylogeny of the Genus Cronius Stimpson, I860, with 567<br />
Reassignment of C. tumidulus and Several American Species ol' Port un us<br />
to the Genus Achelous De Haan, 1833 (Brachyura: Portunidae)<br />
FERNANDO L. MANTELATTO. RAFAEL ROBLES. CHRISTOPH D. SCHUBART<br />
& DARRYL L. FELDER<br />
Index 581<br />
Color Insert
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s<br />
AKIRAASAKURA<br />
<strong>Natural</strong> History Museum & Institute, Chiba, Japan<br />
ABSTRACT<br />
The mating systems of decapod crustaceans are reviewed and classified according to general patterns<br />
of lifestyles and male-female relations. The scheme employs criteria that focus on ecological, life<br />
history, and social determinants of both male and female behavior, and by these criteria nine types of<br />
mating systems are distinguished: (1) Short courtship: Both males and females are free-living (= not<br />
symbiotic with other organisms), and copulation occurs after brief behavioral interactions between a<br />
male and a female. (2) Precopulatory guarding: A male guards a mature female one to several days<br />
before copulation; both males and females are generally free-living. (3) Podding: In some largesize<br />
decapods, aggregations consisting of an extremely large number of individuals are formed, and<br />
mating occurs inside those aggregations. (4) Pair-bonding: In many symbiotic and some free-living<br />
species, males and females are found in a heterosexual pair and are regarded as having a monogamous<br />
mating system. They may live on or inside other organisms such as sponges, corals, molluscs,<br />
polychaetes, sea urchins, ascidians, and algal tubes. (5) Eusocial: In some sponge-dwelling snapping<br />
shrimps, a colony of shrimps contains a single reproductive female and many small individuals<br />
that apparently never breed. (6) Waving display: In many intertidal and semi-terrestrial crabs inhabiting<br />
mudflats or sandy beaches, males conduct visual displays that include species-specific dances<br />
to attract females. (7) Visiting: In some hapalocarcinid crabs, females are sealed inside a coral gall,<br />
and the male crab normally residing outside the gall is assumed to visit the gall for mating. (8)<br />
Reproductive swarm: In some pinnotherid crabs, mating occurs when a female is a free-swimming<br />
instar before she enters her definitive host. (9) Dwarf male mating: In some anomuran sand crabs,<br />
an extremely small male attaches near the gonopore of a free-living female.<br />
1 INTRODUCTION<br />
<strong>Decapod</strong> crustaceans are a large and diverse assemblage of animals. In most decapods, the sexes<br />
live separately and pair briefly as adults. Pairs are formed after a brief display, the sexes remain<br />
together for a relatively short period, the sexes separate after copulation, and the females assume<br />
all further parental duties such as selecting suitable habitat for egg incubation, aeration, and cleaning<br />
(Salmon 1983). However, recent discoveries of often-conspicuous behavior and male-female<br />
relations among decapods have shown that their mating system is highly diverse and is sometimes<br />
quite similar to mating systems of other animals such as birds, mammals, reptiles, and insects (see<br />
Shuster & Wade 2003; Duffy & Thiel 2007 for a review).<br />
As claimed by Emlen & Oring (1977) in their classic work on the relationships among ecological<br />
factors, sexual selection, and the evolution of mating system, sexual selection is the driving force that<br />
underlies the evolution of male-male competition and female choice. However, ecological factors<br />
apparently contribute to the evolution of mating systems as well as to behavioral and morphological<br />
differences between the sexes. From this point of view, much study has been conducted recently on<br />
the evolution of the mating system of decapods (see section 2 below).
122 Asakura<br />
In this paper, I describe the diversity of mating systems of decapods in an attempt to recognize<br />
and classify their general patterns from the viewpoints of the ecological, life history, and social determinants<br />
of both male and female behavior. Historically, there are two ways of describing mating<br />
systems (Shuster & Wade 2003). The first is in behavioral ecology, where mating systems are usually<br />
described in terms of the number of mates per male or female, such as monogamy, polygyny, and<br />
polyandry. The second is in terms of the genetic relationships between mating males and females,<br />
such as random mating, negative assortative mating (outbreeding), and positive assortative mating<br />
(inbreeding). My approach to describing mating systems of decapods is a "recognition of general<br />
pattern" approach, a kind of a combination of these two approaches that captures variation in the<br />
relationship between male and female, from promiscuity to monogamy, as well as the relationship<br />
between male guarding and the female tendency to settle down in certain places or to aggregate, and<br />
the complex nature of eusociality.<br />
Terminology generally follows Duffy & Thiel (2007). Additionally, some basic terms are redefined<br />
here, because these terms are sometimes used in more or less different ways according to taxa,<br />
including birds, mammals, and fish:<br />
• Monogamy (= pair bonding): One male and one female have an exclusive mating relationship.<br />
• Polygamy: One or more males have an exclusive relationship with one or more females. Three<br />
types are recognized: polygyny, where one male has an exclusive relationship with two or more<br />
females; polyandry, where one female has an exclusive relationship with two or more males;<br />
and polygynandry, where two or more males have an exclusive relationship with two or more<br />
females (the numbers of males and females need not be equal, and, in vertebrate species studied<br />
so far, the number of males is usually fewer).<br />
• Promiscuity: Any male within the group mates with any female.<br />
• Eusociality: Multigenerational (cohabitation of different generations), cooperative colonies with<br />
strong reproductive skew (reproductive division of labor, usually a single breeding female) and<br />
cooperative defense of the colony (after Duffy 2003).<br />
• Symbiosis: Here defined simply as dissimilar organisms living together.<br />
2 HISTORY OF STUDY<br />
The first important review of decapod mating systems was Hartnoll's (1969) publication on brachyuran<br />
crabs. He distinguished two types of mating systems. "Soft-female mating" was defined as copulation<br />
occurring immediately after molting of the female, usually preceded by a lengthy pre-molt<br />
courtship behavior including precopulatory guarding by the male. "Hard-female mating" was defined<br />
as mating in which the female copulates during the intermolt stage after a relatively brief<br />
courtship behavior.<br />
Through their intensive study of the harlequin shrimp Hymenocera picta, Wickler & Seibt (see<br />
Reference 16 in Appendix I, Table 10) found that these shrimp form stable heterosexual pairs based<br />
on individual recognition by chemical cues at a distance. Wickler & Seibt discussed several similar<br />
hypotheses, independently developed in research on crustaceans and humans, for the evolution of<br />
monogamy and other mating systems. Individual recognition in the monogamous mating system<br />
was intensively studied in the banded shrimp Stenopus hispidus by Johnson (1969, 1977).<br />
The report by Emlen & Oring (1977) was influential for studies on crustacean mating systems.<br />
They classified the mating system into the following categories:<br />
1. Monogamy<br />
2. Polygyny (subdivided into 2a, resource defense polygyny; 2b, female (or harem) defense<br />
polygyny; and 2c, male dominance polygyny (further subdivided into 2c-1, explosive breeding<br />
assemblages, and 2c-2, leks))
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 123<br />
3. Rapid multiple clutch polygamy<br />
4. Polyandry (subdivided into 4a, resource defense polyandry; and 4b, female access polyandry)<br />
Ridley (1983) intensively reviewed the precopulatory mate guarding behavior in various groups<br />
of animals including tardigrades, crustaceans, arachnids, and anurans, and discussed its evolution.<br />
Work on the behavior of the fiddler crabs (genus Uca) has contributed greatly to our understanding<br />
of the mating systems of brachyuran crabs. These studies include the works of H.O. von Hagen<br />
(e.g., von Hagen 1970), J. Crane (e.g., Crane 1975), J. Christy and his coworkers (e.g., Christy<br />
et al. 2003a, b), M. Salmon and his coworkers (e.g., Salmon & Hyatt 1979), R R. Y. Backwell and<br />
her coworkers (e.g., Backwell et al. 2000), M. Murai and his coworkers (e.g., Murai et al. 2002),<br />
and T. Yamaguchi (e.g., Yamaguchi 2001a, b). Based on the studies of Uca and other brachyurans,<br />
as well as other decapods, Salmon (1983) reported the diversity of behavioral interactions<br />
preceding mating in decapods, and he defined some of the consequences of these interactions in<br />
terms of sexual selection, courtship behavior, and mating systems. The book edited by Reback &<br />
Dunham (1983), which included Salmon's (1983) work, was a landmark in the study of decapod<br />
behavior.<br />
Christy (1987) reviewed the mating systems of brachyuran crabs and classified them, according<br />
to modes of competition among males for females, into three major categories and eight subcategories,<br />
as follows.<br />
1. Female-centered competition, including: la, defense of mobile females following free search;<br />
lb, defense of sedentary females following a restricted search; lc, capture, carrying, and<br />
defense of females at protected mating sites; and Id, attraction and defense of females at<br />
protected mating sites<br />
2. Resource-centered competition, including: 2a, defense of breeding sites; and 2b, defense of<br />
refuges<br />
3. Encounter rate competition, including: 3a, neighborhoods of dominance; and 3b, pure search<br />
and interception<br />
In their book on crustacean sexual biology, Bauer & Martin (1991) introduced developments in<br />
various fields and taxa of crustacean research, including studies on sex attraction, sex recognition,<br />
mating behavior, mating system, and structure and function associated with insemination. Bauer and<br />
his coworkers have extensively studied the mating behavior, mating system, and hermaphroditism<br />
of shrimps (e.g., see Bauer 2004 for a review).<br />
Through their intensive studies on the mating system of the spider crab Inachus and of the extended<br />
maternal care of semi-terrestrial grapsid crabs of Jamaica, Diesel and his coworker revealed<br />
examples of highly specialized mating and social systems in these crabs (see Diesel 1991; Diesel &<br />
Schubart 2D07 for reviews).<br />
Thiel and his students have conducted intensive research on the mating system of rock shrimps<br />
(see Reference 6 in Appendix I, Table 4) and symbiotic anomuran crabs (e.g., Baeza & Thiel 2003).<br />
Based on these studies, Thiel & Baeza (2001) and Baeza & Thiel (2007) reviewed factors affecting<br />
the social behavior of marine crustaceans living symbolically with other invertebrates. Similarly,<br />
Correa & Thiel (2003) reviewed mating systems in caridean shrimp and their evolutionary consequences<br />
for sexual dimorphism and reproductive biology. The book by Duffy & Thiel (2007) on the<br />
evolutionary ecology of social and sexual systems of crustaceans is a monumental landmark that<br />
synthesizes the state of the field in crustacean behavior and sociobiology and places it in a conceptually<br />
based, comparative framework. The relatively recent discovery of eusociality in snapping<br />
shrimp by Duffy has opened the door to a new field in social and mating systems of decapods (see<br />
Duffy 2007 for a review; see also sections 3.5 Eusocial type and 4.5 Evolution of the eusocial type<br />
below for further explanation).
124 Asakura<br />
Asakura (1987, 1990, 1993, 1994, 1995, 1998a, 1998b, 1999, 2001a, b, c), Imazu & Asakura<br />
(1994, 2006), and Nomura & Asakura (1998) reported mating systems and various aspects of sexual<br />
differences in the ecology and behavior of hermit crabs and other decapods.<br />
3 TYPES OF MATING SYSTEMS<br />
3.1 Short courtship type<br />
This type is generally seen in species whose males and females are free living, that is, not symbiotic<br />
with other organisms (Appendix 1, Tables 1, 2). Copulation occurs after a short courtship behavior<br />
by the male, or copulation occurs just after brief behavioral interactions between a male and a<br />
female. This type of courtship includes very different groups of decapods, from the most primitive<br />
group (dendrobranchiate shrimps) to groups specialized for certain habitats such as freshwater crayfishes,<br />
intertidal hermit crabs, and semi-terrestrial and terrestrial brachyuran crabs. It is perhaps the<br />
most widely seen mating system in decapods.<br />
No intensive aggressive behavior between males (for a female) has been reported in species of<br />
dendrobranchiate shrimps of the families Penaeidae and Sicyoniidae, caridean shrimps of the families<br />
Palaemonidae, Hyppolytidae, and Pandalidae, or anomuran sand crabs of the family Hippidae.<br />
In these species, females are generally similar in size to, or larger than, males. On the other hand,<br />
strong aggressive interaction is seen between males in freshwater crayfish species of all three families<br />
(Astacidae, Parastasidae and Cambaridae) as well as in brachyuran crabs of the Grapsoidea and<br />
Gecarcinidae. In these species, the male body and weaponry (chelipeds) are generally larger than<br />
the female.<br />
Among decapods exhibiting this mating system are species whose females molt before copulation<br />
(Appendix 1, Table 1) and those whose females do not molt before copulation (Appendix 1,<br />
Table 2). In species inhabiting terrestrial and semi-terrestrial habitats, females generally copulate<br />
in the hard shell condition; these species include land hermit crabs of the genus Coenobita and<br />
brachyuran crabs of the Grapsoidea and Gecarcinidae.<br />
In penaeid shrimp, the molting condition of copulating females is determined according to the<br />
type of thelycum. The thelycum is the female genital area, i.e., modifications of female thoracic<br />
sternites 7 and 8 (sometimes including thoracic sternite 6) that are related to sperm transfer and<br />
storage. A female with externally deposited spermatophores is said to have an "open thelycum,"<br />
which is formed by modifications of the posterior coxae and sternites to which the spermatophores<br />
attach. Primitive dendrobranchiate shrimps, including species of the families Aristeidae, Solenoceridae,<br />
Benthesicymidae, and the penaeid genus Litopenaeus, have open thelyca. In these species,<br />
females copulate in the hard shell condition. On the other hand, a "closed thelycum" refers to sternal<br />
plates that may (1) enclose a noninvaginated seminal or sperm receptacle, (2) cover a space that<br />
leads to spermathecal opening, or (3) form an external shield guarding the spermathecal openings.<br />
In the most advanced groups, including the penaeoid genera Fenneropenaeus, Penaeus, Farfantepenaeus,<br />
Melicertus, Marsupenaeus, Trachypenaeus, and Xiphopenaeus, females have closed thelyca.<br />
In these species, females molt just before copulation. Since no significant difference is seen in mating<br />
behavior between the open thelycum species and the closed thelycum species, Hartnoll's (1969)<br />
rule, which predicts a lengthy pre-molt courtship behavior associated with soft-female mating and a<br />
relatively brief courtship behavior with hard-female mating, does not hold in the case of the penaeid<br />
shrimps.<br />
A sperm plug, which is believed to preclude subsequent insemination by other males, is known<br />
in some species of Farfantepenaeus, Marsupenaeus, Metapenaeus, and Rimapenaeus (Appendix 1,<br />
Table 3).<br />
In all the above-mentioned taxa, copulation generally continues only for several minutes. After<br />
mating, the male separates from the female and presumably goes on to search for other females.
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 125<br />
The habitat of species that exhibit this mating system varies, ranging from terrestrial through intertidal<br />
to deep water.<br />
3.2 Precopulatory guarding type<br />
This mating system also is generally seen in species whose males and females are free living<br />
(Appendix 1, Table 4). A male guards a mature female for one to several days before copulation.<br />
Generally, males aggressively fight for a female using their cheliped(s) and sometimes also the ambulatory<br />
pereopods. In some species, females always molt prior to mating and copulation; in other<br />
species, females may or may not molt prior to copulation. There are two types of guarding: (1) contact<br />
guarding of hermit crabs and brachyuran crabs, in which a male grasps part of the appendages,<br />
the body, or the shell (in the case of hermit crabs) of a mature female, and (2) non-contact guarding,<br />
as exhibited in Macrobrachium shrimps and Homarus lobsters, in which a male keeps a female<br />
without grasping her. After mating, postcopulatory guarding by a male for a female is sometimes<br />
observed (Appendix 1, Table 5). However, after postcopulatory guarding, or just after copulation,<br />
the male and female separate so that both may later mate with other individuals. Generally, in this<br />
mating system, the body size of males is larger than that of females, or weaponry (chelipeds) is<br />
more developed in males than in females.<br />
Species of the river prawn genus Macrobrachium are well known for the extremely long chelipeds<br />
in males. A male guards a female for one to several days before copulation and fights with<br />
other males using these chelipeds. In some species, such as M. australiense, a male has a nest (a<br />
saucer-shaped depression on the bottom), beckons a female to the nest, and guards and copulates<br />
with her in the nest. In the American lobster Homarus americanus, a male guards a female in his<br />
shelter, which is dug under rocks, boulders, or eelgrass, and the cohabitation of a male and a female<br />
lasts from one to three weeks.<br />
In hermit crabs of the genus Diogenes (Diogenidae) and in many species of the family Paguridae,<br />
al of which have unequal chelipeds in terms of both size and morphology, a male grasps the rim<br />
of the shell inhabited by a mature female by the minor cheliped, guards her for one to several<br />
days before copulation, and fights with other males approaching him using the major cheliped.<br />
In crab-shaped anomurans, the male Paralithodes brevipes conducts both pre-copulatory and postcopulatory<br />
guarding. The male claims a female by grasping her chelae or legs with his chelae, or<br />
he covers the female with his body. Similarly, the male Hapalogaster dentata grasps a female with<br />
his left chela and covers the female with his body; these guarding behaviors occur one to three days<br />
before copulation.<br />
In the brachyuran crab Corystes cassivelaunus (Corystidae), the male carries the female in his<br />
chelae, and, while stationary, holds one or both of the female's chelae in his own and holds her<br />
carapace close to his sternum. Such behavior continues up to several days before copulation. In<br />
species of the Cancridae and Portunidae, males carry the pre-molt female with her carapace or sternum<br />
held against the sternum of the male for a period of days; after this period the male releases<br />
the female so that she molts, and copulation occurs shortly after the molting. In many species in<br />
these two families, the male continues to carry the female after copulation in the pre-molt position<br />
until her integument has partially hardened. Sperm plugs, which are regarded as being produced<br />
by the males to block the females' genital duct to preclude subsequent insemination by other males<br />
(Diesel 1991), also are often reported for species of these families (Appendix 1, Table 6). In Menippe<br />
mercenaria (Xanthidae), the male guards the entrance to the burrow occupied by the pre-molt female,<br />
and they copulate as soon as the female molts. In species of the Majidae and Cheiragonidae,<br />
the male guards the female before copulation in a manner similar to what is seen in the Cancridae<br />
and Portunidae, where the male grasps the ambulatory pereopods, chelipeds, or body of the<br />
female.<br />
Species that exhibit this mating system are from the intertidal through shallow water to deep<br />
waters, but they are not found in terrestrial or semi-terrestrial environments.
126 Asakura<br />
3.3 Podding<br />
In large decapods inhabiting shallow waters, an aggregation consisting of an extremely large number<br />
of individuals in certain places is called a "pod." Podding is regarded as a type of behavior that is<br />
optional and that is associated with different stages in the species' life history, such as molting, mating,<br />
and the incubation period (Appendix 1, Table 7). The pod is also called a "heap" or "mound,"<br />
according to the locality and/or the species.<br />
The function of the pod may vary depending on the condition of the specimens within it (such as<br />
level of maturity, sex, intermolt stage) and possibly on changes in habitat condition, such as water<br />
temperature and presence of predators (Sampedro & Gonzalez-Gurriaran 2004). However, as listed<br />
in Appendix 1, Table 7, pods in some species have the function of facilitating mating, so I will treat<br />
this as a special kind of mass mating in some species.<br />
Stevens (2003) and Stevens et al. (1994), reporting more than 200 pods with a total of 100,000<br />
crabs of the majid Chionoecetes bairdi in an area of only 2 ha off Kodiak Island in Alaska in 1991,<br />
observed that the formation of the pods and mating synchronized with the spring tide. Similar observations<br />
were made for another majid, Hyas lyratus, by Stevens et al. (1992), who reported large<br />
aggregations during the mating season from off Kodiak Island. They found 200 mating pairs (males<br />
grasping females) among 2000 individuals in one pod. The majid crab Loxorhynchus grandis, distributed<br />
along the east coast of North America, often forms large aggregations numbering hundreds.<br />
of animals. The aggregation is composed of crabs of both sexes, and the function is thought to be<br />
the attraction of males for mating (Hobday & Rumsey 1999). DeGoursey & Auster (1992) reported<br />
large mating aggregations in another majid crab, Libinia emarginata, in April and May 1989. Many<br />
mating pairs were found in the aggregations, and the percentage of ovigerous females among all<br />
females increased from 26% on 1 May to 100% on 14 May. Males paired with females were significantly<br />
larger than unpaired males, while the paired and unpaired females were not significantly<br />
different in size. Carlisle (1957) monitored a pod consisting of 60-80 individuals of the majid crab<br />
Maja squinado in shallow waters in the English Channel; 20 were adult males and the rest were<br />
juvenile males and females in equal amounts. He observed crabs molting inside the pod and mating<br />
between intermolt males and postmolt females, which led him to conclude that the main purpose<br />
of podding is to provide protection for newly molted soft crabs against predators and to facilitate<br />
mating. However, later behavioral observations by Hartnoll (1969) indicated that copulation occurs<br />
between a male and a female in the intermolt stage. Furthermore, Sampedro & Gonzalez-Gurriaran<br />
(2004) found that the gonads of females in the pods were in an early stage of development (= not<br />
fully matured) and that the spermathecae were empty, suggesting to them that mating of this species<br />
occurs in deeper waters.<br />
In crab-shaped anomurans, large pods of the red king crab Paralithodes camtschaticus are well<br />
known in the northern Pacific Ocean, with each pod consisting of thousands of crabs in the 2-4<br />
year class (juveniles). Aggregations of adult red king crabs (ovigerous females) also were reported<br />
and are thought to be related to mating (Stone et al. 1993), but detailed surveys have not been<br />
conducted. Dense aggregations of the southern king crab Lithodes santolla have been reported from<br />
Chile (South America); however, the crabs forming these aggregations are juveniles, so this behavior<br />
is not thought be related to mating (Cardenas et al. 2007).<br />
In summary, podding is known only in large species distributed in temperate or boreal waters in<br />
both the Pacific and Atlantic oceans.<br />
3.4 Pair-bonding type<br />
Many species of decapods, in particular those that are symbiotic with other animals, have been<br />
reported as "found in a heterosexual pair" (Appendix 1, Tables 8-12). Most of these are considered
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 111<br />
to have a monogamous mating system, which is well known in birds and mammals. In species whose<br />
males engage in mate-guarding, temporal heterosexual pairing occurs, where the pair is formed<br />
when the female is close to molting or spawning a new batch of unfertilized eggs, and the mateguarding<br />
males abandon the females soon after the eggs are fertilized. However, in pair-bonding<br />
species, males cohabit with females, independent of their reproductive status or of the stage of<br />
development of the brooded embryos. Nevertheless, the observations for the monogamous nature<br />
of these pair-bonding species are often only anecdotal, and how long the pair remains together, and<br />
with whom they mate, is rarely recorded. Some well-documented studies include the formation of<br />
stable pairing and individual recognition (individuals in a pair can recognize each other as mates),<br />
as in the case of the banded shrimp Stenopus hispidus (Reference 8 in Appendix 1, Table 10), the<br />
scarlet cleaner shrimp Lysmata debelius (Reference 12 in Appendix 1, Table 10), and the harlequin<br />
shrimp Hymenocera picta (Reference 16 in Appendix 1, Table 10).<br />
Detailed observations of the monogamous nature of pairing have been made for several species<br />
of snapping shrimps, for example, Alpheus angulatus (Reference 97 in Appendix 1, Table 9),<br />
Alpheus heterochaelis (Reference 99 in Appendix 1, Table 9), Alpheus armatus (Reference 28 in<br />
Appendix 1, Table 9), and Alpheus roquensis (Reference 31 in Appendix 1, Table 9), as well as for<br />
the pontoniid shrimp Pontonia margarita (Reference 45 in Appendix 1, Table 8), the deep-water<br />
sponge-dwelling shrimp Spongicola japonica (Reference 1 in Appendix 1, Table 10), a porcelain<br />
crab Poly onyx gibbesi (Reference 11 in Appendix 1, Table 11), and several species of coral crabs<br />
of the genus Trapezia (References 2-14 in Appendix 1, Table 12). Many pair-bonding species are<br />
known in caridean shrimps of the subfamily Pontoniinae and family Alpheidae, "cleaner" shrimps<br />
of the families Stenopodidae and Spongicolidae, crab-shaped anomurans (family Porcellanidae),<br />
and brachyuran crabs of the family Trapeziidae.<br />
Most of these species are symbiotic with other animals or live in special habitats. Host animals<br />
for these species include sponges, sea anemones, black corals, reef-building corals, gastropods,<br />
opistobranch molluscs, bivalves, polychaetes, crinoid feather stars, sea stars, sea urchins, sea cucumbers,<br />
and ascidians. The special habitats include gastropod shells used by large hermit crabs; tubes<br />
of polychaetes such as Chaetopterus; soft, web-like tubes consisting of filamentous algae, sponges,<br />
and other debris built by shrimp themselves; burrows excavated in hard dead corals; burrows of gobiid<br />
fish; and burrows of the thalassinidean shrimp genus Upogebia. However, free-living species are<br />
also known, such as stenopodid shrimps inhabiting rocky subtidal zones and many alpheid shrimp<br />
species inhabiting rock crevices or found under rubble, around large algae, or in burrows of their<br />
own in mudflats and other soft bottoms.<br />
The following generalizations can be made for almost all of these species. They are territorial,<br />
and they cooperatively defend their habitats (hosts, special habitats, and burrows) against other conspecific<br />
or non-conspecific animals. Thus, the mating system of these species is termed "resourcedefense<br />
monogamy." The pairs are size-matched (— size-assortative pairing); there is strict preference<br />
exerted by either sex for mates of a particular size relative to themselves. Baeza (2008) proposed<br />
two possible explanations for this phenomenon in his study on pontoniid shrimps symbiotic<br />
with bivalves:<br />
1. The two sexes might choose large individuals of the opposite sex as sexual partners and host<br />
companions. In males, a preference for large females should be adaptive, as female size is<br />
positively correlated with fecundity in shrimps. In females, sharing a host with a large male<br />
might result in indirect benefits (i.e., good genes) or direct benefits (increased protection<br />
against predators or competitors).<br />
2. Choice of a certain-size partner could also be a consequence of constraints in the growth rates<br />
of shrimps dictated by host individuals. Space limitations for shrimps in hosts are suggested<br />
by the tight relationship between shrimp and host size, and by the fact that hosts harboring<br />
solitary or no shrimps were among the small hosts.
128 Asakura<br />
These species tend to display low sexual dimorphism in weaponry in terms of cheliped size and morphology<br />
and often in body size. This is in contrast to the large sexual differences in mate-guarding<br />
species in which the weaponry is much more developed and where body size is often much larger<br />
in males than in females. Regarding body size, there is a tendency in pair-bonding shrimp for the<br />
male to be slightly smaller, in terms of body length, and much more slender than its mate female;<br />
in trapeziid crabs the male is often slightly larger than his female mate.<br />
The bathymetric distribution of species with this mating system is generally from intertidal to<br />
shallow water, but a few groups of species, such as those of the Spongicolidae, inhabit deep water.<br />
3.5 Eusociality type<br />
Until the discovery of the eusocial shrimp Zuzalpheus regalis (as Synalpheus regalis) (Duffy 1996),<br />
eusociality was recognized only among social insects, including ants, bees, and wasps (Hymenoptera)<br />
and termites (Isoptera); in gall-making aphids (Hemiptera); in thrips (Thysanoptera); and in<br />
two mammal species, the naked mole rat (Heterocephalus glaber) and the damaraland mole rat<br />
(Cryptomys damarensis). Zuzalpheus regalis lives inside large sponges in colonies of up to >300<br />
individuals, with each colony containing a single reproductive female. Direct-developing juveniles<br />
remain in the natal sponge, and allozyme data indicate that most colony members are full siblings.<br />
Larger members of the colony, most of whom apparently never breed, defend the colony against<br />
heterospecific intruders (Duffy 1996).<br />
Following this initial discovery, Duffy and his coworkers have found several other species<br />
of Zuzalpheus exhibiting monogynous, eusocial colony organization in the western Atlantic<br />
(Appendix 1, Table 13). In the Indo-west Pacific region, Didderen et al. (2006) found a colony<br />
of a sponge-dwelling alpheid shrimp, Synalpheus neptunus neptunus, with one large ovigerous female<br />
or "queen" together with many small individuals, indicating a eusocial colony organization<br />
(Appendix 1, Table 13).<br />
Some 20 species of symbiotic decapod species have been reported as found in a group<br />
(Appendix 1, Tables 14-15). Among them, examples of Synalpheus and Zuzalpheus exhibited more<br />
than 100 individuals in one aggregation, and, in particular in the case of Zuzalpheus brooksi, more<br />
than 1000 individuals were recorded from one sponge. These aggregations are regarded either as<br />
having a non-social structure (Thiel & Baeza 2001) or with the social structure totally unknown.<br />
3.6 Waving display type<br />
In many species of the crab families Ocypodidae, Dotillidae, and Macrophthalmidae, and in species<br />
of the genus Metaplax of the family Varunidae (formerly subfamily Varuninae in the Grapsidae<br />
sensu lato), males perform waving displays using the chelipeds. As in many other territory advertisement<br />
signals in animals, this behavior is commonly thought to have the dual function of simultaneously,<br />
repelling males and attracting females (e.g., Salmon 1987; Crane 1975). These species<br />
typically live in mudflats, tidal creeks, sandbars, and mangrove forests, and each individual has its<br />
own burrow with a small territory around it. They often occur in huge numbers, with thousands of<br />
individuals living in small, adjacent territories, and with males and females living intermixed. The<br />
burrow serves various functions, including a refuge during high tide, an escape from predators, and<br />
the site of mating, oviposition, and incubation.<br />
The behavior and mating systems of fiddler crabs (genus Uca, Ocypodidae) have been intensively<br />
studied (see references in History of Study, above). There are species whose males defend<br />
burrows from which they court females and species whose males wander from their burrows and<br />
court females on the surface (Christy 1987). For the former group of species, the following generalization<br />
is possible (based mainly on P. Backwell and coworkers; see references in History of Study,<br />
above). Males wave their enlarged claw, and, when a female is ready to mate (i.e., she matures),<br />
she leaves her own burrow and wanders through the population of waving males. The female visits
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 129<br />
several males before selecting a mate, and a visit consists of a direct approach to the male. Before<br />
copulation^ both individuals enter the male's burrow, and two behavioral patterns are known: the<br />
male enters his burrow first and the female follows him in, or it happens in the reverse order, i.e.,<br />
the female enters first. The male then gathers up sand or mud to plug the burrow entrance. Mating<br />
occurs in the burrow. On the following day, the male emerges, reseals the burrow entrance with the<br />
female still underground, and leaves the area. The female remains underground for the following<br />
few weeks while she incubates her eggs.<br />
In addition to waving displays, males of some fiddler crab species employ acoustic signals to<br />
attract females. In these species, males attract females during the day first by waving and then by<br />
producing sounds just within their burrows. At night, the males produce sounds at low rates, but<br />
when touched by a female they increase their rate of sound production (Salmon & Atsaides 1968).<br />
Many species of ocypodid crabs build sand structures next to their burrows, some of which function<br />
to attract females for mating, such as pillars (Uca: Christy 1988a, b), hoods (Uca: Zucker 1974,<br />
1981; Christy et al. 2002, 2003a, b), mudballs (Uca: Oliveira et al. 1998), and pyramids (Ocypode:<br />
Linsenmair 1967; Hughes 1973).<br />
3.7 Visiting type<br />
An interesting mating system has been suggested for coral gall crabs (family Cryptochiridae), which<br />
inhabit cavities in scleractinian corals in (usually) shallow water. However, the information is still<br />
anecdotal, based on ecological observations on Hapalocarcinus marsupialis, Troglocarcinus corallicola,<br />
and Opecarcinus hypostegus (Potts 1915; Fize 1956; Kropp & Manning 1987; Takeda &<br />
Tamura 1981; Hiro 1937; Kotb & Hartnoll 2002; Carricart-Ganivet et al. 2004). In H. marsupialis<br />
and T. corallicola, the male crab normally resides outside the gall, which was constructed by the<br />
female, and is thought to visit the gall of the female for mating. The males and females apparently<br />
show promiscuity, and male-male aggressive behavior for a female has not been reported. The female<br />
is much larger than the male and in some species has a soft body with a very large abdomen. On<br />
thfe other hand, the male is usually hard, with a small abdomen. Geographical distribution includes<br />
mostly the tropics (see Wetzer et al. this volume).<br />
In Opecarcinus hypostegus, couples were found sharing cavities; ovigerous females and males<br />
are recorded inhabiting adjoining cavities on colonies of Siderastrea stellata corals (Carricart-<br />
Ganivet et al. 2004). This species may have a mating system different from the above.<br />
3.8 Reproductive swarm type<br />
This mating system is reported only in pinnotherid crabs that are considered parasitic or co-inhabiting<br />
with other animals, including bivalves, gastropods, sea slugs, chitons, polychaetes, echinoderms,<br />
burrowing crustaceans, and sea squirts (Cheng 1967; Gotto 1969). In several species of these crabs,<br />
mating occurs, or is thought to occur, when the female is in the free-swimming stage before she<br />
enters into her definitive host (Appendix 1, Table 16).<br />
The following generalization is possible for these species. Adult females have a soft, membranous<br />
carapace, and generally each one lives by itself within its host animal. These females produce<br />
broods of planktonic larvae. After development, the larvae metamorphose into the "invasive stage"<br />
crab, which is morphologically similar to the later swimming stage in having a flattened shape and<br />
ambulatory legs with dense setae adapted for swimming. Following this stage is a stage designated<br />
as "prehard"; these crabs invade, and live in, the host invertebrate animals. The crab at this stage<br />
is soft, resembling the later posthard stage. These crabs grow and mature into small adults of both<br />
sexes and leave their host to join mating swarms in open water. This stage is called the "hard stage,"<br />
swimming stage, or copulation stage, and it is characterized by a hard body, swimming legs densely<br />
fringed with setae, and a thick fringe of setae along the front of the carapace. They copulate at<br />
this stage, and, in all reported species (see Appendix 1, Table 16), females copulate in the hard
130 Asakura<br />
shell condition. After copulation, each female enters the host animal, but the male dies. The female<br />
becomes soft and grows much larger in the host, and later the female produces eggs fertilized by<br />
sperm from her single mating.<br />
This is a kind of mass mating, with males and females showing promiscuity. In the copulation<br />
stage, no intensive aggressive behavior between males for females has been reported. The males<br />
in this stage are slightly larger than the females, and the morphology is similar between the sexes.<br />
After the female enters the host animal, the female becomes soft and grows much larger and stouter.<br />
The species with this mating system are found generally from intertidal to shallow water where their<br />
host invertebrates occur. In some pinnotherid species, adult crabs are found in a heterosexual pair in<br />
the host animal, although life history and mating systems of these species are mostly unknown.<br />
3.9 Neotenous male type<br />
Extremely small, neotenous males exist in some species of anomuran sand crabs (genus Emerita)<br />
inhabiting wave-exposed sandy beaches in tropical and temperate waters (Appendix 1, Table 17). In<br />
these species, the males become sexually mature soon after their arrival on the beach as a megalopa.<br />
When copulating, a male attaches near one of the female's gonopores, which are located on the<br />
coxae of the third pereopods. Surprisingly, the size of the neotenous males is similar to, or smaller<br />
than, those coxae.<br />
Protandric hermaphroditism is described in detail in Emerita asiatica as it relates to neotenous<br />
males (Subramoniam 1981). The neotenous males occur at 3.5 mm carapace length (CL) and above,<br />
whereas females acquire sexual maturity at 19 mm CL. The neotenous males, as they continue<br />
to grow, gradually lose male functions and reverse sex at about 19 mm CL. In the CL range of<br />
19-22 mm, the male's gonad consists of inactive testicular and active ovarian portions. Androgenic<br />
glands, active in the neotenous males, show signs of degeneration in the larger males and disappear<br />
in the intersexuals.<br />
The male separates from the female after copulation. Aggressive behavior between males is not<br />
reported. As opposed to the female, the neotenous male shows a general simplicity of appendages<br />
associated with its small size. Among decapods, this phenomenon is known only in species of<br />
Emerita.<br />
4 EVOLUTION OF MATING SYSTEMS IN DECAPODA<br />
4.1 Introduction<br />
It is apparent from the above that similar mating systems have evolved independently in different<br />
taxa at different times; i.e., convergent evolution is widespread. Species in ecologically similar<br />
habitats often display patterns that are strikingly comparable. Here I discuss the possible origin and<br />
evolutionary pathway of each mating system and compare them with those of other animals.<br />
4.2 Evolution of the short courtship type and the precopulatory type<br />
These two mating systems are most dominant among decapods. The mode of life is often quite<br />
similar; both males and females are free living (not symbiotic with other organisms), and after<br />
mating the male soon separates from the female. However, the habitat is sometimes different; in<br />
terrestrial and freshwater species, only the short courtship type has been reported. Therefore, a<br />
question arises as to why some groups of species have evolved the prolonged precopulatory mate<br />
guarding, whereas others have not.<br />
Precopulatory mate guarding is known in a very broad range of taxa such as tardigrades, crustaceans,<br />
arachnids, and anurans (Parker 1974; Ridley 1983; Conlan 1991). It is thought to evolve<br />
when male-male competition for females is strong enough and female receptivity is restricted in
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 131<br />
time (Parker 1974; Jormalainen 1998), or even if receptivity is not time-limited but the guarding<br />
costs are low enough (Yamamura 1987). Guarding should be beneficial to the male, if the expected<br />
fitness gain achieved by guarding is greater than that expected by continuing to search for other<br />
females (Parker 1974). Thus, the optimal guarding duration for the male is determined by the encounter<br />
rate of females and the costs of guarding relative to those of searching (Yamamura 1987).<br />
The cost of guarding for males includes decreased mobility and feeding (Adams et al. 1985, 1991;<br />
Robinson & Doyle 1985), an increase in predation risk while guarding (Verrel 1985; Ward 1986), increased<br />
energetic costs associated with carrying females (Sparkes et al. 1996; Plaistow et al. 2003),<br />
and an increase in fighting costs through male-male conflict (Benesh et al. 2007; Yamamura &<br />
Jormalainen 1996). Additionally, a long guarding time decreases future opportunities to mate with<br />
other females (Benesh et al. 2007).<br />
Pelagic dendrobranchiate and caridean shrimps are primarily swimmers, and possibly for that<br />
reason they have not evolved prolonged, elaborate behavioral interactions before copulation. However,<br />
the above-mentioned energetic cost hypothesis (Sparkes et al. 1996; Plaistow et al. 2003) may<br />
be applicable; for males of these species, carrying a swimming female for a long duration requires<br />
much more energy than in benthic species. In fact, all species exhibiting a prolonged precoulatory<br />
guarding period are benthic species.<br />
In all freshwater crayfish studied, the mating system includes a short courtship without a lengthy<br />
precopulatory guarding, even though they have a benthic lifestyle and male-male aggression is<br />
often common. They may live in their burrows separately, or underneath boulders or heaps of fallen<br />
leaves, and these habitats are quite similar to, or virtually the same as, those of shrimps of the genus<br />
Macrobrachium. Why males of Macrobrachium adopt a precopulatory guarding strategy whereas<br />
male crayfish do not is not known.<br />
A similar question arises in intertidal and shallow water decapods. For example, intertidal hermit<br />
crab species exhibiting precopulatory guarding have a tendency toward vastly unequal chelipeds,<br />
with a well-developed major cheliped particularly in males, who use it for fighting with<br />
olher males during guarding. Such species include those of the genera Pagurus (Paguridae) and<br />
Dmgenes (Diogenidae). On the other hand, species of Paguristes have small and similar right and<br />
left chelipeds and execute short courtship mating; males do not aggressively fight with other males.<br />
Species of Calcinus, which conduct short courtship type mating, often have vastly unequal chelipeds,<br />
with the well-developed major cheliped similar to those species that display precopulatory<br />
guarding. However, males of Calcinus species do not aggressively fight with each other during mating.<br />
Further study is needed to clarify the relationship between mating behavior and morphology.<br />
In land hermits and land brachyurans, the above-mentioned predation risk hypothesis (Verrel<br />
1985; Ward 1986) may be applicable to those species where mating system is the short-courtship<br />
type with hard-female mating. Male-male aggression is common in these taxa, but they have never<br />
evolved precopulatory guarding. Prolonged guarding may carry the risk of attack by visual predators<br />
such as birds in a terrestrial environment. In these taxa, a strong connection exists between a<br />
prolonged precopulatory guarding and soft-female mating as well as between a short courtship and<br />
hard-female mating. When marine species adapted to land, the former mating system might have<br />
been lost and changed to the latter, i.e., from soft-female to hard-female, to avoid desiccation and<br />
to deal with the large and often unpredicted fluctuations in availabilities of females in a terrestrial<br />
environment.<br />
The evolution of sperm plugs in species of short-courtship type (penaeid shrimps) and precopulatory<br />
type (brachyuran crabs) is interesting. The sperm plug has virtually the same function as<br />
the copulation plug (= copulatory plug, mating plug) in mammals (rodents, bats, monkeys, koala),<br />
reptiles (snakes and lizards), insects (butterflies, ants, dragonflies, and stinkbugs), spiders, and acanthocephalan<br />
worms (Smith 1984). These plugs, secreted by the male after mating, serve to block the<br />
female tract for some time to prevent further mating by other males.
132 Asakura<br />
4.3 Evolution of the podding type<br />
Why many animal species (e.g., insects, fish, birds, and herbivorous mammals) group together is<br />
one of the most fundamental questions in evolutionary ecology. It is believed that strong selective<br />
pressures lead to aggregation rather than to a solitary existence in most of these groups. These pressures<br />
include protection against predators, increased foraging efficiency, increased ease of assessing<br />
potential mates, and increased information exchange about the location of food (Barta & Giraldeau<br />
2001). Similarly, various ecological reasons for the formation of pods have been proposed, including<br />
protection during molting, location of mates, aiding in food capture, and protection from predation<br />
(see References in Appendix 1, Table 7). Why some species evolved aggregating behavior and others<br />
did not is unknown.<br />
4.4 Evolution of the pair-bonding type<br />
Heterosexual pairing behavior ("social monogamy," Gowaty 1996; Bull et al. 1998; Gillette et al.<br />
2000; Wickler & Seibt 1981) has evolved many times in a broad range of animal taxa, including<br />
mammals, birds, reptiles, amphibians, fish, insects, and crustaceans. For example, a colony of<br />
scleractinian coral sometimes yields a pair of goby fish, alpheid shrimps, and trapeziid crabs. Researchers<br />
interested in social system evolution must look for ecological and physiological factors<br />
(beyond basic sexual differences) that may make social monogamy selectively advantageous to individual<br />
males and/or females. Of particular interest are factors that may consistently correlate with<br />
such behavior across taxonomic groups. Several hypotheses for the evolution of social monogamy<br />
have been developed [see also Mathews (2002b), Baeza (2008), Baeza & Thiel (2007) for a review],<br />
as follows.<br />
Biparental care hypothesis: Kleiman (1977) argued that the advantages of monogamy in mammals<br />
can lead to social monogamy. The hypothesis also implies that both males and females would<br />
suffer significantly reduced or zero fitness if they did not cooperate in caring for the offspring. However,<br />
this is not the case for marine decapods, where only the females care for the fertilized eggs<br />
and where neither parent cares for the larvae.<br />
Extended mate guarding hypothesis: If males are under selection to guard females for some<br />
time before, during, and/or after courtship and mating, they may be forced into partner-exclusive<br />
behavior by some other factor, such as female dispersion (Kleiman 1977; Wickler & Seibt 1981)<br />
or female-female aggression (Wittenberger & Tilson 1980). In other words, monogamy can result<br />
from males guarding females over one or multiple reproductive cycles, because the female's synchronous<br />
receptivity, density, or abundance relative to males renders other male mating strategies<br />
(pure searching) less successful (Parker 1970; Grafen & Ridley 1983).<br />
Territorial cooperation hypothesis: The fact that most monogamous species are territorial leads<br />
to this hypothesis. Territoriality correlates in various ways with social system evolution (Emlen &<br />
Oring 1977; Hixon 1987), and cooperation in territorial defense can lead to individual advantages<br />
in social groups or pairs (Brown 1982; Davies & Houston 1984; Fricke 1986; Clifton 1989, 1990;<br />
Farabaugh et al. 1992). In other words, males and females benefit by sharing a refuge (a territory)<br />
as heterosexual pairs because, for example, the risk of being evicted from the territory by intruders<br />
decreases (Wickler & Seibt 1981).<br />
Recent intensive behavioral studies in various species shrimps have supported the predictions of<br />
the mate-guarding and/or territorial cooperation hypotheses (e.g., in Hymenocera picta, Wickler &<br />
Seibt 1981; Alpheus angulatus, Mathews 2002a, b, 2003; and Alpheus heterochelis, Rahman et al.<br />
2002,2003).<br />
Another hypothesis about social monogamy (Baeza & Thiel 2007) concerns species symbiotic<br />
to other organisms (= host). Baeza & Thiel predicted that monogamy evolved when hosts are<br />
small enough to support few individuals and are relatively rare, and when predation risk away<br />
from the hosts is high. Under these circumstances, movements among hosts are constrained, and
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 133<br />
monopolization of hosts is favored in males and females due to their scarcity and because of the<br />
host's value in offering protection against predators. Because spatial constraints allow only a few<br />
adult symbiotic individuals to cohabit in/on the same host, both adult males and females would<br />
maximize their reproductive success by sharing "their" dwelling with a member of the opposite sex.<br />
This hypothesis was supported by Baeza's (2008) intensive study on a heterosexual pair of Pontonia<br />
margarita, a species symbiotic to the pearl oyster.<br />
However, as mentioned before, most of observations for this mating system are anecdotal,<br />
and further detailed study is needed to clarify actual conditions of monogamous features of those<br />
species.<br />
4.5 Evolution of the eusocial type<br />
Hypotheses explaining how eusociality has evolved include Trophallaxis Theory (Roubaud 1916),<br />
Parental Manipulation Theory (Michener & Brothers 1974), Superorganism Theory (Reeve &<br />
Holldobler 2007), and Inclusive Fitness Theory (Hamilton 1964a, b), of which the last one is most<br />
widely accepted. According to the Inclusive Fitness Theory, eusociality may evolve more easily in<br />
species exhibiting haplodiploidy, which facilitates the operation of kin selection. Although eusocial<br />
mole rats and termites exhibit diploidy, they display high levels of inbreeding by living as a family<br />
in a single burrow, such that colony members share more than 50% of their genes, and therefore<br />
the same model is considered to apply to these species and also to eusocial Zuzalpheus shrimps, in<br />
which all members of a colony share a single sponge.<br />
4.6 Evolution of the waving display type<br />
As compared to terrestrial species, courtship in aquatic species may be short and may not involve<br />
elaborate visual signaling (display) by the males; in aquatic species, chemical or visual cues are<br />
more important stimuli. In species of several genera of semi-terrestrial (-upper intertidal) decapods<br />
including Uca and other ocypodid crabs, visual signalling for prolonged periods is common, and<br />
sounds are often emitted by males to "call" females from their burrows to the surface for mating.<br />
Salmon •& Atsaides (1968) presented ecological arguments to account for these differences in terms<br />
of optimal strategy of distance communications in the terrestrial and aquatic environments. Most<br />
aquatic decapods are nocturnally active and cryptic and live in an acoustically noisy environment,<br />
and this situation virtually eliminates all but the chemical channel for effective distance communication.<br />
On the other hand, visual and acoustic signals are effective in terrestrial species and are well<br />
developed in most terrestrial animals such as insects, birds, mammals, and also ocypodid and other<br />
terrestrial and semi-terrestrial decapods, probably because of the greater visibility in the terrestrial<br />
environment.<br />
Waving displays seen in a variety of semi-terrestrial crabs is a case of convergent evolution<br />
(Kitaura et al. 2002). Grapsid crabs of the genus Metaplax conduct waving displays like species<br />
of the ocypodid crab genera Uca, Macrophthalmus, Scopimera, and Dottila (Kitaura et al. 2002).<br />
Species of Metaplax, unlike other grapsid crabs, which generally live along rocky shores, live in<br />
mud flats and burrow into the mud like many ocypodids. Salmon & Atsaides (1968) proposed the<br />
following factors as advantageous for the evolution of visual signaling in semi-terrestrial crabs: the<br />
substrate, which is flat and relatively free from the vegetational obstructions and other discontinuities;<br />
diurnal activity of the crabs; and the feeding proximity to their shelters, which leads crabs<br />
to live in aggregations so that social contacts are frequent. Therefore, it is assumed that habitat<br />
similarity between Metaplax and ocypodid crabs resulted in convergent evolution of these displays.<br />
A recent molecular phylogenetic analysis suggested that even the waving display in Uca has<br />
multiple origins (Sturmbauer et al. 1996). Indo-west Pacific Uca species have simpler reproductive<br />
social behaviors, are more marine, and were thought to be ancestral to the behaviorally more complex<br />
and more terrestrial American species. It was also thought that the evolution of more complex
134 Asakura<br />
social and reproductive behavior was associated with the colonization of the higher intertidal zones.<br />
However, Sturmbauer et al. (1996) demonstrated that species bearing the set of "derived traits" are<br />
phylogenetically ancestral, suggesting an alternative evolutionary scenario: the evolution of reproductive<br />
behavioral complexity in fiddler crabs may have arisen multiple times during their evolution,<br />
possibly by co-opting of a series of other adaptations for high intertidal living and antipredator<br />
escape.<br />
This mating system is quite similar to male-territory-visiting polygamy (Kuwamura 1996) in<br />
fish, in which many examples are known in intertidal or shallow species; males have a burrow or a<br />
territory, and, when a mature female approaches a male, the male changes the color of part of his<br />
body and/or conducts species-specific courtship displays, after which the female enters the burrow<br />
or territory of the male and spawns (e.g., Miyano et al. 2006). In these fish species, males are brilliantly<br />
colored, as are male Uca species.<br />
4.7 Evolution of the visiting type<br />
A widely recognized tendency among various kinds of animals is that females live in a particular<br />
place and have a narrow home range, whereas males have a comparatively wider home range<br />
(Clutton-Brock et al. 1982). This "visiting type" mating system (seen in cryptochirid crabs) probably<br />
has evolved as one extremity of this tendency, with females living in a very specialized habitat<br />
(inside coral galls).<br />
4.8 Evolution of the reproductive swarm type<br />
Surprisingly, the function of the reproductive swarm in pinnotherid crabs is very similar to that of<br />
the nuptial flight (mating swarm) in ants (Insecta, Formicidae), and indeed their life history is quite<br />
similar. In most species of ants, breeding females and males that mature in their mothers' nest have<br />
wings and, during the breeding season, fly away from their nests and form swarms. Mating occurs<br />
during this period, and the males die shortly afterward. The surviving females land, and each female<br />
digs a burrow for the new nest. As eggs are laid in the burrow, stored sperm, obtained during their<br />
single nuptial flight, is used to fertilize all future eggs produced.<br />
In the pinnotherids, crabs first grow in their host animals (vs. ants in their initial burrow). Then<br />
the crabs with swimming setae leave the hosts and swarm (vs. ants with wings fly away from their<br />
nests and conduct the nuptial flight). Mating occurs during this period (in ants, too), after which the<br />
female crabs enter the hosts, whereas the males die just after the mating (vs. the female ants make<br />
burrows of their own, with males dying just after the mating). As in the case of the ants, the female<br />
crabs reproduce by fertilizing their eggs with sperm from a single mating.<br />
4.9 Evolution of the neotenous male type<br />
The miniaturization of male mole crabs in the anomuran genus Emerita coupled with neoteny is<br />
similar to "dwarf males" (parasitic males, complemental males, miniature males), which are tiny<br />
males often attached to females. This condition has evolved in various groups of animals, including<br />
thoracican barnacles (Yamaguchi et al. 2007), acrothoracican barnacles (Kolbasov 2002), the oyster<br />
Ostreapuelchanas (Castro & Lucas 1987; Pascual 1997), epicaridean isopods (Mizoguchi et al.<br />
2002), an echiuran Bonellia (Berec et al. 2005), anglerfish (Lophiiformes) (Pietsch 2005), blanket<br />
octopus (Tremoctopodidae), argonauts (Argonautidae), football octopus (Ocythoidae), and a deeper<br />
water octopus Haliphron atlanticus (Alloposidae) (Norman et al. 2002). The evolutionary cause for<br />
these phenomena has not been fully studied. The neoteny of male Emerita is considered to be one<br />
rather radical evolutionary solution to the problem of keeping the male and female together in the<br />
harsh and turbulent surf zone environment (Salmon 1983; Subramoniam & Gunamalai 2003).
ACKNOWLEDGEMENTS<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 135<br />
I am deeply grateful to Joel W. Martin (<strong>Natural</strong> History Museum of Los Angeles County, U.S.A.)<br />
for giving me the opportunity to present my work at the symposium on decapod phylogenetics at<br />
the TCS Winter Meeting in San Antonio, Texas. Thanks are also due to the following persons who<br />
provided me with important literature or aided me in my bibliographical survey: Keiichi Nomura<br />
(Kushimoto Marine Park, Japan), Tomomi Saito (Port of Nagoya Public Aquarium), Annie Mercier<br />
(Memorial University of Newfoundland, Canada), Juan P. Carricart-Ganivet (El Colegio de la Frontera<br />
Sur, Unidad Chetumal, Mexico), Jorge Contreras Gardufio (Entomologia Aplicada, Instituto<br />
de Ecologfa, Mexico), Ana Maria S. Pires-Vanin (Instituto Oceanografico, Universidade de Sao<br />
Paulo, Brazil), J. Antonio Baeza (Smithsonian Tropical Research Institute, Republic of Panama),<br />
Martha Nizinski (NOAA/NMFS Systematics Laboratory, Smithsonian Institution, U.S.A.), Panwen<br />
Hsueh (National Chung Hsing University, Taiwan), Estela Anahi Delgado (Undecimar, Facultad<br />
de Ciencias, Uruguay), Michiya Kamio (Georgia State University, U.S.A.), Satoshi Wada<br />
(Hokkaido University, Japan), Yoichi Yusa (Nara Women's University, Japan), Tomoki Sunobe<br />
(Tokyo University of Marine Science and Technology), Charles H. J. M. Fransen (Nationaal Natuurhistorisch<br />
Museum <strong>Natural</strong>is, The Netherlands), E. Gaten (University of Leicester, U.K.), Gil<br />
G. Rosenthal (Texas A&M University, U.S.A.), and Hiromi Watanabe (JAMSTEC, Japan). Special<br />
thanks are due to Raymond Bauer (University of Louisiana Lafayette) and Joel W. Martin for the<br />
careful review of an earlier draft of the manuscript.<br />
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APPENDIX!<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 141<br />
Table 1. Species of the short courtship type, in which females molt before copulation (= soft-female mating<br />
sensu Hartnoll 1969).<br />
DENDROBRANCHIATA<br />
Penaeidae: Marsupenaeus japonicus (1), Melicertus kerathurus (2), Melicertus brasiliensis (3),<br />
Melicertus paulensis (4), Farfantepenaeus aztecus (5), Fenneropenaeus merguiensis(6),Penaeus<br />
monodon (7), Penaeus semisulcatus (8), Trachypenaeus similis (9), Xiphopenaeus sp. (10)*,<br />
Sieyoniidae: Sicyonia dorsalis (11), Sicyonia parri (12), Sicyonia laevigata (13)<br />
PLEOCYEMATA<br />
Caridea<br />
Palaemonidae: Palaemonetes vulgarus (14), Palaemonetes varians (15), Palaemonetes pugio<br />
(16), Palaemon serratus (17), Palaemon elegans (18), Palaemon squilla (19)<br />
Alpheidae: Athanus nitescens (20), Alpheus dentipes (21)<br />
Hippolytidae: Heptacarpus picta (22), Heptacarpus paludicola (23)<br />
Pandalidae: Pandalus dona (24), Pandalus platyceros (25), Pandalus borealis (26)<br />
Crangonidae: Crangon crangon (27), Crangon vulgaris (28)<br />
Astacidea<br />
Nephropidae: Nephrops norvegicus (29)<br />
Palinuridea<br />
Palinuridae: Jasus lalandii (30)*<br />
Anomura<br />
Hippidae: Emerita asiatica (31), Emerita analoga (32)<br />
Diogenidae: Calcinus latens (33), Calcinus seurati (34), Clibanarius tricolor (35), Clibanarius<br />
; antillensis (36), Clibanarius zebra (37), Paguristes cadenati (38), Paguristes tortugae (39),<br />
Paguristes anomalus (40), Paguristes hummi (41), Paguristes oculatus (42)<br />
*Hard-female mating was rarely reported in addition to the soft-female mating. References: (1) Hudinaga<br />
(1942 as Penaeus japonicus), (2) Heldt (1931 as Penaeus caramote), (3) Brisson (1986), (4) de<br />
Saint-Brisson (1985), (5)-(6) Aquacop (1977), (7) Primavera (1979), Aquacop (1977), (8) Browdy<br />
(1989), (9)-(10) Bauer (1991), (11) Bauer (1992, 1996), (12)-(13) Bauer (1991), (14) Burkenroad<br />
(1947), Bauer (1976), (15) Antheunisse et al. (1968), Jefferies (1968), (16) Berg & Sandifer (1984),<br />
Bauer & Abdalla (2001), Caskey & Bauer (2005), (17) Nouvel & Nouvel (1937), Forster (1951),<br />
Bauer (1976), (18) Hoglund (1943), (19) Hoglund (1943), Bauer (1976), (20) Nouvel & Nouvel<br />
(1937), (21) Volz (1938), (22) Bauer (1976), (23) Bauer (1979), (24) Needier (1931), (25) Hoffman<br />
(1973), (26) Carlisle (1959), (27) Nouvel (1939), (28) Lloyd & Young (1947), Havinga (1930),<br />
Bodekke et al. (1991), (29) Farmer (1974), (30) von Bonde (1936), Silberbauer (1971), McKoy<br />
(1979), (3 1) Menon (1933), Subramoniam (1979), (32) MacGinitie (1938), Efford (1965), (33) Hazlett<br />
(1972), (34) Hazlett (1989), (35)-(36) Hazlett (1966), (37) Hazlett (1966, 1989), (38)-(42)<br />
Hazlett (1966).
142 Asakura<br />
Table 2. Species of the short courtship type, in which females do not molt before copulation (= hard-female<br />
mating sensu Hartnoll 1969).<br />
DENDROBRANCHIATA<br />
Penaeoidea: Litopenaeus vannanmei (1), Litopenaeus setiferus (2), Litopenaeus stylirostris (3),<br />
Litopenaeus schmitti (4)<br />
PLEOCYEMATA<br />
Astacidea<br />
Astacidae: Pacifastacus trowbridgii (5), Pacifastacus leniusculus (6), Austropotamobius pallipes<br />
(7), Austropotamobius italicus (8), Austropotamobius torrentium (9), Astacus astacus (10),<br />
Astacus leptodactylus (11)<br />
Parastacidae: Cherax quadricarinatus (12)<br />
Cambaridae: Orconectes nais (13), Orconectes limosus (14), Faxonella clypeata (15),<br />
Orconectes rusticus (16), Orconectes propinquus (17), Orconectes virilis (18), Orconectes<br />
inermis inermis (19), Orconectes pellucidus (20), Cambarus blandingi (21), Cambaroides<br />
japonicus (22), Cambarus immunis (23), Procambarus alleni (24), Procambarus clarkii (25),<br />
Procambarus hayi (26)<br />
Palinuridea<br />
Palinuridae: Panulirus homarus (27)*, Panulirus argus (28)*, Panulirus longipes cygnus (29)<br />
Anomura<br />
Diogenidae: Calcinus verilli (30), Calcinus laevimanus (31), Calcinus seurati (32), Calcinus<br />
elegans (33), Calcinus hazletti (34), Calcinus laurentae (35)<br />
Coenobitidae: Birgus latro (36), Coenobita perlatus (37), Coenobita clypeatus (38), Coenobita<br />
compressus (39)<br />
Brachyura<br />
Leucosiidae: Philyra scabriuscula (40), Ebalia tuberosa (41)<br />
Xanthidae: Lophopanopeus bellus (42), Lophopanopeus diegensis (43), Paraxanthias taylori<br />
(44), Pilumnus hirtellus (45), Xantho incisus (46), Nanopanope sayi (47), Eurypanopeus<br />
depressus (48), Panopeus herbstii (49)<br />
Majidae: Microphrs bicornutus (50), Pw
Table 2. continued.<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 143<br />
* Soft-female mating was rarely reported in addition to the hard-female mating. References: (1)<br />
Yano et al. (1988), Misamore & Browdy (1996), Palacios et al. (2003), (2) Misamore & Browdy<br />
(1996), (3) Aquacop (1977), (4) Bueno (1990), (5) Mason (1970a, b), (6) Lowery & Holdich (1988),<br />
Stebbing et al. (2003), (7) Ingle & Thomas (1974), Brewis & Bowler (1985), Carral et al. (1994),<br />
Villanelli & Gherardi (1998), (8) Galeotti et al. (2007), Rubolini et al. (2006, 2007), (9) Laurent<br />
(1988), (10) Cukerzis (1988), (11) Koksal (1988), (12) Barki & Karplus (1999), (13) Pippit (1977),<br />
(14) Schone (1968), Holdich & Black (2007), (15) Smith (1953), (16) Berrill & Arsenault (1982),<br />
Snedden (1990), Simon & Moore (2007), (17) Tierney & Dunham (1982), (18) Bovbjerg (1953),<br />
Rubenstein & Hazlett (1974), Tierney & Dunham (1982), (19)-(20) Bechler (1981), (21) Pearse<br />
(1909), (22) Kawai & Saito (2001), (23) Tack (1941), (24) Bovbjerg (1956), Mason (1970a, b),<br />
(25) Ameyaw-Akumfi (1981), Corotto et al. (1999), (26) Payne (1972), (27) Berry (1970), Heydon<br />
(1969), (28) Sutcliffe (1952, 1953), Kaestner (1970), Lipcius et al. (1983), Lipcius & Herrnkind<br />
(1987), (29) Chittleborough (1976), Sheard (1949), (30)-(35) Hazlett (1972), (36) Helfman<br />
(1977), (37) Page & Willason (1982), (38) Dunham & Gilchrist (1988), (39) Contreras-Garduno<br />
et al. (2007), (40) Naidu (1954), (41) Schenibri (1983), (42)-(43) Knudsen (1960, 1964), (44)-<br />
(46) Bourdon (1962), (47)-(49) Swartz (1976a, b), (50) Hartnoll (1965a), (51) Vernet-Cornubert<br />
(1958a), (52) Knudsen (1964), (57) Boolootian et al. (1959), Grigg personal communication in<br />
Hartnoll (1969), Knudsen (1964), (54) Berry & Hartnoll (1970), (55) Arakawa (1964), (56) Warner<br />
(1967, 1970), (57) Broekhuysen (1941), (58) Hartnoll (1965b), (59)-(60) Brockerhoff & McLay<br />
(2005a, b)5 (61) Hoestlandt (1948), Peters et al. (1933), (62) Kobayashi & Matsuura (1994), (63)<br />
Schone & Schone (1963), Warner (1967, 1970), (64) Kramer (1967), Schone & Schone (1963),<br />
(65) Brockerhoff & McLay (2005a, b, c), (66) Knudsen (1964), (67) Yaldwyn (1966b), Brockerhoff<br />
(2,002), (68) Knudsen (1964), (69) Brockerhoff & McLay (2005a, b, c), (70) Bovbjerg (1960), Hiatt<br />
(1948), (71) Brockerhoff & McLay (2005a, b), (72) Vernet-Cornubert (1958b), (73) Fukui (1991,<br />
1994), "(74) Hartnoll (1969), (75)-(77) Brockerhoff & McLay (2005a, b), (78) Warner (1967 as<br />
Sesarma ricordt)r(19) Seiple & Salmon (1982), (80)-(81) Hartnoll (1969), (82) von Hagen (1967),<br />
(83) Hartnoll (1969), (84) Seiple & Salmon (1982 as Sesarma cinereum), (85) Hartnoll (1969 as<br />
Sesarma angustipes), (86) Hartnoll (1969 as Sesarma curacaoense), (87) Nye (1977), Beer (1959),<br />
Brockerhoff & McLay (2005a, b), (88) Hicks (1985), (89) Abele et al. (1973), Klassen (1975), Bliss<br />
et al. (1978), (90) Gifford (1962), Henning (1975), (91) Ameyaw-Akumfi (1987).<br />
Table 3. Penaeid shrimp species in which a sperm plug has been reported.<br />
Penaeidae<br />
Rimapenaeus similis (1)<br />
Farfantepenaeus aztecus (2)<br />
Rimapenaeus constrictus (3)<br />
Marsupenaeus japonicus (4)<br />
Metapenaeus joyneri (5)<br />
References: (1) Bauer & Min (1993 as Trachypenaeus similis), (2) Bauer & Min (1993), (3) Costa<br />
& Fransoso (2004), (4) Fuseya (2006), (5) Miyake (1982).
144 Asakura<br />
Table 4. Species of the precopulatory guarding type, in which males guard females before copulation. S =<br />
species in which females molt before copulation. H = species in which females do not molt before copulation.<br />
V = species in which both types (S and H) have been reported. ? = molting condition has not been reported.<br />
CARIDEA<br />
Palaemonidae: Macrobrachium amazonicum [S](l), Macrobrachium rosenbergii [S](2), Macrobrachium<br />
austoraliense [S](3), Macrobrachium nipponense [S](4), Macrobrachium longipes<br />
[S](5)<br />
Rhynchocinetidae: Rhynchocinetes typus [H](6)<br />
ASTACIDEA<br />
Homaridae: Homarus americanus [V](7)<br />
ANOMURA<br />
Diogenidae: Diogenes pugilator [S](8), Diogenes nitidimanus [V](9), Dardanus punctulatus<br />
[?](10), Calcinustibicen [Sl](ll)<br />
Paguridae: Pagurus miamensis [V](12), Pagurus pygmaeus [V](13), Pagurus bonairensis (H](14),<br />
Pagurus marshi [S](15), Pagurus bernhardus [S](16), Pagurus cuanensis [H](17), Pagurus<br />
anachoretus [H](18), Pagurus alatus [H](19), Pagurus marshi [S](20), Pagurus nigrofascia<br />
[S](21), Pagurus lanuginosus [V](22), Pagurus prideauxi [H](23), Pagurus hirsutiuculus<br />
[S](24), Pagurus maculosus [?](25), Pagurus minutus [V](26), Pagurus filholi [V](27), Pagurus<br />
gracilipes [?](28), Pagurus middendorffii [H](29), Pagurus nigrivittatus [V](30), Anapagurus<br />
chiroacanthus [V](31), Anapagurus breriaculeatus [V](32), Pylopagurus sp. sensu Hazlett<br />
(1975)[H](33)<br />
Lithodidae: Paralithodes camtschaticus [S](34), Paralithodes brevipes [S](35), Lithodes maja<br />
[S](36), Lithodes santolla [S](37), Paralomis granulose [S](38), Hapalogaster dentata [S](39)<br />
BRACHYURA<br />
Leucosiidae: Philyrd laevis [H](40)<br />
Majidae: Chionoecetes opilio [S](41), Chionoeceies bairdi [S](42), Macropodia longirostris<br />
[S](43), Macropodia rostrata [S](44)<br />
Hymenosomatidae: Halicarcinus sp.'[S](45), Hymenosoma orbiculare [S](46)<br />
Cancridae: Cancer gracilis [S](47), Cancer irroratus [S](48), Cancer magister [S](49), Cancer<br />
oregonensis [S](50), Cancer pagurus [S](51), Cancer pro ductus [S](52), Cancer borealis<br />
[S](53), Cancer antennarius [S](54)<br />
Cheiragonidae: Telmessus cheiragonus [S](55), Erimacrus isenbeckii [S](56)<br />
Corystidae: Corystes cassivelaunus [H](57)<br />
Portunidae: Callinectes sapidus [S](58), Carcinus maenas [S](59), Macropipes holsatus [S](60),<br />
Ovalipes ocellsatus [S](61), Portunus pelagicus [S](62), Portunus sanguinolentus [S](63),<br />
Portunus puber [S](64), Portunus trituberculatus [S](65), Scylla serrata [S](66)<br />
Xanthidae: Menippe mercenaria [S](67)
Table 4. continued.<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 145<br />
References: (1) Guest (1979), (2) Bhimachar (1965), Rao (1967), Ra'anan & Sagi (1985), Kuris<br />
et al. (1987), (3) Ruello et al. (1973), Lee & Felder (1983), (4) Ogawa et al. (1981), Mashiko (1981),<br />
(5) Shokita (1966), (6) Correa et al. (2000, 2003), Hinojosa &' Thiel (2003), Correa & Thiel (2003a,<br />
b), Diaz & Thiel (2003), Thiel & Hinojosa (2003), Diaz & Thiel (2004), Thiel & Correa (2004), van<br />
Son & Thiel (2006), Dennenmoser & Thiel (2007), (7) Herrick (1909), Templeman (1934, 1936),<br />
McLeese (1970, 1973), Hughes & Matthiessen (1962), Aiken & Waddy (1980), Waddy & Aiken<br />
(1981), Aiken et al. (2004), (8) Bloch (1935), Hazlett (1968), (9) Asakura (1987), (10) Matthews<br />
(1956), (11)-(13) Hazlett (1966), (14)-(17) Hazlett (1968), (18) Hazlett (1968), Hazlett (1975),<br />
(19) Hazlett (1968), (20) Hazlett (1975), (21)-(22) Wada et al. (2007), (23) Hazlett (1968), (24)<br />
MacGinitie (1935), (25) Imazu & Asakura (2006), (26) Imazu & Asakura (2006), Wada et al. (2007),<br />
(27) Imafuku (1986), Goshima et al. (1998), Minouchi & Goshima (1998,2000), Wada et al. (2007),<br />
(28) Imazu & Asakura (2006), (29) Wada et al. (1996, 1999), (30) Wada et al. (2007), (31M32)<br />
Hazlett (1968), (33) Hazlett (1975), (34) Marukawa (1933), Powell & Nickerson (1965a, b), Gray<br />
& Powell (1966), Wallace et al. (1949), McMullen (1969), Matsuura & Takeshita (1976), Takeshita<br />
& Matsuura (1989), (35) Wada et al. (1997, 2000), Sato et al. (2005a, b), (36) Pike & Williamson<br />
(1959), (37)-(38) Lovrich & Vinuesa (1999), (39) Goshima et al. (1995), (40) Schembri (1983),<br />
(41) Watson (1972), (42) Paul (1984), Donaldson & Adams (1989), (43)-(44) Hartnoll (1969),<br />
(45) Lucas personal communication in Hartnoll (1969), (46) Broekhuysen (1955), (47) Knudsen<br />
(1964), (48) Chidchester (1911), Elner & Elner (1980), Elner & Stasko (1978), Haefner Jr. (1976),<br />
(49) Bulter (1960), Cleaver (1949), Snow & Nielsen (1966), (50) Knudsen (1964), (51) Edwards<br />
(1966), (52) Knudsen (1964), (53) Elner et al. (1985), (54) Knudsen (1960), (55) Kamio et al.<br />
(2000, 2002, 2003), (56) Sasaki & Ueda (1992), (57) Hartnoll (1968), (58) Childchester (1911),<br />
Churchill (1919), Hay (1905), Gleeson (1980), Ryan (1966), Gleeson et al. (1984), Christofferson<br />
(1970), Teytaud (1971), Jivoff & Hines (1998), (59) Broekhuysen (1936, 1937), Cheung (1966),<br />
Childchester (1911), Spalding (1942), Veillet (1945), Williamson (1903), Berrill (1982), Berrill &<br />
Arsenault (1982), Jensen (1972), (60) Broekhuysen (1936), (61) Childchester (1911), (62) Delsman<br />
& de Man (1925), Broekhuysen (1936), Fielder & Eales (1972), (63) George (1963), Ryan (1966,<br />
1967a, b), Christofferson (1970, 1978), (64) Duteutre (1930), (65) Oshima (1938), (66) Hill (1975),<br />
(67) Binford (1913), Cheung (1968), Savage (1971), Porter (1960), Wilber (1989).
146 Asakura<br />
Table 5. Duration of guarding time in selected species of decapod crustaceans.<br />
Species<br />
ANOMURA<br />
Lithodidae<br />
Paralithodes brevipes<br />
Paralithodes brevipes<br />
3 males & 3 females<br />
1 male & 5 females<br />
Hapalogaster dentata<br />
BRACHYURA<br />
Cancridae<br />
Cancer pagurus<br />
Conner irroratus<br />
Carcinus maenas<br />
1 male & 1 female<br />
2 or 3 males - 1 female<br />
Majidae<br />
Chionoecets bairdi<br />
Chionoecets opilio<br />
Cheiragonidae<br />
Telmessus cheiragonus<br />
Corystidae<br />
Corystes cassivelaunus<br />
Precopulatory<br />
guarding time<br />
9-84 hrs<br />
(mean 38.9+24.9 hrs)<br />
32.1+44.1 hrs<br />
15.1+20.1 hrs<br />
2-3 days<br />
3-21 days<br />
4.5 days<br />
2-16 days<br />
3-10 days<br />
1-12 days<br />
•7-9 days<br />
11.8 ± 5 SD days<br />
Up to several days<br />
Female condition<br />
when copulating<br />
Soft<br />
Soft<br />
Soft<br />
Soft<br />
Soft<br />
Soft<br />
Soft<br />
Soft<br />
Various<br />
Soft<br />
Soft<br />
Hard<br />
Postcopulatory<br />
guarding time<br />
?<br />
?<br />
?<br />
?<br />
1-12 days<br />
5 days<br />
0-1.5 days<br />
1-3.5 days<br />
?<br />
8 hrs<br />
4.0 ± 6.6 hrs<br />
0<br />
Refere<br />
References: (1) Wada et al. (1997), (2)-(3) Wada et al. (2000), (4) Goshima et al. (1995), (5) Edwa<br />
(1966), (6) Elner & Elner (1980), (7)-(8) Berrill & Arsenault (1982), (9) Donaldson & Adams (198<br />
(10) Watson (1972), (11) Kamio et al. (2003), (12) Hartnoll (1968).<br />
(1)<br />
(2)<br />
(3)<br />
(4)<br />
(5)<br />
(6)<br />
(7)<br />
(8)<br />
(9)<br />
(10)<br />
(11)<br />
(12)
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 147<br />
Table 6. Brachyuran crab species, in which a sperm plug has been reported.<br />
Gancridae<br />
Cancer magister (1)<br />
Cancer irroratus (2)<br />
Cancer pagurus (3)<br />
Geryonidae<br />
Geryon fenneri (4)<br />
Portunidae<br />
Callinectes sapidus (5)<br />
Carcinoplax vestita (6)<br />
Carcinus maenas (7)<br />
Macropipus holsatus (8)<br />
Ovalipes ocellsatus (9)<br />
Portunus sanguinolentus (10)<br />
Necorapuber (11)<br />
Liocarcinus depurator (12)<br />
Cheiragonidae<br />
Telmessus cheiragonus (13)<br />
Eriphiidae<br />
Eriphia smithii (14)<br />
References: (1) Oh & Hankin (2004), (2) Childchester (1911), (3) Edwards (1966), (4) Hinsch<br />
(1988), (5) Childchester (1911), Wenner (1989), Johnson & Oito (1981), Jivoff (1997), (6) Doi &<br />
Watanabe (2006), (7) Broekhuysen (1936, 1937), Spalding (1942), (8) Broekhuysen (1936), (9)<br />
Childchester (1911), (10) George (1963), (11) Gonzalez-Gurriaran & Freire (1994), Norman &<br />
Jones (1993), (12) Abello (1989), (13) Kamio et al. (2003), (14) Tomikawa &Watanabe (1990).<br />
Table 7. Species found in large aggregations called a "pod," "heap," or "mound."<br />
Species<br />
ANOMURA<br />
Lithodidae<br />
Paralithodes camtschaticus<br />
Lithodes santolla<br />
BRACHYURA<br />
Majidac<br />
Maya squinado<br />
Chionoecetesbairdi<br />
Hym lyratus<br />
LojKorhynchus grandis<br />
Libinia emarginata<br />
Number of crabs in<br />
each aggregation Reference<br />
1000 or more<br />
70 ind-m-2 or more<br />
22-50,000 or more<br />
100,000s<br />
2,000<br />
100s<br />
5,000?<br />
References: (1) Dew (1990), Dew et al. (1992), Powell & Nickelson (1965a, b), Powell et al. (1973),<br />
Zhou & Siiirley (1997), Stone et al. (1993), (2) Cardenas et al. (2007), (3) Baal (1953), Le Sueur<br />
(1954), Carlisle (1957), Sampedro & Gonzalez-Gurriaran (2004), (4) Stevens (2003), Stevens et al.<br />
(1994), (5 ) Stevens et al. (1992), (6) Debelius (1999), Hobday & Rumsey (1999), (7) DeGoursey &<br />
Auster (1992), Hinsch (1968).<br />
(1)<br />
(2)<br />
(3)<br />
(4)<br />
(5)<br />
(6)<br />
(7)
148 Asakura<br />
Table 8. Species of the Pontoniinae reported as "found in pair." Species of shrimps with [host animals in<br />
brackets] are listed according to the phyla of the host animals (large capitals).<br />
PORIFERA<br />
Apopontonia dubia [Spongia sp.](l), Onycocaris amakusensis [Callyspongia elegans](2),<br />
Onycocaris oligodentata [purplish sponge](3), Onycocaris spinosa [small sponge](4),<br />
Onycocaridella prima (5)[Mycale sulcata], Onycocaridella monodoa (= Onycocaris monodoa)<br />
[Pavaesperella hidentata](6), Onycocaridites anornodactylus [sponge] (7), Orthopontonia<br />
ornatus [Jaspis stellifera]{%), Periclimenaeus stylirostris [sponge](9), Typton dentatus [Reniera<br />
sp.](10)<br />
CNIDARIA<br />
Antipatharia<br />
Dasycaris zanzibarica [black coral, sea whips] (11)<br />
Actiniaria<br />
Periclimenes brevicarpalis [Cryptodendron adhaesivum](12), Periclimenes colemani<br />
[Asthenosoma intermedium](l3), Periclimenes ornatus [Entacmaea quadricolor, Heteroactis<br />
malu, Parasicyonis actinostroides](\4)<br />
Scleractinia<br />
Anapontonia denticauda [Galaxeafascicularis](\5), Coralliocaris superba [Acropora tubicinaria<br />
and other 15 spp. of Acropora](\6), Jocaste lucina [Acropora tubicinaria](ll), Jocaste japonica<br />
[Acropora sp., Acropora humilis, Acropora variabilis, Acropora tubicinaria, Acropora<br />
nasuta](18), Ischnopontonia lophos [Galaxea fascicularis]{\9), Periclimenes lutescens (20),<br />
Periclimenes koroensis [Fungia actiniformis](21), Philarius imperialis [Acropora sp., Acropora<br />
millepora](22), Vir euphyllius [Euphyllia spp.](23), Vir philippinensis [Plerogyra sinuosa](24)<br />
Scleractinia [in network of fissures on surface of faviid coral]<br />
Ctenopontonia cyphastreophila [Cyphastrea microphthalma](25)<br />
Scleractinia [forming galls or bilocular cyst in corals]<br />
Paratypton siebenrocki [Acropora hyacinthus and other 6 spp. of Acropord\{26)<br />
MOLLUSCA<br />
Opistobranchia<br />
Periclimenes imperator [Hexabranchus marginatus](21)<br />
Bivalvia<br />
Anchistus demani [Tridacna maxima](2S), Anchistus miersi [Tridacna squamosa, Tridacna<br />
maxima](29), Anchistus pectinis [Pecten sp., Pecten albicans], Anchistus custos [Pinna saccata,<br />
Pinna sp.](31), Chernocaris plaunae [Placunaplacenta](32), Conchodytes biunguiculatus<br />
[Pinna bicolor](33), Conchodytes meleagrinea [Meleagrina margaritifera](34), Conchodytes<br />
monodactylus [Pecten sp.,Atrina sp.](35), Conchodytes nipponensis [Pinna sp., Pecten laquetus,<br />
Atrina japonica](36), Conchodytes tridacnae [Tridacna maxima](31), Bruceonia ardeae<br />
(= Pontonia ardeae)[Chama pacifica](3%), Pontonia domestica [Atrina seminuda, Atrina rigida,<br />
Pinna muricata](39), Pontonia mexicana [Pinna cornea, Pinna rigida, Atrina seminuda]{40),<br />
Ascidonia miserabilis (= Pontonia miserabilis)[Spondylus americanus]{4\), lAscidonia<br />
miserabilis (as IPontonia miserabilis)[Spondylus americanus](42), Pontonia pinnae [Pinna<br />
rugosa, Atrina tuberculosa]{43), Pontonia pinnophylax [Pinna rudis, Pinna nobilis]{44),<br />
Pontonia margarita [Pinctada mazatlanica](45), Platypontonia hyotis [Pycnodonta hyotis]{46)
Table 8. continued.<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 149<br />
ECHINODERMATA<br />
Crinoidea: Comatulida<br />
Palaemonellapottsi [Comanthina schlegelii, Comanthus briareus, Stephanometra briareus](47),<br />
Parapontonia nudirostris [Tropiometra afra, Himerometra robustipinna] (48), Periclimenes<br />
alegrias [Lamprometra palmata, Lamprometra klunzingeri, Stephanometra spicata](49),<br />
Periclimenes attenuatus [Comaster multifidus](50), Periclimenes novaecaledoninae<br />
[Lamprometra klunzingeri](51)<br />
Echinoidea<br />
Tuleariocaris holthuisi [Astropyge radiata](52), Tuleariocaris zanzibarica [Astropyge radiata, Diadema<br />
setosum](53)<br />
CHORDATA<br />
Ascidiacea: compound ascidian<br />
Periclimenaeus diplosomatis [Diplosoma lrayneri](54), Periclimenaeus serrula [Leptoclinoides<br />
incertus](55), Periclimenaeus tridentatus [unidentified &scidmn](56),Ascidoniaflavomaculata<br />
(= Pontonia flavomaculata)[Ascidia mentula, Ascidia mammillata, Ascidia involuta, Ascidia<br />
interrupta](51), Odontoma sibogae (= Pontonia sibogae)[Styela whiteleggei, Pyura momus,<br />
Rhopalaea crassa](5&)<br />
Ascidiacea: solitary ascidian<br />
Dasella ansoni [Phallusia depressiuscula](59)<br />
References: (1) Bruce (1983a), (2)-(4) Fujino & Miyake (1969), (5)-(6) Bruce (1981a), (7) Bruce<br />
(1987), (8) Bruce (1982), (9) Bruce & Coombes (1995), (10) Bruce & Coombes (1995), Bruce<br />
(1980a), (11) Gosliner et al. (1996), (12) Bruce & Svoboda (1983), (13) Bruce (1975), (14) Bruce &<br />
Svoboda (1983), Omori et al. (1994), (15) Bruce (1967), (16)-(17) Bruce (1980b), (18) Bruce (1974,<br />
1980b, 1981c), (19) Bruce (1980b, 1981c), Bruce & Coombes (1995), (20) Bruce (1981c), Bruce<br />
& Coombes (1995), (21) Bruce & Svobboda (1984), (22) Bruce & Coombes (1995), (23) Martin<br />
(2007), (24) Bruce & Svoboda (1984), (25) Bruce (1979), (26) Bruce (1980a, b), (27) Bruce (1972a,<br />
1976a), Bruce & Svoboda (1983), Strack (1993), (28) Bruce (1972a), (29) Bruce (1972a), Debelius<br />
(1999), (30) Bruce (1972a), Fujino & Miyake (1967), (31) Bruce (1972a, 1989), Hipeau-Jacquotte<br />
(1973), (32) Bruce (1972a), (33) Bruce (1972a), Hipeau-Jacquotte (1973), (34) Bruce (1973), (35)-<br />
(36) Bruce (1972a), (37) Bruce (1974), (38) Bruce (1981b), Fransen (2002), (39) Bruce (1972a),<br />
Courtney & Couch (1981), Fransen (2002), (40) Bruce (1972a), Criales (1984), Fransen (2002),<br />
(41) Fransen (2002), (42) Criales (1984), (43) Bruce (1972a), (44) Debelius (1999), Richardson et<br />
al. (1997), (45) Baeza (2008), (46) Hipeau-Jacquotte (1971), (47) Bruce & Coombes (1995), Bruce<br />
(1989), (4.8) Bruce (1992), (49) Bruce (1986), Bruce & Coombes (1995), (50) Bruce (1992), (51)<br />
Bruce & Coombes (1995), (52)-(53) Bruce (1967), (54) Bruce (1980b), (55) Bruce & Coombes<br />
(1995), (56) Bruce & Coombes (1995), (57) Monniot (1965), Millar (1971), Fransen (2002), (58)<br />
Bruce (1972b), Fransen (2002), (59) Bruce & Coombes (1995).
150 Asakura<br />
Table 9. Species of the Alpheidae reported as "found in pair." Species of shrimps with [host animals in brackets]<br />
are listed according to the phyla of host animals (large captals) with higher taxa or habitat when known.<br />
PORIFERA<br />
Synalpheus bituberculatus [sponge] (1), Synalpheus hastilicrassus [sponge] (2), Synalpheus<br />
jedanensis [sponge](3), Synalpheus streptodactylus [sponge](4), Synalpheus theano [sponge](5),<br />
Synalpheus fossor [sponge](6), Synalpheus harpagatrus [sponge] (7), Synalpheus nilandensis<br />
[sponge](8), Synalpheus tumidomanus [sponge] (9), Zuzalpheus androsi [Hyattella<br />
intestinalis](lO), Synalpheus couitere [sponge](l 1), Zuzalpheus bousfield [Hymeniacidon<br />
spp.](12), Zuzalpheus carpenteri [Aeglas spp.](13), Zuzalpheus goodei [Xestospongia<br />
wiedenmayeri, Pachypellina podatypa](l4), Zuzalpheus paraneptunus [Hyattella intestinalis,<br />
Oceanapia sp.](15), Zuzalpheus ruetzleri [Hymeniacidon cf. caerulea](16), Zuzalpheus<br />
sanctithomae [Hymeniacidon caerulea etc.](17), Alpheus parvirostris [sponge](18), Alpheus<br />
alcyone [sponge](19), Alpheus aff. eulimene*[sponge](20), Alpheusparalcyone [sponge](21),<br />
Alpheus spongiarum [sponge] (22)<br />
CNIDARIA<br />
Scyphozoa: Coronatae<br />
Synalpheus modestus (23), Synalpheus aff. modestus sensu Nomura & Asakura (1998)<br />
[Stephanoscyphus racemosus](24)<br />
Anthozoa: Gorgonacea<br />
Synalpheus iphinoe [Solenocaulon sp.](25), Synalpheus trispinosus [gorgonacean](26)<br />
Anthozoa: Alcyonacea<br />
Synalpheus neomeris [Dendronephthya](27)<br />
Anthozoa: Actiniaria<br />
Alpheus armatus [Bartholomea annulata](2S), Alpheus immaculatus [Bartholomea annulata](29),<br />
Alpheus polystuctus [Bartholomea annulata](30), Alpheus roquensis [Heteractis lucida](31)<br />
Anthozoa: Scleractinia<br />
Alpheus lottini [reef coral, Pocillopora](32), Alpheus ventrosus (33), Synalpheus charon<br />
[Pocillopora, reef coral](34), Synalpheus scaphoceris [Madracis decactis](35), Racilius<br />
compressus [Galaxea fascicularis](36)<br />
Anthozoa: Scleractinia (in fissures on massive coral)<br />
Alpheus deuteropus [Asteropora, Porites, Acropora, Montipora, Pavona](37)<br />
Anthozoa: Scleractinia (coral borer, in dead coral head)<br />
Alpheus saxidomus (38), Alpheus simus (39), Alpheus schmitti (40), Alpheus idiocheles (41),<br />
Alpheus colluminaus (42)<br />
ANNELIDA<br />
Polychaeta<br />
Alpheus sulcatus [Eurythoe complanata](43)<br />
CRUSTACEA<br />
Shell used by hermit crab<br />
Aretopsis amabilis [Dardanus sanguinolentus, Dardanus megistos, Dardanus guttatus, Dardanus<br />
lagopodes, Clibanarius eurysternus, Calcinus latens](44), Aretopsis manazuruensis [Aniculus<br />
miyakei](45)<br />
In burrow of thalassinidean shrimps<br />
Betaeus longidactylus [Upogebia pugettensis](46), Betaeus harrimani [Upogebiapugettensis](47),<br />
Betaeus ensenadensis [Upogebia pugettensis] (48)<br />
In burrow of mantis shrimp<br />
Athanas squillophilus [Oratosquilla oratoria](49)
Table 9. continued.<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 151<br />
ECHINODERMATA<br />
Crinoidea: Comatulida<br />
Synalpheus carinatus [crinoids](50), Synalpheus comatularum [Comanthus timorensis](51),<br />
Synalpheus demani [criniod](52), Synalpheus stimpsoni [Comaster multibrachiatus, Comaster<br />
multifidus, Comaster gracilis, Comaster alternans](53), Synalpheus odontophorus [crinoid](54)<br />
Echinoidea<br />
Athanas indicus [Echinometra mathaei](55)<br />
ECHIURA<br />
Athanopsis rubricinctuta [Ochetostoma erythrogrammon](56), Betaeus longidactylus [Urechis<br />
sp.](57)<br />
"PISCES" [in burrow of goby fish]<br />
Alpheus bellulus [Tomiyamichthys spp, Amblyeleotris spp.](58), Alpheus purpurilenticularis<br />
[Amblyeleotris steinitzi], (59) Alpheus rapacida [Myersina spp., Vanderhorstia spp., Mahidoria<br />
spp.], (60) Alpheus rapax [Crypto centrus spp.](61)<br />
ALGAE TUBE<br />
Alpheus frontalis [tube of filamentous blue-green algae such as Microcoelus spp.](62), Alpheus<br />
bucephalus [tube of pure algae or algae with sponges and other material](63), Alpheus brevipes<br />
[tube of red filamentous alga](64), Alpheus clypeatus [tube of red filamentous alga<br />
Acrochaetium](65), Alpheus pachychirus [tube of algae](66)<br />
FREE LIVING [crack of rock, under rubble, around large algae, burrow in mudflat]<br />
Alpheopsis chilensis (67), Alpheus normanni (68), Alpheus euphrosyne richardsoni (69), Alpheus<br />
strenuus cremnus (70), Alpheus diadema (71), Alpheus architectus (72), Alpheus amirantei (73),<br />
Alpheus bisincisus (74), Alpheus brevicristatus (75) (might be commensal with goby?), Alpheus<br />
edwardsii (76), Alpheus aff. gracilipes* (77), Alpheus heeia (78), Alpheus'aff. heeia*(19),<br />
Alpheus aff. leviuscuius sp. 1*(80), Alpheus aff. leviusculus sp. 2* ; (81), Alpheus lobidens (82),<br />
Alpheus aff. lobidens sp. 1*(83), Alpheus aff. lobidens sp. 2*(84), Alpheus aff. lobidens sp.<br />
3*(85), Alpheus malleodigitus (86), Alpheus miersi (SI), Alpheus obesomanus (88), Alpheus<br />
pacificus (89), Alpheus aff. pacificus (90), Alpheus paradentipes (91), Alpheus parvirostris (92),<br />
Alpheus polyxo (93), Alpheus serenei (94), Alpheus suluensis (95), Alpheus tenuipes (96),<br />
Alpheus angulatus (97), Alpheus armillatus (98), Alpheus heterochaelis (99), Alpheus floridanus<br />
(100), Alpheus inca (101), Metalpheusparagracilis (102)
152 Asakura<br />
Table 9. continued.<br />
*sensu Nomura & Asakura (1998). References: (1) Banner & Banner (1975), Nomura & Asakura<br />
(1998), (2)-(5) Nomura & Asakura (1998), (6) Didderen et al. (2006), (7) Banner & Banner (1975),<br />
(8)-(9) Nomura & Asakura (1998), (10) Rios & Duffy (2007), (11) Nomura & Asakura (1998), (12)<br />
Rios & Duffy (2007), (13) Macdonald III et al. (2006), Rios & Duffy (2007), (14)-(17) Rios & Duffy<br />
(2007), (18) Banner & Banner (1982), (19)-(27) Nomura & Asakura (1998), (28) Knowlton (1980),<br />
Knowlton & Keller (1982, 1983, 1985), Criales (1984), (29)-(31) Knowlton (1980), Knowlton &<br />
Keller (1982, 1983, 1985), (32) Vannini (1985), Nomura & Asakura (1998), Abele & Patton (1976),<br />
Tsuchiya & Yonaha (1992), (33) Patton (1966), (34) Patton (1966), Nomura & Asakura (1998), (35)<br />
Dardeau (1984, 1986), (36) Bruce (1972c), (37) Banner & Banner (1983), (38) Fischer & Meyer<br />
(1985), Fischer (1980), (39)-(40) Werding (1990), (41) Kropp (1987), Nomura & Asakura (1998),<br />
(42) Banner & Banner (1982), Nomura & Asakura (1998), (43) Banner & Banner (1982), (44)<br />
Bruce (1969), Banner & Banner (1973), Kamezaki & Kamezaki (1986), (45) Suzuki (1971), (46)-<br />
(48) MacGinitie (1937), (49) Hayashi (2002), (50) Bruce (1989), (51) Banner & Banner (1975),<br />
(52) Bruce (1989), Nomura & Asakura (1998), (53) Nomura & Asakura (1998), Van den Spiegel et<br />
al. (1998), (54) Nomura & Asakura (1998), (55) Gherardi (1991), (56) Anker et al. (2005), Berggren<br />
(1991), (57) MacGinitie (1935), (58) Miya & Miyake (1969), Nomura & Asakura (1998), Nomura<br />
(2003), (59) Macnae & Kalk (1962), Karplus (1979), Nomura (2003), (61) Macnae & Kalk (1962),<br />
Nomura (2003), (62) Fishelson (1966), Banner & Banner (1982), (63) Banner & Banner (1982),<br />
Nomura & Asakura (1998), (64)-(65) Banner & Banner (1982), (66) Cowles (1913), Banner &<br />
Banner (1982), (67) Boltana & Thiel (2001), (68) Nolan & Salmon (1970), (69)-(70) Banner &<br />
Banner (1982), (71)-(75) Nomura & Asakura (1998), (76) Nomura & Asakura (1998), Jeng (1994),<br />
(77)-(96) Nomura & Asakura (1998), (97) Mathews (2002a, b, 2003, 2006, 2007), Mathews et<br />
al. (2002), (98) Mathews et al. (2002), (99) Nolan & Salmon (1970), Schein (1975), Obermeier<br />
& Schmitz (2003a, b), Rahman et al. (2001, 2002, 2003, 2005), Schmitz & Herberholz (1998),<br />
Dworschak & Ott (1993), (100) Dworschak & Ott (1993), (101) Boltana & Thiel (2001), (102)<br />
Nomura & Asakura (1998).
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 153<br />
Table 10. Species of shrimps other than Pontoniinae and Alpheidae reported as "found in pair." Species of<br />
shrimps with [host animals in brackets] are listed according to the phyla of host animals (large capitals) with<br />
higher taxa or habitat when known.<br />
SPONGICOLIDAE<br />
PORIFERA<br />
Spongicola japonica [Euplectella oweni](l), Spongicola venusta [Euplectella aspergillum](2),<br />
Spongicola levigata [Euplectella oweni1](3), Spongiocaris semiteres [hexactinellid sponge], (4)<br />
Spongicoloides iheyaensis [Euplectellidae & Hyalonematidae](5), Globospongicola spinulatus<br />
[hexactinellid sponge Semperella sp.](6)<br />
FREE LIVING<br />
Microprosthema validum (7)<br />
STENOPODIDAE<br />
FREE LIVING<br />
Stenopus hispidus ($), Stenopus scutellatus (9), Stenopus tenuirostris (10), Stenopus<br />
zanzibaricus (11)<br />
HIPPOLIYTIDAE<br />
FREE LIVING<br />
Lysmata debelius (12), Lysmata grabhami (13)<br />
CNIDARIA<br />
Actiniaria, Scleractinia<br />
Thor amboinensis'(14)<br />
GNATHOPHYLLIDAE<br />
ECHINODERMATA<br />
Holothuroidea<br />
Pycnocaris chagoae [Holothuria cinerascens](15)<br />
Asteroidea<br />
Hymenocera picta [prey on sea star](16)<br />
References: (1) Saito et al. (2001), (2) Miyake (1982), Hayashi & Ogawa (1987), (3) Hayashi &<br />
Ogawa (1987), (4) Bruce & Baba (1973), (5) Saito et al. (2006), (6) Komai & Saito (2006), (7)<br />
Davie (2002), (8) Johnson (1969,1977), Castro & Jory (1983), Zhang et al. (1998), Yaldwyn (1964,<br />
1966a), (9) Debelius (1999), (10) Bruce (1976b), (11) Gosliner et al. (1996), (12) Rufino & Jones<br />
(2001), Gosliner et al. (1996), (13) Wirtz (1997), Debelius (1999), (14) Stanton (1977), (15) Bruce<br />
(1983b), (16) Seibt & Wickler (1972, 1979, 1981), Wickler & Seibt (1970, 1972, 1981), Seibt<br />
(1973a, b, 1974,1980), Wasserthal & Seibt (1976), Wickler (1973), Kraul & Nelson (1986), Fiedler<br />
(2002).
154 Asakura<br />
Table 11. Species of Thalassinidea and Anomura reported as "found in pair." Species with [host animals or<br />
habitat in brackets] are listed according to the phyla of host animals (in capitals) with higher taxa or habitat<br />
where known.<br />
THALASSINIDEA<br />
Axiidae<br />
FREE LIVING<br />
Axiopsis serratifrons [in burrow in sediments with a higher content of coral rubble](l)<br />
Laomediidae<br />
FREE LIVING<br />
Axianassa australis [in burrow in mud flat](2)<br />
Callianassidae<br />
"PISCES"<br />
Neotrypaea affinis [burrow of blind goby Typhlogobius californiensis](3)<br />
FREE LIVING<br />
Neotrypaea gigas [burrow in mud] (4)<br />
Upogebiidae<br />
PORIFERA<br />
Upogebia synagelas [Agelas sceptrum](5)<br />
CNIDARIA: Scleractinia<br />
Pomatogebia rugosa [inside live colony of Pontes lobata](6), Pomatogebia operculata [inside live<br />
coral colony](7), Upogebia corallifora [inside dead coral colony](8)<br />
FREE LIVING<br />
Upogebia pugettensis [U- or Y-shaped burrow in mudflat](9), Upogebia affinis [burrow in mud](10)<br />
ANOMURA<br />
Porcellanidae<br />
CNIDARIA<br />
Gorgonacea<br />
Aliaporcellana telestophila [Solenocaulon](11)<br />
Pennatulacea<br />
Porcellanella haigae [Cavernularia sp.](12)<br />
Actiniaria<br />
Neopetrolisthes oshimai [Soichactis spp.](13), Neopetrolisthes maculatus [Stychodactyla](l4),<br />
Neopetrolisthes alobatus, Neopetrolisthes spinatus [Heteroactis malu](\5)<br />
ANNELIDA<br />
, Polychaeta [in tube of large polychaete species]<br />
Poly onyx macroheles [Chaetopterus variopedatus](\6), Poly onyx quadriungulatus [Chaetopterus<br />
variopedatus](ll), Polyonyx transversus [Chaetopterus sp.](18), Polyonyx vermicola<br />
[Sasekumaria selangora](19), Polyonyx bella [Chaetopterus variopedatus](20), Polyonyx gibbesi<br />
[Chaetopterus variopedatus](2l), Polyonyx utinomii [Chaetopterus sp.](22), Heteropolyonyx<br />
biforma [Chaetopterus sp.](23), Polyonyx biunguiculatus [Chaetopterus sp.](24)<br />
CRUSTACEA [in shell being used by hermit crab]<br />
Porcellana cancrisocialis [Petrochirus californiensis, Dardanus sinistripes, Aniculus elegans,<br />
Paguristes digueti](25), Porcellana paguriconviva [Petrochirus calif orniensis, Dardanus<br />
sinistripes, Aniculus elegans, Paguristes digueti](26)<br />
ECHINODERMATA<br />
Echinoidea<br />
Clastotoechus vanderhorsti [Echinometra lucunter](27), Clastotoechus vanderhorsti [Echinometra<br />
lucunter](2%)<br />
Asteroida<br />
Minyocerus angustus [Luidia, Astropecten, Tethyaster](29)
Table 11. continued.<br />
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 155<br />
FREE LIVING<br />
Pachycheles rudis [underside of stone, basal portion of large algae](30)<br />
Galatheidae<br />
ECHINODERMATA<br />
Crinoidea<br />
Galathea inflata [Comanthus parvicirrus, Comaster schlehelii](31)<br />
References: (1) Dworschak & Ott (1993), (2) Coelho & Rodrigues (1999), Coelho (2001), (3)-(4)<br />
Meinkoth (1981), (5) Williams (1987), (6) Fonseca & Cortes (1998), (7) Kleeman (1984), Williams<br />
& Ngoc-Ho (1990), Coelho & Rodrigues (1999), Coelho (2001), (8) Williams & Scott (1989), (9)<br />
Jensen (1995), (10) Meinkoth (1981), (11) Ng & Goh (1996), (12) Nakasone & Miyake (1972),<br />
(13) Seibt & Wickler (1971), (14) Debelius (1984), (15) Osawa & Fujita (2001), (16) Gray (1961),<br />
(17) Kudenov & Haig (1974), (18) McNeill & Ward (1930), (19) Ng & Sasekumar (1993), (20)<br />
Hsueh & Huang (1998), (21) Rickner (1975), Williams (1984), Grove & Woodin (1996), (22)-(23)<br />
Osawa (2001), (24) Macnae & Kalk (1962), (25) Glassell (1936), Parente & Hendrickx (2000),<br />
Williams & McDermott (2004), (26) Parente & Hendrickx (2000), Williams & McDermott (2004),<br />
(27) Werding (1983), (28) Werding (1983), Schoppe (1991), (29) Werding (1983), Gore & Shoup<br />
(1968), (30) Meinkoth (1981), (31) Fujita & Baba (1999).<br />
Table 12. Species of brachyuran crabs reported as "found in pair." Species of crabs with [host animals in<br />
brackets] are listed within family or superfamily according to the phyla of host animals (in capitals) with<br />
higher taxa or habitat where known.<br />
XANTHIDAE<br />
CNIDARIA: Scleractinia<br />
Cymo andreossyi \Pocillopora\{\)<br />
TRAPEZHDAE<br />
CNIDARIA<br />
Scleraetinia: Pocillopora<br />
Trapezia areolata (2), Trapezia corallina (3), Trapezia cymodoce (4), Trapezia dentata (5),<br />
Trapezia digitalis (6), Trapezia ferruginea (7), Trapezia flavomaculata (8), Trapezia guttata (10),<br />
Trapezia intermedia (11), Trapezia rufopunctata (12), Trapezia tigrina (13), Trapezia wardi (14)<br />
Antipatharia<br />
Quadrella maculosa [Antipathes] (15), Quadrella spp. [Cirrhipathes abies, Antipathes spp.}(16),<br />
Quadrella reticulata [Antipathes sp.](17)<br />
TETRALIIDAE<br />
CNIDARIA<br />
Scleractinia: Acropora<br />
Tetraliafulva (18), Tetralia nigrolineata (19), Tetralia rubridactyla (20)<br />
CARPILHDAE<br />
FREE LIVING<br />
Carpilius corallinus (21)
156 Asakura<br />
Table 12. continued.<br />
PINNOTHERIDAE<br />
ANNELIDA<br />
Polychaeta [in tube of large polychaetes]<br />
Pinnixa tubicola [terebellids and chaetopterids, Eupolymnia heterobranchia, Amphitrite sp., Eupolymnia<br />
heterobranchia, Neoamphitrite rohusta, Thelepus crispus, Chaetopterus variopedatus](22),<br />
Pinnixa chaetopterana [Chaetopterida spp. Chaetopterus variopedatus, Amphitrite ornata](23),<br />
Pinnixa transversalis [Chaetopterus variopedatus](24)<br />
MOLLUSCA<br />
Bivalvia<br />
Pinnixa faba [Tresus capax, Tresus nuttalli](25), Pinnixa littoralis [Tresus capax](26)<br />
Gastropoda [inside mantle cavity]<br />
Orthotheres turboe [Turbo sp.](27), Orthotheres haliotidis [Haliotis asinina, Haliotis<br />
squamatd\{2%)<br />
SIPUNCULA & ECHIURA<br />
Mortensenella forceps [Ochetostoma erythrogrammon](29)<br />
ECHINODERMATA<br />
Echinoidea<br />
Dissodactylus mellitae [Mellita quinguiesperforata, Echinarachnius parrna, Encope<br />
michelini](30), Dissodactylus crinitichelis [Mellita sexiesperforata](3l)<br />
Holothuroidea<br />
Holotheres halingi (= Pinnotheres halingi) [Holothuria scarba](32), Holotheres semperi (= Pinnotheres<br />
semperi)[Holothuriafursocinerea, Holothuria scabra\{33)<br />
BURROWS OF OTHER ANIMALS<br />
Scleroplax granulata [burrow of echiuroid Urechis caupo, mud shrimps Neotrypaea californiensis,<br />
Neotrypaea gigas, Upogebia pugettensis, Upogebia macginiteorum](34)<br />
GRAPSOIDEA<br />
"REPTILIA": Testudines<br />
Planes minutus [loggerhead sea turtle Caretta caretta, inanimate flotsam](35)<br />
ECHINODERMATA<br />
Echinoidea<br />
Percnon gibbesi [Diadema antillarum](36)<br />
References: (1) Castro (1976), Guinot (1978), Miyake (1983), (2) Miyake (1983), Tsuchiya & Yonaha<br />
(1992), Tsuchiya & Taira (1999), (3) Patton (1966), Miyake (1983), Huber (1985), Gotelli et al.<br />
(1985), Castro (1996), (4) Patton (1966), Tsuchiya & Yonaha (1992), Tsuchiya & Taira (1999), (5)<br />
Patton (1966), Huber (1985), (6) Patton (1966), Preston (1973), Huber (1985,1987), Huber & Coles<br />
(1986), Tsuchiya & Taira (1999), (7) Patton (1966), Preston (1973), Abele & Patton (1976), Finney<br />
& Abele (1981), Miyake (1983), Adams et al. (1985), Huber & Coles (1986), Castro (1978, 1996),<br />
Tsuchiya & Taira (1999), (8) Patton (1966), Preston (1973), Miyake (1983), (9) Gotelli et al. (1985),<br />
Castro (1996), (10) Miyake (1983), Tsuchiya & Yonaha (1992), Tsuchiya & Taira (1999), (11) Preston<br />
(1973), Huber & Coles (1986), Huber (1987), (12)-(13) Huber (1985), (14) Preston (1973),<br />
Miyake (1983), Huber & Coles (1986), (15) Shih & Mok (1996), (16) Tazioli et al. (2007), (17)<br />
Castro (1999), (18) Vytopil & Willis (2001), (19)-(20) Sin (1999), (21) Laughlin (1982), (22) Hart<br />
(1982), Wells (1928), Garth & Abbott (1980), Zmarzly (1992), (23) Gray (1961), Grove & Woodin<br />
(1996), Grove et al. (2000), McDermott (2005), (24) Baeza (1999), (25) Pearce (1965, 1966a), Hart<br />
(1982), Zmarzly (1992), (26) Pearce (1966a), Zmarzly (1992), (27) Sakai (1969), (28) Geiger &<br />
Martin (1999), (29) Anker et al. (2005), (30) Bell & Stancyk (1983), Bell (1984), George & Boone<br />
(2003), (31) Telford (1978), (32) Hamel et al. (1999), (33) Ng & Manning (2003), (34) Anker et al.<br />
(2005), Campos (2006), (35) Dellinger et al. (1997), Frick et al. (2000, 2004, 2006), Carranza et al.<br />
(2003), (36) Hayes et al. (1998).
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 157<br />
Table 13. Eusocial species. All species found inhabiting cavity of sponge.<br />
Alpheidae<br />
Zuzalpheus rathbunae [sponge] (1)<br />
Zuzalpheus elizabethae.(= Synalpheus "rathbunae A")[Lissodendoryx] (2)<br />
Zuzalpheus "paraneptunus small" [sponge] (3)<br />
Zuzalpheus regalis [Xestospongia etc.] (4)<br />
Zuzalpheus filidigitus [Xestospongia etc.] (5)<br />
Zuzalpheus chacei [Aeglas, Hyattella etc.] (6)<br />
Zuzalpheus elizabethae [Lissodendoryx etc.] (7)<br />
Synalpheus neptunus neptunus [sponge] (8)<br />
References: (1) Duffy (2003), (2) Duffy (1996c, 2003), Morrison et al. (2004), (3) Duffy et al.<br />
(2000), Duffy (2003), (4) Duffy (1996a, b), Duffy et al. (2002), Rios & Duffy (2007), (5) Duffy<br />
(1996c), Duffy & Macdonald (1999), Rios & Duffy (2007), (6) Chace (1972), Duffy (1998), Rios<br />
& Duffy (2007),(7) Duffy (1996c), Morrison et al. (2004), Rios & Duffy (2007),(8) Didderen et al.<br />
(2006).<br />
Table 14. Species found in small groups. Species with [host animals] are listed, according to the phyla of host<br />
animals (large capitals) with higher taxa or habitat. One group consists of fewer than 20 individuals on a single<br />
host (species, host, number of individuals found, and reference).<br />
CARIDEA<br />
CNIDARIA<br />
Scyphozoa<br />
Periclimenes holthuisi [Cassiopei]<br />
Actiniaria<br />
Periclimenes holthuisi [sea anemone]<br />
Periclimenes tenuipes [Megalactis, Cryptodendron]<br />
Periclimenes longicarpus [Entacmaea]<br />
Periclimenes anthophilus [Condylactis gigantea]<br />
Scleractinia<br />
Thor marguitae[Porites andrewsi]<br />
Jocaste japonica [Acropora divaricata]<br />
Periclimenes holthuisi [corals]<br />
Periclimenes pederosoni [Antipathe]<br />
Anapontonia denticauda [Galaxea]<br />
ECHINODERMATA<br />
Echinoidea<br />
Gnathophylloides mineri [Tripneustes ventricosus]<br />
GALATHEOIDEA<br />
CNIDARIA<br />
Scleractinia<br />
Lissoporcellana spinuligera [Solenocaulon]<br />
CRUSTACEA: shell used by hermit crab<br />
Porcellana sdyana [Dardanus, Petrochirus, Paguristes]<br />
Max. 8 (various sizes and sexes)(l)<br />
Several individuals (2)<br />
Max. 6 (various sizes and sexes)(3)<br />
Max. 7 (various sizes and sexes)(4)<br />
Up to 9 (5)<br />
10 (2 cf, 5 ov. $, 2 non-ov. 9, 1 juv.)(6)<br />
15 (5 d\ 6 ov. $, 3 non-ov. 9, 1 juv.)(7)<br />
Several individuals (8)<br />
7 (2 cf, 3 ov. 9, 2 non-ov. $)(9)<br />
5(la\l9,3juv.)(10)<br />
Up to 13, with females greatly<br />
outnumbering males (11)<br />
7(ld\3ov. 9, 3juv.)(12)<br />
Max. 11 (several cf, several ov. 9)(13)
158 Asakura<br />
Table 14. continued.<br />
BRACHYURA<br />
MOLLUSCA<br />
Bivalvia<br />
Pinnixafaba [Tresus] More than 3(1 d\ 1 9, fewjuv.)(14)<br />
References: (1) Bruce & Svoboda (1983), (2) Coleman (1991), (3)-(4) Bruce & Svoboda (1983),<br />
(5) Nizinski (1989), (6) Bruce (1978), (7) Bruce (1981b), (8) Coleman (1991), (9) Spotte (1996),<br />
(10) Bruce (1967), (11) Patton et al. (1985), (12) Ng & Goh (1996), (13) Gore (1970), (14) Haig &<br />
Abbott (1980).<br />
Table 15. Species found in large groups. Species with [host animals] are listed, according to the phyla of host<br />
animals (large capitals) with higher taxa or habitat. One group consists of more than 20 individuals on a single<br />
host.<br />
CARIDEA<br />
PORIFERA<br />
Synalpheus dorae [Reiniere]<br />
Synalpheus streptodactylus [sponge]<br />
Synalpheus crosnieri [sponge]<br />
Synalpheus paradoxus [sponge]<br />
Zuzalpheus brooksi [sponge]<br />
Zuzalpheus idios [Hymeniacidon etc.]<br />
Zuzalpheus pectiniger [Spheciospongia]<br />
CNIDARIA<br />
. Scyphozoa<br />
Latreutes anoplonyx [Nemopilema nomurai]<br />
Scleractinia<br />
Coralliocaris macrophthalma [Acropora hyacinthus]<br />
Fennera chacei [Pocillopora]<br />
Periclimenes toloensis [Lytocarpus philippinensis]<br />
ECHINODERMATA<br />
Periclimenes affinis [Heterometra magnipinna]<br />
Periclimenes meyeri [Nemaster grandis]<br />
136 (all cf)(l)<br />
105 (68 cT, 37 ov. 9, several non- ov. $)(2)<br />
147 (144 d\ 3 9)(3)<br />
112 (110 d\ 2 $), 132 (130 cT, 2 ?)(4)<br />
10s to 1000s (5)<br />
Several 10s (including many ov. 9 & juv.)(6)<br />
Few 100s (7)<br />
More than 100 (8)<br />
24 (including 16 ?)(9)<br />
Max. 49 (all adults) (10)<br />
110 (including 43 ov.9)( 11)<br />
64 (including 16 ov. 9)(12)<br />
Max. 25 (various sizes and sexes)(13)<br />
References: (1) Bruce (1988), (2) Banner & Banner (1975, 1982), (3) Banner & Banner (1983), (4)<br />
Banner & Banner (1982), (5)-(7) Rios & Duffy (2007), (8) Hayashi et al. (2003), (9) Bruce (1977),<br />
(10) Gotelli et al. (1985), (11)-(12) Bruce & Coombes (1995), (13) Criales (1984).
The Evolution of Mating Systems in <strong>Decapod</strong> <strong>Crustacean</strong>s 159<br />
Table 16. Selected species of pinnotherid crabs (and their hosts) in which life history has been studied.<br />
MOLLUSCA<br />
Bivalvia<br />
Fabia subquadrata [Modiohis modiolus] (1)<br />
Tumidotheres maculatus (= Pinnotheres maculatus) [Mytilus edulis, Argopecten irradians etc.] (2)<br />
Pinnotheres ostreum [Crass ostrea virginica, Mytilus edulis] (3)<br />
Pinnotheres pisum [Mytilus edulis etc.] (4)<br />
Pinnotheres taichungae [Laternula marilina] (5)<br />
Pinnotheres bidentatus [Laternula marilina] (6)<br />
ANNELIDA: Polychaeta<br />
Tritodynamia horvathi [in tube of Loimia verrucosa] (7)<br />
References: (1) Pearce (1962, 1966b), (2) Pearce (1964), Williams (1984), (3) Christensen & Mc-<br />
Dermott (1958), (4) Atkins (1926), Christensen (1958), Hartnoll (1972), Williams (1984), (5)<br />
Hsueh (2003), (6) Hsueh (2001a, b), (7) Matsuo (1998, 1999), Takahashi et al. (1999).<br />
Table 17. Species in which neotenous males have been reported.<br />
Hippidae<br />
Emerita brasiliensis<br />
Emerita asiatica<br />
Emerita emeritus<br />
Emerita holthuisi<br />
Emerita talpoida<br />
Emerita rathbunae<br />
(i)<br />
(2)<br />
(3)<br />
(4)<br />
• (5)<br />
(6)<br />
References: (1) Delgado & Defeo (2006, 2008), (2) Subramoniam (1981), (3)-(4) Subramoniam &<br />
Gunamalai (2003), (5)-(6) Efford (1967).
160 Asakura<br />
APPENDIX 2:<br />
REFERENCES FOR TABLES OF APPENDIX 1<br />
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decapod crustaceans. /. Biogeogr. 3: 35-47.<br />
Abele, L.G., Robinson, M.H. & Robinson, B. 1973. Observations on sound production by two<br />
species of crabs from Panama (<strong>Decapod</strong>a, Gecarcinidae and Pseudothelphusidae). <strong>Crustacean</strong>a<br />
25: 147-152.<br />
Abello, P. 1989. Reproduction and moulting in Liocarcinus depurator (Linnaeus, 1758) (Brachyura:<br />
Portunidae) in the northwestern Mediterranean Sea. Sci. Mar. 53: 127-134.<br />
Adams, J., Edwards, A.J. & Emberton, H. 1985. Sexual size dimorphism and assortative mating<br />
in the obligate coral commensal Trapezia ferruginea Latreille (<strong>Decapod</strong>a, Xanthidae). <strong>Crustacean</strong>a<br />
48: 188-194.<br />
Aiken, D.E. & Waddy, S.L. 1980. Reproductive biplogy. In: Cobb, J.C. & Phillips, B.F. (eds.), The<br />
Biology and Management of Lobsters. Volume 1: 215-276. New York: Academic Press.<br />
Aiken, D.E., Waddy, S.L. & Mercer, S.M. 2004. Confirmation of external fertilization in the<br />
American lobster, Homarus americanus. J. Crust. Biol. 24: 474-480.<br />
Ameyaw-Akumfl, C. 1981. Courtship in the crayfish Procambarus clarkii (Girad) (<strong>Decapod</strong>a,<br />
Astacidea). <strong>Crustacean</strong>a 40: 57-64.<br />
Ameyaw-Akumfi, C. 1987. Mating in the lagoon crab Cardisoma armatum Herklots. J. Crust. Biol.<br />
7:433-436.<br />
Anker, A., Murina, G.V., Lira, C, Caripe, J.A.V., Palmer, A.R. & Jeng, M.S. 2005. Macrofauna associated<br />
with echiuran burrows: a review with new observations on the innkeeper worm Ochetostoma<br />
erythrogramm on Leuckartana Riippelin, Venezuela. Zool. Stud. 44: 157-190.<br />
Antheunisse, L.J., van den Hoven, N.P. & Jeffries, D.J. 1968. The breeding characters of Palaemonetes<br />
varians (Leach) (<strong>Decapod</strong>a, Palaemonidae). <strong>Crustacean</strong>a 14: 259-270.<br />
Aquacop. 1977. Observations sur la maturation et la reproduction en captivite des crevettes peneides<br />
en milieu tropical. Third Meet. ICES Work. G. Maricult., Brest, France, Actes Colloq. CNEXO<br />
4: 157-178.<br />
Arakawa, K.Y. 1964. On mating behavior of giant Japanese crab, Macrocheira kaempferi De Haan.<br />
Res. Crust. 1:40-46.<br />
Asakura, A. 1987. Population ecology of the sand-dwelling hermit crab, Diogenes nitidimanus<br />
Terao. 3. Mating system. Bull. Mar. Sci. 41: 226-233.<br />
Atkins, D. 1926. The moulting stages of the pea crab {Pinnotheres pisum). J. Mar. Biol. Ass. U.K.<br />
14: 475-493.<br />
Baal, H.J. 1953. Behaviour of spider crabs in the presence of octopuses. Nature 111: 887.<br />
Baeza, J.A. 1999. Indicadores de monogamia en el cangrejo comensal Pinnixa transversalis (Milne<br />
Edwards and Lucas) (<strong>Decapod</strong>a: Brachyura: Pinnotheridae): distribucion poblacional, asociacion.<br />
macho-hembra y dimorfismo sexual. Anal. Mus. Hist. Nat. Valparaeo (Chile) 34: 303-313.<br />
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