Decapod Crustacean Phylogenetics - AToL Decapoda - Natural ...

CRUSTACEAN ISSUES ] 3 

%. m 

II 

Decapod Crustacean Phylogenetics 

edited by 

Joel W. Martin, Keith A. Crandall, and Darryl L. Felder 

£\ CRC Press 

J Taylor & Francis Group

Decapod Crustacean Phylogenetics 

Edited by 

Joel W. Martin 

Natural History Museum of L. A. County 

Los Angeles, California, U.S.A. 

KeithA.Crandall 

Brigham Young University 

Provo,Utah,U.S.A. 

Darryl L. Felder 

University of Louisiana 

Lafayette, Louisiana, U. S. A. 

CRC Press is an imprint of the 

Taylor & Francis Croup, an informa business

CRC Press 

Taylor & Francis Group 

6000 Broken Sound Parkway NW, Suite 300 

Boca Raton, Fl. 33487 2742 

Contents 

Preface 

JOEL W. MARTIN, KEITH A. CRANDALL & DARRYL L. FELDER 

I Overviews of Decapod Phylogeny 

On the Origin of Decapoda 

FREDERICK R. SCHRAM 

Decapod Phylogenetics and Molecular Evolution 15 

ALICIA TOON. MAEGAN FINLEY. JEFFREY STAPLES & KEITH A. CRANDALL 

Development, Genes, and Decapod Evolution 31 

GERHARD SCHOLTZ. ARKHAT ABZHANOV. FREDERIKR ALWES. CATERINA 

BIEFIS & JULIA PINT 

Mitochondrial DNA and Decapod Phylogenies: The Importance of 47 

Pseudogenes and Primer Optimization 

CHRISTOPH D. SCHUBART 

Phylogenetic Inference Using Molecular Data 67 

FERRAN PALERO & KEITH A. CRANDALL 

Decapod Phylogeny: What Can Protein-Coding Genes Tell Us? 89 

K.H. CHU, L.M. TSANG. K.Y. MA. T.Y. CHAN & P.K.L. NG 

Spermatozoal Morphology and Its Bearing on Decapod Phylogeny 101 

CHRISTOPHER TUDGE 

The Evolution of Mating Systems in Decapod Crustaceans 121 

AKIRA ASAKURA 

A Shrimp's Eye View of Evolution: How Useful Are Visual Characters in 183 

Decapod Phylogenetics? 

MEGAN L. PORTER & THOMAS W. CRONIN 

Crustacean Parasites as Phylogenetic Indicators in Decapod Evolution 197 

CHRISTOPHER B. BOYKO & JASON D. WILLIAMS 

The Bearing of Larval Morphology on Brachyuran Phylogeny 221 

PAUL F. CLARK

vi Contents 

II Advances in Our Knowledge of Shrimp-Like Decapods 

Evolution and Radiation of Shrimp-Like Decapods: An Overview 245 

CHARLES H..I.M. ERANSEN & SAMMY DE GRAVE 

A Preliminary Phylogenelic Analysis of the Dendrobranchiata Based on 261 

Morphological Characters 

CAROLINA TAVARES. CRISTIANA SERE.IO & JOEL W. MARTIN 

Phvlogeny of the Infraorder Caridea Based on Mitochondrial and Nuclear 281 

Genes (Crustacea: Decapoda) 

HEATHER D. BRACKEN. SAMMY DE GRAVE & DARRYL L. FEEDER 

III Advances in Our Knowledge of the Thalassinidean 

and Lobster-Like Groups 

Molecular Phylogeny of the Thalassinidea Based on Nuclear and 309 

Mitochondrial Genes 

RAFAEL ROBLES. CHRISTOPHER C. TUDGE, PETER C. DWORSCHAK, GARY C.B. 

POORE & DARRYL L. FBLDER 

Molecular Phylogeny of the Family Callianassidae Based on Preliminary 327 

Analyses of Two Mitochondrial Genes 

DARRYL L. FELDER & RAFAEL ROBLES 

The Timing of the Diversification of the Freshwater Crayfishes 343 

JESSE BREINHOLT. MARCOS PEREZ-LOSADA & KEITH A. CRANDALL 

Phylogeny of Marine Clawed Lobster Families Nephropidae Dana. 1852. 357 

and Thaumastochelidae Bate. 1888, Based on Mitochondrial Genes 

DALE TSHUDY. RAFAEL ROBLES. TIN-YAM CHAN, KA CHAI HO. KA HOU CHU, 

SHANE T. AHYONG & DARRYL L. FELDER 

The Polychelidan Lobsters: Phylogeny and Systematics (Polychelida: 369 

Polychelidae) 

SHANE T. AHYONG 

IV Advances in Our Knowledge of the Anomttra 

Anomuran Phylogeny: New Insights from Molecular Data 399 

SHANE T. AHYONG, KAREEN E. SCHNABHL & ELIZABETH W. MAAS 

V Advances in Our Knowledge of the Brachyura 

Is the Brachyura Podotremata a Monophyletic Group? 417 

GERHARD SCHOLTZ & COLIN L. MCLAY

Contents vii 

Assessing the Contribution of Molecular and Larval Morphological 437 

Characters in a Combined Phylogenetic Analysis of the Supcrfamily 

Majoidea 

KRISTIN M. HUI.TGREN, GUILLERMO GUHRAO, HERNANDO RL. MARQUES & 

EHRRAN P. PALERO 

Molecular Genetic Re-Examination of Subfamilies and Polyphyly in the 457 

Family Pinnotheridae (Crustacea: Decapoda) 

EMMA PALACIOS-THEIL. JOSE A. CUESTA. ERNESTO CAMPOS & DARRYL L. 

FELDER 

Evolutionary Origin of the Gall Crabs (Family Cryptochiridae) Based on 475 

16S rDNA Sequence Data 

REGINA WETZER. JOEL W. MARTIN & SARAH L. BOYCE 

Systematics, Evolution, and Biogeography of Freshwater Crabs 491 

NEIL CUMBERLIDGE & PETER K.L. NG 

Phylogeny and Biogeography of Asian Freshwater Crabs of the Family 509 

Gecarcinucidae (Brachyura: Potamoidea) 

SEBASTIAN KLAUS. DIRK BRANDIS. PETER K.L. NG. DARREN C.J. YEO 

& CHRISTOPH D. SCHUBART 

A Proposal for a New Classification of Porlunoidea and Cancroidea 533 

(Brachyura: Heterotremata) Based on Two Independent Molecular 

Phylogenies 

CHRISTOPH D. SCHUBART & SILKE RRUSCHRL 

Molecular Phylogeny of Western Atlantic Representatives of the Genus 551 

Hexapanopeus (Decapoda: Brachyura: Panopeidae) 

BRENT P. THOMA. CHRISTOPH D. SCHUBART & DARRYL L. FELDER 

Molecular Phylogeny of the Genus Cronius Stimpson, I860, with 567 

Reassignment of C. tumidulus and Several American Species ol' Port un us 

to the Genus Achelous De Haan, 1833 (Brachyura: Portunidae) 

FERNANDO L. MANTELATTO. RAFAEL ROBLES. CHRISTOPH D. SCHUBART 

& DARRYL L. FELDER 

Index 581 

Color Insert

The Evolution of Mating Systems in Decapod Crustaceans 

AKIRAASAKURA 

Natural History Museum & Institute, Chiba, Japan 

ABSTRACT 

The mating systems of decapod crustaceans are reviewed and classified according to general patterns 

of lifestyles and male-female relations. The scheme employs criteria that focus on ecological, life 

history, and social determinants of both male and female behavior, and by these criteria nine types of 

mating systems are distinguished: (1) Short courtship: Both males and females are free-living (= not 

symbiotic with other organisms), and copulation occurs after brief behavioral interactions between a 

male and a female. (2) Precopulatory guarding: A male guards a mature female one to several days 

before copulation; both males and females are generally free-living. (3) Podding: In some largesize 

decapods, aggregations consisting of an extremely large number of individuals are formed, and 

mating occurs inside those aggregations. (4) Pair-bonding: In many symbiotic and some free-living 

species, males and females are found in a heterosexual pair and are regarded as having a monogamous 

mating system. They may live on or inside other organisms such as sponges, corals, molluscs, 

polychaetes, sea urchins, ascidians, and algal tubes. (5) Eusocial: In some sponge-dwelling snapping 

shrimps, a colony of shrimps contains a single reproductive female and many small individuals 

that apparently never breed. (6) Waving display: In many intertidal and semi-terrestrial crabs inhabiting 

mudflats or sandy beaches, males conduct visual displays that include species-specific dances 

to attract females. (7) Visiting: In some hapalocarcinid crabs, females are sealed inside a coral gall, 

and the male crab normally residing outside the gall is assumed to visit the gall for mating. (8) 

Reproductive swarm: In some pinnotherid crabs, mating occurs when a female is a free-swimming 

instar before she enters her definitive host. (9) Dwarf male mating: In some anomuran sand crabs, 

an extremely small male attaches near the gonopore of a free-living female. 

1 INTRODUCTION 

Decapod crustaceans are a large and diverse assemblage of animals. In most decapods, the sexes 

live separately and pair briefly as adults. Pairs are formed after a brief display, the sexes remain 

together for a relatively short period, the sexes separate after copulation, and the females assume 

all further parental duties such as selecting suitable habitat for egg incubation, aeration, and cleaning 

(Salmon 1983). However, recent discoveries of often-conspicuous behavior and male-female 

relations among decapods have shown that their mating system is highly diverse and is sometimes 

quite similar to mating systems of other animals such as birds, mammals, reptiles, and insects (see 

Shuster & Wade 2003; Duffy & Thiel 2007 for a review). 

As claimed by Emlen & Oring (1977) in their classic work on the relationships among ecological 

factors, sexual selection, and the evolution of mating system, sexual selection is the driving force that 

underlies the evolution of male-male competition and female choice. However, ecological factors 

apparently contribute to the evolution of mating systems as well as to behavioral and morphological 

differences between the sexes. From this point of view, much study has been conducted recently on 

the evolution of the mating system of decapods (see section 2 below).

122 Asakura 

In this paper, I describe the diversity of mating systems of decapods in an attempt to recognize 

and classify their general patterns from the viewpoints of the ecological, life history, and social determinants 

of both male and female behavior. Historically, there are two ways of describing mating 

systems (Shuster & Wade 2003). The first is in behavioral ecology, where mating systems are usually 

described in terms of the number of mates per male or female, such as monogamy, polygyny, and 

polyandry. The second is in terms of the genetic relationships between mating males and females, 

such as random mating, negative assortative mating (outbreeding), and positive assortative mating 

(inbreeding). My approach to describing mating systems of decapods is a "recognition of general 

pattern" approach, a kind of a combination of these two approaches that captures variation in the 

relationship between male and female, from promiscuity to monogamy, as well as the relationship 

between male guarding and the female tendency to settle down in certain places or to aggregate, and 

the complex nature of eusociality. 

Terminology generally follows Duffy & Thiel (2007). Additionally, some basic terms are redefined 

here, because these terms are sometimes used in more or less different ways according to taxa, 

including birds, mammals, and fish: 

• Monogamy (= pair bonding): One male and one female have an exclusive mating relationship. 

• Polygamy: One or more males have an exclusive relationship with one or more females. Three 

types are recognized: polygyny, where one male has an exclusive relationship with two or more 

females; polyandry, where one female has an exclusive relationship with two or more males; 

and polygynandry, where two or more males have an exclusive relationship with two or more 

females (the numbers of males and females need not be equal, and, in vertebrate species studied 

so far, the number of males is usually fewer). 

• Promiscuity: Any male within the group mates with any female. 

• Eusociality: Multigenerational (cohabitation of different generations), cooperative colonies with 

strong reproductive skew (reproductive division of labor, usually a single breeding female) and 

cooperative defense of the colony (after Duffy 2003). 

• Symbiosis: Here defined simply as dissimilar organisms living together. 

2 HISTORY OF STUDY 

The first important review of decapod mating systems was Hartnoll's (1969) publication on brachyuran 

crabs. He distinguished two types of mating systems. "Soft-female mating" was defined as copulation 

occurring immediately after molting of the female, usually preceded by a lengthy pre-molt 

courtship behavior including precopulatory guarding by the male. "Hard-female mating" was defined 

as mating in which the female copulates during the intermolt stage after a relatively brief 

courtship behavior. 

Through their intensive study of the harlequin shrimp Hymenocera picta, Wickler & Seibt (see 

Reference 16 in Appendix I, Table 10) found that these shrimp form stable heterosexual pairs based 

on individual recognition by chemical cues at a distance. Wickler & Seibt discussed several similar 

hypotheses, independently developed in research on crustaceans and humans, for the evolution of 

monogamy and other mating systems. Individual recognition in the monogamous mating system 

was intensively studied in the banded shrimp Stenopus hispidus by Johnson (1969, 1977). 

The report by Emlen & Oring (1977) was influential for studies on crustacean mating systems. 

They classified the mating system into the following categories: 

1. Monogamy 

2. Polygyny (subdivided into 2a, resource defense polygyny; 2b, female (or harem) defense 

polygyny; and 2c, male dominance polygyny (further subdivided into 2c-1, explosive breeding 

assemblages, and 2c-2, leks))


3. Rapid multiple clutch polygamy 

4. Polyandry (subdivided into 4a, resource defense polyandry; and 4b, female access polyandry) 

Ridley (1983) intensively reviewed the precopulatory mate guarding behavior in various groups 

of animals including tardigrades, crustaceans, arachnids, and anurans, and discussed its evolution. 

Work on the behavior of the fiddler crabs (genus Uca) has contributed greatly to our understanding 

of the mating systems of brachyuran crabs. These studies include the works of H.O. von Hagen 

(e.g., von Hagen 1970), J. Crane (e.g., Crane 1975), J. Christy and his coworkers (e.g., Christy 

et al. 2003a, b), M. Salmon and his coworkers (e.g., Salmon & Hyatt 1979), R R. Y. Backwell and 

her coworkers (e.g., Backwell et al. 2000), M. Murai and his coworkers (e.g., Murai et al. 2002), 

and T. Yamaguchi (e.g., Yamaguchi 2001a, b). Based on the studies of Uca and other brachyurans, 

as well as other decapods, Salmon (1983) reported the diversity of behavioral interactions 

preceding mating in decapods, and he defined some of the consequences of these interactions in 

terms of sexual selection, courtship behavior, and mating systems. The book edited by Reback & 

Dunham (1983), which included Salmon's (1983) work, was a landmark in the study of decapod 

behavior. 

Christy (1987) reviewed the mating systems of brachyuran crabs and classified them, according 

to modes of competition among males for females, into three major categories and eight subcategories, 

as follows. 

1. Female-centered competition, including: la, defense of mobile females following free search; 

lb, defense of sedentary females following a restricted search; lc, capture, carrying, and 

defense of females at protected mating sites; and Id, attraction and defense of females at 

protected mating sites 

2. Resource-centered competition, including: 2a, defense of breeding sites; and 2b, defense of 

refuges 

3. Encounter rate competition, including: 3a, neighborhoods of dominance; and 3b, pure search 

and interception 

In their book on crustacean sexual biology, Bauer & Martin (1991) introduced developments in 

various fields and taxa of crustacean research, including studies on sex attraction, sex recognition, 

mating behavior, mating system, and structure and function associated with insemination. Bauer and 

his coworkers have extensively studied the mating behavior, mating system, and hermaphroditism 

of shrimps (e.g., see Bauer 2004 for a review). 

Through their intensive studies on the mating system of the spider crab Inachus and of the extended 

maternal care of semi-terrestrial grapsid crabs of Jamaica, Diesel and his coworker revealed 

examples of highly specialized mating and social systems in these crabs (see Diesel 1991; Diesel & 

Schubart 2D07 for reviews). 

Thiel and his students have conducted intensive research on the mating system of rock shrimps 

(see Reference 6 in Appendix I, Table 4) and symbiotic anomuran crabs (e.g., Baeza & Thiel 2003). 

Based on these studies, Thiel & Baeza (2001) and Baeza & Thiel (2007) reviewed factors affecting 

the social behavior of marine crustaceans living symbolically with other invertebrates. Similarly, 

Correa & Thiel (2003) reviewed mating systems in caridean shrimp and their evolutionary consequences 

for sexual dimorphism and reproductive biology. The book by Duffy & Thiel (2007) on the 

evolutionary ecology of social and sexual systems of crustaceans is a monumental landmark that 

synthesizes the state of the field in crustacean behavior and sociobiology and places it in a conceptually 

based, comparative framework. The relatively recent discovery of eusociality in snapping 

shrimp by Duffy has opened the door to a new field in social and mating systems of decapods (see 

Duffy 2007 for a review; see also sections 3.5 Eusocial type and 4.5 Evolution of the eusocial type 

below for further explanation).

124 Asakura 

Asakura (1987, 1990, 1993, 1994, 1995, 1998a, 1998b, 1999, 2001a, b, c), Imazu & Asakura 

(1994, 2006), and Nomura & Asakura (1998) reported mating systems and various aspects of sexual 

differences in the ecology and behavior of hermit crabs and other decapods. 

3 TYPES OF MATING SYSTEMS 

3.1 Short courtship type 

This type is generally seen in species whose males and females are free living, that is, not symbiotic 

with other organisms (Appendix 1, Tables 1, 2). Copulation occurs after a short courtship behavior 

by the male, or copulation occurs just after brief behavioral interactions between a male and a 

female. This type of courtship includes very different groups of decapods, from the most primitive 

group (dendrobranchiate shrimps) to groups specialized for certain habitats such as freshwater crayfishes, 

intertidal hermit crabs, and semi-terrestrial and terrestrial brachyuran crabs. It is perhaps the 

most widely seen mating system in decapods. 

No intensive aggressive behavior between males (for a female) has been reported in species of 

dendrobranchiate shrimps of the families Penaeidae and Sicyoniidae, caridean shrimps of the families 

Palaemonidae, Hyppolytidae, and Pandalidae, or anomuran sand crabs of the family Hippidae. 

In these species, females are generally similar in size to, or larger than, males. On the other hand, 

strong aggressive interaction is seen between males in freshwater crayfish species of all three families 

(Astacidae, Parastasidae and Cambaridae) as well as in brachyuran crabs of the Grapsoidea and 

Gecarcinidae. In these species, the male body and weaponry (chelipeds) are generally larger than 

the female. 

Among decapods exhibiting this mating system are species whose females molt before copulation 

(Appendix 1, Table 1) and those whose females do not molt before copulation (Appendix 1, 

Table 2). In species inhabiting terrestrial and semi-terrestrial habitats, females generally copulate 

in the hard shell condition; these species include land hermit crabs of the genus Coenobita and 

brachyuran crabs of the Grapsoidea and Gecarcinidae. 

In penaeid shrimp, the molting condition of copulating females is determined according to the 

type of thelycum. The thelycum is the female genital area, i.e., modifications of female thoracic 

sternites 7 and 8 (sometimes including thoracic sternite 6) that are related to sperm transfer and 

storage. A female with externally deposited spermatophores is said to have an "open thelycum," 

which is formed by modifications of the posterior coxae and sternites to which the spermatophores 

attach. Primitive dendrobranchiate shrimps, including species of the families Aristeidae, Solenoceridae, 

Benthesicymidae, and the penaeid genus Litopenaeus, have open thelyca. In these species, 

females copulate in the hard shell condition. On the other hand, a "closed thelycum" refers to sternal 

plates that may (1) enclose a noninvaginated seminal or sperm receptacle, (2) cover a space that 

leads to spermathecal opening, or (3) form an external shield guarding the spermathecal openings. 

In the most advanced groups, including the penaeoid genera Fenneropenaeus, Penaeus, Farfantepenaeus, 

Melicertus, Marsupenaeus, Trachypenaeus, and Xiphopenaeus, females have closed thelyca. 

In these species, females molt just before copulation. Since no significant difference is seen in mating 

behavior between the open thelycum species and the closed thelycum species, Hartnoll's (1969) 

rule, which predicts a lengthy pre-molt courtship behavior associated with soft-female mating and a 

relatively brief courtship behavior with hard-female mating, does not hold in the case of the penaeid 

shrimps. 

A sperm plug, which is believed to preclude subsequent insemination by other males, is known 

in some species of Farfantepenaeus, Marsupenaeus, Metapenaeus, and Rimapenaeus (Appendix 1, 

Table 3). 

In all the above-mentioned taxa, copulation generally continues only for several minutes. After 

mating, the male separates from the female and presumably goes on to search for other females.


The habitat of species that exhibit this mating system varies, ranging from terrestrial through intertidal 

to deep water. 

3.2 Precopulatory guarding type 

This mating system also is generally seen in species whose males and females are free living 

(Appendix 1, Table 4). A male guards a mature female for one to several days before copulation. 

Generally, males aggressively fight for a female using their cheliped(s) and sometimes also the ambulatory 

pereopods. In some species, females always molt prior to mating and copulation; in other 

species, females may or may not molt prior to copulation. There are two types of guarding: (1) contact 

guarding of hermit crabs and brachyuran crabs, in which a male grasps part of the appendages, 

the body, or the shell (in the case of hermit crabs) of a mature female, and (2) non-contact guarding, 

as exhibited in Macrobrachium shrimps and Homarus lobsters, in which a male keeps a female 

without grasping her. After mating, postcopulatory guarding by a male for a female is sometimes 

observed (Appendix 1, Table 5). However, after postcopulatory guarding, or just after copulation, 

the male and female separate so that both may later mate with other individuals. Generally, in this 

mating system, the body size of males is larger than that of females, or weaponry (chelipeds) is 

more developed in males than in females. 

Species of the river prawn genus Macrobrachium are well known for the extremely long chelipeds 

in males. A male guards a female for one to several days before copulation and fights with 

other males using these chelipeds. In some species, such as M. australiense, a male has a nest (a 

saucer-shaped depression on the bottom), beckons a female to the nest, and guards and copulates 

with her in the nest. In the American lobster Homarus americanus, a male guards a female in his 

shelter, which is dug under rocks, boulders, or eelgrass, and the cohabitation of a male and a female 

lasts from one to three weeks. 

In hermit crabs of the genus Diogenes (Diogenidae) and in many species of the family Paguridae, 

al of which have unequal chelipeds in terms of both size and morphology, a male grasps the rim 

of the shell inhabited by a mature female by the minor cheliped, guards her for one to several 

days before copulation, and fights with other males approaching him using the major cheliped. 

In crab-shaped anomurans, the male Paralithodes brevipes conducts both pre-copulatory and postcopulatory 

guarding. The male claims a female by grasping her chelae or legs with his chelae, or 

he covers the female with his body. Similarly, the male Hapalogaster dentata grasps a female with 

his left chela and covers the female with his body; these guarding behaviors occur one to three days 

before copulation. 

In the brachyuran crab Corystes cassivelaunus (Corystidae), the male carries the female in his 

chelae, and, while stationary, holds one or both of the female's chelae in his own and holds her 

carapace close to his sternum. Such behavior continues up to several days before copulation. In 

species of the Cancridae and Portunidae, males carry the pre-molt female with her carapace or sternum 

held against the sternum of the male for a period of days; after this period the male releases 

the female so that she molts, and copulation occurs shortly after the molting. In many species in 

these two families, the male continues to carry the female after copulation in the pre-molt position 

until her integument has partially hardened. Sperm plugs, which are regarded as being produced 

by the males to block the females' genital duct to preclude subsequent insemination by other males 

(Diesel 1991), also are often reported for species of these families (Appendix 1, Table 6). In Menippe 

mercenaria (Xanthidae), the male guards the entrance to the burrow occupied by the pre-molt female, 

and they copulate as soon as the female molts. In species of the Majidae and Cheiragonidae, 

the male guards the female before copulation in a manner similar to what is seen in the Cancridae 

and Portunidae, where the male grasps the ambulatory pereopods, chelipeds, or body of the 

female. 

Species that exhibit this mating system are from the intertidal through shallow water to deep 

waters, but they are not found in terrestrial or semi-terrestrial environments.

126 Asakura 

3.3 Podding 

In large decapods inhabiting shallow waters, an aggregation consisting of an extremely large number 

of individuals in certain places is called a "pod." Podding is regarded as a type of behavior that is 

optional and that is associated with different stages in the species' life history, such as molting, mating, 

and the incubation period (Appendix 1, Table 7). The pod is also called a "heap" or "mound," 

according to the locality and/or the species. 

The function of the pod may vary depending on the condition of the specimens within it (such as 

level of maturity, sex, intermolt stage) and possibly on changes in habitat condition, such as water 

temperature and presence of predators (Sampedro & Gonzalez-Gurriaran 2004). However, as listed 

in Appendix 1, Table 7, pods in some species have the function of facilitating mating, so I will treat 

this as a special kind of mass mating in some species. 

Stevens (2003) and Stevens et al. (1994), reporting more than 200 pods with a total of 100,000 

crabs of the majid Chionoecetes bairdi in an area of only 2 ha off Kodiak Island in Alaska in 1991, 

observed that the formation of the pods and mating synchronized with the spring tide. Similar observations 

were made for another majid, Hyas lyratus, by Stevens et al. (1992), who reported large 

aggregations during the mating season from off Kodiak Island. They found 200 mating pairs (males 

grasping females) among 2000 individuals in one pod. The majid crab Loxorhynchus grandis, distributed 

along the east coast of North America, often forms large aggregations numbering hundreds. 

of animals. The aggregation is composed of crabs of both sexes, and the function is thought to be 

the attraction of males for mating (Hobday & Rumsey 1999). DeGoursey & Auster (1992) reported 

large mating aggregations in another majid crab, Libinia emarginata, in April and May 1989. Many 

mating pairs were found in the aggregations, and the percentage of ovigerous females among all 

females increased from 26% on 1 May to 100% on 14 May. Males paired with females were significantly 

larger than unpaired males, while the paired and unpaired females were not significantly 

different in size. Carlisle (1957) monitored a pod consisting of 60-80 individuals of the majid crab 

Maja squinado in shallow waters in the English Channel; 20 were adult males and the rest were 

juvenile males and females in equal amounts. He observed crabs molting inside the pod and mating 

between intermolt males and postmolt females, which led him to conclude that the main purpose 

of podding is to provide protection for newly molted soft crabs against predators and to facilitate 

mating. However, later behavioral observations by Hartnoll (1969) indicated that copulation occurs 

between a male and a female in the intermolt stage. Furthermore, Sampedro & Gonzalez-Gurriaran 

(2004) found that the gonads of females in the pods were in an early stage of development (= not 

fully matured) and that the spermathecae were empty, suggesting to them that mating of this species 

occurs in deeper waters. 

In crab-shaped anomurans, large pods of the red king crab Paralithodes camtschaticus are well 

known in the northern Pacific Ocean, with each pod consisting of thousands of crabs in the 2-4 

year class (juveniles). Aggregations of adult red king crabs (ovigerous females) also were reported 

and are thought to be related to mating (Stone et al. 1993), but detailed surveys have not been 

conducted. Dense aggregations of the southern king crab Lithodes santolla have been reported from 

Chile (South America); however, the crabs forming these aggregations are juveniles, so this behavior 

is not thought be related to mating (Cardenas et al. 2007). 

In summary, podding is known only in large species distributed in temperate or boreal waters in 

both the Pacific and Atlantic oceans. 

3.4 Pair-bonding type 

Many species of decapods, in particular those that are symbiotic with other animals, have been 

reported as "found in a heterosexual pair" (Appendix 1, Tables 8-12). Most of these are considered


to have a monogamous mating system, which is well known in birds and mammals. In species whose 

males engage in mate-guarding, temporal heterosexual pairing occurs, where the pair is formed 

when the female is close to molting or spawning a new batch of unfertilized eggs, and the mateguarding 

males abandon the females soon after the eggs are fertilized. However, in pair-bonding 

species, males cohabit with females, independent of their reproductive status or of the stage of 

development of the brooded embryos. Nevertheless, the observations for the monogamous nature 

of these pair-bonding species are often only anecdotal, and how long the pair remains together, and 

with whom they mate, is rarely recorded. Some well-documented studies include the formation of 

stable pairing and individual recognition (individuals in a pair can recognize each other as mates), 

as in the case of the banded shrimp Stenopus hispidus (Reference 8 in Appendix 1, Table 10), the 

scarlet cleaner shrimp Lysmata debelius (Reference 12 in Appendix 1, Table 10), and the harlequin 

shrimp Hymenocera picta (Reference 16 in Appendix 1, Table 10). 

Detailed observations of the monogamous nature of pairing have been made for several species 

of snapping shrimps, for example, Alpheus angulatus (Reference 97 in Appendix 1, Table 9), 

Alpheus heterochaelis (Reference 99 in Appendix 1, Table 9), Alpheus armatus (Reference 28 in 

Appendix 1, Table 9), and Alpheus roquensis (Reference 31 in Appendix 1, Table 9), as well as for 

the pontoniid shrimp Pontonia margarita (Reference 45 in Appendix 1, Table 8), the deep-water 

sponge-dwelling shrimp Spongicola japonica (Reference 1 in Appendix 1, Table 10), a porcelain 

crab Poly onyx gibbesi (Reference 11 in Appendix 1, Table 11), and several species of coral crabs 

of the genus Trapezia (References 2-14 in Appendix 1, Table 12). Many pair-bonding species are 

known in caridean shrimps of the subfamily Pontoniinae and family Alpheidae, "cleaner" shrimps 

of the families Stenopodidae and Spongicolidae, crab-shaped anomurans (family Porcellanidae), 

and brachyuran crabs of the family Trapeziidae. 

Most of these species are symbiotic with other animals or live in special habitats. Host animals 

for these species include sponges, sea anemones, black corals, reef-building corals, gastropods, 

opistobranch molluscs, bivalves, polychaetes, crinoid feather stars, sea stars, sea urchins, sea cucumbers, 

and ascidians. The special habitats include gastropod shells used by large hermit crabs; tubes 

of polychaetes such as Chaetopterus; soft, web-like tubes consisting of filamentous algae, sponges, 

and other debris built by shrimp themselves; burrows excavated in hard dead corals; burrows of gobiid 

fish; and burrows of the thalassinidean shrimp genus Upogebia. However, free-living species are 

also known, such as stenopodid shrimps inhabiting rocky subtidal zones and many alpheid shrimp 

species inhabiting rock crevices or found under rubble, around large algae, or in burrows of their 

own in mudflats and other soft bottoms. 

The following generalizations can be made for almost all of these species. They are territorial, 

and they cooperatively defend their habitats (hosts, special habitats, and burrows) against other conspecific 

or non-conspecific animals. Thus, the mating system of these species is termed "resourcedefense 

monogamy." The pairs are size-matched (— size-assortative pairing); there is strict preference 

exerted by either sex for mates of a particular size relative to themselves. Baeza (2008) proposed 

two possible explanations for this phenomenon in his study on pontoniid shrimps symbiotic 

with bivalves: 

1. The two sexes might choose large individuals of the opposite sex as sexual partners and host 

companions. In males, a preference for large females should be adaptive, as female size is 

positively correlated with fecundity in shrimps. In females, sharing a host with a large male 

might result in indirect benefits (i.e., good genes) or direct benefits (increased protection 

against predators or competitors). 

2. Choice of a certain-size partner could also be a consequence of constraints in the growth rates 

of shrimps dictated by host individuals. Space limitations for shrimps in hosts are suggested 

by the tight relationship between shrimp and host size, and by the fact that hosts harboring 

solitary or no shrimps were among the small hosts.

128 Asakura 

These species tend to display low sexual dimorphism in weaponry in terms of cheliped size and morphology 

and often in body size. This is in contrast to the large sexual differences in mate-guarding 

species in which the weaponry is much more developed and where body size is often much larger 

in males than in females. Regarding body size, there is a tendency in pair-bonding shrimp for the 

male to be slightly smaller, in terms of body length, and much more slender than its mate female; 

in trapeziid crabs the male is often slightly larger than his female mate. 

The bathymetric distribution of species with this mating system is generally from intertidal to 

shallow water, but a few groups of species, such as those of the Spongicolidae, inhabit deep water. 

3.5 Eusociality type 

Until the discovery of the eusocial shrimp Zuzalpheus regalis (as Synalpheus regalis) (Duffy 1996), 

eusociality was recognized only among social insects, including ants, bees, and wasps (Hymenoptera) 

and termites (Isoptera); in gall-making aphids (Hemiptera); in thrips (Thysanoptera); and in 

two mammal species, the naked mole rat (Heterocephalus glaber) and the damaraland mole rat 

(Cryptomys damarensis). Zuzalpheus regalis lives inside large sponges in colonies of up to >300 

individuals, with each colony containing a single reproductive female. Direct-developing juveniles 

remain in the natal sponge, and allozyme data indicate that most colony members are full siblings. 

Larger members of the colony, most of whom apparently never breed, defend the colony against 

heterospecific intruders (Duffy 1996). 

Following this initial discovery, Duffy and his coworkers have found several other species 

of Zuzalpheus exhibiting monogynous, eusocial colony organization in the western Atlantic 

(Appendix 1, Table 13). In the Indo-west Pacific region, Didderen et al. (2006) found a colony 

of a sponge-dwelling alpheid shrimp, Synalpheus neptunus neptunus, with one large ovigerous female 

or "queen" together with many small individuals, indicating a eusocial colony organization 

(Appendix 1, Table 13). 

Some 20 species of symbiotic decapod species have been reported as found in a group 

(Appendix 1, Tables 14-15). Among them, examples of Synalpheus and Zuzalpheus exhibited more 

than 100 individuals in one aggregation, and, in particular in the case of Zuzalpheus brooksi, more 

than 1000 individuals were recorded from one sponge. These aggregations are regarded either as 

having a non-social structure (Thiel & Baeza 2001) or with the social structure totally unknown. 

3.6 Waving display type 

In many species of the crab families Ocypodidae, Dotillidae, and Macrophthalmidae, and in species 

of the genus Metaplax of the family Varunidae (formerly subfamily Varuninae in the Grapsidae 

sensu lato), males perform waving displays using the chelipeds. As in many other territory advertisement 

signals in animals, this behavior is commonly thought to have the dual function of simultaneously, 

repelling males and attracting females (e.g., Salmon 1987; Crane 1975). These species 

typically live in mudflats, tidal creeks, sandbars, and mangrove forests, and each individual has its 

own burrow with a small territory around it. They often occur in huge numbers, with thousands of 

individuals living in small, adjacent territories, and with males and females living intermixed. The 

burrow serves various functions, including a refuge during high tide, an escape from predators, and 

the site of mating, oviposition, and incubation. 

The behavior and mating systems of fiddler crabs (genus Uca, Ocypodidae) have been intensively 

studied (see references in History of Study, above). There are species whose males defend 

burrows from which they court females and species whose males wander from their burrows and 

court females on the surface (Christy 1987). For the former group of species, the following generalization 

is possible (based mainly on P. Backwell and coworkers; see references in History of Study, 

above). Males wave their enlarged claw, and, when a female is ready to mate (i.e., she matures), 

she leaves her own burrow and wanders through the population of waving males. The female visits


several males before selecting a mate, and a visit consists of a direct approach to the male. Before 

copulation^ both individuals enter the male's burrow, and two behavioral patterns are known: the 

male enters his burrow first and the female follows him in, or it happens in the reverse order, i.e., 

the female enters first. The male then gathers up sand or mud to plug the burrow entrance. Mating 

occurs in the burrow. On the following day, the male emerges, reseals the burrow entrance with the 

female still underground, and leaves the area. The female remains underground for the following 

few weeks while she incubates her eggs. 

In addition to waving displays, males of some fiddler crab species employ acoustic signals to 

attract females. In these species, males attract females during the day first by waving and then by 

producing sounds just within their burrows. At night, the males produce sounds at low rates, but 

when touched by a female they increase their rate of sound production (Salmon & Atsaides 1968). 

Many species of ocypodid crabs build sand structures next to their burrows, some of which function 

to attract females for mating, such as pillars (Uca: Christy 1988a, b), hoods (Uca: Zucker 1974, 

1981; Christy et al. 2002, 2003a, b), mudballs (Uca: Oliveira et al. 1998), and pyramids (Ocypode: 

Linsenmair 1967; Hughes 1973). 

3.7 Visiting type 

An interesting mating system has been suggested for coral gall crabs (family Cryptochiridae), which 

inhabit cavities in scleractinian corals in (usually) shallow water. However, the information is still 

anecdotal, based on ecological observations on Hapalocarcinus marsupialis, Troglocarcinus corallicola, 

and Opecarcinus hypostegus (Potts 1915; Fize 1956; Kropp & Manning 1987; Takeda & 

Tamura 1981; Hiro 1937; Kotb & Hartnoll 2002; Carricart-Ganivet et al. 2004). In H. marsupialis 

and T. corallicola, the male crab normally resides outside the gall, which was constructed by the 

female, and is thought to visit the gall of the female for mating. The males and females apparently 

show promiscuity, and male-male aggressive behavior for a female has not been reported. The female 

is much larger than the male and in some species has a soft body with a very large abdomen. On 

thfe other hand, the male is usually hard, with a small abdomen. Geographical distribution includes 

mostly the tropics (see Wetzer et al. this volume). 

In Opecarcinus hypostegus, couples were found sharing cavities; ovigerous females and males 

are recorded inhabiting adjoining cavities on colonies of Siderastrea stellata corals (Carricart- 

Ganivet et al. 2004). This species may have a mating system different from the above. 

3.8 Reproductive swarm type 

This mating system is reported only in pinnotherid crabs that are considered parasitic or co-inhabiting 

with other animals, including bivalves, gastropods, sea slugs, chitons, polychaetes, echinoderms, 

burrowing crustaceans, and sea squirts (Cheng 1967; Gotto 1969). In several species of these crabs, 

mating occurs, or is thought to occur, when the female is in the free-swimming stage before she 

enters into her definitive host (Appendix 1, Table 16). 

The following generalization is possible for these species. Adult females have a soft, membranous 

carapace, and generally each one lives by itself within its host animal. These females produce 

broods of planktonic larvae. After development, the larvae metamorphose into the "invasive stage" 

crab, which is morphologically similar to the later swimming stage in having a flattened shape and 

ambulatory legs with dense setae adapted for swimming. Following this stage is a stage designated 

as "prehard"; these crabs invade, and live in, the host invertebrate animals. The crab at this stage 

is soft, resembling the later posthard stage. These crabs grow and mature into small adults of both 

sexes and leave their host to join mating swarms in open water. This stage is called the "hard stage," 

swimming stage, or copulation stage, and it is characterized by a hard body, swimming legs densely 

fringed with setae, and a thick fringe of setae along the front of the carapace. They copulate at 

this stage, and, in all reported species (see Appendix 1, Table 16), females copulate in the hard

130 Asakura 

shell condition. After copulation, each female enters the host animal, but the male dies. The female 

becomes soft and grows much larger in the host, and later the female produces eggs fertilized by 

sperm from her single mating. 

This is a kind of mass mating, with males and females showing promiscuity. In the copulation 

stage, no intensive aggressive behavior between males for females has been reported. The males 

in this stage are slightly larger than the females, and the morphology is similar between the sexes. 

After the female enters the host animal, the female becomes soft and grows much larger and stouter. 

The species with this mating system are found generally from intertidal to shallow water where their 

host invertebrates occur. In some pinnotherid species, adult crabs are found in a heterosexual pair in 

the host animal, although life history and mating systems of these species are mostly unknown. 

3.9 Neotenous male type 

Extremely small, neotenous males exist in some species of anomuran sand crabs (genus Emerita) 

inhabiting wave-exposed sandy beaches in tropical and temperate waters (Appendix 1, Table 17). In 

these species, the males become sexually mature soon after their arrival on the beach as a megalopa. 

When copulating, a male attaches near one of the female's gonopores, which are located on the 

coxae of the third pereopods. Surprisingly, the size of the neotenous males is similar to, or smaller 

than, those coxae. 

Protandric hermaphroditism is described in detail in Emerita asiatica as it relates to neotenous 

males (Subramoniam 1981). The neotenous males occur at 3.5 mm carapace length (CL) and above, 

whereas females acquire sexual maturity at 19 mm CL. The neotenous males, as they continue 

to grow, gradually lose male functions and reverse sex at about 19 mm CL. In the CL range of 

19-22 mm, the male's gonad consists of inactive testicular and active ovarian portions. Androgenic 

glands, active in the neotenous males, show signs of degeneration in the larger males and disappear 

in the intersexuals. 

The male separates from the female after copulation. Aggressive behavior between males is not 

reported. As opposed to the female, the neotenous male shows a general simplicity of appendages 

associated with its small size. Among decapods, this phenomenon is known only in species of 

Emerita. 

4 EVOLUTION OF MATING SYSTEMS IN DECAPODA 

4.1 Introduction 

It is apparent from the above that similar mating systems have evolved independently in different 

taxa at different times; i.e., convergent evolution is widespread. Species in ecologically similar 

habitats often display patterns that are strikingly comparable. Here I discuss the possible origin and 

evolutionary pathway of each mating system and compare them with those of other animals. 

4.2 Evolution of the short courtship type and the precopulatory type 

These two mating systems are most dominant among decapods. The mode of life is often quite 

similar; both males and females are free living (not symbiotic with other organisms), and after 

mating the male soon separates from the female. However, the habitat is sometimes different; in 

terrestrial and freshwater species, only the short courtship type has been reported. Therefore, a 

question arises as to why some groups of species have evolved the prolonged precopulatory mate 

guarding, whereas others have not. 

Precopulatory mate guarding is known in a very broad range of taxa such as tardigrades, crustaceans, 

arachnids, and anurans (Parker 1974; Ridley 1983; Conlan 1991). It is thought to evolve 

when male-male competition for females is strong enough and female receptivity is restricted in


time (Parker 1974; Jormalainen 1998), or even if receptivity is not time-limited but the guarding 

costs are low enough (Yamamura 1987). Guarding should be beneficial to the male, if the expected 

fitness gain achieved by guarding is greater than that expected by continuing to search for other 

females (Parker 1974). Thus, the optimal guarding duration for the male is determined by the encounter 

rate of females and the costs of guarding relative to those of searching (Yamamura 1987). 

The cost of guarding for males includes decreased mobility and feeding (Adams et al. 1985, 1991; 

Robinson & Doyle 1985), an increase in predation risk while guarding (Verrel 1985; Ward 1986), increased 

energetic costs associated with carrying females (Sparkes et al. 1996; Plaistow et al. 2003), 

and an increase in fighting costs through male-male conflict (Benesh et al. 2007; Yamamura & 

Jormalainen 1996). Additionally, a long guarding time decreases future opportunities to mate with 

other females (Benesh et al. 2007). 

Pelagic dendrobranchiate and caridean shrimps are primarily swimmers, and possibly for that 

reason they have not evolved prolonged, elaborate behavioral interactions before copulation. However, 

the above-mentioned energetic cost hypothesis (Sparkes et al. 1996; Plaistow et al. 2003) may 

be applicable; for males of these species, carrying a swimming female for a long duration requires 

much more energy than in benthic species. In fact, all species exhibiting a prolonged precoulatory 

guarding period are benthic species. 

In all freshwater crayfish studied, the mating system includes a short courtship without a lengthy 

precopulatory guarding, even though they have a benthic lifestyle and male-male aggression is 

often common. They may live in their burrows separately, or underneath boulders or heaps of fallen 

leaves, and these habitats are quite similar to, or virtually the same as, those of shrimps of the genus 

Macrobrachium. Why males of Macrobrachium adopt a precopulatory guarding strategy whereas 

male crayfish do not is not known. 

A similar question arises in intertidal and shallow water decapods. For example, intertidal hermit 

crab species exhibiting precopulatory guarding have a tendency toward vastly unequal chelipeds, 

with a well-developed major cheliped particularly in males, who use it for fighting with 

olher males during guarding. Such species include those of the genera Pagurus (Paguridae) and 

Dmgenes (Diogenidae). On the other hand, species of Paguristes have small and similar right and 

left chelipeds and execute short courtship mating; males do not aggressively fight with other males. 

Species of Calcinus, which conduct short courtship type mating, often have vastly unequal chelipeds, 

with the well-developed major cheliped similar to those species that display precopulatory 

guarding. However, males of Calcinus species do not aggressively fight with each other during mating. 

Further study is needed to clarify the relationship between mating behavior and morphology. 

In land hermits and land brachyurans, the above-mentioned predation risk hypothesis (Verrel 

1985; Ward 1986) may be applicable to those species where mating system is the short-courtship 

type with hard-female mating. Male-male aggression is common in these taxa, but they have never 

evolved precopulatory guarding. Prolonged guarding may carry the risk of attack by visual predators 

such as birds in a terrestrial environment. In these taxa, a strong connection exists between a 

prolonged precopulatory guarding and soft-female mating as well as between a short courtship and 

hard-female mating. When marine species adapted to land, the former mating system might have 

been lost and changed to the latter, i.e., from soft-female to hard-female, to avoid desiccation and 

to deal with the large and often unpredicted fluctuations in availabilities of females in a terrestrial 

environment. 

The evolution of sperm plugs in species of short-courtship type (penaeid shrimps) and precopulatory 

type (brachyuran crabs) is interesting. The sperm plug has virtually the same function as 

the copulation plug (= copulatory plug, mating plug) in mammals (rodents, bats, monkeys, koala), 

reptiles (snakes and lizards), insects (butterflies, ants, dragonflies, and stinkbugs), spiders, and acanthocephalan 

worms (Smith 1984). These plugs, secreted by the male after mating, serve to block the 

female tract for some time to prevent further mating by other males.

132 Asakura 

4.3 Evolution of the podding type 

Why many animal species (e.g., insects, fish, birds, and herbivorous mammals) group together is 

one of the most fundamental questions in evolutionary ecology. It is believed that strong selective 

pressures lead to aggregation rather than to a solitary existence in most of these groups. These pressures 

include protection against predators, increased foraging efficiency, increased ease of assessing 

potential mates, and increased information exchange about the location of food (Barta & Giraldeau 

2001). Similarly, various ecological reasons for the formation of pods have been proposed, including 

protection during molting, location of mates, aiding in food capture, and protection from predation 

(see References in Appendix 1, Table 7). Why some species evolved aggregating behavior and others 

did not is unknown. 

4.4 Evolution of the pair-bonding type 

Heterosexual pairing behavior ("social monogamy," Gowaty 1996; Bull et al. 1998; Gillette et al. 

2000; Wickler & Seibt 1981) has evolved many times in a broad range of animal taxa, including 

mammals, birds, reptiles, amphibians, fish, insects, and crustaceans. For example, a colony of 

scleractinian coral sometimes yields a pair of goby fish, alpheid shrimps, and trapeziid crabs. Researchers 

interested in social system evolution must look for ecological and physiological factors 

(beyond basic sexual differences) that may make social monogamy selectively advantageous to individual 

males and/or females. Of particular interest are factors that may consistently correlate with 

such behavior across taxonomic groups. Several hypotheses for the evolution of social monogamy 

have been developed [see also Mathews (2002b), Baeza (2008), Baeza & Thiel (2007) for a review], 

as follows. 

Biparental care hypothesis: Kleiman (1977) argued that the advantages of monogamy in mammals 

can lead to social monogamy. The hypothesis also implies that both males and females would 

suffer significantly reduced or zero fitness if they did not cooperate in caring for the offspring. However, 

this is not the case for marine decapods, where only the females care for the fertilized eggs 

and where neither parent cares for the larvae. 

Extended mate guarding hypothesis: If males are under selection to guard females for some 

time before, during, and/or after courtship and mating, they may be forced into partner-exclusive 

behavior by some other factor, such as female dispersion (Kleiman 1977; Wickler & Seibt 1981) 

or female-female aggression (Wittenberger & Tilson 1980). In other words, monogamy can result 

from males guarding females over one or multiple reproductive cycles, because the female's synchronous 

receptivity, density, or abundance relative to males renders other male mating strategies 

(pure searching) less successful (Parker 1970; Grafen & Ridley 1983). 

Territorial cooperation hypothesis: The fact that most monogamous species are territorial leads 

to this hypothesis. Territoriality correlates in various ways with social system evolution (Emlen & 

Oring 1977; Hixon 1987), and cooperation in territorial defense can lead to individual advantages 

in social groups or pairs (Brown 1982; Davies & Houston 1984; Fricke 1986; Clifton 1989, 1990; 

Farabaugh et al. 1992). In other words, males and females benefit by sharing a refuge (a territory) 

as heterosexual pairs because, for example, the risk of being evicted from the territory by intruders 

decreases (Wickler & Seibt 1981). 

Recent intensive behavioral studies in various species shrimps have supported the predictions of 

the mate-guarding and/or territorial cooperation hypotheses (e.g., in Hymenocera picta, Wickler & 

Seibt 1981; Alpheus angulatus, Mathews 2002a, b, 2003; and Alpheus heterochelis, Rahman et al. 

2002,2003). 

Another hypothesis about social monogamy (Baeza & Thiel 2007) concerns species symbiotic 

to other organisms (= host). Baeza & Thiel predicted that monogamy evolved when hosts are 

small enough to support few individuals and are relatively rare, and when predation risk away 

from the hosts is high. Under these circumstances, movements among hosts are constrained, and


monopolization of hosts is favored in males and females due to their scarcity and because of the 

host's value in offering protection against predators. Because spatial constraints allow only a few 

adult symbiotic individuals to cohabit in/on the same host, both adult males and females would 

maximize their reproductive success by sharing "their" dwelling with a member of the opposite sex. 

This hypothesis was supported by Baeza's (2008) intensive study on a heterosexual pair of Pontonia 

margarita, a species symbiotic to the pearl oyster. 

However, as mentioned before, most of observations for this mating system are anecdotal, 

and further detailed study is needed to clarify actual conditions of monogamous features of those 

species. 

4.5 Evolution of the eusocial type 

Hypotheses explaining how eusociality has evolved include Trophallaxis Theory (Roubaud 1916), 

Parental Manipulation Theory (Michener & Brothers 1974), Superorganism Theory (Reeve & 

Holldobler 2007), and Inclusive Fitness Theory (Hamilton 1964a, b), of which the last one is most 

widely accepted. According to the Inclusive Fitness Theory, eusociality may evolve more easily in 

species exhibiting haplodiploidy, which facilitates the operation of kin selection. Although eusocial 

mole rats and termites exhibit diploidy, they display high levels of inbreeding by living as a family 

in a single burrow, such that colony members share more than 50% of their genes, and therefore 

the same model is considered to apply to these species and also to eusocial Zuzalpheus shrimps, in 

which all members of a colony share a single sponge. 

4.6 Evolution of the waving display type 

As compared to terrestrial species, courtship in aquatic species may be short and may not involve 

elaborate visual signaling (display) by the males; in aquatic species, chemical or visual cues are 

more important stimuli. In species of several genera of semi-terrestrial (-upper intertidal) decapods 

including Uca and other ocypodid crabs, visual signalling for prolonged periods is common, and 

sounds are often emitted by males to "call" females from their burrows to the surface for mating. 

Salmon •& Atsaides (1968) presented ecological arguments to account for these differences in terms 

of optimal strategy of distance communications in the terrestrial and aquatic environments. Most 

aquatic decapods are nocturnally active and cryptic and live in an acoustically noisy environment, 

and this situation virtually eliminates all but the chemical channel for effective distance communication. 

On the other hand, visual and acoustic signals are effective in terrestrial species and are well 

developed in most terrestrial animals such as insects, birds, mammals, and also ocypodid and other 

terrestrial and semi-terrestrial decapods, probably because of the greater visibility in the terrestrial 

environment. 

Waving displays seen in a variety of semi-terrestrial crabs is a case of convergent evolution 

(Kitaura et al. 2002). Grapsid crabs of the genus Metaplax conduct waving displays like species 

of the ocypodid crab genera Uca, Macrophthalmus, Scopimera, and Dottila (Kitaura et al. 2002). 

Species of Metaplax, unlike other grapsid crabs, which generally live along rocky shores, live in 

mud flats and burrow into the mud like many ocypodids. Salmon & Atsaides (1968) proposed the 

following factors as advantageous for the evolution of visual signaling in semi-terrestrial crabs: the 

substrate, which is flat and relatively free from the vegetational obstructions and other discontinuities; 

diurnal activity of the crabs; and the feeding proximity to their shelters, which leads crabs 

to live in aggregations so that social contacts are frequent. Therefore, it is assumed that habitat 

similarity between Metaplax and ocypodid crabs resulted in convergent evolution of these displays. 

A recent molecular phylogenetic analysis suggested that even the waving display in Uca has 

multiple origins (Sturmbauer et al. 1996). Indo-west Pacific Uca species have simpler reproductive 

social behaviors, are more marine, and were thought to be ancestral to the behaviorally more complex 

and more terrestrial American species. It was also thought that the evolution of more complex

134 Asakura 

social and reproductive behavior was associated with the colonization of the higher intertidal zones. 

However, Sturmbauer et al. (1996) demonstrated that species bearing the set of "derived traits" are 

phylogenetically ancestral, suggesting an alternative evolutionary scenario: the evolution of reproductive 

behavioral complexity in fiddler crabs may have arisen multiple times during their evolution, 

possibly by co-opting of a series of other adaptations for high intertidal living and antipredator 

escape. 

This mating system is quite similar to male-territory-visiting polygamy (Kuwamura 1996) in 

fish, in which many examples are known in intertidal or shallow species; males have a burrow or a 

territory, and, when a mature female approaches a male, the male changes the color of part of his 

body and/or conducts species-specific courtship displays, after which the female enters the burrow 

or territory of the male and spawns (e.g., Miyano et al. 2006). In these fish species, males are brilliantly 

colored, as are male Uca species. 

4.7 Evolution of the visiting type 

A widely recognized tendency among various kinds of animals is that females live in a particular 

place and have a narrow home range, whereas males have a comparatively wider home range 

(Clutton-Brock et al. 1982). This "visiting type" mating system (seen in cryptochirid crabs) probably 

has evolved as one extremity of this tendency, with females living in a very specialized habitat 

(inside coral galls). 

4.8 Evolution of the reproductive swarm type 

Surprisingly, the function of the reproductive swarm in pinnotherid crabs is very similar to that of 

the nuptial flight (mating swarm) in ants (Insecta, Formicidae), and indeed their life history is quite 

similar. In most species of ants, breeding females and males that mature in their mothers' nest have 

wings and, during the breeding season, fly away from their nests and form swarms. Mating occurs 

during this period, and the males die shortly afterward. The surviving females land, and each female 

digs a burrow for the new nest. As eggs are laid in the burrow, stored sperm, obtained during their 

single nuptial flight, is used to fertilize all future eggs produced. 

In the pinnotherids, crabs first grow in their host animals (vs. ants in their initial burrow). Then 

the crabs with swimming setae leave the hosts and swarm (vs. ants with wings fly away from their 

nests and conduct the nuptial flight). Mating occurs during this period (in ants, too), after which the 

female crabs enter the hosts, whereas the males die just after the mating (vs. the female ants make 

burrows of their own, with males dying just after the mating). As in the case of the ants, the female 

crabs reproduce by fertilizing their eggs with sperm from a single mating. 

4.9 Evolution of the neotenous male type 

The miniaturization of male mole crabs in the anomuran genus Emerita coupled with neoteny is 

similar to "dwarf males" (parasitic males, complemental males, miniature males), which are tiny 

males often attached to females. This condition has evolved in various groups of animals, including 

thoracican barnacles (Yamaguchi et al. 2007), acrothoracican barnacles (Kolbasov 2002), the oyster 

Ostreapuelchanas (Castro & Lucas 1987; Pascual 1997), epicaridean isopods (Mizoguchi et al. 

2002), an echiuran Bonellia (Berec et al. 2005), anglerfish (Lophiiformes) (Pietsch 2005), blanket 

octopus (Tremoctopodidae), argonauts (Argonautidae), football octopus (Ocythoidae), and a deeper 

water octopus Haliphron atlanticus (Alloposidae) (Norman et al. 2002). The evolutionary cause for 

these phenomena has not been fully studied. The neoteny of male Emerita is considered to be one 

rather radical evolutionary solution to the problem of keeping the male and female together in the 

harsh and turbulent surf zone environment (Salmon 1983; Subramoniam & Gunamalai 2003).

ACKNOWLEDGEMENTS 


I am deeply grateful to Joel W. Martin (Natural History Museum of Los Angeles County, U.S.A.) 

for giving me the opportunity to present my work at the symposium on decapod phylogenetics at 

the TCS Winter Meeting in San Antonio, Texas. Thanks are also due to the following persons who 

provided me with important literature or aided me in my bibliographical survey: Keiichi Nomura 

(Kushimoto Marine Park, Japan), Tomomi Saito (Port of Nagoya Public Aquarium), Annie Mercier 

(Memorial University of Newfoundland, Canada), Juan P. Carricart-Ganivet (El Colegio de la Frontera 

Sur, Unidad Chetumal, Mexico), Jorge Contreras Gardufio (Entomologia Aplicada, Instituto 

de Ecologfa, Mexico), Ana Maria S. Pires-Vanin (Instituto Oceanografico, Universidade de Sao 

Paulo, Brazil), J. Antonio Baeza (Smithsonian Tropical Research Institute, Republic of Panama), 

Martha Nizinski (NOAA/NMFS Systematics Laboratory, Smithsonian Institution, U.S.A.), Panwen 

Hsueh (National Chung Hsing University, Taiwan), Estela Anahi Delgado (Undecimar, Facultad 

de Ciencias, Uruguay), Michiya Kamio (Georgia State University, U.S.A.), Satoshi Wada 

(Hokkaido University, Japan), Yoichi Yusa (Nara Women's University, Japan), Tomoki Sunobe 

(Tokyo University of Marine Science and Technology), Charles H. J. M. Fransen (Nationaal Natuurhistorisch 

Museum Naturalis, The Netherlands), E. Gaten (University of Leicester, U.K.), Gil 

G. Rosenthal (Texas A&M University, U.S.A.), and Hiromi Watanabe (JAMSTEC, Japan). Special 

thanks are due to Raymond Bauer (University of Louisiana Lafayette) and Joel W. Martin for the 

careful review of an earlier draft of the manuscript. 

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APPENDIX! 


Table 1. Species of the short courtship type, in which females molt before copulation (= soft-female mating 

sensu Hartnoll 1969). 

DENDROBRANCHIATA 

Penaeidae: Marsupenaeus japonicus (1), Melicertus kerathurus (2), Melicertus brasiliensis (3), 

Melicertus paulensis (4), Farfantepenaeus aztecus (5), Fenneropenaeus merguiensis(6),Penaeus 

monodon (7), Penaeus semisulcatus (8), Trachypenaeus similis (9), Xiphopenaeus sp. (10)*, 

Sieyoniidae: Sicyonia dorsalis (11), Sicyonia parri (12), Sicyonia laevigata (13) 

PLEOCYEMATA 

Caridea 

Palaemonidae: Palaemonetes vulgarus (14), Palaemonetes varians (15), Palaemonetes pugio 

(16), Palaemon serratus (17), Palaemon elegans (18), Palaemon squilla (19) 

Alpheidae: Athanus nitescens (20), Alpheus dentipes (21) 

Hippolytidae: Heptacarpus picta (22), Heptacarpus paludicola (23) 

Pandalidae: Pandalus dona (24), Pandalus platyceros (25), Pandalus borealis (26) 

Crangonidae: Crangon crangon (27), Crangon vulgaris (28) 

Astacidea 

Nephropidae: Nephrops norvegicus (29) 

Palinuridea 

Palinuridae: Jasus lalandii (30)* 

Anomura 

Hippidae: Emerita asiatica (31), Emerita analoga (32) 

Diogenidae: Calcinus latens (33), Calcinus seurati (34), Clibanarius tricolor (35), Clibanarius 

; antillensis (36), Clibanarius zebra (37), Paguristes cadenati (38), Paguristes tortugae (39), 

Paguristes anomalus (40), Paguristes hummi (41), Paguristes oculatus (42) 

*Hard-female mating was rarely reported in addition to the soft-female mating. References: (1) Hudinaga 

(1942 as Penaeus japonicus), (2) Heldt (1931 as Penaeus caramote), (3) Brisson (1986), (4) de 

Saint-Brisson (1985), (5)-(6) Aquacop (1977), (7) Primavera (1979), Aquacop (1977), (8) Browdy 

(1989), (9)-(10) Bauer (1991), (11) Bauer (1992, 1996), (12)-(13) Bauer (1991), (14) Burkenroad 

(1947), Bauer (1976), (15) Antheunisse et al. (1968), Jefferies (1968), (16) Berg & Sandifer (1984), 

Bauer & Abdalla (2001), Caskey & Bauer (2005), (17) Nouvel & Nouvel (1937), Forster (1951), 

Bauer (1976), (18) Hoglund (1943), (19) Hoglund (1943), Bauer (1976), (20) Nouvel & Nouvel 

(1937), (21) Volz (1938), (22) Bauer (1976), (23) Bauer (1979), (24) Needier (1931), (25) Hoffman 

(1973), (26) Carlisle (1959), (27) Nouvel (1939), (28) Lloyd & Young (1947), Havinga (1930), 

Bodekke et al. (1991), (29) Farmer (1974), (30) von Bonde (1936), Silberbauer (1971), McKoy 

(1979), (3 1) Menon (1933), Subramoniam (1979), (32) MacGinitie (1938), Efford (1965), (33) Hazlett 

(1972), (34) Hazlett (1989), (35)-(36) Hazlett (1966), (37) Hazlett (1966, 1989), (38)-(42) 

Hazlett (1966).

142 Asakura 

Table 2. Species of the short courtship type, in which females do not molt before copulation (= hard-female 

mating sensu Hartnoll 1969). 

DENDROBRANCHIATA 

Penaeoidea: Litopenaeus vannanmei (1), Litopenaeus setiferus (2), Litopenaeus stylirostris (3), 

Litopenaeus schmitti (4) 

PLEOCYEMATA 

Astacidea 

Astacidae: Pacifastacus trowbridgii (5), Pacifastacus leniusculus (6), Austropotamobius pallipes 

(7), Austropotamobius italicus (8), Austropotamobius torrentium (9), Astacus astacus (10), 

Astacus leptodactylus (11) 

Parastacidae: Cherax quadricarinatus (12) 

Cambaridae: Orconectes nais (13), Orconectes limosus (14), Faxonella clypeata (15), 

Orconectes rusticus (16), Orconectes propinquus (17), Orconectes virilis (18), Orconectes 

inermis inermis (19), Orconectes pellucidus (20), Cambarus blandingi (21), Cambaroides 

japonicus (22), Cambarus immunis (23), Procambarus alleni (24), Procambarus clarkii (25), 

Procambarus hayi (26) 

Palinuridea 

Palinuridae: Panulirus homarus (27)*, Panulirus argus (28)*, Panulirus longipes cygnus (29) 

Anomura 

Diogenidae: Calcinus verilli (30), Calcinus laevimanus (31), Calcinus seurati (32), Calcinus 

elegans (33), Calcinus hazletti (34), Calcinus laurentae (35) 

Coenobitidae: Birgus latro (36), Coenobita perlatus (37), Coenobita clypeatus (38), Coenobita 

compressus (39) 

Brachyura 

Leucosiidae: Philyra scabriuscula (40), Ebalia tuberosa (41) 

Xanthidae: Lophopanopeus bellus (42), Lophopanopeus diegensis (43), Paraxanthias taylori 

(44), Pilumnus hirtellus (45), Xantho incisus (46), Nanopanope sayi (47), Eurypanopeus 

depressus (48), Panopeus herbstii (49) 

Majidae: Microphrs bicornutus (50), Pw

Table 2. continued. 


* Soft-female mating was rarely reported in addition to the hard-female mating. References: (1) 

Yano et al. (1988), Misamore & Browdy (1996), Palacios et al. (2003), (2) Misamore & Browdy 

(1996), (3) Aquacop (1977), (4) Bueno (1990), (5) Mason (1970a, b), (6) Lowery & Holdich (1988), 

Stebbing et al. (2003), (7) Ingle & Thomas (1974), Brewis & Bowler (1985), Carral et al. (1994), 

Villanelli & Gherardi (1998), (8) Galeotti et al. (2007), Rubolini et al. (2006, 2007), (9) Laurent 

(1988), (10) Cukerzis (1988), (11) Koksal (1988), (12) Barki & Karplus (1999), (13) Pippit (1977), 

(14) Schone (1968), Holdich & Black (2007), (15) Smith (1953), (16) Berrill & Arsenault (1982), 

Snedden (1990), Simon & Moore (2007), (17) Tierney & Dunham (1982), (18) Bovbjerg (1953), 

Rubenstein & Hazlett (1974), Tierney & Dunham (1982), (19)-(20) Bechler (1981), (21) Pearse 

(1909), (22) Kawai & Saito (2001), (23) Tack (1941), (24) Bovbjerg (1956), Mason (1970a, b), 

(25) Ameyaw-Akumfi (1981), Corotto et al. (1999), (26) Payne (1972), (27) Berry (1970), Heydon 

(1969), (28) Sutcliffe (1952, 1953), Kaestner (1970), Lipcius et al. (1983), Lipcius & Herrnkind 

(1987), (29) Chittleborough (1976), Sheard (1949), (30)-(35) Hazlett (1972), (36) Helfman 

(1977), (37) Page & Willason (1982), (38) Dunham & Gilchrist (1988), (39) Contreras-Garduno 

et al. (2007), (40) Naidu (1954), (41) Schenibri (1983), (42)-(43) Knudsen (1960, 1964), (44)- 

(46) Bourdon (1962), (47)-(49) Swartz (1976a, b), (50) Hartnoll (1965a), (51) Vernet-Cornubert 

(1958a), (52) Knudsen (1964), (57) Boolootian et al. (1959), Grigg personal communication in 

Hartnoll (1969), Knudsen (1964), (54) Berry & Hartnoll (1970), (55) Arakawa (1964), (56) Warner 

(1967, 1970), (57) Broekhuysen (1941), (58) Hartnoll (1965b), (59)-(60) Brockerhoff & McLay 

(2005a, b)5 (61) Hoestlandt (1948), Peters et al. (1933), (62) Kobayashi & Matsuura (1994), (63) 

Schone & Schone (1963), Warner (1967, 1970), (64) Kramer (1967), Schone & Schone (1963), 

(65) Brockerhoff & McLay (2005a, b, c), (66) Knudsen (1964), (67) Yaldwyn (1966b), Brockerhoff 

(2,002), (68) Knudsen (1964), (69) Brockerhoff & McLay (2005a, b, c), (70) Bovbjerg (1960), Hiatt 

(1948), (71) Brockerhoff & McLay (2005a, b), (72) Vernet-Cornubert (1958b), (73) Fukui (1991, 

1994), "(74) Hartnoll (1969), (75)-(77) Brockerhoff & McLay (2005a, b), (78) Warner (1967 as 

Sesarma ricordt)r(19) Seiple & Salmon (1982), (80)-(81) Hartnoll (1969), (82) von Hagen (1967), 

(83) Hartnoll (1969), (84) Seiple & Salmon (1982 as Sesarma cinereum), (85) Hartnoll (1969 as 

Sesarma angustipes), (86) Hartnoll (1969 as Sesarma curacaoense), (87) Nye (1977), Beer (1959), 

Brockerhoff & McLay (2005a, b), (88) Hicks (1985), (89) Abele et al. (1973), Klassen (1975), Bliss 

et al. (1978), (90) Gifford (1962), Henning (1975), (91) Ameyaw-Akumfi (1987). 

Table 3. Penaeid shrimp species in which a sperm plug has been reported. 

Penaeidae 

Rimapenaeus similis (1) 

Farfantepenaeus aztecus (2) 

Rimapenaeus constrictus (3) 

Marsupenaeus japonicus (4) 

Metapenaeus joyneri (5) 

References: (1) Bauer & Min (1993 as Trachypenaeus similis), (2) Bauer & Min (1993), (3) Costa 

& Fransoso (2004), (4) Fuseya (2006), (5) Miyake (1982).

144 Asakura 

Table 4. Species of the precopulatory guarding type, in which males guard females before copulation. S = 

species in which females molt before copulation. H = species in which females do not molt before copulation. 

V = species in which both types (S and H) have been reported. ? = molting condition has not been reported. 

CARIDEA 

Palaemonidae: Macrobrachium amazonicum [S](l), Macrobrachium rosenbergii [S](2), Macrobrachium 

austoraliense [S](3), Macrobrachium nipponense [S](4), Macrobrachium longipes 

[S](5) 

Rhynchocinetidae: Rhynchocinetes typus [H](6) 

ASTACIDEA 

Homaridae: Homarus americanus [V](7) 

ANOMURA 

Diogenidae: Diogenes pugilator [S](8), Diogenes nitidimanus [V](9), Dardanus punctulatus 

[?](10), Calcinustibicen [Sl](ll) 

Paguridae: Pagurus miamensis [V](12), Pagurus pygmaeus [V](13), Pagurus bonairensis (H](14), 

Pagurus marshi [S](15), Pagurus bernhardus [S](16), Pagurus cuanensis [H](17), Pagurus 

anachoretus [H](18), Pagurus alatus [H](19), Pagurus marshi [S](20), Pagurus nigrofascia 

[S](21), Pagurus lanuginosus [V](22), Pagurus prideauxi [H](23), Pagurus hirsutiuculus 

[S](24), Pagurus maculosus [?](25), Pagurus minutus [V](26), Pagurus filholi [V](27), Pagurus 

gracilipes [?](28), Pagurus middendorffii [H](29), Pagurus nigrivittatus [V](30), Anapagurus 

chiroacanthus [V](31), Anapagurus breriaculeatus [V](32), Pylopagurus sp. sensu Hazlett 

(1975)[H](33) 

Lithodidae: Paralithodes camtschaticus [S](34), Paralithodes brevipes [S](35), Lithodes maja 

[S](36), Lithodes santolla [S](37), Paralomis granulose [S](38), Hapalogaster dentata [S](39) 

BRACHYURA 

Leucosiidae: Philyrd laevis [H](40) 

Majidae: Chionoecetes opilio [S](41), Chionoeceies bairdi [S](42), Macropodia longirostris 

[S](43), Macropodia rostrata [S](44) 

Hymenosomatidae: Halicarcinus sp.'[S](45), Hymenosoma orbiculare [S](46) 

Cancridae: Cancer gracilis [S](47), Cancer irroratus [S](48), Cancer magister [S](49), Cancer 

oregonensis [S](50), Cancer pagurus [S](51), Cancer pro ductus [S](52), Cancer borealis 

[S](53), Cancer antennarius [S](54) 

Cheiragonidae: Telmessus cheiragonus [S](55), Erimacrus isenbeckii [S](56) 

Corystidae: Corystes cassivelaunus [H](57) 

Portunidae: Callinectes sapidus [S](58), Carcinus maenas [S](59), Macropipes holsatus [S](60), 

Ovalipes ocellsatus [S](61), Portunus pelagicus [S](62), Portunus sanguinolentus [S](63), 

Portunus puber [S](64), Portunus trituberculatus [S](65), Scylla serrata [S](66) 

Xanthidae: Menippe mercenaria [S](67)



References: (1) Guest (1979), (2) Bhimachar (1965), Rao (1967), Ra'anan & Sagi (1985), Kuris 

et al. (1987), (3) Ruello et al. (1973), Lee & Felder (1983), (4) Ogawa et al. (1981), Mashiko (1981), 

(5) Shokita (1966), (6) Correa et al. (2000, 2003), Hinojosa &' Thiel (2003), Correa & Thiel (2003a, 

b), Diaz & Thiel (2003), Thiel & Hinojosa (2003), Diaz & Thiel (2004), Thiel & Correa (2004), van 

Son & Thiel (2006), Dennenmoser & Thiel (2007), (7) Herrick (1909), Templeman (1934, 1936), 

McLeese (1970, 1973), Hughes & Matthiessen (1962), Aiken & Waddy (1980), Waddy & Aiken 

(1981), Aiken et al. (2004), (8) Bloch (1935), Hazlett (1968), (9) Asakura (1987), (10) Matthews 

(1956), (11)-(13) Hazlett (1966), (14)-(17) Hazlett (1968), (18) Hazlett (1968), Hazlett (1975), 

(19) Hazlett (1968), (20) Hazlett (1975), (21)-(22) Wada et al. (2007), (23) Hazlett (1968), (24) 

MacGinitie (1935), (25) Imazu & Asakura (2006), (26) Imazu & Asakura (2006), Wada et al. (2007), 

(27) Imafuku (1986), Goshima et al. (1998), Minouchi & Goshima (1998,2000), Wada et al. (2007), 

(28) Imazu & Asakura (2006), (29) Wada et al. (1996, 1999), (30) Wada et al. (2007), (31M32) 

Hazlett (1968), (33) Hazlett (1975), (34) Marukawa (1933), Powell & Nickerson (1965a, b), Gray 

& Powell (1966), Wallace et al. (1949), McMullen (1969), Matsuura & Takeshita (1976), Takeshita 

& Matsuura (1989), (35) Wada et al. (1997, 2000), Sato et al. (2005a, b), (36) Pike & Williamson 

(1959), (37)-(38) Lovrich & Vinuesa (1999), (39) Goshima et al. (1995), (40) Schembri (1983), 

(41) Watson (1972), (42) Paul (1984), Donaldson & Adams (1989), (43)-(44) Hartnoll (1969), 

(45) Lucas personal communication in Hartnoll (1969), (46) Broekhuysen (1955), (47) Knudsen 

(1964), (48) Chidchester (1911), Elner & Elner (1980), Elner & Stasko (1978), Haefner Jr. (1976), 

(49) Bulter (1960), Cleaver (1949), Snow & Nielsen (1966), (50) Knudsen (1964), (51) Edwards 

(1966), (52) Knudsen (1964), (53) Elner et al. (1985), (54) Knudsen (1960), (55) Kamio et al. 

(2000, 2002, 2003), (56) Sasaki & Ueda (1992), (57) Hartnoll (1968), (58) Childchester (1911), 

Churchill (1919), Hay (1905), Gleeson (1980), Ryan (1966), Gleeson et al. (1984), Christofferson 

(1970), Teytaud (1971), Jivoff & Hines (1998), (59) Broekhuysen (1936, 1937), Cheung (1966), 

Childchester (1911), Spalding (1942), Veillet (1945), Williamson (1903), Berrill (1982), Berrill & 

Arsenault (1982), Jensen (1972), (60) Broekhuysen (1936), (61) Childchester (1911), (62) Delsman 

& de Man (1925), Broekhuysen (1936), Fielder & Eales (1972), (63) George (1963), Ryan (1966, 

1967a, b), Christofferson (1970, 1978), (64) Duteutre (1930), (65) Oshima (1938), (66) Hill (1975), 

(67) Binford (1913), Cheung (1968), Savage (1971), Porter (1960), Wilber (1989).

146 Asakura 

Table 5. Duration of guarding time in selected species of decapod crustaceans. 

Species 

ANOMURA 

Lithodidae 

Paralithodes brevipes 

Paralithodes brevipes 

3 males & 3 females 

1 male & 5 females 

Hapalogaster dentata 

BRACHYURA 

Cancridae 

Cancer pagurus 

Conner irroratus 

Carcinus maenas 

1 male & 1 female 

2 or 3 males - 1 female 

Majidae 

Chionoecets bairdi 

Chionoecets opilio 

Cheiragonidae 

Telmessus cheiragonus 

Corystidae 

Corystes cassivelaunus 

Precopulatory 

guarding time 

9-84 hrs 

(mean 38.9+24.9 hrs) 

32.1+44.1 hrs 

15.1+20.1 hrs 

2-3 days 

3-21 days 

4.5 days 

2-16 days 

3-10 days 

1-12 days 

•7-9 days 

11.8 ± 5 SD days 

Up to several days 

Female condition 

when copulating 

Soft 

Soft 

Soft 

Soft 

Soft 

Soft 

Soft 

Soft 

Various 

Soft 

Soft 

Hard 

Postcopulatory 

guarding time 

? 

? 

? 

? 

1-12 days 

5 days 

0-1.5 days 

1-3.5 days 

? 

8 hrs 

4.0 ± 6.6 hrs 

0 

Refere 

References: (1) Wada et al. (1997), (2)-(3) Wada et al. (2000), (4) Goshima et al. (1995), (5) Edwa 

(1966), (6) Elner & Elner (1980), (7)-(8) Berrill & Arsenault (1982), (9) Donaldson & Adams (198 

(10) Watson (1972), (11) Kamio et al. (2003), (12) Hartnoll (1968). 

(1) 

(2) 

(3) 

(4) 

(5) 

(6) 

(7) 

(8) 

(9) 

(10) 

(11) 

(12)


Table 6. Brachyuran crab species, in which a sperm plug has been reported. 

Gancridae 

Cancer magister (1) 

Cancer irroratus (2) 

Cancer pagurus (3) 

Geryonidae 

Geryon fenneri (4) 

Portunidae 

Callinectes sapidus (5) 

Carcinoplax vestita (6) 

Carcinus maenas (7) 

Macropipus holsatus (8) 

Ovalipes ocellsatus (9) 

Portunus sanguinolentus (10) 

Necorapuber (11) 

Liocarcinus depurator (12) 

Cheiragonidae 

Telmessus cheiragonus (13) 

Eriphiidae 

Eriphia smithii (14) 

References: (1) Oh & Hankin (2004), (2) Childchester (1911), (3) Edwards (1966), (4) Hinsch 

(1988), (5) Childchester (1911), Wenner (1989), Johnson & Oito (1981), Jivoff (1997), (6) Doi & 

Watanabe (2006), (7) Broekhuysen (1936, 1937), Spalding (1942), (8) Broekhuysen (1936), (9) 

Childchester (1911), (10) George (1963), (11) Gonzalez-Gurriaran & Freire (1994), Norman & 

Jones (1993), (12) Abello (1989), (13) Kamio et al. (2003), (14) Tomikawa &Watanabe (1990). 

Table 7. Species found in large aggregations called a "pod," "heap," or "mound." 

Species 

ANOMURA 

Lithodidae 

Paralithodes camtschaticus 

Lithodes santolla 

BRACHYURA 

Majidac 

Maya squinado 

Chionoecetesbairdi 

Hym lyratus 

LojKorhynchus grandis 

Libinia emarginata 

Number of crabs in 

each aggregation Reference 

1000 or more 

70 ind-m-2 or more 

22-50,000 or more 

100,000s 

2,000 

100s 

5,000? 

References: (1) Dew (1990), Dew et al. (1992), Powell & Nickelson (1965a, b), Powell et al. (1973), 

Zhou & Siiirley (1997), Stone et al. (1993), (2) Cardenas et al. (2007), (3) Baal (1953), Le Sueur 

(1954), Carlisle (1957), Sampedro & Gonzalez-Gurriaran (2004), (4) Stevens (2003), Stevens et al. 

(1994), (5 ) Stevens et al. (1992), (6) Debelius (1999), Hobday & Rumsey (1999), (7) DeGoursey & 

Auster (1992), Hinsch (1968). 

(1) 

(2) 

(3) 

(4) 

(5) 

(6) 

(7)

148 Asakura 

Table 8. Species of the Pontoniinae reported as "found in pair." Species of shrimps with [host animals in 

brackets] are listed according to the phyla of the host animals (large capitals). 

PORIFERA 

Apopontonia dubia [Spongia sp.](l), Onycocaris amakusensis [Callyspongia elegans](2), 

Onycocaris oligodentata [purplish sponge](3), Onycocaris spinosa [small sponge](4), 

Onycocaridella prima (5)[Mycale sulcata], Onycocaridella monodoa (= Onycocaris monodoa) 

[Pavaesperella hidentata](6), Onycocaridites anornodactylus [sponge] (7), Orthopontonia 

ornatus [Jaspis stellifera]{%), Periclimenaeus stylirostris [sponge](9), Typton dentatus [Reniera 

sp.](10) 

CNIDARIA 

Antipatharia 

Dasycaris zanzibarica [black coral, sea whips] (11) 

Actiniaria 

Periclimenes brevicarpalis [Cryptodendron adhaesivum](12), Periclimenes colemani 

[Asthenosoma intermedium](l3), Periclimenes ornatus [Entacmaea quadricolor, Heteroactis 

malu, Parasicyonis actinostroides](\4) 

Scleractinia 

Anapontonia denticauda [Galaxeafascicularis](\5), Coralliocaris superba [Acropora tubicinaria 

and other 15 spp. of Acropora](\6), Jocaste lucina [Acropora tubicinaria](ll), Jocaste japonica 

[Acropora sp., Acropora humilis, Acropora variabilis, Acropora tubicinaria, Acropora 

nasuta](18), Ischnopontonia lophos [Galaxea fascicularis]{\9), Periclimenes lutescens (20), 

Periclimenes koroensis [Fungia actiniformis](21), Philarius imperialis [Acropora sp., Acropora 

millepora](22), Vir euphyllius [Euphyllia spp.](23), Vir philippinensis [Plerogyra sinuosa](24) 

Scleractinia [in network of fissures on surface of faviid coral] 

Ctenopontonia cyphastreophila [Cyphastrea microphthalma](25) 

Scleractinia [forming galls or bilocular cyst in corals] 

Paratypton siebenrocki [Acropora hyacinthus and other 6 spp. of Acropord\{26) 

MOLLUSCA 

Opistobranchia 

Periclimenes imperator [Hexabranchus marginatus](21) 

Bivalvia 

Anchistus demani [Tridacna maxima](2S), Anchistus miersi [Tridacna squamosa, Tridacna 

maxima](29), Anchistus pectinis [Pecten sp., Pecten albicans], Anchistus custos [Pinna saccata, 

Pinna sp.](31), Chernocaris plaunae [Placunaplacenta](32), Conchodytes biunguiculatus 

[Pinna bicolor](33), Conchodytes meleagrinea [Meleagrina margaritifera](34), Conchodytes 

monodactylus [Pecten sp.,Atrina sp.](35), Conchodytes nipponensis [Pinna sp., Pecten laquetus, 

Atrina japonica](36), Conchodytes tridacnae [Tridacna maxima](31), Bruceonia ardeae 

(= Pontonia ardeae)[Chama pacifica](3%), Pontonia domestica [Atrina seminuda, Atrina rigida, 

Pinna muricata](39), Pontonia mexicana [Pinna cornea, Pinna rigida, Atrina seminuda]{40), 

Ascidonia miserabilis (= Pontonia miserabilis)[Spondylus americanus]{4\), lAscidonia 

miserabilis (as IPontonia miserabilis)[Spondylus americanus](42), Pontonia pinnae [Pinna 

rugosa, Atrina tuberculosa]{43), Pontonia pinnophylax [Pinna rudis, Pinna nobilis]{44), 

Pontonia margarita [Pinctada mazatlanica](45), Platypontonia hyotis [Pycnodonta hyotis]{46)



ECHINODERMATA 

Crinoidea: Comatulida 

Palaemonellapottsi [Comanthina schlegelii, Comanthus briareus, Stephanometra briareus](47), 

Parapontonia nudirostris [Tropiometra afra, Himerometra robustipinna] (48), Periclimenes 

alegrias [Lamprometra palmata, Lamprometra klunzingeri, Stephanometra spicata](49), 

Periclimenes attenuatus [Comaster multifidus](50), Periclimenes novaecaledoninae 

[Lamprometra klunzingeri](51) 

Echinoidea 

Tuleariocaris holthuisi [Astropyge radiata](52), Tuleariocaris zanzibarica [Astropyge radiata, Diadema 

setosum](53) 

CHORDATA 

Ascidiacea: compound ascidian 

Periclimenaeus diplosomatis [Diplosoma lrayneri](54), Periclimenaeus serrula [Leptoclinoides 

incertus](55), Periclimenaeus tridentatus [unidentified &scidmn](56),Ascidoniaflavomaculata 

(= Pontonia flavomaculata)[Ascidia mentula, Ascidia mammillata, Ascidia involuta, Ascidia 

interrupta](51), Odontoma sibogae (= Pontonia sibogae)[Styela whiteleggei, Pyura momus, 

Rhopalaea crassa](5&) 

Ascidiacea: solitary ascidian 

Dasella ansoni [Phallusia depressiuscula](59) 

References: (1) Bruce (1983a), (2)-(4) Fujino & Miyake (1969), (5)-(6) Bruce (1981a), (7) Bruce 

(1987), (8) Bruce (1982), (9) Bruce & Coombes (1995), (10) Bruce & Coombes (1995), Bruce 

(1980a), (11) Gosliner et al. (1996), (12) Bruce & Svoboda (1983), (13) Bruce (1975), (14) Bruce & 

Svoboda (1983), Omori et al. (1994), (15) Bruce (1967), (16)-(17) Bruce (1980b), (18) Bruce (1974, 

1980b, 1981c), (19) Bruce (1980b, 1981c), Bruce & Coombes (1995), (20) Bruce (1981c), Bruce 

& Coombes (1995), (21) Bruce & Svobboda (1984), (22) Bruce & Coombes (1995), (23) Martin 

(2007), (24) Bruce & Svoboda (1984), (25) Bruce (1979), (26) Bruce (1980a, b), (27) Bruce (1972a, 

1976a), Bruce & Svoboda (1983), Strack (1993), (28) Bruce (1972a), (29) Bruce (1972a), Debelius 

(1999), (30) Bruce (1972a), Fujino & Miyake (1967), (31) Bruce (1972a, 1989), Hipeau-Jacquotte 

(1973), (32) Bruce (1972a), (33) Bruce (1972a), Hipeau-Jacquotte (1973), (34) Bruce (1973), (35)- 

(36) Bruce (1972a), (37) Bruce (1974), (38) Bruce (1981b), Fransen (2002), (39) Bruce (1972a), 

Courtney & Couch (1981), Fransen (2002), (40) Bruce (1972a), Criales (1984), Fransen (2002), 

(41) Fransen (2002), (42) Criales (1984), (43) Bruce (1972a), (44) Debelius (1999), Richardson et 

al. (1997), (45) Baeza (2008), (46) Hipeau-Jacquotte (1971), (47) Bruce & Coombes (1995), Bruce 

(1989), (4.8) Bruce (1992), (49) Bruce (1986), Bruce & Coombes (1995), (50) Bruce (1992), (51) 

Bruce & Coombes (1995), (52)-(53) Bruce (1967), (54) Bruce (1980b), (55) Bruce & Coombes 

(1995), (56) Bruce & Coombes (1995), (57) Monniot (1965), Millar (1971), Fransen (2002), (58) 

Bruce (1972b), Fransen (2002), (59) Bruce & Coombes (1995).

150 Asakura 

Table 9. Species of the Alpheidae reported as "found in pair." Species of shrimps with [host animals in brackets] 

are listed according to the phyla of host animals (large captals) with higher taxa or habitat when known. 

PORIFERA 

Synalpheus bituberculatus [sponge] (1), Synalpheus hastilicrassus [sponge] (2), Synalpheus 

jedanensis [sponge](3), Synalpheus streptodactylus [sponge](4), Synalpheus theano [sponge](5), 

Synalpheus fossor [sponge](6), Synalpheus harpagatrus [sponge] (7), Synalpheus nilandensis 

[sponge](8), Synalpheus tumidomanus [sponge] (9), Zuzalpheus androsi [Hyattella 

intestinalis](lO), Synalpheus couitere [sponge](l 1), Zuzalpheus bousfield [Hymeniacidon 

spp.](12), Zuzalpheus carpenteri [Aeglas spp.](13), Zuzalpheus goodei [Xestospongia 

wiedenmayeri, Pachypellina podatypa](l4), Zuzalpheus paraneptunus [Hyattella intestinalis, 

Oceanapia sp.](15), Zuzalpheus ruetzleri [Hymeniacidon cf. caerulea](16), Zuzalpheus 

sanctithomae [Hymeniacidon caerulea etc.](17), Alpheus parvirostris [sponge](18), Alpheus 

alcyone [sponge](19), Alpheus aff. eulimene*[sponge](20), Alpheusparalcyone [sponge](21), 

Alpheus spongiarum [sponge] (22) 

CNIDARIA 

Scyphozoa: Coronatae 

Synalpheus modestus (23), Synalpheus aff. modestus sensu Nomura & Asakura (1998) 

[Stephanoscyphus racemosus](24) 

Anthozoa: Gorgonacea 

Synalpheus iphinoe [Solenocaulon sp.](25), Synalpheus trispinosus [gorgonacean](26) 

Anthozoa: Alcyonacea 

Synalpheus neomeris [Dendronephthya](27) 

Anthozoa: Actiniaria 

Alpheus armatus [Bartholomea annulata](2S), Alpheus immaculatus [Bartholomea annulata](29), 

Alpheus polystuctus [Bartholomea annulata](30), Alpheus roquensis [Heteractis lucida](31) 

Anthozoa: Scleractinia 

Alpheus lottini [reef coral, Pocillopora](32), Alpheus ventrosus (33), Synalpheus charon 

[Pocillopora, reef coral](34), Synalpheus scaphoceris [Madracis decactis](35), Racilius 

compressus [Galaxea fascicularis](36) 

Anthozoa: Scleractinia (in fissures on massive coral) 

Alpheus deuteropus [Asteropora, Porites, Acropora, Montipora, Pavona](37) 

Anthozoa: Scleractinia (coral borer, in dead coral head) 

Alpheus saxidomus (38), Alpheus simus (39), Alpheus schmitti (40), Alpheus idiocheles (41), 

Alpheus colluminaus (42) 

ANNELIDA 

Polychaeta 

Alpheus sulcatus [Eurythoe complanata](43) 

CRUSTACEA 

Shell used by hermit crab 

Aretopsis amabilis [Dardanus sanguinolentus, Dardanus megistos, Dardanus guttatus, Dardanus 

lagopodes, Clibanarius eurysternus, Calcinus latens](44), Aretopsis manazuruensis [Aniculus 

miyakei](45) 

In burrow of thalassinidean shrimps 

Betaeus longidactylus [Upogebia pugettensis](46), Betaeus harrimani [Upogebiapugettensis](47), 

Betaeus ensenadensis [Upogebia pugettensis] (48) 

In burrow of mantis shrimp 

Athanas squillophilus [Oratosquilla oratoria](49)



ECHINODERMATA 

Crinoidea: Comatulida 

Synalpheus carinatus [crinoids](50), Synalpheus comatularum [Comanthus timorensis](51), 

Synalpheus demani [criniod](52), Synalpheus stimpsoni [Comaster multibrachiatus, Comaster 

multifidus, Comaster gracilis, Comaster alternans](53), Synalpheus odontophorus [crinoid](54) 

Echinoidea 

Athanas indicus [Echinometra mathaei](55) 

ECHIURA 

Athanopsis rubricinctuta [Ochetostoma erythrogrammon](56), Betaeus longidactylus [Urechis 

sp.](57) 

"PISCES" [in burrow of goby fish] 

Alpheus bellulus [Tomiyamichthys spp, Amblyeleotris spp.](58), Alpheus purpurilenticularis 

[Amblyeleotris steinitzi], (59) Alpheus rapacida [Myersina spp., Vanderhorstia spp., Mahidoria 

spp.], (60) Alpheus rapax [Crypto centrus spp.](61) 

ALGAE TUBE 

Alpheus frontalis [tube of filamentous blue-green algae such as Microcoelus spp.](62), Alpheus 

bucephalus [tube of pure algae or algae with sponges and other material](63), Alpheus brevipes 

[tube of red filamentous alga](64), Alpheus clypeatus [tube of red filamentous alga 

Acrochaetium](65), Alpheus pachychirus [tube of algae](66) 

FREE LIVING [crack of rock, under rubble, around large algae, burrow in mudflat] 

Alpheopsis chilensis (67), Alpheus normanni (68), Alpheus euphrosyne richardsoni (69), Alpheus 

strenuus cremnus (70), Alpheus diadema (71), Alpheus architectus (72), Alpheus amirantei (73), 

Alpheus bisincisus (74), Alpheus brevicristatus (75) (might be commensal with goby?), Alpheus 

edwardsii (76), Alpheus aff. gracilipes* (77), Alpheus heeia (78), Alpheus'aff. heeia*(19), 

Alpheus aff. leviuscuius sp. 1*(80), Alpheus aff. leviusculus sp. 2* ; (81), Alpheus lobidens (82), 

Alpheus aff. lobidens sp. 1*(83), Alpheus aff. lobidens sp. 2*(84), Alpheus aff. lobidens sp. 

3*(85), Alpheus malleodigitus (86), Alpheus miersi (SI), Alpheus obesomanus (88), Alpheus 

pacificus (89), Alpheus aff. pacificus (90), Alpheus paradentipes (91), Alpheus parvirostris (92), 

Alpheus polyxo (93), Alpheus serenei (94), Alpheus suluensis (95), Alpheus tenuipes (96), 

Alpheus angulatus (97), Alpheus armillatus (98), Alpheus heterochaelis (99), Alpheus floridanus 

(100), Alpheus inca (101), Metalpheusparagracilis (102)

152 Asakura 


*sensu Nomura & Asakura (1998). References: (1) Banner & Banner (1975), Nomura & Asakura 

(1998), (2)-(5) Nomura & Asakura (1998), (6) Didderen et al. (2006), (7) Banner & Banner (1975), 

(8)-(9) Nomura & Asakura (1998), (10) Rios & Duffy (2007), (11) Nomura & Asakura (1998), (12) 

Rios & Duffy (2007), (13) Macdonald III et al. (2006), Rios & Duffy (2007), (14)-(17) Rios & Duffy 

(2007), (18) Banner & Banner (1982), (19)-(27) Nomura & Asakura (1998), (28) Knowlton (1980), 

Knowlton & Keller (1982, 1983, 1985), Criales (1984), (29)-(31) Knowlton (1980), Knowlton & 

Keller (1982, 1983, 1985), (32) Vannini (1985), Nomura & Asakura (1998), Abele & Patton (1976), 

Tsuchiya & Yonaha (1992), (33) Patton (1966), (34) Patton (1966), Nomura & Asakura (1998), (35) 

Dardeau (1984, 1986), (36) Bruce (1972c), (37) Banner & Banner (1983), (38) Fischer & Meyer 

(1985), Fischer (1980), (39)-(40) Werding (1990), (41) Kropp (1987), Nomura & Asakura (1998), 

(42) Banner & Banner (1982), Nomura & Asakura (1998), (43) Banner & Banner (1982), (44) 

Bruce (1969), Banner & Banner (1973), Kamezaki & Kamezaki (1986), (45) Suzuki (1971), (46)- 

(48) MacGinitie (1937), (49) Hayashi (2002), (50) Bruce (1989), (51) Banner & Banner (1975), 

(52) Bruce (1989), Nomura & Asakura (1998), (53) Nomura & Asakura (1998), Van den Spiegel et 

al. (1998), (54) Nomura & Asakura (1998), (55) Gherardi (1991), (56) Anker et al. (2005), Berggren 

(1991), (57) MacGinitie (1935), (58) Miya & Miyake (1969), Nomura & Asakura (1998), Nomura 

(2003), (59) Macnae & Kalk (1962), Karplus (1979), Nomura (2003), (61) Macnae & Kalk (1962), 

Nomura (2003), (62) Fishelson (1966), Banner & Banner (1982), (63) Banner & Banner (1982), 

Nomura & Asakura (1998), (64)-(65) Banner & Banner (1982), (66) Cowles (1913), Banner & 

Banner (1982), (67) Boltana & Thiel (2001), (68) Nolan & Salmon (1970), (69)-(70) Banner & 

Banner (1982), (71)-(75) Nomura & Asakura (1998), (76) Nomura & Asakura (1998), Jeng (1994), 

(77)-(96) Nomura & Asakura (1998), (97) Mathews (2002a, b, 2003, 2006, 2007), Mathews et 

al. (2002), (98) Mathews et al. (2002), (99) Nolan & Salmon (1970), Schein (1975), Obermeier 

& Schmitz (2003a, b), Rahman et al. (2001, 2002, 2003, 2005), Schmitz & Herberholz (1998), 

Dworschak & Ott (1993), (100) Dworschak & Ott (1993), (101) Boltana & Thiel (2001), (102) 

Nomura & Asakura (1998).


Table 10. Species of shrimps other than Pontoniinae and Alpheidae reported as "found in pair." Species of 

shrimps with [host animals in brackets] are listed according to the phyla of host animals (large capitals) with 

higher taxa or habitat when known. 

SPONGICOLIDAE 

PORIFERA 

Spongicola japonica [Euplectella oweni](l), Spongicola venusta [Euplectella aspergillum](2), 

Spongicola levigata [Euplectella oweni1](3), Spongiocaris semiteres [hexactinellid sponge], (4) 

Spongicoloides iheyaensis [Euplectellidae & Hyalonematidae](5), Globospongicola spinulatus 

[hexactinellid sponge Semperella sp.](6) 

FREE LIVING 

Microprosthema validum (7) 

STENOPODIDAE 

FREE LIVING 

Stenopus hispidus ($), Stenopus scutellatus (9), Stenopus tenuirostris (10), Stenopus 

zanzibaricus (11) 

HIPPOLIYTIDAE 

FREE LIVING 

Lysmata debelius (12), Lysmata grabhami (13) 

CNIDARIA 

Actiniaria, Scleractinia 

Thor amboinensis'(14) 

GNATHOPHYLLIDAE 

ECHINODERMATA 

Holothuroidea 

Pycnocaris chagoae [Holothuria cinerascens](15) 

Asteroidea 

Hymenocera picta [prey on sea star](16) 

References: (1) Saito et al. (2001), (2) Miyake (1982), Hayashi & Ogawa (1987), (3) Hayashi & 

Ogawa (1987), (4) Bruce & Baba (1973), (5) Saito et al. (2006), (6) Komai & Saito (2006), (7) 

Davie (2002), (8) Johnson (1969,1977), Castro & Jory (1983), Zhang et al. (1998), Yaldwyn (1964, 

1966a), (9) Debelius (1999), (10) Bruce (1976b), (11) Gosliner et al. (1996), (12) Rufino & Jones 

(2001), Gosliner et al. (1996), (13) Wirtz (1997), Debelius (1999), (14) Stanton (1977), (15) Bruce 

(1983b), (16) Seibt & Wickler (1972, 1979, 1981), Wickler & Seibt (1970, 1972, 1981), Seibt 

(1973a, b, 1974,1980), Wasserthal & Seibt (1976), Wickler (1973), Kraul & Nelson (1986), Fiedler 

(2002).

154 Asakura 

Table 11. Species of Thalassinidea and Anomura reported as "found in pair." Species with [host animals or 

habitat in brackets] are listed according to the phyla of host animals (in capitals) with higher taxa or habitat 

where known. 

THALASSINIDEA 

Axiidae 

FREE LIVING 

Axiopsis serratifrons [in burrow in sediments with a higher content of coral rubble](l) 

Laomediidae 

FREE LIVING 

Axianassa australis [in burrow in mud flat](2) 

Callianassidae 

"PISCES" 

Neotrypaea affinis [burrow of blind goby Typhlogobius californiensis](3) 

FREE LIVING 

Neotrypaea gigas [burrow in mud] (4) 

Upogebiidae 

PORIFERA 

Upogebia synagelas [Agelas sceptrum](5) 

CNIDARIA: Scleractinia 

Pomatogebia rugosa [inside live colony of Pontes lobata](6), Pomatogebia operculata [inside live 

coral colony](7), Upogebia corallifora [inside dead coral colony](8) 

FREE LIVING 

Upogebia pugettensis [U- or Y-shaped burrow in mudflat](9), Upogebia affinis [burrow in mud](10) 

ANOMURA 

Porcellanidae 

CNIDARIA 

Gorgonacea 

Aliaporcellana telestophila [Solenocaulon](11) 

Pennatulacea 

Porcellanella haigae [Cavernularia sp.](12) 

Actiniaria 

Neopetrolisthes oshimai [Soichactis spp.](13), Neopetrolisthes maculatus [Stychodactyla](l4), 

Neopetrolisthes alobatus, Neopetrolisthes spinatus [Heteroactis malu](\5) 

ANNELIDA 

, Polychaeta [in tube of large polychaete species] 

Poly onyx macroheles [Chaetopterus variopedatus](\6), Poly onyx quadriungulatus [Chaetopterus 

variopedatus](ll), Polyonyx transversus [Chaetopterus sp.](18), Polyonyx vermicola 

[Sasekumaria selangora](19), Polyonyx bella [Chaetopterus variopedatus](20), Polyonyx gibbesi 

[Chaetopterus variopedatus](2l), Polyonyx utinomii [Chaetopterus sp.](22), Heteropolyonyx 

biforma [Chaetopterus sp.](23), Polyonyx biunguiculatus [Chaetopterus sp.](24) 

CRUSTACEA [in shell being used by hermit crab] 

Porcellana cancrisocialis [Petrochirus californiensis, Dardanus sinistripes, Aniculus elegans, 

Paguristes digueti](25), Porcellana paguriconviva [Petrochirus calif orniensis, Dardanus 

sinistripes, Aniculus elegans, Paguristes digueti](26) 

ECHINODERMATA 

Echinoidea 

Clastotoechus vanderhorsti [Echinometra lucunter](27), Clastotoechus vanderhorsti [Echinometra 

lucunter](2%) 

Asteroida 

Minyocerus angustus [Luidia, Astropecten, Tethyaster](29)



FREE LIVING 

Pachycheles rudis [underside of stone, basal portion of large algae](30) 

Galatheidae 

ECHINODERMATA 

Crinoidea 

Galathea inflata [Comanthus parvicirrus, Comaster schlehelii](31) 

References: (1) Dworschak & Ott (1993), (2) Coelho & Rodrigues (1999), Coelho (2001), (3)-(4) 

Meinkoth (1981), (5) Williams (1987), (6) Fonseca & Cortes (1998), (7) Kleeman (1984), Williams 

& Ngoc-Ho (1990), Coelho & Rodrigues (1999), Coelho (2001), (8) Williams & Scott (1989), (9) 

Jensen (1995), (10) Meinkoth (1981), (11) Ng & Goh (1996), (12) Nakasone & Miyake (1972), 

(13) Seibt & Wickler (1971), (14) Debelius (1984), (15) Osawa & Fujita (2001), (16) Gray (1961), 

(17) Kudenov & Haig (1974), (18) McNeill & Ward (1930), (19) Ng & Sasekumar (1993), (20) 

Hsueh & Huang (1998), (21) Rickner (1975), Williams (1984), Grove & Woodin (1996), (22)-(23) 

Osawa (2001), (24) Macnae & Kalk (1962), (25) Glassell (1936), Parente & Hendrickx (2000), 

Williams & McDermott (2004), (26) Parente & Hendrickx (2000), Williams & McDermott (2004), 

(27) Werding (1983), (28) Werding (1983), Schoppe (1991), (29) Werding (1983), Gore & Shoup 

(1968), (30) Meinkoth (1981), (31) Fujita & Baba (1999). 

Table 12. Species of brachyuran crabs reported as "found in pair." Species of crabs with [host animals in 

brackets] are listed within family or superfamily according to the phyla of host animals (in capitals) with 

higher taxa or habitat where known. 

XANTHIDAE 

CNIDARIA: Scleractinia 

Cymo andreossyi \Pocillopora\{\) 

TRAPEZHDAE 

CNIDARIA 

Scleraetinia: Pocillopora 

Trapezia areolata (2), Trapezia corallina (3), Trapezia cymodoce (4), Trapezia dentata (5), 

Trapezia digitalis (6), Trapezia ferruginea (7), Trapezia flavomaculata (8), Trapezia guttata (10), 

Trapezia intermedia (11), Trapezia rufopunctata (12), Trapezia tigrina (13), Trapezia wardi (14) 

Antipatharia 

Quadrella maculosa [Antipathes] (15), Quadrella spp. [Cirrhipathes abies, Antipathes spp.}(16), 

Quadrella reticulata [Antipathes sp.](17) 

TETRALIIDAE 

CNIDARIA 

Scleractinia: Acropora 

Tetraliafulva (18), Tetralia nigrolineata (19), Tetralia rubridactyla (20) 

CARPILHDAE 

FREE LIVING 

Carpilius corallinus (21)

156 Asakura 


PINNOTHERIDAE 

ANNELIDA 

Polychaeta [in tube of large polychaetes] 

Pinnixa tubicola [terebellids and chaetopterids, Eupolymnia heterobranchia, Amphitrite sp., Eupolymnia 

heterobranchia, Neoamphitrite rohusta, Thelepus crispus, Chaetopterus variopedatus](22), 

Pinnixa chaetopterana [Chaetopterida spp. Chaetopterus variopedatus, Amphitrite ornata](23), 

Pinnixa transversalis [Chaetopterus variopedatus](24) 

MOLLUSCA 

Bivalvia 

Pinnixa faba [Tresus capax, Tresus nuttalli](25), Pinnixa littoralis [Tresus capax](26) 

Gastropoda [inside mantle cavity] 

Orthotheres turboe [Turbo sp.](27), Orthotheres haliotidis [Haliotis asinina, Haliotis 

squamatd\{2%) 

SIPUNCULA & ECHIURA 

Mortensenella forceps [Ochetostoma erythrogrammon](29) 

ECHINODERMATA 

Echinoidea 

Dissodactylus mellitae [Mellita quinguiesperforata, Echinarachnius parrna, Encope 

michelini](30), Dissodactylus crinitichelis [Mellita sexiesperforata](3l) 

Holothuroidea 

Holotheres halingi (= Pinnotheres halingi) [Holothuria scarba](32), Holotheres semperi (= Pinnotheres 

semperi)[Holothuriafursocinerea, Holothuria scabra\{33) 

BURROWS OF OTHER ANIMALS 

Scleroplax granulata [burrow of echiuroid Urechis caupo, mud shrimps Neotrypaea californiensis, 

Neotrypaea gigas, Upogebia pugettensis, Upogebia macginiteorum](34) 

GRAPSOIDEA 

"REPTILIA": Testudines 

Planes minutus [loggerhead sea turtle Caretta caretta, inanimate flotsam](35) 

ECHINODERMATA 

Echinoidea 

Percnon gibbesi [Diadema antillarum](36) 

References: (1) Castro (1976), Guinot (1978), Miyake (1983), (2) Miyake (1983), Tsuchiya & Yonaha 

(1992), Tsuchiya & Taira (1999), (3) Patton (1966), Miyake (1983), Huber (1985), Gotelli et al. 

(1985), Castro (1996), (4) Patton (1966), Tsuchiya & Yonaha (1992), Tsuchiya & Taira (1999), (5) 

Patton (1966), Huber (1985), (6) Patton (1966), Preston (1973), Huber (1985,1987), Huber & Coles 

(1986), Tsuchiya & Taira (1999), (7) Patton (1966), Preston (1973), Abele & Patton (1976), Finney 

& Abele (1981), Miyake (1983), Adams et al. (1985), Huber & Coles (1986), Castro (1978, 1996), 

Tsuchiya & Taira (1999), (8) Patton (1966), Preston (1973), Miyake (1983), (9) Gotelli et al. (1985), 

Castro (1996), (10) Miyake (1983), Tsuchiya & Yonaha (1992), Tsuchiya & Taira (1999), (11) Preston 

(1973), Huber & Coles (1986), Huber (1987), (12)-(13) Huber (1985), (14) Preston (1973), 

Miyake (1983), Huber & Coles (1986), (15) Shih & Mok (1996), (16) Tazioli et al. (2007), (17) 

Castro (1999), (18) Vytopil & Willis (2001), (19)-(20) Sin (1999), (21) Laughlin (1982), (22) Hart 

(1982), Wells (1928), Garth & Abbott (1980), Zmarzly (1992), (23) Gray (1961), Grove & Woodin 

(1996), Grove et al. (2000), McDermott (2005), (24) Baeza (1999), (25) Pearce (1965, 1966a), Hart 

(1982), Zmarzly (1992), (26) Pearce (1966a), Zmarzly (1992), (27) Sakai (1969), (28) Geiger & 

Martin (1999), (29) Anker et al. (2005), (30) Bell & Stancyk (1983), Bell (1984), George & Boone 

(2003), (31) Telford (1978), (32) Hamel et al. (1999), (33) Ng & Manning (2003), (34) Anker et al. 

(2005), Campos (2006), (35) Dellinger et al. (1997), Frick et al. (2000, 2004, 2006), Carranza et al. 

(2003), (36) Hayes et al. (1998).


Table 13. Eusocial species. All species found inhabiting cavity of sponge. 

Alpheidae 

Zuzalpheus rathbunae [sponge] (1) 

Zuzalpheus elizabethae.(= Synalpheus "rathbunae A")[Lissodendoryx] (2) 

Zuzalpheus "paraneptunus small" [sponge] (3) 

Zuzalpheus regalis [Xestospongia etc.] (4) 

Zuzalpheus filidigitus [Xestospongia etc.] (5) 

Zuzalpheus chacei [Aeglas, Hyattella etc.] (6) 

Zuzalpheus elizabethae [Lissodendoryx etc.] (7) 

Synalpheus neptunus neptunus [sponge] (8) 

References: (1) Duffy (2003), (2) Duffy (1996c, 2003), Morrison et al. (2004), (3) Duffy et al. 

(2000), Duffy (2003), (4) Duffy (1996a, b), Duffy et al. (2002), Rios & Duffy (2007), (5) Duffy 

(1996c), Duffy & Macdonald (1999), Rios & Duffy (2007), (6) Chace (1972), Duffy (1998), Rios 

& Duffy (2007),(7) Duffy (1996c), Morrison et al. (2004), Rios & Duffy (2007),(8) Didderen et al. 

(2006). 

Table 14. Species found in small groups. Species with [host animals] are listed, according to the phyla of host 

animals (large capitals) with higher taxa or habitat. One group consists of fewer than 20 individuals on a single 

host (species, host, number of individuals found, and reference). 

CARIDEA 

CNIDARIA 

Scyphozoa 

Periclimenes holthuisi [Cassiopei] 

Actiniaria 

Periclimenes holthuisi [sea anemone] 

Periclimenes tenuipes [Megalactis, Cryptodendron] 

Periclimenes longicarpus [Entacmaea] 

Periclimenes anthophilus [Condylactis gigantea] 

Scleractinia 

Thor marguitae[Porites andrewsi] 

Jocaste japonica [Acropora divaricata] 

Periclimenes holthuisi [corals] 

Periclimenes pederosoni [Antipathe] 

Anapontonia denticauda [Galaxea] 

ECHINODERMATA 

Echinoidea 

Gnathophylloides mineri [Tripneustes ventricosus] 

GALATHEOIDEA 

CNIDARIA 

Scleractinia 

Lissoporcellana spinuligera [Solenocaulon] 

CRUSTACEA: shell used by hermit crab 

Porcellana sdyana [Dardanus, Petrochirus, Paguristes] 

Max. 8 (various sizes and sexes)(l) 

Several individuals (2) 

Max. 6 (various sizes and sexes)(3) 


Up to 9 (5) 

10 (2 cf, 5 ov. $, 2 non-ov. 9, 1 juv.)(6) 

15 (5 d\ 6 ov. $, 3 non-ov. 9, 1 juv.)(7) 

Several individuals (8) 

7 (2 cf, 3 ov. 9, 2 non-ov. $)(9) 

5(la\l9,3juv.)(10) 

Up to 13, with females greatly 

outnumbering males (11) 

7(ld\3ov. 9, 3juv.)(12) 

Max. 11 (several cf, several ov. 9)(13)

158 Asakura 


BRACHYURA 

MOLLUSCA 

Bivalvia 

Pinnixafaba [Tresus] More than 3(1 d\ 1 9, fewjuv.)(14) 

References: (1) Bruce & Svoboda (1983), (2) Coleman (1991), (3)-(4) Bruce & Svoboda (1983), 

(5) Nizinski (1989), (6) Bruce (1978), (7) Bruce (1981b), (8) Coleman (1991), (9) Spotte (1996), 

(10) Bruce (1967), (11) Patton et al. (1985), (12) Ng & Goh (1996), (13) Gore (1970), (14) Haig & 

Abbott (1980). 

Table 15. Species found in large groups. Species with [host animals] are listed, according to the phyla of host 

animals (large capitals) with higher taxa or habitat. One group consists of more than 20 individuals on a single 

host. 

CARIDEA 

PORIFERA 

Synalpheus dorae [Reiniere] 

Synalpheus streptodactylus [sponge] 

Synalpheus crosnieri [sponge] 

Synalpheus paradoxus [sponge] 

Zuzalpheus brooksi [sponge] 

Zuzalpheus idios [Hymeniacidon etc.] 

Zuzalpheus pectiniger [Spheciospongia] 

CNIDARIA 

. Scyphozoa 

Latreutes anoplonyx [Nemopilema nomurai] 

Scleractinia 

Coralliocaris macrophthalma [Acropora hyacinthus] 

Fennera chacei [Pocillopora] 

Periclimenes toloensis [Lytocarpus philippinensis] 

ECHINODERMATA 

Periclimenes affinis [Heterometra magnipinna] 

Periclimenes meyeri [Nemaster grandis] 

136 (all cf)(l) 

105 (68 cT, 37 ov. 9, several non- ov. $)(2) 

147 (144 d\ 3 9)(3) 

112 (110 d\ 2 $), 132 (130 cT, 2 ?)(4) 

10s to 1000s (5) 

Several 10s (including many ov. 9 & juv.)(6) 

Few 100s (7) 

More than 100 (8) 

24 (including 16 ?)(9) 

Max. 49 (all adults) (10) 

110 (including 43 ov.9)( 11) 

64 (including 16 ov. 9)(12) 


References: (1) Bruce (1988), (2) Banner & Banner (1975, 1982), (3) Banner & Banner (1983), (4) 

Banner & Banner (1982), (5)-(7) Rios & Duffy (2007), (8) Hayashi et al. (2003), (9) Bruce (1977), 

(10) Gotelli et al. (1985), (11)-(12) Bruce & Coombes (1995), (13) Criales (1984).


Table 16. Selected species of pinnotherid crabs (and their hosts) in which life history has been studied. 

MOLLUSCA 

Bivalvia 

Fabia subquadrata [Modiohis modiolus] (1) 

Tumidotheres maculatus (= Pinnotheres maculatus) [Mytilus edulis, Argopecten irradians etc.] (2) 

Pinnotheres ostreum [Crass ostrea virginica, Mytilus edulis] (3) 

Pinnotheres pisum [Mytilus edulis etc.] (4) 

Pinnotheres taichungae [Laternula marilina] (5) 

Pinnotheres bidentatus [Laternula marilina] (6) 

ANNELIDA: Polychaeta 

Tritodynamia horvathi [in tube of Loimia verrucosa] (7) 

References: (1) Pearce (1962, 1966b), (2) Pearce (1964), Williams (1984), (3) Christensen & Mc- 

Dermott (1958), (4) Atkins (1926), Christensen (1958), Hartnoll (1972), Williams (1984), (5) 

Hsueh (2003), (6) Hsueh (2001a, b), (7) Matsuo (1998, 1999), Takahashi et al. (1999). 

Table 17. Species in which neotenous males have been reported. 

Hippidae 

Emerita brasiliensis 

Emerita asiatica 

Emerita emeritus 

Emerita holthuisi 

Emerita talpoida 

Emerita rathbunae 

(i) 

(2) 

(3) 

(4) 

• (5) 

(6) 

References: (1) Delgado & Defeo (2006, 2008), (2) Subramoniam (1981), (3)-(4) Subramoniam & 

Gunamalai (2003), (5)-(6) Efford (1967).

160 Asakura 

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