Sociality in <Emphasis Type="Italic">Euglossa</Emphasis> (<Emphasis Type="Italic">Euglossa</Emphasis>) <Emphasis Type="Italic">viridissima</Emphasis> Friese (Hymenoptera,Apidae, Euglossini)

Euglossa viridissima is an orchid bee that forms both solitary and multiple female nests, making it a suitable species for the study of factors leading to diverse degrees of sociality in Euglossines. We conducted observations in eight reused nests (where a first generation of bees had been produced) kept in artificial boxes from the Yucatan Peninsula, Mexico. Five nests were reused (reactivated) by a single female (SFN), two nests reused by a mother and one daughter (MFN1) and one nest reused by the mother and two daughters (MFN2). No single nest was reactivated by unrelated females. The number of foraging trips, their duration and the duration of cell provisioning was not different between SFN and MFN. The overall production of cells per female was not different either between both types of nest. However, in MFN although all females did lay eggs, there was a reproductive skew in favor of the mother (95 and 45% of the brood produced in MFN1 and MFN2 respectively). She showed reproductive control of her daughters through oophagy and displaying threatening behavior when the daughters tried to open a cell where she had laid an egg. Brood losses to parasites (Anthrax sp. (Bom-byliidae) and Hoplostelis bivittata (Megachilidae)) were only found in SFN which possibly reflects and advantage of MFN in this respect. Our results coupled with other studies in Euglossa, reveal that a wide range of social behaviors occur in this genus, from solitary and communal to primitive reproductive division of labor. Multiple factors involving different levels of pressure imposed by food availability and parasites may favor such a diverse range of nesting behaviors. Interestingly, female associations in E. viridissima seem a result of kin selection that is enforced by coercion from mother females on their daughters. More studies are needed to shed light upon the social organization of Euglossa and other Euglossines and on their phylogenetic relationships in order to trace the origins of eusociality in Apidae. Received 12 February 2008; revised 25 June 2008; accepted 17 July 2008.     

Nesting biology and social structure of <Emphasis Type="Italic">Euglossa (Euglossa) townsendi</Emphasis> Cockerell (Hymenoptera,Apidae, Euglossini)

Summary. The nesting behavior of the euglossine bee Euglossa townsendi was studied on the campus of the University of São Paulo-Ribeirão Preto, SP, Brazil, from January 1994 to December 1999, using artificial nesting substrate and observation boxes. Twenty-one nests were founded during the study period, with the highest frequencies of nesting occurring during the hot, wet season. Each nest was founded by a single female and, after the completion of the last cell, she spent most of her time in the nest. The males left the nest immediately after emergence and did not return. Some females left the nest within a few days of eclosing, while others stayed in their natal nests and began to reactivate them. Reactivations were performed by a single female, by one or more females in the presence of the mother, by more than one female in the absence of the mother, and by more than one female in the presence of females that participated in the prior reactivation. According to behavior, the females were classified as forager/egg-laying and egg-laying females. The oviposition by egg-laying females was always preceded by oophagy. All of the forager/egg-laying and egg-laying females that were dissected had been inseminated. The behavior displayed by egg-laying females is characteristic of brood parasitism and fits the parental parasitism hypothesis developed as an alternative pathway by which insect sociality could have arisen. The behaviors displayed by E. townsendi, together with those reported for Euglossa cordata, show that both species have bionomic traits that resemble the conditions suggested as precursors of the origin of eusociality.Received 12 June 2003; revised 6 April 2004; accepted 22 April 2004.     

Both cooperation and nest position improve larval survival in the sweat bee, <Emphasis Type="Italic">Lasioglossum</Emphasis> (<Emphasis Type="Italic">Evylaeus</Emphasis>) <Emphasis Type="Italic">baleicum</Emphasis>

Nesting by multiple females in sweat bees raises the question of the benefits associated with grouping. Adult numbers of the sweat bee, Lasioglossum (Evylaeus) baleicum, exhibited variation within a nest population. The effects of adult number and nest position on larval survival were analyzed in a nest aggregation. Both variables independently affected larval survival, with the presence of multiple adults in a nest markedly improving survival rate. Nests located near the periphery of the aggregation of nests suffered high larval mortalities. Increased frequency of cell-inspection in multiple-female nests seems to be associated with a concomitant increase in larval cell protection from external predation. Such predation pressure was assumed, given the existence of an underground ant colony that employs a recruitment system while foraging in the study area.     

Group nesting and individual variation in behavior and physiology in the orchid bee <Emphasis Type="Italic">Euglossa nigropilosa</Emphasis> Moure (Hymenoptera,Apidae)

The Euglossini are a key group for studying the traits that promote or hinder highly social behavior in bees because it is the only tribe in the Apine clade without large colonies or females with distinct life histories, e.g. queens and workers. There have been few studies on behavior of orchid bee females in nests because these nests are not found easily. Taking advantage of the relatively high abundance of Eg. nigropilosa nests at Reserva Natural La Planada, Colombia, we examined social behavior of Eg. nigropilosa individuals in five nests (3 original and 2 reused) for nine months. We report this species to have the largest colonies known for Euglossa, with nests reaching up to 22 individuals, and all nests containing more than one female bee from the same generation. These nests presented many traits that correspond to communal insect colonies. No generational overlap and no cooperative brood care were detected. We examined natural enemies and resource limitation as important factors for group nesting. We examined parasitoid attacks to cells in a nest with females and one without females. We also searched for nesting locations and examined nest re-use as indicators of nest site limitation. Lastly, we examined behavioral and physiological differences among females in the same nest. Such differences could be the bases for evolution of alternative life histories among group living females. We examined extent of ovary development and oviposition rates in similarly aged females in the same nest. We found large variation in reproductive effort of young females. We also examined differences in resin foraging and cell usurpation behaviors. Behavioral specialization was observed, with some individuals bringing only resin to the nest. Inside the nests, bees had territories in which they constructed and defended cells. This territoriality may be a defense against usurpation of provisioned cells by nest mates. Received 10 December 2007; revised 2 May 2008; accepted 7 May 2008.     

Ground nesting in the paper wasp <Emphasis Type="Italic">Polistes aurifer</Emphasis> (Hymenoptera,Vespidae)

Summary Polistes paper wasp species vary in their nest site selection, with some nesting in relatively exposed areas and others in protected cavities. Locating the nest sites of cavity dwelling species in natural habitat can prove difficult, and most behavioral studies on these species are therefore conducted using human-built structures. Since Hungerford and Williams (1912) and Rau (1929) noted the location of several P. fuscatus (Fabricius) nests in rodent burrows, there have been no published accounts of ground nesting Polistes species. I report the occurrence of a large, dense cluster of P. aurifer (Saussure) nests located within cracks in the dried soil of the Santa Monica Mountains of southern California, USA.Received 8 April 2003; revised 10 October 2003; accepted 31 October 2003.     

Solitary and group nesting in the orchid bee <Emphasis Type="Italic">Euglossa hyacinthina</Emphasis> (Hymenoptera,Apidae)

Summary Orchid bees (Euglossini) provide a potentially informative contrast for examining origins of advanced social behaviour in bees because they are the only tribe in the apine clade that do not form large colonies or have queens and workers. We investigated natural nests of Euglossa hyacinthina Dressler, an orchid bee that nests singly or in groups. By comparing the two types of nests, we examined if individuals in a group merely share the nest (are communal) or exhibit a level of social organization where there is reproductive division of labour among the females. Observations are consistent with communal nesting, indicating that all females in group nests are reproductively similar to the solitary nesting females because the provisioning of young, as well as the ovary development and mating status of females sharing nests were not different than that of solitary-nesting females. Also, multiple female nests did not produce a female-biased brood as predicted for nests with reproductive division of labour. We also investigated potential advantages of group nesting vs. individual nesting. We demonstrate that per capita offspring production is lower in nests with more than one female. However, we found that nests with single females were left unattended for longer periods of time during foraging, and that there was a high incidence of natural enemy attack in nests when females were absent. Group and solitary nesting may be advantageous under different conditions.Received 3 December 2002; revised 7 March 2003; accepted 2 April 2003.     

Spatio-temporal variation of colony structure and eusociality level of the Japanese sweat bee<Emphasis Type="Italic"> Lasioglossum</Emphasis> (<Emphasis Type="Italic">Evylaeus</Emphasis>)<Emphasis Type="Italic"> duplex</Emphasis> (Hymenoptera: Halictidae)

Colony structure and eusociality level of the sweat bee Lasioglossum (Evylaeus) duplex were studied in 2001 in Sapporo and Assabu, Hokkaido, northern Japan. Sakagami and his colleagues had also studied this species in Sapporo in 1957–1968. Brood size, sex ratio and queen–worker size dimorphism were geographically and temporally variable, indicating spatio-temporal variation at the eusociality level. Inseminated workers constituted only 7.9% of the populations in 1957–1968 Sapporo but about 60% in 2001 in Sapporo and Assabu. A few of the inseminated workers were believed to leave natal nests for independent colony founding. Thus, partial bivoltinism is likely in this sweat bee species. The presence of workers with developed ovaries and/or corpora lutea suggests the occurrence of worker oviposition.     

Antifungal skin bacteria,embryonic survival,and communal nesting in four-toed salamanders, <Emphasis Type="Italic">Hemidactylium scutatum</Emphasis>

We examined a novel hypothesis for the maintenance of communal nesting in the salamander, Hemidactylium scutatum, namely that communal nests are more likely than solitary nests to be associated with cutaneous antifungal bacteria, which can inhibit fungal infections of embryos. A communal nest contains eggs of two or more females of the same species. The nesting behavior of H. scutatum females and survival of embryos were determined by frequent nest surveys at three ponds. For communal nests, embryonic survival tended to be higher and catastrophic nest failure was lower. Pure bacterial cultures of resident species were obtained from the salamanders’ skins by swabbing and tested against a fungal pathogen of embryos (Mariannaea sp.) in laboratory assays. We found that 27% of females had skin bacteria inhibitory to Mariannaea sp. Communal nests were more likely to have at least one female with antifungal bacteria than were solitary nests. Using a culture-independent assay (denaturing gradient gel electrophoresis of 16S rRNA gene fragments), we found that bacterial species on females and embryos were more similar to each other than they were to bacterial species found in soil within the nest, suggesting that females transmitted skin bacteria to embryos. The presence of anti-Mariannaea skin bacteria identified from the laboratory assays did not prevent fungal presence in field nests. However, once a nest was visibly infected with fungi, presence of anti-Mariannaea bacteria was positively correlated with survival of embryos. Microbe transmission is usually thought to be a cost of group living, but communal nesting in H. scutatum may facilitate the transmission of antifungal bacteria to embryos.     

Nesting biology of the arboreal fungus-growing ant <Emphasis Type="Italic">Cyphomyrmex cornutus</Emphasis> and behavioral interactions with the social-parasitic ant <Emphasis Type="Italic">Megalomyrmex mondabora</Emphasis>

We describe the extraordinary nesting habits of the fungus-growing ant Cyphomyrmex cornutus (Formicidae, Myrmicinae, Attini) and the natural history of Megalomyrmex mondabora (Formicidae, Myrmicinae, Solenopsidini), a social parasite that inhabits nests of C. cornutus and other small attine ants. The study was carried out at two sites on the Atlantic slope of Costa Rica. The C. cornutus nest is an oblong mass of accreted soil, attached to or suspended from low vegetation in wet forest understory. Less than a fourth of the nest volume has chambers and is inhabited by C. cornutus; the remainder is a semi-solid mass of accreted soil often housing a variety of arthropods, including other unspecialized commensal ant species. Five C. cornutus colonies examined were parasitized by M. mondabora. Colonies of M. mondabora inhabited chambers very near those of the host. In laboratory observations, M. mondabora and C. cornutus workers interacted with little aggression despite the consumption of C. cornutus larvae and fungi by M. mondabora. During most interactions, C. cornutus workers responded submissively, whereas M. mondabora appeared indifferent or nonresponsive. Megalomyrmex mondabora parasitizes several other attine species (Cyphomyrmex costatus, Cyphomyrmex salvini, and Apterostigma goniodes), and it appears therefore a relatively unspecialized social parasite with broad attine hostassociation. The size of M. mondabora workers vary with host species, suggesting M. mondabora sensu lato comprises either cryptic species or the host environment affects worker size. Received 14 September 2006; revised 1 February 2007; accepted 6 February 2007.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

Mating System and Sleeping Behaviour of the Male and Female <Emphasis Type="Italic">Centris</Emphasis> (<Emphasis Type="Italic">Paracentris</Emphasis>) <Emphasis Type="Italic">burgdorfi</Emphasis> Friese (Apidae,Centridini)

We describe the mating behaviour of males of Centris (Paracentris) burgdorfi, a solitary, univoltine bee, with a wide distribution in Brazil. We also describe the unusual sleeping sites of males and females. The study was performed during two breeding seasons, in northeastern Brazil, where the species nests in aggregations in petrified dunes. Data on mating behaviour were collected through direct observations of the bees at the nesting-emergence site. Males compete intensely for virgin females in the morning, sometimes killing rivals. The high competition for females near the nesting site makes that the male has to take the female to another place to get genital contact. In the evening, males do not use plants to spend the night, instead they aggregate in sleeping clusters inside old burrows in the nesting-emergence area while females sleep in groups on plants that provide the floral oil used in nest construction.     

Nest-moving by the polydomous ant <Emphasis Type="Italic">Cataglyphis iberica</Emphasis>

In this paper we analyze emigration from nests by the polydomous ant Cataglyphis iberica. Social carrying of workers of this species between different nests of the colony is frequent. In Bellaterra (Barcelona, NE Spain), we monitored field emigration of C. iberica by noting for each nest the migratory behavior of C. iberica workers and, when the nests were attacked by another ant species, Camponotus foreli, we noted the number of C. foreli workers involved in the attacks. Emigration of C. iberica from nests was highly variable. We suggest the main factor determining emigration by this species was attack by workers of C. foreli, so emigration from C. iberica nests was much faster when harassment by C. foreli increased. The system of multiple nests of C. iberica enables this species to abandon attacked nests and to reinstall their population in other nests of the same colony. This reduces risk to the colonies because the route between the different nests is well known by transporter workers.     

Parasitism <Emphasis Type="Italic">versus</Emphasis> mutualism in the ant-garden parabiosis between <Emphasis Type="Italic">Camponotus femoratus</Emphasis> and <Emphasis Type="Italic">Crematogaster levior</Emphasis>

Ant-gardens represent a special type of association between ants and epiphytes. Frequently, two ant species can share the same nest in a phenomenon known as ‘parabiosis’, but the exact nature (i.e., mutualistic or parasitic) of this interaction is the subject of debate. We thus attempted to clarify the mutual costs and benefits for each partner (ants and plants) in the Crematogaster levior/Camponotus femoratus ant-garden parabiosis. The ants’ response to experimental foliar damage to the epiphytes and to the host tree as well as their behavior and interactions during prey capture were investigated to see if the purported parasitic status of Cr. levior could be demonstrated in either the ant-ant or in the ant-plant interactions. The results show that both species take part in protecting the epiphytes, refuting the role of Cr. levior as a parasite of the ant-garden mutualism. During capture of large prey Ca. femoratus took advantage from the ability of Cr. levior to discover prey; by following Cr. levior trails Ca. femoratus workers discover the prey in turn and usurp them during agonistic interactions. Nevertheless, the trade-off between the costs and benefits of this association seems then to be favorable to both species because it is known that Cr. levior benefits from Ca. femoratus building the common carton nests and furnishing protection from vertebrates. Consequently, parabiosis can then be defined as the only mutualistic association existing between ant species, at least in ant-gardens. Received 31 August 2006 ; revised 8 December 2006 ; accepted 12 December 2006     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

Social and genetic structure in colonies of the social wasp <Emphasis Type="Italic">Microstigmus nigrophthalmus</Emphasis>

Microstigmus (Hymenoptera: Crabronidae) is a genus of social apoid wasps which represents an origin of sociality independent from vespoids, but which has so far received little attention. Though group-nesting is widespread in Microstigmus, genetic relatedness has so far been studied in only one species, M. comes. We report on the social biology of M. nigrophthalmus, drawing from behavioural observations and molecular genetic analyses of relatedness and kinship. There was no evidence of distinctive behavioural suites that distinguished reproductive and non-reproductive individuals. Females could mate more than once, but mating frequency was low. Mean relatedness within nests was high, particularly between females (close to 0.5), but pairwise relatedness values were very variable, as nestmates displayed a wide range of relationships. Such high levels of relatedness should be a factor promoting social nesting and cooperative brood care in this species, as females gain only a slight genetic advantage through rearing their own offspring rather than those of nestmates. This study provides the finest analysis of genetic structure so far in an apoid wasp, and indicates that the form of sociality varies greatly between species of Microstigmus.     

Social parasitism of <Emphasis Type="Italic">Polistes dominulus</Emphasis> by <Emphasis Type="Italic">Polistes nimphus</Emphasis> (Hymenoptera,Vespidae)

Summary Interspecific facultative social parasitism is well known in ants and in bumble-bees, but it is rarer in wasps. This form of parasitism is traditionally considered to be an intermediate stage in the evolution of obligate interspecific parasitism, where the parasites are no longer able to nest alone. We report field and experimental observations of a newly discovered facultative parasitic relationship between two closely related free-living Polistes species: P. nimphus and P. dominulus. P. nimphus foundresses sometimes usurp the nests of the larger P. dominulus before worker emergence. The invading queen takes over the nest with abundant abdomen stroking on the nest surface and is accepted by workers if they emerge 6 or more days after usurpation. Morphometric comparisons show that the usurper species, though smaller than its victims, has morphological adaptations consisting of larger heads, mandibles and front femora relative to their body size that may give it an advantage during nest invasion. This strategy is likely to be taken only after the foundress loses her original nest because invading P. nimphus queens have lower reproductive success than they would have had on their own nest. Overall, we found that P. nimphus usurpers use strategies of invasion similar to those of two obligate parasites, suggesting that this may be an example of one of the pathways by which social parasitism evolved.Received 4 April 2003; revised 8 August 2003; accepted 14 September 2003.     

Ambush Predation of Stingless Bees (<Emphasis Type="Italic">Tetragonisca angustula</Emphasis>) by the Solitary-Foraging Ant <Emphasis Type="Italic">Ectatomma tuberculatum</Emphasis>

Social insect colonies are high-value foraging targets for insectivores, prompting the evolution of complex colony defensive adaptations as well as specialized foraging tactics in social insect predators. Predatory ants that forage on other social insects employ a diverse range of behaviors targeted at specific prey species. Here, we describe a solitary foraging strategy of the ant Ectatomma tuberculatum, on nest guards of the stingless bee Tetragonisca angustula. We observed multiple instances of E. tuberculatum ambushing and successfully capturing the hovering and standing guards of T. angustula near nest entrances. The unique hovering behavior of the guard caste of this bee species, an adaptation to frequent cleptoparasitism by other stingless bees, may make these guards particularly vulnerable to ground-based, ambush attacks by E. tuberculatum. Likewise, the behavior of the foraging ants appears to adaptively exploit the defensive formations and activity patterns of these bees. These observations suggest an adaptive and targeted predatory strategy aimed at gathering external guard bees as prey from these heavily fortified nests.     

Nesting biology of the fungus growing ants <Emphasis Type="Italic">Mycetarotes</Emphasis> Emery (Attini,Formicidae)

Summary. Fungus-growing ants of the genus Mycetarotes are among the least studied in the tribe Attini. This report documents nest architecture and worker population numbers for 19 nests of M. parallelus and 5 nests of M. acutus, including the first such report for M. acutus. This new information is integrated with the scant biological information reported on Mycetarotes to date. The resulting picture of Mycetarotes life history, as well as the relative ease with which large numbers of nests can be collected and observed in the field, suggest that Mycetarotes (particularly M. parallelus) is an ideal model system for the study of coevolution of lower-attine ants and their cultivated fungi.Received 30 June 2003; revised 17 December 2003; accepted 23 March 2004.     

Behavioural Interactions Between <Emphasis Type="Italic">Crematogaster brevispinosa rochai</Emphasis> Forel (Hymenoptera: Formicidae) and Two <Emphasis Type="Italic">Nasutitermes</Emphasis> Species (Isoptera: Termitidae)

In the savanna-like Brazilian biome caatinga, the arboreal and polydomous ant Crematogaster brevispinosa rochai can be found cohabiting with two closely related Nasutitermes species (N. corniger and N. ephratae). This ant occupies variably sized portions of the termite nests and maintains a physical separation with its hosts by plugging the cells of the boundary areas with fibrous material. Although all the analysed cohabiting C. b. rochai nests were queenless, they always contained brood, especially from male and female reproductive castes. Interaction experiments between workers of C. b. rochai and workers or soldiers of N. ephratae revealed a low level of aggressiveness between the two species that contrasts with the aggressiveness of both C. b. rochai and N. ephratae in encounters with other ant (Azteca cf. chartifex, Cephalotes pusillus) or termite (M. cf. indistinctus) species. The association could benefit both ants (additional nesting sites, brood rearing places) and termites (protection against predators, dead ants or ant refuses as source of nitrogen).     

Colony structure and sex allocation ratios in the ant <Emphasis Type="Italic">Temnothorax crassispinus</Emphasis>

Summary. We analyzed the impact of ecological parameters, such as nest density and nest site availability, on colony organization and investment patterns in two populations of the ant Temnothorax crassispinus, a parapatric sibling species of the well-studied T. nylanderi (Temnothorax was until recently referred to as Leptothorax (Myrafant); Bolton, 1993). As in T. nylanderi, sex allocation ratios were strongly associated with total sexual reproduction, i. e., nests with large sexual investment produced mainly female sexuals. Furthermore, nest site quality affected sex allocation ratios, with colonies from ephemeral nest sites producing a more male-biased sex allocation ratio than colonies from sturdy nest sites. In contrast to T. nylanderi, workers in colonies of T. crassispinus were mostly fullsisters both in a dense and a sparsely populated area, suggesting that colony fusion and colony usurpation are rare in this species. In addition, the presence of a queen in a local nest unit strongly influenced sex ratio decisions, in that these nests raised a more male biased allocation ratio compared to queenless nests. This also suggests that colony structure is more stable in T. crassispinus than in T. nylanderi. We conclude that sibling species, though often very similar in their morphology and ecological requirements, may nevertheless react very differently to ecological variation.Received 11 December 2003; revised 4 March 2004; accepted 19 April 2004.