Comparative morphology of the antenna cleaner in bees (Apoidea)

The morphology of the antenna cleaners of 175 species of bees representing more than 50 genera was investigated and described comparatively. An ancestral type of antenna cleaner was found in most short-tongued bees (except Oxaeidae and Ctenoplectridae) as well as in Nomadinae, Ceratini, and some Megachilidae. A further ancestral character (shallow notch), however, was only found in some of these bees (but throughout Colletidae and Halictidae). A derived type of antenna cleaner was found in Anthophora, Eucera and related genera, as well as in Melecta, Ancyloscelis, and Ctenoplectra. Several further (derived) types of antenna cleaners were found in Anthophoridae and Apidae. A large variety of antenna cleaners (ancestral as well as derived ones) were found in the Megachilidae. Possible phylogenetic consequences are suggested in Figure 37.     

Die Antennen-Putzeinrichtung der Adephaga (Coleoptera), parallele evolutive Vervollkommnung einer komplexen Struktur

The antenna cleaner of the Adephaga (Coleoptera), parallel evolutionary perfection of a complex structure Independent evolutionary perfection of a complex character ist to be expected in a splitting phylogenetic line, if selective advantages for improving this character persist in the various lines after splitting. Independent perfection leading to very similar results is termed “parallel perfection”. It can either be based on a narrow channelling, due to limitations of the evolutionary substrate (i. e. morphological structures and patterns of behaviour, inherited from the common ancestor), or on very specific functional advantages. The evolution of the antenna cleaner on the protibia of adephagous beetles is analysed under this aspect. The phylogenetic relationships within the Geadephaga are discussed. After defining the terms “anisochaetous” and “isochaetous” somewhat differently from previous usage, the two phylogenetic lines Anisochaeta (Carabidae in a broad sense) and Isochaeta (Metriidae, Ozae-nidae, Paussidae) may be considered sister groups. The Cicindelidae might have split off from the line of the Anisochaeta very early. The Rhysodidae are interpreted as an aberrant offshoot from the primitive adephagous stock. An attempt is made to reconstruct the antenna cleaner of the common ancestor of Anisochaeta and Isochaeta from the sum of plesiomorphous characters. This primitive cleaner might have corresponded in many respects to the array of setae found on an undifferentiated protibia (see comparison with mesotibia). The cleaner of the Cicindelidae, in spite of many plesiomorphous characters cannot be regarded as a model for the common ancestral form, because it already shows some differentiation towards the condition found in the Anisochaeta. The antenna cleaners of highly evolved Anisochaeta and Isochaeta, which correspond surprisingly well in many details, can be traced back to a common basis, but most of their conformities must have evolved independently. The following parallelisms were established: the ventral side of the tibia is shifted to the functional inner side and forms an oblique groove (cleaning channel); the antenna cleaning comb, originally placed transversely, becomes lengthwise oriented and is shifted in proximal direction; several setae are modified so as to form together a pressure clamp; the cleaning channel is completed by a spine-like extension projecting from the tibia; the terminal setal ring forms an additional comb. The gradual perfection of the antenna cleaner can be studied in the more primitive species in both the groups Anisochaeta and Isochaeta. Apparently, the two lines entered different pathways of perfection to begin with, and the cleaners became similar again by convergence later on. This indicates a very slight channelling effect of the common evolutionary substrate. The numerous conformities in the higher evolved genera must result from “parallel selection” (channelling for functional reasons). By observing grooming behaviour in primitive and highly evolved genera, the following advantages connected with the more complex structures could be established: the engagement of the antenna into the cleaning channel is facilitated; the mechanical strain on the antenna during grooming is reduced; the cleaning of antennae and eyes becomes more effective. That the antenna cleaner itself can better be kept clean by a comb of the Mesotibia was proved quantitatively.     

Diversity of native bee visitors of cucurbit crops (Cucurbitaceae) in Yucatán, México

Native bees in Yucatán, México are treatened by agricultural land uses that limit their food resources, alter their reproductive habitats and increase their mortality. Various species may disappear before their importance for regional agricultural productivity and ecosystem maintenance is known. In order to assess their assemblage as visitors of cucurbit crops, we sampled more than 2000 bee specimens on fourteen fields of pumpkin, cucumber, melon and watermelon from five localities between 1995 and 1997; sample units consisted of all bees collected by net sweeping in a given field during 25–30 accumulated hours on separate days. The fourteen samples comprised bees of six families, 29 genera and 58 species. Composition per sample ranged between 10 and 27 species and abundance between 28 and 444 individuals. Seven species (six genera) of Apidae, Anthophoridae and Halictidae comprised around 80% of all the individuals collected. Yet, diversity measures indicated intermediate to high evenness in most sampled bee assemblages, i.e. despite the frequent abundance of some species of Augochlora, Partamona, Ceratina, Trigona or Peponapis, other bee visitors were also relatively important, particularly in small samples. Individual samples of pumpkin bee visitors had significantly different evenness among themselves and to other cucurbit crops. The percent similarity and number of shared species among the fourteen samples were both usually low; lumping data per crop and locality showed, however, higher evenness and more common species than individual samples. Results are discussed in terms of future research priorities: natural history of tropical native bees and strategies to monitor bee community changes for conservation purposes.     

Grooming and pollen manipulation in bees (Apoidea): the nature and evolution of movements involving the foreleg

Three modes of self cleaning occur in insects: nibbling by the maxillae, scraping one structure by another in one direction only, and rubbing back and forth while the respective parts are in continuous contact. This paper describes a comprehensive and comparative account of this behaviour in bees, with special reference to the cleaning of or by the forelegs. Bees, like all Hymenoptera, clean various parts of the head, including the mouthparts and the antennae, with the forelegs. Lower Hymenoptera scrape each antenna with either foreleg; in the species of Aculeata that possess the antenna cleaner (strigil) on the foreleg, only the ipsilateral leg is used. The thoracic dorsum of most bees, as in many sphecoid wasps, is scraped in a forward direction by the middle leg; Triepeolus spp., however, use the hind leg, and the Anthophorinae the foreleg. Some beetles and lacewings clean their forelegs in the mouthparts by nibbling and scraping. Most higher Hymenoptera as a rule scrape the foreleg between the ipsilateral maxilla and the labium; bees, however, clamp the foreleg between the flexed ipsilateral middle leg and then scrape it. An evolution of this behaviour is postulated via several intermediate forms derived from original stepping movements. Halictidae and Andrenidae clamp the foreleg for scraping underneath the middle tibia, whereas all other bees nearly always clamp it underneath the middle basitarsus. Very similar movements are used in various species for transferring pollen, oil, or nest materials from the foreleg to the middle leg. It is argued that the original way of pollen carrying in bees must have been by filling the crop through direct eating or by scraping pollen off the foreleg between the ipsilateral maxilla and the labium. The latter movement is widespread among bees and is homologous to the normal foreleg cleaning in the mouthparts of most other Hymenoptera. The efficiency of this behaviour is enhanced in many lower bees by a comb on the galea, which is the homologue of a similar structure widespread among aculeate wasps. In higher bees, Apidae and Anthophoridae, the galeal comb is replaced by an equifunctional stipes comb. Many bees have neither of these types of maxillary combs.     

A phylogenetic study of the antenna cleaner in Formicidae,Mutillidae, and Tiphiidae (Insecta,Hymenoptera)

Summary The morphology of the tibio-tarsal antenna cleaner (strigilis) of 30 species of Formicidae, 14 species of Mutillidae and 9 species of Tiphiidae was investigated and is described comparatively. In Formicidae, there is a common type of strigilis. The spur has a posterior-dorsal comb and its anterior side is covered with squamae, which usually form a brush. Posteriorly, the basitarsal notch bears a comb and anteriorly specialized paddle-shaped hairs. These characters may be apomorphic for Formicidae. In several Ponerinae and in Myrmecia there is also a velum on the spur. In two species of ants which are strongly parasitic (Teleutomyrmex schneideri and Anergates atratulus ) there are reduced antenna cleaners. Mutillidae (except Myrmosa) have another common type of strigilis: the spur bears a velum with a smooth rim and a clear apex with two rows of teeth. The notch in the basitarsus is usually deeper than that of ants; there is a comb, but no paddle-shaped hairs. The strigilis of Myrmosa females has no velum but there are two prominent rows of teeth on the spur. In the male, the velum is reduced to a slender strip. In Tiphiidae the antenna cleaners show considerable diversity. In Methocha the spur bears a comb but no velum; the spur of Tiphia has a velum with a serrated rim; the spur of Myzinum is equipped with a velum with a smooth rim; in Thynnus the surface of the velum is wrinkled or undulating. An apex (without teeth) is present in all investigated Tiphiidae. The notch of the basitarsus bears a comb except in female Myzinum, where the teeth seem to have fused, thus forming a rim. It is suggested that a velum with a serrated or toothed edge and an apex with two rows of teeth are plesiomorphic for aculeate Hymenoptera. The antenna cleaners in Mutillidae are remarkably similar to those in some bees. This fact is interpreted as partly due to convergence and as partly symplesiomorphic. In several species of ants, there is a specialized cuticular area (bande poreuse) anterior to the comb of the notch, which is characterized by fissures or holes. These are presumably openings of an excretory gland.     

Ground-nesting bees determine the location of their nest relative to a landmark by other than angular size cues

Bees and wasps acquire a visual representation of their nest's environment and use it to locate their nest when they return from foraging trips. This representation contains among other features cues to the distance of near-by landmarks. We worked with two species of ground-nesting bees, Lasioglossum malachurum (Hymenoptera: Halictidae), Dasypoda hirtipes (Hymenoptera: Melittidae) and asked which cues to landmark distance they use during homing. Bees learned to associate a single cylindrical landmark with their nest's location. We subsequently tested returning bees with landmarks of different sizes and thus introduced large discrepancies between the angular size of the landmark as seen from the nest during training and its distance from the nest. The bees' search behaviour and their choice of dummy nest entrances show that both species of ground-nesting bees consistently search for their nest at the learned distance from landmarks. The influence of the apparent size of landmarks on the bees' search and choice behaviour is comparatively weak. We suggest that the bees exploit cues derived from the apparent speed of the landmark's image at their retina for distance evaluation.     

Recent and rapid diversification of the small carpenter bees in eastern North America

Bees fulfil a critical ecological role as pollinators, significantly contributing to the reproductive success of myriad angiosperm species. Although increasingly appreciated for their agricultural contributions, relatively little is yet known about the natural history of the vast majority of the more than 20 000 wild bee species worldwide. The small carpenter bee genus Ceratina occurs globally, and is represented in North America by its most recently diverged subgenera, Ceratinula and Zadontomerus. Recent genetic analysis of eastern Ceratina (Zadontomerus) supports the existence of five closely related, yet genetically distinct species living in sympatry. This phylogeographical study employs molecular barcoding of the most comprehensive specimen collection yet assembled to confirm the identities of these recently diverged eastern North American Ceratina (Zadontomerus) species. Delineation of extant population structure, evolutionary history and known range of this emerging model native pollinator are greatly improved by this study. We consider ecological and behavioural factors potentially contributing to the maintenance of genetic identity among these sympatric species.     

Antennal Asymmetry in Social Behavior of the Australian Stingless Bee, <Emphasis Type="Italic">Tetragonula carbonaria</Emphasis>

The left and right antennae of stingless bees have different roles in learning and recall of olfactory memory. Antennal asymmetry in social behavior is reported here. Approaches and physical contacts were scored in dyads of stingless bees (Tetragonula carbonaria): dyads in which both bees had only their right antennae (left antennae removed) made significantly more physical contacts with each other than dyads in which both bees had only their left antennae. In dyads of one left and one right, it was found, unexpectedly, that the bee with a left antenna approached the bee with the right antenna more often that the other way around, and the bee with the left antenna often attacked (by biting) its hive mate. Hence, the low number of contacts in dyads of bees using their left antennae appears to be due to mutual avoidance. Whereas use of the right antenna stimulates positive contact, the left stimulates avoidance or attack. Via such left-right asymmetries, intact bees may compute behavior directed towards friend and foe. Such antennal asymmetry may have evolved concomitantly with eusocial behavior. We found no evidence that it was associated with significant differences in the number of olfactory or non-olfactory sensilla on the left versus right antenna.     

Thermal effects on activity patterns and behavioural switching in a concourse of foragers on Stachytarpheta mutabilis (Verbenaceae) in Papua New Guinea

Summary 1. The foraging activities of the papilionid butterflies Ornithoptera priamus poseidon and Papilio ulysses, and the solitary bee Amegilla sapiens (Apoidea, Anthophoridae) on the shrub Stachytarpheta mutabilis were studied in highland Papua New Guinea. 2. The insects' activity patterns were analysed at three sites with differing diurnal microclimate variation. O. priamus and A. sapiens foraged in the morning (after a period of basking and wing-whirring) and late afternoon when temperatures were well below daily maxima, whereas P. ulysses showed foraging peaks during the hottest part of the day. 3. Site choice by all 3 species appeared to be determined primarily by temperature, but within the limits imposed by temperature, nectar supplies probably determined which site was visited. 4. P. ulysses showed interspersed foraging and courtship behaviour, and no behavioural switching was observed for this species. At high temperatures, both O. priamus and A. sapiens ceased foraging and showed territorial and courtship behaviour. This behavioural change allowed avoidance of heat stress, and occurred even when nectar supplies were maintained at high levels. 5. Thermal effects on behavioural switching in these insects are compared with related phenomena in other bees and butterflies.     

Adaptive behaviour of honeybees (Apis mellifera) toward beetle invaders exhibiting various levels of colony integration

Social bees generally host fewer nest invaders than do ants and termites. This is potentially explained by the adaptive defensive strategies of host bees when faced with nest invaders exhibiting various levels of colony integration (based on adaptations to the nest habitat and frequency of nest inhabitation). In the present study, experiments are performed to determine the behaviour at the nest entrance of European honeybee guards Apis mellifera L. (Hymenoptera: Apidae) toward beetle invaders of various levels of behavioural integration into colonies. The species used to test this include Aethina tumida Murray (Coleoptera: Nitidulidae), which is regarded as a highly integrated, unwelcome guest (synechthran) or true guest (symphile); Lobiopa insularis Laporte (Coleoptera: Nitidulidae) and Epuraea luteola Erichson (Coleoptera: Nitidulidae) that are accidentals; and Carpophilus humeralis Fabricius (Coleoptera: Nitidulidae), Carpophilus hemipterus L. (Coleoptera: Nitidulidae) and Tribolium castaneum Herbst (Coleoptera: Tenebrionidae), all of which are species that are not integrated into honeybee colonies. The responses of guard bees to a control bead also are noted. In general, bees ignore T. castaneum and E. luteola to a greater extent than other beetle species. Bees make contact with the black glass bead (a non‐aggressive behaviour) more than they do all beetle species. Bees treat A. tumida more defensively than they treat any other beetle species and the level of bee defensiveness varies by colony. These data suggest an adaptive heightened defensive response by bees toward the most integrated colony intruder but a significantly reduced level of response toward invaders representing all other levels of colony integration.     

Tactile learning in the honeybee

Free-flying bees were conditioned on a vertical wall to a vertical tactile pattern consisting of parallel lines of grooves and elevations. The asymptote of the learning curve is reached after approximately 25 rewards. Bees can discriminate the conditioned vertical pattern from a horizontal or diagonal alternative. Angle discrimination is apparent only for relatively coarse tactile cues. The proboscis extension response of fixed bees was used to condition bees to a vertical tactile pattern which was presented to the antennae. The learning curve reaches an asymptote after 4 rewards. After 7 unrewarded extinction trials the conditioned responses are reduced to 50%. Bees show best discrimination for patterns whose edges they can scan with their antennae. The animals show a high degree of generalization by responding to an object irrespective of the trained pattern. Under laboratory conditions fixed bees can discriminate the angles and spatial wavelengths of fine tactile patterns consisting of parallel grooves. Bees can also discriminate forms and sizes of tactile patterns. They do not discriminate between different types of edges and between positive and negative forms. Accepted: 17 September 1998     

A comparative study of cleaning activity of two reef fishes at Fernando de Noronha Archipelago,tropical West Atlantic

Cleaner fishes are usually classified as obligate or facultative cleaners according to their diet and the extent to which their nutritional requirements in the different ontogenetic stages are gained from cleaning. While obligate cleaners clean throughout their lives and ingest mainly food taken from the clients’ body surface, facultative cleaners clean only as juveniles and have a broader diet. In addition, some facultative cleaners may experience a relatively higher predation risk, and thus rarely interact with piscivorous fishes. Despite these acknowledged differences, there are very few studies that compare cleaning activity of obligate and facultative cleaners within the same area. Cleaning activity of the obligate cleaner goby Elacatinus cf. randalli and the facultative cleaner wrasse Thalassoma noronhanum were comparatively examined at Fernando de Noronha Archipelago, tropical West Atlantic. The client assemblage attended by the two cleaners differed, as the goby attended a slightly greater diversity of species (22), mostly piscivores and zoobenthivores, and the wrasse attended fewer species (19), mostly planktivores. Chromis multilineata was the most common client species of both cleaners, although body size (which is expected to be positively correlated to clients’ ectoparasite load) of C. multilineata individuals attended by the goby was larger than that of the individuals attended by the wrasse. Despite such differences, T. noronhanum showed a surprisingly species-rich client assemblage when compared with other cleaners of the genus Thalassoma. In addition, the frequency and time spent on cleaning interactions, as well as the number of client species attended per 10-min period, was similar for both cleaner species, which indicate that they have important yet complimentary ecological roles in the reef community at Fernando de Noronha Archipelago.     

Anthecology of Orchis mascula (Orchidaceae)

Interactions between Orchis mascula L. ssp. mascula and anthophilous insects were studied mainly on the island of Öland, Sweden. The species is nectarless and acts by deceiving various bees (Hymenoptera, Apoidea). Temporary seeking/exploratory drives of bees for food–sources were exploited by means of superiority in floral display. A floral reflectance maximum at 440 nm (blue) largely determines the colour in the visual spectrum of bees. The floral scent is composed to about 90% of mono–terpenes. Nothing suggested mimesis of concurrent food–flowers either in colour or in scent. Anthesis covered a time period in late spring when (a) concurrent food–flowers for bees were rather few, (b) many bumble–bee queens (Bombus Latr. spp.) were inexperienced as regards food–flowers and had no foraging routines, (c) cuckoo bumble–bee females (Psithyrus Lep. spp.) recovery–fed on flowers after hibernation, (d) males of Eucera longicornis (L.) (Anthophoridae) patrolled far out from their nest–area, and when (e) many solitary bee species flew about. The pollinator fauna differed between sites, viz. either Bombus queens, Psithyrus females or E. longicornis males transported the majority of the pollinaria. The pollination system does not seem to be stabilised in the study areas. Floral morphology indicates that the plant's anthecological specialisation to bees as a group implies a graded unspecialisation to each species of bee in the legitimate pollinator group.     

Whale sharks Rhincodon typus get cleaned by the blue-streak cleaner wrasse Labroides dimidiatus and the moon wrasse Thalassoma lunare in the Philippines

Cleaning interactions are essential for healthy marine ecosystem communities. This study reports the first documentation of the whale shark Rhincodon typus cleaning behaviour in the Indo-West Pacific by two wrasse species, the blue-streak cleaner wrasse Labroides dimidiatus and the moon wrasse Thalassoma lunare in Cebu, Philippines. This study documented 36 cleaning interactions with 14 individual whale sharks. The cleaning interactions appear opportunistic rather than targeted by the sharks, unlike that observed in other species of elasmobranchs. Further work should focus on understanding the drivers of these unique cleaning interactions.     

Hunting for healthy microbiomes: determining the core microbiomes of <Emphasis Type="Italic">Ceratina,Megalopta</Emphasis>, and <Emphasis Type="Italic">Apis</Emphasis> bees and how they associate with microbes in bee collected pollen

Social corbiculate bees such as honey bees and bumble bees maintain a specific beneficial core microbiome which is absent in wild bees. It has been suggested that maintaining this microbiome can prevent disease and keep bees healthy. The main aim of our study was to identify if there are any core bacterial groups in the non-corbiculate bees Ceratina and Megalopta that have been previously overlooked. We additionally test for associations between the core bee microbes and pollen provisions to look for potential transmission between the two. We identify three enterotypes in Ceratina samples, with thirteen core bacterial phylotypes in Ceratina females: Rosenbergiella, Pseudomonas, Gilliamella, Lactobacillus, Caulobacter, Snodgrassella, Acinetobacter, Corynebacterium, Sphingomonas, Commensalibacter, Methylobacterium, Massilia, and Stenotrophomonas, plus 19 in pollen (6 of which are shared by bees). Unlike Apis bees, whose gut microbial community differs compared to their pollen, Ceratina adults and pollen largely share a similar microbial composition and enterotype difference was largely explained by pollen age. Megalopta displays a highly diverse composition of microbes throughout all adults, yet Lactobacillus and Saccharibacter were prevalent in 90% of adults as core bacteria. Only Lactobacillus was both a core bee and pollen provision microbe in all three species. The consequences of such diversity in core microbiota between bee genera and their associations with pollen are discussed in relation to identifying potentially beneficial microbial taxa in wild bees to aid the conservation of wild, understudied, non-model bee species.     

Only mothers feed mature offspring in European Ceratina bees

Parental care directed to adult offspring is uncommon in animals. Such parental care has been documented in Xylocopinae bees (Hymenoptera: Apidae). Moreover, some Ceratina bees (Xylocopinae) are known to feed mature siblings, and feeding of mature siblings is achieved by dwarf eldest daughters when mothers died. These daughters are intentionally malnourished by mothers and usually originate from the first brood cell. Here, we examined the pattern of care provided to young adults in three small European carpenter bees: Ceratina (Ceratina) cucurbitina, C. (Euceratina) chalybea, and C. (E.) nigrolabiata. Observations of nest departures and arrivals were performed to study foraging behavior. We detected intensive foraging behavior of mothers in all three studied species. However, we did not observe regular foraging behavior of daughters in any species. The experimental removal of mothers in C. cucurbitina led to the emigration of young adults and did not initiate foraging activity in daughters. We conclude that the feeding of siblings does not occur in these species unlike in the American species C. calcarata. We detected female-biased sex ratios in the first brood cell in C. cucurbitina and C. chalybea. Female offspring in the first brood cell was smaller than other female offspring only in C. cucurbitina. Our results show that a female-biased sex ratio and the small size of daughters in the first brood cell do not provide sufficient evidence for demonstrating the existence of an altruistic daughter and also that the pattern of maternal investment is not exclusively shaped by social interactions.     

Ctenoplectridae and the origin of long-tongued bees*

The Ctenoplectridae (for the genera Ctenoplectra and Ctenoplectrina) are separated from the Melittidae, in which they have commonly been included. Ctenoplectrids differ from melittids in many characters, and are more closely related to the long-tongued families Fideliidae, Megachilidae, Anthophoridae, and Apidae than to the Melittidae and other short-tongued families. Ctenoplectrids are the sister-group of the long-tongued bees (and could be considered as long as long-tongued bees with labial palpi of short-tongued bees); melittids are the sister-group to the ctenoplectrids plus long-tongued bees. Characters are listed that support these relationships. Ctenoplectra species appear to be oil collectors from flowers of curcurbits. Some behavioural observations are included. Ctenoplectrina is a probable parasite in nests of Ctenoplectra, as it lacks the pollen and oil manipulating apparatus.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Caribbean Cleaning Gobies Prefer Client Ectoparasites Over Mucus

If cooperation often involves investment, then what specific conditions prevent selection from acting on cheaters that do not invest? The mutualism between the Indo‐Pacific cleaner wrasse Labroides dimidiatus and its reef fish clients has been a model system to study conflicts of interest and their resolution. These cleaners prefer client mucus over ectoparasites – that is, they prefer to cheat – but punishment and partner switching by clients enforce cooperative behaviour by cleaners. By contrast, clients of Caribbean cleaning gobies (Elacatinus spp.) do not to use punishment or partner switching. Here, we test the hypothesis that the behavioural differences between these two cleaner fish systems are caused by differences in cleaner foraging preferences. In foraging choice experiments, we offered broadstripe cleaning gobies Elacatinus prochilos client‐derived parasitic isopods, client mucus and a control food item. The cleaning gobies significantly preferred ectoparasites over mucus or the control item, which contrasts with cleaner wrasses. We propose that the low level of cleaner–client conflict arising from cleaning goby foraging preferences explains the observed lack of strategic partner control behaviour in the clients of cleaning gobies.     

Multiple cleaner species provide simultaneous services to coral reef fish clients

Cleaning symbioses on tropical coral reefs are typically documented between two species: a single client fish and one or more conspecific cleaners. However, multiple cleaner species living sympatrically in the Caribbean have been anecdotally reported to simultaneously clean the same client. Nothing is known about the patterns and processes driving these interactions, which may differ from those involving a single cleaner species. Here, we used remote underwater videography on three reefs in Honduras to record simultaneous cleaning interactions involving Pederson''s cleaner shrimp (Ancylomenes pedersoni) and cleaner gobies (Elacatinus spp.). A pilot study on adjacent shrimp and goby stations found interactions were always initiated by shrimp. A larger, multi-year dataset shows cleaner gobies joined 28% of all interactions initiated at A. pedersoni cleaning stations with cleaner gobies residing nearby. Client body size significantly predicted simultaneous cleaning interactions, with 45% of interactions simultaneous for clients greater than 20 cm total body length compared with only 8% for clients less than 20 cm. We also found that simultaneous cleaning interactions lasted over twice as long as shrimp-only interactions. We propose these novel multi-species interactions to be an ideal model system to explore broader questions about coexistence, niche overlap and functional redundancy among sympatric cleaner species.