Film Analysis of the Tactile Communication between Cryptocentrus steinitzi (Pisces,Gobiidae) and Alpheus purpurilenticularis (Crustacea,Alpheidae)

The tactile communication between the symbiotic goby Cryptocentrus steinitzi and its burrowing shrimp partner Alpheus purpurilenticularis was investigated by film analysis. Warning signals of the goby which elicit the retreat of the shrimp into its burrow consist mainly of rapid fin flicks transmitted to the shrimp through its long antennae. Five different parameters of the structure of the warning signal were analyzed. The effect of various variables on the response of the shrimp and its latency were examined with a stepwise multiple regression analysis. Feedback mechanisms involved in this communication system are described. The evolution of the tail flick warning signal and a model describing the relationship between the main elements involved in the generation of warning signals are discussed.     

Comparative Ecology of the Gobies <Emphasis Type="Italic">Nes longus</Emphasis> and <Emphasis Type="Italic">Ctenogobius saepepallens</Emphasis>, Both Symbiotic with the Snapping Shrimp <Emphasis Type="Italic">Alpheus floridanus</Emphasis>

Synopsis We re-examined the symbiotic association of the western Atlantic gobiid fishes Nes longus and Ctenogobius saepepallens with the snapping shrimp Alpheus floridanus on the basis of a critical literature review and new data. Our research confirms that N. longus interacts closely with the shrimp and is dependent on it for the cover provided by the burrow that the shrimp constructs; the goby serves as the sentinel at the burrow entrance. Ctenogobius saepepallens is often seen occupying a burrow of the alpheid, and the shrimp will leave the burrow to deposit sediment with the goby at the entrance, even pushing the goby aside at times. However, the shrimp does not make contact with the goby with its antennae, nor does the goby communicate with caudal fin fluttering at the approach of danger. We suggest that their relationship is a first step in an evolutionary process that may lead to the very close mutualistic association exhibited by N. longus and the alpheid, as well as Indo-Pacific shrimp gobies of 13 different genera and their alpheid partners. Nes longus remains close to the burrow entrance; it feeds mainly on small gastropods, decapod crustaceans, ostracods, and isopods. By contrast, C. saepepallens makes longer excusions from the shelter of the burrow; its diet is dominated by benthic copepods, followed by ostracods and lesser amounts of foraminiferans, isopods, and decapod crustaceans. By virture of its greater mobility, it can be more selective in its prey.     

Competition by obligate and facultative mutualists for partners in a shrimp-goby association

Mutualist species compete intra and inter-specifically for the resources provided by their partners. Because obligate mutualists are more reliant than facultative mutualists on the resources that their partners provide, they are expected to compete more strongly for those resources. Here, I examined interference competition in two goby fishes: Nes longus (an obligate mutualist) and Ctenogobius saepepallens (a facultative mutualist). Both gobies associate with the shrimp, Alpheus floridanus. Shrimp provide gobies with refuge from predators (a burrow in the sand), and gobies provide shrimp with a warning signal when predators are near. Using an aquarium experiment, I examined the behavior of a pair of gobies with access to a single shrimp burrow. I used four different goby pairings: large N. longus and small N. longus, large N. longus and small C. saepepallens, large C. saepepallens and small N. longus, and large C. saepepallens and small C. saepepallens. When paired with large N. longus individuals, small gobies of both species were less likely to occupy the single burrow than when paired with large C. saepepallens individuals. In addition, large N. longus individuals were less likely to co-occupy the single burrow with smaller gobies than were large C. saepepallens individuals. These results seem to indicate that large N. longus individuals exclude smaller gobies from burrows, while large C. saepepallens individuals do not. This study adds evidence to the supposition that obligate mutualists in general compete more strongly for mutualist partners than do facultative mutualists.     

Phylogeny and evolution of Indo-Pacific shrimp-associated gobies (Gobiiformes: Gobiidae)

Despite the ubiquity of obligate mutualisms on coral reef ecosystems, little is known about the evolution of many participating species. The shrimp gobies, known primarily from the coral reef habitats of the Indo-Pacific, are small benthic fishes that participate in a remarkable mutualism with alpheid shrimp. In this mutualism, the shrimp build and maintain a burrow that is guarded by the goby, and the shrimp and goby engage in an intricate tactile communication system. The mutualism is obligate for most shrimp gobies as participating species are highly vulnerable to predation when separated from a shrimp partner. We use phylogenetic analysis of nuclear and mitochondrial DNA sequence data to infer evolutionary relationships among shrimp gobies, and between shrimp gobies and their non-mutualistic gobiid relatives. We show that the mutualist shrimp association has arisen twice among gobies, once in a clade composed of Amblyeleotris, Ctenogobiops, and Vanderhorstia, and a second time in a clade including Cryptocentrus, Mahidolia, Tomiamichthys and Stonogobiops. We then compare the evolution of traits within each shrimp goby clade and consider their intrarelationships. We document cryptic diversity among shrimp gobies, with three distinct clades delineated among Mahidolia mysticina specimens captured at the same locality, paired with the same shrimp species. Mahidolia is placed as sister to the Cryptocentrus species Cryptocentrus cinctus; both exhibit pronounced dichromatism, occurring in both brown and yellow (xanthic) forms. We additionally clarify species identities within Amblyeleotris, confirming that widespread similar species Amblyeleotris fasciata, Amblyeleotris steinitzi and Amblyeleotris wheeleri are all distinct. We hypothesize that the flexibility of gobiid gobies and alpheid shrimp to interact with mutualist partners, as well as the apparently highly beneficial nature of mutualism between them, has contributed to the dual evolution of shrimp-association among Indo-Pacific gobies.     

Dynamics of demographically open mutualists: immigration, intraspecific competition, and predation impact goby populations

Although it is now recognized that mutualistic species are common and can have stable populations, the forces controlling their persistence are poorly understood. To better understand the mechanisms that impact the stability of obligate mutualists, I conducted several field experiments within a sandy coral reef lagoon in Moorea, French Polynesia that manipulated densities of fish (gobies) that interact mutualistically with shrimp. Obligate, mutualistic partnerships of gobies and shrimp are common on Indo-Pacific coral reefs and have been shown previously to interact as follows: shrimp construct burrows in which both species reside, and gobies warn shrimp of predators through tactile communication. Augmentation of gobies by up to 100% above ambient densities within 9 m² plots produced no change in overall density of gobies or shrimp because gobies competed intraspecifically for a limited number of shrimp burrows and smaller gobies were outcompeted by larger individuals. I used predators to assess the impact of goby removal on the stability of goby and shrimp populations. First, although surveys taken throughout the lagoon revealed no relationship between goby and predator densities, predators correlated negatively with the proportion of adult gobies and positively with the proportion of small gobies paired with large shrimp. Second, experimental augmentation of predators resulted in a dramatic reduction of adult gobies within predator-addition plots, but had no impact on overall densities as immigrants rapidly replaced the missing adult gobies. Furthermore, goby turnover resulted in an increase in the proportion of small gobies paired with large shrimp because body sizes of gobies and shrimp in a burrow were similar prior to predator introduction, and predators apparently had a greater impact on gobies than shrimp. The mechanisms that prevent expansion (intraspecific competition) and collapse (immigration) of goby-shrimp populations likely contribute to local-scale stability of mutualistic populations in other terrestrial and aquatic environments.     

Burrow morphology and utilization of the goby (Parapocryptes serperaster) in the Mekong Delta,Vietnam

Some fish species living in mudflats construct burrows for dwelling and hiding. The goby Parapocryptes serperaster is a burrowing fish in mudflats of many estuaries in South East Asia. This study was carried out in the Mekong Delta, Vietnam, to examine burrow morphology and usage by this species. Morphology of the burrows constructed by P. serperaster was investigated by resin castings in situ to obtain the physical structure and configuration of each burrow. Fish from the burrows were caught and measured before burrow casts were made. Fish burrows comprised several openings, a few branching tunnels and multi-bulbous chambers. The surface openings were circular, and the shapes of branching tunnels were nearly round. The burrows had interconnected tunnels and various short cul-de-sac side branches. The burrow structure differed between fish sizes, but burrow dimensions were positively correlated with fish size, indicating that larger fish can make larger and more sophisticated burrow. The burrow structure and dimensions were not different between the dry and wet seasons. Laboratory observations showed that P. serperaster used body movements to dig burrows in the sediment. Burrows could provide a low-tide retreat and protection from predators, but were not used for spawning and feeding for this goby species. This study indicates that the burrowing activity of gobies is an important adaptation for living in shallow and muddy habitats.     

Impact of two different flood pulses on planktonic communities of the largest floodplain lakes of the Daugava River (Latvia)

This study investigated whether surface hole counts could be used as a reliable estimate of density of the ghost shrimps Trypaea australiensis Dana 1852 and Biffarius arenosus Poore 1975 (Decapoda, Thalassinidea) in south eastern Australia. The relationship between the number of holes and the number of ghost shrimps was explored in two ways. Resin casts were used to document any changes in the number of burrow openings per shrimp burrow over time. Manual suction pumping (bait pumping) within a given mudflat area was used to directly compare the number of holes on the sediment surface with the number of ghost shrimps occupying the corresponding volume of sediment. Resin casting showed that throughout the year, the burrows of T. australiensis consistently had an average of two openings, whereas the burrows of B. arenosus showed much greater variability over time with two to four openings per burrow. Overall, a significant relationship between the number of holes and the number of ghost shrimps (mixed species populations) was found, with 2.1 burrow openings for each ghost shrimp. However, some temporal and spatial variation was seen in this relationship. We suggest that the hole count method may be reliable in estimating ghost shrimp densities with restricted use and site specific validation based on some limitations found in this study. Handling editor: K. Martens     

Warning Signal Efficacy: Assessing the Effects of Color,Iridescence, and Time of Day in the Field

Warning coloration deters predators from attacking distasteful or toxic prey. Signal features that influence warning color effectiveness are not well understood, and in particular, we know very little about how effective short‐wavelength and iridescent colors are as warning color elements in nature and how warning signal effectiveness changes throughout the day. We tested the effect of these factors on predation risk in nature using specimens of the distasteful pipevine swallowtail butterfly, Battus philenor. B. philenor adults display both iridescent blue and diffusely reflecting orange components in their warning signal. We painted B. philenor wings to create five different model types: all‐black, only‐iridescent‐blue, only‐orange, iridescent‐blue‐and‐orange (intact signal), and matte‐blue‐and‐orange. We placed 25 models in each of 14 replicate field sites for 72 h and checked for attacks three times each day. Model type affected the likelihood of attack; only‐orange models were, the only model attacked significantly less than the all‐black model. Iridescence did not enhance or decrease warning signal effectiveness in our experiment because matte‐blue‐and‐orange models were attacked at the same rate as iridescent‐blue‐and‐orange models. Time of day did not differentially affect model type. Video recordings of attacks revealed that insectivorous birds were responsible. The results of this experiment, when taken with previous work, indicate that the response to blue warning coloration is likely dependent on predator experience and context, but that iridescence per se does not affect warning signals in a natural context.     

Parental care and reproductive behavior of the clown goby, <Emphasis Type="Italic">Microgobius gulosus</Emphasis>, with observations on predator interactions

Describe reproductive behavior and mating system of the clown goby from field observations. Clown gobies exhibit a loosely haremic mating system. Pairs construct burrows at the base of cattails, the roots of which provide structural support and a spawning substrate. Larger males monopolize multiple burrows, each with an individual female. After spawning, males camouflage burrow entrances with sand and females brood developing young for 4 days. Males continue to guard the covered nests in 50% of observed brooding periods. Burrows are also used as shelter from predators. Both sexes confront intruders but only males exhibit a distinct color response to juvenile blue crabs, Callinectes sapidus, the most significant predator. The male color response appeared to mimic the color of adult blue crabs, a known predator of juvenile crabs, perhaps acting as a deterrent. The presence of the predatory blue crab may require one parent to perform deterrent displays, promoting female care in this mating system.     

Monogamous Mating System and Spawning Cycle in the Gobiid Fish, Amblygobius phalaena (Gobiidae)

Reproductive ecology and mating system of the gobiid fish, Amblygobius phalaena, were studied on the coral reef at Sesoko Island, Okinawa, Japan. This goby usually lives in pairs, and maintains territories with several burrows for shelter and spawning. Although a few paired individuals changed partners, most pairs remained together over successive rounds of reproduction. Mate guarding by females appeared to prevent males from mating with other females. Spawnings were synchronous with semilunar periods. Several expected spawnings failed to occur (12%). These may have been caused by the delays in spawning preparation of the paired females or by the disturbance caused by a typhoon. A pair spawned in one of the several burrows within their home ranges. Eggs were deposited on the ceiling of the burrow, and were tended by the paired male for 3–4 days until embryos hatched. The males tended eggs at the expense of their feeding. Aggression toward fishes approaching their burrows were exhibited by the males as well as the females. Because of its low frequency in females, this behavior did not limit their ability to feed.     

Trilobites and minute ovoid pellets in a burrow (Ordovician,llanvirnian, Czech Republic)

Old collections from the Ordovician of the Barrandian area (Czech Republic) have yielded a probable vertical burrow that contains several dozen specimens of the pliomerid trilobite Placo‐paria (Placoparia) cambriensis Hicks, some of them enrolled. The fill of the burrow (classified as ?Skolithos isp.) also contains minute ovoid pellets corresponding to the ichnotaxon Tomaculum cf. problematicum Groom. The enrollment of a portion of the trilobite exoskeletons might reflect a defensive response. The pellets may be interpreted as fecal material or, less probably, as eggs. The exoskeletons and the pellets might have been mechanically trapped into the open burrow, or their concentration may have resulted from biological relationships between the tracemaker and the trilobites.     

Egg production of the burrowing shrimp Callichirus seilacheri (Bott 1955) (Decapoda, Callianassidae) in northern Chile

The thalassinidean shrimp Callichirus seilacheri is a common species in the intertidal zone of the South American Pacific coast. However, our knowledge of its reproductive ecology is rather limited. The present study was carried out between January and December 2003 at Las Machas, northern Chile. Although ovigerous females were encountered almost throughout the study period, they were particularly abundant between May and September when water temperatures were lowest and sediment coverage of the burrow entrances was highest. Females of C. seilacheri produced numerous (17,450 ± 3,796 eggs) and small (0.884 ± 0.080 mm; 0.262 ± 0.054 mm³) eggs when compared to other thalassinidean shrimps for which such information is available. Fecundity was positively correlated with female size; however, correlations were allometric, which might be related to the elasticity of the abdomen. Egg volume increased by 41.2% during embryogenesis, and egg loss during the incubation period was on average 8%. Females inverted on average 14.9% of their dry weight into egg production.     

Seismic signal transmission between burrows of the Cape mole-rat,Georychus capensis

Summary Both seismic and auditory signals were tested for their propagation characteristics in a field study of the Cape mole-rat (Georychus capensis), a subterranean rodent in the family Bathyergidae. This solitary animal is entirely fossorial and apparently communicates with its conspecifics by alternately drumming its hind legs on the burrow floor. Signal production in this species is sexually dimorphic, and mate attraction is likely mediated primarily by seismic signalling between individuals in neighboring burrows. Measurements within, and at various distances away from, natural burrows suggest that seismic signals propagate at least an order of magnitude better than auditory signals. Moreover, using a mechanical thumper which could be triggered from a tape recording of the mole-rat's seismic signals, we established that the vertically-polarized surface wave (Rayleigh wave) propagates with less attenuation than either of the two horizontally-polarized waves. Thus, we tentatively hypothesize that Rayleigh waves subserve intraspecific communication in this species.Abbreviations PPM pulses per min - SB simulated burrow - SD standard deviation - SPL sound pressure level     

Parasite fauna of native and non‐native populations of Neogobius melanostomus (Pallas, 1814) (Gobiidae) in the longitudinal profile of the Danube River

Parasite fauna of round goby Neogobius melanostomus (Pallas, 1814) in the Danube River was investigated in both its native range (two sites in the Bulgarian stretch of the Danube) and non‐native range of distribution (Croatian, Slovak and Austrian stretches) during 2005 and 2006. The aim was to identify possible changes in parasite communities associated with the introduction of a host into the new environment. A total of 29 metazoan parasite species were found to parasitize round goby in the Danube River; twelve of these parasite species were found in both the native and non‐native range of distribution. Introduction of a novel parasite species to the non‐native range via the round goby was not found. Eight parasite species occurred only in the native range and nine species only in the non‐native range of the round goby distribution. Losses of native parasite species in non‐native round goby populations and/or acquiring of novel parasite species in a new environment were not significant. Thirteen parasite taxa were recorded for the first time in round gobies. Three parasite taxa (Diplostomum spp., Pomphorhynchus laevis and Raphidascaris acus) were found in high prevalence and abundance at each sampling site in both the native and non‐native range. Parasite species diversity was assessed for each sampling site and season using three diversity indices (the Shannon, Simpson and Equitability indices), with the highest same‐season values found in a non‐native site in Slovakia (1.38, 0.69 and 0.60, respectively) and the lowest in a native site in Bulgaria (0.28, 0.12 and 0.14, respectively). Species diversity was higher in both non‐native round goby populations (Slovak and Austrian) compared to native Bulgarian populations. However, diversity indices values varied among almost all sampling sites.     

Seismic communication between the burrows of kangaroo rats, Dipodomys spectabilis

Banner-tailed kangaroo rats, Dipodomys spectabilis, footdrum to produce substrate-borne and airborne acoustic energy. Previous studies show that they communicate territorial ownership via airborne footdrumming signals. The research reported here used simulated footdrum patterns generated by an artificial `thumper' to address the question of whether kangaroo rats communicate through seismic components of these acoustic signals. With microphones suspended in sealed burrows, we found that airborne sounds were attenuated by approximately 40 dB as they passed through the burrow wall into the burrow chamber. The substrate-borne vibrations from the thumper yielded sound approximately 40 dB greater in peak amplitude than the attenuated airborne sound. Thus, 99.9% of the peak power of the thumper was transmitted directly through the substrate into the burrow. The rats in sealed burrows timed their responses to playbacks of footdrums from the thumper and a loudspeaker so they did not initiate a drumming sequence during either the seismic or airborne signals. When these signals were masked by loud noise, the rats continued to drum to the seismic signal but drummed randomly during the airborne playback. These results suggest that the sealed burrow provides a quiet place in which D. spectabilis can listen for substrate-borne communications from conspecifics. Accepted: 13 May 1997     

Organic matter distribution in burrows of the thalassinid crustacean Callichirus laurae,Gulf of Aqaba (Red Sea)

Analysis of sediments in and around the large and deep burrows made by the callianassid shrimp Callichirus laurae, Jordan Gulf of Aqaba (Red Sea), showed that organic carbon (OC), humic matter (HM), humic (HA) and fulvic acids (FA) are redistributed both quantitatively and qualitatively by sediment reworking activities:

Additive genetic variation,but not temperature,influences warning signal expression in Amata nigriceps moths (Lepidoptera: Arctiinae)

Many aposematic species show variation in their color patterns even though selection by predators is expected to stabilize warning signals toward a common phenotype. Warning signal variability can be explained by trade‐offs with other functions of coloration, such as thermoregulation, that may constrain warning signal expression by favoring darker individuals. Here, we investigated the effect of temperature on warning signal expression in aposematic Amata nigriceps moths that vary in their black and orange wing patterns. We sampled moths from two flight seasons that differed in the environmental temperatures and also reared different families under controlled conditions at three different temperatures. Against our prediction that lower developmental temperatures would reduce the warning signal size of the adult moths, we found no effect of temperature on warning signal expression in either wild or laboratory‐reared moths. Instead, we found sex‐ and population‐level differences in wing patterns. Our rearing experiment indicated that ~70% of the variability in the trait is genetic but understanding what signaling and non‐signaling functions of wing coloration maintain the genetic variation requires further work. Our results emphasize the importance of considering both genetic and plastic components of warning signal expression when studying intraspecific variation in aposematic species.     

Hidden burrow plugs and their function in the tiger beetle, Cosmodela batesi (Coleoptera, Cicindelidae)

Tiger beetle larvae excavate and live in underground burrows, whose openings they sometimes plug with soil. This study documents the burrow plugging behavior of the tiger beetle, Cosmodela batesi (Fleutiaux), in the field. We also tested the function of burrow plugs in the laboratory. In the field, C. batesi more frequently made a plug when it rained. Most larvae made plugs inside their burrows (rather than at the soil surface), and the use of an endoscope was necessary to detect these sub-surface plugs. In the laboratory, flooding was simulated by artificially introducing water into specially-made arenas. Water filled the entire burrow when there was no plug, whereas plugged burrows maintained air chambers inside. When a plug was broken with a wire, burrows filled up with water. The burrowing and plugging behavior described in this study is likely an important adaptation of C. batesi to its habitat.