Nest association in fishes: evidence for mutualism

Nest association, or the habit of spawning in the nest of another species, is a widespread reproductive strategy among North American fishes, especially minnows. The hosts of nest associates include sunfishes and nest-building minnows. Previous experiments demonstrated that nest associates benefited from the parental care of hosts. The purpose of this study was to determine the effect of nest associates on hosts, and thereby characterize the relationship as parasitic, mutualistic or commensal. Since hosts with associates had higher reproductive success than those without associates in this experiment, I characterize nest association as a mutualistic relationship, at least in some systems. The dilution effect is the proposed mechanism responsible for the higher reproductive success of hosts with associates.     

Ant gardens: interspecific recognition in parabiotic ant species

 In French Guiana, parabiotic societies (natural mixed colonies) are frequently found in ant gardens. Crematogaster limata parabiotica (Myrmicinae), often associated with Camponotus femoratus (Formicinae), was found for the first time in parabiosis with ponerine ants: Pachycondyla goeldii and Odontomachus mayi. A detailed study of the relationships between Cr. l. parabiotica and O. mayi showed that each species is aggressive towards allospecific or conspecific individuals belonging to another colony, but tolerates allospecific individuals from the multi-species society. Studies of cuticular substances of the four ant species were made using gas chromatography. The results showed that each species, living alone or in parabiosis, possesses a specific chemical profile. Thus, the ants are able to recognise nestmate and non-nestmate individuals of the associated species, even though their cuticular profiles are different. The hypothesis that the nestmate allospecific profile is learned is suggested to explain this pattern of recognition. Received: 5 June 1996 / Accepted after revision: 17 October 1996     

Metabolic cold adaptation in Antarctic fishes: evidence from enzymatic activities of brain

To evaluate the concept of metabolic cold adaptation (MCA) in fishes, we compared - in brain, red muscle, and white muscle of Antarctic notothenioid fishes and tropical/subtropical fishes - the activities of two enzymes of ATP-generating pathways, citrate synthase (CS), an indicator of citric acid cycle activity (aerobic metabolism), and lactate dehydrogenase (LDH), an indicator of potential for ATP production through anaerobic glycolysis. Brain was chosen because, unlike locomotory muscle, its metabolic activity is not likely to be influenced by a species' level of activity or nutritional status, so MCA should be readily observed if present. CS and LDH activities in brain exhibited a high level of MCA, but compensation to temperature was not complete (48% for CS; 46% for LDH). CS and LDH activities in red and white muscle varied widely among species, according to the general level of locomotory activity. The 'mode of life'-related enzymatic activities in locomotory muscle show that study of MCA at the level of whole organism metabolism is fraught with difficulties and experimental ambiguities. In contrast, the low variation among species within each group in enzymatic activities in brain, and the large differences between groups in CS and LDH activity, show that brain is an excellent study system for evaluating metabolic compensation to temperature.     

Individual recognition of song repertoires in two wood warblers

We examined recognition of individual neighbors in hooded warblers (Wilsonia citrina) and Kentucky warblers (Oporornis formosus) by comparing responses to playbacks of neighbors' songs from appropriate and inappropriate boundaries of the subjects' territories. Kentucky warblers, which sing one song type each, responded more strongly to the songs of neighbors broadcast from incorrect than from correct boundaries. Hooded warblers use their repertoires of three to nine song types in two distinct modes of singing: repeat mode involves repeated presentation of one song type; mixed mode involves presentation of two or more song types in irregular sequence. Playbacks of neighbors' mixed-mode songs from appropriate and inappropriate boundaries indicated capabilities of individual recognition similar to those reported previously for repeat-mode songs (Godard 1991) and to those of Kentucky warblers. Repertoires of moderate size, therefore, have no pronounced influence on eventual recognition of individual neighbors in hooded warblers.     

Testing hypotheses of exponential growth and size-dependent molting rate in two copepod species

Individual copepodids from nature of the lipidstoring, monocyclic, diapausing Calanus glacialis, and from nature and laboratory culture of the non-storing, polycyclic Eurytemora herdmani (both collected new Halifax, Nova Scotia in 1987 and 1988) were reared in excess food at ca. 3 and 10 °C. Soon after molts, prosome lengths and weights [total dry wt (TW) of E. herdmani; structural wt (SW) and estimated oil-sac wt (OSW) of C. glacialis] were measured. Stage durations were close to published temperature-dependent predictions; C. glacialis (almost all females) did not enter resting stages. Growth of body length was linear and of body weight (TW or SW) was exponential, with no sexual difference in E. herdmani. There were mixed effects of sizes on stage durations of individuals: weakly positive at 3 °C (but not significant at ca. 10°C) for length and SW of C. glacialis and generally weakly negative for E. herdmani, except for TW at 10 °C. Body condition (residuals of log SW vs log length) of C. glacialis at ca. 10°C, unlike length or SW, was negatively related to stage duration. There was no relationship between length-corrected SW and OSW in C. glacialis. The various results appear to suggest that health was more important than allometric constraints on growth rates of individuals. However, copepods reared at temperatures very different from those previously experienced may show long-term adjustments of size, whereas development rates respond immediately. If so, only the weakly positive effects on stage durations of length of C. glacialis at ca. 3 °C and of TW of E. herdmani at ca. 10 °C may illustrate expectations under stable temperatures and adequate food in nature.     

Sensory complementation and the acquisition of predator recognition by salmonid fishes

Following disturbance, some aquatic prey species release chemicals that act as a warning cue and increase vigilance in nearby conspecifics. Such disturbance cues evoke consistent low intensity anti-predator responses. In contrast, alarm cues from injured conspecifics often evoke stronger intensity responses in prey animals. In this study, we test the sensory complement hypothesis, which suggests that multiple cues act in an additive or synergistic fashion to provide additional information for risk assessment by prey. In the first experiment, we showed that juvenile rainbow trout pre-exposed to disturbance cues respond to a given concentration of damage-released alarm cues with a higher intensity of response than the trout that were pre-exposed to cues from undisturbed conspecifics. The two cues acted in an additive fashion. In the second experiment, we demonstrated that disturbance cues alone were not enough to elicit a conditioned response to the odour of a novel predator. We also showed that while disturbance cues elicit an increase in the response of trout to alarm cues, this increase does not translate into a stronger learned response to the predator when the predator odour is paired with alarm cues. Future studies should take into account sensory complementation to avoid underestimating the responses of prey to predators.     

Individual responsiveness to shock and colony-level aggression in honey bees: evidence for a genetic component

The phenotype of the social group is related to phenotypes of individuals that form that society. We examined how honey bee colony aggressiveness relates to individual response of male drones and foraging workers. Although the natural focus in colony aggression has been on the worker caste, the sterile females engaged in colony maintenance and defense, males carry the same genes. We measured aggressiveness scores of colonies and examined components of individual aggressive behavior in workers and haploid sons of workers from the same colony. We describe for the first time, that males, although they have no stinger, do bend their abdomen (abdominal flexion) in a posture similar to stinging behavior of workers in response to electric shock. Individual worker sting response and movement rates in response to shock were significantly correlated with colony scores. In the case of drones, sons of workers from the same colonies, abdominal flexion significantly correlated but their movement rates did not correlate with colony aggressiveness. Furthermore, the number of workers responding at increasing levels of voltage exhibits a threshold-like response, whereas the drones respond in increasing proportion to shock. We conclude that there are common and caste-specific components to aggressive behavior in honey bees. We discuss implications of these results on social and behavioral regulation and genetics of aggressive response.     

Differences in olfactory species recognition in the females of two Australian songbird species

Although birds have recently been shown to possess olfactory abilities and to use chemical cues in communication, limited effort has been made to demonstrate the use of odorants in social contexts. Even less is known regarding the use of odorants in species recognition. The ability to recognize conspecifics should be more pronounced in social species. This study investigated the importance of olfactory cues in species recognition in females of two estrildid finch species with different levels of sociality. Combining odor preference tests with chemical analyses, we surveyed whether female zebra finches and diamond firetails are able to distinguish between the species based on volatile traits and whether individuals exhibit species-specific differences in body odorants. Zebra finches are more social than diamond firetails; nevertheless, both species have an overlapping distribution area. Applying an experimental Y-maze paradigm, we showed that zebra finches can use differences in their species odor fingerprints and displayed a significant preference for the odor of conspecifics over that of heterospecifics, whereas diamond firetails did not reveal a preference. Using gas chromatography and mass spectrometry, we demonstrated that body odorants of the two species were significantly different in relative composition. This finding demonstrates the potential importance of olfactory cues in species recognition, at least in social bird species. Even these two closely related species displayed remarkable differences in their responsiveness to similar chemical cues, which might be caused by species-specific differences in ecology, physiology, or evolution.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Karyotype differentiation between two species of carangid fishes,genusTrachurus (Perciformes: Carangidae)

A karyological study ofTrachurus trachurus andT. mediterraneus (Perciformes: Carangidae) was conducted by standard, fluorochrome staining (CMA₃, mithramycin, quinacrine mustard, DAPI), C-, Ag-NOR, and Alu-I banding methods. The karyotypes of both species consisted of 2n = 48 chromosomes, but of different FN:T. trachurus possessed a chromosome complement of 2 metacentric and 46 acrocentric elements, fundamental number (FN) = 50 andT. mediterraneus, a chromosome complement of 4 metacentric, 4 submetacentric, 14 subtelocentric and 26 acrocentric chromosomes, FN = 70. In neither of the two taxa investigated were heteromorphic sex chromosomes observed. The nucleolar organizer region was interstitially located on the long arm of the Ist pair of chromosomes in both species, intermediate inT. mediterraneus and subterminal inT. trachurus. Constitutive heterochromatin was found in nearly all centromeric and telomeric regions inT. trachurus; inT. mediterraneus it formed less intense telomeric and centromeric bands and thin interstitial bands on eight chromosome pairs. In addition, the C-positive material reacted differently to the digestion with endonuclease Alu-l in the two species. The results are discussed and compared with karyological data known for other species of Carangidae.     

Habitat distribution and species diversity of chaetodontid and pomacentrid fishes near Bimini,Bahamas

The relative abundance of chaetodontid and pomacentrid fishes in 8 habitats, ranging in depth from 0.5 to 17 m, was determined by visual counting of free-swimming fish in the field. Habitat widths and overlaps were determined using weighting factors, developed by Colwell and Futuyma (1971), which account for the different degree of uniqueness of each habitat. The average habitat widths and the average habitat overlaps were greater for the chaetodontids than the pomacentrids. This was due, in part, to the greater use of habitat structure by the pomacentrids, and it is hypothesized that this is because they are the more vulnerable to predation. The juveniles and adults reached peak abundance in different habitats for at least some of the species, and this may further reduce the habitat-overlap values for adults of some pairs of species. It is clear, however, that the ability of the species in these families to coexist is largely due to factors operating within habitats rather than to a large degree of habitat specialization.     

Circadian rhythms in the respiration of eight species of cardinal fishes (Pisces: Apogonidae): comparative analysis and adaptive significance

A comparative study was made of the entrained (12:12 LD) and endogenous (DD) respiratory rhythms of 8 species of cardinal fishes. Volumetric respirometers, designed specifically for long-term measurements of oxygen uptake by small fishes under controlled conditions, were used during standard 5 day experiments. Despite some individual variation, certain species-specific and genus-specific patterns of respiration emerged. Each species showed a persistant (advancing) rhythm of oxygen consumption during 3 days of DD. Light appeared to be the Zeitgeber, with the light-dark transition phasing the nocturnal peaks and the dark-light transition possibly phasing the diurnal ones. Genus-specific and species-specific trends in field behavior, habitat selection, and distribution were studied. There were correlations between the field behavior and the entrained respiratory rhythms of these nocturnally active fishes. Entrained and endogenous rhythms of each species showed characteristic similarities and were interpreted as integral components of a level of organization evolved for nocturnal life in the shallow marine environment.     

Life-history evidence for sibling species in Axiothella rubrocincta (Polychaeta: Maldanidae)

Individual Axiothella rubrocincta Johnson on the Californian coast live in U-shaped tubes, feed on the surface of the sediment, attain densities of 100 m^-2, attain lengths of 140 mm, have maximum oocyte diameters of 220 μm and produce demersal larvae. Individuals from Washington populations live in vertical tubes, feed 10–15 cm below the sediment surface, reach densities of 5000 m^-2, attain lengths of 60 mm, have maximum oocyte diameters of 385 μm and brood their young in their tubes to at least the 7-setiger stage. Individuals from California have a scalloped posterior margin of the cephalic plate, while individuals from Washington have entire margins. There are no other consistent morphological differences between populations. The population from Eagle Cove, Washington has individuals similar in size and oocyte diameter to individuals from California, but which maintain vertical tubes. A. rubrocincta represents a sibling species complex.     

A comparison of absorption and assimilation efficiencies between four species of shallow- and deep-living fishes

We captured two species of deep-sea zoarcids, Melanostigma pammelas and Lycodapus mandibularis, from Monterey Bay California and maintained them in the laboratory. One shallow-water zoarcid, Eucryphycus californicus, and an ecologically and morphologically similar stichaeid fish Xiphister atropurpureus were collected from intertidal and subtidal habitats in Monterey Bay. We investigated the absorption and assimilation efficiencies of these fishes to determine whether deep-sea species have evolved mechanisms to increase their efficiency of food use. Fishes were placed in experimental chambers and fed a known quantity of food. Ammonia excretion was measured and feces were collected daily. Both food and feces were analyzed for water, protein, lipid and ash to determine specific absorption efficiencies. Absorption ranged from 87.7 to 92.4% and assimilation efficiencies from 84.0 to 86.5%. Meal sizes from 0.5 to 4.0% of body weight did not affect the results. No significant differences were found between deep-sea and shallow-water species fed single meals or fed ad libitum for 10 days. This suggests that the selective pressure to maximize absorption and assimilation is universal and is not affected by the productivity of the habitat occupied. However, the relative size of the intestine in the deep-sea species was significantly smaller suggesting that they had a lower metabolic cost to maintain their digestive apparatus. It could not be concluded whether this was the result of pressure to conserve energy or rather the tendency of the shallow-living species to ingest more refractory material (i.e. sediment, algae).     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

Immunocytochemical evidence that symbiotic algae secrete potential recognition signal molecules in hospite

Cnidarian–dinoflagellate symbioses are not well understood at the molecular level. Observed specificity between partners during initiation, establishment, and maintenance of the relationship strongly implies a role for chemical signaling. This report presents biochemical and immunocytochemical evidence for potential signaling molecules, as large molecular weight glycoproteins, secreted by Symbiodinium dinoflagellates both in culture and in symbiosis. Polyclonal antibodies directed against recovered exudate from S. microadriaticum, the natural endosymbiont of Cassiopea xamachana, the upside–down jellyfish, were highly specific in recognizing exudates from Symbiodinium species that can successfully induce developmental metamorphosis in the host but did not recognize exudates from Symbiodinium species that do not. Immunoblot analyses showed S. microadriaticum exudate to be protease sensitive. Release of antigenic material by symbiotic S. microadriaticum was demonstrated through light and electron microscopy using immunogold-labeled anti-S. microadriaticum (anti-Sm-XuLg) antibodies as probes. These secreted, symbiont-derived glycoconjugates may be candidates for interspecific molecular signals.     

Predator biomass,prey density,and species composition effects on group size in recruit coral reef fishes

Group incidence and size are described for recruit parrotfishes, wrasses, and damselfishes on Hawaiian reefs over 3 years (2006–2008) at sites spanning the archipelago (20–28°N, 155–177°W). Coral-poor and coral-rich areas were surveyed at sites with both low (Hawaii Island) and high (Midway Atoll) predator densities, facilitating examination of relations among predator and recruit densities, habitat, and group metrics. Predator and recruit densities varied spatially and temporally, with a sixfold range in total recruit densities among years. Group (≥2 recruits) metrics varied with time and tracked predator and recruit densities and the proportion of schooling species. Groups often included heterospecifics whose proportion increased with group size. A non-saturating relationship between group size and recruit density suggests that the anti-predator benefits of aggregation exceeded competitive costs. Grouping behavior may have overarching importance for recruit survival–even at high recruit densities–and merits further study on Hawaiian reefs and elsewhere.     

Species recognition and species isolation in wandering spiders (Cupiennius spp.; Ctenidae)

Summary One of the roles ascribed to spider courtship is that of mate recognition and reproductive isolation. We describe behavioral mechanisms underlying these functions in three species of wandering spiders of the genus Cupiennius (Ctenidae). C. salei and the two sympatric species C. getazi and C. coccineus discriminate conspecifics and heterospecifics in a multistage process which covers three principal phases of courtship: (1) Chemical phase: male spiders initiate vibratory courtship communication less often upon contact with heterospecific than with conspecific female silk. (2) Vibratory phase: females respond with their own vibration less often and with increased delay to the vibratory courtship signals of heterospecific males than to those of conspecific males. (3) Tactile phase: in heterospecific pairings males and females interrupt the contact phase if their courtship has progressed thus far. The product of the probabilities of passing through each of the different stages of courtship is between 0.44 and 0.88 for conspecific pairings, but zero for most heterospecific pairings. This is to a greater extent due to the behavioral barriers of the vibratory and the tactile phase than to that of the chemical phase. Male taxis on tracks of female silk does not depend on the species-specificity of the silk. Female vibrations are not essential for species recognition: males increase their signal rates, orient toward and approach responding females in both con- and heterospecific pairings. Female attacks on males are similar to attacks on prey; males defend themselves with outstretched front legs and are caught and killed by the female in only 5% of the cases. In all phases of courtship, female C. salei are least and female C. getazi most selective. When female C. coccineus and C. getazi are involved in heterospecific pairings no copulation is attempted. In contrast, 13 copulations of C. salei with males of the other two species were observed; egg sacs never resulted from these copulations. Among males, those of C. getazi were least selective. The likelihood of heterospecific pairs passing through all phases of courtship is higher in case of the allopatric than in that of the sympatric species. This finding applies to the courtship as a whole, but not necessarily to each of its phases.Offprint requests to: F.G. Barth     

On the evidence for species coexistence: a critique of the coexistence program

A major challenge in ecology is to understand how the millions of species on Earth are organized into biological communities. Mechanisms promoting coexistence are one such class of organizing processes, which allow multiple species to persist in the same trophic level of a given web of species interactions. If some mechanism promotes the coexistence of two or more species, each species must be able to increase when it is rare and the others are at their typical abundances; this invasibility criterion is fundamental evidence for species coexistence regardless of the mechanism. In an attempt to evaluate the level of empirical support for coexistence mechanisms in nature, we surveyed the literature for empirical studies of coexistence at a local scale (i.e., species found living together in one place) to determine whether these studies satisfied the invasibility criterion. In our survey, only seven of 323 studies that drew conclusions about species coexistence evaluated invasibility in some way in either observational or experimental studies. In addition, only three other studies evaluated necessary but not sufficient conditions for invasibility (i.e., negative density dependence and a trade-off in performance that influences population regulation). These results indicate that, while species coexistence is a prevalent assumption for why species are able to live together in one place, critical empirical tests of this fundamental assumption of community structure are rarely performed. These tests are central to developing a more robust understanding of the relative contributions of both deterministic and stochastic processes structuring biological communities.