Effects of Female‐Female Aggression in a Sexual/Unisexual Species Complex

The maintenance of sexual reproduction is still a largely unresolved question in evolutionary biology, and one of the most puzzling aspects of this is the coexistence of sexual and asexual females. This often leads to widespread niche overlap, posing an interesting challenge to competitive exclusion theory. In this study, we investigate how the aggressive behaviors between females in a sexual/unisexual mating complex of mollies (Poecilia latipinna and P. formosa) differ, and the effects these behaviors have on both the performer and receiver of aggressive acts. We exposed females to five treatments: (1) alone (control); (2) a large and small sexual female; (3) a large sexual female and a small unisexual female; (4) a small sexual female and a large unisexual female; and (5) a large and small unisexual female. We found that P. formosa females were on average more aggressive over time when compared to their sexual hosts, P. latipinna. The females of P. formosa, however, incurred a higher cost of being aggressive. Large, aggressor females of this species had a lower body fat content than their conspecific recipients. By contrast, P. latipinna females showed no differentiation between aggressors and recipients. Here, we combine life history methodology and behavioral observations to show that female competition can in fact be quantitatively measured and that these aggressive behaviors are costly to perform. The adaptive value of this interspecies aggression is not yet completely understood, but sexual competition, access to resources, and position within the shoal are all possible functions.     

Comparative Analysis of Feeding and Schooling Behaviour of the Cyprinidae Alburnus alburnus (L., 1758), Rutilus rutilus (L., 1758), and Scardinius erythrophthalmus (L., 1758) in a Backwater of the Danube near Vienna

Underwater observations of the three common fish species Alburnus alburnus (L., 1758), Rutilus rutilus (L., 1758) and Scardinius erythrophthalmus (L., 1758) in a small part of the Danube flood area near Vienna have been made from June to October 1985 and 1986. Particular emphasis has been laid on the partitioning of feeding resources and habitat. It has been established that these species prefer different water zones. Alburnus a. and Scardinius e. are “obligate” schoolers (BREDER, 1967) whereas Rutilus r. is a facultative schooler. While feeding Rutilus r. and Scardinius e. form “aggregations” (BREDER, 1959) and both adopt equivalent feeding techniques. Furthermore Alburnus a. is a quick moving species living near the surface and chasing mostly in schools. While inter- and intraspecific aggression is rarely observed for Rutilus r. and Scardinius e., Alburnus a. often show interspecific actions.     

Sequence requirements of the hammerhead RNA self-cleavage reaction.

A previously well-characterized hammerhead catalytic RNA consisting of a 24-nucleotide substrate and a 19-nucleotide ribozyme was used to perform an extensive mutagenesis study. The cleavage rates of 21 different substrate mutations and 24 different ribozyme mutations were determined. Only one of the three phylogenetically conserved base pairs but all nine of the conserved single-stranded residues in the central core are needed for self cleavage. In most cases the mutations did not alter the ability of the hammerhead to assemble into a bimolecular complex. In the few cases where mutant hammerheads did not assemble, it appeared to be the result of the mutation stabilizing an alternate substrate or ribozyme secondary structure. All combinations of mutant substrate and mutant ribozyme were less active than the corresponding single mutations, suggesting that the hammerhead contains few, if any, replaceable tertiary interactions as are found in tRNA. The refined consensus hammerhead resulting from this work was used to identify potential hammerheads present in a variety of Escherichia coli gene sequences.     

Kinetic characterization of two I/II format hammerhead ribozymes.

Five new hammerhead ribozymes were designed that assemble through the formation of helices I and II (I/II format) instead of the more standard assembly through helices I and III (I/III format). The substrate binding and cleavage properties of such hammerheads could potentially be different due to the absence of loop II and the requirement for the entire catalytic core to assemble. Two I/II format hammerheads, HHalpha1 and HHalpha5, which show structural homogeneity on native gels, were characterized kinetically. The association rate constants of both I/II hammerheads are unusually slow compared to the rate of RNA duplex formation. The dissociation rate constants indicate that the hammerhead core destabilizes an uninterrupted RNA helix somewhat less than was observed for I/III hammerheads. Whereas the cleavage rate constant of HHalpha5 is similar to that observed for I/III hammerheads, HHalpha1 cleaves 10-fold faster than any hammerhead previously reported. The temperature and pH dependence of the cleavage rate constant of HHalpha1 are similar to those reported for I/III hammerheads, suggesting a similar mechanism of cleavage.     

Shallow water tidal flat use and associated specialized foraging behavior of the great hammerhead shark (Sphyrna mokarran)

Evidence suggests the great hammerhead shark, Sphyrna mokarran, is vulnerable to a variety of anthropogenic stressors, and is an understudied species of shark due to its cryptic nature and wide-ranging movements. While recognized as both a pelagic-coastal and a highly mobile predator, minimal anecdotal evidence exist describing shallow water habitat use by this species. This report describes six cases in which a great hammerhead shark utilizes an inshore shallow water flats environment (<1.5?m in depth), five of which involve prey capture. These observations permitted identification of two novel behaviors that may allow great hammerheads to inhabit these shallow habitats: a (1) prey-capture technique termed ‘grasp-turning’ that involves burst swimming at tight turning angles while grasping prey and (2) a post-predation recovery period whereby the shark maintains head-first orientation into the current that may facilitate respiration and prey consumption. These behavioral observations provide insights into the natural history of this species.     

Population Differences in the Schooling Behaviour of Newborn Guppies,Poecilia reticulata

Population differences in the schooling behaviour of adult guppies (Poecilia reticulata) from Trinidad have previously been well documented. This paper demonstrates that variation in schooling behaviour is also present in newborn fish. Guppies from the Aripo River of Trinidad display well-developed schooling behaviour from birth. In addition to forming cohesive and polarized groups, these newborn fish also exhibit predator inspection behaviour. Newborn guppies from the Paria River perform inspections but, unlike the Aripo fish, are poor schoolers. Population variation in behaviour at birth does not necessarily reflect the pattern observed in the adult fish. As adults, guppies from the Upper section of the Aripo River have a much lower schooling tendency and are less wary during inspection than the downstream Lower Aripo guppies. At birth, however, the schooling behaviour and inspection behaviour in the two populations is identical. These results suggest that different selective pressures operate on adult and newborn guppies in Trinidad.     

Historical records reveal potential extirpation of four hammerhead sharks (Sphyrna spp.) in Mexican Pacific waters

Populations of hammerhead sharks (Sphyrna and Eusphyra) have declined in many regions of the world. Six of the eight hammerheads known to date are distributed in the Mexican Pacific: S. corona, S. lewini, S. media, S. mokarran, S. tiburo and S. zygaena. These species, with exception of S. corona, were abundant in the Gulf of California in 1960s. I analyze records from fishery-dependent and fishery-independent surveys, and records from ichthyological collections to determine the presence and frequency of hammerheads in the Mexican Pacific. The most frequent hammerheads in fishery-dependent and fishery-independent surveys were S. lewini and S. zygaena. It appears that S. media, S. mokarran and S. tiburo might have been extirpated from the Gulf of California. In the last two decades, records of S. mokarran (n = 61) were restricted to Central and Southern Mexican Pacific, and records of S. tiburo (n = 3) and S. media (n = 3) were restricted to the Southern region. Given the continued fishing pressure, inferred declines and the probable extirpation of populations, S. tiburo and S. media should be reassessed for the IUCN red list as Endangered or Critically Endangered. Sphyrna corona should be reassessed as Endangered or Critically Endangered, because it is endemic to the Eastern Pacific and recent records have been obtained only from Colombian waters. The Endangered status of S. mokarran is confirmed for this region.     

School fidelity in fishes: The yellow perch pattern

School fidelity concerns the cohesiveness of social aggregations over time, as measured by the regularity or repeatability with which known individuals re-occur as members of a school. School fidelity was measured by divers who observed yellow perch (Perca flavescens) marked with individually recognizable tags in Cazenovia Lake, New York. Yellow perch were facultative schoolers, with individual differences in schooling tendency (measured as percentage of time in schools) accounting for different numbers of observations of schooled versus solitary fish. The greatest co-occurrence was found between fish with strong schooling tendencies, but within this group co-occurrence was random. Home ranges overlapped greatly between individuals, yet co-occurrence was more a function of schooling tendency than of overlapping home ranges. Individuals with overlapping home ranges but with weak schooling tendencies seldom co-occurred. Home range size increased directly with schooling tendency: the distances that individuals moved apparently depended on the movements of the schools they joined, with strongly schooling fish remaining in schools longer. Individuals joined and left schools frequently; this also reflects a lack of fidelity between fish. Fish schools that form primarily for predator avoidance may show higher levels of fidelity than do schools that form for foraging or hydrodynamic efficiency.     

Structure–function analysis of the ribozymes of chrysanthemum chlorotic mottle viroid: a loop–loop interaction motif conserved in most natural hammerheads

Loop–loop tertiary interactions play a key role in the folding and catalytic activity of natural hammerhead ribozymes. Using a combination of NMR spectroscopy, site-directed mutagenesis and kinetic and infectivity analyses, we have examined the structure and function of loops 1 and 2 of the (+) and (–) hammerheads of chrysanthemum chlorotic mottle viroid RNA. In both hammerheads, loop 1 is a heptanucleotide hairpin loop containing an exposed U at its 5′ side and an extrahelical U at its 3′-side critical for the catalytic activity of the ribozyme in vitro and for viroid infectivity in vivo, whereas loop 2 has a key opened A at its 3′-side. These structural features promote a specific loop–loop interaction motif across the major groove. The essential features of this tertiary structure element, base pairing between the 5′ U of loop 1 and the 3′ A of loop 2, and interaction of the extrahelical pyrimidine of loop 1 with loop 2, are likely shared by a significant fraction of natural hammerheads.     

Global-scale genetic identification of hammerhead sharks: Application to assessment of the international fin trade and law enforcement

The future status of sharks is an issue of widespread conservation concern due to declines in many species in the face of high levels of exploitation to satisfy market demands for products, especially fins. Substantial declines in the large-bodied hammerhead sharks, Sphyrna lewini, S. mokarran and S. zygaena, even in regions where some management occurs, indicate that informed conservation measures are warranted for these circumglobally distributed species. Despite the importance of assessing shark catch and trade on a species-specific basis to detect potential overexploitation of individual species, achieving this goal for hammerheads has proven elusive due to difficulties in identification of their products. Here, we present the development and application of a diagnostic, streamlined, five-primer multiplex polymerase chain reaction assay utilizing species-specific primers based on nuclear ribosomal ITS2 for the three hammerhead species throughout their global distribution. Application of this assay to investigations of the fin market confirmed the presence of hammerhead fins in the international trade. A study of the world’s largest fin market in Hong Kong revealed a high concordance between specific Chinese-name trade categories and fins from these three species (“Bai Chun” with S. lewini, “Gui Chun” with S. zygaena and “Gu Pian” with S.␣mokarran), and clear species preferences. This concordance information allows the use of market records for monitoring species-specific trends in trade and exploitation rates. The assay is also proving useful for identification of shark body parts in U.S. fisheries law-enforcement activities. Screening of morphologically identified “ S. lewini” from globally distributed areas using this assay with subsequent whole ITS2 sequencing suggests a cryptic species closely related to S. lewini occurs off the SE USA coast.     

Catalytic diversity of extended hammerhead ribozymes

Chimeras of the well-characterized minimal hammerhead 16 and nine extended hammerheads derived from natural viroids and satellite RNAs were constructed with the goal of assessing whether their very different peripheral tertiary interactions modulate their catalytic properties. For each chimera, three different assays were used to determine the rate of cleavage and the fraction of full-length hammerhead at equilibrium and thereby deduce the elemental cleavage ( k 2) and ligation ( k -2) rate constants. The nine chimeras were all more active than minimal hammerheads and exhibited a very broad range of catalytic properties, with values of k 2 varying by 750-fold and k -2 by 100-fold. At least two of the hammerheads exhibited an altered dependence of k obs on magnesium concentration. Since much less catalytic diversity is observed among minimal hammerheads that lack the tertiary interactions, a possible role for the different tertiary interaction is to modulate the hammerhead cleavage properties in viroids. For example, differing hammerhead cleavage and ligation rates could affect the steady state concentrations of linear, circular, and polymeric genomes in infected cells.     

Peripheral regions of natural hammerhead ribozymes greatly increase their self-cleavage activity

Natural hammerhead ribozymes are mostly found in some viroid and viroid-like RNAs and catalyze their cis cleavage during replication. Hammerheads have been manipulated to act in trans and assumed to have a similar catalytic behavior in this artificial context. However, we show here that two natural cis-acting hammerheads self-cleave much faster than trans-acting derivatives and other reported artificial hammerheads. Moreover, modifications of the peripheral loops 1 and 2 of one of these natural hammerheads induced a >100-fold reduction of the self-cleavage constant, whereas engineering a trans-acting artificial hammerhead into a cis derivative by introducing a loop 1 had no effect. These data show that regions external to the central conserved core of natural hammerheads play a role in catalysis, and suggest the existence of tertiary interactions between these peripheral regions. The interactions, determined by the sequence and size of loops 1 and 2 and most likely of helices I and II, must result from natural selection and should be studied in order to better understand the hammerhead requirements in vivo.     

An extra nucleotide in the consensus catalytic core of a viroid hammerhead ribozyme: implications for the design of more efficient ribozymes.

Hammerhead ribozymes catalyze self-cleavage of oligomeric RNAs generated in replication of certain viroid and viroid-like RNAs. Previous studies have defined a catalytic core conserved in most natural hammerheads, but it is still unknown why some present deviations from the consensus. We have addressed this issue in chrysanthemum chlorotic mottle viroid (CChMVd), whose (+) hammerhead has an extra A (A10) between the conserved A9 and the quasi-conserved G10.1. Effects of insertions at this position on hammerhead kinetics have not hitherto been examined. A10 caused a moderate decrease of the trans-cleaving rate constant with respect to the CChMVd (+) hammerhead without this residue, whereas A10-->C and A10-->G substitutions had major detrimental effects, likely because they favor catalytically inactive foldings. By contrast, A10-->U substitution induced a 3-4-fold increase of the rate constant, providing an explanation for the extra U10 present in two natural hammerheads. Because A10 also occupies a singular and indispensable position in the global CChMVd conformation, as revealed by bioassays, these results show that some hammerheads deviate from the consensus due to the involvement of certain residues in critical function(s) other than self-cleavage. Incorporation of the extra U10 into a model hammerhead also caused a similar increase in the rate constant, providing data for a deeper understanding of the hammerhead structural requirements and for designing more efficient ribozymes.     

Levels of diurnal predation on a school of flat-iron herring,Harengula thrissina

Synopsis The incidence of diurnal predation (0800–1600h) on schools of several hundred thousand adult flat-iron herring,Harengula thrissina, was quantified. Nine species of piscine predators perpetrated 653 attacks and 158 captures during 42.5 h of observation. Four predators: cabrilla,Mycteroperca rosacea, cornetfish,Fistularia commersonii, green jack,Caranx caballus, and black skipjack,Euthynnus lineatus, were responsible for 92% of the attacks and 96% of the successes. Among these predators attack rates (attacks × time^–1) and success rates (successes × attacks^–1) varied with respect to time of day, although all predators were active throughout the day. As a consequence, the herring school experienced fairly homogeneous attack and success rates. These rates were estimated as roughly half the crepuscular attack rates but they were sustained over a much longer period. Piscivorous predators, particularly those species frequently found in association with schooling prey, may be focusing their attacks on opportunities, not specific times of day and/or light levels. More direct information needs to be gathered on diurnal as well as crepuscular attack and success rates in order to assess the impact that piscine predators have on schooling prey.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

The biology and conservation status of the large hammerhead shark complex: the great,scalloped, and smooth hammerheads

Hammerhead sharks are among the most intriguing yet imperiled groups of large sharks globally. Until recently, our understanding of their biology, movements, diet, and life histories was challenged by a lack of studies. In recent years there has been a surge of published studies on this group of sharks, incorporating new information on age and growth, behavior, and the threats they face. Here we summarize and compare what is known on the biology and conservation of the three largest species of hammerhead sharks: the great hammerhead (Sphyrna mokarran), the scalloped hammerhead (Sphyrna lewini), and the smooth hammerhead (Sphyrna zygaena). We chose these species since they are the most well-studied of the hammerheads, and also because they are commonly captured in target and non-target fisheries worldwide. Thus, we also discuss population trends and the vulnerabilities of each species, and make recommendations for future studies on these fascinating and complex elasmobranch fishes.     

Complete RNA inverse folding: computational design of functional hammerhead ribozymes

Nanotechnology and synthetic biology currently constitute one of the most innovative, interdisciplinary fields of research, poised to radically transform society in the 21st century. This paper concerns the synthetic design of ribonucleic acid molecules, using our recent algorithm, RNAiFold, which can determine all RNA sequences whose minimum free energy secondary structure is a user-specified target structure. Using RNAiFold, we design ten cis-cleaving hammerhead ribozymes, all of which are shown to be functional by a cleavage assay. We additionally use RNAiFold to design a functional cis-cleaving hammerhead as a modular unit of a synthetic larger RNA. Analysis of kinetics on this small set of hammerheads suggests that cleavage rate of computationally designed ribozymes may be correlated with positional entropy, ensemble defect, structural flexibility/rigidity and related measures. Artificial ribozymes have been designed in the past either manually or by SELEX (Systematic Evolution of Ligands by Exponential Enrichment); however, this appears to be the first purely computational design and experimental validation of novel functional ribozymes. RNAiFold is available at http://bioinformatics.bc.edu/clotelab/RNAiFold/.     

Vulnerability to longline fisheries of three hammerhead shark Sphyrna species in the south‐western and equatorial Atlantic Ocean

Catch and effort data from 29 418 longline sets from Brazilian tuna longline vessels operating in the south‐western and equatorial Atlantic Ocean between 2004 and 2011 were analysed to investigate the distribution, catch rate and size of three species of hammerhead sharks (Sphyrna lewini, Sphyrna mokarran and Sphyrna zygaena). During that period, 6172 hammerhead sharks were caught. Among the elasmobranchs, the highest percentage of hammerhead sharks were caught in 2007, when they accounted for 3·90% of the group, while the lowest value of 0·40% was recorded in 2010. In general, the spatial distribution of the mean catch per unit effort (CPUE) by years and quarters showed a trend of higher catches near the equatorial region and in southern Brazil. The nominal mean CPUE was 0·12 Sphyrna spp. 1000⁻¹ hooks, with the highest value being recorded in 2007 (0·30 Sphyrna spp. 1000⁻¹ hooks). The standardized yearly CPUE estimated by a generalized linear model assuming a zero inflated negative binomial (ZINB) distribution were not much different from nominal values. Of the 205 sexed specimens, 117 were females and 88 were males, resulting in a sex ratio with a predominance of females (1·30:1·00), although not statistically significant. The total length of females ranged from 1200 to 2800 mm and of males from 1100 to 3100 mm. Juvenile hammerhead sharks represented 82 and 54% of the sexed female and male specimens, respectively.     

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.