Patterns of autotomy and regeneration in Hemigrapsus nudus

Leg autotomy and regeneration can have severe impacts on survival and reproduction, and these impacts may be even more pronounced in animals with multifarious legs, such as decapods. Thus, determining the patterns and frequency of autotomy and regeneration could reveal the effects of these processes on the individual and population level. We investigated whether some legs are lost more often than others and if all legs are equally likely to be regenerated. We sampled nearly 500 purple shore crabs (Hemigrapsus nudus) and showed that (1) most animals are found with at least one injured leg, (2) the patterns of autotomy differ between males and females, and (3) successful claw regeneration is unlikely in both males and females. Future work with H. nudus and other grapsid crabs will elucidate how patterns seen here relate to other developmental and ecological factors.     

THE ONTOGENY OF ASYMMETRY IN EARWIG FORCEPS

Fluctuating asymmetry may play an important role in the evolution of naturally selected and secondary sexual traits. However, very little is known about how asymmetries arise or how organisms maintain symmetry during development. Here I propose three mutually exclusive patterns for the development of asymmetries through consecutive growth stages: (1) compensatory growth, in which growth of the shorter side is greatest at the following growth stage; (2) persistent growth, in which growth of the longer side is greatest at the following growth stage; and (3) uncorrelated growth in which growth of the following stage is unrelated to the asymmetry at the previous one. I followed the growth in the forceps of male earwigs through four successive instars. Dyar's rule was used as a null model of insect growth. In the molt from the second to third instar, asymmetries increased through uncorrelated growth and with the magnitude but not the sign expected from Dyar's rule. However, following this, at the molts between instars 3–4 and 4–5, compensatory growth maintained asymmetries at a lower level than expected from Dyar's rule. Although there was no reduction in the absolute magnitude of asymmetry, relative asymmetry did decline. The net growth of forceps length did not follow Dyar's rule. The interpretation of patterns of growth were more sensitive and informative than the interpretation of the relations between asymmetries at consecutive instars.     

Changes in the selection differential exerted on a marine snail during the ontogeny of a predatory shore crab

Empirical estimates of selection gradients caused by predators are common, yet no one has quantified how these estimates vary with predator ontogeny. We used logistic regression to investigate how selection on gastropod shell thickness changed with predator size. Only small and medium purple shore crabs (Hemigrapsus nudus) exerted a linear selection gradient for increased shell‐thickness within a single population of the intertidal snail (Littorina subrotundata). The shape of the fitness function for shell thickness was confirmed to be linear for small and medium crabs but was humped for large male crabs, suggesting no directional selection. A second experiment using two prey species to amplify shell thickness differences established that the selection differential on adult snails decreased linearly as crab size increased. We observed differences in size distribution and sex ratios among three natural shore crab populations that may cause spatial and temporal variation in predator‐mediated selection on local snail populations.     

Marine invasive species: validation of citizen science and implications for national monitoring networks

Approximately 1,000 volunteers assessed the presence of invasive (Carcinus maenas and Hemigrapsus sanguineus) and native crabs within the intertidal zone of seven coastal states of the US, from New Jersey to Maine. Identification of crab species and determination of the gender of the observed crabs was documented at all 52 sites across a 725-km coastal transect. Using quantitative measures of accuracy of data collected by citizen scientists, a significant predictor of a volunteer’s ability was determined and eligibility criteria were set. Students in grade three and seven had the ability to differentiate between species of crabs with over 80% and 95% accuracy, respectively. Determination of gender of the crabs was more challenging and accuracy exceeded 80% for seventh grade students, while 95% accuracy was found for students with at least 2 years of university education. We used the data collected by citizen scientists to create a large-scale standardized database of the distribution and abundance of the native and invasive crabs. Hemigrapsus sanguineus dominated the rocky intertidal zone from Sandy Hook, New Jersey to Boston Harbor, Massachusetts while C. maenas dominated the northern extent of the sampled coastline. A citizen scientist of this monitoring network detected a range expansion of H. sanguineus. We identified obstacles to creating a national monitoring network and proposed recommendations that addressed these issues.     

Pattern and causes of a temperature-dependent gradient of size at terminal moult in snow crab (Chionoecetes opilio) along West Greenland

Geographic variation in size of male and female snow crabs (Chionoecetes opilio) was investigated along the west coast of Greenland to test the hypothesis that size at terminal molt (=adulthood) is temperature dependent. A total of 81,490 snow crabs were collected in small-mesh traps in Disko Bay (68–69°N) and six sites near Sisimiut (66–67°N) in May and June from 2000 to 2005. Average bottom temperature over the study period ranged from −0.8 to 3.2°C across the sampling sites. Mean carapace width was positively correlated with temperature in both sexes, a pattern often described as a converse Jame’s cline. We infer that temperature per se is the causative factor and discount season length, food availability or density as ultimate causes of the cline. Temperature effects on body size of crabs apparently result from a change in the number of instars before terminal molt. This interpretation is supported by size frequency analysis showing that in general crabs were larger at instar in a colder than in a warmer site. We briefly discuss the implications of our findings for population reproductive potential and the effectiveness of a fixed legal size limit in protecting some adult males from exploitation at different temperature regimes.     

Host specificity of <Emphasis Type="Italic">Sacculina carcini</Emphasis>, a potential biological control agent of the introduced European green crab Carcinus maenas in California

The European green crab, Carcinus maenas, is an introduced marine predator established on the west coast of North America. We conducted laboratory experiments on the host specificity of a natural enemy of the green crab, the parasitic barnacle Sacculina carcini, to provide information on the safety of its use as a possible biological control agent. Four species of non-target, native California crabs (Hemigrapsus oregonensis, H. nudus, Pachygrapsus crassipes and Cancer magister) were exposed to infective larvae of S. carcini. Settlement by S. carcini on the four native species ranged from 33 to 53%, compared to 79% for green crabs. Overall, cyprid larvae tended to settle in higher numbers on individual green crabs than on either C. magister or H. oregonensis. However, for C. magister this difference was significant for soft-shelled, but not hard-shelled individuals. Up to 29% of the native crabs arrested early infections by melanizing the rootlets of the parasite. Most native and green crabs settled on by S. carcini became infected, especially when settled on by >3 cyprids. Infected green crabs died at more than twice the rate of uninfected green crabs. In contrast to green crabs, all infected native crabs died without producing an externa (reproductive sac). At high settlement intensities, infected native crabs frequently exhibited neurological symptoms (twitching, loss of movement) before death. These results indicate that use of S. carcini as a biological control agent could result in the death of native crabs. The magnitude of this effect would be proportional to the density of infected green crabs in the environment and the probability that cyprids would contact native crabs in the wild. Potential benefits of biological control should be assessed in relation to these potential non-target effects.     

Morphological integration of fluctuating asymmetry in the mouse mandible

Morphological integration of fluctuating asymmetry (FA) was assessed for nine mandibular characters in random-bred house mice to test the hypothesis that there should be a significant integration or concordance of FAs at the population level, but not at the individual level. FA estimates for each of the nine characters were made for each of 16 subpopulations (two replicates each for mice varying in sex and age) and their correlations indicated a moderate level of integration (index of integration = 0.42). A matrix permutation test of the differences of the correlations within two (‘incisor’ and ‘muscle’) developmentally different character groups versus correlations between groups was significant, indicating the presence of morphological integration. Kendall's coefficient of concordance also indicated a significant population asymmetry parameter for FAs within the two character groups. Correlations among individual FA estimates were generally lower than those calculated from subpopulations, the index of integration being 0.21. The matrix permutation test failed to show significant morphological integration among individual FAs, but Kendall's coefficient of concordance was significant, indicating the presence of an ‘individual asymmetry parameter’ among the nine characters. Principal components analysis and canonical correlation analysis confirmed the overall higher level of integration of FAs among the subpopulations and within the two character groupings.     

Water-borne cues from a shell-crushing predator induce a more massive shell in experimental populations of an intertidal snail

Inducible defenses are important in the life strategies of many taxa. In some species of marine gastropods, water-borne chemical cues from potential predators induce defensive changes in shell form and differences in growth rate. We examined such phenotypic plasticity in the direct-developing snail, Littorina subrotundata (Carpenter, 1864). Among experimental field populations of L. subrotundata exposed to differing intensities of predation by the purple shore crab, Hemigrapsus nudus (Dana, 1851), snails collected from predation-intense environments often had more massive shells than closely related snails from adjacent environments where predation was negligible. Snails collected from both environments were raised in tanks containing cages of H. nudus that were feeding on conspecific snails and compared to a control group raised in the absence of this stimulus. Most snails developed significantly more massive shells in the presence of the crabs suggesting that adaptive phenotypic plasticity may account for some of the variation we observed in the field. In one case, snails from a predation-intense environment did not exhibit a statistically significant amount of plasticity, but instead grew a more massive shell irrespective of the laboratory stimulus. We interpret this as evidence for a genetic difference in the plasticity of shell form among experimental populations, caused by intense selection by H. nudus. There was no statistical difference in the growth rates of snails among treatments.     

The annual cycle of molt in the Cape hare<Emphasis Type="Italic">Lepus capensis</Emphasis> in northern China: pattern,timing and duration

Molt is an important life history event for mammals occurring in temperate and cold zones. In the present paper, I investigate the pattern, timing and duration of seasonal molts of the Cape haresLepus capensis Linnaeus, 1798 in northern China, by tracing and scoring the process of each molt quantitatively. All the seasonal molts in both overwintering and juvenile hares went through a similar order: midback (together with nape), flanks, belly, upper tail, and legs. Yet, there was a relatively confused pattern of replacement during the heaviest molting period of overwintering hares, compared to a relatively uniform pattern among juvenile hares. Overwintering hares experienced a spring and a fall molt. The fall molt was already initiated prior to completion of the spring molt and had a relatively short duration. Juveniles born early in the year (before July), before their fall molt, had undergone a postnatal molt, but those born late (July to September) had not. The juvenile fall molt had already begun when the postnatal molt was at its later stages. These facts suggest that the time budget for fall molt is tight and seasonally constrained. The timing and duration of molts in the overwintering and early-born juvenile hares were independent of individual age. Among overwintering hares, both sexes started spring molt synchronously, but the females were behind the males in late period of this molt and consequently further delayed throughout subsequent fall molt. This may be related to a higher cost of reproduction in females. No sexual difference was found in the molting progress of juvenile hares.     

The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.     

Effect of limb regeneration on size increase at molt of the shore crabs Hemigrapsus oregonensis and Pachygrapsus crassipes.

Size increase at molt is reduced following multiple limb regeneration in the shore crabs, Hemigrapsus oregonensis and Pachygrapsus crassipes. Limb loss per se does not influence postmolt size. Effect of increasing number of regenerating limbs is additive. Postmolt size is programmed early in the premolt period of the preceding instar and is probably not readily influenced by water uptake mechanics at ecdysis. A simple model for growth, molting, and regeneration in heavily calcified Crustacea is developed from the viewpoint of adaptive strategies and energetic considerations.     

The role of nodal roots in prostrate clonal herbs: ‘phalanx’ versus ‘guerrilla’

The influence of nodal rooting on branching was studied in three evolutionarily and morphologically diverse species of prostrate clonal herbs: Tradescantia fluminensis (a monocotyledonous extreme ‘phalanx’ species), Calystegia silvatica (a dicotyledonous extreme ‘guerrilla’ species) and Trifolium repens (a dicotyledonous intermediate species). In all three, branch development from axillary buds is regulated by a positive signal produced by roots together with inhibitory influences from both pre-existing branches and shoot apical buds (apical dominance). Responses to nodal roots are cumulative and increased root activity leads to more vigorous bud outgrowth. In the absence of nodal roots, a single basal root system is unable to maintain continued extension growth of the shoot. We suggest that as individual nodal roots and stem internodes are both short-lived in these nodally-rooting clonal species, the plants’ investment in them is minimal. Thus, in contrast to perennial species lacking nodal roots, individual root systems in prostrate clonal herbs are small and stems have little secondary thickening and development of long-distance transport tissues. Hence the decline in extension growth of the shoot in the absence of nodal roots could be linked to the weak development of long-distance transport tissues in their relatively thin horizontal stems and to resource sharing between primary stems and lateral branches (as suggested by the greater retardation of primary stem growth in the more profusely branched ‘phalanx’ species (Trifolium and Tradescantia) than in the weakly branched ‘guerrilla’ species, Calystegia). These findings are consistent with the view that the long-term persistence of genotypes of nodally-rooting prostrate species is dependent upon them encountering the moist conditions required to facilitate the continual development of new young nodal root systems.     

Fitness, developmental instability, and the ontogeny of fluctuating asymmetry in Daphnia magna

Fluctuating asymmetry (FA) is the most commonly used measure of developmental instability. The relation between FA and individual fitness remains controversial, partly due to limited knowledge on the mechanisms behind variation in FA. To address this, we investigated the associations between FA, growth and reproduction as well as the ontogeny of FA in a clonal population of Daphnia magna . FA was not correlated with growth and reproduction, either at the between- or the within-individual level, in a high ( N  = 48 individuals) or in a low ( N  = 52 individuals) food-quantity regime. There were therefore no indications of functional effects of FA or of phenotypic trade-offs between developmental stability, growth and reproduction. Individual asymmetries varied randomly in sign and magnitude between subsequent molts ( N  = 19 individuals, 9–11 instars), but the levels of FA were generally lowest at intermediate ages. No feedback between right and left sides was detected. This suggests that FA only reflects the most recent growth history, that developmental instability may increase in old age, and that FA depends on processes operating on each side of the body independently. The results also suggest that FA differences within and among individual Daphnia are largely random, with limited biological significance.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 179–192.     

Environmental mediation of intraguild predation between the freshwater invader Gammarus pulex and the native G. duebeni celticus

Invasive species and environmental change often occur simultaneously across a habitat and therefore our understanding of their relative roles in the decline of native species is often poor. Here, the environmental mediation of a critical interspecific interaction, intraguild predation (IGP), was examined between invasive (Gammarus pulex) and native (G. d. celticus) freshwater amphipods. In the laboratory, IGP asymmetries (males preying on congeneric females) were examined in river water sourced from zones where: (1) the invader has completely displaced the native; (2) the two species currently co-exist, and (3) the native currently persists uninvaded. The invader was always a more effective IG predator, but this asymmetry was significantly weaker moving from ‘invader-only water’ through ‘co-existence water’ to ‘native-only water’. The constituent of the water that drives this mediation of IGP was not identified. However, balancing the rigour of laboratory experiments with field derived ‘environment’ has advanced understanding of known patterns in a native species decline, and its co-existence and persistence in the face of an invader.     

Regeneration and molting effects on a proprioceptor organ in the dungeness crab,Cancer magister

Decapoda Crustacea molt in order to grow; some species, such as the Dungeness crab Cancer magister, achieve a very large size. Does sendory neuron hyperplasia in internal proprioceptors accompany this growth? To determine this, neurons in propodite-dactylopodite (PD) chordotonal organs were counted in first walking legs of juvenile (5th through 9th instar) and adult (10th through 13th instar) C. magister. We found that the PD organs of J5 crabs have about 56 neurons; the number increases to about 61 neurons in J6 crabs. Significant hyperplasia now occurs because an average of 79 neurons are found in the PD organs of J7 crabs. Little to no hyperplasia accompanies the several succeeding juvenile and adult molts (ca. 82–86 neurons are present). Because autotomized limbs are regenerated upon molting, we also examined how the number of PD organ neurons in regenerated legs compare with those of pristine legs. Newly regenerated legs (termed 1st stage regenerates) have fewer sensory neurons than do their contralateral pristine partners (65 vs 81); larger regenerated legs which have attained nearly normal size as a result of additional molts (2nd stage regenerates) still have fewer neurons than their pristine partners (69 vs 81). Additionally, in contrast to those of pristine walking legs, the elastic strand of PD organs from 1st stage regenerates is a misshapen sheet containing a cluster of small neurons with no obvious functional organization. Nonetheless, neurophysiological recordings indicate that all the receptor types typical for pristine legs (movement and position cells) are represented. The PD organs of 2nd stage regenerates differentiate to the shape and neuronal organization of pristine legs. © 1994 John Wiley & Sons, Inc.     

A consensus map of rye integrating mapping data from five mapping populations

A consensus map of rye (Secale cereale L.) was constructed using JoinMap 2.0 based on mapping data from five different mapping populations, including ‘UC90’ × ‘E-line’, ‘P87’ × ‘P105’, ‘I_0.1-line’ × ‘I_0.1-line’, ‘E-line’ × ‘R-line’, and ‘Ds2’ × ‘RxL10’. The integration of the five mapping populations resulted in a 779-cM map containing 501 markers with the number of markers per chromosome ranging from 57 on 1R to 86 on 4R. The linkage sizes ranged from 71.5 cM on 2R to 148.7 cM on 4R. A comparison of the individual maps to the consensus map revealed that the linear locus order was generally in good agreement between the various populations, but the 4R orientations were not consistent among the five individual maps. The 4R short arm and long arm assignments were switched between the two population maps involving the ‘E-line’ parent and the other three individual maps. Map comparisons also indicated that marker order variations exist among the five individual maps. However, the chromosome 5R showed very little marker order variation among the five maps. The consensus map not only integrated the linkage data from different maps, but also greatly increased the map resolution, thus, facilitating molecular breeding activities involving rye and triticale.     

Subtle ‘boom and bust’ response of <Emphasis Type="Italic">Macquaria ambigua</Emphasis> to flooding in an Australian dryland river

The ecology of dryland rivers is driven by their highly variable hydrology, particularly flooding regimes, whereby intermittent floods typically generate ‘booms’ of primary and secondary productivity, including massive fish production. We tested these concepts in the Moonie River, Australia, using the percichthyid, Macquaria ambigua, a dryland river species known to display pronounced ‘boom and bust’ abundance patterns in response to floodplain inundation followed by extended periods of low to no channel flow. We expected that body condition (as measured by whole body lipid content) and biomass of M. ambigua would be related to prey biomass, and that these factors would all ‘spike’ following widespread flooding. Instead we found more subtle responses. There were ‘booms’ in biomass of Macrobrachium and zooplankton, two important food items, whereas M. ambigua maintained relatively low but sustained lipid and biomass levels following flooding. It appears that instead of a ‘boom’ in fish biomass, abundant invertebrate food resources and sustained lipid levels contributed to high survivorship of this species during the ‘bust’ period over cool dry months.     

Environment and position of first bud to break on apple shoots affects lateral outgrowth

A study was conducted to determine which bud (terminal or lateral) breaks first, and thereby exerts primigenic dominance, on ‘Granny Smith’ and ‘Golden Delicious’, 1-year-old apple (Malus × domestica Borkh.) shoots grown in two locations in the Western Cape, South Africa, with differing degrees of chilling. Primigenic dominance of laterals was more common in a warm area than a cool area, and more common in ‘Granny Smith’ than ‘Golden Delicious’. Laterals rarely broke before the terminal in ‘Golden Delicious’, and so differences in lateral development due to position of first bud to break were only analyzed in ‘Granny Smith’ shoots from this point on in the study. In ‘Granny Smith’, lateral budbreak and growth was influenced by the position of the first bud to break on the shoot, but did not differ between locations. On ‘Granny Smith’ shoots with primigenic dominance of the terminal, lateral budbreak and growth was suppressed, in accordance with the typical ‘delayed foliation’ commonly observed in warm winter climates. However, when at least one lateral broke before the terminal, lateral budbreak and growth were similar to previous observations in cold winter areas.     

Fluctuating Asymmetry in Body Traits Increases Predation Risks: Tawny Owl Selection Against Asymmetric Woodmice

During the last decade, the study of fluctuating asymmetry (FA) in relation to different fitness aspects has become a popular issue in evolutionary biology. There has been much recent debate in subtle departures from perfect symmetry in bilaterally paired morphological characters, and the extent to which such departure actually reflects aspects of individual quality and fitness. We used data from pellet collection and trapping sessions involving the trophic system Apodemus – Strix aluco, to test the hypothesis that asymmetric woodmice disproportionately fell prey to the tawny owl compared with “normal” woodmice. We found that woodmice preyed on by owls had significantly more asymmetric leg bones than survivors, particularly hind legs, those devoted to jumping. Thus asymmetry in locomotory traits apparently increased predation risks due probably to minor efficiency of asymmetric woodmice in evading predators or to their general low quality. These results suggest that FA affects fitness and consequently may be a good predictor of survival chances for woodmice, i.e. their quality; on the other hand, by removing asymmetric individuals, tawny owls can exert a stabilising selection on prey populations.Co-ordinating editor: M. Klaassen