Neck spine protects Daphnia pulex from predation by Chaoborus, but individuals with longer tail spine are at a greater risk

We tested the prey preference of Chaoborus sp. on Daphnia pulexwith different defensive morphologies. The protective functionof inducible morphological defences, such as neck spine andlonger tail spine, was evaluated Second instar D.pulex individualsof two clones differing in their strength of neck spine inductionwere offered as prey to both Chaoborus obscuripes and Chaoborusflavicans. We used logistic regression analysis to evaluatethe effect of morphometry on the vulnerability of Daphnia. Thepresence of a neck spine and increased total length protectedD.pulex from Chaoborus predation. However, individuals witha longer tail spine were more vulnerable to Chaoborus predationChaoborus obscuripes was able to eat daphnids with a neck spinedue to the larger gape size of this chaoborid. The smaller speciesC.flavicans almost always ate prey with no neck spine.     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Cyclomorphosis as a factor explaining success of a Daphnia hybrid in Tjeukemeer

Recent studies have shown that Daphnia hybrids are intermediate to their parental species for several life history traits, and that they combine advantageous traits of the parental species, thereby decreasing their risk of being preyed upon by planktivorous fish. In this study D. galeata, D. cucullata and their interspecific hybrid are compared with respect to cyclomorphosis and juvenile growth as a potential mechanism to avoid invertebrate predation. Helmets and tail spines are known to reduce predation risk, whereas fast juvenile growth reduces the period that Daphnia are vulnerable to predation. In laboratory cultures, I tested the hypothesis that, with regard to cyclomorphosis, juvenile hybrids are more comparable to the juveniles of D. cucullata, and with respect to growth rate, they are more comparable to D. galeata. The hybrid did not differ significantly from D. galeata in the time needed to reach 0.9 mm body length. In contrast, the relative spine and helmet length of juvenile hybrids were more comparable with those of D. cucullata, whereas in adult instars these characters were more similar to D. galeata. Thus during the first juvenile instars, the hybrids combine the relative long helmet and spine of D. cucullata, with the faster growth of D. galeata. This combination of characteristics might reduce hybrid vulnerability to invertebrate predation in field situations.     

Extreme helmet formation in Daphnia cucullata induced by small-scale turbulence

Although Daphnia cucullata is used as a textbook example forcyclomorphosis, distinct helmet development, as shown in thefield, has not been demonstrated in the laboratory until now.We show for the first time that small-scale turbulence is ableto induce the maximum response of morphological plasticity inD. cucullata. Helmet elongation reached magnitudes as extremeas observed in the field. Different modes of generating small-scaleturbulence caused different levels of helmet formation. Small-scaleturbulence alone may not be responsible for cyclomorphosis innature because field data show that two nearby lakes with similarmorphometry differ in cyclomorphosis patterns, while laboratoryexperiments show that there is no difference in the abilityto form helmets in the clones of each lake. Although helmetformation in D. cucullata is inducible with predator kairomones,helmet elongation is not as strong as that induced by turbulence.We discuss the possible role of helmets under turbulent conditions.     

Observations on the Biology of Fomes annosus, with Particular Reference to East Anglian Pine Plantations: II. Spore Production, Stump Infection, and Saprophytic Activity in Stumps

F. annosus produces conidia abundantly in culture but rarelyunder natural conditions. Sporophorea are formed on stumps anddying trees throughout the year provided that the humidity issufficiently high. Spore discharge in the plantations may occurat any time of year, but slowa down or ceases during very dryweather. It takes place at low temperatures, ceasing only whensporophores freeze. In the laboratory, F. annosus basidiospores germinate in relativehumidity exceeding 92 per cent. on freshly cut, unsterilizedpine wood. Viability is lost rapidly in light at temperaturesabove 15°C. Under humid conditions F. annosus spores will infect freshlycut pine stumps. There is evidence that spores of F. annosus washed down intosoil remain viable for at least a short period and that stumpscovered with soil are infected from this source. F. annosusspores can infect stumps for only a few weeks after felling,colonization after a longer interval probably beingprevdntedby competing fungi. In stumps inoculated with a mixed suspensionof F. annosus and Peniophora gigantea spores, Fomes at firstcolonized the wood but was soon replaced by P. gigantea. Otheraspects of competition are discussed. The incidence of natural infection in stumps and the factorsaffecting it are discussed Growth rates of F. annosus on malt agar and in lengths of pineroot are given. The fungus grows in stump roots at about 1 m.per year, and so closely approaches adjacent trees within ayear of thinning the plantation: In large stumps, F. annosusmay survive 15 or even 30 years after felling. Data are givenwhich suggest that the infective capacity of stumps containingF. annosus is greater in alkaline than in acid mils, while replacementby other fungi is slower. Competition of F. annosus with other fungi during colonizationof stumps is discussed. The parasite can grow along roots alreadyoccupied by certain fungi but not along roots containing others.The probable course of succession in stumps rotted by F. annousis described.     

Seasonality, substrate perference and attachment sites of epizoic diatoms on cladoceran zooplankton

Cladocerans from three lakes in northwest Iowa were examinedfor attached diatoms over a period of 1 year to assess the seasonality,substrate preference and attachment site specificities of epizoicdiatoms. Over 90% of the cladocerans had attached diatoms immediatelyfollowing ice thaw, when the lakes were not yet thermally stratified,surface water temperatures were low(10–15°C)and watertransparency was at its annual high. Mean maximum density ofdiatoms on cladocerans averaged among lakes at this time was3000 cells animal^–1. The dominant epizoic diatom was Synedracyclopum Brutschy, which was found most often on large, pelagicDaphnia pulicaria and Daphnia galeata mendotae. Heavily infestedanimals had diatoms concentrated on the second antennae andanterior ventral margin of the carapace. Animals with <5epizoic diatoms were more likely to have cells attached to thepostabdomen or tail spine. In addition to living epizoically,S.cyclopum was found free living in the plankton, exhibitingtemporal fluctuations in densities similar to epizoic counterpartsHowever, the densities of the free-living cells seldom exceededepizoic densities, suggesting that S.cyclopum is adapted forgrowth on motile cladoceran substrates. ¹Present address: Savannah River Ecology Lab, Drawer E, Aiken,SC 29802, USA ²Present address: Department of Fisheries and Aquatic Sciences,7922 NW 71st St, Gainesville, FL 32606, USA     

Tadpole Locomotion: Axial Movement and Tail Functions in a Largely Vertebraeless Vertebrate

SYNOPSIS.Tadpoles are exceptional among vertebrates in lackingvertebrae along most of their body axis. Their caudal myotomesare also unusually simple for free-living vertebrates. Thisoverall morphological simplicity, in theory, makes tadpolesgood models for exploring how vertebrates control undulatorymovements. We used electromyography (EMG), high speed ciné,computational fluid dynamics (CFD), and mechanical tissue testingto understand how Rana tadpoles regulate their locomotion. Bullfrog (Rana catesbeiana) tadpoles have several patterns ofmuscle activity, each specific to a particular swimming behavior.Ipsilateral muscles in the tail were active either in seriesor simultaneously, depending on the tadpole's velocity, andlinear and angular acceleration. When R. catesbeiana larvaeswam at their natural preferred tail beat frequency, musclesat the caudal end of their tail were inactive. Mechanical testsof tissue further suggest that the preferred tail beat frequencyclosely matches the resonance frequency of the tail thus minimizingthe energetic cost of locomotion. CFD modeling has demonstrated that the characteristically highamplitude oscillations at a tadpole's snout during normal rectilinearlocomotion do not add to drag, as might be supposed, but ratherhelp generate thrust. Mechanical testing of the tadpole tailfin has revealed that the fin is viscoelastic and stiffer insmall rather than large deformations. This property (among others)allows the tail to be light and flexible, yet stiff enough togenerate thrust in the absence of a bony or cartilaginous skeleton. Many recent studies have documented predator-induced polyphenismin tadpole tail shape. We suggest that this developmental plasticityin locomotor structures is more common in tadpoles than in othervertebrates because tadpoles do not need to reform skeletaltissue to change overall caudal shape. Tadpole tail fins and tip, in the absence of any skeleton, arefragile and often scarred by predators. Based on the high incidenceof tail fin injury seen in tadpoles in the wild, we suggestthat the tadpole tail fin and tip may play an ecological rolethat goes beyond serving as a propeller to help tadpoles staybeyond predators' reach. Those soft tissue axial structures,by failing under very small tensile loads, may also allow tadpolesto tear free of a predator's grasp.     

Cyclomorphosis in Daphnia lumholtzi induced by temperature

• 1 Cyclomorphosis is a well known phenomenon in Daphnia that involves a regular, seasonal, or induced change in body allometry. Long helmets and tail spines were induced in laboratory cultures of Daphnia lumholtzi with temperature of 31 °C as the proximal cue (temperature of locally occurring peak abundance in Kentucky Lake). The effect was greater in embryos than juveniles or adults exposed to the temperature cue.
• 2 The temperature cue appears to have a threshold value (animals cultured at 25 or 28 °C did not develop elongated helmets or spines). The helmet and spine length receded both with D. lumholtzi kept at a constant 31 °C temperature and when water temperature was decreased.
• 3 The induced helmet in this experiment (0.66 mm, 1.0 mm animal) was significantly longer than values reported in the literature for induction by planktivorous fish kairomones (0.25 mm, 1.2 mm animal). The strong response to a proximal cue of temperature may require the second weaker chemical cue for maintenance. It is suggested that a synergistic explanation with two cues may be more appropriate for cyclomorphosis induction and maintenance in Daphnia lumholtzi that could be tested with further studies.

     

Life-history responses to Chuoborus of spined and unspined Daphniu pulex

Water-borne chemicals released by the larvae of the predatoryphantom midge Chaoborus are known to induce morphological modificationsin its prey Daphnia pulex: these cladocerans develop neck spineswhich may carry several teeth. Some work has shown that thesemorphological variations enhance the prey's chances of escape.but since these neck teeth are not fixed defence reactions,they are thought to entail some form of cost, such as delayedmaturation and reduced fecundity. In this study. the relationshipbetween morphological and life-history changes in four clonesof Daphnia pulex reared in the presence and absence of Chaoborusflavicans was examined. Special emphasis was placed on the genotypiccomparison of the modifications. While all four clones showeda delay in maturation time in the presence of Chaoborus, theneck spine responses differed markedly among the genotypes:one clone never had any neck teeth, another always producedone single tooth, and two clones produced varying numbers ofteeth per spine (means 2.9 and 4. respectively). These resultsindicate that there is no general pattern of neck teeth productioncorresponding to delayed maturation. What there appears to beis genetic variability in two independent and possibly adaptiveresponses. However, the clone without neck teeth was the onlyone which showed no predator-induced reduction in fecundity.Another common morphological response to Chaoborus was thatjuveniles of all clones developed elongated tail spines.     

Vulnerability of Daphnia middendorffiana to Parabroteas sarsi predation: the role of the tail spine

The predaceous calanoid copepod Parabroteas sarsi and Daphniamiddendorffiana co-exist in South Andes ponds. Daphnia middendorffianajuveniles have a tail spine with negative allometric development.A series ofexperiments was carried out with tailed and tail-removedjuveniles of three different instars. In all cases tested, feedingrates were significantly higher on tail-removed prey. Directobservations showed a higher frequency of unsuccessful attackson spined juveniles when compared with tail-removed juveniles.The proportion of dorsal attacks also increased in spined juveniles.Prey total length was a much better predictor of feeding ratethan prey body size. Three groups of juveniles with equal totallength, but differentage, body size and biomass, showed no significantdifferences in their vulnerability to predation.     

Blue tail and striped body: why do lizards change their infant costume when growing up?

Ontogenetic changes in color and pattern that are not directlyrelated to reproduction are very common yet remain a poorlyunderstood phenomenon. One example is conspicuous colors inthe tails of fish, amphibians, and reptiles that fade out laterin life. We suggest a novel hypothesis: conspicuous tail colorsthat appear only in juveniles compensate for an increased activitylevel, deflecting imminent attacks to the tail. We observedblue-tailed, newly hatched lizards (Acanthodactylus beershebensis)in the field and compared 5 behavioral parameters with thoseof older individuals that had already lost their neonate coloration.In addition, we explored whether tail displays, often assumedto direct a predator's attention to the tail, disappear withthe color change. Striped blue-tailed hatchlings foraged moreactively than 3-week-old juveniles, spent a longer time in openmicrohabitats, and performed deflective tail displays. In comparison,2 other lacertids that do not undergo ontogenetic change didnot switch to safer foraging when growing up. The results suggestthat activity alteration may be a major factor affecting theontogenetic color and pattern change. Active lizards that foragein open habitats increase their probability of attack by ambushpredators. Conspicuous colors and deflection displays may shiftattacks to the expendable tail, increasing the prey's overallprobability of surviving attacks. The persistence of both stripedbody pattern and blue tail fits the active foraging period ofneonates and hence may be appropriate for other species thatdisplay a conspicuous tail accompanied by a striped pattern.     

ASPECTS OF THE LIFE CYCLE, POPULATION DYNAMICS,GROWTH AND SECONDARY PRODUCTION OF THE EDIBLE SNAIL HELIX LUCORUM LINNAEUS, 1758 (GASTROPODA, PULMONATA) IN GREECE

This paper examines the biology and ecology of Helix lucorumL. which lives in mainland Greece, as well as its growth andsecondary production. A demographic study revealed that (a)3 cohorts exist at any time during the year (when adults ofall generations belong to the same cohort) (b) egg-laying andhatching occur during the months of July and August respectively,(c) the most rapid growth takes place during spring. Study ofH. lucorum genitalia in relation to age showed that the snailsaxe sexually mature 3 years after hatching, when the largestdiameter of their shell (D) is equal to or greater than 35 mm Von Bertallanffy's method suggests that Helix lucorum may liveup to 14 years or more in order to reach its possible maximumsize (48.80 mm) The study of relative growth of D in relation to P_s (peristomesurface) of Helix lucorum shows that D grows faster than P_swhen D12.50 mm; juveniles change their growth rate when theirD arrives at 22.05 mm, and growth becomes slower when adultsarrive at 36.27 mm Annual secondary production calculated by the size frequencymethod gave a mean annual density of 3.39 individuals per m²,a mean annual crop (biomass) of 4.04 g-m^–2 and an annualproduction (P) of 5.02 g · m^–2. The annual turnoverratio (P//b) is equal to 1.24 (Received 23 June 1987;     

Symmetrical male sexual ornaments, paternal care, and offspring quality

Simultaneous manipulation of tail length and tail asymmetryin male barn swallows (Hirundo rustica) has revealed that femalesprefer maJes with both long and symmetrical tail ornaments overmales with short and asymmetrical ornaments. Fluctuating asymmetryin tail length has a negative effect on the maneuvering abilityof male barn swallows, and females may prefer males with symmetricaltail ornaments because they thereby acquire more direct fitnessbenefits in terms of paternal care. The least preferred maleswith short tails with high asymmetry performed an absolutelyand relatively larger share of feeding of nestlings than themost preferred males. However, the combined feeding rate ofthe pair was not statistically significantly different betweentreatment groups. Fully grown tarsus length and body mass ofoffspring on day 15 did not differ between treatments. Theseresults indicate that females do not prefer males with symmetricaltail ornaments because such males contribute a relatively orabsolutely larger share of parental duties. Although these resultsdo not explain the basis of female choice for long and symmetricaltails, the results are consistent with a hypothesis that femalesof species with biparental care should invest differentiallyin their offspring relative to the quality of their mates. Theresults are also consistent with a hypothesis that preferredmales have access to mates with superior parenting abilities     

Induction and costs of tail spine elongation in Daphnia hyalina×galeata: reduction of susceptibility to copepod predation

1. Seasonal field data showed a positive correlation between the tail spine length of Daphnia hyalina × galeata and the density of the copepod Acanthocyclops robustus . Laboratory experiments were designed to assess the mechanisms underlying the induction of tail spine elongation and to test whether this morphology reduced predation on Daphnia juveniles.
2. Both the elongated tail spine morph and control spine morph produced progeny with elongated tail spine when exposed to copepods. Instar increments, calculated from individual body length, showed that individual growth decreased in the presence of copepods. This decrease in individual growth was less pronounced in spined morphs relative to control morphs.
3. Copepods exhibited significantly higher feeding rates on control morph juveniles compared to elongated tail spine morph juveniles.
4. All Daphnia matured at instar 5 even though the control morph neonates took 24–30 more hours to mature in the presence of copepods than in the absence of copepods. Enlarged body and spine lengths of progeny coincided with reduced progeny number during the first six reproductive instars. The disadvantage of the reduced offspring number produced per female was balanced by enhanced survivorship of progeny subjected to copepod predation.     

Extreme cyclomorphosis in Daphnia lumholtzi

1. Daphnia lumholtzi, not previously reported in North America, was found in a small reservoir in East Texas in January, 1991, This species possesses extremely long spines and large fornices; an allometric study was performed to detect any temporal differences in specific growth rates of the spines relative to the body. 2. In nature, mature females attained 1.8mm body length, excluding spines, but when the head and tail spines are included, the total length reached a maximum of 5.6mm. 3. Differences in the growth patterns of the head spine and the tail spine relative to the body existed for D. lumholtzi from January to March 1991. Both the head and the tail spines grew at a faster rate than the body during all 3 months although the rates varied between them. The results contradict the invertebrate predation hypothesis (Dodson, 1974) in that D. lumholtzi's head and tail spines continue to grow during adulthood instead of stopping after the juvenile instars. 4. The head spines grew at a constant allometric rate over time while the tail spine grew faster as the temperature increased. Both varied significantly in length over the 3 months, with animals having the shortest spines in February and the longest in March.     

Cell Population Studies in Developing Root Primordia

The proliferation of polyploid and diploid cells was followedduring the development of lateral root primordia of Vicia faba.Presumptive primordial cells were treated with weak solutionsof colchicine, 0.003, 0.006, and 0.0125 per cent, for 3 h. Somecells are induced to become polyploid and they form a markedsub-population. The relative frequencies of diploid and polyploidcells were determined from the time small primordia were presenttill lateral roots were fully emerged. These frequencies havebeen found to change in a regular fashion and the changes reflectdifferent levels of mitotic activity in different parts of aprimordia as it develops. Polyploid cells occupy the centralcore of the primordia and they divide actively except for theperiod just before lateral root emergence. Coincident with thistemporary quiescence of the core cells, the peripheral cells,which remain diploid after treatment with the weaker solutionsof colchicine, continue to divide. These changes in mitoticactivity of the central and peripheral cells are shown bothin the changes in the relative frequencies of polyploid anddiploid mitoses as a primordium grows and in changes in mitoticindex. A microspectrophotometric study indicates that the centralcells are held, temporarily in the G₁ phase of the cell cycle.     

SEXUAL SELECTION IN THE BARN SWALLOW HIRUNDO RUSTICA. III. FEMALE TAIL ORNAMENTS

Male secondary sexual characters are often expressed in females, and the maintenance of the character in females can be due to either direct selection on females favoring the maintenance of the trait, or a correlated response to selection in males. Here I report on determinants of and phenotypic selection on tail length of female barn swallows Hirundo rustica. The homologous trait in males is under strong directional sexual selection. Female tail length was positively associated with several reproductive parameters including total seasonal reproductive success, even when controlling for year and age effects. A change in female tail length from one year to another was often associated with a change in the reproductive parameters correlated with absolute tail length. There was little evidence for an association between female tail length and the duration of the incubation period (only females incubate) and absolute and relative female provisioning rates of offspring, and subsequent size of offspring. Tail length of female barn swallows was positively correlated with that of their mates. Female tail length was a heritable trait as determined from regression of the tail trait of offspring on that of their mother and their father, and there was a positive genetic correlation between the tail trait in males and females. In conclusion, female tail length reliably reflects female reproductive potential, and the trait appears to be under directional selection, which may result in an evolutionary response to selection because of the heritability of the tail trait.     

Pollination Ecophysiology of Mercurialis annua L. (Euphorbiaceae), an Anemophilous Species Flowering all Year Round

Mercurialis annua L. is a dioecious anemophilous species thatflowers all year round in central and southern Italy. The flowersof both sexes are dimorphic: the female flower has a vestigialcalyx; the male flower consists only of a calyx that opens atanthesis. The anthers always dehisce after anthesis. The anthesisof male flowers seems to be temperature dependent, whereas antherdehiscence is related to relative humidity. The pollen grainsvary in volume according to the season: they are smaller whenrelative humidity is low and vice versa. They always decreasein volume after anther dehiscence and have the capacity to varyin volume and reach equilibrium with a changing environment.Viability is high, but may drop suddenly during heavy rain orhail that damage the exposed male flowers. The number of pollengrains per stigma varies from 0 to 300. The data is discussedin relation to the type of pollination and environmental characteristics.Copyright1994, 1999 Academic Press Mercurialis annua, dioecism, anthesis, anther dehiscence, pollen volume, pollen viability, anemophilous pollination, pollination ecology     

Experimental manipulation of tail length in female barn swallows (Hirundo rustica) affects their future reproductive success

Previous studies have shown no significant effect of experimentaltail length manipulation in female barn swallows (Hirundo rustica)at the beginning of a breeding season on reproductive successor behavior during that breeding season. In the present study,we investigate if tail length manipulation had any effect onreproductive performance the following year, the so-called long-termeffect, in contrast to the short-term effects already studied.We found that females with experimentally elongated externaltail feathers at the beginning of a breeding season producedless offspring during the breeding season the following yearthan did females with shortened or unmanipulated tails. Theseresults suggest that tail elongation caused flight deficienciesthat deteriorated the condition of females and eventually reducedreproductive success. The finding of long-term effects but nosignificant short-term effects for female tail elongation suggeststhat female barn swallows have the ability to adjust immediateparental investment. Detrimental effects of long tails in femalesin terms of decreased reproductive success might explain whyfemale tails are not as long as those of males. Finally, femalesmated to long-tailed (sexually attractive) males decreased theirreproductive success the following year more than did femalesmated to short-tailed males, possibly owing to differentialparental effort causing a deterioration of their condition.     

The function of long tails in female barn swallows (Hirundo rustica): an experimental study

The outermost tail feathers in male barn swallows (Hirundo rustica)are the target of a strong directional female mate preference.The tail ornament is also expressed in females, since femaleshave considerably longer tails than juveniles, either due to(1) a strong genetic correlation between the characters in thetwo sexes, or (2) direct sexual selection on females. To discriminatebetween these two hypotheses, we manipulated the length of theoutermost tail feathers in female barn swallows shortly afterarrival by either shortening or elongating the outermost tailfeathers, or maintaining their length among control individuals.Start of laying of the first clutch, reproductive performance,or provisioning of offspring did not show any significant differencesamong treatments. Original female tail length before manipulationwas unrelated to reproductive performance, while male tail lengthexplained some variation in the number of clutches and, to someextent, the total number of eggs laid per year. Females withlonger tails arrived earlier at the breeding grounds. Manipulatedfemale tail length was positively correlated to the tail lengthof their mates. Our results support the correlated responsehypothesis but do not support the sexual selection explanationfor the existence of exaggerated tail feathers in female barnswallows.