Seasonal changes in moult,body mass and reproductive condition in siskins Carduelis spinus exposed to daylength regimes simulating different latitudes

Birds use change in daylength during the year to time events during their annual cycles. Individual Eurasian siskins Carduelis spinus can breed and winter in widely separated areas in different years. Birds at different latitudes will experience different changes in photoperiod. So how does latitude affect photoperiodic control? Our aim in this study was to find whether Siskins caught from the wild in Britain and exposed to different photoperiodic regimes, typical of widely separated latitudes, would differ in the subsequent timing and duration of their moults and associated processes. Siskins were caught in late February and early March, and initially kept outside on natural photoperiods. From the spring equinox (21 March), they were divided into three groups kept under photoperiodic regimes that simulated latitudes 40°, 55° and 70°N respectively. All three groups showed highly significant subsequent changes in body mass, fat scores and cloacal protuberance size. Moult of the primary feathers started during June – August (mean 9 July), and lasted 61–99 days (mean 75 days). Birds that started to moult late in the season had shorter moult durations. All individuals showed lower mass and fat levels during moult than before or after moult. Crucially, there were no significant differences in the timing of these events between the three photoperiodic groups. Apparently these birds did not use prevailing absolute photoperiod or the prevailing rate of change in photoperiod to time moult‐related seasonal events, but used instead some other feature of the annual photoperiod cycle or some form of interval timer linked to photoperiod.     

Presence of sculpins (Cottus gobio) reduces drift and activity of Gammarus pulex (Amphipoda)

Introduction of sculpins into a stream previously devoid of these predators significantly reduced drift rate of Gammarus pulex. The drift of insect larvae was not affected. High amounts of exudates after implantation of sculpins were probably responsible for the low number of drifting G. pulex specimens. Laboratory experiments confirmed reduced locomotory activity of G. pulex when exposed to caged sculpins, an observation that excludes reduced drift activity as a result only of predation.The average size of drifting G. pulex specimens was larger during the night than during the day. This result is in accordance with the hypothesis that large individuals should, in relation to small ones, turn nocturnal because of greater predation risk during daytime. Presence of sculpins did not alter the size composition of drifting G. pulex.     

Influence of diet and photoperiod on development and reproduction of European populations of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae)

The current study examines the effect of photoperiod (16:08 or 12:12 h L:D) and diet (eggs of Ephestia kuehniella Zeller (Lepidoptera: Phycitidae) or the pea aphid Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae)) on the development and reproduction of the multicoloured Asian lady beetle Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). A long-term laboratory population of H. axyridis (since 1998) and a melanic and non-melanic population originating from field collected individuals of H. axyridis in Belgium were used in this study. Long day conditions (16 h photoperiod) shortened development of the field populations with 2–3 days when compared with short day conditions (12 h photoperiod). Oviposition in the field populations was delayed by 1–3 months when reared at a 12 h photoperiod. Dissections indicated that the females were in reproductive diapause. As compared with live pea aphids, a diet consisting of E. kuehniella eggs yielded heavier adult body weights (up to 12%) and increased the number of egg laying days (by 45–169%) for both field populations at a 16 h photoperiod and lengthened adult life span (by 45–92%) under both light regimens. The morph types differed in their response to the foods offered in terms of developmental rate, pre-oviposition period and number of oviposition days. The laboratory and field strains responded differentially to regimens of food and photoperiod. The study indicated a greater nutritional plasticity of the non-melanic morphs which may offer them a competitive advantage that may in part explain the predominance of non-melanic morphs in newly colonized areas.     

Match–Mismatch Experiments Using Photoperiod Expose Developmental Plasticity of Personality Traits

Behavioural differences between individuals are often found to be consistent across contexts and/or over time, although recent studies suggest that ontogenetic processes and learning might influence personality. During ontogeny, environmental influences may play an important role in shaping an individual's personality as well as its physiology. Seasonal changes are ubiquitous and known to influence development. To study developmental plasticity, of behaviour and physiology in the wild cavy (Cavia aperea), we manipulated the photoperiod in a fully crossed match–mismatch design by simulating spring and autumn photoperiod until weaning and subsequently moving half of the animals into the mismatching photoperiod. We found developmental plasticity in behavioural and physiological traits before and after sexual maturation for growth, resting metabolic rate and fearlessness. For fearlessness, changes in response to the opposite photoperiod were more pronounced in males than in females. Exploration and boldness were only influenced by early, but not by late photoperiod. No sex differences were found for these two traits. Even though our treatment changed average trait expression, some behavioural traits proved consistent over time, but physiological traits were not. Fearlessness was consistent only in animals that did not change photoperiod during development, whereas exploration and boldness were consistent over time regardless of photoperiodic treatment. Our study shows that in response to a change in photoperiod personality traits differ substantially in developmental plasticity.     

Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Ecology and population structure of the bayou darter, <Emphasis Type="Italic">Etheostoma rubrum</Emphasis>: disjunct riffle habitats and downstream transport of larvae

The bayou darter, Etheostoma rubrum (Percidae), is endemic to the Bayou Pierre system in Mississippi. Adult and juvenile E. rubrum occupy swift, shallow riffles or runs over coarse gravel and pebble substrata. Habitat requirements of larval and post-larval stages, and the role of downstream dispersal of larvae in colonizing riffles are poorly known. The potential for movement and the high level of habitat specificity for the discontinuous riffle habitat suggest that E. rubrum may comprise a metapopulation of linearly arranged local populations. The greatest population densities of E. rubrum occur in the upper reaches of Bayou Pierre. We hypothesized that metapopulation structure of E. rubrum may include source–sink dynamics, whereby downstream areas are a sink for larvae/early juveniles originating upstream. We tested hypotheses that a transport mechanism, larval drift, occurred in E. rubrum, and that downstream riffles showed characteristics of population sinks. We captured larval stages of E. rubrum in cross-sectional and longitudinal drift nets, and rates of drift tended to increase during the day. Larval E. rubrum (n=19) occurred in samples above and below riffle areas, with no differences among longitudinal drift nets placed above and below riffles. Thus, larval drift is a viable transport mechanism. Relative abundance of adults and juveniles declined from upstream to downstream, and inter-riffle distances increased with cumulative downstream distance. However, both predictions of the source–sink hypothesis were not supported. The distribution of size classes did not change between upstream and downstream riffles nor did the mean size-adjusted body mass.     

Foraging activity of two species of predatory leeches exposed to active and sedentary prey

Summary The foraging activity of the freshwater leeches Nephelopsis obscura and Erpobdella punctata during acclimation to active or sedentary prey, starvation and refeeding was examined by filming over a 24 h cycle. For both species, the time of activity of individual leeches and the total number of leeches active were significantly correlated. Activity in the dark was significantly higher than in the light with the activity of N. obscura in the dark 16.2 times higher in terms of time and 6.7 times higher in terms of numbers of individuals moving than E. punctata. Large N. obscura were significantly more active than medium or small individuals and encountered more sedentary prey. With lower levels of activity E. punctata encounters and feeds on more active prey, and in lotic ecosystems its lower exposure to fish predation and drift is, at least partially, responsible for its numerical abundance over N. obscura. There were no significant differences in the activity of N. obscura or E. punctata fed on active or sedentary prey during either subsequent starvation of 1 to 10 d, or after refeeding, suggesting that unchanged foraging activity provides the optimun net energy gain during periods of change in prey type and availability.     

Seasonal adaptations of the fall webworm <Emphasis Type="Italic">Hyphantria cunea</Emphasis> (Drury) (Lepidoptera: Arctiidae) following its invasion of Japan

The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae), invaded Japan from North America about 60 years ago. Immediately after its invasion – and for the first three decades – its life cycle was bivoltine, two generations per year throughout its entire distribution in Japan. Thereafter, its life cycle shifted to trivoltine in the southwestern areas of Japan. In the present study we examined the life-history traits proposed to be implicated in this event with the aim of clarifying the mechanism of this life-cycle shift. The critical photoperiod for diapause induction, as defined by the photoperiod at which 50% of individuals enter diapause, was shorter in the trivoltine populations than in their bivoltine counterparts. The temperature sensitivity of the photoperiodic response, as defined by the difference in the critical photoperiod between 20 and 25°C, was greater in the trivoltine populations than in the bivoltine ones. The geographic variation in larval and pupal periods was positively correlated to the latitude of the original localities of the populations. The change in the number of larval instars would be one of the main factors accounting for the regional differences in the developmental period. These results suggest that some life-history traits of H. cunea have changed following its invasion of Japan as an adaptive response to local climates.     

Abiotic influences on embryo growth: statoliths as experimental tools in the squid early life history

Statolith size and growth was used to determine the influence of abiotic factors on the growth of Loligo vulgaris and Sepioteuthis australis embryos. Recently spawned egg masses collected from the field were incubated in the laboratory under different levels of light intensity, photoperiod, or short periods of low salinity (30‰). Double tetracycline staining was used to follow statolith growth. In L. vulgaris constant light conditions produced significantly slower growth in the embryonic statoliths and embryos held at summer photoperiod had slower statolith growth than those held at winter photoperiods. However once they hatched out there was no evidence that photoperiod affected statolith growth. After hatching, in all photoperiods statolith growth rates decreased in comparison with late embryonic rates. In S. australis embryos, differences between the high and medium light intensities for summer and intermediate photoperiods were found, suggesting that under summer incubation temperature, longer daylengths at medium light intensity favoured higher statolith growth for this species. In comparison to controls, slower statolith growth in S. australis embryos due to low salinity only occurred when exposed for 72 h. Comparison with previous studies indicates that temperature seems to be the main abiotic factor influencing statolith growth during early stages, however, interactions among all abiotic factors needs to be determined as well as the unknown influence of other isolated factors, e.g., oxygen concentration within the egg mass.     

Reproduction and perennation of Chondrilla juncea L. (skeleton weed) in the Western Australian wheatbelt

Survivorship of first-year plants of Chondrilla juncea was considerably higher on a deep yellow sand than on a yellow duplex soil. Mortality on the latter soil was associated with drying of the Al horizon during early summer. No plants produced seeds in their first year of growth on the duplex soil. However, during both years of the study, first-year individuals of both forms of C. juncea produced viable seeds when growing on the deep sand. Higher population growth rates could be expected on deep sands, owing to a combination of earlier reproduction and greater survivorship through the first summer. It is suggested that whilst the entire wheatbelt might be climatically suitable for C. juncea, the invasive potential of this weed might be restricted by edaphic characteristics over large areas.     

Occurrence of different development time patterns induced by photoperiod in Anagrus atomus (Hymenoptera: Mymaridae), an egg parasitoid of Empoasca vitis (Homoptera: Cicadellidae)

The wasp Anagrus atomus L. parasitizes eggs of the cicadellid Empoasca vitis (Göthe) on grapevines during the vegetative season. In vineyards, abundant parasitoid emergence occurs in the autumn as a result of a developmental delay in part of the A. atomus population. In this context, physiological times required by the parasitoid to develop in E. vitis eggs are recorded during the summer. In early summer, all individuals develop fully, with physiological times in agreement with those reported in the literature. From early August onwards, an increasing proportion of parasitoids show a delay in development involving pre‐imaginal stages. Under transmission microscopy, larvae of A. atomus inside the eggs of E. vitis can be clearly differentiated as ‘light’ and ‘dark’ types, which are associated with normal and delayed development, respectively. The decreasing photoperiod as autumn approaches appears to promote the retardation of growth. This phenomenon can be considered a typical case of risk spreading because, when the availability of E. vitis eggs in vineyards is high (i.e. in early summer), A. atomus shows (i) only a normal developmental pattern that allows a faster population increase, whereas (ii) when E. vitis egg availability begins to be scarce (i.e. from mid summer), an increasing proportion of individuals having delayed development provides better fitness because it allows the parasitoid to synchronize its life history with leafhopper species that lay overwintering eggs on plants other than grapevines.     

Climate change, diapause termination and zooplankton population dynamics: an experimental and modelling approach

1. Earlier spring warming as predicted for climate change will alter combinations of water temperature and photoperiod that act as emergence cues for zooplankton resting stages. As a result, water temperature cue thresholds will be experienced at shorter photoperiods, a variable independent of weather variations. Also, light intensity, another potentially important cue for zooplankton emergence, could decrease in many lakes if symptoms of climate change resemble those of eutrophication. 2. We designed a laboratory experiment to test the effects of three factors, temperature (6, 9 and 12 °C), photoperiod (13L : 11D and 16L : 8D) and light intensity (20 and 35 μE m⁻² s⁻¹) on hatchling abundance and timing of hatching of daphniids (Daphnia ambigua) and rotifers (Keratella spp. and Synchaeta pectinata) from resting eggs. Further, we investigated the implications of potential changes in hatching dynamics, following variations in hatching cues, on zooplankton spring population development using predator–prey simulation models. 3. For hatchling abundance and timing of hatching, photoperiod had a significant effect for D. ambigua but not rotifers. Daphnia ambigua hatchling abundance decreased by 50% when incubated at conditions mimicking early spring (12 °C + 13‐h photoperiod) compared to a later spring (12 °C + 16‐h photoperiod). Light intensity has a significant effect only for S. pectinata, producing greater hatchling abundance at lower light intensity. 4. Simulation models suggest that in contrast to a later spring, an early warming produces a shift in spring zooplankton community composition, from daphniid to rotifer dominance. These patterns are primarily driven by differential zooplankton emergence response with variations in temperature–photoperiod cues. 5. Overall, our laboratory experiments and simulation models suggest that lakes with strong dependence on the ‘resting egg‐bank’, characteristic of many shallow north‐temperate lakes or in years with low winter survivorship of adult zooplankton, may be most susceptible to climate change. Further, fewer large grazers such as daphniids with an earlier spring may result in less control of cyanobacterial blooms in eutrophic lakes.     

Drift of characin larvae,Bryconamericus deuterodonoides,during the dry season from Andean piedmont streams

Synopsis The downstream transport of fish larvae is well known from temperate running waters, but there exists remarkably little information for tropical streams. We sampled drift in two Andean piedmont streams during the dry seasons of 1986–1988. Bryconamericus deuterodonoides (Characidae) larvae were found to be extremely abundant in some drift collections, with peak drift densities exceeding those of temperate streams by as much as an order of magnitude. Drift of fish larvae displayed pronounced diel periodicity, with more than one-quarter of a million larvae collected during the night compared to two individuals during the day. Fish larvae were more abundant during the latter compared to early part of the dry season, even in years when streams were reduced to small isolated pools by the end of the season. These observations indicate that the dry season represents an important period of reproduction for the characid, Bryconamericus deuterodonoides, and possibly other neotropical fish species.     

Black capuchin monkeys dynamically adjust group spread throughout the day

Variation in group spread can affect communication between individuals, synchrony of activity and collective decision-making. Most studies have measured this variation in large time scales, but the variables that influence group spread change as the social unit ranges over different areas throughout the day. Thus, our aim was to analyze the diurnal variation in group spread of a group of black capuchin monkeys (Sapajus nigritus) inhabiting an area within the southeastern Brazilian Atlantic Forest, the Carlos Botelho State Park. We measured group spread as the distance between the coordinates of the location of the two most distant group members every 5 min during entire days. Using time series analysis, we found that the group spread varied within short periods of time (15 min) throughout the day, revolving around the average group spread distance of 36 m. Group members tended to decrease the interindividual distance over time when they were more than 36 m apart, and to increase the distance when they were at a distance of less than 36 m. In addition, group spread was higher when monkeys were eating fruit than during other activities. We conclude that the variation in group spread may reflect competition for discrete food resources and that increasing cohesion several times throughout the day can be a way of resuming social contact between individuals.

     

Geographical variation in photoperiodic induction of larval diapause in the bruchid beetle, Bruchidius dorsalis: polymorphism in overwintering stages

The bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and shows geographical variation of overwintering stages in Japan. Our previous study found that B. dorsalis enters larval diapause in the final instar under short photoperiods. In cooler areas, we observed that most individuals overwinter in the final larval stage in diapause, whereas beetles at different developmental stages (non‐diapausing young instars, diapausing instars, and adults) were overwintering in warmer areas. In this study, we investigated geographical variation in the photoperiodic response for induction of larval diapause at 20 °C (three populations) and 24 °C (two populations) to clarify the overwintering strategy of B. dorsalis. We observed that (1) diapause incidence at 20 °C changed sharply from ca. 100% to 0% with a change in photoperiod in all the populations, (2) critical photoperiod was longer at 20 °C in populations from cooler areas, and (3) critical photoperiod at 24 °C was shorter than at 20 °C and a fraction of the larvae did not enter diapause, even under short photoperiods. Overwintering stages estimated from these results were consistent with those actually observed in the field. This study indicates that the geographical variation of overwintering stages is likely to reflect adaptive diapause induction in each local environment.     

Effect of photoperiod on a winter and on a summer diapause in two species of cranefly (Tipulidae)

Two craneflies, Tipula subnodicornis and Tipula pagana, both undergo diapause in the final larval instar. The species showed differences in the intensity of diapause and in the timing of the photoperiodic reaction during diapause, that could be related to season. Tipula subnodicornis undergoes a winter diapause that is induced and maintained in its early stages by short photoperiod (L:D:6:18). In the laboratory individuals in the early stages of diapause terminated diapause and pupated earlier when they were exposed to daylengths of, or greater than, 12 hr. However, it is suggested that in the field diapause is broken before the natural daylength is long enough to have any accelerating effect on development. Tipula pagana has a summer diapause which is of greater intensity than that of Tipula subnodicornis and some larvae were maintained for 197 days in the laboratory, without pupating, on an 18 hr daylength. Diapause was broken by a L:D;16:8 photoperiod and development was accelerated by a further decrease in daylength. The acceleration in development rate was attended by a decrease in the variance about the mean date of emergence and resulted in a highly synchronised emergence period. It is suggested that this quantitative response to daylength is particularly important to a species that emerges in the autumn when the temperature in the field is falling.     

Short photoperiod reduces the temperature sensitivity of leaf‐out in saplings of Fagus sylvatica but not in horse chestnut

Leaf phenology is one of the most reliable bioindicators of ongoing global warming in temperate and boreal zones because it is highly sensitive to temperature variation. A large number of studies have reported advanced spring leaf‐out due to global warming, yet the temperature sensitivity of leaf‐out has significantly decreased in temperate deciduous tree species over the past three decades. One of the possible mechanisms is that photoperiod is limiting further advance to protect the leaves against potential damaging frosts. However, the “photoperiod limitation” hypothesis remains poorly investigated and experimentally tested. Here, we conducted a photoperiod‐ and temperature‐manipulation experiment in climate chambers on two common deciduous species in Europe: Fagus sylvatica (European beech, a typically late flushing species) and Aesculus hippocastanum (horse chestnut, a typically early flushing species). In agreement with previous studies, we found that the warming significantly advanced the leaf‐out dates by 4.3 and 3.7 days/°C for beech and horse chestnut saplings, respectively. However, shorter photoperiod significantly reduced the temperature sensitivity of beech only (3.0 days/°C) by substantially increasing the heat requirement to avoid leafing‐out too early. Interestingly, the photoperiod limitation only occurs below a certain daylength (photoperiod threshold) when the warming increased above 4°C for beech trees. In contrast, for chestnut, no photoperiod threshold was found even when the ambient air temperature was warmed by 5°C. Given the species‐specific photoperiod effect on leaf phenology, the sequence of the leaf‐out timing among forest tree species may change under future climate warming conditions. Nonphotoperiodic species may benefit from warmer springs by starting the growing season earlier than photoperiodic sensitive species, modifying forest ecosystem structure and functions, but this photoperiod limitation needs to be further investigated experimentally in numerous species.     

A diurnal rhythm of pituitary prolactin activity with diurnal effects of mammalian and teleostean prolactin on total body lipid deposition and liver lipid metabolism in teleost fishes*

Previous investigations have shown that mammalian prolactin has a pronounced effect on lipid deposition in teleost fishes (e.g. Meier et al., 1971; de Vlaming & Sage, 1972). Prolactin can be either lipogenic or lipolytic depending on when during the day it is injected. Studies were initiated to determine whether the diurnal effects of prolactin on lipid metabolism in Fundulus similis vary with season or photoperiod acclimation. Data presented here indicate that the diurnal rhythm of response to prolactin persists in the pre-spawning and post-spawning seasons if F. similis is maintained on a long photoperiod. When animals are acclimated to a short photoperiod, treatment with mammalian prolactin either early or late in the photoperiod causes lipid depletion. Homogenates of the portion of the pituitary containing only prolactin cells cause lipid depletion in F. similis when injected early during a long photoperiod, but stimulate lipogenesis if injected late in the photoperiod; these results imply that fish prolactin may be involved in lipid metabolism. Furthermore, prolactin activity in the pituitary of F. similis varies diurnally. In vitro studies of lipid metabolism in Notemigonous crysoleucas and Ictalurus natalis suggest that the liver may be one of the target organs for prolactin.     

Female reproductive responses to photoperiod and male odours in the subterranean rodentCtenomys talarum

The tuco-tucoCtenomys talarum Thomas, 1898 is a solitary subterranean rodent that breeds seasonally in coastal sand-dune grasslands from Argentina. We assessed whether short-day photoperiod determines the onset of reproduction by an increase in female’s receptivity and whether male odour interacts with photoperiod as a cue by producing a greater response in receptivity. Receptivity was measured by an increase in the percentage of vaginal epithelial cells, particularly cornified cells. Females were trapped during their non-reproductive season (summer-long days) and randomly assigned to different day-length experimental groups: constant summer (CS), early winter (EW), and natural (NA) photoperiod. We examined the vaginal epithelium using vaginal smears during the first experimental phase (120 days) to test the effect of photoperiod. In the second experimental phase (15 days), we presented the females with male-soiled shavings to test the effect of photoperiod and male odor.Ctenomys talarum females showed some evidence of reproductive responsiveness to photoperiodic cuing. The proportion of epithelial cells peaked earlier in females when winter day-length was advanced than in females under natural photoperiod. Contrary to that expected, male odours did not stimulate female receptivity. These results suggest that, while photoperiod may participate in regulating reproduction, the integration of other environmental and social factors, as well as the identification of bodily conditions that lead to differences in individual response needs further consideration.     

A COMPARISON OF DRAGON KELP,EUALARIA FISTULOSA, (PHAEOPHYCEAE) FECUNDITY IN URCHIN BARRENS AND NEARBY KELP BEDS THROUGHOUT THE ALEUTIAN ARCHIPELAGO1

The Aleutian Archipelago coastal ecosystem has undergone a dramatic change in community composition during the past two decades. Following the removal of ～99% of the sea otters, Enhydra lutris, from the ecosystem, changes to the benthic communities resulted in widespread losses to most of the region’s kelp beds and corresponding increases in the prevalence of urchin barrens. Within the urchin barrens, the few kelps that have remained are exposed to elevated light, nutrients and currents, all of which may enhance their physiological condition and thus result in greater fecundity. To explore this further, we examined patterns of sporophyte fecundity in the dominant canopy‐forming kelp, Eualaria fistulosa, in both urchin barrens and in nearby kelp beds at seven Aleutian Islands spanning a range of 800 km. We found that the average weight of E. fistulosa sporophyll bundles was significantly greater on sporophytes occurring in the urchin barrens than in the nearby kelp beds. Furthermore, the average number of zoospores released per cm² of sporophyll area was also significantly greater in individuals from the urchin barrens than the nearby kelp beds. When these two metrics were combined, our results suggest that individual E. fistulosa sporophytes occurring in the urchin barrens may produce as many as three times more zoospores than individual E. fistulosa sporophytes occurring in the nearby kelp beds, and thus they may contribute disproportionately to the following year’s sporophyte recruitment in both urchin barrens and the adjacent kelp beds.