Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

A model for a species with two life history stages and added mortality

A differential equation model of a species with two life stages is constructed, and the behavior of the equilibria investigated when density-independent mortality is continuously imposed. With density dependence largely in recruitment, an increase in either life stage is possible even though mortality is being imposed on both life stages. The equilibrium of a given life stage can increase if density dependence is mainly in that stage.The most effective stage to attack is that which is numerically dominant. With species having a high reproductive potential the most effective way of reducing either stage is to attack the stage with the lesser natural density-independent mortality.Certain parameter values allow multiple positive steady states. Data from Lucilia cuprina appear to conform to some of the predictions of the model.     

Pre-migratory life history stages of juvenile Arctic birds: costs, constraints, and trade-offs

Many young birds on the Arctic tundra are confronted by a challenging task: they must molt their feathers and accumulate fat stores for the autumn migration before climatic conditions deteriorate. Our understanding of the costs and constraints associated with these stages is extremely limited. We investigated post-juvenal molt and premigratory fattening in free-ranging juvenile White-crowned Sparrows (Zonotrichia leucophrys gambelii) on the Arctic tundra. We found evidence for trade-offs between premigratory fat accumulation and molt: heavily molting birds had significantly less fat. Birds increased the rate of fat accumulation as the season progressed, but we found no evidence of a similar increase in rate of molt. Using a controlled captive study to isolate the energetic costs of body feather replacement, we found no difference in fat or size-corrected mass of birds actively growing body feathers as compared to controls. Molting birds, however, consumed 17% more food than controls, suggesting a significant cost of body feather growth. Our results provide evidence of significant costs, constraints, and trade-offs associated with post-juvenal molt and premigratory fat accumulation in young Arctic birds.     

Tissue type matters: selective herbivory on different life history stages of an isomorphic alga

Selective grazing by herbivores can have large effects on the population dynamics and community structure of primary producers. However, the ecological impacts of within-species herbivore preference for tissues of different phases (e.g., ploidy levels) or reproductive status remain relatively poorly known, especially among algae and other species with free-living haploid (gametophyte) and diploid (sporophyte) phases. We tested for herbivore selectivity among tissue types of the isomorphic (identical haploid and diploid free-living stages) red alga Mazzaella flaccida. Laboratory feeding assays demonstrated that the snail Tegula funebralis exhibited more than a threefold preference for gametophyte reproductive tissue over other tissue types, due to morphological differences. In contrast, the urchin Strongylocentrotus purpuratus did not distinguish as clearly between gametophytes and sporophytes; but it did prefer sporophyte reproductive to nonreproductive tissue, due to differences in water-soluble chemicals. Field surveys of grazer damage on M. flaccida blades were consistent with these laboratory preferences, with more damage found on gametophytes than sporophytes and reproductive than nonreproductive tissues. Differential fecundity can contribute to a skew in relative frequencies of phases in the field, and our results suggest that differential grazing by snails may contribute to this pattern and thus play a role in algal population biology.     

Effects of the dinoflagellates Karlodinium veneficum and Prorocentrum minimum on early life history stages of the eastern oyster (Crassostrea virginica)

The bloom-forming dinoflagellates Prorocentrum minimum and Karlodinium veneficum can have detrimental effects on some marine life, including shellfish, but little is known about their effects on early life history stages of bivalves. In the Chesapeake Bay region, blooms of these dinoflagellates overlap with the spawning season of the eastern oyster, Crassostrea virginica. In laboratory experiments, we compared the effects of P. minimum and K. veneficum on the survival and development of embryos and larvae of the eastern oyster. At 10⁴ cells ml⁻¹, P. minimum did not have a negative effect on embryos and larvae in 2-day exposures. The yield of D-hinge larvae was equal to or greater than in control treatments. At 2 × 10⁴ cells ml⁻¹ (approximately equal biomass to the P. minimum treatment) K. veneficum caused significant mortality to oyster embryos within 1 day and almost no embryos developed into D-hinge larvae. This effect was not alleviated by the provision of an alternate food source (Isochrysis sp.). Significant mortality was observed when larvae were exposed to K. veneficum at concentrations of 10⁴ cells ml⁻¹ (approximately 5 ng ml⁻¹ of karlotoxin). The K. veneficum cultures used in these experiments were relatively low in toxin content, more toxic strains could be expected to cause mortality at lower cell concentrations. Survival and maturation of embryos and larvae may be reduced when spawns of the eastern oyster coincide with high bloom densities of K. veneficum.     

Mass mortality of Strongylocentrotus droebachiensis (Echinodermata: Echinoidea) off Nova Scotia,Canada

A mass mortality of Strongylocentrotus droebachiensis, attributed to disease, was monitored in an echinoiddominated barren ground at Eagle Head on the south-western coast of Nova Scotia, Canada, in 1982. Mortality was 70% in a shallow (3 m) nearshore area, resulting in a loss of echinoid biomass of 2 042 g fresh weight m^-2, and 6% in deeper (7 m, 10 m) offshore areas. Echinoid density, size and nutritional condition (gonad index) were highest in the nearshore area. Survivorship was higher in juveniles (<15 mm diameter) than in adults resulting in the formation of a bimodal size distribution in the nearshore area. Mortality began around early October, near the peak of the annual cycle of seawater temperature (15°C), and was arrested by early December (seawater temperature 7°C) when morbid echinoids appeared to recover. In laboratory experiments, time to morbidity of S. droebachiensis exposed to morbid conspecifics increased exponentially with decreasing temperature (20° to 8°C). There was no survival at 20° and 16°C, 20% survival at 12°C and 100% survival at 8°C after 60 d; suggesting a lower temperature limit (between 12° and 8°C) for possible transmission of a pathogenic agent. Morbid laboratory echinoids from experiments at 16°C, and recovering echinoids collected in the nearshore area in early December, showed 100 and 85% survival respectively at <=8°C, and 0 and 15% survival respectively at 16°C, after 30 d. Time to morbidity was not affected significantly by nutritional condition and was similar for juvenile and adult echinoids. Time to morbidity was greater in echinoids exposed to one or three morbid individuals continuously, or seven morbid individuals for 1 h, relative to higher levels of exposure (up to seven morbid individuals continuously). Recent mass mortalities in S. droebachiensis have occurred in years of record high sea surface temperatures. The extent of mortality is correlated with the magnitude and duration of temperatures above a lower limit.     

Sex and microhabitat influence the uptake and allocation of mycosporine-like amino acids to tissues in the purple sea urchin, Strongylocentrotus purpuratus

UVR-absorbing mycosporine-like amino acids (MAAs) were detected in tissues of Strongylocentrotus purpuratus and in ten species of Rhodophyte macroalgae (eight previously untested) collected from intertidal microhabitats in November and January 2006?C2007 in Central California (35°09??N, 120°45??W). In sea urchins, MAA concentrations were higher in ovaries than testes, while epidermal concentrations were similar between sexes. Ovaries and epidermal tissues had similar MAA signatures and broadband UVA/UVB absorbance, while testes had a narrower absorption ranges shifted toward higher energy wavelengths. Sea urchins occupying pits in the substrate exhibited lower MAA concentrations than those outside pits, suggesting adult microhabitat may impact UV protection. Light levels did not influence gonadal MAA concentrations, but correlated with elevated epidermal MAA concentrations for males in the sunniest microhabitat. This study suggests sex and habitat strongly influence MAA concentrations among individual S. purpuratus and that allocation of MAA sunscreens to tissues in response to UVR is sex-dependent.     

Density dependence and density independence during the early life stages of four marine fish stocks

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.     

Reproductive biology and early life history of the hermaphroditic feather star Dorometra sesokonis (Echinodermata: Crinoidea)

Reproduction and larval/post-larval development of the one of the smallest known comatulid Dorometra sesokonis were studied on the coral reefs of Sesoko-jima Island, Okinawa, Japan. Breeding individuals were found every month from May 2003 to April 2004 indicating continuous reproduction. Individuals possessed both mature ovaries and testes, and were therefore characterized as simultaneous hermaphrodites. This species was observed to be an external brooder in which the embryos developed on the surface of parental genital pinnules until the doliolaria larval stage, and subsequently released doliolariae settled on substrata within 4 days after hatching. Larvae then metamorphosed into the post-larval cystidean stage, which lasted for more than a month. The reproductive features of this species are unique among crinoids, and, together with juvenile morphological features observed in adults, can be interpreted as adaptations to their cryptic habitat, which is subject to frequent natural disturbances.     

Growth and mortality of subtidal red sea urchins (Strongylocentrotus franciscanus) at San Nicolas Island,California, USA: problems with models

Red sea urchins, Strongylocentrotus franciscanus, were tagged with tetracycline in 1990 at subtidal sites off San Nicolas Island, California, USA. After one year in the field, the sea urchins were collected and growth increments were measured based on tetracycline marks, which indicated initial slow growth, a maximum rate, and finally a prolonged period of very slow growth. Small red sea urchins (4 cm diam) were estimated to be 3 to 4 yr old, which is much older than has previously been reported. It is estimated that about 12 yr would be required to attain 10 cm diam. Survival has previously been modeled assuming a constant rate. If the population of red sea urchins is assumed to be stable and stationary, annual survival rate was between 71 and 77% yr^-1. Census data for the two years of study have permitted annual survival to be estimated without assuming stable and stationary population structure. Under these conditions, annual survival rate was between 79 and 86% yr^-1. Analysis of transitions in the size distributions from 1990 to 1991 suggested that annual survival may have been sizespecific: 91% yr^-1 for individuals 1.1 to 4.0 cm diam, 82% yr^-1 for individuals 4.1 to 7.0 cm diam, and 63% yr^-1 for those of 7.1 to 10.0 cm diam. An alternative explanation to size-specific survival in our study is sizespecific immigration.     

Effect of thermal stress and total residual chlorination on early life stages of the mummichog Fundulus heteroclitus

Effects of simultaneous short-term (7.5 to 60 min) thermal stress (24° to 34°C) and total residual chlorination (0.05 to 1.0 mg l^-1) on specific development stages of the mummichog Fundulus heteroclitus (Pisces: Cyprinodontidae), were investigated. For the embryonic stages, the total number of successfully hatched larvae was used as the criterion to measure effect. For the larval stages, survival 24 h after exposure was used. In the embryonic stages, temperature was the most important main variable. Only one embryonic stage (gastrula) was confounded by second-order interactions (temperature x duration of exposure x total residual chlorination). Both 0-day and 7-day-old larval stages showed significant higher-order interactions for all combinations of test parameters, suggesting the presence of synergistic effects of the three main experimental variables.Contribution No. 308 to the Gulf Breeze Environmental Research Laboratory. Contribution No. 183 to the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

The structure and sensitivity of the eye of different life history stages of the ontogenetic migrator Gnathophausia ingens

The structure and ultrastructure of the photoreceptors of several life history stages of the ontogenetically migrating lophogastrid crustacean Gnathophausia ingens were examined. The younger instars of this species live in a much brighter light field than the older instars, and this difference is reflected in differences in their visual systems. The shallowest free living individuals (instars 3 and 4) possess a superposition eye with almost no clear zone, which minimizes the light shared between ommatidia and reduces the sensitivity of the eye. A progression to superposition optics with a large clear zone, usually associated with night-active or deep-living species, occurs as the animals move deeper in the water column. Regional differences within the eye are also evident, with a largely nonexistent clear zone in the dorsal region and a large clear zone in the ventral region in the eyes of instar 5 animals, the first instar to move to deeper depths. The deepest living instars (10–12) possess superposition optics with a large clear zone throughout the eye, and are significantly more sensitive to light than the younger, shallower instars.     

Larval durations and recruitment patterns of two Caribbean gobies (Gobiidae): contrasting early life histories in demersal spawners

The population dynamics of small tiger prawns (Penaeus esculentus and P. semisulcatus) were studied at three sites around north-western Groote Eylandt, Gulf of Carpentaria, Australia, between August 1983 and August 1984. Seagrasses typical of open-coastline, reef-flat and river-mouth communities were found in the shallow depths (2.5 m) at these sites. The temperature and salinity of the bottom waters did not differ among the shallowest depths of the three sites and mean values at night ranged from 21.9 to 32.0 °C, and from 30.1 to 37.5% S. Data from fortnightly sampling with beam trawls showed that virtually all post-larvae (90%) were caught in the intertidal and shallow subtidal waters (2.0 m deep). At one site, where the relationship between seagrass biomass, catches and depth could be studied in detail, high catches were confined to seagrass in shallow water, within 200 m of the high-water mark. This was despite the fact that seagrass beds of high biomass (>100 g m^-2 between August and February) were found nearby, in only slightly deeper water (2.5 m). It is likely, therefore, that only the seagrass beds in shallow waters of the Gulf of Carpentaria act as important settlement and nursery areas for tiger prawns. In general, catches of tiger prawn postlarvae (both P. esculentus and P. semisulcatus) and juvenile P. esculentus on the seagrass in the shallowest waters at each site were higher in the tropical prewet (October–December) and wet (January–March) seasons than at other times of the year. Juvenile P. semisulcatus catches were highest in the pre-wet season. While seasonal differences accounted for the highest proportion of variation in catches of tiger prawn postlarvae and juvenile P. semisulcatus, site was the most important factor for juvenile P. esculentus. In each season, catches of juvenile P. esculentus were highest in the shallow, open-coastline seagrass, where the biomass of seagrass was highest. The fact that the type of seagrass community appears to be more important to juvenile P. esculentus than to postlarvae, suggests that characteristics of the seagrass community may affect the survival or emigration of postlarval tiger prawns. Few prawns (<10%) from the seagrass communities in shallow waters exceeded 10.5 mm in carapace length. Despite the intensive sampling, growth was difficult to estimate because postlarvae recruited to the seagrass beds over a long period, and the residence times of juveniles in the sampling area were relatively short (8 wk).     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

Larger foraminifera: Reproduction and early stages of development in Heterostegina depressa

The process of reproduction (multiple fission) of the megalospheric gamont of Heterostegina depressa d'Orbigny, 1826 (Nummulitidae) is described. During this process, the protoplasm separates outside the parent test into the juveniles. The initial chambers receive some of the symbiotic algae from the parent protoplasm. The second chamber develops from the initial chamber by division, not by growth. The newly formed 2-chamber-stage receives a second set of symbiotic algae from the parent residual protoplasm. Thereafter, calcification takes place, and the first ectoplasm sheath is formed. The problem of alternation of generations is discussed. From the very small proportion of microspheric individuals found in Heterostegina species (fossil and recent) it is concluded that gamogony has been greatly reduced.     

Effects of a transitory,low food supply in the early life of larval herring (Clupea harengus) on mortality,growth and digestive capacity

In order to assess possible effects of a transitory, low food supply on later development, three groups of Clyde herring larvae (Clupea harengus L.) were exposed in 1989 to different feeding regimes immediately after yolk resorption. Group 1 received a high daily ration of 80 copepods larvae^–1 for 31 d, Group 2 a low daily ration of 15 copepods larva^–1 for 10 d followed by a high ration (80 copepods larva^–1) for 21 d and Group 3 a low ration of 15 to 20 copepods larva^–1 for 31 d. After 31 d of feeding, digestive capacity, expressed as the sum of trypsin and trypsinogen, was markedly reduced in Group 2 compared to Group 1, while Group 3 had an even lower digestive capacity. After the switch from low to high ration Group 2 exhibited compensatory growth and caught up with Group 1 both in standard length and content of soluble protein. Group 3 had the lowest growth rates. Mortality was equal in Groups 1 and 2, while Group 3 showed an excess mortality of 40% of the start population. Although Group 2 larvae had caught up with Group 1 in growth at the end of the study, content of trypsin and trypsinogen in Group 2 was only half of that found in Group 1. Thus, comparing effects of a short period of food limitation on future growth, mortality and content of digestive enzymes, the study indicates content of trypsin and trypsinogen to be the most sensitive variable for detection of food limitation in the early stages of exogenous feeding.     

Differences in the early life history of the Australasian shortfinned eel Anguilla australis from Australia and New Zealand, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry were examined in juveniles of Australasian shortfinned eel, Anguilla australis, from Australia and New Zealand. Patterns in otolith increment widths were similar in these two stocks. A marked increase in otolith increment width was found from age 138 to 198 d (mean ± SD: 164 ± 18.6 d) from Australia, 161 to 208 d (185 ± 17.3 d) from the western coast of New Zealand and 161 to 211 d (187 ± 18.9 d) from the eastern coast. These changes coincided with a drastic decrease in otolith Sr:Ca ratios, suggesting that metamorphosis began at these ages in each species. The duration of metamorphosis estimated from otolith microstructure was the same in the different stocks, 17 to 41 d (mean ± SD: 27 ± 5.4 d). Ages at recruitment were 186 to 239 d (mean ± SD: 208 ± 17.4 d) from Australia, 214 to 263 d (232 ± 19.8 d) from the western coast of New Zealand and 208 to 266 d (237 ± 20.0 d) from the eastern coast. There was a significant difference in the age between the Australian stock and the two stocks from New Zealand. No significant difference was found in the age between the latter two stocks. The difference in the period for the recruitment between Australian and New Zealand stocks seems to reflect their geographical separation. Received: 1 March 1999 / Accepted: 19 July 1999     

Spore germination and early stages of development in Hypnea musciformis (Rhodophyta,Gigartinales)

The early stages of development of the carpospores and tetraspores of Hypnea muscilormis (Wulfen) Lamouroux have been investigated. Both types of spores germinated immediately after liberation. The spores segmented repeatedly into, 2, 3, 4 cells, etc. until a multicellular ball of cells was produced. The germination pattern was thus of the discal type, i.e., Typus discalis mediatus (Inoh, 1947) or the Dumontia-type in the terminology of Chemin (1937). Subsequently, a relatively expansive attachment holdfast was produced from each sporeling. This was followed by the development of up to 4 or more shoot axes arising from the basal disc. These findings are discussed with reference to an earlier study by the senior author on the developmental biology of other species of Hypnea Lamouroux.     

Prediction of extinction in plants: interaction of extrinsic threats and life history traits

The global extinction of species proceeds through the erosion of local populations. Using a 60-year time series of annual sighting records of plant species, we studied the correlates of local extinction risk associated with a risk of species extinction in the Park Grass Experiment where plants received long-term exposure to nutrient enrichment, soil acidification, and reductions in habitat size. We used multivariate linear models to assess how extrinsic threats and life history traits influence extinction risk. We investigated effects of four extrinsic threats (nitrogen enrichment, productivity, acidification, and plot size) as well as 11 life history traits (month of earliest flowering, flowering duration, stress tolerance, ruderalness [plant species' ability to cope with habitat disturbance], plant height, diaspore mass, seed bank, life form, dispersal mode, apomixis [the ability for a species to reproduce asexuall through seeds], and mating system). Extinction risk was not influenced by plant family. All of the 11 life history traits except life form and all threat variables influenced extinction risk but always via interactions which typically involved one threat variable and one life history trait. We detected comparatively few significant interactions between life history traits, and the interacting traits compensated for each other. These results suggest that simple predictions about extinction risk based on species' traits alone will often fail. In contrast, understanding the interactions between extrinsic threats and life history traits will allow us to make more accurate predictions of extinctions.