Effects of predators on sea urchin density and habitat use in a southern California kelp forest

Grazing sea urchins can reduce kelp abundance and therefore strongly affect kelp forest community structure. Despite the ecological importance of sea urchins, direct field studies on the role that urchin predators play in shaping urchin populations are rare for southern California. We conducted surveys and manipulative experiments within kelp forests near San Diego, CA, (32–51′28″N, 117–16′00″W) from 2006 to 2009 to determine whether predators such as sheephead (Semicossyphus pulcher) and spiny lobsters (Panulirus interruptus) may be linked to purple urchin (Strongylocentrotus purpuratus) and red urchin (Strongylocentrotus franciscanus) distribution and habitat use, as well as purple urchin density-dependent mortality. Purple urchins were less dense and more cryptic inside a local marine protected area (MPA) that contained high predator abundance than in nearby heavily fished areas, whereas red urchins rarely were found outside the MPA. Urchin proportional mortality was inversely density dependent during the day when sheephead were active, despite fish aggregations in plots of high urchin density, but was density independent during the night when lobsters were active. Urchin mortality was reduced under understory algal cover during the day, but not during the night. Examining whether urchin mortality from predation is density dependent and how habitat complexity influences this relationship is imperative because behavioral changes and increases in urchin populations can have vast ecological and economic consequences in kelp forest communities.     

Expansion of a central California kelp forest following the mass mortality of sea urchins

The mass mortality by disease of a localized population of sea urchins, Strongylocencrotus franciscanus, on the seaward side of a kelp forest was followed by the rapid seaward expansion of 4 species of brown algae, Macrocystis pyrifera, Laminaria dentigera, Pterygophora california, and to a lesser extent, Nereocystis leutkeana. One other brown alga, Cystoseira osmundacea, failed to become established in the newly available area. Competition among M. pyrifera, L. dentigera, P. californica, and N. Leutkeana apparently was severe, and within 1 year after the demise of the sea urchins, M. pyrifera formed a dense, nearly monospecific stand. Experimental removal of M. pyrifera demonstrated that the canopy of these plants limited light penetration to levels below that necessary for the growth and survival of other brown and red algae.     

Sexual reproduction in the corallimorpharian sea anemone Corynactis californica in a central California kelp forest

The seasonal cycle of sexual reproduction in the corallimorpharian sea anemone Corynactis californica (Calgren, 1936) was studied for 18 consecutive months (July 1987 to December 1988) at a subtidal area in the Hopkins Marine Refuge (HMLR), Pacific Grove, Monterey Bay, California. Samples were collected, histological sections were prepared, and gametocytes were examined and measured. C. californica grows by multiple or longitudianl fission to form single-sexed clones of various color and size. In female clones, oocytes appeared in late August and early September; they increased in size steadily in fall, and peaked in early December in both years. In male clones, spermatogenesis was synchronous with the female oogenic cycle, and motile sperm were observed in most testes in early December. Spawned gonads were found in both sexes from late November to early December. Spawning was induced in the laboratory in early December, and external fertilization was followed by development of free-swimming larvae. Gametogenesis and spawning are correlated with seasonal increases in seawater temperature and phytoplankton abundance, and we discuss and compare timing of sexual reproduction in this corallimorpharian to those in several actiniarian and scleractinian species.     

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.     

雨雪冰冻灾害后粤北森林各林型凋落物动态

为了考察中国亚热带不同森林类型对雨雪冰冻灾害的响应模式,以粤北天井山3种代表性的林型—针叶林、阔叶林和混交林为对象,于不同森林类型中比较受损森林与未受损森林在凋落物年产量、成分及月际动态方面的差异,从而在凋落物水平上反映不同森林类型在雨雪冰冻灾害后的早期恢复力。研究结果表明,灾后针叶林、阔叶林和混交林的年凋落量分别为0.52、3.21、1.37 t.hm-2,比未受损的同种森林类型年凋落量显著减少,减少程度分别为87.89%、53.46%、76.78%。由此可以看出阔叶林的植被恢复情况最好,说明在凋落物水平上,其灾后恢复的早期阶段恢复力最强。在凋落物成分方面,灾后各森林类型叶凋落物所占比例显著增加,枝凋落物所占比例则显著减少。受损针叶林和阔叶林的凋落物月动态与未受损森林基本一致,但其波动幅度较小;在混交林中,受损和未受损森林其凋落量的季节动态模式则表现出不一致性且为不规则型。根据研究结果,建议在亚热带地区优先考虑种植阔叶林以促进受损森林在类似雨雪冰冻灾害的极端天气后的恢复。     

Day-night differences in the emergence of demersal zooplankton from a sand substrate in a kelp forest

Day-night differences in abundance and biomass of demersal zooplankton in the water column were determined by trapping these animals as they emerged from the sand substrate in a kelp forest (Macrocystis pyrifera) ecosystem off Santa Catalina Island, California, USA. The day and night sampling periods of the 24 June 1979 new moon each lasted 12 h. Abundance and biomass of total demersal zooplankton were significantly higher in night samples. A mean of 2,425±1,168 demersal zooplankton m^-2 24 h^-1 migrated over a diel cycle; 97% of these animals were crustaceans. The mean biomass of demersal zooplankton was 94.2±27.6 mg ash-free dry wt m^-2 24 h^-1. No significant differences were found in either the abundance or biomass of demersal zooplanktion collected in low and high traps, suggesting that most animals collected 25 cm off the bottom can sustain swimming to at least 75 cm and that both traps give comparable estimates of the amount of demersal zooplankton available to planktivorous predators.     

Nutrient-limited growth of juvenile kelp,Macrocystis pyrifera,during the 1982–1984 “El Niño” in southern California

The relative growth rates of juvenile Macrocystis pyrifera in southern California kelp forests were substantially reduced during the El Niño of 1982–1984. The lower growth rates were correlated with increased temperature and decreased nitrogen availability. Fertilization of juvenile plants with slow-release nitrogen-phosphorus fertilizer increased their growth rates to levels previously observed when temperatures were low and nutrient levels were high. The limitation in growth of M. pyrifera by levels of available nutrients during El Niño was in contrast to the usual limitation of growth by irradiance during non-El Niño years. Thus, there was a shift in the relative importance of factors controlling growth of juvenile M. pyrifera during El Niño.     

Morphology and growth of the giant kelp Macrocystis pyrifera in New Zealand and California

At three sites (Papanui Inlet, Aquarium Point and Seacliff) on the Otago coast of New Zealand and at one site (Point Loma) off southern California, morphometric and shortterm growth measurements were made of single fronds of plants of Macrocystis pyrifera (L.) C. Ag. near the time of the autumn equinox, in 1977. Near the apex, internodes and laminae were shorter in wave-exposed Seacliff and California populations than in those of Papanui Inlet and Aquarium Point in strong currents, although lamina thickness was similar. In the immature region near the apex, linear relationships were apparent from double logarithmic plots of stipe distance from apex vs node number, internode fresh weight vs length (different in the two areas), lamina width vs length, fresh weight vs lamina area and immature lamina, pneumatocyst and internode fresh weight vs node number. Morphologically, Seacliff plants resembled those in California more closely than those in the other New Zealand populations. The relative growth rate in length of stipe internodes decreased linearly with the logarithm of the distance from the apex, but was less closely related to node number. Extrapolation gave zero growth at 1.6 m at Aquarium Point and 2.9 m in California, where the internode lengths coincided with a change in their weight/length relationship. Slow relative growth proximal to these points indicated a different type (mainly elongation). It is suggested that the first, apical, type of growth, being apparently unrelated to frond length, would be a useful measure of activity over short time-periods. The relative growth rate in lamina area (and probably volume), while also related to distance from the apex, showed a very different pattern, increasing slightly to a peak at 100 to 200 mm and declining to nil by 1 m at Aquarium Point and 2 m in California. Maximal lamina size was maintained for only 10 to 15 nodes at Aquarium Point. Calculated node initiation frequencies were 2.7 d at Papanui Inlet, 2 d at Aquarium Point, and 1 d in California. At the latter sites it took about 2 mo for laminae and 2^1/2 mo for stipe internodes to mature. Deduced relative growth rates in stipe tissue and pneumatocyst fresh weight decreased from 0.24 (log_e units per day) at the apex to 0.11 at 200 mm, where basal lamina tissue had a rate of 0.29, a doubling in just over 2 d. The production of tissue per frond was estimated as 25 and 36 g fresh wt d^-1 at Aquarium Point and in California, respectively.     

Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.     

Non-indigenous ascidians in southern California harbors and marinas

Southern California's many large harbors form an important coastal ecosystem, yet they are also a␣major destination for thousands of pleasure craft and cargo vessels that have often traveled great distances. Many groups of marine organisms, including ascidians, have now been documented as undergoing range extensions as a consequence of rapid ship-transport between distant harbors phenomenon. This has resulted in a rapid increase in the rate of introductions of non-indigenous species worldwide, yet these effects of boat traffic remain largely unstudied in southern California. Ascidians are sessile marine filter-feeders, hermaphroditic, and often self-fertilizing; many species are tolerant of a wide range of environmental conditions, can reach sexual maturity in just a few weeks, and have a long breeding season. This paper documents the arrival of 14␣non-indigenous species in southern California harbors␣during this century, 13 of which have persisted:␣four prior to the 1960s (Cionaintestinalis, Styelaclava, S.␣plicata, Botryllusschlosseri), another by 1972 (S.␣canopus, formerly S. partita), and 8 since 1983 [C.␣savignyi, Ascidia zara, Ascidia sp., Polyandrocarpa zorritensis, Symplegma brakenhielmi (formerly S. oceania, and S. reptans, Microcosmus squamiger, and Molgula␣manhattensis)]. We estimate the relative abundance and seasonal fluctuations of both non-indigenous and native ascidians in all harbors in southern California from San Diego to Santa Barbara based upon the historical record, our 35 yr of field notes, and our recent surveys carried out during fall 1994, spring and fall 1995, fall 1996 and spring 1997. Possible points of origin of the exotics and predictions on further U.S. Pacific coast range-extensions are included. The concomitant decline in numbers and species of native ascidians in the harbors of southern California during this century is also reviewed. Received: 4 March 1997 / Accepted: 26 September 1997     

Behavioral aspects of competition in a three-species rodent guild of coastal southern California

Summary The influence of vole cycles on the demography, spatial organization, and abundance of potential rodent competitors was studied in three California rodents in the laboratory and in the field. Population densities ofMicrotus californicus were inversely correlated with reproductive success in two potential competitors (Reithrodontomys megalotis andMus musculus). Additionally, high-density vole populations forced these species into suboptimal habitats. During low-density vole populations,Reithrodontomys andMus left their refugia, expanded their usage of habitat, were more active, and their reproductive success was greater. Thus, this work suggests that during vole population peaks, competition with sympatric species of mice is severe. Conversely, when vole populations decline, competition is relaxed. The temporal and spatial organization of the three-species complex supports models of nonequilibrium and fugitive species coexistence.     

Movement and feeding activity of red sea urchins (Strongylocentrotus franciscanus) adjacent to a kelp forest

Movement and feeding were studied in a population of red sea urchins, Strongylocentrotus franciscanus (Agassiz, 1863), found within and immediately seaward of a kelp forest offshore from Santa Cruz, California, USA. Mean sea urchin movements varied from 7.5 cm/day inside the kelp forest to over 50 cm/day at 15 and 100 m outside the kelp forest. The percentage of sea urchins feeding decreased from 66% inside the kelp forest to 16 and 15% at 15 and 100 m outside the kelp forest. These data indicate that movement by these sea urchins is a response to a low food supply.     

Regulation of algal community development in a Macrocystis pyrifera forest

The effects of small and large-scale roughness, overstory development, competition for space with sessile animals, and grazing on algal community development in a subtidal Macrocystis pyrifera forest were examined using specially prepared concrete blocks as substrata. Variation in small-scale roughness (crevices and grooves in the order of 0.1 to 3 mm width and depth) had no significant effects on community composition. However, M. pyrifera colonization, algal diversity, and sessile animal biomass were higher near the upper horizontal edges of blocks and concrete prisms. This “edge” effect may result from a combination of increased spore and larval settlement and enhanced growth of plants and animals associated with the turbulent eddies formed around these obstructions. Natural and experimentally produced variations in the algal overstory demonstrated that the presence of an overstory can reduce algal diversity and cover beneath. Caging experiments suggest that predatory fishes and sea-stars indirectly affect the algal community by removing sessile animals (primarily bryozoans) which compete with the algae for space. The exclusion of grazers resulted in increased growth of Gigartina spp. Selective grazing on this genus may account for its reduced abundance in the study area.     

Foliose algal assemblages and deforested barren areas: phlorotannin content,sea urchin grazing and holdfast community structure in the Aleutian dragon kelp,Eualaria fistulosa

In the Aleutian Archipelago, two distinct organizational states of kelp forest communities exist, foliose algal assemblages and deforested barren areas. The canopy-forming kelp Eualaria fistulosa can be found in both states, although it is much less abundant in the deforested state. In contrast, sea urchins also occur in both states, but they are considerably more abundant in the deforested state. This study determined whether the protective phlorotannin content in E. fistulosa sporophylls that originated in foliose algal assemblages would differ from those that originated in deforested areas. We also examined sea urchin grazing rates on these sporophylls to determine whether there were any differences in grazing rates related to the origin of the sporophylls. Next, we determined whether taxon richness and abundance of E. fistulosa holdfast communities would differ depending on the origin of the holdfast. Our results showed that sporophylls that were collected in barren areas generally had higher phlorotannin content than sporophylls found in foliose algal assemblages, although phlorotannin content varied across the study area. Grazing rates on sporophyll tissue and holdfast community structure did not differ between foliose algal assemblages and barren areas. These results show that phlorotannin content is greatest in areas under high grazing pressure and that phlorotannins may possibly protect E. fistulosa and the holdfast communities from being grazed in barren areas.     

Variations in benthic community structure in a sub-tropical estuary

Abundances of macroinfaunal invertebrates in sand dollar (Dendraster excentricus) beds and in adjacent areas free of sand dollars were studied at 10 intertidal sites in the Pacific Northwest region of the USA and Canada. Each site was sampled once in late summer of 1977. There were no significant differences in overall diversity of species between the beds and adjacent areas; nor did cluster analysis indicate a sand dollar bed community. However, a polychaete (Armandia brevis) was significantly more abundant inside the beds than just outside them. Conversely, in comparison to the beds, adjacent regions contained significantly more specimens of the following species: a bivalve (Transennella tantilla), several tubicolous crustaceans (Corophium spp., Ampelisca agassizi, and Leptochelia savignyi) and two polychaetes (Glycinde polygnatha and Malacoceros arenicola). Sediment characteristics at each study site were not correlated with the presence or absence of sand dollars; moreover, a comparison among the 10 sites revealed no sediment characteristics typical of sand dollar beds. At most of the sites, significantly fewer tube-building invertebrates occurred inside sand dollar beds than outside. The discussion considers the possibility that mechanical disturbance by the sand dollars tends to exclude some macroinvertebrates (especially tubicolous ones) from sand dollar beds.     

Variability over space and time in the genetic structure of the winged kelp Alaria marginata

The genetic structure of Alaria marginata Postels & Ruprecht was investigated spatially and seasonally using amplified-fragment length polymorphisms (AFLP). Using one primer, 206 scoreable bands were produced. Individual plants that were separated by as little as a few decimeters to >100 km could be distinguished, and followed an isolation-by-distance model. Genetic similarity (average April data) ranged from 76% for patches (a few decimeters in diameter), to 71% for individual kelp stands (15 m across) and 67% for a group of stands separated by 185 km. Greater genetic similarity of patches occurred at a wave-sheltered site than at a wave-exposed site. The lower wave action may limit meiospore dispersal, resulting in patches of related sporophytes. Genetic similarities between A. marginata stands indicated a pattern of gene flow that is consistent with local currents. In one stand, genetic similarities were markedly different between seasons. This seasonal pattern may be the result of different age groups dominating the sampled stands over time. The genetic structure of A. marginata comprises local scale (patch and within-stand) heterogeneity and larger scale (between-stands) homogeneity. Received: 30 September 1998 / Accepted: 26 October 1999     

Apparent predation on ichthyoplankton by zooplankton and fishes in nearshore waters of southern California

Plankton collected at discrete depths in Santa Monica Bay, California, USA, during January 1982 were examined for fish eggs and larvae that had been attacked or consumed by zooplankton. The bongo net remained open for only 3 min and samples were preserved within 5 min of capture. Juvenile and adult fishes that had been captured by otter trawl and preserved within 20 min of capture were examined for ingested fish eggs and larvae. Three copepods (Corycaeus anglicus, Labidocera trispinosa, and Tortanus discaudatus), one euphausid larva (Nyctiphanes simplex), one amphipod (Monoculoides sp.), and an unidentified decapod larva were found attached to fish larvae in the preserved plankton samples (attachment to 23% of the fish larvae was observed in one sample). Overall, about 5% of the white croaker (Genyonemus lineatus) larvae and 2% of the northern anchovy (Engraulis mordax) larvae had attached zooplankton predators. Most fish larvae with attached zooplankton predators were small. We found no indication of zooplankton predation on fish eggs. Few fish eggs and larvae were found in the digestive tracts of juvenile or adult fishes, and the ingested fish larvae were relatively large. The discussion considers apparent preyspecificity of the zooplankton predators as well as potential biases that may be associated with preserved samples collected by nets.