Preservation of inherent contractility in isolated gut segments from herbivorous and carnivorous marine fish

Patterns of inherent contractility were investigated in isolated gut segments from three species of New Zealand temperate-water labroid fish: the herbivorous butterfish Odax pullus (Family Odacidae), and the carnivorous banded wrasse Notolabrus fucicola and spotty Notolabrus celidotus (Family Labridae). To maintain the functional viability of gut tissue over extended periods, physiological solutions were formulated for each genus based on biochemical analyses of plasma constituents. A Res-Del perfusion system was used to maintain isolated gut segments under conditions of constant pH, temperature, and laminar flow dynamics. Gut segments exhibited spontaneous contractility over periods of days. Segments from N. fucicola and N. celidotus were continuously active, whereas segments from O. pullus showed discontinuous activity with evidence of daily rhythmicity. Contractions were slight in segments from O. pullus compared to the two Notolabrus species. Contractile strength correlated with gut morphology, with circular and longitudinal muscle layers markedly greater in extent in the Notolabrus species than in O. pullus. The pattern of gut activity in O. pullus is possibly associated with the retention of algal material in the intestine for lengthy periods. This hypothesis is supported by the daily feeding pattern of this species, and the probable involvement of fermentation in its digestive process. Accepted: 22 August 1997     

In situ settlement behaviour of damselfish (Pomacentridae) larvae

Settlement‐stage damselfish (Pomacentridae) larvae of 13 species in seven genera were obtained from light traps at Lizard Island, Great Barrier Reef, Australia. Behaviour, observed in situ by SCUBA divers, of 245 larvae (6–13 mm, L_S; 5–60 individuals per species) released individually within a few m of reefs during the day differed markedly among species. From 0–28% (range among 13 species) of individuals of each species swam away from the adjacent reefs without swimming to the reefs. Of those that swam to a reef, 0–75% settled. For three species, sufficient data were available to test the hypothesis that these percentages did not differ amongst reefs: the hypothesis was rejected in one species. From 0–75% of larvae that reached the reef were eaten, 0–63% subsequently left the reef and 0–60% were still swimming over the reef at the end of the observation period. Swimming speeds of all but one species were greater when swimming away from the reef than toward it. Most species exceeded average current speeds when swimming away from reefs, but not when swimming toward and over them. Average swimming depths were in the upper half of the water column for most species, and were somewhat greater where the water depths were greater. The time the larvae swam over the reef before settling and the distance swum varied greatly among species from 0 to a mean of 5.5 min and 43 m. Settlement habitats chosen differed amongst species, and in some species, they were very specific. Average settlement depth varied among species from 6–13.5 m. In one species, settlement depth varied between reefs. About half of the 53 observed interactions between larvae and reef resident fishes were predation attempts: fishes of eight species (six families) attacked larvae. The other interactions were aggressive approaches by 11 species of resident fishes, all but one of which were pomacentrids. Many of these aggressive interactions discouraged settlement attempts. Larvae of some species experienced no predatory or aggressive interactions, whereas in other species interactions averaged >0.6 per released larva. Species that swam more‐or‐less directly to settlement sites near the reef edge experienced more interactions. Even within the same family, settlement behaviour differed among species in nearly all measures.     

Settlement behaviour of larvae of the Stripey Snapper, <Emphasis Type="Italic">Lutjanus carponotatus</Emphasis> (Teleostei: Lutjanidae)

Larval behaviour is important to dispersal and settlement, but is seldom quantified. Behavioural capabilities of larval Lutjanus carponotatus in both offshore pelagic and reef environments at Lizard Island, Great Barrier Reef were observed in situ to determine if they were sufficient to influence dispersal. Offshore, larvae swam with higher directional precision and faster on the windward side of the island (28 cm.s⁻¹) than on the leeward side (16 cm s⁻¹). Most larvae swam directionally. Mean swimming directions were southerly in the windward area and northerly in the leeward area. Larvae avoided the surface and remained mostly between 3–15 m. Larvae released near reefs were 2–3 times faster swimming away from reefs (19 cm s⁻¹) than swimming toward or over them (6–8 cm s⁻¹). Speed swimming away was similar to that offshore. Of 41 larvae released near reefs, 73% reached the reef, 59% settled, and 13% of those reaching the reef were eaten. Larvae settled onto hard and soft coral (58%), topographic reef features (29%) and sand and rubble (13%). Settlement depth averaged 5.5 m (2–8 m). Before settling larvae spent up to 800 s over the reef (mean 231 s) and swam up to 53 m (mean 14 m). About half of the larvae interacted with reef residents including predatory attacks and aggressive approaches by residents and aggressive approaches by settling larvae. Settlement behaviour of L. carponotatus was more similar to a serranid than to pomacentrids. Settlement-stage larvae of L. carponotatus are behaviourally capable, and have a complex settlement behaviour.     

Reef Odor: A Wake Up Call for Navigation in Reef Fish Larvae

The behavior of reef fish larvae, equipped with a complex toolbox of sensory apparatus, has become a central issue in understanding their transport in the ocean. In this study pelagic reef fish larvae were monitored using an unmanned open-ocean tracking device, the drifting in-situ chamber (DISC), deployed sequentially in oceanic waters and in reef-born odor plumes propagating offshore with the ebb flow. A total of 83 larvae of two taxonomic groups of the families Pomacentridae and Apogonidae were observed in the two water masses around One Tree Island, southern Great Barrier Reef. The study provides the first in-situ evidence that pelagic reef fish larvae discriminate reef odor and respond by changing their swimming speed and direction. It concludes that reef fish larvae smell the presence of coral reefs from several kilometers offshore and this odor is a primary component of their navigational system and activates other directional sensory cues. The two families expressed differences in their response that could be adapted to maintain a position close to the reef. In particular, damselfish larvae embedded in the odor plume detected the location of the reef crest and swam westward and parallel to shore on both sides of the island. This study underlines the critical importance of in situ Lagrangian observations to provide unique information on larval fish behavioral decisions. From an ecological perspective the central role of olfactory signals in marine population connectivity raises concerns about the effects of pollution and acidification of oceans, which can alter chemical cues and olfactory responses.     

A 27-year study of brown trout population dynamics and exploitation in Lake Songsjøen, central Norway

From 1968–1984 (period I), a brown trout Salmo trutta , population in a 70-ha oligotrophic lake in central Norway was exploited using larger mesh gill-nets selectively removing the larger fish. From 1985–1994 (period II), intermediate sized fish were removed using smaller-mesh sizes gill-nets. Fishing mortality and CPUE were correlated positively with effort and numbers of fish >3 years old for period II. The gill-net catchability was correlated negatively with spawner biomass and number of trout >3 years old. The significant positive correlation between natural mortality and stock biomass and spawning stock biomass indicated density-dependent mortality. The significant correlation between spawning stock and recruitment described by the Ricker model, indicated density-dependent recruitment of 1-year-old trout. The fishing regimes in the two periods affected the population dynamics and density differently. Selective removal of smaller fish permitted the larger fish to survive, and was beneficial in reducing fish density and maintaining stocks at low levels, consequently, achieving the expected increase in fish growth rates.     

Hindgut Fermentation in Three Species of Marine Herbivorous Fish

Symbioses with gut microorganisms provides a means by which terrestrial herbivores are able to obtain energy. These microorganisms ferment cell wall materials of plants to short-chain fatty acids (SCFA), which are then absorbed and used by the host animal. Many marine herbivorous fishes contain SCFA (predominantly acetate) in their hindgut, indicative of gut microbial activity, but rates of SCFA production have not been measured. Such information is an important prerequisite to understanding the contribution that gut microorganisms make in satisfying the energy needs of the fish. We have estimated the rates of acetate production in the gut of three species of temperate marine herbivorous fish from northeastern New Zealand: Kyphosus sydneyanus (family Kyphosidae), Odax pullus (family Odacidae), and Aplodactylus arctidens (family Aplodactylidae). Ex vivo preparations of freshly caught fish were maintained with their respiratory and circulatory systems intact, radiolabeled acetate was injected into ligated hindgut sections, and gut fluid was sampled at 20-min intervals for 2 h. Ranges for acetate turnover in the hindguts of the studied species were determined from the slope of plots as the log of the specific radioactivity of acetate versus time and pool size, expressed on a nanomole per milliliter per minute basis. Values were 450 to 570 (K. sydneyanus), 373 to 551 (O. pullus), and 130 to 312 (A. arctidens). These rates are comparable to those found in the guts of herbivorous reptiles and mammals. To determine the contribution of metabolic pathways to the fate of acetate, rates of sulfate reduction and methanogenesis were measured in the fore-, mid-, and hindgut sections of the three fish species. Both rates increased from the distal to proximal end of the hindgut, where sulfate reduction accounted for only a small proportion (<5%) of acetate methyl group transformed to CO₂, and exceeded methanogenesis from acetate by >50-fold. When gut size was taken into account, acetate uptake from the hindgut of the fish species, determined on a millimole per day per kilogram of body weight basis, was 70 (K. sydneyanus), 18 (O. pullus), and 10 (A. arctidens).     

Length-weight relationship of fish species captured around an artificial offshore reef (northern Rio de Janeiro,Brazil)

The present work provides length-weight relationships (LWRs) for fish species captured around artificial offshore reef deployed 5 km from Guaxindiba Port, northern Rio de Janeiro, Brazil. The fish were captured during two sampling periods each year between 1996 and 2017 using bottom gill nets (25 m_length × 3 m_depth; 30 mm mesh). The fish were kept on ice and the biometric data, including total length (cm) and total wet weight (g) were determined in the laboratory. A total of 16 species belonging to 10 families were analyzed with Sciaenidae being the species richest family (5 spp) in the samples. The new value of LWRs for Sphyraena guachancho and new maximum sizes recorded for seven species highlight the scarcity of information on biological aspects of South America coastal fishes. These LWRs should assist fisheries scientist and managers to complement their further studies of population parameters to improve management decisions.     

Spatial variation and effects of habitat on temperate reef fish assemblages in northeastern New Zealand

Reef-associated fishes can respond to changes in habitat structure and the nature of their response can change with different spatial scales of observation. A structured hierarchical mensurative sampling design was used to sample temperate reef fish assemblages in northeastern New Zealand at several spatial scales over 2 years. The three spatial scales examined were tens of meters (transects), hundreds to thousands of meters (sites) and hundreds of kilometers (locations). We tested the hypothesis that fish assemblages differed between kelp forest habitat (relatively dense stands of the kelp, Ecklonia radiata (C. Agardh) J. Agardh, median depth=13.5 m) and barrens habitat (rocky reef dominated by turfing and encrusting red algae and the grazing urchin, Evechinus chloroticus (Valenciennes), median depth=6.7 m). Recently developed multivariate techniques were used to test for and quantify multivariate variation at different spatial scales. There were significant effects of habitat on the spatial distribution of fish assemblages, characterised by greater abundances or frequencies of Parika scaber, Chromis dispilus, Trachurus novaezelandiae, Nemadactylus douglasii, Bodianus unimaculatus, Odax pullus and Pseudolabrus miles in kelp forest habitat, and greater abundances or frequencies of Notolabrus celidotus, Notolabrus fucicola, Girella tricuspidata, Coris sandageri, Chironemus marmoratus, Parma alboscapularis, Scorpis violaceus and Kyphosus sydneyanus in barrens habitat. Some of the more common species, including Upeneichthys lineatus, Scorpis lineolatus and Cheilodactylus spectabilis showed no strong consistent effects of these two differing habitats on their distributions. There was, however, a significant Habitat×Locations interaction: effects of habitat did not occur at all locations. Variability was highest at the scale of individual transects and variability from site to site and from location to location was comparable. Spatial variation was large compared to inter-annual variation, which was minimal, and spatial patterns were consistent in the 2 years examined. Further experiments, including manipulations, are required to understand what mechanisms and processes might be driving these patterns. This study, coupled with results from previous studies, suggests that there may be a dynamic inter-play between effects of habitat on fish and effects of fish on biogenic habitat, such as kelp forests.     

Length‐weight relationships of Acanthopagrus longispinnis (Valenciennes, 1830), Raconda russeliana (Gray, 1831) and Coilia neglecta (Whitehead, 1967) from the Bay of Bengal coast,Bangladesh

Current study reports the length‐weight relationships (LWRs) of three marine fish species collected from the fishermen catches covering three seasons i.e., rainy, autumn and winter between June, 2016 and February, 2017. Fishermen caught the species using beach seine net (3.5 cm mesh size) and set bag net (0.5 cm mesh size) up to 200 m far away from the shoreline. The LWRs for fish species were W = 0.0180TL^3.192 for A. longispinnis, W = 0.0109TL^2.971 for R. russeliana and W = 0.0098TL^2.942 for C. neglecta with their r² values of 0.971, 0.962 and 0.997, respectively.     

Influence of habitat degradation on fish replenishment

Temperature-induced coral bleaching is a major threat to the biodiversity of coral reef ecosystems. While reductions in species diversity and abundance of fish communities have been documented following coral bleaching, the mechanisms that underlie these changes are poorly understood. The present study examined the impacts of coral bleaching on the early life-history processes of coral reef fishes. Daily monitoring of fish settlement patterns found that ten times as many fish settled to healthy coral than sub-lethally bleached coral. Species diversity of settling fishes was least on bleached coral and greatest on dead coral, with healthy coral having intermediate levels of diversity. Laboratory experiments using light-trap caught juveniles showed that different damselfish species chose among healthy, bleached and dead coral habitats using different combinations of visual and olfactory cues. The live coral specialist, Pomacentrus moluccensis, preferred live coral and avoided bleached and dead coral, using mostly visual cues to inform their habitat choice. The habitat generalist, Pomacentrus amboinensis, also preferred live coral and avoided bleached and dead coral but selected these habitats using both visual and olfactory cues. Trials with another habitat generalist, Dischistodus sp., suggested that vision played a significant role. A 20 days field experiment that manipulated densities of P. moluccensis on healthy and bleached coral heads found an influence of fish density on juvenile weight and growth, but no significant influence of habitat quality. These results suggests that coral bleaching will affect settlement patterns and species distributions by influencing the visual and olfactory cues that reef fish larvae use to make settlement choices. Furthermore, increased fish density within the remaining healthy coral habitats could play an important role in influencing population dynamics.     

In situ observation of settlement behaviour in larvae of coral reef fishes at night

The swimming behaviour of 534 coral reef fish larvae from 27 species was explored at Moorea Island (French Polynesia) while they searched for a suitable settlement habitat, on the first night of their lagoon life. Most larvae swam actively (74%) and avoided the bottom (77%). A significant relationship was highlighted between the vertical position of larvae in the water column and the distance they travelled from lagoon entrance to settlement habitat: larvae swimming close to the surface settled farther away on the reef than bottom-dwelling larvae.     

Movements of Fishes Within and Among Fringing Coral Reefs in Barbados

Movement of coral reef fishes across marine reserve boundaries subsequent to their initial settlement from the plankton will affect the ability of no-take reserves to conserve stocks and to benefit adjacent fisheries. However, the mobility of most exploited reef species is poorly known. We tagged 1443 individuals of 35 reef fish species captured in Antillean fish traps in the Barbados Marine Reserve and adjacent non-reserve over a two-month period. Trapping and visual surveys were used to monitor the movements of these fish during the trapping period and the subsequent two months. Estimates of distances moved were corrected for the spatial distribution of sampling effort and for the number of recaptures of individual fish. Recapture rates for individual species ranged from 0–100% (median=38%). Species mobility estimated by recapture and resighting were highly correlated. Most species were strongly site attached, with the majority of recaptures and resightings occurring at the site of tagging. However, only one of 59 tagged jacks (Caranx latus, C. ruber) was ever resighted, suggesting emigration from the study area. All species were occasionally recorded away from the sites where they had been tagged (20–500m), and several species, including surgeonfish, Acanthurus bahianus, A. coeruleus, filefish, Cantherhines pullus, butterflyfish, Chaetodon striatus, angelfish Holocanthus tricolor and parrotfish, Sparisoma viride, ranged widely within reefs. In contrast, few movements were observed between reefs separated by more than 20m of sand and rubble, and no emigration from the Reserve was recorded. Most reef fishes vulnerable to Antillean traps appear sufficiently site-attached to benefit from reserves. However, many species move over a wide enough area to take them out of small reserves on continuous reef. Use of natural home range boundaries could minimize exposure of fishes in reserves to mortality from adjacent fisheries.     

Nocturnal orientation to reefs by late pelagic stage coral reef fishes

 The nocturnal orientation behaviour of the late pelagic stages of two reef fish families (Apogonidae and Pomacentridae) was examined using behavioural cages deployed in the field. The behavioural cages enabled the fish to choose between swimming towards or away from the reef in response to natural cues. Overall, 55% of fish displayed a choice in the experiments, however, the proportion varied between the two families, with 67% of pomacentrids and 27% of apogonids displaying a choice. In both families, of the fish which displayed a choice, the proportion of fish swimming towards the reef was significantly greater than 50%, as random movement would predict (64% of pomacentrids and 67% of apogonids swam towards the reef). This proportion did not vary significantly among four field sites with different current regimes and geographic locations. The results suggest that the late pelagic stages of reef fish display nocturnal orientation behaviour, possibly in response to sound, which may aid in their settlement on reefs. Accepted: 18 August 1997     

Locomotion at –1.0°C: burst swimming performance of five species of Antarctic fish

We investigated the burst swimming performance of five species of Antarctic fish at −1.0°C. The species studied belonged to the suborder, Notothenioidei, and from the families, Nototheniidae and Bathydraconidae. Swimming performance of the fish was assessed over the initial 300 ms of a startle response using surgically attached miniature accelerometers. Escape responses in all fish consisted of a C-type fast start; consisting of an initial pronounced bending of the body into a C-shape, followed by one or more complete tail-beats and an un-powered glide. We found significant differences in the swimming performance of the five species of fish examined, with average maximum swimming velocities (U_max) ranging from 0.91 to 1.39 m s⁻¹ and maximum accelerations (A_max) ranging from 10.6 to 15.6 m s⁻². The cryopelagic species, Pagothenia borchgrevinki, produced the fastest escape response, reaching a U_max and A_max of 1.39 m s⁻¹ and 15.6 m s⁻², respectively. We also compared the body shapes of each fish species with their measures of maximum burst performance. The dragonfish, Gymnodraco acuticeps, from the family Bathdraconidae, did not conform to the pattern observed for the other four fish species belonging to the family Nototheniidae. However, we found a negative relationship between buoyancy of the fish species and burst swimming performance.     

Length-Weight Relationship analysis of four freshwater fish Species from Doyang reservoir at Nagaland,India

The LWRs (length–weight relationships) of four freshwater fish species from medium categorized Doyang reservoir at Wokha District, Nagaland were studied, namely as Garra naganensis Hora, 1921; Devario naganensis Chaudhuri, 1912; Chagunius nicholsi Myers, 1924 and Schistura manipurensis Chaudhuri, 1912. Fish sample were collected in every 15 days interval by cast-nets (1.0 × 1.0 cm mesh size covering the 4.5 m² areas), gill-nets (0.4–5.0 cm mesh size), scoop-nets (mesh size with 0.3 × 0.3 cm to 0.6 × 0.6 cm) and also from the local fishermen in between July 2017 to January 2019. The regression value of slope (b) is fallen into 2.860–3.115 range with significant correlation values (r² ≥ .960).     

Length–weight relationship of five coral reef fish from the Weizhou Island,China

In the present study, the LWRs investigations were conducted for five fish species from Weizhou Island for four times seasonally during the period from April 2017 to January 2018. Samples were obtained using gillnets (25 × 12 m, mesh size 0.5 cm), trawl nets (50 × 12 m, mesh size 0.5 cm) and fishhooks (0.5 × 0.5 × 10 m, mesh size 0.5 cm mesh). Results indicate that mean LWR parameters b for these five coral reef fish varied from 2.847 to 3.525 and r² values varied from 0.972 to 0.988.     

Ontogenetic changes in the critical swimming speed of Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin) larvae and the role of temperature

Most studies on behavioural contributions to dispersal and recruitment during early life history stages of fishes have focused on coral reef species. For cold ocean environments, high variation in seasonal temperature and development times suggest that parallel studies on active behaviour are needed for cold-water species. Thus, we examined the critical swimming speed (U_crit) of marine fish larvae from 2 contrasting species: Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin), a pelagic and bottom spawner respectively. Within-species comparisons showed that sculpin reared at 6 °C had lower initial U_crit values, but a faster U_crit increase through development compared with 3 °C conspecifics, ultimately resulting in faster critical swimming speeds at metamorphosis (10.5 vs. 9.1 cm·s^− 1). In contrast, although cod larvae reared at 10 °C were faster swimmers at first feeding than 6 °C fish, temperature differences were absent after the first week. These results show that temperature influences the trajectory of larval critical swimming speed development, but that the relationship is species-specific. Although 6 °C sculpin and cod of similar length had equivalent U_crit values, the smaller size of cod at hatch (5.3 vs. 10.8 mm for sculpin) resulted in much lower age-specific U_crit values for cod. These data have significant implications for how swimming activity of the two species might affect dispersal, particularly in the first few weeks post-hatch. Overall, our data suggest that temperature during larval development influences the swimming capacity of cold-water marine fishes, and has important ramifications for biophysical models of dispersal.     

The Mangrove Nursery Paradigm Revisited: Otolith Stable Isotopes Support Nursery-to-Reef Movements by Indo-Pacific Fishes

Mangroves and seagrass beds have long been perceived as important nurseries for many fish species. While there is growing evidence from the Western Atlantic that mangrove habitats are intricately connected to coral reefs through ontogenetic fish migrations, there is an ongoing debate of the value of these coastal ecosystems in the Indo-Pacific. The present study used natural tags, viz. otolith stable carbon and oxygen isotopes, to investigate for the first time the degree to which multiple tropical juvenile habitats subsidize coral reef fish populations in the Indo Pacific (Tanzania). Otoliths of three reef fish species (Lethrinus harak, L. lentjan and Lutjanus fulviflamma) were collected in mangrove, seagrass and coral reef habitats and analyzed for stable isotope ratios in the juvenile and adult otolith zones. δ¹³C signatures were significantly depleted in the juvenile compared to the adult zones, indicative of different habitat use through ontogeny. Maximum likelihood analysis identified that 82% of adult reef L. harak had resided in either mangrove (29%) or seagrass (53%) or reef (18%) habitats as juveniles. Of adult L. fulviflamma caught from offshore reefs, 99% had passed through mangroves habitats as juveniles. In contrast, L. lentjan adults originated predominantly from coral reefs (65–72%) as opposed to inshore vegetated habitats (28–35%). This study presents conclusive evidence for a nursery role of Indo-Pacific mangrove habitats for reef fish populations. It shows that intertidal habitats that are only temporarily available can form an important juvenile habitat for some species, and that reef fish populations are often replenished by multiple coastal habitats. Maintaining connectivity between inshore vegetated habitats and coral reefs, and conserving habitat mosaics rather than single nursery habitats, is a major priority for the sustainability of various Indo Pacific fish populations.     

Long-term memory retention in a wild fish species Labroides dimidiatus eleven months after an aversive event

Memory is essential to enhance future survival and reproduction as it helps in storing and retrieving useful information to solve particular environmental problems. However, we lack quantitative evidence on how far animals in the wild can maintain given information for extended periods without reinforcement. Here, we document correlative evidence of cleaner fish Labroides dimidiatus remembering being caught in a barrier net for up to 11 months. In 2015, about 60% of cleaners from one large isolated reef had been used for laboratory experiments and then returned to their site of capture. Eleven months later, 50% of cleaners at the same site showed an unusual hiding response to the placement of the barrier net, in contrast to three control sites where no cleaners had been caught during the last 2 years. The results suggest that a single highly aversive event (i.e., being caught in a barrier net) resulted in cleaners storing long-term crucial information that allowed them to avoid being caught again. Our results further our knowledge of fish cognitive capacities and long-term memory retention.     

Differences in swimming pattern between life cycle stages of the toxic dinoflagellate Alexandrium fundyense

Different life stages of Alexandrium fundyense have different swimming behavior; gametes often are said to “swarm” or “dance” before mating. This behavior was studied, and quantitative measures of these motility patterns in two-dimensions were generated using motion-analysis software applied to video records of individual-cell movements. Behavior, swimming patterns, and growth were studied in two strains of A. fundyense and compared in encystment medium and growth medium. Vegetative cells swam straight, rotating around the apical axis until they hit something and then swam straight in a different direction. Gamete swimming behavior was slower and characterized by frequent direction changes and circular motion. Gametes contacted other cells frequently (>5 cell contacts min⁻¹ cell⁻¹). Zygotes swam slowly when newly formed and later became nearly immobile; these cells continued to contact other cells and also surfaces. The results are in accordance with field observations of long swimming distances for vegetative cells, accumulation in thin layers of gametes, and sinking of developing resting cysts attached to marine snow for zygotes.