Spatiotemporal coupling/decoupling of planktonic larvae and benthic settlement in decapods in the Scottish east coast

Settlement patterns and the relationship between meroplanktonic larvae and settlement in decapods were studied on the Scottish east coast. Artificial settlement substrates (ASS), deployed at two locations (sandy vs. rocky sea substrates), were employed to collect megalopae and newly settled juveniles. Abundance of meroplanktonic larvae was used as an indicator of larval supply. The results showed a clear seasonality in settlement rates, and in some cases, significant differences between sites were detected. Nevertheless, the interference of the ASS with the surrounding habitat limits the study of spatial variability in settlement rates. Significant cross-correlation was found between the abundance of megalopae and juveniles in the collectors and planktonic larval abundance a month earlier. For individual species, this relationship was observed only in Pisidia longicornis. Complexities caused by the great variety of pre- and post-settlement processes, alongside effects of secondary dispersals of early juveniles may have obscured the relationship between meroplanktonic larvae and juveniles in other species.     

Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

<Emphasis Type="Italic">Semibalanus</Emphasis><Emphasis Type="Italic">balanoides</Emphasis> in winter and spring: larval concentration,settlement, and substrate occupancy

This study measured the progression from pelagic larvae to juvenile barnacles, and examined whether recruitment of barnacles, Semibalanus balanoides Linnaeus, at two intertidal sites in contrasting hydrodynamic regimes was determined by pre-settlement or post-settlement processes. The two sites were 1.5 km apart in the vicinity of Woods Hole, Mass., USA. Quantitative plankton samples were taken twice weekly from December 1997 to May 1998 at a nearby site as an estimate of nearshore larval abundance. The presence of S. balanoides nauplii was noted, and cyprids were enumerated and measured. Larval settlement at the two sites [Gansett Point, Buzzards Bay (GP) and Little Harbor, Vineyard Sound (LH)] was estimated from examination of replicate settlement plates exposed for 2 or 3 days throughout the settlement season, and from replicate plots on marked rock quadrats at each site. On both plates and rocks settled cyprids and metamorphs were enumerated. Space occupancy on unmanipulated rock quadrats by all stages from cyprids to adult barnacles was also examined. Settlement occurred from 2 January to 20 May, and major settlement peaks coincided with peaks in pelagic cyprid concentration at LH, but not at GP. Space occupied by juvenile barnacles was close to zero up until late February despite substantial settlement prior to that. At LH, juvenile barnacle cover was zero at the end of the observations; all settlement failed. Almost 100% of settled cyprids failed to metamorphose within 2 days from late January to late March. Then the proportion metamorphosing increased sharply coinciding with a sudden increase of 3°C in water temperature. Observed site differences in space occupancy by juvenile barnacles suggest that while cyprid supply is a necessary condition for barnacle settlement, other factors affecting metamorphosis of settled cyprids and early juvenile mortality determine recruitment.     

Effects of eutrophication and sedimentation on juvenile corals

Settlement of juvenile scleractinian corals was investigated from 1987 to 1990 on eutrophic and less eutrophic fringing reefs on the west coast of Barbados, West Indies. The number of coral recruits and number of recruiting coral species on cement blocks decreased with increasing eutrophication of the reefs. This may suggest lower settlement rates on eutrophic reefs, but could also liave resulted from higher post-settlement mortality, since blocks were examined only once after 3 yr of immersion. Coral settlement rates to artificial plates that were checked monthly were also lower on the more eutrophic reefs. This could result from lower local availability of larvae caused by fewer adult corals and/or lower reproductive rates of corals on eutrophic reefs. However, the ratio of coral recruits to adult coral abundance was considerably lower on eutrophic reefs, suggesting that local coral abundance alone can not explain lower settlement rates on eutrophic reefs. The lower rates on eutrophic reefs may result from a lower probability of coral larvae settlin when present, perhaps because of a limited availability of suitable settlement substrate. Colonization of settlement plates by non-coralline organisms was heavier on eutrophic reefs, and unoccupied space was lower, supporting the suggestion that suitable coral settlement substrate may be limiting on eutrophic reefs. Moreover, coralline algae, which facilitate metamorphosis and settlement of coral larvae, were less abundant on settlement plates on eutrophic reefs.     

Temporal patterns of settlement, recruitment and post-settlement losses in a rocky reef fish assemblage in the South-Western Mediterranean Sea

Post-settlement events can significantly alter the density distribution of settlers and subsequently the adult population structure. The temporal and inter-annual variability of settlement and the effects of mortality on recruitment were investigated across 2 years in the vicinities of Cabo de Palos–Islas Hormigas Marine Reserve by visual census and light trap sampling. Settlement was seasonal with greater species richness and abundance in summer. Although temporal synchronization was observed between larval supply and settlement, densities of settlers could not be predicted from post-larval abundances. Timing of settlement was consistent between years but with high inter-annual variation in abundance. High mortality (~80 %) and general decoupling between post-larval and settlement stages suggest that early mortality is driving such patterns. Nevertheless, indications of habitat-mediated mortality were found for benthic species such as Symphodus and Diplodus, highlighting the importance of habitat in shaping population demography.     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Planktonic larval duration and settlement marks on the otoliths of Mediterranean littoral fishes

The planktonic larval duration (PLD) was estimated for 42 species of littoral fishes from the north-western Mediterranean Sea. Daily increments and settlement marks on the otoliths (sagittae or lapilli) of new settlers and post-settlers were used to determine the larval stage duration. We also used PLD in the new settlers of some species to confirm the accuracy of the settlement marks in post-settlers. The duration of the planktonic larval stage ranges from 9 days in Symphodus ocellatus to 55 days in Xyrichtis novacula and 71 days in Lipophrys trigloides. Species in the same family did not display any clear tendency toward having similar PLDs. On the other hand, larval duration tended to be similar within a genus, with the exception of Lipophrys. Among conspecifics, the time spent in the plankton usually varied only by 2-7 days, except in Aidablennius sphynx, Lipophrys trigloides, Coris julis and Thalassoma pavo. No clear patterns were discernible in genera, with some species that settled in winter and other species that settled in summer, although we observed a certain tendency of individuals of closely related species (e.g. family Sparidae) to have a shorter larval duration in the warmer part of the year than in the colder part of the year. Settlement marks have been observed on the otoliths of all the species studied, and the PLDs in new settlers are an appropriate means of validating settlement marks. A rapid decrease in increment width over settlement (type Ia) is the most common type of mark (66.7% of the total species studied).     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Seasonality,lunar periodicity of settlement and microhabitat association of juvenile humpback red snapper <Emphasis Type="Italic">Lutjanus gibbus</Emphasis> (Lutjanidae) in an Okinawan coral reef

Snappers (Lutjanidae) are one of the important fisheries resources in tropical and subtropical waters. However, there have been few studies clarifying the ecology of newly settled juvenile snappers. The aims of the present study were to clarify the seasonality and lunar periodicity of the larval settlement, and the microhabitat association of newly settled juveniles for Lutjanus gibbus in an Okinawan coral reef. Fifteen coral patches were chosen and underwater visual surveys were conducted during a 2-year period. The larval settlement of the species mainly occurred between May and October during the 2-year survey period. In terms of lunar periodicity of the settlement, the newly settled juveniles were mainly found between the last quarter moon and new moon during the main settlement season. Back calculations of settlement dates based on otolith microstructure showed that most juveniles settled during the last quarter moon and new moon. Stepwise multiple regression analyses revealed that the newly settled juveniles were mainly found on coral patches with a large number of holes on the top of the coral patch and this tendency was mainly consistent during the study period. The volume of holes at the base of coral patches had also a positive effect on the abundance of newly settled juveniles in some cases. The results of the present study suggest that larval settlement has a strong seasonal and lunar periodicity, and newly settled juveniles select coral patches with available refuge spaces.     

Spatial repartition and ontogenetic shifts in habitat use by coral reef fishes (Moorea,French Polynesia)

This study explores the extent to which ontogenetic habitat shifts modify spatial patterns of fish established at settlement in the Moorea Island lagoon (French Polynesia). The lagoon of Moorea Island was divided into 12 habitat zones (i.e. coral seascapes), which were distinct in terms of depth, wave exposure, and substratum composition. Eighty-two species of recently settled juveniles were recorded from March to June 2001. Visual censuses documented changes in the distribution of juveniles of each species over time among the 12 habitats. Two patterns of juvenile habitat use were found among species. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (i.e. recruitment; e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus). Secondly, others settled to several habitats and then disappeared from some habitats through differential mortality and/or post-settlement movement (e.g. 65–70 mm size class for Ctenochaetus striatus, 40–45 mm size class for Epinephelus merra, 50–55 mm size class for Scarus sordidus). A comparison of the spatial distribution of juveniles to that of adults (61 species recorded at both stages) illustrated four patterns of subsequent recruitment in habitat use: (1) an increase in the number of habitats used during the adult stage (e.g. H. hortulanus, Mulloidichthys flavolineatus); (2) a decrease in the number of habitats adults used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis); (3) the use of different habitat types (e.g. Acanthurus triostegus, Caranx melampygus); and (4) no change in habitat use (e.g. Naso litturatus, Stegastes nigricans). Of the 20 most abundant species recorded in Moorea lagoon, 12 species modified the spatial patterns established at settlement by an ontogenetic habitat shift.Communicated by T. Ikeda, Hakodate     

Microhabitat-associated variability in survival and growth of subtidal solitary ascidians during the first 21 days after settlement

Newly settled juvenile ascidians were transplanted into shaded and unshaded sites at two subtidal depths to test the hypothesis that photonegative settlement behavior enhances juvenile survival. Silt, filamentous algae and grazing gastropods (Margarites pupillus) were identified as sources of mortality associated primarily with non-cryptic microhabitats. Silt inflicted heavy mortality on all six species tested. Algal overgrowth was important in shallow water, where it increased mortality and depressed growth rates over a 14-d period. Laboratory experiments demonstrated that gastropods can remove juvenile ascidians from the substratum. The sources of post-settlement mortality investigated may help determine distributional patterns of ascidians in the San Juan Islands, Washington, USA, and may also represent selective pressures that maintain negative phototaxis in the behavioral repertoires of ascidian tadpoles.     

Genetic identity determines risk of post-settlement mortality of a marine fish

Longitudinal sampling of four cohorts of Neopomacentrus filamentosus, a common tropical damselfish from Dampier Archipelago, Western Australia, revealed the evolution of size structure after settlement. Light traps collected premetamorphic individuals from the water column ("settlers") to establish a baseline for each cohort. Subsequently, divers collected benthic juveniles ("recruits") at 1-3-month intervals to determine the relative impacts of post-settlement mortality during the first three months. Growth trajectories for individual fish were back-calculated from otolith records and compared with nonlinear mixed-effects models. Size-selective mortality was detected in all cohorts with the loss of smaller, slower growing individuals. Three months after settlement, recruits showed significantly faster growth as juveniles, faster growth as larvae, and larger sizes as hatchlings. The timing and intensity of post-settlement selection differed among cohorts and was correlated with density at settlement. The cohort with the greatest initial abundance experienced the strongest selective mortality, with most of this mortality occurring between one and two months after settlement when juveniles began foraging at higher positions in the water column. Significant genetic structure was found between settlers and three-month-old recruits in this cohort as a result of natural selection that changed the frequency of mtDNA haplotypes measured at the control region. The extent of this genetic difference was enlarged or reduced by artificially manipulating the intensity of size-based selection, thus establishing a link between phenotype and haplotype. Sequence variation in the control region of the mitochondrial genome has been linked to mitochondrial efficiency and weight gain in other studies, which provides a plausible explanation for the patterns observed here.     

Daily otolith increments and recruitment in two coral-reef wrasses,Thalassoma bifasciatum and Halichoeres bivittatus

Increments on the otoliths of two common coral reef fishes, the bluehead wrasse Thalassoma bifasciatum and the slippery dick Halichoeres bivittatus, were demonstrated by mark-recapture experiments to be daily. Otoliths were marked in two ways; by depriving fish of light, food, and temperature cycles and also by supplemental feeding in the field. Both experiments were performed in late 1980 in the San Blas Islands of Panamá. A mark corresponding to settlement of the planktonic larva onto the reef was found on the otoliths of the bluehead wrasse. This settlement mark was used to calculate the dates of settlement of a collection of juveniles of this species taken from a patch reef in the San Blas Islands of Panamá in 1981. Settlement occurred in short and irregular bursts. The number of daily increments before the settlement mark indicates a planktonic larval life of 40 to 72 d.     

Temperature and salinity interactively impact early juvenile development: a bottleneck in barnacle ontogeny

When juvenile mortality or juvenile growth is impacted by temperature and salinity, these factors have a substantial effect on recruitment success and population dynamics in benthic ecosystems. Using freshly settled cyprids of Amphibalanus improvisus, we investigated the combined effects of temperature (12, 20 and 28 °C) and salinity (5, 15 and 30 psu) on early juvenile stage performance. Mortality as well as size (basal diameter, dry weight, and ash-free dry weight) was monitored for a period of 40 days. Mortality was high (42–63 %) during the first week following attachment, regardless of the temperature and salinity treatments. Subsequently, mortality and size were interactively influenced by temperature and salinity. Highest mortality and lowest size of juveniles occurred at lowest temperature (12 °C) and salinity (5 psu). Apparently, low temperature (12 °C) narrowed the barnacles’ salinity tolerance. Juvenile barnacles constructed more shell material compared to body mass at high temperature and high salinity, while a reverse situation was observed at low temperature and low salinity. Our results demonstrate that environmental changes can directly and/or indirectly alter patterns of survivorship and size. Warming and desalination as predicted for the Baltic Sea in the course of climate change may, however, act antagonistically and compensate each other’s isolated effect on barnacles.     

Estimates of adult and juvenile mortality for labrid fishes at One Tree Reef,Great Barrier Reef

Few estimates of natural mortality have been reported for coral reef fishes, yet this information is essential for predicting the effects of recruitment fluctuations on adult populations. In this study, mortality of adult (10 species) and juvenile (11 species) labrid fishes resident on ten isolated patch reefs in One Tree Lagoon, southern Great Barrier Reef, was estimated by visual censuses covering 30 mo, from November 1981 to April 1984. Numbers of adults of all species declined linearly over this period. Mortality rates differed significantly among species, ranging from 4.9 to 69.5% per annum. Average life-span for fishes that had survived the first year of life varied from 1.6 to 11.5 yr, depending on species. Average juvenile mortality varied among species from 25 to 91.8% per annum and was highest for abundant schooling species. Different cohorts of the same species also experienced significantly different rates of mortality. No clear relationship emerged between the number of new recruits and the number of fishes surviving to one year of age. The influence of recruitment fluctuations on adult populations of labrid fishes may be difficult to predict.     

Disappearance and dispersal of Spermophilus elegans juveniles in relation to behavior

Summary Patterns of disappearance and dispersal of Spermophilus elegans juveniles during the first 6-weeks postemergence were compared for 1977 and 1978 and related to quantitative and qualitative changes in social interaction involving juveniles.Juvenile disappearance (emigration or mortality) and dispersal within the study site varied between the sexes within and between years. Female disappearance and dispersal were significantly greater in 1977, and male losses in 1978 significantly exceeded male losses in 1977. Greater female loss in 1977 resulted in total male — female losses being equivalent, whereas in 1978 juvenile loss was strongly biased toward males by the end of the 6-week period.Greater female loss in 1977 was attributed primarily to increased aggression between female juveniles in that year because of larger average litter size with more females per litter. Increased disappearance of males in 1978 showed no correlation with litter size or relative increase in number of males per litter. Male young interacted with individuals of several age/sex classes, and a possible behavioral influence on male disappearance was increased aggressiveness by yearling males toward juvenile males in 1978.Behavior appeared to act as a proximate factor in juvenile disappearance and dispersal, and the observed differences between how male and female juveniles interacted in 1977 and 1978 were hypothesized to reflect the operation of different selective pressures to increase individual male or female fitness.     

Settlement patterns of newly settled plaice (Pleuronectes platessa) in a non-tidal Swedish fjord in relation to larval supply and benthic predators

In demersal fish species with a pelagic larval stage, settlement patterns may be a consequence of variations in larval supply, habitat selection at settlement, and processes acting between the time of settlement and the time of benthic sampling. This study describes temporal (1994-1998) and spatial variation in plaice (Pleuronectes platessa L.) settlement densities in four semi-isolated nursery areas with similar habitat characteristics, in the non-tidal Gullmarsfjord on the west coast of Sweden. Juvenile abundance varied by a factor of ten, both among years and among nursery grounds. For the 3 years when larval sampling was undertaken (1994-1996) and all nursery areas, there was a significant positive relationship between larval supply and juvenile abundance (linear regression: r²=0.45, n=24, P<0.001). On the southern side of the fjord, a significant positive relationship between larval and juvenile abundance was found in one area (r²=0.62, n=6, P<0.05). The absolute mortality rate of plaice after settlement was related to the initial settlement density (r²=0.95, n=20, P<0.001), and to the abundance of predatory shrimps Crangon crangon (r²=0.44, n=20, P<0.01). Plaice otoliths were found in 6% of the shrimp stomachs analysed from an area with high density (13.3 m^-2) of newly settled plaice. The present study suggests that the density of juvenile plaice was limited by larval supply to the nursery grounds. Consistency in the relative abundance of juveniles among nursery grounds between years also suggested that some nursery areas may be in the settlement shadow of others. The irregular nature of the coastline in combination with larval depletion could thereby cause small-scale (10³-10⁴ m) variation in settlement densities of the same order of magnitude as the inter-annual variability in recruitment to individual nursery grounds.     

Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Population dynamics of the acorn barnacles, Tetraclita squamosa and Tetraclita japonica (Cirripedia: Balanomorpha), in Hong Kong

Tetraclita squamosa and Tetraclita japonica are common, intertidal barnacles. In Hong Kong, the population dynamics of the two species exhibited spatial and temporal variation on two semi-exposed shores. T. squamosa produced egg masses from May–June and annual settlement and recruitment occurred from June–July. In contrast, settlement and recruitment of T. japonica was sparse from March–May, but intensified in July–October. Mature gonads and egg masses were, however, only present from September–November, suggesting the larvae from the two settlement pulses originated from other locations as well as Hong Kong populations. Settlement intensity and post-recruitment mortality of the two species varied between sites, possibly due to spatial variation in free rock space, physical transport of larvae and abundance of food in the water column.Communicated by T. Ikeda, Hakodate     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.