Marine larval ecology

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Marine larval ecology is the study of the factors influencing the dispersing larval stage which is exhibited by many marine invertebrates and fishes. Marine organisms with a larval stage typically release large numbers of larvae into the water column, where they develop before metamorphosing into adults.

Marine larvae are capable of dispersing long distances, although determining their actual dispersal distance is a significant challenge due to their size and the lack of an appropriate tracking method. Understanding dispersal distance is important for fisheries management, effective design of marine reserves, and control of invasive species.

Theories on the evolution of a biphasic life history[edit]

Larval dispersal is one of the most important topics in marine ecology today. Many marine invertebrates and many fishes have a life cycle involving a demersal adult and pelagic larval stage, or pelagic eggs that can be transported long distances.[1] There are several theories behind why these organisms have evolved this biphasic life history:[2]

  • Larvae use a different food source than adults, which decreases competition between life stages.
  • Pelagic larvae can disperse large distances, colonize new territory, and move away from habitat that has become overcrowded or otherwise unsuitable.
  • A long pelagic larval duration can help a species break its parasite cycles.
  • Pelagic larvae avoid benthic predators.

Pelagic larval dispersal can be risky. For example, while larvae do avoid benthic predators, they are still exposed to predators in the water column.

Larval development strategies[edit]

Marine larval development can be classified into three strategies: direct, lecithotrophic, and planktotrophic development. Each strategy has risks in the form of predation and difficulty in finding a safe site to settle.

Direct developers are characterized by larvae that have a similar form to the adult stage of the animal. Direct developers typically very low dispersal potential. These larvae are also known as “crawl-away larvae,” because larvae crawl away from their egg mass after hatching. Some species of frogs and snails hatch this way.

Lecithotrophic larvae have greater dispersal potential than direct developers. Many fish species and some benthic invertebrates have lecithotrophic larvae, which are provided with a source of nutrition to use during their dispersal in the form of a yolk sac. Though some lecithotrophic species are capable of feeding in the water column, many, such as tunicates, are not, and must settle before depleting their food source. Consequently, these species have short pelagic larval durations and do not disperse long distances.

Planktotrophic species have long pelagic larval durations and feed while in the water column leading to large dispersal distances. This ability to disperse is a key adaptation of benthic marine invertebrates.[3] Planktotrophic larvae feed on phytoplankton and small zooplankton, including other larvae. Planktotrophic development is the most common type of larval development, especially among benthic invertebrates.

Because plantotrophic larvae spend a long time in the water column and have low probability of successful recruitment, early researchers developed the “lottery hypothesis”, which states that animals release huge numbers of larvae to increase the chances that at least one will survive, and that larvae cannot influence their probability of success.[4][5][6] This hypothesis views larval survival and successful recruitment as chance events, which numerous studies on larval behavior and ecology have since shown to be false.[7] Though it has been generally disproved, the larval lottery hypothesis represents an important understanding of the difficulties faced by larvae during their time in the water column.

Predator Defense[edit]

Predation is a major threat to marine larvae, which are an important food source for many organisms. Invertebrate larvae in estuaries are particularly at risk because estuaries are nursery grounds for planktivorous fishes. Larvae have evolved strategies to cope with this threat, including direct defense and avoidance.

Direct Defense[edit]

Direct defense can include protective structures and chemical defenses.[8] Most planktivorous fishes are gape-limited predators, meaning their prey is determined by the width of their open mouths, making larger larvae difficult to ingest.[9] One study proved that spines serve a protective function by removing spines from estuarine crab larvae and monitoring differences in predation rates between de-spined and intact larvae.[10] The study also showed that predator defense is also behavioral, as they can keep spines relaxed but erect them in the presence of predators.[10]

Avoidance[edit]

Larvae can avoid predators on small and large spatial scales. Some larvae do this by sinking when approached by a predator. A more common avoidance strategy is to become active at night and remain hidden during the day to avoid visual predators. Most larvae and plankton undertake diel vertical migrations between deeper waters with less light and fewer predators during the day and shallow waters in the photic zone at night, where microalgae is abundant.[11] Estuarine invertebrate larvae avoid predators by developing in the open ocean, where there are fewer predators. This is done using reverse tidal vertical migrations. Larvae use tidal cycles and estuarine flow regimes to aid their departure to the ocean, a process that is well-studied in many estuarine crab species.[12][13][14][15]

An example of reverse tidal migration performed by crab species would begin with larvae being released on a nocturnal spring high tide to limit predation by planktivorous fishes. As the tide begins to ebbs, larvae swim to the surface to be carried away from the spawning site. When the tide begins to flood, larvae swim to the bottom, where water moves more slowly due to the boundary layer. When the tide again changes back to ebb, the larvae swim to the surface waters and resume their journey to the ocean. Depending on the length of the estuary and the speed of the currents, this process can take anywhere from one tidal cycle to several days.[16]

Dispersal and settlement[edit]

The most widely accepted theory explaining the evolution of a pelagic larval stage is the need for long-distance dispersal ability.[17][18] Sessile and sedentary organisms such as barnacles, tunicates, and mussels require a mechanism to move their young into new territory, since they cannot move long distances as adults. Many species have relatively long pelagic larval durations on the order of weeks or months.[19][20] During this time, larvae feed and grow, and many species metamorphose through several stages of development. For example, barnacles molt through six naupliar stages before becoming a cyprid and seeking appropriate settlement substrate.[21]

This strategy can be risky. Some larvae have been shown to be able to delay their final metamorphosis for a few days or weeks, and most species cannot delay it at all.[22][23] If these larvae metamorphose far from a suitable settlement site, they perish. Many invertebrate larvae have evolved complex behaviors and endogenous rhythms to ensure successful and timely settlement.

Many estuarine species exhibit swimming rhythms of reverse tidal vertical migration to aid in their transport away from their hatching site. Individuals can also exhibit tidal vertical migrations to reenter the estuary when they are competent to settle.[24]

As larvae reach their final pelagic stage, they become much more tactile; clinging to anything larger than themselves. One study observed crab postlarvae and found that they would swim vigorously until they encountered a floating object, which they would cling to for the remainder of the experiment.[25] It was hypothesized that by clinging to floating debris, crabs can be transported towards shore due to the oceanographic forces of internal waves,[clarification needed] which carry floating debris shoreward regardless of the prevailing currents.

Once returning to shore, settlers encounter difficulties concerning their actual settlement and recruitment into the population. Space is a limiting factor for sessile invertebrates on rocky shores. Settlers must be wary of adult filter feeders, which cover substrate at settlement sites and eat particles the size of larvae. Settlers must also avoid becoming stranded out of water by waves, and must select a settlement site at the proper tidal height to prevent desiccation and avoid competition and predation. To overcome many of these difficulties, some species rely on chemical cues to assist them in selecting an appropriate settlement site. These cues are usually emitted by adult conspecifics, but some species cue on specific bacterial mats or other qualities of the substrate.[26][27][28]

Larval Sensory Systems[edit]

Although a pelagic larval stage allows many species to increase their dispersal range and decreases the risk of inbreeding,[29] it comes with challenges. Without advanced sensory abilities, larvae are at risk of being washed away without suitable habitat for settlement. Marine larvae have evolved the ability to use a variety of sensory systems at different scales.

Sensory Systems[edit]

Magnetic Fields[edit]

Far from shore, larvae are able to use magnetic fields to orient themselves towards the coast over large spatial scales.[30][31] There is additional evidence that species can recognize anomalies in the magnetic field to return to the same location multiple times throughout their life.[30] Though the mechanisms that these species use is poorly understood, it appears that magnetic fields play an important role in larval orientation offshore, where other cues such as sound and chemicals may be difficult to detect.

Vision[edit]

Phototaxis (ability to differentiate between light and dark areas) is important for the detection of suitable habitat. The ability to perform phototaxis evolved relatively quickly[32] and taxa that lack developed eyes, such as schyphozoans, use phototaxis to find shaded areas to settle away from predators.[33]

Species that produce more complex larvae, such as fish, are able to use full vision[30] to find suitable habitat on small spatial scales. Studies of damselfish have shown that larvae use vision to locate and settle near adults of the same species.[34]

Sound[edit]

Marine larvae use sound and vibrations as a method of locating suitable habitat where they can metamorphose and settle into their juvenile form. This behavior has been seen in fish[34] as well as in the larvae of scleractinian corals.[35] Many families of coral reef fish are particularly attracted to high-frequency sounds produced by invertebrates,[36] which larvae use as an indicator of food availability and complex habitat where they may be protected from predators. It is thought that larvae avoid low frequency sounds because they may be associated with transient fish or predators[36] and is therefore not a reliable indicator of safe habitat.

The spatial range at which larvae detect and use sound waves is still uncertain, though some evidence suggests that it may only be reliable at very small scales.[37] There is concern that changes in community structure in nursery habitats, such as seagrass beds, kelp forests, and mangroves, could lead to a collapse in larval recruitment[38] due to a decrease in sound-producing invertebrates. Other researchers argue that larvae may still successfully find a place to settle even if one cue is unreliable.[39]

Olfaction[edit]

Many marine organisms use olfaction (chemical cues in the form of scent) to locate a safe area to metamorphose at the end of their larval stage.[34] This has been shown in both vertebrates[40] and invertebrates.[41] Research has shown that larvae are able to distinguish between water from the open ocean and water from more suitable nursery habitats such as lagoons[40] and seagrass beds.[42] Chemical cues can be extremely useful for larvae, but may not have a constant presence, as water input can depend on currents and tidal flow.[43]

Human Impacts on Sensory Systems[edit]

Recent research in the field of larval sensory biology has begun focusing more on how human impacts and environmental disturbance affect settlement rates and larval interpretation of different habitat cues. Ocean acidification due to anthropogenic climate change and sedimentation have become areas of particular interest.

Ocean Acidification[edit]

Ocean acidification has been shown to alter the way that pelagic larvae are able to process information[44] and production of the cues themselves.[45] Acidification can alter larval interpretations of sounds, particularly in fish,[46] leading to settlement in suboptimal habitat. Though the mechanism for this process is still not fully understood, some studies indicate that this breakdown may be due to a decrease in size or density of their otoliths.[47] Furthermore, sounds produced by invertebrates that larvae rely on as an indicator of habitat quality can also change due to acidification. For example, snapping shrimp produce different sounds that larvae may not recognize under acidified conditions due to differences in shell calcification.[45]

Hearing is not the only sense that may be altered under future ocean chemistry conditions. Evidence also suggests that larval ability to process olfactory cues was also affected when tested under future pH conditions.[48] Red color cues that coral larvae use to find crustose coralline algae, with which they have a commensal relationship, may also be in danger due to algal bleaching.[49]

Sedimentation[edit]

Sediment runoff, from natural storm events or human development, can also impact larval sensory systems and survival. One study focusing on red soil found that increased turbidity due to runoff negatively influenced the ability of fish larvae to interpret visual cues.[50] More unexpectedly, they also found that red soil can also impair olfactory capabilities.[50]

Self-recruitment[edit]

Marine ecologists are often interested in the degree of self-recruitment in populations. Historically, larvae were considered passive particles that were carried by ocean currents to faraway locations. This led to the belief that all marine populations were demographically open, connected by long distance larval transport. Recent work has shown that many populations are self-recruiting, and that larvae and juveniles are capable of purposefully returning to their natal sites.

Researchers take a variety of approaches to estimating population connectivity and self-recruitment, and several studies have demonstrated their feasibility. Jones et al.[51] and Swearer et al.,[52] for example, investigated the proportion of fish larvae returning to their natal reef. Both studies found higher than expected self-recruitment in these populations using mark, release, and recapture sampling. These studies were the first to provide conclusive evidence of self-recruitment in a species with the potential to disperse far from its natal site, and laid the groundwork for numerous future studies.[53]

Conservation[edit]

Ichthyoplankton have a high mortality rate as they transition their food source from yolk sac to zooplankton.[54] It is proposed that this mortality rate is related to inadequate zooplankton as well as an inability to move through the water effectively at this stage of development, leading to starvation. Many ichthyoplankton use suction to feed. Turgidity of water impairs the organisms’ ability to feed even when there is a high density of prey. Reducing these hydrodynamic constraints on cultivated populations could lead to higher yields for repopulation efforts and has been proposed as a means of conserving fish populations by acting at the larval level.[55]

A network of marine reserves has been initiated for the conservation of the world’s marine larval populations. These areas restrict fishing and therefore increase the number of otherwise fished species. This leads to a healthier ecosystem and affects the number of overall species within the reserve as compared to nearby fished areas; however, the full effect of an increase in larger predator fish on larval populations is not currently known. Also, the potential for utilizing the motility of fish larvae to repopulate the water surrounding the reserve is not fully understood. Marine reserves are a part of a growing conservation effort to combat overfishing; however, reserves still only comprise about 1% of the world’s oceans. These reserves are also not protected from other human-derived threats, such as chemical pollutants, so they cannot be the only method of conservation without certain levels of protection for the water around them as well.[56]

For effective conservation, it is important to understand the larval dispersal patterns of the species in danger, as well as the dispersal of invasive species and predators which could impact their populations. Understanding these patterns is an important factor when creating protocol for governing fishing and creating reserves. A single species may have multiple dispersal patterns. The spacing and size of marine reserves must reflect this variability to maximize their beneficial effect. Species with shorter dispersal patterns are more likely to be affected by local changes and require higher priority for conservation because of the separation of subpopulations.[57]

Implications[edit]

The principles of marine larval ecology can be applied other fields inside and outside the marine realm. Successful fisheries management relies heavily on understanding population connectivity and dispersal distances, which are driven by larvae. Dispersal and connectivity must also be considered when designing natural reserves. If populations are not self-recruiting, reserves may lose their species assemblages. Many invasive species can disperse long distances, including the seeds of land plants and larvae of marine invasives. Understanding the factors influencing their dispersal is key to controlling their spread and managing established populations.

See also[edit]

References[edit]

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