Daphnia Lumholtzi
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''Daphnia lumholtzi'' is a species of small, invasive
water flea The Diplostraca or Cladocera, commonly known as water fleas, are a superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised so far, with many more ...
s that originates in the tropical and subtropical lakes of Africa, Asia, and Australia. As an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
, ''Daphnia lumholtzi'' disrupts aquatic habitats by spreading throughout the warmer waters of lakes and reservoirs.


Description

''Daphnia lumholtzi'' is a small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
that is 2–3 mm in length. It has a large helmet and a long tailspine, usually longer than the length of its body, that fluctuates in size. Its body structure is arched, extending to a sharp point. There are roughly 10 prominent spines on the margin of the abdominal shield covering.


Ecology


Temperature

''Daphnia lumholtzi'' is typically found in the warm, shallow regions of bodies of water with larger surface areas. While most species of ''Daphnia'' see high mortality at temperatures greater than 25 °C, ''D. lumholtzi'' individuals can survive and reproduce at temperatures up to 30 °C, with a thermal optimum occurring at 29 °C. Studies have shown that population density and water surface temperature are positively correlated. Once favorable temperatures are reached, such as those in the late summer, the previously deposited eggs hatch. The lower temperature range of ''D. lumholtzi'' extends to 5 °C, where some individuals are able to survive, with survival increasing significantly at 10 °C and reproduction beginning to occur at 15 °C. This significantly broad temperature range in which ''D. lumholtzi'' can survive has led to it being labeled a
eurytherm A eurytherm is an organism, often an endotherm, that can function at a wide range of ambient temperatures. To be considered a eurytherm, all stages of an organism's life cycle must be considered, including juvenile and larval stages. These wide ...
al species.


Behavior

Both adult and juvenile ''D. lumholtzi'' exhibit a vertical migration pattern, moving upward as the sun sets and downward as the sun rises. This behavior leads to large population densities close to the water surface at night and also occurs in the absence of a suggested predator threat.


Diet

''Daphnia lumhotzi'' mostly feeds on
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
ranging from 1 to 25 micrometers in size, but will also eat foods that contain organic
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
,
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, and
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s which provide an excellent source of nutrients.


Dispersal

''Daphnia lumholtzi'' was originally restricted to the tropical lake and pond regions of southwest Asia, Australia, and most of Africa. The exact location of geographic origin in the United States has not been identified, but scientists believe the introduction of exotic African fish to lakes most likely caused the distribution. It was first detected in Missouri and Texas
reservoir A reservoir (; from French ''réservoir'' ) is an enlarged lake behind a dam. Such a dam may be either artificial, built to store fresh water or it may be a natural formation. Reservoirs can be created in a number of ways, including contro ...
s in 1991 and has since been found in more than 16 states and over 125 lakes and reservoirs. Studies have shown that the ability of ''D. lumholtzi'' to disperse widely is most likely due to human activity. Heavy boat traffic on lakes and reservoirs during warmer seasons when ''D. lumholtzi'' thrive enable them to expand into other nearby bodies of water. The long spines and hairs on eggs act as hooks and enable attachment to boats, facilitating dispersal. The presence of ''D. lumholtzi'' in smaller ponds is atypical; however it is unlikely that non-human dispersal mechanisms, such as smaller
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s moving between bodies of water, have contributed to its widespread distribution.


Reproduction

''Daphnia lumholtzi'' deposits eggs in lake sediment that can remain dormant for long periods of time. The eggs are characterized by long spines and hairs that act as hooks.
Ephippia Ephippia (singular: ephippium) are winter or dry-season eggs of the various species of small crustacean in the order Cladocera (within the Branchiopoda); they are provided with an extra shell layer, which preserves and protects the resting ...
are protective shells that cover the egg until favorable conditions occur, such as warmer temperatures or a larger amount of resources. These ephippia are able to survive in environmental conditions, including oxygen, salinity, and temperature ranges, that are inhospitable to adult daphnids, although exact ranges for ''D. lumholtzi'' ephippia have not been found. This ephippia stage is an example of
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, a state of suspended animation an organism can enter in order to survive a harsh environment. ''D. lumholtzi'' is capable of producing 10 times more
ephippia Ephippia (singular: ephippium) are winter or dry-season eggs of the various species of small crustacean in the order Cladocera (within the Branchiopoda); they are provided with an extra shell layer, which preserves and protects the resting ...
than other daphnid species. In temperatures above the optimal temperature for reproduction, 25 °C, the rate of egg development decreases. In temperatures below 25 °C, egg development slows.


Predators

The main predators of ''D. lumholtzi'' are fish and small invertebrate species. Larger fish are almost always successful in their encounters with ''D. lumholtzi''. Small invertebrate predators are less efficient than large fish in catching ''D. lumholtzi''.


Physiology

Being an invasive species, the
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
of ''D. lumholtzi'' is relatively well studied for a daphnid.


Respiration

''Daphnia lumholtzi'' individuals prefer areas with high levels of dissolved oxygen and avoid areas where oxygen levels are low. Population surveys have found robust ''D. lumholtzi'' populations in water with oxygen saturation levels ranging from 65-163%, while no populations were found in water with saturation levels ranging from 7-50%. The mechanism ''D. lumholtzi'' uses for respiration is very similar to that used by other species of ''Daphnia'', with gas exchange occurring through gills that are fed oxygenated water by appendages on the thorax.


Response to salinity

As with most other members of the order
Cladocera The Diplostraca or Cladocera, commonly known as water fleas, are a superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised so far, with many more ...
, ''D. lumholtzi'' lives in freshwater and is
hyperosmotic In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane- ...
to its environment. ''D. lumholtzi'' is generally found in habitats with a salinity near 0 grams per liter, but can withstand slightly saline water, up to 1.5 grams per liter, for short amounts of time. This ability to survive short bouts of salinity has likely contributed to ''D. lumholtzi''’s ability to invade North America.


As an invasive species

''Daphnia lumholtzi'' exhibits higher survivorship and reproduction in the late summer, under high heat conditions, when compared to other
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s living in these conditions. It has been suggested that ''D. lumholtzi''’s more tropical origins may have enabled it to live in these higher temperatures due to evolutionary changes that resulted in
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s that are better adapted to the heat. This advantage allows them to be a better competitor and ultimately out-compete other species, specifically native
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
species, within the same habitat and come out as a successful invader. In accordance with the
competitive exclusion principle In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the sligh ...
, no other species can inhabit the same late summer
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
as ''D. lumholtzi''; another factor that allows it to have higher survivorship than other ''Daphnia'' species and is ultimately a better invader. It is important to note, however, that high survivorship and reproduction are not the only factors that make ''D. lumholtzi'' an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
. Competition between ''D. lumholtzi'' is increased in habitats that favor the high light intensity of shallow waters. ''D. lumholtzi'' showed greater survivorship than other ''Daphnia'' species (specifically '' D. pulex''), which made them a stronger competitor for light reception and resources in bodies of water receiving high light intensity. It was found to out-compete other species in areas with high light intensity which in turn contributes to its invasive success. ''Daphnia lumholtzi'' is capable of producing 10 times more
ephippia Ephippia (singular: ephippium) are winter or dry-season eggs of the various species of small crustacean in the order Cladocera (within the Branchiopoda); they are provided with an extra shell layer, which preserves and protects the resting ...
than other daphnid species, which can remain dormant until favorable conditions occur. This egg bank gives them an advantage over other species whose eggs cannot withstand
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
or lower temperatures, enabling them to produce more offspring that survive longer. The reproductive rate also increases with a higher concentration of food. Areas exhibiting high food abundance will therefore attract more ''D. lumholtzi'', and in turn result in a higher rate of reproduction. The greater number of offspring puts pressure on the habitat’s resources and other competitors. ''Daphnia lumholtzi'' is highly
plastic Plastics are a wide range of synthetic or semi-synthetic materials that use polymers as a main ingredient. Their plasticity makes it possible for plastics to be moulded, extruded or pressed into solid objects of various shapes. This adaptab ...
, meaning it has the ability to morphologically adapt to factors within the environment by developing structures that enable it to successfully avoid
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. A long tail spine, large helmet, and additional spines on the abdomen are produced in response to predator
kairomone A kairomone (a coinage using the Greek καιρός ''opportune moment'', paralleling pheromone"kairomone, n.". OED Online. September 2012. Oxford University Press. http://www.oed.com/view/Entry/241005?redirectedFrom=kairomone (accessed 3 Octobe ...
s, which are predator hormones, within the water. ''D. lumholtzi'' does not produce these protective structures when there are no predators present, and looks morphologically similar to other ''Daphnia'' species. When predators are detected, ''D. lumholtzi'' responds by producing a tail spine, helmet, abdomen spines for protection; other ''Daphnia'' species do not adapt this way to predator threats. With the development of these morphological features, predators have a more difficult time preying on ''D. lumholtzi''. This excess energy the predators put into eating ''D. lumholtzi'' lessens predator efficiency, making the predators more likely to choose another prey. This prey-switching puts an extra strain on other native zooplankton species, reducing predation on ''D. lumholtzi'' and allowing it to outperform other competitors.


Control

Eradication of ''D. lumholtzi'' is almost impossible once it has invaded a lake or reservoir. ''D. lumholtzi'' is sensitive to various
pesticides Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampric ...
and manmade chemicals, but the introduction of chemicals to natural lakes is often harmful to other species. The focus of most control measures is the prevention of initial invasion. For now, scientists recommend simple practices, such as thorough cleaning of boats and avoiding aquarium water dumps, to slow the spread of the species.


References

{{Taxonbar, from=Q4021132 Cladocera Crustaceans described in 1885 Taxa named by Georg Ossian Sars