The biology and ecology of Spiny Water Flea, Bythotrephes longimanus

Submitted by: Alido, Rommel Simon Rado, Mavreen Siegred Rellosa, Rochel Salatandol, Rissa Sobredo, Audie

Submitted to: Mrs. Venus B. Kinamot

The biology and ecology of Spiny Water Flea, Bythotrephes longimanus

I. Introduction A. Description of Bythotrephes longimanus B. Taxonomic Classification of Bythotrephes longimanus

II. Biology of Bythotrephes longimanus A. Mode of Feeding and Digestion B. Mode of Movement C. Mode of Reproduction D. Growth and Development

III. Ecology of Bythotrephes longimanus A. Distribution and Habitats of Bythotrephes longimanus B. Ecological Impacts of Bythotrephes longimanus

IV. Summary

I. Introduction A. Description of Bythotrephes longimanus
The spiny water flea (Bythotrephes longimanus) is a Eurasian cladoceran invasive in the Laurentian Great Lakes basin of North America. The Great Lakes act as a source for Bythotrephes, which is now rapidly expanding (Lehman, 1987). Bythotrephes belongs to the class Crustacea, a group of animals such as crabs and shrimp that possess a hard exoskeleton (outer shell). Like all other Crustacea, its exoskeleton is molted in order to grow. Bythotrephes is unique for it sheds only the exoskeleton that covers its body, retaining the exoskeleton that covers the tail spine. The tail serves a vital protective function. The body of Bythotrephes is enclosed within a bivalve shell called a carapace that covers the thorax and the abdomen but not the head. The body of Bythotrephes is composed of a head which is dominated by a large black eye, jaws, four pairs of legs and a pair of branched antennae. Adult females are larger than adult males. Juvenile spiny water fleas have one pair of barbs on the tail spine and as they grow larger can have up to four pairs of barbs. The first pair of legs is longer than the rest and is used for catching prey. The other pairs of legs are used to hold the prey while it is being consumed. The body of Bythotrephes is composed of a head which consists of a pair of branched antennae. These pair of swimming antennae, help propel the animals through the water. The antennae are used for swimming. Reproducing females will carry a brood pouch on their back containing developing larvae (Benson, 2005). As a large conspicuous zooplankter, Bythotrephes is subject to heavy predation by young fish. This triggered the evolution of two important adaptations: firstly, a chitinous, barbed spine, that comprises more than 2/3 of the body length and that acts as a defense mechanism and

secondly, there are durable, tough shelled diapause eggs that can pass through fish guts intact. Gut passage means that fish can act as a dispersal vector. Resting eggs in baitfish can be transferred from one lake to another, unintentionally dispersing the spiny water flea. Bythotrephes is a voracious predator, and individuals consume up to 40 smaller invertebrates per day (Liebig, 2005). Tail spine
abdomen

A pair of antenna

Barbs on tail

4 pairs of legs

Dark eye spot

Figure 1. A picture shows the body parts of Bythotrephes longimanus, an invasive crustacean in the Great Lakes (a photograph by Kevin Keeler, U.S. Geological Survey).

B. Taxonomic Classification of Bythotrephes longimanus The spiny water flea (Bythotrephes longimanus) is a predacious zooplankton species that belongs to phylum Arthropoda and a member of the group of organisms known as crustaceans. Shrimp, crayfish, and lobster also belong to this group and the spiny water flea is a relative of another Great Lakes invader - the fishhook water flea. These water fleas are not insects at all, but small cladocerans related to other crustaceans. The distinguishing characteristic that separates the spiny and fishhook water fleas from all other native zooplankton are their tail spines. Both

species reach about ½ inch long but about 70% of the total length is the tail spine. The predatory cladoceran Bythotrephes longimanus (the spiny water flea) has been identified as one of several recent invasive species of great threat to the Great Lakes (Tuchman, 2004). Kingdom Phylum Subphylum Class Subclass Order Suborder Infraorder Family Genus Species Animalia Arthropoda Crustacea Branchiopoda Phyllopoda Diplostraca Cladocera Onychopoda Cercopagididae Bythotrephes longimanus

Figure 2. A diagram shows the taxonomic classification of Bythotrephes longimanus.

II. Biology of Bythotrephes longimanus A. Mode of Feeding and Digestion According to Branstrator (2005), Bythotrephes is a carnivore, which means that it’s feeding on other zooplankton. Bythotrephes eat smaller herbivorous Crustacean, including the common zooplankton, Daphnia. Daphnia, however, are also an important food item for small, juvenile fish such as the bloater chub. The body of Bythotrephes is enclosed within a bivalve shell called a carapace that covers the thorax and the abdomen but not the head, and inside the

carapace are 5 or 6 pairs of feet used to filter the water for food (Tuchman, 2001). Also, Bythotrephes uses their sickle-shaped mandibles to pierce and shred its prey. The animals possess four pairs of legs, the first pair being much longer than the others. These first legs are used for catching prey, whereas the other pairs of limbs are designed for grasping prey while they are being consumed. Bythotrephes ingests no identifiable parts of its prey (it ingests only soft tissues), a simple gut content analysis (looking at what items are in its digestive tract) to determine diet composition was impossible. The researchers analyzed allozyme signals from the gut contents of field-collected Bythotrephes and compared them to the signals from potential prey items in Lake Michigan's offshore zooplankton assemblage to determine which prey were being consumed. Additional studies on the feeding ecology of Bythotrephes also led to the prediction that this invader may be severely decreasing trophic transfer efficiency of energy from herbivorous zooplankton to fish in Lake Michigan. Bythotrephes thus competes directly with young fish for food. Because of their unique reproductive pattern, Bythotrephes can reproduce many times faster than the fish. Their rapid population growth enables Bythotrephes to monopolize the food supply at times, to the eventual detriment of the fish. That’s its big effect in food webs. It reduces the abundance in the diversity of native plankton in lakes, which is used by a variety of fishes that are feeding on the plankton (Caceres, 1999).

Figure 3. An illustration showing small bait fish at the top of aquatic food chain, followed by carnivorous zooplankton, herbivorous zooplankton and phytoplankton. The spiny water flea falls under the categorization of carnivorous zooplankton.

B. Mode of Movement

B. longimanus has a pair of swimming antennae, which propel the animals through the water. Bythotrephes are good swimmers, moving several times their body length in a second. The ability to swim, as opposed to merely drifting with the current, helps Bythotrephes to encounter prey and to move between shallow and deeper lake waters. Bythotrephes has been found to adopt a behavior called diel (daily) vertical migration. Adult females move deeper in the water during daylight hours, where less light penetrates and visibility to fish is reduced. At night, they rise closer to the surface, where there is abundant food and the warmer water helps to quicken metabolism and growth (Sikes, 2002).

A pair of antenna

Figure 4. A picture shows a pair of antenna of B. longimanus that is used for swimming.

C. Mode of Reproduction Bythotrephes longimanus can reproduce with both sexes and parthenogenically (all females). Most of the time females reproduce by parthenogenesis and produce 1-10 eggs that develop into new females without fertilization by males. Maturity is quickly reached. At higher summer temperatures, individuals can reach sexual maturity in about 14 days. A new generation may be produced every two weeks in the summer, sexual reproduction increases their chances of surviving in variable and extreme weather condition (Tuchman, 2001). The sex of the young is determined not by genetics, but by the environment. When conditions deteriorate in the fall, males start being produced. They mate with the females and produce “resting eggs”; that are carried in the brood pouch and then are released to settle on the bottom of the lake, where they go into a “resting phase” through the winter and hatch into females. Eggs can remain dormant for long periods of time and hatch when environmental conditions become more favorable. Overwintering dormant eggs usually hatch when water temperatures reach 4°C. Dormant eggs

can usually survive passage through the digestive tract of fish predators, further enhancing their ability to disperse .The brief sexually reproducing part is to promote genetic diversity (Krieger, 1986).

Eggs

Figure 5. A picture shows female Bythotrephes longimanus with eggs (courtesy by Dave Brenner, Michigan Sea Grant College Program).

D. Growth and Development Life Cycle Stages. Sikes (2002) states that, "Through parthenogenesis the spiny water flea can exhibit explosive population growth, but its ability to produce sexual eggs allows it to increase genetic variability as well as survive and disperse under adverse environmental conditions. Development time till primaparity (1st time mom) is not significantly different for the two modes of reproduction, averaging about 14 days. Sexually reproduced eggs can go into a semi-static metabolic condition called diapause. Through these sexual reproduced "resting eggs", the next generation of B. longimanus can overwinter and hatch usually when temperatures

exceed 4ºC. The spiny water flea can survive a wide range of temperatures, but has lowest mortality between 5ºC and 30ºC. Its development time is temperature dependent and maximized between 20-25ºC without suffering higher mortality. Besides protection from winter conditions, many diapause eggs can also survive passage through fish digestive tract. A female with a full clutch is double her usual weight. This fact causes increased predation on pregnant females above their conspicuous body with a single large eye and long tail spine and thereby further aids in dispersal."

Parthenogenic daughter

Sexual eggs

Parthenogenic son

Figure 6. Alternating asexual and sexual life cycles of Bythotrephes. The top loop illustrates the progression of embryos as they develop asexually. Females hatched from resting eggs can develop as many as four pairs of barbs. The presence of the male in the bottom loop indicates the sexual production of resting eggs (Graphics courtesy of the Ontario Federation of Anglers and Hunters).

III. Ecology of Bythotrephes longimanus A. Habitat and Distribution of Bythotrephes longimanus Habitats of B. longimanus. The spiny water flea is a species of zooplankton that naturally occurs in estuaries, lakes, marine habitats and wetlands. During the daytime, they can sink up to 50 to 60 m. During the night, they accumulate in the surface layer. In its native range it shows a preference for large, deep, clear lakes with relatively low summer bottom temperatures (Liebeg et al., 2005). Scientific study suggests that Bythotrephes has become a permanent member of the Great Lakes ecosystem, and has carved out a niche for itself at some cost to lake fisheries. The quick success of the animal in colonizing all of the Great Lakes raises the possibility that it may soon invade smaller, inland lakes in the Great Lakes basin (Moen, 1997).

Figure 7. A picture shows natural habitat of B. longimanus.

Distribution of B. longimanus. It is believed that the spiny water flea first arrived in North America in the ballast water of Eurasian ships. Research has shown that human recreation involving boats is the principle mechanism of transfer of non-native species between water bodies. Boating is an activity that is extremely vulnerable to “hitch-hiking” by non-native organisms because so many surfaces, nooks, and crannies of nautical gear are immersed in water. This is especially true of the spiny water flea because it produces thick-walled “resting” eggs that can remain dormant for long periods of time and are resistant to environmental extremes. These eggs even survive passage through the digestive tracts of fish. Adult spiny water fleas snagged during boating or fishing may contain resting eggs. These eggs can survive for extended periods after being tangled with downriggers, anchor line, trailer parts, and fishing gear. Consequently, resting eggs are insidious “stowaways” and contribute greatly to the rapid dispersal of this nonnative organism from infested lakes to clean ones (Tuchman et al., 2004). The first recorded occurrence of the spiny water flea in North America was in Lake Ontario in 1982, and by 1987 it was present in all of the Great Lakes. This crustacean’s known range in Ontario now includes over 120 inland lakes and waterways in Michigan, Minnesota, New York, Wisconsin and Ohio. The species has also invaded inland lakes in Ontario, Canada and is also invasive in Belgium and the Netherlands (Moen, 1997). If the proper precautions are not taken, spiny water fleas can be further spread unknowingly by boaters and anglers. It has only been invasive in Ontario for 15 years. Currently, seven new inland lakes were found to have spiny water fleas. It is only a matter of time before the bug makes its presence felt in other lakes.

Figure 8. A map shows the distribution of B. longimanus in the different lakes of America.

B. Ecological Impacts of Bythotrephes longimanus Bythotrephes longimanus is a zooplanktovore that has been established for millennia in large, temperate, nutrient- poor lakes (Grigorovich et al., 1998). However, it is functioning as key regulator of plankton composition (Manca et al., 2000). A healthy global plankton community supplies humankind with services we either cannot do without, e.g. essential fatty acids (Arts et al., 2001). It can also be a source of food for fish including yellow perch, white perch, walley white bass, alewife, bloater chub, Chinook salmon, emerald shiner, spot tail shiner, rainbow smelt, lake herring, lake whitefish and deep water sculpin (Bur et al., 1986). According to Hessen et al. (2011), Bythotrephes is inhabiting 20% of lakes in Norway, contributing to salmonid fish diets, out of its contribution of planktonic biomass (Nilsson, 1979).

On the contrary, in some place in North America, B. longimanus has proven to be a serious threat to pelagic biodiversity in both large and small lakes (Tuchman et al., 2004). It consumes a very large fraction of total zooplankton production (Dumitri et al., 2001), and includes downward migration of its prey into deeper, cooler waters that lower their growth rates (Pangle et al., 2007). This adversely affects the growth rate and survival rate of young fishes due to high rate competition of food supply and apparently affects the abundance of fishes in that certain area where the B. longimanus monopolized. Its damaging effects cascade below its immediate crustacean prey to pelagic rotifers (Hovious et al., 2006) and likely to phytoplankton (Strecker et al., 2011) and also up the food chain to competing macro- invertebrate predators and fish (Foster, 2009). This wide array of effects caused by B. longimanus to the aquatic ecosystem shows that this organism is a nuisance rather a beneficial one.

IV. Summary The spiny water flea (Bythotrephes longimanus) is a predacious zooplankton species that belongs to phylum Arthropoda and a member of the group of organisms known as crustaceans. It is a species of zooplankton that naturally occurs in estuaries, lakes, marine habitats and wetlands. It is believed that the spiny water flea first arrived in North America in the ballast water of Eurasian ships. Research has shown that human recreation involving boats is the principle mechanism of transfer of this non-native species between water bodies. Bythotrephes longimanus is a cladoceran crustacean (water flea) recognizable with its straight tail spine averaging about 70% of its length. Adult individuals have three to four barbs on the spine while juveniles have only one pair. The animal has one large eye that is usually black or red. It also possesses a pair of

swimming antennae and four pairs of legs, of which the first pair is used to catch prey. Mandibles are used for consumption of prey. The spiny water flea preys on smaller planktonic organisms. Its diet consists mostly of zooplankton including Daphnia and smaller crustaceans. Bythotrephes can consume 10–20 prey organisms a day. It may also eat other small organisms it comes across. The spiny water flea is similar to another introduced cladoceran of the same family, the fishhook water flea Cercopagis pengoi, which however has a more slender spine featuring a prominent loop-like hook at its end. There is also variation in the shape of the more robust spine of Bythotrephes itself. Specimens arising from sexually produced eggs have a completely straight and relatively shorter spine. In parthenogenetically produced animals the spine features a kink in the middle. The eggs survive even after being dried out or eaten by fish. The spiny water flea is causing serious concerns in the lakes. The problem is that it feeds on zooplankton and can actually reduce zooplankton species. As zooplankton is the backbone of aquatic food chains, this tiny crustacean presents a serious risk to the ecosystem.

References:

Lehman, J.T. 1987. Palearctic predator invades North American Great Lakes. Oecologia 74:478480. Liebig, J. and A. Benson. 2005. Bythotrephes longimanus. U.S. Geological Survey. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=162

Caceres, Carla E. and John T. Lehman. 1999. Life History and Effects on the Great Lakes of the Spiny Tailed Bythotrephes. Michigan Sea Grant College Program. http://www.seagrant.umn.edu/exotics/spiny.html

Sikes, B.A. 2002. Invader of the Month June 2002: Spiny Water Flea. Institute for Biological Invasions.http://invasions.bio.utk.edu/invaders/flea.html#Native%20Range%20&%20Bio logy

Moen, Sharon. Our Lake has Fleas. 1997. Minnesota Sea Grant College Program. http://www.seagrant.umn.edu/seiche/jun.00/art02.htm

Berg, David J. 1992. The spiny water flea, Bythotrephes cederstroemi: Another unwelcome newcomer to the Great Lakes. Ohio Sea Grant College Program. Publication FS-049. 2pp.

Tuchman, M. L. 2001. Bythotrephes : Invaders of the Great Lakes. http://www.spinywaterflea.umn.edu/seiche/jun.00/art02.htm

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