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   Cercopagis pengoi (crustacean)
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      Sexually produced ossiani morph (on the right) and parthenogenetically
produced pengoi morph (on the left) produced morphs of Cercopagis pengoi
(Photo: Dr. Igor Grigorovich, University of Windsor, Canada) - Click for full size   Sexual female Cecopagis pengoi with a resting egg: anterior region (on the left) and complete body with caudal process (on the right) (Photo: Dr. Igor Grigorovich, University of Windsor, Canada) - Click for full size   Male Cercopagis pengoi: anterior region (on left) and complete body with caudal process (on right) (Photo: Dr. Igor Grigorovich, University of Windsor, Canada) - Click for full size
    Taxonomic name: Cercopagis pengoi (Ostroumov, 1891)
    Synonyms: Cercopagis (Apagis) ossiani (Mordukhai-Boltovskoi 1968)
    Common names: fishhook waterflea (English), Kaspischer Wasserfloh (German)
    Organism type: crustacean
    Cercopagis pengoi is a water flea native to the Ponto-Aralo-Caspian basin in South Eastern Europe, at the meeting point of the Middle East, Europe and Asia. It has spread from its native range and become invasive in some waterways of Eastern Europe and in the Baltic Sea. It has been introduced to the Great Lakes of North America, quickly becoming established and is now increasing its range and abundance. Cercopagis pengoi is a voracious predator and may compete with other planktivorous invertebrates and vertebrates. Through this competition, Cercopagis pengoi has the potential to affect the abundance and condition of zooplanktivorous fish and fish larvae. It also interferes with fisheries by clogging nets and fishing gear.
    Description
    The most pronounced parts of the body are the head, the second pair of antenna, four pairs of thoracic legs (thoracopodsI-IV), abdomen, caudal process, and a brood pouch in females. The head is essentially composed of a large single eye, where the amount of black pigment makes less than one half of the diameter of the eye. The second antenna is a large appendage containing of two branches - the endopod and exopod. The first pair of thoracic legs (thoracopods I) are 3-4 times longer than the second ones. Abdomen length is equal to the length of the rest of the body, and spines are large, equal to 2-3 diameters of caudal process (Mordukhai-Boltovskoi & Rivier, 1987; Rivier 1998). Parthenogenic females of the first generation of C. pengoi that hatch from resting eggs are anatomically distinct from parthenogenic females of following generations. They have a short straight caudal spine unlike the characteristically looped caudal spine of parthenogenically-produced individuals (Simm & Ojaveer 1999). C. pengoi possesses a high degree of regional variability in morphology. Largest adult parthenogentic females are found in the Baltic Sea (mean body length 1.99mm), while those in the Caspian Sea and in Lake Ontario are smaller (1.73 and 1.45mm, respectively). Caudal processes are largest in the instar III parthenogenic females from the Baltic Sea (mean length 9.65mm), medium in Lake Ontario (8.57mm), and shortest in females from the Caspian Sea (7.48mm). However, relative to body length, the length of caudal process is largest in populations from Lake Ontario (mean 5.9mm), medium in the Baltic (4.9mm), and smallest in the Caspian populations (4.3mm) (Grigorovich et al. 2000).
    Similar Species
    Bythotrephes longimanus

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    Occurs in:
    estuarine habitats, lakes, marine habitats, water courses, wetlands
    Habitat description
    Cercopagis pengoi is a euryhaline (having a wide tolerance to salinity) and eurythermic (can tolerate a wide range of temperatures) species, occurring in both brackish (up to 17%) and fresh waters, as well as highly variable temperatures (3 – 38 ºC) (c.f. Gorokhova et al. 2000). Usually highest population densities are found at summer temperatures (16 – 26 ºC) and at salinities of up to 10% (Mordukhai-Boltovskoi & Rivier, 1987; Rivier, 1998). Both in the Caspian Sea (Rivier, 1998) and Lake Ontario (Ojaveer et al. 2001) C. pengoi abundance increases with distance from shore, suggesting that this is a typical pelagic (fish and animals that live in the open sea, away from the sea bottom) species.
    General impacts
    Cercopagis pengoi is a potential competitor with young stages of planktivorous fish for herbivorous zooplankton (Vanderploeg et al. 2002). Several lines of evidence indicate that C. pengoi may affect resident zooplankton communities by selective predation: Lake Ontario (Benoit et al. 2002); Gulf of Riga (Ojaveer et al. 1999, 2004); Gulf of Finland (Uitto et al. 1999). Such changes may result in decreased grazing pressure on phytoplankton and enhanced algal blooms. It is, however, difficult to study food competition between small fish and C. pengoi because of the lack of feeding studies on the latter. On the other hand, zooplanktivorous fish both in the Baltic (Antsulevich and Välipakka 2000; Gorokhova et al. 2004; Ojaveer et al. 2004) and in the Great Lakes (Bushnoe et al. 2003) have been reported to prey on C. pengoi, implying that it has become a new food source, particularly for larger fish. Moreover, this large cladoceran tends to attach to fishing gear, clogs nets and trawls, causing problems and substantial economic losses for fishermen (Leppäkoski & Olenin, 2000; ICES, 2002).
    Notes
    The paucity of ecological studies on C. pengoi is in part due to the difficulty of culturing Cercopagis handling it and using in experiments.
    Geographical range
    Native range: Southern Europe - Ponto-Aralo-Caspian basin, i.e. Caspian, Black and Azov seas and small coastal lakes in this area (Mordukhai-Boltovskoi & Rivier, 1987; Rivier 1998). Known introduced range: waterways and reservoirs in the Eastern Europe, Baltic Sea, Laurentian Great Lakes, Finger Lakes (North America) (Leppäkoski & Olenin, 2000; Therriault et al. 2002; Vanderploeg et al. 2002).
    Introduction pathways to new locations
    Ship ballast water:


    Local dispersal methods
    Boat: On fishing lines.
    Management information
    Preventative measures: Prompted by the explosive increase of ship-borne exotic species in the Great Lakes in May 1993, the USA implemented a regulation that requires inbound vessels to exchange freshwater/estuarine ballast with highly saline oceanic water (Ricciardi and MacIssac 2002). In theory, this procedure should greatly reduce the risk of invasion as freshwater organisms would be purged or killed by seawater and would be replaced by marine organisms that would not survive if released into the freshwaters. However, the recent invasion of C. pengoi causes concern as it was apparently introduced after implementation of ballast water regulations. The authors believe that "without more effective ballast water control, continued invasion and transformation of West European and North American inland waters by Ponto-Caspian species is highly probable."

    The following measures could prevent further spread of this and other species with similar ecology: 1) Do not release bait or bait water into water body or transport from one water body to another. 2) Following good containment measures is important in controlling the spread of adult C. pengoi, but resting eggs are capable of surviving desiccation and freezing (even for periods of several years). 3) Rinsing boat and equipment with hot water (>40°C), high-pressure water spray, or drying boat and equipment for at least 5 days before re-entering water body will help to control the spread of adult C. pengoi. 4) Thoroughly draining and cleaning motor; bilge, transom and live wells; bait buckets; and fishing apparatus and gear will help to control the spread of adult C. pengoi and resting eggs (Crosier and Molloy (UNDATED))

    Nutrition
    Cercopagids capture their prey with the thoracopods I, retain it by thoracopods II-IV, crush its cuticle by mandibles, and suck the prey body contents (Mordukhai-Boltovskoi & Rivier, 1987; Rivier, 1998). Detailed information on feeding behaviour of C. pengoi is not available. Diet includes mainly small crustaceans, both micro- and mesozooplankton (Mordukhai-Boltovskoi, 1968; Rivier, 1998; Laxson et al. 2003; Gorokhova et al. 2005).
    Reproduction
    As many other cladocerans, Cercopagis pengoi is a cyclic parthenogen. It reproduces parthenogenically during the summer and gametogenically later in the year. The parthenogenically-produced young develop in a fluid-filled dorsal brood pouch that ruptures to release the young. In late summer and autumn, parthenogenic females produce eggs that develop into males and gametogenic females, which copulate. Gametogenic reproduction results in resting eggs that are released when the brood pouch ruptures and which overwinter in the sediment. After a refractory period, development proceeds and neonates hatch in spring-summer, depending on local temperatures, to re-found the population (Mordukhai-Boltovskoi & Rivier, 1987; Rivier, 1998). Sexual females are reproductive only at instars II and III, producing 1-4 resting eggs, while parthenogenic females produce between 1 and 24 embryos; average clutch size decreases gradually from instar I to instar III and from early to late stage of embryonic development (Grigorovich et al. 2000).
    Lifecycle stages
    Females (both parthenogenic and gametogenic) and males possess 3 life-history stages (instars), differing by number of spines (barbs) on the caudal process. At each molt, the animal sheds its exoskeleton to the base of the caudal process. A new pair of proximal barbs and the growth of an intercalary segment are inserted between the existing tail spine and the body. The newborn parthenogenic females (instar I) have one pair of barbs on the caudal process, compact oval embryos in the brood pouch without a pointed apex. The second stage (instar II) has two pairs of barbs and the mature stage (instar III) of the parthenogenic female has a large brood pouch with a pointed apex housing embryos. In males at this stage paired penes behind the last thoracic legs and a toothed hook on the first pair of legs are developed (Mordukhai-Boltovskoi & Rivier, 1987; Rivier 1998). Parthenogenic females of the first generation of C. pengoi proceed through 3 moult yielding 4 pairs of proximal barbs on the caudal process unlike the females of following generations that undergo 2 molts to reach adulthood (Simm & Ojaveer 1999).
    This species has been nominated as among 100 of the "World's Worst" invaders
    Reviewed by: Major update under progress
    Elena Gorokhova, Assistant Professor Dept. of Systems Ecology Stockholm University Sweden
    Compiled by: National Biological Information Infrastructure (NBII) & IUCN/SSC Invasive Species Specialist Group (ISSG)
    Last Modified: Tuesday, 28 November 2006


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