Daphnia lumholtzi

Invasion History

First Non-native North American Tidal Record: 1994
First Non-native West Coast Tidal Record: 1999
First Non-native East/Gulf Coast Tidal Record: 1994

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General Invasion History:

Daphnia lumholtzi is a cladoceran native to Southwest Asia, East and North Africa, and Eastern Australia. Its large size and prominent head and tail spines make it conspicuous in plankton samples. In 1991, it was collected in Stockton Lake, Missouri and Lake Fairfield, Texas. A likely mechanism of introduction was an attempt to introduce Nile Perch (Lates niloticus) to Lake Fairfield in 1983. Alternatively, other aquarium fish or plant releases could have brought this cladoceran to North America. Allozyme comparisons, and later microsatellite analysis, indicate that Africa is the likeliest source of D. lumholtzi populations in North America (Havel and Hebert 1993; Havel et al. 2000; Frisch et al. 2012). Subsequently, populations were detected in reservoirs, swamps, and rivers in 22 states from Texas, Florida, New Mexico, and California north to Kansas, Minnesota, Maryland, and the Province of Ontario (Havel and Hebert 1993; Davis and Gobler 2011; Tudorancea 2009; Trebitz et al. 2010; USGS Nonindigenous Aquatic Species Program 2012). Genetic analysis supports the picture of an initial introduction and spread from the Southeastern US, with a later northward expansion (Frisch et al. 2012). Most of the occurrences are in inland reservoirs, suggesting that trailered boats, fishing gear, and natural dispersal by birds may be important local vectors.

North American Invasion History:

Invasion History on the West Coast:

In 1999, Daphnia lumholtizi was collected in several samples in the Sacramento-San Joaquin River Delta, at Clifton Court Forebay and Mildred Island, and is now established there (Orsi 2002). We have no recent information on the distribution of this species in the San Francisco estuary, or in other West Coast estuaries.

Invasion History on the East Coast:

By the late 1990s, Daphnia lumholtzi's range on the Atlantic slope extended from Lake Okeechobee, Florida to reservoirs on the Piedmont of North Carolina (USGS Nonindigenous Aquatic Species Program 2012). In 1999, Daphnia lumholtzi was collected in tidal fresh waters of the James River estuary, Virginia. It re-occurred in 2000, increasing in abundance, but was not found in other Chesapeake tributaries at that time. In the James River, D. lumholtzi was still much less abundant than other Daphnia spp. (Mateja 2000). In 2007-2008, D. lumholtzi was collected in the tidal fresh Transquaking River, Dorchester County, on the Eastern Shore of Maryland (Davis and Gobler 2011). In 2003, it was collected in the St. Johns River system (Florida), in Lake George (tidal, 0.3-0.8 PSU), and Lake Jesup (nontidal). Occurrences were sporadic (2003, 2006) in Lake George, but more regular in small, nontidal, eutrophic Lake Jesup (Havens et al. 2012).

Invasion History on the Gulf Coast:

In 1994, Daphnia lumholtzi was found to be abundant in the tidal lower Atchafalaya River, Louisiana at 19 of 30 sites sampled (Davidson and Kelso 1997). The river system intersects with the Intracoastal Waterway, so that dispersal of D. lumholtzi to many of Louisiana's coastal bayous was likely. In 2002, it was collected in the estuarine headwaters of Mobile Bay, Alabama, at salinities up to 1.5 PSU (deVries et al. 2006).

Invasion History Elsewhere in the World:

In 2000, Daphnia lumholtzi was reported in the Três Irmãos reservoir, on the Tiete River (tributary of the Parana River), in Sao Paulo state, Brazil (Zanata et al. 2003). In 2003, and 2008, it was found in two floodplain lakes of the Upper Parana River, Parana State. It is expected to spread extensively in Brazil (Simões et al. 2009). In Mexico, D. lumholtzi was collected in Sonora (Elías-Gutiérrez et al. 2008), and the Presa El Salto reservoir, in Sinaloa in 2003 (Silva-Briano et al. 2010). The spotty nature of these tropical occurrences could reflect the scarcity of sampling or else a very scattered pattern of dispersal.

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Description

Cladocerans of the genus Daphnia are covered by a bivalve carapace that covers the thorax and abdomen, and terminates in an apical spine. The head projects ventrally, forming a short, downward-pointing beak. The carapace is open ventrally, providing a space through which water can flow, as the animal creates a feeding current with its thoracic appendages. The abdomen is curled downward and backward, ending in the posterior part of the space between the two halves of the carapace. The posterior surface (postabdomen) bears claws, used for cleaning the thoracic appendages. The head has a small, dark compound eye. The antennules are very small, but the two antennae are large and branched, with fringed setae. They are the primary means of locomotion. There are five thoracic appendages, which are biramous and used for filter feeding. The female (males are usually rare) carries eggs and developing embryos in a brood pouch. Morphology of Daphnia can vary seasonally. The anterior and apical spines of many species vary greatly according to environmental conditions, especially those that affect the risk of predation. This pattern of regular seasonal change is called cyclomorphosis. Description based on Barnes 1983.

For most of the year, reproduction is parthenogenetic, so males are usually uncommon. They have a longer antennule and setae than females, and on the first thoracic leg, a copulatory hook. During parthenogenetic reproduction, reproduction is direct, and newborn juveniles are miniature versions of the adults. However, fertilized females produce two large, thick-walled eggs, enclosed in the cast-off brood-chamber, called an ephippium.

Daphnia lumholtzi is morphologically distinctive. Its head-spine (helmet) is larger than that of any native species and its tail spine is equal to, or greater than, its body length. There is a depression (cervical sinus) separating the dorsal base of the head from the rest of the body. The posterior part of the head bears two projecting structures called fornices (fornix = arch or fold), which in this species are projected into sharp points. The ventral carapace bears about 10 sharp spines on each side (Havel and Hebert 1993).

Male D. lumholtzi usually have a head without a helmet, or occasionally with a small spike-shaped crest. The tail spine is about two-thirds the carapace length (Zanata et al 2003). The ephippia (resting egg capsules) of D. lumholtzi have a long point on each end and a dorsal surface covered with fine hairs (Havel and Hebert 1993).

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Taxonomy

Ecology

General:

Cladocerans of the genus Daphnia can develop parthenogenetically from unfertilized eggs in a female's brood pouch. The juveniles have the basic form of miniature adults, and molt and grow as they feed. After a period of growth, parthenogenetic eggs are deposited in the female's brood pouch. For D. lumholtzi at 15-25 C, the eggs were produced at 8.7-4.7 days after birth, and took 1.9- 1.1 days to develop (Tifnouti et al. 1993). When the embryos hatch, the female molts, and a new batch of eggs are deposited in the brood pouch. A female may have several successive broods. Female D. lumholtzi at 20-25 C, on average, had 6.9 - 6.4 broods, consisting of 2.2-2.4 eggs each, and lived for up to 30 days. With parthenogenetic development, and a generation time of 6-9 days, D. lumholtzi is capable of rapid population growth (Tifnouti et al. 1993).

Daphnia spp. can reach high population densities by parthenogenetic reproduction, developing populations that are all female or female-dominated. Environmental factors such as temperature, day-length, the presence of predators, etc., appear to stimulate the production of males. Females, when fertilized, produce large resting eggs, in pairs, enclosed in a capsule formed by modifications of the brood chamber (called an ephippium). The ephippium is cast off when the female molts. It settles into the sediment, but may be stimulated to development by favorable conditions of light and temperature. Sexual reproduction of resting eggs provides a means of surviving winter, droughts, or other adverse conditions, as well as providing new genotypes for potentially altered conditions (Barnes 1983). We are not aware of studies on specific cues affecting sexual reproduction, or hatching of resting eggs in D. lumholtzi.

Daphnia sp. are filter-feeders, creating a current and filtering out phytoplankton in the water column. They have limited capacity for selective feeding (Barnes 1983), but may reduce filtering rates in the presence of large chains of diatoms, or large colonies of toxic organisms, such as cyanobacteria of the genus Microcystis sp. (Gifford et al. 2007; Davis and Gobler 2011).

Daphnia lumholtzi appears to be especially adapted for semi-desert tropical and subtropical areas, prone to wide temperature variations, and some degree of saline conditions (Havel and Hebert 1993; Tifnouti 1993; Work and Gophen 1995; Work and Gophen 1999). It appears to be more tolerant of high temperatures than most other Daphnia (Tifnouti et al. 1993; Fey and Cottingham 2011; Engel and Tollrian 2012). Many temperate Daphnia populations grow rapidly in spring, and then collapse in early or mid- summer. Reasons for this appear complex, and may involve a mixture of temperature responses, predation, and inadequate or toxic phytoplankton. Daphnia lumholtzi appears better suited than many native species to persist in summer or tropical conditions (Work and Gophen 1998; Johnson and Havel 2001; Fey and Cottingham 2011; Engel and Tollrian 2012). This cladoceran has also been found in inland saline lakes (e.g. Lake Texoma, OK-TX, 1.4 ppt; Work and Gophen 1995, Mobile Bay, 1.4 PSU), but the full extent of its salinity tolerance is unknown. Some Daphnia species can tolerate high salinities in inland salt lakes (Hairston et al. 1999), but this genus is a rare straggler in estuarine environments (Johnson and Allen 2005).

Food:

Phytoplankton

Consumers:

Fishes

Competitors:

Daphnia spp.; other zooplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Nontidal Freshwater	None
General Habitat	Unstructured Bottom	None
General Habitat	Canals	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Planktonic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	4	Daphnia lumholtzi survives in ice-covered lakes as resting eggs (Havel et al. 1993). Dense swarms of adult females were observed in a Kentucky reservoir, at a water temperature of 8 C (DBeaver et at. Al. 2018).
Maximum Temperature (ºC)	32	Field (Lake Okeechobee FL, Havens et al. 2012).
Minimum Salinity (‰)	0	This is a freshwater species.
Maximum Salinity (‰)	1.5	Field, Mobile Bay AL (DeVries et al. 2006)
Minimum Reproductive Temperature	15	Experimental (Tifnouti et al. 1993)
Maximum Reproductive Temperature	30	Field (MO, Havel and Graham 2006)
Minimum Length (mm)	1.1	Lake St. Clair (Tudorancea et al. 2009). (The length includes spines.)
Maximum Length (mm)	5.7	Lake Erie (Muzinic 2000) (The length includes spines.)
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Nontidal Limnetic-Oligohaline

General Impacts

Daphnia lumholtzi is a recent arrival in North America, and is still expanding its range. Its impacts are being studied in a number of lakes and reservoirs in the midwestern and southeastern US. The extent of its range and abundance in North American estuaries is not known. Available data suggest abundances were usually low in estuarine habitats (<1000 m-3): Chesapeake Bay (Mateja 2000; Davis and Gobler 2011); St. Johns estuary, FL (Havens et al. 2012); Mobile Bay, AL (de Vries et al. 2006); Atchafalaya River, LA (Davidson et al. 1997); Sacramento-San Joaquin Delta, CA (Orsi 2002). However, many of these records are quite old, and current abundances are not available for most of these estuaries. Experiments in inland fresh lakes and reservoirs suggest that significant impacts are possible.

Competition: Mesocosm experiments in Missouri showed that D. lumholtzi suppressed the increase of the native D. parvula in late summer and fall, but not birth rates, suggesting that the suppression of D. parvula resulted from increased death rates, possibly due to juvenile starvation (Johnson and Havel 2001). However, field surveys and experiments in Missouri reservoirs indicate that relations between D. lumholtzi and native Daphnia are largely complementary, with D. lumholtzi co-occurring with natives, and dominating in summer conditions when native species are normally scarce, while natives dominate in cooler periods, as they had prior to the D. lumholtzi invasion (Havel and Graham 2007). Field data in Lake Okeechobee, FL suggested a similar pattern for D. lumholtzi and D. ambigua (East et al. 1999). In laboratory culture experiments, the native D. pulicularia was the superior competitor in mixed cultures at 20 C, while D. lumholtzi dominated at 28 C (Engel and Tollrien 2012).

The picture becomes more complicated when food is added as a variable. Daphnia lumholtzi grew faster than the native D. pulex at 27 C when fed the green alga Scenedesmus acutus, but when cyanobacteria (Anabaena flosaquae, Microcystis aeruginosa, or both) were added, D. pulex dominated at 27 C. Daphnia lumholtzi's population growth was adversely affected by cyanobacteria at 20 C, too (Fey and Cottingham 2011). Since cyanobacteria are favored by rising temperatures, these conflicting factors make future invasions of D. lumholtzi difficult to predict. In Florida lakes, while D. lumholtzi is widespread, the native D. ambigua is usually dominant, perhaps because smaller Daphnia are more successful on low-quality food, or less vulnerable to visual predators (Havens et al. 2012).

Differential predation also plays a role in zooplankton interactions. In Lake Springfield, IL, the invasion of D. lumholtzi was followed by a change in species composition, with decreased abundance of native cladocerans. Kolar et al. (1998) speculated that competition with native cladocerans in late summer and fall, may affect spring recruitment of the native species. The species co-occur only briefly, but during this time competition, combined with differential predation, may be affecting native cladocerans, which then are overwintering or producing resting eggs, and providing next spring's generations. Predation by fishes may reinforce the effects of competition, since fishes tend to select the less spiny native cladocerans (Kolar et al. 1998). In experiments with mesocosms containing D. lumholtzi, native D. pulex, with and without juvenile Pumpkinseed (Lepomis gibbosus) fish, D. lumholtzi predominated at higher temperature in the presence of fish, because of its higer temperature tolerance, predator defenses, and the increasing rates of predation at higer temperatures (Fey and Herren 2014).

Food/Prey: Initial research suggests that this spiny cladoceran may be altering the available food supply for juvenile fishes, potentially affecting recruitment (Swaffer and O'Brien 1996; Kolar et al.1998). Larval Bluegill Sunfish in laboratory feeding trials preferred the native, shorter-spined D. magna or D. pulex to D. lumholtzi, which were often visually rejected. Visual observation indicated that smallest fish had the greatest trouble ingesting D. lumholtzi (Kolar and Wahl 1998; Swaffer and O'Brien 1996).

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References

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