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1 Abstract

2 The Polyodon spathula is a ram suspension feeding planktivore that resides in fresh

3 water habitats. While there are numerous hypotheses about the external morphology of the Paddlefish,

4 none truly support a definite finding.

5 Context and Content

6 The Paddlefish Polyodon spathula is classified as the following: Kingdom: Animalia, Phylum:

7 Chordata, Class: , Order: , Family: Polyodontidae, Genus: Polyodon,

8 Species: spathula (Ross and Brenneman 2001, Starnes 2016). The order Acipenseriformes includes the

9 (family Acipenseridae) and the (family Polyodontidae; Ross and Brenneman

10 2001). The , Psephurus gladius, is the only other species in the family Polyodontidae

11 (Wilkens and Hofmann 2007) and resides exclusively in ’s River (Ross and Brenneman

12 2001). There are a variety of common names associated with the species such as Paddlefish, Spadefish,

13 Spoonbill Catfish, or Duck-Billed Catfish (Ross and Brenneman 2001). Walbaum portrayed the Paddlefish

14 as a shark in 1792; but was corrected by Lacepedѐ in 1797, who classified Paddlefish in the genus

15 Polyodon (McKinley 1984).

16 General Characteristics

17 Paddlefish are an ancient species and exhibit less derived traits. They are typically characterized

18 by the presence of heterocercal caudal fins and the absence of teeth in adults (Ross and Brenneman

19 2001). Polyodon means ‘many teeth’, while Adult Paddlefish do not have teeth, but do have many gill

20 rakers which they use to filter zooplankton from the water column (Ross and Brenneman 2001) and

21 spathula means ‘flat piece’, referring to the long snout, or rostrum, which makes up around one third of

22 the total body length of adults (Wilkens and Hofmann 2007). For the most part Paddlefish have a

23 smooth shark-like exterior appearance and possess a small amount of scales located on the operculum 24 and the bottom lobe of the tail. The Paddlefish skeleton consists mostly of cartilage with the jaws being

25 the only exception (Ross and Brenneman 2001).

26 Distribution

27 Ross and Brenneman (2001) documented Paddlefish in the: Noxubee River system, Pearl

28 drainages of the Gulf of Mexico Basin, and areas of the Mississippi River Basin. Paddlefish have also been

29 documented throughout the Licking, Tennessee, Mississippi, Ohio, Kentucky, and Cumberland rivers and

30 as far north as the Great Lakes (Burr 1980).

31 Form and Function

32 The main theory pertaining to the rostrum of Paddlefish is that it is used as an electrosensory

33 apparatus for locating plankton in aquatic habitats (Wilkens and Hofmann 2007) While earlier writers

34 suggested that the Paddlefish used the elongate rostrum to dig into the benthic layer to locate items for

35 nourishment, later scientific research confirmed that they feed primarily on plankton (Ross and

36 Brenneman 2001).

37 Paddlefish total length is often measured using eye-fork length, taken from the eye to the fork

38 of the tail (Ross and Brenneman 2001). The rostrum of fingerlings and smaller account for most of

39 the body length which may increase error if measuring total length from the paddle to the fork of the

40 tail. Eye-fork length (EFL) is the best method for determining an accurate total body length (Ross and

41 Brenneman 2001).

42 Ontogeny and Reproduction

43 Paddlefish journey upstream each spring to . Upon locating areas having an average water

44 temperature of 12ᵒC, Paddlefish spawn over coarse gravel (Ross and Brenneman 2001). The timing for

45 spawning varies with location as more northern populations mature later and have differing thermal 46 regimes. There appears to be a positive relationship between seasonally warming waters and adult

47 Paddlefish abundance (Miller et al 2011), suggesting spawning depends on water temperature.

48 Initially, eggs float, but once the fertilization process is completed the eggs sink, sticking to hard

49 surfaces for up to one week. Eggs hatch when the water temperature ranges from 18-21ᵒC (Jennings

50 and Zigler 2000). Maturity rates seem to vary by location, with males tending to mature at age six (Ross

51 and Brenneman 2001). In most cases females mature later in life around age nine, although a study

52 concluded that half matured by age nine and half matured at age ten (Reed et al 1992).

53 Conservation

54 Paddlefish roe provides a domestic alternative to purchasing expensive imported

55 caviar (Clarke-Kolaks et al 2009). However, increased poaching to harvest eggs, driven by the

56 compensation fishers receive for roe, has led to rising concerns Paddlefish conservation. In 2006, fishers

57 could earn as much as $22.50 per pound of roe harvested (Bettoli et al 2009). Although this may be an

58 ongoing problem, management plans have established guidelines for legal harvest regulations, such as

59 regulating areas where harvest is still permitted to prevent over-exploitation. As different habitats can

60 support varying numbers of fish, limits must also vary according to current population status. A

61 commercial harvest of Paddlefish was permitted during the summer of 2006 in Mississippi. Possessing a

62 Paddlefish that was over thirty inches in length was prohibited and the catch limit was two fish per day

63 (Bettoli et al 2009).

64 Paddlefish were plentiful in the Mississippi River drainage before 1900 (Jennings and Zigler 2000

65 However, Paddlefish are listed as vulnerable by the International Union for Conservation of Nature

66 (IUCN) (Grady 2004), largely due to habitat alteration. Paddlefish movement is negatively impacted by

67 the construction of as they increase the difficulty of moving upstream to spawn. Increasing river

68 modification, could significantly reduce spawning success and ultimate result in population decline. 69 Coker (1928) participated in a study to determine whether Paddlefish were injured while passing

70 through the Keokuk located on the upper Mississippi River. The results found that Paddlefish were

71 in fact harmed more often than other species of fish, but Paddlefish were able to move through the

72 system at times unharmed (Coker 1928). River modification can result in loss and degradation of

73 spawning habitats due to the destruction of passageways. Jennings and Zigler (2000) cite river

74 modification (along with pollution and over-exploitation) as they discussed environmental issues relating

75 to declines in Paddlefish abundance. Following modification, Paddlefish populations may be confined to

76 an area, resulting in negative impacts on population size, increased competition for resources, and

77 increased rates of disease transmission and mortality. Piers and other channel construction, while not

78 obstructive, may result changes in Paddlefish movement. Wilkens and Hofmann (2007) noted the

79 Paddlefish avoided structures containing steel, perhaps due to abnormal electric signals. Fish in

80 laboratory trials also avoided metal structures. (Wilkens and Hofmann 2007).

81 The Paddlefishes along with sturgeons, are protected by the Convention on International Trade

82 in Endangered Species of Wild Fauna and Flora. This organization was established in order to prevent

83 over-exploitation of any species due to international trade. As extensive concern for Paddlefish

84 populations began to grow people began to get involved with the conservation of the species, as of

85 March 1992 Paddlefish were listed within the Appendix II section titled “Class Actinopterygii” (Jennings

86 and Zigler 2000; CITES 2015). Currently in the state of Mississippi, the Paddlefish is not considered an

87 endangered species, but is listed by Mississippi Natural Heritage Program as a species of special concern

88 (MNHP 2015; Ross and Brenneman 2001).

89

90 Behavior 91 Paddlefish feed on plankton using a method known as ram suspension feeding in which they

92 open their mouths and continuously swim forward, forcing water to flow through their gill rakers, which

93 filter out plankton (Ross and Brenneman 2001). These fish happen to be the only filter feeders in the

94 family Polyodontidae and have adapted specifically for this method of feeding, as their gill arch and jaw

95 configuration produces a wider gape that allows for successful filter feeding (Wilkens and Hofmann

96 2007).

97 Acknowledgements

98 Upon completion of this paper I would like to acknowledge Jason Bies for being a wonderful

99 mentor and teacher throughout the species review. Also, Dr. Mike Colvin who assisted in providing me

100 with a substantial amount of resources and information that led to the successful completion of literary

101 research to make this paper possible.

102 Literature Cited

103 Bettoli, P. W., J.A. Kerns, G. D. Scholten. 2009. “Status of paddlefish in the United States.” American

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106 Clarke, S. J., J. R. Jackson, and S. E. Lochmann. 2009. “Adult and juvenile paddlefish in the floodplain

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108 Coker, R. E. 1928. “Keokuk Dam and the Fisheries of the Upper Mississippi River.” Bureau of Fisheries.

109 Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). 2015.

110 “Appendices I, II, and III.” Available: https://cites.org/eng/app/appendices.php. (Accessed

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130 Starnes, W.C. 2016. “ and Nomenclature of Polyodon spathula.” Integrated Taxonomic

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