Biological Report 82 (11.35) 

March 1985 

Library 

National Wetlands Research Center 

U. S. Fish and Wildlife Service 

700 Cajundome Boulevard 

Larayette, La. 70506 

TR EL-82-4 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (Gulf of Mexico) 

GRASS SHRIMP 

QL 

155 

.S63 

no. 82- 

11.35 

Fish and Wildlife Service 

U.S. Department of the interior 

Coastal Ecology Group 

Waterways Experiment Station 

U.S. Army Corps of Engineers 
This is one of the first reports to be published in the new "Biological 

Report" series. This technical report series, published by the Research 

and Development branch of the U.S. Fish and Wildlife Service, replaces 

the "FWS/OBS" series published from 1976 to September 1984. The Biological Report series is designed for the rapid publication of reports with 

an application orientation, and it continues the focus of the FWS/OBS 

series on resource management issues and fish and wildlife needs. 
Biological Report 82(11.35) 

TR EL-82-4 

March 1985 

Species Profiles: Life Histories and Environmental Requirements 

of.Coastal Fishes and Invertebrates (Gulf of Mexico) 

GRASS SHRIMP 

Gary Anderson 

Department of Biological Sciences 

University of Southern Mississippi 

Hattiesburg, MS 39406-5018 

Project Manager 

Larry Shanks 

Project Officer 

John Parsons 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

1010 Gause Boulevard 

Slidell, LA 70458 

Performed for 

Coastal Ecology Group 

U.S. Army Corps of Engineers 

Waterways Experiment Station 

Vicksburg, MS 39180 

and 

National Coastal Ecosystems Team 

Division of Biological Services 

Research and Development 

Fish and Wildlife Service 

U.S. Department of the Interior 

Washington, DC 20240 
This series should be referenced as follows: 

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories 

and environmental requirements of coasta fishes and invertebrates. U.S. Fish 

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4. 

This profile should be cited as follows: 

Anderson, G. 1985. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (Gulf of Mexico) -- grass shrimp. 

U.S. Fish Wildl. Serv. Biol. Rep. 82(11.35) U.S. Army Corps of Engineers, TR 

EL-82-4. 19 pp. 
PREFACE 

This species profile is one of a series on coastal aquatic organisms, 

principally fish, of sport, commercial, or ecological importance. The profiles 

are designed to provide coastal managers, engineers, and biologists with a brief 

comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be 

expected to react to environmental changes caused by coastal development. Each 

profile has sections on taxonomy, life history, ecological role, environmental 

requirements, and economic importance, if applicable. A three-ring binder is 

used for this series so that new profiles can be added as they are prepared. This 

project is jointly planned and financed by the U.S. Army Corps of Engineers and 

the U.S. Fish and Wildlife Service. 

Suggestions or questions regarding this report should be directed to one 

of the following addresses. 

Information Transfer Specialist 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

NASA-Slide11 Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

or 

U.S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 

iii 
Multiply !% To Obtain 

millimeters (mn) 0.03937 inches 

centimeters (cm) 0.3937 inches 

meters (m) 3.281 feet 

kilometers (km) 0.6214 miles 

square meters (m2) 

square kilometers (km2) 

hectares (ha) 

10.76 

0.3861 

2.471 

square feet 

square miles 

acres 

liters (1) 0.2642 

cubic meters (m3) 

gallons 

35.31 cubic feet 

cubic meters 0.0008110 acre-feet 

milligrams (mg) 0.00003527 

grams (9) 0.03527 

kilograms (k 

metric tons 9 

) 2.205 

t) 2205.0 

metric tons 1.102 

kilocalories (kcal) 3.968 

Celsius degrees 1.8("C) + 32 Fahrenheit degrees 

inches 25.40 millimeters 

inches 2.54 centimeters 

feet (ft) 0.3048 meters 

fathoms 1.829 meters 

miles (mi) 1.609 kilometers 

nautical miles (nni) 1.852 kilometers 

square feet (ft2) 

acres 

square miles (mi2) 

0.0929 square meters 

0.4047 hectares 

2.590 square kilometers 

gallons (gal) 3.785 liters 

cubic feet (ft3) 0.02831 cubic meters 

acre-feet 1233.0 cubic meters 

CONVERSION TABLE 

Metric to U.S. Customary 

U.S. Customary to Metric 

ounces (oz) 

pounds (lb) 

28.35 

0.4536 

short tons (ton) 0.9072 

British thermal units (Btu) 0.2520 

Fahrenheit degrees 0.5556("F - 32) 

iv 

ounces 

ounces 

pounds 

pounds 

short tons 

British thermal units 

grams 

kilograms 

metric tons 

kilocalories 

Celsius degrees 
CONTENTS 

Page 

iii 

iv 

vi 

1 

2 

2 

4 

4 

4 

4 

5 

5 

65 

6" 

; 

8 

8 

8 

; 

10 

10 

11 

13 

w 

PREFACE ........................................................... 

CONVERSION TABLE .................................................. 

ACKNOWLEDGMENTS ................................................... 

NOMENCLATURE/TAXONOMY/RANGE ....................................... 

MORPHOLOGY/IDENTIFICATION AIDS .................................... 

REASON FOR INCLUSIdN IN SERIES .................................... 

LIFE HISTORY ...................................................... 

Spawning ........................................................ 

Fecundity ....................................................... 

Development of Larvae ........................................... 

Maturation and Life Span ........................................ 

ECOLOGICAL KOLE ................................................... 

Food Habits ..................................................... 

Predation ....................................................... 

Parasites ....................................................... 

Behavioral Ecology .............................................. 

GROWTH CHARACTERISTICS ............................................ 

ENVIRONMENTAL REQUIREMENTS ........................................ 

Aquatic Vegetation ............................................. 

Salinity ........................................................ 

Brackish-Water shrimp ........................................ 

Freshwater shrimp ............................................ 

Water Temperature ............................................... 

Dissolved Oxygen ................................................ 

Physical Factors ................................................ 

Bioassays ....................................................... 

LITERATURE CITED .................................................. 

L 

V 
ACKNOWLEDGMENTS 

I am grateful for reviews and suggestions by Richard Heard, Gulf Coast 

Research Laboratory, Ocean Springs, Mississippi; Henry Tatum, U.S. Army 

Corps of Engineers Waterways Experiment Station, Vicksburg, Mississippi; 

and Steven Ross, Department of Biological Sciences, Universlty of Southern 

Mississippi, Hattiesburg, Mississippi. 

vi 
Figure 1. Grass shrimp. a. Palaemonetes pugio; b-d, second chela of p. 

pugio, P. intermedius and P. vulgaris, respectively; e-h, anterior carapace 

and rostrum of P. paludosu?, P. kadiakensis, P. vulgaris, and Macrobrachium 

ohione, respectTvely; i-j, a<t.ennufe of "brackish water" and "freshwater" 

grass shrimp, respectively. Scale of drawings to specimens in mm is (a) 7 to 

1, (b-d) 16 to 1, (e-g) 4 to 1, (h) 1.6 to 1, and (i-j) 2 to 1. 

GRASS SHRIMP 

NOMENCLATURE/TAXONOMY/RANGE Subphylum.....................Crustacea 

Class......................Malacostraca 

Scientific names . . . . . . . . Palaemonetes Or&r . . . . . . . . . . . . . . . . . . . . . . . . . . 

pugio; f. vulgaris; P. intermedius; 

Decapoda 

P. paludosus; P. kadiaensis. 

Family.....................Palaemonidae 

Common name . . ..y........ Grass shrimp Geographic range and speciation: The 

(Figure 1). 

Other names . . . . . . . . . . . Jumpers, glass 

grass shrimp in the Gulf of Mexico 

shrimp, popcorn shrimp, glass prawns, 

area consist of five species, all 

hardbacks. 

relatively similar in morphological 

- 

characteristics and most with over1 
lapping distribution. Because of 

their similarities, these species are 

often misidentified or lumped as p_ 

vulgaris. 

Although p. paludpsus and p_ 

kadiakensis are prlmarl y freshwater 

species, both are somewhat tolerant 

of mesohaline conditions (Maguire 

1961; Dobkin and Manning 1964; 

Strenth 1976) and may be encountered 

in both freshwater and brackish-water 

tidal marshes (Swingle 1971; 

Christmas and Langley 1973; Heard 

1982). 

MORPHOLOGY/IDENTIFICATION AIDS 

Most of the information summarized in this section is taken from 

Holthuis (1952), Williams (1965), 

Wood (1974), and Heard (1982). 

Several diagnostic features 

distinguish the caridean shrimps, to 

which the genus Palaemonetes belongs, 

from the penaeidean shrimps. In the 

caridean forms, the pleura of the 

second abdominal somite overlap those 

of the first, and the third walking 

legs lack claws or chelae (Figure la). 

Palaemonetes spp. and other members of 

th t -1 

diEtingu?:hid 

Palaemonidae may be 

from members of the 

families Alpheidae, Processidae, and 

Hippolytidae by having the second 

walking leg with an unjointed carpus 

(the segment just proximal to the 

chela). The absence of mandibular 

palps in Palaemonetes distinguishes 

this genus from the other palaemonid 

genera Macrobrachium and Palaemon. 

Furthermore, unlike Palaemonetes, 

Macrobrachium bears hepatic spines 

(tlgure lh) and Palaemon floridanus 

has a prominent striped pattern on the 

carapace and abdomen. 

Other morphological features of 

Palaemonetes, as well as many other 

caridean genera and families, include 

the following: well-developed rostrum 

bearing both dorsal and ventral teeth, 

a smooth carapace and abdomen, rounded 

abdominal pleura 1-4, well-developed 

eyes with globular pigmented corneas, 

well-developed spines on the telson 

(two pairs dorsally, two pairs 

posteriorly), and chelate walking legs 

l-2 (the second legs are stronger than 

the first). Grass shrimp are 

transparent to yellowish brown. Few 

exceed 50 mm in total length. Males 

can be separated from females by the 

presence of the appendix masculina 

attached to the appendix interna on the 

endopod of the second pair of pleopods. 

Also, the endopod of the first pleopod 

is larger in males than in females of 

the same age. Additional details were 

provided by Holthuis (1952) and 

Williams (1965). Selected morphological features of adults of the five 

species of Palaemonetes comnon to 

waters of the Gulf of Mexico are 

compared in Figure la through lj and 

Table 1. However, these features 

cannot be used to distinguish between 

juveniles of the brackish-water forms 

of P. vulgaris, P. intermedius, and P. 

pugio -(Boston Zd Provenzano 198n. 

EZ%%al genitalic details of males may 

also be used to distinguish between the 

five species (Fleming 1969). 

REASON FOR INCLUSION IN SERIES 

Brackish water Palaemonetes are 

among the most widely distributed, 

abundant, and conspicuous of the 

shallow water benthic macroinvertebrates in the estuaries of the Atlantic 

and Gulf Coasts (Wood 1967; Odum and 

Heald 1972; Welsh 1975; Sikora 1977). 

Although grass shrimp have only limited 

value as fish bait (Huner 1979) or food 

for cultured fish or humans, their 

ecological importance is unquestioned. 

Grass shrimp have been extensively 

documented as prey of fishes (Gunter 

1945; Darnell 1958; Diener et al. 1974; 

Overstreet and Heard 1982) and other 

carnivores (Heard 1982) and they are 

instrumental also in transporting 

energy and nutrients between various 

estuarine trophic levels: primary 

producers, decomposers, carnivores, and 

detritivores (Johannes and Satomi 1966; 

2 
. . 

Table 1. Summary of morphological differences between five species of Palaemonetes (grass shrimp) in the 

coastal waters of the Gulf of Mexico (Holthius 1952; Williams 1965). 

--- 

Characters P. vulgaris P. intermedius P. kadiakensis - 

P. pugio P. paludosus 

-- - 

-__l__--- 

Number of teeth 

on dactylus; 

fixed finger of 

second leg 

Rostra1 teeth 

w Upper 

antennular 

flagellum 

Position of 

branchiostegal 

spine 

Approximate 

maximum total 

length (mm) 

Number, size, 

and color of eggs 

2/l 

(Figure Id) 

l/O o/o 

(Figure lc) (Figure lb) 

1 7-10 on dorsal 7-10 on dorsa 1 

surface; 1 on surface; tip 

tip; 4-5 on naked; 2-4 on 

o/o o/o 

(Figure lb) (Figure lb) 

6-8 on dorsal 6-8 on dorsal 

surface; tip surface; tip 

naked; 3-4 on naked; 2-3 on 

8-11 on dorsa 

surface; 1 on 

tip; 3-5 on 

ventral surface ventral surface ventral surface ventral surface ventral surface 

(Figure lg) (Figure la) (Figure le) (Figure If) 

Fused part of the two rami shorter than free part 

(Figure li) 

Fused part of the two rami longer 

than free part (Figure lj) 

On anteroventral part of branchiostegal groove (Figure la, le, lg) 

Considerably 

ventrad to 

branchiostegal 

groove 

(Figure If) 

42 42 50 47 53 

Numerous, small, usually grayish brown Few, large, usually greenish 
Adams and Angelovic 1970; Welsh 1975; 

Morgan 1980). 

LIFE HISTORY 

Spawning 

The spawning season of grass 

shrimp extends from February through 

October but may vary with species and 

geographical location. An exception is 

P. paludosus, which spawns all year in 

'SouThern tlorida (Dobkin 1963). In the 

prespawning female, the ripening 

ovaries are discernable as greenish or 

grayish brown masses of tissue dorsal 

and posterior to the stomach, and 

additional setae develop on the ventral 

surface of the abdomen and thorax. The 

female's molting condition and 

subsequent mating and spawning were 

described for P. vulgaris by Burkenroad 

(1947). Hc reported that after 

molting, the female is receptive to the 

male but that apparently the male 

recognizes her condition only if 

physical contact is made with her 

exoskeleton. 

During copulation, which must 

occur within 7 h after molting, the 

ventral surfaces of the partners are 

positioned so that their genital 

apertures are close together. A 

spermatophore is quickly extruded by 

the male onto the genital sternites of 

the female where it remains until 

oviposition. Before oviposition, 

which occurs within 7 h after sperm 

transfer, part of the spermatophore 

dissolves (probably due to enzymatic 

secretions from the female's oviduct) 

and the spermatozoa are released. Ova 

are fertilized externally as they are 

extruded, then manipulated against the 

pleopods and setae on the ventral 

surface of the abdomen where they 

adhere. 

The eggs hatch 12 to 60 days after 

fertilization, depending on species and 

geographical location. In warmer 

climates, the incubation period is 

usually shorter. Osmotic swelling of 

the inner membrane, struggling by the 

larva (protozoea), and ventilatory 

movements by the female assist in 

freeing the larva from the egg membrane 

(Davis 1965). The female molts again 

within a few days after spawning and 

may produce an additional brood, 

depending on the species or time of 

spawning (Broad and Hubschman 1963; 

Knowlton and Williams 1970; Beck and 

Cowell 1976). 

Fecundity 

The fecundity of p. pugio in Rhode 

Island is greater than that of other 

shrimp populations that have been 

studied. For example, in June, the 

average number of eggs per female was 

486 in Rhode Island (Welsh 1975); 37.2 

in Texas (Wood 1967); and up to 247 in 

South Carolina (Sikora 1977). There 

was a significant positive correlation 

between the length (X) of an ovigerous 

female and the number of eggs (Y). 

Wood (1967) reported the relation to be 

log Y = 1.19 + 0.0408 X. Egg counts of 

P. paludosus in Florida ranged from 

only 8 to 85; log Y = 0.6288 + 0.0294 X 

(Beck and Cowell 1976). 

Development of Larvae 

Larvae are planktonic and feed 

upon zooplankton, algae, and detritus. 

Depending on the species and environmental conditions, there may be from 3 

to 11 morphologically distinct stages 

during larval development. Transition 

from one stage to the next occurs 

during molting. Intraspecific genotypic variability in the number of 

larvae and the morphology of different 

stages may enhance dispersal (Sandifer 

and Smith 1979). 

Larval development has been 

described for each of the five species: 

P. vulaaris and P. ouaio bv Broad 

n95mkadiakGisi?& Br"oad and 

Hubschmani (1w paludosus by 

Dobkin (19631. 'and-' P. intermedius _ 

by Hubs&man 'and Broah(1974). borne 

additional interspecific differences in 

life histories, modified from Hubschman 

4 
and Broad (1974), are shown in Table 2. 

The morphology and behavior of larvae 

and postlarvae differ. Larvae lack 

long appendages and swim almost 

continuously with the head down and the 

dorsal surface oriented toward the 

direction of horizontal movement. The 

duration of larval development may 

range from 11 days to several months 

for P. pugio (Floyd 1977), depending on 

environs1 conditions. The final 

larval stage metamorphoses to a 

postlarva, which closely resembles the 

adult. 

Maturation and Life Span 

Juvenile P. pugio mature when they 

are 1.5 to 2 m??ntmd and about 15 to 

18 mm long; their life span is 6 to 

13 months (Alon and Stancyk 1982). The 

older, overwintering shrimp usually 

spawn early in the year and die by the 

next winter. Most young-of-the-year 

spawn late in the year as adults. 

Postlarvae that survive the fall and 

winter spawn in the following spring. 

ECOLOGICAL ROLE 

Food Habits 

Grass shrimp eat a wide variety of 

aquatic foods. Depending on the 

availability of a particular food they 

may be detritivores, primary consumers, 

or secondary consumers (Odum and Heald 

1972; Morgan 1980). 

Table 2. Spawning season, fecundity, lengths (mm) at hatching and at metamorphosis, and number of larval molts for Palaemonetes spp. in the Gulf of Mexico. 

_--- ----e-e---- _ --- -__ --- 

Fecundity Length (rnn)a 

Spawning (maximum '--AT At - Number of larval 

Species season observed) hatching metamorphosis moltsa 

.- 

P. vulgaris 

P. pugio -- 

P. intermedius 

P. kadiakensis 

P. paludosus - 

April-October 

b 

No data 2.3 6.3 7-11 

March-October' 486 d 2.6 6.3 7-11 

May-September 

b 

129 e 3.5 7.0 6- 8 

February-Octoberf 160' 4.4 7.5 5- 8 

Year roundh 85e 3.8 4.5 3 

aData on length at hatching and metamorphosis and number of larval molts are 

from the following sources: P. vu1 aris and P. 

medius, Hubschman and Broad (i97 

ba,~~w~~~al~~o~rl';i~~~k1~g~~~63). 

&adiak%.l%oair$$ l$u1b9~~t!~a~'(fi~ 

iWood (1967). 

Welsh (1975). 

eBeck and Cowell (1976). 

fWhite (1949). 

;Meehean (1936). 

Dobkin (1963). 
The fundamental role of grass 

shrimp as detritivores is explained by 

Odum and Heald (1972), Welsh (1975), 

and Adams and Angelovic (1970). As 

detritivores, grass shrimp aid in the 

mechanical breakdown of refractory 

organic material such as fibrous plant 

materials, as well as assimilate the 

associated microflora, microfauna, and 

fungi (Adams and Angelovic 1970). The 

assimilation of dissolved organic 

matter sorbed to finely divided 

particulate matter such as clay 

particles is important in grass shrimp 

nutrition (Odum and Heald 1972). 

Although grass shrimp often live among 

aquatic macrophytes (Adams and 

Angelovic 1970; Livingston et al. 

1976; Heck and Orth 1980; Morgan 1980; 

Coen et al. 1981; Gore et al. 1981), 

there is little evidence that the 

macrophyte structure is actually 

consumed. More likely, grass shrimp 

eat and assimilate the epiphytic 

microalgae that coat the plant 

structure (Morgan 1980). Grass shrimp 

also are predators of meiofauna and 

small infaunal polychaetes, 

oligochaetes, nematodes (Sikora 1977; 

Bell and Coull 1978; Chambers 1981), 

epiphytic fauna (Odum and Heald 1972; 

Morgan 1980), and even motile prey such 

as mysids (Morgan 1980). As epibenthic 

predators and sediment disturbers, 

grass shrimp alter infaunal community 

structure (Bell and Coull 1978; Knieb 

and Stiven 1982). For example, in 

North Carolina a sharp decline in the 

abumdance of p. puqio due to predation 

by mumnichogs (Fundulus heteroclitus) 

brought about significant changes in 

i nfaunal composition. 

Predation 

In estuaries, numerous fish species 

and other aquatic carnivores, some of 

which are valuable sport and commercial 

fishes, eat large quantities of grass 

shrimp. Grass shrimp are also eaten by 

forage fishes such as Fundulus spp. 

(Harrington and HarringtonWelsh 

1975; Knieb and Stiven 1982), which in 

turn are preyed upon by larger fishes. 

As prey, grass shrimp play an important 

role in the transfer of energy from the 

producer and decomposer levels to 

higher consumer levels. 

Grass shrimp frequently inhabit 

water near underwater structures and 

are particularly attracted to dense 

stands of underwater macrophytes. 

These stands not only support an 

abundance and diversity of food for 

shrimp, but also provide a refuge from 

predators. Grass shrimp are more prone 

to predation when displaced from 

preferred substrata such as macrophytes 

(Coen et al. 1981; Heck and Thoman 

1981) and oyster shells (Thorp 1976). 

Parasites 

Grass shrimp are hosts for 

numerous species of parasites and 

ectocommensals. The most abundant are 

coccidia (Solangi and Overstreet 1980), 

microsporidians (Overstreet and Weidner 

1974), trematodes (Heard and Overstreet 

1983), isopods (Anderson 1977), and 

leeches (Overstreet 1978). The biology 

of other host-parasite interrelationships involving Palaemonetes spp. were 

summarized by Johnson 71977) and 

Overstreet (1978). Diseases and 

parasites do not appear to be major 

factors in limiting the abundance and 

growth of grass shrimp in the Gulf of 

Mexico. 

Behavioral Ecoloav 

Field studies have shown that the 

movement and distributional patterns of 

grass shrimp may be influenced by both 

photoperiod and tidal cycles. Swimming 

of European species, Palaemonetes 

varians, peaked in morning and evening 

'-(7%DiZnisse et al. 1971). The only 

observation on nocturnal movement of P. 

pugio was documented by Shenker aiid 

Dean (1979), who reported that some 

shrimp are buried in the sediments 

during daylight. These studies, as 

well as that of Sikora (1977), 

indicated that grass shrimp in tidal 

creeks migrate seaward (downstream) or 

drift with the current during ebb tides 

6 
and migrate upstream into tidal creeks 

during incoming tides. 

During laboratory experiments on 

agonistic behavior in P. pugio and P. 

vulgaris, females were dominant 0vZr 

males and large shrimp were dominant 

over smaller ones. In interspecific 

pairings, P. vulgaris generally 

dominated P.- pucjmbers 1981), 

and P. vulgaris isown to displace P. 

pugi; from the preferred shell substratum (Thorp 1976). Such displacement 

may favor coexistence between these two 

sympatric species by reducing the 

future potential for agonistic 

behavior. Distributional differences 

between two sympatric species are 

believed to be caused by displacement 

of p. vulgaris by Palaemon floridanus 

from the preferred red algal substratum 

(Coen et al. 1981). 

GROWTH CHARACTERISTICS 

Because populations of grass 

shrimp may produce more than two broods 

a year, length-frequency distributions 

may be polymodal and growth rates are 

difficult to 

1977). 

characterize (Sikora 

Grass shrimp population growth 

characteristics have been described for 

P. pugio in Texas (Wood 1967), South 

rarolina(Sikora 1977; Alon and Stancyk 

1982), North Carolina (Knowlton and 

Williams 1970), and Rhode Island (Welsh 

1975); and for P. paludosus in Florida 

(Beck and Cowell 19/6). Growth rates 

vary somewhat between species, sexes, 

habitats, and times of year. These 

differences are exemplified in part by 

the different daily growth rates of P. 

pugio from two populations in SouTh 

Carolina (Table 3). 

According to Beck and Cowell 

(1976), P. paludosus grows 3.25 mm per 

month i{ summer and fall. In Rhode 

Island, the length of P. . 

increased rapidly from July th‘$%$ 

September; females had then reached a 

total length of about 3 cm and males 

about 2.6 cm. The growth of grass 

Table 3. Mean daily growth increments 

(mm) of Palaemonetes puqio from two 

habitats in South Carolina (data from 

Alon and Stancyk 1982). 

Season 

Summer Winter 

North Inlet 

female 0.143 + c I.111 0.090 + 0.067 

male ..- - 0.087 F 0 I.060 0.086 r 0.041 

Minim Creek - 

temale 0.133 + 0.109 0.089 + 0.041 

male 0.069 2 0.036 0.068 z 0.042 

shrimp was negligible from September to 

May in Rhode Island (Welsh 1975). In 

contrast, one group of P. pugio in 

South Carolina is born Zd recruited 

in early spring, grows rapidly during 

summer, and dies before winter. A 

second group is recruited in late 

summer, grows rapidly in autumn and 

again the following spring,and usually 

dies by midsummer (Alon and Stancyk 

1982). These age and growth 

characteristics make it difficult to 

tabulate lengths of grass shrimp of 

different ages. 

In the southern and southeastern 

coastal waters of the United States, 

overwintering P. pugio spawn in late 

February thr~gh~rch at water 

temperatures of 15" to 20°C. Juveniles 

from the February-March hatch mature 

and spawn from late summer (JulyAugust) to November. By then the 

length distribution is highly bimodal. 

Shrimp of the summer-fall hatch mature 

from October to February and reproduce 

during the following spring. Shrimp 

hatched in spring grow faster than 

those hatched in summer and fall but 

have a shorter life span. 

In colder coastal waters, growth 

patterns are different. In Rhode 

Island, this species spawns only once 

during May to July in any one year. 

Length distribution is consequently 

unimodal throughout the year (Welsh 

1975). 

7 
Salinity is yet another factor 

that affects growth. In South 

Carolina, p. pugio matures and Spawns 

at a younger age in habitats with 

relatively high salinity than in those 

with relatively low salinity (Alon and 

Stancyk 1982), but specimens collected 

in low salinity waters were smaller 

than those from more saline waters 

(Wood 1967). 

ENVIRONMENTAL REQUIREMENTS 

Aquatic Vegetation 

Grass shrimp, macrophytes, and 

algae live in the same communities and 

are subject to many of the same 

environmental factors (Beck and Cowell 

1976; Thorp 1976; Morgan 1980; Coen et 

al. 1981; Heck and Thoman 1981). 

Studies in Florida showed that P. pugio 

was most abundant in liabm 

characterized by aquatic macrophytes, 

relatively high turbidities, and low 

salinities (Livingston et al. 1976). 

Because grass shrimp depend on 

aquatic vegetation in many coastal 

waters, alterations of estuaries that 

destroy vegetation could seriously 

reduce their abundance. In one 

instance, weirs built in marshes to 

benefit wildlife produced dense masses 

of aquatic vegetation that supported an 

abundance of grass shrimp far higher 

than that in nearby less densely 

vegetated waters (Weaver and Holloway 

1974). In another study in West Bay, 

Texas, dredging, bulkheading, and 

filling of coastal marshes caused a 

permanent loss of intertidal 

vegetation. The reduction of detrital 

input into the adjoining aquatic 

systems caused a noticeable decrease in 

abundance of grass shrimp (Trent et al. 

1976). 

Salinity 

The abundance of freshwater and 

brackish-water grass shrimp in 

estuaries iS clear evidence of their 

survival in a wide range of 

salinities. 

Brackish-water shrimp. Field and 

laboratory studies indicate that P. 

pugio adults tolerate salinities from-0 

to 55 ppt but are most common in 

salinities of 2 ppt to 36 ppt (Wood 

1967; Swingle 1971; Bowler and 

Seidenberg 1971; Christmas and Langley 

1973; Kirby and Knowlton 1976; Morgan 

1980). The 96 h LDso values for adults 

are 0.5 and 44 ppt (Kirby and Knowlton 

1976). 

Attempts to assess salinity 

tolerances of larvae on a broad scale 

have yielded mixed results. McKenney 

and Neff (1979) were able to rear 

nearly 50% of the P. pugio larvae 

exposed to a salinity ?'f-3 ppt, 

whereas, Broad and Hubschman (1962) 

reported poor survival of larvae at 

salinities less than 10 ppt. When 

larvae were exposed to a wide range of 

arbitrarily selected salinities, the 

low and high salinities required to 

kill 50% of the larvae during a 96-h 4 

exposure (LD50) were 16 ppt and 46 ppt, 

respectively (Kirby and Knowlton 1976). 

Optimum salinity for complete larval 

development is 20 to 25 ppt (Floyd 

1977; McKenney and Neff 1979; Knowlton 

and Kirby 1984). Possible reasons for 

the conflicting findings are 

differences in populations in different 

geographic locations, and differences 

in laboratory procedures. 

P. vulgaris is more tolerant of 

high Salinity water and less tolerant 

of low salinity water than is P. pugio 

(Holthuis 1952; Knowlton and -Williams 

1970; Bowler and Seidenberg 1971; Thorp 

and Hoss 1975). Although Nagabushanam 

(1961) reported that salinities as low 

as 3 ppt were lethal to adult P. 

vulgaris, they have been collected in 

freshwater (Swingle 1971; Christmas and 

Langley 1973; Barrett et al. 1978). 

Schoen and Knowlton (1977) reported 

96-h LDao values of 0.8 and 51 ppt for 

adults. Larvae of p. vulgaris are less 

tolerant of low salinity than are 

adults. Larvae thrive at salinities 

8 
between 10 ppt and 30 ppt; the optimum 

is 20 ppt (Sandifer 1973). Larvae do 

not survive at salinity extremes of 5 

and 35 ppt (Knowlton 1965; Schoen and 

Knowlton 1977). Larvae of both P. 

pugio and P. 

mature early, 

vulgaris survive besT, 

and often pass through 

fewer larval stages when salinities are 

near optimum (Sandifer 1973; Floyd 

1977). Interspecific differences in 

salinity tolerance in estuaries are 

likely to segregate P. pugio and P. 

vulgaris (Bowler and sidenberg 1971J, 

although biotic interactions such as 

competition for food and space are 

probably more important for segregation 

and coexistence (Thorp 1976). 

Although P. intermedius is 

euryhaline, it has been collected from 

waters with salinities of 5 to 39 ppt 

(Dobkin and Manning 1964). The larvae 

thrive at salinities of 20 ppt but 

survival is low at salinities less than 

10 ppt (Broad and Hubschman 1962). The 

larvae have been reared at salinities 

near 30 ppt (Hubschman and Broad 1974). 

The two 

paludosus and P. 

kadiakensis, often -live in brackish 

waters (Christmas and Langley 1973). 

P. paludosus was reported in salinities 

of 0 ppt to 10 ppt by Tabb and Manning 

(1961), and one specimen was captured 

in a salinity of 25 ppt (Swingle 1971). 

In a laboratory experiment, P. 

paludosus survived for 7 days at-a 

salinity of 30 ppt but died when the 

salinity was raised to 37 ppt. In 

another experiment, adult P. kadiakensis survived at salinities up to 20 

ppt but died at 25 ppt (Maguire 1961; 

Strenth 1976). Larvae are relatively 

intolerant of salinities higher than 5 

ppt (Hubschman 1975; Strenth 1976) but 

are more tolerant of higher salinities 

after they reach stage IV (Hubschman 

1975). 

The salinity tolerance of grass 

shrimp is summarized in Table 4. 

Water Temperature 

Grass shrimp are eurythermal. In 

coastal waters, P. pugio thrives at 

temperatures of 5"to38"C (Wood 1967; 

Christmas and Langley 1973), but 

survival is greater at 18" to 25°C. 

Growth is most rapid in waters at 

temperatures above 30°C but drops 

rapidly at water temperature below 14°C 

(Wood 1967). This species breeds at 

Table 4. Salinity tolerance limits and optima (ppt) reported by various 

investigators (see text discussion) for adult and larval Palaemonetes. 

-- -----__- -__--_I_-____-__-- -__ 

Form and species Adults Larvae 

Limits Uptimum Limits Optimum 

- 

Freshwater forms 

P. kadiakensis O-25 0 O-10 0 

lJ. paludosus O-30 0 No data 0 

Brackish-water forms 

P. intermedius 5-39 No data 10-30 20 

P. pugio l-55 4-16 3-31 25 

P. vulgar-is 1-51 No data 5-35 20 

9 
temperatures of 22" to 27°C in Rhode 

Island (Sastry and Vargo 1977) and from 

17" to 38°C in Texas (Wood 1967). 

Preferred water temperatures range from 

5" to 35°C for P. vulgaris and P. 

paludosus and from-10' to ST for p. 

paludosus (Christmas and LangEy 

T?j73).Tolerance for warmer waters 

is greater for P. pugio than P. 

vulgaris, but 60th - species are 

relatively tolerant of the heat stress 

of effluents from a power plant (Chung 

1977). The breeding temperatures for 

P. paludosus range from 18" to 33°C 

-(Beck and Cowell 1976). 

Physiological responses of grass 

shrimp to temperature indicate that P. 

vulgaris adults (McFarland and Pickens 

1965)d P. pugio larvae (Sastry and 

Vargo 197q are metabolically active 

over a broad range of water temperatures, exhibiting physiological 

compensation to seasonal temperature 

changes. 

Temperature tolerance data for 

grass shrimp are summarized in Table 

5. 

Dissolved Oxygen 

Dissolved oxygen is another factor 

that helps regulate the distribution 

and abundance of grass shrimp. In 

Louisiana waters. P. vulaaris and P. 

pugio are common i-n F lssol _ ved oxyg& 

(DO) concentrations of 6 to 11 ppm 

(Barrett et al. 1978). In Rhode 

Island, the mortality of P. vulgaris 

was higher than that of P.-jjuglo, when 

both species were cagedtogm in a 

hypoxic microhabitat (DO = 0.3 ppm) 

during a tidal cycle. In the 

laboratory, at oxygen concentrations 

lower than 1 ppm, survival was higher 

in P. pugio than in P. vulgaris at all 

tesr temperatures (IX)' to 30°C). In 

nature, grass shrimp sometimes climb 

out of the water during periods of 

oxygen deficiency, especially during 

warm summer niqhts, but such attempts 

to avoid hypoxi‘a can be effective only 

for a few hours (Pomeroy and Wiegert 

1981). Respiratory studies indicated 

that P. pugio is an oxyconformer; its 

oxygen uI decreases as oxygen 

tension declines from 8 to 2 ppm (Welsh 

1975; Dillon 1983). 

Physical Factors 

Grass shrimp usually inhabit the 

shallows near the water's edge but 

have been reported at depths as great 

* 

as 8 fathoms (Williams 1965). In 

winter, during temperature lows and in 

summer, when water temperatures 

approach seasonal highs, p. pugio moves 

from shallow to relatively deep water 

(Wood 1967). Thorp (1976) noted that 

the extent of the movement of grass 

shrimp among various depths often 

coincides with the distribution of 

Table 5. Temperature tolerance range and optima (OC) reported for adult and 

larval Palaemonetes. 

-__-.-. 

Species 

-.---_ -_--__ --_ --m-Y 

Adults Larvae 

Limits Optimum Limits Optimum 

P. paludosus 10-35b 

5-38a,b 

18-33' No da a !l No data 

v. pug10 

P. vulgaris 5-35b 

18-25a 15-35 pf 20-30f 

No data 20-30e 2oe 

Sources: --- 

-iChristmas and Langley 1973; b - Beck and Cowell 1976; c - Wood 1967; 

d - McKenney and Neff 1979; e - Floyd 1977; f - Sandifer 1973 

10 
oyster shell substrates, which; in some 

waters, are preferred by p. pugio and 

p. vulgaris. Oyster beds provide food 

and protection. 

Grass shrimp are abundant where 

turbidity is relatively high. This is 

particularly true in habitats where 

water currents tend to keep sediments 

suspended, such as in shallow tidal 

creeks or near river mouths. Turbidity 

may provide some degree of protection 

from predators in habitats with little 

vegetation. In less turbid habitats, 

macrophytes afford protection from 

predators. Although Livingston et al. 

(1976) found that grass shrimp abundance was positively correlated with 

turbidity, turbidity may not be an 

environmental requirement. In some 

habitats, grass shrimp have been 

observed in clear water such as is 

often associated with dense aquatic 

vegetation (Weaver and Holloway 1974). 

Grass shrimp tend to avoid fast 

currents and migrate in the direction 

of tidal currents (Antheunisse et al. 

1971; Sikora 1977; Shenker and Dean 

1979). 

Bioassays 

In as much as grass shrimp have 

been recommended for use as bioassay 

test organisms (American Public Health 

Association 1975), much information has 

been published about mortality and 

sublethal effects of various toxicants 

on grass shrimp but only a few are 

mentioned here. Workers evaluating the 

effects of kepone and other insecticides (heptachlor, toxaphene, dieldrin, 

and endosulfan) conclude that P. pugio 

adults were usually less sensiTive and 

showed a lesser tendency to bioconcentrate these toxicants than did the fish 

that were tested (Schimmel and Wilson 

1977; Schimmel et al. 1977). For 

example, the 96-h LDso values for 

kepone were 121 ug/l and 70 ug/l for P. 

u io and for the sheepshead minnow 

Y- Cyprinodon variegatus). Bioconcentration factors were 698 for the shrimp 

and 1,548 for the minnow (Schimmel and 

Wilson 1977). On the other hand, grass 

shrimp are much more sensitive than 

fish to endrin (Tyler-Schroeder 1979), 

dithiocarbamates (Rao et al. 1982), 

various chlorophenols (Rao et al. 

1981>, and DOT and parathion (Sanders 

1972). On the basis of a study of 12 

insecticides, Eisler (1969) concluded 

that grass shrimp are more sensitive to 

organophosphorous insecticides, and 

less sensitive to several organochlorine insecticides at high salinity 

than at low salinity. Low temperatures 

enhanced the survival of shrimp exposed 

to both groups of insecticides. Also, 

for most organophosphorous compounds 

tested, crustaceans were more sensitive 

than marine fishes by several orders of 

magnitude. 

Heavy metals such as cadmium are 

more toxic to grass shrimp than to 

mummichogs, Fundulus heteroclitus 

(Eisler 1971). Concentrations of heavy 

metals (mercury, cadmium, and chromium) 

are acutely toxic to adult grass 

shrimp, usually in the order of 100 to 

1,000 ppb. Except in areas of sewage 

or industrial outfalls, such concentrations far exceed those in estuaries. 

Therefore, grass shrimp are too 

resistant to be of much value in heavy 

metal bioassays (Vernberg et al. 1977). 

Increased temperature or decreased 

salinity increases the sensitivity of 

grass shrimp to heavy metals (Fales 

1978; Sunda et al. 1978). Sublethal 

effects of heavy metals on grass shrimp 

have included loss of a predator 

avoidance response (Barthalmus 1977), 

developmental abnormalities in larvae 

(Shealy and Sandifer 1975), and a 

reduced tolerance to salinity fluctuations (Middaugh and Floyd 1978). 

Attempts have been made by Tatem 

(1976), Tatem et al. (1978), and Dillon 

(1982) to evaluate the effects of 

exposure to petroleum hydrocarbons on 

survival and various physiological 

phenomena in grass shrimp. Several 

conclusions have been drawn from this 

research: (I) concentrations of 

petroleum hydrocarbons normally found 

in saltwater (10 to 20 ppb) and its 

11 
sediments are too low to have acutely 

toxic effects on grass shrimp; (2) when 

sediments and water are exposed 

chronically to oil pollution or during 

a large oil spill, hydrocarbon 

concentrations could cause acute 

toxicity; (3) metabolism, reproduction, 

and growth may be reduced or altered if 

oil concentrations persist near 1 ppm; 

(4) grass shrimp rapidly depurate 

accumulated petroleum hydrocarbons when 

they are returned to oil-free water; 

and (5) the toxicity of oil is 

correlated with the relative proportion 

of the aromatic fraction present. 

Some investigators have evaluated 

the effects of biocides, such as those 

used to remove fouling organisms from 

cooling systems of powerplants 

(generally chlorine), on grass shrimp. 

In a review by Hall et al. (1979) 

it was concluded that the detrimental 

effects of chlorination on grass shrimp 

were negligible under normal powerplant 

operations. 

Additional investigations indicated 

that P. pugio was less sensitive than 

mysid-shrimpto drilling muds but that 

both were several orders of magnitude 

more sensitive than fishes (Conklin et 

al. 1980); grass shrimp were more 

sensitive than pinfish (Lagodon 

rhomboides) to a simulated rm 

eftluent containing phenol, sulfide, 

ammonia, and some other components 

(Hall et al. 1978); and grass shrimp 

were more sensitive than other 

crustaceans to ionizing radiation. 

Radiation affects metabolic pathways 

involved in the synthesis of 

non-essential amino acids used in 

osmotic regulation (Engel et al. 

1974). 

12 
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Academic Press, New York. 

Sanders, H.O. 1972. Toxicity of some 

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But-. Sport Fish. Wildl. Tech. Pap. 

66:1-19. 

Sandifer, P.A. 1973. Effects of 

temperature and salinity on larval 

development of arass ShrimD 

Sandifer, P.A., and T.I.J. Smith. 1979. 

Possible significance of variation 

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Biol. Ecol. 39:55-64. 

Sastry, A.N., and S.L. _Vargo. 1977. 

Variations in the physiological 

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Physiological responses of marine 

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Schimmel, S.C., and A.J. Wilson, Jr. 

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Sci. 18:224-227. 

Schimmel, S.C., J.M. Patrick, Jr., and 

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Pages 241-242 in F.L. Mayer, and 

J.L. Hamelink, eds. Aquatic 

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ASTM STP 634. American Society for 

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Schoen,. R.H., and R.E. Knowlton. 1977. 

Salinity lethal limits and sodium 

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Shealy, M.H., and P.A. Sandifer. 1975. 

Effects of mercury on survival and 

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. 

.- . 

Shenker, J.M., and J.M. Dean. 1979. 

The utilization of an intertidal 

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Smithsonian Contrib. Zool. 228. 

17 
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18 
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coast. 

19 
1027Y ‘101 

REPORT DOCUMENTATION 1’. “c’?‘- ~0. 2. 

PAGE 1 6101. Rep. 82(11.35)* 

c nt* and ~btitl. 

Species profiles: life histories and environmental require 

ments of coastal fishes and invertebrates (Gulf of Mexico)-- 

Department of Biological Sciences 

University of Southern Mississippi 

Hattiesburg, MS 39406-5018 

12 S~onswln~ Or#pnlutlon Npmp and Addmr 

National Coastal Ecosystems Team U.S. Army Corps of Engineers 

Division of Biological Services Waterways Experiment Station 

Fish and Wildlife Service P.O. Box 631 

U.S. Department of Interior Vicksburg, MS 39180 

Washington, DC 20240 

10. l miectlT~sk/Wer* Unit No. 

11. bntmct(C) 01 Gnnt(G) No. 

(C) 

(G) 

*U.S. Army Corps of Engineers Report No. TR EL-82-4 

IL ~r8ct(Umit:2oorro*) 

This species profile summarizes much of the literature published about the biology of 

grass shrimp, Palaemonetes spp. Five species (p. pugio, k. vulgaris, p. intermedius, p. 

paludosus, and P. kadiakensis) are common in coastal waters of the northern Gulf of 

Mexico, although? the last two are primarily freshwater forms. The brackish-water species 

are among the most abundant shallow-water benthic decopods in estuaries and play 

important roles as food for fishes and as transporters of nutrients among various trophic 

levels. They spawn from spring to autumn and the growth of planktonic larvae is normally 

completed in 7 to 12 molts. Postlarvae of 7-10 mm are 1.5 to 2 months old. The life 

span of grass shrimp is 6 to 13 months (maximum size rarely exceeds 50 mm). Palaemonetes 

spp. are eurythermal, euryhaline, and are relatively tolerant to hypoxia and a variety of 

pollutants. Grass shrimp feed on detritus, epiphytes, and meiofauna. They tend to 

concentrate near underwater structures and plants, especially in dense stands of 

underwater macrophytes. 

Estuaries Growth 

Fishes Feeding 

b. IdwWi~n/Own-Ended Twms 

Grass shrimp 

Palaemonetes spp. 

Temperature requirements 

c. COSATI Fidd/Gmup 

Life history 

Habitat requirements 

Spawning 

L Availability Statement 

Unlimited 

1. ANsl-z39.1SI OPTIONAL FORY 272 (67 

(Formwly NTI.%35) 
Speues proiiies : 

Anderson, Gary LO 

2518ir 

Qti55 A3 no.82 

Kesearch Library 

U.S. Department of the Interior 

National Biological Service 

Southern Science Center 

700 Cajundome Blvd. 

* Headquarters. DIVISION of EilologG3 

Services, Washington. DC 

x Eastern Energy and Land Use Team 

Leetown. WV 

l Natlonal Coastal Ecosystems Team 

Slldell. LA 

l Western Energy and Land Use Team 

Ft Collms, CO 

l Locations of Regional OffIces 

REGION 1 

Regional Director 

U.S. Fish and Wildlife Service 

Lloyd Five Hundred Building, Suite 1692 

500 N.E. Multnomah Street 

Portland, Oregon 97232 

REGION 4 

Regional Director 

U.S. Fish and Wildlife Service 

Richard B. Russell Building 

75 Spring Street, SW. 

Atlanta, Georgia 30303 

REGION 2 

Regional Director 

U.S. Fish and Wildlife Service 

P.O. Box 1306 

Albuquerque, New Mexico 87 103 

REGION 3 

Regional Director 

U.S. Fish and Wildlife Service 

Federal Building, Fort Snelling 

Twin Cities, Minnesota 55 I I 1 

REGION 5 

Regional Director 

U.S. Fish and Wildlife Service 

One Gateway Center 

Newton Corner, Massachusetts 02158 

REGION 6 

Regional Director 

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P.O. Box 25486 

Denver Federal Center 

Denver. Colorado 80225 

REGION 7 

Regional Director 

U.S. Fish and Wildlife Service 

101 I E. Tudor Road 

Anchorage, Alaska 99503 
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