Distribution and ecology of Gammarus tigrinus Sexton, 1939
and some other amphipod Crustacea
near Beaufort (North Carolina, U.S.A.)
by
Marion J. van Maren
Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands
&
Duke University Marine Laboratory, Beaufort, North Carolina, U.S.A.
Summary of North America, has at some time been intro-
summer distribution of duced in During 1977 the and ecology am- western Europe and was first described
phipod Crustacea in the coastal plain of north Carolina were from England by Sexton (1939). Gammarus ti- studied. Ecological data were collected in particular on has invaded all Gammarus tigrinus, a North American species, which has grinus rapidly nearly oligohaline
been introduced in western Europe. The present gammarid, waters in the Netherlands, competing successfully
able to endure high water and adapted to a temperatures with the indigenous gammarid species. Its range wide variety of salinities, is found in the more upstream still has been parts of the estuaries in North Carolina. More downstream, extension, proceeding, surveyed by
at it is Gammarus Taxonomie higher salinities, replaced by palustris , members of the Institute of Zoology while low salinities fresh at very or in water Gammarus of the University of Amsterdam, resulting in a fasciatus is commonly met. series of some data are publications & Moreover, given on the distribution and (Nijssen Stock, 1966;
of several other in the Beaufort ecology amphipod species Pinkster & Stock, 1967; Dennert et al., 1968; region. Gras, 1971; Smit, 1974; Pinkster, 1975; Dieleman
Résumé & Pinkster, 1977).
Pendant l'été 1977 la répartition écologique de Crustacés STUDY AREA AND METHODS amphipodes a été étudiée dans la plaine littorale de Caroline
du Nord. Des observations furent effectuées en particulier sur The small town of Beaufort, situated on a tongue l’écologie de Gammarus tigrinus, espèce nord-américaine in- of mainland, is water from the troduite en Europe. Ce Gammare, capable de résister à des separated by sandy offshore known Shackleford Ac- températures élevées et adapté à des salinités très variables, bank, as Bank.
est trouvé dans les des parties plus en amont estuaires de the is cess to open ocean gained by the Beaufort Caroline du Nord. Plus en aval, dans les eaux méso- et Inlet. The various waterways, which communicate polyhalines, Gammarus tigrinus est remplacé par Gammarus
tandis dans les eaux douces et with palustris, que oligohalines directly or indirectly Beaufort, are very com- Gammarus fasciatus est couramment rencontré. in and contain number plicated shape a large of En plus, cette étude a fourni des données sur la répartition shoals, banks and islands. et l'écologie de plusieurs autres espèces d'Amphipodes dans la
région de Beaufort. While offering a variety of sandy and muddy
natural rocks substrates, no true are found in the INTRODUCTION Beaufort region. Hard substrates are formed by During summer 1977 the distribution and ecology jetties, sea walls, pilings, oyster shells and by of amphipod crustaceans was investigated in the man-made miscellaneous objects (bottles, beer coastal of plain North Carolina. The The coastal the present study cans, etc.). area of mainland is
was carried out during a at the Duke Univer- stay characterized by extensive Spartina marshes. The Marine Beaufort. sity Laboratory at Amphipod tidal is for mean range 0.75 m (about 0.9 m spring
were taken in estuaries and freshwater samples tides).
biotopes in the Beaufort to collect data on 1 region The estuary of the Newport River ), a wide the ecology of Gammarus tigrinus in particular. Irrespective of !) the salinity or current, the water types in This species, originally inhabiting the eastern coast the area are locally called "rivers".
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bay with polyhaline salinities, is connected by the
Adam Creek Canal with the Neuse River. In
summer, the freshwater supply of the latter is low
to such an extent that marine influences are still
measurable as far as New Bern, a town situated at
almost 100 km from where the Neuse River
discharges into the Pamlico Sound.
The of the river of the vegetation swamps coastal plain is characterized by the River-cypress
( Taxodium distichum),, growing in shallow water,
often over a calcareous subsoil. This tree has a
trunk the and gradually tapering near base root
formations, which often provide the only hard
in the the substrate more upstream parts of North
Carolina estuaries.
Sampling of the amphipods was carried out with
a dip net. As far as temperature readings have been
thermometer used in made, a mercury was (scale
measured with the aid 0.1°C). Salinities were of a
refractometer. The pH was determined with a pH-
meter "Leeds & Northrop".
The animals were fixed in 4% formaldehyde and preserved in 70% ethanol.
In appendix II the position of the sampling
stations, visited during the present study, is given.
RESULTS
shows the of the Fig. 1 names amphipod species
sampled at different localities in the estuary of
the Neuse River, as well as the salinities measured
these stations. Near at the bridge on highway 43,
of S.W. Askin, gammarids still occurred, but more
upstream, near Kinston, none have been found.
The great majority of the Neuse River stations
presented a sandy substrate. Only in those places
where some hard substrate was available (mostly
the the provided by roots of River-cypress) gam- Fig. 1. Salinities and amphipods at the sampling stations in
in the = maridean Amphipoda were captured wave- the Neuse River (station K Kinston).
washed zone of the river.
The temperature of the water varied from In another tributary, the South River (discharg-
18.5°C in May to 34°C in August, while the pH- ing in the Neuse River about 50 km downstream
of range was 6.3-7.3. New Bern), Gammarus palustris Bousfield,
in Both stations the Trent River, a occurred a of sampled 1969, at salinity 12°/ 00 and, together
of the Neuse Gammarus with Mucro tributary River, yielded gammarus mucronatus (Say, 1818), at
Sexton, 1939 1 and a of These were tigrinus (salinities 2°/ 00 ). salinity 18°/00 . amphipods cap-
Substrate, temperatures and pH in this stream tured amongst submerged parts of the riverbank
Neuse and resemble to a large degree thoseof the River vegetation, on a bottom consisting of clay
estuary. mud.
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Appendix I enumerates the amphipod species boro, and in the Bay River (sta. 61), a tributary
the well the the Pamlico Sound. the investi- found during present study, as as of During present
their this salinities conditions at sampling stations. gation species was sampled at varying
The have been collected: from 0 to once it was found at following amphipods 12°/ 00 (only
22°/oo)> always on sandy substrates, among roots Family GAMMARIDAE (s.l.) of the localities G. River-cypress. In some tigrinus Gammarus tigrinus Sexton, 1939 occurred together with other amphipods: In the
Besides in the Neuse and Trent Rivers, Gammarus Neuse River (figs. 1 & 2), near New Bern, to-
tigrinus was found in the White Oak River (fig. gether with Gammarus daiberi Bousfield, 1969
in the Sound Swans- and Heard 2), discharging Bogue near Corophium aquafuscum & Sikora,
Fig. 2. Distribution of the amphipod species N.W. of Beaufort.
4
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at a of more downstream, some devoid of such a G. 1972, salinity 2°/ 00 ; places, vegetation, pa-
the contained collected shells at a salinity of 12°/ sample also lustris was amongst oyster or tree 00 ,
Corophium lacustre Vanhöffen, 1911, Mucrogam- roots.
marus mucronatus (Say, 1818) and Gammarus While collecting, the present species was easily
its behaviour: palustris Bousfield, 1969; near Cherry Point (sta. recognized by particular the shrimps
6), the latter two occurred together with G. climbed out of the water (in the vial) and were
at a of 10°/ in the White Oak able to "rise to their feet", like tigrinus salinity 00; beachhoppers
do. River, at a salinity of l°/oo> only single specimens (Talitridae) use to
of G. palustris and C. lacustre were sampled to- Ovigerous females and/or precopulae were
gether with G. tigrinus. found from May to August.
in Ovigerous females of G. tigrinus occurred a
sample taken in June (salinity l°/oo> temp. Mucrogammarus mucronatus (Say, 1818)
25.5°C). Mucrogammarus mucronatus occurred sometimes,
together with Gammarus palustris, in the Spartina Gammarus daiberi Bousfield, 1969 marshes, but never at salinities lower than 10°/00 .
On mixed with Gammarus daiberi, a species closely related to muddy substrates, cobbles and
with the Gammarus tigrinus, occurred latter in the shells, the present gammarid was found together
samples from the head of the Neuse River estuary, with Melita nitida Smith, 1873 (viz., the Town-
at salinities varying from 0 to 7°/oo- creek at Beaufort). In that case the vegetation
consisted of Ulva, Fucus vesiculosus or Chaeto-
Gammarus fasciatus Say, 1818 morpha. The salinities measured for Mucrogam-
marus varied from to mucronatus 10 32°/ . Gammarus fasciatus was collected mainly in mod- 00 females of the oc- in Ovigerous present species erately running streams and sometimes slowly in in curred a sample collected May. running or stagnant water. Water temperatures
varied from 15.5 to 27°C, the pH from 5.6 to 7.5, Crangonyx serratus (Embody, 1910) and the salinity values in places where G. fasciatus occurred did not exceed 5°/oo- h lives in shallow Crangonyx serratus seems to prefer stagnant water
small it freshwater streams or small marshy ponds (to- of marshy ponds, although sometimes
with in with gether Crangonyx serratus (Embody, 1910)), occurs running water, not rarely together
in varying depth from 0.3 m to a few meters, Gammarus fasciatus. The salinities at stations with
varied among submerged parts of the marsh vegetation, Cr. serratus from 0 to 5°/oo- When alive,
the shows characteristic on a sandy or muddy bottom. present amphipod a milky
In May and June ovigerous females of G. fas- white colour.
ciatus have been observed. In May ovigerous females of Cr. serratus were
observed.
Gammarus palustris Bousfield, 1969
Gammarus 2 is palustris ) a very common species Crangonyx pseudogracilis Bousfield, 1958
in the meso- and polyhaline waters around Beau- At two localities in freshwater biotopes, besides fort. This gammarid, found at salinities of 8 to Gammarus fasciatus and Crangonyx serratus, Cran- at vide 36°/oo (exceptionally l°/oo, supra), occurred the marsh gonyx pseudogracilis among occurred substrate in the mostly on a sandy Spar- and vegetation, on a bottom of sand mud. Both
tina marshes or sometimes among marine algae samples, collected in May, contained ovigerous Fucus In (viz., vesiculosus, Ulva, Chaetomorpha). females.
2 ) Dr. E. L. Bousfield informed me (in litt., 9 May 1978) Melita nitida 1873 that “Gammarus material consists of Smith, my palustris” two distinct G. the species: true palustris, occurring at higher Melita nitida in lower reach- was found mainly the and the of salinities, a new species of gammarid, description in es of estuaries or the tidal zone along the coast which is soon to be published by him.
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of the mainland of North Carolina. This to be tolerant water tem- amphi- appeared very to high
well Accord- pod was collected in muddy or sandy places, among peratures as (34°C during August!).
marine algae ( Fucus vesiculosus)), in the Spartina ing to Hynes (1955), G. tigrinus seems adapted
under shells. Downstream in the with marshes or oyster to a climate warm summers and springs
Neuse River it occurred with rise in estuary together characterized by a rapid temperature. Such
Gammarus palustris and Corophium lacustre (fig. marine climatic conditions prevail along the coastal
!)• plain of North Carolina: in the Neuse River
In the rise in takes in May ovigerous females of present species estuary a rapid temperature place
were found at a of 32°/ while the water attains salinity 00 . early spring, temperature
rather values high during summer (fig. 3).
Family COROPHIIDAE
Corophium lacustre Vanhöffen, 1911
collected in the Corophium lacustre was wave-
salinities washed zone of rivers, at varying from 1
to on substrates or in burrows in 20°/ oo , sandy
dead tree stumps.
In August ovigerous females were found.
Corophium aquafuscum Heard & Sikora, 1972
Corophium aquafuscum occurred in the Neuse
River (upstream of New Bern) in stagnant water
of the substrate of at a salinity 2°/ consisting 00 ,
sand, debris and tree roots.
Besides the sampling stations in the coastal plain of North Carolina, some waters more inland, near
Chapell Hill, were visited, which yielded no am-
from few of phipods, apart a specimens Hyalella azteca De Saussure, 1857 (family Hyalellidae),
in dammed called collected a creek, Hogans Pond
(Orange County).
Fig. 3. Yearly temperature cycle in the Neuse River near DISCUSSION New Bern (after data by Phibbs, 1969). Fox & Bynum (1975) mention Gammarus tigrinus
be abundant in the of the Steele Steele mention G. to to very oligohaline areas & (1975) tigrinus
and Bous- the Neuse River. Boesch & Diaz (1974) be more successful in southern parts of the field found the in the United States at (1973) present species (more generations per year higher
and of than in the North. oligo- mesohaline waters upper Chesapeake temperatures), Salinity pref-
Bay and its tributary estuaries. The results obtained erence experiments by Dorgelo (1975) proved
the that G. is selective during the present study extend distribution of tigrinus distinctly towards
lower while G. tigrinus southward into the headwaters of salinity at temperatures, at a tempera- of variable. Pamlico Sound and Bogue Sound in North Caro- ture 25°C the results are very Ac- lina. cording to Pinkster (1975), the ability of the
G. shows wide has to at tigrinus a salinity range as present gammarid reproduce high tempera- is been demonstrated also by the experiments of tures, while salinity reduced, explains its success
the Dorgelo (1974, 1975). Not only was it found at in competing indigenous gammarids in the widely different salinities in North Carolina, it Netherlands. Although the mixohaline species
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Gammarus zaddachi Sexton, 1912 and G. duebeni G. tigrinus occurred together with G. daiberi. In duebeni Liljeborg, 1852, can reproduce throughout the Delaware and Chesapeake Bays the latter spe-
the that the is abundant in the year provided salinity high enough, cies is most head of the estuaries
do when the they are unable to so salinity drops at salinities ranging from 1 to 5°/oo (Bousfield,
to the level prevailing in most biotopes inhabited 1969). According to Feeley & Wass (1971) G. by G. tigrinus. Under suboptimal salinity condi- daiberi occupies the niche between G. fasciatus
tions, G. zaddachi and G. d. duebeni are only able and the mesohaline G. tigrinus.
to low reproduce at temperatures. - Gammarus fasciatus was found to occur in the
In North Carolina the salinities established for freshwater creeks discharging in the Neuse and
G. from 0 to The Rivers to Bousfield tigrinus vary 12°/00 . present Newport (fig. 2). According
species was found only in larger streams, often (1958) and Holsinger (1972) G. fasciatus lives
in Never in well in in stagnant or very slowly running waters. lakes as as large rivers, biotopes
has found in smaller it been freshwater creeks, which it was never observed in the Beaufort
where G. More fasciatus was commonly met. region.
in estuaries G. is — The Gammarus in the downstream tigrinus replaced by occurrence of palustris
G. A marshes of palustris. similar distribution pattern was Spartina North Carolina was already established for G. tigrinus in the northern parts recorded by Fox & Bynum (1975) and Cammen of the U.S.A. by Steele & Steele (1972), where the (1976). Also near its northern distribution limit,
of this be limited in distribution gammarid seems to New Hampshire, this gammarid was mentioned
the of G. and to in banks by presence fasciatus upstream, occur Spartina (Gable & Croker,
seawards G. lawrencianus The observations its by Bousfield, 1956, a 1977). concerning presence
in from marine the data very common species the tidal zone New- among algae confirm by Watling &
foundlandto Connecticut. In view of the situation Maurer (1972). The substrate preference experi-
is that in the described above, it interesting to see ments of Van Dolah (in press), who studied G.
does invade Netherlands G. tigrinus neither run- palustris from the salt marshes in the Chesapeake
fresh where ning waters, Gammarus pulex pulex region, indicate a strong behavioural preference
(Linnaeus, 1758) thrives, nor waters of higher of this species for Spartina culms.
in salinities which it is apparently unable to com- Bousfield (1973) records an optimal salinity
with G. zaddachi and G. duebeni for G. of but mentions pete (Pinkster range palustris 5-20°/0 o,
et al., 1977). that it is also able to survive, for shorter periods,
— Gammarus daiberi in low in and Gammarus tigrinus fresh water (at temperatures) or sea-
resemble to such an extent that the former was water (at high temperatures). During summer
as a distinct Al- 1977, it was found at salinities from only recently recognized species. varying 0°/ O o
both can be to though gammarids easily recognized (temp. 20°C) 36°/ 00 (temp. 30°C). Although
in the Delaware and material G occurred low it Chesapeake region, . palustris at very salinities, was
from the southeastern states shows overlapping of never observed in permanent fresh water.
The behaviour characters to a large degree (Bousfield, 1969)- semi-terrestrial of G. palustris,
Thanks the kindness of Dr. E. L. Bousfield is in to to as observed during the present study, agree- check the present samples of Gammarus tigrinus, ment with the observations by Bousfield (1973), the presence of Gammarus daiberi could be de- who mentions it to remain hidden in moisty places tected in the from the debris when samples most upstream (under stones, and among Spartina), parts of the Neuse River estuary. It occurred to- the tide is out.
with gether G. tigrinus at salinities varying from — In agreement with the data by Watling &
0 to This is in with the data Maurer Mucro was 10°/ 00 . agreement (1972), gammarus mucronatus of Ristich et al. (1977), who found G. tigrinus found in the more sheltered places of the tidal
G. and fasciatus to be the most common amphi- zone.
in the On M. the pods freshwater zones of the Hudson River comparing mucronatus from Beau-
estuary, while in the oligo- and mesohalinereaches fort region with specimens from Louisiana (in the
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collection of the Smithsonian Institution in Wash- conditions, the vegetation consisting of River-
dorsal mucronations shown the and ington), the by cypress Spartina.
former animals are weakly developed, while those
in the southern ACKNOWLEDGEMENTS samples possess very pronounced
Bousfield M. I indebted for dorsal elevations. (1969) refers to am to the Netherlands' Organization the
Advancement of Pure Research (Z.W.O.) for providing me mucronatus, originating from the coastal region of with which has allowed the a grant, me to carry out present the southeastern states, as "micromucronate forms". study during the months of May, June and July 1977, as well Barnard also mention the variation & Gray (1968) as to the Duke University (Durham, North Carolina) for a travel Furthermore thanks grant during August. I owe many in development of the pleonal teeth in the present to Prof. John D. Costlow, director of the Duke Marine and it in species state that might reflect variations for and Laboratory, receiving me in his laboratory putting the
facilities physical factors or genetic isolation. A similar of the biological station at my disposal. I want to
express gratitude as well to Dr. J. L. Barnard for allowing of the dorsal is recorded my variability processes by work me to some days in his laboratory with the amphipod Karaman & Pinkster for members of the (1977) of the collection of the Smithsonian Institution samples at my
Prof. Gammarus which like I am also grateful to H. Stock, Drs. F. roeseli-group, are, M. mu- disposal. J. Peeters-Pieters and Dr. S. Pinkster for their critical comments the of cronatus, characterized by presence dorsal on the manuscript of this paper; and to Mr. J. Zaagman for mucronations on the metasome segments. his assistance in making the figures.
— Embody (1910), describing Crangonyx serratus Special thanks are due to Dr. E. L. Bousfield of the Natural for number Museum of checking a the first in Sciences, Ottawa, for time, mentions it from a pond of my samples and for reviewing the manuscript of the Virginia. Bousfield (1958) indicates its occur- present paper. rence in sloughs, ponds and ditches in Virginia,
South Carolina and northern Florida. During the REFERENCES Cr. present investigation serratus was not only- BARNARD, J. L. & W. S. GRAY, 1968. Introduction of an found in stagnant water, but has been observed in amphipod crustacean into the Saltan Sea. Bull. Sth. Calif.
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surface waters of North Carolina, 1966-1967. Bull. Wat. WATUNG, L. & D. MAURER, 1972. Marine shallow water
Carol. of Pollut. Control Div., N. Dep. Wat. Air Resour., amphipods the Delaware Bay area, U.S.A. Crusta-
11 (1): 83. ceana, Suppl. 3: 251-266.
APPENDIX I
Amphipod species and environmental conditions at the sampling stations visited near Beaufort (the position of these stations is given in appendix II).
(A) GAMMARUS TIGRINUS
Station Temp. pH Salinity Substrate Vegetation Remarks (°C) (0/oo)
6 30 10 sand + wood Chaetomorpha among tree roots and in wood burrows
11 18.5 7.1 7 sand + shells + wood in wave-washed zone; among tree roots
12 22 7.1 5 sand + detritus riverbank vegetation few gammarids
6.9 5 sand + wood in wave-washed 13 23 zone; among tree roots
28 7.1 12 sand + 24 clay Spartina in wave-washed zone
6.8 mud + detritus brown 34 25.5 1 Sagittaria slowly running water; ovigerous 9 9
37 24 7.1 2 clay + detritus riverbank vegetation
46 26.5 6.5 1 sand + detritus riverbank vegetation
56 34 6.3 2 sand + detritus + wood riverbank vegetation stagnant water; among tree roots
sand + wood 57 29.5 7.2 0 slowly running water; precopulae
7.8 22 61 33 mud + detritus riverbank vegetation very few gammarids
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Station Temp. pH Salinity Substrate Vegetation Remarks (°C) (%«)
(B) GAMMARUS DAIBERI
11 18.5 7.1 7 sand + shells + wood in wave-washed zone; among tree roots
sand + wood in wave-washed IJ 23 6.9 5 zone; among tree roots
57 29.5 7.2 0 sand + wood slowly running water
61 33 7.8 22 mud + detritus riverbank vegetation very few gammarids
(C) GAMMARUS FASCIATUS
riverbank brown 14 18.5 6.9 0 mud vegetation stagnant water; ovigerous 9 9
sand + detritus + wood 20 15.5 7.5 0 moderately running water; ovigerous
9 9
sand + mud riverbank 21 7.4 0 vegetation moderately running water; ovigerous
9 9
sand mud 22 15 6.8 0 + marsh vegetation moderately running water; ovigerous
9 9
detritus 23 7.1 0 sand + slowly running water
27 20 6.8 0 sand + mud marsh vegetation moderately running water; ovigerous
9 9
29 21 7.0 0 mud marsh vegetation slowly running water
30 20.5 6.9 1 sand + mud marsh vegetation stagnant water; precopulae and ovigerous
9 9
32 22.5 6.7 1 mud + detritus marsh vegetation slowly running water
36 20 6.5 1 sand + mud riverbank vegetation moderately running brown water
marsh 38 20 7.2 0.5 clay vegetation moderately running water; many gam-
marids; precopulae and ovigerous 9 9
sand detritus 39 18.5 6.9 5 + moderately running water; precopulae and ovigerous 9 9
43 6.6 sand marsh brown 23 0.5 vegetation moderately running water; pre- copulae
52b 25 6.6 0 sand + shells moderately running water
55 27 5.6 0 mud marsh vegetation moderately running water
(D) GAMMARUS PALUSTRIS
vesiculosus 1 sand + mud + shells Fucus under shells, on rather dry substrate
2 sand + cobbles Ulva under cobbles
riverbank 3 23 12 clay + detritus vegetation moderately running water detritus riverbank 4 23 12 clay + vegetation slowly running water; precopulae
5 23 12 clay + detritus Chaetomorpha slowly running water
6 30 10 sand + wood in wave-washed Chaetomorpha zone; among tree roots
10 20 8 sand riverbank vegetation in 7.3 wave-washed zone; precopulae mud detritus 17 7.5 32 + Spartina slowly running water; precopulae
18 22 6.9 15 mud + detritus Spartina few slowly running water; very gam-
marids
25 24.5 6.9 15 sand + wood in wave-washed zone; among tree roots; ovigerous 9 9
28 7.1 12 sand 24 + clay Spartina in wave-washed zone; precopulae and
ovigerous 9 9
31 26 6.8 11 sand + wood in wave-washed zone; among tree roots
and other wood
33 6.6 sand + wood in wave-washed wood 25.5 9 zone; among
44a 34 sand S 7.5 partina stagnant water; precopulae 44b 26.5 6.9 29 sand + shells Spartina
46 26.5 6.5 1 sand + detritus riverbank vegetation very few gammarids; slowly running
water
47 sand + + shells Spartina in wave-washed 30 6.3 31 clay zone; precopulae
48 29 6.9 29 clay Spartina, Salicornia stagnant water
mud S 49 34 6.9 32 partina stagnant water Ulva 50 32.5 7.1 31 mud + sand + shells stagnant water
sand 51 29.5 6.6 22 Spartina slowly running water
sand wood 58 30 7.1 21 + among tree roots
59 30 20 sand + detritus + wood Spartina
sand detritus 62 35 7.3 24 + + shells stagnant water; precopulae
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Station Temp. pH Salinity Substrate Vegetation Remarks
(°C) (»/co)
sand 63 30 7.7 36 Spartina precopulae
sand and 64 29.5 7.8 18 Spartina precopulae ovigerous 9 9
(E) MUCROGAMMARUS MUCRONATUS
1 sand + mud + shells Fucus vesiculosus under shells, on a rather dry substrate
2 sand + cobbles Ulva under cobbles
5 23 12 clay + detritus Chaetomorpha slowly running water
10 sand + wood in wave-washed 6 30 Chaetomorpha zone, among tree roots; ovigerous 9 9
7 32 sand + clay + shells Ulva under oyster shells
sand + wood wave-washed 25 24.5 6.9 15 - in zone, among tree roots
28 24 7.1 12 sand + clay Spartina in wave-washed zone
44b 26.5 6.9 29 sand + shells Spartina
47 30 6.3 31 sand + clay + shells Spartina in wave-washed zone
49 34 6.9 32 mud Spartina stagnant water
mud + sand + shells Ulva 50 32.5 7.1 31 stagnant water
(F) CRANGONYX SERRATUS
8 16 6.7 1 clay + detritus stagnant brown water; ovigerous 9 9
15 7.2 0 mud marsh vegetation
mud + detritus 19 15.5 7.1 0 marsh vegetation stagnant water; ovigerous 9 9
20 15.5 7.5 0 sand + detritus + wood moderately running water
22 15 6.8 0 sand + mud marsh vegetation moderately running water
24 6.8 0 mud + detritus stagnant water
detritus 26 21 6.9 0 mud + marsh vegetation stagnant, muddy water
29 21 7.0 0 mud marsh vegetation slowly running water
30 20.5 6.9 1 sand + mud marsh vegetation stagnant water
detritus 32 22.5 6.7 1 mud + marsh vegetation slowly running water
35 21 6.9 1 clay + detritus Sagittaria slowly running water
39 18.5 6.9 5 sand + detritus moderately running water
mud + detritus riverbank 40 19.5 6.6 0.5 vegetation stagnant water
41 20 6.2 1 sand + mud + detritus stagnant water
42 20 6.9 1 sand riverbank vegetation moderately running water
43 23 6.6 0.5 sand marsh vegetation moderately running water
45 21 6.9 1 mud marsh vegetation moderately running water
mud 53 27 5.9 0.5 marsh vegetation stagnant water; few gammarids
54 27 5.6 1 mud moderately running water
55 27 5.6 0 mud marsh vegetation moderately running water
(G) CRANGONYX PSEUDOGRACILIS
6.8 sand + mud marsh 22 15 0 vegetation moderately running water; ovigerous
9 9
sand mud 30 20.5 6.9 1 + marsh vegetation stagnant water
(H) MELITA NITIDA
1 sand + mud + shells Fucus vesiculosus under shells, on a rather dry substrate
sand shells 7 32 + clay + Ulva under oyster shells; ovigerous 9 9
44b 26.5 6.9 29 sand + shells Spartina
47 30 6.3 31 sand + clay + shells Spartina in wave-washed zone
50 32.5 7.1 31 sand + clay + shells Ulva stagnant water
58 30 7.1 21 sand + wood - in wave-washed zone 59 30 20 sand + cobbles + shells Spartina
60 33 6.9 20 sand + mud + wood in wave-washed zone
62 35 7.3 24 sand + detritus + shells - stagnant water; few gammarids
(I) COROPHIUM LACUSTRE
28 24 7.1 12 sand + clay Spartina in wave-washed zone
46 26.5 6.5 1 sand + detritus slowly running water
sand shells wood 52a 31 6.4 11 + + in wave-washed zone
58 30 7.1 21 sand + wood in wave-washed zone
cobbles 59 30 20 sand + + wood Spartina in wave-washed zone; ovigerous 9 9 mud 60 33 6.9 20 sand + + wood in wave-washed zone; ovigerous 9 9
(J) COROPHIUM AQUAFUSCUM
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APPENDIX II
Position of the sampling stations as mentioned in fig. 1 and appendix I. The stations marked with an asterisk are not included in the map (fig. 2).
Station Date Position County
(1977)
1 3-V Beaufort, beach E. of bridge near Pivers Island Carteret
left bank of Carteret 2 3-V Newport River, upstream bridge on highway 70
*3 4-V South River, left bank near village South River Carteret
*4 4-V South River, right bank Carteret
*5 4-V South River, left bank, about 50 m downstream of jetty of D.U.M.L. boat Carteret
6 5-V Neuse River, right bank upstream of landing stage of Cherry Point ferry Carteret
7 8-V Beaufort, Towncreek, midway bridge and town Carteret
8 10-V tributary of Newport River, Newport, near bridge on highway 70 Carteret
of Craven 9 10-V Southwest Prong, N.W. Havelock, highway 70
bank of Riverdale Craven 10 10-V Neuse River, right near Fisher Landing, N.E.
11 10-V Neuse River, right bank near Thurman, S.F.. of New Bern Craven
12 10-V Trent River, left bank, S.W. of New Bern Craven
Neuse left of New Bern Craven 13 10-V River, bank, upstream bridge near
Craven 14 10-V Cahooque Creek, E. of Havelock, near bridge on road to Cherry Point
Mortons Mill N.W. of Craven 15 10-V Pond, Harlowe, near bridge on highway 101
16 10-V Harlowe Creek, near Harlowe Church, N. of Morehead City Carteret
Gale of 24 Carteret 17 13-V Creek, Bogue Sound, upstream bridge on highway
N.E. IS 13-V Pettiford Creek, of Swansboro, upstream of bridge on highway 58 Carteret
small Broad Creek 1124 Carteret 19 13-V stream discharging in (Knoll) near bridge on highway
20 15-V Cedar Swamp, W. of Newport, near bridge on highway 1140 (Roberts Road) Carteret
Carteret 21 17-V Little Deep Creek near Union Point, N.E. of Newport
Creek, of road Carteret 22 17-V Deep N. Newport, near bridge on loop
23 17-V Black Creek near Mill Pond, E. of Newport, near bridge on highway 1154 Carteret
Creek of Carteret 24 17-V Little Swamp, E. Newport, near bridge on highway 1154
25 20-V Adams Creek, left bank, where road from Harlowe ends in Great Neck Craven
Craven 26 20-V Great Neck Creek, S.W. of Type
Grove 1711 Craven 27 25-V Mitchell Creek, near Piney Church, bridge on highway
Craven 28 25-V Neuse River, right bank, near Temple, mouth of Clubfoot Creek
101 Craven 29 25-V Hancock Creek, E. of Havelock, near bridge on highway (New Bern Road)
Craven 30 31-V small stream discharging in Neuse River, near Cherry Point
Craven 31 31-V Neuse River, right bank, near Shade View Beach, downstream of Neuse River ferry
32 31-V East Prong near Havelock, highway 101 Craven
33 3-VI Neuse River, right bank, Flanner Beach, near camping ground of Croatan Craven
National Forest
Trent of 34 3-VI Brice Lees Brook, near River, S.W. New Bern Craven
35 3-VI Ready Brook, near bridge on highway 100 i, S.W. of New Bern Craven/Jones
36 3-VI Island Creek, N.E. of Pollocksville, near bridge on highway 1004 (Island Creek Jones
Road)
37 3-VI Trent River, near Mussy Cove, midway Pollocksville and New Bern Craven/Jones
small left bank of Trent of 38 3-VI stream on River, S.W. New Bern Craven
39 8-VI Jason Brook, W. of Newport, near bridge onby-way (dirt road) of highway 1124 Carteret
Peak of road of 40 8-VI Swamp, W. Newport, near bridge on by-way (dirt 128) Carteret
highway 1124
41 8-VI Mills Swamp, W. of Newport, near bridge on by-way (dirt road 128) of Carteret
highway 1124
42 8-VI Northwest Prong near Holly Springs, N.W. of Newport, bridge on highway 1124 Carteret
small of Carteret 43 8-VI stream near Masontown, N.W. Newport
44a 10-VI beach midway Beaufort and Morehead City Carteret
White Oak Onslow 44b 15-VI River, right bank, near Swansboro, bridge on highway 24
45 15-VI Holland Mill Creek, tributary, N.W. of Swansboro Onslow
Oak 46 15-VI White River, near Stella Onslow/Carteret
Carteret *47 20-VI North River, N.E. of Beaufort, bridge on highway 70
*48 20-VI North River, near Otway, bridge on highway 70 Carteret
*49 20-VI Core Sound, near Smyrna Carteret
*50 20-VI Core Sound, near bridge on highway 70, midway Williston and Davis Carteret
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Station Date Position County
(1977)
51 20-VI water in between Cedar Island and the mainland, connecting Long Bay and Carteret
Core Sound
Neuse left "Minnesott Pamlico 52a 13-VII River, bank, near Arapahoe, Bennett Road, Manor"
52b 13-VII small stream discharging in Neuse River near station 52a Pamlico
Beard U.S. 53 13-VII Creek, discharging in Neuse River, across Cherry Point Naval Pamlico
Reservation
54 13-VII Goose Creek, Reelsboro Fire district, S.E. of New Bern Pamlico
55 13-VII Upper Broad Creek, Tricommunity Fire district Pamlico
*56 13-VII Neuse River, right bank, upstream of New Bern, "Gap Landing" Craven
*57 3-VIII Neuse River, near Askin, N.W. of New Bern Craven
58 9-VIII Neuse River, left bank, midway Minnesott and Janeiro, downstream of Camp Pamlico
Don Lee
59 9-VIII Neuse River, left bank, near Janeiro, bridge of tributary Pamlico
Robert 60 9-VIII Neuse River, left bank, near Oriental, Scott bridge Pamlico
*61 9-VIII Bay River, near Stonewall, bridge on highway 55 Pamlico
*62 9-VIII Bay River, left bank, near Vandemere, N.E. of Bayboro, highway 304 Pamlico
63 11-VIII Newport River, upstream of bridge on highway 70, Morehead City Carteret
64 22-VIII South River, near village South River, downstream of jetty of D.U.M.L. boat Carteret
Received: 8 March 1978
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